White lupin (Lupinus albus) seeds

Datasheet

Description

Click on the "Nutritional aspects" tab for recommendations for ruminants, pigs, poultry, rabbits, horses, fish and crustaceans

Common names

Lupin, lupine, white lupin, white lupine, Egyptian lupin [English]; lupin blanc [French]; weisse lupine [German]; lupino bianco [Italian]; altramuz blanco, chocho, chorcho, entremozo, lupino blanco [Spanish]; tremoceiro, tremoceiro branco, tremoceiro da Beira, tremoço [Portuguese]; الترمس الأبيض [Arabic]; Люпин белый [Russian]; 白羽扇豆 [Chinese]

Species

Lupinus albus L. [Fabaceae]

Related feed(s)

Description

White lupin (Lupinus albus L.) is one of the 200 species of lupins, a genus of multipurpose annual legumes grown throughout the world both for their seeds used in feed and food, and for forage. Lupin seeds can be an alternative to soybean in all livestock species due to their high content in good quality protein (in the 30-40% range). Lupins also contribute to the sustainability of cropping systems (Lucas et al., 2015). The other main cultivated lupin species are the yellow lupin (Lupinus luteus), the blue lupin, or narrow leaf lupin (Lupinus angustifolius), and the pearl lupin (Lupinus mutabilis) (Jansen, 2006). White and yellow lupin seeds provide higher protein than blue lupin seeds, which may be of importance when lupins are used to feed animals (Soya UK, 2017).

Morphology

Lupinus albus is an erect, bushy, annual legume that can reach 1.2 m high, with an indeterminate growing habit. It is many branched and deeply taprooted. The root can grow 70 cm deep. The stems are coarse, branched, slightly silky. The leaves are alternate, medium sized and digitally compound, bearing 5-9 obovate leaflets, 2-6 cm long x 0.5-2 cm broad, smooth on the upper face, hairy on the lower. The inflorescence is a terminal, 3-30 cm long, false raceme that bears many flowers. The flowers are pedicellated, typically papillonaceous, white to violet in colour. The corolla is 15-18 mm x 8-12 mm, the upper lip entire and the lower slightly 3-toothed. The pods are 3-6 seeded, narrowly oblong, laterally compressed, (6-) 9-15 cm long × 1-2 cm wide, yellow in colour. White lupin seeds are large, flat, rectangular or square-shaped with rounded corners, laterally compressed and about 7-16 mm long × 6-12 mm wide × 2-5.5 mm high.The seeds are smooth and white with a salmon/pink tint or with dark brown speckles (Clark, 2014; El Bassam, 2010; Jansen, 2006). White lupin seeds do not shatter at maturity, unlike seeds from other lupin species (Clark, 2014).

Uses

White lupin provides seeds for food and feed, fodder and green manure (Jansen, 2006). The seeds of earlier, alkaloid-rich bitter varieties used to be detoxified by soaking before being cooked for food. They were consumed by low-income classes or during times of scarcity (Jansen, 2006). Modern sweet varieties do not require detoxification and are used in high-value specialties to enrich pastas, cake mixtures, cereals, and other baked goods (Clark, 2014). They can also be pickled and used for snacks.

Due to their good nutrient content (see Nutritional attributes on the "Nutritional aspects" tab), white lupin seeds are used to feed livestock and aquaculture species. The plant may also be grazed during late winter and early spring or cut for forage or silage. As a legume, the white lupin plant is used for green manuring (in Southern Europe, it is traditionnally used in vineyards and olive plantations), and for soil improvement (Jansen, 2006; Duke, 1981). White lupin is a good honey plant and an attractive annual ornamental (Jansen, 2006). White lupin seed was reported to lower blood cholesterol in humans (Fontanari et al., 2012).

Distribution

Lupins (Lupinus spp.) are separated into New world species and Old world species. Of the main cultivated species, white, blue and yellow lupins are Old world species, while pearl lupin is an American species. Lupins occupy almost all kinds of habitats from sea level to altitudes up to 4000 m (Wolko et al., 2010). White lupin originates from South-Eastern Europe and Western Asia. It was cultivated in Greece, Italy and Egypt and Cyprus 2000 years BCE (Terres Univia, 2017; Clark, 2014). It may have been first cultivated in Egypt and later domesticated in Greece, where there is a larger biodiversity and a higher number of wild-growing forms (Clark, 2014). However, the breeding of sweet cultivars did not start until the 1930s (Terres Univia, 2017).

White lupin is a winter growing legume that can be found in the wild on disturbed and poor soils where competition from other species is reduced (Clark, 2014). When cultivated it can be suitable in places too poor for faba bean (Jansen, 2006). It is mainly cultivated in Northern Europe, Russia, arid Australian plains and Andean Highlands of Chile. Spring types can be grown in the Northern Midwestern and in Northeastern USA (Clark, 2014). It is occasionnally grown in Africa, including Kenya, Ethiopia, Tanzania, Zimbabwe, South Africa, and Mauritius (Jansen, 2006).

