Flowering plant

The owering plants (angiosperms), also known as

Angiospermae[4][5] or Magnoliophyta,[6] are the most
diverse group of land plants, with 416 families, approx. 13,164 known genera and a total of c. 295,383
known species.[7] Like gymnosperms, angiosperms are
seed-producing plants; they are distinguished from gymnosperms by characteristics including owers, endosperm
within the seeds, and the production of fruits that contain the seeds. Etymologically, angiosperm means a plant
that produces seeds within an enclosure, in other words,
a fruiting plant. The term angiosperm comes from the
Greek composite word (angeion, case or casing, and
sperma, seed) meaning enclosed seeds, after the enclosed condition of the seeds.
The ancestors of owering plants diverged from
gymnosperms in the Triassic Period, during the range
245 to 202 million years ago (mya), and the rst owering plants are known from 160 mya. They diversied
extensively during the Lower Cretaceous, became
widespread by 120 mya, and replaced conifers as the
dominant trees from 100 to 60 mya.

Angiosperm derived characteristics

Angiosperms dier from other seed plants in several Bud of a pink rose
ways, described in the table. These distinguishing characteristics taken together have made the angiosperms the
most diverse and numerous land plants and the most com- avascular sexual stage, the gametophyte, within the
tissues of the vascular sporophyte. This occurred by
mercially important group to humans.[lower-alpha 1]
spore germination within sporangia rather than spore
release, as in non-seed plants. A current example of how
this might have happened can be seen in the precocious
2 Evolution
spore germination in Selaginella, the spike-moss. The
result for the ancestors of angiosperms was enclosing
Further information: Evolutionary history of plants them in a case, the seed. The rst seed bearing plants,
Flowers
like the ginkgo, and conifers (such as pines and rs), did
not produce owers. The pollen grains (males) of Ginkgo
Fossilized spores suggest that higher plants and cycads produce a pair of agellated, mobile sperm
(embryophytes) have lived on land for at least 475 cells that swim down the developing pollen tube to the
million years.[9] Early land plants reproduced sexually female and her eggs.
with agellated, swimming sperm, like the green algae
from which they evolved. An adaptation to terrestrialization was the development of upright meiosporangia
for dispersal by spores to new habitats. This feature is
lacking in the descendants of their nearest algal relatives, the Charophycean green algae. A later terrestrial
adaptation took place with retention of the delicate,

The apparently sudden appearance of nearly modern

owers in the fossil record initially posed such a problem for the theory of evolution that Charles Darwin
called it an "abominable mystery".[10] However, the fossil record has considerably grown since the time of Darwin, and recently discovered angiosperm fossils such as
Archaefructus, along with further discoveries of fossil
1

2 EVOLUTION
how owers evolved. Some older fossils, such as the upper Triassic Sanmiguelia, have been suggested. Based on
current evidence, some propose that the ancestors of the
angiosperms diverged from an unknown group of gymnosperms in the Triassic period (245202 million years
ago). Fossil angiosperm-like pollen from the Middle Triassic (247.2242.0 Ma) suggests an older date for their
origin.[11] A close relationship between angiosperms and
gnetophytes, proposed on the basis of morphological evidence, has more recently been disputed on the basis of
molecular evidence that suggest gnetophytes are instead
more closely related to other gymnosperms.
The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of whole
genome duplication events.[12] These occurred at 319
million years ago and 192 million years ago. Another
possible whole genome duplication event at 160 million
years ago perhaps created the ancestral line that led to
all modern owering plants.[13] That event was studied
by sequencing the genome of an ancient owering plant,
Amborella trichopoda,[14] and directly addresses Darwins
"abominable mystery.

