Flower

A flower, also known as a bloom or blossom, is

the reproductive structure found in flowering
plants (plants of the division Angiospermae).
Flowers consist of a combination of vegetative
organs – sepals that enclose and protect the
developing flower, petals that attract pollinators,
and reproductive organs that produce
gametophytes, which in flowering plants produce
gametes. The male gametophytes, which produce
sperm, are enclosed within pollen grains produced
in the anthers. The female gametophytes are
contained within the ovules produced in the
carpels.

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.[1] In this way, many
flowering plants have co-evolved with pollinators
to 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.[2]

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 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.[3] Some flowers produce diaspores without
fertilization (parthenocarpy). 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

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 Diagram of flower parts.
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

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 and chlorophyll[9] and may have stipules.
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

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]

In general, there is only one type of stamen, but there are plant
species where the flowers have two types; a "normal" one and one
with anthers that produce sterile pollen meant to attract
pollinators.[13]
Reproductive parts of easter lily
(Lilium longiflorum). 1. Stigma,
Gynoecium 2. Style, 3. Stamens, 4. Filament,
5. Petal
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 are often described as a pistil. Inside the ovary, the ovules
are attached to the placenta by structures called funiculi.[14][15]

Variation
Although this arrangement is considered "typical", plant species show a wide variation in floral
structure.[16] 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.[17] In some families, such as the grasses, 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.[18]

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.[19][20]
Flowers may be directly attached to the plant at their base (sessile—the supporting stalk or stem is highly
reduced or absent).[21] 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.[22] The apex of a flowering stem forms a terminal swelling which is called the torus or
receptacle.[20]

In the majority of species, individual flowers have both pistils and stamens. These flowers are described by
botanists as being perfect, bisexual, or hermaphrodite. In some species of plants, the flowers are imperfect
or unisexual: having only either male (stamens) or female (pistil) parts. If unisexual male and female flowers
appear on the same plant, the species is called monoecious.[23] However, if an individual plant is either
female or male, the species is called dioecious. 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.[24]

Inflorescence
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).[26] An inflorescence may include specialized stems and
modified leaves known as bracts.[27] The calla lily is not a single flower. It
is actually an inflorescence of tiny
flowers pressed together on a central
Floral diagrams and formulae stalk that is surrounded by a large
A floral formula is a way to represent the structure of a flower using petal-like bract.[25]
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.[28] The format of floral
formulae differs in different parts of the world, yet they convey the same information.[29][30][31][32]

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.[33]

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.[34] 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.[35][16] 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).[36][37]

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.[38] 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.[39]

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.[40]

Organ development The ABC model of flower

The ABC model is a simple model that describes the genes responsible for development

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.[41]

Function
The principal purpose of a flower is the reproduction[42] 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.

Pollination
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.[43] 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.[44] Many flowers depend on external factors for pollination,
such as: the wind, water, animals, and especially insects. Larger
animals such as birds, bats, and even some pygmy possums,[45] Grains of pollen sticking to this
[46][47] bee will be transferred to the next
however, can also be employed. To accomplish this, flowers
flower it visits.
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,[48] hence
the study of pollination biology is called anthecology.[49]

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 behavior of the plants.[50] 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.[51]

Though some fit between or outside of these groups,[52] 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).[53] 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.[54] Many flowers rely on simple proximity between flower parts to ensure pollination,
while others have elaborate designs to ensure pollination and prevent self-pollination.[44] Flowers use
animals including: insects (entomophily), birds (ornithophily), bats (chiropterophily), lizards,[47] and even
snails and slugs (malacophilae).[55]

Attraction methods
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.[57] To attract these insects flowers commonly have glands
called nectaries on various parts that attract animals looking for
nutritious nectar.[58] Some flowers have glands called elaiophores,
which produce oils rather than nectar.[59] Birds and bees have color
vision, enabling them to seek out colorful flowers.[60] 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.[61]

