ISSN 1004-5759 CN 62-1105/S
Acta Prataculturae Sinica ›› 2014, Vol. 23 ›› Issue (1): 328-338.DOI: 10.11686/cyxb20140139
• Orignal Article • Previous Articles Next Articles
ZHANG Jia-ning1,2,LIU Kun2
Received:2013-07-28
Online:2014-02-20
Published:2014-02-20
CLC Number:
ZHANG Jia-ning,LIU Kun. Mechanisms for plants detecting the optimum time and place to germinate[J]. Acta Prataculturae Sinica, 2014, 23(1): 328-338.
| Reference:[1]Zong W J, Liu K, Bu H Y, et al. The mode of seed size variation and the effects of seed size on fifty-one species of composite plants in a alpine meadow[J]. Journal of Lanzhou University (Natural Sciences), 2006, 42: 52-55.[2]Zhang D Y. Plant Life History Evolution and Breeding Ecology[M]. Beijing: Science Press, 2004.[3]Cheplick G P. Do seed germination patterns in cleistogamous annual grasses reduce the risk of sibling competition[J]. Journal of Ecology, 1996, 84: 247-255.[4]Lu W H, Wan J J, Yang J J, et al. Review of endozoochory of plant seeds by herbivores[J]. Acta Prataculturae Sinica, 2013, 22(3): 306-313.[5]Lu J H, Lü X, Wu L, et al. Germination responses of three medicinal licorices to saline environments and their suitable ecological regions[J]. Acta Prataculturae Sinica, 2013, 22(2): 195-202.[6]Lu Y M, Su C Q, Li H F. Effects of different salts stress on seed germination and seedling growth of Trifolium repens[J]. Acta Prataculturae Sinica, 2013, 22(4): 123-129.[7]Baskin J M, Baskin C C. A classification system for seed dormancy[J]. Seed Science Research, 2004, 14: 1-16.[8]Vleeshouwers L M, Bouwmeester H J, Karssen C M. Redefining seed dormancy: an attempt to integrate physiology and ecology[J]. Journal of Ecology, 1995, 83: 1031-1037.[9]Baskin C C, Baskin J M. Seeds: Ecology, Biogeography, and Evolution of Dormancy and Germination[M]. San Diego: Academic Press, 1998.[10]Liu K, Baskin J M, Baskin C C, et al. Effect of storage conditions on germination of 489 species from high elevation grassland of the eastern Tibet Plateau and implications for climate change[J]. American Journal of Botany, 2011, 98: 12-19.[11]Washitani I, Masuda M. A comparative study of the germination characteristics of seeds from a moist tall grassland community[J]. Functional Ecology, 1990, 4: 543-457.[12]Grime J P, Mason G, Curtis A V, et al. A comparative study of germination characteristics in a local flora[J]. Journal of Ecology, 1981, 69: 1017-1059.[13]Schütz W. Are germination strategies important for the ability of cespitose wetland sedges (Carex) to grow in forests[J]. Canadian Journal of Botany, 1997, 75: 1692-1699.[14]Fenner M, Thompson K. The Ecology of Seeds[M]. Cambridge: Cambridge University Press, 2005.[15]Simon E W, Minchin A, McMenamin M M, et al. The low temperature limit for seed germination[J]. New Phytologist, 1976, 77: 301-311.[16]Crawford R M M. Studies in Plant Survival[M]. Oxford: Blackwell, 1989.[17]Thanos C A, Georghiou K, Skarou F. Glaucium flavumseed germination: an ecophysiological approach[J]. Annals of Botany, 1989, 63: 121-130. [18]Oberbauer S, Miller P C. Effect of water potential on seed germination[J]. Holarctic Ecology, 1982, 5: 218-220.[19]Singh P, Ibrahim H M, Flury M, et al. Critical water potentials for germination of wheat cultivars in the dryland Northwest USA[J]. Seed Science Research, 2013, 23: 189-198.[20]Parsons F R. Incidence and ecology of very fast germination[J]. Seed Science Research, 2012, 22: 161-167.[21]Liu K, Baskin J M, Baskin C C, et al. Very fast-germinating seeds of desert species are cryptoviparous like[J]. Seed Science Research, 2013, 23: 163-167.[22]Mariko S, Kachi N, Ishikawa S, et al. Germination ecology of coastal plants in relation to salt environment[J]. Ecological Research, 1992, 7: 225-233.