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The soil seed bank is the natural storage of
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s, often dormant, within 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 te ...
of most
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s. The study of soil seed banks started in 1859 when
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
observed the emergence of seedlings using soil samples from the bottom of a lake. The first scientific paper on the subject was published in 1882 and reported on the occurrence of seeds at different soil depths. Weed seed banks have been studied intensely in agricultural science because of their important economic impacts; other fields interested in soil seed banks include
forest regeneration Reforestation (occasionally, reafforestation) is the natural or intentional restocking of existing forests and woodlands (forestation) that have been depleted, usually through deforestation, but also after clearcutting. Management A debat ...
and restoration ecology.


Background

Many
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
have been classified according to the longevity of their seeds in the soil seed bank. Seeds of ''transient'' species remain viable in the soil seed bank only to the next opportunity to
germinate Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
, while seeds of ''persistent'' species can survive longer than the next opportunity—often much longer than one year. Species with seeds that remain viable in the soil longer than five years form the ''long-term'' persistent seed bank, while species whose seeds generally germinate or die within one to five years are called ''short-term'' persistent. A typical long-term persistent species is ''
Chenopodium album ''Chenopodium album'' is a fast-growing weedy annual plant in the genus ''Chenopodium''. Though cultivated in some regions, the plant is elsewhere considered a weed. Common names include lamb's quarters, melde, goosefoot, wild spinach and fat-h ...
'' (Lambsquarters); its seeds commonly remain viable in the soil for up to 40 years and in rare situations perhaps as long as 1,600 years. A species forming no soil seed bank at all (except the dry season between ripening and the first autumnal rains) is ''
Agrostemma githago ''Agrostemma githago'', the common corn-cockle (also written "corncockle") is a herbaceous annual flowering plant in the pink and carnation family Caryophyllaceae. Description It grows with a stem to long with lanceolate leaves. The flowers ar ...
'' (Corncockle), which was formerly a widespread cereal weed.


Seed longevity

Longevity of seeds is very variable and depends on many factors; few species exceed 100 years. In typical soils the longevity of seeds can range from nearly zero (germinating immediately when reaching the soil or even before) to several hundred years. Some of the oldest still-viable seeds were those of Lotus (''Nelumbo nucifera'') found buried in the soil of a pond; these seeds were estimated by
carbon dating Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was dev ...
to be around 1,200 years old. One
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...
of date palm, the
Judean date palm The Judean date palm is a date palm (''Phoenix dactylifera'') grown in Judea. It is not clear whether there was ever a single distinct Judean cultivar, but dates grown in the region have had distinctive reputations for thousands of years, and the ...
, successfully sprouted in 2008 after accidental storage for 2,000 years. One of the longest-running soil seed viability trials was started in
Michigan Michigan () is a state in the Great Lakes region of the upper Midwestern United States. With a population of nearly 10.12 million and an area of nearly , Michigan is the 10th-largest state by population, the 11th-largest by area, and the ...
in 1879 by James Beal. The experiment involved the burying of 20 bottles holding 50 seeds from 21 species. Every five years, a bottle from every species was retrieved and germinated on a tray of sterilized soil which was kept in a growth chamber. Later, after responsibility for managing the experiment was delegated to caretakers, the period between retrievals became longer. In 1980, more than 100 years after the trial was started, seeds of only three species were observed to germinate: moth mullein (''Verbascum blattaria''), common mullein (''Verbascum thapsus'') and
common mallow Common mallow is a common name used for a number of different species of ''Malva'': *''Malva sylvestris'' is the common mallow in Europe and North Africa *''Malva neglecta'' is the common mallow in the United States *'' Malva preissiana'' is a "com ...
(''Malva neglecta'').


