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 modern systems, seeFaboideae.
The five largest genera of the family areAstragalus (over 3,000 species),Acacia (over 1,000 species),Indigofera (around 700 species),Crotalaria (around 700 species), andMimosa (around 400 species), which constitute about a quarter of all legume species. The c. 19,000 known legume species amount to about 7% of flowering plant species.[9][11] Fabaceae is the most common family found in tropical rainforests and dry forests of the Americas and Africa.[12]
Recent molecular and morphological evidence supports the fact that the Fabaceae is a singlemonophyletic family.[13] This conclusion has been supported not only by the degree of interrelation shown by different groups within the family compared with that found among the Leguminosae and their closest relations, but also by all the recentphylogenetic studies based onDNA sequences.[14][15][16] These studies confirm that the Fabaceae are a monophyletic group that is closely related to the familiesPolygalaceae,Surianaceae andQuillajaceae and that they belong to the orderFabales.[17]
Along with thecereals, some fruits and tropical roots, a number of Leguminosae have been a staple human food for millennia and their use is closely related tohuman evolution.[18]
The name 'Fabaceae' comes from the defunct genusFaba, now included inVicia. The term "faba" comes from Latin, and appears to simply mean "bean". Leguminosae is an older name still considered valid,[6] and refers to thefruit of these plants, which are calledlegumes.
Fabaceae range in habit from gianttrees (likeKoompassia excelsa) to smallannualherbs, with the majority being herbaceous perennials. Plants have indeterminate inflorescences, which are sometimes reduced to a single flower. The flowers have a shorthypanthium and a singlecarpel with a shortgynophore, and after fertilization produce fruits that are legumes.
The Fabaceae have a wide variety ofgrowth forms, including trees, shrubs, herbaceous plants, and evenvines orlianas. 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 cauline or foliartendrils. Plants can beheliophytes,mesophytes, orxerophytes.[3][9]
The leaves are usuallyalternate and compound. Most often they are even- or odd-pinnately compound (e.g.Caragana andRobinia respectively), often trifoliate (e.g.Trifolium,Medicago) and rarelypalmately compound (e.g.Lupinus), in the Mimosoideae and the Caesalpinioideae commonly bipinnate (e.g.Acacia,Mimosa). They always havestipules, which can be leaf-like (e.g.Pisum), thornlike (e.g.Robinia) or be rather inconspicuous. Leaf margins are entire or, occasionally,serrate. Both the leaves and the leaflets often have wrinkledpulvini to permitnastic movements. In some species, leaflets have evolved intotendrils (e.g.Vicia).[3][9][18]
Many species have leaves with structures that attractants which protect the plant from herbivore insects (a form ofmutualism).Extrafloral nectaries are common among the Mimosoideae and the Caesalpinioideae, and are also found in some Faboideae (e.g.Vicia sativa). In someAcacia, the modified hollow stipules are inhabited by ants and are known asdomatia.
Many Fabaceae hostbacteria in their roots within structures calledroot nodules. These bacteria, known asrhizobia, have the ability to takenitrogen gas (N2) out of the air and convert it to a form of nitrogen that is usable to the host plant (NO3− orNH3). This process is callednitrogen fixation. The legume, acting as a host, andrhizobia, acting as a provider of usable nitrate, form asymbiotic relationship. Members of thePhaseoleae genusApios form tubers, which can be edible.[19]
In theCaesalpinioideae, the flowers are oftenzygomorphic, as inCercis, or nearly symmetrical with five equal petals, as inBauhinia. The upper petal is the innermost one, unlike in theFaboideae. Some species, like some in the genusSenna, 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 theMimosoideae, the flowers areactinomorphic and arranged in globose inflorescences. 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 theFaboideae, the flowers are zygomorphic, and have aspecialized 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. 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. Various genes in theCYCLOIDEA (CYC)/DICHOTOMA (DICH) family are expressed in the upper (also called dorsal or adaxial) petal; in some species, such asCadia, these genes are expressed throughout the flower, producing a radially symmetrical flower.[20]
The ovary most typically develops into alegume. A legume is asimple dry fruit that usuallydehisces (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 evolvedsamarae,loments,follicles, indehiscent legumes,achenes,drupes, andberries from the basic legume fruit.
