Flowering plants Temporal range: Early Cretaceous — Recent PreЄ Є O S D C P T J K Pg N
Magnolia virginiana Sweet Bay
Scientific classification
Kingdom:	Plantae
Division:	Angiospermae Lindley[1] [P.D. Cantino & M.J. Donoghue][2]
Clades
Amborellaceae Nymphaeales Austrobaileyales Mesangiospermae Ceratophyllaceae Chloranthaceae Eudicotyledoneae (eudicots) Magnoliidae Monocotyledoneae (monocots)
Synonyms
Anthophyta Magnoliophyta Cronquist, Takht. & W.Zimm., 1966

The flowering plants (angiosperms), also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants.^{[citation needed]} Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies (derived characteristics). These characteristics include flowers, endosperm within the seeds, and the production of fruits that contain the seeds.

The ancestors of flowering plants diverged from gymnosperms around 245–202 million years ago, and the first flowering plants known to exist are from 140 million years ago. They diversified enormously during the Lower Cretaceous and became widespread around 100 million years ago, but replaced conifers as the dominant trees only around 60–100 million years ago.

Angiosperm derived characteristics[link]

Bud of a pink rose

• Flowers

The flowers, which are the reproductive organs of flowering plants, are the most remarkable feature distinguishing them from other seed plants. Flowers aid angiosperms by enabling a wider range of adaptability and broadening the ecological niches open to them.^{[clarification needed]} This has allowed flowering plants to largely dominate terrestrial ecosystems.^{[citation needed]}

• Stamens with two pairs of pollen sacs

Stamens are much lighter than the corresponding organs of gymnosperms and have contributed to the diversification of angiosperms through time with adaptations to specialized pollination syndromes, such as particular pollinators. Stamens have also become modified through time^{[clarification needed]} to prevent self-fertilization, which has permitted further diversification, allowing angiosperms eventually to fill more niches.^{[clarification needed]}

• Reduced male parts, three cells

The male gametophyte in angiosperms is significantly reduced in size compared to those of gymnosperm seed plants.^{[citation needed]} The smaller pollen decreases the time^{[clarification needed]} from pollination — the pollen grain reaching the female plant — to fertilization. In gymnosperms, fertilization can occur up to a year after pollination, whereas in angiosperms, fertilization begins very soon after pollination. The shorter time leads to angiosperm plants' setting seeds sooner and faster than gymnosperms, which is a distinct evolutionary advantage.^{[clarification needed]}

• Closed carpel enclosing the ovules (carpel or carpels and accessory parts may become the fruit)

The closed carpel of angiosperms also allows adaptations to specialized pollination syndromes and controls.^{[clarification needed]} This helps to prevent self-fertilization, thereby maintaining increased diversity. Once the ovary is fertilized, the carpel and some surrounding tissues develop into a fruit. This fruit often serves as an attractant to seed-dispersing animals. The resulting cooperative relationship presents another advantage to angiosperms in the process of dispersal.

• Reduced female gametophyte, seven cells with eight nuclei

The reduced female gametophyte, like the reduced male gametophyte, may be an adaptation allowing for more rapid seed set, eventually leading to such flowering plant adaptations as annual herbaceous life-cycles, allowing the flowering plants to fill even more niches.^{[clarification needed]}

• Endosperm

In general, endosperm formation begins after fertilization and before the first division of the zygote. Endosperm is a highly nutritive tissue that can provide food for the developing embryo, the cotyledons, and sometimes the seedling when it first appears.

These distinguishing characteristics taken together have made the angiosperms the most diverse and numerous land plants and the most commercially important group to humans.^{[citation needed]} The major exception to the dominance of terrestrial ecosystems by flowering plants is the coniferous forest.

Evolution[link]

Further information: Evolutionary history of plants – Flowers

Flowers of Malus sylvestris (crab apple)

Fossilized spores suggest that higher plants (embryophytes) have lived on the land for at least 475 million years.^[3] Early land plants reproduced sexually with flagellated, swimming sperm, like the green algae from which they evolved. An adaptation to terrestrialization was the development of upright meiosporangia for dispersal by spores to new habitats. This feature is lacking in the descendants of their nearest algal relatives, the Charophycean green algae. A later terrestrial adaptation took place with retention of the delicate, avascular sexual stage, the gametophyte, within the tissues of the vascular sporophyte. This occurred by spore germination within sporangia rather than spore release, as in non-seed plants. A current example of how this might have happened can be seen in the precocious spore germination in Sellaginella, the spike-moss. The result for the ancestors of angiosperms was enclosing them in a case, the seed. The first seed bearing plants, like the ginkgo, and conifers (such as pines and firs), did not produce flowers. The pollen grains (males) of Ginkgo and cycads produce a pair of flagellated, mobile sperm cells that "swim" down the developing pollen tube to the female and her eggs.

The apparently sudden appearance of relatively modern flowers in the fossil record initially posed such a problem for the theory of evolution that it was called an "abominable mystery" by Charles Darwin.^[4] However, the fossil record has considerably grown since the time of Darwin, and 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. Several groups of extinct gymnosperms, in particular seed ferns, have been proposed as the ancestors of flowering plants, but there is no continuous fossil evidence showing exactly how flowers evolved. Some older fossils, such as the upper Triassic Sanmiguelia, have been suggested. Based on current evidence, some propose that the ancestors of the angiosperms diverged from an unknown group of gymnosperms during the late Triassic (245–202 million years ago). A close relationship between angiosperms and gnetophytes, proposed on the basis of morphological evidence, has more recently been disputed on the basis of molecular evidence that suggest gnetophytes are instead more closely related to other gymnosperms.

