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Phaseolus
P. vulgaris
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Fabales
Family: Fabaceae
Subfamily: Faboideae
Subtribe: Phaseolinae
Genus: Phaseolus
L. (1753)
Type species
Phaseolus vulgaris
L.
Species

See text.

Synonyms[1]
  • Alepidocalyx Piper (1926)
  • Lipusa Alef. (1866)
  • Minkelersia M.Martens & Galeotti (1843)

Phaseolus (bean, wild bean)[2] is a genus of herbaceous to woody annual and perennial vines in the family Fabaceae containing about 70 plant species, all native to the Americas, primarily Mesoamerica.[3][4]

It is one of the most economically important legume genera. Five of the species have been domesticated since pre-Columbian times for their beans: P. acutifolius (tepary bean), P. coccineus (runner bean), P. dumosus (year bean), P. lunatus (lima bean), and P. vulgaris (common bean).[5][6] Most prominent among these is the common bean, P. vulgaris, which today is cultivated worldwide in tropical, semitropical, and temperate climates.

Ecology

[edit]

Phaseolus species are used as food plants by the larvae of some Lepidoptera species, including common swift, garden dart, ghost moth Hypercompe albicornis, H. icasia and the nutmeg.

Taxonomy

[edit]

The generic name Phaseolus was introduced by Linnaeus in 1753,[7] from the Latin phaseolus, a diminutive of phasēlus, in turn borrowed from Greek φάσηλος phasēlos used for either a species labelled as Vigna sinensis, or "calavance" which is an old name for the chickpea; that word itself is of unknown origin.[8][9] The Ancient Greeks probably referred to any bean in a pod as phasēlos including those species, mung bean as well as black gram which were brought to them from Asia during their time; the name extended to the New World beans with the introduction of common bean into Europe via Columbian exchange in the 16th century and later given the name Phaseolus vulgaris.[10]

Species

[edit]

Species have been organized into eight groups based on phylogenetic clades:[11][12][13]

Filiformis group

Leptostachyus group

Lunatus group

Pauciflorus group

Pedicellatus group

Polystachios group

Tuerckheimii group

Vulgaris group

Uncategorized


Previous classifications placed a number of other well-known legume species in this genus, but they were subsequently reassigned to the genus Vigna, sometimes necessitating a change of species name. For example, older literature refers to the mung bean as Phaseolus aureus, whereas more modern sources classify it as Vigna radiata. Similarly, the snail bean Vigna caracalla was discovered in 1753 and in 1970 moved from Phaseolus to Vigna. The modern understanding of Phaseolus indicates a genus endemic only to the New World.[3]

Allergenicity

[edit]

The Phaseolus plant has an OPALS plant allergy scale rating of 4 out of 10, indicating moderate potential to cause allergic reactions, exacerbated by over-use of the same plant throughout a garden. Leaves can cause skin rash and old plants often carry rust.[14]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Phaseolus

View on Grokipedia
from Grokipedia
Phaseolus is a of approximately 70–90 species of herbaceous to woody annual and vines belonging to the family , all native to the from the southward to northern and northern . The genus is monophyletic and primarily distributed in mild temperate to warm regions of the , where thrive in diverse habitats ranging from forests to open areas, often in frost-free environments during their growing period. Five within Phaseolus have been domesticated independently in the , making it a key for understanding and contributing significantly to global as sources of beans, pods, and . These domesticated include the common bean (P. vulgaris), (P. lunatus), runner bean (P. coccineus), year bean (P. polyanthus), and tepary bean (P. acutifolius), which together provide nutritious, protein-rich foods and exhibit ecological benefits such as in soils. The originated around 5–6 million years ago and has been the subject of extensive taxonomic revisions, with current classifications recognizing about 70–90 , most concentrated north of .

