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A classification system for seed dormancy

  • Jerry M. Baskin (a1) and Carol C. Baskin (a1) (a2)

The proposal is made that seed scientists need an internationally acceptable hierarchical system of classification for seed dormancy. Further, we suggest that a modified version of the scheme of the Russian seed physiologist Marianna G. Nikolaeva be adopted. The modified system includes three hierarchical layers – class, level and type; thus, a class may contain levels and types, and a level may contain only types. The system includes five classes of dormancy: physiological dormancy (PD), morphological dormancy (MD), morphophysiological dormancy (MPD), physical dormancy (PY) and combinational dormancy (PY + PD). The most extensive classification schemes are for PD, which contains three levels and five types (in the non-deep level), and MPD, which contains eight levels but no types. PY is not subdivided at all but probably should be, for reasons given. Justifications are presented for not including mechanical dormancy or chemical dormancy in the modified scheme. PD (non-deep level) is the most common kind of dormancy, and occurs in gymnosperms (Coniferales, Gnetales) and in all major clades of angiosperms. Since, first, this is the class and level of dormancy in seeds of wild populations of Arabidopsis thaliana and, secondly, Type 1 (to which seeds of A. thaliana belong) is also common, and geographically and phylogenetically widespread, it seems that biochemical, molecular and genetic studies on seed dormancy in this model species might have rather broad application in explaining the basic mechanism(s) of physiological dormancy in seeds.

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R.D. Amen (1968) A model of seed dormancy. The Botanical Review 34, 131.

Angiosperm Phylogeny Group (APG) (1998) An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden 85, 531553.

C.C. Baskin and J.M. Baskin (1998) Seeds: Ecology, biogeography, and evolution of dormancy and germination. San Diego, Academic Press.

C.C. Baskin , P.L. Chesson and J.M. Baskin (1993) Annual seed dormancy cycles in two desert winter annuals. Journal of Ecology 81, 551556.

C.C. Baskin , O. Zackrisson and J.M. Baskin (2002) Role of warm stratification in promoting germination of seeds of Empetrum hermaphroditum (Empetraceae), a circumboreal species with a stony endocarp. American Journal of Botany 89, 486493.

J.M. Baskin and C.C. Baskin (1972) Ecological life cycle and physiological ecology of seed germination of Arabidopsis thaliana. Canadian Journal of Botany 50, 353360.

J.M. Baskin and C.C. Baskin (1980) Ecophysiology of secondary dormancy in seeds of Ambrosia artemisiifolia. Ecology 61, 475480.

J.M. Baskin and C.C. Baskin (1983) Seasonal changes in the germination responses of buried seeds of Arabidopsis thaliana and ecological interpretation. Botanical Gazette 144, 540543.

J.M. Baskin and C.C. Baskin (1985) The annual dormancy cycle in buried weed seeds: A continuum. BioScience 35, 492498.

J.M. Baskin and C.C. Baskin (1986) Changes in dormancy status of Frasera caroliniensis seeds during overwintering on parent plant. American Journal of Botany 73, 510.

J.M. Baskin , X. Nan and C.C. Baskin (1998) A comparative study of seed dormancy and germination in an annual and a perennial species of Senna (Fabaceae). Seed Science Research 8, 501512.

J.M. Baskin , C.C. Baskin and X. Li (2000) Taxonomy, ecology, and evolution of physical dormancy in seeds. Plant Species Biology 15, 139152.

N.C.W. Beadle (1952) Studies in halophytes. I. The germination of seeds and establishment of seedlings of five species of Atriplex in Australia. Ecology 33, 4962.

N. Beaudoin , C. Serizet , F. Gosti and J. Giraudat (2000) Interactions between abscisic acid and ethylene signaling cascades. The Plant Cell 12, 11031115.

T. Berry and J.D. Bewley (1992) A role of the surrounding fruit tissues in preventing the germination of tomato ( Lycopersicon esculentum ) seeds. A consideration of the osmotic environment and abscisic acid. Plant Physiology 100, 951957.

J.D. Bewley (1997a) Seed germination and dormancy. The Plant Cell 9, 10551066.

J.D. Bewley (1997b) Breaking down the walls – a role for endo-β-mannanase in release from seed dormancy? Trends in Plant Science 2, 464469.