The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of whole genome duplication events.^[5] These occurred in 319 million years ago and 192 million years ago respectively.

The earliest known macrofossil confidently identified as an angiosperm, Archaefructus liaoningensis, is dated to about 125 million years BP (the Cretaceous period),^[6] while pollen considered to be of angiosperm origin takes the fossil record back to about 130 million years BP. However, one study has suggested that the early-middle Jurassic plant Schmeissneria, traditionally considered a type of ginkgo, may be the earliest known angiosperm, or at least a close relative.^[7] In addition, circumstantial chemical evidence has been found for the existence of angiosperms as early as 250 million years ago. Oleanane, a secondary metabolite produced by many flowering plants, has been found in Permian deposits of that age together with fossils of gigantopterids.^[8][9] Gigantopterids are a group of extinct seed plants that share many morphological traits with flowering plants, although they are not known to have been flowering plants themselves.

Recent DNA analysis based on molecular systematics ^[10][11] showed that Amborella trichopoda, found on the Pacific island of New Caledonia, belongs to a sister group of the other flowering plants, and morphological studies ^[12] suggest that it has features that may have been characteristic of the earliest flowering plants.

The orders Amborellales, Nymphaeales, and Austrobaileyales diverged as separate lineages from the remaining angiosperm clade at a very early stage in flowering plant evolution.^[13]

The great angiosperm radiation, when a great diversity of angiosperms appears in the fossil record, occurred in the mid-Cretaceous (approximately 100 million years ago). However, a study in 2007 estimated that the division of the five most recent (the genus Ceratophyllum, the family Chloranthaceae, the eudicots, the magnoliids, and the monocots) of the eight main groups occurred around 140 million years ago.^[14] By the late Cretaceous, angiosperms appear to have dominated environments formerly occupied by ferns and cycadophytes, but large canopy-forming trees replaced conifers as the dominant trees only close to the end of the Cretaceous 65 millions years ago or even later, at the beginning of the Tertiary.^[15] The radiation of herbaceous angiosperm occurred much later.^[16] Yet, many fossil plants recognizable as belonging to modern families (including beech, oak, maple, and magnolia) appeared already at late Cretaceous.

Two bees on a flower head of Creeping Thistle, Cirsium arvense

It is generally assumed that the function of flowers, from the start, was to involve mobile animals in their reproduction processes. That is, pollen can be scattered even if the flower is not brightly colored or oddly shaped in a way that attracts animals; however, by expending the energy required to create such traits, angiosperms can enlist the aid of animals and, thus, reproduce more efficiently.

Island genetics provides one proposed explanation for the sudden, fully developed appearance of flowering plants. Island genetics is believed to be a common source of speciation in general, especially when it comes to radical adaptations that seem to have required inferior transitional forms. Flowering plants may have evolved in an isolated setting like an island or island chain, where the plants bearing them were able to develop a highly specialized relationship with some specific animal (a wasp, for example). Such a relationship, with a hypothetical wasp carrying pollen from one plant to another much the way fig wasps do today, could result in the development of a high degree of specialization in both the plant(s) and their partners. Note that the wasp example is not incidental; bees, which, it is postulated, evolved specifically due to mutualistic plant relationships, are descended from wasps.

Animals are also involved in the distribution of seeds. Fruit, which is formed by the enlargement of flower parts, is frequently a seed-dispersal tool that attracts animals to eat or otherwise disturb it, incidentally scattering the seeds it contains (see frugivory). While many such mutualistic relationships remain too fragile to survive competition and to spread widely, flowering proved to be an unusually effective means of reproduction, spreading (whatever its origin) to become the dominant form of land plant life.

Flower ontogeny uses a combination of genes normally responsible for forming new shoots.^[17] 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 some of them cannot be pollinated in nature. Many modern, domesticated flowers used to be simple weeds, which sprouted only when the ground was disturbed. Some of them tended to grow with human crops, perhaps already having symbiotic companion plant relationships with them, and the prettiest did not get plucked because of their beauty, developing a dependence upon and special adaptation to human affection.^[18]

A few palaeontologists have also come up with a theory that flowering plants, or angiosperms, might have evolved because of dinosaurs; in other words, they believe that dinosaurs "created" flowers. One of the theory's biggest proponents is Robert T. Bakker. He theorizes that herbivorous dinosaurs, with their eating habits, forced plants to find new ways to develop new adaptations, in order to avoid predation by herbivores.^{[citation needed]}

Classification[link]

There are eight groups of living angiosperms:

• Amborella — a single species of shrub from New Caledonia
• Nymphaeales — about 80 species^[21] — water lilies and Hydatellaceae
• Austrobaileyales — about 100 species^[21] of woody plants from various parts of the world
• Chloranthales — several dozen species of aromatic plants with toothed leaves
• Magnoliidae — about 9,000 species,^[21] characterized by trimerous flowers, pollen with one pore, and usually branching-veined leaves — for example magnolias, bay laurel, and black pepper
• Monocotyledonae — about 70,000 species,^[21] characterized by trimerous flowers, a single cotyledon, pollen with one pore, and usually parallel-veined leaves — for example grasses, orchids, and palms
• Ceratophyllum — about 6 species^[21] of aquatic plants, perhaps most familiar as aquarium plants
• Eudicotyledonae — about 175,000 species,^[21] characterized by 4- or 5- merous flowers, pollen with three pores, and usually branching-veined leaves — for example sunflowers, petunia, buttercup, apples and oaks

The exact relationship between these eight groups is not yet clear, although there is agreement that the first three groups to diverge from the ancestral angiosperm were Amborellales, Nymphaeales, and Austrobaileyales.^[22] The term basal angiosperms refers to these three groups. The five other groups form the clade Mesangiospermae. The relationship between the three largest of these groups (magnoliids, monocots and eudicots) remains unclear. Some analyses make the magnoliids the first to diverge, others the monocots.^[20] Ceratophyllum seems to group with the eudicots rather than with the monocots.