Flowers of Malus sylvestris (crab apple)

The earliest known macrofossil condently identied as

an angiosperm, Archaefructus liaoningensis, is dated to
about 125 million years BP (the Cretaceous period),[15]
whereas pollen considered to be of angiosperm origin
takes the fossil record back to about 130 million years BP.
However, one study has suggested that the early-middle
Jurassic plant Schmeissneria, traditionally considered a
type of ginkgo, may be the earliest known angiosperm,
or at least a close relative.[16] In addition, circumstantial
chemical evidence has been found for the existence of angiosperms as early as 250 million years ago. Oleanane, a
secondary metabolite produced by many owering plants,
has been found in Permian deposits of that age together
with fossils of gigantopterids.[17][18] Gigantopterids are
a group of extinct seed plants that share many morphological traits with owering plants, although they are not
known to have been owering plants themselves.
In 2013 owers encased in amber were found and dated
100 million years before present. The amber had frozen
the act of sexual reproduction in the process of taking
place. Microscopic images showed tubes growing out of
pollen and penetrating the owers stigma. The pollen was
sticky, suggesting it was carried by insects.[19]

Flowers and leaves of Oxalis pes-caprae (Bermuda buttercup)

gymnosperms, suggest how angiosperm characteristics

may have been acquired in a series of steps. Several
groups of extinct gymnosperms, in particular seed ferns,
have been proposed as the ancestors of owering plants,
but there is no continuous fossil evidence showing exactly

Recent DNA analysis based on molecular systematics[20][21] showed that Amborella trichopoda, found on
the Pacic island of New Caledonia, belongs to a sister
group of the other owering plants, and morphological
studies[22] suggest that it has features that may have been
characteristic of the earliest owering plants.
The orders Amborellales,
Nymphaeales,
and
Austrobaileyales diverged as separate lineages from
the remaining angiosperm clade at a very early stage in
owering plant evolution.[23]
The great angiosperm radiation, when a great diversity of

3
angiosperms appears in the fossil record, occurred in the
mid-Cretaceous (approximately 100 million years ago).
However, a study in 2007 estimated that the division of
the ve most recent (the genus Ceratophyllum, the family Chloranthaceae, the eudicots, the magnoliids, and the
monocots) of the eight main groups occurred around 140
million years ago.[24] By the late Cretaceous, angiosperms
appear to have dominated environments formerly occupied by ferns and cycadophytes, but large canopy-forming
trees replaced conifers as the dominant trees only close
to the end of the Cretaceous 66 million years ago or even
later, at the beginning of the Tertiary.[25] The radiation
of herbaceous angiosperms occurred much later.[26] Yet,
many fossil plants recognizable as belonging to modern
families (including beech, oak, maple, and magnolia) had
already appeared by the late Cretaceous.

cialized relationship with some specic animal (a wasp,

for example). Such a relationship, with a hypothetical
wasp carrying pollen from one plant to another much the
way g wasps do today, could result in the development
of a high degree of specialization in both the plant(s) and
their partners. Note that the wasp example is not incidental; bees, which, it is postulated, evolved specically
due to mutualistic plant relationships, are descended from
wasps.[27]
Animals are also involved in the distribution of seeds.
Fruit, which is formed by the enlargement of ower parts,
is frequently a seed-dispersal tool that attracts animals
to eat or otherwise disturb it, incidentally scattering the
seeds it contains (see frugivory). Although many such
mutualistic relationships remain too fragile to survive
competition and to spread widely, owering proved to be
an unusually eective means of reproduction, spreading
(whatever its origin) to become the dominant form of land
plant life.
Flower ontogeny uses a combination of genes normally
responsible for forming new shoots.[28] The most primitive owers probably had a variable number of ower
parts, often separate from (but in contact with) each
other. The owers tended to grow in a spiral pattern, to
be bisexual (in plants, this means both male and female
parts on the same ower), and to be dominated by the
ovary (female part). As owers evolved, some variations
developed parts fused together, with a much more specic number and design, and with either specic sexes
per ower or plant or at least ovary-inferior.