[23]Baldwin A H, Mckee K L, Mendelssohn I A. The influence of vegetation, salinity, and inundation on seed banks of oligohaline coastal marshes[J]. American Journal of Botany, 1996, 83: 470-479.[24]Williams K, Meads M V, Sauerbrey D A. The roles of seedling salt tolerance and resprouting in forest zonation on the west coast of Florida, USA[J]. American Journal of Botany, 1998, 85: 1745-1752.[25]Isikawa S. Light sensitivity against germination. I. Photoperiodism in seeds[J]. Botanical Magazine Tokyo, 1954, 67: 51-56.[26]Cumming B G. The dependence of germination on photoperiod, light quality, and temperature in Chenopodiumspp[J]. Canadian Journal of Botany, 1963, 41: 1211-1233.[27]Densmore R V. Effect of day length on germination of seeds collected in Alaska[J]. American Journal of Botany, 1997, 84: 274-278.[28]Zhao D L. Effects of light on seed dormancy and germination[J]. Bulletin of Biology, 1995, 30: 24-25.[29]Black M, Wareing P F. Growth studies in woody species. VII. Photoperiodic control of germination in Betula pubescens Ehrh[J]. Physiologia Plantarum, 1955, 8: 300-316.[30]Stearns F, Olsen J. Interactions of photoperiod and temperature affecting seed germination in Tsuga canadensis[J]. American Journal of Botany, 1958, 45: 53-58.[31]Baskin J M, Baskin C C. Effect of photoperiod on germination of Cyperus inflexusseeds[J]. Botanical Gazette, 1976, 137: 269-273.[32]Bliss D, Smith H. Penetration of light into soil and its role in the control of seed germination[J]. Plant, Cell and Environment, 1985, 8: 475-483.[33]Tester M, Morris C. The penetration of light through soil[J]. Plant, Cell & Environment, 1987, 10: 281-286.[34]Wesson G, Wareing P F. The induction of light sensitivity in weed seeds by burial[J]. Journal of Experimental Botany, 1969, 20: 414-425.[35]Milberg P, Andersson L. Seasonal variation in dormancy and light sensitivity in buried seeds of eight annual weed species[J]. Canadian Journal of Botany, 1997, 75: 1998-2004.[36]Scopel A L, Ballaré C L, Sánchez R A. Induction of extreme light sensitivity in buried weed seeds and its role in the perception of soil cultivations[J]. Plant, Cell and Environment, 1991, 14: 501-508.[37]Górski T, Górska K. Inhibitory effects of full daylight on the germination of Lactuca sativaL[J]. Planta, 1979, 144: 121-124.[38]Pons T L. Seed responses to light[A]. Seeds: The Ecology of Regeneration in Plant Communities[M]. Wallingford: CABI Publishing, 2000: 237-260.[39]Taylorson R B, Borthwick H A. Light filtration by foliar canopies: significance for light controlled weed seed germination[J]. Weed Science, 1969, 17: 48-51.[40]Fenner M. The induction of a light requirement in Bidens pilosaseeds by leaf canopy shade[J]. New Phytologist, 1980, 84: 103-106.[41]Silvertown J W. Leaf canopy-induced seed dormancy in a grassland flora[J]. New Phytologist, 1980, 85: 109-118.[42]Górski T. Germination of seeds in the shadow of plants[J]. Physiologia Plantarum, 1975, 34: 342-346.[43]Górski T, Górska K, Nowicki J. Germination of seeds of various herbaceous species under leaf canopy[J]. Flora, 1977, 166: 249-259.[44]Górski T, Górska K, Rybicki J. Studies on the germination of seeds under leaf canopy[J]. Flora, 1978, 167: 289-299.[45]Ballard L A T. Physical barriers to germination[J]. Seed Science and Technology, 1981, 1: 285-303.[46]Keeley J E. Seed germination and life history syndromes in the California chaparral[J]. Botanical Review, 1991, 57: 81-116.[47]Keeley J E. Seed germination patterns in fire-prone Mediterranean-climate regions[A]. In: Arroyo M T K, Zedler P H, Fox M D. Ecology and Biogeography of Mediterranean Ecosystems in Chile, California and Australia[M]. San Diego: Academic Press, 1995: 239-273.[48]Kilian D, Cowling R M. Comparative seed biology and co-existence of two fynbos shrub species[J]. Journal of Vegetation Science, 1992, 3: 637-646.