History of classification[link]

From 1736, an illustration of Linnaean classification

The botanical term "Angiosperm", from the Ancient Greek αγγείον, angeíon (receptacle, vessel) and σπέρμα, (seed), was coined in the form Angiospermae by Paul Hermann in 1690, as the name of that one of his primary divisions of the plant kingdom. This included flowering plants possessing seeds enclosed in capsules, distinguished from his Gymnospermae, or flowering plants with achenial or schizo-carpic fruits, the whole fruit or each of its pieces being here regarded as a seed and naked. The term and its antonym were maintained by Carolus Linnaeus with the same sense, but with restricted application, in the names of the orders of his class Didynamia. Its use with any approach to its modern scope became possible only after 1827, when Robert Brown established the existence of truly naked ovules in the Cycadeae and Coniferae, and applied to them the name Gymnosperms. From that time onward, as long as these Gymnosperms were, as was usual, reckoned as dicotyledonous flowering plants, the term Angiosperm was used antithetically by botanical writers, with varying scope, as a group-name for other dicotyledonous plants.

Auxanometer: Device for measuring increase or rate of growth in plants

In 1851, Hofmeister discovered the changes occurring in the embryo-sac of flowering plants, and determined the correct relationships of these to the Cryptogamia. This fixed the position of Gymnosperms as a class distinct from Dicotyledons, and the term Angiosperm then gradually came to be accepted as the suitable designation for the whole of the flowering plants other than Gymnosperms, including the classes of Dicotyledons and Monocotyledons. This is the sense in which the term is used today.

In most taxonomies, the flowering plants are treated as a coherent group. The most popular descriptive name has been Angiospermae (Angiosperms), with Anthophyta ("flowering plants") a second choice. These names are not linked to any rank. The Wettstein system and the Engler system use the name Angiospermae, at the assigned rank of subdivision. The Reveal system treated flowering plants as subdivision Magnoliophytina (Frohne & U. Jensen ex Reveal, Phytologia 79: 70 1996), but later split it to Magnoliopsida, Liliopsida, and Rosopsida. The Takhtajan system and Cronquist system treat this group at the rank of division, leading to the name Magnoliophyta (from the family name Magnoliaceae). The Dahlgren system and Thorne system (1992) treat this group at the rank of class, leading to the name Magnoliopsida. The APG system of 1998, and the later 2003^[23] and 2009^[19] revisions, treat the flowering plants as a clade called angiosperms without a formal botanical name. However, a formal classification was published alongside the 2009 revision in which the flowering plants form the Subclass Magnoliidae.^[24]

The internal classification of this group has undergone considerable revision. The Cronquist system, proposed by Arthur Cronquist in 1968 and published in its full form in 1981, is still widely used but is no longer believed to accurately reflect phylogeny. A consensus about how the flowering plants should be arranged has recently begun to emerge through the work of the Angiosperm Phylogeny Group (APG), which published an influential reclassification of the angiosperms in 1998. Updates incorporating more recent research were published as APG II in 2003^[23] and as APG III in 2009.^[19][25]

Monocot (left) and dicot seedlings

Traditionally, the flowering plants are divided into two groups, which in the Cronquist system are called Magnoliopsida (at the rank of class, formed from the family name Magnoliacae) and Liliopsida (at the rank of class, formed from the family name Liliaceae). Other descriptive names allowed by Article 16 of the ICBN include Dicotyledones or Dicotyledoneae, and Monocotyledones or Monocotyledoneae, which have a long history of use. In English a member of either group may be called a dicotyledon (plural dicotyledons) and monocotyledon (plural monocotyledons), or abbreviated, as dicot (plural dicots) and monocot (plural monocots). These names derive from the observation that the dicots most often have two cotyledons, or embryonic leaves, within each seed. The monocots usually have only one, but the rule is not absolute either way. From a diagnostic point of view, the number of cotyledons is neither a particularly handy nor a reliable character.

Recent studies, as by the APG, show that the monocots form a monophyletic group (clade) but that the dicots do not (they are paraphyletic). Nevertheless, the majority of dicot species do form a monophyletic group, called the eudicots or tricolpates. Of the remaining dicot species, most belong to a third major clade known as the Magnoliidae, containing about 9,000 species. The rest include a paraphyletic grouping of primitive species known collectively as the basal angiosperms, plus the families Ceratophyllaceae and Chloranthaceae.

Flowering plant diversity[link]

A poster of twelve different species of flowers of the Asteraceae family

The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000.^[26][27][28] The number of families in APG (1998) was 462. In APG II^[23] (2003) it is not settled; at maximum it is 457, but within this number there are 55 optional segregates, so that the minimum number of families in this system is 402. In APG III (2009) there are 415 families.^[19]

The diversity of flowering plants is not evenly distributed. Nearly all species belong to the eudicot (75%), monocot (23%) and magnoliid (2%) clades. The remaining 5 clades contain a little over 250 species in total, i.e., less than 0.1% of flowering plant diversity, divided among 9 families.

The most diverse families of flowering plants, in their APG circumscriptions, in order of number of species, are:

In the list above (showing only the 10 largest families), the Orchidaceae and Poaceae are monocot families; the others are eudicot families.

