FLOWERS

FLOWERS . The blossom, or reproductive part, of trees, shrubs, and other flora is known as the flower. This part of a growing plant takes on very special and often sacred meanings in every culture and religion of the world. The symbolism of flowers is often determined by a flower's natural properties: its color and smell, where it grows, and the length of its blooming period. While each kind of flower may be assigned a special meaning, flowers in general symbolize beauty and the transitory nature of life. Flowers are often used to represent the cycle of life and are an important part of rituals and ceremonies that celebrate birth, marriage, death, and the promise of regeneration. Flowers also serve as offerings or as a means of communicating with a deity or other sacred being. They are frequently sacred gifts bestowed as signs of welcome or in celebration of victory. Flowers represent certain deities and are associated with cultural beliefs regarding heaven or the afterlife.

It is not just the bloom of a plant that holds these special meanings. Several different plants may be combined to create a "flower" for sacred purposes. In some societies other parts of a living plant may be referred to as a "flower." In Japan, maple leaves are considered to be flowers even though they are not the bloom of the maple tree. And even sea coral was treated as a flower in nineteenth-century Christianity in the United States and was believed to represent heavenly love. Palms and evergreens are often included in the general category of flowers and are used in sacred contexts.

Flowers are used to teach general religious principles. The Japanese myth Mr. Butterfly and His Flowers teaches that all creatures are destined to become Buddhas. In this story a hermit is visited by a number of women who are actually the spirits of the flowers from his garden, which he had left behind in his search for enlightenment. These flowers, as spirits of women, had come to share his Buddhist attainment because flowers too are on the path to enlightenment.

The art of flower arrangement in Japan is called ikebana ("living flowers") and has spiritual significance. The word for flower is hana and includes blossoms, branches, foliage of trees, as well as individual flowers and grasses. The art of arranging these "flowers" expresses the Buddhist ideals of content, calm, and piety. Religious spirit, restraint, serene disposition, and respect for humankind are qualities the flower arranger must possess as he or she strives to portray the growth cycle of the plant from bud to maturity. The Daoist concepts of in (Chin., yin: female, passive, earth, moon, darkness, coolness, silence) and yo (Chin., yang: male, heaven, sun, action, power) must also be combined in the flower arrangement.

The lotus is used in many cultures to stand for the ideal of purity and perfection. The plant grows in muddy water and yet remains pure. The flower itself is most frequently believed to symbolize the oneness of Buddhist instruction and enlightenment.

Flowers and Deities

Flowers are connected to the sacred realm through their association with gods and goddesses. Flora, the Roman goddess of springtime and flowers, brings beauty and fragrance to blossoms, sweetness to honey, and aroma to wine. The Aztec god Xochipilli Cinteotl was one of thirteen day lords. He was the prince of flowers—the god of beauty, love, happiness, and youth. His female twin Xochiquetzal ("flower feather") was also the goddess of love. The Hindu love god, Kāma, is represented riding a parrot with a bow and arrow made of flowers. Ko-no-hana-sa-kura-hime ("the lady who causes trees to bloom") is a supernatural being of Japan, a fairy represented by the cherry blossom. The Japanese tennyo (like the Indian devatā s) are female deities of the sky. They play music and scatter flowers and the aroma of celestial perfume. The tennyo surround pious Buddhists like angels, they appear as decorations in Buddhist temples, and some have their own shrines. The tennyo may be identified with Shintō goddesses. In Zoroastrianism, thirty different species of flowers are associated with the thirty yazata s, or deities, that preside over the thirty days of the month.

Flowers may be created through the actions of gods. Almost every culture credits the presence of all forms of life, including flowers, to the sacred realm. The ancient Greek and Roman religions include several tales of creation. Jupiter, wishing to render Hercules immortal, placed him at the breast of the sleeping Juno. Some drops of her milk fell to the earth from which sprang the white lily.

The Muslims consider the rose a sacred plant that had sprung from the drops of perspiration that fell from the Prophet during his heavenly journey. Among the Indian cultures of Latin America the geranium is believed to have grown from drops of Christ's blood that fell as he ran from Satan. And the lily of the valley is called "Our Lady's Tears," because this plant grew from the tears the Virgin Mary shed at the cross of Christ.

Flowers are also associated with the birth or creation of deities. Ancient Egyptian religion described the fixed stars in the heavens as gods or souls, and also as fields of heavenly flowers and plants, believed to be the dwelling place of the blessed dead. It was from these fields of flowers and souls that the gods were created. Ancient Egyptians also believed that the sun was born every day from a blue lotus in the celestial ocean. In Asia the lotus is the flower on which Brahmā alighted when he sprang from the navel of Viṣṇu. From this beginning Brahmā ordered the existence of all worlds.

Flowers and deities together may protect human birth. In China the bodhisattva Guanyin is known as "the lady who brings children." Sitting on a lotus flower and holding a child in her arms, she is a goddess of fertility and aids in the treatment of all sickness. Her image is found in most homes. Kishimojin (Hariti) is a female divinity of Japan who was converted by the Buddha. She is the protector of children and women in childbirth. This goddess is portrayed standing with a baby at her breast and holding the flower of happiness.

Important symbolism is attached to flowers through their appearance with a sacred being. Many times this connection arises because of particular characteristics of the flower or the season of the year in which the bloom appears and the relation of this blossoming to the religious calendar of the culture. Artists have depicted the angel Gabriel coming to Mary with a spray of lilies in his hand to announce that she will be the mother of Christ. Many flowers are connected to Mary. These flowers all stand for virginity and purity: the annunciation lily, the flowering almond, the madonna lily, the gillyflower, the snow drop, and the rose. The thorns of the rose allude to the suffering of Mary as the mother of Christ. Christ's crown of thorns is believed to have been formed from the acanthus, bramble, or rose-briar. The hawthorn is also believed to be a symbol of Jesus, because it blooms at Christmastime. Because the Easter lily blooms at Eastertide, it is a symbol of Christ's resurrection.

Flowers and Rituals

Flowers as a link between humankind and the deities are presented as offerings to the sacred world, as food for the gods, or even as a reward from the gods. Deities may be appeased and worshiped through the singing of hymns, the anointing of images, the use of lights and incense, and through the offering of foods and flowers. In India flowers are said to have dropped from heaven to express the joy of the gods.

The religions of Latin America are a mixture of sixteenth-century Catholicism and native Indian religions. In many areas the seven most important saint's days are celebrated in festivals lasting three days. The first day of the festival is spent renewing flower decorations and offerings for house and church altars. The flowers used on this day, and throughout the year, are important offerings to the saints and to God. The last day of the festival falls on the saint's day as declared in the Roman Catholic calendar.

