种子形态生理休眠研究进展
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摘要
种子形态生理休眠(MPD)是形态休眠和生理休眠的组合,其种胚一般为线形、匙形或球形,未分化或发育不完全。根据种子萌发对冷(或暖)层积的需求,种胚生长所需要的温度以及对赤霉素的响应,MPD分为非深度简单、中度简单、深度简单、深度简单上胚轴、非深度简单上胚轴、深度简单双重、非深度复杂、中度复杂和深度复杂9种类型。一些物种需要相同的环境条件促进种胚的生长和休眠的打破,而另一些物种需要不同的环境条件来完成这一过程。目前已报道的具有MPD的物种有34目72科植物。本文系统总结了MPD种子的形态结构、休眠打破的环境需求、MPD的起源与进化以及MPD的植物类群。最后,针对国内外的研究不足进行了探讨,并对未来需要进一步开展的研究工作进行了展望。
Seed morphophysiological dormancy( MPD) is a combination of physiological dormancy( PD) and morphological dormancy( MD),which occurs in seeds with undifferentiated or underdeveloped embryos that are linear,spatulate,or globular in shape. Based on cold and/or warm stratification requirements for germination,temperature requirements for embryo growth,and response to GA3,MPD can be classified into nine types,i.e. non-deep simple,intermediate simple,deep simple,deep simple epicotyl,non-deep simple epicotyl,deep simple double,nondeep complex,intermediate complex and deep complex MPD. The embryo growth and dormancy break are promoted by the same environmental conditions for species,whereas in others they require different conditions. The species that have been reported with MPD are about 34 orders and 72 families. We summarized the morphology of MPD seeds,the environmental conditions required to break MPD,the origin and evolution of MPD,and the species with MPD. Last,we summarized the shortcomings of the previous studies and prospected future researches.
Seed morphophysiological dormancy( MPD) is a combination of physiological dormancy( PD) and morphological dormancy( MD),which occurs in seeds with undifferentiated or underdeveloped embryos that are linear,spatulate,or globular in shape. Based on cold and/or warm stratification requirements for germination,temperature requirements for embryo growth,and response to GA3,MPD can be classified into nine types,i.e. non-deep simple,intermediate simple,deep simple,deep simple epicotyl,non-deep simple epicotyl,deep simple double,nondeep complex,intermediate complex and deep complex MPD. The embryo growth and dormancy break are promoted by the same environmental conditions for species,whereas in others they require different conditions. The species that have been reported with MPD are about 34 orders and 72 families. We summarized the morphology of MPD seeds,the environmental conditions required to break MPD,the origin and evolution of MPD,and the species with MPD. Last,we summarized the shortcomings of the previous studies and prospected future researches.
引文
曹帮华,蔡春菊.2006.银杏种子后熟生理与内源激素变化的研究.林业科学,42(2):32-37.
陈瑛.1999.实用中药种子技术手册.北京:人民卫生出版社.
傅飞龙.2017.黄精、多花黄精和滇黄精种子萌发及出苗特点研究(硕士学位论文).北京:中国医学科学院.
付子燕.2013.地下结实植物白番红花种子扩散及萌发特性(硕士学位论文).乌鲁木齐:新疆农业大学.
米热古力·努热拉.2014.两种短命植物的种子休眠与萌发特性及其生活史特征(硕士学位论文).乌鲁木齐:新疆农业大学.
潘睿,孙卫邦.2009.极度濒危植物大果木莲种子的休眠与萌发.植物生理学通讯,45(11):1089-1092.
苏贺.2016.巫山淫羊藿种子休眠机制及解除方法研究(博士学位论文).北京:中国农业大学.
黄玮,孟繁蕴,张文生,等.2008.滇重楼种子休眠机理研究.植物生理科学,24(12):242-246.
景新明,郑光华,裴颜龙,等.1995.野生紫斑牡丹和四川牡丹种子萌发特性及与其致濒的关系.生物多样性,3(2):84-87.