Vascular anatomy[link]

Cross-section of a stem of the angiosperm flax:
1. Pith,
2. Protoxylem,
3. Xylem I,
4. Phloem I,
5. Sclerenchyma (bast fibre),
6. Cortex,
7. Epidermis

The amount and complexity of tissue-formation in flowering plants exceeds that of gymnosperms. The vascular bundles of the stem are arranged such that the xylem and phloem form concentric rings.

In the dicotyledons, the bundles in the very young stem are arranged in an open ring, separating a central pith from an outer cortex. In each bundle, separating the xylem and phloem, is a layer of meristem or active formative tissue known as cambium. By the formation of a layer of cambium between the bundles (interfascicular cambium), a complete ring is formed, and a regular periodical increase in thickness results from the development of xylem on the inside and phloem on the outside. The soft phloem becomes crushed, but the hard wood persists and forms the bulk of the stem and branches of the woody perennial. Owing to differences in the character of the elements produced at the beginning and end of the season, the wood is marked out in transverse section into concentric rings, one for each season of growth, called annual rings.

Among the monocotyledons, the bundles are more numerous in the young stem and are scattered through the ground tissue. They contain no cambium and once formed the stem increases in diameter only in exceptional cases.

The flower, fruit, and seed[link]

Flowers[link]

A collection of flowers forming an inflorescence

The characteristic feature of angiosperms is the flower. Flowers show remarkable variation in form and elaboration, and provide the most trustworthy external characteristics for establishing relationships among angiosperm species. The function of the flower is to ensure fertilization of the ovule and development of fruit containing seeds. The floral apparatus may arise terminally on a shoot or from the axil of a leaf (where the petiole attaches to the stem). Occasionally, as in violets, a flower arises singly in the axil of an ordinary foliage-leaf. More typically, the flower-bearing portion of the plant is sharply distinguished from the foliage-bearing or vegetative portion, and forms a more or less elaborate branch-system called an inflorescence.

There are two kinds of reproductive cells produced by flowers. Microspores, which will divide to become pollen grains, are the "male" cells and are borne in the stamens (or microsporophylls). The "female" cells called megaspores, which will divide to become the egg cell (megagametogenesis), are contained in the ovule and enclosed in the carpel (or megasporophyll).

The flower may consist only of these parts, as in willow, where each flower comprises only a few stamens or two carpels. Usually, other structures are present and serve to protect the sporophylls and to form an envelope attractive to pollinators. The individual members of these surrounding structures are known as sepals and petals (or tepals in flowers such as Magnolia where sepals and petals are not distinguishable from each other). The outer series (calyx of sepals) is usually green and leaf-like, and functions to protect the rest of the flower, especially the bud. The inner series (corolla of petals) is, in general, white or brightly colored, and is more delicate in structure. It functions to attract insect or bird pollinators. Attraction is effected by color, scent, and nectar, which may be secreted in some part of the flower. The characteristics that attract pollinators account for the popularity of flowers and flowering plants among humans.

While the majority of flowers are perfect or hermaphrodite (having both pollen and ovule producing parts in the same flower structure), flowering plants have developed numerous morphological and physiological mechanisms to reduce or prevent self-fertilization. Heteromorphic flowers have short carpels and long stamens, or vice versa, so animal pollinators cannot easily transfer pollen to the pistil (receptive part of the carpel). Homomorphic flowers may employ a biochemical (physiological) mechanism called self-incompatibility to discriminate between self- and non-self pollen grains. In other species, the male and female parts are morphologically separated, developing on different flowers.

Fertilization and embryogenesis[link]

Angiosperm life cycle

Double fertilization refers to a process in which two sperm cells fertilize cells in the ovary. This process begins when a pollen grain adheres to the stigma of the pistil (female reproductive structure), germinates, and grows a long pollen tube. While this pollen tube is growing, a haploid generative cell travels down the tube behind the tube nucleus. The generative cell divides by mitosis to produce two haploid (n) sperm cells. As the pollen tube grows, it makes its way from the stigma, down the style and into the ovary. Here the pollen tube reaches the micropyle of the ovule and digests its way into one of the synergids, releasing its contents (which include the sperm cells). The synergid that the cells were released into degenerates and one sperm makes its way to fertilize the egg cell, producing a diploid (2n) zygote. The second sperm cell fuses with both central cell nuclei, producing a triploid (3n) cell. As the zygote develops into an embryo, the triploid cell develops into the endosperm, which serves as the embryo's food supply. The ovary now will develop into fruit and the ovule will develop into seed.

Fruit and seed[link]

The fruit of the Aesculus or Horse Chestnut tree

As the development of embryo and endosperm proceeds within the embryo sac, the sac wall enlarges and combines with the nucellus (which is likewise enlarging) and the integument to form the seed coat. The ovary wall develops to form the fruit or pericarp, whose form is closely associated with the manner of distribution of the seed.

Frequently, the influence of fertilization is felt beyond the ovary, and other parts of the flower take part in the formation of the fruit, e.g., the floral receptacle in the apple, strawberry, and others.

The character of the seed coat bears a definite relation to that of the fruit. They protect the embryo and aid in dissemination; they may also directly promote germination. Among plants with indehiscent fruits, in general, the fruit provides protection for the embryo and secures dissemination. In this case, the seed coat is only slightly developed. If the fruit is dehiscent and the seed is exposed, in general, the seed-coat is well developed, and must discharge the functions otherwise executed by the fruit.

Economic importance[link]

This unreferenced section requires citations to ensure verifiability.

