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Lucretius
In 1842, Charles Darwin penned his first sketch of On the Origin of Species.[35]
Main article: History of evolutionary thought
Classical times
The proposal that one type of organism could descend from another type goes back
to some of the first pre-Socratic Greek philosophers, such as Anaximander and E
mpedocles.[36] Such proposals survived into Roman times. The poet and philosophe
r Lucretius followed Empedocles in his masterwork De rerum natura (On the Nature
of Things).[37][38]
Medieval
In contrast to these materialistic views, Aristotelianism considered all natural
things as actualisations of fixed natural possibilities, known as forms.[39][40
] This was part of a medieval teleological understanding of nature in which all
things have an intended role to play in a divine cosmic order. Variations of thi
s idea became the standard understanding of the Middle Ages and were integrated
into Christian learning, but Aristotle did not demand that real types of organis
ms always correspond one-for-one with exact metaphysical forms and specifically
gave examples of how new types of living things could come to be.[41]
Pre-Darwinian
In the 17th century, the new method of modern science rejected the Aristotelian
approach. It sought explanations of natural phenomena in terms of physical laws
that were the same for all visible things and that did not require the existence
of any fixed natural categories or divine cosmic order. However, this new appro
ach was slow to take root in the biological sciences, the last bastion of the co
ncept of fixed natural types. John Ray applied one of the previously more genera
l terms for fixed natural types, "species," to plant and animal types, but he st
rictly identified each type of living thing as a species and proposed that each
species could be defined by the features that perpetuated themselves generation
after generation.[42] The biological classification introduced by Carl Linnaeus
in 1735 explicitly recognized the hierarchical nature of species relationships,
but still viewed species as fixed according to a divine plan.[43]
Other naturalists of this time speculated on the evolutionary change of species
over time according to natural laws. In 1751, Pierre Louis Maupertuis wrote of n
atural modifications occurring during reproduction and accumulating over many ge
nerations to produce new species.[44] Georges-Louis Leclerc, Comte de Buffon sug
gested that species could degenerate into different organisms, and Erasmus Darwi
n proposed that all warm-blooded animals could have descended from a single micr
oorganism (or "filament").[45] The first full-fledged evolutionary scheme was Je
an-Baptiste Lamarck's "transmutation" theory of 1809,[46] which envisaged sponta
neous generation continually producing simple forms of life that developed great
er complexity in parallel lineages with an inherent progressive tendency, and po
stulated that on a local level these lineages adapted to the environment by inhe
riting changes caused by their use or disuse in parents.[47][48] (The latter pro
cess was later called Lamarckism.)[47][49][50][51] These ideas were condemned by
established naturalists as speculation lacking empirical support. In particular
, Georges Cuvier insisted that species were unrelated and fixed, their similarit
ies reflecting divine design for functional needs. In the meantime, Ray's ideas
of benevolent design had been developed by William Paley into the Natural Theolo
gy or Evidences of the Existence and Attributes of the Deity (1802), which propo
sed complex adaptations as evidence of divine design and which was admired by Ch
arles Darwin.[52][53][54]
Darwinian revolution
The crucial break from the concept of constant typological classes or types in b
iology came with the theory of evolution through natural selection, which was fo
rmulated by Charles Darwin in terms of variable populations. Partly influenced b
y An Essay on the Principle of Population (1798) by Thomas Robert Malthus, Darwi
n noted that population growth would lead to a "struggle for existence" in which
favorable variations prevailed as others perished. In each generation, many off
spring fail to survive to an age of reproduction because of limited resources. T
his could explain the diversity of plants and animals from a common ancestry thr
ough the working of natural laws in the same way for all types of organism.[55][
56][57][58] Darwin developed his theory of "natural selection" from 1838 onwards
and was writing up his "big book" on the subject when Alfred Russel Wallace sen
t him a version of virtually the same theory in 1858. Their separate papers were
presented together at a 1858 meeting of the Linnean Society of London.[59] At t
he end of 1859, Darwin's publication of his "abstract" as On the Origin of Speci
