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Protein domain
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Pyruvate kinase, a protein from three domains (PDB 1pkn)
A protein domain is a part of protein sequence and structure that can evolve, function, and exist
independently of the rest of the protein chain. Each domain forms a compact three-dimensional structure and
often can be independently stable and folded. Many proteins consist of several structural domains. One
domain may appear in a variety of different proteins. Molecular evolution uses domains as building blocks
and these may be recombined in different arrangements to create proteins with different functions. Domains
vary in length from between about 25 amino acids up to 500 amino acids in length. The shortest domains
such as zinc fingers are stabilized by metal ions or disulfide bridges. Domains often form functional units,
such as the calcium-binding EF hand domain of calmodulin. Because they are independently stable, domains
can be "swapped" by genetic engineering between one protein and another to make chimeric proteins.
• 1 Background
• 2 Domains are units of protein structure
• 2.1 Tertiary structure of domains
• 2.2 Domains have limits on size
• 2.3 Domains and quaternary structure
• 3 Domains as evolutionary modules
• 4 Multidomain proteins
• 4.1 Origin
• 4.2 Types of organisation
• 5 Domains are autonomous folding units
• 5.1 Folding
• 5.2 Advantage of domains in protein folding
• 6 Domains and protein flexibility
• 6.1 Hinges by secondary structures
• 6.2 Helical to extended conformation
• 6.3 Shear motions
• 6.4 Domain motion and functional dynamics in
• 7 Domain definition from structural co-ordinates
• 7.1 Methods
[edit] Background
The concept of the domain was first proposed in 1973 by Wetlaufer after X-ray crystallographic
studies of hen lysozyme [1] and papain [2] and by limited proteolysis studies of immunoglobulins [3][4].
Wetlaufer defined domains as stable units of protein structure that could fold autonomously. In the past
domains have been described as units of:
• compact structure[5]
• function and evolution[6]
• folding [7].
Each definition is valid and will often overlap, i.e. a compact structural domain that is found amongst
diverse proteins is likely to fold independently within its structural environment. Nature often brings several
domains together to form multidomain and multifunctional proteins with a vast number of possibilities [8]. In a
multidomain protein, each domain may fulfil its own function independently, or in a concerted manner with its
neighbours. Domains can either serve as modules for building up large assemblies such as virus particles or
muscle fibres, or can provide specific catalytic or binding sites as found in enzymes or regulatory proteins.
An appropriate example is pyruvate kinase, a glycolytic enzyme that plays an important role in
regulating the flux from fructose-1,6-biphosphate to pyruvate. It contains an all-β regulatory domain, an α/β-
substrate binding domain and an α/β-nucleotide binding domain, connected by several polypeptide linkers [9]
(see figure, right). Each domain in this protein occurs in diverse sets of protein families.
The central α/β-barrel substrate binding domain is one of the most common enzyme folds. It is seen
in many different enzyme families catalysing completely unrelated reactions[10]. The α/β-barrel is commonly
called the TIM barrel named after triose phosphate isomerase, which was the first such structure to be
solved[11]. It is currently classified into 26 homologous families in the CATH domain database [12]. The TIM
barrel is formed from a sequence of β-α-β motifs closed by the first and last strand hydrogen bonding
together, forming an eight stranded barrel. There is debate about the evolutionary origin of this domain. One
study has suggested that a single ancestral enzyme could have diverged into several families[13], while
another suggests that a stable TIM-barrel structure has evolved through convergent evolution [14].
The TIM-barrel in pyruvate kinase is 'discontinuous', meaning that more than one segment of the
polypeptide is required to form the domain. This is likely to be the result of the insertion of one domain into
another during the protein's evolution. It has been shown from known structures that about a quarter of
structural domains are discontinuous.[15][16] The inserted β-barrel regulatory domain is 'continuous', made
up of a single stretch of polypeptide.
Covalent association of two domains represents a functional and structural advantage since there is
an increase in stability when compared with the same structures non-covalently associated [17]. Other,
advantages are the protection of intermediates within inter-domain enzymatic clefts that may otherwise be
unstable in aqueous environments, and a fixed stoichiometric ratio of the enzymatic activity necessary for a
sequential set of reactions [18].

[edit] Domains are units of protein structure

Main article: Protein structure
The primary structure (string of amino acids) of a protein ultimately encodes its uniquely folded 3D
conformation.[19] The most important factor governing the folding of a protein into 3D structure is the
distribution of polar and non-polar side chains.[20] Folding is driven by the burial of hydrophobic side chains
into the interior of the molecule so to avoid contact with the aqueous environment. Generally proteins have a
core of hydrophobic residues surrounded by a shell of hydrophilic residues. Since the peptide bonds
themselves are polar they are neutralised by hydrogen bonding with each other when in the hydrophobic
environment. This gives rise to regions of the polypeptide that form regular 3D structural patterns called
secondary structure. There are two main types of secondary structure: α-helices and β-sheets.
Some simple combinations of secondary structure elements have been found to frequently occur in
protein structure and are referred to as supersecondary structure or motifs. For example, the β-hairpin motif
consists of two adjacent antiparallel β-strands joined by a small loop. It is present in most antiparallel β
structures both as an isolated ribbon and as part of more complex β-sheets. Another common super-
secondary structure is the β-α-β motif, which is frequently used to connect two parallel β-strands. The central
α-helix connects the C-termini of the first strand to the N-termini of the second strand, packing its side chains
against the β-sheet and therefore shielding the hydrophobic residues of the β-strands from the surface.
Structural alignment is an important tool for determining domains.

[edit] Tertiary structure of domains

Several motifs pack together to form compact, local, semi-independent units called domains.[5] The
overall 3D structure of the polypeptide chain is referred to as the protein's tertiary structure. Domains are the
fundamental units of tertiary structure, each domain containing an individual hydrophobic core built from
secondary structural units connected by loop regions. The packing of the polypeptide is usually much tighter
in the interior than the exterior of the domain producing a solid-like core and a fluid-like surface.[21] In fact,
core residues are often conserved in a protein family, whereas the residues in loops are less conserved,
unless they are involved in the protein's function. Protein tertiary structure can be divided into four main
classes based on the secondary structural content of the domain.[22]
• All-α domains have a domain core built exclusively from α-helices. This class is dominated
by small folds, many of which form a simple bundle with helices running up and down.
• All-β domains have a core comprising of antiparallel β-sheets, usually two sheets packed
against each other. Various patterns can be identified in the arrangement of the strands, often giving
rise to the identification of recurring motifs, for example the Greek key motif.[23]
• α+β domains are a mixture of all-α and all-β motifs. Classification of proteins into this class is
difficult because of overlaps to the other three classes and therefore is not used in the CATH domain
• α/β domains are made from a combination of β-α-β motifs that predominantly form a parallel
β-sheet surrounded by amphipathic α-helices. The secondary structures are arranged in layers or

[edit] Domains have limits on size

Domains have limits on size.[24] The size of individual structural domains varies from 36 residues in
E-selectin to 692 residues in lipoxygenase-1,[15] but the majority, 90%, have less than 200 residues[25] with
an average of approximately 100 residues.[26] Very short domains, less than 40 residues, are often
stabilised by metal ions or disulfide bonds. Larger domains, greater than 300 residues, are likely to consist of
multiple hydrophobic cores.[27]

[edit] Domains and quaternary structure

Many proteins have a quaternary structure, which consists of several polypeptide chains that
associate into an oligomeric molecule. Each polypeptide chain in such a protein is called a subunit.
Hemoglobin, for example, consists of two α and two β subunits. Each of the four chains has an all-α globin
fold with a heme pocket.
Domain swapping is a mechanism for forming oligomeric assemblies.[28] In domain swapping, a
secondary or tertiary element of a monomeric protein is replaced by the same element of another protein.
Domain swapping can range from secondary structure elements to whole structural domains. It also
represents a model of evolution for functional adaptation by oligomerisation, e.g. oligomeric enzymes that
have their active site at subunit interfaces.[29]

[edit] Domains as evolutionary modules

Nature is a tinkerer and not an inventor,[30] new sequences are adapted from pre-existing
sequences rather than invented. Domains are the common material used by nature to generate new
sequences, they can be thought of as genetically mobile units, referred to as 'modules'. Often, the C and N
termini of domains are close together in space, allowing them to easily be "slotted into" parent structures
during the process of evolution. Many domain families are found in all three forms of life, Archaea, Bacteria
and Eukarya. Domains that are repeatedly found in diverse proteins are often referred to as modules,
examples can be found among extracellular proteins associated with clotting, fibrinolysis, complement, the
extracellular matrix, cell surface adhesion molecules and cytokine receptors.[31]
Molecular evolution gives rise to families of related proteins with similar sequence and structure.
However, sequence similarities can be extremely low between proteins that share the same structure.
Protein structures may be similar because proteins have diverged from a common ancestor. Alternatively,
some folds may be more favored than others as they represent stable arrangements of secondary structures
and some proteins may converge towards these folds over the course of evolution . There are currently about
45,000 experimentally determined protein 3D structures deposited within the Protein Data Bank (PDB).[32]
However this set contains a lot of identical or very similar structures. All proteins should be classified to
structural families to understand their evolutionary relationships. Structural comparisons are best achieved at
the domain level. For this reason many algorithms have been developed to automatically assign domains in
proteins with known 3D structure, see 'Domain definition from structural co-ordinates'.
The CATH domain database classifies domains into approximately 800 fold families, ten of these
folds are highly populated and are referred to as 'super-folds'. Super-folds are defined as folds for which
there are at least three structures without significant sequence similarity.[33] The most populated is the α/β-
barrel super-fold as described previously.

[edit] Multidomain proteins

The majority of genomic proteins, two-thirds in unicellular organisms and more than 80% in metazoa,
are multidomain proteins created as a result of gene duplication events.[34] Many domains in multidomain
structures could have once existed as independent proteins. More and more domains in eukaryotic
multidomain proteins can be found as independent proteins in prokaryotes.[35] For example, vertebrates
have a multi-enzyme polypeptide containing the GAR synthetase, AIR synthetase and GAR transformylase
modules (GARs-AIRs-GARt; GAR: glycinamide ribonucleotide synthetase/transferase; AIR: aminoimidazole
ribonucleotide synthetase). In insects, the polypeptide appears as GARs-(AIRs)2-GARt, in yeast GARs-AIRs
is encoded separately from GARt, and in bacteria each domain is encoded separately.[36]

[edit] Origin
Multidomain proteins are likely to have emerged from a selective pressure during evolution to create
new functions. Various proteins have diverged from common ancestors by different combinations and
associations of domains. Modular units frequently move about, within and between biological systems
through mechanisms of genetic shuffling:
• transposition of mobile elements including horizontal transfers (between species);[37]
• gross rearrangements such as inversions, translocations, deletions and duplications;
• homologous recombination;
• slippage of DNA polymerase during replication.

[edit] Types of organisation

The simplest multidomain organisation seen in proteins is that of a single domain repeated in
tandem.[38] The domains may interact with each other or remain isolated, like beads on string. The giant
30,000 residue muscle protein titin comprises about 120 fibronectin-III-type and Ig-type domains.[39] In the
serine proteases, a gene duplication event has led to the formation of a two β-barrel domain enzyme.[40] The
repeats have diverged so widely that there is no obvious sequence similarity between them. The active site is
located at a cleft between the two β-barrel domains, in which functionally important residues are contributed
from each domain. Genetically engineered mutants of the chymotrypsin serine protease were shown to have
some proteinase activity even though their active site residues were abolished and it has therefore been
postulated that the duplication event enhanced the enzyme's activity.[40]
Modules frequently display different connectivity relationships, as illustrated by the kinesins and ABC
transporters. The kinesin motor domain can be at either end of a polypeptide chain that includes a coiled-coil
region and a cargo domain.[41] ABC transporters are built with up to four domains consisting of two
unrelated modules, ATP-binding cassette and an integral membrane module, arranged in various
Not only do domains recombine, but there are many examples of a domain having been inserted into
another. Sequence or structural similarities to other domains demonstrate that homologues of inserted and
parent domains can exist independently. An example is that of the 'fingers' inserted into the 'palm' domain
within the polymerases of the Pol I family.[42] Since a domain can be inserted into another, there should
always be at least one continuous domain in a multidomain protein. This is the main difference between
definitions of structural domains and evolutionary/functional domains. An evolutionary domain will be limited
to one or two connections between domains, whereas structural domains can have unlimited connections,
within a given criterion of the existence of a common core. Several structural domains could be assigned to
an evolutionary domain.

[edit] Domains are autonomous folding units

[edit] Folding
Further information: Protein folding
Protein folding - the unsolved problem : Since the seminal work of Anfinsen over forty years ago,[19]
the goal to completely understand the mechanism by which a polypeptide rapidly folds into its stable native
conformation remains elusive. Many experimental folding studies have contributed much to our
understanding, but the principles that govern protein folding are still based on those discovered in the very
first studies of folding. Anfinsen showed that the native state of a protein is thermodynamically stable, the
conformation being at a global minimum of its free energy.
Folding is a directed search of conformational space allowing the protein to fold on a biologically
feasible time scale. The Levinthal paradox states that if an averaged sized protein would sample all possible
conformations before finding the one with the lowest energy, the whole process would take billions of years.
[43] Proteins typically fold within 0.1 and 1000 seconds, therefore the protein folding process must be
directed some way through a specific folding pathway. The forces that direct this search are likely to be a
combination of local and global influences whose effects are felt at various stages of the reaction.[44]
Advances in experimental and theoretical studies have shown that folding can be viewed in terms of
energy landscapes,[45][46] where folding kinetics is considered as a progressive organisation of an
ensemble of partially folded structures through which a protein passes on its way to the folded structure. This
has been described in terms of a folding funnel, in which an unfolded protein has a large number of
conformational states available and there are fewer states available to the folded protein. A funnel implies
that for protein folding there is a decrease in energy and loss of entropy with increasing tertiary structure
formation. The local roughness of the funnel reflects kinetic traps, corresponding to the accumulation of
misfolded intermediates. A folding chain progresses toward lower intra-chain free-energies by increasing its
compactness. The chains conformational options become increasingly narrowed ultimately toward one native

[edit] Advantage of domains in protein folding

The organisation of large proteins by structural domains represents an advantage for protein folding,
with each domain being able to individually fold, accelerating the folding process and reducing a potentially
large combination of residue interactions. Furthermore, given the observed random distribution of
hydrophobic residues in proteins,[47] domain formation appears to be the optimal solution for a large protein
to bury its hydrophobic residues while keeping the hydrophilic residues at the surface.[48][49]
However, the role of inter-domain interactions in protein folding and in energetics of stabilisation of
the native structure, probably differs for each protein. In T4 lysozyme, the influence of one domain on the
other is so strong that the entire molecule is resistant to proteolytic cleavage. In this case, folding is a
sequential process where the C-terminal domain is required to fold independently in an early step, and the
other domain requires the presence of the folded C-terminal domain for folding and stabilisation.[50]
It has been found that the folding of an isolated domain can take place at the same rate or
sometimes faster than that of the integrated domain.[51] Suggesting that unfavourable interactions with the
rest of the protein can occur during folding. Several arguments suggest that the slowest step in the folding of
large proteins is the pairing of the folded domains.[27] This is either because the domains are not folded
entirely correctly or because the small adjustments required for their interaction are energetically
unfavourable,[52] such as the removal of water from the domain interface.

[edit] Domains and protein flexibility

The presence of multiple domains in proteins gives rise to a great deal of flexibility and mobility,
leading to protein domain dynamics. Domain motions can be inferred by comparing different structures of a
protein[citation needed], or they can be directly observed using spectra[53][54] measured by neutron spin
echo spectroscopy. They can also be suggested by sampling in extensive molecular dynamics
trajectories[55]. Domain motions are important for:[56]
• catalysis;
• regulatory activity;
• transport of metabolites;
• formation of protein assemblies; and
• cellular locomotion.
One of the largest observed domain motions is the `swivelling' mechanism in pyruvate phosphate
dikinase. The phosphoinositide domain swivels between two states in order to bring a phosphate group from
the active site of the nucleotide binding domain to that of the phosphoenolpyruvate/pyruvate domain.[57] The
phosphate group is moved over a distance of 45A involving a domain motion of about 100 degrees around a
single residue.In enzymes, the closure of one domain onto another captures a substrate by an induced fit,
allowing the reaction to take place in a controlled way. A detailed analysis by Gerstein led to the classification
of two basic types of domain motion; hinge and shear.[56] Only a relatively small portion of the chain, namely
the inter-domain linker and side chains undergo significant conformational changes upon domain

[edit] Hinges by secondary structures

A study by Hayward[59] found that the termini of α-helices and β-sheets form hinges in a large
number of cases. Many hinges were found to involve two secondary structure elements acting like hinges of
a door, allowing an opening and closing motion to occur. This can arise when two neighbouring strands
within a β-sheet situated in one domain, diverge apart as they join the other domain. The two resulting termini
then form the bending regions between the two domains. α-helices that preserve their hydrogen bonding
network when bent are found to behave as mechanical hinges, storing `elastic energy' that drives the closure
of domains for rapid capture of a substrate.[59]

[edit] Helical to extended conformation

The interconversion of helical and extended conformations at the site of a domain boundary is not
uncommon. In calmodulin, torsion angles change for five residues in the middle of a domain linking α-helix.
The helix is split into two, almost perpendicular, smaller helices separated by four residues of an extended

[edit] Shear motions

Shear motions involve a small sliding movement of domain interfaces, controlled by the amino acid
side chains within the interface. Proteins displaying shear motions often have a layered architecture: stacking
of secondary structures. The interdomain linker has merely the role of keeping the domains in close

[edit] Domain motion and functional dynamics in enzymes

The analysis of the internal dynamics of structurally different, but functionally similar enzymes has
highlighted a common relationship between the positioning of the active site and the two principal protein
sub-domains. In fact, for several members of the hydrolase superfamily, the catalytic site is located close to
the interface separating the two principal quasi-rigid domains[55]. Such positioning appears instrumental for
maintaining the precise geometry of the active site, while allowing for an appreciable functionally-oriented
modulation of the flanking regions resulting from the relative motion of the two sub-domains.

[edit] Domain definition from structural co-ordinates

The importance of domains as structural building blocks and elements of evolution has brought about
many automated methods for their identification and classification in proteins of known structure. Automatic
procedures for reliable domain assignment is essential for the generation of the domain databases,
especially as the number of protein structures is increasing. Although the boundaries of a domain can be
determined by visual inspection, construction of an automated method is not straightforward. Problems occur
when faced with domains that are discontinuous or highly associated.[62] The fact that there is no standard
definition of what a domain really is has meant that domain assignments have varied enormously, with each
researcher using a unique set of criteria.[63]
A structural domain is a compact, globular sub-structure with more interactions within it than with the
rest of the protein.[58] Therefore, a structural domain can be determined by two visual characteristics; its
compactness and its extent of isolation.[64] Measures of local compactness in proteins have been used in
many of the early methods of domain assignment[65][66][67][68] and in several of the more recent methods.

[edit] Methods
One of the first algorithms[65] used a Cα-Cα distance map together with a hierarchical clustering
routine that considered proteins as several small segments, 10 residues in length. The initial segments were
clustered one after another based on inter-segment distances; segments with the shortest distances were
clustered and considered as single segments thereafter. The stepwise clustering finally included the full
protein. Go[68] also exploited the fact that inter-domain distances are normally larger than intra-domain
distances; all possible Cα-Cα distances were represented as diagonal plots in which there were distinct
patterns for helices, extended strands and combinations of secondary structures.
The method by Sowdhamini and Blundell clusters secondary structures in a protein based on their
Cα-Cα distances and identifies domains from the pattern in their dendrograms.[62] As the procedure does
not consider the protein as a continuous chain of amino acids there are no problems in treating discontinuous
domains. Specific nodes in these dendrograms are identified as tertiary structural clusters of the protein,
these include both super-secondary structures and domains. The DOMAK algorithm is used to create the
3Dee domain database.[70] It calculates a 'split value' from the number of each type of contact when the
protein is divided arbitrarily into two parts. This split value is large when the two parts of the structure are
The method of Wodak and Janin[73] was based on the calculated interface areas between two chain
segments repeatedly cleaved at various residue positions. Interface areas were calculated by comparing
surface areas of the cleaved segments with that of the native structure. Potential domain boundaries can be
identified at a site where the interface area was at a minimum. Other methods have used measures of
solvent accessibility to calculate compactness.[25][74][75]
The PUU algorithm[16] incorporates a harmonic model used to approximate inter-domain dynamics.
The underlying physical concept is that many rigid interactions will occur within each domain and loose
interactions will occur between domains. This algorithm is used to define domains in the FSSP domain
Swindells (1995) developed a method, DETECTIVE, for identification of domains in protein structures
based on the idea that domains have a hydrophobic interior. Deficiencies were found to occur when
hydrophobic cores from different domains continue through the interface region.
RigidFinder is a novel method for identification of protein rigid blocks (domains and loops) from two
different conformations. Rigid blocks are defined as blocks where all inter residue distances are conserved
across conformations.
A general method to identify dynamical domains, that is protein regions that behave approximately as
rigid units in the course of structural fluctuations, has been introduced by Potestio et al.[55] and, among other
applications was also used to compare the consistency of the dynamics-based domain subdivisions with
standard structure-based ones. The method, termed PiSQRD, is publicly available in the form of a
webserver[76]. The latter allows users to optimally subdivide single-chain or multimeric proteins into quasi-
rigid domains[55][76] based on the collective modes of fluctuation of the system. By default the latter are
calculated through an elastic network model[77]; alternatively pre-calculated essential dynamical spaces can
be uploaded by the user.
This article incorporates text and figures from George, R. A. (2002) "Predicting Structural Domains in
Proteins" Thesis, University College London, which were contributed by its author.
[edit] See also
• Amino acid
• Binding domain
• Motif domain
• Eukaryotic Linear Motif
• Protein
• Protein structure
• Protein structure prediction
• Protein structure prediction software
• Protein family
• Structural biology
• Structural Classification of Proteins (SCOP)

[edit] Example domains

• Armadillo repeats : named after the β-catenin-like Armadillo protein of the fruit fly Drosophila.
• Basic Leucine zipper domain (bZIP domain) : is found in many DNA-binding eukaryotic
proteins. One part of the domain contains a region that mediates sequence-specific DNA-binding
properties and the Leucine zipper that is required for the dimerization of two DNA-binding regions.
The DNA-binding region comprises a number of basic aminoacids such as arginine and lysine
• Cadherin repeats : Cadherins function as Ca2+-dependent cell-cell adhesion proteins.
Cadherin domains are extracellular regions which mediate cell-to-cell homophilic binding between
cadherins on the surface of adjacent cells.
• Death effector domain (DED) : allows protein-protein binding by homotypic interactions
(DED-DED). Caspase proteases trigger apoptosis via proteolytic cascades. Pro-Caspase-8 and pro-
caspase-9 bind to specific adaptor molecules via DED domains and this leads to autoactivation of
• EF hand : a helix-turn-helix structural motif found in each structural domain of the signaling
protein calmodulin and in the muscle protein troponin-C.
• Immunoglobulin-like domains : are found in proteins of the immunoglobulin superfamily
(IgSF).[78] They contain about 70-110 amino acids and are classified into different categories (IgV,
IgC1, IgC2 and IgI) according to their size and function. They possess a characteristic fold in which
two beta sheets form a "sandwich" that is stabilized by interactions between conserved cysteines and
other charged amino acids. They are important for protein-to-protein interactions in processes of cell
adhesion, cell activation, and molecular recognition. These domains are commonly found in
molecules with roles in the immune system.
• Phosphotyrosine-binding domain (PTB) : PTB domains usually bind to phosphorylated
tyrosine residues. They are often found in signal transduction proteins. PTB-domain binding
specificity is determined by residues to the amino-terminal side of the phosphotyrosine. Examples:
the PTB domains of both SHC and IRS-1 bind to a NPXpY sequence. PTB-containing proteins such
as SHC and IRS-1 are important for insulin responses of human cells.
• Pleckstrin homology domain (PH) : PH domains bind phosphoinositides with high affinity.
Specificity for PtdIns(3)P, PtdIns(4)P, PtdIns(3,4)P2, PtdIns(4,5)P2, and PtdIns(3,4,5)P3 have all
been observed. Given the fact that phosphoinositides are sequestered to various cell membranes
(due to their long lipophilic tail) the PH domains usually causes recruitment of the protein in question
to a membrane where the protein can exert a certain function in cell signalling, cytoskeletal
reorganization or membrane trafficking.
• Src homology 2 domain (SH2) : SH2 domains are often found in signal transduction proteins.
SH2 domains confer binding to phosphorylated tyrosine (pTyr). Named after the phosphotyrosine
binding domain of the src viral oncogene, which is itself a tyrosine kinase. See also: SH3 domain.
• Zinc finger DNA binding domain (ZnF_GATA) : ZnF_GATA domain-containing proteins are
typically transcription factors that usually bind to the DNA sequence [AT]GATA[AG] of promoters.

[edit] References
1. ^ Phillips DC. (1966). "The three-dimensional structure of an enzyme molecule". Scientific
American 215 (5): 78–90. doi:10.1038/255609a0. PMID 5978599.
2. ^ Drenth J, Jansonius JN, Koekoek R, Swen HM, Wolthers BG. (1968). "Structure of papain".
Nature 218 (5145): 929–32. doi:10.1038/218929a0. PMID 5681232.
3. ^ Porter RR. (1973). "Structural studies of immunoglobulins". Science 180 (87): 713–6.
doi:10.1126/science.180.4087.713. PMID 4122075.
4. ^ Edelman GM. (1973). "Antibody structure and molecular immunology". Science 180 (88):
830–40. doi:10.1126/science.180.4088.830. PMID 4540988.
5. ^ a b Richardson J. S. (1981). "The anatomy and taxonomy of protein structure". Adv Protein
Chem 34: 167–339. doi:10.1038/255609a0. PMID 7020376.
6. ^ Bork P. (1991). "Shuffled domains in extracellular proteins". FEBS Lett 286 (1-2): 47–54.
doi:10.1016/0014-5793(91)80937-X. PMID 1864378.
7. ^ Wetlaufer DB. (1973). "Nucleation, rapid folding, and globular intrachain regions in
proteins". Proc Natl Acad Sci USA 70 (3): 697–701. doi:10.1073/pnas.70.3.697. PMID 4351801.
8. ^ Chothia C. (1992). "Proteins. One thousand families for the molecular biologist". Nature
357 (6379): 543–4. doi:10.1038/357543a0. PMID 1608464.
9. ^ George RA, Heringa J. (2002). "An analysis of protein domain linkers: their classification
and role in protein folding". Protein Eng 15 (11): 871–9. doi:10.1093/protein/15.11.871.
PMID 12538906.
10.^ Hegyi H, and Gerstein M. (1999). "The relationship between protein structure and function:
a comprehensive survey with application to the yeast genome". J Mol Biol 288 (1): 147–64.
doi:10.1006/jmbi.1999.2661. PMID 10329133.
11.^ Banner et al.; Bloomer, AC; Petsko, GA; Phillips, DC; Pogson, CI; Wilson, IA; Corran, PH;
Furth, AJ et al. (1975). "Structure of chicken muscle triose phosphate isomerase determined
crystallographically at 2.5 angstrom resolution using amino acid sequence data". Nature 255 (5510):
609–614. doi:10.1038/255609a0. PMID 1134550.
12.^ a b Orengo CA, Michie AD, Jones S, Jones DT, Swindells MB, Thornton JM. (1997).
"CATH--a hierarchic classification of protein domain structures". Structure 5 (8): 1093–108.
doi:10.1016/S0969-2126(97)00260-8. PMID 9309224.
13.^ Copley, R. R. and Bork, P (2000). "Homology among (betaalpha)(8) barrels: implications
for the evolution of metabolic pathways". J Mol Biol 303 (4): 627–641. doi:10.1006/jmbi.2000.4152.
PMID 11054297.
14.^ Lesk AM, Brändén CI, Chothia C. (1989). "Structural principles of alpha/beta barrel
proteins: the packing of the interior of the sheet". Proteins 5 (2): 139–48. doi:10.1002/prot.340050208.
PMID 2664768.
15.^ a b Jones S, Stewart M, Michie A, Swindells MB, Orengo C, Thornton JM. (1998). "Domain
assignment for protein structures using a consensus approach: characterization and analysis".
Protein Sci 7 (2): 233–42. doi:10.1002/pro.5560070202. PMID 9521098. PMC 2143930.
16.^ a b Holm L, Sander C. (1994). "Parser for protein folding units". Proteins 19 (3): 256–68.
doi:10.1002/prot.340190309. PMID 7937738.
17.^ Ghélis C, Yon JM. (1979). "[Conformational coupling between structural units. A decisive
step in the functional structure formation]". C R Seances Acad Sci D 289 (2): 197–9. PMID 117925.
18.^ Ostermeier M, Benkovic SJ. (2000). "Evolution of protein function by domain swapping".
Adv Protein Chem 55: 29–77. PMID 11050932.
19.^ a b ANFINSEN CB, HABER E, SELA M, WHITE FH Jr. (1961). "The kinetics of formation of
native ribonuclease during oxidation of the reduced polypeptide chain". Proc Natl Acad Sci USA 47
(9): 1309–14. doi:10.1073/pnas.47.9.1309. PMID 13683522.
20.^ Cordes, M. H., Davidson, A. R., and Sauer, R. T (1996). "Sequence space, folding and
protein design". Curr Opin Struct Biol 6 (1): 3–10. doi:10.1016/S0959-440X(96)80088-1.
PMID 8696970.
21.^ Zhou, Y., Vitkup, D., and Karplus, M (1999). "Native proteins are surface-molten solids:
application of the Lindemann criterion for the solid versus liquid state". J Mol Biol 285 (4): 1371–1375.
doi:10.1006/jmbi.1998.2374. PMID 9917381.
22.^ Levitt M, Chothia C. (1976). "Structural patterns in globular proteins". Nature 261 (5561):
552–8. doi:10.1038/261552a0. PMID 934293.
23.^ Hutchinson EG, Thornton JM. (1993). "The Greek key motif: extraction, classification and
analysis". Protein Eng 6 (3): 233–45. doi:10.1093/protein/6.3.233. PMID 8506258.
24.^ Savageau MA. (1986). "Proteins of Escherichia coli come in sizes that are multiples of 14
kDa: domain concepts and evolutionary implications". Proc Natl Acad Sci USA 83 (5): 1198–202.
doi:10.1073/pnas.83.5.1198. PMID 3513170. PMC 323042.
25.^ a b c Islam SA, Luo J, Sternberg MJ. (1995). "Identification and analysis of domains in
proteins". Protein Eng 8 (6): 513–25. doi:10.1093/protein/8.6.513. PMID 8532675.
26.^ Wheelan, S. J. and Marchler-Bauer, A. and Bryant, S. H. (2000). "Domain size distributions
can predict domain boundaries". Bioinformatics 16 (7): 613–618. doi:10.1093/bioinformatics/16.7.613.
PMID 11038331.
27.^ a b Garel, J. (1992). "Folding of large proteins: Multidomain and multisubunit proteins". In
Creighton, T., editor, Protein Folding, pages 405-454. W.H. Freeman and Company, New York, first
28.^ Bennett MJ, Schlunegger MP, Eisenberg D. (1995). "3D domain swapping: a mechanism
for oligomer assembly". Protein Sci 4 (12): 2455–68. doi:10.1002/pro.5560041202. PMID 8580836.
PMC 2143041.
29.^ Heringa J, Taylor WR. (1997). "Three-dimensional domain duplication, swapping and
stealing". Curr Opin Struct Biol 7 (3): 416–21. doi:10.1016/S0959-440X(97)80060-7. PMID 9204285.
30.^ Jacob F. (1977). "Evolution and tinkering". Science 196 (4295): 1161–6.
doi:10.1126/science.860134. PMID 860134.
31.^ Campbell ID, Downing AK. (1994). "Building protein structure and function from modular
units". Trends Biotechnol 12 (5): 168–72. doi:10.1016/0167-7799(94)90078-7. PMID 7764899.
33.^ Orengo CA, Jones DT, Thornton JM. (1994). "Protein superfamilies and domain
superfolds". Nature 372 (6507): 631–4. doi:10.1038/372631a0. PMID 7990952.
34.^ Apic, G., Gough, J., and Teichmann, S. A (2001). "Domain combinations in archaeal,
eubacterial and eukaryotic proteomes". J Mol Biol 310 (2): 311–325. doi:10.1006/jmbi.2001.4776.
PMID 11428892.
35.^ Davidson JN, Chen KC, Jamison RS, Musmanno LA, Kern CB. (1993). "The evolutionary
history of the first three enzymes in pyrimidine biosynthesis". Bioessays 15 (3): 157–64.
doi:10.1002/bies.950150303. PMID 8098212.
36.^ Henikoff S, Greene EA, Pietrokovski S, Bork P, Attwood TK, Hood L. (1997). "Gene
families: the taxonomy of protein paralogs and chimeras". Science 278 (5338): 609–14.
doi:10.1126/science.278.5338.609. PMID 9381171.
37.^ Bork, P. and Doolittle, R. F (1992). "Proposed acquisition of an animal protein domain by
bacteria2". Proc Natl Acad Sci USA 89 (19): 8990–8994. doi:10.1073/pnas.89.19.8990.
PMID 1409594.
38.^ Heringa J. (1998). "Detection of internal repeats: how common are they?". Curr Opin Struct
Biol 8 (3): 338–45. doi:10.1016/S0959-440X(98)80068-7. PMID 9666330.
39.^ Politou, A. S., Gautel, M., Improta, S., Vangelista, L., and Pastore, A (1996). "The elastic I-
band region of titin is assembled in a 'modular' fashion by weakly interacting Ig-like domains". J Mol
Biol 255 (4): 604–616. doi:10.1006/jmbi.1996.0050. PMID 8568900.
40.^ a b McLachlan, A. D (1979). "Gene duplications in the structural evolution of chymotrypsin".
J Mol Biol 128 (1): 49–79. doi:10.1016/0022-2836(79)90308-5. PMID 430571.
41.^ Moore JD, Endow SA. (1996). "Kinesin proteins: a phylum of motors for microtubule-based
motility". Bioessays 18 (3): 207–19. doi:10.1002/bies.950180308. PMID 8867735.
42.^ Russell, R. B (1994). "Domain insertion". Protein Eng 7 (12): 1407–1410.
doi:10.1093/protein/7.12.1407. PMID 7716150.
43.^ Levinthal, C. (1968). "Are there pathways for protein folding?". J Chim Phys 65: 44–45.
44.^ Dill KA. (1999). "Polymer principles and protein folding". Protein Sci 8 (6): 1166–80.
doi:10.1110/ps.8.6.1166. PMID 10386867. PMC 2144345.
45.^ Leopold PE, Montal M, Onuchic JN. (1992). "Protein folding funnels: a kinetic approach to
the sequence-structure relationship". Proc Natl Acad Sci USA 89 (18): 8721–5.
doi:10.1073/pnas.89.18.8721. PMID 1528885. PMC 49992.
46.^ Dill KA, Chan HS. (1997). "From Levinthal to pathways to funnels". Nat Struct Biol 4 (1): 10–
9. doi:10.1038/nsb0197-10. PMID 8989315.
47.^ White SH, Jacobs RE. (1990). "Statistical distribution of hydrophobic residues along the
length of protein chains. Implications for protein folding and evolution". Biophys J 57 (4): 911–21.
doi:10.1016/S0006-3495(90)82611-4. PMID 2188687. PMC 1280792.
48.^ George RA, Heringa J. (2002). "SnapDRAGON: a method to delineate protein structural
domains from sequence data". J Mol Biol 316 (3): 839–51. doi:10.1006/jmbi.2001.5387.
PMID 11866536.
49.^ George RA, Lin K, Heringa J. (2005). "Scooby-domain: prediction of globular domains in
protein sequence". Nucleic Acids Res 33 (Web Server issue): W160–3. doi:10.1093/nar/gki381.
PMID 15980446. PMC 1160142.
50.^ Desmadril, M. and Yon, J. M (1981). "Existence of intermediates in the refolding of T4
lysozyme at pH 7.4". Biochem Biophys Res Commun 101 (2): 563–569. doi:10.1016/0006-
291X(81)91296-1. PMID 7306096.
51.^ Teale JM, Benjamin DC. (1977). "Antibody as immunological probe for studying refolding of
bovine serum albumin. Refolding within each domain". J Biol Chem 252 (13): 4521–6. PMID 873903.
52.^ Creighton, T. E. (1983). Proteins: Structures and molecular properties . Freeman, New
York. Second edition.
53.^ Farago B, Li J, Cornilescu G, Callaway DJ, Bu Z (November 2010). "Activation of
Nanoscale Allosteric Protein Domain Motion Revealed by Neutron Spin Echo Spectroscopy".
Biophysical Journal 99 (10): 3473–3482. doi:10.1016/j.bpj.2010.09.058. PMID 21081097.
54.^ Bu Z, Biehl R, Monkenbusch M, Richter D, D.J.E. Callaway (2005). "Coupled protein
domain motion in Taq polymerase revealed by neutron spin-echo spectroscopy.". Proc Natl Acad Sci
USA 102 (49): 17646–17651. doi:10.1073/pnas.0503388102. PMID 16306270.
55.^ a b c d Potestio, R., Pontiggia, F. and Micheletti, C. (2009). "Coarse-grained description of
protein internal dynamics: an optimal strategy for decomposing proteins in rigid subunits.".
Biophysical Journal 96 (12): 4993–5002. doi:10.1016/j.bpj.2009.03.051. PMID 19527659.
PMC 2712024.
56.^ a b Gerstein M, Lesk AM, Chothia C. (1994). "Structural mechanisms for domain
movements in proteins". Biochemistry 33 (22): 6739–49. doi:10.1021/bi00188a001. PMID 8204609.
57.^ Herzberg O et al. (1996). "Swiveling-domain mechanism for enzymatic phosphotransfer
between remote reaction sites". Proc Natl Acad Sci USA 93 (7): 2652–7. doi:10.1073/pnas.93.7.2652.
PMID 8610096. PMC 39685.
58.^ a b Janin, J. and Wodak, S. J (1983). "Structural domains in proteins and their role in the
dynamics of protein function". Prog Biophys Mol Biol 42 (1): 21–78. doi:10.1016/0079-
6107(83)90003-2. PMID 6353481.
59.^ a b Hayward, 1999
60.^ Meador, W. E., Means, A. R., and Quiocho, F. A (1992). "Target enzyme recognition by
calmodulin: 2.4A structure of a calmodulin-peptide complex". Science 257 (5074): 1251–1255.
doi:10.1126/science.1519061. PMID 1519061.
61.^ Ikura, M., Clore, G. M., Gronenborn, A. M., Zhu, G., Klee, C. B., and Bax, A (1992).
"Solution structure of a calmodulin-target peptide complex by multidimensional NMR". Science 256
(5057): 632–638. doi:10.1126/science.1585175. PMID 1585175.
62.^ a b Sowdhamini R, Blundell TL. (1995). "An automatic method involving cluster analysis of
secondary structures for the identification of domains in proteins". Protein Sci 4 (3): 506–20.
doi:10.1002/pro.5560040317. PMID 7795532. PMC 2143076.
63.^ Swindells, M. B (1995). "A procedure for detecting structural domains in proteins". Protein
Sci 4 (1): 103–112. doi:10.1002/pro.5560040113. PMID 7773168.
64.^ Tsai CJ, Nussinov R. (1997). "Hydrophobic folding units derived from dissimilar monomer
structures and their interactions". Protein Sci 6 (1): 24–42. doi:10.1002/pro.5560060104.
PMID 9007974. PMC 2143523.
65.^ a b Crippen, G. M (1978). "The tree structural organisation of proteins". J Mol Biol 126 (3):
315–332. doi:10.1016/0022-2836(78)90043-8. PMID 745231.
66.^ Rossmann MG, Moras D, Olsen KW. (1974). "Chemical and biological evolution of
nucleotide-binding protein". Nature 250 (463): 194–9. doi:10.1038/250194a0. PMID 4368490.
67.^ Rose GD. (1979). "Hierarchic organization of domains in globular proteins". J Mol Biol 134
(3): 447–70. doi:10.1016/0022-2836(79)90363-2. PMID 537072.
68.^ a b Go N, Taketomi H. (1978). "Respective roles of short- and long-range interactions in
protein folding". Proc Natl Acad Sci USA 75 (2): 559–63. doi:10.1073/pnas.75.2.559. PMID 273218.
69.^ a b Holm L, Sander C. (1997). "Dali/FSSP classification of three-dimensional protein folds".
Nucleic Acids Res 25 (1): 231–4. doi:10.1093/nar/25.1.231. PMID 9016542. PMC 146389.
70.^ a b Siddiqui AS, Barton GJ. (1995). "Continuous and discontinuous domains: an algorithm
for the automatic generation of reliable protein domain definitions". Protein Sci 4 (5): 872–84.
doi:10.1002/pro.5560040507. PMID 7663343. PMC 2143117.
71.^ Zehfus, M. H (1997). "Identification of compact, hydrophobically stabilized domains and
modules containing multiple peptide chains". Protein Sci 6 (6): 1210–1219.
doi:10.1002/pro.5560060609. PMID 9194181.
72.^ Taylor WR. (1999). "Protein structural domain identification". Protein Eng 12 (3): 203–16.
doi:10.1093/protein/12.3.203. PMID 10235621.
73.^ Wodak, S. J. and Janin, J (1981). "Location of structural domains in protein". Biochemistry
20 (23): 6544–6552. doi:10.1021/bi00526a005. PMID 7306523.
74.^ Rashin, 1985
75.^ Zehfus MH, Rose GD. (1986). "Compact units in proteins". Biochemistry 25 (19): 5759–65.
doi:10.1021/bi00367a062. PMID 3778881.
76.^ a b Aleksiev, T., Potestio, R., Pontiggia, F., Cozzini, S. and Micheletti, C. (2009). "PiSQRD:
a web server for decomposing proteins into quasi-rigid dynamical domains.". Bioinformatics 25 (20):
2743–4. doi:10.1093/bioinformatics/btp512. PMID 19696046.
77.^ Micheletti, C., Carloni, P. and Maritan, A. Accurate and efficient description of protein
vibrational dynamics: comparing molecular dynamics and gaussian models, Proteins, 55, 635, 2004.
78.^ Barclay A (2003). "Membrane proteins with immunoglobulin-like domains--a master
superfamily of interaction molecules". Semin Immunol 15 (4): 215–23. doi:10.1016/S1044-
5323(03)00047-2. PMID 14690046.

[edit] Key papers

This article's further reading may not follow Wikipedia's content policies or
guidelines. Please improve this article by removing excessive, less relevant or many
publications with the same point of view; or by incorporating the relevant publications into
the body of the article through appropriate citations. (August 2010)
• Bastian, H. C. (1872). The beginnings of life: being some account of the nature, modes of
origin and transformation of lower organisms. Macmillan and Co., England.
• Berman HM et al. (2000). "The Protein Data Bank". Nucleic Acids Res 28 (1): 235–42.
doi:10.1093/nar/28.1.235. PMID 10592235. PMC 102472.
• Branden, C.-I. and Tooze, J. (1991). Introduction to protein structure. Garland, New York.
• Chothia C. (1992). "Proteins. One thousand families for the molecular biologist". Nature 357
(6379): 543–4. doi:10.1038/357543a0. PMID 1608464.
• Das S, Smith TF. (2000). "Identifying nature's protein Lego set". Adv Protein Chem 54: 159–
83. doi:10.1016/S0065-3233(00)54006-6. PMID 10829228.
• Dietmann S, Park J, Notredame C, Heger A, Lappe M, Holm L. (2001). "A fully automatic
evolutionary classification of protein folds: Dali Domain Dictionary version 3". Nucleic Acids Res 29
(1): 55–7. doi:10.1093/nar/29.1.55. PMID 11125048. PMC 29815.
• Dill, Ken A.; Chan, Hue Sun (1997). "From Levinthal to pathways to funnels". Nat Struc Biol 4
(1): 10–9. doi:10.1038/nsb0197-10. PMID 8989315.
• Dyson HJ, Sayre JR, Merutka G, Shin HC, Lerner RA, Wright PE. (1992). "Folding of peptide
fragments comprising the complete sequence of proteins. Models for initiation of protein folding. II.
Plastocyanin". J Mol Biol 226 (3): 819–35. doi:10.1016/0022-2836(92)90634-V. PMID 1507228.
• Fersht AR. (1997). "Nucleation mechanisms in protein folding". Curr Opin Struct Biol 7 (1): 3–
9. doi:10.1016/S0959-440X(97)80002-4. PMID 9032066.
• George DG, Hunt LT, Barker WC. (1996). "PIR-International Protein Sequence Database".
Methods Enzymol 266: 41–59. doi:10.1016/S0076-6879(96)66005-4. PMID 8743676.
• George, R. A. (2002) "Predicting Structural Domains in Proteins". Thesis, University College
• Go M. (1981). "Correlation of DNA exonic regions with protein structural units in
haemoglobin". Nature 291 (5810): 90–2. doi:10.1038/291090a0. PMID 7231530.
• Hadley, C and Jones, D.T. (1999). "A systematic comparison of protein structure
classifications: SCOP, CATH and FSSP". Struct Fold Des 7 (9): 1099. doi:10.1016/S0969-
• Hayward S. (1999). "Structural principles governing domain motions in proteins". Proteins 36
(4): 425–35. doi:10.1002/(SICI)1097-0134(19990901)36:4<425::AID-PROT6>3.0.CO;2-S.
PMID 10450084.
• Heringa J, Argos P. (1991). "Side-chain clusters in protein structures and their role in protein
folding". J Mol Biol 220 (1): 151–71. doi:10.1016/0022-2836(91)90388-M. PMID 2067014.
• Honig B. (1999). "Protein folding: from the levinthal paradox to structure prediction". J Mol
Biol 293 (2): 283–93. doi:10.1006/jmbi.1999.3006. PMID 10550209.
• Kim PS, Baldwin RL. (1990). "Intermediates in the folding reactions of small proteins". Annu
Rev Biochem 59 (1): 631–60. doi:10.1146/ PMID 2197986.
• Larsen TM, Laughlin LT, Holden HM, Rayment I, Reed GH. (1994). "Structure of rabbit
muscle pyruvate kinase complexed with Mn2+, K+, and pyruvate". Biochemistry 33 (20): 6301–9.
doi:10.1021/bi00186a033. PMID 8193145.
• Murvai J, Vlahovicek K, Barta E, Cataletto B, Pongor S. (2000). "The SBASE protein domain
library, release 7.0: a collection of annotated protein sequence segments". Nucleic Acids Res 28 (1):
260–2. doi:10.1093/nar/28.1.260. PMID 10592241. PMC 102474.
• Murzin AG, Brenner SE, Hubbard T, Chothia C. (1995). "SCOP: a structural classification of
proteins database for the investigation of sequences and structures". J Mol Biol 247 (4): 536–40.
doi:10.1016/S0022-2836(05)80134-2. PMID 7723011.
• Nissen P, Hansen J, Ban N, Moore PB, Steitz TA. (2000). "The structural basis of ribosome
activity in peptide bond synthesis". Science 289 (5481): 920–30. doi:10.1126/science.289.5481.920.
PMID 10937990.
• Janin J, Chothia C. (1985). "Domains in proteins: definitions, location, and structural
principles". Methods Enzymol 115: 420–30. doi:10.1016/0076-6879(85)15030-5. PMID 4079796.
• Schultz J, Copley RR, Doerks T, Ponting CP, Bork P. (2000). "SMART: a web-based tool for
the study of genetically mobile domains". Nucleic Acids Res 28 (1): 231–4. doi:10.1093/nar/28.1.231.
PMID 10592234. PMC 102444.
• Siddiqui AS, Dengler U, Barton GJ. (2001). "3Dee: a database of protein structural domains".
Bioinformatics 17 (2): 200–1. doi:10.1093/bioinformatics/17.2.200. PMID 11238081.
• Srinivasarao GY, Yeh LS, Marzec CR, Orcutt BC, Barker WC, Pfeiffer F. (1999). "Database
of protein sequence alignments: PIR-ALN". Nucleic Acids Res 27 (1): 284–5.
doi:10.1093/nar/27.1.284. PMID 9847202. PMC 148157.
• Tatusov RL et al. (2001). "The COG database: new developments in phylogenetic
classification of proteins from complete genomes". Nucleic Acids Res 29 (1): 22–8.
doi:10.1093/nar/29.1.22. PMID 11125040. PMC 29819.
• Taylor WR, Orengo CA. (1989). "Protein structure alignment". J Mol Biol 208 (1): 1–22.
doi:10.1016/0022-2836(89)90084-3. PMID 2769748.
• Yang AS, Honig B. (1995). "Free energy determinants of secondary structure formation: I.
alpha-Helices". J Mol Biol 252 (3): 351–65. doi:10.1006/jmbi.1995.0502. PMID 7563056.
• Yang AS, Honig B. (1995). "Free energy determinants of secondary structure formation: II.
Antiparallel beta-sheets". J Mol Biol 252 (3): 366–76. doi:10.1006/jmbi.1995.0503. PMID 7563057. }

[edit] External links

• The Protein Families (Pfam) database clan browser provides easy access to information
about protein structural domains. A clan contains two or more Pfam families that have arisen from a
single evolutionary origin.

[edit] Structural domain databases

• Conserved Domains at the National Center for Biotechnology website
• 3Dee
• Pawson Lab - Protein interaction domains
• Nash Lab - Protein interaction domains in Signal Transduction
• Definition and assignment of structural domains in proteins .

[edit] Sequence domain databases

• InterPro
• Pfam
• ProDom
• NCBI Conserved Domain Database
• SUPERFAMILY Library of HMMs representing superfamilies and database of (superfamily
and family) annotations for all completely sequenced organisms

[hide]v · d · eProtein domains

BAR • BIR • BZIP • CARD • C1 • C2 • DED • ENTH • FYVE • HEAT • Kringle • LIM • LRR • NACHT •
PAS • PDZ • Pyrin • PH • PX • SH2 • SH3 • SUN • TRIO • WD40 • zinc finger
[hide]v · d · eProtein tertiary structure

General Structural domain · Protein folding · Structure determination methods

All-α folds: Helix bundle · Globin fold · Homeodomain fold · Alpha solenoid

All-β folds: Immunoglobulin domain · Beta barrel · Beta-propeller

α/β folds: TIM barrel · Leucine-rich repeat · Flavodoxin fold · Rossmann fold ·
Thioredoxin fold · Trefoil knot fold

α+β folds: DNA clamp · Ferredoxin fold · Ribonuclease A · SH2-like fold

folds: Conotoxin

←Secondary structure
Quaternary structure→
[hide]v · d · eProteins

Processes Protein biosynthesis · Posttranslational modification · Protein folding · Protein

targeting · Proteome · Protein methods

Structures Protein structure · Protein structural domains · Proteasome

Types List of types of proteins · List of proteins · Membrane protein · Globular protein
(Globulin, Albumin) · Fibrous protein

biochemical families: prot · nucl · carb (glpr, alco, glys) · lipd (fata/i, phld, strd, gllp, eico) · amac/i ·
ncbs/i · ttpy/i
B proteins: BY STRUCTURE: membrane, globular (en, ca, an), fibrous

[hide]v · d · eBiomolecular structure

Protein structure Primary · Secondary · Tertiary · Quaternary · Determination ·

Prediction · Design · Thermodynamics

Nucleic acid Primary · Secondary · Tertiary · Quaternary · Determination ·

structure Prediction · Design · Thermodynamics

See also Protein · Protein domain · Protein engineering · Nucleic acid · DNA ·
RNA · Nucleic acid double helix

Retrieved from ""

Categories: Protein structure | Protein domains


Genetic memory (biology)

From Wikipedia, the free encyclopedia

Jump to: navigation, search

In biology, memory is present if the state of a biological system depends on its past history in
addition to present conditions. If this memory is recorded in the genetic material and stably inherited through
cell division (mitosis or meiosis), it is genetic.
• 1 Molecular biology
• 2 Somatic memory
• 2.1 Cellular memory
• 2.2 In animals
• 2.3 In plants
• 3 Inherited epigenetic memory
• 4 Microbial memory
• 5 Evolution
• 6 Animal behavior
• 7 References
• 8 Further reading

[edit] Molecular biology

In molecular biology, genetic memory resides in the genetic material of the cell and is expressed via
the genetic code used to translate it into proteins.[1][2] The genetic code enables cells to decode the
information needed to construct the protein molecules that make up living cells and therefore record and
store a one-dimensional blueprint for all the parts that make up an organism. This blueprint or genetic
memory in the form of species-specific collections of genes (genotype or genome) is passed from cell to cell
and from generation to generation in the form of DNA molecules. DNA therefore functions as a template for
protein synthesis.[2] Genetic memory can be modified by epigenetic memory, a process by which changes in
gene expression are passed on through mitosis or meiosis through factors other than DNA sequence.

[edit] Somatic memory

Somatic memory is limited to the organism and not passed on to subsequent generations. However,
its mechanism may involve mitotically stable genetic memory. The term applies for cellular memory, animals'
memory, and plants' memory, as described in the following paragraphs.

[edit] Cellular memory

All cells in multicellular organisms are derived from a pluripotent zygote and contain the same
genetic material (with a few exceptions). However, they are capable of recording a history of their
development within the organism leading to their specialized functions and limitations. Cells often employ
epigenetic processes that affect DNA-protein interactions to record this cellular memory in the form of
mitotically stable changes of the genetic material without a change in the DNA sequence itself. This is
typically achieved via changes of the chromatin structure.[3] Examples are methylation patterns of the DNA
molecule itself and proteins involved in packaging DNA, such as histones (also referred to as "histone code").

[edit] In animals
A case of somatic genetic memory is the immunological memory of the adaptive immune response in
vertebrates. The immune system is capable of learning to recognize pathogens and keeping a memory of
this learning process, which is the basis of the success of vaccinations. Antibody genes in B and T
lymphocytes are assembled from separate gene segments, giving each lymphocyte a unique antibody coding
sequence leading to the vast diversity of antibodies in the immune system. If stimulated by an antigen (e.g.
following vaccination or an infection with a pathogen), these antibodies are further fine-tuned via
hypermutation. Memory B cells capable of producing these antibodies form the basis for acquired
immunological memory.[6] Each individual therefore carries a unique genetic memory of its immune system's
close encounters with pathogens. As a somatic memory, this is not passed on to the next generation.

[edit] In plants
Plants that undergo vernalization (promotion of flowering by a prolonged exposure to cold
temperatures) record a genetic memory of winter to gain the competence to flower. The process involves
epigenetically recording the length of cold exposure through chromatin remodeling which leads to mitotically
stable changes in gene expression (the "winter code").[7] This releases the inhibition of flowering initiation
and allows the plants to bloom with the correct timing at the onset of spring. As a somatic memory, this state
is not passed on to subsequent generations but has to be acquired by each individual plant. The process of
vernalization was falsely assumed to be a stably inherited genetic memory passed on to subsequent
generations by the Russian geneticist Trofim Lysenko. Lysenko's claims of genetic memory and efforts to
obtain or fabricate results in proof of it had disastrous effects for Russian genetics in the early 20th century
(also see: Lysenkoism).[8]

[edit] Inherited epigenetic memory

In genetics, genomic imprinting or other patterns of inheritance that are not determined by DNA
sequence alone can form an epigenetic memory that is passed on to subsequent generations through
meiosis. In contrast, somatic genetic memories are passed on by mitosis and limited to the individual, but are
not passed on to the offspring. Both processes include similar epigenetic mechanisms, e.g. involving
histones and methylation patterns.[9][10]

[edit] Microbial memory

In microbes, genetic memory is present in the form of inversion of specific DNA sequences serving
as a switch between alternative patterns of gene expression.[11]

[edit] Evolution
In population genetics and evolution, genetic memory represents the recorded history of adaptive
changes in a species. Selection of organisms carrying genes coding for the best adapted proteins results in
the evolution of species. An example for such a genetic memory is the innate immune response that
represents a recording of the history of common microbial and viral pathogens encountered throughout the
evolutionary history of the species.[12] In contrast to the somatic memory of the adaptive immune response,
the innate immune response is present at birth and does not require the immune system to learn to recognize
In the history of theories of evolution, the proposed genetic memory of an individual's experiences
and environmental influences was a central part of Lamarckism to explain the inheritance of evolutionary

[edit] Animal behavior

In ethology, genetic memory refers the inheritance of instinct in animals and humans.[citation
[edit] References
1. ^ Nirenberg M (1968). "Genetic memory". JAMA 206 (9): 1973–7.
doi:10.1001/jama.206.9.1973. PMID 4880506.
2. ^ a b Marshall Nirenberg (1968): "The genetic code". Nobel Lecture
3. ^ Hirose S (2007). "Crucial roles for chromatin dynamics in cellular memory". J. Biochem.
141 (5): 615–9. doi:10.1093/jb/mvm092. PMID 17416595.
4. ^ Bird A (2002). "DNA methylation patterns and epigenetic memory". Genes Dev. 16 (1): 6–
21. doi:10.1101/gad.947102. PMID 11782440.
5. ^ Turner BM (2002). "Cellular memory and the histone code". Cell 111 (3): 285–91.
doi:10.1016/S0092-8674(02)01080-2. PMID 12419240.
6. ^ Crotty S, Ahmed R (2004). "Immunological memory in humans". Semin. Immunol. 16 (3):
197–203. doi:10.1016/j.smim.2004.02.008. PMID 15130504.
7. ^ Sung S, Amasino RM (2006). "Molecular genetic studies of the memory of winter". J. Exp.
Bot. 57 (13): 3369–77. doi:10.1093/jxb/erl105. PMID 16980591.
8. ^ Amasino R (2004). "Vernalization, competence, and the epigenetic memory of winter".
Plant Cell 16 (10): 2553–9. doi:10.1105/tpc.104.161070. PMID 15466409.
9. ^ Ooi SL, Henikoff S (2007). "Germline histone dynamics and epigenetics". Curr. Opin. Cell
Biol. 19 (3): 257–65. doi:10.1016/ PMID 17467256.
10.^ Gehring M, Henikoff S (2007). "DNA methylation dynamics in plant genomes". Biochim.
Biophys. Acta 1769 (5-6): 276–86. doi:10.1016/j.bbaexp.2007.01.009. PMID 17341434.
11.^ Casadesús J, D'Ari R (2002). "Memory in bacteria and phage". Bioessays 24 (6): 512–8.
doi:10.1002/bies.10102. PMID 12111734.
12.^ Dempsey PW, Vaidya SA, Cheng G (2003). "The art of war: Innate and adaptive immune
responses". Cell. Mol. Life Sci. 60 (12): 2604–21. doi:10.1007/s00018-003-3180-y. PMID 14685686.
[edit] Further reading
• Alan Bullock and Oliver Stallybrass (1977). "Genetic memory". The Harper Dictionary of
Modern Thought. Harper & Row. pp. 258.
Retrieved from ""

Categories: Genetics | Evolutionary biology


Intelligent design movement

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Main article: Intelligent design
The intelligent design movement is a neo-creationist religious campaign for broad social, academic
and political change to promote and support the idea of "intelligent design," which asserts that "certain
features of the universe and of living things are best explained by an intelligent cause, not a possibly
undirected process such as natural selection."[1][2] Its chief activities are a campaign to promote public
awareness of this concept, the lobbying of policymakers to include its teaching in high school science
classes, and legal action, either to defend such teaching or to remove barriers otherwise preventing it.[3][4]
The movement arose out of the previous Christian fundamentalist and evangelistic creation science
movement in the United States,[5] and is driven by a small group of proponents.[6][7]
The overall goal of the intelligent design movement is to "overthrow materialism" and atheism. Its
proponents believe that society has suffered "devastating cultural consequences" from adopting materialism
and that science is the cause of the decay into materialism because it seeks only natural explanations, and is
therefore atheistic. They believe that the theory of evolution implies that humans have no spiritual nature, no
moral purpose, and no intrinsic meaning. They seek to "defeat [the] materialist world view" represented by
the theory of evolution in favor of "a science consonant with Christian and theistic convictions".[3]
To achieve their goal of defeating a materialistic world view, advocates of intelligent design take a
two-pronged approach. Alongside the promotion of intelligent design, proponents also seek to "Teach the
Controversy"; discredit evolution by emphasizing perceived flaws in the theory of evolution, or disagreements
within the scientific community and encourage teachers and students to explore non-scientific alternatives to
evolution, or to critically analyze evolution and the controversy surrounding the teaching of evolution. But the
world's largest general scientific society, the American Association for the Advancement of Science, has
stated that "There is no significant controversy within the scientific community about the validity of evolution."
and that "Evolution is one of the most robust and widely accepted principles of modern science."[8] The
ruling in the Dover trial, Kitzmiller v. Dover Area School District, where the claims of intelligent design
proponents were considered by a United States federal court, stated that "evolution, including common
descent and natural selection, is 'overwhelmingly accepted' by the scientific community."[9][10]
The Discovery Institute[11] is a conservative Christian think tank that drives the intelligent design
movement.[12] The Institute's Center for Science and Culture (CSC) counts most of the leading intelligent
design advocates among its membership, most notably its program advisor Phillip E. Johnson. Johnson is
the architect of the movement's key strategies, the "wedge strategy" and the Teach the Controversy
The Discovery Institute and leading proponents represent intelligent design as a revolutionary
scientific theory.[13][14][15][16] The overwhelming majority of the scientific community,[10] as represented
by the American Association for the Advancement of Science,[17] the National Academy of Sciences[18] and
nearly all scientific professional organizations, firmly rejects these claims, and insist that intelligent design is
not valid science, its proponents having failed to conduct an actual scientific research program.[10] This has
led the movement's critics to state that intelligent design is merely a public relations campaign and a political
According to critics of the intelligent design movement, the movement's purpose is political rather
than scientific or educational. They claim the movement's "activities betray an aggressive, systematic agenda
for promoting not only intelligent design creationism, but the religious worldview that undergirds it."[20]
Intelligent design is an attempt to recast religious dogma in an effort to reintroduce the teaching of biblical
creationism to public school science classrooms; the intelligent design movement is an effort to reshape
American society into a theocracy, primarily through education. As evidence, critics cite the Discovery
Institute's political activities, its "Wedge strategy" and statements made by leading intelligent design
The scientific community's position, as represented by the National Academy of Sciences and the
National Center for Science Education, is that intelligent design is not science, but creationist pseudoscience.
Richard Dawkins, a biologist and professor at Oxford University, compares the intelligent design movement's
demand to "teach the controversy" with the demand to teach flat earthism; acceptable in terms of history, but
not in terms of science. "If you give the idea that there are two schools of thought within science, one that
says the earth is round and one that says the earth is flat, you are misleading children."[21]
• 1 Philosophy
• 2 History of the movement
• 2.1 Origins
• 2.2 Center for the Renewal of Science and Culture
• 2.3 The Wedge strategy
• 2.4 Kansas evolution hearings
• 2.5 Kitzmiller v. Dover Area School District
• 3 Reception by the scientific community
• 4 Structure
• 4.1 The 'big tent' strategy
• 4.2 Obfuscation of religious motivation
• 4.3 Organizations
• 4.3.1 The Center for Science and Culture
• 4.3.2 Other organizations
• 5 Activism
• 5.1 Campaigns
• 5.2 Politics and public education
• 5.3 Higher education
• 5.4 The Web
• 5.5 International
[edit] Philosophy
At the 1999 "Reclaiming America for Christ Conference"[22] called by Reverend D. James Kennedy
of Coral Ridge Ministries, Johnson gave a speech called How the Evolution Debate Can Be Won. [23] In it he
sums up the theological and epistemological underpinnings of intelligent design and its strategy for winning
the battle:
"To talk of a purposeful or guided evolution is not to talk about evolution at all. That is slow
creation. When you understand it that way, you realize that the Darwinian theory of evolution
contradicts not just the Book of Genesis, but every word in the Bible from beginning to end. It
contradicts the idea that we are here because a creator brought about our existence for a
purpose. That is the first thing I realized, and it carries tremendous meaning." -- Phillip Johnson

"I have built an intellectual movement in the universities and churches that we call The Wedge,
which is devoted to scholarship and writing that furthers this program of questioning the
materialistic basis of science. One very famous book that's come out of The Wedge is
biochemist Michael Behe's book, Darwin's Black Box, which has had an enormous impact on
the scientific world." -- Phillip Johnson

"Now the way that I see the logic of our movement going is like this. The first thing you
understand is that the Darwinian theory isn't true. It's falsified by all of the evidence and the logic
is terrible. When you realize that, the next question that occurs to you is, well, where might you
get the truth? When I preach from the Bible, as I often do at churches and on Sundays, I don't
start with Genesis. I start with John 1:1. In the beginning was the word. In the beginning was
intelligence, purpose, and wisdom. The Bible had that right. And the materialist scientists are
deluding themselves." -- Phillip Johnson

[edit] History of the movement

The intelligent design movement grew out of a creationist tradition which argues against evolutionary
theory from a religious standpoint, usually that of evangelical or fundamentalistic Christianity. Although
intelligent design advocates often claim that they are arguing only for the existence of a designer who may or
may not be God, all the movement's leading advocates believe that this designer is God. They frequently
accompany their arguments with a discussion of religious issues, especially when addressing religious
audiences, but elsewhere downplay the religious aspects of their agenda.

[edit] Origins
The modern use of the words "intelligent design", as a term intended to describe a field of inquiry,
began after the Supreme Court of the United States, in the case of Edwards v. Aguillard (1987), ruled that
creationism is unconstitutional in public school science curricula. A Discovery Institute report says that
Charles Thaxton, editor of Of Pandas and People, had picked the phrase up from a NASA scientist, and
thought "That's just what I need, it's a good engineering term".[24] In drafts of the book over one hundred
uses of the root word "creation", such as "creationism" and "creation science", were changed, almost without
exception, to "intelligent design",[25] while "creationists" was changed to "design proponents" or, in one
instance, "cdesign proponentsists". [sic][26] In 1989 Of Pandas and People was published by the Foundation
for Thought and Ethics,[27] with the definition:
Intelligent design means that various forms of life began abruptly through an intelligent agency,
with their distinctive features already intact. Fish with fins and scales, birds with feathers, beaks,
wings, etc.[28]

Pandas was followed in 1991 by Darwin on Trial, a neo-creationist polemic by University of

California, Berkeley law professor emeritus Phillip E. Johnson, that is regarded as a central text of the
movement.[29] Darwin on Trial mentioned Pandas as "'creationist' only in the sense that it juxtaposes a
paradigm of 'intelligent design' with the dominant paradigm of (naturalistic) evolution", but his use of the term
as a focus for his wedge strategy promoting "theistic realism" came later.[page needed] The book was
reviewed by evolutionary biologist Stephen Jay Gould for Scientific American in July 1992, concluding that
the book contains " weighing of evidence, no careful reading of literature on all sides, no full citation of
sources (the book does not even contain a bibliography) and occasional use of scientific literature only to
score rhetorical points."[30] This "devastating" review led to the formation in 1992 or 1993 of an 'Ad Hoc
Origins Committee' of Johnson's supporters, which wrote a letter, circulated to thousands of university
professors, defending the book. Among the 39 signatories were nine who later became members of the
Center for the Renewal of Science and Culture.[31]
During the early 1990s Johnson worked to develop a 'big tent' movement to unify a wide range of
creationist viewpoints in opposition to evolution. In 1992, the first formal meeting devoted to intelligent design
was held in Southern Methodist University. It included a debate between Johnson and Michael Ruse (a key
witness in McLean v. Arkansas) and papers by William A. Dembski, Michael Behe and Stephen C. Meyer. In
1993 Johnson organized a follow-up meeting, including Dembski, Behe, Meyer, Dean H. Kenyon (co-author
of Pandas) and Walter Bradley (co-author with Thaxton and Kenyon of The Mystery of Life's Origin), as well
as two young Earth creationist graduate students, Paul A. Nelson and Jonathan Wells.[32]
[edit] Center for the Renewal of Science and Culture
On 6 December 1993 an article by Meyer was published in the Wall Street Journal, drawing national
attention to the controversy over Kenyon's teaching of creationism. This article also gained the attention of
Discovery Institute co-founder Bruce Chapman. On discovering that Meyer was developing the idea of
starting a scientific research center in conversations with conservative political scientist John G. West,
Chapman invited them to create a unit within the Discovery Institute called the Center for the Renewal of
Science and Culture (later renamed the Center for Science and Culture). This center was dedicated to
overthrowing "scientific materialism" and "fomenting nothing less than a scientific and cultural revolution".[33]
A 1995 conference, on "The Death of Materialism and the Renewal of Culture" served as a blueprint for the
center.[34] By 1996 they had nearly a million dollars in grants, the largest being from Howard Ahmanson, Jr.,
with smaller but still large contributions coming from the Stewardship Foundation established by C. Davis
Weyerhaeuser and the Maclellan Foundation, and appointed their first class of research fellows.[33]

[edit] The Wedge strategy

The Wedge strategy was formulated by Johnson to combat the "evil" of methodological naturalism.
[35] It first came to the general public's attention when a Discovery Institute internal memo now known as the
"Wedge Document" (believed to have been written in 1998) was leaked to the public in 1999. However it is
believed to have been update of an earlier document to be implemented between 1996 and 2001.[36]
The document begins with "the proposition that human beings are created in the image of God is one
of the bedrock principles on which Western civilization was built." and then goes on to outline the
movement's goal to exploit perceived discrepancies within evolutionary theory in order to discredit evolution
and scientific materialism in general. Much of the strategy is directed toward the broader public, as opposed
to the professional scientific community. The stated "governing goals" of the CSC's wedge strategy are:
1. To defeat scientific materialism and its destructive moral, cultural and political legacies
2. To replace materialistic explanations with the theistic understanding that nature and human
beings are created by God.
Critics of intelligent design movement argue that the wedge document and strategy demonstrate that
the intelligent design movement is motivated purely by religion and political ideology and that the Discovery
Institute as a matter of policy obfuscates its agenda. The Discovery Institute's official response was to
characterize the criticism and concern as "irrelevant," "paranoid," and "near-panic" while portraying the
wedge document as a "fund-raising document."[37]
Johnson in his 1997 book Defeating Darwinism by Opening Minds confirmed some of the concerns
voiced by the movement's gainsayers:
"If we understand our own times, we will know that we should affirm the reality of God by
challenging the domination of materialism and naturalism in the world of the mind. With the
assistance of many friends I have developed a strategy for doing this,...We call our strategy the
"wedge." -- Phillip Johnson[38]

[edit] Kansas evolution hearings

The Kansas evolution hearings were a series of hearings held in Topeka, Kansas, United States May
5 to May 12, 2005 by the Kansas State Board of Education and its State Board Science Hearing Committee
to change how evolution and the origin of life would be taught in the state's public high school science
classes. The hearings were arranged by the conservative Christian Board of Education with the intent of
introducing intelligent design into science classes via the Teach the Controversy method.[39][40]
The hearings raised the issues of creation and evolution in public education and were attended by all
the major participants in the intelligent design movement but were ultimately boycotted by the scientific
community over concern of lending credibility to the claim, made by proponents of intelligent design, that
evolution is purportedly the subject of wide dispute within the scientific and science education communities.
The Discovery Institute, hub of the intelligent design movement, played a central role in starting the
hearings by promoting its Critical Analysis of Evolution lesson plan[41] which the Kansas State Board of
Education eventually adopted over objections of the State Board Science Hearing Committee, and
campaigning on behalf of conservative Republican candidates for the Board.[42]
Local science advocacy group Kansas Citizens for Science organized a boycott of the hearings by
mainstream scientists, who accused it of being a kangaroo court and argued that their participation would
lend an undeserved air of legitimacy to the hearings.[43] Board member Kathy Martin declared at the
beginning of the hearings "Evolution has been proven false. ID (Intelligent Design) is science-based and
strong in facts." At their conclusion she proclaimed that evolution is "an unproven, often disproven" theory.
"ID has theological implications. ID is not strictly Christian, but it is theistic," asserted Martin.[44] The
scientific community rejects teaching intelligent design as science; a leading example being the United
States National Academy of Sciences, which issued a policy statement saying "Creationism, intelligent
design, and other claims of supernatural intervention in the origin of life or of species are not science
because they are not testable by the methods of science."[45]
On February 13, 2007, the Board voted 6 to 4 to reject the amended science standards enacted in
[edit] Kitzmiller v. Dover Area School District
In the movement's sole major case, Kitzmiller v. Dover Area School District, it was represented by
the Thomas More Law Center,[47] which had been seeking a test-case on the issue for at least five years.
[48][49] However conflicting agendas resulted in the withdrawal of a number of Discovery Institute (DI)
Fellows as expert witnesses, at the request of DI director Bruce Chapman,[50] and mutual recriminations
with the DI after the case was lost.[51] The Alliance Defense Fund briefly represented the Foundation for
Thought and Ethics (FTE) in its unsuccessful motion to intervene in this case,[52] and prepared amicus
curiae briefs on behalf of the DI and FTE in it.[53] It has also made amicus curiae submissions[54] and
offered to pay for litigation,[55] in other (actual and potential) creationism-related cases. On a far smaller
scale, Larry Caldwell and his wife operate under the name Quality Science Education for All, and have made
a number of lawsuits in furtherance of the movement's anti-evolution agenda. In 2005 they brought at least
three separate lawsuits to further the intelligent design movement's agenda. One was later abandoned, two
were dismissed.[56][57][58]

[edit] Reception by the scientific community

Intelligent design advocates realize that their arguments have little chance of acceptance within the
mainstream scientific community, so they direct them toward politicians, philosophers and the general public.
[59][60][61] What prima facie "scientific" material they have produced has been attacked by critics as
containing factual misrepresentation and misleading, rhetorical and equivocal terminology. A number of
pseudoscientific documentaries that present intelligent design as an increasingly well-supported line of
scientific inquiry have been made.[62][63] The bulk of the material produced by the intelligent design
movement, however, is not intended to be scientific but rather to promote its social and political aims.[64][65]
[66] Polls indicate that intelligent design's main appeal to citizens comes from its link to religious concepts.
An August 2005 poll from The Pew Forum on Religion & Public Life showed 64% of Americans
favoring the teaching of creationism along with evolution in science classrooms, though only 38% favored
teaching it instead of evolution, with the results varying deeply by education level and religiosity. The poll
showed the educated were far less attached to intelligent design than the less educated. Evangelicals and
fundamentalists showed high rates of affiliation with intelligent design while other religious persons and the
secular were much lower.[67]
Scientists responding to a poll overwhelmingly said intelligent design is about religion, not science. A
2002 sampling of 460 Ohio science professors had 91% say it's primarily religion, 93% say there is not "any
scientifically valid evidence or an alternative scientific theory that challenges the fundamental principle of the
theory of evolution," and 97% say that they did not use intelligent design concepts in their own research.[68]
In October and November 2001 the Discovery Institute advertised A Scientific Dissent From
Darwinism listing what they claimed were "100 scientific dissenters" who had signed a statement that "We
are skeptical of claims for the ability of random mutation and natural selection to account for the complexity of
life. Careful examination of the evidence for Darwinian theory should be encouraged."[69] Shortly afterwards
the NCSE described the wording as misleading, noting that a minority of the signatories were biologists and
some of the others were engineers, mathematicians and philosophers, and that some signatories did not fully
support the Discovery Institute's claims. The list was further criticized in a February 2006 New York Times
article[70] which pointed out that only 25% of the signatories by then were biologists and that signatories'
"doubts about evolution grew out of their religious beliefs." In 2003 as a humorous parody of such listings the
NCSE produced the pro-evolution Project Steve list of signatories, all with variations of the name Steve and
most of whom are trained biologists. As of July 31, 2006, the Discovery Institute lists "over 600 scientists",
while Project Steve reported 749 signatories; as of September 30, 2009, 1,112 Steves have signed the
[edit] Structure
[edit] The 'big tent' strategy
The movement's strategy as set forth by Johnson states the replacement of "materialist science" with
"theistic science" as its primary goal; and, more generally, for intelligent design to become "the dominant
perspective in science" and to "permeate our religious, cultural, moral and political life." This agenda is now
being actively pursued by the Center for Science and Culture (CSC), which plays the leading role in the
promotion of intelligent design. Its fellows include most of the leading intelligent design advocates: William A.
Dembski, Michael Behe, Jonathan Wells and Stephen C. Meyer.
Intelligent design has been described by its proponents as a "big tent" belief, one in which all theists
united by a having some kind of creationist belief (but of differing opinions as regards details) can support. If
successfully promoted, it would reinstate creationism in the teaching of science, after which debates
regarding details could resume. In his 2002 article Big Tent: Traditional Creationism and the Intelligent
Design Community,[72] Discovery Institute fellow Paul A. Nelson credits Johnson for the "big tent" approach
and for reviving creationist debate since the Edwards v. Aguillard decision. According to Nelson, "The
promise of the big tent of ID is to provide a setting where Christians and others may disagree amicably and
fruitfully about how best to understand the natural world as well as scripture."
In his presentation to the 1999 Reclaiming America for Christ Conference, How the Evolution Debate
can be Won, Johnson affirmed this "big tent" role for "The Wedge" (without using the term intelligent design):
To talk of a purposeful or guided evolution is not to talk about evolution at all. That is "slow
creation." When you understand it that way, you realize that the Darwinian theory of evolution
contradicts not just the book of Genesis, but every word in the Bible from beginning to end. It
contradicts the idea that we are here because a Creator brought about our existence for a
purpose. That is the first thing I realized, and it carries tremendous meaning. [...]

