Fabaceae – Wikipedia

This article is about Fabaceae s.l. ( or Leguminosae ), as defined by the APG System. For Fabaceae s.s. ( or Papilionaceae ), as defined by less mod systems, see Faboideae family of legume flowering plants

The Fabaceae [ citation needed ] or Leguminosae, [ 6 ] normally known as the legume, pea, or bean family, are a bombastic and economically significant class 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 country implant family in number of species, behind only the Orchidaceae and Asteraceae, with about 765 genus and closely 20,000 know species [ 7 ]. [ 8 ] [ 9 ] [ 1 ]

The five largest genus of the family are Astragalus ( over 3,000 species ), Acacia ( over 1,000 species ), Indigofera ( around 700 species ), Crotalaria ( around 700 species ), and Mimosa ( around 400 species ), which constitute about a quarter of all legume species. The ca. 19,000 know legume species come to about 7 % of unfolding plant species. [ 9 ] [ 10 ] Fabaceae is the most common family found in tropical rainforests and dry forests of the Americas and Africa. [ 11 ] holocene molecular and morphologic evidence supports the fact that the Fabaceae is a single monophyletic family. [ 12 ] This decision has been supported not only by the degree of interrelation shown by different groups within the syndicate compared with that establish among the Leguminosae and their close relations, but besides by all the recent phylogenetic studies based on DNA sequences. [ 13 ] [ 14 ] [ 15 ] These studies confirm that the Fabaceae are a monophyletic group that is closely related to the families Polygalaceae, Surianaceae and Quillajaceae and that they belong to the ordain Fabales. [ 16 ] Along with the cereals, some fruits and tropical roots, a count of Leguminosae have been a staple homo food for millennium and their manipulation is close related to human development. [ 17 ] The family Fabaceae includes a numeral of important agrarian and food plants, including Glycine max ( soy ), Phaseolus ( beans ), Pisum sativum ( pea ), Cicer arietinum ( chickpeas ), Medicago sativa ( alfalfa ), Arachis hypogaea ( peanut ), Ceratonia siliqua ( carob ), and Glycyrrhiza glabra ( licorice ). A numeral of species are besides scraggy pests in different parts of the world, including : Cytisus scoparius ( heather ), Robinia pseudoacacia ( black locust tree ) , Ulex europaeus ( gorse ), Pueraria montana ( kudzu ), and a number of Lupinus species .

etymology [edit ]

The name ‘Fabaceae ‘ comes from the defunct genus Faba, now included in Vicia. The term “ faba ” comes from Latin, and appears to plainly mean “ bean ”. Leguminosae is an older diagnose still considered valid, [ 6 ] and refers to the yield of these plants, which are called legumes .

description [edit ]

Fabaceae scope in habit from colossus trees ( like Koompassia excelsa ) to little annual herb, with the majority being herbaceous perennials. Plants have indeterminate inflorescences, which are sometimes reduced to a single flower. The flowers have a curtly hypanthium and a single carpel with a short gynophore, and after fertilization produce fruits that are legumes .

Growth habit [edit ]

The Fabaceae have a wide variety of growth forms, including trees, shrubs, herbaceous plants, and flush 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. The latter support themselves by means of shoots that twist around a support or through caulescent or foliar tendrils. Plants can be heliophytes, mesophytes, or xerophytes. [ 3 ] [ 9 ]

Leaves [edit ]

The leaves are normally alternate and compound. Most often they are even- or odd- pinnately intensify ( e.g. Caragana and Robinia respectively ), frequently trifoliate ( e.g. Trifolium, Medicago ) and rarely palmately colonial ( e.g. Lupinus ), in the Mimosoideae and the Caesalpinioideae normally bipinnate ( e.g. Acacia, Mimosa ). They constantly have stipules, which can be leaflike ( e.g. Pisum ), thorn-like ( e.g. Robinia ) or be rather inconspicuous. leaf margins are entire or, occasionally, serrate. Both the leaves and the leaflets much have wrinkled pulvini to permit nastic movements. In some species, leaflets have evolved into tendrils ( e.g. Vicia ). [ 3 ] [ 9 ] [ 17 ] many species have leaves with structures that attract ants which protect the plant from herbivore insects ( a form of symbiosis ). Extrafloral nectaries are common among the Mimosoideae and the Caesalpinioideae, and are besides found in some Faboideae ( e.g. Vicia sativa ). In some Acacia, the modified hole stipules are inhabited by ants and are known as domatium .