es explained natural selection in detail and in a way that led to an increasingl
y wide acceptance of concepts of evolution. Thomas Henry Huxley applied Darwin's
ideas to humans, using paleontology and comparative anatomy to provide strong e
vidence that humans and apes shared a common ancestry. Some were disturbed by th
is since it implied that humans did not have a special place in the universe.[60
]
Pangenesis
The mechanisms of reproductive heritability and the origin of new traits remaine
d a mystery. Towards this end, Darwin developed his provisional theory of pangen
esis.[61] In 1865, Gregor Mendel reported that traits were inherited in a predic
table manner through the independent assortment and segregation of elements (lat
er known as genes). Mendel's laws of inheritance eventually supplanted most of D
arwin's pangenesis theory.[62] August Weismann made the important distinction be
tween germ cells that give rise to gametes (such as sperm and egg cells) and the
somatic cells of the body, demonstrating that heredity passes through the germ
line only. Hugo de Vries connected Darwin's pangenesis theory to Weismann's germ
/soma cell distinction and proposed that Darwin's pangenes were concentrated in
the cell nucleus and when expressed they could move into the cytoplasm to change
the cells structure. De Vries was also one of the researchers who made Mendel's
work well-known, believing that Mendelian traits corresponded to the transfer o
f heritable variations along the germline.[63] To explain how new variants origi
nate, de Vries developed a mutation theory that led to a temporary rift between
those who accepted Darwinian evolution and biometricians who allied with de Vrie
s.[48][64][65] In the 1930s, pioneers in the field of population genetics, such
as Ronald Fisher, Sewall Wright and J. B. S. Haldane set the foundations of evol
ution onto a robust statistical philosophy. The false contradiction between Darw
in's theory, genetic mutations, and Mendelian inheritance was thus reconciled.[6
6]
The 'modern synthesis'
In the 1920s and 1930s the so-called modern synthesis connected natural selectio
n, mutation theory, and Mendelian inheritance into a unified theory that applied
generally to any branch of biology. The modern synthesis explained patterns obs
erved across species in populations, through fossil transitions in palaeontology
, and complex cellular mechanisms in developmental biology.[48][67] The publicat
ion of the structure of DNA by James Watson and Francis Crick in 1953 demonstrat
ed a physical mechanism for inheritance.[68] Molecular biology improved our unde
rstanding of the relationship between genotype and phenotype. Advancements were
also made in phylogenetic systematics, mapping the transition of traits into a c
omparative and testable framework through the publication and use of evolutionar
y trees.[69][70] In 1973, evolutionary biologist Theodosius Dobzhansky penned th
at "nothing in biology makes sense except in the light of evolution," because it
has brought to light the relations of what first seemed disjointed facts in nat
ural history into a coherent explanatory body of knowledge that describes and pr
edicts many observable facts about life on this planet.[71]
Further syntheses
Since then, the modern synthesis has been further extended to explain biological
phenomena across the full and integrative scale of the biological hierarchy, fr
om genes to species. This extension, known as evolutionary developmental biology
and informally called "evo-devo," emphasises how changes between generations (e
volution) acts on patterns of change within individual organisms (development).[
72][73][74] Since the beginning of the 21st century and in light of discoveries
made in recent decades, some biologists have argued for an extended evolutionary
synthesis, which would account for the effects of non-genetic inheritance modes
, such as epigenetics, parental effects, ecological and cultural inheritance, an
d evolvability.[75][76]
Heredity
Further information: Introduction to genetics, Genetics, Heredity, and Reaction
norm
This diagram illustrates the twofold cost of sex. If each individual were to con
tribute to the same number of offspring (two), (a) the sexual population remains
the same size each generation, where the (b) Asexual reproduction population do
ubles in size each generation.
The two-fold cost of sex was first described by John Maynard Smith.[112] The fir
st cost is that in sexually dimorphic species only one of the two sexes can bear
young. (This cost does not apply to hermaphroditic species, like most plants an
d many invertebrates.) The second cost is that any individual who reproduces sex
ually can only pass on 50% of its genes to any individual offspring, with even l
ess passed on as each new generation passes.[113] Yet sexual reproduction is the
more common means of reproduction among eukaryotes and multicellular organisms.
