Flower
Encyclopedia
A flower, sometimes known as a bloom or blossom
, is the reproductive structure found in flowering plant
s (plants of the division Magnoliophyta, also called angiosperms). The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs. Flowers may facilitate outcrossing (fusion of sperm and eggs from different individuals in a population) or allow selfing (fusion of sperm and egg from the same flower). Some flowers produce diaspores
without fertilization (parthenocarpy
). Flowers contain sporangia and are the site where gametophytes develop. Flowers give rise to fruit and seeds. Many flowers have evolved to be attractive to animals, so as to cause them to be vectors for the transfer of pollen.
In addition to facilitating the reproduction of flowering plants, flowers have long been admired and used by humans to beautify their environment, but also as objects of romance, ritual, religion, medicine and as a source of food.
on the receptacle. The four main whorls (starting from the base of the flower or lowest node and working upwards) are as follows:
Although the arrangement described above is considered "typical", plant species show a wide variation in floral structure. These modifications have significance in the evolution of flowering plants and are used extensively by botanists to establish relationships among plant species.
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 and/or look like what is typically another part. In some families, like Ranunculaceae
, the petals are greatly reduced and 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 Rose
s are mostly petaloid stamens. Flowers show great variation and plant scientists describe this variation in a systematic way to identify and distinguish species.
Specific terminology is used to describe flowers and their parts. Many flower parts are fused together; fused parts originating from the same whorl are connate
, while fused parts originating from different whorls are adnate
, parts that are not fused are free. When petals are fused into a tube or ring that falls away as a single unit, they are sympetalous (also called gamopetalous.) Connate petals may have distinctive regions: the cylindrical base is the tube, the expanding region is the throat and the flaring outer region is the limb. A sympetalous flower, with bilateral symmetry with an upper and lower lip, is bilabiate. Flowers with connate petals or sepals may have various shaped corolla or calyx including: campanulate, funnelform, tubular, urceolate, salverform or rotate.
Many flowers have a symmetry. When the perianth is bisected through the central axis from any point, symmetrical halves are produced, forming a radial symmetry. These flowers are also known to be actinomorphic or regular, e.g. rose or trillium. When flowers are bisected and produce only one line that produces symmetrical halves the flower is said to be irregular or zygomorphic, e.g. snapdragon or most orchids.
Flowers may be directly attached to the plant at their base (sessile--the supporting stalk or stem is highly reduced or absent). The stem or stalk subtending a flower is called a peduncle
. If a peduncle supports more than one flower, the stems connecting each flower to the main axis are called pedicels
. The apex of a flowering stem forms a terminal swelling which is called the torus or receptacle.
rather than a particular species. The following representations are used:
x: to represent a "variable number"
∞: to represent "many"
A floral formula would appear something like this:
Several additional symbols are sometimes used (see Key to Floral Formulas).
. Some inflorescences are composed of many small flowers arranged in a formation that resembles a single flower. The 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).
An inflorescence may include specialized stems and modified leaves known as bract
s.
tip with compressed internodes, bearing structures that are highly modified leaves
. In essence, a flower develops on a modified shoot or axis from a determinate apical meristem
(determinate meaning the axis grows to a set size).
, 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. 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 bud
s and growing tips to induce a number of different physiological and morphological changes. The first step 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. Once the process begins, even if that cue is removed the stem will continue to develop a flower.
. These gene functions are called A, B and C-gene functions. In the first floral whorl only A-genes are expressed, leading to the formation 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 homeotic mutants in Arabidopsis
thaliana and 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. See also The ABC Model of Flower Development
.
Most genes central in this model belong to the MADS-box
genes and are transcription factors that regulate the expression of the genes specific for each floral organ.
s. Microspores are produced by meiosis inside anthers while megaspores are produced inside ovules, inside an ovary. In fact, 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 (are endosporic).
In the majority of species, individual flowers have both functional carpels and stamens. These flowers are described by botanists as being perfect or bisexual. Some flowers lack one or the other reproductive organ and called imperfect or unisexual If unisex flowers are found on the same individual plant but in different locations, the species is said to be monoecious
. If each type of unisex flower is found only on separate individuals, the plant is dioecious
.
, and this is typically reflected in the morphology of the flowers and the behaviour of the plants. Pollen may be transferred between plants via a number of 'vectors'. Some plants make use of abiotic vectors — namely wind (anemophily
) or, much less commonly, water (hydrophily
). Others use biotic vectors including insects (entomophily
), birds (ornithophily
), bats (chiropterophily) or other animals. Some plants make use of multiple vectors, but many are highly specialised.
Cleistogamous flower
s are self pollinated, after which they may or may not open. Many Viola and some Salvia species are known to have these types of flowers.
The flowers of plants that make use of biotic pollen vectors commonly have glands called nectaries that act as an incentive for animals to visit the flower. Some flowers have patterns, called nectar guide
s, that show pollinators where to look for nectar. Flowers also attract pollinators by scent and color. Still other flowers use mimicry to attract pollinators. Some species of orchids, for example, produce flowers resembling female bees in color, shape, and scent. Flowers are also specialized in shape and have an arrangement of the stamen
s 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). 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.
Anemophilous flower
s use the wind to move pollen from one flower to the next. Examples include grasses, birch trees, ragweed and maples. They have no need to attract pollinators and therefore tend not to be "showy" flowers. Male and female reproductive organs are generally found in separate flowers, the male flowers having a number of long filaments terminating in exposed stamens, and the female flowers having long, feather-like stigmas. Whereas the pollen of animal-pollinated flowers tends to be large-grained, sticky, and rich in protein
(another "reward" for pollinators), anemophilous flower pollen is usually small-grained, very light, and of little nutritional value to animals.
. Flowers have specific designs which encourages the transfer of pollen from one plant to another of the same species. Many plants are dependent upon external factors for pollination, including: wind and animals, and especially insect
s. Even large animals such as birds, bats, and pygmy possum
s can be employed. The period of time during which this process can take place (the flower is fully expanded and functional) is called anthesis.
. Flowers commonly have glands called nectaries on various parts that attract animals looking for nutritious nectar. Bird
s and bee
s have color vision, enabling them to seek out "colorful" flowers. Some flowers have patterns, called nectar guide
s, 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. Flowers also attract pollinators by scent
and some of those scents are pleasant to our sense of smell. Not all flower scents are appealing to humans; a number of flowers are pollinated by insects that are attracted to rotten flesh and have flowers that smell like dead animals, often called Carrion flower
s, including Rafflesia
, the titan arum
, and the North American pawpaw
(Asimina triloba). Flowers pollinated by night visitors, including bats and moths, are likely to concentrate on scent to attract pollinators and most such flowers are white.