[49]Keeley J E, Bond W J. Convergent seed germination in South African fynbos and Californian chaparral[J]. Plant Ecology, 1997, 133: 153-167.[50]Herranz J M, Gerrandis P, Martinez-Sanchez J J. Influence of heat on seed germination of seven Mediterranean Leguminosae species[J]. Plant Ecology, 1998, 136: 95-103.[51]Baskin J M, Baskin C C. New approaches to the study of the evolution of physical and physiological dormancy, the two most common classes of seed dormancy on earth[A]. In: Nicolas G, Bradford K J, Come D, et al. The Biology of Seeds: Recent Research Advances[M]. Wallingford: CABI Publishing, 2003: 371-380.[52]Brown N A C. Promotion of germination of fynbos seeds by plant derived smoke[J]. New Phytologist, 1993, 123: 575-583.[53]Keeley J E, Fotheringham C J. Smoke induced seed germination of California chaparral[J]. Ecology, 1998, 79: 2320-2336.[54]Roche S, Dixon K W, Pate J S. Seed ageing and smoke: partner cues in the amelioration of seed dormancy in selected Australian native species[J]. Australian Journal of Botany, 1997, 45: 783-815.[55]Tieu A, Dixon K W, Meney K A, et al. Interaction of soil burial and smoke on germination patterns in seeds of selected Australian native plants[J]. Seed Science Research, 2001, 11: 69-76.[56]Van de Venter H A, Esterhuizen A D. The effects of factors associated with fire on seed germination of Erica sessilifloraand E. hebgecalys (Ericaceae)[J]. South African Journal of Botany, 1988, 54: 301-304.[57]Flematti G R, Ghisalberti E L, Dixon K W, et al. A compound from smoke that promotes seed germination[J]. Science, 2004, 305: 977.[58]Malakoff D A. Nitrogen oxide pollution may spark seeds growth[J].Science, 1997, 276: 1199.[59]Downes K S, Lamont B B, Light M E, et al. The fire ephemeral Tersonia cyathiflora(Gyrostemonaceae) germinates in response to smoke but not the butenolide 3-methyl-2H-furo[2,3-c]pyran-2-one[J]. Annuals of Botany, 2010, 106: 381-384.[60]Liu K, Baskin J M, Baskin C C, et al. Effect of diurnal fluctuating versus constant temperatures on germination of 445 species from the eastern Tibet Plateau[J]. PLoS ONE, 2013, 8: e69364.[61]Thompson K, Grime J P, Mason G. Seed germination in response to diurnal fluctuations of temperature[J]. Nature, 1997, 267: 147-149.[62]Benech Arnold R L, Ghersa C M, Sanchez R A, et al. The role of fluctuating temperatures in the germination and establishment of Sorghum halepense(L.) Pers. Regulation of germination under leaf canopies[J]. Functonal Ecology, 1988, 3: 311-318.[63]Ren J, Tao L, Liu X M. Effect of sand burial depth on seed germination and seedling emergence of Calligonum L. species[J]. Journal of Arid Environment, 2002, 51: 603-611.[64]Chauhan B S, Gill G, Preston C. Seed germination and seedling emergence of threehorn bedstraw (Galium tricornutum)[J]. Weed Science, 2006, 54: 867-872.[65]Ghersa C M, Benech Arnold R L, Martinez-Ghersa M A. The role of fluctuating temperatures in germination and establishment of Sorghum halepense. Regulation of germination at increasing depths[J]. Functional Ecology, 1992, 6: 460-468.[66]Thompson K, Grime J P. A comparative study of germination responses to diurnally-fluctuating temperatures[J]. Journal of Applied Ecology, 1983, 20: 141-156.[67]Pons T L, Schrder H F J M. Significance of temperature fluctuation and oxygen concentration for germination of the rice field weeds Fimbristylis littoralisand Scirpus juncoides[J]. Oecologia, 1986, 68: 315-319.[68]Steinbauer G P, Grigsby B. Interaction of temperature, light and moistening agent in the germination of weed seeds[J]. Weeds, 1957, 5: 157.[69]Pons T L. Breaking of seed dormancy by nitrate as a gap detection mechanism[J]. Annals of Botany, 1989, 63: 139-143.[70]Hintikka V. Germination ecology of Galeopsis bifida(Lamiaceae) as a pioneer species in forest succession[J]. Silva Fennica, 1987, 21: 301-313.