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Stolon

In biology, a stolon is a slender horizontal branch or connection between parts of an organism, serving to propagate the organism asexually or form colonies.^[1] In botany, a stolon, also known as a runner, is typically an aboveground horizontal stem that grows along or above the soil surface, producing adventitious roots and shoots at its nodes or tip to form genetically identical daughter plants.^[2]^[3] Stolons arise from adventitious buds in the plant's crown zone and feature elongated internodes with rudimentary leaves, enabling rapid vegetative propagation.^[3]^[2] Unlike rhizomes, which are persistent underground stems used for storage and survival, stolons are chiefly aboveground and more ephemeral, adapted for surface spreading.^[2]^[3] This is evident in species like strawberries (Fragaria × ananassa), where stolons extend from the parent, root at nodes, and form new plantlets under longer daylight and warmer conditions.^[4]^[5] In grasses, they enable invasive growth, as in bermudagrass (Cynodon dactylon), zoysia (Zoysia spp.), and buffalo grass (Bouteloua dactyloides), aiding lateral spread and persistence.^[3] White clover (Trifolium repens) also uses stolons for forage spreading.^[2] The term also applies in mycology to horizontal hyphae for fungal spread, in zoology to connections in colonial invertebrates like corals, and in paleontology to fossilized structures indicating ancient clonal growth. The botanical usage is the most common.^[2]

In Botany

Definition and Morphology

A stolon, also known as a runner, is defined in botany as a slender, horizontal stem that grows along or above the soil surface, and in some cases below ground (e.g., in potato), and facilitates vegetative propagation by producing adventitious roots and new shoots at its nodes. This structure allows plants to colonize new areas asexually without relying on seeds, enabling rapid spread in favorable environments. Stolons typically emerge from the base of the parent plant and extend outward, rooting at intervals to form genetically identical daughter plants. Morphologically, stolons exhibit distinct features adapted for horizontal growth and propagation. They possess long internodes, often several centimeters to meters in length, and a thin diameter, usually less than 5 mm, which contrasts with the more robust stems of upright plants. Their growth is plagiotropic, meaning it is oriented horizontally due to gravitropism inhibition, with nodes specialized for the formation of adventitious roots downward and axillary buds upward that develop into new shoots. In cross-section, stolons reveal a simple vascular organization, with vascular bundles arranged in a ring surrounding a central pith (in eudicots) or scattered throughout the ground tissue (in monocots), supporting efficient transport of water, nutrients, and photosynthates along their length. This anatomy, observable in histological studies, underscores their role as modified stems rather than roots or leaves. Stolons differ from related underground structures like rhizomes, which are typically thicker, with shorter internodes and subterranean growth, whereas stolons remain above ground and prioritize elongation for surface coverage. The term "stolon" derives from the Latin word stolo, meaning a sucker or shoot, reflecting its branching propagation function. Developmentally, stolons initiate from axillary buds at the plant's base, where environmental cues trigger their emergence and horizontal orientation. Elongation proceeds through cell division and expansion in the internodes, driven by auxin gradients that polarize transport from the shoot apex toward the nodes, promoting root initiation at those points. Hormonal influences, such as auxin and cytokinin interactions, further modulate this growth pattern.

