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Lupinus albus

Lupinus albus, commonly known as white lupin, is an annual herbaceous legume in the Fabaceae family, native to southeastern Europe and western Asia, characterized by its erect, branched growth up to 120 cm tall, palmately compound leaves with 5-9 leaflets, and white to pale violet flowers arranged in terminal racemes.^[1]^[2]^[3] It produces oblong pods containing 3-7 flattened, reniform seeds that vary from "sweet" (low alkaloid content <0.02%) to "bitter" varieties (up to 4% alkaloids), with the plant featuring a strong taproot and cluster roots that enhance phosphorus uptake in nutrient-poor soils.^[1]^[3] Cultivated for over 2,000 years originating from the Mediterranean Basin and Nile Valley, L. albus thrives in subtropical to temperate climates with optimal temperatures of 15-25°C and annual rainfall of 380-990 mm, exhibiting frost tolerance down to -8°C in some genotypes and serving as a nitrogen-fixing crop that contributes 140-350 kg N/ha to soil fertility.^[1]^[4]^[3] Its seeds are nutrient-dense, boasting 34-44% protein (rich in lysine), 40% dietary fiber, and essential minerals like potassium (8.6-11.1 g/kg) and calcium (2.1-4.7 g/kg), making them valuable for human consumption after processing to remove bitter alkaloids, as well as for livestock fodder, silage, and green manure in crop rotations.^[4]^[3] Beyond agriculture, L. albus holds ecological significance for soil remediation, such as phytostabilizing heavy metals in contaminated sites, and medicinal potential, with studies indicating benefits for cardiovascular health, glucose regulation, and hypertension reduction due to bioactive compounds like γ-conglutin, though bitter varieties pose toxicity risks from quinolizidine alkaloids if unprocessed.^[1]^[4]^[3] Global production, led by Poland, underscores its role as a sustainable protein source amid growing demand for plant-based foods.^[4]

Taxonomy and description

Taxonomic classification

Lupinus albus, commonly known as white lupin, belongs to the kingdom Plantae, phylum Tracheophyta, class Magnoliopsida, order Fabales, family Fabaceae, genus Lupinus, and species L. albus.^[5] Within the Fabaceae, it is placed in the subfamily Faboideae, tribe Genisteae, and subtribe Lupininae.^[5] It is further classified in section Lupinus (or sect. Albus) of the genus.^[6] Accepted synonyms for L. albus include Lupinus termis Forssk. and Lupinus albus var. graecus (Boenn.) Thell., reflecting historical nomenclatural variations.^[7] L. albus stands out among over 200 Lupinus species as the principal annual white-seeded lupin domesticated for low-alkaloid "sweet" varieties, in contrast to the bitter, high-alkaloid profiles of most wild relatives.^[1] Recent phylogeographic and genomic analyses in the 2020s have reinforced L. albus's Mediterranean origin, tracing its domestication to approximately 3000–4000 years ago in the eastern Mediterranean basin through pangenome sequencing and population structure studies.^[8]^[9]

Morphological characteristics

Lupinus albus is an annual herbaceous legume characterized by an erect, branched stem that exhibits a shrubby growth habit, typically reaching heights of 0.3 to 1.2 m, though it can attain up to 1.6 m under optimal conditions.^[1]^[10] The stem is short-hairy and supports sequential branching, contributing to its bushy appearance.^[11] The leaves are alternate and palmately compound, consisting of 5 to 9 digitate leaflets that are nearly smooth on the upper surface and silvery-hairy on the underside.^[1]^[11] Each leaflet measures approximately 2 to 4 cm in length, providing a distinctive foliage structure adapted for photosynthetic efficiency.^[11] Flowers of L. albus are predominantly white, occasionally pale blue to violet, and arranged in dense terminal racemes that can extend up to 30 cm in length.^[1]^[12] The species is largely self-pollinating, with selfing rates of 50 to 85%, though it attracts bees and can undergo cross-pollination.^[1] The fruit is a long, oblong pod, typically 6 to 15 cm in length, containing 3 to 7 large, white, flattened seeds per pod.^[12]^[1] These seeds measure about 7 to 16 mm in length, 6 to 12 mm in width, and 2 to 5.5 mm in thickness, with cultivated "sweet" varieties exhibiting low alkaloid content of 0.01 to 0.03%.^[12]^[1] The root system features a strong taproot that penetrates to depths of 1 to 2 m, facilitating access to deeper soil resources.^[11]^[13] In phosphorus-deficient acidic soils, it develops specialized cluster roots (proteoid roots) that enhance nutrient acquisition through exudation of organic acids.^[1]^[11] As an annual species, L. albus completes its life cycle within 106 to 180 days, with flowering occurring 80 to 120 days after sowing and seed maturation occurring 150 to 180 days after sowing.^[13]^[14] This phenological progression supports its adaptation as a cool-season crop.^[1]

