Flowering plant
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Magnoliopsida - Dicots |
The flowering plants (also called angiosperms) are the dominant and most familiar group of land plants. The flowering plants and the gymnosperms comprise the two groups of seed plants. The flowering plants are distinguished from other seed plants by a series of apomorphies, or derived characteristics.
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[edit] Angiosperm derived characteristics
The flowers of flowering plants are the most remarkable feature distinguishiing them from other seed plants. Flowers aided angiosperms by enabling a wider range of evolutionary relationship and broadening the ecological niches open to them, allowing flowering plants to eventually dominate terrestrial ecosystems.
- Stamens with 2 pairs of pollen sac
Stamens are much lighter than the corresponding microsporophylls 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 been modified through time to prevent self-fertilization, again to increase diversity, allowing angiosperms to eventually fill more niches.
- Reduced male gametophyte, three cells
The reduced male gametophyte in angiosperms may have evolved to decrease the amount of time from pollination, the pollen grain reaching the female plant, to the fertilization of the ovary. In gymnosperms fertilization can occur up to a year after pollination, while in flowering plants the fertilization process begins very soon after pollination, allowing angiosperms, ultimately, to set seeds sooner and faster than gymnosperms.
The closed carpel of angiosperms also allows adaptations to specialized pollination syndromes and controls to prevent self-fertilization, thereby maintaining increased diversity. Once the ovary is fertilized the carpel and some surrounding tissues develop into a fruit, another opportunity for angiosperms to increase their domination of the terrestrial ecosystem with evolutionary adaptations to dispersal mechanisms.
- Reduced female gametophyte, seven cells with eight nuclei
The reduced female gametophyte, like the reduce male gametophyte may be evolutionary adaptations 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 towards their terrestrial domination.
Endosperm formation generally 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 for 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. The major exception to the dominance of terrestrial ecosystems by flowering plants is the coniferous forest.
[edit] Origins
The first undisputed evidence of angiosperms appears in the fossil record in the early Cretaceous period with a rapid diversification in the mid-Cretaceous. Some older fossils, such as the upper Triassic Sanmiguelia, have been suggested to represent early angiosperms. Based on current evidence, it seems 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, suggested on the basis of morphological evidence, has been disputed on the basis of molecular evidence that suggest Gnetophytes are more closely related to other gymnosperms. Fossil plants with some identifiable angiosperm characteristics appear in the Jurassic and early Cretaceous (135-65 million years ago), but in relatively few and primitive forms. The great angiosperm radiation, when a great diversity of angiosperms appear in the fossil record, occurred in the mid-Cretaceous (approximately 100 million years ago). By the late Cretaceous, angiosperms appear to have become the predominant group of land plants, and many fossil plants recognizable as belonging to modern families (including beech, oak, maple, and magnolia) appeared.
[edit] Classification
The botanical term "Angiosperm", from the ancient Greek αγγειον (receptacle) and σπερμα (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, which included flowering plants possessing seeds enclosed in capsules, in contradistinction to 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 only became possible after Robert Brown had established in 1827 the existence of truly naked ovules in the Cycadeae and Coniferae, entitling them to be correctly called Gymnosperms. From that time onwards, so long as these Gymnosperms were, as was usual, reckoned as dicotyledonous flowering plants, the term Angiosperm was used antithetically by botanical writers, but with varying limitation, as a group-name for other dicotyledonous plants.
The advent in 1851 of Hofmeister's discovery of the changes proceeding in the embryo-sac of flowering plants, and his determination of the correct relationships of these with the Cryptogamia, 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, and as including therefore the classes of Dicotyledons and Monocotyledons. This is the sense in which the term is nowadays received and in which it is used here.
In most taxonomic treatments the flowering plants are treated as a coherent group. Usually this takes the form of a taxonomic grouping, or taxon, which will be assigned a rank. For taxa at a rank above the rank of family Art 16 of the ICBN allows either a descriptive name or a name formed from the name of an included family (that in turn is based on a generic name). 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.
A name formed from an included family depends on the rank chosen, with different endings for different ranks. The Reveal system treated it 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. However, the APG system, of 1998, and the APG II system, of 2003, do not treat it as a formal taxon but rather treat it as a clade without a formal botanical name and use the name angiosperms for this clade.
[edit] Internal classification
The internal classification of this group has undergone considerable revision as ideas change about the relationships of the plants that form this group. 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 reflect phylogeny. A general consensus about how the flowering plants should be arranged has recently begun to emerge, through the work of the Angiosperm Phylogeny Group, who published an influential reclassification of the angiosperms in 1998. An update incorporating more recent research was published as APG II in 2003.
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 Art 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 fact that the dicots usually have two cotyledons (embryonic leaves) within each seed, while the monocots usually have only one. From a diagnostic point of view the number of cotyledons is neither a particularly handy nor reliable character.
Recent studies, as by the APG group, show that the monocots are a "good" group (a holophyletic or monophyletic group); this clade is given the name monocots. However, the dicots are not (they are a paraphyletic group). Nevertheless, within the dicots a "good" group does exist, which includes most of the dicots. This clade is called the eudicots or "tricolpates". The name "tricolpates" derives from the type of pollen found throughout this group. The name eudicots is formed by preceding "dicot" by the botanical prefix "eu-" (from the Greek 'eu'= "true"), as the eudicots share the characters traditionally attributed to the dicots, such as flowers with four or five parts (four or five petals, four or five sepals). Separating this group of eudicots from the rest of the (former) dicots leaves a remainder, which sometimes are called informally "palaeodicots" (the prefix "palaeo-" means old, and derives from the classic Greek). As this remainder group is not a "good" group this is a term of convenience only.
