Abstract
Seed dormancy and germination are two closely linked physiological traits that have great impacts on adaptation and survival of seed plants. Seed dormancy strengthen and germination potential are comprehensively influenced by a variety of internal factors and external environment cues. Environmental factors, such as water content, light condition, ambient temperature, and nitrogen availability, act as signal input to determine whether seeds keep in a dormant state or start to germinate. Light, temperature, and nitrogen availability are the most critical environmental factors that have profound impacts on seed dormancy and germination. However, the mechanisms underlying the regulation of seed dormancy and germination by environmental signals are still poorly understood. In this review, we summarize the current knowledge of signal transduction networks linking environmental stimulus to seed dormancy establishment, dormancy break and germination, underscoring the dominating roles of temperature, light, and nitric oxide. We review temperature, light, and nitric oxide signaling pathway separately as well as the integration of these signaling pathways with phytohormone abscisic acid (ABA) and gibberellins (GA) signaling pathway in the context of seed dormancy and germination.
Similar content being viewed by others
Literature Cited
Abeles FB. 1986. Role of ethylene in Lactuca sativa cv `Grand Rapids' seed germination. Plant Physiol 81:780–787
Albertos P, Romero-Puertas MC, Tatematsu K, Mateos I, Sánchez-Vicente I, Nambara E, Lorenzo O. 2015. S-nitrosylation triggers ABI5 degradation to promote seed germination and seedling growth. NAT COMMUN 6:8669
Alboresi A, Gestin C, Leydecker M-T, Bedu M, Meyer C, Truong H-N. 2005. Nitrate, a signal relieving seed dormancy in Arabidopsis. Plant Cell Environ 28:500–512
Ali-Rachedi S, Bouinot D, Wagner M-H, 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
Alonso-Blanco C, Bentsink L, Hanhart CJ, Vries HB-d, Koornneef M. 2003. Analysis of natural allelic variation at seed dormancy loci of Arabidopsis thaliana. Genetics 164:711–729
Arana MV, SÁNchez-Lamas M, Strasser B, Ibarra SE, CerdÁN PD, Botto JF, SÁNchez RA. 2014. Functional diversity of phytochrome family in the control of light and gibberellin-mediated germination in Arabidopsis. Plant Cell Environ 37:2014–2023
Arana MV, Tognacca RS, Estravis-Barcalá M, Sánchez RA, Botto JF. 2017. Physiological and molecular mechanisms underlying the integration of light and temperature cues in Arabidopsis thaliana seeds. Plant Cell Environ 40:3113–3121
Arc E, Chibani K, Grappin P, Jullien M, Godin B, Cueff G, Valot B, Balliau T, Job D, Rajjou L. 2012. Cold stratification and exogenous nitrates entail similar functional proteome adjustments during Arabidopsis seed dormancy release. Journal of Proteome Research 11:5418–5432
Argyris J, Dahal P, Hayashi E, Still DW, Bradford KJ. 2008. Genetic variation for Lettuce seed thermoinhibition is associated with temperature-sensitive expression of abscisic acid, gibberellin, and ethylene biosynthesis, metabolism, and response genes. Plant Physiol 148:926–947
Ashikawa I, Abe F, Nakamura S. 2010. Ectopic expression of wheat and barley DOG1-like genes promotes seed dormancy in Arabidopsis. Plant Sci 179:536–542
Ashikawa I, Mori M, Nakamura S, Abe F. 2014. A transgenic approach to controlling wheat seed dormancy level by using Triticeae DOG1-like genes. Transgenic Res 23:621–629
Bae G, Choi G. 2008. Decoding of light signals by plant phytochromes and their interacting proteins. Annu Rev Plant Biol 59:281–311
Ballaré CL, Scopel AL, Sánchez RA, Radosevich SR. 1992. Photomorphogenic processes in the agricultural environment. Photochem Photobiol 56:777–788
Basbouss-Serhal I, Leymarie J, Bailly C. 2016. Fluctuation of Arabidopsis seed dormancy with relative humidity and temperature during dry storage. J Exp Bot 67:119–130
Baskin CC, Baskin JM. 1998. Seeds: ecology, biogeography, and, evolution of dormancy and germination. Elsevier
Batlla D, Benech-Arnold RL. 2014. Weed seed germination and the light environment: implications for weed management. Weed Biology and Management 14:77–87
Beligni MV, Lamattina L. 2000. Nitric oxide stimulates seed germination and de-etiolation, and inhibits hypocotyl elongation, three light-inducible responses in plants. Planta 210:215–221
Bentsink L, Hanson J, Hanhart CJ, Blankestijn-de Vries H, Coltrane C, Keizer P, El-Lithy M, Alonso-Blanco C, de Andrés MT, Reymond M. 2010. Natural variation for seed dormancy in Arabidopsis is regulated by additive genetic and molecular pathways. Proc Natl Acad Sci 107:4264–4269
Bentsink L, Jowett J, Hanhart CJ, Koornneef M. 2006. Cloning of DOG1, a quantitative trait locus controlling seed dormancy in Arabidopsis. Proc Natl Acad Sci 103:17042–17047
Bentsink L, Koornneef M. 2008. Seed dormancy and germination. In: The Arabidopsis Book. BioOne, pp 1–18
Bethke PC, Gubler F, Jacobsen JV, Jones RL. 2004. Dormancy of Arabidopsis seeds and barley grains can be broken by nitric oxide. Planta 219:847–855