Flower evolution continues to the present day; modern

owers have been so profoundly inuenced by humans
that some of them cannot be pollinated in nature. Many
modern domesticated ower species were formerly simple weeds, which sprouted only when the ground was disturbed. Some of them tended to grow with human crops,
perhaps already having symbiotic companion plant relationships with them, and the prettiest did not get plucked
because of their beauty, developing a dependence upon
Two bees on a ower head of Creeping Thistle, Cirsium arvense and special adaptation to human aection.[29]
It is generally assumed that the function of owers,
from the start, was to involve mobile animals in their
reproduction processes. That is, pollen can be scattered
even if the ower is not brightly colored or oddly shaped
in a way that attracts animals; however, by expending the
energy required to create such traits, angiosperms can enlist the aid of animals and, thus, reproduce more eciently.

A few paleontologists have also proposed that owering

plants, or angiosperms, might have evolved due to interactions with dinosaurs. One of the ideas strongest
proponents is Robert T. Bakker. He proposes that
herbivorous dinosaurs, with their eating habits, provided
a selective pressure on plants, for which adaptations either succeeded in deterring or coping with predation by
herbivores.[30]

Island genetics provides one proposed explanation for the

sudden, fully developed appearance of owering plants. 3 Classication
Island genetics is believed to be a common source of
speciation in general, especially when it comes to radical adaptations that seem to have required inferior tran- There are eight groups of living angiosperms:
sitional forms. Flowering plants may have evolved in an
Amborella, a single species of shrub from New Caleisolated setting like an island or island chain, where the
plants bearing them were able to develop a highly spedonia;

CLASSIFICATION

Nymphaeales, about 80 species,[33] water lilies and

Hydatellaceae;
Austrobaileyales, about 100 species[33] of woody
plants from various parts of the world;
Chloranthales, several dozen species of aromatic
plants with toothed leaves;
Magnoliids, about 9,000 species,[33] characterized by trimerous owers, pollen with one pore,
and usually branching-veined leavesfor example
magnolias, bay laurel, and black pepper;
Monocots, about 70,000 species,[33] characterized
by trimerous owers, a single cotyledon, pollen with
one pore, and usually parallel-veined leavesfor example grasses, orchids, and palms;
Ceratophyllum, about 6 species[33] of aquatic plants,
perhaps most familiar as aquarium plants;
Eudicots, about 175,000 species,[33] characterized
by 4- or 5-merous owers, pollen with three pores,
and usually branching-veined leavesfor example
sunowers, petunia, buttercup, apples, and oaks.
The exact relationship between these eight groups is not
yet clear, although there is agreement that the rst three
groups to diverge from the ancestral angiosperm were
Amborellales, Nymphaeales, and Austrobaileyales.[34]
The term basal angiosperms refers to these three groups.
Among the rest, the relationship between the three broadest of these groups (magnoliids, monocots, and eudicots)
remains unclear. Some analyses make the magnoliids the
rst to diverge, others the monocots.[32] Ceratophyllum
seems to group with the eudicots rather than with the
monocots.

From 1736, an illustration of Linnaean classication.

possible only after 1827, when Robert Brown established the existence of truly naked ovules in the Cycadeae
and Coniferae,[35] and applied to them the name Gymnosperms. From that time onward, as long as these Gymnosperms were, as was usual, reckoned as dicotyledonous
owering plants, the term Angiosperm was used antithetically by botanical writers, with varying scope, as a groupname for other dicotyledonous plants.