娄方芳,赵林森,李宗艳.2007.昆明西山野生黄牡丹种子休眠与萌发特性初步探究.西南林学院学报,27(5):34-37.
盛晋华,张雄杰,李宏义,等.2006.层积对肉苁蓉种子后熟作用的研究.中国种业,(3):23-24.
孙昌高,徐秀瑛,梅捍卫,等.1987.玉兰种子在层积处理期间的形态变化.种子,(1):22-25.
唐安军.2014.中国特有植物凹叶木兰种子的休眠与贮藏行为.植物生理学报,50(1):105-110.
王楠,高静,黄文静,等.2018.赤霉素浸种时长和施用浓度对重度干旱和盐胁迫下黄芪种子萌发的影响.生态学杂志,37(8):2339-2344.
喻梅,周守标,吴晓艳,等.2012.野生鸭儿芹种子休眠特性及破除方法.生态学报,32(4):1347-1354.
张俊杰,韦霄,柴胜丰,等.2018.珍稀濒危植物金丝李种子的休眠机理.生态学杂志,37(5):1371-1381
Albrecht MA,Mccarthy BC.2006.Seed germination and dormancy in the medicinal woodland herbs Collinsonia canadensis L.(Lamiaceae)and Dioscorea villosa L.(Dioscoreaceae).Flora:Morphology,Distribution,Functional Ecology of Plants,201:24-31.
Ali N,Probert R,Hay F,et al.2007.Postdispersal embryo growth and acquisition of desiccation tolerance in Anemone nemorosa L.seeds.Seed Science Research,17:155-163.
Chase MW,Christenhusz MJM,Fay MF,et al.2017.An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants:APG IV.Botanical Journal of the Linnean Society,181:1-20.
Barton LV.1933.Seedling production in tree peony.Contributions from Boyce Thompson Institute,5:451-460.
Barton LV,Chandler C.1957.Physiological and morphological effects of gibberellic acid on epicotyl dormancy of tree peony.Contributions from Boyce Thompson Institute,19:201-214.
Barton LV,Schroeder EM.1942.Dormancy in seeds of Convallaria majalis L.and Smilacina racemosa.Contributions from Boyce Thompson Institute,12:277-300.
Baskin CC,Baskin JM.2004.Classification system for seed dormancy.Seed Science Research,14:1-16.
Baskin CC,Baskin JM.2014.Seeds:Ecology,biogeography and evolution of dormancy and germination,2nd ed.San Diego:Academic Press.
Baskin CC,Chien CT,Chen SY,et al.2008.Germination of Viburnum odoratissimum seeds:A new level of morphophysiological dormancy.Seed Science Research,18:179-184.
Baskin CC,Chien CT,Chen SY,et al.2009.Epicotyl morphophysiological dormancy in seeds of Daphniphyllum glaucescens,a woody member of the Saxifragales.International Journal of Plant Sciences,170:174-181.
Baskin JM,Baskin CC.1984a.Germination ecophysiology of the woodland herb Osmorhiza longistylis(Umbelliferae).American Journal of Botany,71:687-692
Baskin JM,Baskin CC.1984b.Germination ecophysiology of an eastern deciduous forest herb Stylophorum diphyllum.American Midland Naturalist,111:390-399.
Baskin JM,Baskin CC.1985.Seed germination ecophysiology of the woodland spring geophyte Erythronium albidum.Botanical Gazette,146:130-136.
Baskin JM,Baskin CC.1989.Seed germination ecophysiology of Jeffersonia diphylla,a perennial herb of mesic deciduous forests.American Journal of Botany,76:1073-1080.
Baskin JM,Baskin CC.1990.Germination ecophysiology of seeds of the winter annual Chaerophyllum tainturieri:Anew type of morphophysiological dormancy.Journal of Ecology,78:993-1004.
Chase MW,Christenhusz MJM,Fay MF,et al.2016.An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants:APG IV.Botanical Journal of the Linnean Society,181:1-20.