Roots [edit ]

many Fabaceae host bacteria in their roots within structures called root nodules. These bacteria, known as rhizobium, have the ability to take nitrogen boast ( N2 ) out of the air and convert it to a form of nitrogen that is functional to the master of ceremonies plant ( NO3− or NH3 ). This serve is called nitrogen fixation. The legume, acting as a host, and rhizobium, acting as a supplier of useable nitrate, form a symbiotic relationship. Members of the Phaseoleae genus Apios form tubers, which can be edible. [ 18 ]

Flowers [edit ]

“ Pea flower ” redirects here. For the flour produced from peas, see pea flour Wisteria sinensis, Faboideae. Two petals have been removed to show stamens and pistil A flower of, Faboideae. Two petals have been removed to show stamens and pistil The flowers much have five broadly fused sepals and five free petals. They are by and large hermaphroditic and have a short hypanthium, normally cup-shaped. There are normally ten stamens and one elongated superior ovary, with a curved vogue. They are normally arranged in indeterminate inflorescences. Fabaceae are typically entomophilous plants ( i.e. they are pollinated by insects ), and the flowers are normally showy to attract pollinators. In the Caesalpinioideae, the flowers are much zygomorphic, as in Cercis, or closely harmonious with five adequate petals, as in Bauhinia. The upper petal is the inmost one, unlike in the Faboideae. Some species, like some in the genus Senna, have asymmetrical 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 ball-shaped inflorescences. The petals are little and the stamens, which can be more than precisely 10, have long, coloured filaments, which are the showiest part of the flower. All of the flowers in an blossoming open at once. In the Faboideae, the flowers are zygomorphic, and have a specialize social organization. The upper berth petal, called the banner or standard, is big and envelops the pillow of the petals in bud, frequently reflexing when the bloom blooms. The two adjacent petals, the wings, surround the two bottom petals. The two penetrate petals are fused together at the apex ( remaining rid at the basis ), forming a boat-like social organization called the keel. The stamens are always ten in phone number, and their filaments can be fused in diverse configurations, frequently in a group of nine stamens plus one disjoined stamen. assorted genes in the CYCLOIDEA (CYC)/DICHOTOMA (DICH) syndicate are expressed in the upper berth ( besides called abaxial or adaxial ) petal ; in some species, such as Cadia, these genes are expressed throughout the bloom, producing a radially harmonious flower. [ 19 ]

fruit [edit ]

The ovary most typically develops into a legume. A legume is a simpleton dry fruit that normally dehisces ( opens along a wrinkle ) on two sides. A coarse diagnose for this character of fruit is a “ pod ”, although that can besides be applied to a few other yield types. A few species have evolved samarae, loments, follicles, indehiscent legumes, achenes, drupes, and berries from the basic legume fruit .

Physiology and biochemistry [edit ]

The Fabaceae are rarely cyanogenetic. Where they are, the cyanogenetic compounds are derived from tyrosine, phenylalanine or leucine. They frequently contain alkaloids. Proanthocyanidins can be present either as cyanidin or delphinidine or both at the same time. Flavonoids such as kaempferol, quercitin and myricetin are frequently confront. Ellagic acerb has never been found in any of the genus or species analysed. Sugars are transported within the plants in the form of sucrose. C3 photosynthesis has been found in a wide variety of genus. [ 3 ] The family has besides evolved a alone chemistry. many legumes contain toxic [ 20 ] and indigestible substances, antinutrients, which may be removed through versatile processing methods. Pterocarpans are a class of molecules ( derivatives of isoflavonoids ) found only in the Fabaceae. Forisome proteins are found in the sieve tube of Fabaceae ; uniquely they are not dependent on ADT .

