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Polygonum

Polygonum L. is a of approximately 170 of flowering in the family and order , encompassing annual and perennial herbs, shrubs, and subshrubs with a nearly , primarily native to temperate and subtropical regions across , , and . These are distinguished by their prostrate to erect stems, often ribbed or angled and bearing ocreae—papery, cylindrical sheaths at the swollen nodes that disintegrate into fibers—and by alternate, simple leaves with entire margins, varying from linear to ovate or subround in shape. Flowers are small, bisexual, and typically white to pink, with five fused tepals forming a campanulate to urceolate , borne in axillary or terminal spike-like inflorescences of 1–10 flowers each; fruits are achenes, usually three-angled, glabrous, and enclosed or exserted from the . The genus is notable for its ecological diversity, occupying habitats from roadsides and slopes to wetlands, with some species like Polygonum aviculare serving as widespread weeds and others exhibiting medicinal properties due to bioactive compounds such as flavonoids (e.g., quercetin) and phenolic acids, traditionally used for treating conditions like hypertension and inflammation. Certain taxa have been reclassified into segregate genera like Persicaria and Fallopia in modern taxonomy, reflecting phylogenetic revisions based on molecular data.

Description

Morphology

Plants in the genus Polygonum are or rarely shrubs, or , characterized by stems that are prostrate to semi-erect, typically reaching up to 1 m in height. These commonly occur in marshy or wet places, with stems that are branched, slender to stout, and exhibit a jointed appearance due to swollen nodes bearing ocreae. The stems may be glabrous or hairy, often glandular, simple or branched, contributing to their variable growth forms from prostrate mats to upright clusters. Leaves of Polygonum are alternate, , and entire-margined, typically lanceolate to ovate or elliptic in and less than 2 cm long, though some species show broader variation up to 3 cm. They are articulated at the base and often glandular-punctate, with stipules fused to form a distinctive membranous ocrea—a cylindrical or funnel-shaped at the nodes that is silvery, , and frequently or fringed. This ocrea is a key diagnostic trait, enclosing the stem and distinguishing Polygonum from related genera in the family. Flowers are bisexual and small, measuring 2–5 mm, arranged in axillary or clusters such as , racemes, or capitula. The consists of 4–6 greenish-white to petaloid tepals that are not accrescent, surrounding 3–9 stamens and a superior with 2–3 styles ending in capitate stigmas. Fruits are achenes that are trigonous or , 2–4 mm long, black and shiny, and typically enclosed within the persistent , sometimes partly exserted. Morphological variation in Polygonum, such as differences in ocrea fringe, , and flower color, aids in but also underscores the genus's , with the ocrea presence serving as a primary taxonomic marker.

Reproduction

Polygonum species exhibit diverse reproductive strategies, encompassing both sexual and mechanisms that contribute to their adaptability across varied environments. Flowering typically occurs from late spring through autumn, varying by species and geographic region; for instance, flowers from March to November in . Inflorescences are often racemose or paniculate, with flowers arranged in axillary or terminal clusters that facilitate efficient . Pollination in Polygonum is primarily , mediated by small such as bees and flies that are attracted to and rewards in the hermaphroditic flowers. Many , including P. aviculare, are self-compatible and capable of autogamous , though cross- occurs via chasmogamous flowers visited by over 36 taxa in some regions. Following , fertilization involves a tricarpellate, superior that develops into a single-seeded , typically 3-angled and enclosed or exserted from the persistent . , an asexual seed formation bypassing fertilization, has been reported in certain populations of like P. aviculare, leading to larger embryos and in agamospermous fruits. Seed dispersal mechanisms in Polygonum emphasize zoochory, with achenes featuring sticky or hooked segments that adhere to animals, including , mammals, and , facilitating transport. While via elaiosomes is not widespread, some species rely on , , or human-mediated vectors like vehicles for broader dissemination. plays a key role in perennials through rhizomatous growth, enabling clonal spread via extensive that can extend meters and regenerate from fragments; annuals like P. aviculare may achieve limited clonal propagation through stem fragmentation and rooting at nodes. Chromosomally, Polygonum displays a base number of x = 10, with prevalent across the , ranging from diploid (2n = 20) to octoploid (2n = 80) levels, which enhances hybrid vigor and reproductive flexibility in polyploid complexes. This often correlates with shifts toward in some taxa, supporting the 's evolutionary success.

