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Lathyrus

Lathyrus is a genus of approximately 150 species of annual and perennial herbaceous plants in the legume family Fabaceae, commonly known as vetchlings or peavines. These plants are characterized by their often climbing or sprawling growth habit, supported by branched tendrils derived from modified leaflets, with even-pinnate leaves, showy papilionaceous flowers in shades of white, yellow, pink, purple, or blue, and flattened, dehiscent legume pods containing 3–15 seeds. Native primarily to temperate regions of the Northern Hemisphere, including Europe, Asia, North Africa, and North America, the genus has a center of diversity in the Mediterranean and Irano-Turanian regions, with some species extending to tropical East Africa and South America; many are widely naturalized or cultivated globally. The exhibits significant morphological diversity, including variations in winginess, complexity (from 0 to 18 leaflets), and shape, with most diploid (2n = ) and unarmed, though some bear unicellular or multicellular hairs. Economically, Lathyrus hold value as ornamentals, such as the fragrant (L. odoratus), bred for over two centuries into thousands of cultivars with vibrant flowers; as drought- and disease-resistant crops like the grass pea (L. sativus), a protein-rich used in arid regions since times; and as plants including L. cicera and L. hirsutus. However, several , particularly L. sativus, contain β-N-oxalyl-L-α,β-diaminopropionic acid (β-ODAP), a that can cause —a debilitating —upon excessive consumption, limiting their widespread adoption despite nutritional benefits. Taxonomically, Lathyrus belongs to the tribe Fabeae and is the second-largest in the tribe after Vicia, with ongoing revisions reflecting its complex evolutionary history, including rare and adaptations to diverse habitats from coastal dunes to meadows. While many species serve ecological roles in and as food sources, invasive tendencies in some naturalized taxa, like the everlasting (L. latifolius), pose challenges in certain ecosystems.

Taxonomy

Etymology and History

The genus name Lathyrus derives from the lathuros (λάθυρος), an ancient term for a type of vetch or pea-like , as used by the botanist in his Enquiry into Plants around 300 BCE to describe wild such as those resembling Lathyrus sativus or L. maritimus. The etymology's precise origin remains uncertain, but one interpretation links it to Greek la (meaning "very" or "enhance") combined with thouros (stimulant), possibly alluding to the historical use of Lathyrus seeds in medicinal preparations like the purgative "farina ervi." Species of Lathyrus have been recognized and utilized since ancient times in Mediterranean , where such as L. sativus (grass pea) and L. cicera (chickling vetch) were cultivated for food and fodder, as evidenced by archaeological remains and texts from the and islands dating back over 3,500 years. The genus was formally established in modern botany by in his (1753), where he described several species and distinguished Lathyrus from related genera like Pisum (), though early synonyms persisted due to morphological similarities. Throughout the , European botanical explorations significantly expanded the known diversity of Lathyrus, with collectors documenting new species from regions including the Mediterranean, , and , leading to monographic works such as Alvin C. Beal's A Study of the Genus Lathyrus () that synthesized classifications based on morphology and geography. These efforts built on Linnaean foundations, incorporating observations from global expeditions, and paved the way for later taxonomic revisions, such as Frances K. Kupicha's sectional division of the genus into 13 groups in 1983.