The Fabaceae are rarelycyanogenic. Where they are, the cyanogenic compounds are derived fromtyrosine,phenylalanine orleucine. They frequently containalkaloids.Proanthocyanidins can be present either ascyanidin ordelphinidine or both at the same time.Flavonoids such askaempferol,quercitin andmyricetin 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 ofsucrose.C3 photosynthesis has been found in a wide variety of genera.[3] The family has also evolved a unique chemistry. Many legumes contain toxic[21] and indigestible substances,antinutrients, which may be removed through various processing methods.Pterocarpans are a class of molecules (derivatives ofisoflavonoids) found only in the Fabaceae.Forisome proteins are found in the sieve tubes of Fabaceae; uniquely they are not dependent onADT.
The order Fabales contains around 7.3% of eudicot species and the greatest part of this diversity is contained in just one of the four families that the order contains: Fabaceae. This clade also includes the familiesPolygalaceae,Surianaceae andQuillajaceae and its origins date back 94 to 89 million years, although it started its diversification 79 to 74 million years ago.[10] The Fabaceae diversified during thePaleogene to become a ubiquitous part of the modern earth'sbiota, along with many other families belonging to the flowering plants.[13][22]
The Fabaceae have an abundant and diversefossil record, especially for theTertiary period. Fossils of flowers, fruit, leaves, wood andpollen from this period have been found in numerous locations.[23][24][25][26][27][28][29] The earliest fossils that can be definitively assigned to the Fabaceae appeared in the earlyPalaeocene (approximately 65 million years ago).[30] Representatives of the 3 sub-families traditionally recognised as being members of the Fabaceae – Cesalpinioideae, Papilionoideae and Mimosoideae – as well as members of the large clades within these sub-families – such as the genistoides – have been found in periods later, starting between 55 and 50 million years ago.[22] In fact, a wide variety of taxa representing the main lineages in the Fabaceae have been found in the fossil record dating from the middle to the lateEocene, suggesting that the majority of the modern Fabaceae groups were already present and that a broad diversification occurred during this period.[22] Therefore, the Fabaceae started their diversification approximately 60 million years ago and the most important clades separated 50 million years ago.[31] 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.[32][33] The division between Mimosoideae and Faboideae is dated as occurring between 59 and 34 million years ago and the basal group of the Faboideae as 58.6 ± 0.2 million years ago.[34] It has been possible to date the divergence of some of the groups within the Faboideae, even though diversification within each genus was relatively recent. For instance,Astragalus separated from theOxytropis 16 to 12 million years ago. In addition, the separation of theaneuploid species ofNeoastragalus 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.[35][36][37][38]
It has been suggested, based on fossil and phylogenetic evidence, that legumes originally evolved in arid and/or semi-arid regions along theTethys seaway during thePalaeogene Period.[5][39] However, others contend thatAfrica (or even theAmericas) cannot yet be ruled out as the origin of the family.[40][41]
The current hypothesis about the evolution of the genes needed for nodulation is that they were recruited from other pathways after a polyploidy event.[42] Several different pathways have been implicated as donating duplicated genes to the pathways need for nodulation. 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.[43] The pollen tube growth is similar 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 same type of enzymes, pectin-degrading cell wall enzymes.[44] The enzymes needed to reduce nitrogen, nitrogenases, require a substantial input of ATP but at the same time are sensitive to free oxygen. 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.[45] These three genetic pathways are believed to be part of a gene duplication event then recruited to work in nodulation.
Thephylogeny of the legumes has been the object of many studies by research groups from around the world. These studies have used morphology,DNA data (thechloroplastintrontrnL, the chloroplastgenesrbcL andmatK, or the ribosomal spacersITS) andcladistic analysis in order to investigate the relationships between the family's different lineages. Fabaceae is consistently recovered asmonophyletic.[46] The studies further confirmed that the traditional subfamilies Mimosoideae and Papilionoideae were eachmonophyletic but both were nested within the paraphyletic subfamily Caesalpinioideae.[47][46] All the different approaches yielded similar results regarding the relationships between the family's main clades.[10][48][49][50][51][52][53][54][55][excessive citations] 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 Caesapinioideaesensu stricto with the former subfamily Mimosoideae.[4][56] The exact branching order of the different subfamilies is still unresolved.[57]
The Fabaceae are placed in the orderFabales according to most taxonomic systems, including theAPG III system.[2] The family now includes six subfamilies:[4]
The Fabaceae have an essentially worldwide distribution, being found everywhere except Antarctica and the high Arctic.[10] The trees are often found in tropical regions, while the herbaceous plants and shrubs are predominant outside the tropics.[3]
Roots ofVicia with white root nodules visible.Cross-section through a root nodule ofVicia observed through a microscope.