Flowers also attract pollinators by scent, though not all flower scents Ophrys apifera, a bee orchid, which
are appealing to humans; a number of flowers are pollinated by has evolved over many generations
insects that are attracted to rotten flesh and have flowers that smell to mimic a female bee.[56]
like dead animals. These are often called Carrion flowers, including
plants in the genus Rafflesia, and the titan arum.[60] 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.[62] Some plants pollinated by bats have a sonar-reflecting petal above
its flowers, which helps the bat find them,[63] and one species, the cactus Espostoa frutescens, has flowers
that are surrounded by an area of sound-absorbent and wooly hairs called the cephalium, which absorbs the
bat's ultrasound instead.[64]

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.[65][66]

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 an example of coevolution, as the flower and pollinator have
developed together over a long period of time to match each other's needs.[67] 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.[68]

Abiotic pollination
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.[51]
In wind-dispersed (anemophilous) species, the tiny pollen grains are
carried, sometimes many thousands of kilometres,[69] by the wind to
other flowers. Common examples include the grasses, birch trees,
along with many other species in the order Fagales,[70] 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 A grass flower with its long, thin
many thousands of tiny flowers which have comparatively large, filaments and large feathery
feathery stigmas; to increase the chance of pollen being received.[65] stigma.
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.[71][72] 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
forms around a catkin which moves in the wind. Rarer forms of this
involve individual flowers being moveable by the wind (Pendulous), The female flower of Enhalus
or even less commonly; the anthers exploding to release the pollen acoroides, which is pollinated
into the wind.[71] through a combination of
hyphydrogamy and ephydrogamy.
Pollination through water (hydrophily) is a much rarer method,
occurring in only around 2% of abiotically pollinated flowers.[51]
Common examples of this include Calitriche autumnalis, Vallisneria spiralis and some sea-grasses. One
characteristic which most species in this group share is a lack of an exine, or protective layer, around the
pollen grain.[73] Paul Knuth identified two types of hydrophilous pollination in 1906 and Ernst
Schwarzenbach added a third in 1944. Knuth named his two groups 'Hyphydrogamy' and the more
common 'Ephydrogamy'.[74] In hyphydrogamy pollination occurs below the surface of the water and so the
pollen grains are typically negatively buoyant. For marine plants that exhibit this method the stigmas are
usually stiff, while freshwater species have small and feathery stigmas.[75] In ephydrogamy pollination
occurs on the surface of the water and so the pollen has a low density to enable floating, though many also
use rafts, and are hydrophobic. Marine flowers have floating thread-like stigmas and may have adaptations
for the tide, while freshwater species create indentations in the water.[75] The third category, set out by
Schwarzenbach, is those flowers which transport pollen above the water through conveyance. This ranges
from floating plants, (Lemnoideae), to staminate flowers (Vallisneria). Most species in this group have dry,
spherical pollen which sometimes forms into larger masses, and female flowers which form depressions in
the water; the method of transport varies.[75]

Mechanisms
Flowers can be pollinated by two mechanisms; cross-pollination and self-pollination. No mechanism is
indisputably better than the other as they each have their advantages and disadvantages. Plants use one or
both of these mechanisms depending on their habitat and ecological niche.[76]

Cross-pollination
Cross-pollination is the pollination of the carpel by pollen from a different plant of the same species.
Because the genetic make-up of the sperm contained within the pollen from the other plant is different, their
combination will result in a new, genetically distinct, plant, through the process of sexual reproduction.
Since each new plant is genetically distinct, the different plants show variation in their physiological and
structural adaptations and so the population as a whole is better prepared for an adverse occurrence in the
environment. Cross-pollination, therefore, increases the survival of the species and is usually preferred by
flowers for this reason.[44][77]

The principal adaptive function of flowers is the promotion of cross-pollination or outcrossing, a process
that allows the masking of deleterious mutations in the genome of progeny. The masking effect of
outcrossing sexual reproduction is known as “genetic complementation”.[78] This beneficial effect of
outcrossing on progeny is also recognized as hybrid vigor or heterosis. Once outcrossing is established due
to the benefits of genetic complementation, subsequent switching to inbreeding becomes disadvantageous
because it allows expression of the previously masked deleterious recessive mutations, usually referred to as
inbreeding depression. Charles Darwin in his 1889 book The Effects of Cross and Self-Fertilization in the
Vegetable Kingdom[79] at the beginning of chapter XII noted “The first and most important of the
conclusions which may be drawn from the observations given in this volume, is that generally cross-
fertilisation is beneficial and self-fertilisation often injurious, at least with the plants on which I
experimented.”