Bethke PC, Libourel IGL, Jones RL. 2006a. Nitric oxide reduces seed dormancy in Arabidopsis. J Exp Bot 57:517–526
Bethke PC, Libourel IGL, Reinöhl V, Jones RL. 2006b. Sodium nitroprusside, cyanide, nitrite, and nitrate break Arabidopsis seed dormancy in a nitric oxide-dependent manner. Planta 223:805–812
Bewley JD. 1997. Seed germination and dormancy. Plant Cell 9:1055–1066
Bewley JD, Black M. 1994. Seeds: physiology of development and germination. Plenum Press, New York
Bewley JD, Bradford K, Hilhorst H. 2012. Seeds: physiology of development, germination and dormancy. Springer Science & Business Media
Bi C, Ma Y, Wang X-F, Zhang D-P. 2017. Overexpression of the transcription factor NF-YC9 confers abscisic acid hypersensitivity in Arabidopsis. Plant Mol Biol 95:425–439
Boccaccini A, Santopolo S, Capauto D, Lorrai R, Minutello E, Serino G, Costantino P, Vittorioso P. 2014. The DOF protein DAG1 and the DELLA protein GAI cooperate in negatively regulating the AtGA3ox1 gene. Mol Plant 7:1486–1489
Bryant FM, Hughes D, Hassani-Pak K, Eastmond PJ. 2019. Basic LEUCINE ZIPPER TRANSCRIPTION FACTOR 67 transactivates DELAY OF GERMINATION 1 to establish primary seed dormancy in Arabidopsis. Plant Cell tpc.00892.02018
Buijs G, Vogelzang A, Nijveen H, Bentsink L. 2019. Dormancy cycling: translation-related transcripts are the main difference between dormant and non-dormant seeds in the field. The Plant journal : for cell and molecular biology https://doi.org/10.1111/tpj.14626
Burghardt LT, Edwards BR, Donohue K. 2016. Multiple paths to similar germination behavior in Arabidopsis thaliana. New Phytol 209:1301–1312
Canales J, Moyano TC, Gutiérrez RA, Vidal EA. 2014. Nitrogen control of developmental phase transitions in Arabidopsis thaliana. J Exp Bot 65:5611–5618
Cao D, Hussain A, Cheng H, Peng J. 2005. Loss of function of four DELLA genes leads to light- and gibberellin-independent seed germination in Arabidopsis. Planta 223:105–113
Casal JJ, Sánchez RA. 1998. Phytochromes and seed germination. Seed Sci Res 8:317–330
Castillo M-C, Lozano-Juste J, González-Guzmán M, Rodriguez L, Rodriguez PL, León J. 2015. Inactivation of PYR/PYL/RCAR ABA receptors by tyrosine nitration may enable rapid inhibition of ABA signaling by nitric oxide in plants. Science Signaling 8:ra89–ra89
Chahtane H, Kim W, Lopez-Molina L. 2017. Primary seed dormancy: a temporally multilayered riddle waiting to be unlocked. J Exp Bot 68:857–869
Chen Z, Huang Y, Yang W, Chang G, Li P, Wei J, Yuan X, Huang J, Hu X. 2019. The hydrogen sulfide signal enhances seed germination tolerance to high temperatures by retaining nuclear COP1 for HY5 degradation. Plant science : an international journal of experimental plant biology 285:34–43
Chiang GCK, Bartsch M, Barua D, Nakabayashi K, Debieu M, Kronholm I, Koornneef M, Soppe WJJ, Donohue K, De Meaux J. 2011. DOG1 expression is predicted by the seed-maturation environment and contributes to geographical variation in germination in Arabidopsis thaliana. Mol Ecol 20:3336–3349
Chibani K, Ali-Rachedi S, Job C, Job D, Jullien M, Grappin P. 2006. Proteomic analysis of seed dormancy in Arabidopsis. Plant Physiol 142:1493–1510
Chiu RS, Nahal H, Provart NJ, Gazzarrini S. 2012. The role of the Arabidopsis FUSCA3 transcription factor during inhibition of seed germination at high temperature. BMC Plant Biology 12:15
Cho J-N, Ryu J-Y, Jeong Y-M, Park J, Song J-J, Amasino Richard M, Noh B, Noh Y-S. 2012. Control of seed germination by light-induced histone arginine demethylation activity. Developmental Cell 22:736–748
Clack T, Mathews S, Sharrock R. 1994. The phytochrome apoprotein family in Arabidopsis is encoded by five genes: the sequences and expression of PHYD and PHYE. Plant Mol Biol 25:413–427
Contreras S, Bennett M, Tay D. 2009. Temperature during seed development affects weight, germinability and storability of lettuce seeds. Seed Sci Technol 37:398–412
Crawford NM. 1995. Nitrate: nutrient and signal for plant growth. Plant Cell 7:859–868
Cyrek M, Fedak H, Ciesielski A, Guo Y, Sliwa A, Brzezniak L, Krzyczmonik K, Pietras Z, Kaczanowski S, Liu F, Swiezewski S. 2016. Seed dormancy in Arabidopsis is controlled by alternative polyadenylation of DOG1. Plant Physiol 170:947–955
Dechaine JM, Gardner G, Weinig C. 2009. Phytochromes differentially regulate seed germination responses to light quality and temperature cues during seed maturation. Plant Cell Environ 32:1297–1309
Dekkers BJW, He H, Hanson J, Willems LAJ, Jamar DCL, Cueff G, Rajjou L, Hilhorst HWM, Bentsink L. 2016. The Arabidopsis DELAY OF GERMINATION 1 gene affects ABSCISIC ACID INSENSITIVE 5 (ABI5) expression and genetically interacts with ABI3 during Arabidopsis seed development. Plant J 85:451–465
Delledonne M. 2005. NO news is good news for plants. Curr Opin Plant Biol 8:390–396
Di Giammartino Dafne C, Nishida K, Manley James L. 2011. Mechanisms and consequences of alternative polyadenylation. Mol Cell 43:853–866
Dolata J, Guo Y, Kołowerzo A, Smoliński D, Brzyżek G, Jarmołowski A, Świeżewski S. 2015. NTR1 is required for transcription elongation checkpoints at alternative exons in Arabidopsis. Embo J 34:544–558
