Flower

A flower, also known as a bloom or blossom,[1]

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 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 ovary.[2][3] In
some plants, multiple flowers occur singly on a
pedicel (flower stalk), and some are arranged in a
group (inflorescence) on a peduncle
(inflorescence stalk).[2]

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, large petals
with patterns only visible to under ultraviolet
light, attractive scents, and the production of nectar, a food source for pollinators.[4][2] 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.[5]

When pollen from the anther of a flower is transferred to the stigma to another, it 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.[6] Some flowers produce
diaspores without fertilization (parthenocarpy).[7] After fertilization, the ovary of the flower develops into
fruit containing seeds.[2]
Flowers have long been appreciated 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,[8] though it now
refers to flowers only of fruit trees.[9]

Morphology

The morphology of a flower, or its form

and structure,[10] 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 arranged in
whorls around the tip of a short stalk or
axis, called a receptacle.[11][2] The four
main whorls (starting from the base of
the flower or lowest node and working
Diagram of flower parts.
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
monocotyledons, may not be differentiated. If this is the case, then they are described as tepals.[2][12]

Perianth

Calyx
The sepals, collectively called the calyx, are modified leaves that occur on the outermost whorl of the
flower.[2] They are leaf-like, in that they have a broad base, stomata and chlorophyll and may have
stipules. Sepals are often waxy and tough, and grow quickly to protect the flower as it develops.[13][14]
They may be deciduous, but will more commonly grow on to assist in fruit dispersal. If the calyx is fused
it is called gamosepalous.[13]

Corolla
The petals, collectively called 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 colored, shaped, or
scented to encourage pollination.[2][15] 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.[16] In monocotyledonous flowers (e.g.,
Lilium sp.), petals and sepals are indistinguishable and are individually called tepals. Petals also tend to
have patterns only visible under ultraviolet light, which are visible to pollinators but not to humans.[2]

Reproductive

Androecium
The androecium, consisting of 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.[2] 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. [15][16]

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.[17]
Reproductive parts of easter lily
(Lilium longiflorum). 1. Stigma,
Gynoecium 2. Style, 3. Stamens, 4. Filament,
5. Petal
The gynoecium, consisting of one or more 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.[2] 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.[18][19]

Variation
Although this arrangement is considered "typical", plant species show a wide variation in floral
structure.[20] 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.[21] 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 petal-like
stamens; the double flowers of peonies and roses are mostly petaloid stamens.[22]

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. [23][24]
Flowers may be directly attached to the plant at their base (sessile—the supporting stalk or stem is highly
reduced or absent). [2][25] 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. [2][26] The apex of a flowering stem forms a terminal swelling which is called the
torus or receptacle.[2][24]

In the majority of species, individual flowers have both carpels 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 (stamen) or female (carpel) parts. If unisexual male and
female flowers appear on the same plant, the species is called monoecious. [27] However, if an individual
plant is either female or male, the species is called dioecious. Many flowers have floral nectaries, which
are glands that produce a sugary fluid (nectar) used to attract pollinators. They are not considered as an
organ on their own. [28]

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). [2][30] An inflorescence may include
specialized stems and modified leaves known as bracts. [2][31] The calla lily is not a single flower. It
is an inflorescence of tiny flowers
pressed together on a central stalk
Floral diagrams and formulae that is surrounded by a large petal-
A floral formula is a way to represent the structure of a flower like bract.[29]
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.[32]
The format of floral formulae differs in different parts of the world, yet they convey the same
information.[33][34][35][36]

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. [37]

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.[38] 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.[39][20] 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).[40][41]

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 can interpret important endogenous
and environmental cues such as changes in levels of plant hormones and seasonable temperature and
photoperiod changes.[42] 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 several different physiological and morphological changes.[43]

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 the 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 on some environmental cue.[44]

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. Genes are
expressed in only the 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 the B function but the presence of the C
function mimics the fourth whorl, leading to the formation of carpels also in the third whorl. [45]