Among the Sherpas of Nepal, the high gods have achieved salvation and bliss and are utterly fulfilled and self-contained. Following traditional Buddhist beliefs, these gods have obtained enlightenment partly through conquering the delights of the senses. The high gods are unconcerned with humans and must be drawn down to aid humankind through a complex ritual involving sacred offerings. The god is "trapped" or seduced by the various offerings, each designed to appeal to one of the senses. The flower used in this ritual tempts the god to use his sense of smell.

Rites of passage

A rite of passage is a vehicle for transforming an individual, or a group of individuals, from one way of being to another through a series of culturally recognized stages. In most cultures, these transitions are marked or given meaning through the ritual use of flowers.

The earliest evidence for the use of flowers in a rite of passage is connected with a Neanderthal burial in the Shanidar Cave in Iraq. This burial site dates from sixty thousand years ago and reveals that the Neanderthals covered the body of the deceased with at least eight species of flowers.

The association of flowers with rituals of death occurs all over the world. The Greeks and Romans covered the dead and their graves with flowers. The souls of dying Buddhists in Japan are carried upward on a lotus, and the gravestones in cemeteries may rest on carved lotuses. Lotus leaves are also constructed out of gold or silver paper and are carried at Japanese funerals. Tahitians leave bouquets wrapped in ferns by the body after death and then pour floral perfume over the corpse to ease its passage into the sacred afterlife.

In Zorastrianism, the "rite of flowers" is performed by two priests; it invokes the blessings of the sacred and includes vows to the deceased. The priests conduct a complicated exchange of flowers accompanied by prayers and gestures. These flower exchanges symbolize the exchange of life between this world and the world after death. The ritual is also concerned with good and evil and the importance of good thoughts, words, and deeds.

Flowers are used in marriage rituals as an expression of fertility, virginity, purity, and to represent the sacred union of the bride and groom. Christian weddings in the United States include flowers for the altar, a bridal bouquet that represents the bride's fertility and the children that will result from the marriage, and flowers for virtually all the ritual participants. A Roman Catholic bride often lays her bridal bouquet at the foot of a statue of the Virgin Mary at the conclusion of the wedding ceremony as a dedication of her virginity to the mother of Christ.

The marriage ceremony of Java is a syncretism of Hinduism, Islam, and folk religion. This ceremony is completed only when the bride and groom exchange the kembang majang ("blossoming flowers") that represent their virginity. These "flowers" are large composite plants. The stems are from a banana tree trunk, the "blossom" consists of leaves, and the entire "flower" is wrapped in coconut branches.

Flowers and the afterlife or paradise

Not only are flowers used to express cultural beliefs about the changes in the life cycle, but they are also connected to ideas of life after death and paradise. Chinese Buddhists believe that at the hour of death the Buddha will appear to them, and their souls will be placed in a lotus. The souls will remain there until they are cleansed of all impurities, and then they will go to the Land of Extreme Felicity in the West, a paradise of all delights where showers of blossoms fall to the ground. The Aztec paradise was located above the ninth heaven and was called Xochitlicacan, the "place of flowers." The Huichol Indians of northwest Mexico call their paradise Wirikúta, a land of many flowers and much water. It is there that the ancient ones dwell, the ancestors and deities of the Huichol. These ancestor deities are called neyeteurixa, from yeteurixa, a thistle plant that flowers and then becomes the dry burrs found in the everyday world. Through the ingestion of peyote, or the five-petaled "flower," these Indians are able to journey to Wirikúta and to meet and join their ancestors amid the flowers of paradise.

Supernatural Powers of Flowers

Flowers are believed to possess powers that arise from their connection to the sacred realm. Greek and Roman religion held the amaranth as a sacred flower and associated it with immortality. In Switzerland it is believed that if this flower is worn on Ascension Day, it will render the wearer invisible. The peony is valued for medicinal properties and is named in celebration of Apollo, who as Paeon healed the wounds received by the gods in the Trojan War.

The Maya Indians of Zinacantan perform an illness-curing ceremony called "he enters in the flowers." This ritual includes visits to the ancestor/deity mountain shrines that surround the city of Zinacantan, as well as a ceremonial circuit of the sacred Roman Catholic churches within the town itself. This ceremonial circuit is called "great vision," referring to the number of gods visited during the ceremony, or "big flower," named for the large number of flowers necessary for the success of the procession. These flowers include not only the blooms of plants but also the sacred tips of evergreen trees. These tips are considered to be natural crosses, and two are erected alongside a permanent wooden cross to form the calvario, or calvary. This recreation of the three crosses present at Christ's crucifixion is necessary for many rituals, including curing ceremonies.

Perfume, the Essence of Flowers

Flowers may also be present at sacred times in the form of incense or perfume. The aroma of the blooms is believed to reach into the sacred sphere.

The people of Mayotte in the Comoro Islands have two religious systems that exist side by side. The public and male religion is Islam, while the religion of the private domestic sphere of women is a complex system of possession by spirits. Flowers, and especially the perfume made from them, are used to celebrate the sacred in both realms. The flowers are selected for their fragrance, not for their color or form. The cologne made from these flowers is used to mark and give meaning to several occasions: the onset of puberty in women, rituals of curing, and the anointing of the bride and groom during and after their wedding, as well as of the other major participants in the marriage ceremony. Every household keeps a supply of cologne and generously sprinkles the fragrance on body and clothing on all major holidays and Fridays at the weekly mosque service. Cologne is also offered to the mullahs at family rituals, especially during mawlid celebrations of the month of the Prophet's birth. Perfume contacts and pleases the sacred and enables prayers to travel more quickly.

The Zoroastrian Dhup-sarvi ("ceremony of the perfumes") involves the use of fragrant flowers, flower water, and other perfumes. These are passed out to the people, who are assembled to honor the dead, and they symbolize the fragrance and joyful nature of the path that righteous souls take to the afterworld.

The meanings attached to flowers, indeed the definition of flower itself, vary from culture to culture. The religious significance of these blooms is a very important part of the definition of flower for most societies. There is power in all life, especially in this form of nature that almost universally stands for beauty, purity, and the transitory nature of life.

See Also

Gardens; Incense; Lotus; Paradise; Rites of Passage.

