Stolon

Stolons are horizontal stems which grow at the soil surface or below ground. They form new plants at the ends or at the nodes. Stolons are often called runners. Imprecisely they are stems that run atop or just under the ground; more specifically, a stolon is a horizontal shoot from a plant that grows on top of or below the soil surface with the ability to produce new clones of the same plant from buds at the tip. [cite web
title=Stolon
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Stolons are stems

Stolons are similar to normal stems except they produce adventitious roots at the nodes and run horizontally rather than vertically. They also have long internodes with reduced leaves, with the exception of stolons in aqueous plants. Plants with stolons or stolon-like rhizomes are called stoloniferous. A stolon is a plant propagation strategy and the complex of individuals formed by a mother plant and all its clones produced from stolons form a single genetic individual. Runners are a type of stolon that exist above ground and are produced by many plants, such as strawberries. Stolons lack the same type of reduced leaves that rhizomes have at the nodes; stolons have scale-like leaves and new roots are formed at the nodes only while rhizomes typicaly have paper like leaves at the nodes. Typically, stolons have very long internodes that form new plants at the ends. These rise to the soil surface and can produce foliage and flowers. In contrast, rhizomes most often have short internodes with leaf-scars and thin paper-like leaves and root along the under side of the stem. Root formation does not correspond strictly to the nodes but roots can generate from areas around the scar-like nodes as well.

Morphology

Stolons are like long branches of a tree that grow horizontal to the soil surface and produce new plants at the ends, they have nodes and internodes, leaves that are reduced to scales and buds that grow into roots and shoots. Stolons arise from the base of the plant, [cite book|last=Gleason| first=Henry A.| authorlink=Henry Gleason|title=The new Britton and Brown illustrated flora of the Northeastern United States and adjacent Canada, Volume 1| pages=p.ixxiv| location=New York| publisher=Hafner Press| isbn=0-02-845240-2|year=1963] in strawberries the base is above the soil surface, in many bulb-forming species and plants with rhizomes the stolons arise under ground and 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. [cite journal| last=Woolley| first=D. J.| coauthors=P. F. Wareing| month=March| year=1972| title=The role of roots, cytokinins and apical dominance in the control of lateral shoot form in "Solanum andigena"|journa=Planta| volume=105| issue=1| pages=33–42 |doi=10.1007/BF00385161| journal=Planta] Typically after the formation of the new plant the stolon dies away [cite journal| last=Wijesinghe | first=Dushyantha K. | coauthor=Dennis F. Whigham| title=Nutrient foraging in woodland herbs: a comparison of three species of "Uvularia" (Liliaceae) with contrasting belowground morphologies|journal=American Journal of Botany. | year=2001|month=June| volume=88| issue=6| pages=1071–1079.|pmid=11410472 url=http://www.amjbot.org/cgi/content/full/88/6/1071| accessdate=| doi=10.2307/2657090 ] 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. Rhizomes are used as storage structures for nutrients and most often are covered with thin papery leaves (normally a translucent brownish color), while stolons are not covered with reduced leaves but have scales like leaves at the nodes that are most often white in color. Stolons look and act like etiolated stems except for their orientation of growth, instead of stretching upward for light they grow laterally or downward. 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 effected 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 producing fewer stolons. [Cite journal| first= M.| last = Méthy| coauthors = P. Alpert, J. Roy| title = Effects of light quality and quantity on growth of the clonal plant Eichhornia crassipes| journal = Oecologia| url = http://www.springerlink.com/content/r19x274502337562/| accessdate = 2007-05-10| year = 1990| month = September| volume = 84| issue = 2| pages = 265–271 | doi = 10.1007/BF00318283| doi_brokendate = 2008-06-22]

Often the general meaning of stolon (horizontal stems above ground) does not distinguish morphologically the differences between stolons and rhizomes but just differentiates rhizome and stolon from each other strictly by where they are found, with rhizomes being below ground. Though this generalization is imprecise with a number of plants having soil level or above ground rhizomes including "Iris" species and many orchid species. Because of this some botany texts now use the term underground stolon to refer to stolons that move under ground. [Cite web| last=Mosyakin| first=Sergei L.| title="Rumex beringensis"|url= http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250060770| work=Flora of North America| publisher=eFloras.org| accessdate=2005-05-10]

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. [Cite web| last=Henderson| first=Norton C.| title="Iris"|url= http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=116503| work=Flora of North America| publisher=eFloras.org| accessdate=2005-05-10]

Plants with stolons

In some "Cyperus" species the stolons end with the growth of tubers; the tubers are swollen stolons that form new plants [Cite web| last=Tucker| first=Gordon C.|coauthors=Brian G. Marcks, J. Richard Carter| title="Cyperus serotinus"|url= http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=242101127| work=Flora of North America| publisher=eFloras.org| accessdate=2005-05-10] .

Some 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 "Argentina" (silverweed), "Cynodon", "Fragaria", and "Pilosella" (Hawkweeds), "Zoysia japonica", "Ranunculus repens".