evolution, evolution and taxonomy [edit ]

evolution [edit ]

The ordering Fabales contains around 7.3 % of eudicot species and the greatest part of this diversity is contained in equitable one of the four families that the order contains : Fabaceae. This clade besides 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. [ 1 ] In fact, the Fabaceae have diversified during the early tertiary to become a omnipresent region of the modern earth ‘s biota, along with many other families belonging to the blossoming plants. [ 12 ] [ 21 ] The Fabaceae have an abundant and diverse fossil phonograph record, particularly for the Tertiary period. Fossils of flowers, fruit, leaves, forest and pollen from this menstruation have been found in numerous locations. [ 22 ] [ 23 ] [ 24 ] [ 25 ] [ 26 ] [ 27 ] [ 28 ] The earliest fossils that can be definitively assigned to the Fabaceae appeared in the early Palaeocene ( approximately 65 million years ago ). [ 29 ] Representatives of the 3 sub-families traditionally recognised as being members of the Fabaceae – Cesalpinioideae, Papilionoideae and Mimosoideae – deoxyadenosine monophosphate well as members of the boastfully clades within these sub-families – such as the genistoides – have been found in periods late, starting between 55 and 50 million years ago. [ 21 ] In fact, a broad diverseness of taxonomic group representing the independent lineages in the Fabaceae have been found in the fossil record dating from the middle to the former Eocene, suggesting that the majority of the modern Fabaceae groups were already introduce and that a broad diversification occurred during this period. [ 21 ] Therefore, the Fabaceae started their diversification approximately 60 million years ago and the most crucial clades separated 50 million years ago. [ 30 ] The age of the main Cesalpinioideae clades have been estimated as between 56 and 34 million years and the basal group of the Mimosoideae as 44 ± 2.6 million years. [ 31 ] [ 32 ] The division between Mimosoideae and Faboideae is dated as occurring between 59 and 34 million years ago and the radical group of the Faboideae as 58.6 ± 0.2 million years ago. [ 33 ] It has been potential to date the divergence of some of the groups within the Faboideae, even though diversification within each genus was relatively recent. For example, 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 350 species, seems to have diverged in the last 2 million years. [ 34 ] [ 35 ] [ 36 ] [ 37 ] It has been suggested, based on dodo and phylogenetic evidence, that legumes in the first place evolved in arid and/or semi-arid regions along the Tethys seaway during the Palaeogene Period. [ 5 ] [ 38 ] however, others contend that Africa ( or flush the Americas ) can not however be ruled out as the lineage of the syndicate. [ 39 ] [ 40 ] The current guess about the evolution of the genes needed for nodulation is that they were recruited from early pathways after a polyploidy event. [ 41 ] respective unlike pathways have been implicated as donating duplicated genes to the pathways need for nodulation. The independent donors to the pathway were the genes associated with the arbuscular mycorrhiza symbiosis genes, the pollen metro formation genes and the hemoglobin 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. [ 42 ] The pollen tube increase is alike to the infection thread development in that infection threads grow in a polar manner that is similar to a pollen tubes polar growth towards the ovules. Both pathways include the lapp type of enzymes, pectin-degrading cell wall enzymes. [ 43 ] The enzymes needed to reduce nitrogen, nitrogenases, require a significant remark of ATP but at the lapp time are sensible to release oxygen. To meet the requirements of this paradoxical situation, the plants express a type of hemoglobin called leghaemoglobin that is believed to be recruited after a duplication event. [ 44 ] These three genetic pathways are believed to be function of a gene duplication event then recruited to work in nodulation .