Self-pollination
Self-pollination is the pollination of the carpel of a flower by pollen
from either the same flower or another flower on the same plant,[44]
leading to the creation of a genetic clone through asexual
reproduction. This increases the reliability of producing seeds, the
rate at which they can be produced, and lowers the amount energy
needed.[80] But, most importantly, it limits genetic variation. In
addition, self-pollination causes inbreeding depression, due largely
to the expression of recessive deleterious mutations.[81][82]
Clianthus puniceus, the kakabeak.
The extreme case of self-fertilization, when the ovule is fertilized by
pollen from the same flower or plant, occurs in flowers that always
self-fertilize, such as many dandelions.[83] Some flowers are self-pollinated and have flowers that never
open or are self-pollinated before the flowers open; these flowers are called cleistogamous; many species in
the genus Viola exhibit this, for example.[84]

Conversely, many species of plants have ways of preventing self-pollination and hence, 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-incompatible.[23][85] In Clianthus puniceus, self-
pollination is used strategically as an "insurance policy". When a pollinator, in this case a bird, visits C.
puniceus, it rubs off the stigmatic covering and allows for pollen from the bird to enter the stigma. If no
pollinators visit, however, then the stigmatic covering falls off naturally to allow for the flower's own
anthers to pollinate the flower through self-pollination.[80]

Allergies
Pollen is a large contributor to asthma and other respiratory allergies which combined affect between 10 and
50% of people worldwide. This number appears to be growing, as the temperature increases due to climate
change mean that plants are producing more pollen, which is also more allergenic. Pollen is difficult to
avoid, however, because of its small size and prevalence in the natural environment. Most of the pollen
which causes allergies is that produced by wind-dispersed pollinators such as the grasses, birch trees, oak
trees, and ragweeds; the allergens in pollen are proteins which are thought to be necessary in the process of
pollination.[86][87]

Fertilization
Fertilization, also called Synagmy, occurs following pollination, which
is the movement of pollen from the stamen to the carpel. It
encompasses both plasmogamy, the fusion of the protoplasts, and
karyogamy, the fusion of the nuclei. When pollen lands on the stigma
of the flower it begins creating a pollen tube which runs down through
the style and into the ovary. After penetrating the centre-most part of
the ovary it enters the egg apparatus and into one synergid. At this
point the end of the pollen tube bursts and releases the two sperm cells,
one of which makes its way to an egg, while also losing its cell
membrane and much of its protoplasm. The sperm's nucleus then fuses
with the egg's nucleus, resulting in the formation of a zygote, a diploid
(two copies of each chromosome) cell.[88] A floral diagram, with the pollen
tube labelled PG
Whereas in fertilization only plasmogamy, or the fusion of the whole
sex cells, results, in Angiosperms (flowering plants) a process known
as double fertilization, which involves both karyogamy and plasmogamy, occurs. In double fertilization the
second sperm cell subsequently also enters the synergid and fuses with the two polar nuclei of the central
cell. Since all three nuclei are haploid, they result in a large endosperm nucleus which is triploid.[88]

Seed development
Following the formation of zygote it begins to grow through
nuclear and cellular divisions, called mitosis, eventually becoming a
small group of cells. One section of it becomes the embryo, while
the other becomes the suspensor; a structure which forces the
embryo into the endosperm and is later undetectable. Two small
primordia also form at this time, that later become the cotyledon,
which is used as an energy store. Plants which grow out one of
these primordia are called monocotyledons, while those that grow
The fruit of a peach with the seed or
out two are dicotyledons. The next stage is called the Torpedo stage
stone inside.
and involves the growth of several key structures, including: the radicle (embryotic root), the epicotyl
(embryotic stem), and the hypocotyl, (the root/shoot junction). In the final step vascular tissue develops
around the seed.[89]