H.J. Bouwmeester and C.M. Karssen (1992) The dual role of temperature in the regulation of the seasonal changes in dormancy and germination of seeds of Polygonum persicaria L. Oecologia 90, 8894.

C.J. Burrows (1995) Germination behavior of the seeds of the New Zealand species Aristotelia serrata, Coprosma robusta, Cordyline australis, Myrtus obcordata, and Schefflera digitata. New Zealand Journal of Botany 33, 257264.

C.J. Burrows (1999) Germination behavior of seeds of the New Zealand woody species Alseuosmia macrophylla, A. pusilla, Cordyline banksii, Geniostoma rupestre, Myrtus bullata, and Solanum aviculare. New Zealand Journal of Botany 37, 277287.

I. Debeaujon and M. Koornneef (2000) Gibberellin requirement for Arabidopsis seed germination is determined both by testa characteristics and embryonic abscisic acid. Plant Physiology 122, 415424.

L. de Miguel , R.A. Sánchez (1992) Phytochrome-induced germination, endosperm softening and embryo growth potential in Datura ferox seeds: Sensitivity to low water potential and time to escape FR reversal. Journal of Experimental Botany 43, 969974.

M.P.M. Derkx and C.M. Karssen (1994) Are seasonal dormancy patterns in Arabidopsis thaliana regulated by changes in seed sensitivity to light, nitrate and gibberellin? Annals of Botany 73, 129136.

B. Downie and J.D. Bewley (1996) Dormancy in white spruce ( Picea glauca [Moench] Voss) seeds imposed by tissues surrounding the embryo. Seed Science Research 6, 915.

B. Downie , H.W.M. Hilhorst and J.D. Bewley (1997) Endo-β-mannanase activity during dormancy alleviation and germination of white spruce ( Picea glauca ) seeds. Physiologia Plantarum 101, 405415.

W.E. Dyer (1993) Dormancy-associated embryonic mRNAs and proteins in imbibing Avena fatua caryopses. Physiologia Plantarum 88, 201211.

S.A. Fennimore and M.E. Foley (1998) Genetic and physiological evidence for the role of gibberellic acid in the germination of dormant Avena fatua seeds. Journal of Experimental Botany 49, 8994.

M.E. Foley (2001) Seed dormancy: An update on terminology, physiological genetics, and quantitative trait loci regulating germinability. Weed Science 49, 305317.

M.E. Foley and S.A. Fennimore (1998) Genetic basis for seed dormancy. Seed Science Research 8, 173182.

T.A. Forbis and P.K. Diggle (2001) Subnivean embryo development in the alpine herb Caltha leptosepala (Ranunculaceae). Canadian Journal of Botany 79, 635642.

T.A. Forbis , S.K. Floyd , A. de Querioz (2002) The evolution of embryo size in angiosperms and other seed plants: Implications for the evolution of seed dormancy. Evolution 56, 21122125.

G. Garello , P. Barthe , M. Bonelli , J. Bianco-Trinchant , J. Bianco , M.-T. Le Page-Degivry (2000) Abscisic acid-regulated responses of dormant and non-dormant embryos of Helianthus annuus: Role of ABA-inducible proteins. Plant Physiology and Biochemistry 38, 473482.

S.C. Garvin and S.E. Meyer (2003) Multiple mechanisms for seed dormancy regulation in shadscale ( Atriplex confertifolia: Chenopodiaceae). Canadian Journal of Botany 81, 601610.

M. Ghassemian , E. Nambara , S. Cutler , H. Kawaide , Y. Kamiya and P. McCourt (2000) Regulation of abscisic acid signaling by the ethylene response pathway in Arabidopsis. The Plant Cell 12, 11171126.

P.J. Goldmark , J. Curry , C.F. Morris , M.K. Walker-Simmons (1992) Cloning and expression of an embryo-specific mRNA up-regulated in hydrated dormant seeds. Plant Molecular Biology 19, 433441.

P. Grappin , D. Bouinot , B. Sotta , E. Miginiac and M. Jullien (2000) Control of seed dormancy in Nicotiana plumbaginifolia: Post-imbibition abscisic acid synthesis imposes dormancy maintenance. Planta 210, 279285.

S.P.C. Groot and C.M. Karssen (1992) Dormancy and germination of abscisic acid-deficient tomato seeds. Studies with the sitiens mutant. Plant Physiology 99, 952958.

