The Fabaceae or Leguminosae ,  commonly known as the legume , pea , or bean family , are a large and economically important family of flowering plants. It includes trees , shrubs , and perennial or annual herbaceous plants , which are easily recognized by their fruit legume and their compound, stipulate leaves.
Many legumes have characteristic flowers and fruits. The family is widely distributed, and is the third-largest land plant family in number of species, behind only the Orchidaceae and Asteraceae , with about genera and about 19, known species.
The ca. Recent molecular and morphological evidence supports the fact that the Fabaceae is a single monophyletic family. Along with the cereals , some fruits and tropical roots, a number of Leguminosae have been a staple human food for millennia and their use is closely related to human evolution.
The family Fabaceae includes a number of important agricultural and food plants, including Glycine max soybean , Phaseolus beans , Pisum sativum pea , Cicer arietinum chickpeas , Medicago sativa alfalfa , Arachis hypogaea peanut , Ceratonia siliqua carob , and Glycyrrhiza glabra liquorice. A number of species are also weedy pests in different parts of the world, including: Cytisus scoparius broom , Robinia pseudoacacia black locust , Ulex europaeus gorse , Pueraria montana kudzu , and a number of Lupinus species.
The name 'Fabaceae' comes from the defunct genus Faba , now included in Vicia. The term "faba" comes from Latin, and appears to simply mean "bean".
Leguminosae is an older name still considered valid,  and refers to the fruit of these plants, which are called legumes. Fabaceae range in habit from giant trees like Koompassia excelsa to small annual herbs , with the majority being herbaceous perennials.
Plants have indeterminate inflorescences, which are sometimes reduced to a single flower. The flowers have a short hypanthium and a single carpel with a short gynophore , and after fertilization produce fruits that are legumes. The Leguminosae have a wide variety of growth forms , including trees, shrubs, herbaceous plants, and even vines or lianas.
The herbaceous plants can be annuals, biennials , or perennials, without basal or terminal leaf aggregations. Many Legumes have tendrils. They are upright plants, epiphytes , or vines.
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The latter support themselves by means of shoots that twist around a support or through cauline or foliar tendrils. Plants can be heliophytes, mesophytes , or xerophytes. The leaves are usually alternate and compound. Most often they are even- or odd- pinnately compound e. Caragana and Robinia respectively , often trifoliate e. Trifolium , Medicago and rarely palmately compound e. Lupinus , in the Mimosoideae and the Caesalpinioideae commonly bipinnate e. Acacia , Mimosa.
They always have stipules , which can be leaf-like e. Pisum , thorn-like e. Robinia or be rather inconspicuous. Leaf margins are entire or, occasionally, serrate. Both the leaves and the leaflets often have wrinkled pulvini to permit nastic movements.
In some species, leaflets have evolved into tendrils e. Many species have leaves with structures that attract ants which protect the plant from herbivore insects a form of mutualism. Extrafloral nectaries are common among the Mimosoideae and the Caesalpinioideae, and are also found in some Faboideae e. Vicia sativa.
In some Acacia , the modified hollow stipules are inhabited by ants and are known as domatia. Many Fabaceae host bacteria in their roots within structures called root nodules. This process is called nitrogen fixation. The legume, acting as a host, and rhizobia , acting as a provider of usable nitrate, form a symbiotic relationship.
The flowers often have five generally fused sepals and five free petals. They are generally hermaphroditic and have a short hypanthium , usually cup-shaped. There are normally ten stamens and one elongated superior ovary , with a curved style.
They are usually arranged in indeterminate inflorescences. Fabaceae are typically entomophilous plants i.
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In the Caesalpinioideae , the flowers are often zygomorphic , as in Cercis , or nearly symmetrical with five equal petals, as in Bauhinia. The upper petal is the innermost one, unlike in the Faboideae. Some species, like some in the genus Senna , have asymmetric flowers, with one of the lower petals larger than the opposing one, and the style bent to one side.
The calyx, corolla, or stamens can be showy in this group. In the Mimosoideae , the flowers are actinomorphic and arranged in globose inflorescences.
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The petals are small and the stamens, which can be more than just 10, have long, coloured filaments, which are the showiest part of the flower. All of the flowers in an inflorescence open at once. In the Faboideae , the flowers are zygomorphic, and have a specialized structure.
The upper petal, called the banner or standard, is large and envelops the rest of the petals in bud, often reflexing when the flower blooms.