Still other flowers use mimicry to attract pollinators. Some species of orchids, for example, produce flowers resembling female bees in color, shape, and scent. Male bees move from one such flower to another in search of a mate.
Most flowers can be divided between two broad groups of pollination methods:
Entomophilous: flowers 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). 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. Many flowers rely on simple proximity between flower parts to ensure pollination. Others, such as the Sarracenia or lady-slipper orchid
s, have elaborate designs to ensure pollination while preventing self-pollination
.
Anemophilous: flowers use the wind to move pollen from one flower to the next, examples include the grasses
, Birch trees, Ragweed and Maples. They have no need to attract pollinators and therefore tend not to be "showy" flowers. Whereas the pollen of entomophilous flowers tends to be large-grained, sticky, and rich in protein
(another "reward" for pollinators), anemophilous flower pollen is usually small-grained, very light, and of little nutritional value to insect
s, though it may still be gathered in times of dearth. Honeybees and bumblebees actively gather anemophilous corn (maize
) pollen, though it is of little value to them.
Some flowers are self pollinated and use flowers that never open or are self pollinated before the flowers open, these flowers are called cleistogamous. Many Viola species and some Salvia have these types of flowers.
This close relationship compounds the negative effects of extinction
. The extinction of either member in such a relationship would mean almost certain extinction of the other member as well. Some endangered plant species
are so because of shrinking pollinator populations
.
).
by a simple adaptation of their aquatic counterparts: spore
s. 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 abuse 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. The earliest fossil of a flowering plant, Archaefructus liaoningensis
, is dated about 125 million years old.
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, along with further discoveries of fossil gymnosperms, suggest how angiosperm characteristics may have been acquired in a series of steps.
Recent DNA
analysis (molecular systematics) shows that Amborella trichopoda
, found on the Pacific island of New Caledonia, is 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.
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 wasp
s 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.
While there is only hard proof of such flowers existing about 130 million years ago, there is some circumstantial evidence that they did exist up to 250 million years ago. A chemical used by plants to defend their flowers, oleanane
, has been detected in fossil plants that old, including gigantopterid
s, 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. 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".
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.
ic meanings in Western culture. The practice of assigning meanings to flowers is known as floriography. Some of the more common examples include:
Flowers within art are also representative of the female genitalia, 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
.
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.
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 flower art.
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. Apart from the association with Vishnu, the Hindu tradition also considers the lotus to have spiritual significance. For example, it figures in the Hindu stories of creation.
. Around the world, people use flowers for a wide range of events and functions that, cumulatively, encompass one's lifetime:
People therefore grow flowers around their homes, dedicate entire parts of their living space to flower garden
s, pick wildflowers, or buy flowers from florists who depend on an entire network of commercial growers and shippers to support their trade.
Flowers provide less food than other major plants parts (seed
s, fruit
s, root
s, stem
s and leaves
) but they provide several important foods and spice
s. Flower vegetables include broccoli
, cauliflower
and artichoke
. The most expensive spice, saffron
, consists of dried stigmas of a crocus
. Other flower spices are clove
s and caper
s. Hops
flowers are used to flavor beer
. Marigold
flowers are fed to chicken
s to give their egg yolks a golden yellow color, which consumers find more desirable. Dandelion flowers are often made into wine. Bee Pollen
, pollen collected from bees, is considered a health food by some people. Honey
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 few are widely marketed as food. They are often used to add color and flavor to salads. Squash
flowers are dipped in breadcrumbs and fried. Edible flowers include nasturtium, chrysanthemum
, carnation
, cattail, honeysuckle
, chicory
, cornflower
, Canna
, and sunflower
. Some edible flowers are sometimes candied such as daisy
and rose
(you may also come across a candied pansy
).
Flowers can also be made into herbal teas. Dried flowers such as chrysanthemum, rose, jasmine, camomile are infused into tea both for their fragrance and medical properties. Sometimes, they are also mixed with tea
leaves for the added fragrance.
Blossom
In botany, blossom is a term given to the flowers of stone fruit trees and of some other plants with a similar appearance that flower profusely for a period of time in spring...
, is the reproductive structure found in flowering plant
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...
s (plants of the division Magnoliophyta, also called angiosperms). The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs. Flowers may facilitate outcrossing (fusion of sperm and eggs from different individuals in a population) or allow selfing (fusion of sperm and egg from the same flower). Some flowers produce diaspores
Diaspore (botany)
In botany, a diaspore is a plant dispersal unit consisting of a seed or spore plus any additional tissues that assist dispersal. In some seed plants, the diaspore is a seed and fruit together, or a seed and elaiosome. In a few seed plants, the diaspore is most or all of the plant, known as a...
without fertilization (parthenocarpy
Parthenocarpy
In botany and horticulture, parthenocarpy is the natural or artificially induced production of fruit without fertilization of ovules. The fruit is therefore seedless. Stenospermocarpy may also produce apparently seedless fruit, but the seeds are actually aborted while still small...
). Flowers contain sporangia and are the site where gametophytes develop. Flowers give rise to fruit and seeds. Many flowers have evolved to be attractive to animals, so as to cause them to be vectors for the transfer of pollen.
In addition to facilitating the reproduction of flowering plants, flowers have long been admired and used by humans to beautify their environment, but also as objects of romance, ritual, religion, medicine and as a source of food.
Morphology
A stereotypical flower consists of four kinds of structures attached to the tip of a short stalk. Each of these kinds of parts is arranged in a whorlWhorl (botany)
In botany, a whorl is an arrangement of sepals, petals, leaves, or branches in which all the parts are attached at the same point and surround or wrap around the stem.There are four whorls in a general flower...
on the receptacle. The four main whorls (starting from the base of the flower or lowest node and working upwards) are as follows:
- CalyxSepalA sepal is a part of the flower of angiosperms . Collectively the sepals form the calyx, which is the outermost whorl of parts that form a flower. Usually green, sepals have the typical function of protecting the petals when the flower is in bud...
: the outermost whorl consisting of units called sepalSepalA sepal is a part of the flower of angiosperms . Collectively the sepals form the calyx, which is the outermost whorl of parts that form a flower. Usually green, sepals have the typical function of protecting the petals when the flower is in bud...
s; these are typically green and enclose the rest of the flower in the bud stage, however, they can be absent or prominent and petal-like in some species. - CorollaPetalPetals are modified leaves that surround the reproductive parts of flowers. They often are brightly colored or unusually shaped to attract pollinators. Together, all of the petals of a flower are called a corolla. Petals are usually accompanied by another set of special leaves called sepals lying...
: the next whorl toward the apex, composed of units called petalPetalPetals are modified leaves that surround the reproductive parts of flowers. They often are brightly colored or unusually shaped to attract pollinators. Together, all of the petals of a flower are called a corolla. Petals are usually accompanied by another set of special leaves called sepals lying...
s, which are typically thin, soft and colored to attract animals that help the process of pollinationPollinationPollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...