The Red Queen hypothesis has been used to explain the significance of sexual re
production as a means to enable continual evolution and adaptation in response t
o coevolution with other species in an ever-changing environment.[113][114][115]
[116]
Gene flow
Further information: Gene flow
Gene flow is the exchange of genes between populations and between species.[117]
It can therefore be a source of variation that is new to a population or to a s
pecies. Gene flow can be caused by the movement of individuals between separate
populations of organisms, as might be caused by the movement of mice between inl
and and coastal populations, or the movement of pollen between heavy metal toler
ant and heavy metal sensitive populations of grasses.
Gene transfer between species includes the formation of hybrid organisms and hor
izontal gene transfer. Horizontal gene transfer is the transfer of genetic mater
ial from one organism to another organism that is not its offspring; this is mos
t common among bacteria.[118] In medicine, this contributes to the spread of ant
ibiotic resistance, as when one bacteria acquires resistance genes it can rapidl
y transfer them to other species.[119] Horizontal transfer of genes from bacteri
a to eukaryotes such as the yeast Saccharomyces cerevisiae and the adzuki bean w
eevil Callosobruchus chinensis has occurred.[120][121] An example of larger-scal
e transfers are the eukaryotic bdelloid rotifers, which have received a range of
genes from bacteria, fungi and plants.[122] Viruses can also carry DNA between
organisms, allowing transfer of genes even across biological domains.[123]
Large-scale gene transfer has also occurred between the ancestors of eukaryotic
cells and bacteria, during the acquisition of chloroplasts and mitochondria. It
is possible that eukaryotes themselves originated from horizontal gene transfers
between bacteria and archaea.[124]
Mechanisms
These charts depict the different types of genetic selection. On each graph, the
x-axis variable is the type of phenotypic trait and the y-axis variable is the
number of organisms. Group A is the original population and Group B is the popul
ation after selection.
Graph 1 shows directional selection, in which a single extreme phenotype is favo
red.
Graph 2 depicts stabilizing selection, where the intermediate phenotype is favor
ed over the extreme traits.
Graph 3 shows disruptive selection, in which the extreme phenotypes are favored
over the intermediate.
Natural selection within a population for a trait that can vary across a range o
f values, such as height, can be categorised into three different types. The fir
st is directional selection, which is a shift in the average value of a trait ov
er timefor example, organisms slowly getting taller.[131] Secondly, disruptive se
lection is selection for extreme trait values and often results in two different
values becoming most common, with selection against the average value. This wou
ld be when either short or tall organisms had an advantage, but not those of med
ium height. Finally, in stabilising selection there is selection against extreme
trait values on both ends, which causes a decrease in variance around the avera
ge value and less diversity.[125][132] This would, for example, cause organisms
to slowly become all the same height.
A special case of natural selection is sexual selection, which is selection for
any trait that increases mating success by increasing the attractiveness of an o
rganism to potential mates.[133] Traits that evolved through sexual selection ar
e particularly prominent among males of several animal species. Although sexuall
y favoured, traits such as cumbersome antlers, mating calls, large body size and
bright colours often attract predation, which compromises the survival of indiv
idual males.[134][135] This survival disadvantage is balanced by higher reproduc
tive success in males that show these hard-to-fake, sexually selected traits.[13
6]
Natural selection most generally makes nature the measure against which individu
als and individual traits, are more or less likely to survive. "Nature" in this
sense refers to an ecosystem, that is, a system in which organisms interact with
every other element, physical as well as biological, in their local environment
. Eugene Odum, a founder of ecology, defined an ecosystem as: "Any unit that inc
ludes all of the organisms...in a given area interacting with the physical envir
onment so that a flow of energy leads to clearly defined trophic structure, biot
ic diversity and material cycles (ie: exchange of materials between living and n
onliving parts) within the system."[137] Each population within an ecosystem occ
upies a distinct niche, or position, with distinct relationships to other parts
of the system. These relationships involve the life history of the organism, its
position in the food chain and its geographic range. This broad understanding o
f nature enables scientists to delineate specific forces which, together, compri
se natural selection.
