Magnoliophyta
The Angiospermes are the plants with flowers , and thus the plants which carry Fruit S. Angiosperme means “Graine in container” in Greek in opposition to the Gymnosperme S (naked seed) which preceded them in the evolution. One counts some currently more than 200.000 species divided into 12.000 kinds belonging to more than 300 families (against only 700 species of Gymnospermes according to the broadest estimates). One counts more than 165.000 Dicotylédones and more than 35.000 Monocotylédones.
From a Systematic point of view, Angiospermes form a Taxon regarded as Monophylétique. More are the close relatives of Angiospermes the Gnétophyte S (brother group?), the Cycadophyte S and the Coniférophyte S, i.e. mainly the Coniferous S. With these groups, they constitute the plants with seeds (Spermatophyte S).
The angiospermes differ however from the other plants with seeds by the presence of the following characters:
- the condensation of the reproductive bodies in a flower.
- the presence of a ovary wrapping the Ovule S, and which will develop to give a fruit.
- the double fertilization of the ovule, which will give the embryo and its feeder fabric, endosperm.
- the heterogeneous presence of Wood (known as hétéroxylé).
Into traditional Classification the angiospermes are divided into two taxed: the Monocotylédones (Liliopsida, Lilliidae, Moncotyledones, etc) and the Dicotylédones (Magnoliopsida, Magnoliideae, Dicotyledones, etc). This last is regarded today as Paraphylétique.
Of world distribution, Angiospermes represent most of the terrestrial Espèce S vegetable, with more than 220.000 species. Angiospermes dominate the natural landscape in the majority of the tropical or moderated Biotope S, like the Savane or the Forêt. They leave the place with the coniferous trees (Pinophytes) and to the Lichen S in the most cold biotopes. They are not very present in the aquatic environments.
Angiospermes have sizes of a few millimetres to a hundred meters (Eucalyptus).
History
It is undoubtedly Théophraste (v. 372 av. J. - C. - 287 av. J. - C.) which, the first, distinguishes Angiospermes from the Gymnosperme S. John Ray uses, at the end of the 17th century, this difference in its classification which is the first attempt at natural classification of the modern time.
Reproduction of the angiospermes
Asexual reproduction
Angiospermes present the reproductive system most advanced of the plants. In fact besides the reproductive bodies (flowers and fruits) are the most visible characteristics of the group.Angiospermes lay out of a very great number of possibilities of asexual Reproduction (mode of reproduction uniparental but not necessarily clonal). One already quoted various the Parthénogenèse S, but exist also more known processes related to the fragmentation of the thallus (certain graminaceous not selected by the Man needing to be moderately trampled for better spreading by asexual reproduction clonale in the medium). These processes are related to the Totipotence nonapoptotic vegetable cells (i.e. it quasi totality of the vegetable cells, those constituting the various circulatory systems being apoptotic or sowing-apoptotiques). One will be able to quote the exploitation of this potentiality by the Man, in particular the techniques of Bouturage, Marcottage forced or simply the fragmentation of the rhizomes or the tubers. Angiospermes use extensions racinaires or caulinaires as stolons them and the suckers, in particular at Populus , a poplar which can result only from one sucker, a whole wood can actually be one individual. It is also the case at the bamboo which, to compensate for its great slowness to generate flowers (sometimes several tens of years before reaching sexual maturity) reproduces in a superabundant way by emission of rhizomes.
Finally one will be able to also quote the asexual reproduction clonale by apoflorie: formation of buds mitotic called bulblets, which become small individuals developing their cormus (stem and root) on the plant itself before being detached about it. One observes this mode of reproduction at Kalanchoe pinnata in particular (plant crowned of Aztec).
The essential asset of the evolutionary success of Angiospermes with respect to Gymnospermes will have been the massive recourse to the asexual reproduction which makes it possible to multiply the phenotypes adapted to their environment without the individuals necessarily not reaching sexual maturity.
