Flower

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From Wikipedia, the free encyclopedia

For other uses, see Flower (disambiguation).
"Floral" redirects here. For other uses, see Floral (disambiguation).

Flowers or clusters of flowers produced by twelve species of Angiosperms from different families.

Selection of differently constructed flowers at different stages of vascular plant development

A flower, sometimes known as a bloom or blossom, is the reproductive structure

found in flowering plants (plants of the division Angiospermae). Flowers produce
gametophytes, which in flowering plants consist of a few haploid cells which
produce gametes. The "male" gametophyte, which produces non-motile sperm, is
enclosed within pollen grains; the "female" gametophyte is contained within the ovule.
When pollen from the anther of a flower is deposited on the stigma, this is
called pollination. Some flowers may self-pollinate, producing seed using pollen from
the same flower or a different flower of the same plant, but others have mechanisms to
prevent self-pollination and rely on cross-pollination, when pollen is transferred from the
anther of one flower to the stigma of another flower on a different individual of the same
species.
Self-pollination happens in flowers where the stamen and carpel mature at the same
time, and are positioned so that the pollen can land on the flower's stigma. This
pollination does not require an investment from the plant to provide nectar and pollen as
food for pollinators.[1]
Some flowers produce diaspores without fertilization (parthenocarpy). Flowers
contain sporangia and are the site where gametophytes develop.
Most flowering plants depend on animals, such as bees, moths, and butterflies, to
transfer their pollen between different flowers, and have evolved to attract
these pollinators by various strategies, including brightly colored, conspicuous petals,
attractive scents, and the production of nectar, a food source for pollinators.[2] In this
way, many flowering plants have co-evolved with pollinators be mutually dependent on
services they provide to one another—in the plant's case, a means of reproduction; in
the pollinator's case, a source of food.[3] After fertilization, the ovary of the flower
develops into fruit containing seeds.
Flowers have long been appreciated by humans for their beauty and pleasant scents,
and also hold cultural significance as religious, ritual, or symbolic objects, or sources
of medicine and food.

Etymology
Flower is from the Middle English flour, which referred to both the ground grain and the
reproductive structure in plants, before splitting off in the 17th century. It comes
originally from the Latin name of the Italian goddess of flowers, Flora. The early word for
flower in English was blossom,[4] though it now refers to flowers only of fruit trees.[5]

Morphology

Diagram of flower parts.

Main article: Floral morphology

The morphology of a flower, or its form and structure,[6] can be considered in two parts:
the vegetative part, consisting of non-reproductive structures such as petals; and the
reproductive or sexual parts. A stereotypical flower is made up of four kinds of
structures attached to the tip of a short stalk or axis, called a receptacle. Each of these
parts or floral organs is arranged in a spiral called a whorl.[7] The four main whorls
(starting from the base of the flower or lowest node and working upwards) are
the calyx, corolla, androecium, and gynoecium. Together the calyx and corolla make up
the non-reproductive part of the flower called the perianth, and in some cases may not
be differentiated. If this is the case, then they are described as tepals.[8]
Perianth
Main article: Perianth
Calyx
The sepals, collectively called the calyx, are modified leaves that occur on the
outermost whorl of the flower. They are leaf-like, in that they have a broad
base, stomata, stipules, and chlorophyll.[9] Sepals are often waxy and tough, and grow
quickly to protect the flower as it develops.[9][10] They may be deciduous, but will more
commonly grow on to assist in fruit dispersal. If the calyx is fused together it is called
gamosepalous.[9]
Corolla
The petals, together the corolla, are almost or completely fiberless leaf-like structures
that form the innermost whorl of the perianth. They are often delicate and thin, and are
usually coloured, shaped, or scented to encourage pollination. [11] Although similar to
leaves in shape, they are more comparable to stamens in that they form almost
simultaneously with one another, but their subsequent growth is delayed. If the corolla is
fused together it is called sympetalous.[12]
Reproductive
Main article: Plant reproductive morphology

Reproductive parts of Easter Lily (Lilium longiflorum). 1. Stigma, 2. Style, 3. Stamens, 4. Filament, 5. Petal