Taxonomy

Etymology

The genus name Polygonum is derived from the words polús (πολύς, meaning "many") and gónu (γόνυ, meaning "knee" or "joint"), alluding to the numerous swollen joints along the stems of plants in this genus. An alternative etymological interpretation connects it to gónos (γόνος, meaning "seed" or "offspring"), implying "many-seeded," though the jointed-stem reference is more widely accepted. The was formally established by in his 1753 work , where he included 14 , with later designated as the . Common names for Polygonum , such as and knotgrass, directly reflect the knobby, jointed stems that inspired the genus name. Regional variations include "birdweed" for P. aviculare, named for its appeal to seed-eating birds, and "allseed," highlighting the plant's abundant seed production. This nomenclature has ancient roots, with early herbalists like (c. 40–90 CE) describing similar jointed-stemmed in his , contributing to the enduring Greek-derived name.

Classification history

The Polygonum was established by in his (1753), where he included 14 under a broad circumscription that encompassed diverse herbaceous with jointed stems and ocreae. Over the , taxonomic expansions significantly broadened the genus, incorporating over 200 as new collections from temperate regions worldwide revealed morphological similarities, leading to a heterogeneous assemblage that included both annual and perennial forms. In the , efforts to refine the classification began with the recognition of subgenera such as Eupolygonum (encompassing typical knotweeds with persistent ocreae) and Duravia (characterized by North American species with distinct morphology and ocrea structure). Initial segregations occurred around the early 1900s, notably separating aquatic and amphibious species into the genus Mill., based on differences in and adaptations, as detailed in monographic treatments of the amphibious group. Key revisions in the late , led by Louis-P. Ronse De Craene in the 1980s, proposed further segregate genera like Adans. and Reynoutria Houtt., emphasizing floral characters such as morphology and arrangement to delineate monophyletic groups from the polyphyletic Polygonum sensu lato. A pivotal 2015 molecular phylogenetic study by Schuster et al., using and markers, narrowed the core Polygonum to approximately 65 , reassigning many others to genera such as Fallopia, Reynoutria, and Bistorta L., based on evidence of distinct evolutionary lineages within Polygoneae. Post-2020 updates include a 2017 taxonomic revision for Iran, which recognizes five genera (Aconogonon, Bistorta, Fallopia, Persicaria, and Polygonum s.str.) from the former Polygonum s.l., with a total of 25 species resolved through integrated morphological and distributional analyses, though ongoing debates persist regarding unresolved hybrid complexes and infraspecific variation. Recent discoveries as of 2025 include new species such as Polygonum elazigense (2024) from Turkey and Polygonum anatolicum (2025) from Anatolia, highlighting continued taxonomic refinements. Influential regional works, such as the Flora of North America (2005), accept approximately 33 species in the narrowed Polygonum for the region, focusing on sections Polygonum and Duravia. As of 2025, Plants of the World Online lists 171 accepted species under Polygonum, reflecting incomplete global segregations and varying adoption of molecular-based delimitations.