Distribution and habitat

Geographic range

_Lupinus albus is native to the northeastern Mediterranean Basin, with wild populations occurring in southeastern Europe and western Asia, including countries such as Albania, Bulgaria, Greece (including Crete and the East Aegean Islands), Sicily, and Turkey. These wild forms are found in coastal dunes and disturbed areas within this subtropical biome.^[2] The species was likely domesticated in the Aegean region of Greece between 4000 and 2000 BCE, marking the beginning of its cultivation as a legume crop in ancient Mediterranean societies.^[15]^[12] Today, L. albus has been widely introduced and cultivated beyond its native range, particularly in Europe (including Poland, Germany, France, Italy, Portugal, Russia, and Ukraine), Australia (notably in arid plains of Queensland and Western Australia), South America (such as the Andean highlands of Chile and regions in Argentina and Brazil), and parts of North and sub-Saharan Africa (including Egypt, Sudan, Ethiopia, South Africa, Kenya, Tanzania, and Zimbabwe).^[2]^[1]^[16]^[12] The species' spread is limited by its sensitivity to frost in northern latitudes, though ongoing breeding efforts have developed varieties adapted to cooler climates, enabling cultivation in more temperate regions of Europe and beyond.^[17]^[9]

Ecological adaptations

Lupinus albus forms a symbiotic relationship with nitrogen-fixing bacteria, primarily Bradyrhizobium lupini, which inhabit root nodules and convert atmospheric nitrogen into forms usable by the plant, typically fixing 100–250 kg N/ha annually.^[18] This process significantly reduces the need for external nitrogen inputs in natural ecosystems, enhancing soil fertility for subsequent plant growth.^[19] The species thrives in acidic soils with a pH range of 4.5–6.5, particularly well-drained sandy-loam types, where its specialized cluster roots—dense clusters of short lateral roots—facilitate enhanced phosphorus uptake in phosphorus-deficient environments.^[1]^[20] These cluster roots exude organic acids like citrate, solubilizing bound phosphorus and improving nutrient availability without relying on high soil fertility.^[21] Adapted to Mediterranean climates, L. albus performs optimally at temperatures of 15–25°C and requires 400–600 mm of annual rainfall, with its deep taproot system providing drought resistance by accessing subsoil moisture.^[22] In ecosystems, it contributes to soil structure improvement through extensive root networks that prevent erosion and enhance organic matter incorporation during crop rotations, while recent studies highlight its potential for phytoremediation, accumulating heavy metals such as cadmium and zinc from contaminated soils.^[23]^[24] However, in introduced regions, L. albus can exhibit invasive tendencies, outcompeting native vegetation through allelopathic effects from root exudates containing quinolizidine alkaloids that inhibit seed germination and growth of surrounding plants.^[25] This chemical interference alters microbial communities and reduces biodiversity in affected habitats.^[26]

History

Domestication and early uses

Lupinus albus, commonly known as white lupin, was domesticated in the Mediterranean region approximately 3000–4000 years ago, with its center of origin believed to be in southern Greece and western Anatolia.^[8] Primary evidence for early cultivation comes from archaeological finds of seeds in Bronze Age contexts across the Aegean, including sites in Greece and Crete dating to around 2000 BCE, indicating its integration into early agricultural systems as a pulse crop.^[27] These findings suggest that ancient communities in the region selected for larger-seeded forms from wild populations, marking the initial steps toward domestication for food and soil improvement purposes. In ancient Egypt, where it was known as Lupinus termis, the plant was cultivated around 2000 BCE during the Twelfth Dynasty (c. 1991–1802 BCE), as evidenced by seeds discovered in tombs of pharaohs from this period.^[28] Egyptians consumed the bitter seeds after soaking them in water or brine to leach out toxic quinolizidine alkaloids, transforming them into a palatable snack or meal component.^[29] This preparation method was crucial, as wild and early domesticated varieties contained alkaloid levels exceeding 1% of seed dry weight, rendering them unpalatable and potentially harmful without processing.^[30] By the classical period, L. albus had spread to the Roman Empire, where it served dual roles in human diets and animal fodder. Roman naturalist Pliny the Elder, writing in the 1st century CE, described the lupine in his Natural History (Book XVIII, Chapter 36), noting that seeds soaked in warm water could be eaten by humans and that a single modius (about 8.6 liters) sufficed to feed an ox, imparting vigor to livestock.^[31] Early selection efforts focused on "sweet" varieties with reduced bitterness, progressively lowering alkaloid content through breeding; modern cultivars have levels below 0.03%, a stark contrast to wild forms exceeding 1%.^[32] The crop's dissemination continued via Roman trade and expansion, reaching Iberia during the empire's westward growth, where it was valued for its nitrogen-fixing properties in crop rotations.^[33] Following the European discovery of the Americas in 1492, L. albus was introduced to the New World by Spanish and Portuguese colonizers, establishing it as a minor crop in regions like Chile and Peru by the 16th century.^[34]