Sexuée reproduction
The organization of the floral parts
The terms Fleur and Plante are often confused by nonthe botanists. However, it must be noted that the flowers constitute only the reproductive part of certain plants - the plants with flower. The flower consists of several parts, of which the cheesecloth S (constituting the male part), and the Pistil (constituting the female part). A flower hermaphrodite is a flower having at the same time cheesecloths and pistil.
The distribution of the sexes at the plants with flowers
Angiospermes are with reproduction mainly sexuée, with some cases of Parthénogenèse (dandelions).The majority (70%) are hermaphrodite S (the individuals have the two sexes), with in general of the mixed flowers. However, at other species, the male and female flowers are separate on the same individual (species Monoïque S, such as the hazel tree and corn). Other combinations are possible (male flowers hermaphrodites and flowers on the same foot for example).
Some species (approximately 7%) are with sexes separated on separate plant (the individuals are male or female, for example the houx, kiwi, one speaks about species Dioïque), even flower hermaphrodite and male flower on different feet (gynodioïque Espèce). Lastly, one notes more curious reproductive cycles, like the case of the Figuier.
The individuals are not necessarily car-fertilizable S (in other words fecundation is not always possible on the same plant).
Life cycle of Angiospermes
Like all the plants, Angiospermes present an alternation between two states: the state sporophytic and the state gametophytic. However this alternation belongs to a very unbalanced cycle: the gametophytic phase is very reduced in space and time, which corresponds to a better adaptation in the middle of nonwatery life of Angiospermes. Indeed the exposure to the sun increasing the risks of changes, a predominance of the sporophytic phase (on the greatest level of ploïdie) avoids expressing the recessive changes directly. Moreover the sporophyte (for example a tree) is invariably more resistant at the plants.The gamétophytes male and female correspond respectively to the grain of pollen and the embryonic bag at Angiospermes. One would be thus wrong to think that the grain of pollen or that the embryonic bag (itself contained in the ovule , from where confusion) are gamètes: they is indeed completely false since they contain the gamètes or the futures gamètes.
The grain of pollen
See also: Pollen
Generated by the Anthère S, the grain of pollen corresponds to the male gamétophyte. It contains two to three cells and is surrounded by a double formed wall of the most resistant material of the alive world: the Sporopollénine which returns it fossilisable. This wall is consitituée of two noncellular layers: intine, intern who continuous and is secreted by the gamétophyte and the exine, external, which presents apertures and which is secreted by the sporophyte. In the case of a pollen bicellulaire, the grain of pollen contains a vegetative cell (which will germinate to form the pollen tube) and a generating cell (which corresponds to future the gamètes male, it acts of a cell not having achieved its second division of meiosis). In the case of pollen tricellulaire, the generating cell already achieved its second division of meiosis.
The embryonic Bag
It can be monosporé or tétrasporé, i.e. to come from only one mégaspore of the sporophyte or tetrad of spores resulting from the same meiosis. It is a fabric ( not a gamète ) gametophytic ( container the gamète female). It can present 7 or 4 cells. In the case of an embryonic bag with seven cells, are present there:- the oosphere, i.e. the fertilizable cell or the gamète female.
- two synergides, auxiliary cells of fecundation which form a Tore around the oosphere and which will assume the role of hopper to make forward the male gamète. These cells can possibly generate an embryo nonzygotic in certain cases of asexual reproduction.
- Three antipodes located at the other end of the embryonic bag.
- a central cell with two cores (known as polar) which will amalgamate with the male second gamète at the time of the double fertilization for possibly to give endosperm.
Pollination
See also: Pollination
Because of their immobility, the pollination of a terrestrial plant by another geographically distant installation a difficulty. If a possibility is to rely on the wind (pollination anémophile), Angiospermes present the characteristic that them Pollen is very often transported by animal species (pollination zoophile), mainly of the Insectes (pollination entomogame) but also of the Chauve-souris and Marsupiaux (pollination cheiroptérogame, met at the baobab tree) and of the Oiseau X (ornithophilous pollination), which are being attracted by the color, the form, the odor or the nectar of the flowers. What makes it possible to the authors to say In a flower, the petals also take part in the meeting of the gamètes . And indeed the organization of the floral parts plays a very particular part in fecundation, as we will develop a little further.