Agriculture is almost entirely dependent upon angiosperms, which provide virtually all plant-based food, and also provide a significant amount of livestock feed. Of all the families of plants, the Poaceae, or grass family (grains), is by far the most important, providing the bulk of all feedstocks (rice, corn — maize, wheat, barley, rye, oats, pearl millet, sugar cane, sorghum). The Fabaceae, or legume family, comes in second place. Also of high importance are the Solanaceae, or nightshade family (potatoes, tomatoes, and peppers, among others), the Cucurbitaceae, or gourd family (also including pumpkins and melons), the Brassicaceae, or mustard plant family (including rapeseed and the innumerable varieties of the cabbage species Brassica oleracea), and the Apiaceae, or parsley family. Many of our fruits come from the Rutaceae, or rue family (including oranges, lemons, grapefruits, etc.), and the Rosaceae, or rose family (including apples, pears, cherries, apricots, plums, etc.).

In some parts of the world, certain single species assume paramount importance because of their variety of uses, for example the coconut (Cocos nucifera) on Pacific atolls, and the olive (Olea europaea) in the Mediterranean region.

Flowering plants also provide economic resources in the form of wood, paper, fiber (cotton, flax, and hemp, among others), medicines (digitalis, camphor), decorative and landscaping plants, and many other uses. The main area in which they are surpassed by other plants is timber production.

See also[link]

	Plants portal
	Botany portal
	Agriculture and Agronomy portal

• List of garden plants
• List of plants by common name
• List of plant orders
• List of systems of plant taxonomy

References[link]

Further reading[link]

• Cronquist, Arthur (1981). An Integrated System of Classification of Flowering Plants. New York: Columbia Univ. Press. ISBN 0-231-03880-1. 
• Heywood, V. H., Brummitt, R. K., Culham, A. & Seberg, O. (2007). Flowering Plant Families of the World. Richmond Hill, Ontario, Canada: Firefly Books. ISBN 1-55407-206-9. 
• Dilcher, D. (2000). "Toward a new synthesis: Major evolutionary trends in the angiosperm fossil record". Proceedings of the National Academy of Sciences 97 (13): 7030. DOI:10.1073/pnas.97.13.7030. 
• Simpson, M.G. Plant Systematics, 2nd Edition. Elsevier/Academic Press. 2010.
• Raven, P.H., R.F. Evert, S.E. Eichhorn. Biology of Plants, 7th Edition. W.H. Freeman. 2004.

External links[link]

Wikimedia Commons has media related to: Magnoliophyta

Wikispecies has information related to: Magnoliophyta

The Wikibook Dichotomous Key has a page on the topic of Magnoliophyta

• Cole, Theodor C.H.; Hilger, Dr. Harmut H. Angiosperm Phylogeny Poster – Flowering Plant Systamatics
• Cromie, William J. (December 16, 1999). "Oldest Known Flowering Plants Identified By Genes". Harvard University Gazette.
• Watson, L. and Dallwitz, M.J. (1992 onwards). The families of flowering plants: descriptions, illustrations, identification, information retrieval.
• Flowering plant at the Encyclopedia of Life

v t e Botany Subdisciplines of botany Ethnobotany Paleobotany Plant anatomy Plant ecology Plant evo-devo Plant morphology Plant physiology Plants Evolutionary history of plants Algae Bryophyte Pteridophyte Gymnosperm Angiosperm Plant parts Flower Fruit Leaf Meristem Root Stem Stoma Vascular tissue Wood Plant cells Cell wall Chlorophyll Chloroplast Photosynthesis Plant hormone Plastid Transpiration Plant reproduction Alternation of generations Gametophyte Plant sexuality Pollen Pollination Seed Spore Sporophyte Plant taxonomy Botanical name Botanical nomenclature Herbarium IAPT ICN Species Plantarum Glossaries Glossary of botanical terms Glossary of plant morphology terms Category Portal

 This article incorporates text from a publication now in the public domain: Chisholm, Hugh, ed. (1911). Encyclopædia Britannica (11th ed.). Cambridge University Press. vep:Peitsemnižed

Sir David Attenborough
Attenborough in May 2003
Born	(1926-05-08) 8 May 1926 (age 86) Isleworth, London
Nationality	British
Alma mater	Clare College, Cambridge (natural sciences) London School of Economics (social anthropology)
Occupation	Broadcaster Naturalist
Title	Member of the Order of Merit Companion of Honour Commander of the Royal Victorian Order Commander of the Order of the British Empire Fellow of the Royal Society Fellow of Zoological Society of London
Spouse	Jane Elizabeth Ebsworth Oriel (m. 1950–1997, her death)
Children	Robert Attenborough Susan Attenborough

Sir David Frederick Attenborough ( /ˈætənbərə/) OM, CH, CVO, CBE, FRS, FZS, FSA (born 8 May 1926) is a British broadcaster and naturalist.

His career as the face and voice of natural history programmes has endured for more than 50 years. He is best known for writing and presenting the nine Life series, in conjunction with the BBC Natural History Unit, which collectively form a comprehensive survey of all life on the planet. He is also a former senior manager at the BBC, having served as controller of BBC Two and director of programming for BBC Television in the 1960s and 1970s.

Attenborough is widely considered a national treasure in Britain, although he himself does not care for the term.^[1][2][3] He is a younger brother of director, producer and actor Richard Attenborough.^[4]

Early life and family[link]

Attenborough was born in Isleworth, west London, but grew up in College House on the campus of the University College, Leicester, where his father, Frederick, was principal.^[5] He is the middle of three sons (his elder brother, Richard, became an actor and his younger brother, John, an executive at Italian car manufacturer Alfa Romeo).^[6] During World War II his parents also adopted two Jewish refugee girls from Europe.^[7][8]

Attenborough spent his childhood collecting fossils, stones and other natural specimens. He received encouragement in this pursuit at age seven, when a young Jacquetta Hawkes admired his "museum". A few years later, one of his adoptive sisters gave him a piece of amber filled with prehistoric creatures; some 50 years later, it would be the focus of his programme The Amber Time Machine.

