Abstract
The transition between dormancy and germination represents a critical stage in the life cycle of higher plants and is an important ecological and commercial trait. In this review we present current knowledge of the molecular control of this trait in Arabidopsis thaliana, focussing on important components functioning during the developmental phases of seed maturation, after-ripening and imbibition. Establishment of dormancy during seed maturation is regulated by networks of transcription factors with overlapping and discrete functions. Following desiccation, after-ripening determines germination potential and, surprisingly, recent observations suggest that transcriptional and post-transcriptional processes occur in the dry seed. The single-cell endosperm layer that surrounds the embryo plays a crucial role in the maintenance of dormancy, and transcriptomics approaches are beginning to uncover endosperm-specific genes and processes. Molecular genetic approaches have provided many new components of hormone signalling pathways, but also indicate the importance of hormone-independent pathways and of natural variation in key regulatory loci. The influence of environmental signals (particularly light) following after-ripening, and the effect of moist chilling (stratification) are increasingly being understood at the molecular level. Combined postgenomics, physiology and molecular genetics approaches are beginning to provide an unparalleled understanding of the molecular processes underlying dormancy and germination.
MeSH Terms
Abscisic Acid/biosynthesis,physiology
Arabidopsis/embryology,genetics,metabolism
Arabidopsis Proteins/genetics,metabolism,physiology
Gene Expression Profiling
Gene Expression Regulation, Plant
Gene Regulatory Networks
Genome, Plant
Germination/genetics
Gibberellins/biosynthesis,physiology
Indoleacetic Acids/metabolism
Seeds/genetics,growth & development,metabolism
Signal Transduction
Transcription Factors/genetics,metabolism,physiology
Chemicals
Arabidopsis Proteins
Gibberellins
Indoleacetic Acids
Transcription Factors
Abscisic Acid
gibberellic acid
Authors & Affiliations
3 authors, click to expand affiliations / ORCID
Holdsworth Michael J
Department of Agricultural and Environmental Sciences, School of BioSciences, University of Nottingham, Nottingham LE12 5RD, UK.
Bentsink Leónie
Department of Molecular Plant Physiology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands.
Soppe Wim J J
Department of Plant Breeding and Genetics, Max Planck Institute for Plant Breeding Research, 50829, Cologne, Germany.
References (197)
197 references, click to expand
-
Alboresi A, Gestin C, Leydecker MT, Bedu M, Meyer C, Truong HN. 2005. Nitrate, a signal relieving seed dormancy in Arabidopsis. Plant, Cell & Environment 28: 500-512.
-
Ali-Rachedi S, Bouinot D, Wagner MH, Bonnet M, Sotta B, Grappin P, Jullien M. 2004. Changes in endogenous abscisic acid levels during dormancy release and maintenance of mature seeds: studies with the Cape Verde Islands ecotype, the dormant model of Arabidopsis thaliana. Planta 219: 479-488.
-
Almoguera C, Jordano J. 1992. Developmental and environmental concurrent expression of sunflower dry-seed-stored low-molecular-weight heat-shock protein and LEA messenger-RNAs. Plant Molecular Biology 19: 781-792.
-
Alonso-Blanco C, Bentsink L, Hanhart CJ, Vries HBE, Koornneef M. 2003. Analysis of natural allelic variation at seed dormancy loci of Arabidopsis thaliana. Genetics 164: 711-729.
-
Ariizumi T, Steber CM. 2007. Seed germination of GA-insensitive sleepy1 mutants does not require RGL2 protein disappearance in Arabidopsis. Plant Cell 19: 791-804.
-
Bair NB, Meyer SE, Allen PS. 2006. A hydrothermal after-ripening time model for seed dormancy loss in Bromus tectorum L. Seed Science Research 16: 17-28.
-
Baker A, Graham IA, Holdsworth M, Smith SM, Theodolou FL. 2006. Chewing the fat: beta-oxidation in signalling and development. Trends in Plant Science 11: 124-132.
-
Barroco RM, Van Poucke K, Bergervoet JHW, De Veylder L, Groot SPC, Inze D, Engler G. 2005. The role of the cell cycle machinery in resumption of postembryonic development. Plant Physiology 137: 127-140.
-
Baskin JM, Baskin CC. 2004. A classification system for seed dormancy. Seed Science Research 14: 1-16.
-
Bassel GW, Zielinska E, Mullen RT, Bewley JD. 2004. Down-regulation of DELLA genes is not essential for germination of tomato, soybean, and Arabidopsis seeds. Plant Physiology 136: 2782-2789.
-
Bäumlein H, Misera S, Luerssen H, Kolle K, Horstmann C, Wobus U, Müller AJ. 1994. The FUS3 gene of Arabidopsis thaliana is a regulator of gene expression during late embryogenesis. Plant Journal 6: 379-387.
-
Beaudoin N, Serizet C, Gosti F, Giraudat J. 2000. Interactions between abscisic acid and ethylene signaling cascades. Plant Cell 12: 1103-1115.
-
Bentsink L, Jowett J, Hanhart CJ, Koornneef M. 2006. Cloning of DOG1, a quantitative trait locus controlling seed dormancy in Arabidopsis. Proceedings of the National Academy of Sciences, USA 103: 17042-17047.
-
Bentsink L, Koornneef M 2002. Seed dormancy and germination. In: Somerville CR, Meyerowitz EM, eds. The arabidopsis book. Rockville, MD, USA: American Society of Plant Biologists, 1-18.
-
Bethke PC, Libourel IGL, Aoyama N, Chung YY, Still DW, Jones RL. 2007. The Arabidopsis aleurone layer responds to nitric oxide, gibberellin, and abscisic acid and is sufficient and necessary for seed dormancy. Plant Physiology 143: 1173-1188.
-
Bewley JD. 1997a. Breaking down the walls - a role for endo-beta-mannanase in release from seed dormancy? Trends in Plant Science 2: 464-469.
-
Bewley JD. 1997b. Seed germination and dormancy. Plant Cell 9: 1055-1066.
-
Bewley JD, Black M. 1994. Seeds; Physiology of development and germination. New York, NY, USA: Plenum Press.