Classification

The genus Lathyrus is placed within the kingdom Plantae, phylum Tracheophyta, class Magnoliopsida, order Fabales, family Fabaceae, subfamily Faboideae, tribe Fabeae, and genus Lathyrus L. This hierarchical classification reflects its position among the , a diverse characterized by nitrogen-fixing capabilities and compound leaves. The genus was formally established by in (1753), where it was described as including climbing herbs with pinnate leaves and tendrils, and it holds conserved status (nom. cons.) under the International Code of Nomenclature for algae, fungi, and plants to maintain nomenclatural stability. Phylogenetically, Lathyrus belongs to the tribe Fabeae, where it forms a closely related clade with genera such as Vicia (vetches) and Pisum (peas), supported by shared morphological traits like stipitate gynophores and similar floral structures. Molecular studies using chloroplast DNA sequences have confirmed the monophyly of Lathyrus, resolving its relationships within Fabeae and indicating a temperate origin with subsequent diversification. Infrageneric classification, primarily following Kupicha's 1983 revision, divides the genus into 13 sections based on morphological features including seed surface patterns, pod dehiscence, and style twisting; notable sections include Lathyrus (with flattened pods and smooth seeds) and Orobus (with terete stems and pubescent pods), the latter formerly recognized as a distinct genus from which several species were transferred in the 19th and 20th centuries. The taxonomy of Lathyrus is accepted as circumscribed by authoritative sources like and , encompassing approximately species (as of 2025) with no major generic synonyms, though sectional synonyms persist from pre-molecular era classifications. Recent molecular phylogenies have largely validated Kupicha's sections but suggest minor refinements, such as merging or redefining groups like Orobon based on DNA evidence.

Description

Morphological Characteristics

Lathyrus species are herbaceous , either or , that exhibit a range of growth habits including erect, sprawling, or climbing forms, and are unarmed throughout. The stems are typically angled or winged, glabrous or pubescent, and in climbing species can reach heights of up to 3 meters, supported by twining tendrils. Vegetatively, the leaves are alternate and even-pinnate, with 0 to 9 pairs of leaflets (0–18 leaflets) that are entire-margined and glabrous or pubescent on the surfaces; the rachis often ends in a branched or simple , while stipules are present and range from small to foliose, often semisagittate at the base. The roots form a system or arise from rhizomes, featuring nitrogen-fixing nodules formed in symbiosis with rhizobial , which enable atmospheric . Reproductively, the flowers are zygomorphic and papilionaceous, arranged in axillary racemes of 1 to 20 blooms or occasionally solitary; each corolla measures 6–30 mm and displays colors ranging from white and yellow to pink and purple. The fruits are linear, dehiscent legumes that are flattened or terete, typically containing 3–15 seeds per pod.

Growth Forms

Lathyrus species exhibit diverse life cycle strategies adapted to varying environmental pressures, primarily comprising annuals and perennials. Annual species, such as Lathyrus odoratus and L. sativus, complete their life cycle within a single growing season, emphasizing rapid seed production to exploit ephemeral resources in xeric or disturbed habitats. Perennials, including L. latifolius and L. sylvestris, achieve longevity through rhizomatous or taprooted growth, allowing persistence over multiple years and repeated reproduction. Architectural adaptations in Lathyrus reflect functional strategies for resource acquisition and . Climbing species, like many perennials, employ twining tendrils derived from the main leaf axis to ascend supports, facilitating access to in dense vegetation. Erect or prostrate forms predominate in open areas, with stems that are angled, flanged, or winged to enhance stability or dispersal. Seasonal growth patterns, including vegetative elongation and flowering initiation, are modulated by photoperiod, as seen in L. odoratus where longer day lengths (12–24 hours) accelerate bud formation following . Development proceeds through distinct stages, beginning with of hard-coated that exhibit due to impermeable seed coats, as documented in L. maritimus. or natural abrasion breaks this barrier, enabling and emergence, often requiring 7–15 days under optimal conditions. The vegetative phase follows with rapid stem and elongation, supported by tendrils for forms. Reproductive maturity typically occurs within one for annuals, while perennials reach it in 1–2 years, sustaining productivity across seasons.