Phylogeny and taxonomy [edit ]

evolution [edit ]

The evolution of the legumes has been the aim of many studies by research groups from around the populace. These studies have used morphology, DNA data ( the chloroplast intron trnL, the chloroplast genes rbcL and matK, or the ribosomal spacers ITS ) and cladistic analysis in order to investigate the relationships between the family ‘s different lineages. Fabaceae is systematically recovered as monophyletic. [ 45 ] The studies further confirmed that the traditional subfamilies Mimosoideae and Papilionoideae were each monophyletic but both were nested within the paraphyletic subfamily Caesalpinioideae. [ 46 ] [ 45 ] All the different approaches yielded like results regarding the relationships between the syndicate ‘s independent clades. [ 1 ] [ 47 ] [ 48 ] [ 49 ] [ 50 ] [ 51 ] [ 52 ] [ 53 ] [ 54 ] Following extensive discussion in the legume phylogenetics community, the Legume Phylogeny Working Group reclassified Fabaceae into six subfamilies, which necessitated the segregation of four new subfamilies from Caesalpinioideae and merging Caesapinioideae sensu stricto with the former subfamily Mimosoideae. [ 4 ] [ 55 ] The claim branching order of the different subfamilies is still unsolved. [ 56 ]

Fabales
Polygalaceae ( outgroup )
Surianaceae ( outgroup )
Quillajaceae ( outgroup )
Fabaceae
Cercidoideae
Detarioideae
Duparquetioideae

Dialioideae
caesalpinioideae
Faboideae

taxonomy [edit ]

The Fabaceae are placed in the order Fabales according to most taxonomic systems, including the APG III organization. [ 2 ] The family now includes six subfamilies : [ 4 ]

  • Cercidoideae: 12 genera and ~335 species. Mainly tropical. Bauhinia, Cercis.
  • Detarioideae: 84 genera and ~760 species. Mainly tropical. Amherstia, Detarium, Tamarindus.
  • Duparquetioideae: 1 genus and 1 species. West and Central Africa. Duparquetia.
  • Dialioideae: 17 genera and ~85 species. Widespread throughout the tropics. Dialium.
  • Caesalpinioideae: 148 genera and ~4400 species. Pantropical. Caesalpinia, Senna, Mimosa, Acacia. Includes the former subfamily Mimosoideae (80 genera and ~3200 species; mostly tropical and warm temperate Asia and America).
  • Faboideae (Papilionoideae[57]): 503 genera and ~14,000 species. Cosmopolitan. Astragalus, Lupinus, Pisum.

ecology [edit ]

distribution and habitat [edit ]

The Fabaceae have an basically global distribution, being found everywhere except Antarctica and the high Arctic. [ 1 ] The trees are much found in tropical regions, while the herbaceous plants and shrubs are overriding outside the tropics. [ 3 ]

biological nitrogen fixation [edit ]