. - Androecium (from Greek andros oikia: man's house): the next whorl (sometimes multiplied into several whorls), consisting of units called stamenStamenThe stamen is the pollen producing reproductive organ of a flower...
s. Stamens consist of two parts: a stalk called a filamentStamenThe stamen is the pollen producing reproductive organ of a flower...
, topped by an anther where pollenPollenPollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...
is produced by meiosis and eventually dispersed. - GynoeciumGynoeciumGynoecium is most commonly used as a collective term for all carpels in a flower. A carpel is the ovule and seed producing reproductive organ in flowering plants. Carpels are derived from ovule-bearing leaves which evolved to form a closed structure containing the ovules...
(from Greek gynaikos oikia: woman's house): the innermost whorl of a flower, consisting of one or more units called carpelGynoeciumGynoecium is most commonly used as a collective term for all carpels in a flower. A carpel is the ovule and seed producing reproductive organ in flowering plants. Carpels are derived from ovule-bearing leaves which evolved to form a closed structure containing the ovules...
s. The carpel or multiple fused carpels form a hollow structure called an ovary, which produces ovules internally. Ovules are megasporangia and they in turn produce megaspores by meiosis which develop into female gametophytes. These give rise to egg cells. The gynoecium of a flower is also described using an alternative terminology wherein the structure one sees in the innermost whorl (consisting of an ovary, style and stigma) is called a pistil. A pistil may consist of a single carpel or a number of carpels fused together. The sticky tip of the pistil, the stigma, is the receptor of pollen. The supportive stalk, the style, becomes the pathway for pollen tubePollen tubeThe pollen tubes is the male gametophyte of seed plants that acts as a conduit to transport the male sperm cells from the pollen grain, either from the stigma to the ovules at the base of the pistil, or directly through ovule tissue in some gymnosperms .After pollination, the pollen tube...
s to grow from pollen grains adhering to the stigma.
Although the arrangement described above is considered "typical", plant species show a wide variation in floral structure. These modifications have significance in the evolution of flowering plants and are used extensively by botanists to establish relationships among plant species.
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 and/or look like what is typically another part. In some families, like Ranunculaceae
Ranunculaceae
Ranunculaceae are a family of about 1700 species of flowering plants in about 60 genera, distributed worldwide....
, the petals are greatly reduced and 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 Rose
Rose
A rose is a woody perennial of the genus Rosa, within the family Rosaceae. There are over 100 species. They form a group of erect shrubs, and climbing or trailing plants, with stems that are often armed with sharp prickles. Flowers are large and showy, in colours ranging from white through yellows...
s are mostly petaloid stamens. Flowers show great variation and plant scientists describe this variation in a systematic way to identify and distinguish species.
Specific terminology is used to describe flowers and their parts. Many flower parts are fused together; fused parts originating from the same whorl are connate
Connation
Connation in plants is the developmental fusion of organs of the same type, for example, petals to one another to form a tubular corolla. This is in contrast to adnation, the fusion of dissimilar organs.-Terms for connation of flower parts:...
, while fused parts originating from different whorls are adnate
Adnation
Adnation in plants is the "union of unlike parts; organically united or fused with another dissimilar part, e.g. an ovary to a calyx tube, or stamens to petals". This is in contrast to connation, the fusion of similar organs....
, parts that are not fused are free. When petals are fused into a tube or ring that falls away as a single unit, they are sympetalous (also called gamopetalous.) Connate petals may have distinctive regions: the cylindrical base is the tube, the expanding region is the throat and the flaring outer region is the limb. A sympetalous flower, with bilateral symmetry with an upper and lower lip, is bilabiate. Flowers with connate petals or sepals may have various shaped corolla or calyx including: campanulate, funnelform, tubular, urceolate, salverform or rotate.
Many flowers have a symmetry. When the perianth is bisected through the central axis from any point, symmetrical halves are produced, forming a radial symmetry. These flowers are also known to be actinomorphic or regular, e.g. rose or trillium. When flowers are bisected and produce only one line that produces symmetrical halves the flower is said to be irregular or zygomorphic, e.g. snapdragon or most orchids.
Flowers may be directly attached to the plant at their base (sessile--the supporting stalk or stem is highly reduced or absent). The stem or stalk subtending a flower is called a peduncle
Peduncle (botany)
In botany, a peduncle is a stem supporting an inflorescence, or after fecundation, an infructescence.The peduncle is a stem, usually green and without leaves, though sometimes colored or supporting small leaves...
. If a peduncle supports more than one flower, the stems connecting each flower to the main axis are called pedicels
Pedicel (botany)
A pedicel is a stem that attaches single flowers to the main stem of the inflorescence. It is the branches or stalks that hold each flower in an inflorescence that contains more than one flower....
. The apex of a flowering stem forms a terminal swelling which is called the torus or receptacle.
Floral formula
A floral formula is a way to represent the structure of a flower using specific letters, numbers, and symbols. Typically, a general formula will be used to represent the flower structure of a plant familyFamily (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
rather than a particular species. The following representations are used:
- Ca = calyx (sepal whorl; e. g. Ca5 = 5 sepals)
- Co = corolla (petal whorl; e. g., Co3(x) = petals some multiple of three )
- Z = add if zygomorphic (e. g., CoZ6 = zygomorphic with 6 petals)
- A = androecium (whorl of stamens; e. g., A∞ = many stamens)
- G = gynoecium (carpel or carpels; e. g., G1 = monocarpous)
x: to represent a "variable number"
∞: to represent "many"
A floral formula would appear something like this:
- Ca5Co5A10 - ∞G1
Several additional symbols are sometimes used (see Key to Floral Formulas).
Inflorescence
In those species that have more than one flower on an axis, the collective cluster of flowers is termed an inflorescenceInflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...
. Some inflorescences are composed of many small flowers arranged in a formation that resembles a single flower. The common example of this is most members of the very large composite (Asteraceae) group. A single daisy
Bellis perennis
Bellis perennis is a common European species of Daisy, often considered the archetypal species of that name. Many related plants also share the name "Daisy", so to distinguish this species from other daisies it is sometimes qualified as Common Daisy, Lawn Daisy or occasionally English daisy. It is...
or sunflower
Sunflower
Sunflower is an annual plant native to the Americas. It possesses a large inflorescence . The sunflower got its name from its huge, fiery blooms, whose shape and image is often used to depict the sun. The sunflower has a rough, hairy stem, broad, coarsely toothed, rough leaves and circular heads...
, for example, is not a flower but a flower head
Head (botany)
The capitulum is considered the most derived form of inflorescence. Flower heads found outside Asteraceae show lesser degrees of specialization....