Natural selection can act at different levels of organisation, such as genes, ce
lls, individual organisms, groups of organisms and species.[138][139][140] Selec
tion can act at multiple levels simultaneously.[141] An example of selection occ
urring below the level of the individual organism are genes called transposons,
which can replicate and spread throughout a genome.[142] Selection at a level ab
ove the individual, such as group selection, may allow the evolution of cooperat
ion, as discussed below.[143]
Biased mutation
In addition to being a major source of variation, mutation may also function as
a mechanism of evolution when there are different probabilities at the molecular
level for different mutations to occur, a process known as mutation bias.[144]
If two genotypes, for example one with the nucleotide G and another with the nuc
leotide A in the same position, have the same fitness, but mutation from G to A
happens more often than mutation from A to G, then genotypes with A will tend to
evolve.[145] Different insertion vs. deletion mutation biases in different taxa
can lead to the evolution of different genome sizes.[146][147] Developmental or
mutational biases have also been observed in morphological evolution.[148][149]
For example, according to the phenotype-first theory of evolution, mutations ca
n eventually cause the genetic assimilation of traits that were previously induc
ed by the environment.[150][151][152]
Mutation bias effects are superimposed on other processes. If selection would fa
vor either one out of two mutations, but there is no extra advantage to having b
oth, then the mutation that occurs the most frequently is the one that is most l
ikely to become fixed in a population.[153][154] Mutations leading to the loss o
f function of a gene are much more common than mutations that produce a new, ful
ly functional gene. Most loss of function mutations are selected against. But wh
en selection is weak, mutation bias towards loss of function can affect evolutio
n.[155] For example, pigments are no longer useful when animals live in the dark
ness of caves, and tend to be lost.[156] This kind of loss of function can occur
because of mutation bias, and/or because the function had a cost, and once the
benefit of the function disappeared, natural selection leads to the loss. Loss o
f sporulation ability in Bacillus subtilis during laboratory evolution appears t
o have been caused by mutation bias, rather than natural selection against the c
ost of maintaining sporulation ability.[157] When there is no selection for loss
of function, the speed at which loss evolves depends more on the mutation rate
than it does on the effective population size,[158] indicating that it is driven
more by mutation bias than by genetic drift. In parasitic organisms, mutation b
ias leads to selection pressures as seen in Ehrlichia. Mutations are biased towa
rds antigenic variants in outer-membrane proteins.
Genetic drift
Further information: Genetic drift and Effective population size
Homologous bones in the limbs of tetrapods. The bones of these animals have the
same basic structure, but have been adapted for specific uses.
Adaptation is the process that makes organisms better suited to their habitat.[1
92][193] Also, the term adaptation may refer to a trait that is important for an
organism's survival. For example, the adaptation of horses' teeth to the grindi
ng of grass. By using the term adaptation for the evolutionary process and adapt
ive trait for the product (the bodily part or function), the two senses of the w
ord may be distinguished. Adaptations are produced by natural selection.[194] Th
e following definitions are due to Theodosius Dobzhansky:
1. Adaptation is the evolutionary process whereby an organism becom
es better able to live in its habitat or habitats.[195]
2. Adaptedness is the state of being adapted: the degree to which a
n organism is able to live and reproduce in a given set of habitats.[196]
3. An adaptive trait is an aspect of the developmental pattern of t
he organism which enables or enhances the probability of that organism surviving
and reproducing.[197]
Adaptation may cause either the gain of a new feature, or the loss of an ancestr
al feature. An example that shows both types of change is bacterial adaptation t
o antibiotic selection, with genetic changes causing antibiotic resistance by bo
th modifying the target of the drug, or increasing the activity of transporters
