Allium

Allium
Allium sativum[1]
Scientific classification
Kingdom: Plantae
Clade: Angiosperms
Clade: Monocots
Order: Asparagales
Family: Amaryllidaceae
Subfamily: Allioideae
Tribe: Allieae
Genus: Allium
L.
Type species
Allium sativum
L.
Subgenera

see text

Synonyms[2]
Allium flavum (yellow) and Allium carinatum (purple)

Allium is a genus of monocotyledonous flowering plants that includes the cultivated onion, garlic, scallion, shallot and leek as well as chives and hundreds of other wild species.

The generic name Allium is the Latin word for garlic,[3] and Linnaeus first described the genus Allium in 1753. Some sources refer to Greek αλεω (aleo, to avoid) by reason of the smell of garlic.[4] The cooking and consumption of parts of the plants is due to the large variety of flavours and textures of the species. Various Allium have been cultivated from the earliest times and about a dozen species are economically important as crops, or garden vegetables, and an increasing number of species are important as ornamental plants.[4][5] The inclusion of a species to the genus Allium is taxonomically difficult and species boundaries are unclear. Estimates of the number of species have been as low as 260,[6] and as high as 979.[7] Most authorities accept about 750 species.[8] The type species for the genus is Allium sativum.[9]

Allium species occur in temperate climates of the Northern Hemisphere, except for a few species occurring in Chile (such as A. juncifolium), Brazil (A. sellovianum), and tropical Africa (A. spathaceum). They vary in height between 5 cm and 150 cm. The flowers form an umbel at the top of a leafless stalk. The bulbs vary in size between species, from small (around 2–3 mm in diameter) to rather large (8–10 cm). Some species (such as Welsh onion A. fistulosum) develop thickened leaf-bases rather than forming bulbs as such.

Plants of the Allium genus produce chemical compounds (mostly derived from cysteine sulfoxides) that give them a characteristic (alliaceous) onion or garlic taste and odor.[4] Many are used as food plants, though not all members of the genus are equally flavorous. In most cases, both bulb and leaves are edible and the taste may be strong or weak, depending on the species and on ground sulfur (usually as sulfate) content.[4] In the rare occurrence of sulfur-free growth conditions, all Allium species lose their usual pungency altogether.

In the APG III classification system, Allium is placed in the family Amaryllidaceae, subfamily Allioideae (formerly the family Alliaceae).[10] In some of the older classification systems, Allium was placed in Liliaceae.[4][5][11][12][13] Molecular phylogenetic studies have shown this circumscription of Liliaceae is not monophyletic.

Allium is one of about fifty-seven genera of flowering plants with more than 500 species.[14] It is by far the largest genus in the Amaryllidaceae, and also in the Alliaceae in classification systems in which that family is recognized as separate.[6]

Description

Capsule of Allium oreophilum.

Allium species are herbaceous perennials with flowers produced on scapes. They grow from solitary or clustered tunicate bulbs and many have an onion odor and taste. Plants are perennialized by bulbs that reform annually from the base of the old bulb, or are produced on the ends of rhizomes or, in a few species, at the ends of stolons. A small number of species have tuberous roots. The bulbs' outer coats are commonly brown or grey, with a smooth texture, and are fibrous, or with cellular reticulation. The inner coats of the bulbs are membranous.

Many alliums have basal leaves that commonly wither away from the tips downward before or while the plants flower, but some species have persistent foliage. Plants produce from one to 12 leaves, most species having linear, channeled or flat leaf blades. The leaf blades are straight or variously coiled, but some species have broad leaves, including A. victorialis and A. tricoccum. The leaves are sessile, and very rarely narrowed into a petiole.

The flowers are erect or in some species pendent, having six petal-like tepals produced in two whorls. The flowers have one style and six epipetalous stamens; the anthers and pollen can vary in color depending on the species. The ovaries are superior, and three-lobed with three locules.

The fruits are capsules that open longitudinally along the capsule wall between the partitions of the locule.[15][16] The seeds are black, and have a rounded shape.

The terete or flattened flowering scapes are normally persistent. The inflorescences are umbels, in which the outside flowers bloom first and flowering progresses to the inside. Some species produce bulbils within the umbels, and in some species, such as Allium paradoxum, the bulbils replace some or all the flowers. The umbels are subtended by noticeable spathe bracts, which are commonly fused and normally have around three veins.

Some bulbous alliums increase by forming little bulbs or "offsets" around the old one, as well as by seed. Several species can form many bulbils in the flowerhead; in the so-called "tree onion" or Egyptian onion (A. × proliferum) the bulbils are few, but large enough to be pickled.