Chien CT,Chen SY,Baskin JM,et al.2011a.Morphophysiological dormancy in seeds of the ANA grade angiosperm Schisandra arisanensis(Schisandraceae).Plant Species Biology,26:99-104.
Chien CT,Chen SY,Chien TY,et al.2011b.Nondeep simple morphophysiological dormancy in seeds of Ilex maximowicziana from northern(subtropical)and southern(tropical)Taiwan.Ecological Research,26:163-171.
Copete E,Herranz JM,Ferrandis P,et al.2011a.Physiology,morphology and phenology of seed dormancy break and germination in the endemic Iberian species Narcissus hispanicus(Amaryllidaceae).Annals of Botany,107:1003-1016.
Copete E,Herranz JM,Copete MA,et al.2011b.Non-deep complex morphophysiological dormancy in seeds of the Iberian Peninsula endemic geophyte Merendera montana(Colchicaceae).Seed Science Research,21:267-281.
da Silva EAA,de Melo DLB,Davide AC,et al.2007.Germination ecophysiology of Annona crassiflora seeds.Annals of Botany,99:823830.
Finch-Savage WE,Leubner-Metzger G.2006.Seed dormancy and the control of germination.New Phytologist,171:501-523.
Finkelstein R,Reeves W,Ariizumi T,et al.2008.Molecular aspects of seed dormancy.Annual Review of Plant Biology,59:387-415.
Forbis TA,Floyd SK,Queiroz AD.2002.The evolution of embryo size in angiosperms and other seed plants:Implications for the evolution of seed dormancy.Evolution,56:2112-2125.
Hao HP,He Z,Li H,et al.2014.Effect of root length on epicotyl dormancy release in seeds of Paeonia ludlowii,Tibetan peony.Annals of Botany,113:443-452.
Jing XM,Zheng GH.1999.The characteristics in seed germination and dormancy of four wild species of tree peonies and their bearing on endangerment.Acta Phytophysiologica Sinica,25:214-221.
Kharkwal AC,Kushwaha R,Prakash O,et al.2008.An efficient method of propagation of Podophyllum hexandrum:An endangered medicinal plant of the western Himalayas under ex situ conditions.Journal of Natural Medicines,62:211-216.
Kondo T,Miura T,Okubo N,et al.2004.Ecophysiology of deep simple epicotyl morphophysiological dormancy in seeds of Gagea lutea(Liliaceae).Seed Science Research,14:371-378.
Kondo T,Okubo N,Miura T,et al.2002.Ecophysiology of seed germination in Erythronium japonicum(Liliaceae)with underdeveloped embryos.American Journal of Botany,89:1779-1784.
Martin A C.1946.The comparative internal morphology of seeds.American Midland Naturalist,36:513-660.
Mondoni A,Probert R,Rossi G,et al.2009.Habitat-related germination behaviour and emergence phenology in the woodland geophyte Anemone ranunculoides L.(Ranunculaceae)from northern Italy.Seed Science Research,19:137-144.
Née G,Xiang Y,Soppe WJ.2017.The release of dormancy,a wake-up call for seeds to germinate.Current Opinion in Plant Biology,35:8-14.
Nikolaeva MG.1977.Factors controlling the seed dormancy pattern//Khan AA,Ed.The Physiology and Biochemistry of Seed Dormancy and Germination.Amsterdam:North-Holland Publication Co.
Pandey H,Nandi SK,Nadeem M,et al.2000.Chemical stimulation of seed germination in Aconitum heterophyllum Wall.and A.balfourii Stapf:Important Himalayan species of medicinal value.Seed Science Research,28:39-48.
Phartyal SS,Kondo T,Baskin JM,et al.2009.Temperature requirements differ for the two stages of seed dormancy break in Aegopodium podagraria(Apiaceae),a species with deep complex morphophysiological dormancy.American Journal of Botany,96:1086-1095.
Pritchard HW,Wood JA,Manger KR.1993.Influence of temperature on seed germination and the nutritional requirements for embryo growth in Arum maculatum L.New Phytologist,123:801-809.