White lupin grows from sea level up to an altitude of 740 m (Ecocrop, 2017). In Ethiopia, it is cultivated between 1500 and 3000 m altitude (Jansen, 2006). It does better in places where average monthly temperatures during the growing season range from 18°C to 24°C and where rainfall is about 400-1000 mm during the same period. White lupin is tolerant of frost but temperatures of -6°C to -8°C during germination and -3° to -5°C at flowering stage are deleterious to the crop. Moisture deficiency is harmful during the reproductive period (Jansen, 2006). White lupin does well on moderately fertile, well-drained, light or medium textured and mildly acidic or mildly calcareous soils with a pH ranging from 4.5 to 6.5 (-7.5) (Clark, 2014; Jansen, 2006). It does not do well on heavy clay, waterlogged and alkaline soils, though some cultivars have more tolerance of heavy soils and do better on saline soils than other crops (Clark, 2014). Under limiting P conditions, lupins form specialized cluster root structures and/or release P-mobilizing carboxylates that free P from insoluble forms (Lambers et al., 2012).

Worldwide production for all lupin species was about 1 million t in 2014. 63% originates from Australia, which produces mainly blue lupin, though 100,000 t of white lupins were also produced in this country in 2000 (Petterson, 2000). Other producers are Poland, Russia, Germany, Belarus, and Ukraine, totalizing 290,000 tons, mostly yellow lupins (Terres Univia, 2017). France and Mediterranean countries (Italia, Spain, Greece, Egypt) produce about 25,000 tons, likely to be mainly white lupin (FAO, 2017).

Processes

Lupin seeds may undergo several processes.

Physical treatments

Grinding and screening

Lupin seeds can be ground in a roller or hamermill and then screened through sieves. The size of the sieve may vary from 100 µm to 8 mm depending on the quality of flour that is intended: the final product is lupin meal. It has been reported that decreasing the size of particles is more effective for protein degradability of lupin meal than for those of pea meal in the rumen, for example (Carrouée et al., 2003).

Dehulling, grinding and fine screening

The dehulling of lupin seeds is done by cracking them with a plate grinder followed by mechanical removal of the hulls from the cotyledon fraction (dehulled lupins) (Brenes et al., 1993). After dehulling, the dehulled lupins can be ground and screened more or less finely. A French patent from 1990 describes a process combining grinding and screening that results in a lupin flour where 90% of the particles are smaller than 100 µm, and 60% are smaller than 30 µm (Auger et al., 1990).

Dehulling has several important effects on chemical composition. Dehulling reduces white lupin seeds fibre content by 70% (Brenes et al., 1993). Crude protein and fat contents have been reported to increase by approximately 10 to 23% and 16 to 22%, respectively, after dehulling (Nalle, 2009). In white lupin cv. Amiga the dehulling increased the fat content by about 32% (Nalle, 2009).

Heat treatments

Dry heating : Roasting, micronising

Roasting

Lupin seeds can be roasted whole (dry cooking) by direct flame to reach very quickly a temperature of 105°C (Singh et al., 1995). Roasting is reported to improve palatability (for example reduces bitterness) (Nalle, 2009)

Infra-Red heating: micronisation

Lupin seeds could be heated by infra-red rays to a temperature of 135°C for 15 minutes prior to being dehulled and ground to make dehulled micronised lupin meal (Laudadio et al., 2017). This process is called micronisation a term that remains ambiguous as it is also descirbing the process of fine grinding

Wet heating

Autoclaving

Ground seeds can also be autoclaved (steam cooking) for variable durations (10, 20 or 30 min) and variable temperatures over 100°C (Brenes et al., 1993). The effects of steam cooking is to remove heat-labile anti-nutritional factors (ANFs) provided suitable temperature is undergone by the seeds (Nalle, 2009)

Extrusion

Extrusion is a combination of mixing, shearing and wet cooking at high temperature under high pressure that can be applied to ground lupin seeds (Jang-Ho et al., 2012; Nalle, 2009). The effects of extrusion can be variable depending on the composition of the raw material, the degree of moisture and the pressure and temperatures undergone. Extrusion may affect starch in starchy materials, and fats in fatty ones. Extrusion has positive effects on ANFs removal and it improves shelf life by killing pathogens (Nalle, 2009). It has been suggested that the improvement of protein digestibility in extruded grain legumes could be attributed to the removal of ANFs (Nalle, 2009).

Forage management

Yields

The average seed yield of white lupin range from 0.5 to 4 t/ha (Jansen, 2006). In France, winter lupin was reported to yield about 4.5-5.0 t/ha and summer lupin about 3.5-4.0 t/ha (Arvalis, 2014).

Crop management

White lupin seeds intended for sowing should be tested for anthracnose infection and inoculated with Rhizobium lupini if the crop has not been cultivated in the stand before. Sowing rates are variable and depend on climatic conditions, soils, and season type. Sowing rates recommended for winter lupin are 20-30 seeds/m² in France and 50 seeds/m² in the UK (Arvalis, 2014; PGRO, 2014). Seeds should be drilled 30-40 mm deep (no deeper than 50 mm) in a good tilth and moist seedbed in non-compacting soil (PGRO, 2014). In Ethiopia, farmers usually broadcast the seeds of white lupin at 34-40 seeds/m² in July or September (Likawent Yeheyis et al., 2010).

Though white lupin is mainly sown as a sole crop, it can be also grown in association (Arvalis, 2014; PGRO, 2014; Likawent Yeheyis et al., 2010). In organic systems in Europe, white lupin can be cultivated with oats, barley and triticale, which helps controlling weeds and was shown to yield 2.3 t grains/ha (Milleville, 2014). In Ethiopia, white lupin can be cultivated in mixed stands with niger (Guizotia abyssinica), finger millet (Eleusine coracana), potatoes (Solanum tuberosum) or maize (Zea mays) (Likawent Yeheyis et al., 2010).