Modern developments

In the late 19th and early 20th centuries, Lupinus albus experienced a revival in Europe as a protein-rich feed crop, driven by the need to expand cultivation on marginal, sandy soils amid food shortages following World War I, particularly in Germany where it was promoted for its nitrogen-fixing properties and adaptability to poor lands.^[35] Breeding programs emerged shortly after the war in Germany and Poland, focusing on reducing alkaloid content to make seeds safer for livestock and human consumption, with early efforts in Poland targeting early-maturing varieties suitable for local climates.^[36] In Australia, systematic breeding for L. albus began in the 1920s, emphasizing fodder yield and disease tolerance, building on introductions from Europe to support expanding agriculture on acidic, low-fertility soils.^[37] Genetic improvements have accelerated since the mid-20th century, with the development of low-alkaloid "sweet" cultivars that minimize quinolizidine alkaloids to below 0.02% dry weight, enabling broader food and feed applications. Notable examples include the cultivars 'Amiga' and 'Estoril', released in the 2020s, which combine high seed yield (up to 3.5 t/ha) with alkaloid levels under 0.5 g/kg, making them suitable for human nutrition while retaining protein contents of 35-40%.^[38] Recent advancements incorporate genome editing technologies, such as CRISPR/Cas9, to enhance disease resistance against pathogens like anthracnose and improve yield stability; 2025 studies have demonstrated successful targeted mutations in L. albus genes related to phosphorus uptake and stress response, potentially increasing seed production by 20-30% under suboptimal conditions.^[39] Economically, as of 2023, lupin cultivation (including L. albus) covers over 1 million hectares globally, contributing to a total lupin output exceeding 1.6 million tonnes annually and valued for its role in diversifying protein sources amid rising demand for plant-based feeds.^[40] By 2024, global lupin cultivation had expanded further, with EU areas exceeding 400,000 ha, driven by sustainability policies.^[41] In the European Union, its cultivation is promoted as a sustainable alternative to imported soy, aligning with initiatives like the Green Deal for crop rotation, soil health, and reduced synthetic nitrogen use, with incentives for legume integration in farming systems to achieve climate-neutral agriculture by 2050.^[42]^[43] Addressing climate change challenges, breeding efforts have prioritized drought-tolerant lines for Mediterranean regions, where increasing aridity threatens yields; genetic variation among landraces has been exploited to develop cultivars with deeper root systems and enhanced water-use efficiency, sustaining productivity under rainfall deficits of up to 50% during reproductive stages.^[44] These adaptations, informed by phenotypic screening of diverse germplasm, position L. albus as a resilient crop for warming climates, with ongoing pre-breeding programs integrating wild relatives to bolster tolerance to heat and water stress.^[45]

Cultivation

Environmental requirements

_Lupinus albus thrives in cool temperate to subtropical climates, where it is typically sown in autumn or winter to facilitate a spring harvest. It tolerates short-term temperatures down to -5°C to -7°C during vegetative stages but is sensitive to prolonged frost, which can damage germination and flowering. Optimal growth occurs at mean temperatures of 15–25°C, with flowering requiring cool conditions below 20°C and short photoperiods to prevent heat-induced flower abortion.^[1]^[46]^[47] The crop prefers well-drained, light-textured soils such as sands or loams, with a pH range of 5.5–7.0 to support healthy root development and nitrogen fixation. It performs poorly on heavy clay soils prone to compaction or waterlogging, which can lead to root rot and reduced yields. While tolerant of mildly acidic to neutral conditions, high alkalinity or calcareous soils (with more than 3% free lime) may induce iron chlorosis, limiting cultivation in such areas.^[1]^[46]^[47] Water requirements during the growing season range from 400–1,000 mm, with peak demand during flowering and pod filling to avoid drought stress that impairs seed set. Its deep taproot enables efficient water use, making it suitable for rainfed systems in Mediterranean-like environments. However, excessive moisture leading to waterlogging must be avoided to prevent disease. Irrigation can enhance yields by up to 70% in drier conditions, particularly on sandy soils with low retention.^[47]^[48]^[1] Incorporating L. albus into rotations following cereals like wheat or barley helps break disease cycles and replenishes soil nitrogen through symbiotic fixation, improving subsequent crop yields by up to 20% and enhancing soil organic matter. This practice is especially beneficial on marginal lands, where the plant's ability to access phosphorus from deeper soil layers supports its growth without heavy fertilization.^[1]^[46] Limitations include sensitivity to high soil salinity, which reduces germination and growth, and unsuitability for very heavy clays or poorly drained sites. Despite its nitrogen-fixing capacity, L. albus is best suited to marginal but non-extreme lands, avoiding dry regions with less than 380 mm annual rainfall; while tolerant of higher altitudes up to 3,000 m in suitable climates, yields may decline above 500 m in temperate lowlands.^[49]^[47]^[46]