Androecium
The androecium, or stamens, is the whorl of pollen-producing male parts. Stamens
consist typically of an anther, made up of four pollen sacs arranged in two thecae,
connected to a filament, or stalk. The anther contains microsporocytes which
become pollen, the male gametophyte, after undergoing meiosis. Although they exhibit
the widest variation among floral organs, the androecium is usually confined just to one
whorl and to two whorls only in rare cases. Stamens range in number, size, shape,
orientation, and in their point of connection to the flower. [11][12]
Gynoecium
The gynoecium, or the carpels, is the female part of the flower found on the innermost
whorl. Each carpel consists of a stigma, which receives pollen, a style, which acts as a
stalk, and an ovary, which contains the ovules. Carpels may occur in one to several
whorls, and when fused together are often described as a pistil. Inside the ovary,
the ovules are suspended off of pieces of tissue called the placenta.[13][14]
Variation
Although this arrangement is considered "typical", plant species show a wide variation
in floral structure.[15] 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 or look like what is typically another part. [16] In
some families, like Ranunculaceae, the petals are greatly reduced; 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.[17]
Many flowers have symmetry. When the perianth is bisected through the central axis
from any point and symmetrical halves are produced, the flower is said to
be actinomorphic or regular. This is an example of radial symmetry. When flowers are
bisected and produce only one line that produces symmetrical halves, the flower is said
to be irregular or zygomorphic. If, in rare cases, they have no symmetry at all they are
called asymmetric.[18][19]
Flowers may be directly attached to the plant at their base (sessile—the supporting stalk
or stem is highly reduced or absent).[20] The stem or stalk subtending a flower, or
an inflorescence of flowers, is called a peduncle. If a peduncle supports more than one
flower, the stems connecting each flower to the main axis are called pedicels.[21] The
apex of a flowering stem forms a terminal swelling which is called the torus or
receptacle.[19]
In the majority of species, individual flowers have both pistils and stamens. These
flowers are described by botanists as being perfect, bisexual, or hermaphrodite.
However, in some species of plants the flowers are imperfect or unisexual: having only
either male (stamens) or female (pistil) parts. In the latter case, if an individual plant is
either female or male the species is regarded as dioecious. However, where unisexual
male and female flowers appear on the same plant, the species is called monoecious.
[22]
 Many flowers have nectaries, which are glands that produce a sugary fluid used to
attract pollinators. They are not considered as an organ on their own. [23]
Inflorescence
Main article: Inflorescence
The calla lily is not a single flower. It is actually an inflorescence of tiny flowers pressed together on a central
stalk that is surrounded by a large petal-like bract.[24]

In those species that have more than one flower on an axis, the collective cluster of
flowers is called an inflorescence. Some inflorescences are composed of many small
flowers arranged in a formation that resembles a single flower. A 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). [25] An inflorescence may
include specialized stems and modified leaves known as bracts.[26]
Floral diagrams and formulae
Main articles: Floral formula and Floral diagram
A floral formula is a way to represent the structure of a flower using specific letters,
numbers, and symbols, presenting substantial information about the flower in a compact
form. It can represent a taxon, usually giving ranges of the numbers of different organs,
or particular species. Floral formulae have been developed in the early 19th century and
their use has declined since. Prenner et al. (2010) devised an extension of the existing
model to broaden the descriptive capability of the formula. [27] The format of floral
formulae differs in different parts of the world, yet they convey the same information. [28][29]
[30][31]

The structure of a flower can also be expressed by the means of floral diagrams. The
use of schematic diagrams can replace long descriptions or complicated drawings as a
tool for understanding both floral structure and evolution. Such diagrams may show
important features of flowers, including the relative positions of the various organs,
including the presence of fusion and symmetry, as well as structural details. [32]

Development
A flower develops on a modified shoot or axis from a determinate
apical meristem (determinate meaning the axis grows to a set size). It has compressed
internodes, bearing structures that in classical plant morphology are interpreted as
highly modified leaves.[33] Detailed developmental studies, however, have shown that
stamens are often initiated more or less like modified stems (caulomes) that in some
cases may even resemble branchlets.[34][15] Taking into account the whole diversity in the
development of the androecium of flowering plants, we find a continuum between
modified leaves (phyllomes), modified stems (caulomes), and modified branchlets
(shoots).[35][36]
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.[37] 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.[38]

The ABC model of flower development

The first step of the transition 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.[39]
Organ development
Main article: ABC model of flower development
The ABC model is a simple model that describes the genes responsible for the
development of flowers. Three gene activities interact in a combinatorial manner to
determine the developmental identities of the primordia organ within the floral apical
meristem. These gene functions are called A, B, and C. A genes are expressed in only
outer and lower most section of the apical meristem, which becomes a whorl 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
aberrant flowers and mutations in Arabidopsis thaliana and the
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.[40]

Function
See also: Plant reproductive morphology
The principal purpose of a flower is the reproduction of the individual and the species.
All flowering plants are heterosporous, that is, every individual plant produces two types
of spores. Microspores are produced by meiosis inside anthers and megaspores are
produced inside ovules that are within an ovary. 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, i. e., they are endosporic.
In the majority of plant species, individual flowers have both functional carpels and
stamens. Botanists describe these flowers as perfect or bisexual, and the species
as hermaphroditic. In a minority of plant species, their flowers lack one or the other
reproductive organ and are described as imperfect or unisexual. If the individual plants
of a species each have unisexual flowers of both sexes then the species is monoecious.
Alternatively, if each individual plant has only unisexual flowers of the same sex then
the species is dioecious.

