The Fabaceae or Leguminosae, commonly known as the legume, pea, or bean family, is a large and economically important family of flowering plants. The group is the third largest land plant family, behind only the Orchidaceae and Asteraceae, with 730 genera and over 19,400 species. The largest genera are Astragalus (over 2,400 species), Acacia (over 950 species), Indigofera (around 700 species), Crotalaria (around 700 species), and Mimosa (around 500 species).
Plants of this family are found throughout the world, growing in many different environments and climates. A number are important agricultural 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 (licorice), which are among the best known members of Fabaceae.
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 typical fruit of these plants, which are called legumes.
The Fabaceae has an essentially worldwide distribution, being found everywhere except Antarctica and the high arctic.
- Mimosoideae: 80 genera and 3,200 species. Mostly tropical and warm temperate Asia and America. Mimosa, Acacia.
- Caesalpinioideae: 170 genera and 2,000 species, cosmopolitan. Caesalpinia, Senna, Bauhinia, Amherstia.
- Faboideae: 470 genera and 14,000 species, cosmopolitan. Astragalus, Lupinus.
These three subfamilies have been alternatively treated at the family level, as in the Cronquist and Dahlgren systems. However, this choice has not been supported by late 20th and early 21st century evidence, which has shown the Caesalpinioideae to be paraphyletic and the Fabaceae sensu lato to be monophyletic. While the Mimosoideae and the Faboideae are largely monophyletic, the Caesalpinioideae appear to be paraphyletic and the tribe Cercideae is probably sister to the rest of the family. Moreover, there are a number of genera whose placement into the Caesalpinioideae is not always agreed on (e.g. Dimorphandra).
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.
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.
The leaves are usually alternate and compound. Most often they are even- or odd-pinnately compound (e.g. Caragana and Robinia respectively), often trifoliate (e.g. Trifolium, Medicago) and rarely palmately compound (e.g. Lupinus), in the Mimosoideae and the Caesalpinioideae commonly bipinnate (e.g. Acacia, Mimosa). They always have stipules, which can be leaf-like (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 often have wrinkled pulvini to permit nastic movements. In some species, leaflets have evolved into tendrils (e.g. Vicia).
Many species have leaves with structures that attract ants that 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.g. Vicia sativa). In some Acacia, the modified hollow stipules are inhabited by ants.
The flowers always have five generally fused sepals and five free petals. They are generally hermaphrodite, 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.e. they are pollinated by insects), and the flowers are usually showy to attract pollinators.
In the Caesalpinioideae, the flowers are often zygomorphic, as in Cercis, or nearly symmetrical with five equal petals 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. The petals are small and the stamens, which can be more than just ten, have long coloured filaments, which are the most showy 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, 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.
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.
It has been suggested, based on fossil and phylogenetic evidence, that legumes originally evolved in arid and/or semi-arid regions along the Tethys seaway during the early Tertiary. However, others contend that Africa (or even the Americas) cannot yet be ruled out as the origin of the family.
One of the key features of Fabaceae is that some members are able to nodulate. The current hypothesis about the evolution of the genes needed for nodulation is that they were recruited from other pathways after a polyploidy event. 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. 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. The enzymes needed to reduce nitrogen, nitrogenases, are 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. These three genetic pathways are believed to be part of a gene duplication event then recruited to work in nodulation.
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 for supplementing protein where there was not enough meat.
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. Some legume species perform hydraulic lift, which makes them ideal for intercropping.
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 alfalfa, clover, vetch, and Arachis, are sown in pasture and grazed by livestock. Other forage 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 stock feed.
Grain legumes are cultivated for their seeds, and are also called pulses. The seeds are used for human 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. and trees such as carob, mesquite and tamarind.
Bloom legume species include species such as lupin, which are farmed commercially for their blooms as well as being popular in gardens worldwide. Laburnum, Robinia, Gleditsia, Acacia, Mimosa, and Delonix are ornamental trees and shrubs.
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.
The genera included in this family can be viewed on the following three pages:
Acacia baileyana (Wattle)
Dichrostachys cinerea Sickle Bush
Tendrils of Lathyrus odoratus (Sweet pea)
Inflorescence of Lupinus arboreus (Yellow bush lupin)
Pisum sativum (Peas); note the leaf-like stipules
- ^ a b c Wojciechowski, M. F.; Lavin, M.; Sanderson, M. J. (2004). "A phylogeny of legumes (Leguminosae) based on analysis of the plastid matK gene resolves many well-supported subclades within the family". American Journal of Botany 91 (11): 1846–62. doi:10.3732/ajb.91.11.1846. PMID 21652332.