Planting and management

Lupinus albus is typically sown at a seed rate of 100–150 kg/ha to achieve optimal plant density, though rates can vary from 50 to 180 kg/ha depending on soil type and expected germination. Seeds should be planted at a depth of 2–5 cm in firm, well-prepared soil to ensure good establishment, with row spacings of 15–30 cm facilitating mechanical operations and weed suppression. In mild climates with minimal winter frost, autumn sowing from mid-September to late October is preferred for winter-hardy ecotypes, promoting earlier vegetative growth and higher yields compared to spring sowing; in cooler regions, spring planting in March–April is recommended once soil temperatures reach at least 6°C.^[1]^[46]^[50] Due to its ability to fix atmospheric nitrogen through symbiosis with Bradyrhizobium lupini, L. albus requires minimal nitrogen fertilization, typically 0–20 kg N/ha, to avoid suppressing nodulation. Phosphorus and potassium applications should be based on soil tests, with 20–40 kg P/ha and 40–80 kg K/ha recommended for deficient light or sandy soils to support root development and overall vigor. Inoculation of seeds with compatible Rhizobium strains prior to sowing enhances nitrogen fixation efficiency, particularly on soils lacking native symbionts, potentially contributing up to 40 kg N/ha to the crop.^[1]^[46]^[51] Effective weed control is crucial during the early growth stages before canopy closure, as L. albus competes poorly with grasses and broadleaf weeds. Pre-emergence herbicides such as pendimethalin, applied at labeled rates and incorporated shallowly, provide residual control of annual weeds when combined with mechanical tillage or harrowing at 2 km/h. Crop rotation with cereals or non-host crops helps suppress persistent weeds like grasses and reduces herbicide resistance risks, while mechanical cultivation between rows can minimize competition without chemical inputs.^[52]^[53]^[46] Irrigation management focuses on providing supplemental water during dry spells, particularly around flowering and pod fill, to boost seed yields on sandy soils, as L. albus has moderate drought tolerance via its deep taproot. However, excessive moisture must be avoided to prevent waterlogging, which promotes root rot from pathogens like Pleiochaeta setosa; well-drained soils and careful scheduling—aiming for 15–40 cm seasonal rainfall equivalent—are essential.^[1]^[54]^[55] Ongoing monitoring of soil conditions includes regular pH testing, with adjustment using lime if levels fall below 5.5 to mitigate aluminum toxicity, though L. albus thrives in slightly acidic soils (pH 5.5–7.3) and tolerates up to mildly alkaline conditions. Varietal selection should prioritize anthracnose-resistant or early-maturing types, such as FRIEDA or those suited to short growing seasons (110–130 frost-free days), to match regional climates and reduce disease pressure.^[1]^[46]^[49]

Harvesting and yield

Maturity in Lupinus albus is indicated by pods turning light brown and seeds hardening, typically occurring 120–180 days after sowing depending on environmental conditions and variety.^[56] Harvest timing for seed production is optimal when seed moisture content reaches 13–15%, usually in late summer such as August to September in temperate regions.^[1] For forage, green harvest can be performed earlier at 116–130 days to maximize nutritional quality before full pod maturity.^[1] Seed harvest is commonly conducted using direct combining with axial flow threshers to minimize seed damage, with rotor speeds set at 350–400 rpm and concave clearances of 40 mm for optimal performance.^[57] Desiccants, such as diquat or ethephon, are applied 3–4 weeks prior to harvest to promote uniform ripening and reduce pod shattering losses, particularly in regions with variable maturity.^[58] For whole-crop forage, green harvesting involves cutting plants at the ground level and ensiling, which preserves biomass while avoiding seed set.^[59] Under optimal conditions, seed yields range from 1.5 to 4 t/ha, influenced by soil fertility, rainfall, and cultivar selection, with potential up to 5 t/ha in high-input systems.^[57] Forage dry matter yields typically achieve 8–12 t/ha when harvested as whole-crop silage, providing high-protein feed.^[60] Post-harvest, seeds are dried to 12% moisture to prevent mold and stored in cool, dry conditions (below 15°C) where they maintain viability for 2–4 years.^[1] Yield is highly sensitive to abiotic factors; drought stress can reduce seed production by 30–50%, primarily through impaired pod set and seed filling, though cluster roots offer some tolerance in phosphorus-poor soils.^[44] Recent breeding efforts in the 2020s have developed hybrids showing up to 20% higher yields compared to traditional varieties, driven by improved heterosis in pod number and harvest index.^[61]