White lupin is a slow growing legume that needs 11 months to become mature and to be harvested. Winter white lupin should be sown as soon as possible at the end of summer to promote sufficient growth and gain enough cold resistance before winter (Arvalis, 2014). In France, spring lupins should be sown on warm, moist seedbed for better development at higher seed rates (45-60 seeds/m²), with smaller distance between rows than for winter types (Arvalis, 2014; PGRO, 2014).

White lupin is highly susceptible to anthracnose (Colletotrichum gloeosporioides or C. acutatum), a common seed-borne disease in countries with humid summers. Anthracnose spreads rapidly by wind, rain, soil-borne spores, clothing and equipment. It can cause almost total crop loss when the infection is severe and left untreated. However, anthracnose is easy to detect and can be efficiently controlled with fungicide applications (PGRO, 2014).

Harvest

In Europe, lupin is harvested from the end of July to the first half of September for winter lupins and in the second half of September for spring lupins (Terres Inovia, 2019; Métivier et al., 2013). Harvest should begin once seed moisture content is 14% and when air moisture is high to prevent pod drops or seeds shattering (e.g. early morning or at night) and to maximise grain quality (Terres Inovia, 2019; OGTR, 2013). Once harvested, lupin seeds can be either stored dried or ensiled. Lupin seeds can be crimped and ensiled right after harvesting, which enables early harvest (at high moisture content) and provides good preservation of seeds, particularly for the making of home-grown feeds (Strzelecki, 2015; Kemira, 2010).

Environmental impact

N-fixing legume and sustainable P management

Romans used to grow lupins (Lupinus spp.) as a green manuring crop for improving soils (Burtt, 1981). Lupins are N-fixing legumes that are reported to fix 300-400 kg N/ha, in Europe and Australia (Jansen, 2006). Lupins are beneficial to the next crop. Organic rye yielded 0.5 t/ha more when preceded by yellow lupin than when preceded by spring beans (PGRO, 2014).

Lupins are valuable legumes for sustainable P management: in soils depleted in available P, lupins form specialized cluster root structures and/or release P-mobilizing carboxylates that free it from insoluble forms (Lambers et al., 2012).

Soil improver and phytoremediation

Thanks to its deep taproot, the lupin plant improves soil texture and drainage. Lupins can be used for phytoremediation. Lupins may extend the phytoremediation period and increase the bioavailability of metals in polluted soils under recovery (Fumagalli et al., 2014). In mixed cultures of bioenergy crops, lupins (Lupinus albus and Lupinus angustifolius) have the ability to mobilize trace elements during and make these elements available for co-cultured species. Lupines mobilize trace elements by carboxylates and enzymes exudation and by lowering the pH value in the rhizosphere. In a comparison between white and blue lupin for trace element mobilization, white lupin was more effective than blue lupin and thus recommended for phytoremediation (Hentschel et al., 2016).

Nutritional aspects

Nutritional attributes

White lupin seed is rich in protein (30-40% as fed) and richer in fat (about 8-9% as fed) than pea and faba bean. It is richer in protein than blue lupin but poorer than yellow lupin. It is richer in fat than both species (Sujak et al., 2006). Lupins have unique carbohydrate properties characterized by small levels of starch (though the polarimetric method for starch analysis may erroneously report starch levels up to 10%), high levels of soluble and insoluble NSP, and high levels of raffinose oligosaccharides, all of which can affect the utilization of energy and the digestion of other nutrients in the diet (van Barneveld, 1999).

Potential constraints

Alkaloids

White lupins contain bitter quinolizidine alkaloids in variable amounts, from 120 mg/kg to more than 30,000 mg/kg DM. The alkaloid content was reported to raise concern in piglets over a dietary content of 20 mg/kg in Australia and over 120 mg/kg in Poland. Early varieties were rich in alkaloid and not recommended for animal feeding unless they were detoxified by soaking in water. Modern sweet varieties contain less than 200 mg/kg, have a smaller growth and are palatable to stock (Godfrey et al., 1985; Buraczewska et al., 1993).

Oligosaccharides

White lupin seeds contain from 7 to 14% α-galactosides, of which the most important is stachyose (2.8%), followed by saccharose (1.8%), raffinose (0.4%), and verbascose (0.3%) (Zdunczyk et al., 1996; Saini, 1989). The level of oligosaccharides depends on the variety and on conditions of cultivation and harvest (Pisarikova et al., 2009). α-galactosides may have deleterious effects on animal health (causing flatulence), feed intake and growth performance (Cherrière et al., 2003).

Ruminants

White lupin (Lupinus albus) is a valuable lupin species for ruminants due to its high protein and oil contents and it can be used as an alternative to soybean meal. Sweet lupin seeds can be used whole or dehulled, by feed manufacturers and for on-farm feeding (Emile et al., 1991). However, its high ruminal degradability may result in a low metabolisable protein (MP) value. The protein value of lupin can be improved through technological treatments (Poncet et al., 2003).

