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The soil seed bank is the natural storage of seeds, often dormant, within the soil of most ecosystems.^[1] The study of soil seed banks started in 1859 when Charles Darwin 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.^[2] 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 and restoration ecology.

Henry David Thoreau wrote that the contemporary popular belief explaining the succession 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.^[3]

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 plants.^[4] Perennial plants have vegetative propagules 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 on stolons, rhizomes, and bulbs. Moreover, the term soil diaspore bank can be used to include non-flowering plants such as ferns and bryophytes.^{[citation needed]}

Soil seed bank is significant breeding source for vegetation restoration^[5] and species-rich vegetation restoration,^[6] as they provide memories of past vegetation and represent the structure of future population.^[6] Moreover the composition of seed bank is often more stable than the vegetation to environmental changes,^[7] although a chronic N deposition can deplete it.^[8]^[9] In many systems, the density of the soil seed bank is often lower than the vegetation,^[4] and there are a large differences in species composition of the seed bank and the composition of the aboveground vegetation.^[10]^[11]^[12] Additionally, it is a key point that the relationship between soil seed bank and original potential to measure the revegetation potential.^[13]^[14] 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][7],^[15]

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. Soil seed banks are often dominated by pioneer species, those species that are specially adapted to return to an environment first after a disturbance.^[16] Forest ecosystems and wetlands contain a number of specialized plant species forming persistent soil seed banks.^{[citation needed]}

The absence of a soil seed bank impedes the establishment of vegetation during primary succession, while presence of a well-stocked soil seed bank permits rapid development of species-rich ecosystems during secondary succession.^{[citation needed]}

Seed longevity

Many taxa 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, 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.^[17] 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.^{[citation needed]}

Longevity of seeds is very variable and depends on many factors. Seeds buried more deeply tend to be capable of lasting longer.^[18] However, few species exceed 100 years.^[19] 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 to be around 1,200 years old.^[20] One cultivar of date palm, the Judean date palm, successfully sprouted in 2008 after accidental storage for 2,000 years.^[21]

The famous seed longevity experiments

One of the longest-running soil seed viability trials was started in Michigan 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).^[22] Several other experiments have been conducted to determine the long-term longevity of seeds in soil seed banks.

Soil seed bank longevity of seeds in experimental conditions
Species	Time	Comments
Verbascum blattaria	At least 142 years^[23]
Verbascum thapsus	At least 100 years ^[22]
Malva neglecta	At least 100 years
Oenothera biennis	80 years^[24]	10% of seeds sprouted after the 80-year mark
Rumex crispus	80 years	Only 2% of seeds survived to this point.^[24]
Datura stramonium	At least 39 years	Over 90 percent germination rate was reported^[25]
Phytolacca americana	At least 39 years	80-90 percent germination rate was reported^[25]
Solanum nigrum	At least 39 years	Over 80 percent germination rate was reported^[25]
Robinia pseudoacacia	At least 39 years
Ambrosia artemisiifolia	At least 39 years
Potentilla norvegica	At least 39 years
Onopordum acanthium	At least 39 years
Rudbeckia hirta	At least 39 years
Cuscuta polygonorum	At least 39 years
Lespedeza frutescens	At least 39 years
Convolvulus sepium	At least 39 years
Ipomoea lacunosa	At least 39 years
Verbena hastata	At least 39 years
Verbena urticifolia	At least 39 years
Nicotiana tabacum	At least 39 years
Arctium lappa	At least 39 years	Only 1 percent germination was reported.
Boehmeria nivea	At least 39 years
Setaria verticillata	At least 39 years
Trifolium pratense	At least 39 years
Rumex obtusifolius	At least 39 years
Rumex salicifolius	At least 39 years
Chenopodium album	At least 39 years
Chenopodium hybridum	At least 39 years
Abutilon theophrasti	At least 39 years
Leucanthemum vulgare	At least 39 years
Hibiscus militaris	At least 39 years
Hypericum hypericoides	At least 39 years
Sporobolus cryptandrus	At least 39 years
Polygonum scandens	At least 39 years	Germination rate was very low throughout the experiment.
Poa pratensis	At least 39 years
Setaria viridis	At least 39 years
Phalaris arundinacea	30 years	Only 1 percent of seed survived.
Portulaca oleracea	30 years	38 percent of the most deeply buried seeds were viable at 21 years, 1 percent of more shallowly buried seeds are reported sprouting after the 30 year mark.
Polygonum pensylvanicum	30 years
Polygonum persicaria	30 years
Cassia marilandica	30 years
Thlaspi arvense	30 years
Trifolium hybridum	30 years
Ambrosia trifida	21 years
Brassica nigra	21 years
Dracocephalum parviflorum	24.7 years^[26]
Rorippa islandica	24.7 years
Matricaria discoidea	24.7 years
Polygonum aviculare	24.7 years
Helianthus annuus	17 years^[18]
Setaria parviflora	17 years
Cirsium arvense	17 years
Cirsium flodmanii	17 years
Ipomoea hederacea	17 years
Persicaria amphibia	17 years
Amaranthus tuberculatus	17 years
Solanum sarrachoides	17 years
Ambrosia grayii	17 years	Only 1% of seed germinated.
Bassia scoparia	17 years	Only 1% of seed germinated.
Echinochloa crus-galli	17 years	Only 1% of seed germinated.
Amaranthus retroflexus	12 years^[18]
Pyrus calleryana	At least 11 years^[27]