Diversity

Species Diversity

The genus Lathyrus encompasses approximately 183 accepted species worldwide, as documented in the database, reflecting a stable taxonomic consensus into 2025. These species exhibit a predominantly temperate distribution, with the highest concentrations in the : roughly 52 species native to , 30 to , 78 to , alongside smaller numbers in tropical (24 species) and temperate (24 species). Infragenerically, Lathyrus is classified into 13 sections primarily based on morphological traits such as structure, torsion, and characteristics, as established by Kupicha in —a system that integrates geographic patterns and remains widely adopted. This classification highlights regional endemism, particularly in the , where the and Pontic regions serve as a primary center of diversity, hosting numerous species restricted to specific microhabitats like coastal dunes and montane grasslands. Conservation concerns affect a subset of Lathyrus species, with several classified as threatened primarily due to habitat loss from , , and ; notable examples include L. belinensis ( in ) and L. biflorus (rare in ). Overall, endemics in hotspots like the Mediterranean face elevated risks, though global assessments indicate that most species remain of least concern. Trends in taxonomy show limited new discoveries since 2000, with only a handful of species described, such as L. tefennicus (2011, endemic to ), L. alamutensis (2008, ), and L. turcicus (2023, endemic to ), underscoring a maturing understanding of the rather than ongoing rapid .

Notable Species

Lathyrus odoratus, commonly known as the sweet pea, is a scrambling annual climber native to and , prized for its intensely fragrant flowers that bloom in a wide array of colors including pink, purple, white, and red. It typically grows to 1-2.5 meters tall, supported by tendrils, and has been extensively cultivated since the 17th century for ornamental gardens, leading to numerous cultivars selected for enhanced fragrance and flower form. Lathyrus sativus, or , is a drought-tolerant annual widely grown as a pulse crop in and , valued for its high protein content and resilience to harsh environmental conditions such as flooding and poor soils. However, it contains the β-N-oxalyl-L-α,β-diaminopropionic acid (β-ODAP), which can cause —a debilitating neurodegenerative disorder—upon excessive consumption, limiting its use despite breeding efforts to reduce toxin levels. Lathyrus tuberosus, the tuberous or earthnut , is a climbing geophyte with tubers that have served as a historical source in , where they were collected and consumed raw or cooked for their nutty flavor, particularly during times of scarcity. Native from to , , and , it features pinkish-purple flowers and spreads via rhizomatous tubers, though its use has declined with modern agriculture. Lathyrus vernus, known as spring vetchling or spring bitter vetch, is a long-lived found in the of temperate forests across to and , emerging early in spring with yellow to purple flowers before leafing out. It reproduces sexually after 10-15 years and exhibits in response to spring temperatures, contributing to its persistence in shaded, nutrient-poor habitats. Lathyrus japonicus, the beach pea, is a prostrate to climbing perennial adapted to coastal environments, ranging from regions to , , and western , where it stabilizes sandy beaches and dunes with its nitrogen-fixing roots. This early successional species produces pale purple flowers and is sensitive to human disturbance like trampling, making it a focus for in fragile maritime ecosystems such as the Great Lakes shores.

Distribution and Habitat

Geographic Range

The genus Lathyrus is predominantly native to temperate regions of the , with its approximately 150–170 exhibiting a Holarctic distribution pattern characterized by high in extratropical zones. Primary centers of and are centered in southwest and , from where the genus expanded to the during prehistoric times, as evidenced by Neolithic remains dating back to 8200–7500 BCE in the Valley and 6000 BCE in the . Hotspots of species richness occur in the , which hosts 61–76 taxa, including key areas like the , southern-central , and northeastern , and the encompassing the , where alone records 58 species. Secondary centers include western , with around 29 native species primarily in section Orobus, and temperate , supporting about 23 species in section Notolathyrus, mainly in central and . Few species venture into tropical areas, limited to high-altitude populations in , underscoring the genus's adaptation to cooler, temperate climates across and . Biogeographic patterns reveal disjunct distributions, with distinct Asian and American clades reflecting ancient vicariance events, alongside Holarctic dominance that links Eurasian and North American floras. Introduced ranges have expanded globally through human activities, including ornamental escapes and agricultural dissemination; for instance, L. latifolius has naturalized widely in North America, forming invasive populations, while species like L. sativus have been introduced to South Asia, Ethiopia, Australia, and Latin America as crops or fodder. These introductions often blur native boundaries in disturbed habitats across southern Europe, North America, and beyond.