Donohue K, Dorn L, Griffith C, Kim E, Aguilera A, Polisetty CR, Schmitt J. 2005. Environmental and genetic influences on the germination of Arabidopsis thaliana in the field. Evolution 59:740–757
Donohue K, Heschel MS, Butler CM, Barua D, Sharrock RA, Whitelam GC, Chiang GC. 2008. Diversification of phytochrome contributions to germination as a function of seed-maturation environment. New Phytol 177:367–379
Donohue K, Heschel MS, Chiang GCK, Butler CM, Barua D. 2007. Phytochrome mediates germination responses to multiple seasonal cues. Plant Cell Environ 30:202–212
Fedak H, Palusinska M, Krzyczmonik K, Brzezniak L, Yatusevich R, Pietras Z, Kaczanowski S, Swiezewski S. 2016. Control of seed dormancy in Arabidopsis by a cis-acting noncoding antisense transcript. Proc Natl Acad Sci 113:E7846–E7855
Fenner M. 1991. The effects of the parent environment on seed germinability. Seed Sci Res 1:75–84
Finch-Savage WE, Footitt S. 2017. Seed dormancy cycling and the regulation of dormancy mechanisms to time germination in variable field environments. J Exp Bot 68:843–856
Finch-Savage WE, Leubner-Metzger G. 2006. Seed dormancy and the control of germination. New Phytol 171:501–523
Finkelstein R. 2013. Abscisic acid synthesis and response. The Arabidopsis Book e0166
Finkelstein R, Reeves W, Ariizumi T, Steber C. 2008. Molecular aspects of seed dormancy. Annu Rev Plant Biol 59:387–415
Finkelstein RR. 1994. Mutations at two new Arabidopsis ABA response loci are similar to the abi3 mutations. Plant J 5:765–771
Finkelstein RR. 2010. The role of hormones during seed development and germination. In: Plant Hormones. Springer, pp 549–573
Footitt S, Clewes R, Feeney M, Finch-Savage WE, Frigerio L. 2019. Aquaporins influence seed dormancy and germination in response to stress. Plant Cell Environ 42:2325–2339
Footitt S, Douterelo-Soler I, Clay H, Finch-Savage WE. 2011. Dormancy cycling in Arabidopsis seeds is controlled by seasonally distinct hormone-signaling pathways. Proc Natl Acad Sci 108:20236–20241
Footitt S, Huang Z, Clay HA, Mead A, Finch-Savage WE. 2013. Temperature, light and nitrate sensing coordinate Arabidopsis seed dormancy cycling, resulting in winter and summer annual phenotypes. Plant J 74:1003–1015
Footitt S, Müller K, Kermode AR, Finch-Savage WE. 2015. Seed dormancy cycling in Arabidopsis: chromatin remodelling and regulation of DOG1 in response to seasonal environmental signals. Plant J 81:413–425
Gabriele S, Rizza A, Martone J, Circelli P, Costantino P, Vittorioso P. 2010. The Dof protein DAG1 mediates PIL5 activity on seed germination by negatively regulating GA biosynthetic gene AtGA3ox1. Plant J 61:312–323
Gallardo M, Delgado MdM, Sánchez-Calle IM, Matilla AJ. 1991. Ethylene production and 1-aminocyclopropane-1-carboxylic acid conjugation in thermoinhibited Cicer arietinum L. Seeds. Plant Physiol 97:122–127
Giba Z, Grubišić D, Todorović S, Sajc L, Stojaković Đ, Konjević R. 1998. Effect of nitric oxide – releasing compounds on phytochrome – controlled germination of empress tree seeds. Plant Growth Regul 26:175–181
Gibbs Daniel J, Md Isa N, Movahedi M, Lozano-Juste J, Mendiondo Guillermina M, Berckhan S, Marín-de la Rosa N, Vicente Conde J, Sousa Correia C, Pearce SP, Bassel George W, Hamali B, Talloji P, Tomé Daniel FA, Coego A, Beynon J, Alabadí D, Bachmair A, León J, Gray Julie E, Theodoulou Frederica L, Holdsworth Michael J. 2014. Nitric oxide sensing in plants is mediated by proteolytic control of group VII ERF transcription factors. Mol Cell 53:369–379
Gniazdowska A, Dobrzyńska U, Babańczyk T, Bogatek R. 2007. Breaking the apple embryo dormancy by nitric oxide involves the stimulation of ethylene production. Planta 225:1051–1057
Gniazdowska A, Krasuska U, Bogatek R. 2010. Dormancy removal in apple embryos by nitric oxide or cyanide involves modifications in ethylene biosynthetic pathway. Planta 232:1397–1407
Gonai T, Kawahara S, Tougou M, Satoh S, Hashiba T, Hirai N, Kawaide H, Kamiya Y, Yoshioka T. 2004. Abscisic acid in the thermoinhibition of lettuce seed germination and enhancement of its catabolism by gibberellin. J Exp Bot 55:111–118
Graeber K, Linkies A, Steinbrecher T, Mummenhoff K, Tarkowská D, Turečková V, Ignatz M, Sperber K, Voegele A, de Jong H, Urbanová T, Strnad M, Leubner-Metzger G. 2014. DELAY OF GERMINATION 1 mediates a conserved coat-dormancy mechanism for the temperature- and gibberellin-dependent control of seed germination. Proc Natl Acad Sci 111:E3571–E3580
Graeber K, Nakabayashi K, Miatton E, LEUBNER-METZGER G, Soppe WJ. 2012. Molecular mechanisms of seed dormancy. Plant Cell Environ 35:1769–1786
Graeber K, Voegele A, Büttner-Mainik A, Sperber K, Mummenhoff K, Leubner-Metzger G. 2013. Spatiotemporal seed development analysis provides insight into primary dormancy induction and evolution of the Lepidium DELAY OF GERMINATION1 genes. Plant Physiol 161:1903–1917
Groot SPC, Karssen CM. 1987. Gibberellins regulate seed germination in tomato by endosperm weakening: a study with gibberellin-deficient mutants. Planta 171:525–531