Function
The principal purpose of a flower is the reproduction[46] 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. [14] Pollination is the movement of
pollen from the anthers to the stigma. [47] Normally pollen is moved
from one plant to another, known as cross-pollination, but many
plants can self-pollinate. Cross-pollination is preferred because it
allows for genetic variation, which contributes to the survival of the
species. [48] 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,[49] Grains of pollen sticking to this
bee will be transferred to the
however, can also be employed.[50][51] To accomplish this, flowers
next flower it visits.
have specific designs which encourage the transfer of pollen from
one plant to another of the same species. The period during which
this process can take place (when the flower is fully expanded and functional) is called anthesis,[52] hence
the study of pollination biology is called anthecology.[53]

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.[54] Pollen may be
transferred between plants via several '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 specialized.[55]

Though some fit between or outside of these groups,[56] 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). [57] 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. [58] Many flowers rely on simple proximity between flower parts to ensure
pollination, while others have elaborate designs to ensure pollination and prevent self-pollination. [48]
Flowers use animals including: insects (entomophily), birds (ornithophily), bats (chiropterophily),
lizards,[51] and even snails and slugs (malacophilae).[59]

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.[61] To attract these insects flowers commonly
have glands called nectaries on various parts that attract animals
looking for nutritious nectar.[62] Some flowers have glands called
elaiophores, which produce oils rather than nectar.[63] Birds and
bees have color vision, enabling them to seek out colorful
flowers.[64] 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.[65]
Ophrys apifera, a bee orchid, which
Flowers also attract pollinators by scent, though not all flower has evolved over many generations
scents are appealing to humans; several flowers are pollinated by to mimic a female bee.[60]
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. [64] 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.[66]
Some plants pollinated by bats have a sonar-reflecting petal above its flowers, which helps the bat find
them,[67] and one species, the cactus Espostoa frutescens, has flowers that are surrounded by an area of
sound-absorbent and woolly hairs called the cephalium, which absorbs the bat's ultrasound instead.[68]

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 color, shape, and scent. Males move from one flower to the next in
search of a mate, pollinating the flowers. [69][70]

Pollinator relationships
Many flowers have close relationships with one or a few specific pollinating organisms. Many flowers,
for example, attract only one specific species of insect and therefore rely on that insect for successful
reproduction. This close relationship is an example of coevolution, as the flower and pollinator have
developed together over a long period to match each other's needs. [71] 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.[72]

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.[55] In wind-dispersed (anemophilous) species, the tiny pollen
grains are carried, sometimes many thousands of kilometers,[73] by
the wind to other flowers. Common examples include the grasses,
birch trees, along with many other species in the order Fagales,[74]
ragweeds, and many sedges. They do not 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 A grass flower with its long, thin
typically have many thousands of tiny flowers which have filaments and a large feathery
stigma.
comparatively large, feathery stigmas; to increase the chance of
pollen being received. [69] 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.[75][76] 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 The female flower of Enhalus
acoroides, which is pollinated
individual flowers being moveable by the wind (pendulous), or even
through a combination of
less commonly; the anthers exploding to release the pollen into the hyphydrogamy and
wind. [75] ephydrogamy.

Pollination through water (hydrophily) is a much rarer method,

occurring in only around 2% of abiotically pollinated flowers.[55] 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. [77] 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'.[78] 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.[79] 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.[79] 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.[79]

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. [80]

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. [48][81]

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".[82] This beneficial effect of
outcrossing on progeny is also recognized as hybrid vigour or heterosis. Once outcrossing is established
due to the benefits of genetic complementation, subsequent switching to inbreeding becomes
disadvantageous because it allows the 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[83] 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,[48] 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.[84] But, most importantly, it limits genetic variation. In
addition, self-pollination causes inbreeding depression, due largely
to the expression of recessive deleterious mutations.[85][86]
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.[87] 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.[88]
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.[27][89] 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.[84]

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.[90][91]

Fertilization
Fertilization, also called Syngamy, is preceded by 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 center-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.[2][92] A floral diagram, with the pollen
tube labeled PG
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.[92]

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 out two are dicotyledons. The next stage is called the
Torpedo stage 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.[93]
The fruit of a peach with the seed or
stone inside.