Many of the species of Allium have been used as food items throughout their ranges. There are several poisonous species that are somewhat similar in appearance (e.g. in North America, death camas, Toxicoscordion venenosum), but none of these has the distinctive scent of onions or garlic.[17][18]

Allium ursinum (wild garlic) in a Bristol, England, woodland
Selection of cultivated alliums displayed at the BBC Gardeners' World show

Taxonomy

With over 800 species Allium, the sole genus in the Allieae tribe, is one of the largest monocotyledonous genera.,[19] but the precise taxonomy of Allium is poorly understood,[19] with incorrect descriptions being widespread. However, the genus has been shown to be monophyletic, containing three major clades, although some proposed subgenera are not.[19]

Allium includes a number of taxonomic groupings previously considered separate genera (Caloscordum Herb., Milula Prain and Nectaroscordum Lindl.) Allium spicatum had been treated by many authors as Milula spicata, the only species in the monospecific genus Milula. In 2000, it was shown to be embedded in Allium.[20]

History

When Linnaeus[21] formerly described the genus Allium in his Species Plantarum (1753), there were thirty species with this name. He placed Allium in a grouping he referred to as Hexandria monogynia (i.e. six stamens and one pistil)[22] containing 51 genera in all.[23]

Subdivision

Linnaeus originally grouped his 30 species into three alliances, e.g. Foliis caulinis planis. Since then many attempts have been made to divide the growing number of recognised species into infrageneric subgroupings, initially as sections, and then as subgenera further divided into sections.[19] The modern era of phylogenetic analysis dates to 2006 when Friesen, Fritsch and Blattner[24] described a new classification with 15 subgenera, 72 sections and about 780 species based on the nuclear ribosomal gene ITS Some of the subgenera correspond to the once separate genera (Caloscordum, Milula, Nectaroscordum) included in Gilliesieae.[19][25]

Subsequent molecular phylogenetic studies have shown the 2006 classification is a considerable improvement over previous classifications, but some of its subgenera and sections are probably not monophyletic. One of these studies focused on the subgenus Amerallium, which is strongly supported as monophyletic.[26] Another study focused on Allium ampeloprasum and its relatives within the section Allium of subgenus Allium.[8] Sampling in this study was not sufficient to test the monophyly of section Allium.

The major evolutionary lineages correspond to the three major clades. Lineage One (the oldest) with three subgenera are predominantly bulbous, the second, with five subgenera and the third with seven subgenera contain both bulbous and rhizomatous taxa.[19]

Subgenera

The three lineages and fifteen subgenera represent the classification scheme of Friesen et al. (2006)[24] and Li (2010).[19] (number of sections/number of species)

Species

Species include:

Distribution and habitat

The majority of Allium species are native to the Northern Hemisphere, being spread throughout the holarctic region, from dry subtropics to the boreal zone,[19] but predominantly in Asia. Of the latter 138 species occur in China, about a sixth of all species, representing five subgenera.[19] A few species are native to Africa and Central and South America.[15] A single known exception, Allium dregeanum occurs in the Southern Hemisphere (South Africa). There are two centres of diversity, a major one from the Mediterranean Basin to Central Asia and Pakistan, while a minor one is found in western North America.[19]

Ecology

Species grow in various conditions from dry, well-drained mineral-based soils to moist, organic soils; most grow in sunny locations, but a number also grow in forests (e.g., A. ursinum),[4] or even in swamps or water.

Various Allium species are used as food plants by the larvae of the leek moth and onion fly[4] as well as some Lepidoptera including cabbage moth, common swift moth (recorded on garlic), garden dart moth, large yellow underwing moth, nutmeg moth, setaceous Hebrew character moth, turnip moth and Schinia rosea, a moth that feeds exclusively on Allium species.

Cultivation

Many Allium species have been harvested through human history, but only about a dozen are still economically important today as crops or garden vegetables.[4][27]

Ornamental

Many Allium species and hybrids are cultivated as ornamentals.[28] These include A. cristophii and A. giganteum, which are used as border plants for their ornamental flowers, and their "architectural" qualities.[5][29] Several hybrids have been bred, or selected, with rich purple flowers. A. hollandicum 'Purple Sensation' is one of the most popular and has been given an Award of Garden Merit (H4).[30] These ornamental onions produce spherical umbels on single stalks in spring and summer, in a wide variety of sizes and colours, ranging from white (Allium 'Mont Blanc'), blue (A. caeruleum), to yellow (A. flavum) and purple (A. giganteum). By contrast, other species (such as invasive A. triquetrum and A. ursinum) can become troublesome garden weeds.[29][31] The hybrid cultivars 'Beau Regard',[32] 'Gladiator',[33] and 'Globemaster'[34] have gained the Royal Horticultural Society's Award of Garden Merit.