Overview

Description

Phaseolus species are primarily herbaceous vines or erect shrubs, exhibiting either annual or perennial growth habits. Many species are climbing plants that can reach heights of 3–5 meters by twining their stems around supports, while others form bushy, non-climbing habits under 1 meter in height. The leaves are alternate, stipulate, and typically trifoliate, with leaflets that are ovate to lanceolate in shape, often 5–12 cm long and featuring entire or slightly serrated margins. Root systems are fibrous and branched, featuring nodules formed through symbiosis with rhizobial bacteria, enabling biological nitrogen fixation that enhances soil fertility. Reproductive structures include papilionaceous flowers arranged in axillary racemes, which can be 10–35 cm long and bear 10–50 flowers each; corolla colors range from white and pink to red or purple, with individual flowers measuring 0.8–2.5 cm. The fruit is a pod, linear to oblong in shape and 5–30 cm in length (occasionally up to 40 cm), typically containing 4–10 seeds; pods dehisce tardily or remain indehiscent at maturity. Seeds are kidney-shaped (reniform), with a prominent hilum, and vary in size from small (under 5 mm) to large (over 20 mm), featuring diverse colors such as white, black, red, or mottled patterns. Across the genus, morphological variations are notable, particularly in pod texture—which can be smooth, glabrous, or pubescent—and seed characteristics, including size and coloration, reflecting adaptations to diverse habitats. For instance, climbing species often exhibit more vigorous twining stems and longer racemes compared to erect forms, while perennial taxa may develop woody bases or tuberous roots for persistence. These traits provide foundational diversity within the approximately 70 species of the genus.

Etymology

The genus name Phaseolus derives from the Latin phaseolus, a form of phaselus, which originates from the phāsēlos (φάσηλος), denoting a pod-bearing . In classical Greek and Roman texts, such as those by in his Enquiry into Plants and by in , phāsēlos referred to Old World podded plants, a unrelated to the New World beans later classified in this . The precise identification of the referred to by phāsēlos has been subject to scholarly debate. Carl Linnaeus formalized the use of Phaseolus in his 1753 work , applying the classical term to several bean species introduced to via the , thereby broadening its scope beyond its original connotations. This adoption marked a key moment in , aligning the name with the genus's diverse American origins while retaining its linguistic roots. The evolution of common names for Phaseolus species reflects post-1492 cultural exchanges. The English term "bean" stems from Old English bēan, originally specific to the broad bean (Vicia faba), but extended to Phaseolus after the genus's arrival in Europe. In Spanish-speaking regions, "frijol" evolved directly from Latin phaseolus, preserving the ancient derivation. French "haricot," by contrast, derives from Nahuatl ayacotl (a Mesoamerican term for bean) via early Spanish intermediaries, highlighting indigenous influences on European nomenclature.

Taxonomy and Phylogeny

Classification

Phaseolus is classified within the family (Leguminosae), subfamily , tribe , and subtribe Phaseolinae. Within this subtribe, Phaseolus is closely related to sister genera such as and Macroptilium, based on shared morphological and molecular characteristics including pod structure and chloroplast DNA inversions. This placement aligns with the IV (, which organizes the genus under the core in the order . The genus was initially established by in (1753), where it encompassed a broad assemblage of , including Old World species such as the mung bean (Phaseolus aureus). During the 19th and 20th centuries, taxonomic revisions progressively narrowed its scope; for instance, the mung bean was transferred to Vigna radiata in the 1970s as part of efforts to distinguish New World from Old World taxa based on morphological and biogeographical evidence. A pivotal redefinition occurred at the International Legume Conference in (1971), formalized by Maréchal et al. in 1978, which delimited Phaseolus to approximately 50 New World species, excluding pantropical elements reassigned to . Post-1980s molecular studies, including of nuclear and regions, have confirmed and refined this delimitation, recognizing 87 extant , all endemic to the and primarily distributed in . These analyses, building on earlier morphological work, emphasize Phaseolus as a monophyletic group within Phaseolinae, with no Old World representatives. Infrageneric divisions in Phaseolus are primarily informal, based on morphological traits such as shape, structure, and growth habit, rather than formal subgenera. For example, the Vulgaris group encompasses domesticated species like P. vulgaris and P. coccineus, characterized by phaseolin proteins and climbing habits. Modern taxonomic systems, including those informed by APG IV, do not recognize formal subgenera, opting instead for sectional groupings (e.g., sections Paniculati, Phaseoli) to reflect phylogenetic relationships without rigid hierarchies.