Attenborough was educated at Wyggeston Grammar School for Boys in Leicester and then won a scholarship to Clare College, Cambridge in 1945, where he studied geology and zoology and obtained a degree in natural sciences.^[9] In 1947, he was called up for national service in the Royal Navy and spent two years stationed in North Wales and the Firth of Forth.

In 1950, Attenborough married Jane Elizabeth Ebsworth Oriel; the marriage lasted until her death in 1997. The couple had two children, Robert and Susan.^[8] Robert is a senior lecturer in bioanthropology for the School of Archaeology and Anthropology at the Australian National University in Canberra.^[10]

First years at the BBC[link]

After leaving the Navy, Attenborough took a position editing children's science textbooks for a publishing company. He soon became disillusioned with the work, however, and in 1950, he applied for a job as a radio talks producer with the BBC. Although he was rejected for this job, his CV later attracted the interest of Mary Adams, head of the Talks (factual broadcasting) department of the BBC's fledgling television service. Attenborough, like most Britons at that time, did not own a television, and he had seen only one programme in his life.^[11] However, he accepted Adams' offer of a three-month training course, and in 1952 he joined the BBC full-time. Initially discouraged from appearing on camera because Adams thought his teeth were too big,^[12] he became a producer for the Talks department, which handled all non-fiction broadcasts. His early projects included the quiz show Animal, Vegetable, Mineral? and Song Hunter, a series about folk music presented by Alan Lomax.

Attenborough's association with natural history programmes began when he produced and presented the three-part series The Pattern of Animals. The studio-bound programme featured animals from London Zoo, with the naturalist Julian Huxley discussing their use of camouflage, aposematism and courtship displays. Through this programme, Attenborough met Jack Lester, the curator of the zoo's reptile house, and they decided to make a series about an animal-collecting expedition. The result was Zoo Quest, first broadcast in 1954, which Attenborough presented at short notice, due to Lester being taken ill.

In 1957, the BBC Natural History Unit was formally established in Bristol. Attenborough was asked to join it, but declined, not wishing to move from London where he and his young family were settled. Instead he formed his own department, the Travel and Exploration Unit,^[13] which allowed him to continue to front Zoo Quest as well as produce other documentaries, notably the Travellers' Tales and Adventure series.

In the early 1960s, Attenborough resigned from the permanent staff of the BBC to study for a postgraduate degree in social anthropology at the London School of Economics, interweaving his study with further filming.^[14] However, he accepted an invitation to return to the BBC as controller of BBC Two before he could finish the degree.

BBC administration[link]

Attenborough became the controller of BBC Two in March 1965, but had a clause inserted in his contract that would allow him to continue making programmes on an occasional basis. Later the same year, he filmed elephants in Tanzania, and in 1969, he made a three-part series on the cultural history of the Indonesian island of Bali. For the 1971 film A Blank on the Map, he joined the first Western expedition to a remote highland valley in New Guinea to seek out a lost tribe.

BBC Two was launched in 1964, but had struggled to capture the public's imagination. When Attenborough arrived as controller, he quickly abolished the channel's quirky kangaroo mascot and shook up the schedule. With a mission to make BBC Two's output diverse and different from that offered by other networks, he began to establish a portfolio of programmes that defined the channel's identity for decades to come. Under his tenure, music, the arts, entertainment, archaeology, experimental comedy, travel, drama, sport, business, science and natural history all found a place in the weekly schedules. Often, an eclectic mix was offered within a single evening's viewing. Programmes he commissioned included Man Alive, Call My Bluff, Chronicle, Life, One Pair of Eyes, The Old Grey Whistle Test, Monty Python's Flying Circus and The Money Programme. When BBC Two became the first British channel to broadcast in colour in 1967, Attenborough took advantage by introducing televised snooker, as well as bringing rugby league to British television on a regular basis via the BBC2 Floodlit Trophy.

One of his most significant decisions was to order a 13-part series on the history of Western art, to show off the quality of the new UHF colour television service that BBC Two offered. Broadcast to universal acclaim in 1969, Civilisation set the blueprint for landmark authored documentaries, which were informally known as "tombstone" or "sledgehammer" projects. Others followed, including Jacob Bronowski's The Ascent of Man (also commissioned by Attenborough), and Alistair Cooke's America. Attenborough thought that the story of evolution would be a natural subject for such a series. He shared his idea with Chris Parsons, a producer at the Natural History Unit, who came up with the title Life on Earth and returned to Bristol to start planning the series. Attenborough harboured a strong desire to present the series himself, but this would not be possible so long as he remained in a management post.

In 1969, Attenborough was promoted to director of programmes, making him responsible for the output of both BBC channels. His tasks, which included agreeing budgets, attending board meetings and firing staff, were now far removed from the business of filming programmes. When Attenborough's name was being suggested as a candidate for the position of Director General of the BBC in 1972, he phoned his brother Richard to confess that he had no appetite for the job. Early the following year, he left his post to return to full-time programme-making, leaving him free to write and present the planned natural history epic.

Return to broadcasting[link]

After his resignation, Attenborough became a freelance broadcaster and immediately started work on his next project, a pre-arranged trip to Indonesia with a crew from the Natural History Unit. It resulted in the 1973 series Eastwards with Attenborough, which was similar in tone to the earlier Zoo Quests but without the animal-collecting element.

