Sarmentose (botany)
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In
biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
, a stolon ( from
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
'' 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 A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
. Typically,
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
stolons are
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
s (external skeletons).


In botany

In
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
, stolons are
plant stem A stem is one of two main structural axes of a vascular plant, the other being the root. It supports leaf, leaves, flowers and fruits, transports water and dissolved substances between the roots and the shoots in the xylem and phloem, engages ...
s which grow at the
soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
surface or just below ground that form
adventitious roots Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant ...
at the
node In general, a node is a localized swelling (a "knot") or a point of intersection (a vertex). Node may refer to: In mathematics * Vertex (graph theory), a vertex in a mathematical graph *Vertex (geometry), a point where two or more curves, lines ...
s, and new plants from the
bud In botany, a bud is an undeveloped or Plant embryogenesis, embryonic Shoot (botany), shoot and normally occurs in the axil of a leaf or at the tip of a Plant stem, stem. Once formed, a bud may remain for some time in a dormancy, dormant conditi ...
s. Stolons are often called runners.
Rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s, in contrast, are root-like stems that may either grow horizontally at the soil surface or in other orientations underground. Thus, not all horizontal stems are called stolons. Plants with stolons are called stoloniferous. A stolon is a
plant propagation Plant propagation is the process by which new plants grow from various sources, including seeds, Cutting (plant), cuttings, and other plant parts. Plant propagation can refer to both man-made and natural processes. Propagation typically occurs as ...
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

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 ''Stachys sylvatica'', commonly known as hedge woundwort, whitespot, or sometimes as hedge nettle, is a perennial herbaceous plant growing to tall in woodland and unmanaged grassland. In temperate zones of the northern hemisphere it flowers in ...
'' are normal in size. Stolons arise from the base of the plant. In
strawberries The garden strawberry (or simply strawberry; ''Fragaria × ananassa'') is a widely grown hybrid plant cultivated worldwide for its fruit. The genus ''Fragaria'', the strawberries, is in the rose family, Rosaceae. The fruit is appreciated f ...
the base is above the soil surface; in many
bulb In botany, a bulb is a short underground stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs duri ...
-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. Typically after the formation of the new plant the stolon dies away 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. A number of plants have soil-level or above-ground rhizomes, including ''
Iris Iris most often refers to: *Iris (anatomy), part of the eye * Iris (color), an ambiguous color term *Iris (mythology), a Greek goddess * ''Iris'' (plant), a genus of flowering plants * Iris (given name), a feminine given name, and a list of peopl ...
'' species and many
orchid Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Orchids are cosmopolitan plants that are found in almost every habitat on Eart ...
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.
A stolon of
grasses Poaceae ( ), also called Gramineae ( ), is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos, the grasses of natural grassland and species cultivated in ...
is defined as a horizontal stem above or on the soil surface that often roots at the internodes.