Habitat Preferences

Lathyrus species generally thrive in temperate climates characterized by cool winters and moderate summers, with many exhibiting to semi-arid conditions and low temperatures. They prefer well-drained soils ranging from neutral to , often succeeding in loamy or clay-loam textures with levels between 5.8 and 7.5, though some adapt to mildly acidic substrates. As nitrogen-fixing , they are particularly associated with nitrogen-poor soils, enhancing their suitability for marginal lands where soil fertility is low. These plants occupy diverse microhabitats, including woodland edges, open grasslands, montane meadows, riverbanks, and coastal areas. In forested regions, species like L. vernus favor semi-shaded understories and moist forest margins, while grassland and meadow forms such as L. pratensis occur in open, sunny pastures and fallow fields. Coastal species, exemplified by L. japonicus, are found along seashores, dunes, and marshes, contributing to dune stabilization in windy, exposed environments. Many also appear on rocky slopes and roadsides, particularly in Mediterranean and Irano-Turanian hotspots. Adaptations to these habitats include climbing or sprawling growth forms supported by tendrils, which allow species to access in denser , and deep systems in drought-tolerant forms like L. sativus for water access in arid soils. Forest-dwelling species demonstrate , while coastal variants such as L. japonicus exhibit tolerance, enabling survival in saline conditions. rhizomatous or tuberous in many species, such as L. tuberosus, aid drought endurance and nutrient storage in variable environments.

Ecology

Biotic Interactions

Lathyrus species exhibit a range of biotic interactions that facilitate their reproduction and survival in natural ecosystems. Pollination in the genus is predominantly entomophilous, with flowers attracting insects through nectar rewards and visual cues. Many species, such as Lathyrus japonicus and L. odoratus, are primarily pollinated by bees, including bumblebees (Bombus spp.) and mason bees (Osmia sp.), which access the nectar via specialized floral structures like the keel and wings that require sufficient body mass to open. These interactions promote outcrossing, as some species display self-incompatibility mechanisms that prevent self-fertilization, ensuring genetic diversity. Herbivory plays a significant role in Lathyrus , with several species serving as larval host for . For instance, Lathyrus linifolius and L. pratensis are key hosts for wood white butterflies (Leptidea sinapis and L. reali), where larval feeding influences growth rates, decisions, and related to and immunity. Similarly, L. odoratus supports the tailed-blue butterfly (Cupido amyntula). Seed occurs mainly through ballistic dehiscence of mature pods, which explosively eject seeds up to several meters, as observed in species like L. pratensis. Animal-mediated dispersal also contributes, with seeds occasionally ingested and excreted by herbivores, aiding wider distribution. Symbiotic relationships enhance nutrient acquisition in Lathyrus. As , species form nitrogen-fixing nodules with , primarily Rhizobium leguminosarum and R. pisi, enabling atmospheric conversion to for plant use; this is well-documented in L. sativus and other annuals, supporting growth in nitrogen-poor soils. Additionally, arbuscular mycorrhizal fungi (AMF) associate with roots for improved and water uptake, with diverse communities identified in L. cicera and synergistic effects observed when combined with in L. sativus under stress conditions. These mutualisms underscore Lathyrus' adaptation to varied ecosystems.