On his return, he began to work on the scripts for Life on Earth. Due to the scale of his ambition, the BBC decided to partner with an American network to secure the necessary funding. While the negotiations were proceeding he worked on a number of other television projects. He presented a series on tribal art (The Tribal Eye, 1975) and another on the voyages of discovery (The Explorers, 1975). He also presented a BBC children's series about cryptozoology entitled Fabulous Animals (1975), which featured mythical creatures such as the griffin and kraken.^[15] Eventually, the BBC signed a co-production deal with Turner Broadcasting and Life on Earth moved into production in 1976.

Life series[link]

Beginning with Life on Earth in 1979, Attenborough set about creating a body of work which became a benchmark of quality in wildlife film-making and influenced a generation of documentary film-makers. The series also established many of the hallmarks of the BBC's natural history output. By treating his subject seriously and researching the latest discoveries, Attenborough and his production team gained the trust of the scientific community, who responded by allowing him to feature their subjects in his programmes. In Rwanda, for example, Attenborough and his crew were granted privileged access to film Dian Fossey's research group of mountain gorillas. Innovation was another factor in Life on Earth's success: new film-making techniques were devised to get the shots Attenborough wanted, with a focus on events and animals that were hitherto unfilmed. Computerised airline schedules, which had only recently been introduced, enabled the series to be elaborately devised so that Attenborough visited several locations around the globe in each episode, sometimes even changing continents mid-sentence. Although appearing as the on-screen presenter, he consciously restricted his pieces to camera to give his subjects top billing.

The success of Life on Earth prompted the BBC to consider a follow-up, and five years later, The Living Planet was screened. This time, Attenborough built his series around the theme of ecology, the adaptations of living things to their environment. It was another critical and commercial success, generating huge international sales for the BBC. In 1990, The Trials of Life completed the original Life trilogy, looking at animal behaviour through the different stages of life. The series drew strong reactions from the viewing public for its sequences of killer whales hunting sea lions on a Patagonian beach and chimpanzees hunting and violently killing a colobus monkey.

In the 1990s, Attenborough continued to use the "Life" moniker for a succession of authored documentaries. In 1993, he presented Life in the Freezer, the first television series to survey the natural history of Antarctica. Although past normal retirement age, he then embarked on a number of more specialised surveys of the natural world, beginning with plants. They proved a difficult subject for his producers, who had to deliver five hours of television featuring what are essentially immobile objects. The result, The Private Life of Plants (1995), showed plants as dynamic organisms by using time-lapse photography to speed up their growth.

Prompted by an enthusiastic ornithologist at the BBC Natural History Unit, Attenborough then turned his attention to the animal kingdom and in particular, birds. As he was neither an obsessive twitcher, nor a bird expert, he decided he was better qualified to make The Life of Birds (1998) on the theme of behaviour. The order of the remaining "Life" series was dictated by developments in camera technology. For The Life of Mammals (2002), low-light and infrared cameras were deployed to reveal the behaviour of nocturnal mammals. The series contains a number of memorable two shots of Attenborough and his subjects, which included chimpanzees, a blue whale and a grizzly bear. Advances in macro photography made it possible to capture natural behaviour of very small creatures for the first time, and in 2005, Life in the Undergrowth introduced audiences to the world of invertebrates.

At this point, Attenborough realised that he had spent 20 years unconsciously assembling a collection of programmes on all the major groups of terrestrial animals and plants — only reptiles and amphibians were missing. When Life in Cold Blood was broadcast in 2008, he had the satisfaction of completing the set, brought together in a DVD encyclopaedia called Life on Land. In an interview that year, Attenborough was asked to sum up his achievement, and responded:

The evolutionary history is finished. The endeavour is complete. If you'd asked me 20 years ago whether we'd be attempting such a mammoth task, I'd have said "Don't be ridiculous!" These programmes tell a particular story and I'm sure others will come along and tell it much better than I did, but I do hope that if people watch it in 50 years' time, it will still have something to say about the world we live in.^[16]

However, in 2010 Attenborough asserted that his First Life — dealing with evolutionary history before Life on Earth — should also be included within the "Life" series. In the documentary Attenborough's Journey he stated, "This series, to a degree which I really didn't fully appreciate until I started working on it, really completes the set."^[17]

Other documentaries[link]

Alongside the "Life" series, Attenborough has continued to work on other television documentaries, mainly in the natural history genre. He wrote and presented a series on man's influence on the natural history of the Mediterranean basin, The First Eden, in 1987. Two years later, he demonstrated his passion for fossils in Lost Worlds Vanished Lives.

Attenborough narrated every episode of Wildlife on One, a BBC One wildlife series which ran for 253 episodes between 1977 and 2005. At its peak, it drew a weekly audience of eight to ten million, and the 1987 episode "Meerkats United" was voted the best wildlife documentary of all time by BBC viewers.^[18] He has also narrated over 50 episodes of Natural World, BBC Two's flagship wildlife series. (Its forerunner, The World About Us, was created by Attenborough in 1969, as a vehicle for colour television.^[19]) In 1997, he narrated the BBC Wildlife Specials, each focussing on a charismatic species, and screened to mark the Natural History Unit's 40th anniversary.

As a writer and narrator, he continued to collaborate with the BBC Natural History Unit in the new millennium. Alastair Fothergill, a senior producer with whom Attenborough had worked on The Trials of Life and Life in the Freezer, was making The Blue Planet (2001), the Unit's first comprehensive series on marine life. He decided not to use an on-screen presenter due to difficulties in speaking to camera through diving apparatus, but asked Attenborough to narrate the films. The same team reunited for Planet Earth (2006), the biggest nature documentary ever made for television and the first BBC wildlife series to be shot in high definition. In 2011, Fothergill gave Attenborough a more prominent role in Frozen Planet, a major series on the natural history of the polar regions. Attenborough appeared on screen and authored the final episode, in addition to performing voiceover duties.