So did God create us? Or did we create God? That's an issue that unites people across the
theistic world. Even religious, God-believing Jewish people will say, "That's an issue we really
have a stake in, so let's debate that question first. Let us settle that question first. There are
plenty of other important questions on which we may not agree, and we'll have a wonderful time
discussing those questions after we've settled the first one. We will approach those questions in
a better spirit because we have worked together for this important common end." [...]

[The Wedge is] inherently an ecumenical movement. Michael Behe is a Roman Catholic. The
next book that is coming out from Cambridge University Press by one of my close associates is
by an evangelical convert to Greek Orthodoxy. We have a lot of Protestants, too. The point is
that we have this broad-based intellectual movement that is enabling us to get a foothold in the
scientific and academic journals and in the journals of the various religious faiths.

– Phillip Johnson, The Evolution Debate Can Be Won[23]

The Discovery Institute consistently denies allegations that its intelligent design agenda has religious
foundations, and downplays the religious source of much of its funding. In an interview of Stephen C. Meyer
when ABC News'asked about the Discovery Institute's many evangelical Christian donors the institute's
public relations representative stopped the interview saying "I don't think we want to go down that path."[73]
[edit] Obfuscation of religious motivation
Phillip E. Johnson, largely regarded as the leader of the movement, positions himself as a "theistic
realist" against "methodological naturalism" and intelligent design as the method through which God created
life.[74] Johnson explicitly calls for intelligent design proponents to obfuscate their religious motivations so as
to avoid having intelligent design recognized "as just another way of packaging the Christian evangelical
message."[75] Hence intelligent design arguments are carefully formulated in secular terms and intentionally
avoid positing the identity of the designer. Johnson has stated that cultivating ambiguity by employing secular
language in arguments which are carefully crafted to avoid overtones of theistic creationism is a necessary
first step for ultimately introducing the Christian concept of God as the designer. Johnson emphasizes "the
first thing that has to be done is to get the Bible out of the discussion" and that "after we have separated
materialist prejudice from scientific fact" only then can "biblical issues" be discussed.[76] In the foreword to
Creation, Evolution, & Modern Science (2000) Johnson writes "The intelligent design movement starts with
the recognition that 'In the beginning was the Word.' and 'In the beginning God created.' Establishing that
point isn't enough, but it is absolutely essential to the rest of the gospel message."

[edit] Organizations

[edit] The Center for Science and Culture

Main article: Center for Science and Culture
The Center for Science and Culture (CSC), formerly known as the Center for Renewal of Science
and Culture (CRSC), is a division of the Discovery Institute. The Center consists of a tightly knit core of
people who have worked together for almost a decade to advance intelligent design as both a concept and a
movement as necessary adjuncts of its wedge strategy policy. This cadre includes Phillip E. Johnson,
Michael Behe, William A. Dembski and Stephen C. Meyer. They are united by a religious vision which,
although it varies among the members in its particulars and is seldom acknowledged outside of the Christian
press, is predicated on the shared conviction that America is in need of "renewal" which can be
accomplished only by unseating "Godless" materialism and instituting religion as its cultural foundation.
In his keynote address at the "Research and Progress in intelligent design" (RAPID) conference held
in 2002 at Biola University, William A. Dembski described intelligent design's "dual role as a constructive
scientific project and as a means for cultural renaissance." In a similar vein, the movement's hub, the
Discovery Institute's Center for Science and Culture had until 2002 been the "Center for the Renewal of
Science and Culture". Explaining the name change, a spokesperson for the CSC insisted that the old name
was simply too long. However, the change followed accusations that the center's real interest was not
science but reforming culture along lines favored by conservative Christians.
Critics of the movement cite the Wedge Document as confirmation of this criticism and assert that the
movement's leaders, particularly Phillip E. Johnson, view the subject as a culture war: "Darwinian evolution is
not primarily important as a scientific theory but as a culturally dominant creation story ... When there is
radical disagreement in a commonwealth about the creation story, the stage is set for intense conflict, the
kind ... known as 'culture war.' "
Recently the Center for Science and Culture's has moderated its previous overtly theistic mission
statements[77] to appeal to a broader, a more secular audience. It hopes to accomplish this by using less
overtly theistic messages and language.[78] Despite this, the Center for Science and Culture still states as a
goal a redefinition of science, and the philosophy on which it is based, particularly the exclusion of what it
calls the "unscientific principle of materialism", and in particular the acceptance of what it calls "the scientific
theory of intelligent design".
According to Reason magazine, promotional materials from the Discovery Institute acknowledge that
the Ahmanson family donated $1.5 million to the Center for Science and Culture, then known as the Center
for Renewal of Science and Culture, for a research and publicity program to "unseat not just Darwinism but
also Darwinism's cultural legacy". Mr. Ahmanson funds many causes important to the Christian religious
right, including Christian Reconstructionism, whose goal is to place the U.S. "under the control of biblical
law."[79] Until 1995, Ahmanson sat on the board of the Christian reconstructionist Chalcedon Foundation.[80]

[edit] Other organizations

• The Access Research Network (ARN), has become a comprehensive clearinghouse for ID
resources, including news releases, publications, multimedia products and an elementary school
science curriculum. It's stated mission is "providing accessible information on science, technology
and society issues from an intelligent design perspective."[81] Its directors are Dennis Wagner and
CSC Fellows Mark Hartwig, Stephen C. Meyer and Paul Nelson.[82] Its 'Friends of ARN' is also
dominated by CSC Fellows.[81]
• The Intelligent Design and Evolution Awareness Center (IDEA Center) is a Christian[83]
nonprofit organization formed originally as a student club promoting intelligent design at the
University of California, San Diego (UCSD). There are about 25 active chapters of this organization
in the United States, Kenya, Canada, Ukraine, and The Philippines. There have been a total of 35
active chapters formed and several others are currently pending. Six out of the listed 32 chapters in
the USA are located at high schools [84] In December 2008, biologist Allen MacNeill stated, on the
basis of analysis of the webpages of the national organization and local chapters, that it appeared
that the organization is moribund.[85]
• The Intelligent Design Network (IDnet) is a nonprofit organization formed in Kansas to
promote intelligent design. It is based in Shawnee Mission, Kansas. The Intelligent Design Network
was founded by John Calvert, a corporate finance lawyer with a bachelor's degree in geology and
nutritionist William S. Harris. Together, Calvert and Harris have published the article "Intelligent
Design: The Scientific Alternative to Evolution" in the National Catholic Bioethics Quarterly.[86]
Calvert also has written a play about intelligent design in a high school biology class with Daniel
• The Foundation for Thought and Ethics (FTE) is a Christian non-profit organization[88] based
in Richardson, Texas that publishes textbooks and articles promoting intelligent design, abstinence,
and Christian nationism. In addition, the foundation's officers and editors are some of the leading
proponents of intelligent design. The FTE has close associations with the Discovery Institute, hub of
the intelligent design movement and other religious Christian groups.

[edit] Activism
The intelligent design movement primarily campaigns on two fronts: a public relations campaign
meant to influence the popular media and sway public opinion; and an aggressive lobbying campaign to
cultivate support for the teaching of intelligent design amongst policymakers and the wider educational
community. Both these activities are largely funded and directed by the Discovery Institute, from national to
grassroots levels. The movement's first goal is to establish an acceptance of intelligent design at the expense
of evolution in public school science; its long-term goal is no less than the "renewal" of American culture
through the shaping of public policy to reflect conservative Christian values. As the Discovery Institute states,
intelligent design is central to this agenda: "Design theory promises to reverse the stifling dominance of the
materialist worldview, and to replace it with a science consonant with Christian and theistic convictions."
The Discovery Institute has also relied on several polls to indicate the acceptance of intelligent
design. A 2005 Harris poll identified ten percent of adults in the United States as taking what they called the
intelligent design position, that "human beings are so complex that they required a powerful force or
intelligent being to help create them". (64% agreed with the creationist view that "human beings were created
directly by God" and 22% believed that "human beings evolved from earlier species". However, 49%
accepted plant and animal evolution, while 45% did not.)[89] Although some polls commissioned by the
Discovery Institute show more support, these polls have been criticized as suffering from considerable flaws,
such as having a low response rate (248 out of 16,000), being conducted on behalf of an organization with an
expressed interest in the outcome of the poll, and containing leading questions.[90]
Critics of intelligent design and its movement contend that intelligent design is a specific form of
creationism, neo-creationism, a viewpoint rejected by intelligent design advocates. It was bolstered by the
2005 ruling in United States federal court that a public school district requirement for science classes to teach
that intelligent design is an alternative to evolution was a violation of the Establishment Clause of the First
Amendment to the United States Constitution. In Kitzmiller v. Dover Area School District (2005), United
States District Judge John E. Jones III also ruled that intelligent design is not science and is essentially
religious in nature.
In pursuing the goal of establishing intelligent design at the expense of evolution in public school
science, intelligent design groups have threatened and isolated high school science teachers, school board
members and parents who opposed their efforts.[91][92][93] Responding to the well-organized curricular
challenges of intelligent design proponents to local school boards have been disruptive and divisive in the
communities where they've taken place. The campaigns run by intelligent design groups place teachers in
the difficult position of arguing against their employers while the legal challenges to local school districts are
costly and divert scarce funds away from education into court battles. Although these court battles have
almost invariably resulted in the defeat of intelligent design proponents, they are draining and divisive to local
schools. For example, as a result of Kitzmiller v. Dover Area School District trial, the Dover Area School
District was forced to pay $1,000,011 in legal fees and damages for pursuing a policy of teaching the
controversy - presenting intelligent design as an allegedly scientific alternative to evolution. [94]
Leading members of the intelligent design movement are also associated with denialism, both Phillip
Johnson and Jonathan Wells have signed an AIDS denialism petition.[95][96][97][98]

[edit] Campaigns
Main article: Discovery Institute intelligent design campaigns
The Discovery Institute, through its Center for Science and Culture, has formulated a number of
campaigns to promote intelligent design, while discrediting evolutionary biology, which the Institute terms
Prominent Institute campaigns have been to 'Teach the Controversy' and, more recently, to allow
Critical Analysis of Evolution. Other prominent campaigns have claimed that intelligent design advocates
(most notably Richard Sternberg) have been discriminated against, and thus that Academic Freedom bills
are needed to protect academics' and teachers' ability to criticise evolution, and that there is a link from
evolution to ideologies such as Nazism and eugenics. These three claims are all publicised in the pro-ID
movie Expelled: No Intelligence Allowed. Other campaigns have included petitions, most notably A Scientific
Dissent From Darwinism.
The response of the scientific community has been to reiterate that the theory of evolution is
overwhelmingly accepted as a matter of scientific consensus[100] whereas intelligent design has been
rejected by the overwhelming majority of the scientific community (see list of scientific societies explicitly
rejecting intelligent design).

[edit] Politics and public education

Main articles: Intelligent design in politics and creation and evolution in public education
The main battlefield for this culture war has been U.S. regional and state school boards. Courts have
also become involved as those campaigns to include intelligent design or weaken the teaching of evolution in
public school science curricula are challenged on First Amendment grounds.[101] In Kitzmiller v. Dover Area
School District the plaintiffs successfully argued that intelligent design is a form of creationism, and that the
school board policy thus violated the Establishment Clause of the First Amendment.
Intelligent design is an integral part of a political campaign by cultural conservatives, largely from
evangelical religious convictions, that seek to redefine science to suit their own ideological agenda.[102]
Though numerically a minority of Americans,.[67] the politics of intelligent design is based less on numbers
than on intensive mobilization of ideologically committed followers and savvy public relations campaigns.
[103] Political repercussions from the culturally conservative sponsorship of the issue has been divisive and
costly to the effected communities, polarizing and dividing not only those directly charged with educating
young people but entire local communities.
With a doctrine that calls itself science among non-scientists but is rejected by the vast majority of
the real practitioners, an amicable coexistence and collaboration between intelligent design advocates and
upholders of mainstream science education standards is rare. With mainstream scientific and educational
organizations saying the theory of evolution is not "in crisis" or a subject doubted by scientists, nor intelligent
design the emergent scientific paradigm or rival theory its proponents proclaim,[104] "teaching the
controversy" is suitable for classes on politics, history, culture, or theology they say, but not science. By
attempting to force the issue into science classrooms, intelligent design proponents create a charged
environment that forces participants and bystanders alike to declare their positions, which has resulted in
intelligent design groups threatening and isolating high school science teachers, school board members and
parents who opposed their efforts.[91][92][93][105][106]
In a round table discussion entitled "Science Wars: Should Schools Teach Intelligent Design?"[107]
at the American Enterprise Institute on 21 October 2005 and televised on C-SPAN, the Discovery Institute's
Mark Ryland and the Thomas More Law Center's Richard Thompson had a frank disagreement, in which
Ryland claimed the Discovery Institute has always cautioned against the teaching of intelligent design, and
Thompson responded that the institute's leadership had not only advocated the teaching of intelligent design,
but encouraged others to do so, and that the Dover Area School District had merely followed the institute's
calls for action.[51] As evidence, Thompson cited the Discovery Institute's guidebook Intelligent Design in
Public School Science Curricula[108] written by the institute's director and co-founder, Stephen C. Meyer and
David DeWolf, a fellow of the institute, which stated in its closing paragraphs: " Moreover, as the previous
discussion demonstrates, school boards have the authority to permit, and even encourage, teaching about
design theory as an alternative to Darwinian evolution -- and this includes the use of textbooks such as Of
Pandas and People that present evidence for the theory of intelligent design. "

[edit] Higher education

This section needs additional citations for verification.
Please help improve this article by adding reliable references. Unsourced material may be
challenged and removed. (January 2011)
The battle to bring intelligent design and its social and political agenda the high school science
classroom is well established. Bringing intelligent design to higher education is also an active part of
Discovery Institute's strategy, though it has not taken the normal path of emergent scientific paradigms,
through graduate schools and leading professional journals of science. It has been out of the question for
intelligent design to be successfully introduced to the public via higher education venues and gain standing in
such scientific courts as long as the evidence for evolution continues to grow in the view of the scientific
community. The Discovery Institute acknowledges that if intelligent design is to become part of college and
university science curricula, it will come to campus via students, their parents, sympathetic faculty, and the
impositions of consumer-conscious college administrators. To that end the institute has supported 'IDEA'
intelligent design student groups[109] at various campuses,[110] and reports having faculty supporters on
every university campus in this country including the Ivy League schools. Academics who are Discovery
Institute fellows include Robert Kaita of Princeton University, Henry F. Schaefer of the University of Georgia,
Robert Koons and J. Budziszewski of the University of Texas at Austin, and Guillermo Gonzalez of Iowa
State University. Prominent academics who, although not officially associated with the Discovery Institute,
sympathize with its aims, include Alvin Plantinga at Notre Dame, Christopher Macosko at University of
Minnesota, Jed Macosko at Wake Forest University, and Frank Tipler at Tulane University.
A number of religious schools offer Discovery Institute-recommended curricula. Biola University and
Oklahoma Baptist University are listed on the Access Research Network website as "ID Colleges." The
intelligent design and Undergraduate Research Center, ARN’s student division, also recruits and supports
followers at universities. Campus youth ministries play an active role in bringing intelligent design to
university campuses through lectures by intelligent design leaders Phillip Johnson, William Dembski,
Jonathan Wells, Michael Behe and others. This activity takes place outside university science departments.
The few university presses (such as Cambridge and Michigan State) that have published intelligent
design books classify them as philosophy, rhetoric, or public affairs, not science.[111] There are no peer-
reviewed studies supporting intelligent design in the scientific research literature. With the scientific
community as a whole unmoved or unconvinced by proponents' works and rhetoric and the absence of
intelligent design scientific research programs, Dembski conceded that "the scientific research part" of
intelligent design is now "lagging behind" its success in influencing popular opinion.
In 2005 the American Association of University Professors issued a strongly worded statement
asserting that the theory of evolution is nearly universally accepted in the community of scholars, and
criticizing the intelligent design movement's attempts to weaken or undermine the teaching of evolution as
"inimical to principles of academic freedom."[112]
The Discovery Institute organizes on-campus intelligent design conferences across the US for
students. In the beginning, these were generally held at Christian universities and often sponsored by the
administration or other faculty as an official university function. Lateron, Yale and the University of San
Francisco have seen proponents of intelligent design speak on their campuses. Not only did these succeed
in reaching out to a more secular group of students, but the backdrop of prestigious universities achieved a
goal set forth in the Wedge strategy; to lend an aura of academic legitimacy to the proceedings and by
extension, the intelligent design movement. Commenting on the Yale conference, for example, a student
auxiliary of the Access Research Network stated, "Basically, the conference, beside being a statement (after
all we were meeting at Yale University), proved to be very promising." These conferences were not
sponsored by the universities at which they were held. They were sponsored by associated religious
organizations — at Yale, the Rivendell Institute for Christian Thought and Learning.

[edit] The Web

Much of the actual debate over intelligent design between intelligent design proponents and
members of the scientific community has taken place on the Web, primarily blogs and message boards,
instead of the scientific journals and symposia where traditionally much science is discussed and settled. In
promoting intelligent design the actions of its proponents have been more like a political pressure group than
like researchers entering an academic debate as described by movement critic Taner Edis.[113] In the
absence of any verifiable scientific research program and concomitant debates in academic circles,[114] the
most vibrant venues for intelligent design debate are websites such as Pandas Thumb [7], Dembski's blogs
at [8] and [9] and the Discovery Institute's
[10] , often with discussions and their various responses taking place on two or more sites at a time.
[improper synthesis?]
The Web continues to play a central role in the Discovery Institute's strategy of promotion of
intelligent design and it adjunct campaigns. On September 6, 2006, on the center's blog
Discovery Institute staffer Casey Luskin published a post entitled "Putting Wikipedia On Notice About Their
Biased Anti-ID Intelligent Design Entries." There Luskin reprinted a letter from a reader complaining that he
believed Wikipedia's coverage of ID to be "one sided" and that pro-intelligent design editors were censored
and attacked. Along with the letter Luskin published a Wikipedia email address for general information and
urged readers to "to contact Wikipedia to express your feelings about the biased nature of the entries on
intelligent design."[115]

[edit] International
Despite being primarily based in the United States, there have been efforts to introduce pro
Intelligent Design teaching material into educational facilities in other countries. In the United Kingdom, the
group Truth in Science has used material from the Discovery Institute to create free teaching packs which
have been mass-mailed to all UK schools.[116] Shortly after this emerged, government ministers announced
that they regarded intelligent design to be creationism and unsuitable for teaching in the classroom. They
also announced that the teaching of the material in science classes was to be prohibited.[117]

[edit] Criticisms of the movement

One of the most common criticisms of the movement and its leadership is that of intellectual
dishonesty, in the form of misleading impressions created by the use of rhetoric, intentional ambiguity, and
misrepresented evidence.[118] It is alleged that its goal is to lead an unwary public to reach certain
conclusions, and that many have been deceived as a result. Critics of the movement, such as Eugenie Scott,
Robert Pennock and Barbara Forrest, claim that leaders of the Intelligent Design movement, and the
Discovery Institute in particular, knowingly misquote scientists and other experts, deceptively omit contextual
text through ellipsis, and make unsupported amplifications of relationships and credentials. Theologian and
molecular biophysicist Alister McGrath has a number of criticisms of the Intelligent design movement, stating
that "those who adopt this approach make Christianity deeply... vulnerable to scientific progress" and defining
it as just another "god-of-the-gaps" theory. He went on to criticize the movement on theological grounds as
well, stating "It is not an approach I accept, either on scientific or theological grounds."[119]
Critics claim that the institute uses academic credentials and affiliations opportunistically. In 2001,
the Discovery Institute purchased advertisements in three national publications (the New York Review of
Books, the New Republic and the Weekly Standard) to proclaim the adherence of approximately 100
scientists to the following statement: "We are skeptical of claims for the ability of random mutation and
natural selection to account for the complexity of life. Careful examination of the evidence for Darwinian
theory should be encouraged."
Such statements commonly note the institutional affiliations of signatories for purposes of
identification. But this statement strategically listed either the institution that granted a signatory's PhD or the
institutions with which the individual is presently affiliated. Thus the institutions listed for Raymond G. Bohlin,
Fazale Rana, and Jonathan Wells, for example, were the University of Texas, Ohio University, and the
University of California, Berkeley, where they earned their degrees, rather than their current affiliations:
Probe Ministries for Bohlin, The Reasons to Believe Ministry for Rana, and The Discovery Institute's Center
for Science and Culture for Wells. Similarly confusing lists of local scientists were circulated during
controversies over evolution education in Georgia, New Mexico, Ohio, and Texas. In another instance, the
Discovery Institute frequently mentions the Nobel Prize in connection with Henry F. Schaefer, a Discovery
Institute fellow, and chemist at the University of Georgia. Critics allege that Discovery Institute is inflating his
reputation by constantly referring to him as a "five-time nominee for the Nobel Prize" because Nobel Prize
nominations remain confidential for fifty years.
This criticism is not reserved only to the institute; individual intelligent design proponents have been
accused of using their own credentials and those of others in a misleading or confusing fashion. For
example, critics allege William Dembski gratuitously invokes his laurels by boasting of his correspondence
with a Nobel laureate, bragging that one of his books was published in a series whose editors include a
Nobel laureate, and exulting that the publisher of the intelligent design book The Mystery of Life's Origin,
Philosophical Library Inc., also published books by eight Nobel laureates. Critics claim that Dembski
purposefully omits relevant facts which he fails to mention to his audience that in 1986, during the Edwards v.
Aguillard hearings, 72 Nobel laureates endorsed an amicus curiae brief that noted that the "evolutionary
history of organisms has been as extensively tested and as thoroughly corroborated as any biological
Another common criticism is that since no intelligent design research has been published in
mainstream, peer-reviewed scientific journals, the Discovery Institute often misuses the work of mainstream
scientists by putting out lists of articles that allegedly support their arguments for intelligent design drawing
from mainstream scientific literature. Often, the original authors respond that their articles cited by the center
don't support their arguments at all. Many times, the original authors have publicly refuted them for distorting
the meaning of something they've written for their own purposes.
Sahotra Sarkar, a molecular biologist at the University of Texas, has testified that intelligent design
advocates, and specifically the Discovery Institute, have misused his work by misrepresenting its conclusions
to bolster their own claims, has gone on to allege that the extent of the misrepresentations rises to the level
of professional malfeasance:[120]
"When testifying before the Texas State Board of Education in 2003 (in a battle over textbook
adoption that we won hands down), I claimed that my work had been maliciously misused by
members of the Discovery Institute. ... The trouble is that it says nothing of the sort that Meyer
claims. I don't mention Dembski, ID, or "intelligent" information whatever that may be. I don't talk
about assembly instructions. In fact what the paper essentially does is question the value of
informational notions altogether, which made many molecular biologists unhappy, but which is
also diametrically opposed to the "complex specified information" project of the ID
creationists. ... Notice how my work is being presented as being in concordance with ID when
Meyer knows very well where I stand on this issue. If Meyer were an academic, this kind of
malfeasance would rightly earn him professional censure. Unfortunately he's not. He's only the
Director of the Discovery Institute's Center for Science and Culture." -- Sahotra Sarkar

An October 2005 conference called "When Christians and Cultures Clash" was held at the
Pennsylvania Evangelical School of Theology. Attorney Randy Wenger, who is affiliated with the Alliance
Defense Fund, and a close ally of the Discovery Institute, and one of the presenters at the conference
advocated the use of subterfuge for advancing the movement's religious goals: "But even with God’s
blessing, it’s helpful to consult a lawyer before joining the battle. For instance, the Dover area school board
might have had a better case for the intelligent design disclaimer they inserted into high school biology
classes had they not mentioned a religious motivation at their meetings. Give us a call before you do
something controversial like that, I think we need to do a better job at being clever as serpents."[121]

[edit] See also

• Intelligent design in politics
• Kitzmiller v. Dover Area School District
• Expelled: No Intelligence Allowed
• Wedge strategy
• Intelligent Design and Evolution Awareness Center
• Intelligent design network

[edit] External links

• Discovery Institute - Center for Science and Culture
• The Wedge Breaking the Modernist Monopoly on Science by Phillip E. Johnson. Touchstone,
A Journal of Mere Christianity. Volume 12, Issue 4. July/August 1999
• The Wedge at Work: How Intelligent Design Creationism Is Wedging Its Way into the Cultural
and Academic Mainstream Chapter 1 of the book Intelligent Design Creationism and Its Critics by
Barbara Forrest, Ph.D. MIT Press, 2001
• Does "Intelligent Design" Threaten the Definition of Science? John Roach. National
Geographic News. April 27, 2005
• Over 500 Scientists Proclaim Their Doubts About Darwin’s Theory Discovery Institute
February 20, 2006
• Design & the Discriminating Public Gaining a Hearing from Ordinary People by Nancy
Pearcey. Touchstone, A Journal of Mere Christianity. Volume 12, Issue 4. July/August 1999
• The Newest Evolution of Creationism Intelligent design is about politics and religion, not
science. Barbara Forrest Ph.D. From Natural History, April, 2002, page 80
• "Inferior Design" Chris Mooney, The American Prospect Online, 10 August 2005.
• Show Me the Science Daniel C. Dennett. At Tufts University, An editorial originally
published in the New York Times. 28 August 2005
• Should Creationism Be Taught in the Public Schools? (PDF) Robert T. Pennock. March 2002
• Politicized Scholars Put Evolution on the Defensive Jody Wilgoren. Originally published in
the New York Times. 21 August 2005
• Intelligent design opponents invoke US constitution . Donald MacLeod, The Guardian, 18
October 2005.
• Devolution - Why intelligent design isn’t. H. Allen Orr. Annals of Science. New Yorker. May
• Intelligent design network .org
• Intelligent Judging — Evolution in the Classroom and the Courtroom George J. Annas, J.D.,
M.P.H. New England Journal of Medicine. Volume 354:2277-2281, Number 21 May 25, 2006
• The Political Design of Intelligent Design Russell D. Renka, Professor of Political Science,
Southeast Missouri State University November, 2005
• Intelligent Design vs. Evolution debate between paleontologist Peter Ward and Stephen
Meyer co-founder of the Discovery Institute
• Haran Yahya Islamic creationist site
• Intelligent Design Network Australia
• Finnish Intelligent Design site
• Italian Intelligent Design site

[edit] Notes
1. ^ Creationism's Trojan Horse Forrest & Gross, p7
2. ^ Forrest, Barbara (May,2007) (PDF). Understanding the Intelligent Design Creationist
Movement: Its True Nature and Goals. A Position Paper from the Center for Inquiry, Office of Public
Policy. Washington, D.C.: Center for Inquiry, Inc.. Retrieved 2007-08-06 .
3. ^ a b Wedge Strategy Discovery Institute, 1999.
4. ^ Barbara Forrest. 2001. "The Wedge at Work: Intelligent Design Creationism and Its Critics
5. ^ "An Objective Observer Would Know that ID and Teaching About 'Gaps' and 'Problems' in
Evolutionary Theory are Creationist, Religious Strategies that Evolved from Earlier Forms of
Creationism: The concept of intelligent design (hereinafter "ID"), in its current form, came into
existence after the Edwards case was decided in 1987. For the reasons that follow, we conclude that
the religious nature of ID would be readily apparent to an objective observer, adult or child." (page
18) "...we find that ID's religious nature would be further evident to our objective observer because it
directly involves a supernatural designer." (page 24) "A 'hypothetical reasonable observer,' adult or
child, who is 'aware of the history and context of the community and forum' is also presumed to know
that ID is a form of creationism. (page 31) "The evidence at trial demonstrates that ID is nothing less
than the progeny of creationism." (page 31) Context Ruling, Kitzmiller v. Dover Area School District.
Pages 17-35
6. ^ "Q. Has the Discovery Institute been a leader in the intelligent design movement? A. Yes,
the Discovery Institute's Center for Science and Culture. Q. And are almost all of the individuals who
are involved with the intelligent design movement associated with the Discovery Institute? A. All of
the leaders are, yes." Barbara Forrest, 2005, testifying in the Kitzmiller v. Dover Area School District
trial. "Kitzmiller v. Dover Area School District Trial transcript: Day 6 (October 5), PM Session, Part
1.". The TalkOrigins Archive. 2005. Retrieved
• "The Discovery Institute is the ideological and strategic backbone behind the eruption of skirmishes
over science in school districts and state capitals across the country." In: Wilgoren, J (2005-08-21).
"Politicized Scholars Put Evolution on the Defensive" (PDF). The New York Times.
%20Evolution%20on%20the%20Defensive%20-%20New%20York%20Times.pdf. Retrieved 2007-
• "Who is behind the ID movement?". Frequently Asked Questions About "Intelligent Design" .
American Civil Liberties Union. 2005-09-16. Retrieved 2007-07-20.
• Kahn, JP (2005-07-27). "The Evolution of George Gilder. The Author And Tech-Sector Guru Has A
New Cause To Create Controversy With: Intelligent Design". The Boston Globe.
%20-%20News&id=2745. Retrieved 2007-07-19.
• "Who's Who of Intelligent Design Proponents". Science & Religion Guide. Science & Theology
News. November 2005.
command=download&id=602. Retrieved 2007-07-20. (PDF file from Discovery Institute).
• "The engine behind the ID movement is the Discovery Institute." Attie, Alan D.; Elliot Sober, Ronald
L. Numbers, Richard M. Amasino, Beth Cox4, Terese Berceau, Thomas Powell and Michael M. Cox
(2006). "Defending science education against intelligent design: a call to action". Journal of Clinical
Investigation 116:1134–1138. doi:10.1172/JCI28449. A publication of the American Society for
Clinical Investigation.. Retrieved 2007-07-20.
7. ^ "Science and Policy: Intelligent Design and Peer Review". American Association for the
Advancement of Science. 2007. Retrieved 2007-07-19.
8. ^ AAAS Denounces Anti-Evolution Laws American Association for the Advancement of
Science News, February 19, 2006.
9. ^ Ruling, Kitzmiller v. Dover, page 70
10.^ a b c Ruling, Kitzmiller v. Dover page 83
11.^ Defending science education against intelligent design: a call to action Journal of Clinical
Investigation 116:1134-1138 (2006). doi:10.1172/JCI28449. A publication of the American Society for
Clinical Investigation.
12.^ Patricia O’Connell Killen, a religion professor at Pacific Lutheran University in Tacoma
whose work centers around the regional religious identity of the Pacific Northwest, recently wrote that
"religiously inspired think tanks such as the conservative evangelical Discovery Institute" are part of
the "religious landscape" of that area. [1]
13.^ The Wedge Strategy Center for the Renewal of Science and Culture. 1998
14.^ The Design Revolution: Answering the Toughest Questions about Intelligent Design
William A. Dembski. Intervarsity Press, 2004.
15.^ Why scientists dismiss 'intelligent design' Ker Than. MSNBC, September 23, 2005.
16.^ Q&A: Darwin on Trial Margaret Talbot. The New Yorker, November 28, 2005.
17.^ AAAS Board Resolution on Intelligent Design Theory American Association for the
Advancement of Science.
18.^ National Academy of Sciences, 1999 Science and Creationism: A View from the National
Academy of Sciences, Second Edition
19.^ DI's New Talking Point Ed Brayton. Dispatches from the Culture Wars, December 11, 2006.
20.^ The Wedge at Work: How Intelligent Design Creationism Is Wedging Its Way into the
Cultural and Academic Mainstream Barbara Forrest. Chapter 1 of the book Intelligent Design
Creationism and Its Critics (MIT Press, 2001).
21.^ The Evolution Wars Claudia Wallis. TIME magazine. August 15, 2005.
22.^ Reclaim America .org
23.^ a b The Evolution Debate Can Be Won. Phillip Johnson. Truths that Transform.
24.^ Jonathan Witt. Discovery Institute. Evolution News & Views: Dover Judge Regurgitates
Mythological History of Intelligent Design; December 20, 2005 [cited 2007-10-05].
25.^ Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005)., pp. 31 – 33.
26.^ Nick Matzke. National Center for Science Education. NCSE Resource -- 9.0. Matzke
(2006): The Story of the Pandas Drafts; 2006 [cited 2009-11-18]. *Nick Matzke. National Center for
Science Education. Missing Link discovered!; 2006 [archived 2007-01-14; cited 2009-11-18].
27.^ Book thrown at proponents of Intelligent Design Celeste Biever., October
6, 2005,
28.^ Kitzmiller v Dover day 6 a.m. (Barbara Forrest's testimony)
29.^ Stewart, Robert (2007). Intelligent design: William A. Dembski Michael Ruse in dialogue .
Minneapolis, MN: Fortress Press. pp. 2. ISBN 0-8006-6218-0.
30.^ Gould SJ (1992). "Impeaching a Self-Appointed Judge". Scientific American 267 (1). Retrieved 2009-04-01.
31.^ Forrest & Gross(2004), p18
32.^ Numbers(2006) p380
33.^ a b Numbers(2006) pp381-382
34.^ Forrest&Gross(2004) p19
35.^ Numbers(2006) p377
36.^ Forrest&Gross(2004) pp 25-29
37.^ The Wedge Document: So What? Discovery Institute. (PDF file)
38.^ Defeating Darwinism by Opening Minds Phillip Johnson. pg. 91-92, 1997.
39.^ Transcripts, Kansas Evolution Hearings Page 6. Via Talk Origins Archive.
40.^ A Real Monkey Trial Peter Dizikes. Salon, May 13, 2005.
41.^ CSC - Key Resources for Parents and School Board Members
42.^ 6News Lawrence: Some question group's move with elections nearing
43.^ Wichita Eagle, "Scientists Right to Boycott Evolution Hearings," March 30, 2005; "Evolution
Hearings Rejected by Scientists," April 12, 2005.
44.^ Reason Magazine - Unintelligent Design
45.^ [2].
46.^ "Evolution of Kansas science standards continues as Darwin's theories regain
prominence". International Herald-Tribune. 2007-02-13.
Retrieved 2007-02-14.
47.^ Intelligent designer Gordy Slack., October 2005.
48.^ "For years, a lawyer for the Thomas More Law Center in Michigan visited school boards
around the country searching for one willing to challenge evolution by teaching intelligent design, and
to face a risky, high-profile trial." In Intelligent Design Case, a Cause in Search of a Lawsuit Laurie
Goodstein. The New York Times, November 4, 2005.
49.^ "TMLC representatives traveled the country from at least early 2000, encouraging school
boards to teach ID in science classrooms. From Virginia to Minnesota, TMLC recommended the
textbook Of Pandas and People (Pandas) as a supplement to regular biology textbooks, promising to
defend the schools free of charge when the ACLU filed the inevitable lawsuit. Finally, in summer
2004, they found a willing school board in Dover, Pennsylvania, a board known to have been
searching for a way to get creationism inserted into its science classrooms for years. " Kitzmiller et al.
versus Dover Area School District Burt Humburg, Ed Brayton. Skeptic magazine, July–December
50.^ Seattle's Discovery Institute scrambling to rebound after intelligent-design ruling, David
Postman, The Seattle Times, April 26, 2006
51.^ a b Discovery Institute and Thomas More Law Center Squabble in AEI Forum, October
23rd, 2005 National Center for Science Education
52.^ ADF attorneys seek to supply missing link in intelligent design curriculum case Alliance
Defense Fund, May 24, 2005.
53.^ Plaintiffs' Response to Amicus Briefs, Kitzmiller v. Dover Area School District
54.^ Circuit Court Sends 'Textbook Sticker' Case Back to Lower Court at the Wayback Machine
(archived August 22, 2006)., Jim Brown, 2006Agape Press, June 1, 2006
55.^ Focus on religion a central ADF tenet, Michael Moore, Missoulian, 29 February 2004
56.^ Nuisance Lawsuit Against Scott and NCSE Withdrawn, Eugenie Scott, The Pandas Thumb,
September 14, 2005
57.^ Caldwell vs Roseville, NO. CIV. S-05-0061 FCD JFM September 17, 2007
58.^ Court dismisses lawsuit targeting evolution website, Robert Sanders, UC Berkeley News,
15 March 2006
59.^ "Whether educational authorities allow the schools to teach about the controversy or not,
public recognition that there is something seriously wrong with Darwinian orthodoxy is going to keep
on growing. While the educators stonewall, our job is to continue building the community of people
who understand the difference between a science that tests its theories against the evidence, and a
pseudoscience that protects its key doctrines by imposing philosophical rules and erecting legal
barriers to freedom of thought. The Pennsylvania Controversy Phillip E. Johnson, Phillip Johnson's
Weekly Wedge Update. June 11, 2001
60.^ "If the science educators continue to pretend that there is no controversy to teach, perhaps
the television networks and the newspapers will take over the responsibility of informing the public."
Icons of Evolution exposed on CNN Phillip E. Johnson, Phillip Johnson's Weekly Wedge Update.
May 2001
61.^ "If the public school educators will not "teach the controversy," our informal network can do
the job for them. In time, the educators will be running to catch up." Passing the Torch Phillip E.
Johnson, Phillip Johnson's Weekly Wedge Update. April 9, 2002
62.^ Privileged Planet - New science documentary explores Earth’s extraordinary place in the
cosmos Staff, Discovery Institute, August 20, 2004
63.^ Unlocking the Mystery of Life - Documentary reveals growing number of scientific
challenges to Darwinian evolution Stephen C. Meyer and W. Peter Allen. Illustra Media, July 15,
64.^ Ruling - whether ID is science, pg.83 Kitzmiller v. Dover Area School District.
65.^ Ruling - whether ID is science, pg.89 Kitzmiller v. Dover Area School District.
66.^ Ruling - disclaimer, pg. 49 Kitzmiller v. Dover Area School District.
67.^ a b Public Divided on Origins of Life The Pew Forum on Religion & Public Life. August 30,
68.^ Ohio Scientists' Intelligent Design Poll Internet Public Opinion Laboratory, Department of
Political Science University of Cincinnati. October 2002.
69.^ A Scientific Dissent from Darwinism Discovery Institute — Center for Science and Culture
accessed July 27, 2006
70.^ Few Biologists But Many Evangelicals Sign Anti-Evolution Petition Kenneth Chang. The
New York Times, February 21, 2006
71.^ The List of Steves, National Center for Science Education
72.^ Life In The Big Tent: Traditional Creationism And The Intelligent Design Community Paul A.
Nelson. Christian Research Institute. (PDF file)
73.^ Small Group Wields Major Influence in Intelligent Design Debate ABC News, November 9,
74.^ "A theistic realist assumes that the universe and all its creatures were brought into
existence for a purpose by God. Theistic realists expect this "fact" of creation to have empirical,
observable consequences that are different from the consequences one would observe if the
universe were the product of nonrational causes . . . . God always has the option of working through
regular secondary mechanisms, and we observe such mechanisms frequently. On the other hand,
many important questions--including the origin of genetic information and human consciousness--
may not be explicable in terms of unintelligent causes, just as a computer or a book cannot be
explained that way." Phillip Johnson. Reason in the Balance: The Case Against Naturalism in
Science, Law and Education. 1995. InterVarsity Press pg. 208-209.
75.^ "Intelligent Design is an intellectual movement, and the Wedge strategy stops working
when we are seen as just another way of packaging the Christian evangelical message. ... The
evangelists do what they do very well, and I hope our work opens up for them some doors that have
been closed." Phillip Johnson. "Keeping the Darwinists Honest", an interview with Phillip Johnson. In
Citizen Magazine. April 1999.
76.^ "...the first thing that has to be done is to get the Bible out of the discussion. ...This is not to
say that the biblical issues are unimportant; the point is rather that the time to address them will be
after we have separated materialist prejudice from scientific fact." Phillip Johnson. "The Wedge",
Touchstone: A Journal of Mere Christianity. July/August 1999.
77.^ About The CRSC recovered from the Internet Archive.
78.^ About the CSC Discovery Institute
79.^ Avenging angel of the religious right Max Blumenthal., January 1, 2004.
80.^ Theocratic Dominionism Gains Influence Part 3 - No Longer Without Sheep Frederick
Clarkson. The Public Eye Magazine, Vol. 8, No. 1. Political Research Associates, March/June 1994.
81.^ a b Barbara Forrest and Paul R. Gross, Creationism's Trojan Horse. Oxford University
Press, (January 8, 2004) ISBN 0195157427 (pp 165-167)
82.^ "About the Access Research Network". Access Research Network. 2008. Retrieved 2008-05-17.
83.^ "Luskin explained that as a Christian group, 'we wanted to be totally open about who we
thought the designer was.'" Intelligent Design Gains Momentum, Raises Eyebrows on Campuses at
the Wayback Machine (archived September 2, 2006).
85.^ The "Intelligent Design" Movement on College and University Campuses is Dead, Allen
86.^ Intelligent Design: The Scientific Alternative to Evolution , John H. Calvert and William S.
Harris, National Catholic Bioethics Quarterly, Autumn 2003
87.^ The Rule, Daniel Schwabauer and John Calvert
88.^ Kitzmiller v. Dover: July 14 Hearing: Jon A. Buell
89.^ Nearly Two-thirds of U.S. Adults Believe Human Beings Were Created by God The Harris
Poll #52, July 6, 2005.
90.^ Mooney, Chris (2003-09-11). "Polling for ID". Doubt and About. Committee for Skeptical
Inquiry. Archived from the original on February 3, 2007.
Retrieved 2007-02-16.
91.^ a b Testimony, Aralene Callahan Kitzmiller v. Dover Area School District September 27,
92.^ a b Testimony, Julie Smith Kitzmiller v. Dover Area School District September 28, 2005
93.^ a b Defending science education against intelligent design: a call to action Journal of
Clinical Investigation 116:1134-1138 (2006). American Society for Clinical Investigation.
94.^ Dover gets a million-dollar bill Christina Kauffman. The York Dispatch, February 22, 2006
95.^ Is It Science Yet?: Intelligent Design Creationism And The Constitution Matthew J. Brauer,
Barbara Forrest, Steven G. Gey. Washington University Law Quarterly, Volume 83, Number 1, 2005.
(PDF file)
96.^ The Group for the Scientific Reappraisal of the HIV-AIDS Hypothesis
97.^ "His personal peculiarities include membership in the Moonies and support for AIDS
reappraisal - the theory that the HIV is not the primary cause of AIDS" Undercover at the Discovery
Institute Beth Quittman. Seattlest, September 8, 2006.
98.^ "some leading lights of anti-evolution Intelligent Design theory, including ID godfather
Phillip Johnson and Moonie Jonathan Wells, have joined the AIDS denialist camp." AIDS 'Denialism'
Gathers Strange Bedfellows Peter McKnight. Originally published in the Vancouver Sun, June 17,
99.^ Understanding the Intelligent Design Creationist Movement: Its True Nature and Goals. A
Position Paper from the Center for Inquiry, Office of Public Policy Barbara Forrest. May, 2007.
100.^ "99.9 percent of scientists accept evolution" Finding the Evolution in Medicine National
Institutes of Health
101.^ Teaching Evolution: A State-by-State Debate National Public Radio, December 20, 2005.
102.^ The Political Design of Intelligent Design Russell D. Renka, Professor of Political Science.
Southeast Missouri State University. November 16, 2005
103.^ The institute spends more than $1 million a year for research, polls, lobbying and media
pieces that support intelligent design and their Teach the Controversy campaign Battle on Teaching
Evolution Sharpens Peter Slevin Washington Post, March 14, 2005, and is employing the same
Washington, D.C. public relations firm that promoted the Contract with America Politicized Scholars
Put Evolution on the Defensive By Jodi Wilgoren, New York Times, August 21, 2005
104.^ Faculty Association Speaks Out on Three Top Issues American Association of University
Professors. June 17, 2005.
105.^ "Moreover, Board members and teachers opposing the curriculum change and its
implementation have been confronted directly. First, Casey Brown testified that following her
opposition to the curriculum change on October 18, 2004, Buckingham called her an atheist and
Bonsell told her that she would go to hell. Second, Angie Yingling was coerced into voting for the
curriculum change by Board members accusing her of being an atheist and un- Christian. In addition,
both Bryan Rehm and Fred Callahan have been confronted in similarly hostile ways, as have
teachers in the DASD." Ruling, conclusion - Effect of Board’s Actions on Plaintiffs Kitzmiller v. Dover
Area School District.
106.^ In July 2006 a moderator of the blog of intelligent design proponent William A. Dembski,, endorsed bullying the children of the plaintiffs in the Kitzmiller v. Dover Area
School District trial and committing vandalism to drive them out of town and that he intends to publish
their names on the Web to that end.[3][4][5][6]
107.^ Science Wars Should Schools Teach Intelligent Design? Video of American Enterprise
Institute forum that took place during the Kitzmiller case, originally broadcast on C-SPAN
108.^ Intelligent Design in Public School Science Curricula: A Legal Guidebook Access
Research Network
109.^ Intelligent Design and Evolution Awareness
110.^ IDEA chapter locations
111.^ Darwinism, Design, and Public Education, Rhetoric and Public Affairs Series, Michigan
State University Press
112.^ Faculty Association Speaks Out on Three Top Issues at the Wayback Machine (archived
February 10, 2006)., American Association of University Professors, June 17, 2005
113.^ Why ID Fails Taner Edis. 2005.
114.^ The Wedge at Work Chapter 1 of Intelligent Design Creationism and Its Critics . Barbara
Forrest. MIT Press, 2001.
115.^ Putting Wikipedia On Notice About Their Biased Anti-ID Intelligent Design Entries Casey
Luskin., September 6, 2006.
116.^ Revealed: rise of creationism in the UK Guardian Unlimited. November 27, 2006.
117.^ Ministers to ban creationist teaching aids in science lessons Guardian Unlimited.
December 7, 2006.
118.^ "ID supporters present fallacious arguments, use dishonest rhetoric, and often present
non-contemptuous responses as evidence that their theories are gaining acceptance." Leaders and
Followers in the Intelligent Design Movement Jason Rosenhouse. BioScience, Vol. 53 No. 1, January
119.^ McGrath, A: The Dawkins Delusion?, page 30. InterVarsity Press, 2007.
120.^ Fraud from the Discovery Institute Sahotra Sarkar. Sarkar Lab WebLog. December 3,
121.^ "'Bring us your legal issues,' clergy told" Daniel Burke. Lancaster New Era, October 20,

[edit] References
• Forrest, Barbara; Gross, Paul R. (8 January 2004). Creationism's Trojan Horse. Oxford
University Press. ISBN 0195157427.
• Numbers, Ronald (November 30, 2006). The Creationists: From Scientific Creationism to
Intelligent Design, Expanded Edition. Harvard University Press. ISBN 0674023390.
Retrieved from ""

Categories: Intelligent design movement | Intelligent design advocates | Religion and politics |
Religion and education | Discovery Institute campaigns

Evolution of sexual reproduction

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The evolution of sexual reproduction is currently described by several competing scientific
hypotheses. Many groups of organisms, notably the majority of animals and plants, reproduce sexually. The
evolution of sex contains two related, yet distinct, themes: its origin and its maintenance. However, since the
hypotheses for the origins of sex are difficult to test experimentally, most current work has been focused on
the maintenance of sexual reproduction.
It seems that a sexual cycle is maintained because it improves the quality of progeny (fitness),
despite reducing the overall number of offspring (the twofold cost of sex). In order for sex to be evolutionarily
advantageous, it must be associated with a significant increase in the fitness of offspring. One of the most
widely accepted explanations for the advantage of sex lies in the creation of genetic variation. Another
explanation is based on two molecular advantages. First is the advantage of recombinational DNA repair
(promoted during meiosis because homologous chromosomes pair at that time), and second is the
advantage of complementation (also known as hybrid vigor, heterosis or masking of mutations).
For the advantage due to creation of genetic variation, there are three possible reasons this might
happen. First, sexual reproduction can bring together mutations that are beneficial into the same individual
(sex aids in the spread of advantageous traits). Second, sex acts to bring together currently deleterious
mutations to create severely unfit individuals that are then eliminated from the population ( sex aids in the
removal of deleterious genes). Last, sex creates new gene combinations that may be more fit than previously
existing ones, or may simply lead to reduced competition among relatives.
For the advantage due to DNA repair, there is an immediate large benefit to removal of DNA damage
by recombinational DNA repair during meiosis, since this removal allows greater survival of progeny with
undamaged DNA. The advantage of complementation to each sexual partner is avoidance of the bad effects
of their deleterious recessive genes in progeny by the masking effect of normal dominant genes contributed
by the other partner.
The classes of hypotheses based on the creation of variation are further broken down below. It is
important to realise that any number of these hypotheses may be true in any given species (they are not
mutually exclusive), and that different hypotheses may apply in different species. However, a research
framework based on creation of variation has yet to be found that allows one to determine whether the
reason for sex is universal for all sexual species, and, if not, which mechanism is acting in each species.
On the other hand, the maintenance of sex based on DNA repair and complementation applies
widely to all sexual species. This explanation for the maintenance of sex is explored further in Section 6.2.
• 1 Historical perspective
• 2 Two-fold cost of sex
• 3 Promotion of genetic variation
• 4 Spread of advantageous traits
• 4.1 Novel genotypes
• 4.2 Increased resistance to parasites
• 5 Deleterious mutation clearance
• 5.1 Maintenance of mutation-free individuals
• 5.2 Removal of deleterious genes
• 6 Other explanations
• 6.1 Speed of evolution
• 6.2 DNA repair and complementation
• 7 Origin of sexual reproduction
• 8 Mechanistic origin of sexual reproduction
• 8.1 Viral eukaryogenesis
• 8.2 Neomuran revolution
• 9 See also
• 10 References
• 11 Further reading
• 12 External links
[edit] Historical perspective
Modern philosophical-scientific thinking on the problem can be traced back to Erasmus Darwin in the
18th century; it also features in Aristotle's writings. The thread was later picked up by August Weismann in
1889, who understood that the purpose of sex was to generate genetic variation, as is detailed in the majority
of the explanations below. On the other hand, Charles Darwin, also in 1889, concluded that the effects of
hybrid vigor (complementation) “is amply sufficient to account for the ... genesis of the two sexes.” This is
consistent with the repair and complementation hypothesis, given below under “Other explanations.”
Several explanations have been suggested by biologists including W. D. Hamilton, Alexey
Kondrashov, George C. Williams, Harris Bernstein, Carol Bernstein, Michael M. Cox, Frederic A. Hopf and
Richard E. Michod to explain how sexual reproduction is maintained in a vast array of different living

[edit] Two-fold cost of sex

This diagram illustrates the twofold cost of sex. If each individual were to contribute to the same
number of offspring (two), (a) the sexual population remains the same size each generation, where the (b)
asexual population doubles in size each generation.
In all multicellular sexual species, the population consists of two sexes, only one of which is capable
of bearing young (with the exception of simultaneous hermaphrodites). In an asexual species, each member
of the population is capable of bearing young. This implies that an asexual population has an intrinsic
capacity to grow more rapidly each generation. The cost was first described in mathematical terms by John
Maynard Smith.[1] He imagined an asexual mutant arising in a sexual population, half of which comprises
males that cannot themselves produce offspring. With female-only offspring, the asexual lineage doubles its
representation in the population each generation, all else being equal. Often all else is not equal, however, in
which case the realised fitness cost to sex may be much less than this intrinsic twofold cost of producing
males. For example, an asexual mutant arising in a sexual population occupies a niche frozen to that of its
parental genotype because the asexual descendants are genetically self-identical. Analysis of competitive
Lotka-Volterra equations suggests that the asexual lineage may never realise its full twofold advantage in
population growth capacity, if the broader niche of the sexual population confers even a small competitive
An additional cost is that males and females must search for each other in order to mate, and sexual
selection often favours traits that reduce the survival of individuals.[1]
Evidence that the cost is not insurmountable comes from George C. Williams, who noted the
existence of species which are capable of both asexual and sexual reproduction. These species time their
sexual reproduction with periods of environmental uncertainty, and reproduce asexually when conditions are
more favourable. The important point is that these species are observed to reproduce sexually when they
could choose not to, implying that there is a selective advantage to sexual reproduction.[3]
Another disadvantage of sexual reproduction is that a sexually reproducing organism will only be
able to pass on 50% of its genes to each offspring. This is a consequence of the fact that gametes from
sexually reproducing species are haploid.[4]

[edit] Promotion of genetic variation

August Weismann proposed in 1889[5] an explanation for the evolution of sex, where the advantage
of sex is the creation of variation among siblings. It was then subsequently explained in genetics terms by
Fisher[6] and Muller[7] and has been recently summarised by Burt in 2000.[8]
George C. Williams gave an example based around the elm tree. In the forest of this example, empty
patches between trees can support one individual each. When a patch becomes available because of the
death of a tree, other trees' seeds will compete to fill the patch. Since the chance of a seed's success in
occupying the patch depends upon its genotype, and a parent cannot anticipate which genotype is most
successful, each parent will send many seeds, creating competition between siblings. Natural selection
therefore favours parents which can produce a variety of offspring.
A similar hypothesis is named the tangled bank hypothesis after a passage in Charles Darwin's The
Origin of Species:
"It is interesting to contemplate an entangled bank, clothed with many plants of many kinds,
with birds singing on the bushes, with various insects flitting about, and with worms crawling through
the damp earth, and to reflect that these elaborately constructed forms, so different from each other,
and dependent on each other in so complex a manner, have all been produced by laws acting around
The hypothesis, proposed by Michael Ghiselin in his 1974 book, The Economy of Nature and the
Evolution of Sex, suggests that a diverse set of siblings may be able to extract more food from its
environment than a clone, because each sibling uses a slightly different niche. One of the main proponents of
this hypothesis is Graham Bell of McGill University. The hypothesis has been criticised for failing to explain
how asexual species developed sexes. In his book, Evolution and Human Behavior (MIT Press, 2000), John
Cartwright comments:
"Although once popular, the tangled bank hypothesis now seems to face many problems, and
former adherents are falling away. The theory would predict a greater interest in sex among animals
that produce lots of small offspring that compete with each other. In fact, sex is invariably associated
with organisms that produce a few large offspring, whereas organisms producing small offspring
frequently engage in parthenogenesis [asexual reproduction]. In addition, the evidence from fossils
suggests that species go for vast periods of [geologic] time without changing much."
[edit] Spread of advantageous traits
[edit] Novel genotypes

This diagram illustrates how sex might create novel genotypes more rapidly. Two advantageous
alleles A and B occur at random. The two alleles are recombined rapidly in a sexual population (top), but in
an asexual population (bottom) the two alleles must independently arise because of clonal interference.
Sex could be a method by which novel genotypes are created. Since sex combines genes from two
individuals, sexually reproducing populations can more easily combine advantageous genes than can
asexual populations. If, in a sexual population, two different advantageous alleles arise at different loci on a
chromosome in different members of the population, a chromosome containing the two advantageous alleles
can be produced within a few generations by recombination. However, should the same two alleles arise in
different members of an asexual population, the only way that one chromosome can develop the other allele
is to independently gain the same mutation, which would take much longer.
Ronald Fisher also suggested that sex might facilitate the spread of advantageous genes by allowing
them to escape their genetic surroundings, if they should arise on a chromosome with deleterious genes.
But these explanations depend upon the rate of mutation. If favourable mutations are so rare that
each will become fixed in the population before the next arises (bearing in mind that mutation is a Poisson
process), then sexual and asexual populations would evolve at the same rate.
Additionally, these explanations depend upon group selection, which most theorists maintain is a
weak selective force relative to individual selection – sex is still disadvantageous to the individual due to the
twofold cost of sex. Therefore, these explanations do not explain why heterogonic species choose to adopt
sexual reproduction, as George C. Williams pointed out in his balance argument, and hence are insufficient
to explain the evolution of sex.
Supporters of these theories respond to the balance argument that the individuals produced by
sexual and asexual reproduction may differ in other respects too – which may influence the persistence of
sexuality. For example, in water fleas of the genus Cladocera, sexual offspring form eggs which are better
able to survive the winter.

[edit] Increased resistance to parasites

One of the most widely accepted theories to explain the persistence of sex is that it is maintained to
assist sexual individuals in resisting parasites, also known as the Red Queen's Hypothesis.[4][9][10]
When an environment changes, previously neutral or deleterious alleles can become favourable. If
the environment changed sufficiently rapidly (i.e. between generations), these changes in the environment
can make sex advantageous for the individual. Such rapid changes in environment are caused by the co-
evolution between hosts and parasites.
Imagine, for example that there is one gene in parasites with two alleles p and P conferring two types
of parasitic ability, and one gene in hosts with two alleles h and H, conferring two types of parasite
resistance, such that parasites with allele p can attach themselves to hosts with the allele h, and P to H. Such
a situation will lead to cyclic changes in allele frequency - as p increases in frequency, h will be disfavoured.
In reality, there will be several genes involved in the relationship between hosts and parasites. In an
asexual population of hosts, offspring will only have the different parasitic resistance if a mutation arises. In a
sexual population of hosts, however, offspring will have a new combination of parasitic resistance alleles.
In other words, like Lewis Carroll's Red Queen, sexual hosts are continually adapting in order to stay
ahead of their parasites.
Evidence for this explanation for the evolution of sex is provided by comparison of the rate of
molecular evolution of genes for kinases and immunoglobulins in the immune system with genes coding
other proteins. The genes coding for immune system proteins evolve considerably faster.[11][12]
Critics of the Red Queen hypothesis question whether the constantly-changing environment of hosts
and parasites is sufficiently common to explain the evolution of sex.

[edit] Deleterious mutation clearance

Mutations can have many different effects upon an organism. It is generally believed that the majority
of non-neutral mutations are deleterious, which means that they will cause a decrease in the organism's
overall fitness.[13] If a mutation has a deleterious effect, it will then usually be removed from the population
by the process of natural selection. Sexual reproduction is believed to be more efficient than asexual
reproduction in removing those mutations from the genome.[14]
There are two main hypotheses which explain how sex may act to remove deleterious genes from
the genome.

[edit] Maintenance of mutation-free individuals

Main article: Muller's ratchet
In a finite asexual population under the pressure of deleterious mutations, the random loss of
individuals without such mutations is inevitable. This is known as Muller's ratchet. In a sexual population,
however, mutation-free genotypes can be restored by recombination of genotypes containing deleterious
This comparison will only work for a small population - in a large population, random loss of the most
fit genotype becomes unlikely even in an asexual population.
[edit] Removal of deleterious genes

Diagram illustrating different relationships between numbers of mutations and fitness. Kondrashov's
model requires synergistic epistasis, which is represented by the blue line - each mutation has a
disproproportionately large effect on the organism's fitness.
This hypothesis was proposed by Alexey Kondrashov, and is sometimes known as the deterministic
mutation hypothesis.[14] It assumes that the majority of deleterious mutations are only slightly deleterious,
and affect the individual such that the introduction of each additional mutation has an increasingly large effect
on the fitness of the organism. This relationship between number of mutations and fitness is known as
synergistic epistasis.
By way of analogy, think of a car with several minor faults. Each is not sufficient alone to prevent the
car from running, but in combination, the faults combine to prevent the car from functioning.
Similarly, an organism may be able to cope with a few defects, but the presence of many mutations
could overwhelm its backup mechanisms.
Kondrashov argues that the slightly deleterious nature of mutations means that the population will
tend to be composed of individuals with a small number of mutations. Sex will act to recombine these
genotypes, creating some individuals with fewer deleterious mutations, and some with more. Because there
is a major selective disadvantage to individuals with more mutations, these individuals die out. In essence,
sex compartmentalises the deleterious mutations.
There has been much criticism of Kondrashov's theory, since it relies on two key restrictive
conditions. The first requires that the rate of deleterious mutation should exceed one per genome per
generation in order to provide a substantial advantage for sex. While there is some empirical evidence for it
(for example in Drosophila[15] and E. coli[16]), there is also strong evidence against it.[ citation needed]
Secondly, there should be strong interactions among loci (synergistic epistasis), a mutation-fitness relation
for which there is only limited evidence. Conversely, there is also the same amount of evidence that
mutations show no epistasis (purely additive model) or antagonistic interactions (each additional mutation
has a disproportionally small effect).

[edit] Other explanations

[edit] Speed of evolution
Ilan Eshel suggested that sex prevents rapid evolution. He suggests that recombination breaks up
favourable gene combinations more often than it creates them, and sex is maintained because it ensures
selection is longer-term than in asexual populations - so the population is less affected by short-term
changes.[citation needed] This explanation is not widely accepted, as its assumptions are very restrictive.
It has recently been shown in experiments with Chlamydomonas algae that sex can remove the
speed limit on evolution.[17]

[edit] DNA repair and complementation

As discussed in the earlier part of this article, sexual reproduction is conventionally explained as an
adaptation for producing genetic variation through allelic recombination. As acknowledged above, however,
serious problems with this explanation have led many biologists to conclude that the benefit of sex is a major
unsolved problem in evolutionary biology.
An alternative “informational” approach to this problem has led to the view that the two fundamental
aspects of sex, genetic recombination and outcrossing, are adaptive responses to the two major sources of
“noise” in transmitting genetic information. Genetic noise can occur as either physical damage to the genome
(e.g. chemically altered bases of DNA or breaks in the chromosome) or replication errors (mutations)[18][19]
[20] This alternative view is referred to as the repair and complementation hypothesis, to distinguish it from
the traditional variation hypothesis.
The repair and complementation hypothesis assumes that genetic recombination is fundamentally a
DNA repair process, and that when it occurs during meiosis it is an adaptation for repairing the genomic DNA
which is passed on to progeny. Recombinational repair is the only repair process known which can
accurately remove double-strand damages in DNA, and such damages are both common in nature and
ordinarily lethal if not repaired. Recombinational repair is prevalent from the simplest viruses to the most
complex multicellular eukaryotes. It is effective against many different types of genomic damage, and in
particular is highly efficient at overcoming double-strand damages. Studies of the mechanism of meiotic
recombination indicate that meiosis is an adaptation for repairing DNA.[21] These considerations form the
basis for the first part of the repair and complementation hypothesis.
In some lines of descent from the earliest organisms, the diploid stage of the sexual cycle, which was
at first transient, became the predominant stage, because it allowed complementation — the masking of
deleterious recessive mutations (i.e. hybrid vigor or heterosis). Outcrossing, the second fundamental aspect
of sex, is maintained by the advantage of masking mutations and the disadvantage of inbreeding (mating
with a close relative) which allows expression of recessive mutations (commonly observed as inbreeding
depression). This is in accord with Charles Darwin,[22] who concluded that the adaptive advantage of sex is
hybrid vigor; or as he put it, “the offspring of two individuals, especially if their progenitors have been
subjected to very different conditions, have a great advantage in height, weight, constitutional vigor and
fertility over the self fertilised offspring from either one of the same parents.”
However, outcrossing may be abandoned in favor of parthogenesis or selfing (which retain the
advantage of meiotic recombinational repair) under conditions in which the costs of mating are very high. For
instance, costs of mating are high when individuals are rare in a geographic area, such as when there has
been a forest fire and the individuals entering the burned area are the initial ones to arrive. At such times
mates are hard to find, and this favors parthenogenic species.
In the view of the repair and complementation hypothesis, the removal of DNA damage by
recombinational repair produces a new, less deleterious form of informational noise, allelic recombination, as
a by-product. This lesser informational noise generates genetic variation, viewed by some as the major effect
of sex, as discussed in the earlier parts of this article.

[edit] Origin of sexual reproduction

This section requires expansion.
All sexually reproducing organisms derive from a common ancestor which was a single celled
eukaryotic species[citation needed]. Many protists reproduce sexually, as do the multicellular plants, animals,
and fungi. There are a few species which have secondarily lost this feature, such as Bdelloidea and some
parthenocarpic plants.
Organisms need to replicate their genetic material in an efficient and reliable manner. The necessity
to repair genetic damage is one of the leading theories explaining the origin of sexual reproduction. Diploid
individuals can repair a mutated section of its DNA via homologous recombination, since there are two
copies of the gene in the cell and one copy is presumed to be undamaged. A mutation in an haploid
individual, on the other hand, is more likely to become resident, as the DNA repair machinery has no way of
knowing what the original undamaged sequence was.[18] The most primitive form of sex may have been one
organism with damaged DNA replicating an undamaged strand from a similar organism in order to repair
Another theory is that sexual reproduction originated from selfish parasitic genetic elements that
exchange genetic material (that is: copies of their own genome) for their transmission and propagation. In
some organisms, sexual reproduction has been shown to enhance the spread of parasitic genetic elements
(e.g.: yeast, filamentous fungi).[24] Bacterial conjugation, a form of genetic exchange that some sources
describe as sex, is not a form of reproduction, but rather an example of horizontal gene transfer. However, it
does support the selfish genetic element theory, as it is propagated through such a "selfish gene", the F-
plasmid.[23] Similarly, it has been proposed that sexual reproduction evolved from ancient haloarchaea
through a combination of jumping genes, and swapping plasmids.[25]
A third theory is that sex evolved as a form of cannibalism. One primitive organism ate another one,
but rather than completely digesting it, some of the 'eaten' organism's DNA was incorporated into the 'eater'
Sex may also be derived from prokaryotic processes. A comprehensive 'origin of sex as vaccination'
theory proposes that eukaryan sex-as-syngamy (fusion sex) arose from prokaryan unilateral sex-as-infection
when infected hosts began swapping nuclearised genomes containing coevolved, vertically transmitted
symbionts that provided protection against horizontal superinfection by more virulent symbionts. Sex-as-
meiosis (fission sex) then evolved as a host strategy to uncouple (and thereby emasculate) the acquired
symbiont genomes.[26]

[edit] Mechanistic origin of sexual reproduction

Though theories positing benefits that lead to the origin of sex are often problematic, several credible
theories on the evolution of the mechanisms of sexual reproduction have been proposed.

[edit] Viral eukaryogenesis

The viral eukaryogenesis (VE) theory proposes that eukaryotic cells arose from a combination of a
lysogenic virus, an archaeon and a bacterium. This model suggests that the nucleus originated when the
lysogenic virus incorporated genetic material from the archaeon and the bacterium and took over the role of
information storage for the amalgam. The archaeal host transferred much of its functional genome to the
virus during the evolution of cytoplasm but retained the function of gene translation and general metabolism.
The bacterium transferred most of its functional genome to the virus as it transitioned into a mitochondrion.
For these transformations to lead to the eukaryotic cell cycle, the VE hypothesis specifies a pox-like
virus as the lysogenic virus. A pox-like virus is a likely ancestor because of its fundamental similarities with
eukaryotic nuclei. These include a double stranded DNA genome, a linear chromosome with short telomeric
repeats, a complex membrane bound capsid, the ability to produce capped mRNA, and the ability to export
the capped mRNA across the viral membrane into the cytoplasm. The presence of a lysogenic pox-like virus
ancestor explains the development of meiotic division, an essential component of sexual reproduction.[28]
Meiotic division in the VE hypothesis arose because of the evolutionary pressures placed on the
lysogenic virus as a result of its inability to enter into the lytic cycle. This selective pressure resulted in the
development of processes allowing the viruses to spread horizontally throughout the population. The
outcome of this selection was cell-to-cell fusion. (This is distinct from the conjugation methods used by
bacterial plasmids under evolutionary pressure, with important consequences.)[27] The possibility of this kind
of fusion is supported by the presence of fusion proteins in the envelopes of the pox viruses that allow them
to fuse with host membranes. These proteins could have been transferred to the cell membrane during viral
reproduction, enabling cell-to-cell fusion between the virus host and an uninfected cell. The theory proposes
meiosis originated from the fusion between two cells infected with related but different viruses which
recognised each other as uninfected. After the fusion of the two cells, incompatibilities between the two
viruses result in a meiotic-like cell division.[28]
The two viruses established in the cell would initiate replication in response to signals from the host
cell. A mitosis-like cell cycle would proceed until the viral membranes dissolved, at which point linear
chromosomes would be bound together with centromeres. The homologous nature of the two viral
centromeres would incite the grouping of both sets into tetrads. It is speculated that this grouping may be the
origin of crossing over, characteristic of the first division in modern meiosis. The partitioning apparatus of the
mitotic-like cell cycle the cells used to replicate independently would then pull each set of chromosomes to
one side of the cell, still bound by centromeres. These centromeres would prevent their replication in
subsequent division, resulting in four daughter cells with one copy of one of the two original pox-like viruses.
The process resulting from combination of two similar pox viruses within the same host closely mimics
[edit] Neomuran revolution
An alternative theory, proposed by Thomas Cavalier-Smith, was labeled the Neomuran revolution.
The designation "Neomuran revolution" refers to the appearances of the common ancestors of eukaryotes
and archaea. Cavalier-Smith proposes that the first neomurans emerged 850 million years ago. Other
molecular biologists assume that this group appeared much earlier, but Cavalier-Smith dismisses these
claims because they are based on the "theoretically and empirically" unsound model of molecular clocks.
Cavalier-Smith's theory of the Neomuran revolution has implications for the evolutionary history of the cellular
machinery for recombination and sex. It suggests that this machinery evolved in two distinct bouts separated
by a long period of stasis; first the appearance of recombination machinery in a bacterial ancestor which was
maintained for 3 Gy, until the neomuran revolution when the mechanics were adapted to the presence of
nucleosomes. The archaeal products of the revolution maintained recombination machinery that was
essentially bacterial, whereas the eukaryotic products broke with this bacterial continuity. They introduced
cell fusion and ploidy cycles into cell life histories. Cavalier-Smith argues that both bouts of mechanical
evolution were motivated by similar selective forces: the need for accurate DNA replication without loss of

[edit] See also


Books are collections of articles that can be downloaded or ordered in print.

• Asexual reproduction
• Biological reproduction
• Epistasis
• Genetic recombination
• Mutation
• Sexual reproduction

[edit] References
1. ^ a b John Maynard Smith The Evolution of Sex 1978.
2. ^ Doncaster, C. P. et al.; Pound, GE; Cox, SJ (2000). "The ecological cost of sex". Nature
404 (6775): 281–285. doi:10.1038/35005078. PMID 10749210.
3. ^ George C. Williams Sex and Evolution 1975, Princeton University Press, ISBN 0-691-
4. ^ a b Matt Ridley 1995 The Red Queen: Sex and the Evolution of Human Nature 1995
5. ^ Weismann, A. 1889. Essays on heredity and kindred biological subjects. Oxford Univ.
Press, Oxford, UK
6. ^ Fisher, R. A. 1930. The genetical theory of natural selection. Clarendon Press, Oxford, UK
7. ^ Muller, H. J. (1932). "Some genetic aspects of sex". Am. Nat. 8: 118–138.
8. ^ Burt, A. (2000). "Perspective: sex, recombination, and the efficacy of selection—was
Weismann right?". Evolution 54 (2): 337–351. PMID 10937212.
9. ^ Van Valen, L. (1973). "A New Evolutionary Law". Evolutionary Theory 1: 1–30.
10.^ Hamilton, W. D. et al. (1990). "Sexual reproduction as an adaptation to resist parasites".
Proceedings of the National Academy of Sciences 87: 3566–3573. doi:10.1073/pnas.87.9.3566.
11.^ Kuma, K.; Iwabe, N.; Miyata, T. (1995). "Functional constraints against variations on
molecules from the tissue-level - slowly evolving brain-specific genes demonstrated by protein-
kinase and immunoglobulin supergene families". Molecular Biology and Evolution 12 (1): 123–130.
PMID 7877487.
12.^ Wolfe KH & Sharp PM. 1993. Mammalian gene evolution - nucleotide-sequence divergence
between mouse and rat. Journal of molecular evolution 37 (4): 441-456 OCT 1993
13.^ Griffiths et al. 1999. Gene mutations, p197-234, in Modern Genetic Analysis, New York,
W.H. Freeman and Company.
14.^ a b Kondrashov, A. S. (1988). "Deleterious mutations and the evolution of sexual
reproduction". Nature 336: 435–440. doi:10.1038/336435a0.
15.^ Whitlock, M. C.; Bourget, D. (2000). "Factors affecting the genetic load in Drosophila:
synergistic epistasis and correlations among fitness components". Evolution 54 (5): 1654–1660.
PMID 11108592.
16.^ Elena, S. F.; Lenski, R. E. (1997). "Test of synergistic interactions among deleterious
mutations in bacteria". Nature 390 (6658): 395–398. doi:10.1038/37108. PMID 9389477.
17.^ Colegrave, N. (2002). "Sex releases the speed limit on evolution". Nature 420 (6916): 664–
666. doi:10.1038/nature01191. PMID 12478292.
18.^ a b Bernstein H, Byerly HC, Hopf FA, Michod RE (1984). "Origin of sex". J. Theor. Biol. 110
(3): 323–51. doi:10.1016/S0022-5193(84)80178-2. PMID 6209512.
19.^ Bernstein H, Byerly HC, Hopf FA, Michod RE (1985). "Genetic damage, mutation, and the
evolution of sex". Science 229 (4719): 1277–81. doi:10.1126/science.3898363. PMID 3898363.
20.^ Bernstein H, Hopf FA, Michod RE (1987). "The molecular basis of the evolution of sex".
Adv. Genet. 24: 323–70. doi:10.1016/S0065-2660(08)60012-7. PMID 3324702.
21.^ Cox MM (2001). "Historical overview: searching for replication help in all of the rec places".
Proc. Natl. Acad. Sci. U.S.A. 98 (15): 8173–80. doi:10.1073/pnas.131004998. PMID 11459950.
22.^ Darwin C. 1889. The effects of cross and self fertilisation in the vegetable kingdom.
Chapter XII. General Results pp. 436-463. D. Appleton and Company, New York
23.^ a b c Olivia Judson (2002). Dr. Tatiana's sex advice to all creation. New York: Metropolitan
Books. pp. 233–4. ISBN 0-8050-6331-5.
24.^ Hickey D (1982). "Selfish DNA: a sexually-transmitted nuclear parasite". Genetics 101 (3-
4): 519–31. PMID 6293914.
25.^ DasSarma, Shiladitya (2007). "Extreme Microbes". American Scientist. 95. pp. 224–231
26.^ Sterrer W (2002). "On the origin of sex as vaccination". Journal of Theoretical Biology 216
(4): 387–396. doi:10.1006/jtbi.2002.3008. PMID 12151256.
27.^ a b PJ Bell (2001). "Viral eukaryogenesis: was the ancestor of the nucleus a complex DNA
virus?". J Molec Biol 53 (3): 251–6.
28.^ a b c PJ Bell (2006). "Sex and the eukaryotic cell cycle is consistent with a viral ancestry for
the eukaryotic nucleus". J Theor Biol 243 (1): 54–63. doi:10.1016/j.jtbi.2006.05.015. PMID 16846615.
29.^ Cavalier-Smith, Thomas (2006). "Cell evolution and Earth history: stasis and revolution".
Royal Society of Biol Sci 361 (1470): 969–1006. doi:10.1098/rstb.2006.1842. PMID 16754610.

[edit] Further reading

• Bell, Graham (1982). The masterpiece of nature: the evolution and genetics of sexuality.
Berkeley: University of California Press. ISBN 0-520-04583-1.
• Bernstein, Harris; Carol Bernstein (1991). Aging, sex, and DNA repair. Boston: Academic
Press. ISBN 0-12-092860-4.
• Hurst, L.D.; J.R. Peck (1996). "Recent advances in the understanding of the evolution and
maintenance of sex". Trends in Ecology and Evolution 11: 46–52. doi:10.1016/0169-5347(96)81041-
• Levin, Bruce R.; Richard E. Michod (1988). The Evolution of sex: an examination of current
ideas. Sunderland, Mass: Sinauer Associates. ISBN 0-87893-459-6.
• Maynard Smith, John (1978). The evolution of sex. Cambridge, UK: Cambridge University
Press. ISBN 0-521-21887-X.
• Michod, Richard E. (1995). Eros and evolution: a natural philosophy of sex. Reading, Mass:
Addison-Wesley Pub. Co. ISBN 0-201-40754-X.
• "Scientists put sex origin mystery to bed, Wild strawberry research provides evidence on
when gender emerges". MSNBC. Retrieved 2008-11-25.
• Ridley, Mark (1993). Evolution. Oxford: Blackwell Scientific. ISBN 0-632-03481-5.
• Ridley, Mark (2000). Mendel's demon: gene justice and the complexity of life . London:
Weidenfeld & Nicolson. ISBN 0-297-64634-6.
• Ridley, Matt (1995). The Red Queen: sex and the evolution of human nature . New York:
Penguin Books. ISBN 0-14-024548-0.
• Szathmáry, Eörs; John Maynard Smith (1995). The Major Transitions in Evolution. Oxford:
W.H. Freeman Spektrum. ISBN 0-7167-4525-9.
• Taylor, Timothy (1996). The prehistory of sex: four million years of human sexual culture .
New York: Bantam Books. ISBN 0-553-09694-X.
• Williams, George (1975). Sex and evolution. Princeton, N.J: Princeton University Press.
ISBN 0-691-08147-6.