Vicia with white root nodules visible. Roots ofwith white solution nodules visible . Vicia observed through a microscope. Cross-section through a root nodule ofobserved through a microscope. 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 standard atmosphere lacked oxygen. It is alone carried out by Euryarchaeota and good 6 of the more than 50 phylum of bacteria. Some of these lineages co-evolved together with the bloom plants establishing the molecular basis of a mutually beneficial symbiotic relationship. BNF is carried out in nodules that are chiefly located in the settle cortex, although they are occasionally located in the stem as in Sesbania rostrata. The spermatophytes that co-evolved with actinorhizal diazotrophs ( Frankia ) or with rhizobium to establish their symbiotic relationship belong to 11 families contained within the Rosidae clade ( as established by the gene molecular evolution of rbcL, a gene coding for separate of the RuBisCO enzyme in the chloroplast ). This grouping indicates that the predisposition for forming nodules probably lone arise 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 genus within this linage 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 rhizobium. [ 58 ] [ 59 ] The rhizobium and their hosts must be able to recognize each other for nodule formation to start. Rhizobia are particular to particular host species although a rhizobium 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. For exemplar, nodules in Acacia senegal can contain seven species of rhizobium belonging to three unlike genus. The most distinctive characteristics that allow rhizobia to be distinguished apart are the celerity of their growth and the type of ancestor nodule that they form with their server. [ 59 ] Root nodules can be classified as being either indeterminate, cylindrical and much branched, and definitive, ball-shaped with big lenticels. Indeterminate nodules are characteristic of legumes from temperate climates, while determinate nodules are normally found in species from tropical or subtropical climates. [ 59 ] Nodule geological formation is common throughout the Fabaceae. It is found in the majority of its members that entirely form an association with rhizobium, which in turn form an single symbiosis with the Fabaceae ( with the exception of Parasponia, the entirely genus of the 18 Ulmaceae genus that is able of forming nodules ). Nodule constitution is present in all the Fabaceae sub-families, although it is less common in the Caesalpinioideae. All types of nodule formation are present in the subfamily Papilionoideae : indeterminate ( with the meristem retained ), determinate ( without meristem ) and the type included in Aeschynomene. The latter two are thought to be the most modern and specialize type of nodule as they are only present in some lines of the subfamily Papilionoideae. even though nodule formation is common in the two monophyletic subfamilies Papilionoideae and Mimosoideae they besides contain species that do not form nodules. The bearing or absence of nodule-forming species within the three sub-families indicates that nodule formation has arisen several times during the evolution of the Fabaceae and that this ability has been lost in some lineages. For exemplar, within the genus Acacia, a penis of the Mimosoideae, A. pentagona does not form nodules, while other species of the lapp genus readily human body nodules, as is the lawsuit for Acacia senegal, which forms both quickly and slow growing rhizobial nodules .

Chemical ecology [edit ]

A bombastic count of species within many genus of leguminous plants, e.g. Astragalus, Coronilla, Hippocrepis, Indigofera, Lotus, Securigera and Scorpiurus, produce chemicals that derive from the compound 3-nitropropanoic acidic ( 3-NPA, beta-nitropropionic acid ). The loose acid 3-NPA is an irreversible inhibitor of mitochondrial breathing, and frankincense the compound inhibits the tricarboxylic acid bicycle. This inhibition caused by 3-NPA is particularly toxic to nerve cells and represents a very general toxic mechanism suggesting a profound ecological importance due to the big count of species producing this compound and its derivatives. A second and closely related class of secondary metabolites that occur in many species of leguminous plants is defined by isoxazolin-5-one derivatives. These compounds occur in particular together with 3-NPA and relate derivatives at the like time in the like species, as found in Astragalus canadensis and Astragalus collinus. 3-NPA and isoxazlin-5-one derivatives besides occur in many species of leaf beetles ( see defensive structure in insects ). [ 60 ]

Economic and cultural importance [edit ]

Legumes are economically and culturally important plants due to their extraordinary diverseness and abundance, the wide kind of comestible vegetables they represent and ascribable to the kind of uses they can be put to : in gardening and department of agriculture, as a food, for the compounds they contain that have medicative uses and for the vegetable oil and fats they contain that have a variety show of uses. [ 61 ] [ 62 ] [ 63 ] [ 64 ]

food and scrounge [edit ]