Phylogenetic Relationships

Molecular phylogenetic analyses of the genus Phaseolus have revealed a well-supported structure comprising eight major clades, designated as the Filiformis, Leptostachyus, Lunatus, Pauciflorus, Pedicellatus, Polystachios, Tuerckheimii, and Vulgaris clades, with all but five enigmatic species assigned to one of these groups. These clades were delineated using sequence data from nuclear ribosomal ITS/5.8S regions and the plastid trnK locus, providing resolution into two principal sister lineages: the basal A (primarily the Pauciflorus clade) and the more derived B (encompassing the remaining seven clades). Subsequent studies incorporating and nuclear markers have corroborated this framework, highlighting morphological, ecological, and biogeographical correlations among the clades. The evolutionary origins of Phaseolus trace back to a diversification within the , with the stem age of the estimated at approximately 6–8 million years ago and the crown radiation occurring within the last 6 million years, based on calibrations. Basal clades, such as Pauciflorus, are centered in higher-elevation regions of , including Mexico, the , and northern , while derived clades exhibit broader distributions extending from southward to the Andean regions of . This pattern suggests an initial establishment in northern latitudes followed by southward expansion and , with average clade ages around 2 million years reflecting a relatively recent burst of . Evidence from genomic analyses indicates ongoing hybridization and , particularly between wild and domesticated lineages within certain s, such as the Vulgaris where Phaseolus vulgaris exhibits asymmetric from wild Mesoamerican and Andean populations into domesticated forms. These events have contributed to the observed in cultivated beans, with introgressed segments influencing traits like disease resistance. Additionally, while most Phaseolus are diploid (2n=20 or 22), the genus shares an ancient whole-genome duplication event with related approximately 56 million years ago, and rare polyploid variants have been reported in some wild , potentially playing a role in adaptive evolution. events, such as those in the Vulgaris , occurred within this phylogenetic context but are explored further in discussions of cultivation history.

Distribution and Ecology

Native Range

The genus Phaseolus is native exclusively to the , with its primary wild distribution spanning from the , including and , southward through and to northern as far as northern and the Andean regions of , , , and . This Neotropical range also encompasses the Islands, , and the , though the genus is absent from and most of . The southwestern extent begins around southern Chihuahua in , reflecting a broad latitudinal gradient from subtropical to tropical zones. Areas of highest species diversity and endemism are concentrated in two key regions: , particularly the central Mexican highlands, the , and southward to , and the , including the northern Andes from to northern and the central Andes spanning northern , , and northwestern . These hotspots host the majority of the approximately 70 wild Phaseolus species, with alone accounting for significant phylogenetic richness. The two main centers of origin for domestication align with these diversity peaks: (encompassing and ) for P. vulgaris (common bean), P. lunatus (), and P. coccineus (runner bean), and the for separate domestications of P. vulgaris and P. lunatus, consistent with the independent domestications of the five species in the genus. Wild populations thrive in diverse habitats across this range, from arid and semi-arid environments in the southwestern U.S. and to humid tropical forests and montane ecosystems in the . Although Phaseolus species have naturalized outside their native range through escapes from cultivation, establishing feral populations in parts of , , , and beyond, the core wild distribution remains confined to the Neotropics, where genetic diversity and evolutionary history are preserved. This post-Columbian expansion, initiated by European explorers in the , has not altered the foundational American origins of the genus.

Ecological Role

Species in the genus Phaseolus play a significant role in nutrient cycling through symbiotic , forming mutualistic associations with rhizobial such as species in root nodules. This process converts atmospheric into , which the utilize for growth, while the receive carbohydrates from the host; as a result, Phaseolus enhances in legume-dominated ecosystems, contributing to overall availability without synthetic inputs. In natural settings, this symbiosis supports the persistence of Phaseolus in nitrogen-poor soils across Mesoamerican ranges and beyond. Phaseolus species interact with a variety of pollinators and herbivores, influencing their reproductive success and population dynamics. Flowers attract bees and butterflies, with butterflies and moths often being the most frequent visitors, though bees provide more effective pollen transfer due to their foraging behavior. As hosts, wild Phaseolus support Lepidoptera larvae, such as those of cutworm moths (Agrotis spp., including garden dart), which feed on foliage, and various bean beetles like the Mexican bean beetle (Epilachna varivestis) and bean leaf beetle (Ceratoma trifurcata), whose larvae consume roots and pods, thereby integrating into food webs as prey for predators. In ecosystems, Phaseolus provides services such as weed suppression through its climbing or vining habit, which forms dense canopies that shade out competing vegetation in grasslands and forest edges. Wild species contribute to biodiversity in tropical forests and grasslands, where they occupy understory niches and support associated flora and fauna, enhancing habitat complexity. However, wild Phaseolus populations face vulnerability to climate change, particularly drought, which reduces germination and survival rates in aridifying habitats, potentially altering ecosystem dynamics.