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The two adjacent petals, the wings, surround the two bottom petals. The two bottom petals are fused together at the apex remaining free at the base , forming a boat-like structure called the keel. The stamens are always ten in number, and their filaments can be fused in various configurations, often in a group of nine stamens plus one separate stamen.
The ovary most typically develops into a legume. A legume is a simple dry fruit that usually dehisces opens along a seam on two sides. A common name for this type of fruit is a "pod", although that can also be applied to a few other fruit types.
A few species have evolved samarae , loments , follicles , indehiscent legumes, achenes , drupes , and berries from the basic legume fruit.
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The Leguminosae are rarely cyanogenic. Where they are, the cyanogenic compounds are derived from tyrosine , phenylalanine or leucine.
They frequently contain alkaloids.
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Proanthocyanidins can be present either as cyanidin or delphinidine or both at the same time. Flavonoids such as kaempferol , quercitin and myricetin are often present. Ellagic acid has never been found in any of the genera or species analysed.
Sugars are transported within the plants in the form of sucrose. C3 photosynthesis has been found in a wide variety of genera. Many legumes contain toxic and indigestible substances which may be removed through various processing methods.
Pterocarpans are a class of molecules derivatives of isoflavonoids found only in the Fabaceae. Forisome proteins are found in the sieve tubes of Fabaceae; uniquely they are not dependent on ADT. The order Fabales contains around 7. This clade also includes the families Polygalaceae , Surianaceae and Quillajaceae and its origins date back 94 to 89 million years, although it started its diversification 79 to 74 million years ago.
The Fabaceae have an abundant and diverse fossil record, especially for the Tertiary period. Fossils of flowers, fruit, leaves, wood and pollen from this period have been found in numerous locations. For instance, Astragalus separated from the Oxytropis 16 to 12 million years ago. In addition, the separation of the aneuploid species of Neoastragalus started 4 million years ago. Inga , another genus of the Papilionoideae with approximately species, seems to have diverged in the last 2 million years.
The current hypothesis about the evolution of the genes needed for nodulation is that they were recruited from other pathways after a polyploidy event.
The main donors to the pathway were the genes associated with the arbuscular mycorrhiza symbiosis genes, the pollen tube formation genes and the haemoglobin genes. One of the main genes shown to be shared between the arbuscular mycorrhiza pathway and the nodulation pathway is SYMRK and it is involved in the plant-bacterial recognition.
Both pathways include the same type of enzymes, pectin-degrading cell wall enzymes. To meet the requirements of this paradoxical situation, the plants express a type of haemoglobin called leghaemoglobin that is believed to be recruited after a duplication event.
The phylogeny of the legumes has been the object of many studies by research groups from around the world. Fabaceae is consistently recovered as monophyletic. Polygalaceae outgroup. The Fabaceae have an essentially worldwide distribution, being found everywhere except Antarctica and the high arctic.
Biological nitrogen fixation BNF, performed by the organisms called diazotrophs is a very old process that probably originated in the Archean eon when the primitive atmosphere lacked oxygen. It is only carried out by Euryarchaeota and just 6 of the more than 50 phyla of bacteria.
Some of these lineages co-evolved together with the flowering plants establishing the molecular basis of a mutually beneficial symbiotic relationship. BNF is carried out in nodules that are mainly located in the root cortex, although they are occasionally located in the stem as in Sesbania rostrata. The spermatophytes that co-evolved with actinorhizal diazotrophs Frankia or with rhizobia to establish their symbiotic relationship belong to 11 families contained within the Rosidae clade as established by the gene molecular phylogeny of rbcL , a gene coding for part of the RuBisCO enzyme in the chloroplast.
This grouping indicates that the predisposition for forming nodules probably only arose once in flowering plants and that it can be considered as an ancestral characteristic that has been conserved or lost in certain lineages.
However, such a wide distribution of families and genera within this lineage indicates that nodulation had multiple origins. Of the 10 families within the Rosidae, 8 have nodules formed by actinomyces Betulaceae , Casuarinaceae , Coriariaceae , Datiscaceae , Elaeagnaceae , Myricaceae , Rhamnaceae and Rosaceae , and the two remaining families, Ulmaceae and Fabaceae have nodules formed by rhizobia.
The rhizobia and their hosts must be able to recognize each other for nodule formation to commence.
Rhizobia are specific to particular host species although a rhizobia species may often infect more than one host species. This means that one plant species may be infected by more than one species of bacteria.