[edit] External links

• The Origin of Sex: Cosmic Solution to an Ancient Mystery
• Why Sex is Good
• An essay summarising the different theories , dating from around 2001

[hide]v · d · eThe development of phenotype

Key concepts Genotype-phenotype distinction · Norms of reaction · Gene-

environment interaction · Heritability · Quantitative genetics

Dominance relationship · Epistasis · Polygenic inheritance ·

Genetic architecture Pleiotropy · Plasticity · Canalisation · Fitness landscape · Transgressive

influences Epigenetics · Maternal effect · Dual inheritance theory

architecture Segmentation · Modularity

Evolution of genetic
systems Evolvability · Mutational robustness · Evolution of sex

Influential figures C. H. Waddington · Richard Lewontin

Debates Nature versus nurture · Morphogenetic field

List of evolutionary biology topics

[hide]v · d · e Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Multicellular · Sex

Birds · Butterflies · Cephalopods · Dinosaurs · Dolphins and whales ·

Taxa evolution Fungi · Horses · Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants ·
Sirenians (sea cows) · Spiders · Viruses

Modes of
speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from ""

Categories: Evolutionary biology | Sex


From Wikipedia, the free encyclopedia

Jump to: navigation, search

For other uses, see Flower (disambiguation).
A poster with twelve species of flowers or clusters of flowers of different families
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering
plants (plants of the division Magnoliophyta, also called angiosperms). The biological function of a flower is to
effect reproduction, usually by providing a mechanism for the union of sperm with eggs. Flowers may
facilitate outcrossing (fusion of sperm and eggs from different individuals in a population) or allow selfing
(fusion of sperm and egg from the same flower). Some flowers produce diaspores without fertilization
(parthenocarpy). Flowers contain sporangia and are the site where gametophytes develop. Flowers give rise
to fruit and seeds. Many flowers have evolved to be attractive to animals, so as to cause them to be vectors
for the transfer of pollen.
In addition to facilitating the reproduction of flowering plants, flowers have long been admired and
used by humans to beautify their environment but also as objects of romance, ritual, religion, medicine and
as a source of food.
• 1 Morphology
• 2 Development
• 2.1 Flowering transition
• 2.2 Organ development
• 3 Flower specialization and pollination
• 3.1 Floral formula
• 4 Pollination
• 4.1 Attraction methods
• 4.2 Pollination mechanism
• 4.3 Flower-pollinator relationships
• 5 Fertilization and dispersal
• 6 Evolution
• 7 Symbolism
• 8 Usage
• 9 See also
• 10 References
• 11 External links

Diagram showing the main parts of a mature flower

Flowering plants are heterosporous, producing two types of spores. Microspores are produced by
meiosis inside anthers while megaspores are produced inside ovules, inside an ovary. In fact, anthers
typically consist of four microsporangia and an ovule is an integumented megasporangium. Both types of
spores develop into gametophytes inside sporangia. As with all heterosporous plants, the gametophytes also
develop inside the spores (are endosporic).
A flower is a modified stem tip with compressed internodes, bearing structures that are highly
modified leaves.[1] In essence, a flower develops on a modified shoot or axis from a determinate apical
meristem (determinate meaning the axis grows to a set size).
Flowers may be directly attached to the plant at their base (sessile--the supporting stalk or stem is
highly reduced or absent). The stem or stalk subtending a flower is called a peduncle. If a peduncle supports
more than one flower, the stems connecting each flower to the main axis are called pedicels. The apex of a
flowering stem forms a terminal swelling which is called the torus or receptacle. The parts of a flower are
arranged in whorls on the receptacle. The four main whorls (starting from the base of the flower or lowest
node and working upwards) are as follows:
An example of a "perfect flower", this Crateva religiosa flower has both stamens (outer ring) and a
pistil (center).
• Calyx: the outermost whorl consisting of units called sepals; these are typically green and
enclose the rest of the flower in the bud stage, however, they can be absent or prominent and petal-
like in some species.
• Corolla: the next whorl toward the apex, composed of units called petals, which are typically
thin, soft and colored to attract animals that help the process of pollination.
• Androecium (from Greek andros oikia: man's house): the next whorl (sometimes multiplied
into several whorls), consisting of units called stamens. Stamens consist of two parts: a stalk called a
filament, topped by an anther where pollen is produced by meiosis and eventually dispersed.
• Gynoecium (from Greek gynaikos oikia: woman's house): the innermost whorl of a flower,
consisting of one or more units called carpels. The carpel or multiple fused carpels form a hollow
structure called an ovary, which produces ovules internally. Ovules are megasporangia and they in
turn produce megaspores by meiosis which develop into female gametophytes. These give rise to
egg cells. The gynoecium of a flower is also described using an alternative terminology wherein the
structure one sees in the innermost whorl (consisting of an ovary, style and stigma) is called a pistil.
A pistil may consist of a single carpel or a number of carpels fused together. The sticky tip of the
pistil, the stigma, is the receptor of pollen. The supportive stalk, the style, becomes the pathway for
pollen tubes to grow from pollen grains adhering to the stigma.
Although the arrangement described above is considered "typical", plant species show a wide
variation in floral structure. These modifications have significance in the evolution of flowering plants and are
used extensively by botanists to establish relationships among plant species.
In the majority of species, individual flowers have both functional carpels and stamens. These flowers
are described by botanists as being perfect or bisexual. Some flowers lack one or the other reproductive
organ and called imperfect or unisexual If unisex flowers are found on the same individual plant but in
different locations, the species is said to be monoecious. If each type of unisex flower is found only on
separate individuals, the plant is dioecious.
Additional discussions on floral modifications from the basic plan are presented in the articles on
each of the basic parts of the flower. In those species that have more than one flower on an axis, the
collection of flowers is termed an inflorescence. Some inflorescences are composed of many small flowers
arranged in a formation that resembles a single flower. The common example of this is most members of the
very large composite (Asteraceae) group. A single daisy or sunflower, for example, is not a flower but a
flower head—an inflorescence composed of numerous flowers (or florets).
Many flowers have a symmetry, if the perianth is bisected through the central axis from any point,
symmetrical halves are produced—the flower is called regular or actinomorphic, e.g. rose or trillium. When
flowers are bisected and produce only one line that produces symmetrical halves the flower is said to be
irregular or zygomorphic. e.g. snapdragon or most orchids.
Christmas Lillium (Lilium longiflorum). 1. Stigma, 2. Style, 3. Stamens, 4. Filament, 5. Petal
Flowering transition
The transition to flowering is one of the major phase changes that a plant makes during its life cycle.
The transition must take place at a time that is favorable for fertilization and the formation of seeds, hence
ensuring maximal reproductive success. To meet these needs a plant is able to interpret important
endogenous and environmental cues such as changes in levels of plant hormones and seasonable
temperature and photoperiod changes.[2] Many perennial and most biennial plants require vernalization to
flower. The molecular interpretation of these signals is through the transmission of a complex signal known
as florigen, which involves a variety of genes, including CONSTANS, FLOWERING LOCUS C and
FLOWERING LOCUS T. Florigen is produced in the leaves in reproductively favorable conditions and acts in
buds and growing tips to induce a number of different physiological and morphological changes.[3] The first
step is the transformation of the vegetative stem primordia into floral primordia. This occurs as biochemical
changes take place to change cellular differentiation of leaf, bud and stem tissues into tissue that will grow
into the reproductive organs. Growth of the central part of the stem tip stops or flattens out and the sides
develop protuberances in a whorled or spiral fashion around the outside of the stem end. These
protuberances develop into the sepals, petals, stamens, and carpels. Once this process begins, in most
plants, it cannot be reversed and the stems develop flowers, even if the initial start of the flower formation
event was dependent of some environmental cue.[4] Once the process begins, even if that cue is removed
the stem will continue to develop a flower.
Organ development

The ABC model of flower development

The molecular control of floral organ identity determination is fairly well understood. In a simple
model, three gene activities interact in a combinatorial manner to determine the developmental identities of
the organ primordia within the floral meristem. These gene functions are called A, B and C-gene functions. In
the first floral whorl only A-genes are expressed, leading to the formation of sepals. In the second whorl both
A- and B-genes are expressed, leading to the formation of petals. In the third whorl, B and C genes interact
to form stamens and in the center of the flower C-genes alone give rise to carpels. The model is based upon
studies of homeotic mutants in Arabidopsis thaliana and snapdragon, Antirrhinum majus. For example, when
there is a loss of B-gene function, mutant flowers are produced with sepals in the first whorl as usual, but also
in the second whorl instead of the normal petal formation. In the third whorl the lack of B function but
presence of C-function mimics the fourth whorl, leading to the formation of carpels also in the third whorl. See
also The ABC Model of Flower Development.
Most genes central in this model belong to the MADS-box genes and are transcription factors that
regulate the expression of the genes specific for each floral organ.

Flower specialization and pollination

Further information: Pollination syndrome
Flowering plants usually face selective pressure to optimize the transfer of their pollen, and this is
typically reflected in the morphology of the flowers and the behaviour of the plants. Pollen may be transferred
between plants via a number of 'vectors'. Some plants make use of abiotic vectors — namely wind
(anemophily) or, much less commonly, water (hydrophily). Others use biotic vectors including insects
(entomophily), birds (ornithophily), bats (chiropterophily) or other animals. Some plants make use of multiple
vectors, but many are highly specialised.
Cleistogamous flowers are self pollinated, after which they may or may not open. Many Viola and
some Salvia species are known to have these types of flowers.
The flowers of plants that make use of biotic pollen vectors commonly have glands called nectaries
that act as an incentive for animals to visit the flower. Some flowers have patterns, called nectar guides, that
show pollinators where to look for nectar. Flowers also attract pollinators by scent and color. Still other
flowers use mimicry to attract pollinators. Some species of orchids, for example, produce flowers resembling
female bees in color, shape, and scent. Flowers are also specialized in shape and have an arrangement of
the stamens that ensures that pollen grains are transferred to the bodies of the pollinator when it lands in
search of its attractant (such as nectar, pollen, or a mate). In pursuing this attractant from many flowers of the
same species, the pollinator transfers pollen to the stigmas—arranged with equally pointed precision—of all of
the flowers it visits.
Anemophilous flowers use the wind to move pollen from one flower to the next. Examples include
grasses, birch trees, ragweed and maples. They have no need to attract pollinators and therefore tend not to
be "showy" flowers. Male and female reproductive organs are generally found in separate flowers, the male
flowers having a number of long filaments terminating in exposed stamens, and the female flowers having
long, feather-like stigmas. Whereas the pollen of animal-pollinated flowers tends to be large-grained, sticky,
and rich in protein (another "reward" for pollinators), anemophilous flower pollen is usually small-grained,
very light, and of little nutritional value to animals.

Floral formula
A floral formula is a way to represent the structure of a flower using specific letters, numbers, and
symbols. Typically, a general formula will be used to represent the flower structure of a plant family rather
than a particular species. The following representations are used:
Ca = calyx (sepal whorl; e. g. Ca5 = 5 sepals)
Co = corolla (petal whorl; e. g., Co 3(x) = petals some multiple of three )
Z = add if zygomorphic (e. g., CoZ6 = zygomorphic with 6 petals)
A = androecium (whorl of stamens; e. g., A∞ = many stamens)
G = gynoecium (carpel or carpels; e. g., G1 = monocarpous)
x: to represent a "variable number"
∞: to represent "many"
A floral formula would appear something like this:
Ca5Co5A10 - ∞G1
Several additional symbols are sometimes used (see Key to Floral Formulas).
The four main parts of a flower are generally defined by their positions on the receptacle and not by
their function. Many flowers lack some parts or parts may be modified into other functions and/or look like
what is typically another part. In some families, like Ranunculaceae, the petals are greatly reduced and in
many species the sepals are colorful and petal-like. Other flowers have modified stamens that are petal-like,
the double flowers of Peonies and Roses are mostly petaloid stamens.[5] Flowers show great variation and
plant scientists describe this variation in a systematic way to identify and distinguish species.
Specific terminology is used to descried flowers and their parts. Many flower parts are fused
together; fused parts originating from the same whorl are connate, while fused parts originating from different
whorls are adnate, parts that are not fused are free. When petals are fused into a tube or ring that falls away
as a single unit, they are sympetalous (also called gamopetalous.) Petals that are connate may have
distinctive regions: the cylindrical base is the tube, the expanding region is the throat and the flaring outer
region is the limb. A sympetalous flower, with bilateral symmetry with an upper and lower lip, is bilabiate.
Flowers with connate petals or sepals may have various shaped corolla or calyx including: campanulate,
funnelform, tubular, urceolate, salverform or rotate.
Main article: pollination

Grains of pollen sticking to this bee will be transferred to the next flower it visits
Tip of a tulip stamen. Note the grains of pollen
The primary purpose of a flower is reproduction. Since the flowers are the reproductive organs of
plant, they mediate the joining of the sperm, contained within pollen, to the ovules — contained in the ovary.
Pollination is the movement of pollen from the anthers to the stigma. The joining of the sperm to the ovules is
called fertilization. Normally pollen is moved from one plant to another, but many plants are able to self
pollinate. The fertilized ovules produce seeds that are the next generation. Sexual reproduction produces
genetically unique offspring, allowing for adaptation. Flowers have specific designs which encourages the
transfer of pollen from one plant to another of the same species. Many plants are dependent upon external
factors for pollination, including: wind and animals, and especially insects. Even large animals such as birds,
bats, and pygmy possums can be employed. The period of time during which this process can take place (the
flower is fully expanded and functional) is called anthesis.

Attraction methods

A Bee orchid has evolved over many generations to better mimic a female bee to attract male bees
as pollinators.
Plants can not move from one location to another, thus many flowers have evolved to attract animals
to transfer pollen between individuals in dispersed populations. Flowers that are insect-pollinated are called
entomophilous; literally "insect-loving" in Greek. They can be highly modified along with the pollinating
insects by co-evolution. Flowers commonly have glands called nectaries on various parts that attract animals
looking for nutritious nectar. Birds and bees have color vision, enabling them to seek out "colorful" flowers.
Some flowers have patterns, called nectar guides, that show pollinators where to look for nectar; they may be
visible only under ultraviolet light, which is visible to bees and some other insects. Flowers also attract
pollinators by scent and some of those scents are pleasant to our sense of smell. Not all flower scents are
appealing to humans, a number of flowers are pollinated by insects that are attracted to rotten flesh and have
flowers that smell like dead animals, often called Carrion flowers including Rafflesia, the titan arum, and the
North American pawpaw (Asimina triloba). Flowers pollinated by night visitors, including bats and moths, are
likely to concentrate on scent to attract pollinators and most such flowers are white.
Still other flowers use mimicry to attract pollinators. Some species of orchids, for example, produce
flowers resembling female bees in color, shape, and scent. Male bees move from one such flower to another
in search of a mate.

Pollination mechanism
The pollination mechanism employed by a plant depends on what method of pollination is utilized.
Most flowers can be divided between two broad groups of pollination methods:
Entomophilous: flowers attract and use insects, bats, birds or other animals to transfer pollen from
one flower to the next. Often they are specialized in shape and have an arrangement of the stamens that
ensures that pollen grains are transferred to the bodies of the pollinator when it lands in search of its
attractant (such as nectar, pollen, or a mate). In pursuing this attractant from many flowers of the same
species, the pollinator transfers pollen to the stigmas—arranged with equally pointed precision—of all of the
flowers it visits. Many flowers rely on simple proximity between flower parts to ensure pollination. Others,
such as the Sarracenia or lady-slipper orchids, have elaborate designs to ensure pollination while preventing

Anthers detached from a Meadow Foxtail flower.

A grass flower head (Meadow Foxtail) showing the plain coloured flowers with large anthers.
Anemophilous: flowers use the wind to move pollen from one flower to the next, examples include
the grasses, Birch trees, Ragweed and Maples. They have no need to attract pollinators and therefore tend
not to be "showy" flowers. Whereas the pollen of entomophilous flowers tends to be large-grained, sticky,
and rich in protein (another "reward" for pollinators), anemophilous flower pollen is usually small-grained,
very light, and of little nutritional value to insects, though it may still be gathered in times of dearth.
Honeybees and bumblebees actively gather anemophilous corn (maize) pollen, though it is of little value to
Some flowers are self pollinated and use flowers that never open or are self pollinated before the
flowers open, these flowers are called cleistogamous. Many Viola species and some Salvia have these types
of flowers.
Flower-pollinator relationships
Many flowers have close relationships with one or a few specific pollinating organisms. Many flowers,
for example, attract only one specific species of insect, and therefore rely on that insect for successful
reproduction. This close relationship is often given as an example of coevolution, as the flower and pollinator
are thought to have developed together over a long period of time to match each other's needs.
This close relationship compounds the negative effects of extinction. The extinction of either member
in such a relationship would mean almost certain extinction of the other member as well. Some endangered
plant species are so because of shrinking pollinator populations.

Fertilization and dispersal

Main article: biological dispersal
Cassia Fistula. A hermaphrodite flower showing both male and female parts.
Some flowers with both stamens and a pistil are capable of self-fertilization, which does increase the
chance of producing seeds but limits genetic variation. The extreme case of self-fertilization occurs in flowers
that always self-fertilize, such as many dandelions. Conversely, many species of plants have ways of
preventing self-fertilization. Unisexual male and female flowers on the same plant may not appear or mature
at the same time, or pollen from the same plant may be incapable of fertilizing its ovules. The latter flower
types, which have chemical barriers to their own pollen, are referred to as self-sterile or self-incompatible
(see also: Plant sexuality).

Further information: Evolution of flowers

Archaefructus liaoningensis, one of the earliest known flowering plants

Amborella trichopoda, the sister group to the rest of the flowering plants
While land plants have existed for about 425 million years, the first ones reproduced by a simple
adaptation of their aquatic counterparts: spores. In the sea, plants—and some animals—can simply scatter out
genetic clones of themselves to float away and grow elsewhere. This is how early plants reproduced. But
plants soon evolved methods of protecting these copies to deal with drying out and other abuse which is
even more likely on land than in the sea. The protection became the seed, though it had not yet evolved the
flower. Early seed-bearing plants include the ginkgo and conifers. The earliest fossil of a flowering plant,
Archaefructus liaoningensis, is dated about 125 million years old.[6] Several groups of extinct gymnosperms,
particularly seed ferns, have been proposed as the ancestors of flowering plants but there is no continuous
fossil evidence showing exactly how flowers evolved. The apparently sudden appearance of relatively
modern flowers in the fossil record posed such a problem for the theory of evolution that it was called an
"abominable mystery" by Charles Darwin. Recently discovered angiosperm fossils such as Archaefructus,
along with further discoveries of fossil gymnosperms, suggest how angiosperm characteristics may have
been acquired in a series of steps.
Recent DNA analysis (molecular systematics)[7][8] shows that Amborella trichopoda, found on the
Pacific island of New Caledonia, is the sister group to the rest of the flowering plants, and morphological
studies[9] suggest that it has features which may have been characteristic of the earliest flowering plants.
The general assumption is that the function of flowers, from the start, was to involve animals in the
reproduction process. Pollen can be scattered without bright colors and obvious shapes, which would
therefore be a liability, using the plant's resources, unless they provide some other benefit. One proposed
reason for the sudden, fully developed appearance of flowers is that they evolved in an isolated setting like
an island, or chain of islands, where the plants bearing them were able to develop a highly specialized
relationship with some specific animal (a wasp, for example), the way many island species develop today.
This symbiotic relationship, with a hypothetical wasp bearing pollen from one plant to another much the way
fig wasps do today, could have eventually resulted in both the plant(s) and their partners developing a high
degree of specialization. Island genetics is believed to be a common source of speciation, especially when it
comes to radical adaptations which seem to have required inferior transitional forms. Note that the wasp
example is not incidental; bees, apparently evolved specifically for symbiotic plant relationships, are
descended from wasps.
Likewise, most fruit used in plant reproduction comes from the enlargement of parts of the flower.
This fruit is frequently a tool which depends upon animals wishing to eat it, and thus scattering the seeds it
While many such symbiotic relationships remain too fragile to survive competition with mainland
organisms, flowers proved to be an unusually effective means of production, spreading (whatever their actual
origin) to become the dominant form of land plant life.
While there is only hard proof of such flowers existing about 130 million years ago, there is some
circumstantial evidence that they did exist up to 250 million years ago. A chemical used by plants to defend
their flowers, oleanane, has been detected in fossil plants that old, including gigantopterids,[10] which
evolved at that time and bear many of the traits of modern, flowering plants, though they are not known to be
flowering plants themselves, because only their stems and prickles have been found preserved in detail; one
of the earliest examples of petrification.
The similarity in leaf and stem structure can be very important, because flowers are genetically just
an adaptation of normal leaf and stem components on plants, a combination of genes normally responsible
for forming new shoots.[11] The most primitive flowers are thought to have had a variable number of flower
parts, often separate from (but in contact with) each other. The flowers would have tended to grow in a spiral
pattern, to be bisexual (in plants, this means both male and female parts on the same flower), and to be
dominated by the ovary (female part). As flowers grew more advanced, some variations developed parts
fused together, with a much more specific number and design, and with either specific sexes per flower or
plant, or at least "ovary inferior".
Flower evolution continues to the present day; modern flowers have been so profoundly influenced
by humans that many of them cannot be pollinated in nature. Many modern, domesticated flowers used to be
simple weeds, which only sprouted when the ground was disturbed. Some of them tended to grow with
human crops, and the prettiest did not get plucked because of their beauty, developing a dependence upon
and special adaptation to human affection.[12]

Lilies are often used to denote life or resurrection

Flowers are common subjects of still life paintings, such as this one by Ambrosius Bosschaert the
Chinese Jade ornament with flower design, Jin Dynasty (1115-1234 AD), Shanghai Museum.
Many flowers have important symbolic meanings in Western culture. The practice of assigning
meanings to flowers is known as floriography. Some of the more common examples include:
• Red roses are given as a symbol of love, beauty, and passion.
• Poppies are a symbol of consolation in time of death. In the United Kingdom, New Zealand,
Australia and Canada, red poppies are worn to commemorate soldiers who have died in times of war.
• Irises/Lily are used in burials as a symbol referring to "resurrection/life". It is also associated
with stars (sun) and its petals blooming/shining.
• Daisies are a symbol of innocence.
Flowers within art are also representative of the female genitalia, as seen in the works of artists such
as Georgia O'Keeffe, Imogen Cunningham, Veronica Ruiz de Velasco, and Judy Chicago, and in fact in
Asian and western classical art. Many cultures around the world have a marked tendency to associate
flowers with femininity.
The great variety of delicate and beautiful flowers has inspired the works of numerous poets,
especially from the 18th-19th century Romantic era. Famous examples include William Wordsworth's I
Wandered Lonely as a Cloud and William Blake's Ah! Sun-Flower.
Because of their varied and colorful appearance, flowers have long been a favorite subject of visual
artists as well. Some of the most celebrated paintings from well-known painters are of flowers, such as Van
Gogh's sunflowers series or Monet's water lilies. Flowers are also dried, freeze dried and pressed in order to
create permanent, three-dimensional pieces of flower art.
The Roman goddess of flowers, gardens, and the season of Spring is Flora. The Greek goddess of
spring, flowers and nature is Chloris.
In Hindu mythology, flowers have a significant status. Vishnu, one of the three major gods in the
Hindu system, is often depicted standing straight on a lotus flower.[13] Apart from the association with
Vishnu, the Hindu tradition also considers the lotus to have spiritual significance.[14] For example, it figures
in the Hindu stories of creation.[15]

Flower market, Detroit's Eastern Market.

Female hand spreading flowers over a Lingam temple in Varanasi
In modern times, people have sought ways to cultivate, buy, wear, or otherwise be around flowers
and blooming plants, partly because of their agreeable appearance and smell. Around the world, people use
flowers for a wide range of events and functions that, cumulatively, encompass one's lifetime:
• For new births or Christenings
• As a corsage or boutonniere to be worn at social functions or for holidays
• As tokens of love or esteem
• For wedding flowers for the bridal party, and decorations for the hall
• As brightening decorations within the home
• As a gift of remembrance for bon voyage parties, welcome home parties, and "thinking of
you" gifts
• For funeral flowers and expressions of sympathy for the grieving
• For worshiping goddesses. in Hindu culture it is very common to bring flowers as a gift to
People therefore grow flowers around their homes, dedicate entire parts of their living space to flower
gardens, pick wildflowers, or buy flowers from florists who depend on an entire network of commercial
growers and shippers to support their trade.
Flowers provide less food than other major plants parts (seeds, fruits, roots, stems and leaves) but
they provide several important foods and spices. Flower vegetables include broccoli, cauliflower and
artichoke. The most expensive spice, saffron, consists of dried stigmas of a crocus. Other flower spices are
cloves and capers. Hops flowers are used to flavor beer. Marigold flowers are fed to chickens to give their
egg yolks a golden yellow color, which consumers find more desirable. Dandelion flowers are often made into
wine. Bee Pollen, pollen collected from bees, is considered a health food by some people. Honey consists of
bee-processed flower nectar and is often named for the type of flower, e.g. orange blossom honey, clover
honey and tupelo honey.
Hundreds of fresh flowers are edible but few are widely marketed as food. They are often used to
add color and flavor to salads. Squash flowers are dipped in breadcrumbs and fried. Edible flowers include
nasturtium, chrysanthemum, carnation, cattail, honeysuckle, chicory, cornflower, Canna, and sunflower.
Some edible flowers are sometimes candied such as daisy and rose (you may also come across a candied
Flowers can also be made into herbal teas. Dried flowers such as chrysanthemum, rose, jasmine,
camomile are infused into tea both for their fragrance and medical properties. Sometimes, they are also
mixed with tea leaves for the added fragrance.

See also
• Gardening
• Garden
• List of garden plants
• Plant evolutionary developmental biology
• Plant sexuality
• Sowing

1. ^ Eames, A. J. (1961) Morphology of the Angiosperms McGraw-Hill Book Co., New York.
2. ^ Ausín, I., et al. (2005). "Environmental regulation of flowering". Int J Dev Biol 49 (5-6): 689–
705. doi:10.1387/ijdb.052022ia. PMID 16096975.
3. ^ Turck, F., Fornara, F., Coupland, G. (2008). "Regulation and Identity of Florigen:
FLOWERING LOCUS T Moves Centre Stage". Annual Review of Plant Biology 59: 573–594.
doi:10.1146/annurev.arplant.59.032607.092755. PMID 18444908.
4. ^ Searle, I., et al. (2006). "The transcription factor FLC confers a flowering response to
vernalization by repressing meristem competence and systemic signaling in Arabidopsis". Genes &
Dev. 20 (7): 898–912. doi:10.1101/gad.373506. PMID 16600915.
5. ^ Reynolds, Joan; Tampion, John (1983). Double flowers: a scientific study. London:
[Published for the] Polytechnic of Central London Press [by] Pembridge Press. p. 41. ISBN 978-0-
6. ^ "Flowers Modern & Ancient".
Retrieved 2010-08-30.
7. ^ "First Flower". 2007-04-17. Retrieved 2010-08-30.
8. ^ "Amborella not a "basal angiosperm"? Not so fast". doi:10.3732/ajb.91.6.997. Retrieved 2010-08-30.
9. ^ "South Pacific plant may be missing link in evolution of flowering plants".
2006-05-17. Retrieved 2010-
10.^ "Oily Fossils Provide Clues To The Evolution Of Flowers". 2001-04-05. Retrieved 2010-08-30.
11.^ "Age-Old Question On Evolution Of Flowers Answered". 2001-06-15. Retrieved 2010-08-30.
12.^ "Human Affection Altered Evolution of Flowers". Retrieved 2010-08-30.
13.^ "Vishnu". 2009-08-24. Retrieved 2010-08-30.
14.^ "God's Favorite Flower". Hinduism Today. Retrieved 2010-08-30.
15.^ "The Lotus".
mrook.htm. Retrieved 2010-08-30.
• Esau, Katherine (1965) Plant Anatomy (2nd ed.) John Wiley & Sons, New York.

External links
Wikiquote has a collection of quotations related to: Flowers

Look up flower in Wiktionary, the free dictionary.

Wikimedia Commons has media related to: flowers

• Media related to Flowers at Wikimedia Commons

• Native Plant Information Network
• Garden Guide UK - Information on Flowers and Colours
• Israeli researchers bring scent back to our flowers [VIDEO]

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Plant defense against herbivory

From Wikipedia, the free encyclopedia

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Poison ivy produces urushiol to protect the plant from herbivores. In humans this chemical produces
an allergic skin rash, known as urushiol-induced contact dermatitis.
Foxgloves produce several deadly chemicals, namely cardiac and steroidal glycosides. Ingestion can
cause nausea, vomiting, hallucinations, convulsions, or death.
Plant defense against herbivory or host-plant resistance (HPR) describes a range of adaptations
evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants
use several strategies to defend against damage caused by herbivores. Many plants produce secondary
metabolites, known as allelochemicals, that influence the behavior, growth, or survival of herbivores. These
chemical defenses can act as repellents or toxins to herbivores, or reduce plant digestibility.
Other defensive strategies used by plants include escaping or avoiding herbivores in time or in place,
for example by growing in a location where plants are not easily found or accessed by herbivores, or by
changing seasonal growth patterns. Another approach diverts herbivores toward eating non-essential parts,
or enhances the ability of a plant to recover from the damage caused by herbivory. Some plants encourage
the presence of natural enemies of herbivores, which in turn protect the plant. Each type of defense can be
either constitutive (always present in the plant), or induced (produced in reaction to damage or stress caused
by herbivores).
Historically, insects have been the most significant herbivores, and the evolution of land plants is
closely associated with the evolution of insects. While most plant defenses are directed against insects, other
defenses have evolved that are aimed at vertebrate herbivores, such as birds and mammals. The study of
plant defenses against herbivory is important, not only from an evolutionary view point, but also in the direct
impact that these defenses have on agriculture, including human and livestock food sources; as beneficial
'biological control agents' in biological pest control programs; as well as the in the search for plants of
medical importance.
• 1 Evolution of defensive traits
• 1.1 Records of herbivores
• 1.2 Co-evolution
• 2 Types
• 2.1 Chemical defenses
• 2.1.1 Types of chemical defenses
• 2.2 Mechanical defenses
• 2.2.1 Thigmonasty
• 2.3 Mimicry and camouflage
• 2.4 Indirect defenses
• 2.5 Leaf shedding and colour
• 3 Costs and benefits
• 3.1 Optimal defense hypothesis
• 3.2 Carbon:nutrient balance hypothesis
• 3.3 Growth rate hypothesis
• 3.4 Growth-differentiation balance hypothesis
• 4 Importance to humans
• 4.1 Agriculture
• 4.2 Pharmaceutical
• 4.3 Biological pest control
[edit] Evolution of defensive traits

Timeline of plant evolution and the beginnings of different modes of insect herbivory
Main article: Evolutionary history of plants
The earliest land plants evolved from aquatic plants around 450 million years ago (Ma) in the
Ordovician period. These early land plants had no vascular system and required free water for their
reproduction. Vascular plants appeared later and their diversification began in the Devonian era (about 400
Ma). Their reduced dependence on water resulted from adaptations such as protective coatings to reduce
evaporation from their tissues. Reproduction and dispersal of vascular plants in these dry conditions was
achieved through the evolution of specialized seed structures. The diversification of flowering plants
(angiosperms) during the Cretaceous period is associated with the sudden burst of speciation in insects.[1]
This diversification of insects represented a major selective force in plant evolution, and led to selection of
plants that had defensive adaptations. Early insect herbivores were mandibulate and bit or chewed
vegetation; but the evolution of vascular plants lead to the co-evolution of other forms of herbivory, such as
sap-sucking, leaf mining, gall forming and nectar-feeding.[2]

[edit] Records of herbivores

Viburnum lesquereuxii leaf with insect damage; Dakota Sandstone (Cretaceous) of Ellsworth County,
Kansas. Scale bar is 10 mm.
Our understanding of herbivory in geological time comes from three sources: fossilised plants, which
may preserve evidence of defence (such as spines), or herbivory-related damage; the observation of plant
debris in fossilised animal faeces; and the construction of herbivore mouthparts.[3]
Long thought to be a Mesozoic phenomenon, evidence for herbivory is found almost as soon as
fossils which could show it. Within under 20 million years of the first fossils of sporangia and stems towards
the close of the Silurian, around 420 million years ago, there is evidence that they were being consumed.[4]
Animals fed on the spores of early Devonian plants, and the Rhynie chert also provides evidence that
organisms fed on plants using a "pierce and suck" technique.[3] Many plants of this time are preserved with
spine-like enations, which may have performed a defensive role before being co-opted to develop into
During the ensuing 75 million years, plants evolved a range of more complex organs - from roots to
seeds. There was a gap of 50 to 100 million years between each organ evolving, and it being fed upon.[4]
Hole feeding and skeletonisation are recorded in the early Permian, with surface fluid feeding evolving by the
end of that period.[3]

A Plain Tiger Danaus chrysippus caterpillar making a moat to block defensive chemicals of
Calotropis before feeding
[edit] Co-evolution
Herbivores depend on plants for food, and have evolved mechanisms to obtain this food despite the
evolution of a diverse arsenal of plant defenses. Herbivore adaptations to plant defense have been likened to
offensive traits and consist of adaptations that allow increased feeding and use of a host plant.[5]
Relationships between herbivores and their host plants often results in reciprocal evolutionary change, called
co-evolution. When a herbivore eats a plant it selects for plants that can mount a defensive response. In
cases where this relationship demonstrates specificity (the evolution of each trait is due to the other), and
reciprocity (both traits must evolve), the species are thought to have co-evolved.[6] The "escape and
radiation" mechanism for co-evolution presents the idea that adaptations in herbivores and their host plants
have been the driving force behind speciation,[1][7] and have played a role in the radiation of insect species
during the age of angiosperms.[8] Some herbivores have evolved ways to hijack plant defenses to their own
benefit, by sequestering these chemicals and using them to protect themselves from predators.[1]

[edit] Types
Plant defenses can be classified generally as constitutive or induced. Constitutive defenses are
always present in the plant, while induced defenses are produced or mobilized to the site where a plant is
injured. There is wide variation in the composition and concentration of constitutive defenses and these
range from mechanical defenses to digestibility reducers and toxins. Many external mechanical defenses and
large quantitative defenses are constitutive, as they require large amounts of resources to produce and
difficult to mobilize.[9]
Induced defenses include secondary metabolic products, as well as morphological and physiological
changes.[10] An advantage of inducible, as opposed to constitutive defenses, is that they are only produced
when needed, and are therefore potentially less costly, especially when herbivory is variable..[10]
[edit] Chemical defenses

Persimmon, genus Diospyros, has a high tannin content which gives immature fruit, seen above, an
astringent and bitter flavor.
The evolution of chemical defenses in plants is linked to the emergence of chemical substances that
are not involved in the essential photosynthetic and metabolic activities. These substances, secondary
metabolites, are organic compounds that are not directly involved in the normal growth, development or
reproduction of organisms,[11] and often produced as by-products during the synthesis of primary metabolic
products.[12] These secondary metabolites play a major role in defenses against herbivores.[1][11][13]
Secondary metabolites are often characterized as either qualitative or quantitative. Qualitative
metabolites are defined as toxins that interfere with an herbivore’s metabolism, often by blocking specific
biochemical reactions. Qualitative chemicals are present in plants in relatively low concentrations (often less
than 2% dry weight), and are not dosage dependent. They are usually small, water soluble molecules, and
therefore can be rapidly synthesized, transported and stored with relatively little energy cost to the plant.
Qualitative allelochemicals are usually effective against non-adapted specialists and generalist herbivores.
Quantitative chemicals are those that are present in high concentration in plants (5 – 40% dry weight)
and are equally effective against all specialists and generalist herbivores. Most quantitative metabolites are
digestibility reducers that make plant cell walls indigestible to animals. The effects of quantitative metabolites
are dosage dependent and the higher these chemicals’ proportion in the herbivore’s diet, the less nutrition
the herbivore can gain from ingesting plant tissues. Because they are typically large molecules, these
defenses are energetically expensive to produce and maintain, and often take longer to synthesise and
The geranium, for example, produces a unique chemical compound in its petals to defend itself from
Japanese beetles. Within 30 minutes of ingestion the chemical paralyzes the herbivore. While the chemical
usually wears off within a few hours, during this time the beetle is often consumed by its own predators.[2]
• See Toxalbumin
[edit] Types of chemical defenses
Plants have evolved many secondary metabolites involved in plant defense, which are collectively
known as antiherbivory compounds and can be classified into three sub-groups: nitrogen compounds
(including alkaloids, cyanogenic glycosides and glucosinolates), terpenoids, and phenolics.[15]
Alkaloids are derived from various amino acids. Over 3000 known alkaloids exist, examples include
nicotine, caffeine, morphine, colchicine, ergolines, strychnine, and quinine.[16] Alkaloids have
pharmacological effects on humans and other animals. Some alkaloids can inhibit or activate enzymes, or
alter carbohydrate and fat storage by inhibiting the formation phosphodiester bonds involved in their
breakdown.[17] Certain alkaloids bind to nucleic acids and can inhibit synthesis of proteins and affect DNA
repair mechanisms. Alkaloids can also affect cell membrane and cytoskeletal structure causing the cells to
weaken, collapse, or leak, and can affect nerve transmission.[18] Although alkaloids act on a diversity of
metabolic systems in humans and other animals, they almost uniformly invoke an aversively bitter taste.[19]
Cyanogenic glycosides are stored in inactive forms in plant vacuoles. They become toxic when
herbivores eat the plant and break cell membranes allowing the glycosides to come into contact with
enzymes in the cytoplasm releasing hydrogen cyanide which blocks cellular respiration.[20] Glucosinolates
are activated in much the same way as cyanogenic glucosides, and the products can cause gastroenteritis,
salivation, diarrhea, and irritation of the mouth.[19]
The terpenoids, sometimes referred to as isoprenoids, are organic chemicals similar to terpenes,
derived from five-carbon isoprene units. There are over 10,000 known types of terpenoids.[21] Most are
multicyclic structures which differ from one another in both functional groups, and in basic carbon skeletons.
[22] Monoterpenoids, continuing 2 isoprene units, are volatile essential oils such as citronella, limonene,
menthol, camphor, and pinene. Diterpenoids, 4 isoprene units, are widely distributed in latex and resins, and
can be quite toxic. Diterpenes are responsible for making Rhododendron leaves poisonous. Plant steroids
and sterols are also produced from terpenoid precursors, including vitamin D, glycosides (such as digitalis)
and saponins (which lyse red blood cells of herbivores).[23]
Phenolics, sometimes called phenols, consist of an aromatic 6-carbon ring bonded to a hydroxy
group. Some phenols have antiseptic properties, while others disrupt endocrine activity. Phenolics range
from simple tannins to the more complex flavonoids that give plants much of their red, blue, yellow, and white
pigments. Complex phenolics called polyphenols are capable of producing many different types of effects on
humans, including antioxidant properties. Some examples of phenolics used for defense in plants are: lignin,
silymarin and cannabinoids.[24] Condensed tannins, polymers composed of 2 to 50 (or more) flavonoid
molecules, inhibit herbivore digestion by binding to consumed plant proteins and making them more difficult
for animals to digest, and by interfering with protein absorption and digestive enzymes.[25] Silica and lignins,
which are completely indigestible to animals, grind down insect mandibles (appendages necessary for
In addition to the three larger groups of substances mentioned above, fatty acid derivates, amino
acids and even peptides[26] are used as defence. The cholinergic toxine, cicutoxin of water hemlock, is an
polyyne derived from the fatty acid metabolism.[27] β-N-Oxalyl-L-α,β-diaminopropionic acid as simple amino
acid is used by the sweet pea which leads also to intoxication in humans.[28] The synthesis of fluoroacetate
in several plants is an example for the use of small molecules to disturb the metabolism of the herbivore, in
this case the citric acid cycle.[29]
[edit] Mechanical defenses

The thorns on the stem of this raspberry plant, serve as a mechanical defense against herbivory.
Plants have many external structural defenses that discourage herbivory.[30] Depending on the
herbivore’s physical characteristics (i.e. size and defensive armor), plant structural defenses on stems and
leaves can deter, injure, or kill the grazer. Some defensive compounds are produced internally but are
released onto the plant’s surface; for example, resins, lignins, silica, and wax cover the epidermis of
terrestrial plants and alter the texture of the plant tissue. The leaves of holly plants, for instance, are very
smooth and slippery making feeding difficult. Some plants produce gummosis or sap that traps insects.
A plant's leaves and stem may be covered with sharp prickles, spines, thorns, or trichomes- hairs on
the leaf often with barbs, sometimes containing irritants or poisons. Plant structural features like spines and
thorns reduce feeding by large ungulate herbivores (e.g. kudu, impala, and goats) by restricting the
herbivores' feeding rate, or by wearing down the molars as in pears.[31] The structure of a plant, its
branching and leaf arrangement may also be evolved to reduce herbivore impact. The shrubs of New
Zealand have evolved special wide branching adaptations believed to be a response to browsing birds such
as the moas.[32] Similarly, African Acacias have long spines low in the canopy, but very short spines high in
the canopy, which is comparatively safe from herbivores such as giraffes.[33][34]
Coconut palms protect their fruit by surrounding it with multiple layers of armour.
Trees such as coconut and other palms, may protect their fruit by multiple layers of armour, needing
efficient tools to break through to the seed contents, and special skills to climb the tall and relatively smooth

[edit] Thigmonasty
Thigmonastic movements, those that occur in response to touch, are used as a defense in some
plants. The leaves of the sensitive plant, Mimosa pudica, close up rapidly in response to direct touch,
vibration, or even electrical and thermal stimuli. The proximate cause of this mechanical response is an
abrupt change in the turgor pressure in the pulvini at the base of leaves resulting from osmotic phenomena.
This is then spread via both electrical and chemical means through the plant; only a single leaflet need be
This response lowers the surface area available to herbivores, which are presented with the
underside of each leaflet, and results in a wilted appearance. It may also physically dislodge small
herbivores, such as insects.

[edit] Mimicry and camouflage

Some plants mimic the presence of insect eggs on their leaves, dissuading insect species from
laying their eggs there. Because female butterflies are less likely to lay their eggs on plants that already have
butterfly eggs, some species of neotropical vines of the genus Passiflora (Passion flowers) contain physical
structures resembling the yellow eggs of Heliconius butterflies on their leaves, which discourage oviposition
by butterflies.[36]
[edit] Indirect defenses
The large thorn-like stipules of Acacia collinsii are hollow and afford shelter for ants, which in return
protect the plant against herbivores.
Another category of plant defenses are those features that indirectly protect the plant by enhancing
the probability of attracting the natural enemies of herbivores. Such an arrangement is known as mutualism,
in this case of the "enemy of my enemy" variety. One such feature are semiochemicals, given off by plants.
Semiochemicals are a group of volatile organic compounds involved in interactions between organisms. One
group of semiochemicals are allelochemicals; consisting of allomones, which play a defensive role in
interspecies communication, and kairomones, which are used by members of higher trophic levels to locate
food sources. When a plant is attacked it releases allelochemics containing an abnormal ratio of volatiles.[37]
[38] Predators sense these volatiles as food cues, attracting them to the damaged plant, and to feeding
herbivores. The subsequent reduction in the number of herbivores confers a fitness benefit to the plant and
demonstrates the indirect defensive capabilities of semiochemicals. Induced volatiles also have drawbacks,
however; some studies have suggested that these volatiles also attract herbivores.[37]
Plants also provide housing and food items for natural enemies of herbivores, known as “biotic”
defense mechanisms, as a means to maintain their presence. For example, trees from the genus Macaranga
have adapted their thin stem walls to create ideal housing for an ant species (genus Crematogaster), which,
in turn, protects the plant from herbivores.[39] In addition to providing housing, the plant also provides the ant
with its exclusive food source; from the food bodies produced by the plant. Similarly, some Acacia tree
species have developed thorns that are swollen at the base, forming a hollowing structure that acts as
housing. These Acacia trees also produce nectar in extrafloral nectaries on their leaves as food for the ants.
Most plants have endophytes, microbial organisms that live within them. While some cause disease,
others protect plants from herbivores and pathogenic microbes. Endophytes can help the plant by producing
toxins harmful to other organisms that would attack the plant, such as alkaloid producing fungi which are
common in grasses such as tall fescue (Festuca arundinacea).[35]

[edit] Leaf shedding and colour

There have been suggestions that leaf shedding may be a response that provides protection against
diseases and certain kinds of pests such as leaf miners and gall forming insects.[41] Other responses such
as the change of leaf colours prior to fall have also been suggested as adaptations that may help undermine
the camouflage of herbivores.[42] Autumn leaf color has also been suggested to act as an honest warning
signal of defensive commitment towards insect pests that migrate to the trees in autumn.[43][44]

[edit] Costs and benefits

Defensive structures and chemicals are costly as they require resources that could otherwise be
used by plants to maximize growth and reproduction. Many models have been proposed to explore how and
why some plants make this investment in defenses against herbivores.

[edit] Optimal defense hypothesis

The optimal defense hypothesis attempts to explain how the kinds of defenses a particular plant
might use reflect the threats each individual plant faces.[45] This model considers three main factors,
namely: risk of attack, value of the plant part, and the cost of defense.[46][47]
The first factor determining optimal defense is risk: how likely is it that a plant or certain plant parts
will be attacked? This is also related to the plant apparency hypothesis, which states that a plant will invest
heavily in broadly effective defenses when the plant is easily found by herbivores.[48] Examples of apparent
plants that produce generalized protections include long-living trees, shrubs, and perennial grasses.[48]
Unapparent plants, such as short-lived plants of early successional stages, on the other hand, preferentially
invest in small amounts of qualitative toxins that are effective against all but the most specialized herbivores.
The second factor is the value of protection: would the plant be less able to survive and reproduce
after removal of part of its structure by a herbivore? Not all plant parts are of equal evolutionary value, thus
valuable parts contain more defenses. A plant’s stage of development at the time of feeding also affects the
resulting change in fitness. Experimentally, the fitness value of a plant structure is determined by removing
that part of the plant and observing the effect.[49] In general, reproductive parts are not as easily replaced as
vegetative parts, terminal leaves have greater value than basal leaves, and the loss of plant parts mid-
season has a greater negative effect on fitness than removal at the beginning or end of the season.[50][51]
Seeds in particular tend to be very well protected. For example, the seeds of many edible fruits and nuts
contain cyanogenic glycosides such as amygdalin. This results from the need to balance the effort needed to
make the fruit attractive to animal dispersers while ensuring that the seeds are not destroyed by the animal.
The final consideration is cost: how much will a particular defensive strategy cost a plant in energy
and materials? This is particularly important, as energy spent on defense cannot be used for other functions,
such as reproduction and growth. The optimal defense hypothesis predicts that plants will allocate more
energy towards defense when the benefits of protection outweigh the costs, specifically in situations where
there is high herbivore pressure.[54]
[edit] Carbon:nutrient balance hypothesis
The carbon:nutrient balance hypothesis, also known as the environmental constraint hypothesis,
states that the various types of plant defenses are responses to variations in the levels of nutrients in the
environment.[55][56] This hypothesis predicts the Carbon/Nitrogen ratio in plants determines which
secondary metabolites will be synthesized. For example, plants growing in nitrogen-poor soils will use
carbon-based defenses (mostly digestibility reducers), while those growing in low-carbon environments (such
as shady conditions) are more likely to produce nitrogen-based toxins. The hypothesis further predicts that
plants can change their defences in response to changes in nutrients. For example, if plants are grown in
low-nitrogen conditions, then these plants will implement a defensive strategy composed of constitutive
carbon-based defenses. If nutrient levels subsequently increase, by for example the addition of fertilizers,
these carbon-based defenses will decrease.

[edit] Growth rate hypothesis

The growth rate hypothesis, also known as the resource availability hypothesis, states that defense
strategies are determined by the inherent growth rate of the plant, which is in turn determined by the
resources available to the plant. A major assumption is that available resources are the limiting factor in
determining the maximum growth rate of a plant species. This model predicts that the level of defense
investment will increase as the potential of growth decreases.[57] Additionally, plants in resource-poor areas,
with inherently slow-growth rates, tend to have long-lived leaves and twigs, and the loss of plant appendages
may result in a loss of scarce and valuable nutrients.[58]
A recent test of this model involved a reciprocal transplants of seedlings of 20 species of trees
between clay soils (nutrient rich) and white sand (nutrient poor) to determine whether trade-offs between
growth rate and defenses restrict species to one habitat. Seedlings originating from the nutrient-poor sand
had higher levels of constitutive carbon-based defenses, but when they were transplanted into nutrient-rich
clay soils, they experienced higher mortality from herbivory. These finding suggest that defensive strategies
limit the habitats of some plants.[59]

[edit] Growth-differentiation balance hypothesis

The growth-differentiation balance hypothesis states that plant defenses are a result of a tradeoff
between “growth-related processes” and “differentiation-related processes” in different environments.[60]
Differentiation-related processes are defined as “processes that enhance the structure or function of existing
cells (i.e. maturation and specialization).”[45] A plant will produce chemical defenses only when energy is
available from photosynthesis, and plants with the highest concentrations of secondary metabolites are the
ones with an intermediate level of available resources.[60] The GDBH also accounts for tradeoffs between
growth and defense over a resource availability gradient. In situations where resources (e.g. water and
nutrients) limit photosynthesis, carbon supply is predicted to limit both growth and defense. As resource
availability increases, the requirements needed to support photosynthesis are met, allowing for accumulation
of carbohydrate in tissues. As resources are not sufficient to meet the large demands of growth, these carbon
compounds can instead be partitioned into the synthesis of carbon based secondary metabolites (phenolics,
tannins, etc.). In environments where the resource demands for growth are met, carbon is allocated to rapidly
dividing meristems (high sink strength) at the expense of secondary metabolism. Thus rapidly growing plants
are predicted to contain lower levels of secondary metabolites and vice versa. In addition, the tradeoff
predicted by the GDBH may change over time, as evidenced by a recent study on Salix spp. Much support
for this hypothesis is present in the literature, and some scientists consider the GDBH the most mature of the
plant defense hypotheses.
[edit] Importance to humans
[edit] Agriculture
The variation of plant susceptibility to pests was probably known even in the early stages of
agriculture in humans. In historic times, the observation of such variations in susceptibility have provided
solutions for major socio-economic problems. The grape phylloxera was introduced from North America to
France in 1860 and in 25 years it destroyed nearly a third (100,000 km²) of the French grape yards. Charles
Valentine Riley noted that the American species Vitis labrusca was resistant to Phylloxera. Riley, with J. E.
Planchon, helped save the French wine industry by suggesting the grafting of the susceptible but high quality
grapes onto Vitis labrusca root stocks.[61] The formal study of plant resistance to herbivory was first covered
extensively in 1951 by Reginald (R.H.) Painter, who is widely regarded as the founder of this area of
research, in his book Plant Resistance to Insects.[62] While this work pioneered further research in the US,
the work of Chesnokov was the basis of further research in the USSR.[63]
Fresh growth of grass is sometimes high in prussic acid content and can cause poisoning of grazing
livestock. The production of cyanogenic chemicals in grasses is primarily a defense against herbivores.[64]
The human innovation of cooking may have been particularly helpful in overcoming many of the
defensive chemicals of plants. Many enzyme inhibitors in cereal grains and pulses, such as trypsin inhibitors
prevalent in pulse crops, are denatured by cooking, making them digestible.[66][67]
It has been known since the late 17th century that plants contain noxious chemicals which are
avoided by insects. These chemicals have been used by man as early insecticides; in 1690 nicotine was
extracted from tobacco and used as a contact insecticide. In 1773, insect infested plants were treated with
nicotine fumigation by heating tobacco and blowing the smoke over the plants.[68] The flowers of
Chrysanthemum species contain pyrethrin which is a potent insecticide. In later years, the applications of
plant resistance became an important area of research in agriculture and plant breeding, particularly because
they can serve as a safe and low-cost alternative to the use of pesticides.[69] The important role of
secondary plant substances in plant defense was described in the late 1950s by Vincent Dethier and G.S.
Fraenkel.[11][70] The use of botanical pesticides is widespread and notable examples include Azadirachtin
from the neem (Azadirachta indica), d-Limonene from Citrus species, Rotenone from Derris, Capsaicin from
Chili Pepper and Pyrethrum.[71]
Natural materials found in the environment also induce plant resistance as well.[72] Chitosan derived
from chitin induce a plant's natural defense response against pathogens, diseases and insects including cyst
nematodes, both are approved as an biopesticide by the EPA to reduce the dependence on toxic pesticides.
The selective breeding of crop plants often involves selection against the plant's intrinsic resistance
strategies. This makes crop plant varieties particularly susceptible to pests unlike their wild relatives. In
breeding for host-plant resistance, it is often the wild relatives that provide the source of resistance genes.
These genes are incorporated using conventional approaches to plant breeding, but have also been
augmented by recombinant techniques, which allow introduction of genes from completely unrelated
organisms. The most famous transgenic approach is the introduction of genes from the bacterial species,
Bacillus thuringiensis, into plants. The bacterium produces proteins that, when ingested, kill lepidopteran
caterpillars. The gene encoding for these highly toxic proteins, when introduced into the host plant genome,
confers resistance against caterpillars, when the same toxic proteins are produced within the plant. This
approach is controversial, however, due to the possibility of ecological and toxicological side effects.[73]
[edit] Pharmaceutical

Illustration from the 15th century manuscript Tacuinum Sanitatis detailing the beneficial and harmful
properties of Mandrakes.
Many currently available pharmaceuticals are derived from the secondary metabolites plants use to
protect themselves from herbivores, including opium, aspirin, cocaine, and atropine.[74] These chemicals
have evolved to affect the biochemistry of insects in very specific ways. However, many of these biochemical
pathways are conserved in vertebrates, including humans, and the chemicals act on human biochemistry in
ways similar to that of insects. It has therefore been suggested that the study of plant-insect interactions may
help in bioprospecting.[75]
There is evidence that humans began using plant alkaloids in medical preparations as early as 3000
B.C.[17] Although the active components of most medicinal plants have been isolated only recently
(beginning in the early 19th century) these substances have been used as drugs throughout the human
history in potions, medicines, teas and as poisons. For example, to combat herbivory by the larvae of some
Lepidoptera species, Cinchona trees produce a variety of alkaloids, the most familiar of which is quinine.
Quinine is extremely bitter, making the bark of the tree quite unpalatable, it is also an anti-fever agent, known
as Jesuit's bark, and is especially useful in treating malaria.[76]
Throughout history mandrakes (Mandragora officinarum) have been highly sought after for their
reputed aphrodisiac properties. However, the roots of the mandrake plant also contain large quantities of the
alkaloid scopolamine, which, at high doses, acts as a central nervous system depressant, and makes the
plant highly toxic to herbivores. Scopolamine was later found to be medicinal use in pain management before
and during labor; in smaller doses it is used to prevent motion sickness.[77] One of the most well-known
medicinally valuable terpenes is an anticancer drug, taxol, isolated from the bark of the Pacific yew, Taxus
brevifolia, in the early 1960s.[78]

[edit] Biological pest control

Repellent companion planting, defensive live fencing hedges, and "obstructive-repellent"
interplanting, with host-plant resistance species as beneficial 'biological control agents' is a technique in
biological pest control programs for: organic gardening, wildlife gardening, sustainable gardening, and
sustainable landscaping; in organic farming and sustainable agriculture; and in restoration ecology methods
for habitat restoration projects.
[edit] See also
• Alarm signal#Other • List of repellent • Raphide
alarm signals plants • Plant
• Aposematism • List of pest- disease resistance
• Chemical ecology regulating plants • Pollination
• Canavanine • List of beneficial • Phytoalexin
• Druse (botany) weeds • Rapid plant
• Laticifer • List of movement
• Lectin companion plants • Seed

[edit] References
1. ^ a b c d Ehrlich, Paul R.; Peter H. Raven (December 1964). "Butterflies and plants: a study of
coevolution.". Evolution (Society for the Study of Evolution) 18 (4): 586–608. doi:10.2307/2406212.
2. ^ Labandeira, C.C.; D.L. Dilcher, D.R. Davis, D.L. Wagner (1994). "Ninety-seven million
years of angiosperm-insect association: paleobiological insights into the meaning of coevolution"
(PDF). Proceedings of the National Academy of Science of the U.S.A. 91 (25): 12278–82.
doi:10.1073/pnas.91.25.12278. PMID 11607501. PMC 45420.
3. ^ a b c Labandeira, C.C. (1998). "Early History Of Arthropod And Vascular Plant Associations
1". Annual Reviews in Earth and Planetary Sciences 26 (1): 329–377.
4. ^ a b Labandeira, C. (2007). "The origin of herbivory on land: Initial patterns of plant tissue
consumption by arthropods". Insect Science 14 (4): 259–275. doi:10.1111/j.1744-
5. ^ Karban, Richard; Anurag A. Agrawal (November 2002). "Herbivore offense". Annual
Review of Ecology and Systematics 33: 641–664. doi:10.1146/annurev.ecolsys.33.010802.150443.
6. ^ Futuyma, Douglas J.; Montgomery Slatkin (1983). Coevolution. Sunderland,
Massachusetts: Sinauer Associates. ISBN 0-87893-228-3.
7. ^ Thompson, J. (1999). "What we know and do not know about coevolution: insect herbivores
and plants as a test case.". In H. Olff, V. K. Brown, R. H. Drent. Herbivores: between plants and
predators; the 38th symposium of the British Ecological Society in cooperation with the Netherlands
Ecological Society held at the Wageningen Agricultural University, The Netherlands, 1997 . Oxford:
Blackwell Science. pp. 7–30. ISBN 0-632-05155-8.
8. ^ Farrell, Brian D.; Charles Mitter (1994). "Adaptive Radiation in Insects and Plants: Time
and Opportunity". American Zoologist 34 (1): 57–69. doi:10.1093/icb/34.1.57.
9. ^ Traw, Brian M.; Todd E. Dawson (May 2002). "Differential induction of trichomes by three
herbivores of black mustard" (PDF). Oecologia 131 (4): 526–532. doi:10.1007/s00442-002-0924-6. Retrieved 2007-05-27.
10.^ a b Karban, Richard; Anurag A. Agrawal, Marc Mangel (July 1997). "The benefits of induced
defenses against herbivores" (PDF). Ecology (Ecological Society of America) 78 (5): 1351–1355.
doi:10.2307/2266130. Retrieved
11.^ a b c Fraenkel, G. (1959). "The raison d'être of secondary plant substances". Science 129
(3361): 1466–70. doi:10.1126/science.129.3361.1466. PMID 13658975.
12.^ Whittaker, Robert H. (1970). "The biochemical ecology of higher plants". In Ernest
Sondheimer and John B. Simeone. Chemical ecology. Boston: Academic Press. pp. 43–70. ISBN 0-
13.^ Whittaker, Robert H. (1975). Communities and ecosystems. New York: Macmillan. ISBN 0-
14.^ Theis, Nina; Manuel Lerdau (2003). "The evolution of function in plant secondary
metabolites" (PDF). International Journal of Plant Science 164 (3 Suppl.): S93–S102.
doi:10.1086/374190. Archived from the original on 2007-04-18.
.pdf. Retrieved 2007-05-27.
15.^ "Biochemical defenses: secondary metabolites:". Plant Defense Systems & Medicinal
Botany. Retrieved 2007-05-21.
16.^ "Alkaloids: contain a N-containing heterocycle". Plant Defense Systems & Medicinal
Botany. Retrieved 2007-06-26.
17.^ a b Roberts, Margaret F.; Michael Wink (1998). Alkaloids: biochemistry, ecology, and
medicinal applications. New York: Plenum Press. ISBN 0-306-45465-3.
18.^ Sneden, Albert T.. "Alkaloids". Natural Products as Medicinally Useful Agents. Retrieved 2007-05-21.
19.^ a b Rhoades, David F (1979). "Evolution of Plant Chemical Defense against Herbivores". In
Rosenthal, Gerald A., & Janzen, Daniel H. (editors). Herbivores: Their Interaction with Secondary
Plant Metabolites. New York: Academic Press. pp. 3–54. ISBN 0-12-597180-X
20.^ Toxicon Volume 38, Issue 1, January 2000, Pages 11-36 János Vetter Plant cyanogenic
glycosides doi:10.1016/S0041-0101(99)00128-2
21.^ "Terpenoids". Plant Defense Systems & Medicinal Botany. Retrieved 2007-06-26.
22.^ Gershezon, Jonathan; Wolfgang Kreis (1999). "Biochemistry of terpinoids". In Michael
Wink. Biochemistry of plant secondary metabolism. London: Sheffield Academic Press. pp. 222–279.
ISBN 0-8493-4085-3.
23.^ Sneden, Albert T.. "Terpenes". Natural Products as Medicinally Useful Agents. Retrieved 2007-05-21.
24.^ "Phenols". Plant Defense Systems & Medicinal Botany. Retrieved 2007-05-21.
25.^ Van Soest, Peter J. (1982). Nutritional ecology of the ruminant: ruminant metabolism,
nutritional strategies, the cellulolytic fermentation, and the chemistry of forages and plant fibers .
Corvallis, Oregon: O & B Books. ISBN 0-9601586-0-X.
26.^ John W. Hylin (1969). "Toxic peptides and amino acids in foods and feeds". Journal of
Agricultural and Food Chemistry 17 (3): 492–496. doi:10.1021/jf60163a003.
27.^ E. Anet, B. Lythgoe, M. H. Silk, S. Trippett (1953). "Oenanthotoxin and cicutoxin. Isolation
and structures". Journal of the Chemical Society: 309–322. doi:10.1039/JR9530000309.
28.^ Mark V. Barrow; Charles F. Simpson; Edward J. Miller (1974). "Lathyrism: A Review". The
Quarterly Review of Biology 49 (2): 101–128. doi:10.1086/408017. PMID 4601279.
29.^ Donald A. Levin; King, Dennis R. (1991). "The Impact of Fluoroacetate-Bearing Vegetation
on Native Australian Fauna: A Review". Oikos (Blackwell Publishing) 61 (3): 412–430.
30.^ Fernandes GW (1994). "Plant mechanical defenses against insect herbivory.". Revista
Brasileira de Entomologia 38 (2): 421–433 [1].
31.^ Cooper, Susan M.; Norman Owen-Smith (September 1986). "Effects of plant spinescence
on large mammalian herbivores". Oecologia 68 (3): 446–455. doi:10.1007/BF01036753.
32.^ Bond W, Lee W & Craine J (2004). "Plant structural defences against browsing birds: a
legacy of New Zealand's extinct moas.". Oikos 104 (3): 500–508. doi:10.1111/j.0030-
33.^ {{Cite journal|last=Young |first=Truman P. |year=1987 |title=Increased thorn length in
Acacia drepanolobium- an induced response to browsing. |journal=Oecologia |volume=71 |
34.^ {{Cite journal|last=Young |first=Truman P. |coauthors=Bell Okello |year=1998 |
title=Relaxation of an induced defense after exclusion of herbivores: spines on Acacia
drepanolobium. |journal=Oecologia |volume=115 |pages=508–513.
35.^ a b Raven, Peter H.; Ray F. Evert, & Susan E. Eichhorn (2005). Biology of Plants. New
York: W. H. Freeman and Company. ISBN 0-7167-1007-2.
36.^ Williams, Kathy S.; Lawrence E. Gilbert (April 1981). "Insects as selective agents on plant
vegetative morphology: egg mimicry reduces egg-laying by butterflies". Science 212 (4493): 467–469.
doi:10.1126/science.212.4493.467. PMID 17802547.
37.^ a b Dicke, Marcel; Joop J.A. van Loon (December 2000). "Multitrophic effects of herbivore-
induced plant volatiles in an evolutionary context". Entomologia Experimentalis et Applicata 97 (3):
237–249. doi:10.1046/j.1570-7458.2000.00736.x.
38.^ Allmann, S.; Baldwin, I. T. (2010). "Insects Betray Themselves in Nature to Predators by
Rapid Isomerization of Green Leaf Volatiles". Science 329 (5995): 1075.
doi:10.1126/science.1191634. PMID 20798319. edit
39.^ Heil, Martin; Brigitte Fiala, K. Eduard Linsenmair, Gerhard Zotz, Petra Menke (December
1997). "Food body production in Macaranga triloba (Euphorbiaceae): A plant investment in anti-
herbivore defense via symbiotic ant partners". Journal of Ecology (British Ecological Society) 85 (6):
847–861. doi:10.2307/2960606.
40.^ Young, Truman P.; Cynthia H. Stubblefield, Lynne A. Isbell (January 1997). "Ants on
swollen-thorn acacias: species coexistence in a simple system". Oecologia 109 (1): 98–107.
41.^ Williams, Alan G.; Thomas G. Whitham (December 1986). "Premature Leaf Abscission: An
Induced Plant Defense Against Gall Aphids". Ecology (Ecological Society of America) 67 (6): 1619–
1627. doi:10.2307/1939093.
42.^ Lev-Yadun, Simcha; Amots Dafni, Moshe A. Flaishman, Moshe Inbar, Ido Izhaki, Gadi
Katzir, Gidi Ne'eman (October 2004). "Plant coloration undermines herbivorous insect camouflage"
(PDF). BioEssays 26 (10): 1126–1130. doi:10.1002/bies.20112. PMID 15382135.
Retrieved 2007-05-27.
43.^ Archetti, M., M (2000). "The origin of autumn colours by coevolution.". J. Theor. Biol. 205
(4): 625–630. doi:10.1006/jtbi.2000.2089. PMID 10931756.
44.^ Hamilton, W. D.; Brown, S. P. (2001). "Autumn tree colours as a handicap signal." (PDF).
Proc. R. Soc. B 268 (1475): 1489–1493. doi:10.1098/rspb.2001.1672. PMID 11454293.
PMC 1088768.
45.^ a b Stamp, Nancy (March 2003). "Out of the quagmire of plant defense hypotheses".
Quarterly Review of Biology 78 (1): 23–55. doi:10.1086/367580. PMID 12661508.
46.^ Rhoades, D. F.; R. G. Cates. (1974). "Towards a general theory of plant antiherbivore
chemistry". In V. C. Runeckles and E. E. Conn. Recent advances in phytochemistry: proceedings of
the annual meeting of the Phytochemical society of North America. Boston: Academic Press.
pp. 168–213. ISBN 0-12-612408-6.
47.^ Wilf, Peter; Conrad C. Labandeira, Kirk R. Johnson, Phyllis D. Coley, and Asher D. Cutter
(2001). "Insect herbivory, plant defense, and early Cenozoic climate change" (PDF). Proceedings of
the National Academy of Science 98 (11): 6221–6226. doi:10.1073/pnas.111069498.
PMID 11353840. PMC 33449. Retrieved 2007-05-
48.^ a b c Feeny, P. (1976). "Plant apparency and chemical defense.". In James W. Wallace and
Richard L. Mansell. Biochemical interaction between plants and insects: proceedings of the fifteenth
annual meeting of the Phytochemical Society of North America . New York: Plenum Press. pp. 1–40.
ISBN 0-306-34710-5.
49.^ D., McKey (1979). "The distribution of secondary compounds within plants.". In Gerald A.
Rosenthal, Daniel H. Janzen. Herbivores, their interaction with secondary plant metabolites . Boston:
Academic Press. pp. 55–133. ISBN 0-12-597180-X.
50.^ Krischik, V. A.; R. F. Denno. (1983). "Individual, population, and geographic patterns in
plant defense.". In Robert F. Denno, Mark S. McClure. Variable plants and herbivores in natural and
managed systems. Boston: Academic Press. pp. 463–512. ISBN 0-12-209160-4.
51.^ Zangerl, Arthur R.; Claire E. Rutledge (April 1996). "The probability of attack and patterns
of constitutive and induced defense: A test of optimal defense theory". The American Naturalist 147
(4): 599–608. doi:10.1086/285868.
0147(199604)147%3A4%3C599%3ATPOAAP%3E2.0.CO%3B2-D. Retrieved 2007-05-27.
52.^ Swain, Elisabeth; Chun Ping Li, Jonathan E. Poulton (1992). "Development of the Potential
for Cyanogenesis in Maturing Black Cherry (Prunus serotina Ehrh.) Fruits". Plant Physiology 98 (4):
1423–1428. doi:10.1104/pp.98.4.1423. PMID 16668810. PMC 1080367.
53.^ Witmer, M.C. (1998). "Ecological and evolutionary implications of energy and protein
requirements of avian frugivores eating sugary diets". Physiological Zoology 71 (6): 599–610.
PMID 9798248.
54.^ Pennings, Steven C.; Erin L. Siska, Mark D. Bertness (May 2001). "Latitudinal differences
in plant palatability in Atlantic coast salt marshes". Ecology (Ecological Society of America) 82 (5):
1344–1359. doi:10.2307/2679994.
55.^ Bryant, John P.; Stuart Chapin, III, David R. Klein (May 1983). "Carbon/nutrient balance of
boreal plants in relation to vertebrate herbivory". Oikos (Blackwell Publishing) 40 (3): 357–368.
56.^ Tuomi, J.; P. Niemela, F. S. Chapin, III, J. P. Bryant, and S. Siren. (1988). "Defensive
responses of trees in relation to their carbon/nutrient balance.". In William J. Mattson, Jean Levieux,
C. Bernard-Dagan. Mechanisms of woody plant defenses against insects: search for pattern . Berlin:
Springer-Verlag. pp. 57–72. ISBN 0-387-96673-0.
57.^ Colley, Phyllis D.; John P. Bryant, and F. Stuart Chapin III (1985). "Resource availability
and plant antiherbivore defense". Science 230 (4728): 895–899. doi:10.1126/science.230.4728.895.
PMID 17739203.
58.^ Chapin, F. Stuart, III (1980). "The Mineral Nutrition of Wild Plants". Annual Review of
Ecological Systematics 11: 233–260. doi:10.1146/
Retrieved 2007-05-27.
59.^ Fine, Paul V. A.; Italo Mesones, Phyllis D. Coley (July 2004). "Herbivores promote habitat
specialization by trees in Amazonian forests". Science 305 (5684): 663–5.
doi:10.1126/science.1098982. PMID 15286371.
60.^ a b Loomis, W. E. (1981). "Growth and differentiation—an introduction and summary.". In P.
F. Wareing and I. D. J. Phillips. Growth and differentiation in plants. New York: Pergamon Press.
pp. 1–17. ISBN 0-08-026351-8.
Herms, Daniel A.; William J. Mattson (September 1992). "The dilemma of plants: to grow or defend".
Quarterly Review of Biology 67 (3): 283–335. doi:10.1086/417659.
5770(199209)67%3A3%3C283%3ATDOPTG%3E2.0.CO%3B2-A. Retrieved 2007-05-27.
61.^ Polavarapu, Sridhar (2001). "Plant Resistance to insects". Agricultural Entomology & Pest
Management. Rutgers University. Retrieved 2007-
62.^ Painter, Reginald Henry (1951). Insect Resistance in Crop Plants. Lawrence: University of
Kansas Press. OCLC 443998.
63.^ Chesnokov, Pavel G. (1953). Methods of Investigating Plant Resistance to Pests .
Jerusalem: Israel Program for Scientific Translations. OCLC 3576157.
64.^ Gleadow, Roslyn M.; Ian E. Woodrow (2002). "Constraints on effectiveness of cyanogenic
glycosides in herbivore defense". Journal of Chemical Ecology 28 (7): 1301–13.
doi:10.1023/A:1016298100201. PMID 12199497.
65.^ Vough, Lester R.; E. Kim Cassel (July 2002). "Prussic Acid Poisoning of Livestock: Causes
and Prevention (ExEx 4016)" (PDF). Extension Extra. South Dakota State University Extension
66.^ Grant, G; Linda J. More, Norma H. McKenzie, Arpad Pusztai (1982). "The effect of heating
on the haemagglutinating activity and nutritional properties of bean (Phaseolus vulgaris) seeds".
Journal of the Science of Food and Agriculture 33 (12): 1324–6. doi:10.1002/jsfa.2740331220.
PMID 7166934.
67.^ Jean-Louis (1999). "Natural Toxins in Raw Foods and How Cooking Affects Them". Is
Cooked Food Poison?. Beyond Vegetarianism.
cooked-1g.shtml. Retrieved 2007-05-22.
68.^ George W. (2004). The Pesticide Book. Willoughby: MeisterPro. ISBN 1-892829-11-8.
69.^ Michael Smith, C. (2005). Plant Resistance to Arthropods: Molecular and Conventional
Approaches. Berlin: Springer. ISBN 1-4020-3701-5.
70.^ Dethier, V. G. (March 1954). "Evolution of feeding preferences in phytophagous insects".
Evolution (Society for the Study of Evolution) 8 (1): 33–54. doi:10.2307/2405664.
71.^ Russ, Karen. "Less toxic insecticides". Clemson University Home & Garden Information
Center. Retrieved 2007-05-27.
72.^ "Linden, J., Stoner, R., Knutson, K. Gardner-Hughes, C. “Organic Disease Control
Elicitors”. Agro Food Industry Hi-Te (p12-15 Oct 2000)".
73.^ van Emden, H.F. (November 1999). "Transgenic Host Plant Resistance to Insects—Some
Reservations". Annals of the Entomological Society of America 92 (6): 788–797. Retrieved
74.^ Ghosh, B. (2000). "Polyamines and plant alkaloids". Indian Journal of Experimental Biology
38 (11): 1086–91. PMID 11395950.
75.^ Eisner, Thomas (March 1990). "Prospecting for nature's chemical riches". Chemoecology 1
(1): 38–40. doi:10.1007/BF01240585.
76.^ Albert T. Sneden. "The Quinine Alkaloids" (PDF). Medicinal Chemistry and Drug Design . Retrieved 2007-05-23.
77.^ Albert T. Sneden. "The Tropane Alkaloids" (PDF). Medicinal Chemistry and Drug Design. Retrieved 2007-05-23.
78.^ Albert T. Sneden. "Taxol (Paclitaxe)" (PDF). Medicinal Chemistry and Drug Design . Retrieved 2007-05-23.

[edit] Further references

• Robert S. Fritz and Ellen L. Simms (editors) (1992). Plant resistance to herbivores and
pathogens: ecology, evolution, and genetics. Chicago: University of Chicago Press. ISBN 0-226-
• Hartley, Sue (2010) The 300 Million Years War: Plant Biomass v Herbivores Royal Institution
Christmas Lecture.
• Howe, H. F., and L. C. Westley. 1988. Ecological relationships of plants and animals. Oxford
University Press, Oxford, UK.
• Pierre Jolivet, (1998). Interrelationship Between Insects and Plants . Boca Raton: CRC.
ISBN 1-57444-052-7.
• Richard Karban and Ian T. Baldwin (1997). Induced responses to herbivory. Chicago:
University of Chicago Press. ISBN 0-226-42495-2.
• Martin R. Speight, Mark D. Hunter, Allan D. Watt (1999). Ecology of insects: concepts and
applications. Oxford: Blackwell Science. ISBN 0-86542-745-3.
• John N. Thompson (1994). The coevolutionary process. Chicago: University of Chicago
Press. ISBN 0-226-79759-7.
• Wiens, D. (1978). "Mimicry in plants". Evolutionary Biology 11: 365–403.

[edit] External links

• Bruce A. Kimball Evolutionary Plant Defense Strategies Life Histories and Contributions to
Future Generations
• Plant Defense Systems & Medicinal Botany
• Herbivore Defenses of Senecio viscosus L.

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This article is about the group of pteridophyte plants. For other uses, see Fern (disambiguation).
Ferns (Pteridophyta)
Temporal range: Mid Devonian[1]—Recent
Athyrium filix-femina unrolling young frond
Scientific classification

Kingdom: Plantae

Division: Pteridophyta


• †Cladoxylopsida
• Psilotopsida
• Equisetopsida (alias Sphenopsida)
• Marattiopsida
• Polypodiopsida (alias Pteridopsida,
A fern is any one of a group of about 12,000 species of plants.[3] Unlike mosses, they have xylem
and phloem (making them vascular plants). They have stems, leaves, and roots like other vascular plants.
Ferns do not have either seeds or flowers (they reproduce via spores).
By far the largest group of ferns are the leptosporangiate ferns, but ferns as defined here (also called
monilophytes) include horsetails, whisk ferns, marattioid ferns, and ophioglossoid ferns. The term
pteridophyte also refers to ferns (and possibly other seedless vascular plants; see classification section
below). A pteridologist is a specialist in the study of ferns and lycophytes.
Ferns first appear in the fossil record 360 million years ago in the Carboniferous but many of the
current families and species did not appear until roughly 145 million years ago in the late Cretaceous (after
flowering plants came to dominate many environments).
Ferns are not of major economic importance, but some are grown or gathered for food, as
ornamental plants, or for remediating contaminated soils. Some are significant weeds. They also featured in
mythology, medicine, and art.
• 1 Life cycle
• 2 Fern ecology
• 3 Fern structure
• 4 Evolution and classification
• 5 Smith's Classification
• 6 Uses
• 7 Cultural connotations
• 8 Misunderstood names
• 9 Gallery
• 10 See also
• 11 References
• 12 External links
[edit] Life cycle

Gametophyte (thalloid green mass) and sporophyte (ascendent frond) of Onoclea sensibilis
Ferns are vascular plants differing from lycophytes by having true leaves (megaphylls). They differ
from seed plants (gymnosperms and angiosperms) in their mode of reproduction—lacking flowers and seeds.
Like all other vascular plants, they have a life cycle referred to as alternation of generations, characterized by
a diploid sporophytic and a haploid gametophytic phase. Unlike the gymnosperms and angiosperms, the
ferns' gametophyte is a free-living organism.
Life cycle of a typical fern:
1. A sporophyte (diploid) phase produces haploid spores by meiosis.
2. A spore grows by mitosis into a gametophyte, which typically consists of a photosynthetic
3. The gametophyte produces gametes (often both sperm and eggs on the same prothallus) by
4. A mobile, flagellate sperm fertilizes an egg that remains attached to the prothallus.
5. The fertilized egg is now a diploid zygote and grows by mitosis into a sporophyte (the typical
"fern" plant).