Environmental significance

Soil seed banks are a crucial part of the rapid re-vegetation of sites disturbed by wildfire, catastrophic weather, agricultural operations, and timber harvesting, a natural process known as
secondary succession Secondary succession is the secondary ecological succession of a plant's life. As opposed to the first, primary succession, secondary succession is a process started by an event (e.g. forest fire, harvesting, hurricane, etc.) that reduces an a ...
. Soil seed banks are often dominated by
pioneer species Pioneer species are hardy species that are the first to colonize barren environments or previously biodiverse steady-state ecosystems that have been disrupted, such as by wildfire. Pioneer flora Some lichens grow on rocks without soil, so ...
, those species that are specially adapted to return to an environment first after a disturbance.
Forest ecosystem Forest ecology is the scientific study of the interrelated patterns, processes, flora, fauna and ecosystems in forests. The management of forests is known as forestry, silviculture, and forest management. A forest ecosystem is a natural woodland u ...
s and
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The ...
s contain a number of specialized plant species forming persistent soil seed banks. Before the advent of herbicides, a good example of a persistent seed bank species was ''
Papaver rhoeas ''Papaver rhoeas'', with common names including common poppy, corn poppy, corn rose, field poppy, Flanders poppy, and red poppy, is an annual herbaceous species of flowering plant in the poppy family Papaveraceae. It is a temperate native with ...
'', sometimes so abundant in agricultural fields in Europe that it could be mistaken for a crop. The absence of a soil seed bank impedes the establishment of vegetation during
primary succession Primary succession is the beginning step of ecological succession after an extreme disturbance, which usually occurs in an environment devoid of vegetation and other organisms. These environments are typically lacking in soil, as disturbances l ...
, while presence of a well-stocked soil seed bank permits rapid development of species-rich ecosystems during
secondary succession Secondary succession is the secondary ecological succession of a plant's life. As opposed to the first, primary succession, secondary succession is a process started by an event (e.g. forest fire, harvesting, hurricane, etc.) that reduces an a ...
.


Population densities and diversity

The mortality of seeds in the soil is one of the key factors for the persistence and density fluctuations of plant populations, especially for annual plants. Studies on the genetic structure of ''Androsace septentrionalis'' populations in the seed bank compared to those of established plants showed that diversity within populations is higher below ground than above ground. There are indications that mutations are more important for species forming a persistent seed bank compared to those with only transient seeds. The increase of species richness in a plant community due to a species-rich and abundant soil seed bank is known as the ''storage effect''. Species of ''
Striga ''Striga'', commonly known as witchweed, is a genus of parasitic plants that occur naturally in parts of Africa, Asia, and Australia. It is currently classified in the family Orobanchaceae, although older classifications place it in the Scrophul ...
'' (witchweed) are known to leave some of the highest seed densities in the soil compared to other plant
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 nomenclat ...
; this is a major factor that aids their invasive potential. Each plant has the capability to produce between 90,000 and 450,000 seeds, although a majority of these seeds are not viable. It has been estimated that only two witchweeds would produce enough seeds required to refill a seed bank after seasonal losses.


Associated ecosystem processes

The term soil
diaspore Diaspore , also known as diasporite, empholite, kayserite, or tanatarite, is an aluminium oxide hydroxide mineral, α-AlO(OH), crystallizing in the orthorhombic system and isomorphous with goethite. It occurs sometimes as flattened crystals, bu ...
bank can be used to include non-flowering plants such as
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s and
bryophyte The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in ...
s. In addition to seeds,
perennial plant A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
s have vegetative
propagule In biology, a propagule is any material that functions in propagating an organism to the next stage in its life cycle, such as by dispersal. The propagule is usually distinct in form from the parent organism. Propagules are produced by organisms ...
s to facilitate forming new plants, migration into new ground, or reestablishment after being top-killed. These propagules are collectively called the 'soil bud bank', and include dormant and adventitious buds on stolons,
rhizomes 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 hori ...
, and bulbs.


The relationship between soil seed bank and aboveground vegetation

Soil seed bank is significant breeding source for vegetation restoration and species-rich vegetation restoration, as they provide memories of past vegetation and represent the structure of future population. Soil seed bank did not very differ in overall seed density or species diversity, and there was little relative between the species composition of the seed bank and the composition of the aboveground vegetation. These two facts could lead to the conclusion that the species composition of the aboveground vegetation and the soil bank can differ. Additionally, it is a key point that the relationship between soil seed bank and original potential to measure the revegetation potential. In endangered habitats, such as mudflats, rare and critically endangered species may be present in high densities within the soil seed bank and survive between 50 years or a century.


References

{{reflist Plant reproduction Seeds Gene banks Ecological restoration Biology terminology Habitat Environmental terminology Biorepositories