Use as an alternative to soybean meal

White lupin seeds can be included in diets up to 20-25% (DM) in dairy cows (Benchaar et al., 1991; Emile et al., 1991; Brunschwig et al., 2002) and growing steers (Brunschwig et al., 2002). A higher level (32%) could be used for double-muscled Belgian Blue bulls (Froidmont et al., 2008). Up to 29% (DM) could be used in growing lambs (Kung et al., 1991), 15-20% in dairy ewes and 20-30% in dairy goats (Brunschwig et al., 2002).

White lupin seeds should be ground before feeding to high-producing dairy cows, though they can be eaten unground by fattening cattle (Emile et al., 1991). In dairy cows and fattening cattle, lupin seeds resulted in performance similar to that obtained with a cereal-soybean meal concentrate. However, they tended to decrease milk protein content and had mixed effects on milk fat content (Emile et al., 1991; White et al., 2007).

Intake of dry matter and milk protein content tend to be lower for lupin-supplemented dairy cows than for soybean meal fed cows (Robinson et al., 1993). Milk fat had higher concentrations of long-chain fatty acids (Singh et al., 1995). A shift of the fatty acid profile of milk (increase in C18:1, decrease in C12:0-C16:0) towards guidelines for improved cardiovascular health in human populations has been observed (White et al., 2007).

Grinding and flattening

N solubility (Moss et al., 2001) and in situ protein degradability of lupin (Jahn et al., 1999) both increase with the fineness of grinding, and the protein value can be controlled through substrate particle size by grinding. The effective degradability of N is lower with coarse grinding (5-mm screen) than with fine grinding (through a 1.0 or 1.5-mm screen) (63 vs. 93-95%). This increases the amount of protein digested in the intestine, and thus the MP value, although the whole tract apparent digestibility of CP is unchanged (Kibelolaud et al., 1991). In vivo measurements of digestive flows in fistulated bulls show that lupin seed should be coarsely ground or flattened to obtain a mean particle size between 2.0 and 4.2 mm for cattle, which provides the highest level of digestible protein in the small intestine. With finer particles, proteins are highly degraded, and with insufficient grinding level (6 mm), a higher ruminal degradation of lupin protein is also observed, probably due to more intense rumination (Froidmont et al., 2008).

Thermal treatments

Dry roasting

Roasting increases the undegraded protein fraction of lupin seed (+70 g/kg DM) without increasing the ADF-N fraction, which indicates a minimal protein damage due to roasting (Singh et al., 1995). However, an increase in ADF-N could be observed with roasting at 300°C during 1 to 4 min (Zaman et al., 1995). In situ N disappearance of whole lupin or dehulled lupin decreases as roasting temperature increases from 130 to 175°C (Kung et al., 1991). In situ ruminal incubation followed by in vitro enzyme digestibility trials suggests that dry roasting whole lupin seeds can shift the digestion of protein from the rumen to the lower gastrointestinal tract without depressing the digestion of rumen-undegraded protein. Among the tested conditions (110, 130 or 150°C for 15, 30 or 45 min), dry roasting at 150°C for 45 min gave the best results (Yu et al., 1999c). A similar study conducted under different dry roasting conditions (100, 118 and 136°C for 3, 7, 15 and 30 min) lead to similar conclusions: rumen undegraded protein increased (+108 g/kg DM between raw and roasted at 136°C/15 min), due to decreased soluble fraction and degradation rate. Estimated microbial protein based on available energy in the rumen is only slightly reduced, and true digestibility of dietary protein is unchanged (88% with the mobile bags technique), so that the value of true digested protein in the small intestine is increased by dry roasting from 119 to 197 g/kg DM (Yu et al., 1999a). However, the potential damaging effects of processing on individual amino acids, especially on the first limiting amino acids, remains unclear (Yu et al., 1999a).

In dairy cows, roasting, while decreasing lupin protein degradability, does not always have an effect on milk production. Cows fed roasted lupins produced more milk (with higher milk fat, protein and lactose yields) than those fed raw lupins (Singh et al., 1995), but roasting had no effect on DM intake or milk protein content (Robinson et al., 1993). Roasting increased protection of lupin oil from ruminal hydrogenation, as evidenced by increased concentrations of long-chain fatty acids in milk from cows supplemented with roasted lupins (Robinson et al., 1993).

In lambs, roasting decreased ruminal in situ protein disappearance but had no effect on growth rate and feed efficiency, which were similar to that obtained with a soybean meal-based diet, suggesting that whole or dehulled lupins can replace soybean meal as the sole protein supplement for growing lambs (Kung et al., 1991).

Extrusion

Extrusion decreases ruminal N in situ degradability by up to 20-30 percentage points (Cros et al., 1991; Benchaar et al., 1992; Kibelolaud et al., 1993; Aufrère et al., 2001; Rémond et al., 2003; Solanas et al., 2005; Barchiesi-Ferrari et al., 2011). This effect depends on extrusion temperature (110°C to 195°C) or moisture (dry to 20% humidity). It is linked to a reduction in the soluble fraction (Kibelolaud et al., 1993), and to an increase in the slowly degradable fraction (Barchiesi-Ferrari et al., 2011).

Extrusion can also increase the intestinal digestibility of by-pass protein (Solanas et al., 2005). Both effects contribute to a positive impact of extrusion on the amount of truly digestible protein in the intestine (Benchaar et al., 1992; Cros et al., 1991; Kibelolaud et al., 1993). The inclusion of a source of carbohydrates before extrusion at 140°C increases this response (Solanas et al., 2005).