-
Bove J, Lucas P, Godin B, Oge L, Jullien M, Grappin P. 2005. Gene expression analysis by cDNA-AFLP highlights a set of new signaling networks and translational control during seed dormancy breaking in Nicotiana plumbaginifolia. Plant Molecular Biology 57: 593-612.
-
Brady SM, McCourt P. 2003. Hormone cross-talk in seed dormancy. Journal of Plant Growth Regulation 22: 25-31.
-
Braybrook SA, Stone SL, Park S, Bui AQ, Le BH, Fischer RL, Goldberg RB, Harada JJ. 2006. Genes directly regulated by LEAFY COTYLEDON2 provide insight into the control of embryo maturation and somatic embryogenesis. Proceedings of the National Academy of Sciences, USA 103: 3468-3473.
-
Busk PK, Pages M. 1998. Regulation of abscisic acid-induced transcription. Plant Molecular Biology 37: 425-435.
-
Cadman CSC, Toorop PE, Hilhorst HWM, Finch-Savage WE. 2006. Gene expression profiles of Arabidopsis Cvi seeds during dormancy cycling indicate a common underlying dormancy control mechanism. Plant Journal 46: 805-822.
-
Cao DN, Cheng H, Wu W, Soo HM, Peng JR. 2006. Gibberellin mobilizes distinct DELLA-dependent transcriptomes to regulate seed germination and floral development in Arabidopsis. Plant Physiology 142: 509-525.
-
Cao DN, Hussain A, Cheng H, Peng JR. 2005. Loss of function of four DELLA genes leads to light- and gibberellin-independent seed germination in Arabidopsis. Planta 223: 105-113.
-
Carrera E, Holman T, Medhurst A, Dietrich D, Footitt S, Theodoulou FL, Holdsworth MJ. 2008. Seed after-ripening is a discrete developmental pathway associated with specific gene networks in Arabidopsis. Plant Journal 53: 214-224.
-
Carrera E, Holman T, Medhurst A, Peer W, Schmuths H, Footitt S, Theodoulou FL, Holdsworth MJ. 2007. Gene expression profiling reveals defined functions of the ABC transporter COMATOSE late in phase II of germination. Plant Physiology 143: 1669-1679.
-
Chen F, Bradford KJ. 2000. Expression of an expansin is associated with endosperm weakening during tomato seed germination. Plant Physiology 124: 1265-1274.
-
Chen F, Nonogaki H, Bradford KJ. 2002. A gibberellin-regulated xyloglucan endotransglycosylase gene is expressed in the endosperm cap during tomato seed germination. Journal of Experimental Botany 53: 215-223.
-
Chen Y, Ji FF, Xie H, Liang JS, Zhang JH. 2006. The regulator of G-protein signaling proteins involved in sugar and abscisic acid signaling in Arabidopsis seed germination. Plant Physiology 140: 302-310.
-
Chibani K, Ali-Rachedi S, Job C, Job D, Jullien M, Grappin P. 2006. Proteomic analysis of seed dormancy in Arabidopsis. Plant Physiology 142: 1493-1510.
-
Chiwocha SDS, Cutler AJ, Abrams SR, Ambrose SJ, Yang J, Ross ARS, Kermode AR. 2005. The etr1-2 mutation in Arabidopsis thaliana affects the abscisic acid, auxin, cytokinin and gibberellin metabolic pathways during maintenance of seed dormancy, moist-chilling and germination. Plant Journal 42: 35-48.
-
Choi HI, Hong JH, Ha JO, Kang JY, Kim SY. 2000. ABFs, a family of ABA-responsive element binding factors. Journal of Biological Chemistry 275: 1723-1730.
-
Colucci G, Apone F, Alyeshmerni N, Chalmers D, Chrispeels MJ. 2002. GCR1, the putative Arabidopsis G protein-coupled receptor gene is cell cycle-regulated, and its overexpression abolishes seed dormancy and shortens time to flowering. Proceedings of the National Academy of Sciences, USA 99: 4736-4741.
-
Comai L, Harada JJ. 1990. Transcriptional activities in dry seed nuclei indicate the timing of the transition from embryogeny to germination. Proceedings of the National Academy of Sciences, USA 87: 2671-2674.
-
Curaba J, Moritz T, Blervaque R, Parcy F, Raz V, Herzog M, Vachon G. 2004. AtGA3ox2, a key gene responsible for bioactive gibberellin biosynthesis, is regulated during embryogenesis by LEAFY COTYLEDON2 and FUSCA3 in Arabidopsis. Plant Physiology 136: 3660-3669.
-
Debeaujon I, Koornneef M. 2000. Gibberellin requirement for Arabidopsis seed germination is determined both by testa characteristics and embryonic abscisic acid. Plant Physiology 122: 415-424.
-
Debeaujon I, Léon-Kloosterziel KM, Koornneef M. 2000. Influence of the testa on seed dormancy, germination, and longevity in Arabidopsis. Plant Physiology 122: 403-413.
-
Dickinson CD, Evans RP, Nielsen NC. 1988. RY repeats are conserved in the 5′-flanking regions of legume seed-protein genes. Nucleic Acids Research 16: 371-371.
-
Donohue K, Dorn L, Griffith C, Kim E, Aguilera A, Polisetty CR, Schmitt J. 2005. The evolutionary ecology of seed germination of Arabidopsis thaliana: variable natural selection on germination timing. Evolution 59: 758-770.
-
Dubreucq B, Berger N, Vincent E, Boisson M, Pelletier G, Caboche M, Lepiniec L. 2000. The Arabidopsis AtEPR1 extensin-like gene is specifically expressed in endosperm during seed germination. Plant Journal 23: 643-652.
-
Dugas DV, Bartel B. 2004. MicroRNA regulation of gene expression in plants. Current Opinions in Plant Biology 7: 512-520.
-
Dure L, Waters L. 1965. Long-lived messenger RNA: evidence from cotton seed germination. Science 147: 410-412.
-
Eastmond PJ, Germain V, Lange PR, Bryce JH, Smith SM, Graham IA. 2000. Postgerminative growth and lipid catabolism in oilseeds lacking the glyoxylate cycle. Proceedings of the National Academy of Sciences, USA 97: 5669-5674.
-
Ellerstrom M, Stalberg K, Ezcurra I, Rask L. 1996. Functional dissection of a napin gene promoter: identification of promoter elements required for embryo and endosperm-specific transcription. Plant Molecular Biology 32: 1019-1027.