Bibliography

One of the best examples of the nineteenth-century use of flowers by Christians is Andrew Joseph Ambauen's The Floral Apostles, or, What the Flowers Say to Thinking Man (Milwaukee, Wis., 1900). The use and meaning of flowers in religious practices, especially the historical development of their role in ritual, is well described by Jacques Duchesne-Guillemin in Symbols and Values in Zoroastrianism: Their Survival and Renewal (New York, 1966). For an excellent description of the Nyingmawa (Rñiṅ-ma-pa) sect of Tibetan Buddhist beliefs and practices, see Sherry Ortner's Sherpas through Their Rituals (New York, 1978). This book is primarily concerned with the underlying beliefs of hospitality that shape the lives of the Sherpas of Nepal and includes a consideration of flowers and their religious significance. The use of flowers as offerings to Mayan gods and Catholic saints is explored by Evon Z. Vogt in Zinacantan: A Maya Community in the Highlands of Chiapas (Cambridge, Mass., 1969). This is an extensive study of the Tzotzil-speaking Indians of Guatemala and includes a thorough description of the importance of flowers in ritual. Clifford Geertz's seminal work The Religion of Java (New York, 1960) describes the syncretism of Hindu, Islamic, and folk beliefs that constitute Javanese religion. He includes in this work an excellent consideration of flowers as part of the life-cycle rituals and in other religious celebrations. Michael Lambek presents a fascinating account of the importance of flowers and perfume among the people of Mayotte in the Comoro Islands in his Human Spirits: A Cultural Account of Trance in Mayotte (New York, 1981). Flowers and the perfume made from flower petals are essential ingredients for the two religious belief systems among the Mayotte: Islam and a native folk religion that centers around trance and possession by spirits. Barbara G. Myerhoff describes two different types of "flowers" in her consideration of Huichol Indian culture. In her book Peyote Hunt: The Sacred Journey of the Huichol Indians (Ithaca, N.Y., 1974), flowers represent the Huichol ancestor/deities in paradise and refer to the peyote, the five-petaled flower that enables the contemporary Huichol to journey to this paradise.

New Sources

Coffey, Timothy. The History and Folklore of North American Wildflowers. Boston, 1994.

Hielmeyer, Marine. The Language of Flowers: Symbols and Myths. New York, 2001.

Innes, Miranda, and Clay Perry. Medieval Flowers. London, 1997.

Laufer, Geraldine Adamich. Tussie-Mussies: The Language of Flowers. New York, 2000.

Ward, Bobby. A Contemplation upon Flowers: Garden Plants in Myth and Literature. Portland, Ore., 1999.

Wells, Diana. 100 Flowers and How They Got Their Names. New York, 1997.

Pamela R. Frese (1987)

Revised Bibliography

Flowers

An enormous diversity of size, shape, and complexity exists among the flowers of the quarter-million species of angiosperms . Flower size varies over a thousandfold, with Rafflesia (Rafflesiaceae) flowers as large as 1 meter in diameter dwarfing the minuscule flowers of Wolffia (Lemnaceae), which measure less than 1 millimeter across. The number of floral organs also varies, with the complex flowers of the Tambourissa (Monimiaceae) species having more than one thousand organs while the simple flowers of the Chloranthaceae may consist of just a few. The coevolution of angiosperms with their animal pollinators is a driving force in the generation of flower diversity. The end product of pollination is the formation of a viable seed, therefore ensuring that the species will be perpetuated. Exclusive pollinator-flower relationships ensure that pollen will not be wasted by delivery to flowers of a different species.

Definition and Flower Parts

Despite the enormous diversity in the number, size, and shape of floral organs within the angiosperms, they all are built of four basic organ types (sepals , petals, stamens, and carpels ) whose relative positions are invariant. The flower is an assemblage of sterile and fertile (reproductive) parts borne on a shoot or axis called the receptacle. The sterile parts include the sepals (collectively called the calyx) and the petals (collectively called the corolla). The sepals and petals together constitute the perianth. In a typical flower the sepals are green, and they enclose and protect the young flower before it opens. The petals, whose function is to attract pollinators, exhibit an assortment of colors, shapes, and sizes. In flowers in which the sepals and petals are indistinguishable from each other, such as tulips (Liliaceae), the perianth parts are called tepals .

The reproductive parts can be divided into the androecium and gynoecium . The androecium is composed of stamens, the male floral organs. Stamens usually have an apical anther, in which pollen develops, and a basal filament connecting the anther to the receptacle. One or more carpels constitute the gynoecium. Carpels are made of several functional tissues that facilitate pollination and protect developing ovules and seeds. The stigma on which the pollen germinates, and the style, through which the pollen tube grows toward the ovules, are examples of tissues that are intimately associated with pollination. The ovary, which houses the ovules, provides protection for both the developing ovules and seeds. In addition, the ovary often develops into a fruit that facilitates seed dispersal. The formation of a protected chamber in which the ovules and seeds develop is one of the defining features of angiosperms. The term angiosperm is derived from the Greek angio (a capsule-like covering) and sperm (seed).

Function of Flowers

The function of the flower is to facilitate the reproduction of the organism. Cells within the pollen and embryo sac are haploid and are derived from the diploid cells that develop within the anthers and ovules, respectively. In angiosperms, pollination results in double fertilization. The egg cell nucleus fuses with a sperm nucleus to produce the zygote while the other sperm cell nucleus fuses with the two polar nuclei to form the triploid endosperm . The endosperm acts as a food supply for the developing embryo. After fertilization, the ovule with the developing embryo becomes a seed and the ovary becomes the fruit that houses the seed(s).

Diversity of Flowers

Among the quarter-million species of angiosperms, there are many variations of the generalized flower yielding an immense diversity of floral patterns. The diversity is due to variations in the number, symmetry, size, and fusion of floral parts. While most flowers contain both stamens and carpels (and are referred to as hermaphroditic), other flowers are unisexual. These may be either staminate flowers (missing the carpels) or carpellate flowers (missing the stamens). Species bearing both carpellate and staminate flowers on a single plant are referred to as monoecious ("one house"; maize [Gramineae] and oak trees [Fagaceae], for example). In contrast, dioecious ("two houses") species bear staminate and carpellate flowers on different plants (willow [Salicaceae], for example). Monoecism and dioecism, both of which have evolved multiple times within the angiosperms, provide a mechanism to promote outbreeding. Many other species have evolved more subtle mechanisms to promote cross-pollination. For example, plants may be self-incompatible; that is, they discriminate between self and nonself pollen and consequently reject their own pollen. Alternatively, a difference in the timing of maturation of the androecium and gynoecium in hermaphroditic flowers favors outbreeding.

Two conspicuous characters that contribute to floral diversity are the number and size of floral organs. Angiosperms are often divided into two groups based on their cotyledon number: monocotyledons (monocots, meaning one cotyledon) and dicotyledons (dicots, meaning two cotyledons). One of the characters that distinguish monocots from dicots is the number of appendages within a whorl . Monocots usually have flowers with floral parts in multiples of three, whereas the dicots often have floral parts in multiples of four or five. Organ size can also vary enormously. For example, in the species Lepidium (Brassicaeae), the petals are microscopic, but in the genus Camellia (Theaceae), some species have petals 15 centimeters long.