Extrusion at 195°C did not alter the amino acid (AA) profile of whole lupin seeds, but did affect markedly the AA profile of the lupin protein that escapes ruminal digestion (Cros et al., 1991; Benchaar et al., 1993). The increase in intestinal disappearance varies substantially among AA (Benchaar et al., 1993). Based on estimated AA chemical score (test-to-milk ratio), the absorbed protein shows a higher protein quality in extruded than in raw lupin seed (Cros et al., 1991).

Extrusion was shown to affect the MP supply in dairy cows (Benchaar et al., 1991; Benchaar et al., 1994) and in bulls (Froidmont et al., 2008). It resulted in a decrease in protein (and to a lesser extent OM) ruminal true digestibility, in an increase in duodenal non-ammonia N and dietary N flows, and in an increase in AA true digestibility and subsequently in the amount of AA absorbed in the small intestine (up to 700 g/d in dairy cows). The effect on ruminal NH₃-N and VFA concentrations and on the efficiency of microbial synthesis seems less clear. In sheep (whethers), the increase in dietary protein at the duodenum was balanced by a decrease in microbial flow, so that the total non-ammonia N flow and the profile of apparently absorbed AA did not differ between raw and extruded lupin (Rémond et al., 2003).

In growing/fattening bulls, extrusion shifted lupin protein digestion from rumen to small intestine, but was less efficient than grinding (4.2 mm) for improving the nutritional value of lupin seeds, and grinding was also simpler and cheaper (Froidmont et al., 2008).

Other processes

Autoclaving at 120°C for 30 min reduced both the soluble fraction and the fractional rate of protein degradation of the slowly degraded fraction, and subsequently the effective degradability of protein (Aguilera et al., 1992). NaOH addition did not affect protein degradability (Jahn et al., 1999).

Limits of the in situ approach to assess the protein value of lupin

It is recognized that the in situ technique underestimates the protein value of lupin seed due to its high soluble protein fraction. Although this fraction is assumed to be fully degraded in the effective degradability calculation, a non negligible part may actually escape ruminal degradation as proteins, peptides or amino-acids and contribute to dietary non-ammonia N flow at duodenum as quantified in vivo on whethers (Aufrère et al., 2001; Rémond et al., 2003).

Pigs

White lupin is generally not included in pig diets above 15-20% of the diet, due to its depressing effect on feed intake and growth at higher dietary level (Froidmont et al., 2006; van Barneveld, 1999; Edwards et al., 1998; Donovan et al., 1993; Kelly et al., 1990). White lupin seeds have a high energy value for pigs, being rich in digestible lipids and carbohydrates (Aguilera et al., 1985). However, some of the soluble and non soluble polysaccharides (NSP) and oligosaccharides present in the lupin seed have deleterious effects on digestible energy and on animal health as they are readily fermented in the large intestine (van Barneveld, 1999). Including lupin in pig diets tend to decrease digestibility at the end of the intestine and it has been recommended to use the net energy value of lupin seed rather than the digestible energy value in order to optimize ration efficiency (van Barneveld, 1999). It should be noted that data on the effect of lupin on production efficiency in pigs has not always been consistent (Pisarikova et al., 2009).

Early results showed that barley-based diets containing 27-30% white lupin seeds had good fibre digestibility (60%) and that white lupin crude fibre was more digestible than that of soybean meal (Aguilera et al., 1985; Florence, 1965; Farries et al., 1968 cited by Aguilera et al., 1985). The addition of 30% white lupin seeds in barley-based diets increased pigs tolerance to high fibre basal diets (Aguilera et al., 1985; Barnett et al., 1981).

White lupin seeds may contain small amounts of deleterious alkaloids that hamper pig health. No more than 0.012% of alkaloids from white lupin seeds can be tolerated in pig diets (Buraczewska et al., 1993).

Raw lupin seeds

Piglets

The recommended inclusion level of white lupin seeds in piglet diets varies according to authors. In Portugal, white lupin seeds were included at 30% of the diet of weaned piglets (28 day-old) during 11 days without negative effect on body weight gain and feed conversion ratio. However, their lipase and trypsin activity was significantly reduced (by 45% for trypsin activity) (Pereira et al., 2005). In France, white lupin seeds were recommended at only 10% for weaned piglets over 12 kg (Royer et al., 2005). When compared to other lupin species such as yellow lupin or blue lupin, it was reported that white lupin had lower DM and protein digestibilities in young pigs (Gdala et al., 1996).

Growing and fattening pigs

In a growing pig experiment comparing a diet containing 20% lupin seeds to a 15% soybean meal-based diet, N retention and growth rates were lower for the lupin-based diet, resulting in a longer fattening period and a lower average daily gain (Froidmont et al., 2006). This result is in agreement with former observations with fattening pigs (34-110 kg BW) where animals were reported to have decreased feed intake and daily gain when lupin was included at 20%. This also resulted in inferior back fat consistency, and it was thus suggested to limit lupin inclusion to 10% (Zettl et al., 1995). Other experiments have been more positive: in experiments where white lupin protein replaced 50% or 100% of soybean meal protein, average daily gain was non-significantly lower by 1.5 to 6.2% compared with soybean in the starter diet, but performance and feed efficiency were higher compared to the control in grower and finisher diets. The authors concluded that soybean protein could be completely supplemented by lupin in fattening diets, provided that the diets are balanced with limiting amino acids (Pisarikova et al., 2009).