-
Eshed Y, Baum SF, Bowman JL. 1999. Distinct mechanisms promote polarity establishment in carpels of Arabidopsis. Cell 99: 199-209.
-
Ezcurra I, Ellerstrom M, Wycliffe P, Stalberg K, Rask L. 1999. Interaction between composite elements in the napA promoter: both the B-box ABA-responsive complex and the RY/G complex are necessary for seed-specific expression. Plant Molecular Biology 40: 699-709.
-
Ezcurra I, Wycliffe P, Nehlin L, Ellerstrom M, Rask L. 2000. Transactivation of the Brassica napus napin promoter by ABI3 requires interaction of the conserved B2 and B3 domains of ABI3 with different cis-elements: B2 mediates activation through an ABRE, whereas B3 interacts with an RY/G-box. Plant Journal 24: 57-66.
-
Fait A, Angelovici R, Less H, Ohad I, Urbanczyk-Wochniak E, Fernie AR, Galili G. 2006. Arabidopsis seed development and germination is associated with temporally distinct metabolic switches. Plant Physiology 142: 839-854.
-
Finch-Savage WE, Cadman CS, Toorop PE, Lynn JR, Hilhorst HW. 2007. Seed dormancy release in Arabidopsis Cvi by dry after-ripening, low temperature, nitrate and light shows common quantitative patterns of gene expression directed by environmentally specific sensing. Plant Journal 51: 60-78.
-
Finch-Savage WE, Leubner-Metzger G. 2006. Seed dormancy and the control of germination. New Phytologist 171: 501-523.
-
Finkelstein RR, Gampala SSL, Rock CD. 2002. Abscisic acid signaling in seeds and seedlings. Plant Cell 14: S15-S45.
-
Finkelstein RR, Lynch TJ. 2000. The Arabidopsis abscisic acid response gene ABI5 encodes a basic leucine zipper transcription factor. Plant Cell 12: 599-609.
-
Footitt S, Marquez J, Schmuths H, Baker A, Theodoulou FL, Holdsworth M. 2006. Analysis of the role of COMATOSE and peroxisomal beta-oxidation in the determination of germination potential in Arabidopsis. Journal of Experimental Botany 57: 2805-2814.
-
Footitt S, Slocombe SP, Larner V, Kurup S, Wu YS, Larson T, Graham I, Baker A, Holdsworth M. 2002. Control of germination and lipid mobilization by COMATOSE, the Arabidopsis homologue of human ALDP. EMBO Journal 21: 2912-2922.
-
Fujii H, Verslues PE, Zhu JK. 2007. Identification of two protein kinases required for abscisic acid regulation of seed germination, root growth, and gene expression in Arabidopsis. Plant Cell 19: 485-494.
-
Gallardo K, Job C, Groot SPC, Puype M, Demol H, Vandekerckhove J, Job D. 2001. Proteomic analysis of Arabidopsis seed germination and priming. Plant Physiology 126: 835-848.
-
Gallardo K, Job C, Groot SPC, Puype M, Demol H, Vandekerckhove J, Job D. 2002. Proteomics of Arabidopsis seed germination. A comparative study of wild-type and gibberellin-deficient seeds. Plant Physiology 129: 823-837.
-
Gao Y, Zeng Q, Guo J, Cheng J, Ellis BE, Chen JG. 2007. Genetic characterization reveals no role for the reported ABA receptor, GCR2, in ABA control of seed germination and early seedling development in Arabidopsis. Plant Journal 52: 1001-1013.
-
Ghassemian M, Nambara E, Cutler S, Kawaide H, Kamiya Y, McCourt P. 2000. Regulation of abscisic acid signaling by the ethylene response pathway in Arabidopsis. Plant Cell 12: 1117-1126.
-
Giraudat J, Hauge BM, Valon C, Smalle J, Parcy F, Goodman HM. 1992. Isolation of the Arabidopsis Abi3 gene by positional cloning. Plant Cell 4: 1251-1261.
-
Griffiths J, Murase K, Rieu I, Zentella R, Zhang ZL, Powers SJ, Gong F, Phillips AL, Hedden P, Sun TP et al . 2006. Genetic characterization and functional analysis of the GID1 gibberellin receptors in Arabidopsis. Plant Cell 18: 3399-3414.
-
Groot SPC, Karssen CM. 1992. Dormancy and germination of abscisic acid-deficient tomato seeds- studies with the sitiens mutant. Plant Physiology 99: 952-958.
-
Gutierrez L, Van Wuytswinkel O, Castelain M, Bellini C. 2007. Combined networks regulating seed maturation. Trends in Plant Science 12: 294-300.
-
Van Hengel AJ, Roberts K. 2003. AtAGP30, an arabinogalactan-protein in the cell walls of the primary root, plays a role in root regeneration and seed germination. Plant Journal 36: 256-270.
-
Heschel MS, Selby J, Butler C, Whitelam GC, Sharrock RA, Donohue K. 2007. A new role for phytochromes in temperature-dependent germination. New Phytologist 174: 735-741.
-
Hilhorst HWM, Karssen CM. 1992. Seed dormancy and germination - the role of abscisic-acid and gibberellins and the importance of hormone mutants. Plant Growth Regulation 11: 225-238.
-
Hill A, Nantel A, Rock CD, Quatrano RS. 1996. A conserved domain of the viviparous-1 gene product enhances the DNA binding activity of the bZIP protein EmBP-1 and other transcription factors. Journal of Biological Chemistry 271: 3366-3374.
-
Himmelbach A, Yang Y, Grill E. 2003. Relay and control of abscisic acid signaling. Current Opinions in Plant Biology 6: 470-479.
-
Hobo T, Kowyama Y, Hattori T. 1999. A bZIP factor, TRAB1, interacts with VP1 and mediates abscisic acid-induced transcription. Proceedings of the National Academy of Sciences, USA 96: 15348-15353.
-
Hoekstra FA, Golovina EA, Buitink J. 2001. Mechanisms of plant desiccation tolerance. Trends in Plant Science 6: 431-438.
-
Holdsworth MJ, Finch-Savage WE, Grappin P, Job D. 2008. Post-genomics dissection of seed dormancy and germination. Trends In Plant Science 13: 7-13.
