# Stolon

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Horizontal connections between organisms

*[Ficinia spiralis](/source/Ficinia_spiralis)* (pīngao) spreads by forming stolons in the sand.

*[Argentina anserina](/source/Argentina_anserina)* (common silverweed) showing red stolons

In [biology](/source/Biology), **a stolon** ([/ˈstoʊlɒn/](https://en.wikipedia.org/wiki/Help:IPA/English) [ⓘ](https://en.wikipedia.org/wiki/File:LL-Q1860_(eng)-Naomi_Persephone_Amethyst_(NaomiAmethyst)-stolon.wav) from [Latin](/source/Latin) *[stolō](https://en.wiktionary.org/wiki/stolo)*, genitive *stolōnis* – "branch"), also known as a **runner**, is a horizontal connection between parts of an organism. It may be part of the organism, or of its [skeleton](/source/Skeleton). Typically, [animal](/source/Animal) stolons are [exoskeletons](/source/Exoskeleton) (external skeletons).

## In botany

In [botany](/source/Botany), stolons are [plant stems](/source/Plant_stem) which grow at the [soil](/source/Soil) surface or just below ground that form [adventitious roots](/source/Adventitious_roots) at the [nodes](/source/Node_(botany)), and new plants from the [buds](/source/Bud).[1][2] Stolons are often called **runners**. [Rhizomes](/source/Rhizome), in contrast, are root-like stems that may either grow horizontally at the soil surface or in other orientations underground.[1] Thus, not all horizontal stems are called stolons. Plants with stolons are called **stoloniferous**.

A stolon is a [plant propagation](/source/Plant_propagation) strategy and the complex of individuals formed by a mother plant and all its [clones](/source/Cloning) produced from stolons form a single genetic individual, a [genet](/source/Genet_(biology)).[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*]

### Morphology

These stolons from the [corm](/source/Corm) of a *[Crocosmia](/source/Crocosmia)* are stems that emerged from axillary buds at the nodes of the [tunic leaves](/source/Bulb).

Stolons may have long or short [internodes](/source/Internode_(botany)). The leaves along the stolon are usually very small, but in a few cases such as *[Stachys sylvatica](/source/Stachys_sylvatica)* are normal in size.[3]

Stolons arise from the base of the plant.[4] In [strawberries](/source/Strawberry) the base is above the soil surface; in many [bulb](/source/Bulb)-forming species and plants with rhizomes, the stolons remain underground and form shoots that rise to the surface at the ends or from the nodes. The nodes of the stolons produce roots, often all around the node and hormones produced by the roots cause the stolon to initiate shoots with normal leaves.[5] Typically after the formation of the new plant the stolon dies away[6] in a year or two, while rhizomes persist normally for many years or for the life of the plant, adding more length each year to the ends with active growth. The horizontal growth of stolons results from the interplay of different hormones produced at the growing point and hormones from the main plant, with some studies showing that stolon and rhizome growth are affected by the amount of shady light the plant receives with increased production and branching from plants exposed to mixed shade and sun, while plants in all day sun or all shade produce fewer stolons.[7]

A number of plants have soil-level or above-ground rhizomes, including *[Iris](/source/Iris_(plant))* species and many [orchid](/source/Orchid) species.

T. Holm (1929) restricted the term rhizome to a horizontal, usually subterranean, stem that produces roots from its lower surface and green leaves from its apex, developed directly from the plumule of the embryo. He recognized stolons as axillary, subterranean branches that do not bear green leaves but only membranaceous, scale-like ones.[8]

A stolon of [grasses](/source/Poaceae) is defined as a horizontal stem above or on the soil surface that often roots at the internodes.[9]

### Plants with stolons

['Lipstick' hybrid strawberry](/source/Fragaria_%C3%97_Comarum_hybrids) (*Comarum palustre* × *Fragaria × ananassa*) growing new plants using [vegetative propagation](/source/Vegetative_propagation) along stolons

*[Iris pseudacorus](/source/Iris_pseudacorus)*

In some *[Cyperus](/source/Cyperus)* species the stolons end with the growth of tubers; the tubers are swollen stolons that form new plants.[10]