Gu D, Chen C-Y, Zhao M, Zhao L, Duan X, Duan J, Wu K, Liu X. 2017. Identification of HDA15-PIF1 as a key repression module directing the transcriptional network of seed germination in the dark. Nucleic Acids Res 45:7137–7150
Gu D, Ji R, He C, Peng T, Zhang M, Duan J, Xiong C, Liu X. 2019. Arabidopsis Histone Methyltransferase SUVH5 Is a Positive Regulator of Light-Mediated Seed Germination. Front Plant Sci 10:
Gualberti G, Papi M, Bellucci L, Ricci I, Bouchez D, Camilleri C, Costantino P, Vittorioso P. 2002. Mutations in the Dof Zinc Finger Genes DAG2 and DAG1 influence with opposite effects the germination of Arabidopsis seeds. Plant Cell 14:1253–1263
Gubler F, Millar AA, Jacobsen JV. 2005. Dormancy release, ABA and pre-harvest sprouting. Curr Opin Plant Biol 8:183–187
He H, de Souza Vidigal D, Snoek LB, Schnabel S, Nijveen H, Hilhorst H, Bentsink L. 2014. Interaction between parental environment and genotype affects plant and seed performance in Arabidopsis. J Exp Bot 65:6603–6615
Hennig L, Stoddart WM, Dieterle M, Whitelam GC, Schäfer E. 2002. Phytochrome E controls light-induced germination of Arabidopsis. Plant Physiol 128:194–200
Heschel MS, Butler CM, Chiang GCK, Wheeler A, Sharrock RA, Whitelam GC, Donohue K. 2008. New roles of phytochromes during seed germination. Int J Plant Sci 169:531–540
Heschel MS, Selby J, Butler C, Whitelam GC, Sharrock RA, Donohue K. 2007. A new role for phytochromes in temperature-dependent germination. New Phytol 174:735–741
Holdsworth MJ, Bentsink L, Soppe WJJ. 2008. Molecular networks regulating Arabidopsis seed maturation, after-ripening, dormancy and germination. New Phytol 179:33–54
Holman TJ, Jones PD, Russell L, Medhurst A, Úbeda Tomás S, Talloji P, Marquez J, Schmuths H, Tung S-A, Taylor I, Footitt S, Bachmair A, Theodoulou FL, Holdsworth MJ. 2009. The N-end rule pathway promotes seed germination and establishment through removal of ABA sensitivity in Arabidopsis. Proc Natl Acad Sci 106:4549–4554
Huo H, Wei S, Bradford KJ. 2016. DELAY OF GERMINATION1 (DOG1) regulates both seed dormancy and flowering time through microRNA pathways. Proc Natl Acad Sci 113:E2199–E2206
Iuchi S, Suzuki H, Kim Y-C, Iuchi A, Kuromori T, Ueguchi-Tanaka M, Asami T, Yamaguchi I, Matsuoka M, Kobayashi M, Nakajima M. 2007. Multiple loss-of-function of Arabidopsis gibberellin receptor AtGID1s completely shuts down a gibberellin signal. Plant J 50:958–966
Jiang Z, Xu G, Jing Y, Tang W, Lin R. 2016. Phytochrome B and REVEILLE1/2-mediated signalling controls seed dormancy and germination in Arabidopsis. NAT COMMUN 7:12377
Jung J-H, Domijan M, Klose C, Biswas S, Ezer D, Gao M, Khattak AK, Box MS, Charoensawan V, Cortijo S, Kumar M, Grant A, Locke JCW, Schäfer E, Jaeger KE, Wigge PA. 2016. Phytochromes function as thermosensors in Arabidopsis. Science 354:886–889
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
Kendall SL, Hellwege A, Marriot P, Whalley C, Graham IA, Penfield S. 2011. Induction of dormancy in Arabidopsis summer annuals requires parallel regulation of DOG1 and hormone metabolism by low temperature and CBF transcription factors. Plant Cell 23:2568–2580
Khanna R, Huq E, Kikis EA, Al-Sady B, Lanzatella C, Quail PH. 2004. A novel molecular recognition motif necessary for targeting photoactivated phytochrome signaling to specific basic helix-loop-helix transcription factors. Plant Cell 16:3033–3044
Kim DH, Yamaguchi S, Lim S, Oh E, Park J, Hanada A, Kamiya Y, Choi G. 2008. SOMNUS, a CCCH-type zinc finger protein in Arabidopsis, negatively regulates light-dependent seed germination downstream of PIL5. Plant Cell 20:1260–1277
Kim J, Kang H, Park J, Kim W, Yoo J, Lee N, Kim J, Yoon T-y, Choi G. 2016. PIF1-interacting transcription factors and their binding sequence elements determine the in vivo targeting sites of PIF1. Plant Cell 28:1388–1405
Kim SY, Warpeha KM, Huber SC. 2019. The brassinosteroid receptor kinase, BRI1, plays a role in seed germination and the release of dormancy by cold stratification. J Plant Physiol 241:153031–153031
Kim W, Lee Y, Park J, Lee N, Choi G. 2013. HONSU, a protein phosphatase 2C, regulates seed dormancy by inhibiting ABA signaling in Arabidopsis. Plant Cell Physiol 54:555–572
Kinoshita N, Berr A, Belin C, Chappuis R, Nishizawa NK, Lopez-Molina L. 2010. Identification of growth insensitive to ABA3 (gia3), a recessive mutation affecting ABA signaling for the control of early post-germination growth in Arabidopsis thaliana. Plant Cell Physiol 51:239–251
Koornneef M, Bentsink L, Hilhorst H. 2002. Seed dormancy and germination. Curr Opin Plant Biol 5:33–36
Koornneef M, Reuling G, Karssen CM. 1984. The isolation and characterization of abscisic acid-insensitive mutants of Arabidopsis thaliana. Physiol Plantarum 61:377–383
Kowalczyk J, Palusinska M, Wroblewska-Swiniarska A, Pietras Z, Szewc L, Dolata J, Jarmolowski A, Swiezewski S. 2017. Alternative polyadenylation of the sense transcript controls antisense transcription of DELAY OF GERMINATION 1 in Arabidopsis. Mol Plant 10:1349–1352