-
Hooks MA, Turner JE, Murphy EC, Johnston KA, Burr S, Jaroslawski S. 2007. The Arabidopsis ALDP protein homologue COMATOSE is instrumental in peroxisomal acetate metabolism. Biochemical Journal 406: 399-406.
-
Hughes DW, Galau GA. 1989. Temporally modular gene-expression during cotyledon development. Genes and Development 3: 358-369.
-
Hugouvieux V, Kwak JM, Schroeder JI. 2001. An mRNA cap binding protein, ABH1, modulates early abscisic acid signal transduction in Arabidopsis. Cell 106: 477-487.
-
Ishibashi N, Yamauchi D, Minamikawa T. 1990. Stored mRNA in cotyledons of Vigna unguiculata seeds: nucleotide sequence of cloned cDNA for a stored mRNA and induction of its synthesis by precocious germination. Plant Molecular Biology 15: 59-64.
-
Job C, Rajjou L, Lovigny Y, Belghazi M, Job D. 2005. Patterns of protein oxidation in Arabidopsis seeds and during germination. Plant Physiology 138: 790-802.
-
Johnston CA, Temple BR, Chen JG, Gao YJ, Moriyama EN, Jones AM, Siderovski DP, Willard FS. 2007. Comment on ‘A G protein-coupled receptor is a plasma membrane receptor for the plant hormone abscisic acid’. Science 318: 914.
-
Kadosh D, Struhl K. 1998. Targeted recruitment of the Sin3-Rpd3 histone deacetylase complex generates a highly localized domain of repressed chromatin in vivo. Molecular and Cellular Biology 18: 5121-5127.
-
Kagaya Y, Toyoshima R, Okuda R, Usui H, Yamamoto A, Hattori T. 2005. LEAFY COTYLEDON1 controls seed storage protein genes through its regulation of FUSCA3 and ABSCISIC ACID INSENSITIVE3. Plant Cell Physiology 46: 399-406.
-
Karssen CM, Brinkhorst Van der Swan DLC, Breekland AE, Koornneef M. 1983. Induction of dormancy during seed development by endogenous Abscisic-Acid - studies on Abscisic-Acid deficient genotypes of Arabidopsis thaliana (L.) Heynh. Planta 157: 158-165.
-
Katagiri T, Ishiyama K, Kato T, Tabata S, Kobayashi M, Shinozaki K. 2005. An important role of phosphatidic acid in ABA signaling during germination in Arabidopsis thaliana. Plant Journal 43: 107-117.
-
Keith K, Kraml M, Dengler NG, McCourt P. 1994. fusca3: a heterochronic mutation affecting late embryo development in Arabidopsis. Plant Cell 6: 589-600.
-
Kepczynski J, Kepczynska E. 1997. Ethylene in seed dormancy and germination. Physiologia Plantarum 101: 720-726.
-
Kohler C, Hennig L, Spillane C, Pien S, Gruissem W, Grossniklaus U. 2003. The Polycomb-group protein MEDEA regulates seed development by controlling expression of the MADS-box gene PHERES1. Genes and Development 17: 1540-1553.
-
Koornneef M. 1978. Gibberellin sensitive mutants in Arabidopsis thaliana. Arabidopsis Information Service 15: 17-20.
-
Koornneef M, Elgersma A, Hanhart CJ, Vanloenenmartinet EP, Vanrijn L, Zeevaart JAD. 1985. A gibberellin insensitive mutant of Arabidopsis thaliana. Physiologia Plantarum 65: 33-39.
-
Koornneef M, Jorna ML, Brinkhorst Van der Swan DLC, Karssen CM. 1982. The isolation of abscisic acid (ABA) deficient mutants by selection of induced revertants in nongerminating gibberellin sensitive lines of Arabidopsis thaliana (L.) Heynh. Theoretical and Applied Genetics 61: 385-393.
-
Koornneef M, Karssen CM 1994. Seed dormancy and germination. In: Meyerowitz EM, Somerville CR, eds. Arabidopsis. Cold Spring Harbor, NY, USA: Cold Spring Harbor Laboratory Press, 313-334.
-
Koornneef M, Reuling G, Karssen CM. 1984. The isolation and characterization of abscisic acid insensitive mutants of Arabidopsis thaliana. Physiologia Plantarum 61: 377-383.
-
Koornneef M, Van der Veen JH. 1980. Induction and analysis of gibberellin-sensitive mutants in Arabidopsis thaliana (L.) Heynh. Theoretical and Applied Genetics 58: 257-263.
-
Kroj T, Savino G, Valon C, Giraudat J, Parcy F. 2003. Regulation of storage protein gene expression in Arabidopsis. Development 130: 6065-6073.
-
Kucera B, Cohn MA, Leubner-Metzger G. 2005. Plant hormone interactions during seed dormancy release and germination. Seed Science Research 15: 281-307.
-
Kuligowski J, Ferrand M, Chenou E. 1991. Stored mRNA in early embryos of a fern Marsilea vestita: a paternal and maternal origin. Molecular Reproduction and Development 30: 27-33.
-
Kurup S, Jones HD, Holdsworth MJ. 2000. Interactions of the developmental regulator ABI3 with proteins identified from developing Arabidopsis seeds. Plant Journal 21: 143-155.
-
Kushiro T, Okamoto M, Nakabayashi K, Yamagishi K, Kitamura S, Asami T, Hirai N, Koshiba T, Kamiya Y, Nambara E. 2004. The Arabidopsis cytochrome P450CYP707A encodes ABA 8′-hydroxylases: key enzymes in ABA catabolism. EMBO Journal 23: 1647-1656.
-
Kwong RW, Bui AQ, Lee H, Kwong LW, Fischer RL, Goldberg RB, Harada JJ. 2003. LEAFY COTYLEDON1-LIKE defines a class of regulators essential for embryo development. Plant Cell 15: 5-18.
-
Lara P, Onate-Sanchez L, Abraham Z, Ferrandiz C, Diaz I, Carbonero P, Vicente-Carbajosa J. 2003. Synergistic activation of seed storage protein gene expression in Arabidopsis by ABI3 and two bZIPs related to OPAQUE2. Journal of Biological Chemistry 278: 21003-21011.
