A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Magnoliophyta, also called angiosperms). The biological function of a flower is to mediate the union of male sperm with female ovum in order to produce seeds. The process begins with pollination, is followed by fertilization, leading to the formation and dispersal of the seeds. For the higher plants, seeds are the next generation, and serve as the primary means by which individuals of a species are dispersed across the landscape. The grouping of flowers on a plant is called the inflorescence. In addition to serving as the reproductive organs of flowering plants, flowers have long been admired and used by humans, mainly to beautify their environment but also as a source of food. Flower specialization and pollination Further information: Pollination syndrome Flowering plants usually face selective pressure to optimise the transfer of their pollen, 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 flowers 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 guides, 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 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. Callistemon citrinus flowers. Anemophilous flowers 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. Evolution Further information: Evolutionary history of plants#Evolution of flowers Archaefructus liaoningensis, one of the earliest known flowering plants 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 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.[6] 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)[7][8] show 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[9] suggest that it has features which may have been characteristic of the earliest flowering plants. Amborella buds The general assumption is that the function of flowers, from the start, was to involve other 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 animals and spread, flowers proved to be an unusually effective means of production, spreading (whatever their actual origin) to become the dominant form of land plant life. Lomatium parryi, a plant that used to be consumed by early Native Americans 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 gigantopterids[10], 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.[11] 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.[12] Symbolism Lilies are often used to denote life or resurrection Flowers are common subjects of still life paintings, such as this one by Ambrosius Bosschaert the Elder Chinese Jade ornament with flower design, Jin Dynasty (1115-1234 AD), Shanghai Museum. Many flowers have important symbolic meanings in Western culture. 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. * Poppies are a symbol of consolation in time of death. In the UK, 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 "resurrection/life". It is also associated with stars (sun) and its petals blooming/shining. * Daisies 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, 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.[13] Apart from the association with Vishnu, the Hindu tradition also considers the lotus to have spiritual significance.[14] For example, it figures in the Hindu stories of creation.[15]