Species Diversity

Overall Diversity

The genus Phaseolus encompasses approximately 87 accepted , predominantly herbaceous , many of which are climbing vines adapted to various tropical and subtropical habitats. These exhibit a range of growth forms, from annuals to short-lived perennials, but the herbaceous climbing habit predominates, facilitating their ecological roles in diverse ecosystems across the . Phylogenetic analyses have organized most Phaseolus species into eight informal groups—Filiformis, Leptostachyus, Lunatus, Pauciflorus, Pedicellatus, Polystachios, Tuerckheimii, and Vulgaris—based on shared morphological traits such as seed coat patterns, pod structure, and flower characteristics, with approximately five remaining ungrouped or enigmatic. These groupings reflect evolutionary divergences within the , supported by molecular and morphological data, and highlight the of Phaseolus in response to varied environmental pressures. In terms of geographic diversity, most Phaseolus species are found in and (Mesoamerica), representing the primary center of origin and highest , with significant numbers occurring in and fewer in northern regions. is particularly elevated in montane regions, such as the Sierra Madre and Talamanca ranges, where habitat specificity and isolation have driven speciation in specialized microhabitats.

Notable Species

Phaseolus vulgaris, commonly known as the common bean, is one of the most important domesticated worldwide, serving as a crop in many regions due to its high protein content and versatility in diets. It was independently domesticated approximately 8,000 years ago in two distinct centers: (modern-day ) and the (northern and ), leading to diverse landraces adapted to various environments. Notable varieties include kidney beans, characterized by their large, red, kidney-shaped seeds, and black beans, with small, shiny black seeds, both widely used in culinary applications such as soups, stews, and salads. Under favorable irrigated conditions, seed yields typically range from 1 to 3 tons per , contributing to its economic significance in global . Phaseolus lunatus, the , is distinguished by its large, flat seeds, which can measure up to several centimeters in length, making it a valuable source of carbohydrates and proteins in traditional cuisines. occurred independently in two regions: the and , with archaeological evidence dating back several thousand years, resulting in a wide array of seed colors and sizes across its cultivars. Wild forms contain cyanogenic glycosides, such as and lotaustralin, which release toxic upon tissue damage, serving as a defense mechanism against herbivores but requiring processing like soaking or cooking in domesticated varieties to ensure safety. Phaseolus coccineus, known as the runner bean, is a vigorous climbing vine prized for both its edible pods and seeds as well as its ornamental scarlet flowers, often grown in gardens for aesthetic appeal. Introduced to in the early 16th century following the , it quickly became popular in temperate regions due to its relative cold tolerance compared to other Phaseolus species, allowing cultivation in cooler climates where it can overwinter as a in mild areas. This adaptability, combined with its dual use as a and nitrogen-fixing , underscores its significance in small-scale farming and home . Phaseolus acutifolius, the tepary bean, is renowned for its exceptional drought resistance, thriving in arid environments with annual rainfall as low as 200-400 mm, far surpassing the tolerance of the common bean. Native to the and , where it has been cultivated by for millennia, this species produces the smallest seeds in the , with 100-seed weights often below 10 grams, enabling rapid maturation and efficient use during dry spells. Its resilience to heat and low moisture makes it a promising crop for climate-challenged in desert regions. Phaseolus dumosus (syn. P. polyanthus), or the year bean, is a rare Mesoamerican species endemic to a narrow region in and southern , where it grows as an annual or short-lived perennial in highland ecosystems. Positioned as an intermediate form between wild and fully domesticated beans, it exhibits partial loss of pod dehiscence and larger seeds than its wild relatives, traits that facilitated its limited cultivation by indigenous farmers as a supplementary crop alongside and squash. Its close genetic ties to P. vulgaris and P. coccineus highlight its potential as a genetic resource for breeding drought- and disease-resistant varieties, though its restricted distribution limits broader use.