-
Lee SC, Cheng H, King KE, Wang WF, He YW, Hussain A, Lo J, Harberd NP, Peng JR. 2002. Gibberellin regulates Arabidopsis seed germination via RGL2, a GAI/RGA-like gene whose expression is up-regulated following imbibition. Genes and Development 16: 646-658.
-
Lefebvre V, North H, Frey A, Sotta B, Seo M, Okamoto M, Nambara E, Marion-Poll A. 2006. Functional analysis of Arabidopsis NCED6 and NCED9 genes indicates that ABA synthesized in the endosperm is involved in the induction of seed dormancy. Plant Journal 45: 309-319.
-
Léon-Kloosterziel KM, Gil MA, Ruijs GJ, Jacobsen SE, Olszewski NE, Schwartz SH, Zeevaart JAD, Koornneef M. 1996. Isolation and characterization of abscisic acid-deficient Arabidopsis mutants at two new loci. Plant Journal 10: 655-661.
-
Leubner-Metzger G. 2003. Functions and regulation of beta-1,3-glucanases during seed germination, dormancy release and after-ripening. Seed Science Research 13: 17-34.
-
Leubner-Metzger G. 2005. Beta-1,3-glucanase gene expression in low-hydrated seeds as a mechanism for dormancy release during tobacco after-ripening. Plant Journal 41: 133-145.
-
Leubner-Metzger G, Petruzzelli L, Waldvogel R, Vogeli-Lange R, Meins F. 1998. Ethylene-responsive element binding protein (EREBP) expression and the transcriptional regulation of class I beta-1,3-glucanase during tobacco seed germination. Plant Molecular Biology 38: 785-795.
-
Leung, J, Merlot S, Giraudat. 1997. The Arabidopsis ABSCISIC ACID-INSENSITIVE2 (ABI2) and ABI1 genes encode homologous protein phosphatases 2C involved in abscisic acid signal transduction. Plant Cell 9: 759-771.
-
Leymarie J, Bruneaux E, Gibot-Leclerc S, Corbineau F. 2007. Identification of transcripts potentially involved in barley seed germination and dormancy using cDNA-AFLP. Journal of Experimental Botany 58: 425-437.
-
Li HC, Chuang K, Henderson JT, Rider SD, Bai Y, Zhang H, Fountain M, Gerber J, Ogas J. 2005. PICKLE acts during germination to repress expression of embryonic traits. Plant Journal 44: 1010-1022.
-
Liu PP, Koizuka N, Homrichhausen TM, Hewitt JR, Martin RC, Nonogaki H. 2005a. Large-scale screening of Arabidopsis enhancer-trap lines for seed germination-associated genes. Plant Journal 41: 936-944.
-
Liu PP, Koizuka N, Martin RC, Nonogaki H. 2005b. The BME3 (Blue Micropylar End 3) GATA zinc finger transcription factor is a positive regulator of Arabidopsis seed germination. Plant Journal 44: 960-971.
-
Liu PP, Montgomery TA, Fahlgren N, Kasschau KD, Nonogaki H, Carrington JC. 2007. Repression of AUXIN RESPONSE FACTOR10 by microRNA160 is critical for seed germination and post-germination stages. Plant Journal 52: 133-146.
-
Liu XG, Yue YL, Li W, Ma LG. 2007b. Response to comment on “A G protein-coupled receptor is a plasma membrane receptor for the plant hormone abscisic acid”. Science 318: 914.
-
Liu XG, Yue YL, Li B, Nie YL, Li W, Wu WH, Ma LG. 2007a. A G protein-coupled receptor is a plasma membrane receptor for the plant hormone abscisic acid. Science 315: 1712-1716.
-
Liu Y, Koornneef M, Soppe WJJ. 2007. The absence of histone H2B monoubiquitination in the Arabidopsis hub1 (rdo4) mutant reveals a role for chromatin remodeling in seed dormancy. Plant Cell 19: 433-444.
-
Lopez-Molina L, Mongrand S, Chua NH. 2001. A postgermination developmental arrest checkpoint is mediated by abscisic acid and requires the AB15 transcription factor in Arabidopsis. Proceedings of the National Academy of Sciences, USA 98: 4782-4787.
-
Lopez-Molina L, Mongrand S, Kinoshita N, Chua NH. 2003. AFP is a novel negative regulator of ABA signaling that promotes ABI5 protein degradation. Genes and Development 17: 410-418.
-
Lopez-Molina L, Mongrand B, McLachlin DT, Chait BT, Chua NH. 2002. ABI5 acts downstream of ABI3 to execute an ABA-dependent growth arrest during germination. Plant Journal 32: 317-328.
-
Lotan T, Ohto M, Yee KM, West MAL, Lo R, Kwong RW, Yamagishi K, Fischer RL, Goldberg RB, Harada JJ. 1998. Arabidopsis LEAFY COTYLEDON1 is sufficient to induce embryo development in vegetative cells. Cell 93: 1195-1205.
-
Luerssen K, Kirik V, Herrmann P, Misera S. 1998. FUSCA3 encodes a protein with a conserved VP1/ABI3-like B3 domain which is of functional importance for the regulation of seed maturation in Arabidopsis thaliana. Plant Journal 15: 755-764.
-
Mansfield SG, Briarty LG. 1996. The dynamics of seedling and cotyledon cell development in Arabidopsis thaliana during reserve mobilization. International Journal of Plant Sciences 157: 280-295.
-
Masubelele NH, Dewitte W, Menges M, Maughan S, Collins C, Huntley R, Nieuwland J, Scofield S, Murray JAH. 2005. D-type cyclins activate division in the root apex to promote seed germination in Arabidopsis. Proceedings of the National Academy of Sciences, USA 102: 15694-15699.
-
McGinnis KM, Thomas SG, Soule JD, Strader LC, Zale JM, Sun TP, Steber CM. 2003. The Arabidopsis SLEEPY1 gene encodes a putative F-box subunit of an SCF E3 ubiquitin ligase. Plant Cell 15: 1120-1130.
-
Meinke DW, Franzmann LH, Nickle TC, Yeung EC. 1994. Leafy cotyledon mutants of Arabidopsis. Plant Cell 6: 1049-1064.
