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Lupinus albus L.

Protologue
Sp. pl. 2: 721 (1753).
Family
Papilionaceae (Leguminosae - Papilionoideae, Fabaceae)
Chromosome number
2n = 50
Vernacular names
White lupin, Egyptian lupin (En). Lupin blanc, lupin (Fr). Tremoceiro, tremoceiro branco, tremoceiro da Beira, tremoço (Po).
Origin and geographic distribution
White lupin originates from south-eastern Europe and western Asia where wild types still occur. It is known to have been cultivated since ancient times in Greece, Italy, Egypt and Cyprus. The importance of white lupin has fluctuated often during the history of its cultivation; at present it has almost disappeared in central Europe, while it is becoming more widely grown in the Americas. Today it is a traditional minor pulse crop, grown around the Mediterranean and the Black Sea, and in the Nile valley, extending to Sudan and Ethiopia. It is also occasionally grown elsewhere, e.g. in Kenya, Tanzania, Zimbabwe, South Africa, Mauritius, United States and South America (mainly Brazil and Chile).
Uses
White lupin is traditionally cultivated for human consumption, green manuring and as forage. Before consumption, seeds are first soaked for 1–3 days in running water to remove the bitter, toxic alkaloids, then cooked and eaten as a pulse or pickled in brine and served as a snack. In Ethiopia a high-quality spirit (‘araki’) is distilled from fermented seeds. In general, consumption of white lupin seeds is restricted to low-income classes and to times of drought, because of their bitter taste. Modern sweet cultivars have very low alkaloid contents, and their seeds do not require laborious detoxification; they are a promising nutritive pulse and can be used as a rich additive for human food and livestock feed products.
White lupin plants are fed to livestock as fresh or dry fodder. In southern Europe it is a traditional green manure crop in vineyards and olive plantations. White lupin is a good honey plant and an attractive annual ornamental. In traditional medicine it is used for various ailments, e.g. as an anthelmintic, carminative, deobstruent, diuretic and pectoral. Lupin meal mixed with honey or vinegar is used as a cure for worms, while infusions or poultices are applied to treat boils and skin complaints. Burning seeds are used as an insect repellant.
Production and international trade
No specific statistics are available for Lupinus albus. About 2 million ha are cultivated with lupin (all species) worldwide, of which 60% is mainly for seed production and 40% for forage and green manure. The major producer of lupin seed is Australia, with about 1.4 million t/year from 1.2 million ha in the early 1990s, mainly Lupinus angustifolius L. for livestock feed.
Properties
Mature, raw Lupinus albus seeds contain per 100 g edible portion: water 10.4 g, energy 1552 kJ (371 kcal), protein 36.2 g, fat 9.7 g, carbohydrate 40.4 g, Ca 176 mg, Mg 198 mg, P 440 mg, Fe 4.4 mg, Zn 4.8 mg, vitamin A 23 IU, thiamin 0.64 mg, riboflavin 0.22 mg, niacin 2.2 mg, vitamin B6 0.36 mg, folate 355 μg and ascorbic acid 4.8 mg. The essential amino-acid composition per 100 g edible portion is: tryptophan 289 mg, lysine 1933 mg, methionine 255 mg, phenylalanine 1435 mg, threonine 1331 mg, valine 1510 mg, leucine 2743 mg and isoleucine 1615 mg. The principal fatty acids are per 100 g edible portion: oleic acid 3558 mg, linoleic acid 1995 mg, palmitic acid 742 mg, linolenic acid 446 mg, stearic acid 316 mg and eicosenoic acid 255 mg (USDA, 2005). The seed coat makes up about 15% of the seed weight. The net protein utilization for humans is 77% and the protein fraction is low in lysine and methionine. The levels of antinutritional compounds such as condensed tannins and trypsin inhibitors are lower than in other pulses. Suspensions of ground seed of white lupin have shown hypoglycaemic effects in rats.
The major alkaloids of white lupin are lupanine, 13-hydroxylupanine and sparteine. The pharmacological effects of these alkaloids are that they block ganglionic transmission, decrease cardiac contractility and contract uterine smooth muscle. ‘Sweet lupin’ is defined as having less than 200 mg alkaloids/kg; it can be consumed without special precautions. In bitter cultivars the alkaloids, which are water-soluble, can be soaked out from seeds in running water. Processing techniques such as sprouting and fermentation into tempeh also strongly reduce the alkaloid content. Best control is achieved by chemical extraction, which currently is not economically feasible.