Fruit development
The ovary, inside which the seed is forming from the ovule, grows into a fruit. All the other main floral
parts wither and 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 dry and fleshy fruits. 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.[93]

Seed dispersal

The kererū, Hemiphaga A samara from a maple tree with Acaena novae-zelandiae uses
[95]
novaeseelandiae, is an important its distinctive "wings." epizoochory to disperse its
disperser of seeds in New seeds.[96]
[94]
Zealand.

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.[97] In Angiosperms (flowering
plants) seeds are dispersed away from the plant so as to not force competition between the mother and the
daughter plants,[98] as well as to enable the colonization of new areas. They are often divided into two
categories, though many plants fall in between or in one or more of these:[99]

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.[99][100][101]

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.[101] Some seeds are specially
adapted either to last in the gizzard of animals or even to germinate better after passing through
them.[102][103] They can be eaten by birds (ornithochory), bats (chiropterochory), rodents, primates, ants
(myrmecochory),[104] non-bird sauropsids (saurochory), mammals in general (mammaliochory),[102] and
even fish.[105] 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.[106]

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.[107] 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.[106][108]

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.[109]

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.[101][100] 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. Plants such as tumbleweeds detach the entire shoot to let the seeds roll 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 spaces within
the seeds.[106] These plants fall into three categories: ones where seeds are dispersed on the surface of
water currents, under the surface of water currents, and by rain landing on a plant.[110]

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,[111] or
sometimes in the creation of creeping diaspores.[106] Because of the
relatively small distances that these methods can disperse their seeds,
they are often paired with an external vector.[108]

The lichen Usnea angulata,

which uses hydrochory, is a
Evolution weed in New Zealand.[110]

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 Hura crepitans disperses its
seeds ballistically and is
had not yet evolved the flower. Early seed-bearing plants include the
hence commonly called the
ginkgo and conifers.
"dynamite tree".[111]

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.
Archaefructus liaoningensis, one of
Recently discovered angiosperm fossils such as Archaefructus,
the earliest known flowering plants
along with further discoveries of fossil gymnosperms, suggest
how 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.[112][113] 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.[114] In 2018, scientists reported that the earliest flowers began
about 180 million years ago.[115]

Recent DNA analysis (molecular systematics)[116] 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.[117]

Besides the hard proof of flowers in or shortly before the Cretaceous,[118][119] there is some
circumstantial evidence of flowers as much as 250 million years ago. A chemical used by plants to defend
their flowers, oleanane, has been detected in fossil plants that old, including gigantopterids,[120] 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.[121] The most primitive
Amborella trichopoda may have flowers are thought to have had a variable number of flower parts,
characteristic features of the earliest often separate from (but in contact with) each other. The flowers
flowering plants
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.[122]

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
Reflectance spectra for the flowers
same species). Color allows a flowering plant to be more specific
of several varieties of rose. A red
about the pollinator it seeks to attract. The color model used by rose absorbs about 99.7% of light
human color reproduction technology (CMYK) relies on the across a broad area below the red
modulation of pigments that divide the spectrum into broad areas wavelengths of the spectrum,
of absorption. Flowering plants by contrast are able to shift the leading to an exceptionally pure red.
transition point wavelength between absorption and reflection. If it A yellow rose will reflect about 5% of
is assumed that the visual systems of most pollinators view the blue light, producing an unsaturated
yellow (a yellow with a degree of
visible spectrum as circular then it may be said that flowering
white in it).
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 color that they produce. Color may be modulated by shifting the transition point between absorption
and reflection and in this way a flowering plant may specify which pollinator it seeks to attract. Some
flowering plants also have a limited ability to modulate areas of absorption. This is typically not as
precise as control over wavelength. Humans observers will perceive this as degrees of saturation (the
amount of white in the color).