-
Mitchum MG, Yamaguchi S, Hanada A, Kuwahara A, Yoshioka Y, Kato T, Tabata S, Kamiya Y, Sun TP. 2006. Distinct and overlapping roles of two gibberellin 3-oxidases in Arabidopsis development. Plant Journal 45: 804-818.
-
Monke G, Altschmied L, Tewes A, Reidt W, Mock HP, Bäumlein H, Conrad U. 2004. Seed-specific transcription factors ABI3 and FUS3: molecular interaction with DNA. Planta 219: 158-166.
-
Müller K, Tintelnot S, Leubner-Metzger G. 2006. Endosperm-limited Brassicaceae seed germination: abscisic acid inhibits embryo-induced endosperm weakening of Lepidium sativum (cress) and endosperm rupture of cress and Arabidopsis thaliana. Plant Cell Physiology 47: 864-877.
-
Nakabayashi K, Okamoto M, Koshiba T, Kamiya Y, Nambara E. 2005. Genome-wide profiling of stored mRNA in Arabidopsis thaliana seed germination: epigenetic and genetic regulation of transcription in seed. Plant Journal 41: 697-709.
-
Nakajima M, Shimada A, Takashi Y, Kim YC, Park SH, Ueguchi-Tanaka M, Suzuki H, Katoh E, Iuchi S, Kobayashi M et al . 2006. Identification and characterization of Arabidopsis gibberellin receptors. Plant Journal 46: 880-889.
-
Nakamura S, Lynch TJ, Finkelstein RR. 2001. Physical interactions between ABA response loci of Arabidopsis. Plant Journal 26: 627-635.
-
Nambara E, Marion-Poll A. 2003. ABA action and interactions in seeds. Trends in Plant Science 8: 213-217.
-
Ni DA, Wang LJ, Ding CH, Xu ZH. 2001. Auxin distribution and transport during embryogenesis and seed germination of Arabidopsis. Cell Research 11: 273-278.
-
Nishimura N, Yoshida T, Kitahata N, Asami T, Shinozaki K, Hirayama T. 2007. ABA-Hypersensitive Germination1 encodes a protein phosphatase 2C, an essential component of abscisic acid signaling in Arabidopsis seed. Plant Journal 50: 935-949.
-
Nishimura N, Yoshida T, Murayama M, Asami T, Shinozaki K, Hirayama T. 2004. Isolation and characterization of novel mutants affecting the abscisic acid sensitivity of Arabidopsis germination and seedling growth. Plant Cell Physiology 45: 1485-1499.
-
Ogas J, Cheng JC, Sung ZR, Somerville C. 1997. Cellular differentiation regulated by gibberellin in the Arabidopsis thaliana pickle mutant. Science 277: 91-94.
-
Ogas J, Kaufmann S, Henderson J, Somerville C. 1999. PICKLE is a CHD3 chromatin-remodeling factor that regulates the transition from embryonic to vegetative development in Arabidopsis. Proceedings of the National Academy of Sciences, USA 96: 13839-13844.
-
Ogawa M, Hanada A, Yamauchi Y, Kuwalhara A, Kamiya Y, Yamaguchi S. 2003. Gibberellin biosynthesis and response during Arabidopsis seed germination. Plant Cell 15: 1591-1604.
-
Oh E, Kim J, Park E, Kim JI, Kang C, Choi G. 2004. PIL5, a phytochrome-interacting basic helix-loop-helix protein, is a key negative regulator of seed germination in Arabidopsis thaliana. Plant Cell 16: 3045-3058.
-
Oh E, Yamaguchi S, Hu JH, Yusuke J, Jung B, Paik I, Lee HS, Sun TP, Kamiya Y, Choi G. 2007. PIL5, a phytochrome-interacting bHLH protein, regulates gibberellin responsiveness by binding directly to the GAI and RGA promoters in Arabidopsis seeds. Plant Cell 19: 1192-1208.
-
Oh E, Yamaguchi S, Kamiya Y, Bae G, Chung WI, Choi G. 2006. Light activates the degradation of PIL5 protein to promote seed germination through gibberellin in Arabidopsis. Plant Journal 47: 124-139.
-
Okamoto M, Kuwahara A, Seo M, Kushiro T, Asami T, Hirai N, Kamiya Y, Koshiba T, Nambara E. 2006. CYP707A1 and CYP707A2, which encode abscisic acid 8 ¢-hydroxylases, are indispensable for proper control of seed dormancy and germination in Arabidopsis. Plant Physiology 141: 97-107.
-
Ooms JJJ, Léon-Kloosterziel KM, Bartels D, Koornneef M, Karssen CM. 1993. Acquisition of desiccation tolerance and longevity in seeds of Arabidopsis thaliana (a comparative-study using abscisic acid-insensitive abi3 mutants). Plant Physiology 102: 1185-1191.
-
Oracz K, Bouteau HEM, Farrant JM, Cooper K, Belghazi M, Job C, Job D, Corbineau F, Bailly C. 2007. ROS production and protein oxidation as a novel mechanism for seed dormancy alleviation. Plant Journal 50: 452-465.
-
Pandey S, Chen JG, Jones AM, Assmann SM. 2006. G-protein complex mutants are hypersensitive to abscisic acid regulation of germination and postgermination development. Plant Physiology 141: 243-256.
-
Parcy F, Giraudat J. 1997. Interactions between the ABI1 and the ectopically expressed ABI3 genes in controlling abscisic acid responses in Arabidopsis vegetative tissues. Plant Journal 11: 693-702.
-
Parcy F, Valon C, Raynal M, Gaubiercomella P, Delseny M, Giraudat J. 1994. Regulation of gene expression programs during Arabidopsis seed development- Roles of the ABI3 locus and of endogenous Abscisic Acid. Plant Cell 6: 1567-1582.
-
Peeters AJM, Blankestijn-de Vries H, Hanhart CJ, Leon-Kloosterziel KM, Zeevaart JAD, Koornneef M. 2002. Characterization of mutants with reduced seed dormancy at two novel rdo loci and a further characterization of rdo1 and rdo2 in Arabidopsis. Physiologia Plantarum 115: 604-612.
