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The soil seed bank is the natural storage of
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s, often dormant, within 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 ...
of most
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
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 Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
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 Agricultural science (or agriscience for short) is a broad multidisciplinary field of biology that encompasses the parts of exact, natural, economic and social sciences that are used in the practice and understanding of agriculture. Professio ...
because of their important economic impacts; other fields interested in soil seed banks include forest regeneration and
restoration ecology Ecological restoration, or ecosystem restoration, is the process of assisting the recovery of an ecosystem that has been degraded, damaged, destroyed or transformed. It is distinct from Conservation movement, conservation in that it attempts t ...
.
Henry David Thoreau Henry David Thoreau (born David Henry Thoreau; July 12, 1817May 6, 1862) was an American naturalist, essayist, poet, and philosopher. A leading Transcendentalism, transcendentalist, he is best known for his book ''Walden'', a reflection upon sim ...
wrote that the contemporary popular belief explaining the
succession Succession is the act or process of following in order or sequence. Governance and politics *Order of succession, in politics, the ascension to power by one ruler, official, or monarch after the death, resignation, or removal from office of ...
of a logged forest, specifically to trees of a dissimilar species to the trees cut down, was that seeds either spontaneously generated in the soil, or sprouted after lying dormant for centuries. However, he dismissed this idea, noting that heavy nuts unsuited for distribution by wind were distributed instead by animals.


The ecological importance of seed bank

The seed bank is one of the key factors for the persistence and density fluctuations of plant populations, especially for
annual plant An annual plant is a plant that completes its life cycle, from germination to the production of seeds, within one growing season, and then dies. Globally, 6% of all plant species and 15% of herbaceous plants (excluding trees and shrubs) are ...
s.
Perennial plant In horticulture, the term perennial (''wikt:per-#Prefix, per-'' + ''wikt:-ennial#Suffix, -ennial'', "through the year") is used to differentiate a plant from shorter-lived annual plant, annuals and biennial plant, biennials. It has thus been d ...
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, which are analogous to seed bank in their persistence ability under disturbance. These propagules are collectively called the 'soil bud bank', and include dormant and adventitious
buds 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 ...
on
stolons In biology, a stolon ( 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 ar ...
,
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
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 ...
s. Moreover, the term soil
diaspore Diaspore ()also called diasporite, empholite, kayserite, or tanatariteis an aluminium hydroxide oxide mineral, α-AlO(OH), crystallizing in the orthorhombic system and isomorphous with goethite. It occurs sometimes as flattened crystals, but ...
bank can be used to include non-flowering plants such as
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s and
bryophyte Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s. 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. Moreover the composition of seed bank is often more stable than the vegetation to environmental changes, although a chronic N deposition can deplete it. In many systems, the density of the soil seed bank is often lower than the vegetation, and there are a large differences in
species composition Relative species abundance is a component of biodiversity and is a measure of how common or rare a species is relative to other species in a defined location or community.Hubbell, S. P. 2001. ''The unified neutral theory of biodiversity and biogeog ...
of the seed bank and the composition of the aboveground vegetation. 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, the composition of the seed bank is often more stable than the vegetation to environmental changes 7], 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 resilient species that are the first to colonize barren environments, or to repopulate disrupted biodiverse steady-state ecosystems as part of ecological succession. Various kinds of events can create good conditions for pi ...
, those species that are specially adapted to return to an environment first after a disturbance. Forest ecosystems and
wetland A wetland is a distinct semi-aquatic ecosystem whose groundcovers are flooded or saturated in water, either permanently, for years or decades, or only seasonally. Flooding results in oxygen-poor ( anoxic) processes taking place, especially ...
s contain a number of specialized plant species forming persistent soil seed banks. The absence of a soil seed bank impedes the establishment of vegetation during
primary succession Primary succession is the beginning step of ecological succession where species known as pioneer species colonize an uninhabited site, which usually occurs in an environment devoid of vegetation and other organisms. In contrast, secondary su ...
, 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 ...
.


Seed longevity

Many
taxa In biology, a taxon (back-formation from ''taxonomy''; : 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 name and ...
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 flowering plant, angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the sp ...
, 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'' (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'' (Corncockle), which was formerly a widespread cereal weed. Longevity of seeds is very variable and depends on many factors. Seeds buried more deeply tend to be capable of lasting longer. However, 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 ...
to be around 1,200 years old. One
cultivar A cultivar is a kind of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and which retains those traits when Plant propagation, propagated. Methods used to propagate cultivars include division, root a ...
of
date palm ''Phoenix dactylifera'', commonly known as the date palm, is a flowering-plant species in the palm family Arecaceae, cultivated for its edible sweet #Fruits, fruit called dates. The species is widely cultivated across North Africa, northern A ...
, the Judean date palm, successfully sprouted in 2008 after accidental storage for 2,000 years.


The famous seed longevity experiments

One of the longest-running soil seed viability trials was started in
Michigan Michigan ( ) is a peninsular U.S. state, state in the Great Lakes region, Great Lakes region of the Upper Midwest, Upper Midwestern United States. It shares water and land boundaries with Minnesota to the northwest, Wisconsin to the west, ...
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 (''Malva neglecta''). Several other experiments have been conducted to determine the long-term longevity of seeds in soil seed banks.


Other studies

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 (; : 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 ...
; 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. Before the advent of herbicides, a good example of a persistent seed bank species was '' Papaver rhoeas'', sometimes so abundant in agricultural fields in Europe that it could be mistaken for a crop. 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.


References

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