—an inflorescence composed of numerous flowers (or florets).
An inflorescence may include specialized stems and modified leaves known as bract
Bract
In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis, or cone scale. Bracts are often different from foliage leaves. They may be smaller, larger, or of a different color, shape, or texture...
s.
Development
A flower is a modified stemPlant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
tip with compressed internodes, bearing structures that are highly modified leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
. In essence, a flower develops on a modified shoot or axis from a determinate apical meristem
Meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....
(determinate meaning the axis grows to a set size).
Flowering 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 seedsSEEDS
SEEDS is a voluntary organisation registered under the Societies Act of India....
, 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
Temperature
Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...
and photoperiod changes. Many perennial and most biennial plants require vernalization
Vernalization
Vernalization is the acquisition of a plant's ability to flower or germinate in the spring by exposure to the prolonged cold of winter...
to flower. The molecular interpretation of these signals is through the transmission of a complex signal known as florigen
Florigen
Florigen is the term used to describe the hypothesized hormone-like molecules responsible for controlling and/or triggering flowering in plants. Florigen is produced in the leaves and acts in the shoot apical meristem of buds and growing tips. It is known to be graft-transmissible and even...
, 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 bud
Bud
In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of the stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be specialized to develop flowers or short shoots, or may have...
s and growing tips to induce a number of different physiological and morphological changes. The first step 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. Once the process begins, even if that cue is removed the stem will continue to develop a flower.
Organ development
The molecular control of floral organ identity determination is fairly well understood. In a simple model, three gene activities interact in a combinatorial manner to determine the developmental identities of the organ primordia within the floral meristemMeristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....
. These gene functions are called A, B and C-gene functions. In the first floral whorl only A-genes are expressed, leading to the formation 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 homeotic mutants in Arabidopsis
Arabidopsis
Arabidopsis is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress , one of the model organisms used for studying plant biology and the first plant to have its entire genome sequenced...
thaliana and snapdragon, Antirrhinum majus
Antirrhinum majus
Antirrhinum majus is a species of plants belonging to the genus Antirrhinum...
. 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. See also The ABC Model of Flower Development
The ABC Model of Flower Development
The ABC model of flower development in angiosperms was formulated by Enrico Coen and Elliot Meyerowitz in 1991. This model is built on the observation of mutants with defects in floral organ development...
.
Most genes central in this model belong to the MADS-box
MADS-box
The MADS box is a conserved sequence motif found in genes which comprise the MADS-box gene family. The MADS box encodes the DNA-binding MADS domain. The MADS domain binds to DNA sequences of high similarity to the motif CC[A/T]6GG termed the CArG-box. MADS-domain proteins are generally...
genes and are transcription factors that regulate the expression of the genes specific for each floral organ.
Floral function
The principal purpose of a flower is the reproduction of the individual and the species. All flowering plants are heterosporous, producing two types of sporeSpore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
s. Microspores are produced by meiosis inside anthers while megaspores are produced inside ovules, inside an ovary. In fact, 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 (are endosporic).
In the majority of species, individual flowers have both functional carpels and stamens. These flowers are described by botanists as being perfect or bisexual. Some flowers lack one or the other reproductive organ and called imperfect or unisexual If unisex flowers are found on the same individual plant but in different locations, the species is said to be monoecious
Plant sexuality
Plant sexuality covers the wide variety of sexual reproduction systems found across the plant kingdom. This article describes morphological aspects of sexual reproduction of plants....
. If each type of unisex flower is found only on separate individuals, the plant is dioecious
Plant sexuality
Plant sexuality covers the wide variety of sexual reproduction systems found across the plant kingdom. This article describes morphological aspects of sexual reproduction of plants....
.
Flower specialization and pollination
Flowering plants usually face selective pressure to optimize the transfer of their pollenPollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...
, and this is typically reflected in the morphology of the flowers and the behaviour of the plants. Pollen may be transferred between plants via a number of 'vectors'. Some plants make use of abiotic vectors — namely wind (anemophily
Anemophily
Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Anemophilous plants may be either gymnosperms or angiosperms ....
) or, much less commonly, water (hydrophily
Hydrophily
Hydrophily is a fairly uncommon form of pollination whereby pollen is distributed by the flow of waters, particularly in rivers and streams. Hydrophilous species fall into two categories: those that distribute their pollen to the surface of water, and those that distribute it beneath the...
). Others use biotic vectors including insects (entomophily
Entomophily
Entomophily is a form of pollination whereby pollen is distributed by insects. Several insect are reported to be responsible for the pollination of many plant species, particularly bees, Lepidoptera , wasps, flies, ants and beetles. Some plant species co-evolved with a particular pollinator, such...
), birds (ornithophily
Ornithophily
Ornithophily or Bird Pollination is the pollination of flowering plants by birds. This coevolutionary association is derived from insect pollination and is particularly well developed in some parts of the world, especially in the tropics and on some island chains. The association involves several...
), bats (chiropterophily) or other animals. Some plants make use of multiple vectors, but many are highly specialised.
Cleistogamous flower
Cleistogamy
Cleistogamy or automatic self-pollination describes the trait of certain plants to propagate by using non-opening, self-pollinating flowers...
s are self pollinated, after which they may or may not open. Many Viola and some Salvia species are known to have these types of flowers.
The flowers of plants that make use of biotic pollen vectors commonly have glands called nectaries that act as an incentive for animals to visit the flower. Some flowers have patterns, called nectar guide
Nectar guide
Nectar guides are patterns seen in some species of flowers, guiding pollinators to their rewards. Rewards commonly take the form of nectar, pollen, or both, but various plants produce oil, resins, scents, or waxes...
s, that show pollinators where to look for nectar. Flowers also attract pollinators by scent and color. Still other flowers use mimicry to attract pollinators. Some species of orchids, for example, produce flowers resembling female bees in color, shape, and scent. Flowers are also specialized in shape and have an arrangement of the stamen
Stamen
The stamen is the pollen producing reproductive organ of a flower...
s 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). 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.
Anemophilous flower
Anemophily
Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Anemophilous plants may be either gymnosperms or angiosperms ....
s use the wind to move pollen from one flower to the next. Examples include grasses, birch trees, ragweed and maples. They have no need to attract pollinators and therefore tend not to be "showy" flowers. Male and female reproductive organs are generally found in separate flowers, the male flowers having a number of long filaments terminating in exposed stamens, and the female flowers having long, feather-like stigmas. Whereas the pollen of animal-pollinated flowers tends to be large-grained, sticky, and rich in protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
(another "reward" for pollinators), anemophilous flower pollen is usually small-grained, very light, and of little nutritional value to animals.