Fruit development
The ovary, inside which the seed is forming from the ovule, grows into a fruit. All the other main floral parts
die during this development, including: the style, stigma, sepals, stamens, and petals. The fruit contains
three structures: the exocarp, or outer layer, the mesocarp, or the fleshy part, and the endocarp, or innermost
layer, while the fruit wall is called the pericarp. The size, shape, toughness, and thickness varies among
different fruit. This is because it is directly connected to the method of seed dispersal; that being the purpose
of fruit - to encourage or enable the seed's dispersal and protect the seed while doing so.[89]

Seed dispersal

The kererū, Hemiphaga A samara from a maple tree with Acaena novae-zelandiae uses
[91]
novaeseelandiae, is an important its distinctive "wings." epizoochory to disperse its
disperser of seeds in New seeds.[92]
Zealand.[90]

Following the pollination of a flower, fertilization, and finally the development of a seed and fruit, a
mechanism is typically used to disperse the fruit away from the plant.[93] In Angiosperms (flowering plants)
seeds are dispersed away from the plant so as to not force competition between the mother and the daughter
plants,[94] as well as to enable the colonisation of new areas. They are often divided into two categories,
though many plants fall in between or in one or more of these:[95]

Allochory
In allochory, plants use an external vector, or carrier, to transport their seeds away from them. These can be
either biotic (living), such as by birds and ants, or abiotic (non-living), such as by the wind or
water.[95][96][97]

Biotic vectors
Many plants use biotic vectors to disperse their seeds away from them. This method falls under the umbrella
term zoochory, while endozoochory, also known as fruigivory, refers specifically to plants adapted to grow
fruit in order to attract animals to eat them. Once eaten they go through typically go through animal's
digestive system and are dispersed away from the plant.[97] Some seeds are specially adapted either to last
in the gizzard of animals or even to germinate better after passing through them.[98][99] They can be eaten
by birds (ornithochory), bats (chiropterochory), rodents, primates, ants (myrmecochory),[100] non-bird
sauropsids (saurochory), mammals in general (mammaliochory),[98] and even fish.[101] Typically their fruit
are fleshy, have a high nutritional value, and may have chemical attractants as an additional "reward" for
dispersers. This is reflected morphologically in the presence of more pulp, an aril, and sometimes an
elaiosome (primarily for ants), which are other fleshy structures.[102]

Epizoochory occurs in plants whose seeds are adapted to cling on to animals and be dispersed that way,
such as many species in the genus Acaena.[103] Typically these plants seed's have hooks or a viscous
surface to easier grip to animals, which include birds and animals with fur. Some plants use mimesis, or
imitation, to trick animals into dispersing the seeds and these often have specially adapted colors.[102][104]

The final type of zoochory is called synzoochory, which involves neither the digestion of the seeds, nor the
unintentional carrying of the seed on the body, but the deliberate carrying of the seeds by the animals. This
is usually in the mouth or beak of the animal (called Stomatochory), which is what is used for many birds
and all ants.[105]

Abiotic vectors
In abiotic dispersal plants use the vectors of the wind, water, or a
mechanism of their own to transport their seeds away from them.[97][96]
Anemochory involves using the wind as a vector to disperse plant's seeds.
Because these seeds have to travel in the wind, they are almost always
small — sometimes even dust-like, have a high surface-area-to-volume
ratio, and are produced in a large number — sometimes up to a million. The lichen Usnea angulata,
which uses hydrochory, is a
Plants such as tumbleweeds detach the entire shoot to let the seeds roll
weed in New Zealand.[106]
away with the wind. Another common adaptation are wings, plumes or
balloon-like structures that let the seeds stay in the air for longer and hence
travel farther.

In hydrochory plants are adapted to disperse their seeds through bodies of

water and so typically are buoyant and have a low relative density with
regards to the water. Commonly seeds are adapted morphologically with
hydrophobic surfaces, small size, hairs, slime, oil, and sometimes air Hura crepitans disperses its
spaces within the seeds.[102] These plants fall into three categories: ones seeds ballistically and is
where seeds are dispersed on the surface of water currents, under the hence commonly called the
surface of water currents, and by rain landing on a plant.[106] "dynamite tree".[107]

Autochory
In autochory, plants create their own vectors to transport the seeds away from them. Adaptations for this
usually involve the fruits exploding and forcing the seeds away ballistically, such as in Hura crepitans,[107]
or sometimes in the creation of creeping diaspores.[102] Because of the relatively small distances that these
methods can disperse their seeds, they are often paired with an external vector.[104]

Evolution
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 damage 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.