Fecundation
See also: double fertilization
The Fécondation is done by germination of pollen on the Pistil, without need for moisture as in the case of the Fougère S (character which they divide with other Spermaphytes).
At Angiospermes fecundation shows three essential characteristics:
- it is Siphonogame: the grain of pollen aperture having two or three cells sees its vegetative cell germinating and forming a pollen tube which will convey, using its cytosquelette, the male gamète (case of pollen tricellulaire) where the generating cell (case of pollen bicellulaire) which will carry out its second division of meiosis there. In this the siphonogamy corresponds to a total stamping from water for fecundation.
- it is Anisogame: the gamète female is most massive by far of both (the grain of pollen often being very dehydrated, the cells which it contains have a very reduced cytoplasmic space). This implies that the embryo systematically has the cytoplasmic characters of his/her mother. The transmission of the genes chloroplastic and mitochondriaux (which play a big role in certain cases of resistance to the pesticides like the atrazine or also in cases of sterility male) is not mendelienne. The mother transmits her cytoplasm in block and thus her genome mitochondrial and chloroplastic.
- it is double: the male gamètes conveyed by the grain of pollen will amalgamate their cores (caryomixie) with those of two different cells. A first caryomixie between gamète male and oosphere will give the embryo, another between the male second gamète and the cores of the ovule will lead to a triploid cell which will form the Albumen seed.
The seed
See also: Seed
Once fertilized, the oosphere will develop in embryo. The ovule containing the embryonic bag which contained it thus will form seed (as for the ovary, it will form the true Fruit, the unit granulates and fruit being called diaspore). One determines three principal categories of seeds according to their stock room of the reserves:
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albuminous seeds or seeds with Endosperm. The two products of the double fertilization develop to with it together: the polarized zygote gives the embryo and the triploid core (3x) of the central cell divides very quickly within the same cell (Coenocyte) then takes place a cellular stage of bulk-heading to form a feeder fabric: endosperm. The coconut for example presents a “milk” which corresponds in fact to the coenocytic phase of its development. Observed under the microscope, the “coconut milk” reveals that it is in fact a cytoplasm continuum: it is a giant cell having a very great number of cores. Once cellularized, this coenocyte will form the Coprah.
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exalbuminées seeds, without endosperm. Divisions of the triploid core stop very quickly but the embryo develops normally starting from the zygote. It will store its reserves in its Cotylédon S. Exemples: Rosacées (apple tree), Fabacées (or Leguminous ), Composed, Juglandées (to drown).
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seeds with Périsperme, without endosperm. Divisions of the triploid core also stop very quickly. The embryo does not store its reserves itself, it is the Nucelle which will take up this duty while becoming the périsperme. Examples: Musacées (banana), Nymphéacées, Chénopodiacées (beet, spinach).
The seed can be more or less dehydrated. Longevity being proportional to the degree of dehydration, certain seeds (in particular for the plants of the intertropical areas without unfavourable season) are to it little (example: broad bean of cocoa), others are it on extreme levels (water mass lower than 1% of the total mass at Nelumbo , the lotus of Asia whose longevity is about the thousand of years.)
Fruit
See also: Fruit
August 1st
The dispersion of the offspring
The fruit unit and seed are called diaspore, the fruit is a structure only ovarienne (this is why one speaks in many cases about “false fruits”, apple and the strawberry by forming part) while the seed comes from the ovule.The seed and the fruit correspond to a certain form of Viviparité. The embryo of Angiospermes develops initially protected and supplied with his/her relative female, then is released, in a state of slowed down life (Diapause), provided with food protections and reserves (fruit). Certain fruits present in addition to the characteristics allowing dispersion at important distances. One can quote the brushes of the dandelions, which make that the wind transports seeds on tens of meters. Let us quote also the setting with contribution of mammals for transport (by fixing in the hairs or, less important it should be admitted in the socks of campers). Another mode of dispersion is observed when animals consume certain fruits: they can then release nonedible seed and its envelopes far from his/her relative (before or after intestinal transit time, according to). It is in addition clear that an imitation and a coévolution can be in question in the adaptation of the plants to flowers to fauna able to disperse their offspring. One knows for example certain seeds whose dormancy is specifically raised by digestive enzymes of specific animals (for example some Acacia S African).