Y. Gutterman (1993) Seed germination in desert plants. Berlin, Springer-Verlag.

H.W.M. Hilhorst (1995) A critical update on seed dormancy. I. Primary dormancy. Seed Science Research 5, 6173.

H.W.M. Hilhorst (1998) The regulation of secondary dormancy. The membrane hypothesis revisited. Seed Science Research 8, 7790.

H.W.M. Hilhorst and B. Downie (1995) Primary dormancy in tomato ( Lycopersicon esculentum cv. Moneymaker): Studies with the sitiens mutant. Journal of Experimental Botany 47, 8997.

H.W.M. Hilhorst and C.M. Karssen (1992) Seed dormancy and germination: The role of abscisic acid and gibberellins and the importance of hormone mutants. Plant Growth Regulation 11, 225238.

M. Holdsworth , S. Kurup and R. McKibbin (1999) Molecular and genetic mechanisms regulating the transition from embryo development to germination. Trends in Plant Science 4, 275280.

S.B. Jarvis , M.A. Taylor , M.R. MacLeod and H.V. Davies (1996) Cloning and characterisation of the cDNA clones of three genes that are differentially expressed during dormancy-breakage in the seeds of Douglas fir ( Pseudotsuga menziesii ). Journal of Plant Physiology 147, 559566.

S.B. Jarvis , M.A. Taylor , J. Bianco , F. Corbineau and H.V. Davies (1997) Dormancy-breakage in seeds of Douglas fir ( Pseudotsuga menziesii (Mirb.) Franco). Support for the hypothesis that LEA gene expression is essential for the process. Journal of Plant Physiology 151, 457464.

R.R. Johnson , H.J. Cranston , M.E. Chaverra and W.E. Dyer (1995) Characterization of cDNA clones for differentially expressed genes in embryos of dormant and nondormant Avena fatua L. caryopses. Plant Molecular Biology 28, 113122.

C.M. Karssen and E. Lacka (1986) A revision of the hormone balance theory of seed dormancy: Studies on gibberellin and/or abscisic acid-deficient mutants of Arabidopsis thaliana. pp. 315323. in M. Bopp Plant growth substances 1985. Berlin, Springer-Verlag.

J. Kępczyński , E. Kępczyńska (1997) Ethylene in seed dormancy and germination. Physiologia Plantarum 101, 720726.

D. Koller (1957) Germination-regulating mechanisms in some desert seeds. I. Atriplex dimorphostegia Kar. et Kir. Ecology 38, 113.

M. Koornneef , L. Bentsink and H. Hilhorst (2002) Seed dormancy and germination. Current Opinion in Plant Biology 5, 3336.

G. Leubner-Metzger (2003) Functions and regulation of β-1,3-glucanases during seed germination, dormancy release and after-ripening. Seed Science Research 13, 1734.

B. Li and M.E. Foley (1994) Differential polypeptide patterns in imbibed dormant and after-ripened Avena fatua embryos. Journal of Experimental Botany 45, 275279.

B. Li and M.E. Foley (1995) Cloning and characterization of differentially expressed genes in imbibed dormant and afterripened Avena fatua embryos. Plant Molecular Biology 29, 823831.

B. Li and M.E. Foley (1997) Genetic and molecular control of seed dormancy. Trends in Plant Science 2, 384389.

X. Li , J.M. Baskin and C.C. Baskin (1999) Physiological dormancy and germination requirements of seeds of several North American Rhus species (Anacardiaceae). Seed Science Research 9, 237245.

O. Lorenzo , C. Nicolás , G. Nicolás and D. Rodriquez (2002) GA 3 -induced expression of a new functional AAA-ATPase (FsA1) is correlated with the onset of germination in Fagus sylvatica L. seeds. Plant and Cell Physiology 43, 2734.

B. Mandák , P. Pyšek (2001) The effect of light quality, nitrate concentration and presence of bracteoles on germination of different fruit types in heterocarpous Atriplex sagittata. Journal of Ecology 89, 149158.

A.J. Matilla (2000) Ethylene in seed formation and germination. Seed Science Research 10, 111126.

A. Meisert (2002) Physical dormancy in Geraniaceae seeds. Seed Science Research 12, 121128.

A. Meisert , D. Schulz and H. Lehman (1999) Structural features underlying hardseededness in Geraniaceae. Plant Biology 1, 311314.