that pump the drug out of the cell.[198] Other striking examples are the bacteri
a Escherichia coli evolving the ability to use citric acid as a nutrient in a lo
ng-term laboratory experiment,[199] Flavobacterium evolving a novel enzyme that
allows these bacteria to grow on the by-products of nylon manufacturing,[200][20
1] and the soil bacterium Sphingobium evolving an entirely new metabolic pathway
that degrades the synthetic pesticide pentachlorophenol.[202][203] An interesti
ng but still controversial idea is that some adaptations might increase the abil
ity of organisms to generate genetic diversity and adapt by natural selection (i
ncreasing organisms' evolvability).[204][205][206][207][208]
A baleen whale skeleton, a and b label flipper bones, which were adapted from fr
ont leg bones: while c indicates vestigial leg bones, suggesting an adaptation f
rom land to sea.[209]
Adaptation occurs through the gradual modification of existing structures. Conse
quently, structures with similar internal organisation may have different functi
ons in related organisms. This is the result of a single ancestral structure bei
ng adapted to function in different ways. The bones within bat wings, for exampl
e, are very similar to those in mice feet and primate hands, due to the descent
of all these structures from a common mammalian ancestor.[210] However, since al
l living organisms are related to some extent,[211] even organs that appear to h
ave little or no structural similarity, such as arthropod, squid and vertebrate
eyes, or the limbs and wings of arthropods and vertebrates, can depend on a comm
on set of homologous genes that control their assembly and function; this is cal
led deep homology.[212][213]
During evolution, some structures may lose their original function and become ve
stigial structures.[214] Such structures may have little or no function in a cur
rent species, yet have a clear function in ancestral species, or other closely r
elated species. Examples include pseudogenes,[215] the non-functional remains of
eyes in blind cave-dwelling fish,[216] wings in flightless birds,[217] the pres
ence of hip bones in whales and snakes,[209] and sexual traits in organisms that
reproduce via asexual reproduction.[218] Examples of vestigial structures in hu
mans include wisdom teeth,[219] the coccyx,[214] the vermiform appendix,[214] an
d other behavioural vestiges such as goose bumps[220][221] and primitive reflexe
s.[222][223][224]
However, many traits that appear to be simple adaptations are in fact exaptation
s: structures originally adapted for one function, but which coincidentally beca
me somewhat useful for some other function in the process.[225] One example is t
he African lizard Holaspis guentheri, which developed an extremely flat head for
hiding in crevices, as can be seen by looking at its near relatives. However, i
n this species, the head has become so flattened that it assists in gliding from
tree to treean exaptation.[225] Within cells, molecular machines such as the bac
terial flagella[226] and protein sorting machinery[227] evolved by the recruitme
nt of several pre-existing proteins that previously had different functions.[177
] Another example is the recruitment of enzymes from glycolysis and xenobiotic m
etabolism to serve as structural proteins called crystallins within the lenses o
f organisms' eyes.[228][229]
An area of current investigation in evolutionary developmental biology is the de
velopmental basis of adaptations and exaptations.[230] This research addresses t
he origin and evolution of embryonic development and how modifications of develo
pment and developmental processes produce novel features.[231] These studies hav
e shown that evolution can alter development to produce new structures, such as
embryonic bone structures that develop into the jaw in other animals instead for
ming part of the middle ear in mammals.[232] It is also possible for structures
that have been lost in evolution to reappear due to changes in developmental gen
es, such as a mutation in chickens causing embryos to grow teeth similar to thos
e of crocodiles.[233] It is now becoming clear that most alterations in the form
of organisms are due to changes in a small set of conserved genes.[234]
Coevolution
Common garter snake (Thamnophis sirtalis sirtalis) has evolved resistance to the
defensive substance tetrodotoxin in its amphibian prey.