Variations in fusion, arrangement, and symmetry of floral organs provide further diversity. Fusion between organs of the same whorl (coalescence) and fusion of organs from separate whorls (adnation) is common among many families of angiosperms. For instance, coalescence is seen in snapdragon (Antirrhinum, Scrophulariaceae) with the petals fused at the base, and adnation in tobacco (Nicotiana, Solanaceae) with stamens fused to the petals. Position of organ attachment to the receptacle also influences flower architecture. If the corolla and stamens attach to the receptacle below the ovary, the flower is referred to as having a superior ovary (e.g., Liriodendron tulipifera, Magnoliaceae). In contrast, having the corolla and stamens attach to the receptacle above the ovary produces a flower with an inferior ovary (e.g., Iris, Iridaceae). In radially symmetric flowers, termed actinomorphic, all organs within any particular whorl are identical and positioned equidistant from other organs within the whorl (e.g., California poppy Eschscholzia californica, Papaveraceae). Flowers in which organs in a particular whorl differ from other organs in the same whorl are referred to as zygomorphic (e.g., most orchids, Orchidaceae).

While all of the mechanisms generating diversity contribute to their interactions with pollinators, the fusion and asymmetry of floral organs has allowed the evolution of fascinating and often bizarre plant-insect interactions. For example, some species of orchids attract potential pollinators with insect pheromones , luring the insect into a maze constructed of fused petals and stamens in which there is one entrance and one exit. In navigating the maze, the insect both delivers to the stigma of the gynoecium pollen from another flower and picks up a load of pollen to be distributed to another flower.

Evolution of Flowers

While angiosperms are prevalent in the fossil record from the mid-Cretaceous (approximately one hundred million years ago), it is thought that they may have evolved substantially earlier, perhaps as far back as two hundred million years ago. The closest extant relatives of the angiosperms are the gymnosperms, of which conifers are members. Conifers do not have flowers but rather produce female and male cones consisting of scales bearing exposed ovules and pollen sacs, respectively. It's intriguing to consider what evolutionary processes occurred to produce the complex assemblage of floral organs of extant angiosperms. Charles Darwin, upon thinking about this question, stated that flowers are an "abominable mystery." The answer to this question, however, may be found in comparative genetic studies. For example, some of the important regulatory genes promoting floral organ development are also found in conifers. Understanding their function in the cones of conifers may allow scientists to model evolutionary changes that occurred resulting in the formation of early flowers.

It is not clear which features were present in the flowers of the earliest angiosperms. Although by the mid-Cretaceous period angiosperm flowers were already quite diverse, a number of key features of extant flowers had not yet appeared. Fossil flowers from the Cretaceous often have organs that are spirally arranged, a perianth that does not have a distinct calyx and corolla, relatively few stamens, and multiple carpels that are not fused together. The attractiveness of the mainly fly- and beetle-pollinated flowers was due to the androceium, which was composed of anthers attached to showy, leaflike structures. The stigma and style of the early individually fused carpels ran down along the side of the ovary instead of being at the top, as seen in most extant carpels.

There are a number of major evolutionary trends when comparing these Cretaceous flowers to their modern counterparts. For example, in many modern flowers the perianth is differentiated into a distinct calyx and corolla. The evolution of the corolla facilitated the reduction in stature of the androecium such that the stamens are composed of anthers attached to slender filaments rather than large, leaflike appendages. In addition, fusion of organs occurred along with the establishment of zygomorphic flowers, creating flowers with deep, open, funnel-shaped flowers, both innovations allowing the evolution of elaborate pollination strategies. Another evolutionary trend is the formation of a gynoecium made up of multiple fused carpels that have only one stigma and style situated at the apical end of the ovary. This has been hypothesized to provide selection at the level of the male gametophyte , which must grow through these structures to effect fertilization.

Coevolution with Pollinators

The early seed-producing plants (such as conifers) utilize wind to move pollen from the staminate cones to the female ovule-bearing cones. To ensure that enough viable seeds are generated, copious quantities of pollen need to be produced. This process requires the expenditure of large amounts of stored resources and is not very efficient. Utilizing insects to transfer pollen to other flowers enables angiosperms to produce less pollen and still maintain a high fecundity in comparison with wind-pollinated plants. In contrast to flowers of early angiosperms, extant flowers have evolved highly attractive characters to ensure that a specific pollinator continues to visit flowers from a specific species of plant. Flowers provide special sources of food for their pollinators to induce them to visit similar flowers. In addition to pollen and other edible floral parts, nectaries provide nectar, a high-energy food source for animals that can sometimes contain amino acids, proteins, lipids, antioxidants , and alkaloids.

Many of the modifications that have evolved in angiosperm flowers are adaptations to promote constancy in pollinator visitation. However, not all angiosperm flowers require animal pollinators. The grasses, which evolved from insect-pollinated flowers, are wind-pollinated. The flowers of grasses are small with reduced or absent petals, and they produce large amounts of pollen.

The Development of Flowers

A basic floral ground plan exists that defines the relationship between organ type and position in all angiosperm species. Because of the constancy in the relative positions of floral organ types, it is hypothesized that a common genetic program to specify floral organ identity is utilized during the development of all flowers. Floral organs are ultimately derived from primordia that arise from the flanks of the flower meristem . It is thought that cells within the flower meristem assess their position relative to other cells and differentiate into the appropriate floral organ based on this positional information.

To clarify how the identity of flower organ primordia is specified, researchers have taken a genetic approach. Mutations affecting flowers and their organs provide a powerful means for studying the genetic interactions involved in their development. Differences in the development of mutant versus normal (wild-type) plants reveal the function of the mutated gene. To carry out this work, researchers use two model plant systems, Arabidopsis (Brassicaceae) and Antirrhinum (Scrophulariaceae), and screen plants that have induced mutations in their genome . Studies have focused on a particular set of homeotic mutations. In homeotic mutants, normal organs develop in the positions where organs of another type are typically found. Specifically, the Arabidopsis floral homeotic mutations result in transformations of one floral organ type into another floral organ type.

It is of interest to note that several homeotic mutants exist in commonly cultivated garden plants. For example, a wild rose flower has five petals, and yet some of the hybrid tea roses have many times that number. Similar situations exist for camellias and carnations. These hybrid varieties represent changes in the genetic constitution of the plant to yield alterations in the floral architecture, which some people find attractive.

see also Angiosperms; Genetic Mechanisms and Development; Identification of Plants; Inflorescence; Interactions, Plant-Insect; Pollination Biology.

Stuart F. Baum

John L. Bowman

Bibliography

Bentley, Barbara, and Thomas Elias, eds. The Biology of Nectaries. New York: Columbia University Press, 1983.

Campbell, Neil A. Biology, 3rd ed. Redwood City, CA: The Benjamin/Cummings Publishing Company, Inc., 1993.

Crepet, W. L., and E. M. Friis. "The Evolution of Insect Pollination in Angiosperms."In The Origins of Angiosperms and Their Biological Consequences, eds. Else Marie Friis, William G. Chaloner, and Peter R. Crane. Cambridge: Cambridge University Press, 1987.