Pollination
Main article: Pollination

A Tūī, Prosthemadera novaeseelandiae, feeding on flax flower nectar, with yellow pollen on its forehead
Grains of pollen sticking to this bee will be transferred to the next flower it visits.

The primary purpose of the flower is reproduction.[41] Since the flowers are the
reproductive organs of the plant, they mediate the joining of the sperm, contained within
pollen, to the ovules — contained in the ovary.[10] Pollination is the movement of pollen
from the anthers to the stigma.[42] Normally pollen is moved from one plant to another,
known as cross-pollination, but many plants are able to self-pollinate. Cross-pollination
is preferred because it allows for genetic variation, which contributes to the survival of
the species.[43] Many flowers are dependent, then, upon external factors for pollination,
such as: the wind, water, animals, and especially insects. Larger animals such as birds,
bats, and even some pygmy possums,[44] however, can also be employed.[45][46] To
accomplish this, flowers have specific designs which encourage the transfer of pollen
from one plant to another of the same species. The period of time during which this
process can take place (when the flower is fully expanded and functional) is
called anthesis,[47] hence the study of pollination biology is called anthecology.[48]
Flowering plants usually face evolutionary 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.[49] Pollen may be transferred between plants via a number of
'vectors,' or methods. Around 80% of flowering plants make use of biotic, or living
vectors. Others use abiotic, or non-living, vectors and some plants make use of multiple
vectors, but most are highly specialised.[50]
Though some fit between or outside of these groups, [51] most flowers can be divided
between the following two broad groups of pollination methods:
Biotic pollination
Flowers that use biotic vectors 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).[52] 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.[53] Many flowers rely on simple proximity between flower parts
to ensure pollination, while others have elaborate designs to ensure pollination and
prevent self-pollination.[43] Flowers use animals including: insects (entomophily), birds
(ornithophily), bats (chiropterophily), lizards,[46] and even snails and slugs (malacophilae).
[54]
Attraction methods

Ophrys apifera, a bee orchid, which has evolved over many generations to mimic a female bee. [55]

Plants cannot move from one location to another, thus many flowers have evolved to
attract animals to transfer pollen between individuals in dispersed populations. Most
commonly, flowers are insect-pollinated, known as entomophilous; literally "insect-
loving" in Greek.[56] To attract these insects flowers commonly have glands
called nectaries on various parts that attract animals looking for nutritious nectar.[57] Birds
and bees have color vision, enabling them to seek out "colorful" flowers.[58] 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.[59]
Flowers also attract pollinators by scent, though 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. These are often called Carrion flowers,
including plants in the genus Rafflesia, and the titan arum.[58] Flowers pollinated by night
visitors, including bats and moths, are likely to concentrate on scent to attract pollinators
and so most such flowers are white.[60]
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. Other flowers use mimicry or pseudocopulation to attract
pollinators. Many orchids for example, produce flowers resembling female bees or
wasps in colour, shape, and scent. Males move from one flower to the next in search of
a mate, pollinating the flowers.[61][62]
Pollinator relationships
Further information: Pollination syndrome
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 an example
of coevolution, as the flower and pollinator have developed together over a long period
of time to match each other's needs.[63] This close relationship compounds the negative
effects of extinction, however, since the extinction of either member in such a
relationship would almost certainly mean the extinction of the other member as well. [64]
Abiotic pollination
Main articles: Anemophily and Hydrophily

A Grass flower with its long, thin filaments and large feathery stigma.

The female flower of Enhalus acoroides, which is pollinated through a combination of Hyphydrogamy and
Ephydrogamy.

Flowers that use abiotic, or non-living, vectors use the wind or, much less
commonly, water, to move pollen from one flower to the next. [50] In wind-dispersed
(anemophilous) species, the tiny pollen grains are carried, sometimes many thousands
of kilometres,[65] by the wind to other flowers. Common examples include
the grasses, birch trees, along with many other species in the order fagales,
[66]
 ragweeds, and many sedges. They have no need to attract pollinators and therefore
tend not to grow large, showy, or colorful flowers, and do not have nectaries, nor a
noticeable scent. Because of this, plants typically have many thousands of tiny flowers
which have comparatively large, feathery stigmas; to increase the chance of pollen
being received.[61] Whereas the pollen of entomophilous flowers is usually large, sticky,
and rich in protein (to act as a "reward" for pollinators), anemophilous flower pollen is
typically small-grained, very light, smooth, and of little nutritional value to insects.[67][68] In
order for the wind to effectively pick up and transport the pollen, the flowers typically
have anthers loosely attached to the end of long thin filaments, or pollen f