Phylogeny

Polygonum belongs to the subfamily Polygonoideae within the family and is classified in the Polygoneae. Molecular phylogenetic analyses place the genus as sister to the Duma-Atraphaxis (DAP ), which together forms a well-supported monophyletic group within the . Infrageneric of Polygonum recognizes several sections, including section Polygonum (cosmopolitan weedy species), section Duravia (North American endemics with prostrate habits), and section Pseudomollia. The core Polygonum , excluding aquatic segregates such as and Aconogonon (now in Koenigia), is monophyletic based on morphological and molecular data. Molecular evidence from ribosomal ITS and matK sequences supports the of the core Polygonum , with strong bootstrap values in maximum likelihood analyses. Many lineages within Polygonum exhibit polyploid origins, often involving allopolyploidy through interspecific hybridization, as evidenced by incongruent and phylogenies and elevated numbers (e.g., 2n=40–60 in the P. aviculare complex). Key evolutionary events include a radiation in temperate zones, as inferred from fossil-calibrated phylogenies of Polygoneae. Hybridization events have driven the formation of allopolyploids, contributing to species diversification and adaptive success in disturbed habitats. Recent studies using complete chloroplast genomes confirm the separation of Koenigia from Polygonum, with Koenigia forming a distinct clade sister to Fallopia in tribe Polygoneae. Analyses also reveal unresolved polytomies among Asian Polygonum species, indicating rapid diversification and potential reticulate evolution in that region.

Distribution and ecology

Geographic range

The genus Polygonum exhibits a nearly , though it is predominantly centered in north-temperate regions of the , with highest species diversity occurring in and . Globally, the genus comprises approximately 171 accepted , many of which are native to temperate zones across these areas. In , particularly , diversity is elevated, with estimates suggesting around 80 , reflecting the region's extensive temperate and montane habitats that support a wide array of annual and perennial forms. hosts about 33 , representing a significant portion of the genus's variation, including endemics restricted to the continent. Fewer are native to southern continents, such as (with scattered occurrences in ) and , where distributions are often disjunct and limited to higher elevations or coastal areas. Regional hotspots underscore the genus's concentration in specific biogeographic zones. The serves as a key center of diversity, hosting numerous species adapted to Mediterranean climates, while features high endemism in montane and environments. In western North America, section Duravia is entirely endemic, comprising around 20 species confined to arid and semi-arid regions from the southward. These patterns highlight a Holarctic dominance, with evolutionary origins tied to northern temperate floras, and occasional disjunct populations in southern hemispheres likely resulting from ancient vicariance events. Many Polygonum species have expanded beyond native ranges as widespread weeds in temperate and subtropical zones, facilitated by human activities such as trade and since the post-1500s era. For instance, P. aviculare has become pantropically distributed through unintentional dispersal, appearing in disturbed sites across North and South , , , and . These introductions have led to rapid range expansions in non-native temperate regions, where species often thrive in anthropogenic habitats like roadsides and fields, contributing to their global ubiquity today.

Habitats and adaptations

Species of the genus Polygonum predominantly occupy ruderal habitats, including disturbed soils, roadsides, agricultural fields, and riverbanks, where they thrive as opportunistic colonizers in nitrogen-rich and compacted ground. These environments often result from human activities or natural disturbances, allowing Polygonum to exploit open spaces with minimal competition. Certain species establish readily along disturbed riparian zones that experience periodic flooding but remain unsaturated during the growing season. Physiological adaptations enable Polygonum species to persist in these challenging conditions. Perennial taxa develop extensive rhizomatous systems and deep roots that enhance drought tolerance by accessing subsurface water reserves. Coastal species like Polygonum maritimum exhibit notable salt tolerance, maintaining growth under saline conditions through mechanisms that regulate uptake and osmotic balance in fore-dune habitats exposed to spray. Annual species demonstrate rapid rates, enabling quick establishment in ephemeral, disturbed sites before competitors dominate. In , Polygonum often functions as a pioneer , initiating recovery on bare or eroded substrates. Certain lead primary succession in volcanic or alluvial deposits, stabilizing through dense networks that prevent and facilitate later-stage . Additionally, interactions with arbuscular mycorrhizal fungi enhance uptake, particularly , in nutrient-poor soils; fungal hyphae extend the root system's reach, improving acquisition of immobile nutrients and supporting overall vigor. Under stress, Polygonum employs defensive strategies to maintain competitive edges. Allelopathy via root exudates inhibits neighboring plant germination and growth, as observed in Polygonum aviculare, where phenolic compounds in exudates suppress seedling development of co-occurring species. Certain taxa also hyperaccumulate , aiding survival in contaminated sites. The genus spans temperate to subtropical climates, with distributions influenced by regional variability. In the , Polygonum species occur from to altitudes exceeding 4000 m, adapting to cooler conditions through specialized reproductive traits.