Pests and diseases

Insect pests

Lupinus albus is susceptible to several key insect pests that can compromise plant health and reduce productivity. Among these, aphids such as Acyrthosiphon pisum (pea aphid) colonize stems and leaves, feeding on sap and excreting honeydew that promotes sooty mold growth, which impairs photosynthesis.^[62]^[63] Similarly, lupin flower thrips (Frankliniella spp., including F. occidentalis) target blooms and young foliage, rasping tissues and causing distortion, discoloration, and reduced flower set.^[64]^[65] Seed weevils of the genus Sitona (e.g., S. griseus and S. gressorius, now classified under Charagmus) pose a significant threat, with adults notching leaves and scarring seed pods, while larvae bore into roots and nodules, disrupting nitrogen fixation and weakening plant vigor.^[66]^[46] These pests collectively lead to yield losses of up to 40% in untreated fields, with aphids also acting as vectors for viruses like bean yellow mosaic virus.^[46]^[63] Effective management relies on integrated pest management (IPM) strategies, including the use of resistant varieties such as certain L. albus cultivars that show tolerance to aphids.^[63]^[67] Neem-based sprays provide targeted control for aphids and thrips by disrupting feeding and reproduction, while encouraging beneficial insects like ladybugs (Coccinellidae) and lacewings enhances natural predation.^[63]^[67] For weevils, cultural practices such as border treatments and promoting ground-dwelling predators like carabid beetles are recommended over broad-spectrum insecticides.^[46]^[67] Pest pressure varies regionally, with Sitona weevils causing more severe damage in Europe—particularly in Poland and Mediterranean areas—compared to Australia, where aphids predominate but L. albus exhibits greater resistance.^[66]^[68]^[63]

Fungal pathogens

_Lupinus albus is susceptible to several fungal pathogens that can significantly impact yield, particularly in regions with favorable environmental conditions for disease development. Among the most destructive is anthracnose, caused by the hemibiotrophic fungus Colletotrichum lupini, which has emerged as a global threat since the late 20th century. This seed- and air-borne pathogen infects all growth stages, leading to stem twisting, necrotic lesions on stems, leaves, and pods, and ultimately pod rot that reduces seed quality and quantity. The fungus spreads internationally via contaminated seeds and locally through rain splash during the growing season, with high inoculum levels capable of causing total crop loss even from low initial infection rates.^[69]^[70] Root rots pose another major challenge, primarily driven by soil-borne fungi such as Fusarium solani and Phytophthora spp., which thrive in wet, poorly drained soils. Fusarium solani induces damping-off in seedlings and root decay in mature plants, resulting in wilting, yellowing foliage, and stunted growth as the vascular system is compromised. Similarly, Phytophthora root rot manifests as sudden patch death in spring, with rotten taproots, sparse lateral roots, and dark lesions on the stem base; waterlogging exacerbates zoospore release and infection. These rots weaken plant vigor, predisposing roots to secondary invasions by nematodes and other pathogens in saturated conditions.^[71]^[72] Rust, incited by the obligate parasite Uromyces lupinicola, is a common foliar disease in cool, humid climates. It produces small light-brown spots on leaves that develop into orange uredinial pustules on both surfaces, leading to chlorosis, premature defoliation, and reduced photosynthesis in severe epidemics. The pathogen's lifecycle involves wind-dispersed urediniospores that infect under prolonged leaf wetness, with rapid buildup during wet weather favoring outbreaks in dense canopies. While less devastating than anthracnose, rust can compound losses when co-occurring with other stresses.^[73]^[74] Effective management of these fungal pathogens relies on integrated strategies to minimize inoculum and limit spread. Planting certified, disease-free seeds is essential to prevent introduction of seed-borne pathogens like C. lupini, while crop rotation intervals of 3–4 years reduce soil inoculum from root rot agents. Fungicides such as azoxystrobin provide protective and curative control against foliar diseases including anthracnose and rust when applied preventively during flowering and podding stages, significantly lowering infection incidence. In recent years, studies from the 2020s have highlighted the clonal population structure of C. lupini, with four distinct lineages—including a globally dominant one—indicating limited genetic diversity that may accelerate adaptation and reduce efficacy of standard treatments in Mediterranean lupin fields.^[1]^[54]^[75]^[76]

