Stolon

ⓘ
In biology, a stolon (/ˈstoʊlɒn/ from Latin stolō, 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. Typically, animal stolons are exoskeletons (external skeletons).

Ficinia spiralis (pīngao) spreads by forming stolons

in the sand.

Argentina anserina (common silverweed) showing

red stolons
In botany
In botany, stolons are plant stems which grow at the soil surface or just below ground that form
adventitious roots at the nodes, and new plants from the buds.[1][2] Stolons are often called runners.
Rhizomes, 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 strategy and the complex of individuals formed by a mother plant and
all its clones produced from stolons form a single genetic individual, a genet.

Morphology

These stolons from the corm of a Crocosmia are

stems that emerged from axillary buds at the
nodes of the tunic leaves.

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

Stolons arise from the base of the plant.[4] In strawberries the base is above the soil surface; in many
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 species and many 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 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 (Comarum palustre × Fragaria × ananassa) growing new plants using
vegetative propagation along stolons
 Iris pseudacorus

In some Cyperus species the stolons end with the growth of tubers; the tubers are swollen stolons
that form new plants.[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 Riccia, Argentina
(silverweed), Cynodon, Fragaria, and Pilosella (Hawkweeds), Zoysia japonica, 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, Mentha,[12] and
Stachys. Several species of Irises have stolons attached to their rhizomes,[13] including Iris
stolonifera.

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.[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.

In potatoes, the stolons[15] start to grow within 10 days of plants emerging above ground, with 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).

Hydrilla use stolons that produce tubers to spread themselves and to survive dry periods in aquatic
habitats [18]
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. A number of
bulbous species produce stolons, such as Erythronium propullans. Flowering plants often produce no
stolons.[19]

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 tissue to keep the parent plant alive from one year to the next.[21]

In mycology
In mycology, a stolon is defined as an occasionally septate hypha, which connects 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.

In zoology

The Cnidarian fish parasite Polypodium hydriforme

has a stolon stage of interconnected medusoids.
 Members of the genus Myrianida in
the Polychaeta, sometimes known as
"trainworms", form stolons containing
eggs or sperm.

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

Some colonial Cnidaria develop as stolons with interconnected medusoid structures that later
separate.

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

The worm Megasyllis nipponica 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 in the Ediacaran
period.[25][26]
See also

Offshoot (plant) – Lateral growth

branching from plant stem
Root – Basal organ of a vascular plant
Sprigging – Method for plant propagation
Vegetative reproduction – Asexual
method of reproduction in plants

References

1. Hickey, M.; King, C. (2001). The

Cambridge Illustrated Glossary of
Botanical Terms. Cambridge University
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3. Goebel, K. E. R. (1969) [1905].
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the Archegoniatae and Spermaphyta.
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4. Gleason, Henry A. (1963). The new
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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.
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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" ([Link]
[Link]/10.2307%2F2657090) . American
Journal of Botany. 88 (6): 1071–1079.
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1990). "Effects of light quality and
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Eichhornia crassipes". Oecologia. 84
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Carter, J. R. "Cyperus serotinus" (http://
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"Sarmentose" ([Link]
g/wiki/1911_Encyclop%C3%A6dia_Brita
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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
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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.
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0..285V) . doi:10.1111/j.1399-
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18. "Hydrilla in the Catawba River Basin" (ht
tp://[Link]/aquati
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19. weakley, Alan S. "Flora of the Carolinas,
Virginia, and Georgia, and Surrounding
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20. Harris, Peter. "Field and Hedge
bindweeds Convolvulus arvensis L. and
Calystegia sepium (L.) R. Br" ([Link]
[Link]/web/20070926235000/ht
tp://[Link]/lethbridge/weedbio/pla
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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" ([Link]
article/10.1007/BF00045274) . Plant
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22. Levinton, Jeffrey S. "Marine Biology."
3rd Edition. Oxford Press. 2008.
23. Pleijel, Fredrik (16 September 2016).
"Runner up: Evolutionary Biology.
"Polychaetous worm with engine and
wagons" " ([Link]
2016/09/royal-society-photo-contest-wi
nners-capture-breathtaking-details-of-o
ur-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."
24. Rayne, Elizabeth (8 December 2023).
"Worm's rear end develops its own
head, wanders off to mate" ([Link]
[Link]/science/2023/12/bizarre-
worm-can-detach-its-own-butt-and-mak
e-it-swim-away/) . 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?"
25. Mitchell et al. (2015),
doi:10.1038/nature14646
26. Peterson et al. (2003), Integr Comp Biol,
43:127-36
Retrieved from
"[Link]
title=Stolon&oldid=1333521170"

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