Kozarewa I, Cantliffe DJ, Nagata RT, Stoffella PJ. 2006. High maturation temperature of Lettuce seeds during development increased ethylene production and germination at elevated temperatures. J Am Soc Hortic Sci 131:564–570
Kushiro T, Okamoto M, Nakabayashi K, Yamagishi K, Kitamura S, Asami T, Hirai N, Koshiba T, Kamiya Y, Nambara E. 2004. The Arabidopsis cytochrome P450 CYP707A encodes ABA 8′-hydroxylases: key enzymes in ABA catabolism. Embo J 23:1647–1656
Lee KP, Piskurewicz U, Turečková V, Carat S, Chappuis R, Strnad M, Fankhauser C, Lopez-Molina L. 2012. Spatially and genetically distinct control of seed germination by phytochromes A and B. Gene Dev 26:1984–1996
Lee KP, Piskurewicz U, Turečková V, Strnad M, Lopez-Molina L. 2010. A seed coat bedding assay shows that RGL2-dependent release of abscisic acid by the endosperm controls embryo growth in Arabidopsis dormant seeds. Proc Natl Acad Sci 107:19108–19113
Lee N, Park J, Kim K, Choi G. 2015. The transcriptional coregulator LEUNIG_HOMOLOG inhibits light-dependent seed germination in Arabidopsis. Plant Cell 27:2301–2313
Lee S, Cheng H, King KE, Wang W, He Y, Hussain A, Lo J, Harberd NP, Peng J. 2002. Gibberellin regulates Arabidopsis seed germination via RGL2, a GAI/RGA-like gene whose expression is up-regulated following imbibition. Gene Dev 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 J 45:309–319
Legris M, Klose C, Burgie ES, Rojas CCR, Neme M, Hiltbrunner A, Wigge PA, Schäfer E, Vierstra RD, Casal JJ. 2016. Phytochrome B integrates light and temperature signals in Arabidopsis. Science 354:897–900
Leivar P, Quail PH. 2011. PIFs: pivotal components in a cellular signaling hub. Trends Plant Sci 16:19–28
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 J 10:655–661
Leubner-Metzger G, Fründt C, Meins F. 1996. Effects of gibberellins, darkness and osmotica on endosperm rupture and class I β-1,3-glucanase induction in tobacco seed germination. Planta 199:282-288
Li X, Chen T, Li Y, Wang Z, Cao H, Chen F, Li Y, Soppe W, Li W, Liu Y. 2019. ETR1/RDO3 regulates seed dormancy by relieving the inhibitory effect of the ERF12-TPL complex on DELAY OF GERMINATION1 expression. Plant Cell tpc.00449.02018
Lim S, Park J, Lee N, Jeong J, Toh S, Watanabe A, Kim J, Kang H, Kim DH, Kawakami N, Choi G. 2013. ABA-INSENSITIVE3, ABA-INSENSITIVE5, and DELLAs interact to activate the expression of SOMNUS and other high-temperature-inducible genes in imbibed seeds in Arabidopsis. Plant Cell 25:4863–4878
Lin P-C, Hwang S-G, Endo A, Okamoto M, Koshiba T, Cheng W-H. 2007. Ectopic expression of ABSCISIC ACID 2/GLUCOSE INSENSITIVE 1 in Arabidopsis promotes seed dormancy and stress tolerance. Plant Physiol 143:745–758
Linkies A, Müller K, Morris K, Turečková V, Wenk M, Cadman CSC, Corbineau F, Strnad M, Lynn JR, Finch-Savage WE, Leubner-Metzger G. 2009. Ethylene interacts with abscisic acid to regulate endosperm rupture during germination: A comparative approach using Lepidium sativum and Arabidopsis thaliana. Plant Cell 21:3803–3822
Liu J, Hasanuzzaman M, Wen H, Zhang J, Peng T, Sun H, Zhao Q. 2019. High temperature and drought stress cause abscisic acid and reactive oxygen species accumulation and suppress seed germination growth in rice. Protoplasma 256:1217–1227
Liu S-J, Xu H-H, Wang W-Q, Li N, Wang W-P, Møller IM, Song S-Q. 2015. A proteomic analysis of rice seed germination as affected by high temperature and ABA treatment. Physiol Plantarum 154:142–161
Liu X, Hu P, Huang M, Tang Y, Li Y, Li L, Hou X. 2016. The NF-YC–RGL2 module integrates GA and ABA signalling to regulate seed germination in Arabidopsis. Nature Communications 7:12768
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
Liu Y, Shi L, Ye N, Liu R, Jia W, Zhang J. 2009. Nitric oxide-induced rapid decrease of abscisic acid concentration is required in breaking seed dormancy in Arabidopsis. New Phytol 183:1030–1042
Lozano-Juste J, Colom-Moreno R, León J. 2011. In vivo protein tyrosine nitration in Arabidopsis thaliana. J Exp Bot 62:3501–3517
Ma Z, Marsolais F, Bykova NV, Igamberdiev AU. 2016. Nitric oxide and reactive oxygen species mediate metabolic changes in barley seed embryo during germination. Front Plant Sci 7:138–138
Majee M, Kumar S, Kathare PK, Wu S, Gingerich D, Nayak NR, Salaita L, Dinkins R, Martin K, Goodin M, Dirk LMA, Lloyd TD, Zhu L, Chappell J, Hunt AG, Vierstra R, Huq E, Downie AB. 2018. KELCH F-BOX protein positively influences Arabidopsis seed germination by targeting PHYTOCHROME-INTERACTING FACTOR1. Proc Natl Acad Sci 115:E4120–E4129
Martel C, Blair LK, Donohue K. 2018. PHYD prevents secondary dormancy establishment of seeds exposed to high temperature and is associated with lower PIL5 accumulation. J Exp Bot 69:3157–3169
Matakiadis T, Alboresi A, Jikumaru Y, Tatematsu K, Pichon O, Renou J-P, Kamiya Y, Nambara E, Truong H-N. 2009. The Arabidopsis abscisic acid catabolic Gene CYP707A2 plays a key role in nitrate control of seed dormancy. Plant Physiol 149:949–960
Mathews S, Sharrock RA. 1997. Phytochrome gene diversity. Plant Cell Environ 20:666–671