Saunders AP.1918.A method of hastening germination of hard coated seeds.American Peony Society Bulletin,6:17-19.
Stokes P.1952.A physiological study of embryo development in Heracleum sphondylium L.II.The effect of temperature on after ripening.Annals of Botany,16:571-576.
Takagi H.2001.Breaking of two types of dormancy in seeds of Polygonatum odoratum used as vegetables.Journal of the Japanese Society for Horticultural Science,70:416-423.
Tipirdamaz R,Gomurgen AN.2000.The effects of temperature and gibberellic acid on germination of Eranthis hyemalis(L.)Salisb.seeds.Turkish Journal of Botany,24:143-145.
Vandelook F,Van Assche JA.2008.Deep complex morphophysiological dormancy in Sanicula europaea(Apiaceae)fits a recurring pattern of dormancy types in genera with an ArctoTertiary distribution.Botany,86:1370-1377.
Wang H,Van Staden J.2002.Seedling establishment characteristics of Paeonia ostii var.lishizhenii.South African Journal of Botany,68:382-385.
Willis CG,Baskin CC,Baskin JM,et al.2014.The evolution of seed dormancy:Environmental cues,evolutionary hubs,and diversification of the seed plants.New Phytologist,203:300-309.
Zilis MR,Meyer MM.1976.Rapid in vitro germination of immature,dormant embryos.International Plant Propagators’Society Combined Proceedings,26:272-275.
陈瑛.1999.实用中药种子技术手册.北京:人民卫生出版社.
傅飞龙.2017.黄精、多花黄精和滇黄精种子萌发及出苗特点研究(硕士学位论文).北京:中国医学科学院.
付子燕.2013.地下结实植物白番红花种子扩散及萌发特性(硕士学位论文).乌鲁木齐:新疆农业大学.
米热古力·努热拉.2014.两种短命植物的种子休眠与萌发特性及其生活史特征(硕士学位论文).乌鲁木齐:新疆农业大学.
潘睿,孙卫邦.2009.极度濒危植物大果木莲种子的休眠与萌发.植物生理学通讯,45(11):1089-1092.
苏贺.2016.巫山淫羊藿种子休眠机制及解除方法研究(博士学位论文).北京:中国农业大学.
黄玮,孟繁蕴,张文生,等.2008.滇重楼种子休眠机理研究.植物生理科学,24(12):242-246.
景新明,郑光华,裴颜龙,等.1995.野生紫斑牡丹和四川牡丹种子萌发特性及与其致濒的关系.生物多样性,3(2):84-87.
娄方芳,赵林森,李宗艳.2007.昆明西山野生黄牡丹种子休眠与萌发特性初步探究.西南林学院学报,27(5):34-37.
盛晋华,张雄杰,李宏义,等.2006.层积对肉苁蓉种子后熟作用的研究.中国种业,(3):23-24.
孙昌高,徐秀瑛,梅捍卫,等.1987.玉兰种子在层积处理期间的形态变化.种子,(1):22-25.
唐安军.2014.中国特有植物凹叶木兰种子的休眠与贮藏行为.植物生理学报,50(1):105-110.
王楠,高静,黄文静,等.2018.赤霉素浸种时长和施用浓度对重度干旱和盐胁迫下黄芪种子萌发的影响.生态学杂志,37(8):2339-2344.
喻梅,周守标,吴晓艳,等.2012.野生鸭儿芹种子休眠特性及破除方法.生态学报,32(4):1347-1354.
张俊杰,韦霄,柴胜丰,等.2018.珍稀濒危植物金丝李种子的休眠机理.生态学杂志,37(5):1371-1381
Albrecht MA,Mccarthy BC.2006.Seed germination and dormancy in the medicinal woodland herbs Collinsonia canadensis L.(Lamiaceae)and Dioscorea villosa L.(Dioscoreaceae).Flora:Morphology,Distribution,Functional Ecology of Plants,201:24-31.