[edit] Fern ecology

Ferns at Muir Woods, California

The stereotypic image of ferns growing in moist shady woodland nooks is far from being a complete
picture of the habitats where ferns can be found growing. Fern species live in a wide variety of habitats, from
remote mountain elevations, to dry desert rock faces, to bodies of water or in open fields. Ferns in general
may be thought of as largely being specialists in marginal habitats, often succeeding in places where various
environmental factors limit the success of flowering plants. Some ferns are among the world's most serious
weed species, including the bracken fern growing in the Scottish highlands, or the mosquito fern ( Azolla)
growing in tropical lakes, both species forming large aggressively spreading colonies. There are four
particular types of habitats that ferns are found in: moist, shady forests; crevices in rock faces, especially
when sheltered from the full sun; acid wetlands including bogs and swamps; and tropical trees, where many
species are epiphytes (something like a quarter to a third of all fern species[4]).
Many ferns depend on associations with mycorrhizal fungi. Many ferns only grow within specific pH
ranges; for instance, the climbing fern (Lygodium) of eastern North America will only grow in moist, intensely
acid soils, while the bulblet bladder fern ( Cystopteris bulbifera), with an overlapping range, is only found on
The spores are rich in lipids, protein and calories, so some vertebrates eat these. The European
woodmouse (Apodemus sylvaticus) has been found to eat the spores of Culcita macrocarpa and the bullfinch
(Pyrrhula murina) and the New Zealand lesser short-tailed bat (Mystacina tuberculata) also eat fern spores.

[edit] Fern structure

Ferns at the Royal Melbourne Botanical Gardens

Tree ferns, probably Dicksonia antarctica, growing in Nunniong, Australia

Like the sporophytes of seed plants, those of ferns consist of:
• Stems: Most often an underground creeping rhizome, but sometimes an above-ground
creeping stolon (e.g., Polypodiaceae), or an above-ground erect semi-woody trunk (e.g.,
Cyatheaceae) reaching up to 20 m in a few species (e.g., Cyathea brownii on Norfolk Island and
Cyathea medullaris in New Zealand).
• Leaf: The green, photosynthetic part of the plant. In ferns, it is often referred to as a frond, but
this is because of the historical division between people who study ferns and people who study seed
plants, rather than because of differences in structure. New leaves typically expand by the unrolling
of a tight spiral called a crozier or fiddlehead. This uncurling of the leaf is termed circinate vernation.
Leaves are divided into three types:
• Trophophyll: A leaf that does not produce spores, instead only producing sugars by
photosynthesis. Analogous to the typical green leaves of seed plants.
• Sporophyll: A leaf that produces spores. These leaves are analogous to the scales of
pine cones or to stamens and pistil in gymnosperms and angiosperms, respectively. Unlike
the seed plants, however, the sporophylls of ferns are typically not very specialized, looking
similar to trophophylls and producing sugars by photosynthesis as the trophophylls do.
• Brophophyll: A leaf that produces abnormally large amounts of spores. Their leaves
are also larger than the other leaves but bear a resemblance to trophophylls.
• Roots: The underground non-photosynthetic structures that take up water and nutrients from
soil. They are always fibrous and are structurally very similar to the roots of seed plants.
The gametophytes of ferns, however, are very different from those of seed plants. They typically
consist of:
• Prothallus: A green, photosynthetic structure that is one cell thick, usually heart or kidney
shaped, 3–10 mm long and 2–8 mm broad. The prothallus produces gametes by means of:
• Antheridia: Small spherical structures that produce flagellate sperm.
• Archegonia: A flask-shaped structure that produces a single egg at the bottom,
reached by the sperm by swimming down the neck.
• Rhizoids: root-like structures (not true roots) that consist of single greatly elongated cells,
water and mineral salts are absorbed over the whole structure. Rhizoids anchor the prothallus to the
One difference between sporophytes and gametophytes might be summed up by the saying that
"Nothing eats ferns, but everything eats gametophytes." This is an over-simplification, but it is true that
gametophytes are often difficult to find in the field because they are far more likely to be food than are the

[edit] Evolution and classification

Ferns first appear in the fossil record in the early-Carboniferous period. By the Triassic, the first
evidence of ferns related to several modern families appeared. The "great fern radiation" occurred in the late-
Cretaceous, when many modern families of ferns first appeared.
One problem with fern classification is the problem of cryptic species. A cryptic species is a species
that is morphologically similar to another species, but differs genetically in ways that prevent fertile
interbreeding. A good example of this is the currently designated species Asplenium trichomanes, the
maidenhair spleenwort. This is actually a species complex that includes distinct diploid and tetraploid races.
There are minor but unclear morphological differences between the two groups, which prefer distinctly
differing habitats. In many cases such as this, the species complexes have been separated into separate
species, thus raising the number of overall fern species. Possibly many more cryptic species are yet to be
discovered and designated.
Ferns have traditionally been grouped in the Class Filices, but modern classifications assign them
their own phylum or division in the plant kingdom, called Pteridophyta, also known as Filicophyta. The group
is also referred to as Polypodiophyta, (or Polypodiopsida when treated as a subdivision of tracheophyta
(vascular plants), although Polypodiopsida sometimes refers to only the leptosporangiate ferns). The term
"pteridophyte" has traditionally been used to describe all seedless vascular plants, making it synonymous
with "ferns and fern allies". This can be confusing since members of the fern phylum Pteridophyta are also
sometimes referred to as pteridophytes. The study of ferns and other pteridophytes is called pteridology, and
one who studies ferns and other pteridophytes is called a pteridologist.
Traditionally, three discrete groups of plants have been considered ferns: two groups of
eusporangiate ferns—families Ophioglossaceae (adders-tongues, moonworts, and grape-ferns) and
Marattiaceae—and the leptosporangiate ferns. The Marattiaceae are a primitive group of tropical ferns with a
large, fleshy rhizome, and are now thought to be a sibling taxon to the main group of ferns, the
leptosporangiate ferns. Several other groups of plants were considered "fern allies": the clubmosses,
spikemosses, and quillworts in the Lycopodiophyta, the whisk ferns in Psilotaceae, and the horsetails in the
Equisetaceae. More recent genetic studies have shown that the Lycopodiophyta are more distantly related to
other vascular plants, having radiated evolutionarily at the base of the vascular plant clade, while both the
whisk ferns and horsetails are as much "true" ferns as are the Ophioglossoids and Marattiaceae. In fact, the
whisk ferns and Ophioglossoids are demonstrably a clade, and the horsetails and Marattiaceae are arguably
another clade. Molecular data—which remain poorly constrained for many parts of the plants' phylogeny —
have been supplemented by recent morphological observations supporting the inclusion of Equisetaceae
within the ferns, notably relating to the construction of their sperm, and peculiarities of their roots.[2]
However, there are still differences of opinion about the placement of the Equisetum species (see
Equisetopsida for further discussion). One possible means of treating this situation is to consider only the
leptosporangiate ferns as "true" ferns, while considering the other three groups as "fern allies". In practice,
numerous classification schemes have been proposed for ferns and fern allies, and there has been little
consensus among them.
A 2006 classification by Smith et al. is based on recent molecular systematic studies, in addition to
morphological data. Their phylogeny is a consensus of a number of studies, and is shown below (to the level
of orders).[2]















Their classification based on this phylogeny divides extant ferns into four classes:
• Psilotopsida (whisk ferns and ophioglossoid ferns), about 92 species[6]
• Equisetopsida (horsetails), about 15 species[6]
• Marattiopsida, about 150 species[6]
• Polypodiopsida (leptosporangiate ferns), over 9000 species[6]
The last group includes most plants familiarly known as ferns. Modern research supports older ideas
based on morphology that the Osmundaceae diverged early in the evolutionary history of the
leptosporangiate ferns; in certain ways this family is intermediate between the eusporangiate ferns and the
leptosporangiate ferns.

[edit] Smith's Classification

The complete classification scheme proposed by Smith et al. (2006) is shown below (alternative
names in brackets).[2]
• Class Psilotopsida
• Order Ophioglossales
• Family Ophioglossaceae (incl. Botrychiaceae, Helminthostachyaceae)
• Order Psilotales
• Family Psilotaceae (incl. Tmesipteridaceae)
• Class Equisetopsida [=Sphenopsida]
• Order Equisetales
• Family Equisetaceae
• Class Marattiopsida
• Order Marattiales
• Family Marattiaceae (incl. Angiopteridaceae, Christenseniaceae,
Danaeaceae, Kaulfussiaceae)
• Class Pteridopsida [=Filicopsida, Polypodiopsida]
• Order Osmundales
• Family Osmundaceae
• Order Hymenophyllales
• Family Hymenophyllaceae (incl. Trichomanaceae)
• Order Gleicheniales
• Family Gleicheniaceae (incl. Dicranopteridaceae, Stromatopteridaceae)
• Family Dipteridaceae (incl. Cheiropleuriaceae)
• Family Matoniaceae
• Order Schizaeales
• Family Lygodiaceae
• Family Anemiaceae (incl. Mohriaceae)
• Family Schizaeaceae
• Order Salviniales
• Family Marsileaceae (incl. Pilulariaceae)
• Family Salviniaceae (incl. Azollaceae)
• Order Cyatheales
• Family Thyrsopteridaceae
• Family Loxsomataceae
• Family Culcitaceae
• Family Plagiogyriaceae
• Family Cibotiaceae
• Family Cyatheaceae (incl. Alsophilaceae, Hymenophyllopsidaceae)
• Family Dicksoniaceae (incl. Lophosoriaceae)
• Family Metaxyaceae
• Order Polypodiales
Adiantum lunulatum from Family Pteridaceae
• Family Lindsaeaceae (incl. Cystodiaceae, Lonchitidaceae)
• Family Saccolomataceae
• Family Dennstaedtiaceae (incl. Hypolepidaceae, Monachosoraceae,
• Family Pteridaceae (incl. Pellaeaceae, Adiantaceae, Ceratopteridaceae,
• Family Aspleniaceae
• Family Thelypteridaceae
• Family Woodsiaceae (incl. Athyriaceae, Cystopteridaceae)
• Family Blechnaceae (incl. Stenochlaenaceae)
• Family Onocleaceae
• Family Dryopteridaceae (incl. Aspidiaceae, Bolbitidaceae,
Elaphoglossaceae, Hypodematiaceae, Peranemataceae)
• Family Oleandraceae
• Family Davalliaceae
• Family Lomariopsidaceae (incl. Nephrolepis)
• Family Polypodiaceae (incl. Drynariaceae, Grammitidaceae,
Gymnogrammitidaceae, Loxogrammaceae, Platyceriaceae, Pleurisoriopsidaceae)
• Family Tectariaceae

[edit] Uses
Ferns are not as important economically as seed plants but have considerable importance in some
societies. Some ferns are used for food, including the fiddleheads of bracken, Pteridium aquilinum, ostrich
fern, Matteuccia struthiopteris, and cinnamon fern, Osmunda cinnamomea. Diplazium esculentum is also
used by some tropical peoples as food. Tubers from the King Fern or para (Ptisana salicina) are a traditional
food in New Zealand and the South Pacific. Fern tubers were used for food 30,000 years ago in Europe.[7][8]
Fern tubers were used by the Guanches to make gofio in the Canary Islands. Licorice fern rhizomes were
chewed by the natives of the Pacific Northwest for their flavor.
Ferns of the genus Azolla are very small, floating plants that do not resemble ferns. Called mosquito
fern, they are used as a biological fertilizer in the rice paddies of southeast Asia, taking advantage of their
ability to fix nitrogen from the air into compounds that can then be used by other plants.
Many ferns are grown in horticulture as landscape plants, for cut foliage and as houseplants,
especially the Boston fern (Nephrolepis exaltata) and other members of the genus Nephrolepis. The Bird's
Nest Fern (Asplenium nidus) is also popular, as is the staghorn ferns (genus Platycerium). Perennial (also
known as hardy) ferns planted in gardens in the northern hemisphere also have a considerable following.
Several ferns are noxious weeds or invasive species, including Japanese climbing fern (Lygodium
japonicum), mosquito fern and sensitive fern (Onoclea sensibilis). Giant water fern (Salvinia molesta) is one
of the world's worst aquatic weeds. The important fossil fuel coal consists of the remains of primitive plants,
including ferns.
Ferns have been studied and found to be useful in the removal of heavy metals, especially arsenic,
from the soil. Other ferns with some economic significance include:
• Dryopteris filix-mas (male fern), used as a vermifuge, and formerly in the US Pharmacopeia;
also, this fern accidentally sprouting in a bottle resulted in Nathaniel Bagshaw Ward's 1829 invention
of the terrarium or Wardian case
• Rumohra adiantiformis (floral fern), extensively used in the florist trade
• Microsorum pteropus (Java fern), one of the most popular freshwater aquarium plants.
• Osmunda regalis (royal fern) and Osmunda cinnamomea (cinnamon fern), the root fiber
being used horticulturally; the fiddleheads of O. cinnamomea are also used as a cooked vegetable
• Matteuccia struthiopteris (ostrich fern), the fiddleheads used as a cooked vegetable in North
• Pteridium aquilinum or Pteridium esculentum (bracken), the fiddleheads used as a cooked
vegetable in Japan and are believed to be responsible for the high rate of stomach cancer in Japan.
It is also one of the world's most important agricultural weeds, especially in the British highlands, and
often poisons cattle and horses.
• Diplazium esculentum (vegetable fern), a source of food for some native societies
• Pteris vittata (brake fern), used to absorb arsenic from the soil
• Polypodium glycyrrhiza (licorice fern), roots chewed for their pleasant flavor
• Tree ferns, used as building material in some tropical areas
• Cyathea cooperi (Australian tree fern), an important invasive species in Hawaii
• Ceratopteris richardii, a model plant for teaching and research, often called C-fern
[edit] Cultural connotations

Blätter des Manns Walfarn. by Alois Auer, Vienna: Imperial Printing Office, 1853
Ferns figure in folklore, for example in legends about mythical flowers or seeds.[9] In Slavic folklore,
ferns are believed to bloom once a year, during the Ivan Kupala night. Although alleged to be exceedingly
difficult to find, anyone who sees a "fern flower" is thought to be guaranteed to be happy and rich for the rest
of their life. Similarly, Finnish tradition holds that one who finds the "seed" of a fern in bloom on Midsummer
night will, by possession of it, be guided and be able to travel invisibly to the locations where eternally blazing
Will o' the wisps called aarnivalkea mark the spot of hidden treasure. These spots are protected by a spell
that prevents anyone but the fern-seed holder from ever knowing their locations.[10]
"Pteridomania"' is a term for the Victorian era craze of fern collecting and fern motifs in decorative art
including pottery, glass, metals, textiles, wood, printed paper, and sculpture "appearing on everything from
christening presents to gravestones and memorials." The fashion for growing ferns indoors led to the
development of the Wardian case, a glazed cabinet that would exclude air pollutants and maintain the
necessary humidity.[11]

Barnsley fern created using chaos game, through an Iterated function system (IFS).
The dried form of ferns was also used in other arts, being used as a stencil or directly inked for use in
a design. The botanical work, The Ferns of Great Britain and Ireland , is a notable example of this type of
nature printing. The process, patented by the artist and publisher Henry Bradbury, impressed a specimen on
to a soft lead plate. The first publication to demonstrate this was Alois Auer's The Discovery of the Nature

[edit] Misunderstood names

Several non-fern plants are called "ferns" and are sometimes confused with true ferns. These
• "Asparagus fern"—This may apply to one of several species of the monocot genus
Asparagus, which are flowering plants.
• "Sweetfern"—A flowering shrub of the genus Comptonia.
• "Air fern"—A group of animals called hydrozoan that are distantly related to jellyfish and
corals. They are harvested, dried, dyed green, and then sold as a "plant" that can "live on air". While
it may look like a fern, it is merely the skeleton of this colonial animal.
• "Fern bush"—Chamaebatiaria millefolium—a rose family shrub with fern-like leaves.
In addition, the book Where the Red Fern Grows has elicited many questions about the mythical "red
fern" named in the book. There is no such known plant, although there has been speculation that the oblique
grape-fern, Sceptridium dissectum, could be referred to here, because it is known to appear on disturbed
sites and its fronds may redden over the winter.
[edit] Gallery

Fern leaf, probably

Tree fern, probably T
Adiantum lunulatum Blechnum nudum
A tree fern Dicksonia antarctica Dicksonia
unrolling a new frond

"Filicinae" from Ernst Leaf of fern with spor
Unidentified tree Tree Fern Spores Rotorua,
Haeckel's Kunstformen der
fern in Oaxaca San Diego, CA
Natur, 1904
Ferns in one of
many natural Coast F
Nature prints in The Fern fronds
Redwood undergrowth forest can
Ferns of Great Britain and A young, newly-
settings Santa Cruz, CA. Franklin,
Ireland used fronds to formed fern frond
produce the plates

[edit] See also

• Fern spike
• Fiddlehead greens
• Pteridomania

[edit] References
1. ^ Wattieza, Stein, W. E., F. Mannolini, L. V. Hernick, E. Landling, and C. M. Berry. 2007.
"Giant cladoxylopsid trees resolve the enigma of the Earth's earliest forest stumps at Gilboa", Nature
(19 April 2007) 446:904–907.
2. ^ a b c d Smith, A.R.; Pryer, K.M.; Schuettpelz, E.; Korall, P.; Schneider, H.; Wolf, P.G. (2006).
"A classification for extant ferns". Taxon 55 (3): 705–731. doi:10.2307/25065646. Retrieved 2008-02-12.
3. ^ Chapman, Arthur D. (2009). Numbers of Living Species in Australia and the World . Report
for the Australian Biological Resources Study. Canberra, Australia. September 2009.
4. ^ Schuettpelz, Eric. "Fern Phylogeny Inferred from 400 Leptosporangiate Species and Three
Plastid Genes," contained in "The Evolution and Diversification of Epiphytic Ferns." Doctoral
dissertation, Duke University. 2007.
5. ^ Walker, Matt (19 February 2010). "A mouse that eats ferns like a dinosaur". BBC Earth
News. Retrieved 20
February 2010.
6. ^ a b c d Eric Schuettpelz (2007). "table 1". The evolution and diversification of epiphytic ferns.
Duke University PhD thesis.
7. ^ "Stone Age humans liked their burgers in a bun", Sonia Van Gilder Cooke, New Scientist,
23 Oct. 2010, p. 18.
8. ^ "Thirty thousand-year-old evidence of plant food processing" by Anna Revedin et al.,
PNAS, published online Oct. 18, 2010.
9. ^ May, Lenore Wile (1978). "The economic uses and associated folklore of ferns and fern
allies". The Botanical Review 44 (4): 491–528. doi:10.1007/BF02860848
11.^ * Boyd, Peter D. A. (2002-01-02). Pteridomania - the Victorian passion for ferns. Revised:
web version. Antique Collecting 28, 6, 9–12.. Retrieved
• Pryer, Kathleen M., Harald Schneider, Alan R. Smith, Raymond Cranfill, Paul G. Wolf, Jeffrey
S. Hunt and Sedonia D. Sipes. 2001. Horsetails and ferns are a monophyletic group and the closest
living relatives to seed plants. Nature 409: 618–622 (abstract here).
• Pryer, Kathleen M., Eric Schuettpelz, Paul G. Wolf, Harald Schneider, Alan R. Smith and
Raymond Cranfill. 2004. Phylogeny and evolution of ferns (monilophytes) with a focus on the early
leptosporangiate divergences. American Journal of Botany 91:1582–1598 (online abstract here).
• Moran, Robbin C. (2004). A Natural History of Ferns. Portland, OR: Timber Press. ISBN 0-
• Lord, Thomas R. (2006). Ferns and Fern Allies of Pennsylvania. Indiana, PA: Pinelands
Press. [1]

[edit] External links

Wikispecies has information related to: Pteridophyta

Wikimedia Commons has media related to: Pteridophyta

• Tree of Life Web Project: Filicopsida

• A classification of the ferns and their allies
• A fern book bibliography
• Register of fossil Pteridophyta
• L. Watson and M.J. Dallwitz (2004 onwards). The Ferns (Filicopsida) of the British Isles.
• Ferns and Pteridomania in Victorian Scotland
• Non-seed plant images at
• "American Fern Society"
• "British Pteridological Society"
• Checklist of Ferns and Lycophytes of the World

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Animal echolocation
From Wikipedia, the free encyclopedia
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A depiction of the ultrasound signals emitted by a bat, and the echo from a nearby object.
Echolocation, also called biosonar, is the biological sonar used by several animals, most notably
microchiropteran bats and odontocetes (toothed whales and dolphins), but has also been demonstrated in
simpler form in other groups such as shrews, one genus of megachiropteran bats (Rousettus) and two cave
dwelling bird groups, the so called cave swiftlets in the genus Aerodramus (formerly Collocalia) and the
unrelated Oilbird Steatornis caripensis. The term echolocation was coined by Donald Griffin, whose work with
Robert Galambos was the first to conclusively demonstrate its existence in bats in 1938.[1][2]. However, long
before that, the Italian 18th century scientist Lazzaro Spallanzani had, by means of a series of elaborate
experiments concluded that bats navigated by hearing and not vision[3]. Echolocation in odontocetes was not
properly described before two decades later, by Schevill and McBride[4].
Echolocating animals emit calls out to the environment and listen to the echoes of those calls that
return from various objects in the environment. They use these echoes to locate, range, and identify the
objects. Echolocation is used for navigation and for foraging (or hunting) in various environments.
• 1 Basic principle
• 2 Bats
• 2.1 Calls and ecology
• 2.2 Acoustic features
• 2.2.1 FM Signal Advantages
• 2.2.2 CF signal advantages
• 2.2.3 Acoustic environments of FM and CF
• 2.3 Neural mechanisms in the brain
• 2.4 Inner ear and primary sensory neurons
• 2.5 Inferior colliculus
• 2.6 Auditory cortex
• 3 Toothed whales
• 4 Oilbirds and swiftlets
• 5 Shrews and tenrecs
• 6 See also
• 7 References
• 8 External links
[edit] Basic principle
Echolocation is the same as active sonar, using sounds made by the animal itself. Ranging is done
by measuring the time delay between the animal's own sound emission and any echoes that return from the
environment. The relative intensity of sound received at each ear as well as the time delay between arrival at
the two ears provide information about the horizontal angle (azimuth) from which the reflected sound waves
arrive.[5] Unlike some man-made sonars that relies on many extremely narrow beams and many receivers to
localize a target (multibeam sonar), animal echolocation has only one transmitter and two receivers (the
ears). Echolocating animals have two ears positioned slightly apart. The echoes returning to the two ears
arrive at different times and at different loudness levels, depending on the position of the object generating
the echoes. The time and loudness differences are used by the animals to perceive distance and direction.
With echolocation, the bat or other animal can see not only where it is going but also how big another animal
is, what kind of animal it is, and other features.
[edit] Bats

Spectrogram of Pipistrellus Bat vocalizations. Detail is shown as the pulses transition to a faster
repetition rate. The bat appears to use a hybrid pulse which combines a sharp falling frequency chirp with an
extended constant frequency tail. Such a waveform may offer combined benefits of range estimation as well
as Doppler shift detection. Spectrogram generated with Fatpigdog's PC based Real Time FFT Spectrum
Pipistrellus Pulses

Recording of Pipistrellus
bat approaching its prey.

Problems listening to this file? See media help.

Microbats use echolocation to navigate and forage, often in total darkness. They generally emerge
from their roosts in caves, attics, or trees at dusk and hunt for insects into the night. Their use of echolocation
allows them to occupy a niche where there are often many insects (that come out at night since there are
fewer predators then) and where there is less competition for food, and where there are fewer other species
that may prey on the bats themselves.
Microbats generate ultrasound via the larynx and emit the sound through the open mouth or, much
more rarely, the nose. The latter is most pronounced in the horseshoe bats (Rhinolophus spp.). Microbat
calls (help·info) range in frequency from 14,000 to well over 100,000 Hz, mostly beyond the range of the
human ear (typical human hearing range is considered to be from 20 Hz to 20,000 Hz). Bats may estimate
the elevation of targets by interpreting the interference patterns caused by the echoes reflecting from the
tragus, a flap of skin in the external ear[6]. There are two hypotheses about the evolution of echolocation in
bats. The first suggests that laryngeal echolocation evolved twice in Chiroptera, once in Yangochiroptera and
once in the rhinolophoids.[7][8] The second proposes that laryngeal echolocation had a single origin in
Chiroptera, was subsequently lost in the family Pteropodidae (all megabats), and later evolved as a system
of tongue-clicking in the genus Rousettus.[9]
Individual bat species echolocate within specific frequency ranges that suit their environment and
prey types. This has sometimes been used by researchers to identify bats flying in an area simply by
recording their calls with ultrasonic recorders known as 'bat detectors'. However echolocation calls are not
always species specific and some bats overlap in the type of calls they use so recordings of echolocation
calls cannot be used to identify all bats. In recent years researchers in several countries have developed 'bat
call libraries' that contain recordings of local bat species that have been identified known as 'reference calls'
to assist with identification.
Since the 1970s there has been an ongoing controversy among researchers as to whether bats use
a form of processing known from radar termed coherent cross-correlation. Coherence means that the phase
of the echolocation signals is used by the bats, while cross-correlation just implies that the outgoing signal is
compared with the returning echoes in a running process. Today most - but not all - researchers believe that
they use cross-correlation, but in an incoherent form, termed a filter bank receiver.
When searching for prey they produce sounds at a low rate (10-20/sec). During the search phase the
sound emission is coupled to respiration, which is again coupled to the wingbeat. This coupling appears to
dramatically conserve energy as there is little to no additional energetic cost of echolocation to flying bats
(Speakman and Racey 1991). After detecting a potential prey item, microbats increase the rate of pulses,
ending with the terminal buzz, at rates as high as 200/sec. During approach to a detected target, the duration
of the sounds is gradually decreased, as is the energy of the sound.

[edit] Calls and ecology

Bats belonging to the suborder Microchiroptera (microbats) occupy a diverse set of ecological
conditions - they can be found living in environments as different as Europe and Madagascar, and hunting for
food sources as different as insects, frogs, nectar, fruit, and blood. Additionally, the characteristics of an
echolocation call are adapted to the particular environment, hunting behavior, and food source of the
particular bat. However, this adaptation of echolocation calls to ecological factors is constrained by the
phylogenetic relationship of the bats, leading to a process known as descent with modification, and resulting
in the diversity seen in the Microchiropteran suborder today. (Jones and Teeling 2006; Grinnell 1995; Zupanc
2004; Fenton 1995; Neuweiler 2003; Simmons and Stein 1980)

[edit] Acoustic features

Describing the diversity of bat echolocation calls requires examination of the frequency and temporal
features of the calls. It is the variations in these aspects that produce echolocation calls suited for different
acoustic environments and hunting behaviors. (Fenton 2005; Jones and Teeling 2006; Zupanc 2004;
Simmons and Stein 1980; Hiryu et al. 2007)
• Frequency Modulation and Constant Frequency: Echolocation calls can be composed of two
different types of frequency structures: frequency modulated (FM) sweeps, and constant frequency
(CF) tones. A particular call can consist of one, the other, or both structures. An FM sweep is a
broadband signal – that is, it contains a downward sweep through a range of frequencies. A CF tone
is a narrowband signal: the sound stays constant at one frequency throughout its duration.
• Intensity: Echolocation calls have been measured at intensities anywhere between 60 and
110 decibels. Certain microbat species can modify their call intensity mid-call, lowering the intensity
as they approach objects that reflect sound strongly. This prevents the returning echo from deafening
the bat (Hiryu et al. 2007). Additionally, the so-called "whispering bats" have adapted low-amplitude
echolocation so that their prey, moths, which are able to hear echolocation calls, are less able to
detect and avoid an oncoming bat (Fullard 1997).
• Harmonic composition: Calls can be composed of one frequency, or multiple frequencies
comprising a harmonic series. In the latter case, the call is usually dominated by a certain harmonic
("dominant" frequencies are those present at higher intensities than other harmonics present in the
• Call duration: A single echolocation call (a call being a single continuous trace on a sound
spectrogram, and a series of calls comprising a sequence or pass) can last anywhere from 0.2 to 100
milliseconds in duration, depending on the stage of prey-catching behavior that the bat is engaged in.
For example, the duration of a call usually decreases when the bat is in the final stages of prey
capture – this enables the bat to call more rapidly without overlap of call and echo. Reducing duration
comes at the cost of having less total sound available for reflecting off objects and being heard by the
• Pulse interval: The time interval between subsequent echolocation calls (or pulses)
determines two aspects of a bat's perception. First, it establishes how quickly the bat's auditory
scene information is updated. For example, bats increase the repetition rate of their calls (that is,
decrease the pulse interval) as they home in on a target. This allows the bat to get new information
regarding the target's location at a faster rate when it needs it most. Secondly, the pulse interval
determines the maximum range that bats can detect objects. This is because bats can only keep
track of the echoes from one call at a time; as soon as they make another call they stop listening for
echoes from the previously made call (Wilson and Moss 2004). For example, a pulse interval of 100
ms (typical of a bat searching for insects) allows sound to travel in air roughly 34 meters so a bat can
only detect objects as far away as 17 meters (the sound has to travel out and back). With a pulse
interval of 5 ms (typical of a bat in the final moments of a capture attempt), the bat can only detect
objects up to 85 cm away. Therefore the bat constantly has to make a choice between getting new
information updated quickly and detecting objects far away.
[edit] FM Signal Advantages

Echolocation call produced by Pipistrellus pipistrellus, an FM bat. The ultrasonic call has been
"heterodyned" - multiplied by a constant frequency to produce frequency subtraction, and thus an audible
sound - by a bat detector. A key feature of the recording is the increase in the repetition rate of the call as the
bat nears its target - this is called the "terminal buzz".
The major advantage conferred by an FM signal is extremely precise range discrimination, or
localization, of the target. J.A. Simmons demonstrated this effect with a series of elegant experiments that
showed how bats using FM signals could distinguish between two separate targets even when the targets
were less than half a millimeter apart. This amazing ability is due to the broadband sweep of the signal,
which allows for better resolution of the time delay between the call and the returning echo, thereby
improving the cross correlation of the two. Additionally, if harmonic frequencies are added to the FM signal,
then this localization becomes even more precise. (Jones and Teeling 2006; Zupanc 2004; Simmons and
Stein 1980; Grinnell 1995)
One possible disadvantage of the FM signal is a decreased operational range of the call. Because
the energy of the call is spread out among many frequencies, the distance at which the FM-bat can detect
targets is limited (Fenton 1995). This is in part because any echo returning at a particular frequency can only
be evaluated for a brief fraction of a millisecond, as the fast downward sweep of the call does not remain at
any one frequency for long (Grinnell 1995).
[edit] CF signal advantages
The structure of a CF signal is adaptive in that it allows the CF-bat to detect both the velocity of a
target, and the fluttering of a target's wings as Doppler shifted frequencies. A Doppler shift is an alteration in
sound wave frequency, and is produced in two relevant situations: when the bat and its target are moving
relative to each other, and when the target's wings are oscillating back and forth. CF-bats must compensate
for Doppler shifts, lowering the frequency of their call in response to echoes of elevated frequency - this
ensures that the returning echo remains at the frequency to which the ears of the bat are most finely tuned.
The oscillation of a target's wings also produces amplitude shifts, which gives a CF-bat additional help in
distinguishing a flying target from a stationary one. (Schnitzler and Flieger 1983; Zupanc 2004; Simmons and
Stein 1980; Grinnell 1995; Neuweiler 2003; Jones and Teeling 2006)
Additionally, because the signal energy of a CF call is concentrated into a narrow frequency band,
the operational range of the call is much greater than that of an FM signal. This relies on the fact that echoes
returning within the narrow frequency band can be summed over the entire length of the call, which maintains
a constant frequency for up to 100 milliseconds (Grinnell 1995; Fenton 1995)

[edit] Acoustic environments of FM and CF signals

• FM: An FM component is excellent for hunting prey while flying in close, cluttered
environments. Two aspects of the FM signal account for this fact: the precise target localization
conferred by the broadband signal, and the short duration of the call. The first of these is essential
because in a cluttered environment, the bats must be able to resolve their prey from large amounts of
background noise. The 3D localization abilities of the broadband signal enable the bat to do exactly
that, providing it with what Simmons and Stein (1980) call a "clutter rejection strategy." This strategy
is further improved by the use of harmonics, which, as previously stated, enhance the localization
properties of the call. The short duration of the FM call is also best in close, cluttered environments
because it enables the bat to emit many calls extremely rapidly without overlap. This means that the
bat can get an almost continuous stream of information – essential when objects are close, because
they will pass by quickly – without confusing which echo corresponds to which call. (Neuweiler 2003;
Simmons and Stein 1980; Jones and Teeling 2006; Fenton 1995)
• CF: A CF component is often used by bats hunting for prey while flying in open, clutter-free
environments, or by bats that wait on perches for their prey to appear. The success of the former
strategy is due to two aspects of the CF call, both of which confer excellent prey-detection abilities.
First, the greater working range of the call allows bats to detect targets present at great distances – a
common situation in open environments. Second, the length of the call is also suited for targets at
great distances: in this case, there is a decreased chance that the long call will overlap with the
returning echo. The latter strategy is made possible by the fact that the long, narrowband call allows
the bat to detect Doppler shifts, which would be produced by an insect moving either towards or
away from a perched bat. (Neuweiler 2003; Simmons and Stein 1980; Jones and Teeling 2006;
Fenton 1995)

[edit] Neural mechanisms in the brain

Because bats use echolocation to orient themselves and to locate objects, their auditory systems are
adapted for this purpose, highly specialized for sensing and interpreting the stereotyped echolocation calls
characteristic of their own species. This specialization is evident from the inner ear up to the highest levels of
information processing in the auditory cortex.
[edit] Inner ear and primary sensory neurons
Both CF and FM bats have specialized inner ears which allow them to hear sounds in the ultrasonic
range, far outside the range of human hearing. Although in most other aspects, the bat's auditory organs are
similar to those of most other mammals, certain bats (horseshoe bats, Rhinolophus spp. and the moustached
bat, Pteronotus parnelii) with a constant frequency (CF) component to their call (known as high duty cycle
bats) do have a few additional adaptations for detecting the predominant frequency (and harmonics) of the
CF vocalization. These include a narrow frequency "tuning" of the inner ear organs, with an especially large
area responding to the frequency of the bat's returning echoes (Neuweiler 2003).
The basilar membrane within the cochlea contains the first of these specializations for echo
information processing. In bats that use CF signals, the section of membrane that responds to the frequency
of returning echoes is much larger than the region of response for any other frequency. For example, in
Rhinolophus ferrumequinum, the horseshoe bat, there is a disproportionately lengthened and thickened
section of the membrane that responds to sounds around 83 kHz, the constant frequency of the echo
produced by the bat's call. This area of high sensitivity to a specific, narrow range of frequency is known as
an "acoustic fovea" (Schuller and Pollack 1979).
Odontocetes (toothed whales and dolphins) have similar cochlear specializations to those found in
bats. Odontocetes also have the highest neural investment of any cochleae reported to date with ratios of
greater than 1500 ganglion cells/mm of basilar membrane.
Further along the auditory pathway, the movement of the basilar membrane results in the stimulation
of primary auditory neurons. Many of these neurons are specifically "tuned" (respond most strongly) to the
narrow frequency range of returning echoes of CF calls. Because of the large size of the acoustic fovea, the
number of neurons responding to this region, and thus to the echo frequency, is especially high (Carew
[edit] Inferior colliculus
In the Inferior colliculus, a structure in the bat's midbrain, information from lower in the auditory
processing pathway is integrated and sent on to the auditory cortex. As George Pollak and others showed in
a series of papers in 1977, the interneurons in this region have a very high level of sensitivity to time
differences, since the time delay between a call and the returning echo tells the bat its distance from the
target object. Especially interesting is that while most neurons respond more quickly to stronger stimuli,
collicular neurons maintain their timing accuracy even as signal intensity changes.
These interneurons are specialized for time sensitivity in several ways. First, when activated, they
generally respond with only one or two action potentials. This short duration of response allows their action
potentials to give a very specific indication of the exact moment of the time when the stimulus arrived, and to
respond accurately to stimuli that occur close in time to one another. In addition, the neurons have a very low
threshold of activation – they respond quickly even to weak stimuli. Finally, for FM signals, each interneuron
is tuned to a specific frequency within the sweep, as well as to that same frequency in the following echo.
There is specialization for the CF component of the call at this level as well. The high proportion of neurons
responding to the frequency of the acoustic fovea actually increases at this level (Carew 2001, Pollak 1977,
Zupanc 2004).

[edit] Auditory cortex

The auditory cortex in bats is quite large in comparison with other mammals (Anderson 1995).
Various characteristics of sound are processed by different regions of the cortex, each providing different
information about the location or movement of a target object. Most of the existing studies on information
processing in the auditory cortex of the bat have been done by Nobuo Suga on the mustached bat,
Pteronotus parnellii. This bat's call has both CF tone and FM sweep components.
Suga and his colleagues have shown that the cortex contains a series of "maps" of auditory
information, each of which is organized systematically based on characteristics of sound such as frequency
and amplitude. The neurons in these areas respond only to a specific combination of frequency and timing
(sound-echo delay), and are known as combination-sensitive neurons.
The systematically organized maps in the auditory cortex respond to various aspects of the echo
signal, such as its delay and its velocity. These regions are composed of "combination sensitive" neurons
that require at least two specific stimuli to elicit a response. The neurons vary systematically across the
maps, which are organized by acoustic features of the sound and can be two dimensional. The different
features of the call and its echo are used by the bat to determine important characteristics of their prey. The
maps include:
Sketch of the regions of the auditory cortex in a bat's brain
• FM-FM area: This region of the cortex contains FM-FM combination-sensitive neurons.
These cells respond only to the combination of two FM sweeps: a call and its echo. The neurons in
the FM-FM region are often referred to as "delay-tuned," since each responds to a specific time delay
between the original call and the echo, in order to find the distance from the target object (the range).
Each neuron also shows specificity for one harmonic in the original call and a different harmonic in
the echo. The neurons within the FM-FM area of the cortex of Pteronotus are organized into
columns, in which the delay time is constant vertically but increases across the horizontal plane. The
result is that range is encoded by location on the cortex, and increases systematically across the FM-
FM area (Suga et al. 1975, Suga et al. 1979, Neuweiler 2003, Carew 2001).
• CF-CF area: Another kind of combination-sensitive neuron is the CF-CF neuron. These
respond best to the combination of a CF call containing two given frequencies – a call at 30 kHz
(CF1) and one of its additional harmonics around 60 or 90 kHz (CF2 or CF3) – and the corresponding
echoes. Thus, within the CF-CF region, the changes in echo frequency caused by the Doppler shift
can be compared to the frequency of the original call to calculate the bat's velocity relative to its
target object. As in the FM-FM area, information is encoded by its location within the map-like
organization of the region. The CF-CF area is first split into the distinct CF1-CF2 and CF1-CF3 areas.
Within each area, the CF1 frequency is organized on an axis, perpendicular to the CF2 or CF3
frequency axis. In the resulting grid, each neuron codes for a certain combination of frequencies that
is indicative of a specific velocity (Suga et al. 1975, Suga et al. 1987, Carew 2001).
• DSCF area: This large section of the cortex is a map of the acoustic fovea, organized by
frequency and by amplitude. Neurons in this region respond to CF signals that have been Doppler
shifted (in other words, echoes only) and are within the same narrow frequency range to which the
acoustic fovea responds. For Pteronotus, this is around 61 kHz. This area is organized into columns,
which are arranged radially based on frequency. Within a column, each neuron responds to a
specific combination of frequency and amplitude. Suga's studies have indicated that this brain region
is necessary for frequency discrimination (Suga et al. 1975, Suga et al. 1987, Carew 2001).

[edit] Toothed whales

Diagram illustrating sound generation, propagation and reception in a toothed whale. Outgoing
sounds are red and incoming ones are green
Biosonar is valuable to Toothed whales (suborder odontoceti), including dolphins, porpoises, river
dolphins, killer whales and sperm whales, because they live in an underwater habitat that has favourable
acoustic characteristics and where vision is extremely limited in range due to absorption or turbidity.
Echolocation evolved in these aquatic mammals during the late Eocene (around 30 million years ago) maybe
as an adaptation to diel migrating cephalopods.[10][11] Since its advent, there has been adaptive radiation
especially in the Delphinidae family (dolphins) in which echolocation has become extremely derived.[12]
One specific type of echolocation, narrow-band high frequency (NBHF) clicks, evolved at least four
times in groups of odontocetes, including pygmy sperm whale (Kogiidae) and porpoise (Phocoenidae)
families, Pontoporia blainvillei, the genus Cephalorhynchus, and part of the genus Lagenorhynchus.[13][14]
These high frequency clicks likely evolved as adaptation of predator avoidance, as they inhabit areas that
have many killer whales and the signals are inaudible to killer whales due to the absence of energy below
100 kHz. Another reason for variation in echolocation frequencies is habitat. Shallow waters, where many of
these species live, tend to have more debris; a more directional transmission reduces clutter in reception.[14]
Toothed whales emit a focused beam of high-frequency clicks in the direction that their head is
pointing. Sounds are generated by passing air from the bony nares through the phonic lips.[15] These
sounds are reflected by the dense concave bone of the cranium and an air sac at its base. The focused beam
is modulated by a large fatty organ known as the 'melon'. This acts like an acoustic lens because it is
composed of lipids of differing densities. Most toothed whales use clicks in a series, or click train, for
echolocation, while the sperm whale may produce clicks individually. Toothed whale whistles do not appear
to be used in echolocation. Different rates of click production in a click train give rise to the familiar barks,
squeals and growls of the bottlenose dolphin. A click train with a repetition rate over 600 per second is called
a burst pulse. In bottlenose dolphins, the auditory brain response resolves individual clicks up to 600 per
second, but yields a graded response for higher repetition rates.
It has been suggested that some smaller toothed whales may have their tooth arrangement suited to
aid in echolocation. The placement of teeth in the jaw of a bottlenose dolphin, as an example, are not
symmetrical when seen from a vertical plane, and this asymmetry could possibly be an aid in the dolphin
sensing if echoes from its biosonar are coming from one side or the other.[16][17]. However, this idea lacks
experimental support.
Echoes are received using complex fatty structures around the lower jaw as the primary reception
path, from where they are transmitted to the middle ear via a continuous fat body (Ketten 1992,2000). Lateral
sound may be received though fatty lobes surrounding the ears with a similar density to water. Some
researchers believe that when they approach the object of interest, they protect themselves against the
louder echo by quietening the emitted sound. In bats this is known to happen, but here the hearing sensitivity
is also reduced close to a target.
Before the echolocation abilities of "porpoises" were officially discovered, Jacques Yves Cousteau
suggested that they might exist. In his first book, The Silent World (1953, pp. 206–207), he reported that his
research vessel, the Élie Monier, was heading to the Straits of Gibraltar and noticed a group of porpoises
following them. Cousteau changed course a few degrees off the optimal course to the center of the strait, and
the porpoises followed for a few minutes, then diverged toward mid-channel again. It was obvious that they
knew where the optimal course lay, even if the humans didn't. Cousteau concluded that the cetaceans had
something like sonar, which was a relatively new feature on submarines.

[edit] Oilbirds and swiftlets

Oilbirds and some species of swiftlet are known to use a relatively crude form of echolocation
compared to that of bats and dolphins. These nocturnal birds emit calls while flying and use the calls to
navigate through trees and caves where they live.[18][19]

[edit] Shrews and tenrecs

Main article: Shrews#Echolocation
Terrestrial mammals other than bats known to echolocate include two genera ( Sorex and Blarina) of
shrews and the tenrecs of Madagascar.[20] These include the wandering shrew (Sorex vagrans), the
common or Eurasian shrew (Sorex araneus), and the short-tailed shrew (Blarina brevicauda).[21] The nature
of shrew sounds unlike those of bats are low amplitude, broadband, multi-harmonic and frequency
modulated.[21] They contain no ‘echolocation clicks’ with reverberations and would seem to be used for
simple, close range spatial orientation. In contrast to bats, shrews use echolocation only to investigate their
habitat rather than additionally to pinpoint food.[21]

[edit] See also

• Human echolocation

[edit] References
1. ^ Yoon, Carol Kaesuk. "Donald R. Griffin, 88, Dies; Argued Animals Can Think", The New
York Times, November 14, 2003. Accessed July 16, 2010.
2. ^ D. R. Griffin (1958). Listening in the dark. Yale Univ. Press, New York.
3. ^ S. Dijkgraaf (1949). Spallanzani und die Fledermäuse. Experientia 5:90-92.
4. ^ Schevill, W.E. and McBride, A.F. 1956. Evidence for echolocation by cetaceans. Deep Sea
Research 3:153-154.
5. ^ Jones G. (2005). "Echolocation". Current Biology 15 (13): 484–488.
doi:10.1016/j.cub.2005.06.051. PMID 16005275.
6. ^ Muller, R. (2004). "A numerical study of the role of the tragus in the big brown bat.". JASA
116: 3701–3712. doi:10.1121/1.1815133
7. ^ Teeling et al. 2000. Molecular evidence regarding the origin of echolocation and flight in
bats. Nature 403: 188–192.
8. ^ "Order Chiroptera (Bats)". Animal Diversity Web. Retrieved 2007-12-
9. ^ Springer et al. 2001. Integrated fossil and molecular data reconstruct bat echolocation.
Proceedings of the National Academy of Sciences 98: 6241–6246.
10.^ Fordyce, R.E., and L.G. Barnes, (1994). "The evolutionary history of whales and
dolphins."Annual Review of Earth and Planetary Sciences 22:419-455.
11.^ Lindberg, D.R., and N.D. Pyenson, (2007). "Things that go bump in the night: evolutionary
interactions between cephalopods and cetaceans in the tertiary." Lethaia 40:335-343.
12.^ McGowen, M.R., M. Spaulding, and J. Gatesy, (2009). "Divergence date estimation and a
comprehensive molecular tree of extant cetaceans." Molecular Phylogenetics and Evolution 53:891-
13.^ Morisaka, T., and R.C. Connor, (2007). "Predation by killer whales ( Orcinus orca) and the
evolution of whistle loss and narrow-band high frequency clicks in odontocetes." Journal of
Evolutionary Biology 20:1439-1458.
14.^ a b Kyhn, L.A., F.H. Jensen, K. Beedholm, J. Tougaard, M. Hansen, and P.T. Madsen,
(2010). "Echolocation in sympatric Peale's dolphins (Lagenorhynchus australis) and Commerson's
dolphins (Cephalorhynchus commersonii) producing narrow-band high-frequency clicks." Journal of
Experimental Biology 213:1940-1949.
15.^ Cranford, T.W., (2000). "In Search of Impulse Sound Sources in Odontocetes." In Hearing
by Whales and Dolphins (Springer Handbook of Auditory Research series), W.W.L. Au, A.N. Popper
and R.R. Fay, Eds. Springer-Verlag, New York.
16.^ Goodson, A.D., and Klinowska, M.A., (1990). "A proposed echolocation receptor for the
Bottlenose Dolphin (Tursiops truncatus): modeling the receive directivity from tooth and lower jaw
geometry." In Sensory Abilities of Cetaceans vol 196 ed J A Thomas and R A Kastelein (New York:
Plenum) pp 255–67 (NATO ASI Series A)
17.^ Dobbins, P. (2007). "Dolphin sonar—modeling a new receiver concept." Bioinspired
Biomimicry 2 (2007) 19–29
18.^ Jon Fjeldså and Niels Krabbe (1990). Birds of the High Andes: a manual to the birds of the
temperate zone of the Andes and Patagonia, South America . Apollo Books. p. 232.
ISBN 9788788757163.
19.^ Marshall Cavendish Corp. (2000). Exploring Life Science. Marshall Cavendish. p. 547.
ISBN 9780761471424.
20.^ Thomas E. Tomasi, "Echolocation by the Short-Tailed Shrew Blarina brevicauda", Journal
of Mammalogy, Vol. 60, No. 4 (Nov., 1979), pp. 751–759.
21.^ a b c Siemers BM, Schauermann G, Turni H, von Merten S. (2009). Why do shrews twitter?
Communication or simple echo-based orientation. Biol Lett. 5(5):593-6. doi:10.1098/rsbl.2009.0378
PMID 19535367
• Hiryu, S. et al. 2007. Echo-intensity compensation in echolocating bats ( Pipistrellus
abramus)during flight measured by a telemetry microphone. J. Acoust. Soc. Am. 121(3): .
• Schnitzler, H.U. and Flieger, E. 1983. Detection of oscillating target movements by
echolocation in the Greater Horseshoe bat. J. Comp. Physiology. 153: 385-391.
• Zupanc, G.K.H. 2004. Behavioral Neurobiology: An Integrative Approach. Oxford University
Press: Oxford, UK.
• Simmons, J.A. and Stein, R.A. 1980. Acoustic Imaging in bat sonar: echolocation signals and
the evolution of echolocation. J. Comp. Physiol. A. 135: 61-84.
• Neuweiler, G. 2003. Evolutionary aspects of bat echolocation. J. Comp. Physiol. A. 189: 245-
• Jones, G. and Teeling, E.C. 2006. The evolution of echolocation in bats. Trends in Ecology
and Evolution. 21(3): 149-156.
• Fenton, M.B. 1995. Natural History and Biosonar Signals. In: Hearing in Bats. Popper, A.N.
and Fay, R.R. (eds.). Springer Verlag. New York. pp. 37–86.
• Grinnell, Alan D. (1995). Hearing in Bats: An Overview. In: Hearing in Bats. Popper, A.N. and
Fay, R.R. (eds.). Springer Verlag. New York. pp. 1–36.
• Reynolds JE III & Rommel SA (1999), Biology of Marine Mammals, Smithsonian Institution
Press, ISBN . Authoritative work on marine mammals with in depth sections on marine mammal
acoustics written by eminent experts in the field.
• Au, Whitlow W. L. (1993). The Sonar of Dolphins. New York: Springer-Verlag. Provides a
variety of findings on signal strength, directionality, discrimination, biology and more.
• Pack, Adam A. & Herman, Louis M. (1995). "Sensory integration in the bottlenosed dolphin:
Immediate recognition of complex shapes across the senses of echolocation and vision", J.
Acoustical Society of America, 98(2), 722-733. Shows evidence for the sensory integration of shape
information between echolocation and vision, and presents the hypothesis of the existence of the
mental representation of an "echoic image".
• Anderson, J.A. (1995) An Introduction to Neural Networks. MIT Press.
• Carew, T. (2001). Behavioral Neurobiology. Sinauer Associates, Inc., USA.
• Hopkins, C. (2007). Echolocation II. BioNB 424 Neuroethology Powerpoint presentation.
Cornell University, Ithaca NY.
• Ketten. D.R. (1992) The Marine mammal ear: Specializations for aquatic audition and
echolocation. In: The Evolutionary Biology of Hearing, D. Webster, R. Fay, and A. Popper (eds.),
Springer-Verlag, pp. 717–750.
• Ketten, D.R. (2000) Cetacean Ears. In: Hearing by Whales and Dolphins. W. Au, R. Fay, and
A. Popper (eds.), SHAR Series for Auditory Research, Springer-Verlag, pp. 43–108.
• Wilson, W. & C. Moss (2004) in Echolocation in Bats and Dolphins, J. Thomas, C. Moss, eds.
pp. 22-.
• Moss C. & Sinha S. (2003). Neurobiology of Echolocation in Bats. Current Opinion in
Neurobiology, 13(6), 751-758.
• Pollak G. et al. (1977). Echo-detecting characteristics of neurons in inferior colliculus of
unanesthetized bats. Science, 196: 675-678.
• Schuller, G. & Pollack, G. (1979). Disproportionate frequency representation in the inferior
colliculus of Doppler-compensating greater horseshoe bats: Evidence of an acoustic fovea. Journal
of Comparative Physiology A. 132: 47-54.
• Speakman J., Racey, P. No cost of echolocation for bats in flight. Nature, 350: 421-423
• Suga, N., Niwa H., Taniguchi I., Margoliash D. (1987). The personalized auditory cortex of
the mustached bat: adaptation for echolocation. Journal of Neurophysiology, 58: 643-654.
• Suga N., O'Neill W.E. (1979). Neural axis representing target range in the auditory cortex of
the mustache bat. Science, 206: 351-353.
• Suga N, Simmons JA and Jen PH. (1975) Peripheral specialization for fine analysis of
doppler-shifted echoes in the auditory system of the "CF-FM" bat Pteronotus parnellii. Journal of
Experimental Biology, 63: 161-192.
[edit] External links
• The British Library Sound Archive 150,000 recordings of over 10,000 species, including
many echolocation recordings
• International Bioacoustics Council links to many bioacoustics resources
• Listen to Nature has bat and swiftlet sonar signals
• Bat Ecology & Bioacoustics Lab
• Keep your ears peeled article on

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Mustang (horse)
From Wikipedia, the free encyclopedia

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This article is about the horse. For other uses, see Mustang (disambiguation).
Mustang adopted from the BLM

Distinguishing features: Small, compact, good bone, very hardy

Country of origin: North America

Horse (Equus ferus caballus)

A Mustang is a free-roaming feral horse of the North American west that first descended from horses
brought to the Americas by the Spanish. Mustangs are often referred to as wild horses, but the more correct
term is feral horses.
In 1971, the United States Congress recognized Mustangs as “living symbols of the historic and
pioneer spirit of the West, which continue to contribute to the diversity of life forms within the Nation and
enrich the lives of the American people.” Today, Mustang herds vary in the degree to which they can be
traced to original Iberian horses. Some contain a greater genetic mixture of ranch stock and more recent
breed releases, while others are relatively unchanged from the original Iberian stock, most strongly
represented in the most isolated populations.
Today, the Mustang population is managed and protected by the Bureau of Land Management.
Controversy surrounds the sharing of land and resources by the free ranging Mustangs with the livestock of
the ranching industry, and also with the methods with which the federal government manages the wild
population numbers. Many methods of population management are used, including the adoption by private
individuals of horses taken from the range.
• 1 Etymology and usage
• 2 History
• 3 Ancestry
• 4 Mustangs today
• 5 Land use controversies
• 6 Management and adoption
• 7 See also
• 8 Notes
• 9 References
• 10 Further reading

[edit] Etymology and usage

Mustangs are often referred to as wild horses but, since all free-roaming horses now in the Americas
descended from horses that were once domesticated, the more correct term is feral horses.[1]
The English word "mustang" comes from the Mexican Spanish word mestengo, derived from Spanish
mesteño, meaning "stray livestock animal".[2] The Spanish word in turn may possibly originate from the Latin
expression animalia mixta (mixed beasts), referring to beasts of uncertain ownership, which were distributed
in shepherd councils, known as mestas in medieval Spain.[3] A mestengo was any animal distributed in
those councils, and by extension any feral animal.

[edit] History

Free-roaming Mustangs
Today, the only true wild horse is the Przewalski's Horse, native to Mongolia. However, the horse
family Equidae and the genus Equus evolved in North America. Studies using ancient DNA as well as DNA
of recent individuals shows there once were two closely related horse species in North America, the "wild
horse" (Equus ferus) and the "Stilt-legged Horse;" which is taxonomically assigned to various names.[4][5]
Thus, primitive horses lived in North America in prehistoric times. However, the entire equus genus died out
at the end of the last ice age around 10-12,000 years ago, possibly due to a changing climate or the impact
of newly arrived human hunters.[6] Thus at the beginning of the Columbian Exchange, there were no equids
in the Americas at all. Horses first returned with the Conquistadors, beginning with Columbus, who imported
horses from Spain to the West Indies on his second voyage in 1493.[7] Domesticated horses came to the
mainland with the arrival of Cortés in 1519.[8]
The first Mustangs descended from Iberian horses[9] brought to Mexico and Florida. Most of these
horses were of Andalusian, Arabian and Barb ancestry. Some of these horses escaped or were stolen by
Native Americans, and rapidly spread throughout western North America.
Native Americans quickly adopted the horse as a primary means of transportation. Horses replaced
the dog as a travois puller and greatly improved success in battles, trade, and hunts, particularly bison hunts.
Starting in the colonial era and continuing with the westward expansion of the 1800s, horses
belonging to explorers, traders and settlers that escaped or were purposely released joined the gene pool of
Spanish-descended herds. It was also common practice for western ranchers to release their horses to
locate forage for themselves in the winter and then recapture them, as well as any additional Mustangs, in
the spring. Some ranchers also attempted to "improve" wild herds by shooting the dominant stallions and
replacing them with pedigreed animals.
By 1900 North America had an estimated two million free-roaming horses.[9] Since 1900, the
Mustang population has been reduced drastically. Mustangs were viewed as a resource that could be
captured and used or sold (especially for military use) or slaughtered for food, especially pet food. The
controversial practice of mustanging was dramatized in the John Huston film The Misfits, and the abuses
linked to certain capture methods, including hunting from airplanes and poisoning, led to the first federal wild
free-roaming horse protection law in 1959.[11] This statute, known as the "Wild Horse Annie Act," prohibited
the use of motor vehicles for hunting wild horses and burros.[12] Protection was increased further by the Wild
Free-Roaming Horse and Burro Act of 1971.[13]
The 1971 Act provided for protection of certain previously established herds of horses and burros.
Today, the Bureau of Land Management is the primary authority that oversees the protection and
management of Mustang herds on public lands,[14] while the United States Forest Service administers
additional wild horse or burro territories.[15]

[edit] Ancestry

Mustangs in Utah
Historically, many of the Native American tribes bred their horses carefully to improve them for their
purposes. Among the most capable horse-breeding people of North America were the Comanche, the
Shoshoni, and the Nez Perce. The last in particular became master horse breeders, and developed one of
the first truly American breeds: the Appaloosa. Most other tribes did not practice extensive amounts of
selective breeding, though they sought out desirable horses through capture, trade and theft, and quickly
traded away or otherwise eliminated those with undesirable traits.[ citation needed]
In some modern mustang herds there is clear evidence of other domesticated horse breeds having
become intermixed with feral herds. Some herds show the signs of the introduction of Thoroughbred or other
light racehorse-types into herds, a process that also led in part to the creation of the American Quarter Horse.
[citation needed] Other herds show signs of the intermixing of heavy draft horse breeds turned loose in an
attempt to create work horses. Other, more isolated herds, retain a strong influence of original Spanish stock.
Some breeders of domestic horses consider the Mustang herds of the west to be inbred and of
inferior quality.[citation needed] However, supporters of the Mustang argue that the animals are merely small
due to their harsh living conditions and that natural selection has eliminated many traits that lead to
weakness or inferiority. Some mustang supporters also maintain that some "inbreeding" actually
concentrates the traits of hardiness and durability, making the mustang a valuable genetic resource.[ citation
needed] Regardless of these debates, the Mustang of the modern west has several different breeding
populations today which are genetically isolated from one another and thus have distinct traits traceable to
particular herds. These herds vary in the degree to which they can be traced to original Iberian horses. Some
contain a greater genetic mixture of ranch stock and more recent breed releases, others are relatively
unchanged from the original Iberian stock.[citation needed]
Two researchers have advanced an argument that Mustangs should be legally classified as "wild"
rather than "feral." They argue that, due to the presence of Equus ferus ferus on the North American
continent till the end of the Pleistocene era, horses were once native animals and should still be considered
as native animals, and therefore defined as "wild,"[16] and not viewed as an exotic species that draws
resources and attention away from true native species.[17]
[edit] Mustangs today

Nevada's State Quarter, featuring the Mustang

Mustang in NW Nevada mountains
Today, free-roaming horses are protected under United States law, but have disappeared from
several states where there were once established populations.[18] A few hundred free-roaming horses
survive in Alberta and British Columbia. The BLM considers roughly 26,000 individuals a manageable
number,[19] but the feral Mustang population in February 2010 was 33,700 horses and 4,700 burros.[19]
More than half of all Mustangs in North America are found in Nevada (which features the horses on its State
Quarter in commemoration of this), with other significant populations in Montana, Wyoming and Oregon.[18]
Another 34,000 horses are in holding facilities.[19]
[edit] Land use controversies

Mustangs at a water source

Controversy surrounds the presence of feral Mustang herds. Supporters argue that Mustangs are
part of the natural heritage of the American West, whose history predates modern land use practices, and
thus the animals have an inherent right of inhabitation.[20] However, other people remain vehemently
opposed to their presence, arguing that the animals degrade rangeland and compete with livestock and wild
species for forage.[21]
The debate as to what degree Mustangs and cattle compete for forage is multifaceted. One group of
opponents, primarily cattle and sheep ranchers and those who depend on the livestock industry, argue
essentially that feral horses degrade rangeland and compete with private livestock for public land forage.[22]
The environmentalist community is split over the position of the Mustang within the North American
ecosystem. This debate centers on the potential classification of Mustangs as either an introduced species
such as cattle, or as a reintroduced native species due to the prehistoric presence of horses in North
America, albeit with a gap of thousands of years between their extinction and reintroduction from European
Researchers note that most current Mustang herds live in arid areas which cattle cannot fully utilize
due to the lack of water sources. Horses are better adapted by evolution to such climates.[24]; they may
range nine times as far from water sources as cattle, traveling as much as 50 miles a day.[25] This allows
them to utilize areas not grazed by cattle. In addition, horses are "hindgut fermenters," meaning that they
digest nutrients by means of the cecum rather than by a multi-chambered stomach. In practical effect, horses
can obtain adequate nutrition from poorer forage than can cattle, surviving in areas where cattle will starve.

[edit] Management and adoption

Mustangs at the Palomino Valley Adoption Center
The Bureau of Land Management (BLM) is tasked with protecting, managing, and controlling wild
horses and burros under the authority of the 1971 Wild Free-Roaming Horses and Burros Act to ensure that
healthy herds thrive on healthy rangelands and as multiple-use mission under the 1976 Federal Land Policy
and Management Act. Under the 1971 Act, shooting or poisoning Mustangs in the wild is illegal, and doing so
can be prosecuted as a criminal felony.
Healthy adult Mustangs have few natural predators aside from mountain lions,[26] and to a lesser
extent, grizzly bears and wolves.[27] The mountain lion is well-known for predation on feral horses, and the
larger members of the species may hunt both horses and moose. They are very effective predators that kill
by either leaping onto an animal or chasing it down in a sprint, then grabbing the prey with their front claws
and biting the neck, either at the windpipe or the spine.[28]
Where there is natural balance of predators and prey, Mustang numbers tend to stay in balance.
However, in many areas, natural predators have been eliminated from the ecosystem.[27] Without some form
of population control, Mustang herd sizes can multiply rapidly, doubling as fast as every four years.[12] To
maintain population balance, (or, some argue, to make room for cattle[29]) one of the BLM’s key
responsibilities under the 1971 law is to determine an appropriate management level (AML) of wild horses
and burros in areas of public rangelands dedicated specifically for them.
Control of the population to within AML is achieved through a capture program. There are strict
guidelines for techniques used to round up Mustangs. One method uses a tamed horse, called a "Judas
horse," which has been trained to lead wild horses into a pen or corral. Once the Mustangs are herded into
an area near the holding pen, the Judas horse is released. Its job is then to move to the head of the herd and
lead them into a confined area. Already confused and looking for direction, the wild horses instinctively follow
this animal.[citation needed]
Most horses that are captured are offered for adoption to individuals or groups willing and able to
provide humane, long-term care after payment of an adoption fee that ranges from $25 to $125.[30] In order
to prevent the later sale of mustangs as horse meat, adopted mustangs are still protected under the Act, and
cannot be sold in the first year except when certain very specific criteria are met. As of 2010, nearly 225,000
Mustangs have been adopted.[19]
Because there is a much larger pool of captured horses than of prospective adoptive owners, a
number of efforts have been made to reduce the number of horses in holding facilities. At present, there are
about 34,000 Mustangs in holding facilities and long-term grassland pastures.[19] The BLM has publicly
considered euthanasia as a possible solution to overpopulation.[31] In January 2005, a controversial
amendment was attached to an appropriation bill before the United States Congress by former Senator
Conrad Burns, dubbed the "Burns rider."[32] This modified the adoption program to allow the sale (with the
result usually being slaughter) of captured horses that are "more than 10 years of age", or that were "offered
unsuccessfully for adoption at least three times."[33] In 2009, Secretary of the Interior Ken Salazar proposed
the creation of federal wild horse preserves in the midwest, where non-reproducing animals would be kept.
[34] Another approach to placing excess animals has been advanced by Madeleine Pickens, wife of oil
magnate T. Boone Pickens, who seeks to create a private sanctuary in northern Nevada.[34] There are also
increased efforts to assist with finding appropriate adoption homes. One example is a promotional
competition that gives trainers 100 days to gentle and train 100 mustangs, which are then adopted through
an auction.[35]
[edit] See also
Wikimedia Commons has media related to: Feral horses from America

• Colonial Spanish Horse

• Feral horse
• Kiger Mustang
• Kleppe v. New Mexico
• Spanish Mustang
• Wild horse, including
• Przewalski's horse
• Tarpan

[edit] Notes
1. ^ The American Museum of Natural History The Nature of Horses
2. ^ "Mustang". Retrieved 2009-03-31.
3. ^ Websters Third International Dictionary Unabridged
4. ^ Weinstock, J.; et al. (2005). "Evolution, systematics, and phylogeography of Pleistocene
horses in the New World: a molecular perspective". PLoS Biology 3 (8): e241.
doi:10.1371/journal.pbio.0030241. PMID 15974804. PMC 1159165.
document&doi=10.1371%2Fjournal.pbio.0030241&ct=1. Retrieved 2008-12-19.
5. ^ Orlando, L.; et al. (2008). "Ancient DNA Clarifies the Evolutionary History of American Late
Pleistocene Equids". Journal of Molecular Evolution 66 (5): 533–538. doi:10.1007/s00239-008-9100-
x. PMID 18398561.
6. ^ "Ice Age Horses May Have Been Killed Off by Humans" National Geographic News, May 1,
7. ^ Bennett, Deb. Conquerors: The Roots of New World Horsemanship. Amigo Publications,
1998, pp. 150 ISBN 0-9658533-0-6
8. ^ Henry, Marguerite and Wesley Dennis. All About Horses. Random House, 1962. ASIN:
9. ^ a b Dobie, The Mustangs
10.^ "Seeds of Change." Corpus Christi Museum, Science and History educational resources.
Accessed June 1, 2007 at
11.^ "Wild Horse Annie Act". Retrieved 2010-08-09.
12.^ a b Mangum, "The Mustang Dilemma," p. 77
13.^ ""Background Information on HR297"" (PDF). Retrieved 2010-08-09.
14.^ Mangum, "The Mustang Dilemma," p. 75
15.^ "Wild Horse and Burro Territories". Retrieved 2009-01-
16.^ "The Surprising History of America's Wild Horses". 2008-07-24. Retrieved 2010-08-09.
17.^ "Wild Horses". Retrieved 2010-08-09.
18.^ a b "National Summary, FY2007" (PDF). Retrieved 2010-08-09.
19.^ a b c d e Mangum, "The Mustang Dilemma", p.77
20.^ The American Wild Horse Preservation Campaign
21.^ see, e.g. National Academy of Sciences Report, 1982
General Accounting Office Report, 1991
22.^ "Bellisle, Martha. "Legislative battle brews over Nevada's wild horses" ''Associated press''
reprinted at I.G.H.A. / HorseAid's Bureau of Land Management News. Web site accessed May 11,
2007". Retrieved 2010-08-09.
23.^ "Wild Horses as Native North American Wildlife" Jay F. Kirkpatrick, Ph.D. and Patricia M.
24.^ "Wild Horses and the Ecosystem". Retrieved 2010-08-09.
25.^ a b Budiansky, Stephen. The Nature of Horses. Free Press, 1997. ISBN 0-684-82768-9
26.^ John W. Turner, Jr. and Michael L. Morrison (2008). "Influence of Predation by Mountain
Lions on Numbers and Survivorship of a Feral Horse Population". The Southwestern Naturalist Vol.
46, No.2 pp. 183-190. Retrieved 2008-08-29.
27.^ a b "FAQ'S On BLM Wild Horses" Wild Mustang Coalition. Web site accessed December 9,
28.^ French, Brett. "Ferocious appetites: Study finds mountain lions may be eating more than
previously believed." Billings Gazette. December 9, 2010. Accessed December 9, 2010.
29.^ "On BLM's wild horse management practices". Retrieved 2010-08-09.
30.^ "Adoption". New Mexican Mustang & Burro Association. Retrieved 2010-12-2.
31.^ ""Plan to kill wild horses runs into trouble" ''Associated Press,'' July 7, 2008" (PDF). Retrieved 2010-08-09.
32.^ "The Story Behind the Burns Amendment". Retrieved 2010-08-09.
33.^ "Burns amendment". 2004-12-06. Retrieved 2010-08-09.
34.^ a b Mangum, "The Mustang Dilemma", p.78
35.^ The Extreme Mustang Makeover

[edit] References
• Dobie, Frank (1952). The Mustangs (Seventh printing ed.). Boston: Little, Brown and
Company. LC no. 52-6802.
• Mangum, A.J. (December 2010). "The Mustang Dilemma". Western Horseman: 70–80.

[edit] Further reading

• Nimmo, D. G., & Miller, K. K. (2007) Ecological and human dimensions of management of
feral horses in Australia: A review. Wildlife Research, 34, 408-417
• Text of Wild Free-Roaming Horse and Burro Act of 1971
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Temporal range: Late Cambrian–Recent[1]
Air-breathing land gastropod Helix pomatia, the Roman snail

Scientific classification [ e ]

Kingdom: Animalia

Phylum: Mollusca

Cuvier, 1795[2]


"Paleozoic uncertain …"

"Basal taxa …"
clade Patellogastropoda
clade Vetigastropoda
clade Cocculiniformia
clade Neritimorpha
clade Caenogastropoda
clade Heterobranchia
The Gastropoda or gastropods are a large taxonomic class within the molluscs, a group of animals
that are more commonly known as snails and slugs. The class includes snails and slugs of all kinds and all
sizes: huge numbers of marine snails and sea slugs, as well as freshwater snails and freshwater limpets, and
the terrestrial (land) snails and slugs. The class Gastropoda contains a vast total of named species, second
only to the insects in overall number. The fossil history of this class goes all the way back to the Late
Cambrian. There are 611 families of gastropods, of which 202 families are extinct, being found only in the
fossil record.[3]
Gastropoda (previously known as univalves and sometimes spelled Gasteropoda) are a major part of
the phylum Mollusca and are the most highly diversified class in the phylum, with 60,000 to 80,000[3][4] living
snail and slug species. The anatomy, behavior, feeding and reproductive adaptations of gastropods vary very
significantly from one clade or group to another. Therefore, it is difficult or impossible to make more than a
few general statements that are valid for all gastropods.
The class Gastropoda has an extraordinary diversification of habitats. Representatives live in
gardens, in woodland, in deserts, and on mountains; in small ditches, great rivers and lakes; in estuaries,
mudflats, the rocky intertidal, the sandy subtidal, in the abyssal depths of the oceans including the
hydrothermal vents, and numerous other ecological niches, including parasitic ones.
Although the name "snail" can be, and often is, applied to all the members of this class, commonly
this word means only those species with an external shell large enough that the soft parts can withdraw
completely into it. Those gastropods without a shell, and those with only a very reduced or internal shell, are
usually known as slugs.
The marine shelled species of gastropod include edible species such as abalone, conches,
periwinkles, whelks, and numerous other sea snails that produce seashells which are coiled in the adult
stage, even though in some cases the coiling may not be very visible, for example in cowries. There are also
a number of families of species such as all the various limpets, where the shell is coiled only in the larval
stage, and is a simple conical structure after that.
• 1 Etymology
• 2 Diversity
• 3 Habitat
• 4 Anatomy
• 4.1 The shell
• 4.2 Body wall
• 4.3 Digestive system
• 4.4 Respiratory system
• 4.5 Circulatory system
• 4.6 Excretory system
• 4.7 Sensory organs and nervous system
• 4.8 Reproductive system
• 5 Life cycle
• 6 Feeding behavior
• 7 Genetics
• 8 Geological history
• 9 Taxonomy
• 10 References
• 11 External links
[edit] Etymology
Look up gastropod or univalve in Wiktionary, the free dictionary.

The word "gastropod" is derived from the Ancient Greek words γαστήρ (gastér, stem: gastr-)
"stomach", and πούς (poús, stem: pod-) "foot", hence stomach-foot. This is an anthropomorphic name, based
on the fact that to humans it appears as if snails and slugs crawl on their bellies. In reality, snails and slugs
have their stomach, the rest of their digestive system and all the rest of their viscera in a hump on the
opposite, dorsal side of the body. In most gastropods this visceral hump is covered by, and contained within,
the shell.
In the scientific literature, gastropods were described under the vernacular (French) name
"gasteropodes" by Georges Cuvier in 1795.[2] The name was later Latinized.
The earlier name univalve means "one valve", and contrasts with bivalve ("two valves").

[edit] Diversity
At all taxonomic levels, gastropods are second only to the insects in terms of their diversity.[5]
Gastropods form the class of molluscs with the greatest numbers. However the estimated total
number of gastropod species varies widely, depending on the cited sources. The number of gastropod
species can be deduced from estimates of the number of described species of Mollusca with accepted
names: about 85,000[6] (minimum 50,000,[6] maximum 120,000[6]). But an estimate of the total number of
Mollusca, including undescribed species, is about 200,000 species.[6] The estimate of 85,000 molluscs
includes 24,000 described species of terrestrial gastropods.[6]
Different estimations (from different sources) for aquatic gastropods give about 30,000 species[7] of
marine gastropods and about 5,000 species of freshwater and brackish gastropods.[7] Total number of
recent species of freshwater snails is about 4,000.[8]
The number of prehistoric (fossil) species of gastropods is at least 15,000 species.[9]

[edit] Habitat
Some of the more familiar and better-known gastropods are terrestrial (the land snails and slugs) and
some live in freshwater, but more than two thirds of all named species live in a marine environment.
Gastropods have a worldwide distribution from the near Arctic and Antarctic zones to the tropics.
They have become adapted to almost every kind of existence on earth, having colonized every medium
available except the air.
In habitats where there is not enough calcium carbonate to build a really solid shell, such as on some
acidic soils on land, there are still various species of slugs, and also some snails with a thin translucent shell,
mostly or entirely composed of the protein conchiolin.
Snails such as Sphincterochila boissieri and Xerocrassa seetzeni have adapted to desert conditions,
other snails have adapted to an existence in ditches, near deepwater hydrothermal vents, the pounding surf
of rocky shores, caves, and many other diverse areas.
[edit] Anatomy
The anatomy of a common air-breathing land snail such as Helix aspersa. Note that much of this
anatomy does not apply to gastropods in other clades or groups.
The anatomy of an aquatic snail with a gill, a male prosobranch gastropod. Note that much of this
anatomy does not apply to gastropods in other clades.
Light yellow - body
Brown - shell and operculum
Green - digestive system
Light purple - gills
Yellow - osphradium
Red - heart
Pink -
Dark violet -
1. foot
2. cerebral ganglion
3. pneumostome
4. upper commissura
5. osphradium
6. gills
7. pleural ganglion
8. atrium of heart
9. visceral ganglion
10. ventricle
11. foot
12. operculum
13. brain
14. mouth
15. tentacle (chemosensory, 2 or 4)
16. eye
17. penis (everted, normally internal)
18. esophageal nerve ring
19. pedal ganglion
20. lower commissura
21. vas deferens
22. pallial cavity / mantle cavity / respiratory cavity
23. parietal ganglion
24. anus
25. hepatopancreas
26. gonad
27. rectum
28. nephridium
Snails are distinguished by an anatomical process known as torsion, where the visceral mass of the
animal rotates 180° to one side during development, such that the anus is situated more or less above the
head. This process is unrelated to the coiling of the shell, which is a separate phenomenon. Torsion is
present in all gastropods, but the opisthobranch gastropods are secondarily de-torted to various degrees.[10]
Torsion occurs in two mechanistic stages. The first is muscular, and the second is mutagenetic. The
effects of torsion are primarily physiological - the organism develops an asymmetrical nature with the majority
of growth occurring on the left side. This leads to the loss of right-paired appendages (e.g. ctenidia (comb-
like respiratory apparatus), gonads, nephridia, etc.). Furthermore, the anus becomes redirected to the same
space as the head. This is speculated to have some evolutionary function, as prior to torsion, when retracting
into the shell, first the posterior end would get pulled in, and then the anterior. Now, the front can be retracted
more easily, perhaps suggesting a defensive purpose.
However, this "rotation hypothesis" is being challenged by the "asymmetry hypothesis" in which the
gastropod mantle cavity originated from one side only of a bilateral set of mantle cavities.[12]
Gastropods typically have a well-defined head with two or four sensory tentacles with eyes, and a
ventral foot, which gives them their name (Greek gaster, stomach, and poda, feet). The foremost division of
the foot is called the propodium. Its function is to push away sediment as the snail crawls. The larval shell of
a gastropod is called a protoconch.

[edit] The shell

Main article: Gastropod shell
The shell of Zonitoides nitidus, a small land snail, has dextral coiling, which is typical (but not
universal) in gastropod shells.
Upper image: dorsal view of the shell, showing the apex
Central image: lateral view showing the spire and aperture of the shell
Lower image: basal view showing the umbilicus
Most shelled gastropods have a one piece shell, typically coiled or spiraled. This coiled shell usually
opens on the right-hand side (as viewed with the shell apex pointing upward). Numerous species have an
operculum, which in many species acts as a trapdoor to close the shell. This is usually made of a horn-like
material, but in some molluscs it is calcareous. In the land slugs, the shell is reduced, internal, or absent, and
the body is streamlined.

[edit] Body wall

Some sea slugs are very brightly colored. This serves either as a warning, when they are poisonous
or contain stinging cells, or to camouflage them on the brightly-colored hydroids, sponges and seaweeds on
which many of the species are found.
Lateral outgrowths on the body of nudibranchs are called cerata. These contain a part of digestive
gland, which is called the diverticula.