Biological nitrogen fixation (BNF, performed by the organisms calleddiazotrophs) is a very old process that probably originated in theArchean eon when the primitiveatmosphere lackedoxygen. It is only carried out byEuryarchaeota and just 6 of the more than 50phyla ofbacteria. Some of these lineages co-evolved together with theflowering plants establishing the molecular basis of a mutually beneficialsymbiotic relationship. BNF is carried out in nodules that are mainly located in the root cortex, although they are occasionally located in the stem as inSesbania rostrata. Thespermatophytes that co-evolved withactinorhizal diazotrophs (Frankia) or withrhizobia to establish their symbiotic relationship belong to 11 families contained within theRosidaeclade (as established by the gene molecular phylogeny ofrbcL, a gene coding for part of theRuBisCO enzyme in thechloroplast). 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 byactinomyces (Betulaceae,Casuarinaceae,Coriariaceae,Datiscaceae,Elaeagnaceae,Myricaceae,Rhamnaceae andRosaceae), and the two remaining families,Ulmaceae and Fabaceae have nodules formed by rhizobia.[59][60]
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. For example, nodules inAcacia senegal can contain seven species of rhizobia belonging to three different genera. The most distinctive characteristics that allow rhizobia to be distinguished apart are the rapidity of their growth and the type of root nodule that they form with their host.[60] Root nodules can be classified as being either indeterminate, cylindrical and often branched, and determinate, spherical with prominent lenticels. Indeterminate nodules are characteristic of legumes from temperate climates, while determinate nodules are commonly found in species from tropical or subtropical climates.[60]
Nodule formation is common throughout the Fabaceae. It is found in the majority of its members that only form an association with rhizobia, which in turn form an exclusive symbiosis with the Fabaceae (with the exception ofParasponia, the only genus of the 18 Ulmaceae genera that is capable of forming nodules). Nodule formation 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 themeristem retained), determinate (without meristem) and the type included inAeschynomene. The latter two are thought to be the most modern and specialised type of nodule as they are only present in some lines of the subfamily Papilionoideae. Even though nodule formation is common in the twomonophyletic subfamilies Papilionoideae and Mimosoideae they also contain species that do not form nodules. The presence 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 example, within the genusAcacia, a member of the Mimosoideae,A. pentagona does not form nodules, while other species of the same genus readily form nodules, as is the case forAcacia senegal, which forms both rapidly and slow growing rhizobial nodules.
A large number of species within many genera of leguminous plants, e.g.Astragalus,Coronilla,Hippocrepis,Indigofera,Lotus,Securigera andScorpiurus, produce chemicals that derive from the compound 3-nitropropanoic acid (3-NPA,beta-nitropropionic acid). The free acid 3-NPA is anirreversible inhibitor of mitochondrialrespiration, and thus the compound inhibits thetricarboxylic acid cycle. This inhibition caused by 3-NPA is especially toxic to nerve cells and represents a very general toxic mechanism suggesting a profound ecological importance due to the big number 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 related derivatives at the same time in the same species, as found inAstragalus canadensis andAstragalus collinus. 3-NPA and isoxazlin-5-one derivatives also occur in many species of leaf beetles (seedefense in insects).[61]
Legumes are economically and culturally important plants due to their extraordinary diversity and abundance, the wide variety of edible vegetables they represent and due to the variety of uses they can be put to: in horticulture and agriculture, as a food, for the compounds they contain that have medicinal uses and for the oil and fats they contain that have a variety of uses.[62][63][64][65]
Thehistory of legumes is tied in closely with that of human civilization, appearing early inAsia, theAmericas (thecommon bean, several varieties) andEurope (broad beans) by 6,000BCE, where they became a staple, essential as a source of protein.