[71]Scharenbroch B C, Bockheim J G. Impacts of forest gaps on soil properties and processes in old growth northern hardwood-hemlock forests[J]. Plant Soil, 2007, 294: 219-233.[72]Toole V T. Germination requirements of the seed of some introduced and native range grasses[J]. Proceeding Association Official Seed Analysts, 1938, 30: 227-243.[73]Hilton J R. The influence of light and potassium nitrate on the dormancy and germination of Avena fatua L. (wild oat) seed and its ecological significance[J]. New Phytologist, 1984, 96: 31-34.[74]Bell D T, King L A, Plummer J A. Ecophysiological effects of light quality and nitrate on seed germination in species from Western Australia[J]. Australian Journal of Ecology, 1999, 24: 2-10.[75]Saini H S, Bassi P S, Spencer M S. Seed germination in Chnopodium album L.: Further evidence for dependence of the effects of growth regulators on nitrate availability[J]. Plant Cell Environment, 1985, 8: 707-711.[76]Carmona R, Murdoch A J. Interactions of temperature and dormancy-relieving compounds on the germination of weed seeds[J]. Seed Science Research, 1995, 5: 227-236.[77]Finch-Savage W E, Leubner-Metzger G. Seed dormancy and the control of germination[J]. New Phytologist, 2006, 171: 501-523.[78]Donohue K, Dorn L, Griffith C, et al. Environmental and genetic influences on the germination of Arabidopsis thalianain the field[J]. Evolution, 2005, 59: 740-757.[79]Footitt S, Douterelo-Soler I, Clay H, et al. Dormancy cycling in Arabidopsisseeds is controlled by seasonally distinct hormone signaling pathways[J]. Proceedings of National Academy of Sciences of the United States of America, 2011, 108: 20236-20241.[80]Footitt S, Huang Z, Clay H, et al. Temperature, light and nitrate sensing coordinate Arabidopsisseed dormancy cycling, resulting in winter and summer annual phenotypes[J].The Plant Journal, 2013, 74: 1003-1015.[81]Cadman C S C, Toorop P E, Hilhorst H W M, et al. Gene expression profiles of Arabidopsis Cvi seeds during dormancy cycling indicate a common underlying dormancy control mechanism[J]. The Plant Journal, 2006, 46: 805-822.[82]Oh E, Kang H, Yamaguchi S, et al. Genome-wide analysis of genes targeted by PHYTOCHROME INTERACTING FACTOR 3-LIKE5 during seed germination in Arabidopsis[J]. Plant Cell, 2009, 21:403-419.[83]Matakiadis T, Alboresi A, Jikumaru Y, et al. The Arabidopsis abscisic acid catabolic gene CYP707A2 plays a key role in nitrate control of seed dormancy[J]. Plant Physiology, 2009, 149: 949-960.参考文献:[1]宗文杰，刘坤，卜海燕，等. 高寒草甸51种菊科植物种子大小变异及其对种子萌发的影响研究[J]. 兰州大学学报（自然科学版）, 2006, 42: 52-55.[2]张大勇. 植物生活史进化与繁殖生态学[M].北京: 科学出版社， 2004.[3]Cheplick G P. Do seed germination patterns in cleistogamous annual grasses reduce the risk of sibling competition[J]. Journal of Ecology, 1996, 84: 247-255.[4]鲁为华，万娟娟，杨洁晶，等. 草食动物对植物种子的消化道传播研究进展[J]. 草业学报，2013, 22（3）：306-313.[5]陆嘉惠，吕新，吴玲，等. 三种药用甘草种子对盐渍环境的萌发响应及其适宜生态种植区[J]. 草业学报，2013, 22（2）：195-202.[6]卢艳敏，苏长青，李会芬. 不同盐胁迫对白三叶种子萌发及幼苗生长的影响[J]. 草业学报，2013, 22（4）：123-129.[7]Baskin J M, Baskin C C. A classification system for seed dormancy[J]. Seed Science Research, 2004, 14: 1-16.[8]Vleeshouwers L M, Bouwmeester H J, Karssen C M. Redefining seed dormancy: an attempt to integrate physiology and ecology[J]. Journal of Ecology, 1995, 83: 1031-1037.[9]Baskin C C, Baskin J M. Seeds: Ecology, Biogeography, and Evolution of Dormancy and Germination[M]. San Diego: Academic Press, 1998.[10]Liu K, Baskin J M, Baskin C C,et al. Effect of storage conditions on germination of 489 species from high elevation grassland of the eastern Tibet Plateau and implications for climate change[J]. American Journal of Botany, 2011, 98: 12-19.[11]Washitani I, Masuda M. A comparative study of the germination characteristics of seeds from a moist tall grassland community[J]. Functional Ecology, 1990, 4: 543-457.