The history of legumes is tied in closely with that of human civilization, appearing early in Asia, the Americas ( the common bean, several varieties ) and Europe ( broad beans ) by 6,000 BCE, where they became a staple, essential as a source of protein. Their ability to fix atmospheric nitrogen reduces fertilizer costs for farmers and gardeners who grow legumes, and means that legumes can be used in a snip rotation to replenish soil that has been depleted of nitrogen. Legume seeds and leaf have a relatively higher protein content than non-legume materials, ascribable to the extra nitrogen that legumes receive through the action. Legumes are normally used as natural fertilizers. Some legume species perform hydraulic lift, which makes them ideal for intercropping. [ 65 ] Farmed legumes can belong to numerous classes, including scrounge, granulate, blooms, pharmaceutical/industrial, fallow/green manure and timbre species, with most commercially farm species filling two or more roles simultaneously. There are of two broad types of eatage legumes. Some, like alfalfa, clover, vetch, and Arachis, are sown in crop and grazed by livestock. early scrounge legumes such as Leucaena or Albizia are woody shrub or tree species that are either broken down by livestock or regularly cut by humans to provide fodder. granulate legumes are cultivated for their seeds, and are besides called pulses. The seeds are used for homo and animal consumption or for the production of oils for industrial uses. Grain legumes include both herbaceous plants like beans, lentils, lupins, peas and peanuts. [ 66 ] and trees such as carob, mesquite and tamarind. Lathyrus tuberosus, once extensively cultivated in Europe, forms tubers used for human consumption. [ 67 ] [ 68 ] Bloom legume species include species such as lupine, which are farmed commercially for their blooms, and frankincense are popular in gardens worldwide. Laburnum, Robinia, Gleditsia ( honey locust tree ), Acacia, Mimosa, and Delonix are cosmetic trees and shrubs. Industrial farmed legumes include Indigofera, cultivated for the production of indigo, Acacia, for mumble arabic, and Derris, for the insecticide action of rotenone, a compound it produces. Fallow or green manure legume species are cultivated to be tilled back into the land to exploit the high nitrogen levels found in most legumes. numerous legumes are farmed for this determination, including Leucaena, Cyamopsis and Sesbania. assorted legume species are farmed for timber production global, including numerous Acacia species, Dalbergia species, and Castanospermum australe. Melliferous plants offer ambrosia to bees and other insects to encourage them to carry pollen from the flowers of one plant to others thereby ensuring pollination. Some legume species such as alfalfa, ashen clover, sweet clover and respective Prosopis species are dear nectar providers. many plants in the family Fabaceae are an crucial source of pollen for the bumblebee species Bombus hortorum. This bee species is particularly fond of one species in particular ; Trifolium pratense, besides known as red clover, is a popular food source in the diet of Bombus hortorum. [ 69 ]

Industrial uses [edit ]

Natural gums [edit ]

Natural gums are vegetable exudates that are released as the result of damage to the plant such as that resulting from the attack of an insect or a natural or artificial cut. These exudates contain heterogenous polysaccharides formed of different sugars and normally containing uronic acids. They form syrupy colloidal solutions. There are different species that produce gums. The most significant of these species belong to the Fabaceae. They are widely used in the pharmaceutical, cosmetic, food, and fabric sectors. They besides have interesting therapeutic properties ; for example gingiva arabic is antitussive and anti-inflammatory. The most well known gums are tragacanth ( Astragalus gummifer ), gumwood arabic ( Acacia senegal ) and guar gum ( Cyamopsis tetragonoloba ). [ 70 ]

Dyes [edit ]

respective species of Fabaceae are used to produce dyes. The heartwood of logwood, Haematoxylon campechianum, is used to produce crimson and purple dyes. The histological stain called haematoxylin is produced from this species. The wood of the Brazilwood tree ( Caesalpinia echinata ) is besides used to produce a loss or purple dye. The Madras thorn ( Pithecellobium dulce ) has red fruit that are used to produce a yellow dye. [ 71 ] Indigo dye is extracted from the anil plant Indigofera tinctoria that is native to Asia. In Central and South America dyes are produced from two species in the same genus : anil and Maya blue from Indigofera suffruticosa and Natal indigo from Indigofera arrecta. yellow dyes are extracted from Butea monosperma, normally called flare of the forest and from dyer ‘s woodwaxen, ( Genista tinctoria ). [ 72 ]

Ornamentals [edit ]

Legumes have been used as ornamental plants throughout the global for many centuries. Their huge diverseness of heights, shapes, leaf and flower color means that this kin is normally used in the design and establish of everything from little gardens to bombastic parks. [ 17 ] The trace is a tilt of the independent cosmetic legume species, listed by subfamily .

emblematic Fabaceae [edit ]

image gallery [edit ]

References [edit ]

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