Abiotic Responses

Lathyrus species exhibit varied temperature sensitivities depending on their ecological niches, with montane and temperate s often showing growth limitations under elevated temperatures. For instance, in Lathyrus vernus, a common in cooler habitats, growth during early is primarily constrained by low temperatures. Studies on related species like Lathyrus sativus (grass pea) indicate that high temperatures during reproductive stages can significantly reduce accumulation and yield, highlighting vulnerability in warming scenarios for montane-adapted taxa. Conversely, many Lathyrus species demonstrate notable frost tolerance, hardy in USDA zones 3–9 (surviving down to approximately -35°C in zone 3), as seen in Lathyrus latifolius (zones 4–9) and L. vernus (zones 3–9), which harden off through physiological adjustments to endure winter conditions without significant damage. Drought resilience is a prominent trait among xerophytic Lathyrus species, particularly L. sativus, which maintains productivity in arid environments through adaptive physiological mechanisms. This species exhibits high water-use efficiency, with roots extending deeply to access soil moisture, enabling survival under prolonged dry spells where other legumes fail. A key response to water deficit involves rapid stomatal closure mediated by abscisic acid (ABA), which reduces transpiration by 50–70% while preserving photosynthetic rates longer than in sensitive crops, thereby minimizing biomass loss. Xerophytic accessions of L. sativus show less yield reduction under moderate drought than less adapted varieties, underscoring their role in water-limited habitats. Responses to other abiotic stressors, such as extremes, further illustrate the genus's environmental adaptability. Lathyrus species generally avoid highly acidic soils (pH <5.5), where aluminum inhibits elongation and uptake; optimal occurs at pH 6.0–7.5, as in L. odoratus and L. sativus, with extending to mildly alkaline conditions up to pH 8.0 in well-drained loams. These preferences align with their native habitats in neutral to soils, where extremes disrupt balance.

Cultivation and Uses

Ornamental Cultivation

Lathyrus species, particularly the annual (L. odoratus) and its hybrids, are widely cultivated as ornamental climbers and for their fragrant, colorful flowers. These add vertical interest to gardens, often trained on trellises or obelisks to showcase their cascading blooms in , , , , and bicolors. Popular hybrids derived from L. odoratus dominate ornamental use due to their enhanced flower size, scent, and vigor compared to wild forms. Propagation of ornamental Lathyrus primarily occurs through seed sowing, with optimal timing in late winter or early indoors, or directly outdoors after the last frost. Seeds should be soaked in water for 24 hours to soften the hard coat and nicked with a if needed to promote , which typically occurs in 7-15 days at temperatures around 15-20°C. For climbers like L. odoratus hybrids, young plants require sturdy support structures such as netting, canes, or wire frames installed at planting to prevent damage to tender stems as they reach heights of 1.5-2 meters. Ideal growing conditions for ornamental Lathyrus include full sun exposure for at least six hours daily to maximize flowering, paired with fertile, well-drained loamy enriched with or well-rotted to a depth of 30-60 cm. Regular watering is essential to maintain consistent , especially during dry spells, but overwatering should be avoided to prevent ; aim for deep watering at the base in the morning. For bushier forms, pinch out the growing tips when plants reach 10-15 cm in height to encourage lateral branching and more abundant blooms. The ornamental cultivation of Lathyrus traces back to the late , but Victorian-era breeding in the transformed L. odoratus into a horticultural favorite through selective hybridization for expanded color ranges and larger flowers. Breeders like Henry Eckford, often called the "Father of the Sweet Pea," introduced over 100 Grandiflora varieties by 1901, featuring plain-petaled blooms with intensified fragrance and hues from soft pastels to vivid scarlets. Modern varieties build on this legacy, incorporating traits like wavey-edged "Spencer" petals from early 20th-century developments and ongoing selections for extended bloom periods. Disease management in ornamental Lathyrus focuses on preventing common issues like (Erysiphe pisi), which thrives in humid, poorly ventilated conditions and appears as white powdery patches on leaves. Selecting resistant hybrids, such as those derived from crosses with L. belinensis, provides qualitative resistance governed by a single dominant , where fungal spores collapse post-germination on treated plants. Cultural practices, including good air circulation via spacing plants 20-30 cm apart and avoiding overhead watering, further reduce mildew incidence in susceptible L. odoratus cultivars.