[12]Grime J P, Mason G, Curtis A V,et al. A comparative study of germination characteristics in a local flora[J]. Journal of Ecology, 1981, 69: 1017-1059.[13]Schütz W. Are germination strategies important for the ability of cespitose wetland sedges（Carex) to grow in forests[J]. Canadian Journal of Botany, 1997, 75: 1692-1699.[14]Fenner M, Thompson K. The Ecology of Seeds[M]. Cambridge: Cambridge University Press, 2005.[15]Simon E W, Minchin A, McMenamin M M,et al. The low temperature limit for seed germination[J]. New Phytologist, 1976, 77: 301-311.[16]Crawford R M M. Studies in Plant Survival[M]. Oxford: Blackwell, 1989.[17]Thanos C A, Georghiou K, Skarou F. Glaucium flavum seed germination: an ecophysiological approach[J]. Annals of Botany, 1989, 63: 121-130. [18]Oberbauer S, Miller P C. Effect of water potential on seed germination[J]. Holarctic Ecology, 1982, 5: 218-220.[19]Singh P, Ibrahim H M, Flury M,et al. Critical water potentials for germination of wheat cultivars in the dryland Northwest USA[J]. Seed Science Research, 2013, 23: 189-198.[20]Parsons F R. Incidence and ecology of very fast germination[J]. Seed Science Research, 2012, 22: 161-167.[21]Liu K, Baskin J M, Baskin C C,et al. Very fast-germinating seeds of desert species are cryptoviparous-like[J]. Seed Science Research, 2013, 23: 163-167.[22]Mariko S, Kachi N, Ishikawa S,et al. Germination ecology of coastal plants in relation to salt environment[J]. Ecological Research, 1992, 7: 225-233.[23]Baldwin A H, Mckee K L, Mendelssohn I A. The influence of vegetation, salinity, and inundation on seed banks of oligohaline coastal marshes[J]. American Journal of Botany, 1996, 83: 470-479.[24]Williams K, Meads M V, Sauerbrey D A. The roles of seedling salt tolerance and resprouting in forest zonation on the west coast of Florida, USA[J]. American Journal of Botany, 1998, 85: 1745-1752.[25]Isikawa S. Light sensitivity against germination. I. Photoperiodism in seeds[J]. Botanical Magazine Tokyo, 1954, 67: 51-56.[26]Cumming B G. The dependence of germination on photoperiod, light quality, and temperature in Chenopodium spp[J]. Canadian Journal of Botany, 1963, 41: 1211-1233.[27]Densmore R V. Effect of day length on germination of seeds collected in Alaska[J]. American Journal of Botany, 1997, 84: 274-278.[28]赵笃乐. 光对种子休眠与萌发的影响[J]. 生物学通报, 1995, 30: 24-25.[29]Black M, Wareing P F. Growth studies in woody species. VII. Photoperiodic control of germination in Betula pubescensEhrh[J]. Physiologia Plantarum, 1955, 8: 300-316.[30]Stearns F, Olsen J. Interactions of photoperiod and temperature affecting seed germination in Tsuga canadensis[J]. American Journal of Botany, 1958, 45: 53-58.[31]Baskin J M, Baskin C C. Effect of photoperiod on germination of Cyperus inflexus seeds[J]. Botanical Gazette, 1976, 137: 269-273.[32]Bliss D, Smith H. Penetration of light into soil and its role in the control of seed germination[J]. Plant, Cell and Environment, 1985, 8: 475-483.[33]Tester M, Morris C. The penetration of light through soil[J]. Plant, Cell & Environment, 1987, 10: 281-286.[34]Wesson G, Wareing P F. The induction of light sensitivity in weed seeds by burial[J]. Journal of Experimental Botany, 1969, 20: 414-425.[35]Milberg P, Andersson L. Seasonal variation in dormancy and light sensitivity in buried seeds of eight annual weed species[J]. Canadian Journal of Botany, 1997, 75: 1998-2004.[36]Scopel A L, Ballaré C L, Sánchez R A. Induction of extreme light sensitivity in buried weed seeds and its role in the perception of soil cultivations[J]. Plant, Cell and Environment, 1991, 14: 501-508.[37]Górski T, Górska K. Inhibitory effects of full daylight on the germination of Lactuca sativa L[J]. Planta, 1979, 144: 121-124.