Cultivation and Uses

Domestication History

The domestication of Phaseolus species, particularly the common bean () and lima bean (), occurred independently in multiple centers across the pre-Columbian , marking one of the earliest examples of plant cultivation in the . Archaeological and genetic evidence indicates that these processes began approximately 10,000 to 8,000 years ago, with the earliest signs in the Andean region of . For P. vulgaris, domestication events took place separately in (centered in the Lerma-Santiago basin of west-central ) and the southern (Peru to northern ), while P. lunatus underwent dual domestication in and the (western to ). These independent origins are supported by distinct gene pools identified through molecular markers and chloroplast DNA analyses, reflecting geographic isolation and human selection pressures in diverse agroecological niches. Key domestication traits were selectively favored to enhance yield and usability, including non-shattering pods to prevent seed loss during harvest, larger seed size for improved and storage, reduced for synchronized germination, and diminished —such as lower levels of cyanogenic glycosides in P. lunatus—to make the beans safer for consumption. These changes are evident in archaeobotanical remains from key sites: in the , early domesticated P. vulgaris seeds dating to around 8,600–7,000 calibrated years (cal BP) were found in the Nanchoc Valley of northern , with additional evidence from Guitarrero (~5,000 cal BP) showing larger, non-shattering forms; in , P. vulgaris remains from the Valley (~2,300–2,400 cal BP) and (~8,300 cal BP) exhibit similar modifications, while linguistic evidence from Proto-Mayan languages suggests bean cultivation predated these finds by about 1,100 years. For P. lunatus, archaeological records from Mexican and Ecuadorian sites confirm parallel trait evolution around 5,000–8,000 years ago. Following the European contact in , Phaseolus species spread rapidly to the via the , with P. vulgaris introduced to by Spanish explorers in the early . Historical accounts note that encountered and likely transported beans during his 1519–1521 conquest of , leading to cultivation in and by the 1520s; by 1542, the first European illustrations of the common bean appeared in herbal texts, and the crop adapted to Mediterranean and African climates through selection. This global dissemination diversified diets and agriculture in the , with beans establishing as staples in regions like and within decades.

Agricultural Practices

Phaseolus crops, particularly the common bean (Phaseolus vulgaris), thrive in warm climates with average temperatures ranging from 15°C to 30°C, as these conditions support optimal growth and pod development. The plants require well-drained, fertile soils with a pH between 6.0 and 7.5 to prevent nutrient deficiencies and root issues, with sandy loam textures preferred for good aeration and water retention. Germination occurs best at soil temperatures above 18°C, and seedlings emerge within 7-10 days at around 16-20°C, though frost sensitivity necessitates planting after the last freeze. Planting densities typically range from 20 to 40 plants per square meter for bush varieties, allowing efficient light capture and yield maximization while minimizing competition. Crop rotation plays a key role in sustainable cultivation, with Phaseolus species often rotated with cereals like or to leverage the ' nitrogen-fixing capabilities via rhizobial , thereby enhancing for subsequent non-legume crops without synthetic inputs. systems, such as the traditional Mesoamerican , integrate beans with and squash, where beans climb maize stalks for support and contribute nitrogen, improving overall system productivity and resilience. Breeding programs focus on developing hybrid varieties with enhanced disease resistance, such as to anthracnose (Colletotrichum lindemuthianum), through marker-assisted selection and pyramiding of resistance genes from wild relatives. Efforts also target drought tolerance via conventional breeding and genetic transformation trials, including the introduction of genes like HVA1 from barley to confer abiotic stress resistance, though widespread GMO adoption remains limited. These varieties, including upright architectures for mechanical harvest, have improved yields under variable conditions while maintaining adaptability across domesticated species like P. lunatus. Major challenges include pests such as (Aphis fabae) and Mexican bean beetles (Epilachna varivestis), which defoliate plants and can cause up to 37% yield losses if unmanaged. Diseases like (Uromyces appendiculatus), anthracnose, and bean common mosaic virus (BCMV) further threaten production, with BCMV leading to stunting and mosaic symptoms that reduce pod quality. exacerbates these issues, with rising temperatures and erratic rainfall projected to decrease common yields by 20-50% in tropical regions by mid-century due to heat stress during flowering and increased pressure. and resilient cultivars are essential to mitigate these impacts.