Functions and Ecological Role

Stolons serve a primary reproductive function in plants through asexual propagation, producing genetically identical clonal offspring that promote uniformity within populations and enable rapid colonization of suitable habitats. This vegetative mode of reproduction allows plants to bypass sexual processes, facilitating efficient spread in favorable conditions without reliance on pollinators or seed dispersal mechanisms.^[6] Ecologically, stolons enhance resource allocation by enabling foraging in heterogeneous or patchy environments, where connected ramets can translocate nutrients, water, and carbohydrates to support growth in resource-poor areas. This clonal integration also confers drought tolerance, as stolons and their root connections allow water and assimilates to be shared between ramets, buffering stress and improving survival in arid conditions. Furthermore, stolons contribute to invasiveness in non-native species, where clonal fragments store higher levels of sugars and starch compared to native counterparts, enhancing regeneration and competitive dominance post-disturbance.^[7]^[8]^[9] Hormonal regulation is central to stolon development, with auxin (indole-3-acetic acid, IAA) promoting internode elongation and adventitious root initiation at stolon tips. Gibberellins, particularly GA3, drive the growth phase by stimulating cell expansion and meristem activity. Abscisic acid (ABA) levels rise during transitions to tuber formation or dormancy, inhibiting further elongation, while the IAA/ABA ratio influences stolon size, with higher ratios favoring larger structures as shown in recent studies on potato and strawberry.^[10]^[11]^[12] Environmental factors modulate stolon responses through hormonal pathways; elevated CO2 concentrations enhance growth by increasing auxin-mediated elongation and soluble carbohydrate accumulation, thereby boosting photosynthetic efficiency and resource availability. Sucrose-induced stolon initiation, as revealed by 2025 transcriptomic analyses, involves coordinated hormonal signaling, including upregulated auxin and gibberellin pathways that trigger meristem differentiation under high sugar conditions.^[13]

Examples and Applications

Common stoloniferous plants include the wild strawberry (Fragaria vesca), which produces long, above-ground stolons that root at nodes to form new plants, facilitating fruit production in cultivated systems.^[14] The spider plant (Chlorophytum comosum) similarly develops arching stolons bearing plantlets, making it a popular houseplant for easy propagation.^[14] Creeping bentgrass (Agrostis stolonifera) spreads via prostrate stolons, contributing to dense turf in lawns and golf courses.^[15] In the potato (Solanum tuberosum), stolons elongate underground to swell into tubers, serving as the primary harvestable organ.^[16] In agriculture, stolons enable efficient vegetative propagation; for instance, protocols using stolon explants from the fern Hypolepis punctata have achieved high-efficiency in vitro regeneration, with up to 75.56% green globular body induction and 98.89% sporophyte regeneration rates, supporting ornamental and medicinal cultivation.^[17] Timing of stolon removal in everbearing strawberries like 'Albion' optimizes daughter plant production and quality; stolon removal every 7 days resulted in 16 daughter plants per mother plant with larger individual size, while removal every 63 days yielded 102 daughter plants but reduced individual dry weight to 0.51 g per plant, with less frequent removals enhancing total biomass accumulation.^[18] Stolons play a key role in conservation and ecology, particularly in grassland restoration, where stoloniferous grasses like bermudagrass (Cynodon dactylon) promote rapid vegetative spread and soil stabilization, reducing runoff by 32% and sediment yield by 60% in degraded sites.^[19] However, C. dactylon's invasiveness requires management, such as persistent manual removal of stolons and rhizomes or competitive exclusion through dense native planting, to prevent dominance in natural areas.^[20] Genotypic variations in stolon traits are evident in zoysiagrass (Zoysia spp.) progeny, where initiation rates range from 2.2 to 8.6 stolons per week and elongation from 18.8 to 65.1 mm per week, influencing establishment speed and turf coverage across cultivars.

In Mycology

Structure and Development

In mycology, a stolon is defined as a horizontal hypha that connects sporangiophores or fruiting bodies in certain fungi, particularly within the Mucoromycota (formerly Zygomycota), and is often associated with rhizoids that provide anchorage to the substrate.^[21]^[22] These structures facilitate the lateral spread of the fungal thallus across surfaces. Stolons are particularly characteristic of fungi in the order Mucorales, such as Rhizopus species.^[23] Structurally, fungal stolons can be aerial or bound to the substrate, appearing as stouter, slightly arched hyphae with a larger diameter compared to other mycelial elements; they are typically multinucleate and coenocytic, lacking septa in actively growing regions, though some may develop incomplete septa.^[24]^[21] In mucoromycetes such as Rhizopus stolonifer, stolons exhibit branching patterns that allow for extensive colonization, with nodes where rhizoids—short, root-like hyphae—emerge to anchor the fungus and absorb nutrients.^[22]^[24] Developmentally, stolons emerge from the mycelium at points of contact with the substrate, elongating horizontally through apical tip growth before arching to form new nodes upon recontacting the surface, from which additional stolons and sporangiophores differentiate to support reproductive structures.^[24]^[21] This process enables rapid lateral expansion of the colony. For instance, endophytic studies on the orchid Epipogium aphyllum have noted the absence of fungal colonization in its stolons, highlighting differences in substrate preferences between fungal and plant structures.^[25] Unlike plant stolons, which are macroscopic, stem-like organs capable of producing adventitious roots and shoots, fungal stolons are microscopic, filamentous hyphae composed of chitin rather than cellulose, serving primarily for vegetative spread and anchorage without vascular tissue or photosynthetic function.^[24]^[22]