Their ability tofix atmospheric nitrogen reducesfertilizer costs for farmers and gardeners who grow legumes, and means that legumes can be used in acrop rotation to replenish soil that has been depleted ofnitrogen. Legume seeds and foliage have a comparatively higherprotein content than non-legume materials, due to the additional nitrogen that legumes receive through the process. Legumes are commonly used as natural fertilizers. Some legume species performhydraulic lift, which makes them ideal forintercropping.[66]
Farmed legumes can belong to numerous classes, includingforage,grain, blooms, pharmaceutical/industrial, fallow/green manure and timber species, with most commercially farmed species filling two or more roles simultaneously.
There are of two broad types of forage legumes. Some, likealfalfa,clover,vetch, andArachis, are sown inpasture and grazed by livestock. Other forage legumes such asLeucaena orAlbizia are woody shrub or tree species that are either broken down by livestock or regularly cut by humans to providefodder.
Grain legumes are cultivated for theirseeds, and are also calledpulses. The seeds are used for human and animal consumption or for the production ofoils for industrial uses. Grain legumes include both herbaceous plants likebeans,lentils,lupins,peas andpeanuts,[67] and trees such ascarob,mesquite andtamarind.
Lathyrus tuberosus, once extensively cultivated in Europe, forms tubers used for human consumption.[68][69]
Fallow orgreen manure legume species are cultivated to be tilled back into the soil to exploit the high nitrogen levels found in most legumes. Numerous legumes are farmed for this purpose, includingLeucaena,Cyamopsis andSesbania.
Melliferous plants offernectar tobees and other insects to encourage them to carry pollen from theflowers of one plant to others thereby ensuring pollination. Many Fabaceae species are important sources of pollen and nectar for bees, including for honey production in the beekeeping industry. Example Fabaceae such asalfalfa, and various clovers includingwhite clover andsweet clover, are important sources of nectar and honey for theWestern honey bee.[70]
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 heterogeneouspolysaccharides formed of different sugars and usually containinguronic acids. They form viscous colloidal solutions. There are different species that produce gums. The most important of these species belong to the Fabaceae. They are widely used in the pharmaceutical, cosmetic, food, and textile sectors. They also have interesting therapeutic properties; for examplegum arabic isantitussive andanti-inflammatory.[medical citation needed] The most well known gums aretragacanth (Astragalus gummifer), gum arabic (Acacia senegal) andguar gum (Cyamopsis tetragonoloba).[71]
Several species of Fabaceae are used to produce dyes. The heartwood of logwood,Haematoxylon campechianum, is used to produce red and purple dyes. Thehistological stain calledhaematoxylin is produced from this species. The wood of the Brazilwood tree (Caesalpinia echinata) is also used to produce a red or purple dye. The Madras thorn (Pithecellobium dulce) has reddish fruit that are used to produce a yellow dye.[72] Indigo dye is extracted from the indigo plantIndigofera tinctoria that is native to Asia. In Central and South America dyes are produced from two species in the same genus: indigo andMaya blue fromIndigofera suffruticosa and Natal indigo fromIndigofera arrecta. Yellow dyes are extracted fromButea monosperma, commonly called flame of the forest and from dyer's greenweed, (Genista tinctoria).[73]
Legumes have been used as ornamental plants throughout the world for many centuries. Their vast diversity of heights, shapes, foliage and flower colour means that this family is commonly used in the design and planting of everything from small gardens to large parks.[18] The following is a list of the main ornamental legume species, listed by subfamily.
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^Gilbert Vargas Ulate. 1997. Geografía turística de Costa Rica. EUNED, 180 p.ISBN9977-64-900-6, 9789977649009.
^"Lei Nº 6.607, de 7 de dezembro de 1978. O Presidente da República, faço saber que o Congresso Nacional decreta e eu sanciono a seguinte Lei:Art. 1º- É declarada Árvore Nacional a leguminosa denominada Pau-Brasil (Caesalpinia echinata, Lam), cuja festa será comemorada, anualmente, quando o Ministério da Educação e Cultura promoverá campanha elucidativa sobre a relevância daquela espécie vegetal na História do Brasil."
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