Further information: Coevolution
Interactions between organisms can produce both conflict and cooperation. When t
he interaction is between pairs of species, such as a pathogen and a host, or a
predator and its prey, these species can develop matched sets of adaptations. He
re, the evolution of one species causes adaptations in a second species. These c
hanges in the second species then, in turn, cause new adaptations in the first s
pecies. This cycle of selection and response is called coevolution.[235] An exam
ple is the production of tetrodotoxin in the rough-skinned newt and the evolutio
n of tetrodotoxin resistance in its predator, the common garter snake. In this p
redator-prey pair, an evolutionary arms race has produced high levels of toxin i
n the newt and correspondingly high levels of toxin resistance in the snake.[236
]
Cooperation
Further information: Co-operation (evolution)
Not all co-evolved interactions between species involve conflict.[237] Many case
s of mutually beneficial interactions have evolved. For instance, an extreme coo
peration exists between plants and the mycorrhizal fungi that grow on their root
s and aid the plant in absorbing nutrients from the soil.[238] This is a recipro
cal relationship as the plants provide the fungi with sugars from photosynthesis
. Here, the fungi actually grow inside plant cells, allowing them to exchange nu
trients with their hosts, while sending signals that suppress the plant immune s
ystem.[239]
Coalitions between organisms of the same species have also evolved. An extreme c
ase is the eusociality found in social insects, such as bees, termites and ants,
where sterile insects feed and guard the small number of organisms in a colony
that are able to reproduce. On an even smaller scale, the somatic cells that mak
e up the body of an animal limit their reproduction so they can maintain a stabl
e organism, which then supports a small number of the animal's germ cells to pro
duce offspring. Here, somatic cells respond to specific signals that instruct th
em whether to grow, remain as they are, or die. If cells ignore these signals an
d multiply inappropriately, their uncontrolled growth causes cancer.[240]
Such cooperation within species may have evolved through the process of kin sele
ction, which is where one organism acts to help raise a relative's offspring.[24
1] This activity is selected for because if the helping individual contains alle
les which promote the helping activity, it is likely that its kin will also cont
ain these alleles and thus those alleles will be passed on.[242] Other processes
that may promote cooperation include group selection, where cooperation provide
s benefits to a group of organisms.[243]
Speciation
Main article: Speciation
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Homo erectus
Neanderthal
Homo sapiens
Earlier apes
Possibly bipedal
Earliest bipedal
Earliest exit
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Earliest cooking
Earliest clothes
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Axis scale: millions of years.
Also see: Life timeline and Nature timeline
Life timeline
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Single-celled
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Land life
Dinosaurs
Mammals
Flowers
Earliest water
Earliest life
LHB meteorites
Earliest oxygen
Atmospheric oxygen
Oxygen crisis
Earliest sexual reproduction
Ediacara biota
Cambrian explosion
Earliest humans
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Quaternary
Evolutionary tree showing the divergence of modern species from their common anc
estor in the centre.[294] The three domains are coloured, with bacteria blue, ar
chaea green and eukaryotes red.
Prokaryotes inhabited the Earth from approximately 34 billion years ago.[295][296
] No obvious changes in morphology or cellular organisation occurred in these or
ganisms over the next few billion years.[297] The eukaryotic cells emerged betwe
en 1.62.7 billion years ago. The next major change in cell structure came when ba
cteria were engulfed by eukaryotic cells, in a cooperative association called en
dosymbiosis.[298][299] The engulfed bacteria and the host cell then underwent co
evolution, with the bacteria evolving into either mitochondria or hydrogenosomes
.[300] Another engulfment of cyanobacteriallike organisms led to the formation
of chloroplasts in algae and plants.[301]
The history of life was that of the unicellular eukaryotes, prokaryotes and arch
aea until about 610 million years ago when multicellular organisms began to appe
ar in the oceans in the Ediacaran period.[295][302] The evolution of multicellul
arity occurred in multiple independent events, in organisms as diverse as sponge
s, brown algae, cyanobacteria, slime moulds and myxobacteria.[303] In January 20
16, scientists reported that, about 800 million years ago, a minor genetic chang
e in a single molecule called GKPID may have allowed organisms to go from a sin
gle cell organism to one of many cells.[304]
Soon after the emergence of these first multicellular organisms, a remarkable am
ount of biological diversity appeared over approximately 10 million years, in an
event called the Cambrian explosion. Here, the majority of types of modern anim
als appeared in the fossil record, as well as unique lineages that subsequently
became extinct.[305] Various triggers for the Cambrian explosion have been propo
sed, including the accumulation of oxygen in the atmosphere from photosynthesis.