In 1851, Hofmeister discovered the changes occurring in

the embryo-sac of owering plants, and determined the
correct relationships of these to the Cryptogamia. This
Based on the 4th version of the Angiosperm Phylogeny xed the position of Gymnosperms as a class distinct
Group classication.[1]
from Dicotyledons, and the term Angiosperm then gradually came to be accepted as the suitable designation for the
whole of the owering plants other than Gymnosperms,
3.2 History of classication
including the classes of Dicotyledons and MonocotyleThe botanical term Angiosperm, from the Ancient dons. This is the sense in which the term is used today.
Greek , angeon (bottle, vessel) and , In most taxonomies, the owering plants are treated as
(seed), was coined in the form Angiospermae by Paul a coherent group. The most popular descriptive name
Hermann in 1690, as the name of one of his primary has been Angiospermae (Angiosperms), with Anthophyta
divisions of the plant kingdom. This included ower- (owering plants) a second choice. These names are
ing plants possessing seeds enclosed in capsules, dis- not linked to any rank. The Wettstein system and the
tinguished from his Gymnospermae, or owering plants Engler system use the name Angiospermae, at the aswith achenial or schizo-carpic fruits, the whole fruit or signed rank of subdivision. The Reveal system treated
each of its pieces being here regarded as a seed and naked. owering plants as subdivision Magnoliophytina (Frohne
The term and its antonym were maintained by Carl Lin- & U. Jensen ex Reveal, Phytologia 79: 70 1996), but later
naeus with the same sense, but with restricted applica- split it to Magnoliopsida, Liliopsida, and Rosopsida. The
tion, in the names of the orders of his class Didynamia. Takhtajan system and Cronquist system treat this group at
Its use with any approach to its modern scope became the rank of division, leading to the name Magnoliophyta

3.1

APG IV classication

3.3

Flowering plant diversity

Auxanometer: Device for measuring increase or rate of growth

in plants

(from the family name Magnoliaceae). The Dahlgren system and Thorne system (1992) treat this group at the rank
of class, leading to the name Magnoliopsida. The APG
system of 1998, and the later 2003[36] and 2009[31] revisions, treat the owering plants as a clade called angiosperms without a formal botanical name. However,
a formal classication was published alongside the 2009
revision in which the owering plants form the Subclass
Magnoliidae.[37]

Monocot (left) and dicot seedlings

(plural monocotyledons), or abbreviated, as dicot (plural

dicots) and monocot (plural monocots). These names derive from the observation that the dicots most often have
two cotyledons, or embryonic leaves, within each seed.
The monocots usually have only one, but the rule is not
absolute either way. From a broad diagnostic point of
view, the number of cotyledons is neither a particularly
The internal classication of this group has undergone
handy nor a reliable character.
considerable revision. The Cronquist system, proposed
by Arthur Cronquist in 1968 and published in its full form Recent studies, as by the APG, show that the monocots
in 1981, is still widely used but is no longer believed to form a monophyletic group (clade) but that the dicots
accurately reect phylogeny. A consensus about how the do not (they are paraphyletic). Nevertheless, the maowering plants should be arranged has recently begun to jority of dicot species do form a monophyletic group,
emerge through the work of the Angiosperm Phylogeny called the eudicots or tricolpates. Of the remaining dicot
Group (APG), which published an inuential reclassi- species, most belong to a third major clade known as the
cation of the angiosperms in 1998. Updates incorporat- magnoliids, containing about 9,000 species. The rest ining more recent research were published as APG II in clude a paraphyletic grouping of primitive species known
collectively as the basal angiosperms, plus the families
2003[36] and as APG III in 2009.[31][38]
Ceratophyllaceae and Chloranthaceae.
Traditionally, the owering plants are divided into two
groups, which in the Cronquist system are called Magnoliopsida (at the rank of class, formed from the family
name Magnoliaceae) and Liliopsida (at the rank of class, 3.3 Flowering plant diversity
formed from the family name Liliaceae). Other descriptive names allowed by Article 16 of the ICBN include The number of species of owering plants is estimated
Dicotyledones or Dicotyledoneae, and Monocotyledones to be in the range of 250,000 to 400,000.[39][40][41] This
or Monocotyledoneae, which have a long history of use. compares to around 12,000 species of moss[42] or 11,000
In English a member of either group may be called species of pteridophytes,[43] showing that the owering
a dicotyledon (plural dicotyledons) and monocotyledon plants are much more diverse. The number of families in