[38]Pons T L. Seed responses to light[A]. Seeds: The Ecology of Regeneration in Plant Communities[M]. Wallingford: CABI Publishing, 2000: 237-260.[39]Taylorson R B, Borthwick H A. Light filtration by foliar canopies: significance for light controlled weed seed germination[J]. Weed Science, 1969, 17: 48-51.[40]Fenner M. The induction of a light requirement in Bidens pilosa seeds by leaf canopy shade[J]. New Phytologist, 1980, 84: 103-106.[41]Silvertown J W. Leaf-canopy-induced seed dormancy in a grassland flora[J]. New Phytologist, 1980, 85: 109-118.[42]Górski T. Germination of seeds in the shadow of plants[J]. Physiologia Plantarum, 1975, 34: 342-346.[43]Górski T, Górska K, Nowicki J. Germination of seeds of various herbaceous species under leaf canopy[J]. Flora, 1977, 166: 249-259.[44]Górski T, Górska K, Rybicki J. Studies on the germination of seeds under leaf canopy[J]. Flora, 1978, 167: 289-299.[45]Ballard L A T. Physical barriers to germination[J]. Seed Science and Technology, 1981, 1: 285-303.[46]Keeley J E. Seed germination and life history syndromes in the California chaparral[J]. Botanical Review, 1991, 57: 81-116.[47]Keeley J E. Seed germination patterns in fire-prone Mediterranean-climate regions[A]. In: Arroyo M T K, Zedler P H, Fox M D. Ecology and Biogeography of Mediterranean Ecosystems in Chile, California and Australia[M]. San Diego: Academic Press, 1995: 239-273.[48]Kilian D, Cowling R M. Comparative seed biology and co-existence of two fynbos shrub species[J]. Journal of Vegetation Science, 1992, 3: 637-646.[49]Keeley J E, Bond W J. Convergent seed germination in South African fynbos and Californian chaparral[J]. Plant Ecology, 1997, 133: 153-167.[50]Herranz J M, Gerrandis P, Martinez-Sanchez J J. Influence of heat on seed germination of seven Mediterranean Leguminosae species[J]. Plant Ecology, 1998, 136: 95-103.[51]Baskin J M, Baskin C C. New approaches to the study of the evolution of physical and physiological dormancy, the two most common classes of seed dormancy on earth[A]. In: Nicolas G, Bradford K J, Come D,et al. The Biology of Seeds: Recent Research Advances[M]. Wallingford: CABI Publishing, 2003: 371-380.[52]Brown N A C. Promotion of germination of fynbos seeds by plant-derived smoke[J]. New Phytologist, 1993, 123: 575-583.[53]Keeley J E, Fotheringham C J. Smoke induced seed germination of California chaparral[J]. Ecology, 1998, 79: 2320-2336.[54]Roche S, Dixon K W, Pate J S. Seed ageing and smoke: partner cues in the amelioration of seed dormancy in selected Australian native species[J]. Australian Journal of Botany, 1997, 45: 783-815.[55]Tieu A, Dixon K W, Meney K A,et al. Interaction of soil burial and smoke on germination patterns in seeds of selected Australian native plants[J]. Seed Science Research, 2001, 11: 69-76.[56]Van de Venter H A, Esterhuizen A D. The effects of factors associated with fire on seed germination of Erica sessiliflora and E. hebgecalys (Ericaceae)[J]. South African Journal of Botany, 1988, 54: 301-304.[57]Flematti G R, Ghisalberti E L, Dixon K W,et al. A compound from smoke that promotes seed germination[J]. Science, 2004, 305: 977.[58]Malakoff D A. Nitrogen oxide pollution may spark seeds growth[J].Science, 1997, 276: 1199.[59]Downes K S, Lamont B B, Light M E,et al. The fire ephemeral Tersonia cyathiflora (Gyrostemonaceae) germinates in response to smoke but not the butenolide 3-methyl-2H-furo[2,3-c]pyran-2-one[J]. Annuals of Botany, 2010, 106: 381-384.[60]Liu K, Baskin J M, Baskin C C,et al. Effect of diurnal fluctuating versus constant temperatures on germination of 445 species from the eastern Tibet Plateau[J]. PLoS ONE, 2013, 8: e69364.[61]Thompson K, Grime J P, Mason G. Seed germination in response to diurnal fluctuations of temperature[J]. Nature, 1997, 267: 147-149.