Some [species](/source/Species) of crawling plants can also sprout adventitious roots, but are not considered stoloniferous: a stolon is sprouted from an existing stem and can produce a full individual. Examples of plants that extend through stolons include some species from the [genera](/source/Genus) *[Riccia](/source/Riccia)*, *[Argentina](/source/Argentina_(plant))* (silverweed), *[Cynodon](/source/Cynodon)*, *[Fragaria](/source/Strawberry)*, and *[Pilosella](/source/Pilosella)* (Hawkweeds), *[Zoysia](/source/Zoysia) japonica*, *[Ranunculus repens](/source/Ranunculus_repens)*. Plants with long, slender stolons are referred to as *sarmentose* plants.[11]

Other plants with stolons below the soil surface include many grasses, *[Ajuga](/source/Ajuga)*, *[Mentha](/source/Mentha)*,[12] and *[Stachys](/source/Stachys)*. Several species of [Irises](/source/Iris_(plant)) have stolons attached to their rhizomes,[13] including *[Iris stolonifera](/source/Iris_stolonifera)*.

[Lily-of-the-valley](/source/Lily-of-the-valley) (*Convallaria majalis*) has rhizomes that grow stolon-like stems called stoloniferous rhizomes or leptomorph rhizomes. A number of plants have stoloniferous rhizomes including [Asters](/source/Aster_(genus)).[14] These stolon-like rhizomes are long and thin, with long internodes and indeterminate growth with lateral buds at the node, which mostly remain dormant.[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*]

In [potatoes](/source/Potatoes), the stolons[15] start to grow within 10 days of plants emerging above ground, with [tubers](/source/Tubers) usually beginning to form on the end of the stolons.[16] The tubers are modified stolons[17] that hold food reserves, with a few buds that grow into stems. Since it is *not* a rhizome it does not generate roots, but the new stem growth that grows to the surface produces roots. See also [BBCH-scale (potato)](/source/BBCH-scale_(potato)).

*[Hydrilla](/source/Hydrilla)* use stolons that produce tubers to spread themselves and to survive dry periods in aquatic habitats.[18]

*[Erythronium](/source/Erythronium)*, commonly called Trout Lily, have white stolons growing from the bulb. Most run horizontally, either underground or along the surface of the ground under [leaf litter](/source/Leaf_litter). A number of [bulbous](/source/Bulb) species produce stolons, such as *[Erythronium propullans](/source/Erythronium_propullans)*. Flowering plants often produce no stolons.[19]

*[Convolvulus arvensis](/source/Convolvulus_arvensis)* is a weed species in agriculture that spreads by under ground stolons that produce rhizomes.[20]

In studies on grass species, with plants that produce stolons or rhizomes and plants that produce both stolons and rhizomes, morphological and physiological differences were noticed. Stolons have longer internodes and function as means of seeking out light and are used for propagation of the plant, while rhizomes are used as storage organs for carbohydrates and the maintenance of [meristem](/source/Meristem) tissue to keep the parent plant alive from one year to the next.[21]

## In mycology

In [mycology](/source/Mycology), a stolon is defined as an occasionally septate [hypha](/source/Hypha), which connects [sporangiophores](/source/Sporangiophore) together. Root-like structures called [rhizoids](/source/Rhizoid) may appear on the stolon as well, anchoring the hyphae to the [substrate](/source/Substrate_(biology)). The stolon is commonly found in [bread molds](/source/Bread_mold), and are seen as horizontally expanding across the mold.

## In zoology

The Cnidarian fish parasite *[Polypodium hydriforme](/source/Polypodium_hydriforme)* has a stolon stage of interconnected medusoids.

Members of the genus *Myrianida* in the [Polychaeta](/source/Polychaete), sometimes known as "trainworms", form stolons containing eggs or sperm.

Some [bryozoans](/source/Bryozoa) form colonies through the connection of individual units by stolons. Other colonies include sheets and erect colonies.[22]

Some colonial [Cnidaria](/source/Cnidaria) develop as stolons with interconnected medusoid structures that later separate.[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*]

Some worm-like animals, such as certain [Polychaeta](/source/Polychaete) in the genus *Myrianida*, form stolons containing [eggs](/source/Egg) or [sperm](/source/Sperm) which trail behind the main body before detaching to mate with other stolons.[23]

The worm *[Megasyllis nipponica](https://en.wikipedia.org/w/index.php?title=Megasyllis_nipponica&action=edit&redlink=1)* takes this to an extreme, developing stolons with their own eyes, antennae, gut, and brain, which detach, seek out, and mate with an opposite-sex stolon to produce fertilized eggs.[24]

## In palaeontology

Stolon-based reproduction is thought to have been used by [Rangeomorphs](/source/Rangeomorpha) in the [Ediacaran](/source/Ediacaran) period.[25][26]

## See also

Wikimedia Commons has media related to [Plant stolons](https://commons.wikimedia.org/wiki/Plant_stems#Stolons).