When white lupin is fed as dried forage, lupinosis can occur. This disease is caused by the ingestion of toxins, known as phomopsins, produced by the fungus Diaporthe toxica that colonizes lupin plants. It is primarily a disease of sheep but can also occur in other livestock and is characterized by severe liver damage, which results in loss of appetite and condition, lethargy, jaundice and often death. Most of the problem can be solved by using Diaporthe -resistant cultivars such as ‘Kiev’ and ‘Ultra’; where other cultivars are used, overfeeding must be avoided.
Botany
Annual, erect, branched, bushy, short-hairy herb up to 120 cm tall, with a strong taproot. Leaves alternate, digitately compound with 5–9 leaflets; stipules linear to narrowly triangular, up to 1 cm adnate to the base of the petiole; petiole 3.5–7(–12) cm long; leaflets obovate, 2–6 cm × 0.5–2 cm, cuneate at base, rounded and mucronate at apex, nearly glabrous above, hairy beneath. Inflorescence a terminal false raceme 3–30 cm long, many-flowered, lower flowers alternate, upper ones in whorls; peduncle short or absent. Flowers bisexual, papilionaceous; pedicel 1–2 mm long; calyx 8–14 mm long, densely hairy outside, tube c. 4 mm long, 2-lipped, upper lip entire, lower lip entire or slightly 3-toothed; corolla white to violet-blue, standard obovate, 15–18 mm × 8–12 mm, margins partly reflexed, wings obovate, 13–17 mm × 6–10 mm, keel ladle-shaped, 12–15 mm × 4 mm, beaked; stamens 10, all joined into a tube; ovary superior, 1-celled, style c. 7.5 mm long with a ring of small hairs below the stigma. Fruit a narrowly oblong, laterally compressed pod 6–15 cm × 1–2 cm, bulging over the seeds, shortly hairy but glabrescent, yellow, 3–6-seeded. Seeds rectangular or square with rounded corners, laterally compressed, 7–16 mm × 6–12 mm × 2–5 mm, more or less smooth, white variably tinged salmon pink or mottled dark brown. Seedling with epigeal germination.
Lupinus comprises about 200 species, mostly American; only 12 species are native to the Old World. In tropical Africa 3 native or naturalized species occur and another 9 species have been introduced. Many Lupinus spp. are ornamental garden plants, and 4 species are cultivated on a larger scale as agricultural crops.
Lupinus albus represents a crop-weed complex with wide variability in wild and cultivated types. Both types have been classified as subspecies although for the cultivated types a classification into cultivar groups and cultivars would be more appropriate. The wild type is distinguished as subsp. graecus (Boiss. & Spruner) Franco & P.Silva (synonym: Lupinus graecus Boiss. & Spruner) and can be found in south-eastern Europe and western Asia. The corolla is dark violet-blue, pods are 6–8 cm × 1–1.5 cm and shatter seeds at maturity, and seeds are small, 7–10 mm × 6–8 mm × 2–3 mm, mottled dark brown with impermeable seed coat. The cultivated types are distinguished as subsp. albus (synonym: Lupinus termis Forssk.), with corolla white, pods 9–15 cm × 1.5–2 cm with non-shattering seeds at maturity, and seeds 10–14 mm × 8–12 mm × 3–5 mm, pinkish white or white with permeable seed coat. In Ethiopia 2 types of cultivated plants are found: a large-seeded type as grown in Egypt and Sudan, but also a small-seeded type with small leaves. In northern parts of the distribution area of white lupin, in South Africa and in the Americas mostly sweet (low-alkaloid) modern cultivars are grown, whereas in the Mediterranean region and eastern Africa bitter landraces prevail.
White lupin is mainly self-pollinating, but 5–10% outcrossing can occur. White lupin nodulates effectively with Bradyrhizobium bacteria. Atmospheric nitrogen fixation rates up to 400 kg N per ha have been observed in Europe and Australia.
Ecology
Wild white lupin prefers disturbed sites and poor soils, where there is less competition from other species. White lupin is usually grown at mean monthly temperatures during the growing season of 15–25°C, the optimum being 18–24°C. Higher temperatures and moisture stress hinder flowering and pod setting. White lupin is cold-tolerant, but temperatures of –6 to –8°C are harmful at germination, temperatures of –3 to –5°C at flowering. A rainfall of 400–1000 mm during the growing period is optimal for yield. Lupin species are drought-tolerant due to their deep roots, but are sensitive to moisture deficiency during the reproductive period.