CLASSIFICATION

Nearly all species belong to the eudicot (75%), monocot (23%), and magnoliid (2%) clades. The remaining 5
clades contain a little over 250 species in total; i.e. less
than 0.1% of owering plant diversity, divided among 9
families. The 42 most-diverse of 443 families of owering plants by species,[45] in their APG circumscriptions,
are
1. Asteraceae or Compositae (daisy family): 22,750
species;
2. Orchidaceae (orchid family): 21,950;
3. Fabaceae or Leguminosae (bean family): 19,400;
4. Rubiaceae (madder family): 13,150;[46]
5. Poaceae or Gramineae (grass family): 10,035;
6. Lamiaceae or Labiatae (mint family): 7,175;
7. Euphorbiaceae (spurge family): 5,735;
8. Melastomataceae or Melastomaceae (melastome
family): 5,005;
A poster of twelve dierent species of owers of the Asteraceae
family

9. Myrtaceae (myrtle family): 4,625;

10. Apocynaceae (dogbane family): 4,555;
11. Cyperaceae (sedge family): 4,350;
12. Malvaceae (mallow family): 4,225;
13. Araceae (arum family): 4,025;
14. Ericaceae (heath family): 3,995;
15. Gesneriaceae (gesneriad family): 3,870;
16. Apiaceae or Umbelliferae (parsley family): 3,780;
17. Brassicaceae or Cruciferae (cabbage family): 3,710:
18. Piperaceae (pepper family): 3,600;
19. Acanthaceae (acanthus family): 3,500;
20. Rosaceae (rose family): 2,830;
21. Boraginaceae (borage family): 2,740;
22. Urticaceae (nettle family): 2,625;
23. Ranunculaceae (buttercup family): 2,525;

Lupinus pilosus

24. Lauraceae (laurel family): 2,500;

25. Solanaceae (nightshade family): 2,460;
[36]

APG (1998) was 462. In APG II (2003) it is not settled; at maximum it is 457, but within this number there
are 55 optional segregates, so that the minimum number
of families in this system is 402. In APG III (2009) there
are 415 families.[31][44]

26. Campanulaceae (bellower family): 2,380;

The diversity of owering plants is not evenly distributed.

29. Caryophyllaceae (pink family): 2,200;

27. Arecaceae (palm family): 2,361;

28. Annonaceae (custard apple family): 2,220;

7
30. Orobanchaceae (broomrape family): 2,060;
31. Amaranthaceae (amaranth family): 2,050;
32. Iridaceae (iris family): 2,025;
33. Aizoaceae or Ficoidaceae (ice plant family): 2,020;
34. Rutaceae (rue family): 1,815;
35. Phyllanthaceae (phyllanthus family): 1,745;
36. Scrophulariaceae (gwort family): 1,700;
37. Gentianaceae (gentian family): 1,650;
38. Convolvulaceae (bindweed family): 1,600;
39. Proteaceae (protea family): 1,600;
40. Sapindaceae (soapberry family): 1,580;

bundles of the stem are arranged such that the xylem and
phloem form concentric rings.
In the dicotyledons, the bundles in the very young stem
are arranged in an open ring, separating a central pith
from an outer cortex. In each bundle, separating the
xylem and phloem, is a layer of meristem or active formative tissue known as cambium. By the formation of
a layer of cambium between the bundles (interfascicular
cambium), a complete ring is formed, and a regular periodical increase in thickness results from the development
of xylem on the inside and phloem on the outside. The
soft phloem becomes crushed, but the hard wood persists and forms the bulk of the stem and branches of the
woody perennial. Owing to dierences in the character
of the elements produced at the beginning and end of the
season, the wood is marked out in transverse section into
concentric rings, one for each season of growth, called
annual rings.