Ali N,Probert R,Hay F,et al.2007.Postdispersal embryo growth and acquisition of desiccation tolerance in Anemone nemorosa L.seeds.Seed Science Research,17:155-163.
Chase MW,Christenhusz MJM,Fay MF,et al.2017.An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants:APG IV.Botanical Journal of the Linnean Society,181:1-20.
Barton LV.1933.Seedling production in tree peony.Contributions from Boyce Thompson Institute,5:451-460.
Barton LV,Chandler C.1957.Physiological and morphological effects of gibberellic acid on epicotyl dormancy of tree peony.Contributions from Boyce Thompson Institute,19:201-214.
Barton LV,Schroeder EM.1942.Dormancy in seeds of Convallaria majalis L.and Smilacina racemosa.Contributions from Boyce Thompson Institute,12:277-300.
Baskin CC,Baskin JM.2004.Classification system for seed dormancy.Seed Science Research,14:1-16.
Baskin CC,Baskin JM.2014.Seeds:Ecology,biogeography and evolution of dormancy and germination,2nd ed.San Diego:Academic Press.
Baskin CC,Chien CT,Chen SY,et al.2008.Germination of Viburnum odoratissimum seeds:A new level of morphophysiological dormancy.Seed Science Research,18:179-184.
Baskin CC,Chien CT,Chen SY,et al.2009.Epicotyl morphophysiological dormancy in seeds of Daphniphyllum glaucescens,a woody member of the Saxifragales.International Journal of Plant Sciences,170:174-181.
Baskin JM,Baskin CC.1984a.Germination ecophysiology of the woodland herb Osmorhiza longistylis(Umbelliferae).American Journal of Botany,71:687-692
Baskin JM,Baskin CC.1984b.Germination ecophysiology of an eastern deciduous forest herb Stylophorum diphyllum.American Midland Naturalist,111:390-399.
Baskin JM,Baskin CC.1985.Seed germination ecophysiology of the woodland spring geophyte Erythronium albidum.Botanical Gazette,146:130-136.
Baskin JM,Baskin CC.1989.Seed germination ecophysiology of Jeffersonia diphylla,a perennial herb of mesic deciduous forests.American Journal of Botany,76:1073-1080.
Baskin JM,Baskin CC.1990.Germination ecophysiology of seeds of the winter annual Chaerophyllum tainturieri:Anew type of morphophysiological dormancy.Journal of Ecology,78:993-1004.
Chase MW,Christenhusz MJM,Fay MF,et al.2016.An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants:APG IV.Botanical Journal of the Linnean Society,181:1-20.
Chien CT,Chen SY,Baskin JM,et al.2011a.Morphophysiological dormancy in seeds of the ANA grade angiosperm Schisandra arisanensis(Schisandraceae).Plant Species Biology,26:99-104.
Chien CT,Chen SY,Chien TY,et al.2011b.Nondeep simple morphophysiological dormancy in seeds of Ilex maximowicziana from northern(subtropical)and southern(tropical)Taiwan.Ecological Research,26:163-171.
Copete E,Herranz JM,Ferrandis P,et al.2011a.Physiology,morphology and phenology of seed dormancy break and germination in the endemic Iberian species Narcissus hispanicus(Amaryllidaceae).Annals of Botany,107:1003-1016.
Copete E,Herranz JM,Copete MA,et al.2011b.Non-deep complex morphophysiological dormancy in seeds of the Iberian Peninsula endemic geophyte Merendera montana(Colchicaceae).Seed Science Research,21:267-281.
da Silva EAA,de Melo DLB,Davide AC,et al.2007.Germination ecophysiology of Annona crassiflora seeds.Annals of Botany,99:823830.
Finch-Savage WE,Leubner-Metzger G.2006.Seed dormancy and the control of germination.New Phytologist,171:501-523.
Finkelstein R,Reeves W,Ariizumi T,et al.2008.Molecular aspects of seed dormancy.Annual Review of Plant Biology,59:387-415.