Heywood, V. H. Flowering Plants of the World. New York: Oxford University Press,1993.

Meyerowitz, Elliot M. "The Genetics of Flower Development." Scientific American 271 (1994): 56-57, 60-65.

Raven, Peter H., Ray F. Evert, and Susan E. Eichhorn. Biology of Plants. New York:Worth Publishers, Inc., 1986.

Rost, Thomas L., Michael G. Barbour, C. Ralph Stocking, and Terence M. Murphy. Plant Biology. Belmont, CA: Wadsworth Publishing Co., 1998.

FLOWERS

Almost all the very rich and variegated flora of Ereẓ Israel are flowering plants (Phanerogamae), and most of them have an attractively colored corolla. In Israel flowers bloom all year, in the cold and rainy season as well as in the burning heat of summer, but mainly during the spring. In this respect Israel differs from those countries where plants almost entirely cease blooming in winter and burst forth in spring in a blaze of flowers and greenery. The biblical "month of Abib" ("spring"; Ex. 13:4; et al.) refers to the time when the grains of corn in the ear are still tender. In point of fact, there are two seasons of the year in Israel: "Cold and heat, summer and winter" (Gen. 8:22); "Thou hast made summer and winter" (Ps. 74:17). The winter season begins after the early rains have fallen (October–November). Shortly afterward the ground is covered with a blanket of green grass. Appearing soon after the early rains, the first flowers bloom, mainly those of bulbous plants such as species of colchicum with their pinkish-white flowers and the crocus and saffron with their white ones. Masses of yellow dandelions and calendula appear a few weeks later. Once more the color of the fields changes, this time to the bright pink of the thousands of silene. In January the fields are covered with the blood-red flowers of the anemone, other colors of it – white, pink, and violet – growing in Galilee and on the Carmel. Their red is replaced shortly after by the fiery red flowers of the ranunculus, and here and there are to be seen the beautiful red blooms of the tulip, which in the 1930s and 1940s dominated the landscape of the Sharon and the mountains but have been greatly diminished as a result of ruthless picking. The red species of poppy, however, which bring the season of abundant flowering to a close, have not been affected in this way.

Such is the main cycle of the landscape's changing hues in Israel's Mediterranean areas until the arrival of summer. In addition to these there are hundreds of species of other flowers, some of them the most beautiful in the country, as well as several endemic species that are among the prettiest in the world, such as the cyclamen, conspicuous by its delicate flowers and picturesque leaves, which appears among the rocks as early as December. The fragrant narcissus is found in the valleys and prominent in March–May are many species of terrestrial orchids, not inferior in beauty despite their small flowers to their congener, the tropical epiphytic orchids. This period is rich in the blooming of floral species of the Iridaceae family, such as those of the gladiolus and iris, as well as flowers of the Liliaceae family, to which the Lilium candidum belong. Because of expanding agricultural settlement and the intensive picking of blooms, the areas of these beautiful flowers have been greatly reduced, but their number has been increasing year by year since the passing of the State of Israel's nature protection laws.

In the desert areas of Israel – the Negev and Aravah – flowering begins after the first rain and here also are many splendid flowers, particularly of the Liliaceae and Iridaceae families as well as species of the Salvia and crucifers. To the last belongs the desert Mantur which in winter covers the rocky hammada with a purple carpet of millions of flowers. The flowering period of the desert flora is short. Annuals have a brief existence, the entire life cycle of some – sprouting, growing, flowering, and seeding – lasting no more than ten weeks. In this way desert flora ensure their survival before the advent of the long, dry summer months. The Mediterranean single-season flora, too, have a brief span of life. Taking advantage of the rainy season, they grow rapidly, and flower for a few days, only to disappear suddenly with the coming of the first hot sharav ("sirocco") winds of spring. This phenomenon has found expression in many metaphors, such as those describing the end of man's life: "The grass withereth, the flower fadeth" (Isa. 40:7), and his short life: "In the morning it flourisheth, and groweth up; in the evening it is cut down, and withereth" (Ps. 90:6). The wicked, too, are likened to masses of single-season flowers that suddenly flourish in all their brilliance but with the coming of the hot sharav wind disappear without a trace (ibid. 92:8).

Besides wild annuals or bulbous and tuberous flora, there are many wild perennials – beautiful flowering shrubs and trees. Cyclical changes mirrored in the blossoms' hues on the dominant shrubs and trees can be distinguished in the color of the landscape. The first tree to bloom in Israel, when foresttrees are still shedding their leaves is the *almond, covered with a white mantle of blossoms (some strains of the cultivated almond have pinkish flowers). Shortly after, in January, with the blossoming of the Calycotome shrub, the prevailing color of the woods changes to yellow. Next the rockrose (Cistus) shrubs bloom with their large pink and white flowers. The purple flowers of the Judas tree (Cercis) are conspicuous in the woods in March. Then yellow, the color of the spartium shrubs, once again becomes the predominant hue, to be replaced by the white of the flowers of the styrax and the hawthorn (Crataegus). The great majority of the country's shrubs and trees blossom in winter and spring, and only a few of them in the dry summer – the season when the eucalyptus flowers. Soon after the rains the various species of *citrus bloom, and the air is filled with the scent of their blossoms. The end of summer, with the approach of the rainy season, sees a revival in some species of flora known as "the harbingers of winter"; prominent among these is the *squill with its white flower, like an erect candle, which comes out of the bulb during the last months of summer. The Pancriatum– "lily of Sharon" – blooms in this season, its large, fragrant flowers visible from afar in the desolate landscape. Two species of the colchicum flourish at the end of summer. lt is, at first sight, surprising that the Bible and talmudic literature seldom refer to the use of flowers for decorative purposes, but in ancient times the emphasis was laid on aromatic flora, and it is their fragrance which is emphasized. The picking of flowers is referred to in the Bible only once: "My beloved is gone down to his garden, to the beds of spices, to feed in the gardens, and to gather lilies" (Song 6:2). The Mishnah, too, speaks of the picking of lilies, but in a cemetery (Toh. 3:7). Mention is made of a "rose garden" which existed in Jerusalem since the days of the prophets (bk 82b). According to the Mishnah, figs grew there (Ma'as 2:5). But here, too, it is doubtful whether this garden was for decorative purposes or whether the fragrant roses were not used in the preparation of perfumes (see *Rose). The flowers mentioned in the Talmud, such as the saffron, jasmin, and narcissus, are chiefly mentioned as aromatic and medicinal flora.