Economic Importance

Phaseolus species, particularly P. vulgaris, represent a cornerstone of global , with dry production reaching approximately 28 million tonnes in 2022, primarily from cultivation on approximately 37 million hectares worldwide. Leading producers include , , and , which together account for a significant portion of output, followed by and ; P. vulgaris dominates this production, comprising the vast majority of dry beans harvested. This scale underscores the crop's role as a key , contributing to systems in both developing and developed regions, with annual global yields reflecting steady growth driven by demand for affordable protein sources. Economically, Phaseolus beans drive a market valued at around USD 9 billion in 2023, encompassing production, processing, and trade, with major exports from countries like Myanmar, Argentina, and the United States supporting international supply chains. The primary use is as dry beans, which serve as a nutrient-dense staple providing 20-25% protein content, making them an essential vegetarian protein source in diverse diets; secondary applications include fresh pods and sprouts for culinary versatility, while non-food roles encompass ornamental varieties in horticulture and green manure for soil improvement in sustainable farming. Trade volumes highlight their accessibility, with smallholder farmers benefiting from cash crop opportunities that enhance household incomes in regions like sub-Saharan Africa and Latin America. Culturally, Phaseolus beans hold profound significance as dietary staples, integral to traditional cuisines that reflect historical and regional identities; in Mexico, refried beans (frijoles refritos) embody Mesoamerican heritage, often accompanying tortillas and rice as a daily essential. In the Andean region, dishes like Chilean porotos granados—a stew of cranberry beans, corn, and squash—exemplify indigenous agricultural synergies, providing sustenance and cultural continuity for communities. Beyond cuisine, these beans bolster food security for over 300 million people, particularly smallholders reliant on them for nutrition and economic stability amid climate challenges.

Health and Safety

Allergenicity

Phaseolus species exhibit low to moderate allergenic potential in humans, primarily through IgE-mediated responses to specific proteins. The Ogren Plant Allergy Scale (OPALS) assigns the genus a rating of 4 out of 10 for pollen-related risk, signifying moderate potential that can increase with excessive planting of the same species in landscapes. Moderate occurs with other family members due to shared homologous proteins, though clinical reactions vary widely among sensitized individuals. Key allergens include the vicilin-like phaseolin (approximately 47-50 ) and such as phytohemagglutinin (PHA), both of which are heat-stable and capable of eliciting strong IgE binding. These proteins provoke reactions, manifesting as cutaneous symptoms like urticaria and , gastrointestinal distress, and respiratory manifestations including wheezing or in severe cases. For instance, phaseolin and PHA were isolated and confirmed as major IgE-reactive components in patients experiencing following ingestion. , while also implicated in non-allergic toxicities, contribute to allergic sensitization via their glycoprotein-binding properties. Allergy prevalence to Phaseolus is relatively low and varies by population and region. rates to common bean have been reported as high as 22% in certain at-risk groups such as and patients in , but clinical reactions remain uncommon globally, with common bean considered a moderate sensitizer compared to or soy. Reactions are more frequently reported in atopic individuals, but true anaphylactic events remain uncommon globally.

Toxicity

The primary toxin in Phaseolus species, particularly in raw or undercooked kidney beans (Phaseolus vulgaris), is (PHA), a that causes acute gastrointestinal distress including , , and upon ingestion. Symptoms can occur within 1-3 hours of consumption, with as few as four to five raw beans sufficient to trigger poisoning in humans due to the toxin's hemagglutinating and enterotoxic effects. In animal studies, diets containing 1% raw kidney beans have led to mortality in rats within weeks, underscoring PHA's potency, though human cases are typically non-fatal and resolve within hours to days. PHA is heat-labile and can be fully inactivated by proper cooking, specifically soaked beans for at least 10 minutes at 100°C, which denatures the and eliminates its . Soaking beans overnight prior to reduces cooking time and further aids in reduction, while methods like slow cooking without prior may fail to reach sufficient temperatures, exacerbating risks. Another toxic compound occurs in wild varieties of lima beans (Phaseolus lunatus), where cyanogenic glycosides such as and lotaustralin are present and can release (HCN) upon enzymatic during tissue damage or digestion. This cyanogenesis serves as a defense mechanism against herbivores but poses risks in high concentrations, potentially causing cellular asphyxiation; however, domesticated cultivars have significantly reduced levels, with commercial varieties limited to less than 200 mg HCN/kg to ensure safety. Adherence to safety guidelines—such as thorough soaking and boiling—is essential to mitigate these risks, as evidenced by rare but notable outbreaks, including over 50 suspected incidents of PHA poisoning in the UK between 1976 and 1989, where undercooked beans led to symptoms in affected individuals.

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