Among the monocotyledons, the bundles are more numerous in the young stem and are scattered through
the ground tissue. They contain no cambium and once
42. Araliaceae (Aralia or ivy family): 1,450.
formed the stem increases in diameter only in exceptional
Of these, the Orchidaceae, Poaceae, Cyperaceae, Are- cases.
caceae, and Iridaceae are monocot families; Piperaceae,
Lauraceae, and Annonaceae are magnoliid dicots; the rest
of the families are eudicots.
5 The ower, fruit, and seed
41. Cactaceae (cactus family): 1,500;

Vascular anatomy

76

Cross-section of a stem of the angiosperm ax:

1. Pith,
2. Protoxylem,
3. Xylem I,
4. Phloem I,
5. Sclerenchyma (bast bre),
6. Cortex,
7. Epidermis

5.1 Flowers
Main articles: Flower and Plant reproductive morphology
The characteristic feature of angiosperms is the ower.
Flowers show remarkable variation in form and elaboration, and provide the most trustworthy external characteristics for establishing relationships among angiosperm
species. The function of the ower is to ensure fertilization of the ovule and development of fruit containing
seeds. The oral apparatus may arise terminally on a
shoot or from the axil of a leaf (where the petiole attaches
to the stem). Occasionally, as in violets, a ower arises
singly in the axil of an ordinary foliage-leaf. More typically, the ower-bearing portion of the plant is sharply
distinguished from the foliage-bearing or vegetative portion, and forms a more or less elaborate branch-system
called an inorescence.
There are two kinds of reproductive cells produced by
owers. Microspores, which will divide to become
pollen grains, are the male cells and are borne in the
stamens (or microsporophylls). The female cells called
megaspores, which will divide to become the egg cell
(megagametogenesis), are contained in the ovule and enclosed in the carpel (or megasporophyll).

The ower may consist only of these parts, as in willow,

where each ower comprises only a few stamens or two
carpels. Usually, other structures are present and serve
The amount and complexity of tissue-formation in ow- to protect the sporophylls and to form an envelope atering plants exceeds that of gymnosperms. The vascular tractive to pollinators. The individual members of these

5 THE FLOWER, FRUIT, AND SEED

is eected by color, scent, and nectar, which may be secreted in some part of the ower. The characteristics that
attract pollinators account for the popularity of owers
and owering plants among humans.
While the majority of owers are perfect or
hermaphrodite (having both pollen and ovule producing parts in the same ower structure), owering
plants have developed numerous morphological and
physiological mechanisms to reduce or prevent selffertilization. Heteromorphic owers have short carpels
and long stamens, or vice versa, so animal pollinators
cannot easily transfer pollen to the pistil (receptive part
of the carpel). Homomorphic owers may employ a
biochemical (physiological) mechanism called selfincompatibility to discriminate between self and non-self
pollen grains. In other species, the male and female parts
are morphologically separated, developing on dierent
owers.

5.2 Fertilization and embryogenesis

Main articles: Fertilization and Plant embryogenesis
Double fertilization refers to a process in which two

A collection of owers forming an inorescence

surrounding structures are known as sepals and petals (or

tepals in owers such as Magnolia where sepals and petals
are not distinguishable from each other). The outer series
(calyx of sepals) is usually green and leaf-like, and functions to protect the rest of the ower, especially the bud.
The inner series (corolla of petals) is, in general, white
or brightly colored, and is more delicate in structure. It
functions to attract insect or bird pollinators. Attraction

Angiosperm life cycle

sperm cells fertilize cells in the ovary. This process begins when a pollen grain adheres to the stigma of the pistil
(female reproductive structure), germinates, and grows a
long pollen tube. While this pollen tube is growing, a haploid generative cell travels down the tube behind the tube
nucleus. The generative cell divides by mitosis to produce
two haploid (n) sperm cells. As the pollen tube grows, it
makes its way from the stigma, down the style and into the
ovary. Here the pollen tube reaches the micropyle of the
ovule and digests its way into one of the synergids, releas-

9
ing its contents (which include the sperm cells). The synergid that the cells were released into degenerates and one
sperm makes its way to fertilize the egg cell, producing a
diploid (2n) zygote. The second sperm cell fuses with
both central cell nuclei, producing a triploid (3n) cell. As
the zygote develops into an embryo, the triploid cell develops into the endosperm, which serves as the embryos
food supply. The ovary will now develop into a fruit and
the ovule will develop into a seed.

