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 hereditar ...
, stolons (from
Latin
Latin (, or , ) is a classical language belonging to the 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 the power ...
'' stolō'', genitive ''stolōnis'' – "branch"), also known as runners, are horizontal connections between organisms. They may be part of the organism, or of its skeleton; typically,
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
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 science (or plant sciences), 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 "bot ...
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 ...
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, line ...
s, and new plants from the buds. 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. 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
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'' 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 a ...
is defined as a horizontal stem above or on the soil surface that often roots at the internodes.
Plants with stolons
In some '' Cyperus'' 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 ...
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, t ...
'' (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'', and '' Pilosella'' (Hawkweeds), '' Zoysia japonica'', '' Ranunculus repens''. Plants with long, slender stolons are referred to as ''sarmentose'' plants.
Other plants with stolons below the soil surface include many grasses, '' Ajuga'', '' Mentha'', and '' Stachys''. 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. I ...
''.
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 stolons start to grow within 10 days of plants emerging above ground, with tubers 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) The BBCH-scale (potato) identifies the phenological development stages of a potato (''Solanum tuberosum''). It is a plant species-specific version of the BBCH-scale.
{, class="wikitable" style="clear"
!rowspan="2", Growth stage!!colspan="2", Code ...
.
''
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'', 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 du ...
ous species produce stolons, such as '' Erythronium propullans''. Flowering plants often produce no stolons.
'' Convolvulus arvensis'' 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 tissue to keep the parent plant alive from one year to the next.
In mycology
In mycology, 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 together. Root-like structures called rhizoids 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 (locomotion), the surface over which an organism lo ...
. The stolon is commonly found in
bread mold
A mold () or mould () is one of the structures 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. Not ...
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 invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a ...
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 Fresh water, freshwater and Marine habitats, marine environments, predominantly the latter.
Their distinguishing feature is cnidocyt ...
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 they trail behind the main body.
Palaeontology
Stolon based reproduction is thought to have been used by Rangeomorphs 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