Sows

Lupins are more suitable to sows than to growing pigs, as sows can better ferment lupin carbohydrates and get more energy from them. Lupin seed hulls have a higher energy values for sows than for growing pigs (Noblet, 1997). However, it was recommended to include lupin at levels below 20% of the diet since they may result in high levels of gas production and could compromise sow health (van Barneveld, 1999).

Processing

Dehulling

It has been suggested that white lupin kernels have a higher NSP content than lupin hulls, which may explain why dehulled lupin seeds were not readily consumed by growing pigs and did not result in satisfactory animal performance (Ferguson et al., 2003). Data from the literature concerning the effect of seed dehulling on nutrient digestibility and performance characteristics for the main lupin species are rather inconsistent: some trials found that dehulling had a negative effect or no effect on performance (Fernández et al., 1995; King et al., 2000), while other trials reported better performance or/and higher nutritional values (Flis et al., 1996; Flis et al., 1997; Noblet et al., 1998). In 2020, a review reported that dehulled white lupin could valuably replace soybean meal in pigs diets at all stages of growth (starter, growers and finishers) (Struti et al., 2020).

Extrusion

Extrusion of white lupin seeds had no effect on feed intake, average daily gain and feed conversion ratio for 28 day-old weaned piglets fed on soybean meal based diet and 17% white lupin seeds (Prandini et al., 2005).

Enzyme or chemical processing

White lupin seeds treated with phosphoric acid and hot water in fattening pigs at 6% of the diet, partially replacing soybean meal or meat meal, increased weight gain and carcass quality (Herold et al., 1991). The addition of an α-galactosidase with or without a mixture of cellulase and hemicellulase to the diet of weaned piglets had no effect on the ileal digestibility of protein and amino acids but increased NDF ileal digestibility when α-galactosidase was associated to cellulase and hemicellulase (Pires et al., 2007).

Lupin seeds hulls

Metabolizable energy value of lupin seeds hulls was reported to be as high as 14 MJ/kg in sows and 7 MJ/kg in growing pigs (Noblet, 1997).

Poultry

The nutritional value and suitability of lupin seeds for poultry feeding vary greatly according to the species (white, blue, yellow lupin), to the variety (sweet or bitter), to seed composition and to processes (Jeroch et al., 2016). The protein content, the amino acid profile and the fatty acid profile of lupin seeds suggest that they are valuable raw materials for poultry feeding, but these seeds remain poorly used since they are poorly available for feed manufacturers or poultry producers. The interest of lupines for poultry feeding must be discussed according to their ME value, amino acids digestibility and to their anti-nutritional factors content (alkaloid, oligosaccharides, …). According to the variety and the cultivar, their ME value can be higher than 10 MJ/kg DM, but values as low as 8.3 MJ/Kg DM were observed for seeds whose oligosaccharides content (mainly raffinose) was high (Kaczmarek et al., 2016).

Oligosaccharides

Since the advent of sweet varieties with low alkaloid content, oligosaccharides have become the main issue when using lupin in diets that are well balanced for energy, amino acids (methionine supply) and minerals. Alpha-galactosides are not digested in the upper part of the gastro intestinal tract of the birds and they are only fermented in the caeca and produce short-chain fatty acids, poorly used in poultry. High levels of oligosaccharides are known to produce negative effects on digestive processes (increased digesta viscosity, reduced digestibility, sticky droppings…) and consequently reduce performance.

When high amounts of lupins (30%) are fed, ME value and amino acid digestibility are negatively correlated with the level of raffinose and water extract viscosity of lupin seeds and this could be not only due to a reduced diet quality but also to endogenous losses caused by white lupin inclusion (Kaczmarek et al., 2016). However, small amounts of lupins have been shown to affect favourably fermentation processes and stimulate gut microbiota in turkey (Zdunczyk et al., 1996).

Dehulling

The nutritional value of lupin seeds can be improved by dehulling, as this process reduces the non starch polysaccharide (NSP) content: dehulling significantly improves dry matter digestibility, protein digestibility, starch digestibility, and AME (Nalle, 2009). Dehulling of white lupin seeds (low alkaloid, cv. Amiga) increased the AME values by 15 to 18%, whilst the digestibility of protein improved by 7% (Brenes et al., 2003; Brenes et al., 1993). This higher AME could be partially explained by the removal of fibrous hulls and, a reduction in soluble and total non-starch polysaccharides (Nalle, 2009). In some cases, dehulling also increases amino acid digestibility (Brenes et al., 1993, Nalle et al., 2010).

Extrusion

Other processes like extrusion were not found to be effective in improving lupin seed nutritive value for poultry. Extruded lupin seeds resulted in a lower feed intake compared to the control and to raw lupin seeds in broilers (Diaz et al., 2006). The lower feed intake could be attributed to the higher bulkiness of extruded lupin seeds in comparison to the control diet or to the raw lupin seed. Extrusion was also reported to decrease the AME value of sweet lupin (Lupinus angustifolius) (8.61 MJ/kg vs. 7.52 MJ/kg). This decrease of AME was also attributed to the increased bulkiness associated with extrusion with a resulting decrease in energy intake (Breytenbach, 2005).