5.3

Fruit and seed

(a structure within the ovary that is located within the pistil at the center of the ower) (see diagram labeled Angiosperm lifecycle). A diploid cell (megaspore mother
cell) in the ovule undergoes meiosis (involving two successive cell divisions) to produce four cells (megaspores
or female gametes) with haploid nuclei.[47] One of these
four cells (megaspore) then undergoes three successive
mitotic divisions to produce an immature embryo sac
(megagametocyte) with eight haploid nuclei. Next, these
nuclei are segregated into separate cells by cytokinesis to
producing 3 antipodal cells, 2 synergid cells and an egg
cell. Two polar nuclei are left in the central cell of the
embryo sac.

Main articles: Seed and Fruit

Pollen is also produced by meiosis in the male anAs the development of embryo and endosperm proceeds
ther (microsporangium). During meiosis, a diploid microspore mother cell undergoes two successive meiotic
divisions to produce 4 haploid cells (microspores or male
gametes). Each of these microspores, after further mitoses, becomes a pollen grain (microgametophyte) containing two haploid generative (sperm) cells and a tube
nucleus. When a pollen grain makes contact with the
female stigma, the pollen grain forms a pollen tube that
grows down the style into the ovary. In the act of fertilization, a male sperm nucleus fuses with the female egg
nucleus to form a diploid zygote that can then develop
into an embryo within the newly forming seed. Upon
germination of the seed, a new plant can grow and mature.
The fruit of the Aesculus or Horse Chestnut tree

within the embryo sac, the sac wall enlarges and combines with the nucellus (which is likewise enlarging) and
the integument to form the seed coat. The ovary wall develops to form the fruit or pericarp, whose form is closely
associated with the manner of distribution of the seed.
Frequently, the inuence of fertilization is felt beyond the
ovary, and other parts of the ower take part in the formation of the fruit, e.g., the oral receptacle in the apple,
strawberry, and others.

The adaptive function of meiosis is currently a matter of

debate. A key event during meiosis in a diploid cell is the
pairing of homologous chromosomes and homologous recombination (the exchange of genetic information) between homologous chromosomes. This process promotes
the production of increased genetic diversity among
progeny and the recombinational repair of damages in the
DNA to be passed on to progeny. To explain the adaptive function of meiosis in owering plants, some authors
emphasize diversity [48] and others emphasize DNA repair [49]

The character of the seed coat bears a denite relation

to that of the fruit. They protect the embryo and aid in
dissemination; they may also directly promote germination. Among plants with indehiscent fruits, in general,
the fruit provides protection for the embryo and secures
dissemination. In this case, the seed coat is only slightly
developed. If the fruit is dehiscent and the seed is exposed, in general, the seed-coat is well developed, and
must discharge the functions otherwise executed by the
fruit.

7 Apomixis

8 Economic importance

Meiosis

Apomixis (reproduction via asexually formed seeds) is

found naturally in about 2.2% of angiosperm genera [50]
One type of apomixis, gametophytic apomixis found in a
dandelion species [51] involves formation of an unreduced
embryo sac due to incomplete meiosis (apomeiosis) and
development of an embryo from the unreduced egg inside
the embryo sac, without fertilization (parthenogenesis).