Role in Fungal Reproduction

In fungal reproduction, particularly among saprotrophic molds in the phylum Mucoromycota (formerly Zygomycota) such as Rhizopus stolonifer, stolons serve as horizontal hyphal runners that facilitate vegetative growth and support the development of reproductive structures. These structures anchor the mycelium to the substrate via associated rhizoids at nodal points, while enabling the transport of nutrients and water absorbed from the environment to upright sporangiophores. This nutrient distribution is crucial for sustaining energy-intensive processes like spore production, allowing the fungus to expand its colony laterally across organic substrates without relying solely on spore dispersal.^[26]^[27] Stolons play a key role in asexual reproduction by connecting multiple sporangia, which are sac-like structures at the tips of sporangiophores that produce and release sporangiospores. These spores, formed mitotically within the sporangia, enable rapid propagation in favorable conditions, with stolons positioning reproductive sites optimally for spore release and germination. Additionally, stolons contribute to vegetative propagation through fragmentation; breakage of the stolon due to mechanical stress or environmental factors results in independent segments that can develop into new mycelial colonies, promoting clonal expansion.^[26]^[28] While stolons have an indirect supportive function in sexual reproduction by linking compatible hyphae of opposite mating types, their primary contribution remains to asexual mechanisms, which predominate in nutrient-rich, ephemeral environments. Direct research on stolon-specific roles in fungal reproduction has been limited since 2020, with most studies focusing on broader mycelial networks in saprotrophic molds rather than stolons per se; however, their established function in substrate colonization underscores an ecological importance in resource acquisition and decomposition. For instance, in bioremediation contexts, fungal mycelial systems including stolons aid in breaking down organic pollutants, though stolon contributions are not distinctly quantified in recent phytoremediation analyses.^[26]^[27]

In Zoology

In Colonial Invertebrates

In colonial invertebrates, stolons are horizontal, tube-like structures that connect individual modules within a colony, enabling asexual growth and resource sharing. A prominent example occurs in hydrozoans (class Hydrozoa, phylum Cnidaria), where stolons, often called hydrorhizae, form a network of hollow, ectodermal tubes adhering to the substrate. These originate from the basal disc of a founding polyp and facilitate the budding of new polyps or hydrocladia (feeding branches), allowing colonies to spread across surfaces like rocks or algae. Structurally, hydrozoan stolons are covered by a non-living chitinous exoskeleton called perisarc, enclosing coelenteron extensions for nutrient transport and gas exchange. This modular system supports rapid colonization and resilience to fragmentation, as seen in species like Hydractinia echinata, where stolons interconnect gastrozooids and gonozooids for efficient foraging and reproduction in intertidal zones.^[29] In colonial invertebrates, particularly bryozoans (also known as ectoprocts), stolons are defined as horizontal, tubular extensions of the body wall that serve as connectors between individual zooids, facilitating colony growth through asexual budding.^[30] These structures are characteristic of stoloniferous colony forms, where zooids arise separately along the stolon rather than in compact, adjacent arrays typical of non-stoloniferous bryozoans.^[31] Stolons enable the modular construction of colonies, allowing for efficient expansion in marine environments.^[32] Structurally, stolons in bryozoans are typically hollow tubes composed of chitinous material; for instance, in ctenostome bryozoans, they feature a thin chitinous wall enclosing coelomic fluid, with proximal regions often exhibiting cuticular wrinkles for flexibility and attachment to substrates.^[31] They connect to autozooids (feeding units) via septal pore plates, permitting the exchange of coelomic fluid and nutrients through a stolonal funiculus.^[31] In some forms, such as endolithic bryozoans, stolons include specialized tubulets—small tubes extending toward the substrate surface—to aid in boring and colony establishment within hard substrates.^[33] These elongated kenozooids (non-feeding modules) often branch orthogonally, with distal expansions containing muscles for controlled growth along surfaces like polychaete tubes or rocks.^[34] The primary function of stolons is to support asexual reproduction through stoloniferous budding, where new zooids or satellite colonies develop from buds along the stolon, enabling rapid horizontal colony expansion and repair of damaged sections.^[31] This modular growth strategy allows bryozoan colonies to adapt to heterogeneous substrates, outcompete other sessile organisms, and recover from partial mortality, contributing to their resilience in dynamic marine habitats.^[32] In fouling communities, stolons promote the spread of colonies across artificial surfaces, influencing biodiversity and biofouling dynamics.^[35] Representative examples include the ctenostome Hypophorella expansa, where stolons form a network within host polychaete tubes, bearing lateral autozooids that alternate sides for efficient two-dimensional dispersal, and the cheilostome Bugulina stolonifera (formerly Bugula stolonifera), in which rhizoid-like stolons spread across substrates to bud secondary upright colonies, playing a key role in overwintering and seasonal recolonization.^[34]^[36] These structures underscore the adaptive modularity of bryozoan colonies in marine ecosystems.^[35]