[edit] Digestive system

Main articles: Digestive system of gastropods and Radula
The radula of a gastropod is usually adapted to the food that a species eats. The simplest gastropods
are the limpets and abalones, herbivores that use their hard radula to rasp at seaweeds on rocks.
Many marine gastropods are burrowers, and have a siphon that extends out from the mantle edge.
Sometimes the shell has a siphonal canal to accommodate this structure. A siphon enables the animal to
draw water into their mantle cavity and over the gill. They use the siphon primarily to "taste" the water to
detect prey from a distance. Gastropods with siphons tend to be either predators or scavengers.

[edit] Respiratory system

Main articles: Respiratory system of gastropods and Respiratory system#Anatomy of respiratory
system in invertebrates
Almost all marine gastropods breathe with a gill, but many freshwater species, and the majority of
terrestrial species, have a pallial lung. Gastropods with a lung belong to one group with common descent, the
Pulmonata, however, gastropods with gills are paraphyletic. The respiratory protein in almost all gastropods
is hemocyanin, but a pulmonate family Planorbidae have hemoglobin as respiratory protein.
In one large group of sea slugs, the gills are arranged as a rosette of feathery plumes on their backs,
which gives rise to their other name, nudibranchs. Some nudibranchs have smooth or warty backs and have
no visible gill mechanism, such that respiration may likely take place directly through the skin.

[edit] Circulatory system

Main article: Circulatory system of gastropods
Gastropods have open circulatory system and the transport fluid is hemolymph. Hemocyanin is
present in the hemolymph as the respiratory pigment.
[edit] Excretory system
Main article: Excretory system of gastropods
The primary organs of excretion in gastropods are nephridia, which produce either ammonia or uric
acid as a waste product. The nephridium also plays an important role in maintaining water balance in
freshwater and terrestrial species. Additional organs of excretion, at least in some species, include
pericardial glands in the body cavity, and digestive glands opening into the stomach.

[edit] Sensory organs and nervous system

The upper pair of tentacles on the head of Helix pomatia have eye spots, but the main sensory
organs of the snail are sensory receptors for olfaction, situated in the epithelium of the tentacles.
Main articles: Sensory organs of gastropods and Nervous system of gastropods
Sensory organs of gastropods include olfactory organs, eyes, statocysts and mechanoreceptors.[13]
Gastropods have no hearing.[13]
In terrestrial gastropods (land snails and slugs), the olfactory organs, located on the tips of the 4
tentacles, are the most important sensory organ,[13] The chemosensory organs of opisthobranch marine
gastropods are called rhinophores.
The majority of gastropods have simple visual organs, eye spots, that are situated either at the tip of
the tentacles or the base of the tentacles. However "eyes" in gastropods range from these simple ocelli which
cannot process an image being only able to distinguish light and dark, to more complex pit eyes, and even to
lens eyes.[14] In land snails and slugs, vision is not the most important sense, because they are mainly
nocturnal animals.[13]
The nervous system of gastropods includes the peripheral nervous system and the central nervous
system. The central nervous system consist of ganglia connected by nerve cells. It includes paired ganglia:
the cerebral ganglia, pedal ganglia, osphradial ganglia, pleural ganglia, parietal ganglia and the visceral
ganglia. There are sometimes also buccal ganglia.[13]

[edit] Reproductive system

Main article: Reproductive system of gastropods
Courtship is a part of mating behavior in some gastropods including some of the Helicidae. Again, in
some land snails, an unusual feature of the reproductive system of gastropods is the presence and utilization
of love darts.
In many marine gastropods other than the opisthobranchs, there are separate sexes; most land
gastropods however are hermaphrodites.

[edit] Life cycle

A 9-hour-old trochophore of Haliotis asinina

sf - shell field
mating behaviour of Elysia timida
The main aspects of the life cycle of gastropods include:
• Egg laying and the eggs of gastropods
• The Embryonic development of gastropods
• The larvae or larval stadium: some gastropods may be trochophore and/or veliger
• Estivation and hibernation (each of these are present in some gastropods only)
• The growth of gastropods
• Courtship of gastropods and mating of gastropods: fertilisation is internal or external
according to the species. External fertilisation is common in marine gastropods.
[edit] Feeding behavior
Marine gastropods include some that are herbivores, detritus feeders, predatory carnivores,
scavengers, parasites, and also a few ciliary feeders, in which the radula is reduced or absent. In some
species that have evolved into endoparasites, such as Parenteroxenos doglieli, many of the standard
gastropod features are strongly reduced or absent.
A few sea slugs are herbivores and some are carnivores. Many have distinct dietary preferences and
regularly occur in close association with their food species.
Some predatory carnivorous gastropods include, for example: Cone shells, Testacella, Daudebardia,
Ghost slug and others.

[edit] Genetics
Gastropods exhibit an important degree of variation in mt gene organization when compared to other
animals.[15] Main events of gene rearrangement occurred at the origin of Patellogastropoda and
Heterobranchia, whereas fewer changes occurred between the ancestors of Vetigastropoda (only tRNAs D,
C and N) and Caenogastropoda (a large single inversion, and translocations of the tRNAs D and N).[15]
Within Heterobranchia, gene order seems to be relatively conserved and gene rearrangements are mostly
related with transposition of tRNA genes.[15]
[edit] Geological history

Fossil gastropod and attached mytilid bivalves on a Jurassic limestone bedding plane of the Matmor
Formation in southern Israel.
Helix aspersa: a European pulmonate land snail that has been accidentally introduced in many
countries throughout the world.
See also: :fr:Gastropoda (classification phylogénétique) and List of marine gastropod genera in the
fossil record
The first gastropods were exclusively marine, with the earliest representatives of the group appearing
in the Late Cambrian (Chippewaella, Strepsodiscus). Early Cambrian forms like Helcionella and Scenella are
no longer considered gastropods, and the tiny coiled Aldanella of earliest Cambrian time is probably not even
a mollusk. By the Ordovician period the gastropods were a varied group present in a range of aquatic
habitats. Commonly, fossil gastropods from the rocks of the early Palaeozoic era are too poorly preserved for
accurate identification. Still, the Silurian genus Poleumita contains fifteen identified species. Fossil
gastropods were less common during the Palaeozoic era than bivalves.
Most of the gastropods of the Palaeozoic era belong to primitive groups, a few of which still survive
today. By the Carboniferous period many of the shapes we see in living gastropods can be matched in the
fossil record, but despite these similarities in appearance the majority of these older forms are not directly
related to living forms. It was during the Mesozoic era that the ancestors of many of the living gastropods
One of the earliest known terrestrial (land-dwelling) gastropods is Maturipupa, which is found in the
Coal Measures of the Carboniferous period in Europe, but relatives of the modern land snails are rare before
the Cretaceous period, when the familiar Helix first appeared.

Cepaea nemoralis: another European pulmonate land snail, which has been introduced to many
other countries
In rocks of the Mesozoic era, gastropods are slightly more common as fossils, their shells are often
well preserved. Their fossils occur in ancient beds deposited in both freshwater and marine environments.
The "Purbeck Marble" of the Jurassic period and the "Sussex Marble" of the early Cretaceous period, which
both occur in southern England, are limestones containing the tightly packed remains of the pond snail
Rocks of the Cenozoic era yield very large numbers of gastropod fossils, many of these fossils being
closely related to modern living forms. The diversity of the gastropods increased markedly at the beginning of
this era, along with that of the bivalves.
Certain trail-like markings preserved in ancient sedimentary rocks are thought to have been made by
gastropods crawling over the soft mud and sand. Although these trails are of debatable origin, some of them
do resemble the trails made by living gastropods today.
Gastropod fossils may sometimes be confused with ammonites or other shelled cephalopods. An
example of this is Bellerophon from the limestones of the Carboniferous period in Europe, the shell of which
is planispirally coiled and can be mistaken for the shell of a cephalopod.
Gastropods are one of the groups that record the changes in fauna caused by the advance and
retreat of the Ice Sheets during the Pleistocene epoch.
[edit] Taxonomy

A group of fossil Turritella cingulifera from the Pliocene of Cyprus.

The taxonomy of the Gastropoda is under constant revision, and more and more of the old taxonomy
is being abandoned as the results of DNA studies slowly become clearer. Nevertheless a few of the older
terms such as "opisthobranch" and "prosobranch" are still sometimes used in a descriptive way.
The taxonomy of the Gastropoda as shown in various texts can differ in major ways, and on-going
revisions of the higher taxonomic levels are to be expected in the near future.
In the older classification there were four subclasses[16]:
• Opisthobranchia (gills to the right and behind the heart).
• Gymnomorpha (no shell)
• Prosobranchia (gills in front of the heart).
• Pulmonata (with a lung instead of gills)
Main article: Taxonomy of the Gastropoda (Ponder & Lindberg, 1997)
According to newer insights based on DNA sequencing, the taxonomy of the Gastropoda must be
rewritten in terms of strictly monophyletic groups. Integrating these findings into a working taxonomy will
continue to be a challenge in coming years. At present, it is impossible to give a classification of the
Gastropoda that has consistent ranks and also reflects current usage.
Convergent evolution, which appears to exist at especially high frequency in the Gastropoda class,
may account for the observed differences between the phylogenies, which are obtained from morphological
data and more recent gene sequences studies.
Main articles: Taxonomy of the Gastropoda (Bouchet & Rocroi, 2005) and Changes in the taxonomy
of gastropods since 2005
Bouchet & Rocroi (2005)[3][17] made changes in systematics, resulting in a taxonomy that is a step
closer to the evolutionary history of the phylum.
This classification system is based partly on the older systems of classification and partly on new
cladistic research. In the past, the taxonomy of gastropods was largely based on phenetic morphological
characters of the taxa. The recent advances are more based on molecular characters through research of
DNA[18] and RNA. This has made the taxonomical ranks and their hierarchy controversial. The debate about
these issues is not likely to end soon.
In this taxonomy, Bouchet, Rocroi et al. have used unranked clades for taxa above the rank of
superfamily (replacing the ranks suborder, order, superorder and subclass), while using the traditional
Linnaean approach for all taxa below the rank of superfamily. Whenever monophyly has not been tested, or
is known to be paraphyletic or polyphyletic, the term "group" or "informal group" has been used. The
classification of families into subfamilies is often not well resolved, and should be regarded as the best
possible hypothesis.
In 2004 Brian Simison and David R. Lindberg showed possible diphyletic origins of the Gastropoda
based on mitochondrial gene order and amino acid sequence analyses of complete genes.[19]

[edit] References
This article incorporates CC-BY-2.0 text from the reference.[15]
1. ^ 'Latest Early Cambrian', per Landing, E.; Geyer, G.; Bartowski, K. E. (March 2002). "Latest
Early Cambrian Small Shelly Fossils, Trilobites, and Hatch Hill Dysaerobic Interval on the Quebec
Continental Slope". Journal of Paleontology 76 (2): 287–305. doi:10.1666/0022-
3360(2002)076<0287:LECSSF>2.0.CO;2. edit; see section in article for first 'concrete evidence' in
Late Cambrian.
2. ^ a b (French) Cuvier G. (1795). "Second mémoire sur l'organisation et les rapports des
animaux à sang blanc, dans lequel on traite de la structure des Mollusques et de leur division en
ordres, lu à la Société d'histoire naturelle de Paris, le 11 Prairial, an III". Magazin Encyclopédique, ou
Journal des Sciences, des Lettres et des Arts 2: 433-449. page 448.
3. ^ a b c Bouchet P. & Rocroi J.-P. (Ed.); Frýda J., Hausdorf B., Ponder W., Valdes A. & Warén
A. 2005. Classification and nomenclator of gastropod families . Malacologia: International Journal of
Malacology, 47(1-2). ConchBooks: Hackenheim, Germany. ISBN 3-925919-72-4. 397 pp.
4. ^ Britannica online: abundance of the Gastropoda
5. ^ McArthur, A.G.; M.G. Harasewych (2003). "Molecular systematics of the major lineages of
the Gastropoda.". Molecular Systematics and Phylogeography of Mollusks . Washington: Smithsonian
Books. pp. 140–160.
6. ^ a b c d e Chapman, A.D. (2009). Numbers of Living Species in Australia and the World, 2nd
edition. Australian Biological Resources Study, Canberra. Accessed 12 January 2010. ISBN 978 0
642 56860 1 (printed); ISBN 978 0 642 56861 8 (online).
7. ^ a b "gastropod". (2010). In Encyclopædia Britannica. Retrieved March 05, 2010, from
Encyclopædia Britannica Online.
8. ^ Strong E. E., Gargominy O., Ponder W. F. & Bouchet P. (2008). "Global Diversity of
Gastropods (Gastropoda; Mollusca) in Freshwater". Hydrobiologia 595: 149-166. doi:10.1007/s10750-007-9012-6.
9. ^ (Spanish) Nájera J. M. (1996). "Moluscos del suelo como plagas agrícolas y
cuarentenarias". X Congreso Nacional Agronómico / II Congreso de Suelos 1996 51-56. PDF
10.^ Kay, A.; Wells, F. E.; Poder, W. F. (1998). "Class Gastropoda". In Beesley, P. L.; Ross, G.
J. B.; Wells, A.. Mollusca: The Southern Synthesis. Fauna of Australia. CSIRO Publishing. pp. 565–
604. ISBN 0 643 05756 0.
11.^ Brusca, R. C.; Brusca, G. J. (2003). "Phylum Mollusca". Invertebrates. Sinauer Associates,
Inc.. pp. 701–769. ISBN 0-87893-097-3.
12.^ Louise R. Page (2006). "Modern insights on gastropod development: Reevaluation of the
evolution of a novel body plan". Integrative and Comparative Biology 46 (2): 134–143.
13.^ a b c d e Chase R.: Sensory Organs and the Nervous System. in Barker G. M. (ed.): The
biology of terrestrial molluscs. CABI Publishing, Oxon, UK, 2001, ISBN 0-85199-318-4. 1-146, cited
pages: 179-211.
14.^ Götting, Klaus-Jürgen (1994). "Schnecken". In Becker, U., Ganter, S., Just, C. &
Sauermost, R.. Lexikon der Biologie. Heidelberg: Spektrum Akademischer Verlag. ISBN 3-86025-
15.^ a b c d Cunha R. L., Grande C. & Zardoya R. (23 August 2009). "Neogastropod phylogenetic
relationships based on entire mitochondrial genomes". BMC Evolutionary Biology 2009, 9: 210.
16.^ Paul Jeffery. Suprageneric classification of class Gastropoda. The Natural History
Museum, London, 2001.
17.^ Poppe G.T. & Tagaro S.P. 2006. The new classification of Gastropods according to
Bouchet & Rocroi, 2005. Visaya, février 2006: 10 pp. http://www.journal-
18.^ Elpidio A. Remigio and Paul D.N. Hebert (2003). "Testing the utility of partial COI
sequences for phylogenetic (full text on line)". Molecular Phylogenetics and Evolution 29 (3): 641–
647. doi:10.1016/S1055-7903(03)00140-4. PMID 14615199.
19.^ - Unitas malacologica, Newsletter number 21 december 2004 - a .pdf file

[edit] External links

Wikimedia Commons has media related to: Gastropoda
Gastropods portal

• Gastropod reproductive behavior

• Reconstructions of fossil gastropods
• 2004 Linnean taxonomy of gastropods
• Webster S. J. & Fiorito G. (October 2001) "Socially guided behaviour in non-insect
invertebrates". Animal Cognition 4(2): 69–79. doi: 10.1007/s100710100108 - An article about social
learning also in gastropods.
Retrieved from ""

Categories: Gastropods

From Wikipedia, the free encyclopedia

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"Archea" redirects here. For the geologic eon, see Archean. For the spider family, see Archaeidae.
Temporal range: Paleoarchean – Recent
Halobacteria sp. strain NRC-1, each cell about 5 μm long
Scientific classification

Woese, Kandler & Wheelis, 1990

Kingdoms and phyla

The Archaea (/ɑrˈkiːə/ ( listen) ar-KEE-ə) are a group of single-celled microorganisms. A single
individual or species from this domain is called an archaeon (sometimes spelled "archeon"). They have no
cell nucleus nor any other membrane-bound organelles within their cells. In the past they were viewed as an
unusual group of bacteria and named archaebacteria, but since the Archaea have an independent
evolutionary history and show many differences in their biochemistry from other forms of life, they are now
classified as a separate domain in the three-domain system. In this system the phylogenetically distinct
branches of evolutionary descent are the Archaea, Bacteria and Eukaryota. Archaea are divided into four
recognized phyla, but many more phyla may exist. Of these groups the Crenarchaeota and the
Euryarchaeota are most intensively studied. Classification is still difficult, since the vast majority have never
been studied in the laboratory and have only been detected by analysis of their nucleic acids in samples from
the environment. Although archaea have, in the past, been classed with bacteria as prokaryotes (or Kingdom
Monera), this classification is regarded by some as outdated.[1]
Archaea and bacteria are quite similar in size and shape, although a few archaea have very unusual
shapes, such as the flat and square-shaped cells of Haloquadratum walsbyi. Despite this visual similarity to
bacteria, archaea possess genes and several metabolic pathways that are more closely related to those of
eukaryotes: notably the enzymes involved in transcription and translation. Other aspects of archaean
biochemistry are unique, such as their reliance on ether lipids in their cell membranes. The archaea exploit a
much greater variety of sources of energy than eukaryotes: ranging from familiar organic compounds such as
sugars, to using ammonia, metal ions or even hydrogen gas as nutrients. Salt-tolerant archaea (the
Halobacteria) use sunlight as an energy source and other species of archaea fix carbon; however, unlike
plants and cyanobacteria, no species of archaea is known to do both. Archaea reproduce asexually and
divide by binary fission, fragmentation, or budding; in contrast to bacteria and eukaryotes, no known species
form spores.
Initially, archaea were seen as extremophiles that lived in harsh environments, such as hot springs
and salt lakes, but they have since been found in a broad range of habitats, including soils, oceans, and
marshlands. Archaea are particularly numerous in the oceans, and the archaea in plankton may be one of
the most abundant groups of organisms on the planet. Archaea are now recognized as a major part of Earth's
life and may play roles in both the carbon cycle and the nitrogen cycle. No clear examples of archaeal
pathogens or parasites are known, but they are often mutualists or commensals. One example is the
methanogens that inhabit the gut of humans and ruminants, where their vast numbers aid digestion.
Methanogens are used in biogas production and sewage treatment, and enzymes from extremophile archaea
that can endure high temperatures and organic solvents are exploited in biotechnology.
• 1 Classification
• 1.1 New domain
• 1.2 Current classification
• 1.2.1 Species
• 2 Origin and evolution
• 2.1 Archaea and eukaryotes
• 3 Morphology
• 4 Structure, composition development, operation
• 4.1 Membranes
• 4.2 Wall and flagella
• 5 Metabolism
• 6 Genetics
• 7 Reproduction
• 8 Ecology
• 8.1 Habitats
• 8.2 Role in chemical cycling
• 8.3 Interactions with other organisms
• 8.3.1 Mutualism
• 8.3.2 Commensalism
• 9 Significance in technology and industry
[edit] Classification
[edit] New domain
For much of the 20th century, prokaryotes were regarded as a single group of organisms and
classified based on their biochemistry, morphology and metabolism. For example, microbiologists tried to
classify microorganisms based on the structures of their cell walls, their shapes, and the substances they
consume.[2] However, a new approach was proposed in 1965,[3] using the sequences of the genes in these
organisms to work out which prokaryotes are genuinely related to each other. This approach, known as
phylogenetics, is the main method used today.

Archaea were first found in extreme environments, such as volcanic hot springs.
Archaea were first classified as a separate group of prokaryotes in 1977 by Carl Woese and George
E. Fox in phylogenetic trees based on the sequences of ribosomal RNA (rRNA) genes.[4] These two groups
were originally named the Archaebacteria and Eubacteria and treated as kingdoms or subkingdoms, which
Woese and Fox termed Urkingdoms. Woese argued that this group of prokaryotes is a fundamentally
different sort of life. To emphasize this difference, these two domains were later renamed Archaea and
Bacteria.[5] The word archaea comes from the Ancient Greek ἀρχαῖα, meaning "ancient things".[6]
At first, only the methanogens were placed in this new domain, and the archaea were seen as
extremophiles that exist only in habitats such as hot springs and salt lakes. By the end of the 20th century,
microbiologists realized that archaea is a large and diverse group of organisms that are widely distributed in
nature and are common in much less extreme habitats, such as soils and oceans.[7] This new appreciation of
the importance and ubiquity of archaea came from using the polymerase chain reaction to detect prokaryotes
in samples of water or soil from their nucleic acids alone. This allows the detection and identification of
organisms that cannot be cultured in the laboratory, which generally remains difficult.[8][9]

[edit] Current classification

Further information: Biological classification and Systematics
The classification of archaea, and of prokaryotes in general, is a rapidly moving and contentious field.
Current classification systems aim to organize archaea into groups of organisms that share structural
features and common ancestors.[10] These classifications rely heavily on the use of the sequence of
ribosomal RNA genes to reveal relationships between organisms (molecular phylogenetics).[11] Most of the
culturable and well-investigated species of archaea are members of two main phyla, the Euryarchaeota and
Crenarchaeota. Other groups have been tentatively created. For example, the peculiar species
Nanoarchaeum equitans, which was discovered in 2003, has been given its own phylum, the Nanoarchaeota.
[12] A new phylum Korarchaeota has also been proposed. It contains a small group of unusual thermophilic
species that shares features of both of the main phyla, but is most closely related to the Crenarchaeota.[13]
[14] Other recently detected species of archaea are only distantly related to any of these groups, such as the
Archaeal Richmond Mine Acidophilic Nanoorganisms (ARMAN), which were discovered in 2006[15] and are
some of the smallest organisms known.[16]

The ARMAN are a new group of archaea recently discovered in acid mine drainage.

[edit] Species
The classification of archaea into species is also controversial. Biology defines a species as a group
of related organisms. The familiar exclusive breeding criterion (organisms that can breed with each other but
not with others), is of no help because archaea reproduce asexually.[17]
Archaea show high levels of horizontal gene transfer between lineages. Some researchers suggest
that individuals can be grouped into species-like populations given highly similar genomes and infrequent
gene transfer to/from cells with less-related genomes, as in the genus Ferroplasma.[18] On the other hand,
studies in Halorubrum found significant genetic transfer to/from less-related populations, limiting the
criterion's applicability.[19] A second concern is to what extent such species designations have practical
Current knowledge on genetic diversity is fragmentary and the total number of archaean species
cannot be estimated with any accuracy.[11] Estimates of the number of phyla range from 18 to 23, of which
only 8 have representatives that have been cultured and studied directly. Many of these hypothesized groups
are known from a single rRNA sequence, indicating that the diversity among these organisms remains
obscure.[21] The Bacteria also contain many uncultured microbes with similar implications for

[edit] Origin and evolution

Further information: Timeline of evolution
Although probable prokaryotic cell fossils date to almost 3.5 billion years ago, most prokaryotes do
not have distinctive morphologies and fossil shapes cannot be used to identify them as Archaea.[23] Instead,
chemical fossils of unique lipids are more informative because such compounds do not occur in other
organisms.[24] Some publications suggest that archaean or eukaryotic lipid remains are present in shales
dating from 2.7 billion years ago;[25] such data have since been questioned.[26] Such lipids have also been
detected in Precambrian formations. The oldest such traces come from the Isua district of west Greenland,
which include Earth's oldest sediments, formed 3.8 billion years ago.[27] The archaeal lineage may be the
most ancient that exists on earth.[28]
Phylogenetic tree showing the relationship between the archaea and other forms of life. Eukaryotes
are colored red, archaea green and bacteria blue. Adapted from Ciccarelli et al.[29]
Woese argued that the bacteria, archaea, and eukaryotes represent separate lines of descent that
diverged early on from an ancestral colony of organisms.[30][31] A few biologists, however, argue that the
Archaea and Eukaryota arose from a group of bacteria.[32] In any case it is thought that viruses and archaea
began relationships approximately two billion years ago, and that co-evolution may have been occurring
between members of these groups.[33] It is possible that the last common ancestor of the bacteria and
archaea was a thermophile, which raises the possibility that lower temperatures are "extreme environments"
in archaeal terms, and organisms that live in cooler environments appeared only later.[34] Since the Archaea
and Bacteria are no more related to each other than they are to eukaryotes, the term prokaryote's only
surviving meaning is "not a eukaryote", limiting its value.[35]

[edit] Archaea and eukaryotes

The relationship between archaea and eukaryotes remains problematic. Aside from the similarities in
cell structure and function that are discussed below, many genetic trees group the two.
Complicating factors include claims that the relationship between eukaryotes and the archaeal
phylum Euryarchaeota is closer than the relationship between the Euryarchaeota and the phylum
Crenarchaeota[36] and the presence of archaean-like genes in certain bacteria, such as Thermotoga
maritima, from horizontal gene transfer.[37] The leading hypothesis is that the ancestor of the eukaryotes
diverged early from the Archaea,[38][39] and that eukaryotes arose through fusion of an archaean and
eubacterium, which became the nucleus and cytoplasm; this accounts for various genetic similarities but runs
into difficulties explaining cell structure.[40]
[edit] Morphology

The sizes of prokaryotic cells relative to other cells and biomolecules (logarithmic scale)
Individual archaea range from 0.1 micrometers (μm) to over 15 μm in diameter, and occur in various
shapes, commonly as spheres, rods, spirals or plates.[41] Other morphologies in the Crenarchaeota include
irregularly shaped lobed cells in Sulfolobus, needle-like filaments that are less than half a micrometer in
diameter in Thermofilum, and almost perfectly rectangular rods in Thermoproteus and Pyrobaculum.[42]
Haloquadratum walsbyi are flat, square archaea that live in hypersaline pools.[43] These unusual shapes are
probably maintained both by their cell walls and a prokaryotic cytoskeleton. Proteins related to the
cytoskeleton components of other organisms exist in archaea,[44] and filaments form within their cells,[45]
but in contrast to other organisms, these cellular structures are poorly understood.[46] In Thermoplasma and
Ferroplasma the lack of a cell wall means that the cells have irregular shapes, and can resemble amoebae.
Some species form aggregates or filaments of cells up to 200 μm long.[41] These organisms can be
prominent in biofilms.[48] Notably, aggregates of Thermococcus coalescens cells fuse together in culture,
forming single giant cells.[49] Archaea in the genus Pyrodictium produce an elaborate multicell colony
involving arrays of long, thin hollow tubes called cannulae that stick out from the cells' surfaces and connect
them into a dense bush-like agglomeration.[50] The function of these cannulae is not settled, but they may
allow communication or nutrient exchange with neighbors.[51] Multi-species colonies exist, such as the
"string-of-pearls" community that was discovered in 2001 in a German swamp. Round whitish colonies of a
novel Euryarchaeota species are spaced along thin filaments that can range up to 15 centimetres (5.9 in)
long; these filaments are made of a particular bacteria species.[52]

[edit] Structure, composition development, operation

Archaea and bacteria have generally similar cell structure, but cell composition and organization set
the archaea apart. Like bacteria, archaea lack interior membranes and organelles.[35] Like bacteria, archaea
cell membranes are usually bounded by a cell wall and they swim using one or more flagella.[53] Structurally,
archaea are most similar to gram-positive bacteria. Most have a single plasma membrane and cell wall, and
lack a periplasmic space; the exception to this general rule is Ignicoccus, which possess a particularly large
periplasm that contains membrane-bound vesicles and is enclosed by an outer membrane.[54]
[edit] Membranes

Membrane structures. Top, an archaeal phospholipid: 1, isoprene chains; 2, ether linkages; 3, L-

glycerol moiety; 4, phosphate group. Middle, a bacterial or eukaryotic phospholipid: 5, fatty acid chains; 6,
ester linkages; 7, D-glycerol moiety; 8, phosphate group. Bottom: 9, lipid bilayer of bacteria and eukaryotes;
10, lipid monolayer of some archaea.
Archaeal membranes are made of molecules that differ strongly from those in other life forms,
showing that archaea are related only distantly to bacteria and eukaryotes.[55] In all organisms cell
membranes are made of molecules known as phospholipids. These molecules possess both a polar part that
dissolves in water (the phosphate "head"), and a "greasy" non-polar part that does not (the lipid tail). These
dissimilar parts are connected by a glycerol moiety. In water, phospholipids cluster, with the heads facing the
water and the tails facing away from it. The major structure in cell membranes is a double layer of these
phospholipids, which is called a lipid bilayer.
These phospholipids are unusual in four ways:
• Bacteria and eukaryotes have membranes composed mainly of glycerol-ester lipids, whereas
archaea have membranes composed of glycerol-ether lipids.[56] The difference is the type of bond
that joins the lipids to the glycerol moiety; the two types are shown in yellow in the figure at the right.
In ester lipids this is an ester bond, whereas in ether lipids this is an ether bond. Ether bonds are
chemically more resistant than ester bonds. This stability might help archaea to survive extreme
temperatures and very acidic or alkaline environments.[57] Bacteria and eukaryotes do contain some
ether lipids, but in contrast to archaea these lipids are not a major part of their membranes.
• The stereochemistry of the glycerol moiety is the reverse of that found in other organisms.
The glycerol moiety can occur in two forms that are mirror images of one another, called the right-
handed and left-handed forms; in chemistry these are called enantiomers. Just as a right hand does
not fit easily into a left-handed glove, a right-handed glycerol molecule generally cannot be used or
made by enzymes adapted for the left-handed form. This suggests that archaea use entirely different
enzymes for synthesizing phospholipids than do bacteria and eukaryotes. Such enzymes developed
very early in life's history, suggesting an early split from the other two domains.[55]
• Archaeal lipid tails are chemically different from other organisms. Archaeal lipids are based
upon the isoprenoid sidechain and are long chains with multiple side-branches and sometimes even
cyclopropane or cyclohexane rings.[58] This is in contrast to the fatty acids found in other organisms'
membranes, which have straight chains with no branches or rings. Although isoprenoids play an
important role in the biochemistry of many organisms, only the archaea use them to make
phospholipids. These branched chains may help prevent archaean membranes from leaking at high
• In some archaea the lipid bilayer is replaced by a monolayer. In effect, the archaea fuse the
tails of two independent phospholipid molecules into a single molecule with two polar heads; this
fusion may make their membranes more rigid and better able to resist harsh environments.[60] For
example, the lipids in Ferroplasma are of this type, which is thought to aid this organism's survival in
its highly acidic habitat.[61]

[edit] Wall and flagella

Further information: Cell wall
Most archaea (but not Thermoplasma and Ferroplasma) possess a cell wall.[47] In most archaea the
wall is assembled from surface-layer proteins, which form an S-layer.[62] An S-layer is a rigid array of protein
molecules that cover the outside of the cell (like chain mail).[63] This layer provides both chemical and
physical protection, and can prevent macromolecules from contacting the cell membrane.[64] Unlike
bacteria, archaea lack peptidoglycan in their cell walls.[65] Methanobacteriales do have cell walls containing
pseudopeptidoglycan, which resembles eubacterial peptidoglycan in morphology, function, and physical
structure, but pseudopeptidoglycan is distinct in chemical structure; it lacks D-amino acids and N-
acetylmuramic acid.[64]
Archaea flagella operate like bacterial flagella—their long stalks are driven by rotatory motors at the
base. These motors are powered by the proton gradient across the membrane. However, archaeal flagella
are notably different in composition and development.[53] The two types of flagella evolved from different
ancestors. The bacterial flagellum shares a common ancestor with the type III secretion system,[66][67] while
archaeal flagella appear to have evolved from bacterial type IV pili.[68] In contrast to the bacterial flagellum,
which is hollow and is assembled by subunits moving up the central pore to the tip of the flagella, archaeal
flagella are synthesized by adding subunits at the base.[69]

[edit] Metabolism
Further information: Microbial metabolism
Archaea exhibit a great variety of chemical reactions in their metabolism and use many sources of
energy. These reactions are classified into nutritional groups, depending on energy and carbon sources.
Some archaea obtain energy from inorganic compounds such as sulfur or ammonia (they are lithotrophs).
These include nitrifiers, methanogens and anaerobic methane oxidisers.[70] In these reactions one
compound passes electrons to another (in a redox reaction), releasing energy to fuel the cell's activities. One
compound acts as an electron donor and one as an electron acceptor. The energy released generates
adenosine triphosphate (ATP) through chemiosmosis, in the same basic process that happens in the
mitochondrion of eukaryotic cells.[71]
Other groups of archaea use sunlight as a source of energy (they are phototrophs). However,
oxygen–generating photosynthesis does not occur in any of these organisms.[71] Many basic metabolic
pathways are shared between all forms of life; for example, archaea use a modified form of glycolysis (the
Entner–Doudoroff pathway) and either a complete or partial citric acid cycle.[72] These similarities to other
organisms probably reflect both early origins in the history of life and their high level of efficiency.[73]
Nutritional types in archaeal metabolism
Source of
Nutritional type Source of carbon Examples

Phototrophs Sunlight Halobacteria

Inorganic Ferroglobus,
Lithotrophs compounds or carbon
compounds Methanobacteria or Pyrolobus

Organic Pyrococcus, Sulfolobus
Organotrophs compounds or carbon
compounds or Methanosarcinales
Some Euryarchaeota are methanogens living in anaerobic environments such as swamps. This form
of metabolism evolved early, and it is even possible that the first free-living organism was a methanogen.[74]
A common reaction involves the use of carbon dioxide as an electron acceptor to oxidize hydrogen.
Methanogenesis involves a range of coenzymes that are unique to these archaea, such as coenzyme M and
methanofuran.[75] Other organic compounds such as alcohols, acetic acid or formic acid are used as
alternative electron acceptors by methanogens. These reactions are common in gut-dwelling archaea. Acetic
acid is also broken down into methane and carbon dioxide directly, by acetotrophic archaea. These
acetotrophs are archaea in the order Methanosarcinales, and are a major part of the communities of
microorganisms that produce biogas.[76]
Bacteriorhodopsin from Halobacterium salinarum. The retinol cofactor and residues involved in
proton transfer are shown as ball-and-stick models.[77]
Other archaea use CO2 in the atmosphere as a source of carbon, in a process called carbon fixation
(they are autotrophs). This process involves either a highly modified form of the Calvin cycle[78] or a recently
discovered metabolic pathway called the 3-hydroxypropionate/4-hydroxybutyrate cycle.[79] The
Crenarchaeota also use the reverse Krebs cycle while the Euryarchaeota also use the reductive acetyl-CoA
pathway.[80] Carbon–fixation is powered by inorganic energy sources. No known archaea carry out
photosynthesis.[81] Archaeal energy sources are extremely diverse, and range from the oxidation of
ammonia by the Nitrosopumilales[82][83] to the oxidation of hydrogen sulfide or elemental sulfur by species
of Sulfolobus, using either oxygen or metal ions as electron acceptors.[71]
Phototrophic archaea use light to produce chemical energy in the form of ATP. In the Halobacteria,
light-activated ion pumps like bacteriorhodopsin and halorhodopsin generate ion gradients by pumping ions
out of the cell across the plasma membrane. The energy stored in these electrochemical gradients is then
converted into ATP by ATP synthase.[41] This process is a form of photophosphorylation. The ability of these
light-driven pumps to move ions across membranes depends on light-driven changes in the structure of a
retinol cofactor buried in the center of the protein.[84]

[edit] Genetics
Further information: Plasmid, Genome
Archaea usually have a single circular chromosome,[85] the size of which may be as great as
5,751,492 base pairs in Methanosarcina acetivorans,[86] the largest known archaean genome. One-tenth of
this size is the tiny 490,885 base-pair genome of Nanoarchaeum equitans, the smallest archaean genome
known; it is estimated to contain only 537 protein-encoding genes.[87] Smaller independent pieces of DNA,
called plasmids, are also found in archaea. Plasmids may be transferred between cells by physical contact,
in a process that may be similar to bacterial conjugation.[88][89]

Sulfolobus infected with the DNA virus STSV1.[90] Bar is 1 micrometer.

Archaea can be infected by double-stranded DNA viruses that are unrelated to any other form of
virus and have a variety of unusual shapes, including bottles, hooked rods, or teardrops.[91] These viruses
have been studied in most detail in thermophilics, particularly the orders Sulfolobales and Thermoproteales.
[92] A single-stranded DNA virus that infects halophilic archaea was identified in 2009.[93] Defenses against
these viruses may involve RNA interference from repetitive DNA sequences that are related to the genes of
the viruses.[94][95]
Archaea are genetically distinct from bacteria and eukaryotes, with up to 15% of the proteins
encoded by any one archaeal genome being unique to the domain, although most of these unique genes
have no known function.[96] Of the remainder of the unique proteins that have an identified function, most
are involved in methanogenesis. The proteins that archaea, bacteria and eukaryotes share form a common
core of cell function, relating mostly to transcription, translation, and nucleotide metabolism.[97] Other
characteristic archaean features are the organization of genes of related function—such as enzymes that
catalyze steps in the same metabolic pathway into novel operons, and large differences in tRNA genes and
their aminoacyl tRNA synthetases.[97]
Transcription and translation in archaea resemble these processes in eukaryotes more than in
bacteria, with the archaean RNA polymerase and ribosomes being very close to their equivalents in
eukaryotes.[85] Although archaea only have one type of RNA polymerase, its structure and function in
transcription seems to be close to that of the eukaryotic RNA polymerase II, with similar protein assemblies
(the general transcription factors) directing the binding of the RNA polymerase to a gene's promoter.[98]
However, other archaean transcription factors are closer to those found in bacteria.[99] Post-transcriptional
modification is simpler than in eukaryotes, since most archaean genes lack introns, although there are many
introns in their transfer RNA and ribosomal RNA genes,[100] and introns may occur in a few protein-
encoding genes.[101][102]

[edit] Reproduction
Further information: Asexual reproduction
Archaea reproduce asexually by binary or multiple fission, fragmentation, or budding; meiosis does
not occur, so if a species of archaea exists in more than one form, all have the same genetic material.[41]
Cell division is controlled in a cell cycle; after the cell's chromosome is replicated and the two daughter
chromosomes separate, the cell divides.[103] Details have only been investigated in the genus Sulfolobus,
but here that cycle has characteristics that are similar to both bacterial and eukaryotic systems. The
chromosomes replicate from multiple starting-points (origins of replication) using DNA polymerases that
resemble the equivalent eukaryotic enzymes.[104] However, the proteins that direct cell division, such as the
protein FtsZ, which forms a contracting ring around the cell, and the components of the septum that is
constructed across the center of the cell, are similar to their bacterial equivalents.[103]
Both bacteria and eukaryotes, but not archaea, make spores.[105] Some species of Haloarchaea
undergo phenotypic switching and grow as several different cell types, including thick-walled structures that
are resistant to osmotic shock and allow the archaea to survive in water at low salt concentrations, but these
are not reproductive structures and may instead help them reach new habitats.[106]

[edit] Ecology
[edit] Habitats
Archaea exist in a broad range of habitats, and as a major part of global ecosystems,[7] may
contribute up to 20% of earth's biomass.[107] The first-discovered archaeans were extremophiles.[70]
Indeed, some archaea survive high temperatures, often above 100 °C (212 °F), as found in geysers, black
smokers, and oil wells. Other common habitats include very cold habitats and highly saline, acidic, or alkaline
water. However, archaea include mesophiles that grow in mild conditions, in marshland, sewage, the oceans,
and soils.[7]
Image of plankton (light green) in the oceans; archaea form a major part of oceanic life.
Extremophile archaea are members of four main physiological groups. These are the halophiles,
thermophiles, alkaliphiles, and acidophiles.[108] These groups are not comprehensive or phylum-specific,
nor are they mutually exclusive, since some archaea belong to several groups. Nonetheless, they are a
useful starting point for classification.
Halophiles, including the genus Halobacterium, live in extremely saline environments such as salt
lakes and outnumber their bacterial counterparts at salinities greater than 20–25%.[70] Thermophiles grow
best at temperatures above 45 °C (113 °F), in places such as hot springs; hyperthermophilic archaea grow
optimally at temperatures greater than 80 °C (176 °F).[109] The archaeal Methanopyrus kandleri Strain 116
grows at 122 °C (252 °F), the highest recorded temperature of any organism.[110]
Other archaea exist in very acidic or alkaline conditions.[108] For example, one of the most extreme
archaean acidophiles is Picrophilus torridus, which grows at pH 0, which is equivalent to thriving in 1.2 molar
sulfuric acid.[111]
This resistance to extreme environments has made archaea the focus of speculation about the
possible properties of extraterrestrial life.[112] Some extremophile habitats are not dissimilar to those on
Mars,[113] leading to the suggestion that viable microbes could be transferred between planets in
Recently, several studies have shown that archaea exist not only in mesophilic and thermophilic
environments but are also present, sometimes in high numbers, at low temperatures as well. For example,
archaea are common in cold oceanic environments such as polar seas.[115] Even more significant are the
large numbers of archaea found throughout the world's oceans in non-extreme habitats among the plankton
community (as part of the picoplankton).[116] Although these archaea can be present in extremely high
numbers (up to 40% of the microbial biomass), almost none of these species have been isolated and studied
in pure culture.[117] Consequently, our understanding of the role of archaea in ocean ecology is rudimentary,
so their full influence on global biogeochemical cycles remains largely unexplored.[118] Some marine
Crenarchaeota are capable of nitrification, suggesting these organisms may affect the oceanic nitrogen
cycle,[119] although these oceanic Crenarchaeota may also use other sources of energy.[120] Vast numbers
of archaea are also found in the sediments that cover the sea floor, with these organisms making up the
majority of living cells at depths over 1 meter below the ocean bottom.[121][122]

[edit] Role in chemical cycling

Further information: Biogeochemical cycle
Archaea recycle elements such as carbon, nitrogen and sulfur through their various habitats.
Although these activities are vital for normal ecosystem function, archaea can also contribute to human-made
changes, and even cause pollution.
Archaea carry out many steps in the nitrogen cycle. This includes both reactions that remove
nitrogen from ecosystems, such as nitrate-based respiration and denitrification, as well as processes that
introduce nitrogen, such as nitrate assimilation and nitrogen fixation.[123][124] Archaean involvement in
ammonia oxidation reactions was recently discovered. These reactions are particularly important in the
oceans.[125][126] The archaea also appear to be crucial for ammonia oxidation in soils. They produce nitrite,
which other microbes then oxidize to nitrate. Plants and other organisms consume the latter.[127]
In the sulfur cycle, archaea that grow by oxidizing sulfur compounds release this element from rocks,
making it available to other organisms. However, the archaea that do this, such as Sulfolobus, produce
sulfuric acid as a waste product, and the growth of these organisms in abandoned mines can contribute to
acid mine drainage and other environmental damage.[128]
In the carbon cycle, methanogen archaea remove hydrogen and are important in the decay of
organic matter by the populations of microorganisms that act as decomposers in anaerobic ecosystems,
such as sediments, marshes and sewage treatment works.[129] However, methane is one of the most
abundant greenhouse gases in Earth's atmosphere, constituting 18% of the global total.[130] It is 25 times
more potent as a greenhouse gas than carbon dioxide.[131] Methanogens are the primary source of
atmospheric methane, and are responsible for most of the world's yearly methane emissions.[132] As a
consequence, these archaea contribute to global greenhouse gas emissions and global warming.

[edit] Interactions with other organisms

Further information: Biological interaction
Methanogenic archaea form a symbiosis with termites.
The well-characterized interactions between archaea and other organisms are either mutual or
commensal. As of 2007, no clear examples of archaeal pathogens or parasites were known.[133][134]
However, a relationship has been proposed between some species of methanogens and infections in the
mouth,[135][136] and Nanoarchaeum equitans may be a parasite of another species of archaea, since it only
survives and reproduces within the cells of the Crenarchaeon Ignicoccus hospitalis,[137] and appears to offer
no benefit to its host.[138]

[edit] Mutualism
One well-understood example of mutualism is the interaction between protozoa and methanogenic
archaea in the digestive tracts of animals that digest cellulose, such as ruminants and termites.[139] In these
anaerobic environments, protozoa break down plant cellulose to obtain energy. This process releases
hydrogen as a waste product, but high levels of hydrogen reduce energy production. When methanogens
convert hydrogen to methane, protozoa benefit from more energy.[140]
In anaerobic protozoa such as Plagiopyla frontata, archaea reside inside the protozoa and consume
hydrogen produced in their hydrogenosomes.[141][142] Archaea also associate with larger organisms. For
example, the marine archaean Cenarchaeum symbiosum lives within (is an endosymbiont of) the sponge
Axinella mexicana.[143]

[edit] Commensalism
Archaea can also be commensals, benefiting from an association without helping or harming the
other organism. For example, the methanogen Methanobrevibacter smithii is by far the most common
archaean in the human flora, making up about one in ten of all the prokaryotes in the human gut.[144] In
termites and in humans, these methanogens may in fact be mutualists, interacting with other microbes in the
gut to aid digestion.[145] Archaean communities also associate with a range of other organisms, such as on
the surface of corals,[146] and in the region of soil that surrounds plant roots (the rhizosphere).[147][148]

[edit] Significance in technology and industry

Further information: Biotechnology
Extremophile archaea, particularly those resistant either to heat or to extremes of acidity and
alkalinity, are a source of enzymes that function under these harsh conditions.[149][150] These enzymes
have found many uses. For example, thermostable DNA polymerases, such as the Pfu DNA polymerase
from Pyrococcus furiosus, revolutionized molecular biology by allowing the polymerase chain reaction to be
used in research as a simple and rapid technique for cloning DNA. In industry, amylases, galactosidases and
pullulanases in other species of Pyrococcus that function at over 100 °C (212 °F) allow food processing at
high temperatures, such as the production of low lactose milk and whey.[151] Enzymes from these
thermophilic archaea also tend to be very stable in organic solvents, allowing their use in environmentally
friendly processes in green chemistry that synthesize organic compounds.[150] This stability makes them
easier to use in structural biology. Consequently the counterparts of bacterial or eukaryotic enzymes from
extremophile archaea are often used in structural studies.[152]
In contrast to the range of applications of archaean enzymes, the use of the organisms themselves in
biotechnology is less developed. Methanogenic archaea are a vital part of sewage treatment, since they are
part of the community of microorganisms that carry out anaerobic digestion and produce biogas.[153] In
mineral processing, acidophilic archaea display promise for the extraction of metals from ores, including gold,
cobalt and copper.[154]
Archaea host a new class of potentially useful antibiotics. A few of these archaeocins have been
characterized, but hundreds more are believed to exist, especially within Haloarchaea and Sulfolobus.[155]
These compounds differ in structure from bacterial antibiotics, so they may have novel modes of action. In
addition, they may allow the creation of new selectable markers for use in archaeal molecular biology.[156]

[edit] See also

• Aerobic methane production
• List of Archaea genera
• List of sequenced archaeal genomes
• The Surprising Archaea (book)

[edit] References
1. ^ Pace NR (May 2006). "Time for a change". Nature 441 (7091): 289. doi:10.1038/441289a.
PMID 16710401.
2. ^ Staley JT (2006). "The bacterial species dilemma and the genomic-phylogenetic species
concept". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 361 (1475): 1899–909.
doi:10.1098/rstb.2006.1914. PMID 17062409. PMC 1857736.
3. ^ Zuckerkandl E, Pauling L (1965). "Molecules as documents of evolutionary history". J.
Theor. Biol. 8 (2): 357–66. doi:10.1016/0022-5193(65)90083-4. PMID 5876245.
4. ^ Woese C, Fox G (1977). "Phylogenetic structure of the prokaryotic domain: the primary
kingdoms". Proc Natl Acad Sci USA 74 (11): 5088–90. doi:10.1073/pnas.74.11.5088. PMID 270744.
5. ^ Woese CR, Kandler O, Wheelis ML (1990). "Towards a natural system of organisms:
proposal for the domains Archaea, Bacteria, and Eucarya". Proc. Natl. Acad. Sci. U.S.A. 87 (12):
4576–9. doi:10.1073/pnas.87.12.4576. PMID 2112744. PMC 54159.
6. ^ archaea. (2008). In Merriam-Webster Online Dictionary. Retrieved July 1, 2008, from
7. ^ a b c DeLong EF (1998). "Everything in moderation: archaea as 'non-extremophiles'". Curr.
Opin. Genet. Dev. 8 (6): 649–54. doi:10.1016/S0959-437X(98)80032-4. PMID 9914204.
8. ^ Theron J, Cloete TE (2000). "Molecular techniques for determining microbial diversity and
community structure in natural environments". Crit. Rev. Microbiol. 26 (1): 37–57.
doi:10.1080/10408410091154174. PMID 10782339.
9. ^ Schmidt TM (2006). "The maturing of microbial ecology" (PDF). Int. Microbiol. 9 (3): 217–23.
PMID 17061212.
10.^ Gevers D, Dawyndt P, Vandamme P, et al. (2006). "Stepping stones towards a new
prokaryotic taxonomy". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 361 (1475): 1911–6.
doi:10.1098/rstb.2006.1915. PMID 17062410. PMC 1764938.
11.^ a b Robertson CE, Harris JK, Spear JR, Pace NR (2005). "Phylogenetic diversity and
ecology of environmental Archaea". Curr. Opin. Microbiol. 8 (6): 638–42.
doi:10.1016/j.mib.2005.10.003. PMID 16236543.
12.^ Huber H, Hohn MJ, Rachel R, Fuchs T, Wimmer VC, Stetter KO. (2002). "A new phylum of
Archaea represented by a nanosized hyperthermophilic symbiont". Nature 417 (6884): 27–8.
doi:10.1038/417063a. PMID 11986665.
13.^ Barns SM, Delwiche CF, Palmer JD, Pace NR (1996). "Perspectives on archaeal diversity,
thermophily and monophyly from environmental rRNA sequences". Proc. Natl. Acad. Sci. U.S.A. 93
(17): 9188–93. doi:10.1073/pnas.93.17.9188. PMID 8799176. PMC 38617.
14.^ Elkins JG, Podar M, Graham DE, et al. (June 2008). "A korarchaeal genome reveals
insights into the evolution of the Archaea". Proc. Natl. Acad. Sci. U.S.A. 105 (23): 8102–7.
doi:10.1073/pnas.0801980105. PMID 18535141. PMC 2430366.
15.^ Baker, B.J., Tyson, G.W., Webb, R.I., Flanagan, J., Hugenholtz, P. and Banfield, J.F.
(2006). "Lineages of acidophilic Archaea revealed by community genomic analysis. Science".
Science 314 (6884): 1933–1935. doi:10.1126/science.1132690. PMID 17185602.
16.^ Baker BJ, Comolli LR, Dick GJ, et al. (May 2010). "Enigmatic, ultrasmall, uncultivated
Archaea". Proc. Natl. Acad. Sci. U.S.A. 107 (19): 8806–11. doi:10.1073/pnas.0914470107.
PMID 20421484.
17.^ de Queiroz K (2005). "Ernst Mayr and the modern concept of species". Proc. Natl. Acad.
Sci. U.S.A. 102 Suppl 1: 6600–7. doi:10.1073/pnas.0502030102. PMID 15851674. PMC 1131873.
18.^ Eppley JM, Tyson GW, Getz WM, Banfield JF (2007). "Genetic exchange across a species
boundary in the archaeal genus ferroplasma". Genetics 177 (1): 407–16.
doi:10.1534/genetics.107.072892. PMID 17603112. PMC 2013692.
19.^ Papke RT, Zhaxybayeva O, Feil EJ, Sommerfeld K, Muise D, Doolittle WF (2007).
"Searching for species in haloarchaea". Proc. Natl. Acad. Sci. U.S.A. 104 (35): 14092–7.
doi:10.1073/pnas.0706358104. PMID 17715057. PMC 1955782.
20.^ Kunin V, Goldovsky L, Darzentas N, Ouzounis CA (2005). "The net of life: reconstructing
the microbial phylogenetic network". Genome Res. 15 (7): 954–9. doi:10.1101/gr.3666505.
PMID 15965028. PMC 1172039.
21.^ Hugenholtz P (2002). "Exploring prokaryotic diversity in the genomic era". Genome Biol. 3
(2): REVIEWS0003. doi:10.1186/gb-2002-3-2-reviews0003. PMID 11864374. PMC 139013.
22.^ Rappé MS, Giovannoni SJ (2003). "The uncultured microbial majority". Annu. Rev.
Microbiol. 57: 369–94. doi:10.1146/annurev.micro.57.030502.090759. PMID 14527284.
23.^ Schopf J (2006). "Fossil evidence of Archaean life" (PDF). Philos Trans R Soc Lond B Biol
Sci 361 (1470): 869–85. doi:10.1098/rstb.2006.1834. PMID 16754604. PMC 1578735.
24.^ Chappe B, Albrecht P, Michaelis W (July 1982). "Polar Lipids of Archaebacteria in
Sediments and Petroleums". Science 217 (4554): 65–66. doi:10.1126/science.217.4554.65.
PMID 17739984.
25.^ Brocks JJ, Logan GA, Buick R, Summons RE (1999). "Archean molecular fossils and the
early rise of eukaryotes". Science 285 (5430): 1033–6. doi:10.1126/science.285.5430.1033.
PMID 10446042.
26.^ Rasmussen B, Fletcher IR, Brocks JJ, Kilburn MR (October 2008). "Reassessing the first
appearance of eukaryotes and cyanobacteria". Nature 455 (7216): 1101–4. doi:10.1038/nature07381.
PMID 18948954.
27.^ Hahn, Jürgen; Pat Haug (1986). "Traces of Archaebacteria in ancient sediments". System
Applied Microbiology 7 (Archaebacteria '85 Proceedings): 178–83.
28.^ Wang M, Yafremava LS, Caetano-Anollés D, Mittenthal JE, Caetano-Anollés G (2007).
"Reductive evolution of architectural repertoires in proteomes and the birth of the tripartite world".
Genome Res. 17 (11): 1572–85. doi:10.1101/gr.6454307. PMID 17908824.
29.^ Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork P (2006). "Toward
automatic reconstruction of a highly resolved tree of life". Science 311 (5765): 1283–7.
doi:10.1126/science.1123061. PMID 16513982.
30.^ Woese CR, Gupta R (1981). "Are archaebacteria merely derived 'prokaryotes'?". Nature
289 (5793): 95–6. doi:10.1038/289095a0. PMID 6161309.
31.^ Woese C (1998). "The universal ancestor". Proc. Natl. Acad. Sci. U.S.A. 95 (12): 6854–9.
doi:10.1073/pnas.95.12.6854. PMID 9618502. PMC 22660.
32.^ Gupta RS (2000). "The natural evolutionary relationships among prokaryotes". Crit. Rev.
Microbiol. 26 (2): 111–31. doi:10.1080/10408410091154219. PMID 10890353.
33.^ C. Michael Hogan. 2010. Virus. Encyclopedia of Earth. Editors: Cutler Cleveland and
Sidney Draggan
34.^ Gribaldo S, Brochier-Armanet C (2006). "The origin and evolution of Archaea: a state of the
art". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 361 (1470): 1007–22. doi:10.1098/rstb.2006.1841.
PMID 16754611. PMC 1578729.
35.^ a b Woese CR (1 March 1994). "There must be a prokaryote somewhere: microbiology's
search for itself". Microbiol. Rev. 58 (1): 1–9. PMID 8177167. PMC 372949.
36.^ Lake JA (January 1988). "Origin of the eukaryotic nucleus determined by rate-invariant
analysis of rRNA sequences". Nature 331 (6152): 184–6. doi:10.1038/331184a0. PMID 3340165.
37.^ Nelson KE, Clayton RA, Gill SR, et al. (1999). "Evidence for lateral gene transfer between
Archaea and bacteria from genome sequence of Thermotoga maritima". Nature 399 (6734): 323–9.
doi:10.1038/20601. PMID 10360571.
38.^ Gouy M, Li WH (May 1989). "Phylogenetic analysis based on rRNA sequences supports
the archaebacterial rather than the eocyte tree". Nature 339 (6220): 145–7. doi:10.1038/339145a0.
PMID 2497353.
39.^ Yutin N, Makarova KS, Mekhedov SL, Wolf YI, Koonin EV (May 2008). "The deep archaeal
roots of eukaryotes". Mol. Biol. Evol. 25 (8): 1619–30. doi:10.1093/molbev/msn108. PMID 18463089.
PMC 2464739.
40.^ Lake JA. (1988). "Origin of the eukaryotic nucleus determined by rate-invariant analysis of
rRNA sequences". Nature 331 (6152): 184–6. doi:10.1038/331184a0. PMID 3340165.
41.^ a b c d Krieg, Noel (2005). Bergey's Manual of Systematic Bacteriology. US: Springer.
pp. 21–6. ISBN 978-0-387-24143-2.
42.^ Barns, Sue and Burggraf, Siegfried. (1997) Crenarchaeota. Version 01 January 1997. in
The Tree of Life Web Project
43.^ Walsby, A.E. (1980). "A square bacterium". Nature 283 (5742): 69–71.
44.^ Hara F, Yamashiro K, Nemoto N, et al. (2007). "An actin homolog of the archaeon
Thermoplasma acidophilum that retains the ancient characteristics of eukaryotic actin". J. Bacteriol.
189 (5): 2039–45. doi:10.1128/JB.01454-06. PMID 17189356. PMC 1855749.
45.^ Trent JD, Kagawa HK, Yaoi T, Olle E, Zaluzec NJ (1997). "Chaperonin filaments: the
archaeal cytoskeleton?". Proc. Natl. Acad. Sci. U.S.A. 94 (10): 5383–8. doi:10.1073/pnas.94.10.5383.
PMID 9144246. PMC 24687.
46.^ Hixon WG, Searcy DG (1993). "Cytoskeleton in the archaebacterium Thermoplasma
acidophilum? Viscosity increase in soluble extracts". BioSystems 29 (2–3): 151–60. doi:10.1016/0303-
2647(93)90091-P. PMID 8374067.
47.^ a b Golyshina OV, Pivovarova TA, Karavaiko GI, et al. (1 May 2000). "Ferroplasma
acidiphilum gen. nov., sp. nov., an acidophilic, autotrophic, ferrous-iron-oxidizing, cell-wall-lacking,
mesophilic member of the Ferroplasmaceae fam. nov., comprising a distinct lineage of the Archaea".
Int. J. Syst. Evol. Microbiol. 50 Pt 3 (3): 997–1006. PMID 10843038.
48.^ Hall-Stoodley L, Costerton JW, Stoodley P (2004). "Bacterial biofilms: from the natural
environment to infectious diseases". Nat. Rev. Microbiol. 2 (2): 95–108. doi:10.1038/nrmicro821.
PMID 15040259.
49.^ Kuwabara T, Minaba M, Iwayama Y, et al. (November 2005). "Thermococcus coalescens
sp. nov., a cell-fusing hyperthermophilic archaeon from Suiyo Seamount". Int. J. Syst. Evol.
Microbiol. 55 (Pt 6): 2507–14. doi:10.1099/ijs.0.63432-0. PMID 16280518.
50.^ Nickell S, Hegerl R, Baumeister W, Rachel R (2003). "Pyrodictium cannulae enter the
periplasmic space but do not enter the cytoplasm, as revealed by cryo-electron tomography". J.
Struct. Biol. 141 (1): 34–42. doi:10.1016/S1047-8477(02)00581-6. PMID 12576018.
51.^ Horn C, Paulmann B, Kerlen G, Junker N, Huber H (15 August 1999). "In vivo observation
of cell division of anaerobic hyperthermophiles by using a high-intensity dark-field microscope". J.
Bacteriol. 181 (16): 5114–8. PMID 10438790. PMC 94007.
52.^ Rudolph C, Wanner G, Huber R (May 2001). "Natural communities of novel archaea and
bacteria growing in cold sulfurous springs with a string-of-pearls-like morphology". Appl. Environ.
Microbiol. 67 (5): 2336–44. doi:10.1128/AEM.67.5.2336-2344.2001. PMID 11319120.
53.^ a b Thomas NA, Bardy SL, Jarrell KF (2001). "The archaeal flagellum: a different kind of
prokaryotic motility structure". FEMS Microbiol. Rev. 25 (2): 147–74. doi:10.1111/j.1574-
6976.2001.tb00575.x. PMID 11250034.
54.^ Rachel R, Wyschkony I, Riehl S, Huber H (March 2002). "The ultrastructure of Ignicoccus:
evidence for a novel outer membrane and for intracellular vesicle budding in an archaeon" (PDF).
Archaea 1 (1): 9–18. doi:10.1155/2002/307480. PMID 15803654. PMC 2685547.
55.^ a b Koga Y, Morii H (2007). "Biosynthesis of ether-type polar lipids in archaea and
evolutionary considerations". Microbiol. Mol. Biol. Rev. 71 (1): 97–120. doi:10.1128/MMBR.00033-06.
PMID 17347520. PMC 1847378.
56.^ De Rosa M, Gambacorta A, Gliozzi A (1 March 1986). "Structure, biosynthesis, and
physicochemical properties of archaebacterial lipids". Microbiol. Rev. 50 (1): 70–80. PMID 3083222.
PMC 373054.
57.^ Albers SV, van de Vossenberg JL, Driessen AJ, Konings WN (September 2000).
"Adaptations of the archaeal cell membrane to heat stress". Front. Biosci. 5: D813–20.
doi:10.2741/albers. PMID 10966867.
58.^ Damsté JS, Schouten S, Hopmans EC, van Duin AC, Geenevasen JA (October 2002).
"Crenarchaeol: the characteristic core glycerol dibiphytanyl glycerol tetraether membrane lipid of
cosmopolitan pelagic crenarchaeota". J. Lipid Res. 43 (10): 1641–51. doi:10.1194/jlr.M200148-
JLR200. PMID 12364548.
59.^ Koga Y, Morii H (November 2005). "Recent advances in structural research on ether lipids
from archaea including comparative and physiological aspects". Biosci. Biotechnol. Biochem. 69
(11): 2019–34. doi:10.1271/bbb.69.2019. PMID 16306681.
60.^ Hanford MJ, Peeples TL (January 2002). "Archaeal tetraether lipids: unique structures and
applications". Appl. Biochem. Biotechnol. 97 (1): 45–62. doi:10.1385/ABAB:97:1:45.
PMID 11900115.
61.^ Macalady JL, Vestling MM, Baumler D, Boekelheide N, Kaspar CW, Banfield JF (October
2004). "Tetraether-linked membrane monolayers in Ferroplasma spp: a key to survival in acid".
Extremophiles 8 (5): 411–9. doi:10.1007/s00792-004-0404-5. PMID 15258835.
62.^ Sára M, Sleytr UB (2000). "S-Layer proteins". J. Bacteriol. 182 (4): 859–68.
doi:10.1128/JB.182.4.859-868.2000. PMID 10648507. PMC 94357.
63.^ Engelhardt H, Peters J (1998). "Structural research on surface layers: a focus on stability,
surface layer homology domains, and surface layer-cell wall interactions". J Struct Biol 124 (2–3):
276–302. doi:10.1006/jsbi.1998.4070. PMID 10049812.
64.^ a b Kandler, O; König, H (1998). "Cell wall polymers in Archaea (Archaebacteria)" (PDF).
Cellular and Molecular Life Sciences (CMLS) 54 (4): 305–308. doi:10.1007/s000180050156.
65.^ Howland, John L. (2000). The Surprising Archaea: Discovering Another Domain of Life .
Oxford: Oxford University Press. p. 32. ISBN 0-19-511183-4.
66.^ Gophna U, Ron EZ, Graur D (July 2003). "Bacterial type III secretion systems are ancient
and evolved by multiple horizontal-transfer events". Gene 312: 151–63. doi:10.1016/S0378-
1119(03)00612-7. PMID 12909351.
67.^ Nguyen L, Paulsen IT, Tchieu J, Hueck CJ, Saier MH (April 2000). "Phylogenetic analyses
of the constituents of Type III protein secretion systems". J. Mol. Microbiol. Biotechnol. 2 (2): 125–44.
PMID 10939240.
68.^ Ng SY, Chaban B, Jarrell KF (2006). "Archaeal flagella, bacterial flagella and type IV pili: a
comparison of genes and posttranslational modifications". J. Mol. Microbiol. Biotechnol. 11 (3–5):
167–91. doi:10.1159/000094053. PMID 16983194.
69.^ Bardy SL, Ng SY, Jarrell KF (February 2003). "Prokaryotic motility structures". Microbiology
(Reading, Engl.) 149 (Pt 2): 295–304. doi:10.1099/mic.0.25948-0. PMID 12624192.
70.^ a b c Valentine DL (2007). "Adaptations to energy stress dictate the ecology and evolution
of the Archaea". Nat. Rev. Microbiol. 5 (4): 316–23. doi:10.1038/nrmicro1619. PMID 17334387.
71.^ a b c Schäfer G, Engelhard M, Müller V (1 September 1999). "Bioenergetics of the
Archaea". Microbiol. Mol. Biol. Rev. 63 (3): 570–620. PMID 10477309. PMC 103747.
72.^ Zillig W (December 1991). "Comparative biochemistry of Archaea and Bacteria". Curr.
Opin. Genet. Dev. 1 (4): 544–51. doi:10.1016/S0959-437X(05)80206-0. PMID 1822288.
73.^ Romano A, Conway T (1996). "Evolution of carbohydrate metabolic pathways". Res
Microbiol 147 (6–7): 448–55. doi:10.1016/0923-2508(96)83998-2. PMID 9084754.
74.^ Koch A (1998). "How did bacteria come to be?". Adv Microb Physiol 40: 353–99.
doi:10.1016/S0065-2911(08)60135-6. PMID 9889982.
75.^ DiMarco AA, Bobik TA, Wolfe RS (1990). "Unusual coenzymes of methanogenesis". Annu.
Rev. Biochem. 59: 355–94. doi:10.1146/ PMID 2115763.
76.^ Klocke M, Nettmann E, Bergmann I, et al. (May 2008). "Characterization of the
methanogenic Archaea within two-phase biogas reactor systems operated with plant biomass". Syst.
Appl. Microbiol. 31 (3): 190–205. doi:10.1016/j.syapm.2008.02.003. PMID 18501543.
77.^ Based on PDB 1FBB. Data published in Subramaniam S, Henderson R (August 2000).
"Molecular mechanism of vectorial proton translocation by bacteriorhodopsin". Nature 406 (6796):
653–7. doi:10.1038/35020614. PMID 10949309.
78.^ Mueller-Cajar O, Badger MR (August 2007). "New roads lead to Rubisco in
archaebacteria". Bioessays 29 (8): 722–4. doi:10.1002/bies.20616. PMID 17621634.
79.^ Berg IA, Kockelkorn D, Buckel W, Fuchs G (December 2007). "A 3-hydroxypropionate/4-
hydroxybutyrate autotrophic carbon dioxide assimilation pathway in Archaea". Science (journal) 318
(5857): 1782–6. doi:10.1126/science.1149976. PMID 18079405.
80.^ Thauer RK (December 2007). "Microbiology. A fifth pathway of carbon fixation". Science
(journal) 318 (5857): 1732–3. doi:10.1126/science.1152209. PMID 18079388.
81.^ Bryant DA, Frigaard NU (November 2006). "Prokaryotic photosynthesis and phototrophy
illuminated". Trends Microbiol. 14 (11): 488–96. doi:10.1016/j.tim.2006.09.001. PMID 16997562.
82.^ Könneke M, Bernhard AE, de la Torre JR, Walker CB, Waterbury JB, Stahl DA (September
2005). "Isolation of an autotrophic ammonia-oxidizing marine archaeon". Nature 437 (7058): 543–6.
doi:10.1038/nature03911. PMID 16177789.
83.^ Francis CA, Beman JM, Kuypers MM (May 2007). "New processes and players in the
nitrogen cycle: the microbial ecology of anaerobic and archaeal ammonia oxidation". ISME J 1 (1):
19–27. doi:10.1038/ismej.2007.8. PMID 18043610.
84.^ Lanyi JK (2004). "Bacteriorhodopsin". Annu. Rev. Physiol. 66: 665–88.
doi:10.1146/annurev.physiol.66.032102.150049. PMID 14977418.
85.^ a b Allers T, Mevarech M (2005). "Archaeal genetics - the third way". Nat. Rev. Genet. 6 (1):
58–73. doi:10.1038/nrg1504. PMID 15630422.
86.^ Galagan JE, Nusbaum C, Roy A, et al. (April 2002). "The genome of M. acetivorans reveals
extensive metabolic and physiological diversity". Genome Res. 12 (4): 532–42.
doi:10.1101/gr.223902. PMID 11932238.
87.^ Waters E, et al. (2003). "The genome of Nanoarchaeum equitans: insights into early
archaeal evolution and derived parasitism". Proc. Natl. Acad. Sci. U.S.A. 100 (22): 12984–8.
doi:10.1073/pnas.1735403100. PMID 14566062. PMC 240731.
88.^ Schleper C, Holz I, Janekovic D, Murphy J, Zillig W (1 August 1995). "A multicopy plasmid
of the extremely thermophilic archaeon Sulfolobus effects its transfer to recipients by mating". J.
Bacteriol. 177 (15): 4417–26. PMID 7635827. PMC 177192.
89.^ Sota M; Top EM (2008). "Horizontal Gene Transfer Mediated by Plasmids". Plasmids:
Current Research and Future Trends. Caister Academic Press. ISBN 978-1-904455-35-6.
90.^ Xiang X, Chen L, Huang X, Luo Y, She Q, Huang L (2005). "Sulfolobus tengchongensis
spindle-shaped virus STSV1: virus-host interactions and genomic features". J. Virol. 79 (14): 8677–
86. doi:10.1128/JVI.79.14.8677-8686.2005. PMID 15994761. PMC 1168784.
91.^ Prangishvili D, Forterre P, Garrett RA (2006). "Viruses of the Archaea: a unifying view".
Nat. Rev. Microbiol. 4 (11): 837–48. doi:10.1038/nrmicro1527. PMID 17041631.
92.^ Prangishvili D, Garrett RA (2004). "Exceptionally diverse morphotypes and genomes of
crenarchaeal hyperthermophilic viruses". Biochem. Soc. Trans. 32 (Pt 2): 204–8.
doi:10.1042/BST0320204. PMID 15046572.
93.^ Pietilä MK, Roine E, Paulin L, Kalkkinen N, Bamford DH (March 2009). "An ssDNA virus
infecting archaea; A new lineage of viruses with a membrane envelope". Mol. Microbiol. 72 (2): 307–
19. doi:10.1111/j.1365-2958.2009.06642.x. PMID 19298373.
94.^ Mojica FJ, Díez-Villaseñor C, García-Martínez J, Soria E (2005). "Intervening sequences of
regularly spaced prokaryotic repeats derive from foreign genetic elements". J. Mol. Evol. 60 (2): 174–
82. doi:10.1007/s00239-004-0046-3. PMID 15791728.
95.^ Makarova KS, Grishin NV, Shabalina SA, Wolf YI, Koonin EV (2006). "A putative RNA-
interference-based immune system in prokaryotes: computational analysis of the predicted
enzymatic machinery, functional analogies with eukaryotic RNAi, and hypothetical mechanisms of
action". Biol. Direct 1: 7. doi:10.1186/1745-6150-1-7. PMID 16545108.
96.^ Graham DE, Overbeek R, Olsen GJ, Woese CR (2000). "An archaeal genomic signature".
Proc. Natl. Acad. Sci. U.S.A. 97 (7): 3304–8. doi:10.1073/pnas.050564797. PMID 10716711.
PMC 16234.
97.^ a b Gaasterland T (1999). "Archaeal genomics". Curr. Opin. Microbiol. 2 (5): 542–7.
doi:10.1016/S1369-5274(99)00014-4. PMID 10508726.
98.^ Werner F (September 2007). "Structure and function of archaeal RNA polymerases". Mol.
Microbiol. 65 (6): 1395–404. doi:10.1111/j.1365-2958.2007.05876.x. PMID 17697097.
99.^ Aravind L, Koonin EV (1999). "DNA-binding proteins and evolution of transcription
regulation in the archaea". Nucleic Acids Res. 27 (23): 4658–70. doi:10.1093/nar/27.23.4658.
PMID 10556324. PMC 148756.
100.^ Lykke-Andersen J, Aagaard C, Semionenkov M, Garrett RA (September 1997). "Archaeal
introns: splicing, intercellular mobility and evolution". Trends Biochem. Sci. 22 (9): 326–31.
doi:10.1016/S0968-0004(97)01113-4. PMID 9301331.
101.^ Watanabe Y, Yokobori S, Inaba T, et al. (January 2002). "Introns in protein-coding genes
in Archaea". FEBS Lett. 510 (1–2): 27–30. doi:10.1016/S0014-5793(01)03219-7. PMID 11755525.
102.^ Yoshinari S, Itoh T, Hallam SJ, et al. (August 2006). "Archaeal pre-mRNA splicing: a
connection to hetero-oligomeric splicing endonuclease". Biochem. Biophys. Res. Commun. 346 (3):
1024–32. doi:10.1016/j.bbrc.2006.06.011. PMID 16781672.
103.^ a b Bernander R (1998). "Archaea and the cell cycle". Mol. Microbiol. 29 (4): 955–61.
doi:10.1046/j.1365-2958.1998.00956.x. PMID 9767564.
104.^ Kelman LM, Kelman Z (2004). "Multiple origins of replication in archaea". Trends
Microbiol. 12 (9): 399–401. doi:10.1016/j.tim.2004.07.001. PMID 153371581.
105.^ Onyenwoke RU, Brill JA, Farahi K, Wiegel J (2004). "Sporulation genes in members of the
low G+C Gram-type-positive phylogenetic branch ( Firmicutes)". Arch. Microbiol. 182 (2–3): 182–92.
doi:10.1007/s00203-004-0696-y. PMID 15340788.
106.^ Kostrikina NA, Zvyagintseva IS, Duda VI. (1991). "Cytological peculiarities of some
extremely halophilic soil archaeobacteria". Arch. Microbiol. 156 (5): 344–49.
107.^ DeLong EF, Pace NR (2001). "Environmental diversity of bacteria and archaea". Syst.
Biol. 50 (4): 470–8. doi:10.1080/106351501750435040. PMID 12116647.
108.^ a b Pikuta EV, Hoover RB, Tang J (2007). "Microbial extremophiles at the limits of life".
Crit. Rev. Microbiol. 33 (3): 183–209. doi:10.1080/10408410701451948. PMID 17653987.
109.^ Madigan MT, Martino JM (2006). Brock Biology of Microorganisms (11th ed.). Pearson.
p. 136. ISBN 0-13-196893-9.
110.^ Takai K, Nakamura K, Toki T, Tsunogai U, Miyazaki M, Miyazaki J, Hirayama H,
Nakagawa S, Nunoura T, Horikoshi K (2008). "Cell proliferation at 122°C and isotopically heavy CH4
production by a hyperthermophilic methanogen under high-pressure cultivation". Proc Natl Acad Sci
USA 105 (31): 10949–54. doi:10.1073/pnas.0712334105. PMID 18664583.
111.^ Ciaramella M, Napoli A, Rossi M (February 2005). "Another extreme genome: how to live
at pH 0". Trends Microbiol. 13 (2): 49–51. doi:10.1016/j.tim.2004.12.001. PMID 15680761.
112.^ Javaux EJ (2006). "Extreme life on Earth—past, present and possibly beyond". Res.
Microbiol. 157 (1): 37–48. doi:10.1016/j.resmic.2005.07.008. PMID 16376523.
113.^ Nealson KH (January 1999). "Post-Viking microbiology: new approaches, new data, new
insights". Orig Life Evol Biosph 29 (1): 73–93. doi:10.1023/A:1006515817767. PMID 11536899.
114.^ Davies PC (1996). "The transfer of viable microorganisms between planets". Ciba Found.
Symp. 202: 304–14; discussion 314–7. PMID 9243022.
115.^ López-García P, López-López A, Moreira D, Rodríguez-Valera F (July 2001). "Diversity of
free-living prokaryotes from a deep-sea site at the Antarctic Polar Front". FEMS Microbiol. Ecol. 36
(2–3): 193–202. PMID 11451524.
116.^ Karner MB, DeLong EF, Karl DM (2001). "Archaeal dominance in the mesopelagic zone of
the Pacific Ocean". Nature 409 (6819): 507–10. doi:10.1038/35054051. PMID 11206545.
117.^ Giovannoni SJ, Stingl U. (2005). "Molecular diversity and ecology of microbial plankton".
Nature 427 (7057): 343–8. doi:10.1038/nature04158. PMID 16163344.
118.^ DeLong EF, Karl DM (September 2005). "Genomic perspectives in microbial
oceanography". Nature 437 (7057): 336–42. doi:10.1038/nature04157. PMID 16163343.
119.^ Konneke M, Bernhard AE, de la Torre JR, Walker CB, Waterbury JB, Stahl DA. (2005).
"Isolation of an autotrophic ammonia-oxidizing marine archaeon". Nature 437 (7057): 543–6.
doi:10.1038/nature03911. PMID 16177789.
120.^ Agogué, H; Brink, M; Dinasquet, J; Herndl, GJ (2008). "Major gradients in putatively
nitrifying and non-nitrifying Archaea in the deep North Atlantic". Nature 456 (7223): 788–791.
doi:10.1038/nature07535. PMID 19037244.
121.^ Teske A, Sørensen KB (January 2008). "Uncultured archaea in deep marine subsurface
sediments: have we caught them all?". ISME J 2 (1): 3–18. doi:10.1038/ismej.2007.90.
PMID 18180743.
122.^ Lipp JS, Morono Y, Inagaki F, Hinrichs KU (July 2008). "Significant contribution of
Archaea to extant biomass in marine subsurface sediments". Nature 454 (7207): 991–4.
doi:10.1038/nature07174. PMID 18641632.
123.^ Cabello P, Roldán MD, Moreno-Vivián C (November 2004). "Nitrate reduction and the
nitrogen cycle in archaea". Microbiology (Reading, Engl.) 150 (Pt 11): 3527–46.
doi:10.1099/mic.0.27303-0. PMID 15528644.
124.^ Mehta MP, Baross JA (December 2006). "Nitrogen fixation at 92 degrees C by a
hydrothermal vent archaeon". Science (journal) 314 (5806): 1783–6. doi:10.1126/science.1134772.
PMID 17170307.
125.^ Francis CA, Beman JM, Kuypers MM (May 2007). "New processes and players in the
nitrogen cycle: the microbial ecology of anaerobic and archaeal ammonia oxidation". ISME J 1 (1):
19–27. doi:10.1038/ismej.2007.8. PMID 18043610.
126.^ Coolen MJ, Abbas B, van Bleijswijk J, et al. (April 2007). "Putative ammonia-oxidizing
Crenarchaeota in suboxic waters of the Black Sea: a basin-wide ecological study using 16S
ribosomal and functional genes and membrane lipids". Environ. Microbiol. 9 (4): 1001–16.
doi:10.1111/j.1462-2920.2006.01227.x. PMID 17359272.
127.^ Leininger S, Urich T, Schloter M, et al. (August 2006). "Archaea predominate among
ammonia-oxidizing prokaryotes in soils". Nature 442 (7104): 806–9. doi:10.1038/nature04983.
PMID 16915287.
128.^ Baker, B. J; Banfield, J. F (2003). "Microbial communities in acid mine drainage". FEMS
Microbiology Ecology 44 (2): 139–152. doi:10.1016/S0168-6496(03)00028-X. PMID 19719632.
129.^ Schimel J (August 2004). "Playing scales in the methane cycle: from microbial ecology to
the globe". Proc. Natl. Acad. Sci. U.S.A. 101 (34): 12400–1. doi:10.1073/pnas.0405075101.
PMID 15314221. PMC 515073.
130.^ "EDGAR 3.2 Fast Track 2000".
Retrieved 2008-06-26.
131.^ "Annual Greenhouse Gas Index (AGGI) Indicates Sharp Rise in Carbon Dioxide and
Methane in 2007". 2008-04-23. Archived from the original on May 14, 2008.
Retrieved 2008-06-26.
132.^ "Trace Gases: Current Observations, Trends, and Budgets". Climate Change 2001.
United Nations Environment Programme.
133.^ Eckburg P, Lepp P, Relman D (2003). "Archaea and their potential role in human
disease". Infect Immun 71 (2): 591–6. doi:10.1128/IAI.71.2.591-596.2003. PMID 12540534.
134.^ Cavicchioli R, Curmi P, Saunders N, Thomas T (2003). "Pathogenic archaea: do they
exist?". Bioessays 25 (11): 1119–28. doi:10.1002/bies.10354. PMID 14579252.
135.^ Lepp P, Brinig M, Ouverney C, Palm K, Armitage G, Relman D (2004). "Methanogenic
Archaea and human periodontal disease". Proc Natl Acad Sci USA 101 (16): 6176–81.
doi:10.1073/pnas.0308766101. PMID 15067114.
136.^ Vianna ME, Conrads G, Gomes BP, Horz HP (April 2006). "Identification and
quantification of archaea involved in primary endodontic infections". J. Clin. Microbiol. 44 (4): 1274–
82. doi:10.1128/JCM.44.4.1274-1282.2006. PMID 16597851. PMC 1448633.
137.^ Waters E, Hohn MJ, Ahel I, et al. (October 2003). "The genome of Nanoarchaeum
equitans: insights into early archaeal evolution and derived parasitism". Proc. Natl. Acad. Sci. U.S.A.
100 (22): 12984–8. doi:10.1073/pnas.1735403100. PMID 14566062. PMC 240731.
138.^ Jahn U, Gallenberger M, Paper W, et al. (March 2008). "Nanoarchaeum equitans and
Ignicoccus hospitalis: new insights into a unique, intimate association of two archaea". J. Bacteriol.
190 (5): 1743–50. doi:10.1128/JB.01731-07. PMID 18165302. PMC 2258681.
139.^ Chaban B, Ng SY, Jarrell KF (February 2006). "Archaeal habitats—from the extreme to the
ordinary". Can. J. Microbiol. 52 (2): 73–116. doi:10.1139/w05-147. PMID 16541146.
140.^ Schink B (June 1997). "Energetics of syntrophic cooperation in methanogenic
degradation". Microbiol. Mol. Biol. Rev. 61 (2): 262–80. PMID 9184013.
141.^ Lange, M; Westermann, P; Ahring, BK (2005). "Archaea in protozoa and metazoa".
Applied Microbiology and Biotechnology 66 (5): 465–474. doi:10.1007/s00253-004-1790-4.
PMID 15630514.
142.^ van Hoek AH, van Alen TA, Sprakel VS, et al. (1 February 2000). "Multiple acquisition of
methanogenic archaeal symbionts by anaerobic ciliates". Mol. Biol. Evol. 17 (2): 251–8.
PMID 10677847.
143.^ Preston, C.M; Wu, K.Y; Molinski, T.F; Delong, E.F (1996). "A psychrophilic crenarchaeon
inhabits a marine sponge: Cenarchaeum symbiosum gen. nov., sp. nov". Proc Natl Acad Sci USA 93
(13): 6241–6. doi:10.1073/pnas.93.13.6241. PMID 8692799.
144.^ Eckburg PB, Bik EM, Bernstein CN, et al. (June 2005). "Diversity of the human intestinal
microbial flora". Science 308 (5728): 1635–8. doi:10.1126/science.1110591. PMID 15831718.
145.^ Samuel BS, Gordon JI (June 2006). "A humanized gnotobiotic mouse model of host-
archaeal-bacterial mutualism". Proc. Natl. Acad. Sci. U.S.A. 103 (26): 10011–6.
doi:10.1073/pnas.0602187103. PMID 16782812.
146.^ Wegley, L; Yu; Breitbart; Casas; Kline; Rohwer (2004). "Coral-associated Archaea" (PDF).
Marine Ecology Progress Series 273: 89–96. doi:10.3354/meps273089.
147.^ Chelius MK, Triplett EW (April 2001). "The Diversity of Archaea and Bacteria in
Association with the Roots of Zea mays L". Microb. Ecol. 41 (3): 252–63.
doi:10.1007/s002480000087. PMID 11391463.
148.^ Simon HM, Dodsworth JA, Goodman RM (October 2000). "Crenarchaeota colonize
terrestrial plant roots". Environ. Microbiol. 2 (5): 495–505. doi:10.1046/j.1462-2920.2000.00131.x.
PMID 11233158.
149.^ Breithaupt H (2001). "The hunt for living gold. The search for organisms in extreme
environments yields useful enzymes for industry". EMBO Rep. 2 (11): 968–71. doi:10.1093/embo-
reports/kve238. PMID 11713183.
150.^ a b Egorova K, Antranikian G (2005). "Industrial relevance of thermophilic Archaea". Curr.
Opin. Microbiol. 8 (6): 649–55. doi:10.1016/j.mib.2005.10.015. PMID 16257257.
151.^ Synowiecki J, Grzybowska B, Zdziebło A (2006). "Sources, properties and suitability of
new thermostable enzymes in food processing". Crit Rev Food Sci Nutr 46 (3): 197–205.
doi:10.1080/10408690590957296. PMID 16527752.
152.^ Jenney FE, Adams MW (January 2008). "The impact of extremophiles on structural
genomics (and vice versa)". Extremophiles 12 (1): 39–50. doi:10.1007/s00792-007-0087-9.
PMID 17563834.
153.^ Schiraldi C, Giuliano M, De Rosa M (2002). "Perspectives on biotechnological
applications of archaea" (PDF). Archaea 1 (2): 75–86. doi:10.1155/2002/436561. PMID 15803645.
PMC 2685559.
154.^ Norris PR, Burton NP, Foulis NA (2000). "Acidophiles in bioreactor mineral processing".
Extremophiles 4 (2): 71–6. doi:10.1007/s007920050139. PMID 10805560.
155.^ O'Connor EM, Shand RF (January 2002). "Halocins and sulfolobicins: the emerging story
of archaeal protein and peptide antibiotics". J. Ind. Microbiol. Biotechnol. 28 (1): 23–31.
doi:10.1038/sj/jim/7000190. PMID 11938468.
156.^ Shand RF; Leyva KJ (2008). "Archaeal Antimicrobials: An Undiscovered Country". In
Blum P (ed.). Archaea: New Models for Prokaryotic Biology. Caister Academic Press. ISBN 978-1-
[edit] Further reading
• Howland, John L. (2000). The Surprising Archaea: Discovering Another Domain of Life.
Oxford: Oxford University Press. ISBN 0-19-511183-4.
• Martinko JM, Madigan MT (2005). Brock Biology of Microorganisms (11th ed.). Englewood
Cliffs, N.J: Prentice Hall. ISBN 0-13-144329-1.
• Garrett RA, Klenk H (2005). Archaea: Evolution, Physiology and Molecular Biology.
WileyBlackwell. ISBN 1-4051-4404-1.
• Cavicchioli R (2007). Archaea: Molecular and Cellular Biology. American Society for
Microbiology. ISBN 1-55581-391-7.
• Blum P (editor) (2008). Archaea: New Models for Prokaryotic Biology. Caister Academic
Press. ISBN 978-1-904455-27-1.
• Lipps G (2008). "Archaeal Plasmids". Plasmids: Current Research and Future Trends .
Caister Academic Press. ISBN 978-1-904455-35-6.
• Sapp, Jan (2009). The New Foundations of Evolution. On the Tree of Life . New York: Oxford
University Press. ISBN 019538850X.
• Schaechter, M (2009). Archaea (Overview) in The Desk Encyclopedia of Microbiology, 2nd
edition. San Diego and London: Elsevier Academic Press. ISBN 13: 978-0-12-374980-2.