[edit] Flowering plant diversity
The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000. The number of families in APG (1998) was 462. In APG II (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.
The most diverse families of flowering plants, in order of number of species, are:
- Orchidaceae (orchid family): 25,000 or more species
- Asteraceae or Compositae (daisy family): 20,000 species
- Fabaceae or Leguminosae (pea family): 17,000
- Rubiaceae (madder family): 13,183
- Poaceae or Gramineae (grass family): 9,000
- Euphorbiaceae (spurge family): 5,000
- Malvaceae (mallow family): 4,300
- Cyperaceae (sedge family): 4,000
- Araceae (aroid family): 3700
In the list above (showing only the 9 largest families), the Orchidaceae, Poaceae, Cyperaceae and Araceae are monocot families; the others are dicot families.
[edit] Plant anatomy
The amount and complexity of tissue-formation in flowering plants far exceeds that found in Gymnosperms. The vascular bundles of the stem are arranged such that the xylem and phloem stand side by side on the same radius. 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 it by the development of xylem on the inside and phloem on the outside. The soft phloem soon becomes crushed, but the hard wood persists, and forms the great 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. In the smaller group, the Monocotyledons, the bundles are more numerous in the young stem and scattered through the ground tissue. Moreover they contain no cambium and the stem once formed increases in diameter only in exceptional cases.
[edit] The flower, fruit, and seed
[edit] Flowers
The characteristic feature of angiosperms is the flower, which shows remarkable variation in form and elaboration, and provides the most trustworthy external characteristics for establishing relationships among angiosperm species. The function of the flower is that of ensuring 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. Occasionally, as in violet, a flower arises singly in the axil of an ordinary foliage-leaf. However, 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.
The reproductive cells produced by flowers are of two kinds, microspores which will divide to become pollen grains, are the "male" cells and are borne in the stamens (or microsporophylls), and the "female" cells called megaspores, which will divide to become the egg-cell (in a process called megagametogenesis), 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, however, other structures are present and serve both to protect the sporophylls and to form an envelope attractive to pollinating insects. The individual members of these surrounding structures are called 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 in the bud. The inner series (corolla of petals) is generally white or brightly coloured, and more delicate in structure, and functions in attracting a particular insect or bird by agency of which pollination is effected. This attraction involves colour and scent, and frequently also nectar which is secreted in some part of the flower. These 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 male and female 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.
[edit] Fertilization and embryogenesis
Double fertilization refers to a process in flowering plants during reproduction, in which two sperm cells fertilize two cells in the ovary. The pollen grain adheres to the stigma of the carpel (female reproductive structure) and grows a pollen tube that penetrates the ovum through a tiny pore called a micropyle. Two sperm cells are released into the ovary through this tube. One of the two sperm cells fertilizes the egg cell, forming a diploid zygote or embryo, also called the ovule. The other sperm cell fuses with two haploid polar nuclei in the center of the embryo sac. The resulting cell is triploid (3n). This triploid cell divides through mitosis and forms the endosperm, a nutrient-rich tissue inside the fruit. When seed develops without fertilization, the process is known as apomixis.
[edit] Fruit and seed
As the development of embryo and endosperm proceeds within the embryo-sac, its wall enlarges and commonly absorbs the substance of the nucellus (which is likewise enlarging) to near its outer limit, and combines with it and the integument to form the seed-coat; or the whole nucellus and even the integument may be absorbed. The ovary wall has developed to form the fruit or pericarp, the structure of which 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, as the floral receptacle in the apple, strawberry and others. The character of the seed-coat bears a definite relation to that of the fruit. Their function is the twofold one of protecting the embryo and of aiding in dissemination; they may also directly promote germination. If the fruit is a dehiscent one and the seed is therefore soon exposed, the seed-coat has to provide for the protection of the embryo and may also have to secure dissemination. On the other hand, indehiscent fruits discharge these functions for the embryo, and the seed-coat is only slightly developed.
[edit] Economic importance
Agriculture is almost entirely dependent on angiosperms, either directly or indirectly through livestock feed. Of all the families of flowering plants, the Poaceae, or grass family, is by far the most important, providing the bulk of all feedstocks (rice, corn (maize), wheat, barley, rye, oats, millet, sugar cane, sorghum), with the Fabaceae, or legume family, 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 cabbage), and the Apiaceae, or parsley family. Many of our fruits come from the Rutaceae, or rue family, and the Rosaceae (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. An example is the coconut (Cocos nucifera) on Pacific atolls. Another example is the olive (Olea europaea) in the Mediterranean.
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, many other uses.
[edit] See also
[edit] References and external links
- Angiosperm Phylogeny Group (2003). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141: 399-436. Available online.
- Angiosperms – Tree of Life Web Project
- Cronquist, Arthur. (1981) An Integrated System of Classification of Flowering Plants. Columbia Univ. Press, New York.
- Dilcher, D. 2000. Toward a new synthesis: Major evolutionary trends in the angiosperm fossil record. PNAS [Proceedings of the National Academy of Sciences of the United States of America] 97: 7030-7036 (available online here)
- Oldest Known Flowering Plants Identified By Genes, William J. Cromie, Harvard Gazette, December 16, 1999.
- Stevens, P.F. (2001 onwards). Angiosperm Phylogeny Website at Missouri Botanical Garden.
- L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants: descriptions, illustrations, identification, information retrieval. http://delta-intkey.com
- Simpson, M.G. Plant Systematics. Elsevier Academic Press. 2006.
- Raven, P.H., R.F. Evert, S.E. Eichhorn. Biology of Plants, 7th Edition. W.H. Freeman. 2004.
This article incorporates text from the Encyclopædia Britannica Eleventh Edition, a publication now in the public domain.