In Segmented Worms

In segmented worms, particularly within the polychaete family Syllidae, stolonization represents a distinctive reproductive strategy involving the formation of a posterior stolon—a specialized segment dedicated to gamete production—that detaches from the parent body through a process known as schizogamy, or asexual fission followed by sexual reproduction.^[37] This mechanism allows the benthic "stock" or parent worm to remain in its habitat while the stolon serves as a mobile reproductive unit, often participating in synchronized swarming events for mating.^[38] Unlike vegetative stolons in botany or mycology, which facilitate clonal growth, syllid stolons are temporary and epitokous, prioritizing gamete dispersal over long-term survival.^[39] The stolonization process begins with the differentiation of posterior segments, where gonad primordia emerge early, followed by sex-specific development of testes or ovaries.^[39] In Megasyllis nipponica, this unfolds across six stages: initial gut kinking and primordia formation (Stage 1), sex determination and gamete maturation (Stage 2), eye development (Stage 3), antenna formation (Stage 4), elongation of swimming setae (Stage 5), and final detachment via muscular vibration (Stage 6).^[39] Gene expression patterns, such as peaks in vasa and piwi during early gametogenesis and rising nanos levels later, underpin this progression.^[39] Environmental cues, including lunar phases and moonlight, trigger synchronous stolonization in many syllids, coordinating population-level swarming for enhanced fertilization success; for instance, exposure to moonlight induces the process in M. nipponica and related species.^[38] Post-detachment, the stock regenerates its posterior end, potentially undergoing multiple cycles.^[37] Structurally, the stolon forms as an autonomous, worm-like appendage with a simplified digestive tube, a dorsally positioned cerebral ganglion in its anterior segment, paired enlarged eyes, short antennae, and elongated swimming notochaetae for pelagic locomotion.^[39] It lacks a functional mouth, pharynx, or proventricle, emphasizing its short-lived role in reproduction rather than feeding.^[39] This contrasts sharply with the stock's complex anatomy, highlighting the stolon's adaptation for gamete release and evasion of benthic predators during swarming.^[39] The post-detachment stolon swims actively, spawning upon encountering opposite-sex individuals, after which it typically dies.^[38] Within the Syllidae, stolonization enables prolific swarming reproduction, as seen in genera like Myrianida, where chains of multiple stolons develop sequentially from the stock, facilitating mass mating aggregations in coastal waters. In Myrianida species, this chained formation amplifies reproductive output, with stolons detaching in succession to form dense swarms synchronized by lunar cues. Similarly, in Megasyllis nipponica, the Japanese green syllid, single stolons detach for epitokous swimming and spawning, contributing to the family's diverse strategies for ensuring gene flow across marine environments.^[39] This fission-based approach parallels colonial budding in certain invertebrates but yields free-swimming, ephemeral units rather than persistent attached colonies.^[37]