Viral and bacterial diseases

Lupinus albus is susceptible to several viral and bacterial diseases that can significantly impact plant health and yield. Among the viral pathogens, bean yellow mosaic virus (BYMV), a member of the Potyvirus genus, is a primary concern. BYMV is primarily transmitted by aphids in a non-persistent manner, leading to symptoms such as mosaic patterns on leaves, chlorosis, leaf malformation, and overall stunted growth in infected plants.^[77] This virus can also be seed-borne in L. albus, allowing it to establish primary infection foci within crops from contaminated planting material.^[78] Infections often result in yield losses ranging from 20% to 50%, with more severe reductions possible under high aphid pressure or widespread infection.^[79] Cucumber mosaic virus (CMV), belonging to the Cucumovirus genus, also affects L. albus, though its impact is generally less pronounced compared to other lupin species due to the plant's immunity to seed transmission of this virus.^[80] CMV is transmitted by aphids similarly to BYMV and persists in numerous weed hosts, serving as reservoirs that facilitate spread to crops.^[81] In L. albus, infections can cause mottling, stunting, and reduced seed quality, particularly when plants are infected early in development, though outbreaks are sporadic and depend on vector abundance.^[77] On the bacterial front, Pseudomonas syringae pv. lupini causes brown spot disease, manifesting as necrotic brown spots on leaves and stems, especially under cool, moist conditions that favor bacterial proliferation.^[82] This pathogen spreads through splashing rain or overhead irrigation, entering plants via wounds, and is more prevalent in temperate growing regions.^[83] Unlike viral infections, bacterial diseases like this one are not seed-borne in L. albus, but contaminated equipment or debris can contribute to dissemination. Management of these viral and bacterial diseases emphasizes prevention over cure. For viruses, using certified virus-free seed is critical to minimize seed-borne introduction, particularly for BYMV.^[84] In-field practices include rogueing infected plants to reduce inoculum sources and deploying reflective mulches to deter aphid vectors.^[85] Bacterial infections lack effective curative bactericides, so control relies on sanitation measures such as crop rotation, removing plant debris, and avoiding overhead watering to limit moisture on foliage.^[86] Recent studies in 2024 and 2025 have highlighted emerging potyvirus complexes, including BYMV variants, in Australian lupin-growing regions, underscoring the need for ongoing surveillance and resistant cultivars to address evolving threats.^[84]

Uses

Human nutrition

Lupinus albus seeds, known as white lupin, have been consumed by humans for thousands of years, particularly in Mediterranean regions where they are prepared as traditional snacks. Dehulled seeds are typically soaked and boiled to remove bitter alkaloids, making them edible as a simple boiled or pickled product; in Egypt, this preparation is known as "termis," a popular street food enjoyed for its nutty flavor after brining in saltwater.^[87]^[88] Fermentation methods, such as bacterial or fungal processing, further enhance digestibility and flavor, while ground seeds are incorporated into lupin flour for baking breads and pasta, providing a gluten-free alternative in local cuisines.^[89]^[90] To reduce antinutrients like alkaloids and phytic acid, traditional debittering involves prolonged soaking in water followed by cooking, which significantly lowers toxicity while preserving nutritional value; modern industrial processes employ aqueous extraction or extrusion cooking to produce high-purity protein isolates suitable for fortification.^[91]^[90] In contemporary applications, L. albus features in innovative products such as lupin tofu, which substitutes up to 40% of soybean without compromising texture or acceptability, alongside meat analogues, extruded snacks, and protein-enriched beverages that leverage its neutral taste and emulsifying properties.^[92]^[93] As of 2025, lupin protein isolates have gained approval for export to the Chinese market, expanding their availability in plant-based products.^[94] Consumption of L. albus is rising in vegan and plant-based diets due to its complete amino acid profile and high protein content of 30–40%, which supports muscle maintenance and satiety comparable to animal sources.^[95]^[96] Culturally, it remains a staple festival food in Mediterranean countries like Greece and Italy, where brined seeds are shared during celebrations for their affordability and nutritional density.^[88] However, due to structural similarities in proteins, L. albus can trigger allergic reactions in approximately 20% of peanut-allergic individuals via cross-reactivity, necessitating clear labeling in products.^[97]^[98]

Animal feed

_Lupinus albus, particularly low-alkaloid sweet varieties, serves as a valuable protein and energy source in livestock diets, with the whole plant used for forage, silage, or hay, and seeds as a grain concentrate incorporated at 20–30% of the total diet to leverage its high oil content for energy.^[12]^[29] This legume's nitrogen-fixing properties also support sustainable feed production by reducing reliance on imported soy.^[99] In ruminants, L. albus enhances rumen degradable protein, promoting efficient fermentation and nutrient utilization. For dairy cows, inclusion levels up to 25% of dry matter have improved milk yield, with one study showing an increase of 2.8 kg/day when replacing 75% of soybean meal, alongside maintained milk composition.^[100]^[12] In sheep, supplementation at around 300 g dry matter per day boosts feed intake, digestibility, average daily gain, and carcass quality without differences among raw, soaked, or roasted forms.^[101] For non-ruminants, processed L. albus seeds (e.g., dehulled or extruded to minimize alkaloids) are suitable at 15–25% inclusion, supporting growth in pigs and poultry. In growing pigs, up to 20% replaces soybean meal effectively, improving performance and meat quality in breeds like Iberian pigs.^[102]^[12] Poultry, including broilers and laying hens, tolerate 15–30% after processing, with no anti-thyroid effects in sweet varieties and comparable weight gain, feed efficiency, and egg production to soy-based diets.^[103]^[29] High fiber in whole seeds can limit digestibility, particularly in non-ruminants, necessitating balancing with energy-dense grains to prevent digestive disturbances.^[12]^[29] Trials in the 2020s highlight L. albus's role in organic poultry feed, where 20–25% inclusion as a soy alternative sustains hen performance and reduces import dependency, aligning with sustainable farming goals.^[104]^[99]