- [Offshoot (plant)](/source/Offshoot_(plant)) – Lateral growth branching from plant stem

- [Root](/source/Root) – Basal organ of a vascular plant

- [Sprigging](/source/Sprigging) – Method for plant propagation

- [Vegetative reproduction](/source/Vegetative_reproduction) – Asexual method of reproduction in plants

## References

1. ^ [***a***](#cite_ref-Hickey&King_1-0) [***b***](#cite_ref-Hickey&King_1-1) Hickey, M.; King, C. (2001). *The Cambridge Illustrated Glossary of Botanical Terms*. Cambridge University Press.

1. **[^](#cite_ref-2)** ["Stolon"](http://dictionary.reference.com/browse/stolon). Dictionary.com. Retrieved 2007-05-07.

1. **[^](#cite_ref-3)** Goebel, K. E. R. (1969) [1905]. *Organography of plants, especially of the Archegoniatae and Spermaphyta*. New York: Hofner publishing company.

1. **[^](#cite_ref-4)** [Gleason, Henry A.](/source/Henry_Gleason) (1963). *The new Britton and Brown illustrated flora of the Northeastern United States and adjacent Canada, Volume 1*. New York: Hafner Press. p. ixxiv. [ISBN](/source/ISBN_(identifier)) [0-02-845240-2](https://en.wikipedia.org/wiki/Special:BookSources/0-02-845240-2). {{[cite book](https://en.wikipedia.org/wiki/Template:Cite_book)}}: ISBN / Date incompatibility ([help](https://en.wikipedia.org/wiki/Help:CS1_errors#invalid_isbn_date))

1. **[^](#cite_ref-5)** Woolley, D. J.; P. F. Wareing (March 1972). "The role of roots, cytokinins and apical dominance in the control of lateral shoot form in *Solanum andigena*". *Planta*. **105** (1): 33–42. [Bibcode](/source/Bibcode_(identifier)):[1972Plant.105...33W](https://ui.adsabs.harvard.edu/abs/1972Plant.105...33W). [doi](/source/Doi_(identifier)):[10.1007/BF00385161](https://doi.org/10.1007%2FBF00385161). [PMID](/source/PMID_(identifier)) [24477700](https://pubmed.ncbi.nlm.nih.gov/24477700). [S2CID](/source/S2CID_(identifier)) [6832024](https://api.semanticscholar.org/CorpusID:6832024).

1. **[^](#cite_ref-6)** Wijesinghe, Dushyantha K.; Dennis F. Whigham (June 2001). ["Nutrient foraging in woodland herbs: a comparison of three species of *Uvularia* (Liliaceae) with contrasting belowground morphologies"](https://doi.org/10.2307%2F2657090). *American Journal of Botany*. **88** (6): 1071–1079. [doi](/source/Doi_(identifier)):[10.2307/2657090](https://doi.org/10.2307%2F2657090). [JSTOR](/source/JSTOR_(identifier)) [2657090](https://www.jstor.org/stable/2657090). [PMID](/source/PMID_(identifier)) [11410472](https://pubmed.ncbi.nlm.nih.gov/11410472).

1. **[^](#cite_ref-7)** Méthy, M.; P. Alpert; J. Roy (September 1990). "Effects of light quality and quantity on growth of the clonal plant Eichhornia crassipes". *Oecologia*. **84** (2): 265–271. [Bibcode](/source/Bibcode_(identifier)):[1990Oecol..84..265M](https://ui.adsabs.harvard.edu/abs/1990Oecol..84..265M). [doi](/source/Doi_(identifier)):[10.1007/BF00318283](https://doi.org/10.1007%2FBF00318283). [PMID](/source/PMID_(identifier)) [28312764](https://pubmed.ncbi.nlm.nih.gov/28312764). [S2CID](/source/S2CID_(identifier)) [1279519](https://api.semanticscholar.org/CorpusID:1279519).