In Paleontology

Fossil Evidence

Fossil evidence for stolon-like structures primarily comes from Ediacaran rangeomorphs, enigmatic soft-bodied organisms that exhibited modular growth patterns suggestive of clonal propagation via horizontal runners or stolons. These structures are preserved as impressions of branching networks and filamentous connections between fronds, indicating asexual reproduction through extension from a parent organism. Key examples include Fractofusus andersoni from the Mistaken Point assemblage in Newfoundland, Canada, where distal extensions of the central axis are interpreted as stolons facilitating colony expansion, dated to approximately 567.63 ± 0.66 Ma. Similarly, horizontal branches and connecting filaments observed in clusters of Charnia and related fronds at the same site suggest stolon-mediated dispersal, with impressions showing parent-child groupings up to several meters across.^[40]^[41] Preservation of these stolon networks is rare due to the delicate, non-mineralized nature of rangeomorph bodies, relying on exceptional conditions in deep-marine ash beds that captured fine details as positive epirelief or hyporelief casts. Body fossils dominate, with trace-like impressions of stolons appearing as linear or branching traces on bedding planes, often in dense assemblages covering large surfaces. The Avalon biota, including Mistaken Point and Charnwood Forest sites, provides the richest record, spanning ~575–560 Ma, while the White Sea biota in Russia yields comparable examples in Dickinsonia and Charnia clusters, dated to ~558–550 Ma, showing similar horizontal extensions interpreted as stolons. No major discoveries specific to stolon structures have been reported between 2023 and 2025 beyond refinements in existing assemblages.^[40]^[41]^[42] Beyond the Ediacaran, potential stolon-like structures appear in early Cambrian colonial fossils, such as the interconnected tubular frameworks of archaeocyathid reefs (~530–520 Ma), though their interpretation as stolons remains debated due to the calcified nature differing from soft-bodied precursors. These records are primarily body fossils from reefal deposits in Siberia and Antarctica, with horizontal interconnections possibly representing modular growth akin to later colonial invertebrates. Such evidence underscores the transition from Ediacaran networks to more rigid Cambrian forms, with modern bryozoan stolons providing interpretive analogs for understanding attachment and propagation.^[43]^[44]

Evolutionary Significance

Stolons played a pivotal role in fostering modularity among early multicellular organisms, particularly during the Ediacaran period (approximately 635–541 million years ago), by enabling asexual reproduction through runner-like structures that facilitated rapid horizontal spread and colonial formation. In Precambrian oceans, this mechanism allowed organisms such as rangeomorph fronds to produce closely spaced clusters, often interpreted as "conga lines," enhancing survival by promoting efficient resource exploitation and evasion of localized threats in nutrient-scarce environments.^[45]^[46]^[47] These stolon networks marked a key evolutionary transition from solitary, simple forms to interconnected colonies, providing a modular architecture that could scale in complexity and adaptability. In Ediacaran biota, such as rangeomorphs, stolons supported propagule detachment and reattachment, laying groundwork for the diversification of colonial strategies that preceded the Cambrian explosion of bilaterian animals around 541 million years ago.^[48]^[49] The implications of stolon-mediated modularity extended to greater resilience against environmental perturbations, including potential extinction events, by allowing persistent vegetative propagation without reliance on sexual reproduction. This trait parallels developments in fungal and plant evolution, where stolons and analogous structures promoted clonal persistence and resource foraging, contributing to the colonization of terrestrial habitats.^[50]^[51] Research on stolon evolution remains limited in updates from 2023 to 2025, with ongoing studies highlighting potential indirect connections to early fungal terrestrialization, where fungi predated plants by up to 1.4 billion years and facilitated soil formation through hyphal networks akin to stolons.^[52]^[53]

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