Species

Accepted species

The genus Polygonum comprises approximately 170 accepted species worldwide, primarily annual and perennial herbs characterized by their cosmopolitan distribution and adaptation to diverse habitats. As of 2025, this includes 171 accepted species. These species are classified into several infrageneric sections, with Section Polygonum (also known as Avicularia) being the most widespread, encompassing nearly cosmopolitan annuals that often thrive as weeds in disturbed areas. Section Duravia, in contrast, includes North American perennials adapted to alpine and subalpine environments. Notable species in Section Polygonum include P. aviculare (common knotgrass), a highly variable annual weed distributed globally from temperate to subtropical regions, often found in roadsides, fields, and waste places. Another example is P. dentoceras (Small's jointweed), a rare U.S. endemic restricted to the central ridges of , where it grows in sandy soils and faces threats from habitat loss. In Section Duravia, P. douglasii (Douglas' knotweed) represents a mat-forming native to western , occurring in dry, rocky slopes and meadows from to . Australian diversity is exemplified by P. plebeium (in Section Polygonum), an annual herb endemic to temperate southeastern , typically in grasslands and woodlands. Species diversity within Polygonum is reflected in morphological variations, particularly in achene shape (ranging from trigonous to ) and ochrea size (from short and membranous to elongated and lacerate), which aid in taxonomic delimitation and to environmental stresses. Recent taxonomic revisions have added to this count, including P. uysalii ( Polygonum), a new species described in 2024 from southwestern , distinguished by its pubescent stems and compact inflorescences in Mediterranean-Irano-Turanian transition zones.

Formerly included species

Many species formerly classified under the genus Polygonum L. in the broad sense (Polygonum s.l.) have been transferred to other genera within the family based on molecular phylogenetic analyses that demonstrated the of Polygonum s.l.. These studies, utilizing genes such as matK, ndhF, and rbcL, along with ribosomal ITS regions, revealed distinct clades corresponding to separate evolutionary lineages, necessitating a revised to reflect monophyletic groups. Morphological characters, including differences in (e.g., vs. , herbaceous vs. woody), structure, and the presence or absence of certain ocreae features, further supported these segregations. As a result, the core Polygonum in the strict sense (Polygonum s.str.) has been narrowed to approximately 170 , primarily or with terete stems and simple leaves, down from over 300 species previously included. A significant portion of the reclassifications involved around 100–150 aquatic or perennial species moved to Persicaria Mill., which encompasses mostly wetland-adapted herbs with ocreae that are often fringed or lacerate. For example, Polygonum amphibium L. was transferred to Persicaria amphibia (L.) Delarbre, reflecting its amphibious habit and distinct achene morphology. Similarly, approximately 12–15 woody climbing species were reassigned to Fallopia Adans., characterized by twining stems and paniculate inflorescences; notable transfers include Polygonum convolvulus L. to Fallopia convolvulus (L.) Á.Löve & D.Löve. The Japanese knotweeds, previously under Polygonum sect. Tiniaria, numbering about 5–6 species, were placed in Reynoutria Houtt., distinguished by their robust rhizomes and dioecious flowers, such as P. cuspidatum Siebold & Zucc. becoming Reynoutria japonica Houtt. Additional genera received smaller but taxonomically important transfers. The alpine bistorts, around 40–50 species of rhizomatous perennials with cylindrical spikes, were segregated into Bistorta L., exemplified by P. bistorta L. now as Bistorta officinalis Delarbre. Buckwheats, comprising about 30 species with trimerous flowers and often lacking ocreae, form Fagopyrum Mill., including the cultivated P. fagopyrum L. transferred to Fagopyrum esculentum Moench. High-altitude taxa, roughly 40–60 species of cushion-forming herbs, were assigned to Koenigia L. emend. T.M. Schust. & Reveal, such as P. alpina All. becoming Koenigia alpina (All.) T.M. Schust. & Reveal. These changes, driven by evidence of non-monophyly from post-2011 phylogenetic updates, have clarified evolutionary relationships but left approximately 20 Asian endemic species in uncertain placements pending further genomic and morphological studies.