Other studies

Species of Striga (witchweed) are known to leave some of the highest seed densities in the soil compared to other plant genera; this is a major factor that aids their invasive potential.^[28] Each plant has the capability to produce between 90,000 and 450,000 seeds, although a majority of these seeds are not viable.^[29] It has been estimated that only two witchweeds would produce enough seeds required to refill a seed bank after seasonal losses.^[30] 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.^{[citation needed]}

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.^{[citation needed]}

References

^ Jack Dekker (1997). "The Soil Seed Bank". Agronomy Department, Iowa State University. Retrieved 10 December 2015.
^ Christoffoleti, P. J.; Caetano, R. S. X. (July 17, 1998). "Soil seed banks". Scientia Agricola. 55: 74–78. doi:10.1590/S0103-90161998000500013.
^ Mcartney, Eugene S. (1931). "Forest Succession and Folklore". The Classical Weekly. 25 (6): 47–48. doi:10.2307/4389644. JSTOR 4389644.
^ ^a ^b DeMalach, Niv; Kigel, Jaime; Sternberg, Marcelo (2023-03-01). "Contrasting dynamics of seed banks and standing vegetation of annuals and perennials along a rainfall gradient". Perspectives in Plant Ecology, Evolution and Systematics. 58: 125718. arXiv:2301.12696. doi:10.1016/j.ppees.2023.125718. ISSN 1433-8319.
^ Lu, Z.J., Li, L.F., Jiang, M.X., Huang, H.D., and Bao, D.C., Can the soil seed bank contribute to revegetation of the drawdown zone in the Three Gorges reservoir region? Plant Ecol., 2010, vol. 209, no. 1, pp. 153–165.
^ ^a ^b Fisher, Judith L.; Loneragan, William A.; Dixon, Kingsley; Veneklaas, Erik J. (2009-02-01). "Soil seed bank compositional change constrains biodiversity in an invaded species-rich woodland". Biological Conservation. 142 (2): 256–269. doi:10.1016/j.biocon.2008.10.019. ISSN 0006-3207.
^ DeMalach, Niv; Kigel, Jaime; Sternberg, Marcelo (March 2021). Dalling, James (ed.). "The soil seed bank can buffer long-term compositional changes in annual plant communities". Journal of Ecology. 109 (3): 1275–1283. arXiv:2010.15693. doi:10.1111/1365-2745.13555. ISSN 0022-0477.
^ Eskelinen, Anu; Elwood, Elise; Harrison, Susan; Beyen, Eva; Gremer, Jennifer R. (December 2021). "Vulnerability of grassland seed banks to resource-enhancing global changes". Ecology. 102 (12): e03512. doi:10.1002/ecy.3512. ISSN 0012-9658. PMID 34358331.
^ Basto, Sofía; Thompson, Ken; Phoenix, Gareth; Sloan, Victoria; Leake, Jonathan; Rees, Mark (2015-02-04). "Long-term nitrogen deposition depletes grassland seed banks". Nature Communications. 6 (1): 6185. doi:10.1038/ncomms7185. ISSN 2041-1723.
^ Sanderson, M.A., Goslee, S.C., Klement, K.D., and Soder, K.J., Soil seed bank composition in pastures of diverse mixtures of temperate forages, Agron. J., 2007, vol. 99, no. 6, p. 1514.
^ White, S.; Bork, E.; Karst, J.; Cahill, J. (2012-11-21). "Similarity between grassland vegetation and seed bank shifts with altered precipitation and clipping, but not warming". Community Ecology. 13 (2): 129–136. doi:10.1556/comec.13.2012.2.1. ISSN 1588-2756.
^ Hopfensperger, K.N., A review of similarity between seed bank and standing vegetation across ecosystems, Oikos, 2007, vol. 116, pp. 1438–1448.