[62]Benech Arnold R L, Ghersa C M, Sanchez R A,et al. The role of fluctuating temperatures in the germination and establishment of Sorghum halepense (L.) Pers. Regulation of germination under leaf canopies[J]. Functonal Ecology, 1988, 3: 311-318.[63]Ren J, Tao L, Liu X M. Effect of sand burial depth on seed germination and seedling emergence of Calligonum L. species[J]. Journal of Arid Environment, 2002, 51: 603-611.[64]Chauhan B S, Gill G, Preston C. Seed germination and seedling emergence of threehorn bedstraw（Galium tricornutum)[J]. Weed Science, 2006, 54: 867-872.[65]Ghersa C M, Benech Arnold R L, Martinez-Ghersa M A. The role of fluctuating temperatures in germination and establishment of Sorghum halepense. Regulation of germination at increasing depths[J]. Functional Ecology, 1992, 6: 460-468.[66]Thompson K, Grime J P. A comparative study of germination responses to diurnally-fluctuating temperatures[J]. Journal of Applied Ecology, 1983, 20: 141-156.[67]Pons T L, Schrder H F J M. Significance of temperature fluctuation and oxygen concentration for germination of the rice field weeds Fimbristylis littoralis and Scirpus juncoides[J]. Oecologia, 1986, 68: 315-319.[68]Steinbauer G P, Grigsby B. Interaction of temperature, light and moistening agent in the germination of weed seeds[J]. Weeds, 1957, 5: 157.[69]Pons T L. Breaking of seed dormancy by nitrate as a gap detection mechanism[J]. Annals of Botany, 1989, 63: 139-143.[70]Hintikka V. Germination ecology of Galeopsis bifida (Lamiaceae) as a pioneer species in forest succession[J]. Silva Fennica, 1987, 21: 301-313.[71]Scharenbroch B C, Bockheim J G. Impacts of forest gaps on soil properties and processes in old growth northern hardwood-hemlock forests[J]. Plant Soil, 2007, 294: 219-233.[72]Toole V T. Germination requirements of the seed of some introduced and native range grasses[J]. Proceeding Association Official Seed Analysts, 1938, 30: 227-243.[73]Hilton J R. The influence of light and potassium nitrate on the dormancy and germination of Avena fatua L. (wild oat) seed and its ecological significance[J]. New Phytologist, 1984, 96: 31-34.[74]Bell D T, King L A, Plummer J A. Ecophysiological effects of light quality and nitrate on seed germination in species from Western Australia[J]. Australian Journal of Ecology, 1999, 24: 2-10.[75]Saini H S, Bassi P S, Spencer M S. Seed germination in Chnopodium album L.: Further evidence for dependence of the effects of growth regulators on nitrate availability[J]. Plant Cell Environment, 1985, 8: 707-711.[76]Carmona R, Murdoch A J. Interactions of temperature and dormancy-relieving compounds on the germination of weed seeds[J]. Seed Science Research, 1995, 5: 227-236.[77]Finch-Savage W E, Leubner-Metzger G. Seed dormancy and the control of germination[J]. New Phytologist, 2006, 171: 501-523.[78]Donohue K, Dorn L, Griffith C,et al. Environmental and genetic influences on the germination of Arabidopsis thaliana in the field[J]. Evolution, 2005, 59: 740-757.[79]Footitt S, Douterelo-Soler I, Clay H,et al. Dormancy cycling in Arabidopsis seeds is controlled by seasonally distinct hormone-signaling pathways[J]. Proceedings of National Academy of Sciences of the United States of America, 2011, 108: 20236-20241.[80]Footitt S, Huang Z, Clay H,et al. Temperature, light and nitrate sensing coordinate Arabidopsis seed dormancy cycling, resulting in winter and summer annual phenotypes[J].The Plant Journal, 2013, 74: 1003-1015.[81]Cadman C S C, Toorop P E, Hilhorst H W M,et al. Gene expression profiles of Arabidopsis Cvi seeds during dormancy cycling indicate a common underlying dormancy control mechanism[J]. The Plant Journal, 2006, 46: 805-822.[82]Oh E, Kang H, Yamaguchi S,et al. 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