[edit] External links

Wikispecies has information related to: Archaea
Wikimedia Commons has media related to: Archaea

Look up Archaea in Wiktionary, the free dictionary.

• Introduction to the Archaea, ecology, systematics and morphology
• Oceans of Archaea – E.F. DeLong, ASM News, 2003
• NCBI taxonomy page on Archaea
• Genera of the domain Archaea – list of Prokaryotic names with Standing in Nomenclature
• Tree of Life illustration showing how Archaea relates to other lifeforms
• Shotgun sequencing finds nanoorganisms – discovery of the ARMAN group of archaea
• Browse any completed archaeal genome at UCSC
• Comparative Analysis of Archaeal Genomes (at DOE's IMG system)

[hide]v · d · eArchaea classification

Domain : Archaea - Bacteria - Eukaryota

Crenarchaeota Acidilobales · Desulfurococcales · Sulfolobales · Thermoproteales

Euryarchaeota Archaeoglobi · Halobacteria · Methanobacteria · Methanococci ·

Methanomicrobia · Methanopyri · Thermococci · Thermoplasmata

Thaumarchaeota Cenarchaeales · Nitrosopumilales

Other Korarchaeota · Nanoarchaeota

[hide]v · d · eExtremophiles

Types Acidophile · Alkaliphile · Capnophile ·

Endolith · Halophile · Hypolith · Lipophile ·
Lithoautotroph · Lithophile · Methanogen ·
Metalotolerant · Oligotroph · Osmophile ·
Piezophile · Polyextremophile · Psychrophile ·
Radioresistant · Thermophile/Hyperthermophile ·
Thermoacidophile · Xerophile
aurantiacus · Deinococcus
radiodurans · Deinococcus-
Bacteria Thermus · Snottite · Thermus
aquaticus · Thermus
thermophilus · Spirochaeta
Notable americana · GFAJ-1
Pyrococcus furiosus ·
Archaea Strain 121 · Pyrolobus fumarii

Paralvinella sulfincola ·
Animalia Pompeii worm · Tardigrada

Related Abiogenic petroleum origin ·

articles Acidithiobacillales · Acidobacteria · Acidophiles in
acid mine drainage · Archaeoglobaceae · Berkeley
Pit · Blood Falls · Crenarchaeota · Grylloblattidae ·
Halobacteria · Halobacterium · Helaeomyia
petrolei · Hydrothermal vent · Methanopyrus ·
Movile Cave · Radioresistance · Radiotrophic
fungus · Rio Tinto · Taq polymerase ·
Thermostability · Thermotogae

Retrieved from ""

Categories: Archaea | Extremophiles | Microbiology


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Temporal range: Cambrian-Recent,[1] 525–0 Ma
Individual organisms from each major vertebrate group. Clockwise, starting from top left:
Fire Salamander, Saltwater Crocodile, Southern Cassowary, Black-and-rufous Giant Elephant
Shrew, Ocean Sunfish

Scientific classification [ e ]
Kingdom: Animalia

Phylum: Chordata

(unranked): Craniata

Cuvier, 1812

Simplified grouping (see text)

• Fishes
• Tetrapods
Vertebrates are members of the subphylum Vertebrata, chordates with backbones and spinal
columns. About 58,000 species of vertebrates have been currently described.[2] Vertebrata is the largest
subphylum of chordates, and contains many familiar groups of large land animals. Vertebrates are the
animals from the groups of jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and
birds. Extant vertebrates range in size from the carp species Paedocypris, at as little as 7.9 mm (0.3 inch), to
the Blue Whale, at up to 33 m (110 ft). Vertebrates make up about 5% of all described animal species; the
rest are invertebrates, which lack backbones.
The vertebrates traditionally include the hagfish, which do not have proper vertebrae, though their
closest living relatives, the lampreys, do have vertebrae.[3] For this reason, the vertebrate subphylum is
sometimes referred to as "Craniata", as all members do possess a cranium.
• 1 Etymology
• 2 Anatomy and morphology
• 3 Evolutionary history
• 4 Classification
• 4.1 Traditional classification
• 4.2 Phylogenetic relationships
• 5 See also
• 6 References
• 7 Bibliography
• 8 External links

[edit] Etymology
The word vertebrate derives from the Latin word vertebratus (Pliny), meaning joint of the spine.[4] It
is closely related to the word vertebra, which refers to any of the bones or segments of the spinal column.[5]

[edit] Anatomy and morphology

All vertebrates are built along the basic chordate body plan: a stiff rod running through the length of
the animal (vertebral column or notochord), with a hollow tube of nervous tissue (the spinal cord) above it
and the gastrointestinal tract below. In all vertebrates the mouth is found at or right below the anterior end of
the animal, while the anus opens to the exterior before the end of the body. The remaining part of the body
continuing aft of the anus forms a tail with vertebrae and spinal cord, but no gut.
The defining characteristic of a vertebrate is the vertebral column, in which the notochord (a stiff rod
of uniform composition) has been replaced by a segmented series of stiffer elements (vertebrae) separated
by mobile joints (intervertebral discs, derived embryonically and evolutionarily from the notochord). However,
a few vertebrates have secondarily lost this anatomy, retaining the notochord into adulthood, as in the
sturgeon and the Latimeria. Jawed vertebrates are typified by paired appendages (fins or legs, which may be
secondarily lost), but this is not part of the definition of vertebrates as a whole.
Fossilized skeleton of Diplodocus, showing an extreme example of the backbone that characterizes
the vertebrates. Exhibited at the Museum für Naturkunde (Museum of Natural Science), Berlin.

[edit] Evolutionary history

Vertebrates originated about 525 million years ago during the Cambrian explosion, which was an
event of massive rise in organism diversity that occurred in the Cambrian period. The earliest known
vertebrate is believed to be the Myllokunmingia.[1] Molecular analysis since 1999 have suggested that the
hagfishes are most closely related to lampreys, and so also are vertebrates. Others consider them a sister
group of vertebrates in the common taxon of Craniata.[3][6] Another early vertebrate is Haikouichthys
ercaicunensis, also from the Chengjiang fauna 524 million years ago. All of these groups lacked a jaw in the
common sense.
The first jawed vertebrates appeared in the Ordovician and became common in the Devonian, often
known as the "Age of Fishes". The two groups of bony fishes, the actinopterygii and sarcopterygii, evolved
and became common. The Devonian also saw the demise of virtually all jawless fishes, save for lampreys
and hagfish, as well as the rise of the first labyrinthodonts, transitional between fish and amphibians. The
Placodermi, a group of fishes that dominated much of the late Silurian and the majority of the Devonian
period, also became extinct at the end of the Devonian.
The reptiles appeared from labyrinthodont stock in the subsequent Carboniferous period. The
anapsid and synapsid reptiles were common during the late Paleozoic, while the diapsids became dominant
during the Mesozoic. The dinosaurs gave rise to the birds in the Jurassic. The demise of the dinosaurs at the
end of the Cretaceous promoted expansion of the mammals, which had developed from the therapsids, a
group of synapsid reptiles, during the Late Triassic Period. The post-dinosaur world have seen great
diversification of bony fishes, frogs, birds and mammals.
Over half of all living vertebrate species (about 32,000 species) are fishes (non-tetrapod craniates), a
diverse set of lineages that inhabit all the world’s aquatic ecosystems, from snow minnows (Cypriniformes) in
Himalayan lakes at elevations over 4,600 meters (15,000 feet) to flatfishes (order Pleuronectiformes) in the
Challenger Deep, the deepest ocean trench at about 11,000 meters (36,000 feet). Fishes of myriad varieties
are the main predators in most of the world’s water bodies, both freshwater and marine. The rest of the
vertebrate species are tetrapods, a single lineage that includes amphibians (frogs, with more than 5,800
species; salamanders, with about 580 species; and caecilians, with about 175 species); mammals (with over
5,400 species); and reptiles and birds (with more than 18,000 species). Tetrapods dominate the megafauna
of most terrestrial environments (including fossorial and arboreal realms) and also include many partially or
fully aquatic groups (e.g., sea snakes, penguins, cetaceans).

[edit] Classification
There are several ways of classifying animals. Evolutionary systematics relies on anatomy,
physiology and evolutionary history, which is determined through similarities in anatomy and, if possible, the
genetics of organisms. Phylogenetic classification is based solely on phylogeny. Evolutionary systematics
gives an overview; phylogenetic systematics gives detail. The two systems are thus complementary rather
than opposed.[7]

[edit] Traditional classification

Conventional classification has living vertebrates grouped into seven classes based on traditional
interpretations of gross anatomical and physiological traits. This classification is the one most commonly
encountered in school textbooks, overviews, non-specialist, and popular works:[8]
• Subphylum Vertebrata
• Class Agnatha (jawless fish)
• Class Chondrichthyes (cartilaginous fishes)
• Class Osteichthyes (bony fishes)
• Class Amphibia (amphibians)
• Class Reptilia (reptiles)
• Class Aves (birds)
• Class Mammalia (mammals)
While this traditional classification is orderly most of the groups are paraphyletic, i.e. do not contain
all descendants of the class's common ancestor. Descendants of the first reptiles do for instance include the
birds and mammals as well as reptiles. Quite a few scientists working with vertebrates use a classification
based purely on phylogeny, organized by their known evolutionary history and sometimes disregarding the
conventional interpretations of their anatomy and physiology. An example based on Janvier (1981, 1997),
Shu et al. (2003), and Benton (2004)[9] is given here:
• Subphylum Vertebrata
• Superclass Agnatha or Cephalaspidomorphi (lampreys and other jawless fishes,
some ancestral to other vertebrates)
• Infraphylum Gnathostomata (vertebrates with jaws)
• Class †Placodermi (extinct armoured fishes)
• Class Chondrichthyes (cartilaginous fishes)
• Class †Acanthodii (extinct spiny "sharks")
• Superclass Osteichthyes (bony fishes)
• Class Actinopterygii (ray-finned bony fishes)
• Class Sarcopterygii (lobe-finned fishes, some ancestral to tetrapods)
• Superclass Tetrapoda (four-limbed vertebrates)
• Class Amphibia (amphibians, some ancestral to the amniotes)
• Class †Synapsida (extinct mammal-like "reptiles", some ancestral to
mammals, sometimes classed with Reptilia)
• Class Mammalia (mammals)
• Class Reptilia (reptiles, some ancestral to birds)
• Class Aves (birds)
Most of the classes listed are not "complete" taxa: the agnathans have given rise to the jawed
vertebrates; the bony fishes have given rise to the land vertebrates; the traditional "amphibians" have given
rise to the reptiles (traditionally including the mammal-like "reptiles"), which in turn have given rise to the
birds and mammals.

[edit] Phylogenetic relationships

In phylogenetic taxonomy, the relationships between animals are not typically divided into ranks, but
illustrated as a nested "family tree" known as a cladogram. Phylogenetic groups are given definitions based
on their relationship to one another, rather than purely on physical traits such as the presence of a backbone.
This nesting pattern is often combined with traditional taxonomy (as above), in a practice known as
evolutionary taxonomy.
The cladogram presented below is based on studies compiled by Philippe Janvier and others for the
Tree of Life Web Project.[10]





(ray-finned fishes)




Dipnoi (lungfishes)



[edit] See also
• Invertebrate
• Marine vertebrates

[edit] References
1. ^ a b Shu et al.; Luo, H-L.; Conway Morris, S.; Zhang, X-L.; Hu, S-X.; Chen, L.; Han, J.; Zhu,
M. et al. (November 4, 1999). "Lower Cambrian vertebrates from south China". Nature 402: 42–46.
2. ^ Jonathan E.M. Baillie, et al. (2004). "A Global Species Assessment". World Conservation
3. ^ a b Kuraku et al.; Hoshiyama, D; Katoh, K; Suga, H; Miyata, T (December 1999).
"Monophyly of Lampreys and Hagfishes Supported by Nuclear DNA–Coded Genes". Journal of
Molecular Evolution 49 (6): 729. doi:10.1007/PL00006595. PMID 10594174.
4. ^ Douglas Harper, Historian. "vertebrate". Online Etymology Dictionary.
5. ^ Douglas Harper, Historian. "vertebra". Online Etymology Dictionary.
6. ^ Nicholls, Henry (10 September 2009). "Mouth to Mouth". Nature 461 (7261): 164–166.
doi:10.1038/461164a. PMID 19741680.
7. ^ Hildebran, M. & Gonslow, G. (2001): Analysis of Vertebrate Structure. 5th edition. John
Wiley & Sons, Inc. New York, page 33: Comment: The problem of naming sister groups
8. ^ Romer, A.S. (1949): The Vertebrate Body. W.B. Saunders, Philadelphia. (2nd ed. 1955; 3rd
ed. 1962; 4th ed. 1970)
9. ^ Benton, Michael J. (2004-11-01). Vertebrate Palaeontology (Third ed.). Blackwell
Publishing. pp. 455 pp.. ISBN 0632056371/978-0632056378.
10.^ Janvier, Philippe. 1997. Vertebrata. Animals with backbones. Version 01 January 1997
(under construction). in The Tree of Life Web Project,

[edit] Bibliography
• Kardong, Kenneth V. (1998). Vertebrates: Comparative Anatomy, Function, Evolution
(second ed.). USA: McGraw-Hill. pp. 747 pp.. ISBN 0-07-115356-X/0-697-28654-1.
• "Vertebrata". Integrated Taxonomic Information System.
Retrieved 6 August 2007.
[edit] External links
Wikispecies has information related to: Vertebrata

• Tree of Life
• Tunicates and not cephalochordates are the closest living relatives of vertebrates
• Vertebrate Pests chapter in United States Environmental Protection Agency and University of
Florida/Institute of Food and Agricultural Sciences National Public Health Pesticide Applicator
Training Manual

[hide]v · d · eExtant phyla of kingdom Animalia by subkingdom

Parazoa Porifera (Calcarea, Demospongiae, Hexactinellida) · Placozoa (Trichoplax)

Mesozoa Orthonectida · Rhombozoa

Eumetazoa Radiata Ctenophora · Cnidaria (Anthozoa, Hydrozoa, Scyphozoa, Cubozoa, Stauroz

Bilateria Protostomia Ecdysozoa Cycloneuralia: Scalidophora (K

Loricifera, Priapulida) · Nematoida (Ne
Panarthropoda: Onychophora

Platyzoa Rotifera · Aca
Spiralia Cycliophora

(Sipuncula, N
Mollusca, An
Entoprocta, P

Deuterostomia Hemichordata · Echinoderma

Craniata (Vertebrata, Myxini
Cephalochordata · Tunicata

Basal/disputed Acoelomorpha (Acoela, Nemertodermatida) · Chae

[hide]v · d · eExtant Chordata classes by subphylum

Kingdom Animalia · Subkingdom Eumetazoa · (unranked) Bilateria · Superphylum Deuterostomia

Urochordata Ascidiacea (Ascidians) · Thaliacea · Appendicularia (Larvaceans) ·

(Tunicates) Sorberacea

(Lancelets) Leptocardii

Myxini (Hagfish) · Hyperoartia (Lampreys) · Chondrichthyes

Craniata (Cartilaginous fish) · Actinopterygii (Ray-finned fish) · Sarcopterygii (Lobe-
finned fish) · Amphibia (Amphibians) · Sauropsida (Reptiles) · Aves (Birds) ·
Mammalia (Mammals)
Retrieved from ""

Categories: Vertebrates


From Wikipedia, the free encyclopedia
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For other uses, see Amphibian (disambiguation).
Temporal range: Late Devonian–present
Strawberry Poison-dart Frog, Oophaga pumilio

Scientific classification [ e ]

Kingdom: Animalia

Phylum: Chordata

Superclass: Tetrapoda

Linnaeus, 1758

Subclasses and Orders

Order Temnospondyli – extinct

Subclass Lepospondyli – extinct
Subclass Lissamphibia
Order Anura
Order Caudata
Order Gymnophiona
Amphibians (class Amphibia, from Amphi- meaning "on both sides" and -bios meaning "life"), such as
frogs, salamanders, and caecilians, are ectothermic (or cold-blooded) animals that metamorphose from a
juvenile water-breathing form, either to an adult air-breathing form, or to a paedomorph that retains some
juvenile characteristics. Mudpuppies, for example, retain juvenile gills in adulthood. The three modern orders
of amphibians are Anura (frogs and toads), Caudata (salamanders and newts), and Gymnophiona
(caecilians, limbless amphibians that resemble snakes), and in total they numbers approximately 6,500
species.[1] Many amphibians lay their eggs in water. Amphibians are superficially similar to reptiles, but
reptiles are amniotes, along with mammals and birds. The study of amphibians is called batrachology.
Amphibians are ecological indicators,[2] and in recent decades there has been a dramatic decline in
amphibian populations around the globe. Many species are now threatened or extinct.
Amphibians evolved in the Devonian Period and were top predators in the Carboniferous and
Permian Periods, but many lineages were wiped out during the Permian–Triassic extinction. One group, the
metoposaurs, remained important predators during the Triassic, but as the world became drier during the
Early Jurassic they died out, leaving a handful of relict temnospondyls like Koolasuchus and the modern
orders of Lissamphibia.
• 1 Etymology
• 2 Evolutionary history
• 3 Taxonomic history
• 3.1 Respiration
• 3.2 Reproduction
• 4 Conservation
• 5 See also
• 6 References
• 7 Further reading
• 8 External links

[edit] Etymology
Amphibian is derived from the Ancient Greek term ἀμφίβιος amphíbios which means both kinds of
life, amphi meaning “both” and bio meaning life. The term was initially used for all kinds of combined natures.
Eventually it was used to refer to animals that live both in the water and on land.[3]

[edit] Evolutionary history

Main article: Labyrinthodontia
See also: List of prehistoric amphibians
The first major groups of amphibians developed in the Devonian Period from fish similar to the
modern coelacanth and lungfish which had evolved multi-jointed leg-like fins that enabled them to crawl
along the sea bottom. These amphibians were as much as one to five meters in length. However,
amphibians never developed the ability to live their entire lives on land, having to return to water to lay their
shell-less eggs.
In the Carboniferous Period, the amphibians moved up in the food chain and began to occupy the
ecological position currently occupied by crocodiles. These amphibians were notable for eating the mega
insects on land and many types of fishes in the water. During the Triassic Period, the better land-adapted
proto-crocodiles began to compete with amphibians, leading to their reduction in size and importance in the

[edit] Taxonomic history

Traditionally, amphibians have included all tetrapod vertebrates that are not amniotes. They are
divided into three subclasses, of which two are only known as extinct subclasses:
• Subclass Labyrinthodontia† (diverse Paleozoic and early Mesozoic group)
• Subclass Lepospondyli† (small Paleozoic group, sometimes included in the
• Subclass Lissamphibia (frogs, toads, salamanders, newts, etc.)
Of these only the last subclass includes recent species.
With the phylogenetic classification Labyrinthodontia has been discarded as it is a paraphyletic group
without unique defining features apart from shared primitive characteristics. Classification varies according to
the preferred phylogeny of the author, whether they use a stem-based or node-based classification.
Generally amphibians are defined as the group that includes the common ancestors of all living amphibians
(frogs, salamanders and caecilians) and all their descendants. This may also include extinct groups like the
temnospondyls (traditionally placed in the subclass “Labyrinthodontia”), and the Lepospondyls. This means
that cladistic nomenclature list a large number of basal Devonian and Carboniferous tetrapod groups,
undoubtedly were “amphibians” in biology, that are formally placed in Amphibia in Linnaean taxonomy, but
not in cladistic taxonomy.
All recent amphibians are included in the subclass Lissamphibia, superorder Salientia, which is
usually considered a clade (which means that it is thought that they evolved from a common ancestor apart
from other extinct groups), although it has also been suggested that salamanders arose separately from a
temnospondyl-like ancestor.[4]
Authorities also disagree on whether Salientia is a Superorder that includes the order Anura, or
whether Anura is a sub-order of the order Salientia. Practical considerations seem to favor using the former
arrangement now. The Lissamphibia, superorder Salientia, are traditionally divided into three orders, but an
extinct salamander-like family, the Albanerpetontidae, is now considered part of the Lissamphibia, besides
the superorder Salientia. Furthermore, Salientia includes all three recent orders plus a single Triassic proto-
frog, Triadobatrachus.
Class Amphibia
• Subclass Lissamphibia
• Family Albanerpetontidae — Jurassic to Miocene (extinct)
• Superorder Salientia
• Genus Triadobatrachus — Triassic (extinct)
• Order Anura (frogs and toads): Jurassic to recent — 5,602 recent species in
48 families
• Order Caudata or Urodela (salamanders, newts): Jurassic to recent — 571
recent species in 9 families
• Order Gymnophiona or Apoda (caecilians): Jurassic to recent — 174 recent
species in 3 families
The actual number of species partly also depends on the taxonomic classification followed, the two
most common classifications being the classification of the website AmphibiaWeb, University of California
(Berkeley) and the classification by herpetologist Darrel Frost and The American Museum of Natural History,
available as the online reference database Amphibian Species of the World.[5] The numbers of species cited
above follow Frost.

[edit] Respiration
The lungs in amphibians are primitive compared to that of the amniotes, possessing few internal
septa, large alveoli and therefore a slow diffusion rate of oxygen into the blood. Ventilation is accomplished
by buccal pumping. However, most amphibians are able to exchange gasses with the water or air via their
skin. To enable sufficient cutaneous respiration, the surface of their highly vascularized skin must remain
moist in order for the oxygen to diffuse at a sufficient rate. Because oxygen concentration in the water
increases at both low temperatures and high flow rates, aquatic amphibians in these situations can rely
primarily on cutaneous respiration, as in the Titicaca water frog or hellbender salamanders. In air, where
oxygen is more concentrated, some small species can rely solely on cutaneous gas exchange, most
famously the plethodontid salamanders which have neither lungs nor gills. Many aquatic salamanders and all
tadpoles have gills in their larval stage, with some (such as the axolotl) retaining gills as aquatic adults.
[edit] Reproduction

Caecilian from the San Antonio zoo

For the purpose of reproduction most amphibians require fresh water. A few (e.g. Fejervarya raja)
can inhabit brackish water and even survive (though not thrive) in seawater, but there are no true marine
amphibians. Several hundred frog species in adaptive radiations (e.g., Eleutherodactylus, the Pacific
Platymantines, the Australo-Papuan microhylids, and many other tropical frogs), however, do not need any
water for breeding in the wild. They reproduce via direct development, an ecological and evolutionary
adaptation that has allowed them to be completely independent from free-standing water. Almost all of these
frogs live in wet tropical rainforests and their eggs hatch directly into miniature versions of the adult, passing
through the tadpole stage within the egg. Reproductive success of many amphibians is dependent not only
on the quantity of rainfall, but the seasonal timing.[6]
Several species have also adapted to arid and semi-arid environments, but most of them still need
water to lay their eggs. Symbiosis with single celled algae that lives in the jelly-like layer of the eggs has
evolved several times. The larvae of frogs (tadpoles or polliwogs) breathe with exterior gills at the start, but
soon a pouch is formed that covers the gills and the front legs. Lungs are also formed quite early to assist in
breathing. Newt larvae have large external gills that gradually disappear and the larvae of newts are quite
similar to the adult form from early age on.
Frogs and toads however have a tadpole stage, which is a totally different organism that is a grazing
algae or ongrowth or filtering plankton until a certain size has been reached, where metamorphosis sets in.
This metamorphosis lasts typically only 24 hours and consists of:
• The disappearance of the gill pouch, making the front legs visible.
• The transformation of the jaws into the big jaws of predatory frogs (most tadpoles are
scraping of algae or are filter feeders)
• The transformation of the digestive system: the long spiral gut of the larva is being replaced
by the typical short gut of a predator.
• An adaptation of the nervous system for stereoscopic vision, locomotion and feeding
• A quick growth and movement of the eyes to higher up the skull and the formation of eyelids.
• Formation of skin glands, thickening of the skin and loss of the lateral line system
• An eardrum is developed to lock the middle ear.
The disappearance of the tail is somewhat later (occurs at higher thyroxin levels) and after the tail
has been resorbed the animals are ready to leave the water. The material of the tail is being used for a quick
growth of the legs. The disappearance of the larval structures is a regulated process called apoptosis.
The transformation of newts when leaving the water is reversible except for the loss of the external
gills. When the animals enter the water again for reproduction changes are driven by prolactin, when they
return to the land phase by thyroxin

[edit] Conservation
Main article: Decline in amphibian populations

The Golden Toad of Monteverde, Costa Rica was among the first casualties of amphibian declines.
Formerly abundant, it was last seen in 1989.
Dramatic declines in amphibian populations, including population crashes and mass localized
extinction, have been noted in the past two decades from locations all over the world, and amphibian
declines are thus perceived as one of the most critical threats to global biodiversity. A number of causes are
believed to be involved, including habitat destruction and modification, over-exploitation, pollution, introduced
species, climate change, endocrine-disrupting pollutants, destruction of the ozone layer (ultraviolet radiation
has shown to be especially damaging to the skin, eyes, and eggs of amphibians), and diseases like
chytridiomycosis. However, many of the causes of amphibian declines are still poorly understood, and are a
topic of ongoing discussion. A global strategy to stem the crisis has been released in the form of the
Amphibian Conservation Action Plan (available at Developed by over 80 leading
experts in the field, this call to action details what would be required to curtail amphibian declines and
extinctions over the next 5 years - and how much this would cost. The Amphibian Specialist Group of the
World Conservation Union (IUCN) is spearheading efforts to implement a comprehensive global strategy for
amphibian conservation. Amphibian Ark is an organization that was formed to implement the ex-situ
conservation recommendations of this plan, and they have been working with zoos and aquaria around the
world encouraging them to create assurance colonies of threatened amphibians. One such project is the
Panama Amphibian Rescue and Conservation Project that built on existing conservation efforts in Panama to
create a country-wide response to the threat of chytridiomycosis rapidly spreading into eastern Panama[7]
On January 21, 2008, Evolutionarily Distinct and Globally Endangered (EDGE), as given by chief
Helen Meredith, identified nature's most endangered species: "The EDGE amphibians are amongst the most
remarkable and unusual species on the planet and yet an alarming 85% of the top 100 are receiving little or
no conservation attention." The top 10 endangered species (in the List of endangered animal species)
include: the Chinese giant salamander, a distant relative of the newt, the tiny Gardiner's Seychelles, the
limbless Sagalla caecilian, South African ghost frogs, lungless Mexican salamanders, the Malagasy rainbow
frog, Chile's Darwin frog (Rhinoderma rufum) and the Betic Midwife Toad.[8][9][10][11]

[edit] See also

• Vegetal rotation
[edit] References
1. ^ Amphibian diversity and life history.
3. ^ "Amphibious definition".
q=amphibious&db=luna. Retrieved 2009-04-07.
4. ^ Carroll, 2007
5. ^ Amphibian Species of the World The online database by Darrel Frost and The American
Museum of Natural History
6. ^ C.Michael Hogan. 2010. Abiotic factor. Encyclopedia of Earth. eds Emily Monosson and C.
Cleveland. National Council for Science and the Environment. Washington DC
7. ^ Panama Amphibian Rescue and Conservation Project
8. ^ Lovell, Jeremy (2008-01-20). "Reuters, Giant newt, tiny frog identified as most at risk". Retrieved 2009-04-07.
9. ^ Sample, Ian (2008-01-20). "Drive to save weird and endangered amphibians". The
Guardian (London). Retrieved
10.^ "/environment, images of the species". London: Guardian. 2008-01-18.
picture=332110244. Retrieved 2009-04-07.
11.^ "/environment, Gallery: the world's strangest amphibians". London: Guardian. 2008-01-18. Retrieved 2009-04-
[edit] Further reading
• Carroll, Robert L. (1988). Vertebrate Paleontology and Evolution. New York: W.H. Freeman
& Co..
• Carroll, Robert L. (2009). The Rise of Amphibians: 365 Million Years of Evolution . Baltimore:
The Johns Hopkins University Press. ISBN 978-0-8018-9140-3.
• Duellman, William E.; Linda Trueb (1994). Biology of Amphibians. Johns Hopkins University
Press. ISBN 978-0801847806.
• Frost, Darrel R.; Taran Grant, Julián Faivovich, Raoul H. Bain, Alexander Haas, Célio F.B.
Haddad, Rafael O. De Sá, Alan Channing, Mark Wilkinson, Stephen C. Donnellan, Christopher J.
Raxworthy, Jonathan A. Campbell, Boris L. Blotto, Paul Moler, Robert C. Drewes, Ronald A.
Nussbaum, John D. Lynch, David M. Green, Ward C. Wheeler (March 2006). "The Amphibian Tree
of Life". Bulletin of the American Museum of Natural History 297: 1–291. doi:10.1206/0003-
• Pounds, J. Alan; Martín R. Bustamante, Luis A. Coloma, Jamie A. Consuegra, Michael P. L.
Fogden, Pru N. Foster, Enrique La Marca, Karen L. Masters, Andrés Merino-Viteri, Robert
Puschendorf, Santiago R. Ron, G. Arturo Sánchez-Azofeifa, Christopher J. Still and Bruce E. Young
(January 2006). "Widespread amphibian extinctions from epidemic disease driven by global
warming". Nature 439 (7073): 161–167. doi:10.1038/nature04246. PMID 16407945.
• San Mauro, Diego; Miguel Vences, Marina Alcobendas, Rafael Zardoya and Axel Meyer
(May 2005). "Initial diversification of living amphibians predated the breakup of Pangaea". American
Naturalist 165 (5): 590–599. doi:10.1086/429523. PMID 15795855.
• San Mauro, Diego (2010). "A multilocus timescale for the origin of extant amphibians".
Molecular Phylogenetics and Evolution 56 (2): 554–561. doi:10.1016/j.ympev.2010.04.019.
PMID 20399871.
• Solomon Berg Martin, Biology
• Stuart, Simon N.; Janice S. Chanson, Neil A. Cox, Bruce E. Young, Ana S. L. Rodrigues,
Debra L. Fischman, Robert W. Waller (December 2004). "Status and trends of amphibian declines
and extinctions worldwide". Science 306 (5702): 1783–1786. doi:10.1126/science.1103538.
PMID 15486254.
• S.N.Stuart, M.Hoffmann, J.S.Chanson, N.A.Cox, R.J.Berridge, P.Ramani, B.E. Young
(editors), Collective work. (September 2008). Threatened Amphibians of the World. Published by
Lynx Edicions, in association with IUCN-The World Conservation Union, Conservation International
and NatureServe.. ISBN 978-84-96553-41-5.
lynx/MON0017-threatened-amphibians-world.html. 776 pages

[edit] External links

Wikispecies has information related to: Amphibia

The Wikibook Dichotomous Key has a page on the topic of

• Save the Frogs!
• Amphibian Specialist Group
• AmphibiaWebEcuador
• Amphibians photos Brazil
• Amphibian Ark
• AmphibiaWeb
• Global Amphibian Assessment
• Amphibians of central Europe
• USGS--Online Guide for the Identification of Amphibians in North America north of Mexico
• General amphibian biology information - Living UnderWorld
• Panama Amphibian Rescue and Conservation Project
• Atlanta Botanical Garden Amphibian Conservation Program
• Amphibian vocalisations on Archival Sound Recordings
• Froglife (British conservation charity)

[hide]v · d · eExtant Chordata classes by subphylum

Kingdom Animalia · Subkingdom Eumetazoa · (unranked) Bilateria · Superphylum Deuterostomia

Urochordata Ascidiacea (Ascidians) · Thaliacea · Appendicularia (Larvaceans) ·

(Tunicates) Sorberacea

Cephalochordata Leptocardii

Myxini (Hagfish) · Hyperoartia (Lampreys) · Chondrichthyes

Craniata (Cartilaginous fish) · Actinopterygii (Ray-finned fish) · Sarcopterygii (Lobe-
finned fish) · Amphibia (Amphibians) · Sauropsida (Reptiles) · Aves (Birds) ·
Mammalia (Mammals)

[hide]v · d · eExtant amphibian positions by subclass

Kingdom Animalia · Phylum Chordata · Subphylum Craniata · Superclass Tetrapoda

Lissamphibia Anura · Caudata (Urodela) · Gymnophiona (Apoda)

Retrieved from ""

Categories: Amphibians

From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is about the aquatic animal. For the porous cleaning tool, see Sponge (material). For
other uses, see Sponge (disambiguation).

Fossil range: Ediacaran–Recent
Scientific classification
Domain: Eukaryota

Kingdom: Animalia

Phylum: Porifera*
Grant in Todd, 1836

Included groups
Sponges are animals of the phylum Porifera (meaning "pore bearer"; pronounced /pɒˈrɪfərə/). Their
bodies consist of jelly-like mesohyl sandwiched between two thin layers of cells. While all animals have
unspecialized cells that can transform into specialized cells, sponges are unique in having some specialized
cells that can transform into other types, often migrating between the main cell layers and the mesohyl in the
process. Sponges do not have nervous, digestive or circulatory systems. Instead, most rely on maintaining a
constant water flow through their bodies to obtain food and oxygen and to remove wastes, and the shapes of
their bodies are adapted to maximize the efficiency of the water flow. All are sessile aquatic animals and,
although there are freshwater species, the great majority are marine (salt water) species, ranging from tidal
zones to depths exceeding 8,800 metres (5.5 mi).
While most of the approximately 5,000–10,000 known species feed on bacteria and other food
particles in the water, some host photosynthesizing micro-organisms as endosymbionts and these alliances
often produce more food and oxygen than they consume. A few species of sponge that live in food-poor
environments have become carnivores that prey mainly on small crustaceans.[1]
Most species use sexual reproduction, releasing sperm cells into the water and meeting ova that in
species are release and others are retained the "mother". The fertilized eggs form larvae which swim off in
search of places to settle. Sponges are known for regenerating from fragments that are broken off, although
this only works if the fragments include the right types of cells. A few species reproduce by budding. When
conditions deteriorate, for example as temperatures drop, many freshwater species and a few marine ones
produce gemmules, "survival pods" of unspecialized cells that remain dormant until conditions improve and
then either form completely new sponges or re-colonize the skeletons of their parents.
The mesohyl functions as an endoskeleton in most sponges, and is the only skeleton in soft sponges
that encrust hard surfaces such as rocks. More commonly the mesohyl is stiffened by mineral spicules, by
spongin fibers or both. Demosponges use spongin, and in many species silica spicules and in some species
calcium carbonate exoskeletons. Demosponges constitute about 90% of all known sponge species, including
all freshwater ones, and have the widest range of habitats. Calcareous sponges, which have calcium
carbonate spicules and in some species calcium carbonate exoskeletons, are restricted to relatively shallow
marine waters where production of calcium carbonate is easiest. The fragile glass sponges, with "scaffolding"
of silica spicules, are restricted to polar regions and the ocean depths where predators are rare. Fossils of all
of these types have been found in rocks dated from 580 million years ago. In addition Archaeocyathids,
whose fossils are common in rocks from 530 to 490 million years ago, are now regarded as a type of sponge.
It is generally thought that the sponge's closest single-celled relatives are choanoflagellates, which
strongly resemble the cells that sponges use to drive their water flow systems and capture most of their food.
It is also generally agreed that sponges do not form a monophyletic group, in other words do not include all
and only the descendants of a common ancestor, because it is thought that Eumetazoa (more complex
animals) are descendants of a sub-group of sponges. However it is uncertain which group of sponges is
closest to Eumetazoa, as both calcareous sponges and a sub-group of demosponges called
Homoscleromorpha have been nominated by different researchers. In addition a study in 2008 suggested
that the earliest animals may have been similar to modern comb jellies.
The few species of demosponge that have entirely soft fibrous skeletons with no hard elements have
been used by humans over thousands of years for several purposes, including as padding and as cleaning
tools. However by the 1950s these had been over-fished so heavily that the industry almost collapsed, and
most sponge-like materials are now synthetic. Sponges and their microscopic endosymbionts are now being
researched as possible sources of medicines for treating a wide range of diseases. Dolphins have been
observed using sponges as tools while foraging.
• 1 Distinguishing features
• 2 Basic structure
• 2.1 Cell types
• 2.2 Glass sponges' syncytia
• 2.3 Water flow and body structures
• 2.4 Skeleton
• 3 Classes
• 4 Vital functions
• 4.1 Movement
• 4.2 Respiration, feeding and excretion
• 4.3 Carnivorous sponges
• 4.4 Endosymbionts
• 4.5 "Immune" system
• 4.6 Reproduction
• 4.6.1 Asexual
• 4.6.2 Sexual
• 4.6.3 Life cycle
• 4.7 Coordination of activities
• 5 Ecology
• 5.1 Habitats
Distinguishing features
Further information: Cnidaria and Ctenophore
Sponges constitute the phylum Porifera, and have been defined as sessile metazoans (multi-celled
animals) that have water intake and outlet openings connected by chambers lined with choanocytes, cells
with whip-like flagella. However, a few carnivorous sponges have lost these water flow systems and the
choanocytes.[2] All known living sponges can remold their bodies, as most types of their cells can move
within their bodies and a few can change from one type to another.[2][3]
Like cnidarians (jellyfish, etc.) and ctenophores (comb jellies), and unlike all other known metazoans,
sponges' bodies consist of a non-living jelly-like mass sandwiched between two main layers of cells.[4][5]
Cnidarians and ctenophores have simple nervous systems, and their cell layers are bound by internal
connections and by being mounted on a basement membrane (thin fibrous mat, also known as "basal
lamina").[5] Sponges have no nervous systems, their middle jelly-like layers have large and varied
populations of cells, and some types of cell in their outer layers may move into the middle layer and change
their functions.[3]
Cnidarians and

Nervous system No Yes, simple

No , except that Yes: inter-cell

Cells in each layer
Homoscleromorpha have basement connections; basement
bound together
membranes.[6] membranes
Number of
cells in middle "jelly" Many Few

Cells in outer
layers can move
Yes No
inwards and change

Basic structure
Cell types Water flow
A sponge's body is hollow and is held in shape by the mesohyl, a
jelly-like substance made mainly of collagen and reinforced by a dense
network of fibers also made of collagen. The inner surface is covered
with choanocytes, cells with cylindrical or conical collars surrounding one
flagellum per choanocyte. The wave-like motion of the whip-like flagella
drives water through the sponge's body. All sponges have ostia,
channels leading to the interior through the mesohyl, and in most
sponges these are controlled by tube-like porocytes that form closable
inlet valves. Pinacocytes, plate-like cells, form a single-layered external
skin over all other parts of the mesohyl that are not covered by

Main cell types of

choanocytes, and the pinacocytes also digest food particles that are too large to enter the ostia,[3][4] while
those at the base of the animal are responsible for anchoring it.[4]
Other types of cell live and move within the mesohyl:[3][4]
• Lophocytes are amoeba-like cells that move slowly through the mesohyl and secrete
collagen fibres.
• Collencytes are another type of collagen-producing cell.
• Rhabdiferous cells secrete polysaccharides that also form part of the mesohyl.
• Oocytes and spermatocytes are reproductive cells.
• Sclerocytes secrete the mineralized spicules ("little spines") that form the skeletons of many
sponges and in some species provide some defense against predators.
• In addition to or instead of sclerocytes, demosponges have spongocytes that secrete a form
of collagen that polymerizes into spongin, a thick fibrous material that stiffens the mesohyl.
• Myocytes ("muscle cells") conduct signals and cause parts of the animal to contract.
• "Grey cells" act as sponges' equivalent of an immune system.
• Archaeocytes (or amoebocytes) are amoeba-like cells that are totipotent, in other words each
is capable of transformation into any other type of cell. They also have important roles in feeding and
in clearing debris that block the ostia.
Glass sponges' syncytia
Glass sponges present a distinctive variation on this basic
plan. Their spicules, which are made of silica, form a scaffolding-like
Spicules framework between whose rods the living tissue is suspended like a
syncitium cobweb that contains most of the cell types.[3] This tissue is a
syncytium that in some ways behaves like many cells that share a
Choanosyncitium single external membrane, and in others like a single cell with
and collar bodies multiple nuclei. The mesohyl is absent or minimal. The syncytium's
showing interior cytoplasm, the soupy fluid that fills the interiors of cells, is organised
into "rivers" that transport nuclei, organelles ("organs" within cells)
Water flow and other substances.[9] Instead of choanocytes they have further
syncytia, known as choanosyncytia, which form bell-shaped
chambers which water enters via perforations. The insides of these
chambers are lined with "collar bodies", each consisting of a collar
and flagellum but without a nucleus of its own. The motion of the
flagella sucks water through passages in the "cobweb" and expels it
via the open ends of the bell-shaped chambers.[3]
Some types of cells have a single nucleus and membrane
each, but are connected to other single-nucleus cells and to the main
syncytium by "bridges" made of cytoplasm. The sclerocytes that build
spicules have multiple nuclei, and in glass sponge larvae they are
The glass sponge
connected to other tissues by cytoplasm bridges; such connections
between sclerocytes have not so far been found in adults, but this
may simply reflect the difficulty of investigating such small-scale features. The bridges are controlled by
"plugged junctions" that
apparently permit some
substances to pass while
blocking others.[9]

Water flow and body

Most sponges work
rather like chimneys: they take
in water at the bottom and eject
it from the osculum ("little
mouth") at the top. Since
ambient currents are faster at
the top, the suction effect that
they produce does some of the
work for free. Sponges can
control the water flow by various
combinations of wholly or
partially closing the osculum

Porifera body structures[10]

and ostia (the intake pores) and varying the beat of the flagella, and may shut it down if there is a lot of sand
or silt in the water.[3]
Although the layers of pinacocytes and choanocytes resemble the epithelia of more complex
animals, they are not bound tightly by cell-to-cell connections or a basal lamina (thin fibrous sheet
underneath). The flexibility of these layers and re-modeling of the mesohyl by lophocytes allow the animals to
adjust their shapes throughout their lives to take maximum advantage of local water currents.[3]
The simplest body structure in sponges is a tube or vase shape known as "asconoid", but this
severely limits the size of the animal. If it is simply scaled up, the ratio of its volume to surface area
increases, because surface increases as the square of length or width while volume increases proportionally
to the cube. The amount of tissue that needs food and oxygen is determined by the volume, but the pumping
capacity that supplies food and oxygen depends on the area covered by choanocytes. Asconoid sponges
seldom exceed 1 millimetre (0.039 in) in diameter.[3]
Some sponges overcome this limitation by adopting the "syconoid" structure, in which the body wall
is pleated. The inner pockets of the pleats are lined with choanocytes, which connect to the outer pockets of
the pleats by ostia. This increase in the number of choanocytes and hence in pumping capacity enables
syconoid sponges to grow up to a few centimeters in diameter. The "leuconid" pattern boosts pumping
capacity further by filling the interior almost completely with mesohyl that contains a network of chambers
lined with choanocytes and connected to each other and to the water intakes and outlet by tubes. Leuconid
sponges grow to over 1 metre (3.3 ft) in diameter, and the fact that growth in any direction increases the
number of choanocyte chambers enables them to take a wider range of forms, for example "encrusting"
sponges whose shapes follow those of the surfaces to which they attach. All freshwater and most shallow-
water marine sponges have leuconid bodies. The networks of water passages in glass sponges are similar to
the leuconid structure.[3] In all three types of structure the cross-section area of the choanocyte-lined regions
is much greater than that of the intake and outlet channels. This makes the flow slower near the choanocytes
and thus makes it easier for them to trap food particles.[3] For example in Leuconia, a small leuconoid
sponge about 10 centimetres (3.9 in) tall
and 1 centimetre (0.39 in) in diameter,
water enters each of more than 80,000
intake canals at 6 cm per minute.
However, because Leuconia has more
than 2 million flagellated chambers and other cells in
whose combined diameter is much
greater than that of the canals, water flow
through chambers slows to 3.6 cm per
hour, making it easy for choanocytes to
capture food. All the water is expelled
through a single osculum at about 8.5 cm
per second, fast enough to carry waste
products some distance away.[11] Seabed / rock
Water flow
In zoology a skeleton is any fairly
rigid structure of an animal, irrespective
of whether it has joints and irrespective of
whether it is biomineralized. The mesohyl
functions as an endoskeleton in most
sponges, and is the only skeleton in soft

Sponge with calcium carbonate skeleton[3]

sponges that encrust hard surfaces such as rocks. More commonly the mesohyl is stiffened by mineral
spicules, by spongin fibers or both. Spicules may be made of silica or calcium carbonate, and vary in shape
from simple rods to three-dimensional "stars" with up to six rays. Spicules are produced by sclerocyte cells,
[3] and may be separate, connected by joints, or fused.[2]
Some sponges also secrete exoskeletons that lie completely outside their organic components. For
example sclerosponges ("hard sponges") have massive calcium carbonate exoskeletons over which the
organic matter forms a thin layer with choanocyte chambers in pits in the mineral. These exoskeletons are
secreted by the pinacocytes that form the animals' skins.[3]

Sponges are divided into classes mainly according to the composition of their skeletons:[4]
Type of Spicules[ Spongin Massive Body
cells[4] 4] fibers[4] exoskeleton[12] form[4]

Calcite Common. Asconoid,
nucleus, single
Calcarea May be individual Never Made of calcite if syconoid or
or large masses present. leuconoid

Mostly Silica
Glass sponges syncytia in all May be individual Never Never Leuconoid
species or fused

Demosponges Single Silica In many In some Leuconoid

nucleus, single species.
external species Made of aragonite if
membrane present.[2][12]

Vital functions

Spongia officinalis, "the kitchen sponge", is dark grey when alive

Although adult sponges are fundamentally sessile animals, some marine and freshwater species can
move across the bottom at speeds of 1–4 millimetres (0.039–0.16 in) per day, as a result of amoeba-like
movements of pinacocytes and other cells. A few species can contract their whole bodies, and many can
close their oscula and ostia.[3]
Respiration, feeding and excretion
Sponges do not have distinct circulatory, respiratory, digestive, and excretory systems – instead the
water flow system supports all these functions. They filter food particles out of the water flowing through
them. Particles larger than 50 micrometers cannot enter the ostia and pinacocytes consume them by
phagocytosis (engulfing and internal digestion). Particles from 0.5 μm to 50 μm are trapped in the ostia,
which taper from the outer to inner ends. These particles are consumed by pinacocytes or by archaeocytes
which partially extrude themselves through the walls of the ostia. Bacteria-sized particles, below
0.5 micrometers, pass through the ostia and are caught and consumed by choanocytes.[3] Since the
smallest particles are by far the most common, choanocytes typically capture 80% of a sponge's food supply.
[12] Archaeocytes transport food packaged in vesicles from cells that directly digest food to those that do not.
At least one species of sponge has internal fibers that function as tracks for use by nutrient-carrying
archaeocytes,[3] and these tracks also move inert objects.[4]
It used to be claimed that glass sponges could live on nutrients dissolved in sea water and were very
averse to silt.[13] However a study in 2007 found no evidence of this and concluded that they extract bacteria
and other micro-organisms from water very efficiently (about 79%) and process suspended sediment grains
to extract such prey.[14] Collar bodies digest food and distribute it wrapped in vesicles that are transported
by dynein "motor" molecules along bundles of microtubules that run throughout the syncytium.[3]
Sponges' cells absorb oxygen by diffusion from the water flow system, into which carbon dioxide and
other soluble waste products such as ammonia also diffuse. Archeocytes remove mineral particles that
threaten to block the ostia, transport them through the mesohyl and generally dump them into the outgoing
water current, although some species incorporate them into their skeletons.[3]
Carnivorous sponges
A few species that live in waters where the supply of food particles is very poor prey on crustaceans
and other small animals. Most belong to the family Cladorhizidae, but a few members of the Guitarridae and
Esperiopsidae are also carnivores.[15] In most cases little is known about how they actually capture prey,
although some species are thought to use either sticky threads or hooked spicules.[15][16] Most carnivorous
sponges live in deep waters, up to 8,840 metres (5.49 mi),[17] and the development of deep-ocean
exploration techniques is expected to lead to the discovery of several more.[3][15] However one species has
been found in Mediterranean caves at depths of 17–23 metres (56–75 ft), alongside the more usual filter
feeding sponges. The cave-dwelling predators capture crustaceans under 1 millimetre (0.039 in) long by
entangling them with fine threads, digest them by enveloping them with further threads over the course of a
few days, and then return to their normal shape; there is no evidence that they use venom.[17]
Most known carnivorous sponges have completely lost the water flow system and choanocytes.
However the genus Chondrocladia uses a highly modified water flow system to inflate balloon-like structures
that are used for capturing prey.[15][18]
Freshwater sponges often host green algae as endosymbionts within archaeocytes and other cells,
and benefit from nutrients produced by the algae. Many marine species host other photosynthesizing
organisms, most commonly cyanobacteria but in some cases dinoflagellates. Symbiotic cyanobacteria may
form a third of the total mass of living tissue in some sponges, and some sponges gain 48% to 80% of their
energy supply from these micro-organisms.[3] In 2008 a University of Stuttgart team reported that spicules
made of silica conduct light into the mesohyl, where the photosynthesizing endosymbionts live.[19] Sponges
that host photosynthesizing organisms are most common in waters with relatively poor supplies of food
particles, and often have leafy shapes that maximize the amount of sunlight they collect.[4]
A recently-discovered carnivorous sponge that lives near hydrothermal vents hosts methane-eating
bacteria, and digests some of them.[4]

"Immune" system
Sponges do not have the complex immune systems of most other animals. However they reject
grafts from other species but accept them from other members of their own species. In a few marine species,
gray cells play the leading role in rejection of foreign material. When invaded, they produce a chemical that
stops movement of other cells in the affected area, thus preventing the intruder from using the sponge's
internal transport systems. If the intrusion persists, the grey cells concentrate in the area and release toxins
that kill all cells in the area. The "immune" system can stay in this activated state for up to three weeks.[4]


The freshwater sponge Spongilla lacustris

Sponges have three asexual methods of reproduction: after fragmentation; by budding; and by
producing gemmules. Fragments of sponges may be detached by currents or waves, and perhaps by
predators. They use the mobility of their pinacocytes and choanocytes and reshaping of the mesohyl to re-
attach themselves to a suitable surface and then rebuild themselves as small but functional sponges over the
course of several days. The same capabilities enable sponges that have been squeezed through a fine cloth
to regenerate.[3] A sponge fragment can only regenerate if it contains both collencytes to produce mesohyl
and archeocytes to produce all the other cell types.[12] A very few species reproduce by budding.[3]
Gemmules are "survival pods" which a few marine sponges and many freshwater species produce
by the thousands when dying and which some, mainly freshwater species, regularly produce in autumn.
Spongocytes make gemmules by wrapping shells of spongin, often reinforced with spicules, round clusters of
archeocytes that are full of nutrients.[3] Freshwater gemmules may also include phytosynthesizing
symbionts.[20] The gemmules then become dormant, and in this state can survive cold, drying out, lack of
oxygen and extreme variations in salinity.[3] Freshwater gemmules often do not revive until the temperature
drops, stays cold for a few months and then reaches a near-"normal" level.[20] When a gemmule germinates,
the archeocytes round the outside of the cluster transform into pinacocytes, a membrane over a pore in the
shell bursts, the cluster of cells slowly emerges, and most of the remaining archeocytes transform into other
cell types needed to make a functioning sponge. Gemmules from the same species but different individuals
can join forces to form one sponge.[3] Some gemmules are retained within the parent sponge, and in spring it
can be difficult to tell whether an old sponge has revived or been "recolonized" by its own gemmules.[20]

Most sponges are hermaphrodites (function as both sexes simultaneously), although sponges have
no gonads (reproductive organs). Sperm are produced by choanocytes or entire choanocyte chambers that
sink into the mesohyl and form spermatic cysts while eggs are formed by transformation of archeocytes, or of
choanocytes in some species. Each egg generally acquires a yolk by consuming "nurse cells". During
spawning, sperm burst out of their cysts and are expelled via the osculum. If they contact another sponge of
the same species, the water flow carries them to choanocytes that engulf them but, instead of digesting them,
metamorphose to an ameboid form and carry the sperm through the mesohyl to eggs, which in most cases
engulf the carrier and its cargo.[3]
A few species release fertilized eggs into the water, but most retain the eggs until they hatch. There
are four types of larvae, but all are balls of cells with an outer layer of cells whose flagellae or cilia enable the
larvae to move. After swimming for a few days the larvae sink and crawl until they find a place to settle. Most
of the cells transform into archeocytes and then into the types appropriate for their locations in a miniature
adult sponge.[3]
Glass sponge embryos start by dividing into separate cells, but once 32 cells have formed they
rapidly transform into larvae that externally are ovoid with a band of cilia round the middle that they use for
movement, but internally have the typical glass sponge structure of spicules with a cobweb-like main
syncitium draped around and between them and choanosyncytia with multiple collar bodies in the center. The
larvae then leave their parents' bodies.[21]

Life cycle
Sponges in temperate regions live for at most a few years, but some tropical species and perhaps
some deep-ocean ones may live for 200 years or more. Some calcified demosponges grow by only
0.2 millimetres (0.0079 in) per year and, if that rate is constant, specimens 1 metre (3.3 ft) wide must be
about 5,000 years old. Some sponges start sexual reproduction when only a few weeks old, while others wait
until they are several years old.[3]

Coordination of activities
Adult sponges lack neurons or any other kind of nervous tissue. However most species have the
ability to perform movements that are coordinated all over their bodies, mainly contractions of the
pinacocytes, squeezing the water channels and thus expelling excess sediment and other substances that
may cause blockages. Some species can contract the osculum independently of the rest of the body.
Sponges may also contract in order to reduce the area that is vulnerable to attack by predators. In cases
where two sponges are fused, for example if there is a large but still unseparated bud, these contraction
waves slowly become coordinated in both of the "Siamese twins". The coordinating mechanism is unknown,
but may involve chemicals similar to neurotransmitters.[22] However glass sponges rapidly transmit electrical
impulses through all parts of the syncytium, and use this to halt the motion of their flagella if the incoming
water contains toxins or excessive sediment.[3] Myocytes are thought to be responsible for closing the
osculum and for transmitting signals between different parts of the body.[4]
Sponges contain genes very similar to those that contain the "recipe" for the post-synaptic density,
an important signal-receiving structure in the neurons of all other animals. However in sponges these genes
are only activated in "flask cells" that appear only in larvae and may provide some sensory capability while
the larvae are swimming. This raises questions about whether flask cells represent the predecessors of true
neurons or are evidence that sponges' ancestors had true neurons but lost them as they adapted to a sessile

Euplectella aspergillum, a glass sponge known as "Venus' Flower Basket"

Sponges are worldwide in their distribution, from the polar regions to the tropics.[12] Most live in
quiet, clear waters, because sediment stirred up by waves or currents would block their pores, making it
difficult for them to feed and breathe.[13] The greatest numbers of sponges are usually found on firm
surfaces such as rocks, but some sponges can attach themselves to soft sediment by means of a root-like
Sponges are more abundant but less diverse in temperate waters than in tropical waters, possibly
because organisms that prey on sponges are more abundant in tropical waters.[25] Glass sponges are the
most common in polar waters and in the depths of temperate and tropical seas, as their very porous
construction enables them to extract food from these resource-poor waters with the minimum of effort.
Demosponges and calcareous sponges are abundant and diverse in shallower non-polar waters.[26]
The different classes of sponge live in different ranges of habitat:
Type of
Water type[4] Depth[4]

less than 100 metres (330

Calcarea Marine Hard

Soft or firm
Glass sponges Marine Deep
Marine, brackish; Inter-tidal to abyssal;[4] a
Demosponges and about 150 freshwater carnivorous demosponge has been Any
species[3] found at 8,840 metres (5.49 mi)[17]

As primary producers
Sponges with photosynthesizing endosymbionts produce up to three times more oxygen than they
consume, as well as more organic matter than they consume. Such contributions to their habits' resources
are significant along Australia 's Great Barrier Reef but relatively minor in the Caribbean.[12]

Holes made by clionaid sponge (producing the trace Entobia) after the death of a modern bivalve
shell of species Mercenaria mercenaria, from North Carolina

Close-up of the sponge boring Entobia in a modern oyster valve. Note the chambers which are
connected by short tunnels.
Many sponges shed spicules, forming a dense carpet several meters deep that keeps away
echinoderms which would otherwise prey on the sponges.[12] They also produce toxins that prevent other
sessile organisms such as bryozoans or sea squirts from growing on or near them, making sponges very
effective competitors for living space.
A few species, such as the Caribbean fire sponge Tedania ignis, cause a severe rash in humans who
handle them.[3] Turtles and some fish feed mainly on sponges. It is often said that sponges produce
chemical defenses against such predators.[3] However an experiment showed that there is no relationship
between the toxicity of chemicals produced by sponges and how they taste to fish, which would diminish the
usefulness of chemical defenses as deterrents. Predation by fish may even help to spread sponges by
detaching fragments.[4]
Glass sponges produce no toxic chemicals, and live in very deep water where predators are rare.[13]

Sponge flies, also known as spongilla-flies (Neuroptera, Sisyridae), are specialist predators of
freshwater sponges. The female lays her eggs on vegetation overhanging water. The larvae hatch and drop
into the water where they seek out sponges to feed on. They use their elongated mouthparts to pierce the
sponge and suck the fluids within. The larvae of some species cling to the surface of the sponge while others
take refuge in the sponge's internal cavities. The fully grown larvae leave the water and spin a cocoon in
which to pupate.[27]

The Caribbean chicken-liver sponge Chondrilla nucula secretes toxins that kill coral polyps, allowing
the sponges to grow over the coral skeletons.[3] Others, especially in the family Clionaidae, use corrosive
substances secreted by their archeocytes to tunnel into rocks, corals and the shells of dead molluscs.[3]
Sponges may remove up to 1 metre (3.3 ft) per year from reefs, creating visible notches just below low-tide
Caribbean sponges of the genus Aplysina suffer from Aplysina red band syndrome. This causes
Aplysina to develop one or more rust-colored bands, sometimes with adjacent bands of necrotic tissue
(dead). These lesions may completely encircle branches of the sponge. The disease appears to be
contagious (spread by physical contact). The rust-colored bands are caused by a cyanobacterium, but it is
unknown whether this organism actually causes the disease.[28]

Collaboration with other organisms

In addition to hosting photosynthesizing endosymbionts,[3] sponges are noted for their wide range of
collaborations with other organisms. The relatively large encrusting sponge Lissodendoryx colombiensis is
most common on rocky surfaces, but has extended its range into seagrass meadows by letting itself be
surrounded or overgrown by seagrass sponges, which are distasteful to the local starfish and therefore
protect Lissodendoryx against them; in return the seagrass sponges get higher positions away from the sea-
floor sediment.[29]
Shrimps of the genus Synalpheus form colonies in sponges, and each shrimp species inhabits a
different sponge species, making Synalpheus one of the most diverse crustacean genera.[30]
Evolutionary history
Fossil record
Traces of the chemical 24-
isopropylcholestane have been found in rocks formed
1,800 million years ago.[31] This is a stable derivative
of 24-isopropylcholesterol, which is thought to be
produced by demosponges but not by eumetazoans 7
("true animals", i.e. cnidarians and bilaterians). Since
choanoflagellates are thought to be animals' closest 1: Gap
single-celled relatives, a team of scientists examined 2: Central cavity
the biochemistry and genes of one choanoflagellate 3 Internal wall 4: Pore
species. They concluded that this species could not (all walls have pores)
produce 24-isopropylcholesterol but that investigation 5 Septum 6 Outer wall
of a wider range of choanoflagellates would be
7 Holdfast
necessary in order to prove that the fossil 24-
isopropylcholestane could only have been produced
by demosponges.[32]
Silica spicules like those of demosponges
Fossil sponge have been reported from Nevada in rocks dated
Raphidonema around 750 million years ago.[33] Well-preserved
faringdonense from fossil sponges from about 580 million years ago in
Cretaceous rocks in the Ediacaran period have been found in the

Doushantuo Formation. These fossils, which include spicules, pinacocytes, porocytes, archeocytes,
sclerocytes and the internal cavity, have been classified as demosponges. Fossils of glass sponges have
been found from around 540 million years ago in rocks in Australia, China and Mongolia.[34] Calcium
carbonate spicules of calcareous sponges have been found in Early Cambrian rocks from about 530 to 523
million years ago in Australia. Other probable demosponges have been found in the Early Cambrian
Chengjiang fauna, from 525 to 520 million years ago.[35] Freshwater sponges appear to be much younger,
as the earliest known fossils date from the Mid-Eocene period about 48 to 40 million years ago.[34] Although
about 90% of modern sponges are demosponges, fossilized remains of this type are less common than
those of other types because their skeletons are composed of relatively soft spongin that does not fossilize
Archaeocyathids, which some classify as a type of coralline sponge, are common in the Cambrian
period from about 530 million years ago, but apparently died out by the end of the Cambrian 490 million
years ago.[35]

Family tree
A choanoflagellate
In the 1990s sponges were widely regarded as a monophyletic group, in other words all of them
descended from a common ancestor that was itself a sponge, and as the "sister-group" to all other
metazoans (multi-celled animals), which themselves form a monophyletic group. On the other hand some
1990s analyses also revived the idea that animals' nearest evolutionary relatives are choanoflagellates,
single-celled organisms very similar to sponges' choanocytes – which would imply that most Metazoa evolved
from very sponge-like ancestors and therefore that sponges may not be monophyletic, as the same sponge-
like ancestors may have given rise both to modern sponges and to non-sponge members of Metazoa.[37]




(jellyfish, etc.)
Analyses since 2001 have concluded that Eumetazoa (more complex than sponges) are more
closely related to particular groups of sponges than to the rest of the sponges. Such conclusions imply that
sponges are not monophyletic, because the last common ancestor of all sponges would also be a direct
ancestor of the Eumetazoa, which are not sponges. A study in 2001 based on comparisons of ribosome DNA
concluded that the most fundamental division within sponges was between glass sponges and the rest, and
that Eumetazoa are more closely related to Calcareous sponges, those with calcium carbonate spicules, than
to other types of sponge.[37] In 2007 one analysis based on comparisons of RNA and another based mainly
on comparison of spicules concluded that demosponges and glass sponges are more closely related to each
other than either is to calcareous sponges, which in turn are more closely related to Eumetazoa.[34][39]
Other anatomical and biochemical evidence links the Eumetazoa with Homoscleromorpha, a sub-
group of demosponges. A comparison in 2007 of nuclear DNA, excluding glass sponges and comb jellies,
concluded that: Homoscleromorpha are most closely related to Eumetazoa; calcareous sponges are the next
closest; the other demosponges are evolutionary "aunts" of these groups; and the chancelloriids, bag-like
animals whose fossils are found in Cambrian rocks, may be sponges.[38] The sperm of Homoscleromorpha
share with those of Eumetazoa features that those of other sponges lack. In both Homoscleromorpha and
Eumetazoa layers of cells are bound together by attachment to a carpet-like basal membrane composed
mainly of "type IV" collagen, a form of collagen not found in other sponges – although the spongin fibers that
reinforce the mesohyl of all demosponges is similar to "type IV" collagen.[6]
A comb jelly
The analyses described above concluded that sponges are closest to the ancestors of all Metazoa, in
other words of all multi-celled animals including both sponges and more complex groups. However, another
comparison in 2008 of 150 genes in each of 21 genera, ranging from fungi to humans but including only two
species of sponge, suggested that comb jellies (ctenophora) are the most basal lineage of the Metazoa
included in the sample. If this is correct, either modern comb jellies developed their complex structures
independently of other Metazoa, or sponges' ancestors were more complex and all known sponges are
drastically simplified forms. The study recommended further analyses using a wider range of sponges and
other simple Metazoa such as Placozoa.[40] The results of such an analysis, published in 2009, suggest that
a return to the previous view may be warranted. 'Family trees' constructed using a combination of all
available data - morphological, developmental and molecular - concluded that the sponges are in fact a
monophyletic group, and with the cnidarians form the sister group to the bilaterians.[41]
Archaeocyathids are very common fossils in rocks from the Early Cambrian about 530 to 520 million
years ago but are not found after the Late Cambrian. It has been
suggested that they were produced by: sponges; cnidarians; algae;
foraminiferans; a completely separate phylum of animals,
Archaeocyatha; or even a completely separate kingdom of life, labelled
Archaeata or Inferibionta. Since the 1990s archaeocyathids have been
regarded as a distinctive group of sponges.[42]
It is difficult to fit chancelloriids into classifications of sponges or
more complex animals. An analysis in 1996 concluded that they were
closely related to sponges on the grounds that the detailed structure of
chancellorid sclerites ("armor plates") is similar to that of fibers of

Halkieriid sclerite
spongin, a collagen protein, in modern keratose (horny) demosponges such as Darwinella.[44] However
another analysis in 2002 concluded that chancelloriids are not sponges and may be intermediate between
sponges and more complex animals, among other reasons because their skins were thicker and more tightly-
connected than those of sponges.[45] In 2008 a detailed analysis of chancelloriids' sclerites concluded that
they were very similar to those of halkieriids, mobile bilaterian animals that looked like slugs in chain mail and
whose fossils are found in rocks from the very Early Cambrian to the Mid Cambrian. If this is correct, it would
create a dilemma, as it is extremely unlikely that totally unrelated organisms could have developed such
similar sclerites independently, but the huge difference in the structures of their bodies makes it hard to see
how they could be closely related.[43]

Levels in the Linnean taxonomy.