Forbis TA,Floyd SK,Queiroz AD.2002.The evolution of embryo size in angiosperms and other seed plants:Implications for the evolution of seed dormancy.Evolution,56:2112-2125.
Hao HP,He Z,Li H,et al.2014.Effect of root length on epicotyl dormancy release in seeds of Paeonia ludlowii,Tibetan peony.Annals of Botany,113:443-452.
Jing XM,Zheng GH.1999.The characteristics in seed germination and dormancy of four wild species of tree peonies and their bearing on endangerment.Acta Phytophysiologica Sinica,25:214-221.
Kharkwal AC,Kushwaha R,Prakash O,et al.2008.An efficient method of propagation of Podophyllum hexandrum:An endangered medicinal plant of the western Himalayas under ex situ conditions.Journal of Natural Medicines,62:211-216.
Kondo T,Miura T,Okubo N,et al.2004.Ecophysiology of deep simple epicotyl morphophysiological dormancy in seeds of Gagea lutea(Liliaceae).Seed Science Research,14:371-378.
Kondo T,Okubo N,Miura T,et al.2002.Ecophysiology of seed germination in Erythronium japonicum(Liliaceae)with underdeveloped embryos.American Journal of Botany,89:1779-1784.
Martin A C.1946.The comparative internal morphology of seeds.American Midland Naturalist,36:513-660.
Mondoni A,Probert R,Rossi G,et al.2009.Habitat-related germination behaviour and emergence phenology in the woodland geophyte Anemone ranunculoides L.(Ranunculaceae)from northern Italy.Seed Science Research,19:137-144.
Née G,Xiang Y,Soppe WJ.2017.The release of dormancy,a wake-up call for seeds to germinate.Current Opinion in Plant Biology,35:8-14.
Nikolaeva MG.1977.Factors controlling the seed dormancy pattern//Khan AA,Ed.The Physiology and Biochemistry of Seed Dormancy and Germination.Amsterdam:North-Holland Publication Co.
Pandey H,Nandi SK,Nadeem M,et al.2000.Chemical stimulation of seed germination in Aconitum heterophyllum Wall.and A.balfourii Stapf:Important Himalayan species of medicinal value.Seed Science Research,28:39-48.
Phartyal SS,Kondo T,Baskin JM,et al.2009.Temperature requirements differ for the two stages of seed dormancy break in Aegopodium podagraria(Apiaceae),a species with deep complex morphophysiological dormancy.American Journal of Botany,96:1086-1095.
Pritchard HW,Wood JA,Manger KR.1993.Influence of temperature on seed germination and the nutritional requirements for embryo growth in Arum maculatum L.New Phytologist,123:801-809.
Saunders AP.1918.A method of hastening germination of hard coated seeds.American Peony Society Bulletin,6:17-19.
Stokes P.1952.A physiological study of embryo development in Heracleum sphondylium L.II.The effect of temperature on after ripening.Annals of Botany,16:571-576.
Takagi H.2001.Breaking of two types of dormancy in seeds of Polygonatum odoratum used as vegetables.Journal of the Japanese Society for Horticultural Science,70:416-423.
Tipirdamaz R,Gomurgen AN.2000.The effects of temperature and gibberellic acid on germination of Eranthis hyemalis(L.)Salisb.seeds.Turkish Journal of Botany,24:143-145.
Vandelook F,Van Assche JA.2008.Deep complex morphophysiological dormancy in Sanicula europaea(Apiaceae)fits a recurring pattern of dormancy types in genera with an ArctoTertiary distribution.Botany,86:1370-1377.
Wang H,Van Staden J.2002.Seedling establishment characteristics of Paeonia ostii var.lishizhenii.South African Journal of Botany,68:382-385.
Willis CG,Baskin CC,Baskin JM,et al.2014.The evolution of seed dormancy:Environmental cues,evolutionary hubs,and diversification of the seed plants.New Phytologist,203:300-309.
Zilis MR,Meyer MM.1976.Rapid in vitro germination of immature,dormant embryos.International Plant Propagators’Society Combined Proceedings,26:272-275.