Also: Pericarp, Akène, Drupe, Fruit parthenocarpic, fleshy Fruit, dry Fruit indéhiscent
Specificity of the natural conditions
Being fixed and in nonaqueous medium, Angiospermes have to face a very great number of selective pressures. it is in particular for this reason which they are much diversified than Thallophytes (algae) which, they, develop within a medium very little changing. In fact Angiospermes represent the subphylum more diversified of Chlorobiontes (green line). This diversification necessarily passes by a genotypic diversity. Thus the descent of a angiosperme in wild conditions always tends not to be homogeneous, and thus to present a very broad range of different genotypes.To maintain this genetic diversity, the plants have recourse to the cross-fertilization (mode of reproduction sexuée at the species whose individuals interfécondent themselves systematically, in opposition to self-fertilization who was selected by the Man, we will see for which reasons) which makes impossible to circumvent the broad diffusion of pollen. Only one descent thus counts a very great number of different genotypes which will lead to phenotypes of which some certainly will be very well adapted to the medium (the others will die). The most adapted having the asexual reproduction then to multiply before to have reached sexual maturity. One knows various modes supporting the cross-fertilization (being opposed to self-fertilization).
- Systems of (car) Incompatibility Sporophytiques and Gamétophytiques (LOCATED and SIG). They are reactions of rejection of the autopollen (pollen coming from the oneself ). It was seen, the grain of pollen is composed of two to three cells and is surrounded of the sporoderme primarily composed of the most resistant material at the Alive one: the sporopollenine . this sporoderme is composed of two layers: intine and the exine from which the origins are very different. The exine (which presents the apertures) was synthesized by the sporophyte, and thus presents certain molecules (RNAses and glycoprotéines) which result from the expression of alleles sporophytic. These proteins are recognized when the autopollen arrives at the level of the mark, and they start the autoincompatibility Sporophytique (by recognition of molecules synthesized by the sporophyte). This rejection meets in the case of pollen tricellulaire.
L' autoincompatibility Gamétophytique rests on the same principle, but is based on the recognition of substances synthesized by the male gamétophyte (i.e. the grain of pollen at Angiospermes). These substances are contained in the intine and are recognized at the time of the germination of the pollen which is then rejected. The LOCATED one meets at species with pollen bicellulaire.
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shifts of maturity. At certain species the male parts of the flower are mature before the female parts (protandrie) or after (protogynie, rarer), which reduces the number of autofécondations considerably.
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floral dimorphism: the species having developed this system present two types of individuals in equal proportions in their population. Individuals with longistylées flowers (long style, mark with the top of cheesecloths, small grains of pollen and large stigmatic papillae) and individuals with brévistylées flowers (short style, cheesecloths with the top of the mark but coarse grains of pollen and small stigmatic papillae). This dimorphism prevents the autofecondation because in both cases the autopollen cannot penetrate the mark of the style, either because it is too large, or because the style is too high.
Anatomy and growth
Angiospermes are in majority Phototrophe S, drawing to them energy from solar light and them Carbone of atmospheric carbon dioxide. Some species are partially parasitic (GUI), others are completely, or are to it Saprophyte S and, in these the last two cases, are deprived of Chlorophylle. Let us quote the case of Mycotrophe S, Angiospermes without chlorophyl living in symbiotic relation with mushrooms (certain orchises, and the Monotrope S).
Angiospermes, like other Spermatophytes, develop a important system racinaire enabling them to draw from the ground the Eau and the Rock salt which they need. The root S are often the seat of Symbiose S with the Bactérie S of the ground, in particular for the metabolism of the Azote. Certain species, alive frequently on very poor grounds, supplement their contribution nitrogenizes some by the capture of animals (plants carnivores).