^ Lu, Z.J., Li, L.F., Jiang, M.X., Huang, H.D., and Bao, D.C., Can the soil seed bank contribute to revegetation of the drawdown zone in the Three Gorges reservoir region? Plant Ecol., 2010, vol. 209, no. 1, pp. 153–165
^ Wang, Yongcui; Jiang, Deming; Toshio, Oshida; Zhou, Quanlai (2013-09-01). "Recent advances in soil seed bank research". Contemporary Problems of Ecology. 6 (5): 520–524. doi:10.1134/S1995425513050181. ISSN 1995-4263. S2CID 255553677.
^ Poschlod, Peter; Rosbakh, Sergey (2018). "Mudflat species: Threatened or hidden? An extensive seed bank survey of 108 fish ponds in Southern Germany". Biological Conservation. 225: 154–163. doi:10.1016/j.biocon.2018.06.024. S2CID 91872044.
^ Tang, Yong; Cao, Min; Fu, Xianhui (2006). "Soil Seedbank in a Dipterocarp Rain Forest in Xishuangbanna, Southwest China". Biotropica. 38 (3): 328–333. doi:10.1111/j.1744-7429.2006.00149.x. S2CID 53974012.
^ "Iowa State University: College of Agriculture and Life Science: Lambsquarters".
^ ^a ^b ^c Burnside, Orvin C.; Wilson, Robert G.; Weisberg, Sanford; Hubbard, Kenneth G. (1996). "Seed Longevity of 41 Weed Species Buried 17 Years in Eastern and Western Nebraska". Weed Science. 44 (1): 74–86. doi:10.1017/S0043174500093589. S2CID 82721189.
^ Ken Thompson, Jan P. Bakker, and Renée M. Bekker. 1997. The soil seed banks of north west Europe : methodology, density and longevity. New York : Cambridge University Press. p. 276
^ J. Derek Bewley; Michael Black; Peter Halmer (2006). The Encyclopedia of Seeds: Science, Technology and Uses. CABI. pp. 14–15. ISBN 978-0-85199-723-0.
^ Fountain, Henry (2008-06-17). "Date Seed of Masada is Oldest Ever to Sprout". New York Times. Retrieved December 9, 2021.
^ ^a ^b Frank W. Telewski. "Research & Teaching". Department of Plant Biology, Michigan State University. Retrieved 10 December 2015.
^ "Unearthing a scientific mystery". msutoday.msu.edu. Michigan State University.
^ ^a ^b Darlington, H.T.; Steinbauer, G.P. (1961). "The Eighty-Year Period for Dr. Beal's Seed Viability Experiment". American Journal of Botany. 48 (4): 321–325. doi:10.1002/j.1537-2197.1961.tb11645.x.
^ ^a ^b ^c Brown, E.; Toole, E.H. (1946). "Final Results of the Duvel Buried Seed Experiment". Journal of Agricultural Research. 72 (6).
^ Conn, Jeffrey S.; Werdin-Pfisterer, Nancy R. (2010). "Variation in Seed Viability and Dormancy of 17 Weed Species after 24.7 Years of Burial: The Concept of Buried Seed Safe Sites". Weed Science. 58 (3): 209–215. doi:10.1614/WS-D-09-00084.1. S2CID 9103710.
^ Serota, Tziporah H.; Culley, Theresa M. (2019). "Seed Germination and Seedling Survival of Invasive Callery Pear (Pyrus calleryana Decne.) 11 Years After Fruit Collection". Castanea. 84 (1): 47. doi:10.2179/0008-7475.84.1.47. S2CID 191180173.
^ Ross, Merrill A.; Lembi, Carole A. (2008). Applied Weed Science: Including the Ecology and Management of Invasive Plants. Prentice Hall. p. 22. ISBN 978-0-13-502814-8.
^ Faiz F. Bebawi; Robert E. Eplee; Rebecca S. Norris (March 1984). "Effects of Seed Size and Weight on Witchweed (Striga asiatica) Seed Germination, Emergence, and Host-Parasitization". Weed Science. 32 (2): 202–205. doi:10.1017/S0043174500058811. JSTOR 4043831. S2CID 89078686.
^ Daniel M. Joel; Jonathan Gressel; Lytton J. Musselman (2013). Parasitic Orobanchaceae: Parasitic Mechanisms and Control Strategies. Springer Science & Business Media. p. 394. ISBN 978-3-642-38146-1.