The flower was a common motif in ancient Hebrew art: the ornamentation of the candlestick was in the form of "a calyx and petals" (Ex. 25:33). On the brim of the "sea" in the Temple were embellishments "like the brim of a cup, like the flower of a lily" (I Kings 7:26), while the boards of cedar in it were "carved with knops and open flowers" (ibid. 6:18). Josephus tells that the crown worn by the high priest was in the form of the calyx of the Hyoscyamus flower. Apparently the passage, "Woe to the crown of pride of the drunkards of Ephraim, and to the fading flower of his glorious beauty" (Isa. 28:1), alludes to floral wreaths. In apocalyptic literature it is stated that virgins wore floral chaplets, as did those celebrating Tabernacles and the victor in battle (Il Bar. 10:13; Jub. 16:30; iv Macc. 17:15). The Talmud refers several times to chaplets of roses (Shab. 152a; bm 84a). It is also related that bridegrooms wore wreaths of roses and myrtle (Sot. 9:14; Tosef. 15:8) and that non-Jews garlanded the idols on their festivals with crowns of roses and corn (Av. Zar. 4:2; tj, Av. Zar. 4:2, 43d).

Flowers of the Bible

Only three flowers are mentioned by name in the Bible, the shoshan or shoshannah ("lily" or "rose"), shoshannat haamakim (shoshannah "of the valleys"), and ḥavaẓẓelet ha-Sharon ("rose" or "lily" of the Sharon (Valley)). The complex question of the identification of the shoshan or shoshannah has provoked more studies than any other flora mentioned in the Bible, there being scarcely a beautiful flower found in lsrael (and even beyond its borders) that has not been suggested. Symbolizing in the Bible beauty and fragrance, it is most probably to be identified with the Lilium candidum– the white (madonna) lily. Abraham Ibn Ezra (in his commentary on Song 2:1) had this flower in mind when hestated that shoshan, shoshannah is derived from shesh ("six"), "since it always has six white petals as well as a pistil and long stamens which likewise number six." On the basis of this identification and doubtless also because of its delightful smell, Ibn Ezra declared, contrary to the generally accepted view, that the expression "his lips are as lilies" (ibid. 5:13) refers "to scent and not to appearance," that is, not to the red color of the lips but to their sweet odor. The other descriptions of the word in the Bible fit in with "lily" (Song 6:2–3; 4:5; 5:13; 7:3). Previously doubt was cast on this identification on the grounds that it was not proved that in ancient times the lily grew wild in Ereẓ Israel, but this large, beautiful, and scented bloom is to be found in woods in the Carmel and Galilee areas. To see the lily's lovely fragrant flowers blooming at the beginning of summer among the various thorns that then dominate the landscape is an enchanting experience, and hence "as a lily among thorns, so is my love among the daughters" (Song 2:1–2). It is this passage that was responsible for the incorrect identification of shoshannah as a rose, having been explained as referring to a rose amid its thorny stems. The rose was not found in Ereẓ Israel in biblical times, however, and although two species of rose grow wild in the country, they are neither beautiful nor fragrant, nor do the name shoshan and its biblical description fit the rose. It must, however, be pointed out that its identification as a rose already appears in the Midrash which speaks of the "red shoshannah" and of "a shoshannah of a rose" (Lev. R. 23:3; Song R. 7:3, no. 2).

Although the identification of shoshan/shoshannah as a lily is almost certain, it is difficult to identify the shoshannat ha-amakim mentioned with it in the Song of Songs since the white lily does not grow specifically in valleys. Of the many suggestions put forward in identifying it, the most likely appears to be the narcissus (Narcissus tazetta), a fragrant flower with six enveloping petals that flourishes particularly in valleys with a heavy soil. Ḥavaẓẓelet ha-Sharon is mentioned in the same verse (Song 2:1) and also in the vision of the flowering of the desolate land which shall "blossom as the ḥavaẓẓelet" and to which "the excellency of Carmel and Sharon" shall be given (Isa. 35:1–2). The ḥavaẓẓelet, as also the shoshannah, is identified by the Septuagint as κρίνον, that is, a lily. The Targum on Song of Songs identifies it with a narcissus, while various exegetes have identified it with the country's beautiful flowers, such as the iris or rose (Ibn Ezra), the colchicum (Loew), the tulip, as well as other flowers of bulbous plants, since the word is very probably connected with baẓal ("bulb"). lt is generally accepted that ḥavazzelet ha-Sharon is to be identified with the Pancratium maritimum, a bulbous plant with white, highly scented flowers which blooms at the end of summer in the coastal lowland; thus it appropriately symbolizes the flowering of the desolate land and its transformation into "the excellency of Carmel and Sharon."

[Jehuda Feliks]

Ceremonial Use

Man's awareness of the fragrance of flowers is an occasion for him to say the blessing, "Blessed art Thou, O Lord … who createst fragrant plants" (Ber. 43b). Yet flowers and plants were not generally used in synagogal or Jewish home ceremonies. On *Shavuot, however, it is customary to decorate the synagogue with fragrant grass, flowers, and branches. A threefold reason is given for this custom: the branches are a reminder that Shavuot is also the "Day of Judgment" for trees (rh 1:2); the fragrant grass is symbolic of the people of Israel assembled around Mount Sinai for the giving of the Torah (Ex. 34:3); and the flowers are a symbol for the betrothal of Israel to the Torah. The decorating of synagogues with flowers on Shavuot was opposed by some authorities on grounds of its similarity to the Christian practice (see *Ḥukkat ha-Goi). In modern times on Shavuot, synagogues are sometimes also adorned with sheaves of wheat, etc., symbolic of Shavuot as the festival of the wheat harvest and the offering of the *first fruits (bikkurim; see also Bik. 3:3). In the U.S. the custom has grown of having flowers at most family events, On Simḥat Torah in some congregations, a ḥuppah ("bridal canopy") made of plants and flowers is placed on the bimah ("platform"), and on Sukkot, the sukkah is embellished with fruits, flowers, and plants. Traditional Jewish mourning customs admit neither wreaths nor flowers at funerals or on tombstones (although in modern times, this custom is frequently disregarded). The planting of trees and shrubs around the synagogue building was the cause of heated debates a century and a half ago. Orthodox rabbinical authorities strongly objected to the landscaping of synagogue grounds, based on Deuteronomy 16:21 (see also Maim. Yad, Avodat Kokhavim 6:9). This objection, motivated by fear of innovation and reform, subsided in the course of time and yielded to the desire for an aesthetically appropriate setting for the synagogue.

bibliography:

J. Feliks, Olam ha-Ẓome'aḥ ha-Mikra'i (1968²), 232–43; Loew, Flora, 2 (1924), 144ff; Z. Avidov and I. Harpaz, Plants of Israel (1968).