Matilla AJ. 2000. Ethylene in seed formation and germination. Seed Sci Res 10:111–126
Mendel RR. 2007. Biology of the molybdenum cofactor. J Exp Bot 58:2289–2296
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 J 45:804–818
Molitor AM, Bu Z, Yu Y, Shen W-H. 2014. Arabidopsis AL PHD-PRC1 complexes promote seed germination through H3K4me3-to-H3K27me3 chromatin state switch in repression of seed developmental genes. PLoS Genet 10:e1004091
Monteiro HP, Arai RJ, Travassos LR. 2008. Protein tyrosine phosphorylation and protein tyrosine nitration in redox signaling. Antioxidants & Redox Signaling 10:843–890
Moreau M, Lindermayr C, Durner J, Klessig DF. 2010. NO synthesis and signaling in plants – where do we stand? Physiol Plantarum 138:372–383
Mortensen SA, Grasser KD. 2014. The seed dormancy defect of Arabidopsis mutants lacking the transcript elongation factor TFIIS is caused by reduced expression of the DOG1 gene. Febs Lett 588:47–51
Mortensen SA, Sønderkær M, Lynggaard C, Grasser M, Nielsen KL, Grasser KD. 2011. Reduced expression of the DOG1 gene in Arabidopsis mutant seeds lacking the transcript elongation factor TFIIS. Febs Lett 585:1929–1933
Müller K, Bouyer D, Schnittger A, Kermode AR. 2012. Evolutionarily conserved histone methylation dynamics during seed life-cycle transitions. PLoS ONE 7:e51532
Nakabayashi K, Bartsch M, Ding J, Soppe WJJ. 2015. Seed dormancy in Arabidopsis requires self-binding ability of DOG1 protein and the presence of multiple isoforms generated by alternative splicing. PLoS Genet 11:e1005737
Nakabayashi K, Bartsch M, Xiang Y, Miatton E, Pellengahr S, Yano R, Seo M, Soppe WJJ. 2012. The time required for dormancy release in Arabidopsis is determined by DELAY OF GERMINATION1 protein levels in freshly harvested seeds. Plant Cell 24:2826–2838
Nakamura S, Abe F, Kawahigashi H, Nakazono K, Tagiri A, Matsumoto T, Utsugi S, Ogawa T, Handa H, Ishida H. 2011. A wheat homolog of MOTHER OF FT AND TFL1 acts in the regulation of germination. Plant Cell 23:3215–3229
Nakashima K, Fujita Y, Kanamori N, Katagiri T, Umezawa T, Kidokoro S, Maruyama K, Yoshida T, Ishiyama K, Kobayashi M, Shinozaki K, Yamaguchi-Shinozaki K. 2009. Three Arabidopsis SnRK2 protein kinases, SRK2D/SnRK2.2, SRK2E/SnRK2.6/OST1 and SRK2I/SnRK2.3, involved in ABA signaling are essential for the control of seed development and dormancy. Plant Cell Physiol 50:1345–1363
Nambara E, Marion-Poll A. 2005. Abscisic acid biosynthesis and catabolism. Annu Rev Plant Biol 56:165–185
Née G, Kramer K, Nakabayashi K, Yuan B, Xiang Y, Miatton E, Finkemeier I, Soppe WJJ. 2017. DELAY OF GERMINATION1 requires PP2C phosphatases of the ABA signalling pathway to control seed dormancy. NAT COMMUN 8:72
Nishimura N, Tsuchiya W, Moresco JJ, Hayashi Y, Satoh K, Kaiwa N, Irisa T, Kinoshita T, Schroeder JI, Yates JR, Hirayama T, Yamazaki T. 2018. Control of seed dormancy and germination by DOG1-AHG1 PP2C phosphatase complex via binding to heme. NAT COMMUN 9:2132
Nonogaki H, Bassel GW, Bewley JD. 2010. Germination-still a mystery. Plant Sci 179:574–581
Ogawa M, Hanada A, Yamauchi Y, Kuwahara A, Kamiya Y, Yamaguchi S. 2003. Gibberellin biosynthesis and response during Arabidopsis seed germination. Plant Cell 15:1591–1604
Oh E, Kang H, Yamaguchi S, Park J, Lee D, Kamiya Y, Choi G. 2009. Genome-wide analysis of genes targeted by PHYTOCHROME INTERACTING FACTOR 3-LIKE5 during seed germination in Arabidopsis. Plant Cell 21:403–419
Oh E, Kim J, Park E, Kim J-I, 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 J, Yusuke J, Jung B, Paik I, Lee HS, Sun T, 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 W-I, Choi G. 2006. Light activates the degradation of PIL5 protein to promote seed germination through gibberellin in Arabidopsis. Plant J 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 Physiol 141:97–107
Osuna D, Prieto P, Aguilar M. 2015. Control of seed germination and plant development by carbon and nitrogen availability. Front Plant Sci 6:1023
Paik I, Chen F, Ngoc Pham V, Zhu L, Kim J-I, Huq E. 2019. A phyB-PIF1-SPA1 kinase regulatory complex promotes photomorphogenesis in Arabidopsis. NAT COMMUN 10:4216–4216
Papi M, Sabatini S, Altamura MM, Hennig L, Schäfer E, Costantino P, Vittorioso P. 2002. Inactivation of the phloem-specific Dof Zinc Finger Gene DAG1 affects response to light and integrity of the testa of Arabidopsis seeds. Plant Physiol 128:411–417
Park J, Lee N, Kim W, Lim S, Choi G. 2011. ABI3 and PIL5 collaboratively activate the expression of SOMNUS by directly binding to its promoter in imbibed Arabidopsis seeds. Plant Cell 23:1404–1415
Pawłowski TA. 2007. Proteomics of European beech (Fagus sylvatica L.) seed dormancy breaking: Influence of abscisic and gibberellic acids. Proteomics 7:2246–2257
Piskurewicz U, Jikumaru Y, Kinoshita N, Nambara E, Kamiya Y, Lopez-Molina L. 2008. The gibberellic acid signaling repressor RGL2 inhibits Arabidopsis seed germination by stimulating abscisic acid synthesis and ABI5 activity. Plant Cell 20:2729–2745