Pollination
The primary purpose of a flower is reproduction. Since the flowers are the reproductive organs of plant, they mediate the joining of the sperm, contained within pollen, to the ovules — contained in the ovary. Pollination is the movement of pollen from the anthers to the stigma. The joining of the sperm to the ovules is called fertilization. Normally pollen is moved from one plant to another, but many plants are able to self pollinate. The fertilized ovules produce seeds that are the next generation. Sexual reproduction produces genetically unique offspring, allowing for adaptationAdaptation
An adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....
. Flowers have specific designs which encourages the transfer of pollen from one plant to another of the same species. Many plants are dependent upon external factors for pollination, including: wind and animals, and especially insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s. Even large animals such as birds, bats, and pygmy possum
Pygmy possum
The pygmy possums are a family of small possums that together form the marsupial family Burramyidae. There are five extant species of pygmy possum, grouped into two genera. Four of the species are endemic to Australia, with one species also co-occurring in Papua New Guinea and Indonesia.Pygmy...
s can be employed. The period of time during which this process can take place (the flower is fully expanded and functional) is called anthesis.
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. Flowers that are insect-pollinated are called entomophilous; literally "insect-loving" in Greek. They can be highly modified along with the pollinating insects by co-evolutionCo-evolution
In biology, coevolution is "the change of a biological object triggered by the change of a related object." Coevolution can occur at many biological levels: it can be as microscopic as correlated mutations between amino acids in a protein, or as macroscopic as covarying traits between different...
. Flowers commonly have glands called nectaries on various parts that attract animals looking for nutritious nectar. Bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s and bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...
s have color vision, enabling them to seek out "colorful" flowers. Some flowers have patterns, called nectar guide
Nectar guide
Nectar guides are patterns seen in some species of flowers, guiding pollinators to their rewards. Rewards commonly take the form of nectar, pollen, or both, but various plants produce oil, resins, scents, or waxes...
s, that show pollinators where to look for nectar; they may be visible only under ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...
light, which is visible to bees and some other insects. Flowers also attract pollinators by scent
Odor
An odor or odour is caused by one or more volatilized chemical compounds, generally at a very low concentration, that humans or other animals perceive by the sense of olfaction. Odors are also commonly called scents, which can refer to both pleasant and unpleasant odors...
and some of those scents are pleasant to our sense of smell. Not all flower scents are appealing to humans; a number of flowers are pollinated by insects that are attracted to rotten flesh and have flowers that smell like dead animals, often called Carrion flower
Carrion flower
Carrion flowers or stinking flowers are flowers that emit an odor that smells like rotting flesh. Carrion flowers attract mostly scavenging flies and beetles as pollinators. Some species may trap the insects temporarily to ensure the gathering and transfer of pollen.- Amorphophallus :Many plants in...
s, including Rafflesia
Rafflesia
Rafflesia is a genus of parasitic flowering plants. It contains approximately 28 species , all found in southeastern Asia, on the Malay Peninsula, Borneo, Sumatra, Thailand and the Philippines.Rafflesia was found in the Indonesian rain forest by an Indonesian guide working for Dr...
, the titan arum
Titan arum
The titan arum or Amorphophallus titanum is a flowering plant with the largest unbranched inflorescence in the world...
, and the North American pawpaw
Asimina triloba
Asimina triloba, the pawpaw, paw paw, paw-paw, or common pawpaw, is a species of Asimina in the same plant family as the custard-apple, cherimoya, sweetsop, ylang-ylang and soursop...
(Asimina triloba). Flowers pollinated by night visitors, including bats and moths, are likely to concentrate on scent to attract pollinators and most such flowers are white.
Still other flowers use mimicry to attract pollinators. Some species of orchids, for example, produce flowers resembling female bees in color, shape, and scent. Male bees move from one such flower to another in search of a mate.
Pollination mechanism
The pollination mechanism employed by a plant depends on what method of pollination is utilized.Most flowers can be divided between two broad groups of pollination methods:
Entomophilous: flowers 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). 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. Many flowers rely on simple proximity between flower parts to ensure pollination. Others, such as the Sarracenia or lady-slipper orchid
Lady's Slipper
Lady's slipper orchids, lady slipper orchids or slipper orchids are the orchids in the subfamily Cypripedioidea, which includes the genera Cypripedium, Mexipedium, Paphiopedilum, Phragmipedium and Selenipedium...
s, have elaborate designs to ensure pollination while preventing self-pollination
Self-pollination
Self-pollination is a form of pollination that can occur when a flower has both stamen and a carpel in which the cultivar or species is self fertile and the stamens and the sticky stigma of the carpel contact each other in order to accomplish pollination...
.
Anemophilous: flowers use the wind to move pollen from one flower to the next, examples include the grasses
Poaceae
The Poaceae is a large and nearly ubiquitous family of flowering plants. Members of this family are commonly called grasses, although the term "grass" is also applied to plants that are not in the Poaceae lineage, including the rushes and sedges...
, Birch trees, Ragweed and Maples. They have no need to attract pollinators and therefore tend not to be "showy" flowers. Whereas the pollen of entomophilous flowers tends to be large-grained, sticky, and rich in protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
(another "reward" for pollinators), anemophilous flower pollen is usually small-grained, very light, and of little nutritional value to insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s, though it may still be gathered in times of dearth. Honeybees and bumblebees actively gather anemophilous corn (maize
Maize
Maize known in many English-speaking countries as corn or mielie/mealie, is a grain domesticated by indigenous peoples in Mesoamerica in prehistoric times. The leafy stalk produces ears which contain seeds called kernels. Though technically a grain, maize kernels are used in cooking as a vegetable...
) pollen, though it is of little value to them.
Some flowers are self pollinated and use flowers that never open or are self pollinated before the flowers open, these flowers are called cleistogamous. Many Viola species and some Salvia have these types of flowers.
Flower-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 often given as an example of coevolution, as the flower and pollinator are thought to have developed together over a long period of time to match each other's needs.This close relationship compounds the negative effects of extinction
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and recover may have been lost before this point...
. The extinction of either member in such a relationship would mean almost certain extinction of the other member as well. Some endangered plant species
Endangered species
An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...
are so because of shrinking pollinator populations
Pollinator decline
The term pollinator decline refers to the reduction in abundance of pollinators in many ecosystems worldwide during the end of the twentieth century....
.
Fertilization and dispersal
Some flowers with both stamens and a pistil are capable of self-fertilization, which does increase the chance of producing seeds but limits genetic variation. The extreme case of self-fertilization occurs in flowers that always self-fertilize, such as many dandelions. Conversely, many species of plants have ways of preventing 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-sterile or self-incompatible (see also: Plant sexualityPlant sexuality
Plant sexuality covers the wide variety of sexual reproduction systems found across the plant kingdom. This article describes morphological aspects of sexual reproduction of plants....