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,

Archaefructus liaoningensis, one of
along with further discoveries of fossil gymnosperms, suggest how
the earliest known flowering plants
angiosperm characteristics may have been acquired in a series of
steps. An early fossil of a flowering plant, Archaefructus
liaoningensis from China, is dated about 125 million years old.[108][109] Even earlier from China is the
125–130 million years old Archaefructus sinensis. In 2015 a plant (130 million-year-old Montsechia vidalii,
discovered in Spain) was claimed to be 130 million years old.[110] In 2018, scientists reported that the
earliest flowers began about 180 million years ago.[111]

Recent DNA analysis (molecular systematics)[112] shows that

Amborella trichopoda, found on the Pacific island of New
Caledonia, is the only species in the sister group to the rest of the
flowering plants, and morphological studies suggest that it has
features which may have been characteristic of the earliest
flowering plants.[113]

Besides the hard proof of flowers in or shortly before the

Cretaceous,[114][115] there is some circumstantial evidence of
Amborella trichopoda may have flowers as much as 250 million years ago. A chemical used by
characteristic features of the earliest plants to defend their flowers, oleanane, has been detected in fossil
flowering plants
plants that old, including gigantopterids,[116] 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.[117] 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".

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 contains.

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.

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.[118]

Colour
Many flowering plants reflect as much light as possible within the range of visible wavelengths of the
pollinator the plant intends to attract. Flowers that reflect the full range of visible light are generally
perceived as white by a human observer. An important feature of white flowers is that they reflect equally
across the visible spectrum. While many flowering plants use white to attract pollinators, the use of color is
also widespread (even within the same species). Color allows a flowering plant to be more specific about
the pollinator it seeks to attract. The color model used by human color reproduction technology (CMYK)
relies on the modulation of pigments that divide the spectrum into broad areas of absorption. Flowering
plants by contrast are able to shift the transition point wavelength between absorption and reflection. If it is
assumed that the visual systems of most pollinators view the visible spectrum as circular then it may be said
that flowering plants produce color by absorbing the light in one region of the spectrum and reflecting the
light in the other region. With CMYK, color is produced as a
function of the amplitude of the broad regions of absorption.
Flowering plants by contrast produce color by modifying the
frequency (or rather wavelength) of the light reflected. Most flowers
absorb light in the blue to yellow region of the spectrum and reflect
light from the green to red region of the spectrum. For many species
of flowering plant, it is the transition point that characterizes the
Reflectance spectra for the flowers
color that they produce. Color may be modulated by shifting the
of several varieties of rose. A red
transition point between absorption and reflection and in this way a rose absorbs about 99.7% of light
flowering plant may specify which pollinator it seeks to attract. across a broad area below the red
Some flowering plants also have a limited ability to modulate areas wavelengths of the spectrum, leading
of absorption. This is typically not as precise as control over to an exceptionally pure red. A yellow
wavelength. Humans observers will perceive this as degrees of rose will reflect about 5% of blue
saturation (the amount of white in the color). light, producing an unsaturated
yellow (a yellow with a degree of
white in it).

Classical taxonomy
In plant taxonomy, which is the study of the classification and
identification of plants, the morphology of plant's flowers are used
extensively – and have been for thousands of years. Although the
history of plant taxonomy extends back to at least around 300 B.C.
with the writings of Theophrastus,[120] the foundation of the modern
science is based on works in the 18th and 19th centuries.[121]