For a long time sponges were assigned to a separate subkingdom, Parazoa ("beside the animals"),
separate from the Eumetazoa which formed the rest of the kingdom Animalia.[42] They are now classified as
a phylum within Animalia, and divided into classes mainly according to the composition of their skeletons:[2]
• Hexactinellida (glass sponges) have silicate spicules, the largest of which have six rays and
may be individual or fused.[2] The main components of their bodies are syncytia in which large
numbers of cell share a single external membrane.[12]
• Calcarea have skeletons made of calcite, a form of calcium carbonate, which may form
separate spicules or large masses. All the cells have a single nucleus and membrane.[12]
• Most Demospongiae have silicate spicules or spongin fibers or both within their soft tissues.
However a few also have massive external skeletons made of aragonite, another form of calcium
carbonate.[2][12] All the cells have a single nucleus and membrane.[12]
• Archeocyatha are known only as fossils from the Cambrian period.[42]
In the 1970s sponges with massive calcium carbonate skeletons were assigned to a separate class,
Sclerospongiae, otherwise known as "coralline sponges".[46] However in the 1980s it was found that these
were all members of either the Calcarea or the Demospongiae.[47]
So far scientific publications have identified about 9,000 poriferan species,[12] of which: about 400
are glass sponges; about 500 are calcareous species; and the rest are demosponges.[3] However some
types of habitat, such as vertical rock and cave walls and galleries in rock and coral boulders, have been
investigated very little, even in shallow seas.[12]

By dolphins
A report in 1997 described use of sponges as a tool by bottlenose dolphins in Shark Bay. A dolphin
will attach a marine sponge to its rostrum, which is presumably then used to protect it when searching for
food in the sandy sea bottom.[48] The behaviour, known as sponging, has only been observed in this bay,
and is almost exclusively shown by females. A study in 2005 concluded that mothers teach the behaviour to
their daughters, and that all the sponge-users are closely related, suggesting that it is a fairly recent

By humans

Display of natural sponges for sale on Kalymnos in Greece

Main article: Sponge (tool)
The calcium carbonate or silica spicules of most sponge genera make them too rough for most uses,
but two genera, Hippospongia and Spongia, have soft, entirely fibrous skeletons. Early Europeans used soft
sponges for many purposes, including padding for helmets, portable drinking utensils and municipal water
filters. Until the invention of synthetic sponges, they were used as cleaning tools, applicators for paints and
ceramic glazes and discreet contraceptives. However by the mid-20th century, over-fishing brought both the
animals and the industry close to extinction.[50] See also sponge diving.
Many objects with sponge-like textures are now made of substances not derived from poriferans.
Synthetic sponges include personal and household cleaning tools, breast implants,[51] and contraceptive
sponges.[52] Typical materials used are cellulose foam, polyurethane foam, and less frequently, silicone
The luffa "sponge", also spelled loofah, which is commonly sold for use in the kitchen or the shower,
is not derived from an animal but from the fibrous "skeleton" of a gourd (Cucurbitaceae).[53]

Antibiotic compounds
Sponges have medicinal potential due to the presence in sponges themselves or their microbial
symbionts of chemicals that may be used to control viruses, bacteria, tumors and fungi.[54][55]

See also
• Aaptos kanuux
• Randolph Kirkpatrick
• Sponge reef
• Sponge Reef Project
1. ^ J. Vacelet and E. Duport. Prey capture and digestion in the carnivorous sponge
Asbestopluma hypogea (Porifera: Demospongiae).
2. ^ a b c d e f g Hooper, J. N. A., Van Soest, R. W. M., and Debrenne, F. (2002). "Phylum
Porifera Grant, 1836". In Hooper, J. N. A., and Van Soest, R. W. M.. Systema Porifera: A Guide to
the Classification of Sponges. New York: Kluwer Academic/Plenum. pp. 9–14. ISBN 9780306472602.
Retrieved 2008-11-06.
3. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al Ruppert, E. E., Fox, R.
S., and Barnes, R. D. (2004). Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 76–97.
ISBN 0030259827.
4. ^ a b c d e f g h i j k l m n o p q r s t Bergquist, P. R., (1998). "Porifera". In Anderson, D.T.,.
Invertebrate Zoology. Oxford University Press. pp. 10–27. ISBN 0195513681.
5. ^ a b c Hinde, R. T., (1998). "The Cnidaria and Ctenophora". In Anderson, D.T.,. Invertebrate
Zoology. Oxford University Press. pp. 28–57. ISBN 0195513681.
6. ^ a b Exposito, J-Y., Cluzel, C., Garrone, R., and Lethias, C. (2002). "Evolution of collagens".
The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology = 268
(3): 302–316. doi:10.1002/ar.10162. PMID 12382326.
7. ^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). Invertebrate Zoology (7 ed.). Brooks /
Cole. p. 82. ISBN 0030259827.
8. ^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). Invertebrate Zoology (7 ed.). Brooks /
Cole. pp. 83. ISBN 0030259827. Fig. 5-7
9. ^ a b Leys, S. P. (2003). "The significance of syncytial tissues for the position of the
Hexactinellida in the Metazoa". Integrative and Comparative Biology 43 (1): 19–27.
10.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). Invertebrate Zoology (7 ed.). Brooks /
Cole. p. 78. ISBN 0030259827.
11.^ C. Hickman, C .P. (Jr.), Roberts, L. S., and Larson, A. (2001). Integrated Principles of
Zoology (11 ed.). New York: McGraw-Hill. p. 247. ISBN 9780072909616.
12.^ a b c d e f g h i j k l m n o Bergquist, P. R. (2001). "Porifera (Sponges)". Encyclopedia of Life
Sciences. John Wiley & Sons, Ltd.. doi:10.1038/npg.els.0001582.
13.^ a b c Krautter, M. (1998). "Ecology of siliceous sponges: Application to the environmental
interpretation of the Upper Jurassic sponge facies (Oxfordian) from Spain" (PDF). Cuadernos de
Geología Ibérica 24: 223–239. ISSN 0378-102X. Retrieved 2008-
11-10. [dead link]
14.^ Yahel, G., Whitney, F., Reiswig, H. M., Eerkes-Medrano, D. I., and Leys, S.P. (April 20,
2007). "In situ feeding and metabolism of glass sponges (Hexactinellida, Porifera) studied in a deep
temperate fjord with a remotely operated submersible". Limnology and oceanography 52 (1): 428–
440. doi:10.4319/lo.2007.52.1.0428. ISSN 0024-3590.
aModele=afficheN&cpsidt=18876170. Retrieved 2008-11-02.
15.^ a b c d Vacelet, J. (2008). "A new genus of carnivorous sponges (Porifera: Poecilosclerida,
Cladorhizidae) from the deep N-E Pacific, and remarks on the genus Neocladia" (PDF). Zootaxa
1752: 57–65. Retrieved 2008-10-31.
16.^ Watling, L. (2007). "Predation on copepods by an Alaskan cladorhizid sponge". Journal of
the Marine Biological Association of the UK 87: 1721–1726 doi=10.1017/S0025315407058560.
17.^ a b c Vacelet, J., and Boury-Esnault, N. (January 1995). "Carnivorous sponges". Nature
373: 333–335. doi:10.1038/373333a0.
18.^ Vacelet, J., and Kelly, M. (September 2008). "New species from the deep Pacific suggest
that carnivorous sponges date back to the Early Jurassic". Nature Precedings.
doi:10.1038/npre.2008.2327.1 (inactive 2010-03-19). Retrieved 2008-10-31.
19.^ News report at Brümmer, F., Pfannkuchen, M., Baltz, A., Hauser, T., and Thiel, V. (2008).
"Light inside sponges". Journal of Experimental Marine Biology and Ecology 367 (2): 61–64.
doi:10.1016/j.jembe.2008.06.036. . News report at "Nature's 'fibre optics' experts". BBC. 2008-11-10. Retrieved 2008-11-10. .
20.^ a b c Smith, D. G., and Pennak, R. W. (2001). Pennak's Freshwater Invertebrates of the
United States: Porifera to Crustacea (4 ed.). John Wiley and Sons. pp. 47–50. ISBN 0471358371. Retrieved
21.^ Leys, S., Cheung, E., and Boury-Esnault, N. (2006). "Embryogenesis in the glass sponge
Oopsacas minuta: Formation of syncytia by fusion of blastomeres". Integrative and Comparative
Biology 46 (2): 104–117. doi:10.1093/icb/icj016.
22.^ Nickel, M. (December 2004). "Kinetics and rhythm of body contractions in the sponge
Tethya wilhelma (Porifera: Demospongiae)". Journal of Experimental Biology 207 (Pt 26): 4515–4524.
doi:10.1242/jeb.01289. PMID 15579547.
23.^ Sakarya, O., Armstrong, K. A., Adamska, M., Adamski, M., Wang, I., et al. (2007). "A Post-
Synaptic Scaffold at the Origin of the Animal Kingdom". PLoS ONE 2 (6): e506.
doi:10.1371/journal.pone.0000506. PMID 17551586.
24.^ Weaver, J. C., Aizenberg, J., and Fantner, G .E. et al. (April 2007). "Hierarchical assembly
of the siliceous skeletal lattice of the hexactinellid sponge Euplectella aspergillum". Journal of
Structural Biology 158 (1): 93–106. doi:10.1016/j.jsb.2006.10.027. PMID 17175169.
25.^ Ruzicka, R., and Gleason, D. F. (2008). "Latitudinal variation in spongivorous fishes and
the effectiveness of sponge chemical defenses" (PDF). Oecologia 154 (4): 785–794.
doi:10.1007/s00442-007-0874-0. PMID 17960425.
home/Gleason/Ruzicka&Gleasonfulltext.pdf. Retrieved 2008-11-11.
26.^ Gage, J. D., and Tyler, P. A. (1996). Deep-sea Biology: A Natural History of Organisms at
the Deep-Sea Floor. Cambridge University Press. pp. 91–93. ISBN 0521336651.
id=kHZO5igKhsAC&dq=sponge+porifera+geographical+distribution+temperature. Retrieved 2008-
27.^ Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual
Animals, Greenwood Publishing Group.
28.^ Olson, J. B., Gochfeld, D. J., and Slattery, M. (July 2006). " Aplysina red band syndrome: a
new threat to Caribbean sponges". Diseases of aquatic organisms 71 (2): 163–8.
doi:10.3354/dao071163. PMID 16956064. News report at New disease threatens sponges (Practical
29.^ Wulff, J. L (2008). "Collaboration among sponge species increases sponge diversity and
abundance in a seagrass meadow". Marine Ecology 29: 193–204. doi:10.1111/j.1439-
30.^ Duffy, J. E. (1996). "Species boundaries, specialization, and the radiation of sponge-
dwelling alpheid shrimp". Biological Journal of the Linnean Society 58 (3): 307–324.
svr=www. Retrieved 2008-11-11.
31.^ Nichols, S., and Wörheide, G. (2005). "Sponges: New Views of Old Animals". Integrative
and Comparative Biology 45 (2): 333–334. doi:10.1093/icb/45.2.333.
32.^ Kodner, R. B., Summons, R. E., Pearson, A., King, N., and Knoll, A. H. (July 2008). "Sterols
in a unicellular relative of the metazoans". Proceedings of the National Academy of Sciences 105
(29): 9897–9902. doi:10.1073/pnas.0803975105. PMID 18632573.
33.^ Reitner, J., and Wörheide, G. (2002). "Non-Lithistid Fossil Demospongiae – Origins of their
Palaeobiodiversity and Highlights in History of Preservation". In Hooper, J. N. A., and Van Soest, R.
W. M. (PDF). Systema Porifera: A Guide to the Classification of Sponges . New York: Kluwer
Academic Plenum.
Retrieved 2008-11-04.
34.^ a b c Müller, W. E. G., Li, J., Schröder, H. C., Qiao, L., and Wang, X. (2007). "The unique
skeleton of siliceous sponges (Porifera; Hexactinellida and Demospongiae) that evolved first from the
Urmetazoa during the Proterozoic: a review". Biogeosciences 4: 219–232. doi:10.5194/bg-4-219-
35.^ a b Li, C-W., Chen, J-Y., and Hua, T-E. (February 1998). "Precambrian Sponges with
Cellular Structures". Science 279 (5352): 879–882. doi:10.1126/science.279.5352.879.
PMID 9452391.
36.^ "Demospongia". University of California Museum of Paleontology. Retrieved 2888-11-27.
37.^ a b c Borchiellini, C., Manuel, M., Alivon, E., Boury-Esnault, N., Vacelet J., and Le Parco, Y.
(2001). "Sponge paraphyly and the origin of Metazoa". Journal of Evolutionary Biology 14 (1): 171–
179. doi:10.1046/j.1420-9101.2001.00244.x.
38.^ a b Sperling, E.A.; Pisani, D. and Peterson, K.J. (2007). "Poriferan paraphyly and its
implications for Precambrian paleobiology" (PDF). Journal of the Geological Society of London 286:
355–368. doi:10.1144/SP286.25.,%20Pisani%20and
%20Peterson.pdf. Retrieved 2008-11-04.
39.^ Medina, M., Collins, A. G., Silberman, J. D., and Sogin, M. L. (August 2001). "Evaluating
hypotheses of basal animal phylogeny using complete sequences of large and small subunit rRNA".
Proceedings of the National Academy of Sciences 98 (17): 9707–9712. doi:10.1073/pnas.171316998.
PMID 11504944.
40.^ Dunn, C. W., Hejnol, A., et al. (2008). "Broad phylogenomic sampling improves resolution
of the animal tree of life". Nature 452 (7188): 745–9. doi:10.1038/nature06614. PMID 18322464.
41.^ Schierwater, B.; Eitel, M.; Jakob, W.; Osigus, J.; Hadrys, H.; Dellaporta, L.; Kolokotronis,
O.; Desalle, R.; Eitel M, Jakob W, Osigus H-J, Hadrys H, et al. (Jan 2009). "Concatenated Analysis
Sheds Light on Early Metazoan Evolution and Fuels a Modern “Urmetazoon” Hypothesis" (Free full
text). PLoS Biology 7 (1): e20. doi:10.1371/journal.pbio.1000020. ISSN 1544-9173. PMID 19175291.
PMC 2631068. edit
42.^ a b c Rowland, S. M. (2001). "Archaeocyatha: A history of phylogenetic interpretation".
Journal of Paleontology 75: 1065–1078. doi:10.1666/0022-3360(2001)075<1065:AAHOPI>2.0.CO;2.;col1.
Retrieved 2008-11-05.
43.^ a b Porter, S. M (2008). "Skeletal microstructure indicates Chancelloriids and Halkieriids are
closely related". Palaeontology 51 (4): 865–879. doi:10.1111/j.1475-4983.2008.00792.x.
44.^ Butterfield, N. J., and C. J. Nicholas (1996). "Burgess Shale-type preservation of both non-
mineralizing and "shelly" Cambrian organisms from the Mackenzie Mountains, northwestern
Canada". Journal of Paleontology 70: 893–899.
45.^ Janussen, D., Steiner, M., and Zhu, M-Y. (July 2002). "New Well-preserved Scleritomes of
Chancelloridae from the Early Cambrian Yuanshan Formation (Chengjiang, China) and the Middle
Cambrian Wheeler Shale (Utah, USA) and paleobiological implications". Journal of Paleontology 76
(4): 596–606. doi:10.1666/0022-3360(2002)076<0596:NWPSOC>2.0.CO;2. Retrieved 2008-08-04. Free full
text without images at Janussen, Dorte (2002). "(as above)". Journal of Paleontology.;col1.
Retrieved 2008-08-04.
46.^ Hartman, W. D., and Goreau, T. F. (1970). "Jamaican coralline sponges: Their morphology,
ecology and fossil relatives". Symposium of the Zoological Society of London 25: 205–243. (cited by
47.^ J. Vacelet (1985). "Coralline sponges and the evolution of the Porifera". In Conway Morris,
S., George, J. D., Gibson, R., and Platt, H. M.. The Origins and Relationships of Lower Invertebrates .
Oxford University Press. pp. 1–13. ISBN 019857181.
48.^ Smolker, R. A., et al. (1997). "Sponge-carrying by Indian Ocean bottlenose dolphins:
Possible tool-use by a delphinid". Ethology 103: 454–465.
49.^ Krutzen M, Mann J, Heithaus M.R., Connor R. C, Bejder L, Sherwin W.B. (2005). "Cultural
transmission of tool use in bottlenose dolphins". Proceedings of the National Academy of Sciences
102 (25): 8939–8943. doi:10.1073/pnas.0500232102. PMID 15947077. . News report at Dolphin
Moms Teach Daughters to Use Tools, publisher National Geographic).
50.^ McClenachan, L. (2008). "Social conflict, Over-fishing and Disease in the Florida Sponge
Fishery, 1849-1939". In Starkey, D. J. Holm, P., and Barnard, M.. Oceans Past: Management
Insights from the History of Marine Animal Populations. Earthscan. pp. 25–27. ISBN 1844075273. Retrieved 2008-11-12.
51.^ Jacobson, N. (2000). Cleavage. Rutgers University Press. p. 62. ISBN 0813527155. Retrieved 2008-11-
52.^ "Sponges". Cervical Barrier Advancement Society. 2004. Retrieved 2006-09-17.
53.^ Porterfield, W. M. (July 1955). "Loofah — The sponge gourd". Economic Botany 9 (3): 211–
223. doi:10.1007/BF02859814.
54.^ Imhoff, J. F., and Stöhr, R. (2003). "Sponge-Associated Bacteria". In Müller, W. E. G..
Sponges (Porifera): Porifera. Springer. pp. 43–44. ISBN 354000968X.
55.^ Teeyapant, R., Woerdenbag, H. J., Kreis, P., Hacker, J., Wray, V., Witte, L., and Proksch P.
(1993). "Antibiotic and cytotoxic activity of brominated compounds from the marine sponge Verongia
aerophoba". Zeitschrift für Naturforschung. C, Journal of biosciences 48: 939–45.

Further reading
• Bergquist, Patricia R. (1978). Sponges. London: Hutchinson. ISBN 0091318203. .
• Hickman, C., Jr.; Roberts, L. & Larson, A. (2003). Animal Diversity (3rd ed.). New York:
McGraw-Hill. ISBN 0072349034. .
• Template:Pll icon, Collection Jurassic fossils sponges
External links
Wikimedia Commons has media related to: Sponge

Wikispecies has information related to: Porifera

The Wikibook Dichotomous Key has a page on the topic of

• Water flow and feeding in the phylum Porifera (sponges) - Flash animations of sponge body
structures, water flow and feeding
• Bioerosion website at The College of Wooster
• Carsten's Spongepage, Information on the ecology and the biotechnological potential of
sponges and their associated bacteria.
• History of Tarpon Springs, Florida sponge industry
• Nature's 'fibre optics' experts
• The Sponge Reef Project
• Queensland Museum FAQ about sponges
• Sponge Guide from Queensland Museum, John Hooper
• Sponge Guide for Britain and Ireland, Bernard Picton, Christine Morrow & Rob van Soest
• World Porifera database, the world list of extant sponges, includes a searchable database.
[hide]v · d · eExtant phyla of kingdom Animalia by subkingdom

Parazoa Porifera (Calcarea, Demospongiae, Hexactinellida) · Placozoa (Trichoplax)

Mesozoa Orthonectida · Rhombozoa

Eumetazoa Radiata Ctenophora · Cnidaria (Anthozoa, Hydrozoa, Scyphozoa, Cubozoa, Stauroz

Bilateria Protostomia Cycloneuralia: Scalidophora (K

Loricifera, Priapulida) · Nematoida (Ne
Ecdysozoa Nematomorpha)
Panarthropoda: Onychophora

Spiralia Platyzoa Platy

Rotifera · Aca

(Sipuncula, N
Mollusca, An
Entoprocta, P

Hemichordata · Echinoderma
Craniata (Vertebrata, Myxini
Cephalochordata · Tunicata

Basal/disputed Acoelomorpha (Acoela, Nemertodermatida) · Chae

Retrieved from ""

Categories: Poriferans

From Wikipedia, the free encyclopedia

Jump to: navigation, search

Temporal range: 580–0 Ma
g Ediacaran–Recent
Pacific sea nettles, Chrysaora fuscescens

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Hatschek, 1888

Anthozoa—corals and sea anemones

Cubozoa—box jellyfish, sea wasps
Hydrozoa—hydroids, hydra-like animals
Scyphozoa—true jellyfish
Staurozoa—stalked jellyfish
Unranked, may not be
Cnidaria (pronounced /naɪˈdɛəriə/ with a silent c) is a phylum containing over 9,000 species of
animals found exclusively in aquatic and mostly marine environments. Their distinguishing feature is
cnidocytes, specialized cells that they use mainly for capturing prey. Their bodies consist of mesoglea, a non-
living jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. They
have two basic body forms: swimming medusae and sessile polyps, both of which are radially symmetrical
with mouths surrounded by tentacles that bear cnidocytes. Both forms have a single orifice and body cavity
that are used for digestion and respiration. Many cnidarian species produce colonies that are single
organisms composed of medusa-like or polyp-like zooids, or both. Cnidarians' activities are coordinated by a
decentralized nerve net and simple receptors. Several free-swimming Cubozoa and Scyphozoa possess
balance-sensing statocysts, and some have simple eyes. Not all cnidarians reproduce sexually. Many have
complex lifecycles with asexual polyp stages and sexual medusae, but some omit either the polyp or the
medusa stage.
Cnidarians were for a long time grouped with Ctenophores in the phylum Coelenterata, but
increasing awareness of their differences caused them to be placed in separate phyla. Cnidarians are
classified into four main groups: sessile Anthozoa (sea anemones, corals, and sea pens (sea anemones are
not sessile but only move 3-4 inces an hour); swimming Scyphozoa (jellyfish); Cubozoa (box jellies); and
Hydrozoa, a diverse group that includes all the freshwater cnidarians as well as many marine forms, and has
both sessile members such as Hydra and colonial swimmers such as the Portuguese Man o' War. Staurozoa
have recently been recognised as a class in their own right rather than a sub-group of Scyphozoa, and there
is debate about whether Myxozoa and Polypodiozoa are cnidarians or closer to bilaterians (more complex
Most cnidarians prey on organisms ranging in size from plankton to animals several times larger than
themselves, but many obtain much of their nutrition from endosymbiotic algae, and a few are parasites. Many
are preyed upon by other animals including starfish, sea slugs, fish and turtles. Coral reefs, whose polyps are
rich in endosymbiotic algae, support some of the world's most productive ecosystems, and protect vegetation
in tidal zones and on shorelines from strong currents and tides. While corals are almost entirely restricted to
warm, shallow marine waters, other cnidarians live in the depths, in polar seas and in freshwater.
Fossil cnidarians have been found in rocks formed about 580 million years ago, and other fossils
show that corals may have been present shortly before 490 million years ago and diversified a few million
years later. Fossils of cnidarians that do not build mineralized structures are very rare. Scientists currently
think that cnidarians, ctenophores and bilaterians are more closely related to calcareous sponges than these
are to other sponges, and that anthozoans are the evolutionary "aunts" or "sisters" of other cnidarians, and
the most closely related to bilaterians. Recent analyses have concluded that cnidarians, although considered
more "primitive" than bilaterians, have a wider range of genes.
Jellyfish stings killed several hundred people in the 20th century, and cubozoans are particularly
dangerous. On the other hand, some large jellyfish are considered a delicacy in eastern and southern Asia.
Coral reefs have long been economically important as providers of fishing grounds, protectors of shore
buildings against currents and tides, and more recently as centers of tourism. However, they are vulnerable
to over-fishing, mining for construction materials, pollution, and damage caused by tourism.
• 1 Classification
• 2 Ecology
• 3 History
• 4 Distinguishing features
• 5 Description
• 5.1 Main cell layers
• 5.2 Cnidocytes
• 5.3 Basic body forms
• 5.4 Colonial forms
• 5.5 Skeletons
• 5.6 Locomotion
• 5.7 Nervous system and senses
• 5.8 Feeding and excretion
• 5.9 Respiration
• 5.10 Regeneration
• 6 Reproduction
• 6.1 Sexual
• 6.2 Asexual
• 7 Evolutionary history
• 7.1 Fossil record
[edit] Classification
Cnidarians were for a long time grouped with Ctenophores in the phylum Coelenterata, but
increasing awareness of their differences caused them to be placed in separate phyla. Cnidarians are
classified into four main groups: sessile Anthozoa (sea anemones, corals, sea pens); swimming Scyphozoa
(jellyfish); Cubozoa (box jellies); and Hydrozoa, a diverse group that includes all the freshwater cnidarians as
well as many marine forms, and has both sessile members such as Hydra and colonial swimmers such as
the Portuguese Man o' War. Staurozoa have recently been recognised as a class in their own right rather
than a sub-group of Scyphozoa, and there is debate about whether Myxozoa and Polypodiozoa are
cnidarians or closer to bilaterians.
Modern cnidarians are generally classified into four classes:[4]
Hydrozoa Scyphozoa Cubozoa Anthozoa

Number of
2,700 200 20 6,000

Hydra, Box
Examples Jellyfish anemones, corals,
siphonophores jellies
sea pens

Cells found in
No Yes Yes Yes
No Yes Yes Yes
in exodermis

Yes, except for

Medusa In some
Stauromedusae if they Yes No
phase in life cycle species
are scyphozoans

Number of
medusae produced Many Many One
per polyp
Stauromedusae, small sessile cnidarians with stalks and no medusa stage, have traditionally been
classified as members of the Scyphozoa, but recent research suggests they should be regarded as a
separate class, Staurozoa.[5]
The Myxozoa, microscopic parasites, were first classified as protozoans,[6] but recently as heavily
modified cnidarians, and more closely related to Hydrozoa and Scyphozoa than to Anthozoa.[7] However
other recent research suggests that Polypodium hydriforme, a parasite within the egg cells of sturgeon, is
closely related to the Myxozoa and that both Polypodium and the Myxozoa are intermediate between
cnidarians and bilaterian animals.[8]
Some researchers classify the extinct conulariids as cnidarians, while others propose that they form
a completely separate phylum.[9]
[edit] Ecology

Coral reefs support rich ecosystems

Many cnidarians are limited to shallow waters because they depend on endosymbiotic algae for
much of their nutrients. The life cycles of most have polyp stages, which are limited to locations that offer
stable substrates. Nevertheless major cnidarian groups contain species that have escaped these limitations.
Hydrozoans have a worldwide range: some, such as Hydra, live in freshwater; Obelia appears in the coastal
waters of all the oceans; and Liriope can form large shoals near the surface in mid-ocean. Among
anthozoans, a few scleractinian corals, sea pens and sea fans live in deep, cold waters, and some sea
anemones inhabit polar seabeds while others live near hydrothermal vents over 10 kilometres (6.2 mi) below
sea-level. Reef-building corals are limited to tropical seas between 30°N and 30°S with a maximum depth of
46 metres (151 ft), temperatures between 20°C and 28°C, high salinity and low carbon dioxide levels.
Stauromedusae, although usually classified as jellyfish, are stalked, sessile animals that live in cool to Arctic
waters.[10] Cnidarians range in size from Hydra, 5–20 millimetres (0.20–0.79 in) long,[11] to the Lion's mane
jellyfish, which may exceed 2 metres (6.6 ft) in diameter and 75 metres (246 ft) in length.[12]
Prey of cnidarians ranges from plankton to animals several times larger than themselves.[10][13]
Some cnidarians are parasites, mainly on jellyfish but a few are major pests of fish.[10] Others obtain most of
their nourishment from endosymbiotic algae or dissolved nutrients.[4] Predators of cnidarians include: sea
slugs, which can incorporate nematocysts into their own bodies for self-defense;[14] starfish, notably the
crown of thorns starfish, which can devastate corals;[10] butterfly fish and parrot fish, which eat corals;[15]
and marine turtles, which eat jellyfish.[12] Some sea anemones and jellyfish have a symbiotic relationship
with some fish; for example clown fish live among the tentacles of sea anemones, and each partner protects
the other against predators.[10]
Coral reefs form some of the world's most productive ecosystems. Common coral reef cnidarians
include both Anthozoans (hard corals, octocorals, anemones) and Hydrozoans (fire corals, lace corals) The
endosymbiotic algae of many cnidarian species are very effective primary producers, in other words
converters of inorganic chemicals into organic ones that other organisms can use, and their coral hosts use
these organic chemicals very efficiently. In addition reefs provide complex and varied habitats that support a
wide range of other organisms.[16] "Fringing" reefs just below low-tide level also have a mutually beneficial
relationship with mangrove forests at high-tide level and sea grass meadows in between: the reefs protect
the mangroves and seagrass from strong currents and waves that would damage them or erode the
sediments in which they are rooted, while the mangroves and seagrass protect the coral from large influxes
of silt, fresh water and pollutants. This additional level of variety in the environment is beneficial to many
types of coral reef animals, which for example may feed in the sea grass and use the reefs for protection or
[edit] History
Fossil cnidarians have been found in rocks formed about 580 million years ago, and other fossils
show that corals may have been present shortly before 490 million years ago and diversified a few million
years later. Fossils of cnidarians that do not build mineralized structures are very rare. Scientists currently
think that cnidarians, ctenophores and bilaterians are more closely related to calcareous sponges than these
are to other sponges, and that anthozoans are the evolutionary "aunts" or "sisters" of other cnidarians, and
the most closely related to bilaterians. Recent analyses have concluded that cnidarians, although considered
more "primitive" than bilaterians, have a wider range of genes.

[edit] Distinguishing features

Further information: Sponge, Ctenophore, and Bilateria
Cnidarians form an animal phylum that is more complex than sponges, about as complex as
ctenophores (comb jellies), and less complex than bilaterians, which include almost all other animals.
However, both cnidarians and ctenophores are more complex than sponges as they have: cells bound by
inter-cell connections and carpet-like basement membranes; muscles; nervous systems; and some have
sensory organs. Cnidarians are distinguished from all other animals by having cnidocytes that fire like
harpoons and are used mainly to capture prey but also as anchors in some species.[4]
Like sponges and ctenophores, cnidarians have two main layers of cells that sandwich a middle layer
of jelly-like material, which is called the mesoglea in cnidarians; more complex animals have three main cell
layers and no intermediate jelly-like layer. Hence, cnidarians and ctenophores have traditionally been
labelled diploblastic, along with sponges.[4][18] However, both cnidarians and ctenophores have a type of
muscle that, in more complex animals, arises from the middle cell layer.[19] As a result some recent text
books classify ctenophores as triploblastic,[20] and it has been suggested that cnidarians evolved from
triploblastic ancestors.[19]
Sponges[21][22] Cnidarians[4][18] Bilateria[4]

Cnidocytes No Yes No

Colloblasts No Yes No

and circulatory No Yes

Number of Two[4] or
Two, with jelly-like layer between them Three
main cell layers Three[19][20]

Cells in No, except that

each layer bound Homoscleromorpha have Yes: inter-cell connections; basement membranes
together basement membranes.[23]

No Yes
Number of
cells in middle Many Few
"jelly" layer

Cells in
outer layers can (Not
Yes No
move inwards and applicable)
change functions

Nervous Simple to
No Yes, simple
system complex

Mostly Mostly Mostly

Muscles None
epitheliomuscular myoepithelial myocytes

[edit] Description
[edit] Main cell layers
Cnidaria are diploblastic animals, in other words they have two main cell layers, while more complex
animals are triploblasts having three main layers. The two main cell layers of cnidarians form epithelia that
are mostly one cell thick, and are attached to a fibrous basement membrane, which they secrete. They also
secrete the jelly-like mesoglea that separates the layers. The layer that faces outwards, known as the
ectoderm ("outside skin"), generally contains the following types of cells:[4]
• Epitheliomuscular cells whose bodies form part of the epithelium but whose bases extend to
form muscle fibers in parallel rows.[24] The fibers of the outward-facing cell layer generally run at
right angles to the fibers of the inward-facing one. In Anthozoa (anemones, corals, etc.) and
Scyphozoa (jellyfish), the mesoglea also contains some muscle cells.[18]
• Cnidocytes, the harpoon-like "nettle cells" that give the phylum Cnidaria its name. These
appear between or sometimes on top of the muscle cells.[4]
• Nerve cells. Sensory cells appear between or sometimes on top of the muscle cells,[4] and
communicate via synapses (gaps across which chemical signals flow) with motor nerve cells, which
lie mostly between the bases of the muscle cells.[18]
• Interstitial cells, which are unspecialized and can replace lost or damaged cells by
transforming into the appropriate types. These are found between the bases of muscle cells.[4]
In addition to epitheliomuscular, nerve and interstitial cells, the inward-facing gastroderm ("stomach
skin") contains gland cells that secrete digestive enzymes. In some species it also contains low
concentrations of cnidocytes, which are used to subdue prey that is still struggling.[4][18]
The mesoglea contains small numbers of amoeba-like cells,[18] and muscle cells in some species.[4]
However the number of middle-layer cells and types are much lower than in sponges.[18]
[edit] Cnidocytes

A hydra's nematocyst, before firing.

"trigger" cilium[18]
Firing sequence of the cnida in a hydra's nematocyst[18]
Operculum (lid)
"Finger" that turns inside out
/ / / Barbs
Victim's skin
Victim's tissues
These "nettle cells" function as harpoons, since their payloads remain connected to the bodies of the
cells by threads. Three types of cnidocytes are known:[4][18]
• Nematocysts inject venom into prey, and usually have barbs to keep them embedded in the
victims. Most species have nematocysts.[4]
• Spirocysts do not penetrate the victim or inject venom, but entangle it by means of small
sticky hairs on the thread. Only members of the class Anthozoa (sea anemones and corals) have
• Ptychocysts are not used for prey capture — instead the threads of discharged ptychocysts
are used for building protective tubes in which their owners live. Ptychocysts are found only in the
order Cerianthria, tube anemones.[18]
The main components of a cnidocyte are:[4][18]
• A cilium (fine hair) which projects above the surface and acts as a trigger. Spirocysts do not
have cilia.
• A tough capsule, the cnida, which houses the thread, its payload and a mixture of chemicals
which may include venom or adhesives or both. ("cnida" is derived from the Greek word κνίδη, which
means "nettle"[25])
• A tube-like extension of the wall of the cnida that points into the cnida, like the finger of a
rubber glove pushed inwards. When a cnidocyte fires, the finger pops out. If the cell is a venomous
nematocyte, the "finger"'s tip reveals a set of barbs that anchor it in the prey.
• The thread, which is an extension of the "finger" and coils round it until the cnidocyte fires.
The thread is usually hollow and delivers chemicals from the cnida to the target.
• An operculum (lid) over the end of the cnida. The lid may be a single hinged flap or three
flaps arranged like slices of pie.
• The cell body which produces all the other parts.
It is difficult to study the firing mechanisms of cnidocytes as these structures are small but very
complex. At least four hypotheses have been proposed:[4]
• Rapid contraction of fibers round the cnida may increase its internal pressure.
• The thread may be like a coiled spring that extends rapidly when released.
• In the case of Chironex (the "sea wasp"), chemical changes in the cnida's contents may
cause them to expand rapidly by polymerization.
• Chemical changes in the liquid in the cnida make it a much more concentrated solution, so
that osmotic pressure forces water in very rapidly to dilute it. This mechanism has been observed in
nematocysts of the class Hydrozoa, sometimes producing pressures as high as 140 atmospheres,
similar to that of scuba air tanks, and fully extending the thread in as little as 2 milliseconds
(0.002 second).[18]
Cnidocytes can only fire once, and about 25% of a hydra's nematocysts are lost from its tentacles
when capturing a brine shrimp. Used cnidocytes have to be replaced, which takes about 48 hours. To
minimise wasteful firing, two types of stimulus are generally required to trigger cnidocytes: their cilia detect
contact, and nearby sensory cells "smell" chemicals in the water. This combination prevents them from firing
at distant or non-living objects. Groups of cnidocytes are usually connected by nerves and, if one fires, the
rest of the group requires a weaker minimum stimulus than the cells that fire first.[4][18]
[edit] Basic body forms


Gastroderm (Endoderm) Oral end of actinodiscus polyp, with close-up of the

Mesoglea Adult cnidarians appear as either swimming medusae or
sessile polyps. Both are radially symmetrical, like a wheel and a
Digestive cavity tube respectively. Since these animals have no heads, their
ends are described as "oral" (nearest the mouth) and "aboral"
(furthest from the mouth). Most have fringes of tentacles
equipped with cnidocytes around their edges, and medusae
generally have an inner ring of tentacles around the mouth. The
mesoglea of polyps is usually thin and often soft, but that of
medusae is usually thick and springy, so that it returns to its

Medusa (left) and polyp (right)[18]

original shape after muscles around the edge have contracted to squeeze water out, enabling medusae to
swim by a sort of jet propulsion.[18]

[edit] Colonial forms

Tree-like polyp colony[18]

Cnidaria produce a variety of colonial forms, each of which is one organism but consists of polyp-like
zooids. The simplest is a connecting tunnel that runs over the substrate (rock or seabed) and from which
single zooids sprout. In some cases the tunnels form visible webs, and in others they are enclosed in a fleshy
mat. More complex forms are also based on connecting tunnels but produce "tree-like" groups of zooids. The
"trees" may be formed either by a central zooid that functions as a "trunk" with later zooids growing to the
sides as "branches", or in a zig-zag shape as a succession of zooids, each of which grows to full size and
then produces a single bud at an angle to itself. In many cases the connecting tunnels and the "stems" are
covered in periderm, a protective layer of chitin.[18] Some colonial forms have other specialized types of
zooid, for example, to pump water through their tunnels.[10]
Siphonophores form complex colonies that consist of: an upside-down polyp that forms a central
stem with a gas-filled float at the top; one or more sets of medusa-like zooids that provide propulsion; leaf-
like bracts that give some protection to other parts; sets of tentacles that bear nematocytes that capture prey;
other tentacles that act as sensors; near the base of each set of tentacles, a polyp-like zooid that acts as a
stomach for the colony; medusa-like zooids that serve as gonads. Although some of these zooids resemble
polyps or medusae in shape, they lack features that are not relevant to their specific functions, for example
the swimming "medusae" have no digestive, sensory or reproductive cells. The best-known siphonophore is
the Portuguese Man o' War (Physalia physalis).[10][26][27]

[edit] Skeletons
In medusae the only supporting structure is the mesoglea. Hydra and most sea anemones close their
mouths when they are not feeding, and the water in the digestive cavity then acts as a hydrostatic skeleton,
rather like a water-filled balloon. Other polyps such as Tubularia use columns of water-filled cells for support.
Sea pens stiffen the mesoglea with calcium carbonate spicules and tough fibrous proteins, rather like
In some colonial polyps a chitinous periderm gives support and some protection to the connecting
sections and to the lower parts of individual polyps. Stony corals secrete massive calcium carbonate
exoskeletons. A few polyps collect materials such as sand grains and shell fragments, which they attach to
their outsides. Some colonial sea anemones stiffen the mesoglea with sediment particles.[18]
[edit] Locomotion

Chrysaora quinquecirrha ("sea nettle") swimming

Medusae swim by a form of jet propulsion: muscles, especially inside the rim of the bell, squeeze
water out of the cavity inside the bell, and the springiness of the mesoglea powers the recovery stroke. Since
the tissue layers are very thin, they provide too little power to swim against currents and just enough to
control movement within currents.[18]
Hydras and some sea anemones can move slowly over rocks and sea or stream beds by various
means: creeping like snails, crawling like inchworms, or by somersaulting. A few can swim clumsily by
waggling their bases.[18]
[edit] Nervous system and senses
Cnidaria have no brains or even central nervous systems. Instead they have decentralized nerve
nets consisting of : sensory neurons that generate signals in response to various types of stimulus, such as
odors; motor neurons that tell muscles to contract; all connected by "cobwebs" of intermediate neurons. As
well as forming the "signal cables", intermediate neurons also form ganglia that act as local coordination
centers. The cilia of the cnidocytes detect physical contact. Nerves inform cnidocytes when odors from prey
or attackers are detected and when neighbouring cnidocytes fire. Most of the communications between nerve
cells are via chemical synapses, small gaps across which chemicals flow. As this process is too slow to
ensure that the muscles round the rim of a medusa's bell contract simultaneously in swimming the neurons
which control this communicate by much faster electrical signals across gap junctions.[18]
Medusae and complex swimming colonies such as siphonophores and chondrophores sense tilt and
acceleration by means of statocysts, chambers lined with hairs which detect the movements of internal
mineral grains called statoliths. If the body tilts in the wrong direction, the animal rights itself by increasing the
strength of the swimming movements on the side that is too low. They also have ocelli ("little eyes"), which
can detect the direction from which light is coming. Box jellies have camera eyes, although these probably do
not form images, and their lenses simply produce a clearer indication of the direction from which light is

[edit] Feeding and excretion

Cnidarians feed in several ways: predation, absorbing dissolved organic chemicals, filtering food
particles out of the water, and obtaining nutrients from symbiotic algae within their cells. Most obtain the
majority of their food from predation but some, including the corals Hetroxenia and Leptogorgia, depend
almost completely on their endosymbionts and on absorbing dissolved nutrients.[4] Cnidaria give their
symbiotic algae carbon dioxide, some nutrients and a place in the sun.[18]
Predatory species use their cnidocytes to poison or entangle prey, and those with venomous
nematocysts may start digestion by injecting digestive enzymes. The "smell" of fluids from wounded prey
makes the tentacles fold inwards and wipe the prey off into the mouth. In medusae the tentacles round the
edge of the bell are often short and most of the prey capture is done by "oral arms", which are extensions of
the edge of the mouth and are often frilled and sometimes branched to increase their surface area. Medusae
often trap prey or suspended food particles by swimming upwards, spreading their tentacles and oral arms
and then sinking. In species for which suspended food particles are important, the tentacles and oral arms
often have rows of cilia whose beating creates currents that flow towards the mouth, and some produce nets
of mucus to trap particles.[4]
Once the food is in the digestive cavity, gland cells in the gastroderm release enzymes that reduce
the prey to slurry, usually within a few hours. This circulates through the digestive cavity and, in colonial
cnidarians, through the connecting tunnels, so that gastroderm cells can absorb the nutrients. Absorption
may take a few hours, and digestion within the cells may take a few days. The circulation of nutrients is
driven by water currents produced by cilia in the gastroderm or by muscular movements or both, so that
nutrients reach all parts of the digestive cavity.[18] Nutrients reach the outer cell layer by diffusion or, for
animals or zooids such as medusae which have thick mesogleas, are transported by mobile cells in the
Indigestible remains of prey are expelled through the mouth. The main waste product of cells' internal
processes is ammonia, which is removed by the external and internal water currents.[18]
[edit] Respiration
There are no respiratory organs, and both cell layers absorb oxygen from and expel carbon dioxide
into the surrounding water. When the water in the digestive cavity becomes stale it must be replaced, and
nutrients that have not been absorbed will be expelled with it. Some Anthozoa have ciliated grooves on their
tentacles, allowing them to pump water out of and into the digestive cavity without opening the mouth. This
improves respiration after feeding and allows these animals, which use the cavity as a hydrostatic skeleton,
to control the water pressure in the cavity without expelling undigested food.[4]
Cnidaria that carry photosynthetic symbionts may have the opposite problem, an excess of oxygen,
which may prove toxic. The animals produce large quantities of antioxidants to neutralize the excess oxygen.

[edit] Regeneration
All cnidarians can regenerate, allowing them to recover from injury and to reproduce asexually.
Medusae have limited ability to regenerate, but polyps can do so from small pieces or even collections of
separated cells. This enables corals to recover even after apparently being destroyed by predators.[4]
[edit] Reproduction
[edit] Sexual
In the Cnidaria sexual reproduction often involves a complex life
cycle with both polyp and medusa stages. For example in Scyphozoa
(jellyfish) and Cubozoa (box jellies) a larva swims until it finds a good site,
and then becomes a polyp. This grows normally but then absorbs its
tentacles and splits horizontally into a series of disks that become juvenile
medusae, a process called strobilation. The juveniles swim off and slowly
grow to maturity, while the polyp re-grows and may continue strobilating
periodically. The adults have gonads in the gastroderm, and these release
ova and sperm into the water in the breeding season.[4][18]
Shortened forms of this life cycle are common, for example some
oceanic scyphozoans omit the polyp stage completely, and cubozoan 12
polyps produce only one medusa. Hydrozoa have a variety of life cycles.
Some have no polyp stages and some (e.g. hydra) have no medusae. In 13
some species the medusae remain attached to the polyp and are 14
responsible for sexual reproduction; in extreme cases these reproductive
zooids may not look much like medusae. Anthozoa have no medusa
stage at all and the polyps are responsible for sexual reproduction.[4]
Spawning is generally driven by environmental factors such as
changes in the water temperature, and their release is triggered by
lighting conditions such as sunrise, sunset or the phase of the moon.
Many species of Cnidaria may spawn simultaneously in the same location, so that there are too many ova
and sperm for predators to eat more than a tiny percentage — one famous example is the Great Barrier Reef,
where at least 110 corals and a few non-cnidarian invertebrates produce enough to turn the water cloudy.
These mass spawnings may produce hybrids, some of which can settle and form polyps, but it is not known
how long these can survive. In some species the ova release chemicals that attract sperm of the same
The fertilized eggs develop into larvae by dividing until there are enough cells to form a hollow
sphere (blastula) and then a depression forms at one end (gastrulation) and eventually become the digestive
cavity. However in cnidarians the depression forms at the end further from the yolk (at the animal pole), while
in bilaterians it forms at the other end (vegetal pole).[18] The larvae, called planulae, swim or crawl by means
of cilia.[4] They are cigar-shaped but slightly broader at the "front" end, which is the aboral, vegetal-pole end
and eventually attaches to a substrate if the species has a polyp stage.[18]
Anthozoan larvae either have large yolks or are capable of feeding on plankton, and some already
have endosymbiotic algae that help to feed them. Since the parents are immobile, these feeding capabilities
extend the larvae's range and avoid overcrowding of sites. Scyphozoan and hydrozoan larvae have little yolk
and most lack endosymbiotic algae, and therefore have to settle quickly and metamorphose into polyps.
Instead these species rely on their medusae to extend their ranges.[18]

[edit] Asexual
All known cnidaria can reproduce asexually by various means, in addition to regenerating after being
fragmented. Hydrozoan polyps only bud, while the medusae of some hydrozoans can divide down the
middle. Scyphozoan polyps can both bud and split down the middle. In addition to both of these methods,
Anthozoa can split horizontally just above the base.[4][18]
[edit] Evolutionary history
[edit] Fossil record

The fossil coral Cladocora from Pliocene rocks in Cyprus

The earliest widely accepted animal fossils are rather modern-looking cnidarians, possibly from
around 580 million years ago, although fossils from the Doushantuo Formation can only be dated
approximately.[28] The identification of some of these as embryos of animals has been contested, but other
fossils from these rocks strongly resemble tubes and other mineralized structures made by corals.[29] Their
presence implies that the cnidarian and bilaterian lineages had already diverged.[30] Although the Ediacaran
fossil Charnia used to be classified as a jellyfish or sea pen,[31] more recent study of growth patterns in
Charnia and modern cnidarians has cast doubt on this hypothesis,[32][33] and there are now no bona-fide
cnidarian body fossils in the Ediacaran. Few fossils of cnidarians without mineralized skeletons are known
from more recent rocks, except in lagerstätten that preserved soft-bodied animals.[34]
A few mineralized fossils that resemble corals have been found in rocks from the Cambrian period,
and corals diversified in the Early Ordovician.[34] These corals, which were wiped out in the Permian-
Triassic extinction about 251 million years ago,[34] did not dominate reef construction since sponges and
algae also played a major part.[35] During the Mesozoic era rudist bivalves were the main reef-builders, but
they were wiped out in the Cretaceous-Tertiary extinction 65 million years ago,[36] and since then the main
reef-builders have been scleractinian corals.[34]

[edit] Family tree

Further information: Phylogeny


(comb jellies)
(sea anemones
and corals)
(Hydra, siphonophores,

(box jellies)


(jellyfish, excluding




Other Bilateria
(more complex)
Family tree of Cnidaria and the origins of animals[2][37][38]
It is difficult to reconstruct the early stages in the evolutionary "family tree" of animals using only
morphology (their shapes and structures), because the large differences between Porifera (sponges),
Cnidaria plus Ctenophora (comb jellies), Placozoa and Bilateria (all the more complex animals) make
comparisons difficult. Hence reconstructions now rely largely or entirely on molecular phylogenetics, which
groups organisms according to similarities and differences in their biochemistry, usually in their DNA or RNA.
It is now generally thought that the Calcarea (sponges with calcium carbonate spicules) are more
closely related to Cnidaria, Ctenophora (comb jellies) and Bilateria (all the more complex animals) than they
are to the other groups of sponges.[37][40][41] In 1866 it was proposed that Cnidaria and Ctenophora were
more closely related to each other than to Bilateria and formed a group called Coelenterata ("hollow guts"),
because Cnidaria and Ctenophora both rely on the flow of water in and out of a single cavity for feeding,
excretion and respiration. In 1881 it was proposed that Ctenophora and Bilateria were more closely related to
each other, since they shared features that Cnidaria lack, for example muscles in the middle layer (mesoglea
in Ctenophora, mesoderm in Bilateria). However more recent analyses indicate that these similarities are
rather vague, and the current view, based on molecular phylogenetics, is that Cnidaria and Bilateria are more
closely related to each other than either is to Ctenophora. This grouping of Cnidaria and Bilateria has been
labelled "Planulozoa" because it suggests that the earliest Bilateria were similar to the planula larvae of
Within the Cnidaria, the Anthozoa (sea anemones and corals) are regarded as the sister-group of the
rest, which suggests that the earliest cnidarians were sessile polyps with no medusa stage. However it is
unclear how the other groups acquired the medusa stage, since Hydrozoa form medusae by budding from
the side of the polyp while the other Medusozoa do so by splitting them off from the tip of the polyp. The
traditional grouping of Scyphozoa included the Staurozoa, but morphology and molecular phylogenetics
indicate that Staurozoa are more closely related to Cubozoa (box jellies) than to other "Scyphozoa".
Similarities in the double body walls of Staurozoa and the extinct Conulariida suggest that they are closely
related. The position of Anthozoa nearest the beginning of the cnidarian family tree also implies that
Anthozoa are the cnidarians most closely related to Bilateria, and this is supported by the fact that Anthozoa
and Bilateria share some genes that determine the main axes of the body.[2][42]
However in 2005 Katja Seipel and Volker Schmid suggested that cnidarians and ctenophores are
simplified descendants of triploblastic animals, since ctenophores and the medusa stage of some cnidarians
have striated muscle, which in bilaterians arises from the mesoderm. They did not commit themselves on
whether bilaterians evolved from early cnidarians or from the hypothesized triploblastic ancestors of
In molecular phylogenetics analyses from 2005 onwards, important groups of developmental genes
show the same variety in cnidarians as in chordates.[43] In fact cnidarians, and especially anthozoans (sea
anemones and corals), retain some genes that are present in bacteria, protists, plants and fungi but not in

[edit] Interaction with humans

Jellyfish stings killed about 1,500 people in the 20th century,[45] and cubozoans are particularly
dangerous. On the other hand, some large jellyfish are considered a delicacy in eastern and southern Asia.
Coral reefs have long been economically important as providers of fishing grounds, protectors of shore
buildings against currents and tides, and more recently as centers of tourism. However, they are vulnerable
to over-fishing, mining for construction materials, pollution, and damage caused by tourism.
Beaches protected from tides and storms by coral reefs are often the best places for housing in
tropical countries. Reefs are an important food source for low-technology fishing, both on the reefs
themselves and in the adjacent seas.[46] However despite their great productivity reefs are vulnerable to
over-fishing, because much of the organic carbon they produce is exhaled as carbon dioxide by organisms at
the middle levels of the food chain and never reaches the larger species that are of interest to fishermen.[16]
Tourism centered on reefs provides much of the income of some tropical islands, attracting photographers,
divers and sports fishermen. However human activities damage reefs in several ways: mining for
construction materials; pollution, including large influxes of fresh water from storm drains; commercial fishing,
including the use of dynamite to stun fish and the capture of young fish for aquariums; and tourist damage
caused by boat anchors and the cumulative effect of walking on the reefs.[46] Coral, mainly from the Pacific
Ocean has long been used in jewellery, and demand rose sharply in the 1980s.[47]
Some large jellyfish species have been used in Chinese cuisine at least
since 200 AD, and are now fished in the seas around most of South East Asia.
Japan is the largest single consumer of edible jellyfish, importing at first only from
China but now from all of South East Asia as prices rose in the 1970s. This fishing
industry is restricted to daylight hours and calm conditions in two short seasons, from
March to May and August to November.[48] The commercial value of jellyfish food
products depends on the skill with which they are prepared, and "Jellyfish Masters"
guard their trade secrets carefully. Jellyfish is very low in cholesterol and sugars, but
cheap preparation can introduce undesirable amounts of heavy metals.[49]
The "sea wasp" Chironex fleckeri has been described as the world's most
venomous animal and is held responsible for 67 deaths, although it is difficult to
identify the animal as it is almost transparent. Most stingings by C. fleckeri cause
only mild symptoms.[50] Seven other box jellies can cause a set of symptoms called
Irukandji syndrome,[51] which takes about 30 minutes to develop,[52] and from a few
hours to two weeks to disappear.[53] Hospital treatment is usually required, and
there have been a few deaths.[51]

dangerous "sea wasp"
Chironex fleckeri
[edit] Notes
1. ^ Classes in Medusozoa based on "The Taxonomicon - Taxon: Subphylum Medusozoa".
Universal Taxonomic Services.
Retrieved 2009-01-26.
2. ^ a b c d Collins, A.G. (2002). "Phylogeny of Medusozoa and the Evolution of Cnidarian Life
Cycles" (PDF). Journal of Evolutionary Biology 15 (3): 418–432. doi:10.1046/j.1420-
9101.2002.00403.x. Retrieved 2008-
3. ^ Subphyla Anthozoa and Medusozoa based on "The Taxonomicon - Taxon: Phylum
Cnidaria". Universal Taxonomic Services.
id=11551. Retrieved 2007-07-10.
4. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af Hinde, R.T., (1998). "The Cnidaria
and Ctenophora". In Anderson, D.T.,. Invertebrate Zoology. Oxford University Press. pp. 28–57.
ISBN 0195513681.
5. ^ Collins, A.G., Cartwright, P., McFadden, C.S., and Schierwater, B. (2005). "Phylogenetic
Context and Basal Metazoan Model Systems". Integrative and Comparative Biology 45 (4): 585–594.
6. ^ Štolc, A. (1899). "Actinomyxidies, nouveau groupe de Mesozoaires parent des
Myxosporidies". Bull. Int. L'Acad. Sci. Bohème 12: 1–12.
7. ^ E. Jímenez-Guri; Philippe, H; Okamura, B; Holland, PW (July 2007). " Buddenbrockia is a
cnidarian worm". Science 317 (116): 116–118. doi:10.1126/science.1142024. PMID 17615357.
8. ^ Zrzavý, J. and Hypša, V. (2003). "Myxozoa, Polypodium, and the origin of the Bilateria: The
phylogenetic position of "Endocnidozoa" in light of the rediscovery of Buddenbrockia". Cladistics 19
(2): 164–169. doi:10.1111/j.1096-0031.2003.tb00305.x.
9. ^ "The Conulariida". University of California Museum of Paleontology. Retrieved 2008-11-27.
10.^ a b c d e f g Shostak, S. (2006). "Cnidaria (Coelenterates)". Encyclopedia of Life Sciences.
John Wiley & Sons. doi:10.1038/npg.els.0004117.
11.^ Blaise, C., and Férard, J-F. (2005). Small-scale Freshwater Toxicity Investigations: Toxicity
Test Methods. Springer. p. 398. ISBN 140203119X.
id=Ibew5SLx2oMC&dq=hydra+size+length. Retrieved 2008-11-21.
12.^ a b Safina, C. (2007). Voyage of the Turtle: In Pursuit of the Earth's Last Dinosaur .
Macmillan. p. 154. ISBN 0805083189.
id=dQD883dAv6YC&pg=PA154&dq=cnidaria+turtle. Retrieved 2008-11-21.
13.^ Cowen, R. (2000). History of Life (3 ed.). Blackwell. p. 54. ISBN 0632044446. Retrieved 2008-11-21.
14.^ Frick, K (2003). "Predator Suites and Flabellinid Nudibranch Nematocyst Complements in
the Gulf of Maine.". In: SF Norton (ed). Diving for Science...2003. Proceedings of the American
Academy of Underwater Sciences (22nd Annual Scientific Diving Symposium). http://archive.rubicon- Retrieved 2008-07-03.
15.^ Choat, J.H. and Bellwood, D.R. (1998). Paxton, J.R. and Eschmeyer, W.N.. ed.
Encyclopedia of Fishes. San Diego: Academic Press. pp. 209–211. ISBN 0-12-547665-5.
16.^ a b Barnes, R.S.K., and Mann, K.H. (1991). Fundamentals of Aquatic Ecology. Blackwell
Publishing. pp. 217–227. ISBN 0632029838.
id=mOZZlzgdTrwC&pg=PA227&dq=%22Coral+Reef%22+productivity. Retrieved 2008-11-26.
17.^ Hatcher, B.G. Johannes, R.E., and Robertson, A.J. (1989). "Conservation of Shallow-water
Marine Ecosystems". Oceanography and Marine Biology: An Annual Review: Volume 27. Routledge.
p. 320. ISBN 0080377181.
%22Coral+Reef%22+mangrove+%22seagrass%22. Retrieved 2008-11-21.
18.^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag Ruppert, E.E., Fox, R.S., and
Barnes, R.D. (2004). Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 111–124. ISBN 0030259827.
19.^ a b c d Seipel, K., and Schmid, V. (June 2005). "Evolution of striated muscle: Jellyfish and
the origin of triploblasty". Developmental Biology 282 (1): 14–26. doi:10.1016/j.ydbio.2005.03.032.
PMID 15936326.
20.^ a b c Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). Invertebrate Zoology (7 ed.).
Brooks / Cole. pp. 182–195. ISBN 0030259827.
21.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). Invertebrate Zoology (7 ed.). Brooks /
Cole. pp. 76–97. ISBN 0030259827.
22.^ Bergquist, P.R., (1998). "Porifera". In Anderson, D.T.,. Invertebrate Zoology. Oxford
University Press. pp. 10–27. ISBN 0195513681.
23.^ Exposito, J-Y., Cluzel, C., Garrone, R., and Lethias, C. (2002). "Evolution of collagens".
The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology 268 (3):
302–316. doi:10.1002/ar.10162. PMID 12382326.
24.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Introduction to Metazoa". Invertebrate
Zoology (7 ed.). Brooks / Cole. pp. 103–104. ISBN 0030259827.
25.^ Trumble, W., and Brown, L. (2002). "Cnida". Shorter Oxford English Dictionary. Oxford
University Press.
26.^ Bhamrah, H.S., and Juneja, K. (2002). A Textbook of Invertebrates. Anmol Publications.
pp. 278–280. ISBN 8126104163.
id=05So4shx9tIC&pg=PA279&dq=siphonophore+siphonophora. Retrieved 2008-11-17.
27.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). Invertebrate Zoology (7 ed.). Brooks /
Cole. pp. 167–170. ISBN 0030259827.
28.^ Chen, J-Y. (2000). "Putative phosphatized embryos from the Doushantuo Formation of
China". Proceedings of the National Academy of Sciences 97: 4457–4462.
doi:10.1073/pnas.97.9.4457. PMID 10781044. Retrieved
29.^ Xiao, S., Yuan, X., and Knoll, A.H. (2000). "Eumetazoan fossils in terminal Proterozoic
phosphorites?". Proceedings of the National Academy of Sciences 97 (25): 13684–13689.
doi:10.1073/pnas.250491697. PMID 11095754.
30.^ Chen, J.-Y., Oliveri, P., Gao, F., Dornbos, S.Q., Li, C-W., Bottjer, D.J. and Davidson, E.H.
(August 2002). "Precambrian Animal Life: Probable Developmental and Adult Cnidarian Forms from
Southwest China" (PDF). Developmental Biology 248 (1): 182–196. doi:10.1006/dbio.2002.0714.
PMID 12142030.'s/Chen%20et%20al.%202002.pdf. Retrieved
31.^ Donovan, Stephen K., Lewis, David N. (2001). "Fossils explained 35. The Ediacaran biota"
(abstract). Geology Today 17 (3): 115–120. doi:10.1046/j.0266-6979.2001.00285.x.
32.^ Antcliffe, J.B.; Brasier, M. D. (2007). "Charnia and sea pens are poles apart". Journal of the
Geological Society 164 (1): 49–51. doi:10.1144/0016-76492006-080.
33.^ Antcliffe, J.B.; Brasier, Martin D. (2007). "Charnia At 50: Developmental Models For
Ediacaran Fronds". Palaeontology 51 (1): 11–26. doi:10.1111/j.1475-4983.2007.00738.x.
34.^ a b c d "Cnidaria: Fossil Record". University of California Museum of Paleontology. Retrieved 2008-11-27.
35.^ Copper, P. (January 1994). "Ancient reef ecosystem expansion and collapse". Coral Reefs
13 (1): 3–11. doi:10.1007/BF00426428.
36.^ "The Rudists". University of California Museum of Paleontology. Retrieved 2008-11-27.
37.^ a b Borchiellini, C., Manuel, M., Alivon, E., Boury-Esnault, N., Vacelet J., and Le Parco, Y.
(2001). "Sponge paraphyly and the origin of Metazoa". Journal of Evolutionary Biology 14 (1): 171–
179. doi:10.1046/j.1420-9101.2001.00244.x.
38.^ a b Wallberg, A., Thollesson, M., , Farris, J.S., and Jondelius, U. (2004). "The phylogenetic
position of the comb jellies (Ctenophora) and the importance of taxonomic sampling". Cladistics 20:
558–578. doi:10.1111/j.1096-0031.2004.00041.x.
39.^ Halanych, K.M. (December 2004). "The New View of Animal Phylogeny" (PDF). Annual
Review of Ecology, Evolution, and Systematics 35: 229–256.
doi:10.1146/annurev.ecolsys.35.112202.130124. Retrieved 2008-11-27.
40.^ Medina, M., Collins, A.G., Silberman, J.D., and Sogin, M.L. (August 2001). "Evaluating
hypotheses of basal animal phylogeny using complete sequences of large and small subunit rRNA".
Proceedings of the National Academy of Sciences 98 (17): 9707–9712. doi:10.1073/pnas.171316998.
PMID 11504944.
41.^ Müller, W.E.G., Li, J., Schröder, H.C., Qiao, L., and Wang, X. (2007). "The unique skeleton
of siliceous sponges (Porifera; Hexactinellida and Demospongiae) that evolved first from the
Urmetazoa during the Proterozoic: a review". Biogeosciences 4: 219–232. doi:10.5194/bg-4-219-
42.^ Marques, A.C., and Collins, A.G. (2004). "Cladistic analysis of Medusozoa and cnidarian
evolution". Invertebrate Biology 123 (1): 23–42. doi:10.1111/j.1744-7410.2004.tb00139.x. Retrieved 2008-11-27.
43.^ Miller, D.J., Ball, E.E., and Technau, U. (October 2005). "Cnidarians and ancestral genetic
complexity in the animal kingdom". Trends in Genetics 21 (10): 536–539.
doi:10.1016/j.tig.2005.08.002. PMID 16098631.
44.^ Technau, U., Rudd, S., and Maxwell, P (December 2005). "Maintenance of ancestral
complexity and non-metazoan genes in two basal cnidarians". Trends in Genetics 21 (12): 633–639.
doi:10.1016/j.tig.2005.09.007. PMID 16226338.
45.^ Williamson, J.A., Fenner, P.J., Burnett, J.W., and Rifkin, J. (1996). Venomous and
Poisonous Marine Animals: A Medical and Biological Handbook. UNSW Press. pp. 65–68.
ISBN 0868402796.
id=YsZ3GryFIzEC&pg=PA75&lpg=PA75&dq=mollusc+venom+fatal. Retrieved 2008-10-03.
46.^ a b Clark, J.R. (1998). Coastal Seas: The Conservation Challenge. Blackwell. pp. 8–9.
ISBN 0632049553.
%22+productivity. Retrieved 2008-11-28.
47.^ Cronan, D.S., (1991). Marine Minerals in Exclusive Economic Zones . Springer. pp. 63–65.
ISBN 041229270X.
Retrieved 2008-11-28.
48.^ Omori, M. and Nakano, E. (2001). "Jellyfish fisheries in southeast Asia". In Purcell, J.E.,.
Jellyfish Blooms: Ecological and Societal Importance. Springer. pp. 19–26. ISBN 0792369645.
49.^ Hsieh, Y-H.P. Leong, F-M., and Rudloe, J. (2001). "Jellyfish as food". In Purcell, J.E.,.
Jellyfish Blooms: Ecological and Societal Importance. Springer. pp. 11–17. ISBN 0792369645.
50.^ Greenberg, M.I., Hendrickson, R.G., Silverberg, M., Campbell, C., and Morocco, A. (2004).
"Box Jellyfish Envenomation". Greenberg's Text-atlas of Emergency Medicine. Lippincott Williams &
Wilkins. p. 875. ISBN 0781745861.
51.^ a b Little, M., Pereira, P., Carrette, T., and Seymour, J. (2006). "Jellyfish Responsible for
Irukandji Syndrome". QJM (Quarterly Journal of Medicine) 99 (6): 425–427.
doi:10.1093/qjmed/hcl057. PMID 16687419.
52.^ Barnes, J. (1964). "Cause and effect in Irukandji stingings". Medical Journal of Australia 14:
897–904. PMID 14172390.
53.^ Grady J, Burnett J (2003). "Irukandji-like syndrome in South Florida divers". Annals of
Emergency Medicine 42 (6): 763–6. doi:10.1016/S0196-0644(03)00513-4. PMID 14634600.
[edit] Further reading
[edit] Books
• Arai, M.N. (1997). A Functional Biology of Scyphozoa. London: Chapman & Hall [p. 316].
ISBN 0-412-45110-7.
• Ax, P. (1999). Das System der Metazoa I. Ein Lehrbuch der phylogenetischen Systematik.
Gustav Fischer, Stuttgart-Jena: Gustav Fischer. ISBN 3-437-30803-3.
• Barnes, R.S.K., P. Calow, P. J. W. Olive, D. W. Golding & J. I. Spicer (2001). The
invertebrates—a synthesis. Oxford: Blackwell. 3rd edition [chapter 3.4.2, p. 54]. ISBN 0-632-04761-5.
• Brusca, R.C., G.J. Brusca (2003). Invertebrates. Sunderland, Mass.: Sinauer Associates.
2nd edition [chapter 8, p. 219]. ISBN 0-87893-097-3.
• Dalby, A. (2003). Food in the Ancient World: from A to Z. London: Routledge.
• Moore, J.(2001). An Introduction to the Invertebrates. Cambridge: Cambridge University
Press [chapter 4, p. 30]. ISBN 0-521-77914-6.
• Schäfer, W. (1997). Cnidaria, Nesseltiere. In Rieger, W. (ed.) Spezielle Zoologie. Teil 1.
Einzeller und Wirbellose Tiere. Stuttgart-Jena: Gustav Fischer. Spektrum Akademischer Verl.,
Heidelberg, 2004. ISBN 3-8274-1482-2.
• Werner, B. 4. Stamm Cnidaria. In: V. Gruner (ed.) Lehrbuch der speziellen Zoologie. Begr.
von Kaestner. 2 Bde. Stuttgart-Jena: Gustav Fischer, Stuttgart-Jena. 1954, 1980, 1984, Spektrum
Akad. Verl., Heidelberg-Berlin, 1993. 5th edition. ISBN 3-334-60474-8.
[edit] Journal articles
• D. Bridge, B. Schierwater, C. W. Cunningham, R. DeSalle R, L. W. Buss: Mitochondrial DNA
structure and the molecular phylogeny of recent cnidaria classes. in: Proceedings of the Academy of
Natural Sciences of Philadelphia. Philadelphia USA 89.1992, p. 8750. ISSN 0097-3157
• D. Bridge, C. W. Cunningham, R. DeSalle, L. W. Buss: Class-level relationships in the
phylum Cnidaria—Molecular and morphological evidence. in: Molecular biology and evolution. Oxford
University Press, Oxford 12.1995, p. 679. ISSN 0737-4038
• D. G. Fautin: Reproduction of Cnidaria. in: Canadian Journal of Zoology. Ottawa Ont.
80.2002, p. 1735. (PDF, online) ISSN 0008-4301
• G. O. Mackie: What's new in cnidarian biology? in: Canadian Journal of Zoology. Ottawa Ont.
80.2002, p. 1649. (PDF, online) ISSN 0008-4301
• P. Schuchert: Phylogenetic analysis of the Cnidaria. in: Zeitschrift für zoologische Systematik
und Evolutionsforschung. Paray, Hamburg-Berlin 31.1993, p. 161. ISSN 0044-3808
• G. Kass-Simon, A. A. Scappaticci Jr.: The behavioral and developmental physiology of
nematocysts. in: Canadian Journal of Zoology. Ottawa Ont. 80.2002, p. 1772. (PDF, online)
ISSN 0044-3808
• J. Zrzavý (2001). "The interrelationships of metazoan parasites: a review of phylum- and
higher-level hypotheses from recent morphological and molecular phylogenetic analyses" (PDF).
Folia Parasitologica 48 (2): 81–103. PMID 11437135. Archived from the original on 2007-10-25. Retrieved
[edit] External links
Wikispecies has information related to: Cnidaria

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Look up Cnidaria in Wiktionary, the free dictionary.

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• Cnidaria - Guide to the Marine Zooplankton of south eastern Australia , Tasmanian
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• A Cnidaria homepage maintained by University of California, Irvine
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Categories: Venomous animals | Cnidarians

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Temporal range: 245–90 Ma
g Middle Triassic - Late Cretaceous
Mounted fossil of Temnodontosaurus trigonodon, Staatliches Museum für Naturkunde Stuttgart

Scientific classification [ e ]

Kingdom: Animalia

Phylum: Chordata

Class: Reptilia

Superorder: †Ichthyopterygia

(unranked): †Eoichthyosauria
Blainville, 1835


Ichthyosaurs (Greek for "fish lizard" - ιχθυς/ichthys meaning "fish" and σαυρος/sauros meaning
"lizard") were giant marine reptiles that resembled fish and dolphins. Ichthyosaurs thrived during much of the
Mesozoic era; based on fossil evidence, they first appeared approximately 245 million years ago (mya) and
disappeared about 90 million years ago, about 25 million years before the dinosaurs became extinct. During
the middle Triassic Period, ichthyosaurs evolved from as-yet unidentified land reptiles that moved back into
the water, in a development parallel to that of the ancestors of modern-day dolphins and whales. They were
particularly abundant in the Jurassic Period, until they were replaced as the top aquatic predators by
plesiosaurs in the Cretaceous Period. They belong to the order known as Ichthyosauria or Ichthyopterygia
('fish flippers' - a designation introduced by Sir Richard Owen in 1840, although the term is now used more
for the parent clade of the Ichthyosauria).
• 1 Description
• 2 Reproduction
• 3 Feeding
• 4 History of discoveries
• 5 Evolutionary history
• 6 Taxonomy
• 7 Phylogeny
• 8 Geologic formations
• 9 References
• 10 External links
[edit] Description

CGI restoration of Ichthyosaurus

Ichthyosaurs averaged 2–4 metres (6.6–13 ft) in length (although a few were smaller, and some
species grew much larger), with a porpoise-like head and a long, toothed snout. Built for speed, like modern
tuna, some ichthyosaurs appear also to have been deep divers, like some modern whales (Motani, 2000). It
has been estimated that ichthyosaurs could swim at speeds up to 40 kilometres per hour (25 mph).[1] Similar
to modern cetaceans such as whales and dolphins, they were air-breathing.
According to weight estimates by Ryosuke Motani[2] a 2.4 meters (8 ft) Stenopterygius weighed
around 163–168 kilograms (360–370 lb) whilst a 4 meters (13 ft) Ophthalmosaurus icenicus weighed 930–950
kilograms (1.03–1.05 short tons).
Although ichthyosaurs looked like fish, they were not. Biologist Stephen Jay Gould said the
ichthyosaur was his favorite example of convergent evolution, where similarities of structure are analogous
not homologous, for this group:
converged so strongly on fishes that it actually evolved a dorsal fin and tail in just the right place
and with just the right hydrological design. These structures are all the more remarkable
because they evolved from nothing — the ancestral terrestrial reptile had no hump on its back or
blade on its tail to serve as a precursor.

Ichthyosaur 'paddle' (Charmouth Heritage Coast Centre).

In fact the earliest reconstructions of ichthyosaurs omitted the dorsal fin, which had no hard skeletal
structure, until finely-preserved specimens recovered in the 1890s from the Holzmaden lagerstätten in
Germany revealed traces of the fin. Unique conditions permitted the preservation of soft tissue impressions.
Ichthyosaur vertebra from the Sundance Formation (Jurassic) of Natrona County, Wyoming. Note the
characteristic "figure-8" cross-section for this variety. Scale in mm.
Ichthyosaurs had fin-like limbs, which were possibly used for stabilization and directional control,
rather than propulsion, which would have come from the large shark-like tail. The tail was bi-lobed, with the
lower lobe being supported by the caudal vertebral column, which was "kinked" ventrally to follow the
contours of the ventral lobe.
Apart from the obvious similarities to fish, the ichthyosaurs also shared parallel developmental
features with dolphins, lamnid sharks, and tunas. This gave them a broadly similar appearance, possibly
implied similar activity levels (including thermoregulation), and presumably placed them broadly in a similar
ecological niche.
[edit] Reproduction

Adult and juvenile Stenopterygius quadriscissus

They were viviparous (bore live young). Some adult fossils have even been found containing fetuses.
Although they were reptiles and descended from egg-laying ancestors, viviparity is not as unexpected as it
first appears. Air-breathing marine creatures must either come ashore to lay eggs, like turtles and some sea
snakes, or else give birth to live young in surface waters, like whales and dolphins. Given their streamlined
bodies, heavily adapted for fast swimming, it would have been difficult for ichthyosaurs to move far enough
on land to lay eggs.
Crocodiles, most sea turtles and some lizards, control offspring gender by manipulating the
temperature of the eggs' environment. I.e., they do not have of distinct sex chromosomes. Live-bearing
reptiles do not regulate sex through incubation temperature. A recent study examined 94 living species of
reptiles, birds and mammals and found that the genetic control of sex appears to be crucial to live birth and
that genetics likely controlled gender in ichthyosaurs, mosasaurs and other extinct marine reptiles.[4]
[edit] Feeding

Ichthyosaurus acutirostris with ammonoids

For their food, many of the fish-shaped ichthyosaurs relied heavily on ancient cephalopod kin of
squids called belemnites. Some early ichthyosaurs had teeth adapted for crushing shellfish. They also most
likely fed on fish, and a few of the larger species had heavy jaws and teeth that indicated they fed on smaller
reptiles. Ichthyosaurs ranged so widely in size, and survived for so long, that they are likely to have had a
wide range of prey. Typical ichthyosaurs have very large eyes, protected within a bony ring, suggesting that
they may have hunted at night.
[edit] History of discoveries

Diagram of the skeletal anatomy of an Ichthyosaurus communis found by Mary Anning, from an 1824
paper by Conybeare
The nominate genus Ichthyosaurus had first been described in 1699 from fossil fragments
discovered in Wales.
The first fossil vertebrae were published twice in 1708 as tangible mementos of the Universal
Deluge. The first complete ichthyosaur fossil was found in 1811 by Mary Anning in Lyme Regis, along what is
now called the Jurassic Coast. She subsequently discovered three separate species.
In 1905, the Saurian Expedition led by John C. Merriam of the University of California and financed
by Annie Alexander, found 25 specimens in central Nevada, which during the Triassic was under a shallow
ocean. Several of the specimens are now in the collection of the University of California Museum of
Paleontology. Other specimens are embedded in the rock and visible at Berlin–Ichthyosaur State Park in Nye
County. In 1977, the Triassic ichthyosaur Shonisaurus became the State Fossil of Nevada. Nevada is the
only state to possess a complete skeleton, 17 m (55 ft) of this extinct marine reptile. In 1992, Canadian
ichthyologist Dr. Elizabeth Nicholls (Curator of Marine Reptiles at the Royal Tyrrell Museum) uncovered the
largest known specimen, a 23 m-long (75 ft) example.
[edit] Evolutionary history

Grippia longirostris from early Triassic of Spitsbergen

The earliest ichthyosaurs, looking more like finned lizards than the familiar fish or dolphin forms, are
known from the Early and Early-Middle (Olenekian and Anisian) Triassic strata of Canada, China, Japan, and
Spitsbergen in Norway. These primitive forms included the genera Chaohusaurus, Grippia, and
These very early proto-ichthyosaurs, which are now classified as Ichthyopterygia rather than as
ichthyosaurs proper (Motani 1997, Motani et al. 1998), quickly gave rise to true ichthyosaurs sometime
around the boundary between the Early Triassic and Middle Triassic. These later diversified into a variety of
forms, including the sea serpent like Cymbospondylus, which reached 10 metres, and smaller more typical
forms like Mixosaurus. By the Late Triassic, ichthyosaurs consisted of both classic Shastasauria and more
advanced, "dolphin"-like Euichthyosauria (Californosaurus, Toretocnemus) and Parvipelvia (Hudsonelpidia,
Macgowania). Experts disagree over whether these represent an evolutionary continuum, with the less
specialised shastosaurs a paraphyletic grade that was evolving into the more advanced forms (Maisch and
Matzke 2000), or whether the two were separate clades that evolved from a common ancestor earlier on
(Nicholls and Manabe 2001).
Shonisaurus popularis.
During the Carnian and Norian, shastosaurs reached huge sizes. Shonisaurus popularis, known from
a number of specimens from the Carnian of Nevada, was 15 metres long. Norian shonisaurs are known from
both sides of the Pacific. Himalayasaurus tibetensis and Tibetosaurus (probably a synonym) have been
found in Tibet. These large (10 to 15 metres long) ichthyosaurs probably belong to the same genus as
Shonisaurus (Motani et al., 1999; Lucas, 2001, pp. 117–119). While the gigantic Shonisaurus sikanniensis,
whose remains were found in the Pardonet formation of British Columbia by Elizabeth Nicholls, reached as
much as 21 metres in length - the largest marine reptile known to date.
These giants (along with their smaller cousins) seemed to have disappeared at the end of the Norian.
Rhaetian (latest Triassic) ichthyosaurs are known from England, and these are very similar to those of the
Early Jurassic. Like the dinosaurs, the ichthyosaurs and their contemporaries the plesiosaurs survived the
end-Triassic extinction event, and immediately diversified to fill the vacant ecological niches of the earliest
Ichthyosaurus sp. fossil
The Early Jurassic, like the Late Triassic, was the heyday of the ichthyosaurs, which are represented
by four families and a variety of species, ranging from one to ten metres in length. Genera include
Eurhinosaurus, Ichthyosaurus, Leptonectes, Stenopterygius, and the large predator Temnodontosaurus,
along with the persistently primitive Suevoleviathan, which was little changed from its Norian ancestors. All
these animals were streamlined, dolphin-like forms, although the more primitive animals were perhaps more
elongated than the advanced and compact Stenopterygius and Ichthyosaurus.
Ichthyosaurs were still common in the Middle Jurassic, but had now decreased in diversity. All
belonged to the single clade Ophthalmosauria. Represented by the 4 metre long Ophthalmosaurus and
related genera, they were very similar to Ichthyosaurus, and had attained a perfect "tear-drop" streamlined
form. The eyes of Ophthalmosaurus were huge, and it is likely that these animals hunted in dim and deep
water (Motani 2000).
Platypterygius kiprijanovi - Albian-Cenomanian of Kursk region (Russia)
Ichthyosaurs seemed to decrease in diversity even further with the Cretaceous. Only three genera
are known, Caypullisaurus, Maiaspondylus, and Platypterygius, although they had a worldwide distribution.
This last ichthyosaur genus became extinct during the Cenomanian-Turonian extinction event early in the
Late Cretaceous (as did the pliosaurs). Interestingly, less hydrodynamically efficient animals like mosasaurs
and long-necked plesiosaurs flourished. It could be that the ichthyosaurian over-specialisation was a
contributing factor to their extinction, possibly being unable to 'keep up' with the fast swimming and highly
evasive new teleost fish, which had become dominant at this time, against which the sit-and-wait ambush
strategies of the mosasaurs proved superior (Lingham-Soliar 1999).