1. **[^](#cite_ref-8)** Henderson, Norton C. ["Iris"](http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=116503). *Flora of North America*. eFloras.org. Retrieved 2005-05-10.

1. **[^](#cite_ref-9)** Aldous, David E.; Chivers, Ian H. (2002-06-13). [*Sports Turf and Amenity Grasses: A Manual for Use and Identification*](https://books.google.com/books?id=bfFuyRs_aMQC&pg=PP13). Landlinks Press. [ISBN](/source/ISBN_(identifier)) [978-0-643-09960-9](https://en.wikipedia.org/wiki/Special:BookSources/978-0-643-09960-9).

1. **[^](#cite_ref-10)** Tucker, Gordon C.; Marcks, Brian G.; Carter, J. R. ["Cyperus serotinus"](http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=242101127). *Flora of North America*. eFloras.org. Retrieved 2005-05-10.

1. **[^](#cite_ref-11)** [Chisholm, Hugh](/source/Hugh_Chisholm), ed. (1911). ["Sarmentose"](https://en.wikisource.org/wiki/1911_Encyclop%C3%A6dia_Britannica/Sarmentose). *[Encyclopædia Britannica](/source/Encyclop%C3%A6dia_Britannica_Eleventh_Edition)*. Vol. 24 (11th ed.). Cambridge University Press. p. 220.

1. **[^](#cite_ref-12)** Aflatuni, Abbas; Uusitalo, J.; Ek, S.; Hohtola, A. (February 2005). "Variation in the Amount of Yield and in the Extract Composition Between Conventionally Produced and Micropropagated Peppermint and Spearmint". *Journal of Essential Oil Research*. **17** (1): 66–70. [doi](/source/Doi_(identifier)):[10.1080/10412905.2005.9698833](https://doi.org/10.1080%2F10412905.2005.9698833). [ISSN](/source/ISSN_(identifier)) [1041-2905](https://search.worldcat.org/issn/1041-2905). [S2CID](/source/S2CID_(identifier)) [97042181](https://api.semanticscholar.org/CorpusID:97042181).

1. **[^](#cite_ref-13)** ["Home"](http://encyclopaedia.alpinegardensociety.net/plants/Iris). *Alpine Garden Society*. Retrieved 8 November 2020.

1. **[^](#cite_ref-14)** Jones, Almut G. (January 1978). "Observations on Reproduction and Phenology in Some Perennial Asters". *American Midland Naturalist*. **99** (1): 184–97. [doi](/source/Doi_(identifier)):[10.2307/2424942](https://doi.org/10.2307%2F2424942). [JSTOR](/source/JSTOR_(identifier)) [2424942](https://www.jstor.org/stable/2424942).

1. **[^](#cite_ref-15)** Visser, Richard G. F.; Vreugdenhil, Dick; Hendriks, Theo; Evert Jacobsen (February 1994). "Gene expression and carbohydrate content during stolon to tuber transition in potatoes (*Solanum tuberosum*)". *Physiologia Plantarum*. **90** (2): 285–92. [Bibcode](/source/Bibcode_(identifier)):[1994PPlan..90..285V](https://ui.adsabs.harvard.edu/abs/1994PPlan..90..285V). [doi](/source/Doi_(identifier)):[10.1111/j.1399-3054.1994.tb00389.x](https://doi.org/10.1111%2Fj.1399-3054.1994.tb00389.x). [S2CID](/source/S2CID_(identifier)) [86752575](https://api.semanticscholar.org/CorpusID:86752575).

1. **[^](#cite_ref-16)** Monaco Educational Service. ["Introduction to stems"](https://web.archive.org/web/20050413012738/http://www.personal.psu.edu/faculty/w/x/wxm15/Online/Botany/Stems/stem_lecture_01.htm). *Botany*. Archived from [the original](http://www.personal.psu.edu/faculty/w/x/wxm15/Online/Botany/Stems/stem_lecture_01.htm) on 2005-04-13. Retrieved 2005-05-10.