White lupin is adapted to well-drained, mildly acid or neutral soils of light to medium texture, with pH 4.5–7.5. Growth is hampered on heavy clay and waterlogged soils, while calcareous or alkaline soils induce chlorosis and reduce growth, frequently precluding cultivation. The accepted maximum soil level of CaCO3 is 3–5 g/100 g. Some cultivars of white lupin are more tolerant to soil salinity and heavy soils than most other crops.
In Ethiopia white lupin is grown at 1500–3000 m altitude, on soils too poor for a good faba bean crop.
Management
White lupin is propagated by seed. The 1000-seed weight ranges from 70 g in some Kenyan populations to more than 1 kg in modern seed cultivars. Seed can easily be stored for 2–4 years under normal conditions; longer storage is possible at lower temperatures. In areas with mild winters such as the Mediterranean region, seed is sown broadcast or drilled from mid September to late October. The seed rate is 50–180 kg/ha, the seeding depth 2.5–5 cm. In Ethiopia seed is sown in the main rainy season (July–September). White lupin is often grown intercropped with cereals or forage legumes, or in rotation with cereals. Weed control is essential until a closed canopy is formed. White lupin is sensitive to P deficiency, but its roots can make more P available through acidification of the rhizosphere, a property from which also associated crops benefit. Wheat intercropped with white lupin has access to a larger pool of P, Mn and N than sole-cropped wheat. Inoculation of the soil with Bradyrhizobium bacteria is beneficial, giving up to a 5-fold increase in yield and a higher protein content of the seed. A well-known commercially available inoculant strain is the Australian WU425.
The major diseases of white lupin are root rot and brown leaf spot caused by Pleiochaeta setosa, anthracnose (Colletotrichum acutatum), resulting in early plant death through stem breakage, and rust (Uromyces lupinicolus). Sources of resistance to anthracnose have been found in Ethiopian landraces, but resistant cultivars are not yet available. Bean Yellow Mosaic Virus (BYMV) is the major virus disease; it is transmitted by aphids and by seed. No sources of resistance have yet been identified. White lupin is immune to Cucumber Mosaic Virus (CMV), a major disease of other Lupinus spp. Major pests are the bean seedling maggot (Delia platura, synonym: Phorbia platura) causing seedlings to wilt and die, beetle and moth larvae (e.g. Agriotes and Agrotis spp. killing seedlings), slugs (attacking leaves), thrips (Frankliniella spp., attacking flower buds and leaves), mirid bugs (attacking young pods) and budworms (e.g. Helicoverpa armigera feeding on pod and seed). In Ethiopia harvesting is in December. Seed yields are 500–4000 kg/ha.
Genetic resources and breeding
Major germplasm collections of white lupin are available in France (INRA, Station d’Amélioration des Plantes Fourragères, Lusignan, 1400 accessions), the United Kingdom (University of Reading, Reading, 1100 accessions), Australia (Western Australian Department of Agriculture, South Perth, 890 accessions) and Spain (Servicio de Investigación y Desarrollo Tecnológico, Guadajira, 690 accessions). In tropical Africa small collections are held in Ethiopia (International Livestock Research Institute (ILRI), Addis Ababa, Ethiopia, 25 accessions) and Kenya (National Genebank of Kenya, Crop Plant Genetic Resources Centre, KARI, Kikuyu, 20 accessions).
Major objectives in breeding of white lupin are to produce rapid-growing, alkaloid-free, disease-resistant (particularly against anthracnose), high-yielding, high-pH-tolerant, frost-tolerant dwarf cultivars, well adapted to local ecological conditions. It appears that bitter cultivars tolerate cold and disease stress better than sweet ones. The level of cross-pollination may limit the relevance of sweet white lupin cultivars in regions where also bitter weedy or cultivated types are present, because pollen of the latter would reintroduce the bitter character in farm-saved sowing seed. Sweet cultivars, however, are a prerequisite for any further breeding advancement. Commercial cultivars are pure lines bred through pedigree selection. Some well-known cultivars of white lupin are: ‘Eldo’, ‘Kiev’, ‘Multolupa’ and ‘Ultra’. From Ethiopia ‘Bahar Dar’ is known.
Prospects
White lupin is a promising annual legume crop for human consumption, green manuring and forage. The composition of the seed and especially the high protein content makes white lupin highly suitable for livestock diets as a protein-rich product in intensive farming systems. The low level of antinutritional factors facilitates a direct on-farm use of white lupin in self-sustained systems. Since it often can grow on land unsuitable for other crops (too saline, heavy, acid or poor), the development of cultivars adapted to tropical African conditions is highly recommended. Much can be learned from the excellent results obtained with Lupinus angustifolius in Australia.