[edit] Taxonomy
• Superorder Ichthyopterygia
• Family Grippidae
• Chaohusaurus
• Grippia
• Family Utatsusauridae
• Utatsusaurus
• Cymbospondylus
• Family Mixosauridae
• Mixosaurus
• Suborder Merriamosauriformes
• Guanlingsaurus
• (unranked) Merriamosauria
• Infraorder Shastasauria
• Family Shastasauridae
• Himalayasaurus
• Shastasaurus
• Shonisaurus
• Tibetosaurus
• Infraorder Ichthyosauria (=Euichthyosauruia)
• Family Ichthyosauridae
• Ichthyosaurus
• Family Leptopterygiidae
• Eurhinosaurus
• Excalibosaurus
• Leptonectes
• Family Temnodontosauridae
• Temnodontosaurus
• Family Ophthalmosauridae
• Aegirosaurus
• Arthropterygius
• Brachypterygius
• Caypullisaurus
• Nannopterygius
• Ophthalmosaurus
• Platypterygius
• Family Stenopterygiidae
• ?Chacaicosaurus
• Stenopterygius
• Family Toretocnemidae
• Californosaurus
• Qianichthyosaurus
• Toretocnemus
• Family Suevoleviathanidae
• Suevoleviathan
• Infraorder Parvipelvia
• Hudsonelpidia
• Macgowania

[edit] Phylogeny
The following cladogram is based on Motani (1999):[5]











Cladogram based on Maisch and Matzke (2000)[6] and Maisch and Matzke (2003)[7] with clade
names following Maisch (2010)[8]:






[edit] Geologic formations
The following is a list of geologic formations with ichthyosaur fossils.

Name Age Location Notes

Franciscan Formation • USA

Hosselkus Limestone • USA

• Arge
Los Molles Formation

Oxford Clay • UK

• Germ
Solnhofen Limestone

• Cana
Sulphur Mountain Formation

Sundance Formation • USA

[edit] References
1. ^
2. ^
3. ^ Martill D.M. 1993. Soupy Substrates: A Medium for the Exceptional Preservation of
Ichthyosaurs of the Posidonia Shale (Lower Jurassic) of Germany. Kaupia - Darmstädter Beiträge
zur Naturgeschichte 2: 77-97
4. ^ Carroll, Sean B. (March 22, 2010). "For Extinct Monsters of the Deep, a Little Respect".
The New York Times.
5. ^ Motani, Ryosuke (1999). "Phylogeny of the Ichthyopterygia". Journal of Vertebrate
Paleontology 19: 472–495.
6. ^ Michael W. Maisch and Andreas T. Matzke (2000). "The Ichthyosauria". Stuttgarter
Beiträge zur Naturkunde: Serie B 298: 159. http://www.naturkundemuseum-
7. ^ Michael W. Maisch and Andreas T. Matzke (2003). "Observations on Triassic ichthyosaurs.
Part XII. A new Lower Triassic ichthyosaur genus from Spitzbergen". Neues Jahrbuch für Geologie
und Paläontologie Abhandlungen 229: 317–338.
8. ^ Michael W. Maisch (2010). "Phylogeny, systematics, and origin of the Ichthyosauria – the
state of the art". Palaeodiversity 3: 151–214.
• Ellis, Richard, (2003) Sea Dragons - Predators of the Prehistoric Oceans. University Press of
Kansas ISBN 0-7006-1269-6
• Stephen Jay Gould, "Bent out of Shape" in Eight Little Piggies.
• Lingham-Soliar, T. (1999): A functional analysis of the skull of Goronyosaurus nigeriensis
(Squamata: Mosasauridae) and Its Bearing on the Predatory Behavior and Evolution of the Enigmatic
Taxon. N. Jb. Geol. Palaeont. Abh. 2134 (3):355-74
• Maisch, M. W. & Matzke, A. T. (2000) The ichthyosauria. Stuttgarter Beitraege zur
Naturkunde. Serie B. Geologie und Palaeontologie . 2000; (298):1-159.
• McGowan, Christopher (1992) Dinosaurs, Spitfires and Sea Dragons, Harvard University
Press, ISBN 0-674-20770-X
• McGowan, Christopher & Motani, Ryosuke, (2003) Ichthyopterygia, Handbook of
Paleoherpetology, Part 8, Verlag Dr. Friedrich Pfeil
• Motani, R. (1997), Temporal and spatial distribution of tooth implantation in ichthyosaurs, in
JM Callaway & EL Nicholls (eds.), Ancient Marine Reptiles. Academic Press. pp. 81–103.
• Motani, R. (2000), Rulers of the Jurassic Seas, Scientific American vol.283, no. 6
• Motani, R., Minoura, N. & Ando, T. (1998), Ichthyosaurian relationships illuminated by new
primitive skeletons from Japan. Nature 393:255-257.
• Motani, R., Manabe, M., and Dong, Z-M, (1999) The status of Himalayasaurus tibetensis
(Ichthyopterygia) pdf, Paludicola2(2):174-181.
• Nicholls, E. L. & Manabe, M. 2001. A new genus of ichthyosaur from the Late Triassic
Pardonet Formation of British Columbia: bridging the Triassic-Jurassic gap. Canadian Journal of
Earth Sciences 38:983-1002.
[edit] External links
Wikimedia Commons has media related to: Ichthyosauria

• USMP Berkeley's ichthyosaur introduction

• Ryosuke Motani's detailed Ichthyosaur homepage, with vivid graphics
• Eureptilia: Ichthyosauria - Palaeos
• Ichthyosauria - cladogram (Mikko's Phylogeny Archive)
• Hauff Museum, Germany - exhibiting the finds of Holzmaden
Retrieved from ""

Categories: Ichthyosaurs | Natural history of Nevada


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Temporal range: Early Ordovician–Recent[1]
Glycera sp.
Scientific classification

Kingdom: Animalia

Superphylum: Lophotrochozoa

Lamarck, 1809

Classes and subclasses

Class Polychaeta (paraphyletic?)

Class Clitellata (see below)
Oligochaeta – earthworms, etc.
Hirudinea – leeches
Class Myzostomida
Class Archiannelida (polyphyletic)
The annelids (also called "ringed worms"), formally called Annelida (from French annelés "ringed
ones", ultimately from Latin anellus "little ring"[2]), are a large phylum of segmented worms, with over 17,000
modern species including ragworms, earthworms and leeches. They are found in marine environments from
tidal zones to hydrothermal vents, in freshwater, and in moist terrestrial environments. Although most
textbooks still use the traditional division into polychaetes (almost all marine), oligochaetes (which include
earthworms) and leech-like species, research since 1997 has radically changed this scheme, viewing
leeches as a sub-group of oligochaetes and oligochaetes as a sub-group of polychaetes. In addition, the
Pogonophora, Echiura and Sipuncula, previously regarded as separate phyla, are now regarded as sub-
groups of polychaetes. Annelids are considered members of the Lophotrochozoa, a "super-phylum" of
protostomes that also includes molluscs, brachiopods, flatworms and nemerteans.
The basic annelid form consists of multiple segments, each of which has the same sets of organs
and, in most polychaetes, a pair of parapodia that many species use for locomotion. Septa separate the
segments of many species, but are poorly-defined or absent in some, and Echiura and Sipuncula show no
obvious signs of segmentation. In species with well-developed septa, the blood circulates entirely within
blood vessels, and the vessels in segments near the front ends of these species are often built up with
muscles to act as hearts. The septa of these species also enable them to change the shapes of individual
segments, which facilitates movement by peristalsis ("ripples" that pass along the body) or by undulations
that improve the effectiveness of the parapodia. In species with incomplete septa or none, the blood
circulates through the main body cavity without any kind of pump, and there is a wide range of locomotory
techniques – some burrowing species turn their pharynges inside out to drag themselves through the
Although many species can reproduce asexually and use similar mechanisms to regenerate after
severe injuries, sexual reproduction is the normal method in species whose reproduction has been studied.
The minority of living polychaetes whose reproduction and lifecycles are known produce trochophore larvae,
which live as plankton and then sink and metamorphose into miniature adults. Oligochaetes are full
hermaphrodites and produce a ring-like cocoon round their bodies, in which the eggs and hatchlings are
nourished until they are ready to emerge.
Earthworms support terrestrial food chains both as prey and by aerating and enriching soil. The
burrowing of marine polychaetes, which may constitute up to a third of all species in near-shore
environments, encourages the development of ecosystems by enabling water and oxygen to penetrate the
sea floor. In addition to improving soil fertility, annelids serve humans as food and as bait. Scientists observe
annelids to monitor the quality of marine and fresh water. Although blood-letting is no longer in favor with
doctors, some leech species are regarded as endangered species because they have been over-harvested
for this purpose in the last few centuries. Ragworms' jaws are now being studied by engineers as they offer
an exceptional combination of lightness and strength.
Since annelids are soft-bodied, their fossils are rare – mostly jaws and the mineralized tubes that
some of the species secreted. Although some late Ediacaran fossils may represent annelids, the oldest
known fossil that is identified with confidence comes from about 518 million years ago in the early Cambrian
period. Fossils of most modern mobile polychaete groups appeared by the end of the Carboniferous, about
299 million years ago. Scientists disagree about whether some body fossils from the mid Ordovician, about
472 to 461 million years ago, are the remains of oligochaetes, and the earliest certain fossils of the group
appear in the Tertiary period, which began 65 million years ago.
• 1 Classification and diversity
• 2 Distinguishing features
• 3 Description
• 3.1 Segmentation
• 3.2 Body wall, chetae and parapodia
• 3.3 Nervous system and senses
• 3.4 Coelom, locomotion and circulatory system
• 3.5 Respiration
• 3.6 Feeding and excretion
• 3.7 Reproduction and life cycle
• 3.7.1 Asexual reproduction
• 3.7.2 Sexual reproduction
• 4 Ecological significance
• 5 Interaction with humans
• 6 Evolutionary history
• 6.1 Fossil record
• 6.2 Family tree
• 7 References
• 8 Further reading
• 9 External links
[edit] Classification and diversity
There are over 17,000 living annelid species,[3] ranging in size from microscopic to the Australian
giant Gippsland earthworm, which can grow up to 3 metres (9.8 ft) long.[4][5] Although research since 1997
has radically changed scientists' views about the evolutionary family tree of the annelids,[6][7] most
textbooks use the traditional classification into the following sub-groups:[4][8]
• Polychaetes (about 12,000 species[3]). As their name suggests, they have multiple chetae
("hairs") per segment. Polychaetes have parapodia that function as
limbs, and nuchal organs ("nuchal" means "on the neck") that are
thought to be chemosensors.[4] Most are marine animals, although
a few species live in fresh water and even fewer on land.[9]
• Clitellates (about 5,000 species[3]). These have few or no
chetae per segment, and no nuchal organs or parapodia. However,
they have a unique reproductive organ, the ring-shaped clitellum
("pack saddle") round their bodies, which produces a cocoon that
stores and nourishes fertilized eggs until they hatch.[8][10] The
clitellates are sub-divided into:[4]
• Oligochaetes ("with few hairs"), which includes
earthworms. Oligochaetes have a sticky pad in the roof of
the mouth.[4] Most are burrowers that feed on wholly or
partly decomposed organic materials.[9]
• Hirudinea, whose name means "leech-shaped" and
whose best known members are leeches.[4] Marine species
are mostly blood-sucking parasites, mainly on fish, while

An earthworm's
most freshwater species are predators.[9] They have suckers at both ends of their bodies,
and use these to move rather like inchworms.[11]
The Archiannelida, minute annelids that live in the spaces between grains of sediment, were treated
as a separate class because of their simple body structure, but are now regarded as polychaetes.[8] Some
other groups of animals have been classified in various ways, but are now widely regarded as annelids:
• Pogonophora / Siboglinidae were first discovered in 1914, and their lack of a recognizable
gut made it difficult to classify them. They have been classified as a separate phylum, Pogonophora,
or as two phyla, Pogonophora and Vestimentifera. More recently they have been re-classified as a
family, Siboglinidae, within the polychaetes.[9][12]
• The Echiura have a checkered taxonomic history: in the 19th century they were assigned to
the phylum "Gephyrea", which is now empty as its members have been assigned to other phyla; the
Echiura were next regarded as annelids until the 1940s, when they were classified as a phylum in
their own right; but a molecular phylogenetics analysis in 1997 concluded that Echiurans are
• Myzostomida live on crinoids and other echinoderms, mainly as parasites. In the past they
have been regarded as close relatives of the trematode flatworms or of the tardigrades, but in 1998 it
was suggested that they are a sub-group of polychaetes.[9] However, another analysis in 2002
suggested that myzostomids are more closely related to flatworms or to rotifers and

[edit] Distinguishing features

No single feature distinguishes Annelids from other invertebrate phyla, but they have a distinctive
combination of features. Their bodies are long, with segments that are divided externally by shallow ring-like
constrictions called annuli and internally by septa ("partitions") at the same points, although in some species
the septa are incomplete and in a few cases missing. Most of the segments contain the same sets of organs,
although sharing a common gut, circulatory system and nervous system makes them inter-dependent.[4][8]
Their bodies are covered by a cuticle (outer covering) that does not contain cells but is secreted by cells in
the skin underneath, is made of tough but flexible collagen[4] and does not molt[14] – on the other hand
arthropods' cuticles are made of the more rigid α-chitin,[4][15] and molt until the arthropods reach their full
size.[16] Most annelids have closed circulatory systems, where the blood makes its entire circuit via blood
Summary of distinguishing features
Recently merged into
Echiura[1 Sipuncula[ Arth
7] 18] 20]

External Only in a few Yes,

Yes no no
segmentation species in mites

Repetition of In pr
Yes no no Yes
internal organs forms

Septa In most
no no No No
between segments species

Cuticle collagen collagen collagen none α-


no;[14] but some
polychaetes molt
Molting no[23] no[23] no[23] Ye
their jaws, and
leeches molt their

Coelom; but
this is reduced or
2 2 coeloms,
missing in many Coelom only in
Body cavity coeloms, main main and in Hem
leeches and some proboscis
and in proboscis tentacles

Circulatory Closed in
outflow, return via Open Closed O
system most species
branched vein
[edit] Description
[edit] Segmentation Prostomium
Most of an annelid's body consists of segments that are practically
identical, having the same sets of internal organs and external chaetae (Greek Peristomium
χαιτα, meaning "hair") and, in some species, appendages. However, the
frontmost and rearmost sections are not regarded as true segments as they do
not contain the standard sets of organs and do not develop in the same way as Growth zone
the true segments. The frontmost section, called the prostomium (Greek προ-
meaning "in front of" and στομα meaning "mouth") contains the brain and Pygidium
sense organs, while the rearmost, called the pygidium (Greek πυγιδιον,
meaning "little tail") contains the anus, generally on the underside. The first Anus
section behind the prostomium, called the peristomium (Greek περι- meaning
"around" and στομα meaning "mouth"), is regarded by some zoologists as not
a true segment, but in some polychaetes the peristomium has chetae and
appendages like those of other segments.[4]
The segments develop one at a time from a growth zone just ahead of
the pygidium, so that an annelid's youngest segment is just in front of the
growth zone while the peristomium is the oldest. This pattern is called
teloblastic growth.[4] Some groups of annelids, including all leeches,[11] have
fixed maximum numbers of segments, while others add segments throughout
their lives.[8]

Segments of an
The phylum's name is derived from the Latin word
annelus, meaning "little ring".[3]

[edit] Body wall, chetae and parapodia

Annelids' cuticles are made of collagen fibers, usually
in layers that spiral in alternating directions so that the fibers
cross each other. These are secreted by the one-cell deep
epidermis (outermost skin layer). A few marine annelids that
live in tubes lack cuticles, but their tubes have a similar
structure, and mucus-secreting glands in the epidermis
protect their skins.[4] Under the epidermis is the dermis,
which is made of connective tissue, in other words a 10 Coelom
combination of cells and non-cellular materials such as
collagen. Below this are two layers of muscles, which develop
from the lining of the coelom (body cavity): circular muscles
make a segment longer and slimmer when they contract,
while under them are longitudinal muscles, usually four
distinct strips,[14] whose contractions make the segment
shorter and fatter.[4] Some annelids also have oblique
internal muscles that connect the underside of the body to
each side.[14]
The chetae ("hairs") of annelids project out from the
epidermis to provide traction and other capabilities. The
simplest are unjointed and form paired bundles near the top

Cross-section through a typical

and bottom of each side of each segment. The parapodia ("limbs") of annelids that have them often bear
more complex chetae at their tips – for example jointed, comb-like or hooked.[4] Chetae are made of
moderately flexible β-chitin and are formed by follicles, each of which has a chaetoblast ("hair-forming") cell
at the bottom and muscles that can extend or retract the cheta. The chetoblasts produce chetae by forming
microvilli, fine hair-like extensions that increase the area available for secreting the cheta. When the cheta is
complete, the microvilli withdraw into the chetoblast, leaving parallel tunnels that run almost the full length of
the cheta.[4] Hence annelids' chetae are structurally different from the setae ("bristles") of arthropods, which
are made of the more rigid α-chitin, have a single internal cavity, and are mounted on flexible joints in shallow
pits in the cuticle.[4]
Nearly all polychaetes have parapodia that function as limbs, while other major annelid groups lack
them. Parapodia are unjointed paired extensions of the body wall, and their muscles are derived from the
circular muscles of the body. They are often supported internally by one or more large, thick chetae. The
parapodia of burrowing and tube-dwelling polychaetes are often just ridges whose tips bear hooked chetae.
In active crawlers and swimmers the parapodia are often divided into large upper and lower paddles on a
very short trunk, and the paddles are generally fringed with chetae and sometimes with cirri (fused bundles of
cilia) and gills.[14]

[edit] Nervous system and senses

The brain generally forms a ring round the pharynx (throat), consisting of a pair of ganglia (local
control centers) above and in front of the pharynx, linked by nerve cords either side of the pharynx to another
pair of ganglia just below and behind it.[4] The brains of polychaetes are generally in the prostomium, while
those of clitellates are in the peristomium or sometimes the first segment behind the peristomium.[24] In
some very mobile and active polychaetes the brain is enlarged and more complex, with visible hindbrain,
midbrain and forebrain sections.[14] The rest of the central nervous system is generally "ladder-like",
consisting of a pair of nerve cords that run through the bottom part of the body and have in each segment
paired ganglia linked by a transverse connection. From each segmental ganglion a branching system of local
nerves runs into the body wall and then encircles the body.[4] However, in most polychaetes the two main
nerve cords are fused, and in the tube-dwelling genus Owenia the single nerve chord has no ganglia and is
located in the epidermis.[8][25]
As in arthropods, each muscle fiber (cell) is controlled by more than one neuron, and the speed and
power of the fiber's contractions depends on the combined effects of all its neurons. Vertebrates have a
different system, in which one neuron controls a group of muscle fibers.[4] Most annelids' longitudinal nerve
trunks include giant axons (the output signal lines of nerve cells). Their large diameter decreases their
resistance, which allows them to transmit signals exceptionally fast. This enables these worms to withdraw
rapidly from danger by shortening their bodies. Experiments have shown that cutting the giant axons
prevents this escape response but does not affect normal movement.[4]
The sensors are primarily single cells that detect light, chemicals, pressure waves and contact, and
are present on the head, appendages (if any) and other parts of the body.[4] Nuchal ("on the neck") organs
are paired, ciliated structures found only in polychaetes, and are thought to be chemosensors.[14] Some
polychaetes also have various combinations of ocelli ("little eyes") that detect the direction from which light is
coming and camera eyes or compound eyes that can probably form images.[25] The compound eyes
probably evolved independently of arthropods' eyes.[14] Some tube-worms use ocelli widely spread over
their bodies to detect the shadows of fish, so that they can quickly withraw into their tubes.[25] Some
burrowing and tube-dwelling polychaetes have statocysts (tilt and balance sensors) that tell them which way
is down.[25] A few polychaete genera have on the undersides of their heads palps that are used both in
feeding and as "feelers", and some of these also have antennae that are structurally similar but probably are
used mainly as "feelers".[14]
[edit] Coelom, locomotion and circulatory system
Most annelids have a pair of coeloms (body cavities) in each segment, separated from other
segments by septa and from each other by vertical mesenteries. Each septum forms a sandwich with
connective tissue in the middle and mesothelium (membrane that serves as a lining) from the preceding and
following segments on either side. Each mesentery is similar except that the mesothelium is the lining of
each of the pair of coeloms, and the blood vessels and, in polychaetes, the main nerve cords are embedded
in it.[4] The mesothelium is made of modified epitheliomuscular cells,[4] in other words their bodies form part
of the epithelium but their bases extend to form muscle fibers in the body wall.[26] The mesothelium may
also form radial and circular muscles on the septa, and circular muscles around the blood vessels and gut.
Parts of the mesothelium, especially on the outside of the gut, may also form chloragogen cells that perform
similar functions to the livers of vertebrates: producing and storing glycogen and fat; producing the oxygen-
carrier hemoglobin; breaking down proteins; and turning nitrogenous waste products into ammonia and urea
to be excreted.[4]

Peristalsis moves this "worm" to the right

Many annelids move by peristalsis (waves of contraction and expansion that sweep along the body),
[4] or flex the body while using parapodia to crawl or swim.[27] In these animals the septa enable the circular
and longitudinal muscles to change the shape of individual segments, by making each segment a separate
fluid-filled "balloon".[4] However, the septa are often incomplete in annelids that are semi-sessile or that do
not move by peristalsis or by movements of parapodia – for example some move by whipping movements of
the body, some small marine species move by means of cilia (fine muscle-powered hairs) and some
burrowers turn their pharynges (throats) inside out to penetrate the sea-floor and drag themselves into it.[4]
The fluid in the coeloms contains coelomocyte cells that defend the animals against parasites and
infections. In some species coelomocytes may also contain a respiratory pigment – red hemoglobin in some
species, green chlorocruorin in others[14] – and provide oxygen transport within their segments. Respiratory
pigment is also dissolved in the blood plasma. Species with well-developed septa generally also have blood
vessels running all long their bodies above and below the gut, the upper one carrying blood forwards while
the lower one carries it backwards. Networks of capillaries (fine blood vessels) in the body wall and around
the gut transfer blood between the main blood vessels and to parts of the segment that need oxygen and
nutrients. Both of the major vessels, especially the upper one, can pump blood by contracting. In some
annelids the forward end of the upper blood vessel is enlarged with muscles to form a heart, while in the
forward ends of many earthworms some of the vessels that connect the upper and lower main vessels
function as hearts. Species with poorly-developed or no septa generally have no blood vessels and rely on
the circulation within the coelom for delivering nutrients and oxygen.[4]
However, leeches and their closest relatives have a body structure that is very uniform within the
group but significantly different from that of other annelids, including other members of the Clitellata.[11] In
leeches there are no septa, the connective tissue layer of the body wall is so thick that it occupies much of
the body, and the two coeloms are widely separated and run the length of the body. They function as the
main blood vessels, although they are side-by-side rather than upper and lower. However, they are lined with
mesothelium, like the coeloms and unlike the blood vessels of other annelids. Leeches generally use suckers
at their front and rear ends to move like inchworms. The anus is on the upper surface of the pygidium.[11]
[edit] Respiration
In some annelids, including earthworms, all respiration is via the skin. However, many polychaetes
and some clitellates (the group to which earthworms belong) have gills associated with most segments, often
as extensions of the parapodia in polychaetes. The gills of tube-dwellers and burrowers usually cluster
around whichever end has the stronger water flow.[14]

[edit] Feeding and excretion

Feeding structures in the mouth region vary widely, and have little correlation with the animals' diets.
Many polychaetes have a muscular pharynx that can be everted (turned inside out to extend it). In these
animals the foremost few segments often lack septa so that, when the muscles in these segments contract,
the sharp increase in fluid pressure from all these segments everts the pharynx very quickly. Two families,
the Eunicidae and Phyllodocidae, have evolved jaws, which can be used for seizing prey, biting off pieces of
vegetation, or grasping dead and decaying matter. On the other hand some predatory polychaetes have
neither jaws nor eversible pharynges. Selective deposit feeders generally live in tubes on the sea-floor and
use palps to find food particles in the sediment and then wipe them into their mouths. Filter feeders use
"crowns" of palps covered in cilia that wash food particles towards their mouths. Non-selective deposit
feeders ingest soil or marine sediments via mouths that are generally unspecialized. Some clitellates have
sticky pads in the roofs of their mouths, and some of these can evert the pads to capture prey. Leeches often
have an eversible proboscis, or a muscular pharynx with two or three teeth.[14]
The gut is generally an almost straight tube supported by the mesenteries (vertical partitions within
segments), and ends with the anus on the underside of the pygidium.[4] However, in members of the tube-
dwelling family Siboglinidae the gut is blocked by a swollen lining that houses symbiotic bacteria, which can
make up 15% of the worms' total weight. The bacteria convert inorganic matter – such as hydrogen sulfide
and carbon dioxide from hydrothermal vents, or methane from seeps – to organic matter that feeds
themselves and their hosts, while the worms extend their palps into the gas flows to absorb the gases
needed by the bacteria.[14]
Annelids with blood vessels use metanephridia to remove soluble waste products, while those
without use protonephridia.[4] Both of these systems use a two-stage filtration process, in which fluid and
waste products are first extracted and these are filtered again to re-absorb any re-usable materials while
dumping toxic and spent materials as urine. The difference is that protonephridia combine both filtration
stages in the same organ, while metanephridia perform only the second filtration and rely on other
mechanisms for the first – in annelids special filter cells in the walls of the blood vessels let fluids and other
small molecules pass into the coelomic fluid, where it circulates to the metanephridia.[28] In annelids the
points at which fluid enters the protonephridia or metanephridia are on the forward side of a septum while the
second-stage filter and the nephridiopore (exit opening in the body wall) are in the following segment. As a
result the hindmost segment (before the growth zone and pygidium) has no structure that extracts its wastes,
as there is no following segment to filter and discharge them, while the first segment contains an extraction
structure that passes wastes to the second, but does not contain the structures that re-filter and discharge
[edit] Reproduction and life cycle

[edit] Asexual reproduction

This sabellid tubeworm is budding

Polychaetes can reproduce asexually, by dividing into two or more pieces or by budding off a new
individual while the parent remains a complete organism.[4][29] Some oligochaetes, such as Aulophorus
furcatus, seem to reproduce entirely asexually, while others reproduce asexually in summer and sexually in
autumn. Asexual reproduction in oligochaetes is always by dividing into two or more pieces, rather than by
budding.[8][30] However, leeches have never been seen reproducing asexually.[8][31]
Most polychaetes and oligochaetes also use similar mechanisms to regenerate after suffering
damage. Two polychaete genera, Chaetopterus and Dodecaceria, can regenerate from a single segment,
and others can regenerate even if their heads are removed.[8][29] Annelids are the most complex animals
that can regenerate after such severe damage.[32] On the other hand leeches cannot regenerate.[31]
[edit] Sexual reproduction
It is thought that annelids were originally animals with
two separate sexes, which released ova and sperm into the
water via their nephridia.[4] The fertilized eggs develop into Apical tuft (cilia)
trochophore larvae, which live as plankton.[34] Later they sink Prototroch (cilia)
to the sea-floor and metamorphose into miniature adults: the
part of the trochophore between the apical tuft and the Stomach
prototroch becomes the prostomium (head); a small area
round the trochophore's anus becomes the pygidium (tail- Mouth
piece); a narrow band immediately in front of that becomes the Metatroch (cilia)
growth zone that produces new segments; and the rest of the
trochophore becomes the peristomium (the segment that Mesoderm
contains the mouth).[4]
However, the lifecycles of most living polychaetes,
which are almost all marine animals, are unknown, and only /// = cilia
about 25% of the 300+ species whose lifecycles are known
follow this pattern. About 14% use a similar external
fertilization but produce yolk-rich eggs, which reduce the time
the larva needs to spend among the plankton, or eggs from
which miniature adults emerge rather than larvae. The rest
care for the fertilized eggs until they hatch – some by producing
jelly-covered masses of eggs which they tend, some by
attaching the eggs to their bodies and a few species by
keeping the eggs within their bodies until they hatch. These

Trochophore larva[33]
species use a variety of methods for sperm transfer; for example, in some the females collect sperm released
into the water, while in others the males have penes that inject sperm into the female.[34] There is no
guarantee that this is a representative sample of polychaetes' reproductive patterns, and it simply reflects
scientists' current knowledge.[34]
Some polychaetes breed only once in their lives, while others breed almost continuously or through
several breeding seasons. While most polychaetes remain of one sex all their lives, a significant percentage
of species are full hermaphrodites or change sex during their lives. Most polychaetes whose reproduction
has been studied lack permanent gonads, and it is uncertain how they produce ova and sperm. In a few
species the rear of the body splits off and becomes a separate individual that lives just long enough to swim
to a suitable environment, usually near the surface, and spawn.[34]
Most mature clitellates (the group that includes earthworms and leeches) are full hermaphrodites,
although in a few leech species younger adults function as males and become female at maturity. All have
well-developed gonads, and all copulate. Earthworms store their partners' sperm in spermathecae ("sperm
stores") and then the clitellum produces a cocoon that collects ova from the ovaries and then sperm from the
spermathecae. Fertilization and development of earthworm eggs takes place in the cocoon. Leeches' eggs
are fertilized in the ovaries, and then transferred to the cocoon. In all clitellates the cocoon also either
produces yolk when the eggs are fertilized or nutrients while they are developing. All clitellates hatch as
miniature adults rather than larvae.[34]

[edit] Ecological significance

Charles Darwin's book The Formation of Vegetable Mould through the Action of Worms (1881)
presented the first scientific analysis of earthworms' contributions to soil fertility.[35] Some burrow while
others live entirely on the surface, generally in moist leaf litter. The burrowers loosen the soil so that oxygen
and water can penetrate it, and both surface and burrowing worms help to produce soil by mixing organic and
mineral matter, by accelerating the decomposition of organic matter and thus making it more quickly
available to other organisms, and by concentrating minerals and converting them to forms that plants can use
more easily.[36][37] Earthworms are also important prey for birds ranging in size from robins to storks, and
for mammals ranging from shrews to badgers, and in some cases conserving earthworms may be essential
for conserving endangered birds.[38]
Marine annelids may account for over one-third of bottom-dwelling animal species round coral reefs
and in tidal zones.[35] Burrowing species increase the penetration of water and oxygen and water into the
sea-floor sediment, which encourages the growth of populations of bacteria and small animals alongside
their burrows.[39]
Although blood-sucking leeches do little direct harm to their victims, some transmit flagellates that
can be very dangerous to their hosts. Some small tube-dwelling oligochaetes transmit myxosporean
parasites that cause whirling disease in fish.[35]

[edit] Interaction with humans

Earthworms make a significant contribution to soil fertility.[35] The rear end of the Palolo worm, a
marine polychaete that tunnels through coral, detaches in order to spawn at the surface, and the people of
Samoa regard these spawning modules as a delicacy.[35] Anglers sometimes find that worms are more
effective bait than artificial flies, and worms can be kept for several days in a tin lined with damp moss.[40]
Ragworms are commercially important as bait and as food sources for aquaculture, and there have been
proposals to farm them in order to reduce over-fishing of their natural populations.[39] Some marine
polychaetes' predation on molluscs causes serious losses to fishery and aquaculture operations.[35]
Scientists study aquatic annelids to monitor the oxygen content, salinity and pollution levels in fresh
and marine water.[35]
Accounts of the use of leeches for the medically dubious practise of blood-letting have come from
China around 30 AD, India around 200 AD, ancient Rome around 50 AD and later throughout Europe. In the
19th century medical demand for leeches was so high that some areas' stocks were exhausted and other
regions imposed restrictions or bans on exports, and Hirudo medicinalis is treated as an endangered species
by both IUCN and CITES. More recently leeches have been used to assist in microsurgery, and their saliva
has provided anti-inflammatory compounds and several important anticoagulants, one of which also prevents
tumors from spreading.[35]
Ragworms' jaws are strong but much lighter than the hard parts of many other organisms, which are
biomineralized with calcium salts. These advantages have attracted the attention of engineers. Investigations
showed that ragworm jaws are made of unusual proteins that bind strongly to zinc.[41]

[edit] Evolutionary history

[edit] Fossil record
Since annelids are soft-bodied, their fossils are rare.[42] Polychaetes' fossil record consists mainly of
the jaws that some species had and the mineralized tubes that some secreted.[43] Some Ediacaran fossils
such as Dickinsonia in some ways resemble polychaetes, but the similarities are too vague for these fossils
to be classified with confidence.[44] The small shelly fossil Cloudina, from 549 to 542 million years ago, has
been classified by some authors as an annelid, but by others as a cnidarian (i.e. in the phylum to which
jellyfish and sea anemones belong).[45] Until 2008 the earliest fossils widely accepted as annelids were the
polychaetes Canadia and Burgessochaeta, both from Canada's Burgess Shale, formed about 505 million
years ago in the early Cambrian.[46] Myoscolex, found in Australia and a little older than the Burgess Shale,
was possibly an annelid. However, it lacks some typical annelid features and has features which are not
usually found in annelids and some of which are associated with other phyla.[46] Then Simon Conway Morris
and John Peel reported Phragmochaeta from Sirius Passet, about 518 million years old, and concluded that
it was the oldest annelid known to date.[44] There has been vigorous debate about whether the Burgess
Shale fossil Wiwaxia was a mollusc or an annelid.[46] Polychaetes diversified in the early Ordovician, about
488 to 474 million years ago. It is not until the early Ordovician that the first annelid jaws are found, thus the
crown-group cannot have appeared before this date and probably appeared somewhat later.[1] By the end of
the Carboniferous, about 299 million years ago, fossils of most of the modern mobile polychaete groups had
appeared.[46] Many fossil tubes look like those made by modern sessile polychaetes, but the first tubes
clearly produced by polychaetes date from the Jurassic, less than 199 million years ago.[46]
The earliest good evidence for oligochaetes occurs in the Tertiary period, which began 65 million
years ago, and it has been suggested that these animals evolved around the same time as flowering plants
in the early Cretaceous, from 130 to 90 million years ago.[47] A trace fossil consisting of a convoluted burrow
partly filled with small fecal pellets may be evidence that earthworms were present in the early Triassic period
from 251 to 245 million years ago.[47][48] Body fossils going back to the mid Ordovician, from 472 to 461
million years ago, have been tentatively classified as oligochaetes, but these identifications are uncertain and
some have been disputed.[47][49]

[edit] Family tree

some "Canalipalpata"

Sipuncula, previously a separate






"Scolecida" and

some "Scolecida"
Echiura, previously a separate

some "Scolecida"

some "Canalipalpata"

Siblonginidae, previously phylum Pogonophora

some "Canalipalpata"

Annelid groups and phyla incorporated into Annelida (2007; simplified).[6]

Highlights major changes to traditional classifications.
Traditionally the annelids have been divided into two major groups, the polychaetes and clitellates. In
turn the clitellates were divided into oligochaetes, which include earthworms, and hirudinomorphs, whose
best-known members are leeches.[4] For many years there was no clear arrangement of the approximately
80 polychaete families into higher-level groups.[6] In 1997 Greg Rouse and Kristian Fauchald attempted a
"first heuristic step in terms of bringing polychaete systematics to an acceptable level of rigour", based on
anatomical structures, and divided polychaetes into:[51]
• Scolecida, less than 1,000 burrowing species that look rather like earthworms.[52]
• Palpata, the great majority of polychaetes, divided into:
• Canalipalpata, which are distinguished by having long grooved palps that they use
for feeding, and most of which live in tubes.[52]
• Aciculata, the most active polychaetes, which have parapodia reinforced by internal
spines (aciculae).[52]
Also in 1997 Damhnait McHugh, using molecular phylogenetics to compare similarities and
differences in one gene, presented a very different view, in which: the clitellates were an off-shoot of one
branch of the polychaete family tree; the pogonophorans and echiurans, which for a few decades had been
regarded as a separate phyla, were placed on other branches of the polychaete tree.[53] Subsequent
molecular phylogenetics analyses on a similar scale presented similar conclusions.[54]
In 2007 Torsten Struck and colleagues compared 3 genes in 81 taxa, of which 9 were outgroups,[6]
in other words not considered closely related to annelids but included to give an indication of where the
organisms under study are placed on the larger tree of life.[55] For a cross-check the study used an analysis
of 11 genes (including the original 3) in 10 taxa. This analysis agreed that clitellates, pogonophorans and
echiurans were on various branches of the polychaete family tree. It also concluded that the classification of
polychaetes into Scolecida, Canalipalpata and Aciculata was useless, as the members of these alleged
groups were scattered all over the family tree derived from comparing the 81 taxa. In addition, it also placed
sipunculans, generally regarded at the time as a separate phylum, on another branch of the polychaete tree,
and concluded that leeches were a sub-group of oligochaetes rather than their sister-group among the
clitellates.[6] Rouse accepted the analyses based on molecular phylogenetics,[8] and their main conclusions
are now the scientific consensus, although the details of the annelid family tree remain uncertain.[7]
In addition to re-writing the classification of annelids and 3 previously independent phyla, the
molecular phylogenetics analyses undermine the emphasis that decades of previous writings placed on the
importance of segmentation in the classification of invertebrates. Polychaetes, which these analyses found to
be the parent group, have completely segmented bodies, while polychaetes' echiurans and sipunculan
offshoots are not segmented and pogonophores are segmented only in the rear parts of their bodies. It now
seems that segmentation can appear and disappear much more easily in the course of evolution than was
previously thought.[6][53] The 2007 study also noted that the ladder-like nervous system, which is associated
with segmentation, is less universal than previously thought in both annelids and arthropods.[6]
Annelids are members of the
protostomes, one of the two major
superphyla of bilaterian animals – the
other is the deuterostomes, which Protostomia
includes vertebrates.[54] Within the
protostomes, annelids used to be
grouped with arthropods under the
super-group Articulata ("jointed
animals"), as segmentation is obvious in
most members of both phyla. However, Lophotrochozoa
the genes that drive segmentation in
arthropods do not appear to do the
same in annelids. Arthropods and
annelids both have close relatives that
are unsegmented. It is at least as easy Annelida
to assume that they evolved segmented
bodies independently as it is to assume
that the ancestral protostome or Sipuncula
bilaterian was segmented and that
segmentation disappeared in many
descendant phyla.[54] The current view Mollusca
is that annelids are grouped with
molluscs, brachiopods and several other
phyla that have lophophores (fan-like Phoronida
feeding structures) and/or trochophore and Brachiopoda


larvae as members of Lophotrochozoa.[56] Bryzoa may be the most basal phylum (the one that first became
distinctive) within the Lophotrochozoa, and the relationships between the other members are not yet known.
[54] Arthropods are now regarded as members of the Ecdysozoa ("animals that molt"), along with some phyla
that are unsegmented.[54][57]
The "Lophotrochozoa" hypothesis is also supported by the fact that many phyla within this group,
including annelids, molluscs, nemerteans and flatworms, follow a similar pattern in the fertilized egg's
development. When their cells divide after the 4-cell stage, descendants of these 4 cells form a spiral pattern.
In these phyla the "fates" of the embryo's cells, in other words the roles their descendants will play in the
adult animal, are the same and can be predicted from a very early stage.[58] Hence this development pattern
is often described as "spiral determinate cleavage".[59]

[edit] References
1. ^ a b Budd, G. E.; Jensen, S. (2000). "A critical reappraisal of the fossil record of the
bilaterian phyla". Biological reviews of the Cambridge Philosophical Society 75 (2): 253–95.
doi:10.1017/S000632310000548X. PMID 10881389. edit
2. ^ Unabridged: Annelida. Based on the Random House Dictionary, Random
House, Inc., 2009.
3. ^ a b c d e Rouse, G.W. (2002). "Annelida (Segmented Worms)". Encyclopedia of Life
Sciences. John Wiley & Sons, Ltd.. doi:10.1038/npg.els.0001599.
4. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al Ruppert, E.E., Fox,
R.S., and Barnes, R.D. (2004). "Annelida". Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 414–420.
ISBN 0030259827.
5. ^ Lavelle, P. (July 1996). "Diversity of Soil Fauna and Ecosystem Function". Biology
International 33.
46/010008126.pdf. Retrieved 2009-04-20.
6. ^ a b c d e f g h i Struck, T.H.; Schult, N., Kusen, T., Hickman, E., Bleidorn. C., McHugh, D.,
and Halanych, K.M. (2007). "Annelid phylogeny and the status of Sipuncula and Echiura". BMC
Evolutionary Biology 7 (57): 57. doi:10.1186/1471-2148-7-57. PMID 17411434.
7. ^ a b Hutchings, P. (2007). "Book Review: Reproductive Biology and Phylogeny of Annelida".
Integrative and Comparative Biology 47: 788. doi:10.1093/icb/icm008.
8. ^ a b c d e f g h i j k l Rouse, G. (1998). "The Annelida and their close relatives". In Anderson,
D.T.. Invertebrate Zoology. Oxford University Press. pp. 176–179. ISBN 0195513681.
9. ^ a b c d e Rouse, G. (1998). "The Annelida and their close relatives". In Anderson, D.T..
Invertebrate Zoology. Oxford University Press. pp. 179–183. ISBN 0195513681.
10.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Annelida". Invertebrate Zoology (7 ed.).
Brooks / Cole. p. 459. ISBN 0030259827.
11.^ a b c d Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Annelida". Invertebrate Zoology
(7 ed.). Brooks / Cole. pp. 471–482. ISBN 0030259827.
12.^ a b c Halanych, K.M.; Dahlgren, T.G., and McHugh, D. (2002). "Unsegmented Annelids?
Possible Origins of Four Lophotrochozoan Worm Taxa". Integrative and Comparative Biology 42 (3):
678–684. doi:10.1093/icb/42.3.678.
13.^ McHugh, D. (July 1997). "Molecular evidence that echiurans and pogonophorans are
derived annelids". Proceedings of the National Academy of Sciences of the United States of America
94 (15): 8006–8009. doi:10.1073/pnas.94.15.8006. PMID 9223304. PMC 21546. Retrieved 2009-04-02.
14.^ a b c d e f g h i j k l m n o Rouse, G. (1998). "The Annelida and their close relatives". In
Anderson, D.T.. Invertebrate Zoology. Oxford University Press. pp. 183–196. ISBN 0195513681.
15.^ Cutler, B. (August, 1980). "Arthropod cuticle features and arthropod monophyly". Cellular
and Molecular Life Sciences 36 (8): 953. doi:10.1007/BF01953812. Retrieved 2008-09-25.
16.^ a b Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Introduction to Arthropoda".
Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 523–524. ISBN 0030259827.
17.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Echiura and Sipuncula". Invertebrate
Zoology (7 ed.). Brooks / Cole. pp. 490–495. ISBN 0030259827.
18.^ Anderson, D.T., (1998). "The Annelida and their close relatives". In Anderson, D.T.,.
Invertebrate Zoology. Oxford University Press. pp. 183–196. ISBN 0195513681.
19.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Nemertea". Invertebrate Zoology (7
ed.). Brooks / Cole. pp. 271–282. ISBN 0030259827.
20.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Arthropoda". Invertebrate Zoology (7
ed.). Brooks / Cole. pp. 518–521. ISBN 0030259827.
21.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Onychophora and Tardigrada".
Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 505–510. ISBN 0030259827.
22.^ Paxton, H. (June 2005). "Molting polychaete jaws—ecdysozoans are not the only molting
animals". Evolution & Development 7 (4): 337–340. doi:10.1111/j.1525-142X.2005.05039.x.
PMID 15982370.
23.^ a b c Nielsen, C. (September 2003,). "Proposing a solution to the Articulata–Ecdysozoa
controversy". Zoologica Scripta 32 (5): 475–482. doi:10.1046/j.1463-6409.2003.00122.x.
%20vs%20ecdiso.pdf. Retrieved 2009-03-11.
24.^ Jenner, R.A. (2006). "Challenging received wisdoms: Some contributions of the new
microscopy to the new animal phylogeny". Integrative and Comparative Biology 46 (2): 93–103.
25.^ a b c d Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Annelida". Invertebrate Zoology
(7 ed.). Brooks / Cole. pp. 425–429. ISBN 0030259827.
26.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Introduction to Metazoa". Invertebrate
Zoology (7 ed.). Brooks / Cole. pp. 103–104. ISBN 0030259827.
27.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Annelida". Invertebrate Zoology (7 ed.).
Brooks / Cole. pp. 423–425. ISBN 0030259827.
28.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Introduction to Bilateria". Invertebrate
Zoology (7 ed.). Brooks / Cole. pp. 196–224. ISBN 0030259827.
29.^ a b Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Annelida". Invertebrate Zoology (7
ed.). Brooks / Cole. pp. 434–441. ISBN 0030259827.
30.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Annelida". Invertebrate Zoology (7 ed.).
Brooks / Cole. pp. 466–469. ISBN 0030259827.
31.^ a b Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Annelida". Invertebrate Zoology (7
ed.). Brooks / Cole. pp. 477–478. ISBN 0030259827.
32.^ Hickman, Cleveland; Roberts L. Keen S. Larson A. Eisenhour D (2007). Animal Diversity (4
th ed.). New York: Mc Graw Hill. p. 204. ISBN 978-0-07-252844-2.
33.^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Mollusca". Invertebrate Zoology (7 ed.).
Brooks / Cole. pp. 290-291. ISBN 0030259827.
34.^ a b c d e Rouse, G. (1998). "The Annelida and their close relatives". In Anderson, D.T..
Invertebrate Zoology. Oxford University Press. pp. 196–202. ISBN 0195513681.
35.^ a b c d e f g h Siddall, M.E.; Borda, E., and Rouse, G.W. (2004). "Towards a tree of life for
Annelida". In Cracraft, J., and Donoghue, M.J.. Assembling the tree of life. Oxford University Press
US. pp. 237–248. ISBN 0195172345.
id=6lXTP0YU6_kC&pg=PA237&dq=annelid+food+eat. Retrieved 2009-04-02.
36.^ New, T.R. (2005). Invertebrate conservation and agricultural ecosystems . Cambridge
University Press. pp. 44–46. ISBN 0521532019.
id=bwqGf_JK3HcC&pg=PA44&dq=annelid+ecosystem#PPA45,M1. Retrieved 2009-04-02.
37.^ Nancarrow, L.; Taylor, J.H. (1998). The worm book. Ten Speed Press. pp. 2–6.
ISBN 0898159946.
id=U9uQVXCzmGcC&pg=PA139&dq=annelid+ecosystem#PPA6,M1. Retrieved 2009-04-02.
38.^ Edwards, C.A.; Bohlen, P.J. (1996). "Earthworm ecology: communities". Biology and
ecology of arthworms. Springer. pp. 124–126. ISBN 0412561603.
id=ad4rDwD_GhsC&pg=PT7&dq=earliest+oligochaete+clitellate+fossil+. Retrieved 2009-04-12.
39.^ a b Scaps, P. (February 2002). "A review of the biology, ecology and potential use of the
common ragworm Hediste diversicolor". Hydrobiologia 470 (1–3): 203–218.
40.^ Sell, F.E. (re-published 2008). "The humble worm – with a difference". Practical Fresh
Water Fishing. Read Books. pp. 14–15. ISBN 1443761575.
id=t_8tfWmRmmQC&pg=PA14&dq=worm+fishing+angler+bait. Retrieved 2009-04-02.
41.^ "Rags to riches". The Economist. July 2008.
story_id=11785227&CFID=14891129&CFTOKEN=52298400. Retrieved 2009-04-20.
42.^ Rouse, G. (1998). "The Annelida and their close relatives". In Anderson, D.T.. Invertebrate
Zoology. Oxford University Press. p. 202. ISBN 0195513681.
43.^ Briggs, D.E.G.; Kear, A.J. (1993). "Decay and preservation of polychaetes; taphonomic
thresholds in soft-bodied organisms". Paleobiology 19 (1): 107–135. Retrieved 2009-04-12.
44.^ a b Conway Morris, S.; Peel, J.S. (2008). "The earliest annelids: Lower Cambrian
polychaetes from the Sirius Passet Lagerstätte, Peary Land, North Greenland". Acta Palaeontologica
Polonica 53 (1): 137–148. doi:10.4202/app.2008.0110. Retrieved 2009-04-12.
45.^ Miller, A.J. (unpublished; produced in 2004). "A Revised Morphology of Cloudina with
Ecological and Phylogenetic Implications". Retrieved 2009-04-12.
46.^ a b c d e Dzik, J. (2004). "Anatomy and relationships of the Early Cambrian worm
Myoscolex". Zoologica Scripta 33 (1): 57–69. doi:10.1111/j.1463-6409.2004.00136.x.
47.^ a b c Humphreys, G.S. (2003). "Evolution of terrestrial burrowing invertebrates". In Roach,
I.C.. Advances in Regolith. CRC LEME. pp. 211–215. ISBN 0731552210. Retrieved 2009-04-13.
48.^ Retallack, G.J. (1997). "Palaeosols in the upper Narrabeen Group of New South Wales as
evidence of Early Triassic palaeoenvironments without exact modern analogues". Australian Journal
of Earth Sciences 44: 185–201. doi:10.1080/08120099708728303. Retrieved 2009-04-13.
49.^ Conway Morris, S.; Pickerill, R.K. and Harland, T.L. (1982). "A possible annelid from the
Trenton Limestone (Ordovician) of Quebec, with a review of fossil oligochaetes and other annulate
worms". Canadian Journal of Earth Sciences 19: 2150–2157. doi:10.1139/e82-189 (inactive 2010-01-
calyLang=eng&journal=cjes&volume=19&year=0&issue=11&msno=e82-189. Retrieved 2009-04-13.
50.^ A group of worms classified by some as polychaetes and by others as clitellates, see
Rouse & Fauchald (1997) "Cladistics and polychaetes"
51.^ Rouse, G.W.; Fauchald, K. (1997). "Cladistics and polychaetes". Zoologica Scripta 26 (2):
139–204. doi:10.1111/j.1463-6409.1997.tb00412.x.
52.^ a b c Rouse, G.W.; Pleijel, F. and McHugh, D. (August 2002). "Annelida. Annelida.
Segmented worms: bristleworms, ragworms, earthworms, leeches and their allies". The Tree of Life
Web Project. Tree of Life Project. Retrieved 2009-04-13.
53.^ a b McHugh, D. (1997). "Molecular evidence that echiurans and pogonophorans are derived
annelids". Proceedings of the National Academy of Sciences of the United States of America 94 (15):
8006–8009. doi:10.1073/pnas.94.15.8006. PMID 9223304. PMC 21546. Retrieved 2009-04-13.
54.^ a b c d e f Halanych, K.M.. (2004). "The new view of animal phylogeny". Annual Review of
Ecology, Evolution, and Systematics 35: 229–256. doi:10.1146/annurev.ecolsys.35.112202.130124. Retrieved 21009-04-17.
55.^ "Reading trees: A quick review". University of California Museum of Paleontology. Retrieved 2009-04-13.
56.^ Dunn et al., CW; Hejnol, A; Matus, DQ; Pang, K; Browne, WE; Smith, SA; Seaver, E;
Rouse, GW et al. (2008). "Broad phylogenomic sampling improves resolution of the animal tree of
life". Nature 452 (7188): 745–749. doi:10.1038/nature06614. PMID 18322464.
57.^ Aguinaldo, A. M. A.; J. M. Turbeville, L. S. Linford, M. C. Rivera, J. R. Garey, R. A. Raff, &
J. A. Lake (1997). "Evidence for a clade of nematodes, arthropods and other moulting animals".
Nature 387 (6632): 489–493. doi:10.1038/387489a0. PMID 9168109.
58.^ Shankland, M.; Seaver, E.C. (April 2000). "Evolution of the bilaterian body plan: What have
we learned from annelids?". Proceedings of the National Academy of Sciences of the United States
of America 97 (9): 4434–4437. doi:10.1073/pnas.97.9.4434. PMID 10781038. PMC 34316. Retrieved 2009-04-20.
59.^ Pearson, R.D. (2003). "The Determined Embryo". In Hall, B.K., Pearson, R.D., and Müller,
G.B.. Environment, Development, and Evolution. MIT Press. pp. 67–69. ISBN 0-262-08319-1. Retrieved

[edit] Further reading

• Dales, R. P. 1967. Annelids, 2nd edition. London: Hutchinson University Library.
• "Annelid Fossils" (Web page). The Virtual Fossil Museum. 2006. Retrieved May 20, 2006. – Descriptions
and images of annelid fossils from Mazon Creek and the Utah House Range.

Wikimedia Commons has media related to: Annelida

Wikispecies has information related to: Annelida

The Wikibook Dichotomous Key has a page on the topic of

[edit] External links
• Polychaete Larva – Guide to the Marine Zooplankton of south eastern Australia , Tasmanian
Aquaculture & Fisheries Institute
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Natural selection
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For other uses, see Natural Selection (disambiguation).
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Biology portal ·v · d · e
Natural selection is the process by which traits become more or less common in a population due to
consistent effects upon the survival or reproduction of their bearers. It is a key mechanism of evolution.
The natural genetic variation within a population of organisms may cause some individuals to survive
and reproduce more successfully than others in their current environment. For example, the peppered moth
exists in both light and dark colours in the United Kingdom, but during the industrial revolution many of the
trees on which the moths rested became blackened by soot, giving the dark-colored moths an advantage in
hiding from predators. This gave dark-colored moths a better chance of surviving to produce dark-colored
offspring, and in just a few generations the majority of the moths were dark. Factors which affect reproductive
success are also important, an issue which Charles Darwin developed in his ideas on sexual selection.
Natural selection acts on the phenotype, or the observable characteristics of an organism, but the
genetic (heritable) basis of any phenotype which gives a reproductive advantage will become more common
in a population (see allele frequency). Over time, this process can result in adaptations that specialize
populations for particular ecological niches and may eventually result in the emergence of new species. In
other words, natural selection is an important process (though not the only process) by which evolution takes
place within a population of organisms. As opposed to artificial selection, in which humans favor specific
traits, in natural selection the environment acts as a sieve through which only certain variations can pass.
Natural selection is one of the cornerstones of modern biology. The term was introduced by Darwin
in his influential 1859 book On the Origin of Species,[1] in which natural selection was described as
analogous to artificial selection, a process by which animals and plants with traits considered desirable by
human breeders are systematically favored for reproduction. The concept of natural selection was originally
developed in the absence of a valid theory of heredity; at the time of Darwin's writing, nothing was known of
modern genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical and
molecular genetics is termed the modern evolutionary synthesis. Natural selection remains the primary
explanation for adaptive evolution.
• 1 General principles
• 1.1 Nomenclature and usage
• 1.2 Fitness
• 1.3 Types of selection
• 1.4 Sexual selection
• 1.5 Examples of natural selection
• 2 Evolution by means of natural selection
• 2.1 Speciation
• 3 Historical development
• 3.1 Pre-Darwinian theories
• 3.2 Darwin's theory
• 3.3 Modern evolutionary synthesis
• 4 Impact of the idea
• 4.1 Cell and molecular biology
• 4.2 Social and psychological theory
• 4.3 Information and systems theory
• 5 Genetic basis of natural selection
• 5.1 Genotype and phenotype
• 5.2 Directionality of selection
• 5.3 Selection and genetic variation
[edit] General principles

Darwin's illustrations of beak variation in the finches of the Galápagos Islands, which hold 13 closely
related species that differ most markedly in the shape of their beaks. The beak of each species is suited to its
preferred food, suggesting that beak shapes evolved by natural selection.
Natural variation occurs among the individuals of any population of organisms. Many of these
differences do not affect survival (such as differences in eye color in humans), but some differences may
improve the chances of survival of a particular individual. A rabbit that runs faster than others may be more
likely to escape from predators, and algae that are more efficient at extracting energy from sunlight will grow
faster. Something that increases an animal's survival will often also include its reproductive rate; however,
sometimes there is a trade-off between survival and current reproduction. Ultimately, what matters is total
lifetime reproduction of the animal.
If the traits that give these individuals a reproductive advantage are also heritable, that is, passed
from parent to child, then there will be a slightly higher proportion of fast rabbits or efficient algae in the next
generation. This is known as differential reproduction. Even if the reproductive advantage is very slight, over
many generations any heritable advantage will become dominant in the population. In this way the natural
environment of an organism "selects" for traits that confer a reproductive advantage, causing gradual
changes or evolution of life. This effect was first described and named by Charles Darwin.
The concept of natural selection predates the understanding of genetics, the mechanism of heredity
for all known life forms. In modern terms, selection acts on an organism's phenotype, or observable
characteristics, but it is the organism's genetic make-up or genotype that is inherited. The phenotype is the
result of the genotype and the environment in which the organism lives (see Genotype-phenotype
This is the link between natural selection and genetics, as described in the modern evolutionary
synthesis. Although a complete theory of evolution also requires an account of how genetic variation arises in
the first place (such as by mutation and sexual reproduction) and includes other evolutionary mechanisms
(such as genetic drift and gene flow), natural selection appears to be the most important mechanism for
creating complex adaptations in nature.

[edit] Nomenclature and usage

The term natural selection has slightly different definitions in different contexts. It is most often
defined to operate on heritable traits, because these are the traits that directly participate in evolution.
However, natural selection is "blind" in the sense that changes in phenotype (physical and behavioral
characteristics) can give a reproductive advantage regardless of whether or not the trait is heritable (non
heritable traits can be the result of environmental factors or the life experience of the organism).
Following Darwin's primary usage[1] the term is often used to refer to both the evolutionary
consequence of blind selection and to its mechanisms.[2][3] It is sometimes helpful to explicitly distinguish
between selection's mechanisms and its effects; when this distinction is important, scientists define "natural
selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce",
without regard to whether the basis of the selection is heritable. This is sometimes referred to as "phenotypic
natural selection".[4]
Traits that cause greater reproductive success of an organism are said to be selected for, whereas
those that reduce success are selected against. Selection for a trait may also result in the selection of other
correlated traits that do not themselves directly influence reproductive advantage. This may occur as a result
of pleiotropy or gene linkage.[5]

[edit] Fitness
Main article: Fitness (biology)
The concept of fitness is central to natural selection. Broadly, individuals which are more "fit" have
better potential for survival, as in the well-known phrase "survival of the fittest". However, as with natural
selection above, the precise meaning of the term is much more subtle, and Richard Dawkins manages in his
later books to avoid it entirely. (He devotes a chapter of his book, The Extended Phenotype, to discussing the
various senses in which the term is used). Modern evolutionary theory defines fitness not by how long an
organism lives, but by how successful it is at reproducing. If an organism lives half as long as others of its
species, but has twice as many offspring surviving to adulthood, its genes will become more common in the
adult population of the next generation.
Though natural selection acts on individuals, the effects of chance mean that fitness can only really
be defined "on average" for the individuals within a population. The fitness of a particular genotype
corresponds to the average effect on all individuals with that genotype. Very low-fitness genotypes cause
their bearers to have few or no offspring on average; examples include many human genetic disorders like
cystic fibrosis.
Since fitness is an averaged quantity, it is also possible that a favorable mutation arises in an
individual that does not survive to adulthood for unrelated reasons. Fitness also depends crucially upon the
environment. Conditions like sickle-cell anemia may have low fitness in the general human population, but
because the sickle-cell trait confers immunity from malaria, it has high fitness value in populations which
have high malaria infection rates.

[edit] Types of selection

Natural selection can act on any heritable phenotypic trait, and selective pressure can be produced
by any aspect of the environment, including sexual selection and competition with members of the same or
other species. However, this does not imply that natural selection is always directional and results in adaptive
evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants.
The unit of selection can be the individual or it can be another level within the hierarchy of biological
organisation, such as genes, cells, and kin groups. There is still debate about whether natural selection acts
at the level of groups or species to produce adaptations that benefit a larger, non-kin group. Likewise, there
is debate as to whether selection at the molecular level prior to gene mutations and fertilization of the zygote
should be ascribed to conventional natural selection because traditionally natural selection is an
environmental and exterior force that acts upon a phenotype typically after birth. Some science journalists
distinguish natural selection from gene selection by informally referencing selection of mutations as "pre-
Selection at a different level such as the gene can result in an increase in fitness for that gene, while
at the same time reducing the fitness of the individuals carrying that gene, in a process called intragenomic
conflict. Overall, the combined effect of all selection pressures at various levels determines the overall fitness
of an individual, and hence the outcome of natural selection.

The life cycle of a sexually reproducing organism. Various components of natural selection are
indicated for each life stage.[7]
Natural selection occurs at every life stage of an individual. An individual organism must survive until
adulthood before it can reproduce, and selection of those that reach this stage is called viability selection. In
many species, adults must compete with each other for mates via sexual selection, and success in this
competition determines who will parent the next generation. When individuals can reproduce more than
once, a longer survival in the reproductive phase increases the number of offspring, called survival selection.
The fecundity of both females and males (for example, giant sperm in certain species of Drosophila)
[8] can be limited via "fecundity selection". The viability of produced gametes can differ, while intragenomic
conflicts such as meiotic drive between the haploid gametes can result in gametic or "genic selection".
Finally, the union of some combinations of eggs and sperm might be more compatible than others; this is
termed compatibility selection.

[edit] Sexual selection

Main article: Sexual selection
It is useful to distinguish between "ecological selection" and "sexual selection". Ecological selection
covers any mechanism of selection as a result of the environment (including relatives, e.g. kin selection,
competition, and infanticide), while "sexual selection" refers specifically to competition for mates.[9]
Sexual selection can be intrasexual, as in cases of competition among individuals of the same sex in
a population, or intersexual, as in cases where one sex controls reproductive access by choosing among a
population of available mates. Most commonly, intrasexual selection involves male–male competition and
intersexual selection involves female choice of suitable males, due to the generally greater investment of
resources for a female than a male in a single offspring. However, some species exhibit sex-role reversed
behavior in which it is males that are most selective in mate choice; the best-known examples of this pattern
occur in some fishes of the family Syngnathidae, though likely examples have also been found in amphibian
and bird species.[10]
Some features that are confined to one sex only of a particular species can be explained by selection
exercised by the other sex in the choice of a mate, for example, the extravagant plumage of some male birds.
Similarly, aggression between members of the same sex is sometimes associated with very distinctive
features, such as the antlers of stags, which are used in combat with other stags. More generally, intrasexual
selection is often associated with sexual dimorphism, including differences in body size between males and
females of a species.[11]
[edit] Examples of natural selection
Resistance to antibiotics is increased though the survival of individuals which are immune to the
effects of the antibiotic, whose offspring then inherit the resistance, creating a new population of resistant
A well-known example of natural selection in action is the development of antibiotic resistance in
microorganisms. Since the discovery of penicillin in 1928 by Alexander Fleming, antibiotics have been used
to fight bacterial diseases. Natural populations of bacteria contain, among their vast numbers of individual
members, considerable variation in their genetic material, primarily as the result of mutations. When exposed
to antibiotics, most bacteria die quickly, but some may have mutations that make them slightly less
susceptible. If the exposure to antibiotics is short, these individuals will survive the treatment. This selective
elimination of maladapted individuals from a population is natural selection.
These surviving bacteria will then reproduce again, producing the next generation. Due to the
elimination of the maladapted individuals in the past generation, this population contains more bacteria that
have some resistance against the antibiotic. At the same time, new mutations occur, contributing new genetic
variation to the existing genetic variation. Spontaneous mutations are very rare, and advantageous mutations
are even rarer. However, populations of bacteria are large enough that a few individuals will have beneficial
mutations. If a new mutation reduces their susceptibility to an antibiotic, these individuals are more likely to
survive when next confronted with that antibiotic.
Given enough time, and repeated exposure to the antibiotic, a population of antibiotic-resistant
bacteria will emerge. This new changed population of antibiotic-resistant bacteria is optimally adapted to the
context it evolved in. At the same time, it is not necessarily optimally adapted any more to the old antibiotic
free environment. The end result of natural selection is two populations that are both optimally adapted to
their specific environment, while both perform substandard in the other environment.
The widespread use and misuse of antibiotics has resulted in increased microbial resistance to
antibiotics in clinical use, to the point that the methicillin-resistant Staphylococcus aureus (MRSA) has been
described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing
drugs.[12] Response strategies typically include the use of different, stronger antibiotics; however, new
strains of MRSA have recently emerged that are resistant even to these drugs.[13]
This is an example of what is known as an evolutionary arms race, in which bacteria continue to
develop strains that are less susceptible to antibiotics, while medical researchers continue to develop new
antibiotics that can kill them. A similar situation occurs with pesticide resistance in plants and insects. Arms
races are not necessarily induced by man; a well-documented example involves the spread of a gene in the
butterfly Hypolimnas bolina suppressing male-killing activity by Wolbachia bacteria parasites on the island of
Samoa, where the spread of the gene is known to have occurred over a period of just five years [14]

[edit] Evolution by means of natural selection

Main articles: Evolution and Darwinism
A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation, is the
presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of
mutations, recombinations and alterations in the karyotype (the number, shape, size and internal
arrangement of the chromosomes). Any of these changes might have an effect that is highly advantageous or
highly disadvantageous, but large effects are very rare. In the past, most changes in the genetic material
were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in a
synonymous substitution. However, recent research suggests that many mutations in non-coding DNA do
have slight deleterious effects.[15][16] Although both mutation rates and average fitness effects of mutations
are dependent on the organism, estimates from data in humans have found that a majority of mutations are
slightly deleterious.[17]

The exuberant tail of the peacock is thought to be the result of sexual selection by females. This
peacock is an albino; selection against albinos in nature is intense because they are easily spotted by
predators or are unsuccessful in competition for mates.
By the definition of fitness, individuals with greater fitness are more likely to contribute offspring to the
next generation, while individuals with lesser fitness are more likely to die early or fail to reproduce. As a
result, alleles which on average result in greater fitness become more abundant in the next generation, while
alleles which generally reduce fitness become rarer. If the selection forces remain the same for many
generations, beneficial alleles become more and more abundant, until they dominate the population, while
alleles with a lesser fitness disappear. In every generation, new mutations and re-combinations arise
spontaneously, producing a new spectrum of phenotypes. Therefore, each new generation will be enriched
by the increasing abundance of alleles that contribute to those traits that were favored by selection,
enhancing these traits over successive generations.
Some mutations occur in so-called regulatory genes. Changes in these can have large effects on the
phenotype of the individual because they regulate the function of many other genes. Most, but not all,
mutations in regulatory genes result in non-viable zygotes. Examples of nonlethal regulatory mutations occur
in HOX genes in humans, which can result in a cervical rib[18] or polydactyly, an increase in the number of
fingers or toes.[19] When such mutations result in a higher fitness, natural selection will favor these
phenotypes and the novel trait will spread in the population.

X-ray of the left hand of a ten year old boy with polydactyly.
Established traits are not immutable; traits that have high fitness in one environmental context may
be much less fit if environmental conditions change. In the absence of natural selection to preserve such a
trait, it will become more variable and deteriorate over time, possibly resulting in a vestigial manifestation of
the trait, also called evolutionary baggage. In many circumstances, the apparently vestigial structure may
retain a limited functionality, or may be co-opted for other advantageous traits in a phenomenon known as
preadaptation. A famous example of a vestigial structure, the eye of the blind mole rat, is believed to retain
function in photoperiod perception.[20]

[edit] Speciation
Main article: Speciation
Speciation requires selective mating, which result in a reduced gene flow. Selective mating can be
the result of 1. Geographic isolation, 2. Behavioral isolation, or 3. Temporal isolation. For example, a change
in the physical environment (geographic isolation by an extrinsic barrier) would follow number 1, a change in
camouflage for number 2 or a shift in mating times (i.e., one species of deer shifts location and therefore
changes its "rut") for number 3.[citation needed]
Over time, these subgroups might diverge radically to become different species, either because of
differences in selection pressures on the different subgroups, or because different mutations arise
spontaneously in the different populations, or because of founder effects – some potentially beneficial alleles
may, by chance, be present in only one or other of two subgroups when they first become separated. A
lesser-known mechanism of speciation occurs via hybridization, well-documented in plants and occasionally
observed in species-rich groups of animals such as cichlid fishes.[21] Such mechanisms of rapid speciation
can reflect a mechanism of evolutionary change known as punctuated equilibrium, which suggests that
evolutionary change and particularly speciation typically happens quickly after interrupting long periods of
Genetic changes within groups result in increasing incompatibility between the genomes of the two
subgroups, thus reducing gene flow between the groups. Gene flow will effectively cease when the distinctive
mutations characterizing each subgroup become fixed. As few as two mutations can result in speciation: if
each mutation has a neutral or positive effect on fitness when they occur separately, but a negative effect
when they occur together, then fixation of these genes in the respective subgroups will lead to two
reproductively isolated populations. According to the biological species concept, these will be two different

[edit] Historical development

Main articles: History of evolutionary thought, Inception of Darwin's theory, and Development of
Darwin's theory
The modern theory of natural selection derives from the work of Charles Darwin in the nineteenth
[edit] Pre-Darwinian theories
Several ancient philosophers expressed the idea that nature produces a huge variety of creatures,
apparently randomly, and that only those creatures survive that manage to provide for themselves and
reproduce successfully; well-known examples include Empedocles[22] and his intellectual successor,
Lucretius,[23] while related ideas were later refined by Aristotle.[24] The struggle for existence was later
described by Al-Jahiz, who argued that environmental factors influence animals to develop new
characteristics to ensure survival.[25][26][27]
Abu Rayhan Biruni described the idea of artificial selection and argued that nature works in much the
same way.[28] Similar ideas were later expressed by Nasir al-Din Tusi[29] and Ibn Khaldun.[30][31] Such
classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis[32] and others,
including Charles Darwin's grandfather Erasmus Darwin. While these forerunners had an influence on
Darwinism, they later had little influence on the trajectory of evolutionary thought after Charles Darwin.
Until the early 19th century, the prevailing view in Western societies was that differences between
individuals of a species were uninteresting departures from their Platonic idealism (or typus) of created kinds.
However, the theory of uniformitarianism in geology promoted the idea that simple, weak forces could act
continuously over long periods of time to produce radical changes in the Earth's landscape. The success of
this theory raised awareness of the vast scale of geological time and made plausible the idea that tiny,
virtually imperceptible changes in successive generations could produce consequences on the scale of
differences between species.
Early 19th century evolutionists such as Jean Baptiste Lamarck suggested the inheritance of
acquired characteristics as a mechanism for evolutionary change; adaptive traits acquired by an organism
during its lifetime could be inherited by that organism's progeny, eventually causing transmutation of species.
[33] This theory has come to be known as Lamarckism and was an influence on the anti-genetic ideas of the
Stalinist Soviet biologist Trofim Lysenko.[34]

[edit] Darwin's theory

In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for
adaptation and speciation. He defined natural selection as the "principle by which each slight variation [of a
trait], if useful, is preserved".[35] The concept was simple but powerful: individuals best adapted to their
environments are more likely to survive and reproduce. As long as there is some variation between them,
there will be an inevitable selection of individuals with the most advantageous variations. If the variations are
inherited, then differential reproductive success will lead to a progressive evolution of particular populations
of a species, and populations that evolve to be sufficiently different eventually become different species.[36]
Darwin's ideas were inspired by the observations that he had made on the Beagle voyage, and by
the work of a political economist, the Reverend Thomas Malthus, who in An Essay on the Principle of
Population, noted that population (if unchecked) increases exponentially whereas the food supply grows only
arithmetically; thus inevitable limitations of resources would have demographic implications, leading to a
"struggle for existence".[37] When Darwin read Malthus in 1838 he was already primed by his work as a
naturalist to appreciate the "struggle for existence" in nature and it struck him that as population outgrew
resources, "favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The
result of this would be the formation of new species."[38]
Here is Darwin's own summary of the idea, which can be found in the fourth chapter of the Origin:
If during the long course of ages and under varying conditions of life, organic beings vary at all
in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the
high geometrical powers of increase of each species, at some age, season, or year, a severe struggle
for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of
all organic beings to each other and to their conditions of existence, causing an infinite diversity in
structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary
fact if no variation ever had occurred useful to each being's own welfare, in the same way as so many
variations have occurred useful to man. But if variations useful to any organic being do occur,
assuredly individuals thus characterised will have the best chance of being preserved in the struggle
for life; and from the strong principle of inheritance they will tend to produce offspring similarly
characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection.
Once he had his theory "by which to work", Darwin was meticulous about gathering and refining
evidence as his "prime hobby" before making his idea public. He was in the process of writing his "big book"
to present his researches when the naturalist Alfred Russel Wallace independently conceived of the principle
and described it in an essay he sent to Darwin to forward to Charles Lyell. Lyell and Joseph Dalton Hooker
decided (without Wallace's knowledge) to present his essay together with unpublished writings which Darwin
had sent to fellow naturalists, and On the Tendency of Species to form Varieties; and on the Perpetuation of
Varieties and Species by Natural Means of Selection was read to the Linnean Society announcing co-
discovery of the principle in July 1858.[39] Darwin published a detailed account of his evidence and
conclusions in On the Origin of Species in 1859. In the 3rd edition of 1861 Darwin acknowledged that others
— notably William Charles Wells in 1813, and Patrick Matthew in 1831 — had proposed similar ideas, but had
neither developed them nor presented them in notable scientific publications.[40]
Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding,
which he called "artificial selection"; in his early manuscripts he referred to a 'Nature' which would do the
selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly
formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that [it] has
been the main, but not exclusive means of modification."[41] In a letter to Charles Lyell in September 1860,
Darwin regretted the use of the term "Natural Selection", preferring the term "Natural Preservation".[42]
For Darwin and his contemporaries, natural selection was essentially synonymous with evolution by
natural selection. After the publication of On the Origin of Species, educated people generally accepted that
evolution had occurred in some form. However, natural selection remained controversial as a mechanism,
partly because it was perceived to be too weak to explain the range of observed characteristics of living
organisms, and partly because even supporters of evolution balked at its "unguided" and non-progressive
nature,[43] a response that has been characterized as the single most significant impediment to the idea's
However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert
Spencer introduced the term survival of the fittest, which became a popular summary of the theory.[45] The
fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to
natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer, of the Survival of
the Fittest, is more accurate, and is sometimes equally convenient."[46] Although the phrase is still often
used by non-biologists, modern biologists avoid it because it is tautological if "fittest" is read to mean
"functionally superior" and is applied to individuals rather than considered as an averaged quantity over

[edit] Modern evolutionary synthesis

Main article: Modern evolutionary synthesis
Natural selection relies crucially on the idea of heredity, but it was developed long before the basic
concepts of genetics. Although the Austrian monk Gregor Mendel, the father of modern genetics, was a
contemporary of Darwin's, his work would lie in obscurity until the early 20th century. Only after the
integration of Darwin's theory of evolution with a complex statistical appreciation of Gregor Mendel's 're-
discovered' laws of inheritance did natural selection become generally accepted by scientists.
The work of Ronald Fisher (who developed the required mathematical language and The Genetical
Theory of Natural Selection),[2] J.B.S. Haldane (who introduced the concept of the "cost" of natural
selection),[48] Sewall Wright (who elucidated the nature of selection and adaptation),[49] Theodosius
Dobzhansky (who established the idea that mutation, by creating genetic diversity, supplied the raw material
for natural selection: see Genetics and the Origin of Species),[50] William Hamilton (who conceived of kin
selection), Ernst Mayr (who recognised the key importance of reproductive isolation for speciation: see
Systematics and the Origin of Species)[51] and many others formed the modern evolutionary synthesis. This
synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today.

[edit] Impact of the idea

Darwin's ideas, along with those of Adam Smith and Karl Marx, had a profound influence on 19th
century thought. Perhaps the most radical claim of the theory of evolution through natural selection is that
"elaborately constructed forms, so different from each other, and dependent on each other in so complex a
manner" evolved from the simplest forms of life by a few simple principles. This claim inspired some of
Darwin's most ardent supporters—and provoked the most profound opposition. The radicalism of natural
selection, according to Stephen Jay Gould,[52] lay in its power to "dethrone some of the deepest and most
traditional comforts of Western thought". In particular, it challenged long-standing beliefs in such concepts as
a special and exalted place for humans in the natural world and a benevolent creator whose intentions were
reflected in nature's order and design.
[edit] Cell and molecular biology
In the 19th century, Wilhelm Roux, a founder of modern embryology, wrote a book entitled « Der
Kampf der Teile im Organismus » (The struggle of parts in the organism) in which he suggested that the
development of an organism results from a Darwinian competition between the parts of the embryo,
occurring at all levels, from molecules to organs. In recent years, a modern version of this theory has been
proposed by Jean-Jacques Kupiec. According to this cellular Darwinism, stochasticity at the molecular level
generates diversity in cell types whereas cell interactions impose a characteristic order on the developing

[edit] Social and psychological theory

The social implications of the theory of evolution by natural selection also became the source of
continuing controversy. Friedrich Engels, a German political philosopher and co-originator of the ideology of
communism, wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind when he
showed that free competition, the struggle for existence, which the economists celebrate as the highest
historical achievement, is the normal state of the animal kingdom".[53] Interpretation of natural selection as
necessarily 'progressive', leading to increasing 'advances' in intelligence and civilisation, was used as a
justification for colonialism and policies of eugenics, as well as broader sociopolitical positions now described
as Social Darwinism. Konrad Lorenz won the Nobel Prize in Physiology or Medicine in 1973 for his analysis