Toxicity

Dogs and cats are very susceptible to poisoning after the consumption of certain species.[4][35]

Uses

The genus includes many economically important species. These include onions (A. cepa), French shallots (A. oschaninii), leeks (A. ampeloprasum), scallions (various Allium species), and herbs such as garlic (A. sativum) and chives (A. schoenoprasum). Some have been used as traditional medicines.[19]

References

  1. ↑ 1793 illustration from William Woodville: "Medical botany", London, James Phillips, 1793, Vol. 3, Plate 168: Allium sativum (Garlic). Hand-coloured engraving
  2. ↑ Kew World Checklist of Selected Plant Families
  3. ↑ Quattrocchi 1999, vol. 1 p. 91.
  4. 1 2 3 4 5 6 7 8 9 Eric Block (2010). Garlic and Other Alliums: The Lore and the Science. Royal Society of Chemistry. ISBN 978-0-85404-190-9.
  5. 1 2 3 Dilys Davies (1992). Alliums: The Ornamental Onions. Timber Press. ISBN 0-88192-241-2.
  6. 1 2 Knud Rahn. 1998. "Alliaceae" pages 70-78. In: Klaus Kubitzki (editor). The Families and Genera of Vascular Plants volume III. Springer-Verlag: Berlin;Heidelberg, Germany. ISBN 978-3-540-64060-8
  7. ↑ The Plant List, for genus Allium
  8. 1 2 Hirschegger, Pablo; Jaške, Jernej; Trontelj, Peter; Bohanec, Borut (2010). "Origins of Allium ampeloprasum horticultural groups and a molecular phylogeny of the section Allium (Allium; Alliaceae)".". Molecular Phylogenetics and Evolution 54 (2): 488–497. doi:10.1016/j.ympev.2009.08.030.
  9. ↑ Allium In: Index Nominum Genericorum. In: Regnum Vegetabile (see External links below).
  10. ↑ Chase, M.W.; Reveal, J.L. & Fay, M.F. (2009), "A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae", Botanical Journal of the Linnean Society 161 (2): 132–136, doi:10.1111/j.1095-8339.2009.00999.x
  11. ↑ James L. Brewster, "Onions and Other Alliums" (Wallingford: CABI Publishing, 2008)
  12. ↑ Haim D. Rabinowitch, Leslie Currah, "Allium Crop Sciences: Recent Advances" (Wallingford: CABI Publishing, 2002)
  13. ↑ Penny Woodward, "Garlic and Friends: The History, Growth and Use of Edible Alliums" (South Melbourne: Hyland House, 1996)
  14. ↑ Frodin, David G. (2004). "History and concepts of big plant genera". Taxon 53 (3): 753–776. doi:10.2307/4135449.
  15. 1 2 "Allium in Flora of China @". Efloras.org. Retrieved 2012-12-11.
  16. ↑ "Allium in Flora of North America @". Efloras.org. Retrieved 2012-12-11.
  17. ↑ Peterson, R.P. 1982. A Field Guide to Edible Wild Plants: Eastern and central North America. Houghton Mifflin, Boston.
  18. ↑ Gibbons, E. 1962. Stalking the wild asparagus. David McKay, New York.
  19. 1 2 3 4 5 6 7 8 9 10 11 Li et al. 2010.
  20. ↑ Friesen et al 2000.
  21. ↑ Linnaeus 1753, Allium I pp. 294–301.
  22. ↑ Linnaeus Sexual System 2015.
  23. ↑ Linnaeus 1753, Hexandria monogynia I pp. 285–332.
  24. 1 2 Friesen, Fritsch & Blattner 2006.
  25. ↑ Sykorova 2006.
  26. ↑ Nguyen et al 2008.
  27. ↑ Gualtiero Simonetti (1990). Stanley Schuler, ed. Simon & Schuster's Guide to Herbs and Spices. Simon & Schuster, Inc. ISBN 0-671-73489-X.
  28. ↑ Anthony Huxley, Mark Griffiths, and Margot Levy (1992). The New Royal Horticultural Society Dictionary of Gardening. The Macmillan Press,Limited: London. The Stockton Press: New York. ISBN 978-0-333-47494-5 (set).
  29. 1 2 Brickell, Christopher (Editor-in-chief),The Royal Horticultural Society A–Z Encyclopedia of Garden Plants, p.95, Dorling Kindersley, London, 1996, ISBN 0-7513-0303-8
  30. ↑ RHS Plant Finder 2009–2010, p68, Dorling Kindersley, London, 2009, ISBN 978-1-4053-4176-9
  31. ↑ Lloyd, Christopher & Rice, Graham, (1991) Garden Flowers From Seed, p45, Viking, ISBN 0-670-82455-0
  32. ↑ "RHS Plant Selector Allium 'Beau Regard' AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2012-12-11.
  33. ↑ "RHS Plant Selector Allium 'Gladiator' AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2012-12-11.
  34. ↑ "RHS Plant Selector Allium 'Globemaster' AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2012-12-11.
  35. ↑ Cope RB. Toxicology Brief: Allium species poisoning in dogs and cats. Veterinary Medicine 2005

Bibliography

Books

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External links

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