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,[124] the foundation of the
modern science is based on works in the 18th and 19th
centuries.[125]

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
Carl Linnaeus's method for
latter of which is still used around the world today.[125][note 1] He
classifying plants focused solely identified 24 classes, based mainly on the number, length and union
on the structure and nature of the of the stamens.
flowers.[123]
The first ten classes follow the number of stamens directly (Octandria have 8 stamens etc.),[123] 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,[123] was used
extensively for the following seven decades, before being replaced by the system of another botanist.[126]

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.[126]

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.[127] It is the most highly regarded and
deemed the "best system of classification," in some settings.[128]

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.[128][129]

Symbolism
Many flowers have important symbolic meanings in Western
culture.[130] 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.[131]
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,[132] as seen in the works of artists such as Georgia
O'Keeffe, Imogen Cunningham, Veronica Ruiz de Velasco, and
Judy Chicago, and in fact in Asian and western classical art. Many
cultures around the world have a marked tendency to associate
flowers with femininity.

Flowers are common subjects of still The great variety of delicate and beautiful flowers has inspired the
life paintings, such as this one by works of numerous poets, especially from the 18th–19th century
Ambrosius Bosschaert the Elder 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".[133]

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.[134] Apart from the association with Vishnu,
the Hindu tradition also considers the lotus to have spiritual significance.[135] For example, it figures in
the Hindu stories of creation.[136]

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".[137]

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
 As a gift of remembrance for bon voyage parties,
welcome-home parties, and "thinking of you" gifts
For funeral flowers and expressions of sympathy for the
grieving
For worship. In Christianity, chancel flowers often adorn
churches.[138] In Hindu culture, adherents commonly
bring flowers as a gift to temples[139]
Flowers like jasmine have been used as a replacement for
traditional tea in China for centuries. Most recently many other
Brazilian sailors pay floral tribute to
herbs and flowers used traditionally across the world are gaining British naval flag officer Thomas
importance to preapare a range of floral tea. Cochrane in Westminster Abbey,
1901
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.[140]

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 coloring agent in Georgian
cuisine. Flowers of the dandelion and elder are often made into wine. Bee A woman spreading flowers
pollen, pollen collected from bees, is considered a health food by some over a lingam in a temple in
people. Honey consists of bee-processed flower nectar and is often named Varanasi
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.[141] Edible flowers such as daisy, rose, and violet are
sometimes candied.[142] 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.[143]

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.[144] 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 ancient Greeks, as recorded
in Euripides's play The Phoenician Women, placed a crown of flowers on
the head of the deceased;[145] they also covered tombs with wreaths and
flower petals. Flowers were widely used in ancient Egyptian burials,[146]
and the Mexicans to this day use flowers prominently in their Day of the
Dead celebrations[147] in the same way that their Aztec ancestors did.

View of the Tampere

Central Square during the
Tampere Floral Festival in
July 2007.

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 Eastern Market
to their teachers. To give yellow flowers in a romantic relationship means
breakup in Russia. Nowadays, flowers are often given away in the form of
a flower bouquet.[148][149][150]

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.[123]

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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.
Esau, Katherine (1965). Plant Anatomy (https://archive.org/details/plantanatomy00esau_0)
(2nd ed.). New York: John Wiley & Sons. ISBN 978-0-471-24455-4.
Greyson, R.I. (1994). The Development of Flowers (https://archive.org/details/developmento
fflo0000grey). 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 (https://web.archive.org/web/20090510230926/http://wildfl
ower.utexas.edu/)
Flower Database (https://www.flower-db.com/en)
10 Famous Flower Paintings in History (https://www.belart-gallery.com/post/10-famous-flowe
r-paintings)

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