In 2009, he co-wrote and narrated Life, a ten-part series focussing on extraordinary animal behaviour,^[20] and narrated Nature's Great Events, which showed how seasonal changes trigger major natural spectacles.^[21]

By the turn of the millennium, Attenborough's authored documentaries were adopting a more overtly environmentalist stance. In State of the Planet (2000), he used the latest scientific evidence and interviews with leading scientists and conservationists to assess the impact of man's activities on the natural world. He later turned to the issues of global warming (The Truth about Climate Change, 2006) and human population growth (How Many People Can Live on Planet Earth?, 2009). He also contributed a programme which highlighted the plight of endangered species to the BBC's Saving Planet Earth project in 2007, the 50th anniversary of the Natural History Unit.

Attenborough is also forging a new partnership with Sky, working on documentaries for the broadcaster's new 3D network, Sky 3D. Their first collaboration was Flying Monsters 3D, a film about pterosaurs which debuted on Christmas Day of 2010.^[22] A second film, The Bachelor King 3D, followed a year later, and further collaborations are planned.

Current projects[link]

Attenborough celebrates his 60th year in broadcasting in 2012, and continues to work on a number of television, film and radio projects. The anniversary will be marked by the BBC with the screening of Attenborough: 60 Years in the Wild, a three part series which looks at how wildlife filmmaking techniques, discoveries in the field of biological sciences and our relationship with the natural world have all developed over the past 60 years.^[23]

Further collaborations with Sky 3D are also in the pipeline. First will come Kingdom of Plants 3D, a three-part series filmed at the Royal Botanical Gardens, Kew which debuts on 26 May 2012 and will be simulcast on Sky Atlantic HD.^[24] A series on the wildlife of the Galapagos Islands has also been announced^[25] and the Sky 3D film The Bachelor King 3D, which Attenborough wrote and narrated, will get a theatrical release in 2012. All Attenborough's 3D projects are being co-produced by Atlantic Productions, as is a second series of First Life, which will explore the origins of the mammals.^[26]

2012 will also herald the third series of David Attenborough's Life Stories for BBC Radio 4 and the new commission David Attenborough's Natural Curiosities on Eden.^[27]

Other work[link]

From 1983, Attenborough worked on two environmentally themed musicals with the WWF and writers Peter Rose and Anne Conlon. Yanomamo was the first, about the Amazon rainforest, and the second, Ocean World, premiered at the Royal Festival Hall in 1991. They were both narrated by Attenborough on their national tour, and recorded on to audio cassette. Ocean World was also filmed for Channel 4 and later released.

In 1990, he highlighted the case of Mahjoub Sharif as part of the BBC's Prisoners of Conscience series.^[28]

In January 2009, the BBC commissioned Attenborough to provide a series of 20 ten-minute monologues covering the history of nature. Entitled David Attenborough's Life Stories, they are broadcast on Radio 4 in the Friday night slot vacated by Alistair Cooke's Letter from America.^[29] Part of Radio 4's A Point of View strand, the talks are also available as podcasts.^[30]

He appeared in the 2009 Children's Prom at the BBC Promenade Concerts and in the Last Night of the Proms on 12 September 2009, playing a floor polisher in Sir Malcolm Arnold's "A Grand, Grand Overture" (after which he was "shot" by Rory Bremner, who was playing the gun).

Attenborough also serves on the advisory board of BBC Wildlife magazine.

Achievements, awards and recognition[link]

Attenborough's contribution to broadcasting and wildlife film-making has brought him international recognition. He has been called "the great communicator, the peerless educator"^[31] and "the greatest broadcaster of our time".^[32] His programmes are often cited as an example of what public service broadcasting should be, even by critics of the BBC, and have influenced a generation of wildlife film-makers.^[33]

Honorary titles[link]

In 2010 it was reported that Attenborough has been given 29 honorary degrees by British universities, more than any other person.^[34] In 1980, he was honoured by the Open University, with whom he has had a close association throughout his career. He also has honorary Doctor of Science awards from the University of Cambridge (1984), the University of Oxford (1988)^[35], the University of Bath (1981)^[36] and the University of St Andrews (2011).^[37] In 2006, the two eldest Attenborough brothers returned to their home city to receive the title of Distinguished Honorary Fellows of the University of Leicester, "in recognition of a record of continuing distinguished service to the University."^[38] David Attenborough was previously awarded an honorary Doctor of Letters degree by the university in 1970, and was made an honorary Freeman of the City of Leicester in 1990.

Attenborough has received the title Honorary Fellow from Clare College, Cambridge (1980), the Zoological Society of London (1998), the Linnean Society (1999), the Institute of Biology (2000) and the Society of Antiquaries (2007).

Recognition[link]

Attenborough has featured as the subject of a number of BBC television programmes. Life on Air (2002) examined the legacy of his work and Attenborough the Controller (2002) focused on his time in charge of BBC Two. He also featured prominently in The Way We Went Wild (2004), a series about natural history television presenters, and 100 Years of Wildlife Films (2007), a special programme marking the centenary of the nature documentary. In 2006, British television viewers were asked to vote for their Favourite Attenborough Moments for a UKTV poll to coincide with the broadcaster's 80th birthday. The winning clip showed Attenborough observing the mimicry skills of the superb lyrebird.

Attenborough was named as the most trusted celebrity in Britain in a 2006 Reader's Digest poll,.^[39] and the following year he won The Culture Show's Living Icon Award.^[40] He has also been named among the 100 Greatest Britons in a 2002 BBC poll and is one of the top ten "Heroes of Our Time" according to New Statesman magazine.^[41]