Piskurewicz U, Turečková V, Lacombe E, Lopez-Molina L. 2009. Far-red light inhibits germination through DELLA-dependent stimulation of ABA synthesis and ABI3 activity. Embo J 28:2259–2271
Piterková J, Luhová L, Hofman J, Petřivalský M, Novák O, Turečková V, Fellner M. 2012. Nitric oxide is involved in light-specific responses of tomato during germination under normal and osmotic stress conditions. Ann Bot-london 110:767–776
Preston J, Tatematsu K, Kanno Y, Hobo T, Kimura M, Jikumaru Y, Yano R, Kamiya Y, Nambara E. 2009. Temporal expression patterns of hormone metabolism genes during imbibition of Arabidopsis thaliana seeds: a comparative study on dormant and non-dormant accessions. Plant Cell Physiol 50:1786–1800
Quail PH. 2002. Phytochrome photosensory signalling networks. Nat Rev Mol Cell Bio 3:85
Raz V, Bergervoet J, Koornneef M. 2001. Sequential steps for developmental arrest in Arabidopsis seeds. Development 128:243–252
Reed JW, Nagatani A, Elich TD, Fagan M, Chory J. 1994. Phytochrome A and phytochrome B have overlapping but distinct functions in Arabidopsis development. Plant Physiol 104:1139–1149
Reynolds T, Thompson PA. 1971. Characterisation of the high temperature inhibition of germination of Lettuce (Lactuca sativa). Physiol Plantarum 24:544–547
Santopolo S, Boccaccini A, Lorrai R, Ruta V, Capauto D, Minutello E, Serino G, Costantino P, Vittorioso P. 2015. DOF AFFECTING GERMINATION 2 is a positive regulator of light-mediated seed germination and is repressed by DOF AFFECTING GERMINATION 1. BMC Plant Biology 15:72
Sarath G, Bethke PC, Jones R, Baird LM, Hou G, Mitchell RB. 2006. Nitric oxide accelerates seed germination in warm-season grasses. Planta 223(6):1154–1164
Sawada Y, Aoki M, Nakaminami K, Mitsuhashi W, Tatematsu K, Kushiro T, Koshiba T, Kamiya Y, Inoue Y, Nambara E, Toyomasu T. 2008. Phytochrome- and gibberellin-mediated regulation of abscisic acid metabolism during germination of photoblastic Lettuce seeds. Plant Physiol 146:1386–1396
Sen S. 2010. S-Nitrosylation process acts as a regulatory switch for seed germination in wheat. American Journal of Plant Physiology 5:122–132
Seo M, Hanada A, Kuwahara A, Endo A, Okamoto M, Yamauchi Y, North H, Marion-Poll A, Sun T-p, Koshiba T, Kamiya Y, Yamaguchi S, Nambara E. 2006. Regulation of hormone metabolism in Arabidopsis seeds: phytochrome regulation of abscisic acid metabolism and abscisic acid regulation of gibberellin metabolism. Plant J 48:354–366
Seo M, Nambara E, Choi G, Yamaguchi S. 2009. Interaction of light and hormone signals in germinating seeds. Plant Mol Biol 69:463–472
Shen H, Moon J, Huq E. 2005. PIF1 is regulated by light-mediated degradation through the ubiquitin-26S proteasome pathway to optimize photomorphogenesis of seedlings in Arabidopsis. Plant J 44:1023–1035
Shi H, Zhong S, Mo X, Liu N, Nezames CD, Deng XW. 2013. HFR1 sequesters PIF1 to govern the transcriptional network underlying light-initiated seed germination in Arabidopsis. Plant Cell 25:3770–3784
Shinomura T, Nagatani A, Chory J, Furuya M. 1994. The induction of seed germination in Arabidopsis thaliana is regulated principally by phytochrome B and secondarily by phytochrome A. Plant Physiol 104:363–371
Shinomura T, Nagatani A, Hanzawa H, Kubota M, Watanabe M, Furuya M. 1996. Action spectra for phytochrome A- and B-specific photoinduction of seed germination in Arabidopsis thaliana. Proc Natl Acad Sci 93:8129–8133
Shu K, Chen Q, Wu Y, Liu R, Zhang H, Wang P, Li Y, Wang S, Tang S, Liu C, Yang W, Cao X, Serino G, Xie Q. 2016a. ABI4 mediates antagonistic effects of abscisic acid and gibberellins at transcript and protein levels. Plant J 85:348–361
Shu K, Liu X-d, Xie Q, He Z-h. 2016b. Two faces of one seed: hormonal regulation of dormancy and germination. Mol Plant 9:34–45
Shu K, Zhang H, Wang S, Chen M, Wu Y, Tang S, Liu C, Feng Y, Cao X, Xie Q. 2013. ABI4 regulates primary seed dormancy by regulating the biogenesis of abscisic acid and gibberellins in Arabidopsis. PLoS Genet 9:1
Song Q, Cheng S, Chen Z, Nie G, Xu F, Zhang J, Zhou M, Zhang W, Liao Y, Ye J. 2019. Comparative transcriptome analysis revealing the potential mechanism of seed germination stimulated by exogenous gibberellin in Fraxinus hupehensis. BMC plant biology 19:199–199
Tamura N, Yoshida T, Tanaka A, Sasaki R, Bando A, Toh S, Lepiniec L, Kawakami N. 2006. Isolation and characterization of high temperature-resistant germination mutants of Arabidopsis thaliana. Plant Cell Physiol 47:1081–1094
Tang W, Ji Q, Huang Y, Jiang Z, Bao M, Wang H, Lin R. 2013. FAR-RED ELONGATED HYPOCOTYL3 and FAR-RED IMPAIRED RESPONSE1 transcription factors integrate light and abscisic acid signaling in Arabidopsis. Plant Physiol 163:857–866
Toh S, Imamura A, Watanabe A, Nakabayashi K, Okamoto M, Jikumaru Y, Hanada A, Aso Y, Ishiyama K, Tamura N, Iuchi S, Kobayashi M, Yamaguchi S, Kamiya Y, Nambara E, Kawakami N. 2008. High temperature-induced abscisic acid biosynthesis and its role in the inhibition of gibberellin action in Arabidopsis seeds. Plant Physiol 146:1368–1385
Toh S, Kamiya Y, Kawakami N, Nambara E, McCourt P, Tsuchiya Y. 2012. Thermoinhibition uncovers a role for strigolactones in Arabidopsis seed germination. Plant Cell Physiol 53:107–117