[306]
About 500 million years ago, plants and fungi colonised the land and were soon f
ollowed by arthropods and other animals.[307] Insects were particularly successf
ul and even today make up the majority of animal species.[308] Amphibians first
appeared around 364 million years ago, followed by early amniotes and birds arou
nd 155 million years ago (both from "reptile"like lineages), mammals around 129
million years ago, homininae around 10 million years ago and modern humans arou
nd 250,000 years ago.[309][310][311] However, despite the evolution of these lar
ge animals, smaller organisms similar to the types that evolved early in this pr
ocess continue to be highly successful and dominate the Earth, with the majority
of both biomass and species being prokaryotes.[188]
Applications
Main articles: Applications of evolution, Selective breeding, and Evolutionary c
omputation
Concepts and models used in evolutionary biology, such as natural selection, hav
e many applications.[312]
Artificial selection is the intentional selection of traits in a population of o
rganisms. This has been used for thousands of years in the domestication of plan
ts and animals.[313] More recently, such selection has become a vital part of ge
netic engineering, with selectable markers such as antibiotic resistance genes b
eing used to manipulate DNA. Proteins with valuable properties have evolved by r
epeated rounds of mutation and selection (for example modified enzymes and new a
ntibodies) in a process called directed evolution.[314]
Understanding the changes that have occurred during an organism's evolution can
reveal the genes needed to construct parts of the body, genes which may be invol
ved in human genetic disorders.[315] For example, the Mexican tetra is an albino
cavefish that lost its eyesight during evolution. Breeding together different p
opulations of this blind fish produced some offspring with functional eyes, sinc
e different mutations had occurred in the isolated populations that had evolved
in different caves.[316] This helped identify genes required for vision and pigm
entation.[317]
Many human diseases are not static phenomena, but capable of evolution. Viruses,
bacteria, fungi and cancers evolve to be resistant to host immune defences, as
well as pharmaceutical drugs.[318][319][320] These same problems occur in agricu
lture with pesticide[321] and herbicide[322] resistance. It is possible that we
are facing the end of the effective life of most of available antibiotics[323] a
nd predicting the evolution and evolvability[324] of our pathogens and devising
strategies to slow or circumvent it is requiring deeper knowledge of the complex
forces driving evolution at the molecular level.[325]
In computer science, simulations of evolution using evolutionary algorithms and
artificial life started in the 1960s and were extended with simulation of artifi
cial selection.[326] Artificial evolution became a widely recognised optimisatio
n method as a result of the work of Ingo Rechenberg in the 1960s. He used evolut
ion strategies to solve complex engineering problems.[327] Genetic algorithms in
particular became popular through the writing of John Henry Holland.[328] Pract
ical applications also include automatic evolution of computer programmes.[329]
Evolutionary algorithms are now used to solve multidimensional problems more ef
ficiently than software produced by human designers and also to optimise the des
ign of systems.[330]
Social and cultural responses
Further information: Social effects of evolutionary theory, 1860 Oxford evolutio
n debate, Creationevolution controversy, and Objections to evolution
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Evolution on In Our Time at the BBC. (listen now)
"Evolution". New Scientist. Retrieved 2011-05-30.
"Evolution Resources from the National Academies". Washington, D.C.: Nat
ional Academy of Sciences. Retrieved 2011-05-30.
"Understanding Evolution: your one-stop resource for information on evol
ution". University of California, Berkeley. Retrieved 2011-05-30.
"Evolution of Evolution 150 Years of Darwin's 'On the Origin of Species'
". Arlington County, VA: National Science Foundation. Retrieved 2011-05-30.
Human Timeline (Interactive) Smithsonian, National Museum of Natural His
tory (August 2016).
Experiments concerning the process of biological evolution
Lenski, Richard E. "Experimental Evolution". Michigan State University.
Retrieved 2013-07-31.
Chastain, Erick; Livnat, Adi; Papadimitriou, Christos; Vazirani, Umesh (
July 22, 2014). "Algorithms, games, and evolution". Proc. Natl. Acad. Sci. U.S.A
. Washington, D.C.: National Academy of Sciences. 111 (29): 1062010623. Bibcode:2
014PNAS..11110620C. doi:10.1073/pnas.1406556111. ISSN0027-8424. Retrieved 2015-01
-03.
Online lectures
Carroll, Sean B. "The Making of the Fittest". Retrieved 2011-05-30.
Stearns, Stephen C. "Principles of Evolution, Ecology and Behavior". Ret
rieved 2011-08-30.
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Evolutionary biology
Evolutionary history of life
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