Industrial and environmental applications

_Lupinus albus proteins, extracted from seeds or hulls, have been utilized in the development of biodegradable bioplastics and biomaterials, such as edible films and cellular scaffolds for biomedical and food packaging applications. These protein isolates contribute to the mechanical strength and barrier properties of the films, offering an eco-friendly alternative to synthetic polymers. Seed oil from L. albus, comprising 6–13% of seed weight, is rich in unsaturated fatty acids like oleic and erucic acids, making it suitable for incorporation into cosmetic formulations for skin barrier repair and soothing effects. While specific applications in adhesives remain underexplored for lupin proteins, their structural similarities to other plant proteins suggest potential in bio-based adhesives, though further research is needed. As a green manure, Lupinus albus enhances soil fertility through symbiotic nitrogen fixation, incorporating 100–200 kg N/ha into the soil upon incorporation, which reduces the need for synthetic fertilizers in subsequent crops. In crop rotations with wheat, L. albus has been shown to increase cereal yields by 15–20% due to improved nitrogen availability and soil structure, with nitrogen fertilization requirements lowered by 23–31 kg N/ha. This nitrogen-fixing capacity, primarily via root nodules hosting rhizobia bacteria, supports sustainable farming by minimizing nutrient leaching and enhancing organic matter. Lupinus albus demonstrates potential in phytoremediation, particularly for accumulating heavy metals such as zinc (Zn) and cadmium (Cd) primarily in its roots, aiding in the stabilization of contaminated soils. Field trials in the 2020s have explored its use in mine reclamation, where rotations with metal-accumulating plants leverage L. albus's tolerance to acidic, metal-rich tailings to prevent erosion and facilitate site restoration, with root uptake reaching up to 2 g/kg dry matter for Cd in young plants. The plant shows potential for significant removal of toxic metals like Cd and Zn from wastewater under low pH conditions, further supporting its role in phytostabilization strategies for polluted sites. Biomass from L. albus holds promise for biofuel production, particularly biogas via anaerobic digestion, yielding up to 532 L CH₄/kg volatile solids when co-digested with organic wastes, though its high food and feed value limits prioritization for energy applications. Additionally, ornamental varieties of L. albus are cultivated for aesthetic purposes in gardens, while as a cover crop, it excels in erosion control on marginal lands by stabilizing soil with its deep taproot system and dense growth, promoting biodiversity and preventing nutrient runoff in degraded areas.

Nutritional profile

Chemical composition

Lupinus albus seeds are characterized by a high macronutrient content on a dry weight basis, with protein comprising 30–40% of the total composition, making it a valuable plant-based protein source. This protein is particularly rich in lysine but relatively low in sulfur-containing amino acids such as methionine and cysteine. Carbohydrates account for 40–50%, predominantly in the form of dietary fiber (around 30–40%), including oligosaccharides and non-starchy polysaccharides, with low starch content (3–10%), while lipids range from 5–13%, primarily unsaturated fatty acids like oleic acid (approximately 50%), linoleic acid (20%), and linolenic acid (10%).^[4]^[105]^[88]^[12] Micronutrients in L. albus seeds include substantial levels of minerals such as potassium (1–1.5%), phosphorus (0.3–0.5%), and magnesium (0.1–0.2%), along with trace elements like manganese and zinc. Vitamins present are primarily from the B-group, including thiamine, niacin, and riboflavin, as well as vitamin E in the form of γ-tocopherol. These nutrient profiles contribute to the overall nutritional density of the seeds.^[4]^[106] Antinutrients in L. albus include quinolizidine alkaloids, which are present at 0.01–0.1% in sweet varieties, phytic acid at 0.4–1.2%, and low levels of tannins; these compounds can affect nutrient bioavailability but are significantly reduced through processing methods such as soaking or debittering. A 2025 review confirms that unprocessed bitter varieties retain high alkaloid levels, underscoring the need for proper debittering to avoid toxicity risks.^[4]^[107]^[108] The chemical composition exhibits variability depending on cultivar and processing; for instance, dehulling removes the hull, which constitutes about 20–25% of the whole seed and eliminates up to 70% of the fiber content, resulting in kernels with higher relative concentrations of protein and lipids. Recent analyses from 2023–2024 highlight cultivar-specific differences, such as varying mineral levels across genotypes like Multitalia and Luxor.^[12]^[106]^[105] Compared to soybeans, L. albus has a similar or slightly higher protein content but a lower proportion of methionine, though it excels in lysine abundance.^[88]