Enzymes

Attempts have also been made to reduce the negative effects of oligosaccharides by adding exogenous carbohydrases, which resulted in better performance of birds receiving diets containing large amounts (50 and 70%) of untreated (raw) white lupins only. The birds receiving diets containing the same amount of treated (autoclaved, dehulled) lupins did not improve their growth performance (Brenes et al., 1993).

Broilers

Results about the use of white lupin in broiler diets are not consistent. Trials conducted prior to 2000 reported that very high levels of lupin seeds could be safely included in broiler diets: from 20-30% to 50%-70% (Brenes et al., 1993; Castanon et al., 1990; Bekric et al., 1990). When methionine was provided, inclusion rate could reach 40% (Watkins et al., 1988; Buraczewska et al., 1993). Experimental recommendations were as high as 30-40% in balanced broiler diets (Olver, 1987; Uzu, 1983). And, at this level (40%) of inclusion no difference in growth performance, feed efficiency or carcass characteristics were observed in broilers up to 8 weeks of age (Olver, 1987).

More recent trials are more inconsistent and suggest that raw lupin seed levels should be included at lower level in broilers diets. Some experiments indicated that up to 20% inclusion, raw lupin seeds had no adverse effect on average daily gain of birds or on FCR (Nalle et al., 2012; Moschini et al., 2005). this is in accordance with prior observations from Roth-Maier in 1994 (Roth-Maier et al., 1994). When 25-45% white lupins were added to broiler diet, no difference in final body weight could be observed but feed intake and feed conversion ratio deteriorated by about 6%. Moreover, above 25% lupine seeds in the diet, the faecal DM of broilers dropped below 32% , the faeces became watery and sticky, and abdominal feathers were glued together or had been lost. It was concluded that no more than 20% white lupins should be included in broiler diets (Roth-Maier et al., 1994b). A recent trial reported a depressing effect on growth at 20% inclusion rate, and a degraded feed conversion ratio at 15% inclusion rate (Kaczmarek et al., 2016).

In German and polish context, maximum recommended levels of raw lupin seeds in well-balanced diets (particularly for methionine) were set at 5%-10 % in starter diets and at up to 15-20% in older broilers (Jeroch et al., 2016). Under commercial management of broilers, the recommended level of lupin seeds remained lower than 10% because of the occurrence of wet droppings and high gut viscosity (Farrell et al., 1999).

Extruded lupin seeds

Extruded lupin and raw lupin seeds were included at 30% in broilers diet over growing (1-21 days) and finishing (22-42 days) periods. The inclusion of extruded lupin had depressive effect on growth on both periodes, but not on the overall period (Diaz et al., 2006). Extruded lupin seeds resulted in lower feed intake than the control or the raw lupin seeds (Diaz et al., 2006).

Dehulled-micronised lupin seeds

It was possible to feed broiler chickens (14-49 d) on a diet containing 24% dehulled-micronised lupin seeds without adverse effect on their growth rate or FCR compared to control diet (soybean meal based diet). In addition, the inclusion of dehulled-micronised lupin seeds had positive effects on meat quality and on fatty acids profile (Laudadio et al., 2011).

Laying hens

Laying hens, could be fed over 3 months with up to 30% white lupin seeds when the diet was supplemented with methionine or with methionine and enzymes (200 mg Roxazyme) (Roth-Maier et al., 1994a; Prinsloo et al., 1992). White lupin levels up to 30% did not reduce performance while yolk colour and poly-unsaturated fatty acids profiles were improved (Krawczyk et al., 2015).

Turkeys

Up to 40% dehulled-micronised lupine seeds could be used in growing turkeys diet without compromising their growth rate while the same level of whole lupine seed decreased animal growth rate. This could be explained by the lower fibre content of the diet containing dehulled lupine seeds (Halvorson et al., 1983).

Ducks

Pekin ducklings could receive 40% white lupin seed meal without deleterious effect on growth, feed conversion or carcass quality (Petterson, 2000).

Guinea fowls

Guinea fowls could be fed on 24% dehulled-micronised lupin as the main source of protein to replace soybean meal without any adverse effect on growth rates, dressing percentage or breast and thigh muscle. The abdominal fat was reduced by the lupine seed based diet, meat lipid content and cholesterol were reduced and the phospholipids were in higher concentration. The use of lupin had positive effects on polyunsaturated fatty acids (PUFA) in breast meat while decreasing saturated fatty acids: two desirable characters for human health (Tufarelli et al., 2015)

Rabbits

Seeds

White lupin seeds have been recommended for rabbit feeding for a long time (Benoit et al., 1948). The usual recommended inclusion rate in balanced diets for growing rabbits varies from 10 to 20% depending on the trial (Battaglini et al., 1991; Mesini, 1997; Sarhan, 1999; Seroux, 1984; Volek et al., 2008; Volek et al., 2009). However, higher inclusion rates in the 40-50% range have been used in experiments without depressing growth performance (Fekete et al., 1986; Kelly et al., 1990). White lupin seeds could be used successfully in the diet of lactating rabbit does. Compared to a soybean-sunflower meal control diet, a diet with 25% white lupin seeds increased the 1-31 d. milk production of the does by 11%, most due to the higher lipid content of the lupin diet: 4.1% vs. 2.3% (Volek et al., 2014). A comparison of 6 French cultivars of white lupin seeds differing in composition did not result in differences in growth performance, and the cultivars were found of similar nutritive value (Lebas, 1986).