Also: Méristème, Liber, Phloem, Xylème, root, Mode of ramification of the plants with flower, Sheet
Phylogeny and classification
See also: Contenu=Voir [[Classification of Cronquist]] for traditional classification and [[Classification APG II]] for phylogenic classification
The oldest known fossils go back to a little more than 100 million years. It is thus about a young group, which quickly diversified. This group developed in an already rich environment, underwent a pressure of selection on behalf of many other groups (in particular animal) like the insects. This makes it possible to include/understand the astonishing adequacy which can exist between certain today angiospermes and certain animals (see concept of Coévolution). Angiospermes are traditionally classified in forty orders. One distinguishes various orders from old divergences, and two great units, the Monocotylédone S, which include inter alia the graminaceous (Blé, Maïs, Bambou), the date palm S or the plants with bulbs (Jonquille, Oignon, Poireau), and the true Dicotylédone S, which include the Chêne as well as the Potato, the Ortie S or the Artichaut S.
The history of botanical classifications shows that the systematicians try to approach an increasingly precise description of the vegetable groups according to the family ties between species. The last systems of classification published are that of Arthur Cronquist in 1981, used in many works, and more recently, that the Classification APG II (proposed by the Angiosperm Phylogeny Group) in 2003 founded on molecular phylogeny.
Importance for the man
One could not minimize the importance of Angiospermes for the human society. Not only they provide the majority of our Aliment S, directly (cereals, vegetables, fruits) or indirectly (poultry, animal feed), but still they provide, with certain Poacées (Bamboo…) and Euphorbiacées (Hévéa) a broad raw material diversity of which the use continued since the Neolithic era. Thus, they provide construction materials (wood, bamboo) that it is for the dwelling, transport (Bateau X, Brouette S), furniture or the tools; textile fibers (Cotton, flax), fuel (wood), fibers of cellulose (Paper), medicinal substances, dyes natural, elements of decorations (house plants, Gardening), etcThe selection and the intensive culture of Angiospermes are the field of the Agronomie, the Agriculture or the Sylviculture. And precisely, the Man completed a very rigorous work of selection to preserve the characters more the interesting one. Vast majority of the domesticated species present of the characters which the natural selection disadvantages. Indeed the Man primarily seeks to fix interesting characters (large fruits, beautiful flowers, many seeds, large tubers…) and in that it must be opposed to the very strong genetic mixing which carry out naturally Angiospermes. Also the Man thus tends to circumvent the systems supporting the cross-fertilization and to select the rare systems supporting self-fertilization (as the system baseds on the anatomy of the organized flower so that the autofecondation is obligatory: rolled up Hull, Cleistogamie selected at the corns Triticum and Chasmogamie). Have regard to the economic interest that the crop plants cause, the Man completed many research tasks to answer problems of output: “which are the means of obtaining a homogeneous descent of the same sex at dioïques species” when, at asparaguses for example, the male feet are systematically more productive, which gave place to the creation of the “super asparagus males” (a kind of equivalent of a genotype YY at the Man) able to give to a descent 100% male. Important work was undertaken with regard to the family of Moracées (Hop and Hemp) which it also generates an economic interest (legal or not) obvious.
It results from these processes of selection that the plants thus chosen by the Man are often homozygotes for all their couples of alleles (because born from a mode of self-fertilization maintained on a very great number of generations), even which they constitute quite simply a clone (together of individuals sharing the same genotype) as it was the case at the cultivated Potato Solanum tuberosum of which all the European cultures were in fact a clone at the 19th century. This involved a homogeneous answer vis-a-vis the mildew ( Phytophtora infestans ) and harvests were invariably destroyed. Polymorphism being the only response of the systems alive to the epidemics and the selective pressures, the monomorphism imposed by the Man for potato caused the destruction of harvests and the last great famine of the history in Europe, in particular in Ireland or almost 50% of the population found death, which supported a broad wave of immigration towards the United States.
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