Carl Linnaeus (1707–1778), was a Swedish botanist who spent most

of his working life as a professor of natural history. His landmark
1757 book Species Plantarum lays out his system of classification as
well as the concept of binomial nomenclature, the latter of which is
still used around the world today.[121][note 1] He identified 24 classes,
Carl Linnaeus's method for
based mainly on the number, length and union of the stamens.
classifying plants focused solely on
the structure and nature of the
The first ten classes follow the number of stamens directly
flowers.[119] (Octandria have 8 stamens etc.),[119] while class eleven has 11–20
stamens and classes twelve and thirteen have 20 stamens; differing
only in their point of attachment. The next five classes deal with the
length of the stamens and the final five with the nature of the reproductive capability of the plant; where the
stamen grows; and if the flower is concealed or exists at all (such as in ferns). This method of classification,
despite being artificial,[119] was used extensively for the following seven decades, before being replaced by
the system of another botanist.[122]

Antoine Laurent de Jussieu (1748–1836) was a French botanist whose 1787 work Genera plantarum:
secundum ordines naturales disposita set out a new method for classifying plants; based instead on natural
characteristics. Plants were divided by the number, if any, of cotyledons, and the location of the
stamens.[122]
The next most major system of classification came in the late 19th century from the botanists Joseph Dalton
Hooker (1817–1911) and George Bentham (1800–1884). They built on the earlier works of de Jussieu and
Augustin Pyramus de Candolle and devised a system which is still used in many of the world's herbaria.

Plants were divided at the highest level by the number of cotyledons and the nature of the flowers, before
falling into orders (families), genera, and species. This system of classification was published in their
Genera plantarum in three volumes between 1862 and 1883.[123] It is the most highly regarded and
deemed the "best system of classification," in some settings.[124]

Following the development in scientific thought after Darwin's On the Origin of Species, many botanists
have used more phylogenetic methods and the use of genetic sequencing, cytology, and palynology has
become increasingly common. Despite this, morphological characteristics such as the nature of the flower
and inflorescence still make up the bedrock of plant taxonomy.[124][125]

Symbolism
Many flowers have important symbolic meanings in Western
culture.[126] 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.[127]
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 Lilies are often used to denote life or
"resurrection/life". It is also associated with stars (sun) resurrection
and its petals blooming/shining.
Daisies are a symbol of innocence.
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 floral art.

Flowers within art are also representative of the female

genitalia,[128] as seen in the works of artists such as Georgia
Flowers are common subjects of still
O'Keeffe, Imogen Cunningham, Veronica Ruiz de Velasco, and
life paintings, such as this one by
Judy Chicago, and in fact in Asian and western classical art. Many
Ambrosius Bosschaert the Elder
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.

Their symbolism in dreams has also been discussed, with possible interpretations including "blossoming
potential".[129]

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.[130] Apart from the association with Vishnu,
the Hindu tradition also considers the lotus to have spiritual significance.[131] For example, it figures in the
Hindu stories of creation.[132]

Human use
History shows that flowers have been used by humans for
thousands of years, to serve a variety of purposes. An early
example of this is from about 4,500 years ago in Ancient
Egypt, where flowers would be used to decorate women's hair.
Flowers have also inspired art time and time again, such as in
Monet's Water Lilies or William Wordsworth's poem about
daffodils entitled: "I Wandered Lonely as a Cloud".[133]

In modern times, people have sought ways to cultivate, buy,

Chancel flowers, placed upon the altar of
wear, or otherwise be around flowers and blooming plants,
St. Arsatius's Church in Ilmmünster
partly because of their agreeable appearance and smell. Around
the world, people use flowers to mark important events in their
lives:

For new births or christenings

As a corsage or boutonniere worn at social functions or
for holidays
As tokens of love or esteem
For wedding flowers for the bridal party, and as
decorations for wedding venues
As brightening decorations within the home
Brazilian sailors pay floral tribute to
As a gift of remembrance for bon voyage parties,
British naval flag officer Thomas
welcome-home parties, and "thinking of you" gifts
Cochrane in Westminster Abbey,
For funeral flowers and expressions of sympathy for the 1901
grieving
For worship. In Christianity, chancel flowers often adorn
churches.[134] In Hindu culture, adherents commonly bring flowers as a gift to temples[135]
Flowers like jasmine have been used as a replacement for traditional tea in China for centuries. Most
recently many other herbs and flowers used traditionally across the world are gaining importance to
preapare a range of floral tea.
People therefore grow flowers around their homes, dedicate parts of their
living space to flower gardens, pick wildflowers, or buy commercially-
grown flowers from florists. Flower production and trade supports
developing economies through their availability as a fair trade product.[136]