While the presence of alkaloids and other antinutritional factors in lupin seeds makes processing or enzymes useful for their optimal utilisation in pigs and poultry (Cheeke et al., 1989; Brenes et al., 1993), rabbits are little sensitive to lupin alkaloids (Cheeke et al., 1989). Washing the seeds or the addition of an enzyme cocktail failed to improve the nutritive value of white lupin seeds for growing rabbits (Falcao-e-Cunha et al., 2008). However, extrusion of imported Australian white lupin seeds (16% of the diet) improved growth rate by 10% compared to the control diet without lupin in an Italian study (Battaglini et al., 1991).

In practical conditions, white lupin seeds are a valuable source of proteins with a high content of digestible energy: 13.5 to 16.0 MJ/kg DM (Fekete et al., 1986; Lebas, 1986; Maertens et al., 2002). This high energy content results from the relatively low fibre content and, more specifically, from the high lipid content despite a negligible starch content (Petterson, 2000). These lipids are relatively rich in alpha-linolenic acid C18:3, that represents 9-10% of total fatty acids (Chiofalo et al., 2012). This can be useful to improve the quality of rabbit meat for human consumption through a better omega 6 to omega 3 ratio (Volek et al., 2011), and can also partly explain the improvement of the health condition of rabbits fed a lupin-based diet compared to that of rabbits fed a soybean meal-based diet (Colin et al., 2012; Uhlířová et al., 2016). However, white lupin seeds are not very rich in lysine (4.9 g/16 g N), which only covers the growing requirements. Like other legume seeds, white lupin is strongly deficient in sulphur-containing amino acids, covering only 65% of rabbit needs (Lebas, 2013), which makes necessary the supplementation with ingredients such as cereals or cereal byproducts (wheat bran...), or with synthetic methionine.

Hulls

Lupin hulls represent 22% of the whole grain (Petterson, 2000). In growing rabbits, replacing wheat bran (10% inclusion rate) with 5% white lupin hulls and 5% barley did not affect growth rate (52.4 g/d on average), feed intake (155.1 g/d), feed conversion ratio (2.97), energy digestibility and nutrient digestibility (protein and NDF). As a consequence, white lupin hulls can be considered as a suitable source of fibre for rabbits (Volek et al., 2013).

Fish

White lupin seeds are considered to be a potential protein source in aquaculture, particularly as a substitute for fish meal or soybean meal.

Salmonids

Rainbow trout (Oncorhynchus mykiss)

Recommended inclusion levels are very variable, ranging from 20% to 50% (Hernández et al., 2016; Borquez et al., 2011; Bangoula et al., 1993). White lupin seeds included in rainbow trout (5 g BW) diets at 25% (DM) to replace fish meal were found to be a better protein source than soybean meal for the formulation of low-phosphorus loading diets, without affecting feed acceptability and growth performance (Hernández et al., 2016). In rainbow trouts (75 g or 50 g) fed on diets containing 20% or 30% raw or extruded white lupin seeds during 2 weeks, apparent digestibility coefficients (ADC) of the diets were reduced as the level of lupin increased (Bangoula et al., 1993). White lupin seeds could be included at up to 20% in commercial extruded diets without modifying dietary ADCs, feed intake or animal performance (Bórquez et al., 2011). In a longer experiment (83 days), raw white lupin included at 20% resulted in deleterious effects and mortality, but trouts fed extruded white lupin seeds had higher growth rate, feed intake, and feed conversion ratio which indicated that antinutritional factors were reduced through the extrusion process (Bangoula et al., 1993). Rainbow trouts (54 g BW) fed during 11 weeks on diets containing up to 50% white lupin seeds showed no significant effects on growth, feed utilization, ADC or whole-body composition. However, increasing levels of dietary lupin led to histological changes in the digestive tract (slight lipid infiltration into hepatocytes and enterocytes) and in muscle fatty acid profile (Borquez et al., 2011). Feeding rainbow trout juveniles on 35% white lupin seeds as fish meal replacer had no deleterious effect on the expression of immunological genes (Hernandez et al., 2013).

Atlantic salmon (Salmo salar L.)

The inclusion of white lupin seeds in young Atlantic salmons (90 g BW) diet depressed growth rate and feed efficiency significantly when compared to the control diet based on fish meal (Salini et al., 2014).

Latids

Barramundi (Lates calcarifer)

White lupin seed meals from 3 different cultivars were included in the diets of tropical barramundi fish at 30% as a substitute for fish meal. The lupin-based diets had higher protein digestibility than the control diet. Both DM matter and energy digestibilities were superior to other lupin species such as blue lupin. White lupin seeds were considered a suitable source of protein for carnivorous tropical barramundi (Tabrett et al., 2012).

Crustaceans

Black tiger shrimp (Penaeus monodon)

It was possible to replace up to 75% fishmeal with lupin meal in black tiger shrimp (Penaeus monodon) juveniles diet without affecting growth, survival, feed conversion ratio (FCR), protein efficiency ratio (PER), and apparent net protein utilization (ANPU), and body composition (Sudaryono et al., 1999).

Nutritional tables

Tables of chemical composition and nutritional value

Avg: average or predicted value; SD: standard deviation; Min: minimum value; Max: maximum value; Nb: number of values (samples) used