Plants with stolons

In some ''
Cyperus ''Cyperus'' is a large genus of about 700 species of sedges, distributed throughout all continents in both tropical and temperate regions. Description They are annual or perennial plants, mostly aquatic and growing in still or slow-moving ...
'' species the stolons end with the growth of tubers; the tubers are swollen stolons that form new plants. Some
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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 Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
''
Riccia ''Riccia'' is a genus of liverworts in the order Marchantiales. Description These plants are small and thalloid, i.e. not differentiated into root, stem and leaf. Depending on species, the thallus may be strap-shaped and about 0.5 to 4 m ...
'', ''
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
'' (silverweed), ''
Cynodon ''Cynodon'', from Ancient Greek κύων (''kúōn''), meaning "dog", and ὀδούς (''odoús''), meaning "tooth", is a genus of plants in the grass family. It is native to warm temperate to tropical regions of the Old World, as well as bei ...
'', ''
Fragaria ''Fragaria'' () is a genus of flowering plants in the rose family, Rosaceae, commonly known as strawberries for their edible fruits. There are more than 20 described species and many Hybrid plant, hybrids and cultivars. The most common strawberr ...
'', and ''
Pilosella ''Pilosella'' is a genus of flowering plants in the family Asteraceae. It includes approximately 250 species native to temperate Eurasia and northwestern Africa. Some sources include it within the genus ''Hieracium''. Species , Plants of the Wor ...
'' (Hawkweeds), ''
Zoysia ''Zoysia'' (;"Zoysia."
entry at CollinsDictionary.com. Retrieved 5 September 2021.
, -, ...
japonica'', ''
Ranunculus repens ''Ranunculus repens'', the creeping buttercup, is a flowering plant in the buttercup family Ranunculaceae, native to Europe, Asia and northwestern Africa. Habitat It is a very common weed of agricultural land and gardens, spreading quickly by it ...
''. Plants with long, slender stolons are referred to as ''sarmentose'' plants. Other plants with stolons below the soil surface include many grasses, ''
Ajuga ''Ajuga'' , also known as bugleweed,Bailey, L.H.; Bailey, E.Z.; the staff of the Liberty Hyde Bailey Hortorium. 1976. ''Hortus third: A concise dictionary of plants cultivated in the United States and Canada''. Macmillan, New York. ground pine, c ...
'', ''
Mentha ''Mentha'', also known as mint (from Greek , Linear B ''mi-ta''), is a genus of flowering plants in the mint family, Lamiaceae. It is estimated that 13 to 24 species exist, but the exact distinction between species is unclear. Hybridization oc ...
'', and ''
Stachys ''Stachys'' is a genus of plants, one of the largest in the mint family Lamiaceae.Harley, R. M., et al. 2004. "Labiatae". pages 167–275. In: Kubitzki, K. (editor) and J. W. Kadereit (volume editor). ''The Families and Genera of Vascular Plants ...
''. Several species of Irises have stolons attached to their rhizomes, including ''
Iris stolonifera ''Iris stolonifera'' is a plant species in the genus '' Iris''; it is also in the subgenus '' Iris'', and in the section ''Regelia''. It is a rhizomatous perennial, from the mountains of Turkestan, between Tajikistan, Uzbekistan and Afghanistan. ...
''.
Lily-of-the-valley Lily of the valley (''Convallaria majalis'' ), sometimes written lily-of-the-valley, is a woodland flowering plant with sweetly scented, pendent, bell-shaped white flowers borne in sprays in spring. It is native throughout the cool temperate No ...
(''Convallaria majalis'') has rhizomes that grow stolon-like stems called stoloniferous rhizomes or leptomorph rhizomes. A number of plants have stoloniferous rhizomes including
Asters ''Aster'' is a genus of perennial flowering plants in the family Asteraceae. Its circumscription has been narrowed, and it now encompasses around 170 species, all but one of which are restricted to Eurasia; many species formerly in ''Aster'' are ...
. 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 potato () is a starchy tuberous vegetable native to the Americas that is consumed as a staple food in many parts of the world. Potatoes are underground stem tubers of the plant ''Solanum tuberosum'', a perennial in the nightshade famil ...
, the stolons start to grow within 10 days of plants emerging above ground, with
tubers Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reprod ...
usually beginning to form on the end of the stolons. The tubers are modified stolons 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 ''Hydrilla'' (waterthyme) is a genus of aquatic plant, usually treated as containing just one species, ''Hydrilla verticillata'', though some botanists divide it into several species. It is native to the cool and warm waters of the Old World in A ...
'' use stolons that produce tubers to spread themselves and to survive dry periods in aquatic habitats. ''
Erythronium ''Erythronium'', the fawn lily, trout lily, dog's-tooth violet or adder's tongue, is a genus of Eurasian and North American plants in the Liliaceae, lily family, most closely related to tulips. The name Erythronium derives from Ancient Greek () ...
'', 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 Plant litter (also leaf litter, tree litter, soil litter, litterfall, or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that has fallen to the ground. This detritus or dead organic material and its constituen ...
. A number of
bulb In botany, a bulb is a short underground stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs duri ...
ous species produce stolons, such as ''
Erythronium propullans ''Erythronium propullans'', the Minnesota dwarf trout lily, Minnesota adder's tongue or Minnesota fawnlily, is a rare plant endemic to the Cannon River and North Fork Zumbro River watersheds in Rice County, Goodhue County and the extreme nort ...
''. Flowering plants often produce no stolons. ''
Convolvulus arvensis ''Convolvulus arvensis'', or field bindweed, is a species of bindweed in the Convolvulaceae native to Europe and Asia. It is a rhizomatous and climbing or creeping herbaceous perennial plant with stems growing to in length. It is usually found a ...
'' is a weed species in agriculture that spreads by under ground stolons that produce rhizomes. 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 In cell biology, the meristem is a structure composed of specialized tissue found in plants, consisting of stem cells, known as meristematic cells, which are undifferentiated cells capable of continuous cellular division. These meristematic c ...
tissue to keep the parent plant alive from one year to the next.


In mycology

In
mycology Mycology is the branch of biology concerned with the study of fungus, fungi, including their Taxonomy (biology), taxonomy, genetics, biochemistry, biochemical properties, and ethnomycology, use by humans. Fungi can be a source of tinder, Edible ...
, a stolon is defined as an occasionally septate
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
, which connects
sporangiophore {{Short pages monitor thumb , right , A photomicrograph of a sporangiophore together. Root-like structures called
rhizoid Rhizoids are protuberances that extend from the lower epidermal cells of bryophytes and algae. They are similar in structure and function to the root hairs of vascular land plants. Similar structures are formed by some fungi. Rhizoids may be un ...
s may appear on the stolon as well, anchoring the hyphae to the
substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...
. The stolon is commonly found in
bread mold A mold () or mould () is one of the structures that certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi ...
s, and are seen as horizontally expanding across the mold.


In zoology

Some
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
ns form colonies through the connection of individual units by stolons. Other colonies include sheets and erect colonies. Some colonial
Cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
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
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
s or
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
which trail behind the main body before detaching to mate with other stolons. 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.


In palaeontology

Stolon-based reproduction is thought to have been used by
Rangeomorphs The rangeomorphs are a group of Ediacaran fossils. Ediacarans are the oldest large fossil organisms on earth, and many are not self-evidently related to anything else that has ever lived. However, some Ediacarans clearly resemble each other. Pale ...
in the
Ediacaran The Ediacaran ( ) is a geological period of the Neoproterozoic geologic era, Era that spans 96 million years from the end of the Cryogenian Period at 635 Million years ago, Mya to the beginning of the Cambrian Period at 538.8 Mya. It is the last ...
age.Peterson et al. (2003), ''Integr Comp Biol'', 43:127-36


See also

* * * *


References

{{reflist Plant reproduction Plant stem morphology