FLOWERS

FLOWERS. Throughout history, flowers, like seeds, leaves, and stems, have contributed to human cookery. But edible flowers, when added to food, provide more than sustenance and flavoring. They add form and color. They are an exception to the rule, a spark of interest, and a spectacle that cannot last. Whole fresh flowers connect people to food in a way that nothing else does. Just as beauty is associated with good in children's fairy tales, so too with food: flowers add to the pleasure of eating. Flowers have long been an essential link in the human food chain. The joining of flower pollen with flower ovules is a starting point in the cycle of life. More than we imagine, life depends on a sequence that starts with flowers and progresses to pollination, seeds, and plants. Without the rebirth of plants, without continuous replacement, life would cease to exist. To sustain life, there must be birth and growth.

Flower Biology

Flowers are the blossoms of plants and the reproductive organs of angiosperms. Edible and nonedible flowers alike have a common concentric structure of distinct parts that, beginning at the base of the flower and proceeding up to the center, include the stem, ovary, sepal, petal, stamen (filament and anther), and pistil (style and stigma). Flower petals are showy, fragrant, and often fully flavored. Sepals, on the other hand, are leaflike structures that enclose the flower before it blooms and, unlike petals, are not valued as a source of food, flavor, or color. Stamens are the male parts, and they produce pollen. Anther sacs hold pollen, and in the case of pine trees, so much pollen is produced in the early spring that it accumulates in masses on lake shorelines and can be scooped up, dried, and used as an ingredient in bread and soups. Pollen is a complete food, rich in proteins, carbohydrates, vitamins, minerals, and enzymes. It can tone, detoxify, and balance the human organism.

Pistils are the female flower part. When pollen grains light on pistils, they absorb moisture, grow tubes, penetrate ovaries, and, finally, connect to ovules to fertilize seeds. Flower nectaries, or nectar glands, secrete nectar, which is used by bees to form honey. Nectar glands are located at the base of the ovary and above the anther.

History

In Roman times rose petals were used to flavor cooked brains, sweet marjoram flowers were baked in hash, and safflower petals were used for a boiled sauce. Roses and violets were added to wine to enhance flavor.

Later, in the Middle Ages, rose petals were used to flavor cakes, creams, and confectionery. Both orange blossom and rose petal water are flavorings made from flowers. Since the third and fourth centuries c.e., rose water has been made by steeping petals and then distilling the water. Middle Eastern and Indian sweets such as shola, baklava, firni, and halvah are flavored with rose water. It is also used to flavor Middle Eastern beverages such as lassi and sherbet. Flower use varies from culture to culture and age to age. While in America today roses are used more as a decoration than a flavoring, dried rosebuds are used as a condiment in Asian cookery.

Symbolism and Healing

Flowers are a symbol of life and a source of birth and healing. For example, when placed on a wedding cake flowers signify new life, and at times of sickness and death they comfort the grieving. During the Easter season the passionflower is a symbol of the holy passion, the suffering of Jesus Christ. In ancient Greece the rose symbolized love, beauty, and happiness, and during the Roman era, roses were associated with Venus, the goddess of love.

Edible flowers are used with various foods to mark events such as graduation, marriage, and retirement. Christians associate flowers with Christmas, Easter, Ascension, and Epiphany. States and nations have adopted flowers as emblems. For example, the emblem of the Netherlands is the edible tulip, and Illinois, New Jersey, Rhode Island, and Wisconsin have adopted the violet as their state flower. Four states—Georgia, Iowa, New York, and North Dakota—have adopted roses. Florida adopted the orange blossom, and Hawaii the hibiscus.

But in addition to their symbolic and spiritual uses, flowers are consumed for their healing properties. Flowers from the great scarlet poppy contain alkaloids such as thebaine, which is a source of codeine. The unripe pods of opium poppies are used to make many alkaloids including morphine, thebaine, narcotine, and codeine. The list of flowers used as medicine is extensive, and it includes arnica used as an anti-inflammatory analgesic and hawthorn used as an antispasmodic, cardiac, and vasodilator. The marsh mallow is a diuretic, antitussive, and demulcent. Passionflowers are a sedative. Rosemary is used as a tonic, diaphoretic, antiseptic, and astringent. And finally, due to their astringent qualities, some flowers, including nasturtiums, roses, and yarrow, are used as bath oils.

Preparation and Consumption

While flowers are often used fresh, they are also preserved for later use when they are stored dried, freeze-dried, candied, crystallized, or even frozen in ice cubes.

Flowers or flower parts are eaten as sweeteners, vegetables, flavorings, beverages, and garnishes. In terms of quantity, the most widely used flower today is the hop, a conelike flower or strobilus that is dried and used to flavor beer and ale and is also an antimicrobial agent. Squash blossoms are served stuffed, fried, or deep-fried. The great variety of flower foods is typified in honey, a sweetener made by bees from flower nectar. Cauliflower, broccoli, and pickled capers are flower buds. Pansy and lilac flowers are crystallized and then used for cake, cookie, and pastry decorations. Lavender, chamomile, lilac, and jasmine flowers are used to make herbal teas. Hibiscus flowers are boiled and sweetened to become agua de jamaica or jamaica water, a Mexican beverage made like tea, but served like iced tea or fruit juice. Violets, mimosa, and forget-me-nots are used to flavor confectionery.

Today, out-of-season blossoms, as well as dried flowers and flowers made of marzipan and frosting, are common on wedding and birthday cakes. Flowers are stored and shipped fresh, pressed, dried, and crystallized. Some institutions even use flowers of plastic, silk, paper, wood shavings, and wire to decorate food. This pursuit of flowers is epitomized by upscale restaurant chefs who order a box of mixed fresh flowers, and then use them indiscriminately, either whole or in parts, as the finishing touch to elegantly served dishes from medallions of venison to creamy custards with Grand Marnier.

The stigmata of the fall-blooming saffron crocus provide an essential spice for the bouillabaisse of southern France, the paella of Spain, the risotto of Italy, and the pilaf and biriani of India. Saffron, which is native to Asia Minor, adds an orange-yellow color to these dishes and gives them a spicy, pungent, and bitter flavor. Today, some of the finest saffron is produced in Spain. Saffron is costly because it must be handpicked; it takes four thousand stigmata to yield one ounce of powdered saffron.

Issues

The use of edible flowers as food also raises a number of concerns. First, culinary flowers must be free from insecticides and herbicides. In one sense, assuring toxin-free flowers is easy because fresh flower buds and flowers grow quickly. On some plants it takes only a few days for new buds and flowers to form. On the other hand, commercial flowers are often sprayed to keep them pest-free and visually attractive.

But of even greater concern than pesticides is the loss of historical species. As a result of evolution and environmental degradation, there has been a loss of species and genetic diversity. This, however, is somewhat offset by natural evolution and, in the case of flowers, the constant breeding of new and more beautiful varieties. A third problem is the use of personal or regional nomenclature that makes it difficult to trace the use of flowers in history. Cultures from Native Americans to tribal Africans have celebrated food with flowers, but these traditions are largely lost.