).
Evolution
While land plants have existed for about 425 million years, the first ones reproducedPlant sexuality
Plant sexuality covers the wide variety of sexual reproduction systems found across the plant kingdom. This article describes morphological aspects of sexual reproduction of plants....
by a simple adaptation of their aquatic counterparts: spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
s. In the sea, plants—and some animals—can simply scatter out genetic clones
Clones
Clones is a small town in western County Monaghan, in the 'border area' of the Republic of Ireland. The area is part of the Border Region, earmarked for economic development by the Irish Government due to its currently below-average economic situation...
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 abuse which is even more likely on land than in the sea. The protection became the seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...
, though it had not yet evolved the flower. Early seed-bearing plants include the ginkgo
Ginkgo
Ginkgo , also spelled gingko and known as the Maidenhair Tree, is a unique species of tree with no close living relatives...
and conifers. The earliest fossil of a flowering plant, Archaefructus liaoningensis
Archaefructus
Archaefructus is an extinct genus of herbaceous aquatic seed plants with 3 known species. Fossil material assigned to this genus originates from the Yixian Formation in northeastern China, originally dated as late Jurassic but now thought to be approximately 125 million years old, or early...
, is dated about 125 million years old.
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
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...
. Recently discovered angiosperm fossils such as Archaefructus, along with further discoveries of fossil gymnosperms, suggest how angiosperm characteristics may have been acquired in a series of steps.
Recent DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...
analysis (molecular systematics) shows that Amborella trichopoda
Amborella
Amborella is a genus of rare understory shrubs or small trees endemic to the island of New Caledonia. The genus consists of only a single species, Amborella trichopoda, and is the only member of the family Amborellaceae. Wood of Amborella lacks the vessels characteristic of most flowering plants...
, found on the Pacific island of New Caledonia, is 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.
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 wasp
Fig wasp
Fig wasps are wasps of the family Agaonidae which pollinate figs or are otherwise associated with figs, a coevolutional relationship that has been developing for at least 80 million years...
s 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
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...
, 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
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
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.
While there is only hard proof of such flowers existing about 130 million years ago, there is some circumstantial evidence that they did exist up to 250 million years ago. A chemical used by plants to defend their flowers, oleanane
Oleanane
Oleanane is a natural triterpene. It forms the central core for a wide variety of chemical compounds found in flowering plants which are referred to collectively as oleanane triterpenes.Some oleanane triterpenes have a suppressing effect on insect pests...
, has been detected in fossil plants that old, including gigantopterid
Gigantopterid
Gigantopterids is the name given to fossils of a group of plants existing in the Late Permian, some . Gigantopterids were among the most advanced land plants of the Paleozoic and disappeared soon after the massive Permian–Triassic extinction event...
s, 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
Petrified wood
Petrified wood is the name given to a special type of fossilized remains of terrestrial vegetation. It is the result of a tree having turned completely into stone by the process of permineralization...
.
The similarity in leaf
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
and stem
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
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. 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
Ovary (plants)
In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule and is located above or below or at the point of connection with the base of the petals and sepals...
(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".
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.
Symbolism
Many flowers have important symbolSymbol
A symbol is something which represents an idea, a physical entity or a process but is distinct from it. The purpose of a symbol is to communicate meaning. For example, a red octagon may be a symbol for "STOP". On a map, a picture of a tent might represent a campsite. Numerals are symbols for...
ic meanings in Western culture. The practice of assigning meanings to flowers is known as floriography. Some of the more common examples include:
- Red roseRoseA rose is a woody perennial of the genus Rosa, within the family Rosaceae. There are over 100 species. They form a group of erect shrubs, and climbing or trailing plants, with stems that are often armed with sharp prickles. Flowers are large and showy, in colours ranging from white through yellows...
s are given as a symbol of love, beauty, and passion. - PoppiesPoppyA poppy is one of a group of a flowering plants in the poppy family, many of which are grown in gardens for their colorful flowers. Poppies are sometimes used for symbolic reasons, such as in remembrance of soldiers who have died during wartime....
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. - IrisesIris (plant)Iris is a genus of 260-300species of flowering plants with showy flowers. It takes its name from the Greek word for a rainbow, referring to the wide variety of flower colors found among the many species...
/Lily are used in burials as a symbol referring to "resurrection/life". It is also associated with stars (sun) and its petals blooming/shining. - DaisiesAsteraceaeThe Asteraceae or Compositae , is an exceedingly large and widespread family of vascular plants. The group has more than 22,750 currently accepted species, spread across 1620 genera and 12 subfamilies...
are a symbol of innocence.
Flowers within art are also representative of the female genitalia, as seen in the works of artists such as Georgia O'Keeffe
Georgia O'Keeffe
Georgia Totto O'Keeffe was an American artist.Born near Sun Prairie, Wisconsin, O'Keeffe first came to the attention of the New York art community in 1916, several decades before women had gained access to art training in America’s colleges and universities, and before any of its women artists...
, Imogen Cunningham
Imogen Cunningham
Imogen Cunningham was an American photographer known for her photography of botanicals, nudes and industry.-Life and career:...
, Veronica Ruiz de Velasco
Veronica Ruiz de Velasco
Veronica Ruiz de Velasco is a neo-figurative painter of Mexican origin living in the United States. She was a disciple of Teodulo Romulo, Rufino Tamayo, Jean Dubuffet, and Gilberto Aceves Navarro.-Early life:...
, and Judy Chicago
Judy Chicago
Judy Chicago is a feminist artist, author, and educator.Chicago has been creating artwork since the mid 1960s. Her earliest forays into the art world coincided with the rise of Minimalism, which she eventually abandoned in favor of art she believed to have greater content and relevance...
, and in fact in Asian and western classical art. Many cultures around the world have a marked tendency to associate flowers with femininity
Female
Female is the sex of an organism, or a part of an organism, which produces non-mobile ova .- Defining characteristics :The ova are defined as the larger gametes in a heterogamous reproduction system, while the smaller, usually motile gamete, the spermatozoon, is produced by the male...
.
The great variety of delicate and beautiful flowers has inspired the works of numerous poets, especially from the 18th-19th century Romantic
Romantic poetry
Romanticism, a philosophical, literary, artistic and cultural era which began in the mid/late-1700s as a reaction against the prevailing Enlightenment ideals of the day , also influenced poetry...
era. Famous examples include William Wordsworth
William Wordsworth
William Wordsworth was a major English Romantic poet who, with Samuel Taylor Coleridge, helped to launch the Romantic Age in English literature with the 1798 joint publication Lyrical Ballads....