1. **[^](#cite_ref-17)** Hartmann, Hudson Thomas; Dale E Kester (1983). *Plant propagation : principles and practices*. Englewood Cliffs: Prentice-Hall. p. 508. [ISBN](/source/ISBN_(identifier)) [0-13-681007-1](https://en.wikipedia.org/wiki/Special:BookSources/0-13-681007-1).

1. **[^](#cite_ref-18)** ["Hydrilla in the Catawba River Basin"](http://www.weedscience.ncsu.edu/aquaticweeds/catawba.pdf) (PDF). NCSU Aquatic Weed Management Program. Retrieved 2005-05-10.

1. **[^](#cite_ref-19)** weakley, Alan S. ["Flora of the Carolinas, Virginia, and Georgia, and Surrounding Areas, Part 6"](https://web.archive.org/web/20060304021820/http://www.herbarium.unc.edu/WeakleysFloraPart6.pdf) (PDF). p. 808. Archived from [the original](http://www.herbarium.unc.edu/WeakleysFloraPart6.pdf) (PDF) on 2006-03-04. Retrieved 2005-05-10.

1. **[^](#cite_ref-20)** Harris, Peter. ["Field and Hedge bindweeds *Convolvulus arvensis* L. and *Calystegia sepium* (L.) R. Br"](https://web.archive.org/web/20070926235000/http://res2.agr.ca/lethbridge/weedbio/plant/convolvulus_e.htm). *Classical Biological Control of Weeds*. Agriculture and Agri-Food Canada, Lethbridge Research Centre. Archived from [the original](http://res2.agr.ca/lethbridge/weedbio/plant/convolvulus_e.htm) on 2007-09-26. Retrieved 2005-05-10.

1. **[^](#cite_ref-21)** Pierdominici, Maria Grazia; Ming Dong (January 1995). ["Morphology and growth of stolons and rhizomes in three clonal grasses, as affected by different light supply"](https://link.springer.com/article/10.1007/BF00045274). *Plant Ecology*. **116** (1): 25–32. [doi](/source/Doi_(identifier)):[10.1007/BF00045274](https://doi.org/10.1007%2FBF00045274). [S2CID](/source/S2CID_(identifier)) [6281632](https://api.semanticscholar.org/CorpusID:6281632).

1. **[^](#cite_ref-22)** Levinton, Jeffrey S. "Marine Biology." 3rd Edition. Oxford Press. 2008.

1. **[^](#cite_ref-23)** Pleijel, Fredrik (16 September 2016). ["Runner up: Evolutionary Biology. "Polychaetous worm with engine and wagons""](https://designyoutrust.com/2016/09/royal-society-photo-contest-winners-capture-breathtaking-details-of-our-rapidly-changing-world/). *Royal Society Photo Contest Winners Capture Breathtaking Details of our Rapidly Changing World*. Retrieved 2023-12-08. This trainworm (*Myrianida pinnigera*), which is 35 mm from head to tail, lives on the sea floor. Its front end, the trainworm's engine, is followed by a row of carriages called 'stolons' that increase in size towards the worm's tail end. The carriages are the worm's swimming sexual organs. When the trainworm is mature, the last carriage in the train lets go and detaches. It swims up the water column to reproduce. The carriages, unlike the engine, all lack a gut and are full of either sperm or eggs. In the water column the well-developed sensory organs seek out another stolon to mate with. After mating the male stolons die. The females survive a short time to shelter the embryos, which are carried in their bellies. Meanwhile, the trainworm on the sea floor continues to produce stolon carriages.

1. **[^](#cite_ref-24)** Rayne, Elizabeth (8 December 2023). ["Worm's rear end develops its own head, wanders off to mate"](https://arstechnica.com/science/2023/12/bizarre-worm-can-detach-its-own-butt-and-make-it-swim-away/). *[Ars Technica](/source/Ars_Technica)*. Retrieved 2023-12-08. This appendage will detach from the main body and swim away, carrying gonads that will merge with those from other disembodied rear ends and give rise to a new generation. Wait, *what in the science fiction B-movie alien star system is this thing?*

1. **[^](#cite_ref-25)** Mitchell et al. (2015), doi:10.1038/nature14646

1. **[^](#cite_ref-26)** Peterson et al. (2003), *Integr Comp Biol*, **43**:127-36

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Adapted from the Wikipedia article [Stolon](https://en.wikipedia.org/wiki/Stolon) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Stolon?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.