Toledo-Ortiz G, Huq E, Quail PH. 2003. The Arabidopsis basic/helix-loop-helix transcription factor family. Plant Cell 15:1749–1770
Topham AT, Taylor RE, Yan D, Nambara E, Johnston IG, Bassel GW. 2017. Temperature variability is integrated by a spatially embedded decision-making center to break dormancy in <em>Arabidopsis</em> seeds. Proc Natl Acad Sci 114:6629–6634
Toyomasu T, Yamane H, Murofushi N, Inoue Y. 1994. Effects of exogenously applied gibberellin and red light on the endogenous levels of abscisic acid in photoblastic Lettuce seeds. Plant Cell Physiol 35:127–129
Vaistij FE, Barros-Galvão T, Cole AF, Gilday AD, He Z, Li Y, Harvey D, Larson TR, Graham IA. 2018. MOTHER-OF-FT-AND-TFL1 represses seed germination under far-red light by modulating phytohormone responses in Arabidopsis thaliana. Proc Natl Acad Sci 115:8442–8447
Vaistij FE, Gan Y, Penfield S, Gilday AD, Dave A, He Z, Josse E-M, Choi G, Halliday KJ, Graham IA. 2013. Differential control of seed primary dormancy in Arabidopsis ecotypes by the transcription factor SPATULA. Proc Natl Acad Sci 110:10866–10871
Wang P, Du Y, Hou Y-J, Zhao Y, Hsu C-C, Yuan F, Zhu X, Tao WA, Song C-P, Zhu J-K. 2015a. Nitric oxide negatively regulates abscisic acid signaling in guard cells by S-nitrosylation of OST1. Proc Natl Acad Sci 112:613–618
Wang P, Zhu J-K, Lang Z. 2015b. Nitric oxide suppresses the inhibitory effect of abscisic acid on seed germination by S-nitrosylation of SnRK2 proteins. Plant Signal Behav 10:e1031939
Wang W-Q, Song B-Y, Deng Z-J, Wang Y, Liu S-J, Møller IM, Song S-Q. 2015. Proteomic analysis of Lettuce seed germination and thermoinhibition by sampling of individual seeds at germination and removal of storage proteins by polyethylene glycol fractionation. Plant Physiol 167:1332–1350
Weitbrecht K, Müller K, Leubner-Metzger G. 2011. First off the mark: early seed germination. J Exp Bot 62:3289–3309
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
Xi W, Liu C, Hou X, Yu H. 2010. MOTHER OF FT AND TFL1 regulates seed germination through a negative feedback loop modulating ABA signaling in Arabidopsis. Plant Cell 22:1733–1748
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
Yamauchi Y, Takeda-Kamiya N, Hanada A, Ogawa M, Kuwahara A, Seo M, Kamiya Y, Yamaguchi S. 2007. Contribution of gibberellin deactivation by AtGA2ox2 to the suppression of germination of dark-imbibed Arabidopsis thaliana seeds. Plant Cell Physiol 48:555–561
Yan A, Chen Z. 2017. The pivotal role of abscisic acid signaling during transition from seed maturation to germination. Plant Cell Rep 36:689–703
Yan A, Wu M, Yan L, Hu R, Ali I, Gan Y. 2014. AtEXP2 is involved in seed germination and abiotic stress tesponse in Arabidopsis. PLoS ONE 9:e85208
Yan D, Easwaran V, Chau V, Okamoto M, Ierullo M, Kimura M, Endo A, Yano R, Pasha A, Gong Y, Bi Y-M, Provart N, Guttman D, Krapp A, Rothstein SJ, Nambara E. 2016. NIN-like protein 8 is a master regulator of nitrate-promoted seed germination in Arabidopsis. NAT COMMUN 7:13179–13179
Yang W, Chen Z, Huang Y, Chang G, Li P, Wei J, Yuan X, Huang J, Hu X. 2019. Powerdress as the novel regulator enhances Arabidopsis seeds germination tolerance to high temperature stress by histone modification of SOM locus. Plant science : an international journal of experimental plant biology 284:91–98
Yatusevich R, Fedak H, Ciesielski A, Krzyczmonik K, Kulik A, Dobrowolska G, Swiezewski S. 2017. Antisense transcription represses Arabidopsis seed dormancy QTL DOG1 to regulate drought tolerance. Embo Rep 18:2186–2196
Zentella R, Zhang ZL, Park M, Thomas SG, Endo A, Murase K, Fleet CM, Jikumaru Y, Nambara E, Kamiya Y. 2007. Global analysis of DELLA direct targets in early gibberellin signaling in Arabidopsis. Plant Cell 19:3037–3057
Zhao Ml. 2015. Arabidopsis histone demethylases LDL1 and LDL2 control primary seed dormancy by regulating DELAY OF GERMINATION 1 and ABA signaling-related genes. Front Plant Sci 6:159
Zheng J, Chen F, Wang Z, Cao H, Li X, Deng X, Soppe WJJ, Li Y, Liu Y. 2012. A novel role for histone methyltransferase KYP/SUVH4 in the control of Arabidopsis primary seed dormancy. New Phytol 193:605–616
Zhu L, Bu Q, Xu X, Paik I, Huang X, Hoecker U, Deng XW, Huq E. 2015. CUL4 forms an E3 ligase with COP1 and SPA to promote light-induced degradation of PIF1. NAT COMMUN 6:7245
Funding
Authors thank the research grant (NTU-MSE-Facile Non-T) to Z.C. and NIE AcRF funding (RI 8/16 CZ) to support A.Y.
Author information
Authors and Affiliations
Contributions
Z.C. conceived and initiated the work. A.Y. and Z.C. wrote and revised the manuscript.
Corresponding author
Ethics declarations
Conflict of interest
The authors declare no conflict of interest.
Rights and permissions
About this article
Cite this article
Yan, A., Chen, Z. The Control of Seed Dormancy and Germination by Temperature, Light and Nitrate. Bot. Rev. 86, 39–75 (2020). https://doi.org/10.1007/s12229-020-09220-4
Published:
Issue Date:
DOI: https://doi.org/10.1007/s12229-020-09220-4