Fabaceae
Fabaceae Temporal range:
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Kudzu (Pueraria lobata) | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Clade: | Rosids |
Order: | Fabales |
Family: | Fabaceae nom. cons.).[3]
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Type genus | |
Faba (now included in Vicia) Mill.
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Subfamilies[4] | |
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Diversity | |
730 genera and 19,400 species | |
Fabaceae distribution map. Legumes are found in four major biomes: tropical forest, temperate, grass, and succulent.[5] | |
Synonyms | |
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The Fabaceae (
The five largest genera 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 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 single monophyletic 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 recent phylogenetic studies based on DNA sequences.[14][15][16] 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 order Fabales.[17]
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.[18]
The family Fabaceae includes a number of plants that are common in agriculture including Glycine max (
Etymology
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,[6] and refers to the fruit of these plants, which are called legumes.
Description
Fabaceae range in habit from giant
Growth habit
The Fabaceae 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. 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.[3][9]
Leaves
The leaves are usually
Many species have leaves with structures that attract
Roots
Many Fabaceae host bacteria in their roots within structures called root nodules. These bacteria, known as rhizobia, have the ability to take nitrogen gas (N2) out of the air and convert it to a form of nitrogen that is usable to the host plant ( NO3− or NH3 ). 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. Members of the Phaseoleae genus Apios form tubers, which can be edible.[19]
Flowers
The
In the
In the
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. 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 the CYCLOIDEA (CYC)/DICHOTOMA (DICH) family are expressed in the upper (also called dorsal or adaxial) petal; in some species, such as Cadia, these genes are expressed throughout the flower, producing a radially symmetrical flower.[20]
Fruit
The ovary most typically develops into a
Physiology and biochemistry
The Fabaceae are rarely
Evolution, phylogeny and taxonomy
Evolution
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 families
The Fabaceae have an abundant and diverse
It has been suggested, based on fossil and phylogenetic evidence, that legumes originally evolved in arid and/or semi-arid regions along the
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.
Phylogeny and taxonomy
Phylogeny
The phylogeny of the legumes has been the object of many studies by research groups from around the world. 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 consistently recovered as monophyletic.[46] The studies further confirmed that the traditional subfamilies Mimosoideae and Papilionoideae were each monophyletic 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] 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][56] The exact branching order of the different subfamilies is still unresolved.[57]
Fabales |
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Taxonomy
The Fabaceae are placed in the order Fabales according to most taxonomic systems, including the APG III system.[2] The family now includes six subfamilies:[4]
- Cercidoideae: 12 genera and ~335 species. Mainly tropical. Bauhinia, Cercis.
- 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).
- Pisum.
Ecology
Distribution and habitat
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]
Biological nitrogen fixation
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 in
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 of Parasponia, 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 the meristem retained), determinate (without meristem) and the type included in Aeschynomene. 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 two monophyletic 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 genus Acacia, 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 for Acacia senegal, which forms both rapidly and slow growing rhizobial nodules.
Chemical ecology
A large number of species within many genera of leguminous plants, e.g.
Economic and cultural importance
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]
Food and forage
The
, 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 crop rotation to replenish soil that has been depleted of nitrogen. Legume seeds and foliage have a comparatively higher protein 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 perform hydraulic lift, which makes them ideal for intercropping.[66]
Farmed legumes can belong to numerous classes, including forage, 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, like
Grain legumes are cultivated for their
.Lathyrus tuberosus, once extensively cultivated in Europe, forms tubers used for human consumption.[68][69]
Bloom legume species include species such as
Industrial farmed legumes include Indigofera, cultivated for the production of indigo, Acacia, for gum 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 soil to exploit the high nitrogen levels found in most legumes. Numerous legumes are farmed for this purpose, including Leucaena, Cyamopsis and Sesbania.
Various legume species are farmed for timber production worldwide, including numerous
Melliferous plants offer
Industrial uses
Natural gums
Dyes
Several species of Fabaceae are used to produce dyes. The heartwood of logwood,
Ornamentals
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.
- Subfamily Caesalpinioideae:
- Subfamily Mimosoideae: Paraserianthes lophantha, Prosopis chilensis.[74]
- Subfamily Faboideae: Tipuana tipu, Wisteria sinensis.[74]
Emblematic Fabaceae
- The Cockspur Coral Tree (
- The Elephant ear tree (Enterolobium cyclocarpum) is the national tree of Costa Rica, by Executive Order of 31 August 1959.[76]
- The Brazilwood tree (
- The Golden wattle Acacia pycnantha is Australia's national flower.[78]
- The Hong Kong Orchid tree
Image gallery
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Acacia baileyana (Wattle)
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Loments of Alysicarpus vaginalis
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Calliandra emarginata
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Cassia leptophylla tree
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Dichrostachys cinerea Sickle Bush
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Delonix regia tree
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Tendrils ofLathyrus odoratus(Sweet pea)
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Inflorescence of Lupinus arboreus (Yellow bush lupin)
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Pisum sativum(Peas); note the leaf-like stipules
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Kashubian vetch – Kashubia
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Cytisus scoparius (Scotch broom)
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Senna pendula (Easter cassia)
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Hosackia stipularis (Stipulate Lotus)
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Lupinus manus (Sky Lupine)
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Vigna caracalla(snail vine) flowers
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Lupinus succulentus (Arroyo Lupine Succulent)
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Lupinus stiversii (Harlequin Lupine)
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Virgilia oroboides(Cape lilac) mauve flowers
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