Edible and Agricultural Uses

Lathyrus sativus, commonly known as grass pea, serves as a key drought-resistant pulse crop in regions like and , where it thrives under harsh conditions such as low rainfall and poor soils, acting as an "insurance crop" during famines and environmental stresses. In , its cultivation is particularly valued for tolerance to , , waterlogging, and low , contributing significantly to local food security. Similarly, L. tuberosus, or tuberous vetchling, has been utilized as a through its edible tubers, which provide a nutrient-dense option in times of scarcity, especially in historical contexts. Nutritionally, L. sativus seeds are high in protein, containing approximately 27% on a dry weight basis, making it a valuable complement to cereal-based diets by providing essential that enhance overall when consumed together. This balance is crucial in staple diets across and , where grass pea supplements the lysine-deficient proteins in grains like or . The tubers of L. tuberosus are also protein-rich, offering up to 25% protein content, alongside vitamins and , positioning it as a resilient source in marginal lands. In agricultural practices, Lathyrus species like L. sativus are integrated into crop s to leverage their nitrogen-fixing capabilities through , which enrich and reduce the need for synthetic fertilizers, thereby supporting sustainable farming systems. This process improves and nutrient availability for subsequent crops, such as cereals, while minimizing and pressures in rotation sequences. Yields for L. sativus typically from 0.8 to 2 tons per hectare under rainfed conditions, varying by region and management, with higher outputs in optimal settings like parts of exceeding 2 tons per hectare. Historically, Lathyrus cultivation dates back to the period in the , with archaeological evidence of consumption in Late Bronze and sites in , indicating its role as a staple alongside early cereals. Breeding efforts for L. sativus have focused on developing low-toxin varieties to mitigate health risks from beta-ODAP while preserving agronomic traits, using classical crossing with low-toxin relatives like L. cicer and mutational techniques to achieve ODAP levels below 0.06%. As of 2024, the developed varieties like BARI-3 with 0.086% β-ODAP. These advancements, including gene-editing informed by sequencing, aim to integrate grass pea into modern as a climate-resilient crop for food and in arid and semi-arid zones. Such varieties enhance its viability in diverse agroecosystems, promoting and reducing reliance on high-input crops.

Toxicity and Health Risks

Certain species of Lathyrus, particularly L. sativus (grass pea), contain the β-N-oxalyl-L-α,β-diaminopropionic acid (β-ODAP) in their seeds, which is responsible for the neuromuscular disorder known as . This non-protein is present in varying concentrations across approximately 21 Lathyrus species, with L. sativus exhibiting notably high levels, often ranging from 0.5% to 2.5% of seed dry weight. β-ODAP acts primarily by overstimulating glutamate receptors in the , leading to and degeneration of motor neurons. Health risks from Lathyrus consumption include both acute and chronic effects. Acute exposure, typically from moderate ingestion, can cause gastrointestinal distress such as and . Chronic overconsumption, especially when L. sativus forms a staple in the diet, results in —a progressive, irreversible spastic paraparesis characterized by lower limb weakness, stiffness, and eventual . Historical epidemics underscore these dangers; for instance, a major outbreak in during 1944–1945 affected thousands due to widespread reliance on Lathyrus seeds amid food shortages, with many cases leading to permanent . As of 2023, surveys in report a population-level prevalence of 2.4% neurolathyrism in grass pea-dependent areas. Mitigation strategies focus on reducing β-ODAP levels through . Soaking in for 12–24 hours before cooking can leach out up to 50% of the toxin, while boiling soaked further reduces it by 65–70%. Safe consumption limits are generally set at β-ODAP concentrations below 0.15% in or no more than 500 mg per day per person for 2–3 months to avoid neurotoxic effects. In veterinary contexts, Lathyrus species pose risks to ; for example, ingestion of L. hirsutus by can induce stringhalt-like abnormalities, while L. sylvestris consumption in sheep leads to neurologic disease associated with , such as and depression.

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