Uses

Medicinal applications

Polygonum species have been employed in across various cultures, particularly in and , for their , , and properties. In European folk medicine, (common knotweed) is traditionally used as a and for treating urinary tract and conditions, while in Asian , species like P. hydropiper address pain and allergies in and . In , P. multiflorum (now reclassified as multiflora), known as He Shou Wu, is valued for promoting hair growth, tonifying the liver and kidneys, and exhibiting anti-aging effects. Similarly, P. capitatum is utilized in for urologic disorders, including urinary tract . The pharmacological potential of Polygonum species stems from their rich array of bioactive compounds, including such as and , anthraquinones like , and . , found in the roots of several species, contributes to effects by promoting intestinal motility. In P. cuspidatum (Japanese knotweed, now ), stilbenes such as and polydatin are prominent, offering antioxidant and anti-inflammatory benefits. These compounds, particularly and phenolics like and caffeic acids, underpin the genus's traditional applications in and digestive ailments. Modern research has substantiated many traditional uses, with a 2024 comprehensive review highlighting the antioxidant and anticancer potentials of Polygonum extracts through and studies. like demonstrate free radical scavenging, while from P. cuspidatum shows promise in inhibiting proliferation. Clinical trials on P. cuspidatum extracts containing have reported reduced and in athletes after six weeks of supplementation, supporting cardiovascular health. Additionally, extracts of P. cuspidatum accelerate in models by enhancing TGF-β1 expression. Extracts of P. aviculare accelerate cutaneous in models by activating the Wnt/β-catenin pathway. Studies also explore P. cuspidatum for and due to its anti-inflammatory anthraquinones. Despite these benefits, safety concerns exist, particularly with long-term use. P. multiflorum has been associated with liver and toxicity, including idiosyncratic reported in clinical cases worldwide, attributed to anthraquinones like . Some contain oxalates, which may exacerbate kidney issues in susceptible individuals, and emodin can cause at high doses. Contraindications include , as laxative effects from anthraquinones may pose risks, and caution is advised for those with liver conditions. methods, such as steaming P. multiflorum, can reduce while preserving efficacy.