See also Herbs and Spices ; Presentation of Food ; Weddings .

BIBLIOGRAPHY

Kowalchik, Claire, and William H. Hylton. Rodale's Illustrated Encyclopedia of Herbs. Emmaus, Pa.: Rodale Press, 1987.

Morse, Kitty. Edible Flowers: A Kitchen Companion with Recipes. Berkeley, Calif.: Ten Speed Press, 1995.

Sohn, Mark F. "From Anise Hyssop to Zucchini: Edible Flowers From Home Gardens." Appalachian News-Express, July 23, 1997.

Sohn, Mark F. Southern Country Cooking. Iowa City: Penfield Press, 1992.

Sohn, Mark F. Mountain Country Cooking: A Gathering of the Best Recipes from the Smokies to the Blue Ridge. New York: St. Martin's, 1996.

Weaver, William Woys. Heirloom Vegetable Gardening. NewYork: Henry Holt, 1997.

Mark F. Sohn

Popular Garden Flowers and Their Flavors

ANISE HYSSOP: sweet, anise, or licorice; used to flavor red meats and as an herb

CARNATION: mild to sweet; use petals only in salads

CHIVE: onion smell, peppery, savory; use as garnish, flavoring for meats, salads, sauces

CHRYSANTHEMUM: savory, mild to strong, and bitter(some are poisonous)

DAISY: sweet and savory; flower buds are pickled, flowers used on salads, used to make wine

DANDELION: savory; a bitter herb mentioned in the Old Testament: used in salads and wine, the greens are eaten as a vegetable and sold canned

DAY LILY: savory (can be toxic)

FUCHSIA: mild flavor; use whole as garnish on salads and cooked fish

GRAPE HYACINTH: grape flavor, slightly sour, bitter aftertaste (can be toxic)

HOLLYHOCK: mild; use as garnish or container (can be toxic)

HIBISCUS: slightly acidic, sweet; used in tropical fruit salads

HONEYSUCKLE: sweet; use in salads and with fish

LILAC: strong smell; sweet and savory; crystallize the flowers to use on cakes or confectionary; used in tea blends

MARIGOLD: savory; small four-and five-petal varieties best; use in soups

NASTURTIUM: savory, peppery, piquant, like water-cress; used in salads and used fresh as a garnish for hot vegetables

PANSY: sweet and savory; crystallized to use on cakes; used fresh on salads

POPPY, EUROPEAN AND CALIFORNIA: mild; use in salads for color

RED CLOVER: mild, like hay; used with wild herb salads; add florets to salads

ROSE: sweet to bitter; herbal teas; use old varieties; also used to decorate cold platters of meat and fish

ROSE PETALS: sweet to bitter, crystallize; add to salads; garnish plates

SCENTED GERANIUM: tastes like variety, either lemon or rose used to make tea and an herb in pastry

SQUASH: savory; batter and fry into fritters; serve as vegetable; also served stuffed

SUNFLOWER: member of daisy family; use unopened flowers like artichokes or use the bitter petals; garnish for granola made with sunflower seeds; use with pasta salads

TULIP: mild, sweet to bitter; use with asparagus or rhubarb

VIOLET: sweet; use in salad, as a garnish for poke sallet, an egg dish that includes cooked poke; use crystallized on cakes

Flowers

Flowers are the typically showy reproductive organs of angiosperms (flowering plants). Their diverse blooms generate countless horticultural products; functionally flowers are essential for sexual reproduction. Flowers may occur individually on the plant or together with other flowers forming inflorescences. Their flowers and inflorescences are: (1) highly adapted to the plant's ecology and pollination strategy; (2) highly variable among the flowering plants; (3) critically important to plant identification; and (4) subject to their own rich and specialized terminology.

Morphologically and evolutionarily, the flower is regarded as a terminal shoot with variably leaflike lateral appendages . The stem of the flower, known as a pedicel, is connected to the other floral parts via the receptacle. From the outside rim of the receptacle, the first appendages are typically green, leaflike sepals , collectively called the calyx. The next whorl is composed of variously colored and often complex petals, collectively called the corolla.

The androecium forms the next whorl, formed of stamens, each made of a filament and anther. Anther sacs are filled with pollen at maturity. Anthers release the pollen timed to match the receptivity of the female of either the same flower or others of the same species.

The center of the flower is occupied by the gynoecium, which is formed of one or more carpels. Carpels may fuse to form a compound pistil or may remain separate and unfused. The carpel is divided into three regions, from the tip: the stigma, a receptive surface for pollen; the style, a specialized region for pollen tube elongation; and the ovary, a location where immature seeds called ovules occur, containing the female gametophyte or embryo sac. Meiosis occurs deep within the anther to produce many haploid male gametophytes (which become pollen) and deep within the ovule to produce the female gametophyte (or embryo sac).

The group of angiosperms known as monocotyledons typically have floral organs in multiples of three, whereas dicotyledons have floral organs in multiples of four or five. Fusion of floral organs often occurs obscuring these parts and increasing the diversity of floral form. Similar organs fuse in the example of the corolla tube (as in the morning glory), but dissimilar organs fuse in the example of apple flowers.

The largest flowers are species of Rafflesia, a carrion flower measuring up to 2 meters (6 feet) wide. This flower, native to the tropical rain forests of Sumatra, has no leaves or aboveground organs except for the flower. Living up to its name, Rafflesia attracts and is pollinated by carrion flies.

The smallest flower is a duckweed. The smallest duckweeds are members of the genus Wolffia, with the whole plant measuring less than 1 millimeter (.039 inch) across. The flower of this plant is reduced to one pistil and one anther, which are borne on different plants. Other floral parts are vestigial or fail to form.

Organs may also be modified to perform different functions. For example, roses have five petals (as do most dicots), but some or all of the numerous stamens form petal-like staminodes, sterile petal-like stamens that give horticultural roses the appearance of having many more petals. Symmetry of individual flowers may be radial (forming mirror images around the center), or bilateral (forming mirror images along only one plane).

The flower is the focus of considerable metabolic activity for the plant, producing reproductive organs with high energy content. Flowers attract insects and other animal pollinators to visit the flower. In return, pollinators are often provided with a food reward of pollen or nectar to assure this continued relationship. After the flower is fertilized, floral organs, including petals, stigma, and style, die back while seeds form inside the carpels. The tissues surrounding the ovary are dramatically modified to become the fruit.

see also Angiosperms; Fruits; Pollination and Fertilization; Seeds

Scott D. Russell

Bibliography

Uno, Gordon E., Richard Storey, and Randy Moore. Principles of Botany. Boston: Mc-Graw-Hill, 2001.

Young, Paul G., and Jacquelyn Giuffre. The Botany Coloring Book. New York: Harper Perennial, 1982.