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In
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
, stolons (from
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
'' stolō'', genitive ''stolōnis'' – "branch"), also known as runners, are horizontal connections between organisms. They may be part of the organism, or of its
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
; typically,
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
stolons are
external skeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the l ...
s.


In botany

In
botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
, stolons are
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
s which grow at the
soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former ...
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 embryonic shoot and normally occurs in the axil of a leaf or at the tip of a stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be spec ...
s. Stolons are often called runners.
Rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow h ...
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 a variety of sources: seeds, cuttings, and other plant parts. Plant propagation can also refer to the man-made or natural dispersal of seeds. Propagation typically occurs as a step i ...
strategy and the complex of individuals formed by a mother plant and all its
clones Clone or Clones or Cloning or Cloned or The Clone may refer to: Places * Clones, County Fermanagh * Clones, County Monaghan, a town in Ireland Biology * Clone (B-cell), a lymphocyte clone, the massive presence of which may indicate a patholog ...
produced from stolons form a single genetic individual, a
genet Genet or Genêt may refer to: Aircraft *Armstrong Siddeley Genet, aircraft engine * Armstrong Siddeley Genet Major, aircraft engine Animals and plants * Genet (biology), a colony of plants, fungi or bacteria that come from a single genetic sourc ...
.


Morphology

Stolons may or may not have long 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 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. 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 (mythology), a Greek goddess * ''Iris'' (plant), a genus of flowering plants * Iris (color), an ambiguous color term Iris or IRIS may also refer to: Arts and media Fictional ent ...
'' 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. Along with the Asteraceae, they are one of the two largest families of flowering ...
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 () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns ...
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 In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
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 ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomencla ...
''
Riccia ''Riccia'' is a genus of liverworts in the order Marchantiales. These plants are small and thalloid, that is not differentiated into root, stem and leaf. Depending on species, the thallus may be strap-shaped and about 0.5 to 4 mm wide wit ...
'', ''
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
'' (silverweed), ''
Cynodon ''Cynodon'' 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 being cultivated and naturalized in the New World and on many oceanic islands. The genus name comes from G ...
'', ''
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 hybrids and cultivars. The most common strawberries grown com ...
'', and ''
Pilosella ''Pilosella'' is a genus of flowering plants in the family Asteraceae. Some sources include it within the genus '' Hieracium''. Species , Plants of the World Online Plants of the World Online (POWO) is an online database published by the Ro ...
'' (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 i ...
''. Plants with long, slender stolons are referred to as ''sarmentose'' plants. Other plants with stolons below the soil surface include many grasses, '' Ajuga'', ''
Mentha ''Mentha'' (also known as mint, from Greek , Linear B ''mi-ta'') is a genus of plants in the family Lamiaceae (mint family). The exact distinction between species is unclear; it is estimated that 13 to 24 species exist. Hybridization occurs n ...
'', 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''. 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. 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 food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae. Wild potato species can be found from the southern United ...
, the stolons start to grow within 10 days of plants emerging above ground, with
tubers Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
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 ...
'' 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 lily family, most closely related to tulips. The name Erythronium derives from Ancient Greek () "red" in ...
'', 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
bulb In botany, a bulb is structurally a short 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''. Flowering plants often produce no stolons. ''
Convolvulus arvensis ''Convolvulus arvensis'', the field bindweed, is a species of bindweed that is rhizomatous and is in the morning glory family (Convolvulaceae), native to Europe and Asia. It is a climbing or creeping herbaceous perennial plant with stems growing ...
'' 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 The meristem is a type of tissue found in plants. It consists of undifferentiated cells (meristematic cells) capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells conti ...
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 fungi, including their genetic and biochemical properties, their taxonomy and their use to humans, including as a source for tinder, traditional medicine, food, and entheogens, as ...
, 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 or ...
, which connects sporangiophores 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 u ...
s 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

Some
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
ns form colonies through 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 animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
develop as stolons with interconnected medusoid structures that later separate. Some worm-like animals, such as certain
Polychaeta Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are m ...
in the genus ''Myrianida'', form stolons containing eggs or sperm which they trail behind the main body.


Palaeontology

Stolon based reproduction is thought to have been used by
Rangeomorphs The rangeomorphs are a form taxon of frondose Ediacaran fossils that are united by a similarity to ''Rangea''. Some researchers, such as Pflug and Narbonne, suggest that a natural taxon Rangeomorpha may include all similar-looking fossils. Rang ...
in the
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and t ...
age.Peterson et al. (2003), ''Integr Comp Biol'', 43:127-36


See also

* * * *


References

{{reflist Plant morphology Plant reproduction