Flowers provide less food than other major plant parts (seeds, fruits, roots,
stems and leaves), but still provide several important vegetables 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; dried and ground marigold flowers
are also used as a spice and colouring agent in Georgian cuisine. Flowers of
the dandelion and elder are often made into wine. Bee pollen, pollen A woman spreading flowers
over a lingam in a temple in
collected from bees, is considered a health food by some people. Honey
Varanasi
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 only few are widely
marketed as food. They are often added to salads as garnishes.
Squash blossoms are dipped in breadcrumbs and fried. Some edible
flowers include nasturtium, chrysanthemum, carnation, cattail,
Japanese honeysuckle, chicory, cornflower, canna, and
sunflower.[137] Edible flowers such as daisy, rose, and violet are
sometimes candied.[138] Flowers collected for worship of
Hindu deities in morning, in West
Flowers such as chrysanthemum, rose, jasmine, Japanese Bengal.
honeysuckle, and chamomile, chosen for their fragrance and
medicinal properties, are used as tisanes, either mixed with tea or on
their own.[139]

Flowers have been used since prehistoric times in funeral rituals: traces of
pollen have been found on a woman's tomb in the El Miron Cave in
Spain.[140] Many cultures draw a connection between flowers and life and
death, and because of their seasonal return flowers also suggest rebirth,
which may explain why many people place flowers upon graves. The
View of the Tampere Central
ancient Greeks, as recorded in Euripides's play The Phoenician Women, Square during the Tampere
placed a crown of flowers on the head of the deceased;[141] they also Floral Festival in July 2007.
covered tombs with wreaths and flower petals. Flowers were widely used
in ancient Egyptian burials,[142] and the Mexicans to this day use flowers
prominently in their Day of the Dead celebrations[143] in the same way that their Aztec ancestors did.
 Eight Flowers, a painting by artist Qian Xuan, 13th century, Palace Museum, Beijing.

Giving
The flower-giving tradition goes back to prehistoric times when flowers
often had a medicinal and herbal attributes. Archaeologists found in several
grave sites remnants of flower petals. Flowers were first used as sacrificial
and burial objects. Ancient Egyptians and later Greeks and Romans used
flowers. In Egypt, burial objects from the time around 1540 BC were
found, which depicted red poppy, yellow Araun, cornflower and lilies.
Records of flower giving appear in Chinese writings and Egyptian
hieroglyphics, as well as in Greek and Roman mythology. The practice of
giving a flower flourished in the Middle Ages when couples showed
affection through flowers.

The tradition of flower-giving exists in many forms. It is an important part Flower market – Detroit's
of Russian culture and folklore. It is common for students to give flowers to Eastern Market
their teachers. To give yellow flowers in a romantic relationship means
break-up in Russia. Nowadays, flowers are often given away in the form of
a flower bouquet.[144][145][146]

See also
Floral color change
Flower preservation
Garden
List of garden plants
Plant evolutionary developmental biology
Plant reproductive morphology
 Sowing

Notes
1. His earlier works: Systema Naturae (1735) and Genera plantarum (1737) were also
influential in the field.[119]

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Further reading
Buchmann, Stephen (2016). The Reason for Flowers: Their History, Culture, Biology, and
How They Change Our Lives. Scribner. ISBN 978-1-4767-5553-3.
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(2nd ed.). New York: John Wiley & Sons. ISBN 978-0-471-24455-4.
Greyson, R.I. (1994). The Development of Flowers ([Link]
lo0000grey). Oxford University Press. ISBN 978-0-19-506688-3.
Leins, P. & Erbar, C. (2010). Flower and Fruit. Stuttgart: Schweizerbart Science Publishers.
ISBN 978-3-510-65261-7.
Sattler, R. (1973). Organogenesis of Flowers. A Photographic Text-Atlas. University of
Toronto Press. ISBN 978-0-8020-1864-9.

External links
Quotations related to Flowers at Wikiquote
Native Plant Information Network ([Link]
[Link]/)
Flower Database ([Link]

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