-
Penfield S, Gilday AD, Halliday KJ, Graham IA. 2006a. DELLA-mediated cotyledon expansion breaks coat-imposed seed dormancy. Current Biology 16: 2366-2370.
-
Penfield S, Josse EM, Kannangara R, Gilday AD, Halliday KJ, Graham IA. 2005. Cold and light control seed germination through the bHLH transcription factor SPATULA. Current Biology 15: 1998-2006.
-
Penfield S, Li Y, Gilday AD, Graham S, Graham IA. 2006b. Arabidopsis ABA INSENSITIVE4 regulates lipid mobilization in the embryo and reveals repression of seed germination by the endosperm. Plant Cell 18: 1887-1899.
-
Penfield S, Rylott EL, Gilday AD, Graham S, Larson TR, Graham IA. 2004. Reserve mobilization in the Arabidopsis endosperm fuels hypocotyl elongation in the dark, is independent of abscisic acid, and requires PHOSPHOENOLPYRUVATE CARBOXYKINASE1. Plant Cell 16: 2705-2718.
-
Peng JR, Richards DE, Hartley NM, Murphy GP, Devos KM, Flintham JE, Beales J, Fish LJ, Worland AJ, Pelica F et al . 1999. ‘Green revolution’ genes encode mutant gibberellin response modulators. Nature 400: 256-261.
-
Perruc E, Kinoshita N, Lopez-Molina L. 2007. The role of chromatin-remodeling factor PKL in balancing osmotic stress responses during Arabidopsis seed germination. Plant Journal 52: 927-936.
-
Rajjou L, Gallardo K, Debeaujon I, Vandekerckhove J, Job C, Job D. 2004. The effect of alpha-amanitin on the Arabidopsis seed proteome highlights the distinct roles of stored and neosynthesized mRNAs during germination. Plant Physiology 134: 1598-1613.
-
Raz V, Bergervoet JHW, Koornneef M. 2001. Sequential steps for developmental arrest in Arabidopsis seeds. Development 128: 243-252.
-
Razem FA, El-Kereamy A, Abrams SR, Hill RD. 2006. The RNA-binding protein FCA is an abscisic acid receptor. Nature 439: 290-294.
-
Reidt W, Wohlfarth T, Ellerstrom M, Czihal A, Tewes A, Ezcurra I, Rask L, Bäumlein H. 2000. Gene regulation during late embryogenesis: the RY motif of maturation-specific gene promoters is a direct target of the FUS3 gene product. Plant Journal 21: 401-408.
-
Ren CW, Kermode AR. 2000. An increase in pectin methyl esterase activity accompanies dormancy breakage and germination of yellow cedar seeds. Plant Physiology 124: 231-242.
-
Reyes JL, Chua NH. 2007. ABA induction of miR159 controls transcript levels of two MYB factors during Arabidopsis seed germination. Plant Journal 49: 592-606.
-
Rider SD, Henderson JT, Jerome RE, Edenberg HJ, Romero-Severson J, Ogas J. 2003. Coordinate repression of regulators of embryonic identity by PICKLE during germination in Arabidopsis. Plant Journal 35: 33-43.
-
Riefler M, Novak O, Strnad M, Schmulling T. 2006. Arabidopsis cytokinin receptor mutants reveal functions in shoot growth, leaf senescence, seed size, germination, root development, and cytokinin metabolism. Plant Cell 18: 40-54.
-
Rock CD. 2000. Pathways to abscisic acid-regulated gene expression. New Phytologist 148: 357-396.
-
Rundlett SE, Carmen AA, Suka N, Turner BM, Grunstein M. 1998. Transcriptional repression by UME6 involves deacetylation of lysine 5 of histone H4 by RPD3. Nature 392: 831-835.
-
Russell L, Larner V, Kurup S, Bougourd S, Holdsworth MJ. 2000. The Arabidopsis COMATOSE locus regulates germination potential. Development 127: 3759-3767.
-
Saez A, Robert N, Maktabi MH, Schroeder JI, Serrano R, Rodriguez PL. 2006. Enhancement of abscisic acid sensitivity and reduction of water consumption in Arabidopsis by combined inactivation of the protein phosphatases type 2C ABI1 and HAB1. Plant Physiology 141: 1389-1399.
-
Saito S, Hirai N, Matsumoto C, Ohigashi H, Ohta D, Sakata K, Mizutani M. 2004. Arabidopsis CYP707As encode (+)-abscisic acid 8′-hydroxylase, a key enzyme in the oxidative catabolism of abscisic acid. Plant Physiology 134: 1439-1449.
-
Seo M, Hanada A, Kuwahara A, Endo A, Okamoto M, Yamauchi Y, North H, Marion-Poll A, Sun TP, Koshiba T et al . 2006. Regulation of hormone metabolism in Arabidopsis seeds: phytochrome regulation of abscisic acid metabolism and abscisic acid regulation of gibberellin metabolism. Plant Journal 48: 354-366.
-
Shen QX, Zhang PN, Ho THD. 1996. Modular nature of abscisic acid (ABA) response complexes: composite promoter units that are necessary and sufficient for ABA induction of gene expression in barley. Plant Cell 8: 1107-1119.
-
Shen YY, Wang XF, Wu FQ, Du SY, Cao Z, Shang Y, Wang XL, Peng CC, Yu XC, Zhu SY et al . 2006. The Mg-chelatase H subunit is an abscisic acid receptor. Nature 443: 823-826.
-
Shirley BW, Kubasek WL, Storz G, Bruggemann E, Koornneef M, Ausubel FM, Goodman HM. 1995. Analysis of Arabidopsis mutants deficient in flavonoid biosynthesis. Plant Journal 8: 659-671.
-
Singh DP, Jermakow AM, Swain SM. 2002. Gibberellins are required for seed development and pollen tube growth in Arabidopsis. Plant Cell 14: 3133-3147.
-
Sitrit Y, Hadfield KA, Bennett AB, Bradford KJ, Downie AB. 1999. Expression of a polygalacturonase associated with tomato seed germination. Plant Physiology 121: 419-428.
-
Smalle J, Kurepa J, Yang PZ, Emborg TJ, Babiychuk E, Kushnir S, Vierstra RD. 2003. The pleiotropic role of the 26S proteasome subunit RPN10 in Arabidopsis growth and development supports a substrate-specific function in abscisic acid signaling. Plant Cell 15: 965-980.