's I Wandered Lonely as a Cloud
I Wandered Lonely as a Cloud
"I Wandered Lonely as a Cloud" is a poem by William Wordsworth.It was inspired by an April 15, 1802 event in which Wordsworth and his sister, Dorothy, came across a "long belt" of daffodils...
and William Blake
William Blake
William Blake was an English poet, painter, and printmaker. Largely unrecognised during his lifetime, Blake is now considered a seminal figure in the history of both the poetry and visual arts of the Romantic Age...
's Ah! Sun-Flower.
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
Vincent van Gogh
Vincent Willem van Gogh , and used Brabant dialect in his writing; it is therefore likely that he himself pronounced his name with a Brabant accent: , with a voiced V and palatalized G and gh. In France, where much of his work was produced, it is...
's sunflowers
Sunflowers (series of paintings)
Sunflowers are the subject of two series of still life paintings by the Dutch painter Vincent van Gogh. The earlier series executed in Paris in 1887 gives the flowers lying on the ground, while the second set executed a year later in Arles shows bouquets of sunflowers in a vase...
series or Monet
Claude Monet
Claude Monet was a founder of French impressionist painting, and the most consistent and prolific practitioner of the movement's philosophy of expressing one's perceptions before nature, especially as applied to plein-air landscape painting. . Retrieved 6 January 2007...
's water lilies. Flowers are also dried, freeze dried and pressed in order to create permanent, three-dimensional pieces of flower art.
The Roman goddess of flowers, gardens, and the season of Spring is Flora. The Greek goddess of spring, flowers and nature is Chloris
Chloris
thumb|250px|right| "As she talks, her lips breathe spring roses:I was Chloris, who am now called Flora." [[Ovid]]There are many stories in Greek mythology about figures named Chloris...
.
In Hindu
Hindu
Hindu refers to an identity associated with the philosophical, religious and cultural systems that are indigenous to the Indian subcontinent. As used in the Constitution of India, the word "Hindu" is also attributed to all persons professing any Indian religion...
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
Lotus (plant)
Lotus identifies various plant taxa:* Nelumbo, a genus of aquatic plants with showy flowers** Nelumbo nucifera, the Sacred or Indian lotus** Nelumbo lutea, the American or Yellow lotus...
flower. Apart from the association with Vishnu, the Hindu tradition also considers the lotus to have spiritual significance. For example, it figures in the Hindu stories of creation.
Usage
In modern times, people have sought ways to cultivate, buy, wear, or otherwise be around flowers and blooming plants, partly because of their agreeable appearance and smellOdor
An odor or odour is caused by one or more volatilized chemical compounds, generally at a very low concentration, that humans or other animals perceive by the sense of olfaction. Odors are also commonly called scents, which can refer to both pleasant and unpleasant odors...
. Around the world, people use flowers for a wide range of events and functions that, cumulatively, encompass one's lifetime:
- For new births or ChristeningInfant baptismInfant baptism is the practice of baptising infants or young children. In theological discussions, the practice is sometimes referred to as paedobaptism or pedobaptism from the Greek pais meaning "child." The practice is sometimes contrasted with what is called "believer's baptism", or...
s - As a corsageCorsageCorsage refers to a bouquet of flowers worn on a woman's dress or worn around her wrist.* Corsages are often worn by the mothers and grandmothers of the bride and groom at a wedding ceremony in some countries....
or boutonniereBoutonniereA boutonnière is a floral decoration worn by men, typically a single flower or bud. The word comes from the French word for buttonhole, which is the British term....
to be worn at social functions or for holidays - As tokens of love or esteem
- For wedding flowers for the bridal party, and decorations for the hall
- As brightening decorations within the home
- As a gift of remembrance for bon voyage parties, welcome home parties, and "thinking of you" gifts
- For funeralFuneralA funeral is a ceremony for celebrating, sanctifying, or remembering the life of a person who has died. Funerary customs comprise the complex of beliefs and practices used by a culture to remember the dead, from interment itself, to various monuments, prayers, and rituals undertaken in their honor...
flowers and expressions of sympathySympathySympathy is a social affinity in which one person stands with another person, closely understanding his or her feelings. Also known as empathic concern, it is the feeling of compassion or concern for another, the wish to see them better off or happier. Although empathy and sympathy are often used...
for the grieving - For worshiping goddesses. in HinduHinduHindu refers to an identity associated with the philosophical, religious and cultural systems that are indigenous to the Indian subcontinent. As used in the Constitution of India, the word "Hindu" is also attributed to all persons professing any Indian religion...
culture it is very common to bring flowers as a gift to templeTempleA temple is a structure reserved for religious or spiritual activities, such as prayer and sacrifice, or analogous rites. A templum constituted a sacred precinct as defined by a priest, or augur. It has the same root as the word "template," a plan in preparation of the building that was marked out...
s.
People therefore grow flowers around their homes, dedicate entire parts of their living space to flower garden
Flower garden
A flower garden is any garden where flowers are grown for decorative purposes. Because flowers bloom at varying times of the year, and some plants are annual, dying each winter, the design of flower gardens can take into consideration to maintain a sequence of bloom and even of consistent color...
s, pick wildflowers, or buy flowers from florists who depend on an entire network of commercial growers and shippers to support their trade.
Flowers provide less food than other major plants parts (seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...
s, fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
s, root
Root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...
s, stem
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
s and leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
) but they provide several important foods and spice
Spice
A spice is a dried seed, fruit, root, bark, or vegetative substance used in nutritionally insignificant quantities as a food additive for flavor, color, or as a preservative that kills harmful bacteria or prevents their growth. It may be used to flavour a dish or to hide other flavours...
s. Flower vegetables include broccoli
Broccoli
Broccoli is a plant in the cabbage family, whose large flower head is used as a vegetable.-General:The word broccoli, from the Italian plural of , refers to "the flowering top of a cabbage"....
, cauliflower
Cauliflower
Cauliflower is one of several vegetables in the species Brassica oleracea, in the family Brassicaceae. It is an annual plant that reproduces by seed...
and artichoke
Artichoke
-Plants:* Globe artichoke, a partially edible perennial thistle originating in southern Europe around the Mediterranean* Jerusalem artichoke, a species of sunflower with an edible tuber...
. The most expensive spice, saffron
Saffron
Saffron is a spice derived from the flower of Crocus sativus, commonly known as the saffron crocus. Crocus is a genus in the family Iridaceae. Each saffron crocus grows to and bears up to four flowers, each with three vivid crimson stigmas, which are each the distal end of a carpel...
, consists of dried stigmas of a crocus
Crocus
Crocus is a genus in the iris family comprising about 80 species of perennials growing from corms. Many are cultivated for their flowers appearing in autumn, winter, or spring...