Health benefits and risks

Consumption of Lupinus albus seeds provides notable health benefits, primarily through its rich content of dietary fiber, proteins, and bioactive peptides that support metabolic and cardiovascular health. The soluble and insoluble fibers in lupin exert hypocholesterolemic effects by binding bile acids and reducing low-density lipoprotein (LDL) cholesterol levels, with clinical trials demonstrating significant reductions of approximately 5–10% in hypercholesterolemic individuals after regular intake.^[109]^[110] Additionally, lupin's low glycemic index, around 15, promotes stable blood glucose levels, making it suitable for managing diabetes; this is enhanced by peptide inhibitors like γ-conglutin, which stimulate glucose uptake and glycogen synthesis in muscle cells.^[4]^[111]^[112]^[113] Lupinus albus also contributes to cardiovascular protection, as its high potassium content (approximately 1,000 mg per 100 g) helps regulate blood pressure by counteracting sodium effects and promoting vasodilation.^[4] Clinical studies between 2017 and 2025 have shown that lupin-enriched diets lead to modest reductions in body weight and obesity markers, such as waist circumference, in overweight participants, attributing these outcomes to increased satiety from fiber and protein.^[114]^[115] Despite these advantages, Lupinus albus poses certain risks, particularly when seeds are not properly processed to remove bitter alkaloids. Unprocessed or bitter varieties can cause toxicity due to quinolizidine alkaloids like lupanine, resulting in symptoms such as nausea, vomiting, tremors, and mydriasis at doses exceeding 0.5 g/kg body weight of alkaloids-laden seeds.^[4]^[108] Allergenicity is another concern, with IgE-mediated reactions including anaphylaxis reported in sensitized individuals, often cross-reacting with peanut allergies.^[116]^[117] Furthermore, phytic acid in lupin seeds can chelate minerals like iron, zinc, and calcium, potentially inhibiting their absorption and contributing to deficiencies in high-consumption diets.^[118]^[119] To mitigate risks, consumption should be limited to 20–30 g per day of raw seed equivalent (or processed equivalents like flour), ensuring debittering through soaking or cooking; pregnant women are advised to avoid high doses due to limited safety data on alkaloid exposure during gestation.^[120]^[95] Recent systematic reviews, including a 2022 analysis of clinical trials, confirm lupin's benefits for hypertension management through blood pressure lowering, though efficacy varies by cultivar due to differences in alkaloid and nutrient profiles.^[121]^[4]

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Antidiabetic effects and mechanisms of action of γ-conglutin from ...
γ-conglutin from lupin possesses anti-diabetic properties. γ-conglutin peptides promote glucose uptake and glycogen synthesis in human muscle cells.
[111]
(PDF) Effects of lupin-enriched foods on body composition and ...
Aug 9, 2025 · However, such a diet may provide cardiovascular health benefits in terms of insulin sensitivity and blood pressure.
[112]
Lupins and Health Outcomes: A Systematic Literature Review - MDPI
Unintended weight loss occurred in 25% of studies. Whole lupin demonstrated more consistent beneficial effects for satiety, glycaemic control and blood pressure ...
[113]
Lupin poisoning: a review - Frontiers
Apr 8, 2025 · Poisoning occurred in all age groups and sexes equally. Most frequent symptoms were bilateral mydriasis (n = 25), xerostomia (n = 25), blurred ...
[114]
Occupational IgE-mediated allergy after exposure to lupine seed flour
The ingestion of lupine seed flour (LSF) has been reported as a cause of allergic reactions, particularly in patients sensitized to peanut, ...Missing: Lupin | Show results with:Lupin
[115]
Lupin allergy and lupin sensitization among patients with suspected ...
Clinically relevant lupin allergy often occurs in patients without atopic background or other food allergies, although lupin sensitization most commonly seems ...
[116]
Effect of instantaneous controlled pressure drop on the phytate ...
Phytic acid reduces mineral availability and hinders zinc and iron absorption in humans (Greiner, 2001; Greiner et al., 2001; Frokiaer, Barkholt, & Bagger ...
[117]
Effect of different debittering processes on mineral and phytic acid ...
Bitter lupin seeds cannot be consumed directly since its high toxic alkaloid content. Cooking and soaking are effective processes for removing these toxic ...
[118]
Lupin - Uses, Side Effects, and More - WebMD
The most common adverse effects are gas, bloating, and stomach pain. Lupin is LIKELY UNSAFE when products with toxic levels of alkaloids (sometimes called ...
[119]
Lupins and Health Outcomes: A Systematic Literature Review
Jan 13, 2022 · Whole lupin demonstrated more consistent beneficial effects for satiety, glycaemic control and blood pressure than lupin protein or lupin fibre.Missing: 2020-2025 | Show results with:2020-2025