- ^ "GRIN-CA". http://pgrc3.agr.ca/cgi-bin/npgs/html/family.pl?440. Retrieved 2002-09-01.
- ^ a b Schrire, B. D.; Lewis, G. P.; Lavin, M. (2005). "Biogeography of the Leguminosae". In Lewis, G; Schrire, G.; Mackinder, B. et al.. Legumes of the world. Kew, England: Royal Botanic Gardens. pp. 21–54. ISBN 1900347806. http://www.kewbooks.com/asps/ShowDetails.asp?id=506.
- ^ a b Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website Version 9, June 2008 Mobot.org
- ^ Wiktionary. "Faba". Searched November, 2011. http://en.wiktionary.org/wiki/faba
- ^ International Code of Botanical Nomenclature Art. 18.5 (Vienna Code)
- ^ NOTE: The subfamilial name Papilionoideae for Faboideae is approved by the International Code of Botanical Nomenclature, as is 'Leguminosae' for the Fabaceae sensu lato.
- ^ a b c Martin F. Wojciechowski, Johanna Mahn, and Bruce Jones (2006). "Fabaceae". The Tree of Life Web Project. http://tolweb.org/Fabaceae/21093/2006.06.14.
- ^ Hélène L. Citerne, R. Toby Pennington, and Quentin C. B. Cronk (August 8, 2006). "An apparent reversal in floral symmetry in the legume Cadia is a homeotic transformation". PNAS 103 (32): 12017–12020. doi:10.1073/pnas.0600986103. PMC 1567690. PMID 16880394. http://www.pnas.org/content/103/32/12017.full
- ^ Schrire, B. D.; Lavin, M.; Lewis, G. P. (2005). "Global distribution patterns of the Leguminosae: insights from recent phylogenies". In Friis, I; Balslev, H.. Plant diversity and complexity patterns: local, regional and global dimensions. Biologiske Skrifter. 55. Viborg, Denmark: Special-Trykkeriet Viborg A/S. pp. 375–422. ISBN 8773043044.
- ^ Pan, Aaron D.; Jacobs, Bonnie F.; Herendeen, Patrick S. (2010). "Detarieae sensu lato (Fabaceae) from the Late Oligocene (27.23 Ma) Guang River flora of north-western Ethiopia". Botanical Journal of the Linnean Society 163: 44. doi:10.1111/j.1095-8339.2010.01044.x.
- ^ Doyle, J. J.; Luckow, MA (2003). "The Rest of the Iceberg. Legume Diversity and Evolution in a Phylogenetic Context". Plant Physiology 131 (3): 900–10. doi:10.1104/pp.102.018150. PMC 1540290. PMID 12644643. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1540290.
- ^ Yokota, Keisuke; Hayashi, Makoto (2011). "Function and evolution of nodulation genes in legumes". Cellular and Molecular Life Sciences 68 (8): 1341–51. doi:10.1007/s00018-011-0651-4. PMID 21380559.
- ^ Markmann, Katharina; Giczey, Gábor; Parniske, Martin (2008). "Functional Adaptation of a Plant Receptor- Kinase Paved the Way for the Evolution of Intracellular Root Symbioses with Bacteria". PLoS Biology 6 (3): e68. doi:10.1371/journal.pbio.0060068. PMC 2270324. PMID 18318603. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2270324.
- ^ Rodríguez-Llorente, Ignacio D.; Pérez-Hormaeche, Javier; Mounadi, Kaoutar El; Dary, Mohammed; Caviedes, Miguel A.; Cosson, Viviane; Kondorosi, Adam; Ratet, Pascal et al. (2004). "From pollen tubes to infection threads: Recruitment ofMedicagofloral pectic genes for symbiosis". The Plant Journal 39 (4): 587–98. doi:10.1111/j.1365-313X.2004.02155.x. PMID 15272876.
- ^ Downie, J. Allan (2005). "Legume Haemoglobins: Symbiotic Nitrogen Fixation Needs Bloody Nodules". Current Biology 15 (6): R196–8. doi:10.1016/j.cub.2005.03.007. PMID 15797009.
- ^ Sprent, Janet I. (2009). Legume Nodulation: A Global Perspective. Ames, Iowa: Wiley-Blackwell. p. 12. ISBN 1405181753. http://www.sprentland.com/index.php?pr=Janet. Preview available at Google Books.
- ^ The gene bank and breeding of grain legumes (lupine, vetch, soya and beah) / B.S. Kurlovich and S.I. Repyev (Eds.), - St. Petersburg, The N.I. Vavilov Institute of Plant Industry, 1995, 438p. - (Theoretical basis of plant breeding. V.111)