. Other flower spices are clove
Clove
Cloves are the aromatic dried flower buds of a tree in the family Myrtaceae. Cloves are native to the Maluku islands in Indonesia and used as a spice in cuisines all over the world...
s and caper
Caper
Capparis spinosa, the caper bush, is a perennial winter-deciduous species that bears rounded, fleshy leaves and large white to pinkish-white flowers. A caper is also the pickled bud of this plant...
s. Hops
Hops
Hops are the female flower clusters , of a hop species, Humulus lupulus. They are used primarily as a flavoring and stability agent in beer, to which they impart a bitter, tangy flavor, though hops are also used for various purposes in other beverages and herbal medicine...
flowers are used to flavor beer
Beer
Beer is the world's most widely consumed andprobably oldest alcoholic beverage; it is the third most popular drink overall, after water and tea. It is produced by the brewing and fermentation of sugars, mainly derived from malted cereal grains, most commonly malted barley and malted wheat...
. Marigold
Calendula
Calendula , pot marigold, is a genus of about 12–20 species of annual or perennial herbaceous plants in the daisy family Asteraceae, native to the area from Macaronesia east through the Mediterranean region to Iran...
flowers are fed to chicken
Chicken
The chicken is a domesticated fowl, a subspecies of the Red Junglefowl. As one of the most common and widespread domestic animals, and with a population of more than 24 billion in 2003, there are more chickens in the world than any other species of bird...
s to give their egg yolks a golden yellow color, which consumers find more desirable. Dandelion flowers are often made into wine. Bee Pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...
, pollen collected from bees, is considered a health food by some people. Honey
Honey
Honey is a sweet food made by bees using nectar from flowers. The variety produced by honey bees is the one most commonly referred to and is the type of honey collected by beekeepers and consumed by humans...
consists of bee-processed flower nectar and is often named for the type of flower, e.g. orange
Orange (fruit)
An orange—specifically, the sweet orange—is the citrus Citrus × sinensis and its fruit. It is the most commonly grown tree fruit in the world....
blossom honey, clover
Clover
Clover , or trefoil, is a genus of about 300 species of plants in the leguminous pea family Fabaceae. The genus has a cosmopolitan distribution; the highest diversity is found in the temperate Northern Hemisphere, but many species also occur in South America and Africa, including at high altitudes...
honey and tupelo
Tupelo
The tupelo , black gum, or pepperidge tree, genus Nyssa , is a small genus of about 9 to 11 species of trees with alternate, simple leaves...
honey.
Hundreds of fresh flowers are edible but few are widely marketed as food. They are often used to add color and flavor to salads. Squash
Squash (fruit)
Squashes generally refer to four species of the genus Cucurbita, also called marrows depending on variety or the nationality of the speaker...
flowers are dipped in breadcrumbs and fried. Edible flowers include nasturtium, chrysanthemum
Chrysanthemum
Chrysanthemums, often called mums or chrysanths, are of the genus constituting approximately 30 species of perennial flowering plants in the family Asteraceae which is native to Asia and northeastern Europe.-Etymology:...
, carnation
Carnation
Dianthus caryophyllus is a species of Dianthus. It is probably native to the Mediterranean region but its exact range is unknown due to extensive cultivation for the last 2,000 years. It is the wild ancestor of the garden carnation.It is a herbaceous perennial plant growing to 80 cm tall...
, cattail, honeysuckle
Honeysuckle
Honeysuckles are arching shrubs or twining vines in the family Caprifoliaceae, native to the Northern Hemisphere. There are about 180 species of honeysuckle, 100 of which occur in China; Europe, India and North America have only about 20 native species each...
, chicory
Chicory
Common chicory, Cichorium intybus, is a somewhat woody, perennial herbaceous plant usually with bright blue flowers, rarely white or pink. Various varieties are cultivated for salad leaves, chicons , or for roots , which are baked, ground, and used as a coffee substitute and additive. It is also...
, cornflower
Cornflower
Centaurea cyanus is a small annual flowering plant in the family Asteraceae, native to Europe. "Cornflower" is also erroneously used for chicory, and more correctly for a few other Centaurea species; to distinguish C...
, Canna
Canna (plant)
Canna is a genus of nineteen species of flowering plants. The closest living relations to cannas are the other plant families of the order Zingiberales, that is the gingers, bananas, marantas, heliconias, strelitzias, etc.Canna is the only genus in the family Cannaceae...
, and sunflower
Sunflower
Sunflower is an annual plant native to the Americas. It possesses a large inflorescence . The sunflower got its name from its huge, fiery blooms, whose shape and image is often used to depict the sun. The sunflower has a rough, hairy stem, broad, coarsely toothed, rough leaves and circular heads...
. Some edible flowers are sometimes candied such as daisy
Asteraceae
The Asteraceae or Compositae , is an exceedingly large and widespread family of vascular plants. The group has more than 22,750 currently accepted species, spread across 1620 genera and 12 subfamilies...
and rose
Rose
A rose is a woody perennial of the genus Rosa, within the family Rosaceae. There are over 100 species. They form a group of erect shrubs, and climbing or trailing plants, with stems that are often armed with sharp prickles. Flowers are large and showy, in colours ranging from white through yellows...
(you may also come across a candied pansy
Pansy
The Pansy is a large group of hybrid plants cultivated as garden flowers. Pansies are derived from Viola species Viola tricolor hybridized with other viola species, these hybrids are referred to as Viola × wittrockiana or less commonly Viola tricolor hortensis...
).
Flowers can also be made into herbal teas. Dried flowers such as chrysanthemum, rose, jasmine, camomile are infused into tea both for their fragrance and medical properties. Sometimes, they are also mixed with tea
Camellia sinensis
Camellia sinensis is the species of plant whose leaves and leaf buds are used to produce Chinese tea. It is of the genus Camellia , a genus of flowering plants in the family Theaceae. White tea, green tea, oolong, pu-erh tea and black tea are all harvested from this species, but are processed...
leaves for the added fragrance.
See also
Gardening- GardenGardenA garden is a planned space, usually outdoors, set aside for the display, cultivation, and enjoyment of plants and other forms of nature. The garden can incorporate both natural and man-made materials. The most common form today is known as a residential garden, but the term garden has...
- List of garden plants
- Plant evolutionary developmental biologyPlant evolutionary developmental biologyEvolutionary developmental biology refers to the study of developmental programs and patterns from an evolutionary perspective. It seeks to understand the various influences shaping the form and nature of life on the planet. Evo-devo arose as a separate branch of science rather recently. An early...
- Plant sexualityPlant sexualityPlant sexuality covers the wide variety of sexual reproduction systems found across the plant kingdom. This article describes morphological aspects of sexual reproduction of plants....
- SowingSowingSowing is the process of planting seeds.-Plants which are usually sown:Among the major field crops, oats, wheat, and rye are sowed, grasses and legumes are seeded, and maize and soybeans are planted...