Flowering plants generate gametes using a specialized cell Agriculture is almost entirely dependent on angiosperms,
division called meiosis. Meiosis takes place in the ovule which provide virtually all plant-based food, and also pro-

10

11

vide a signicant amount of livestock feed. Of all the

families of plants, the Poaceae, or grass family (grains),
is by far the most important, providing the bulk of all
feedstocks (rice, corn maize, wheat, barley, rye, oats,
pearl millet, sugar cane, sorghum). The Fabaceae, or
legume family, comes in second place. Also of high
importance are the Solanaceae, or nightshade family
(potatoes, tomatoes, and peppers, among others), the
Cucurbitaceae, or gourd family (also including pumpkins
and melons), the Brassicaceae, or mustard plant family
(including rapeseed and the innumerable varieties of the
cabbage species Brassica oleracea), and the Apiaceae,
or parsley family. Many of our fruits come from the
Rutaceae, or rue family (including oranges, lemons,
grapefruits, etc.), and the Rosaceae, or rose family (including apples, pears, cherries, apricots, plums, etc.).
In some parts of the world, certain single species assume
paramount importance because of their variety of uses,
for example the coconut (Cocos nucifera) on Pacic atolls,
and the olive (Olea europaea) in the Mediterranean region.

REFERENCES

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See also
List of garden plants
List of plant orders
List of plants by common name
List of systems of plant taxonomy

10

Notes

[1] The major exception to the dominance of terrestrial

ecosystems by owering plants is the coniferous forest.

11

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EXTERNAL LINKS

Takhtajan, A. (June 1964). The Taxa of the

Higher Plants above the Rank of Order. Taxon.
13 (5): 160164. doi:10.2307/1216134. JSTOR
10.2307/1216134.
Simpson, M.G. Plant Systematics, 2nd Edition. Elsevier/Academic Press. 2010.
Raven, P.H., R.F. Evert, S.E. Eichhorn. Biology of
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Sattler, R. 1973. Organogenesis of Flowers. A Photographic Text-Atlas. University of Toronto Press.
Zeng, Liping; Zhang, Qiang; Sun, Renran; Kong,
Hongzhi; Zhang, Ning; Ma, Hong (24 September 2014). Resolution of deep angiosperm phylogeny using conserved nuclear genes and estimates
of early divergence times. Nature Communications.
5 (4956). doi:10.1038/ncomms5956.
Cole, Theodor C.H.; Hilger, Dr. Harmut H.
Angiosperm Phylogeny Poster Flowering Plant
Systematics
Cromie, William J. (December 16, 1999). Oldest
Known Flowering Plants Identied By Genes. Harvard University Gazette.
Watson, L. and Dallwitz, M.J. (1992 onwards). The
families of owering plants: descriptions, illustrations, identication, information retrieval.
Flowering plant at the Encyclopedia of Life

13 External links

Cronquist, Arthur (October 1960). The divisions This article incorporates text from a publication now
and classes of plants. The Botanical Review. 26 in the public domain: Chisholm, Hugh, ed. (1911).
(4): 425482. doi:10.1007/BF02940572.
"article name needed ". Encyclopdia Britannica (11th ed.).
Cronquist, Arthur (1981). An Integrated System Cambridge University Press.
of Classication of Flowering Plants. New York:
Columbia Univ. Press. ISBN 0-231-03880-1.
Dahlgren, R. M. T. (February 1980). A revised
system of classication of the angiosperms. Botanical Journal of the Linnean Society. 80 (2): 91124.
doi:10.1111/j.1095-8339.1980.tb01661.x.
Heywood, V. H., Brummitt, R. K., Culham, A. &
Seberg, O. (2007). Flowering Plant Families of the
World. Richmond Hill, Ontario, Canada: Firey
Books. ISBN 1-55407-206-9.
Dilcher, D. (2000). Toward a new synthesis: Major
evolutionary trends in the angiosperm fossil record.
Proceedings of the National Academy of Sciences. 97
(13): 7030. doi:10.1073/pnas.97.13.7030.
Pooja (2004). Angiosperms. New Delhi: Discovery.
ISBN 9788171417889. Retrieved 7 January 2016.

13

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Text and image sources, contributors, and licenses

Text

Flowering plant Source: https://en.wikipedia.org/wiki/Flowering_plant?oldid=759838610 Contributors: AxelBoldt, Mav, Bryan Derksen,

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