C.F. Morris , R.J. Anderberg , P.J. Goldmark , M.K. Walker-Simmons (1991) Molecular cloning and expression of abscisic acid-responsive genes in embryos of dormant wheat seeds. Plant Physiology 95, 814821.

D.A. Morrison , K. McClay , C. Porter and S. Rish (1998) The role of the lens in controlling heat-induced breakdown of testa-imposed dormancy in native Australian legumes. Annals of Botany 82, 3540.

C. Nicolás , G. Nicolás and D. Rodriquez (1996) Antagonistic effects of abscisic acid and gibberellic acid on the breaking of dormancy of Fagus sylvatica seeds. Physiologia Plantarum 96, 244250.

C.B. Osmond , O. Björkman and D.J. Anderson (1980) Physiological processes in plant ecology – Toward a synthesis with Atriplex. Berlin, Springer-Verlag.

N. Pallais (1995a) Storage factors control germination and seedling establishment of freshly harvested true potato seed. American Potato Journal 72, 427436.

C. Ren and A.R. Kermode (1999) Analyses to determine the role of the megagametophyte and other seed tissues in dormancy maintenance of yellow cedar ( Chamaecyparis nootkatensis ) seeds: Morphological, cellular and physiological changes following moist chilling and during germination. Journal of Experimental Botany 50, 14031419.

R.A. Sánchez , L. de Miguel (1997) Phytochrome promotion of mannan-degrading enzyme activities in the micropylar endosperm of Datura ferox seeds requires the presence of the embryo and gibberellin synthesis. Seed Science Research 7, 2733.

G.M. Simpson (1990) Seed dormancy in grasses. Cambridge, Cambridge University Press.

A.L. Takhtajan (1980) Outline of the classification of flowering plants (Magnoliophyta). The Botanical Review 46, 225359.

K. Thompson , R.M. Ceriani , J.P. Bakker and R.M. Bekker (2003) Are seed dormancy and persistence in soil related? Seed Science Research 13, 97100.

R.F. Thorne (2000) The classification and geography of flowering plants: Dicotyledons of the class Angiospermae. The Botanical Review 66, 441647.

P.F. Threadgill , J.M. Baskin and C.C. Baskin (1981) Dormancy in seeds of Frasera caroliniensis (Gentianaceae). American Journal of Botany 68, 8086.

W. Van der Schaar , C. Alonso-Blanco , K.M. Léon-Kloosterziel , R.C. Jansen , J.W. Van Ooijen and M. Koornneef (1997) QTL analysis of seed dormancy in Arabidopsis using recombinant inbred lines and MQM mapping. Heredity 79, 190200.

A. Vegis (1964) Dormancy in higher plants. Annual Review of Plant Physiology 15, 185224.

L.M. Vleeshouwers , H.J. Bouwmeester and C.M. Karssen (1995) Redefining seed dormancy: An attempt to integrate physiology and ecology. Journal of Ecology 83, 10311037.

J.L. Walck , C.C. Baskin and J.M. Baskin (1999) Seeds of Thalictrum mirabile (Ranunculaceae) require cold stratification for loss of nondeep simple morphophysiological dormancy. Canadian Journal of Botany 77, 17691776.

P.F. Wareing and P.F. Saunders (1971) Hormones and dormancy. Annual Review of Plant Physiology 22, 261288.

G.E. Welbaum , T. Tissaoui and K.J. Bradford (1990) Water relations of seed development and germination in muskmelon ( Cucumis melo L.). III. Sensitivity of germination to water potential and abscisic acid during development. Plant Physiology 92, 10291037.

G.E. Welbaum , W.J. Muthui , J.H. Wilson , R.L. Grayson and R.D. Fell (1995) Weakening of muskmelon perisperm envelope tissue during germination. Journal of Experimental Botany 46, 391400.

F.W. Went (1949) Ecology of desert plants. II. The effect of rain and temperature on germination and growth. Ecology 30, 113.

C.N. White and C.J. Rivin (2000) Gibberellins and seed development in maize. II. Gibberellin synthesis inhibition enhances abscisic acid signaling in cultured embryos. Plant Physiology 122, 10891097.

C.N. White , W.M. Proebsting , P. Hedden and C.J. Rivin (2000) Gibberellins and seed development in maize. I. Evidence that gibberellin/abscisic acid balance governs germination versus maturation pathways. Plant Physiology 122, 10811088.

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Seed Science Research
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