Culinary and agricultural roles

Several species of Polygonum have been utilized in culinary contexts, particularly during times of scarcity. The seeds of P. aviculare (prostrate ) are edible raw or ground into , serving as a or substitute in various regions. Young shoots of perennial species, such as P. sachalinense (giant ), are harvested in spring and can be eaten raw in salads or cooked as an alternative, though they possess an acidic, sometimes bitter flavor that may require preparation to enhance palatability. In , young shoots and leaves of P. aviculare are also consumed, often boiled or added to teas and dishes. In , Polygonum species have historical ties to cultivation through relatives like (Fagopyrum esculentum, formerly classified as Polygonum fagopyrum), which is grown for its grain-like seeds processed into nutritious used in breads, noodles, and pancakes worldwide. Certain Polygonum species, including giant , have been employed as and soil binders to prevent and improve in traditional farming systems. However, they are not widely adopted as dedicated cover crops due to their weedy nature. As forage, Polygonum provides moderate nutritional value for ; P. aviculare is grazed by in some rangelands and used as pig fodder in , offering protein and energy during early growth stages. Its abundance supports , with seeds consumed by and small mammals, but high seed production and persistence complicate weed in pastures and croplands. While oxalates in forages generally limit intake to avoid in ruminants, specific oxalate levels in Polygonum are not well-documented as prohibitive, though can exacerbate issues. Economically, reclassified species like remain significant for sustainable farming, with F. esculentum valued for its rapid growth and role in crop rotations that enhance without nitrogen fertilizers, indirectly supporting nitrogen-fixing companions like clovers through residue incorporation. Some Polygonum show potential in eco-friendly practices, such as associating with soil microbes for nutrient cycling, though direct is absent as they are non-leguminous. Despite these uses, Polygonum poses agricultural challenges as an invasive , forming dense mats that reduce crop yields by competing for light, water, and nutrients in over 60 affected , including grains and . Recent studies highlight allelopathic effects, where extracts from species like P. chinense inhibit in crops such as and ryegrass through compounds like loliolide and dehydrovomifoliol, exacerbating yield losses in invaded fields. Rhizomatous spread and flood dispersal further amplify control difficulties, necessitating integrated management to mitigate economic impacts.

Cultural significance

References in literature

In classical literature, the Greek physician and botanist Pedanius Dioscorides referenced knotgrass (Polygonum aviculare) in his seminal work De Materia Medica (circa 50–70 CE), noting its use as a medicinal plant. This mention established knotgrass as a staple in ancient pharmacopeias, with its jointed stems inspiring the genus name Polygonum, derived from Greek terms for "many knees." During the , alluded to knotgrass in (Act 3, Scene 2, circa 1595–1596), where mocks as a "minimus of hindering knot-grass made," invoking a that the plant could stunt growth or shrink the body, often tied to its use in charms for transformation or diminishment. In medieval and early modern herbal texts, extolled P. aviculare in his Complete Herbal (1653), praising its efficacy in "staunching of blood" from wounds, , or fluxes, recommending the juice drunk in wine or applied externally to halt and promote in ulcers. Modern literature has incorporated knotweeds into ecological narratives, with Japanese knotweed (, formerly Polygonum cuspidatum) serving as a for unchecked and environmental disruption; for instance, Sam Knight's "The Day of the Knotweed" (2015) describes its spread in as a resilient intruder altering landscapes, echoing broader themes of human impact on nature. In Asian cultural motifs, knotweeds appear in , with itadori (Japanese knotweed) associated with mischievous spirits in traditional . Botanical illustrations of Polygonum species in 19th-century texts, such as William Curtis's Botanical Magazine (e.g., P. griffithii, 1917 edition), blended scientific depiction with literary appreciation, portraying the plants as elegant subjects in accompaniments that highlighted their ornamental and narrative allure.

Symbolic and historical mentions

Polygonum species, particularly Japanese knotweed (, formerly Polygonum cuspidatum), have come to symbolize and in contemporary due to their aggressive invasive growth, which allows them to thrive in disturbed environments and even penetrate structures. This tenacity mirrors themes of unstoppable natural forces, often invoked in discussions of environmental adaptation and human-nature conflicts. In historical contexts, roots of have been utilized by Dong communities in to produce red dyes for clothing and food, a practice passed down orally across generations and reflecting cultural traditions predating synthetic alternatives. Archaeological evidence from Neolithic sites in Eastern reveals seeds of erect knotweed (Polygonum erectum), indicating early associations with human settlements as a potential domesticate or persistent , with morphological changes suggesting cultivation by around 3000 years ago. In modern times, Japanese knotweed serves as an emblem of invasiveness, frequently featured in UK media and policy discussions as a destructive force threatening and , leading to legal restrictions under the Wildlife and Countryside Act. This cultural notoriety extends to 2025 art exhibits, such as "Artists Stand Up for Invasive Species" at various galleries, where knotweed-inspired works explore themes of and ecological disruption in urban settings.

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