-
Steadman KJ, Crawford AD, Gallagher RS. 2003. Dormancy release in Lolium rigidum seeds is a function of thermal after-ripening time and seed water content. Functional Plant Biology 30: 345-352.
-
Steber CM, McCourt P. 2001. A role for brassinosteroids in germination in Arabidopsis. Plant Physiology 125: 763-769.
-
Stone SL, Kwong LW, Yee KM, Pelletier J, Lepiniec L, Fischer RL, Goldberg RB, Harada JJ. 2001. LEAFY COTYLEDON2 encodes a B3 domain transcription factor that induces embryo development. Proceedings of the National Academy of Sciences, USA 98: 11806-11811.
-
Sun TP, Gubler F. 2004. Molecular mechanism of gibberellin signaling in plants. Annual Review of Plant Biology 55: 197-223.
-
Suzuki M, Kao CY, McCarty DR. 1997. The conserved B3 domain of VIVIPAROUS1 has a co-operative DNA binding activity. Plant Cell 9: 799-807.
-
Suzuki M, Wang HHY, McCarty DR. 2007. Repression of the LEAFY COTYLEDON 1/B3 regulatory network in plant embryo development by VP1/ABSCISIC ACID INSENSITIVE 3-LIKE B3 genes. Plant Physiology 143: 902-911.
-
Tanaka M, Kikuchi A, Kamada H. 2008. The Arabidopsis histone deacetylases HDA6 and HDA19 contribute to the repression of embryonic properties after germination. Plant Physiology 146: 149-161.
-
Theodoulou FL, Job K, Slocombe SP, Footitt S, Holdsworth M, Baker A, Larson TR, Graham IA. 2005. Jasmonic acid levels are reduced in COMATOSE ATP-binding cassette transporter mutants. Implications for transport of jasmonate precursors into peroxisomes. Plant Physiology 137: 835-840.
-
Tiedemann J, Rutten T, Mönke G, Vorwieger A, Rolletschek H, Meissner D, Milkowski C, Petereck S, Mock HP, Zank T et al . 2008. Dissection of a complex seed phenotype: novel insights of FUSCA3 regulated developmental processes. Developmental Biology in press.
-
To A, Valon C, Savino G, Guilleminot J, Devic M, Giraudat J, Parcy F. 2006. A network of local and redundant gene regulation governs Arabidopsis seed maturation. Plant Cell 18: 1642-1651.
-
Toyomasu T, Kawaide H, Mitsuhashi W, Inoue Y, Kamiya Y. 1998. Phytochrome regulates gibberellin biosynthesis during germination of photoblastic lettuce seeds. Plant Physiology 118: 1517-1523.
-
Tyler L, Thomas SG, Hu JH, Dill A, Alonso JM, Ecker JR, Sun TP. 2004. DELLA proteins and gibberellin-regulated seed germination and floral development in Arabidopsis. Plant Physiology 135: 1008-1019.
-
Ullah H, Chen JG, Wang SC, Jones AM. 2002. Role of a heterotrimeric G protein in regulation of Arabidopsis seed germination. Plant Physiology 129: 897-907.
-
Uno Y, Furihata T, Abe H, Yoshida R, Shinozaki K, Yamaguchi-Shinozaki K. 2000. Arabidopsis basic leucine zipper transcription factors involved in an abscisic acid-dependent signal transduction pathway under drought and high-salinity conditions. Proceedings of the National Academy of Sciences, USA 97: 11632-11637.
-
Vicient CM, Bies-Etheve N, Delseny M. 2000. Changes in gene expression in the leafy cotyledon1 (lec1) and fusca3 (fus3) mutants of Arabidopsis thaliana L. Journal of Experimental Botany 51: 995-1003.
-
Weber H, Borisjuk L, Wobus U. 2005. Molecular physiology of legume seed development. Annual Review of Plant Biology 56: 253-279.
-
Weiss D, Ori N. 2007. Mechanisms of cross talk between gibberellin and other hormones. Plant Physiology 144: 1240-1246.
-
West MAL, Harada JJ. 1993. Embryogenesis in higher plants - an overview. Plant Cell 5: 1361-1369.
-
Wilkinson MD, McKibbin RS, Bailey PC, Flintham JE, Gale MD, Lenton JR, Holdsworth MJ. 2002. Use of comparative molecular genetics to study pre harvest sprouting in wheat. Euphytica 126: 27-33.
-
Willige BC, Ghosh S, Nill C, Zourelidou M, Dohmann EMN, Maier A, Schwechheimer C. 2007. The DELLA domain of GA INSENSITIVE mediates the interaction with the GA INSENSITIVE DWARF1A gibberellin receptor of Arabidopsis. Plant Cell 19: 1209-1220.
-
Wobus U, Weber H. 1999. Sugars as signal molecules in plant seed development. Biological Chemistry 380: 937-944.
-
Xiong LM, Gong ZZ, Rock CD, Subramanian S, Guo Y, Xu WY, Galbraith D, Zhu JK. 2001. Modulation of abscisic acid signal transduction and biosynthesis by an Sm-like protein in Arabidopsis. Developmental Cell 1: 771-781.
-
Yamaguchi S, Smith MW, Brown RGS, Kamiya Y, Sun TP. 1998. Phytochrome regulation and differential expression of gibberellin 3 beta-hydroxylase genes in germinating Arabidopsis seeds. Plant Cell 10: 2115-2126.
-
Yamauchi Y, Ogawa M, Kuwahara A, Hanada A, Kamiya Y, Yamaguchi S. 2004. Activation of gibberellin biosynthesis and response pathways by low temperature during imbibition of Arabidopsis thaliana seeds. Plant Cell 16: 367-378.
-
Yoine M, Ohto MA, Onai K, Mita S, Nakamura K. 2006. The lba1 mutation of UPF1 RNA helicase involved in nonsense-mediated mRNA decay causes pleiotropic phenotypic changes and altered sugar signalling in Arabidopsis. Plant Journal 47: 49-62.
-
Zhang XR, Garreton V, Chua NH. 2005. The AIP2 E3 ligase acts as a novel negative regulator of ABA signaling by promoting ABI3 degradation. Genes and Development 19: 1532-1543.