Other plants with stolons below the soil surface include many grasses, "Ajuga", "Mentha", [cite journal| last=Aflatuni| first=Abbas| coauthors=J. Uusitalo, S. Ek, A. Hohtola| month=January/February| year=2005| title=Variation in the Amount of Yield and in the Extract Composition Between Conventionally Produced and Micropropagated Peppermint and Spearmint| journal=Journal of Essential Oil Research| volume=17 |issue=1| pages=66–70|issn=1041-2905|url=http://findarticles.com/p/articles/mi_qa4091/is_200501/ai_n9474312| accessdate=2005-05-10] and "Stachys".

Lily-of-the-valley ("Convallaria majalis") which has rhizomes that grow stolon-like stems called stoloniferous rhizomes or leptomorph rhizomes. A number of plants have stoloniferous rhizomes including Asters [cite journal |last=Jones |first=Almut G. |year=1978 |month=January |title=Observations on Reproduction and Phenology in Some Perennial Asters |journal=American Midland Naturalist |volume=99 |issue=1 |pages=184–97 |doi=10.2307/2424942] These stolon-like rhizomes are long and thin, with long internodes and indeterminate growth with lateral buds at the node that mostly remain dormant.

In potatoes, the stolons [cite journal |last=Visser |first=Richard G. F. |coauthors=Dick Vreugdenhil, Theo Hendriks, Evert Jacobsen |year=1994 |month=February |title=Gene expression and carbohydrate content during stolon to tuber transition in potatoes ("Solanum tuberosum") |journal=Physiologia Plantarum |volume=90 |issue=2 |pages=285–92 |doi=10.1111/j.1399-3054.1994.tb00389.x] start to grow within 10 days of plants emerging above ground, with tubers usually beginning to form on the end of the stolons. [cite web| authos=Monaco Educational Service| title=Introduction to stems| url=http://www.personal.psu.edu/faculty/w/x/wxm15/Online/Botany/Stems/stem_lecture_01.htm| work=Botany| accessdate=2005-05-10] The tubers are modified stolons [cite book |last=Hartmann |first=Hudson Thomas |coauthor=Dale E Kester |title=Plant propagation : principles and practices |year=1983 |publisher=Prentice-Hall |location=Englewood Cliffs|isbn=0136810071 |pages=p.508] 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)

"Hydrilla" use stolons that produce tubers to spread themselves and survive dry periods in aquatic habitats. [cite web| title=Hydrilla in the Catawba River Basin|url=http://www.weedscience.ncsu.edu/aquaticweeds/catawba.pdf |publisher= NCSU Aquatic Weed Management Program| accessdate=2005-05-10]

"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. Flowering plants often produce no stolons. [Cite web|last=weakley| first=Alan S.| title=Flora of the Carolinas, Virginia, and Georgia, and Surrounding Areas, Part 6| url=http://www.herbarium.unc.edu/WeakleysFloraPart6.pdf| format=PDF| pages=p.808| accessdate=2005-05-10] A number of bulbous forming species produce stolons with "Erythronium propullans" producing one stolon below the soil surface on the midway point of the stem on blooming plants that produces a new bulb. On non blooming plants 1 to 3 stolons are produce directly from the bulbs, each ending in a new clone. [Cite web| last=Allen| first=Geraldine A.| coauthor=Kenneth R. Robertson| title="Erythronium propullans"|url= http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=242101601| work=Flora of North America| publisher=eFloras.org| accessdate=2005-05-10]

"Convolvulus arvensis" is a weed species in agriculture that spreads by under ground stolons that produce rhizomes. [cite web| last=Harris| first=Peter| title=Field and Hedge bindweeds "Convolvulus arvensis" L. and "Calystegia sepium" (L.) R. Br.| url=http://res2.agr.ca/lethbridge/weedbio/plant/convolvulus_e.htm| work= Classical Biological Control of Weeds |publisher=Agriculture and Agri-Food Canada, Lethbridge Research Centre| accessdate=2005-05-10]

In studies on grass species, with plants that produce stolons or rhizomes and plants that produce both stolons and rhizomes; morphological and physiological differences where noticed. Stolons have longer internodes and function as means of seeking out light and were used for propagation of the plant, while rhizomes are used as storage organs for carbohydrates and the maintenance of meristem tissue to keep the parent plant alive from one year to the next. [cite journal |last=Pierdominici |first=Maria Grazia |coauthors=Ming Dong |year=1995 |month=January |title= Morphology and growth of stolons and rhizomes in three clonal grasses, as affected by different light supply |journal=Plant Ecology |volume=116 |issue=1 |pages=25–32 |doi=10.1007/BF00045274|url=http://www.springerlink.com/content/p2672v2617k61473/| accessdate=2005-05-10 |doi_brokendate= 2008-06-22]

Mycology

In mycology, a stolon is defined as an occasionally septate hyphae, which connect sporangiophores together. Root-like structures called rhizoids may appear on the stolon as well, anchoring the hyphae to the substrate. The stolon is commonly found in bread molds, and are seen as horizontally expanding across the mold.

ee also

*Root
*Vegetative reproduction

References