Major references
• Cowling, W.A., Buirchell, B.J. & Tapia, M.E., 1998. Lupin. Lupinus L. Promoting the conservation and use of underutilized and neglected crops. 23. Institute of Plant Genetics and Crop Plant Research (IPK), Gatersleben, Germany / International Plant Genetic Resources Institute (IPGRI), Rome, Italy. 105 pp.
• Gladstones, J.S., Atkins, C.A. & Hamblin, J. (Editors), 1998. Lupins as crop plants: biology, production and utilization. CAB International, Oxon, United Kingdom. 465 pp.
• Huyghe, C., 1997. White lupin (Lupinus albus L.). Field Crops Research 53: 147–160.
• van Santen, E., Wink, M., Weissmann, S. & Römer, P. (Editors), 2000. Lupin, an ancient crop for the new millennium: proceedings of the 9th international lupin conference, Klink/Müritz, Germany, 20–24 June, 1999. International Lupin Association, Canterbury, New Zealand. 481 pp.
• Westphal, E., 1974. Pulses in Ethiopia, their taxonomy and agricultural significance. Agricultural Research Reports 815. Centre for Agricultural Publishing and Documentation, Wageningen, Netherlands. 263 pp.
Other references
• al-Zaid, M.M., Hassan, M.A.M., Badir, N. & Gumaa, K.A., 1991. Evaluation of blood glucose lowering activity of three plant diet additives. International Journal of Pharmacognosy 29(2): 81 88.
• Duke, J.A., 1981. Handbook of legumes of world economic importance. Plenum Press, New York, United States, and London, United Kingdom. 345 pp.
• Hanelt, P. & Institute of Plant Genetics and Crop Plant Research (Editors), 2001. Mansfeld’s encyclopedia of agricultural and horticultural crops (except ornamentals). 1st English edition. Springer Verlag, Berlin, Germany. 3645 pp.
• Haq, N., 1993. Lupins (Lupinus species). In: Williams, J.T. (Editor). Pulses and vegetables. Chapman and Hall, London, United Kingdom. pp. 103–130.
• Hill, G.D., 1998. Diseases of lupins. In: Allen, D.J. & Lenné, J.M. (Editors). The pathology of food and pasture legumes. CAB International, Wallingford, United Kingdom. pp. 559–589.
• Kay, D.E., 1979. Food legumes. Crops and Product Digest No 3. Tropical Products Institute, London, United Kingdom. 435 pp.
• López-Bellido, L. & Fuentes, M., 1997. Lupinus L. In: Faridah Hanum, I. & van der Maesen, L.J.G. (Editors). Plant Resources of South-East Asia No 11. Auxiliary plants. Backhuys Publishers, Leiden, Netherlands. pp. 180–184.
• Thulin, M., 1989. Fabaceae (Leguminosae). In: Hedberg, I. & Edwards, S. (Editors). Flora of Ethiopia. Volume 3. Pittosporaceae to Araliaceae. The National Herbarium, Addis Ababa University, Addis Ababa, Ethiopia and Department of Systematic Botany, Uppsala University, Uppsala, Sweden. pp. 49–251.
• UC SAREP, undated. Cover crop database. [Internet] Sustainable Agriculture Research and Education Program, University of California, Davis, California, United States. http://www.sarep.ucdavis.edu/ccrop/. Accessed July 2005.
• USDA, 2005. USDA national nutrient database for standard reference, release 18. [Internet] U.S. Department of Agriculture, Agricultural Research Service, Nutrient Data Laboratory, Beltsville, Maryland, United States. http://www.nal.usda.gov/fnic/foodcomp. Accessed September 2005.
Author(s)
P.C.M. Jansen
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands


Editors
M. Brink
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
G. Belay
Ethiopian Agricultural Research Organization, Debre Zeit Center, P.O. Box 32, Debre Zeit, Ethiopia
Associate editors
J.M.J. de Wet
Department of Crop Sciences, Urbana-Champaign, Turner Hall, 1102 South Goodwin Avenue, Urbana, IL 61801, United States
O.T. Edje
Faculty of Agriculture, University of Swaziland, P.O. Luyengo, Luyengo, Swaziland
E. Westphal
Ritzema Bosweg 13, 6706 BB Wageningen, Netherlands
General editors
R.H.M.J. Lemmens
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
L.P.A. Oyen
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
Photo editor
A. de Ruijter
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands

Correct citation of this article:
Jansen, P.C.M., 2006. Lupinus albus L. In: Brink, M. & Belay, G. (Editors). PROTA 1: Cereals and pulses/Céréales et légumes secs. [CD-Rom]. PROTA, Wageningen, Netherlands.