克隆植物中的同步雌雄异熟现象研究进展
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摘要
很多植物具有克隆生长的习性,克隆生长对植物的交配格局和长期进化具有重要意义。克隆生长通常伴随有较多的花展示,但花粉的有效散布又会受到克隆规模的限制,进而形成分株间的同株异花授粉,造成近交衰退或花粉贴现。同步雌雄异熟是雌雄异熟的一种特殊形式,同时满足2个条件:1)单个花朵内为雌雄异熟;2)不同花朵间在同一时段内处于同一性别阶段。该种现象可以有效阻止分株内或分株间同株异花授粉,并可能是一种进化选择的特殊繁殖保障机制。目前其在克隆植物中的表现形式和作用机理尚未明确。在梳理近20a来相关文献的基础上,以克隆植物中的同步雌雄异熟现象为核心议题,结合克隆植物种群结构的特殊性探讨该现象对克隆植物的进化意义,并对该现象的类群分布情况、研究方法以及主要结论进行了分析,认为未来的研究应该注重4个方面:1)采用统一的同步性度量体系,并使其扩展到对雌性和雄性阶段的分别描述;2)重视目标物种交配系统的研究,尤其是自交亲和性的重要意义;3)重视克隆多样性格局对交配过程的影响,并采取恰当的方法予以量化;4)加强风媒、水媒等非生物因素传粉方式对克隆植物交配过程影响的研究。
Clonal growth is a common trait in plants.It has profound effect in the mating pattern and long term evolution of plant population but is currently seldom explored.Clonal growth is usually associated with mass flowering,restricted pollen dispersal,and geitonogamous self-pollination,which can result in inbreeding depression and pollen discounting.However,mating costs can be reduced by differences in the relative timing of female and male function(dichogamy).Synchronous dichogamy,a subtype of dichogamy,fulfills:1)dichogamy within one flower;2)having the same sexual phase between different flowers at a given time.It's a potentially effective mechanism to limit inter/intra ramet geitonogamy,as well as a unique way to provide reproductive assurance.In this review,we surveyed related literatures of the past two decades.We focused on the concepts and evolutionary significance of synchronous dichogamy in clonal plants.A summary of acquired information on systematic distribution,research methods and ecological significance of synchronous dichogamy in clonal plants was presented.We recommend future research efforts to address such aspects:1)universal and effective metrics to evaluate synchronicity of different sex function separately;2)the ecological implication of selfing compatibility in synchronous dichogamy;3)the interaction of clonal diversity and mating patterns with synchronicity taken into consideration;4)abiotic pollination and synchronous dichogamy in clonal plants.
Clonal growth is a common trait in plants.It has profound effect in the mating pattern and long term evolution of plant population but is currently seldom explored.Clonal growth is usually associated with mass flowering,restricted pollen dispersal,and geitonogamous self-pollination,which can result in inbreeding depression and pollen discounting.However,mating costs can be reduced by differences in the relative timing of female and male function(dichogamy).Synchronous dichogamy,a subtype of dichogamy,fulfills:1)dichogamy within one flower;2)having the same sexual phase between different flowers at a given time.It's a potentially effective mechanism to limit inter/intra ramet geitonogamy,as well as a unique way to provide reproductive assurance.In this review,we surveyed related literatures of the past two decades.We focused on the concepts and evolutionary significance of synchronous dichogamy in clonal plants.A summary of acquired information on systematic distribution,research methods and ecological significance of synchronous dichogamy in clonal plants was presented.We recommend future research efforts to address such aspects:1)universal and effective metrics to evaluate synchronicity of different sex function separately;2)the ecological implication of selfing compatibility in synchronous dichogamy;3)the interaction of clonal diversity and mating patterns with synchronicity taken into consideration;4)abiotic pollination and synchronous dichogamy in clonal plants.
引文
[1]董鸣,于飞海.克隆植物生态学术语和概念[J].植物生态学报.2007,31(4):689-694.
[2]DONG M,YU F,ALPERT P.Ecological consequences of plant clonality[J].Annals of Botany,2014,114(2):367-379.
[3]李甜江,李根前,韦宇,等.克隆植物中国沙棘人工林生长动态研究[J].西北林学院学报,2007,22(4):30-34.LI T J,LI G Q,WEI Y,et al.Growth dynamics of plantation of clonal tree species Hippophae rhamniodes L.subsp sinensis[J].Journal of Northwest Forestry University,2007,22(4):30-34.(in Chinese)
[4]VALLEJO-MARIN M,DORKEN M E,BARRETT S C.The ecological and evolutionary consequencesof clonality for plant mating[J].The Annual Review of Ecology,Evolution,and Systematics,2010,41(3):193-213.
[5]王宁,李卫芳,周兵,等.中国入侵克隆植物入侵性、克隆方式及地理起源[J].生物多样性,2016,24(1):12-19.WANG N,LI W F,ZHOU B,et al.Invasiveness,clonal form and geographical origin of invasive clonal plant species in China[J].Biodiversity,2016,24(1):12-19.(in Chinese)
[6]安锋,阚丽艳,谢贵水,等.海南外来植物入侵的现状与对策[J].西北林学院学报,2007,22(5):193-197.AN F,KAN L Y,XIE G S,et al.Alien invasion plants in Hainan Island and contral countermeasures[J]Journal of Northwest Forestry University,2007,22(5):193-197.
[7]BARRETT S C.Influences of clonality on plant sexual reproduction[J].Proceedings of the National Academy of Sciences,2015,112(29):8859-8866.
[8]BARRETT S C H,HARDER L D.The ecology of mating and its evolutionary consequences in seed plants[J].Annual Review of Ecology Evolution&Systematics,2016,48(1):1-23.
[9]LLOYD D G,WEBB C J.The avoidance of interference between the presentation of pollen and stigmas in angiosperms I.dichogamy[J].New Zealand Journal of Botany,1986,24:135-162.
[10]CETINBAS A,UNAL M.An overview of dichogamy in angiosperms[J].Research in Plant Biology,2014,4(5):9-27.
[11]予茜,张彦文,郭友好.传粉生物学常用术语释译[J].植物分类学报,2008,46(1):96-102.YU Q,ZHANG Y W,GUO Y H.Translation and elucidation of common terms in pollination biology[J].Journal of Systematics and Evolution,2008,46(1):96-102.(in Chinese)
[12]DE KROON H,HUBER H,STUEFER J F,et al.A modular concept of phenotypic plasticity in plants[J].New Phytologist,2005,166(1):73-82.
[13]CHARPENTIER A.Consequences of clonal growth for plant mating[J].Evolutionary Ecology,2002,15(3):521-530.
[14]ECKERT C G,DORKEN M E,BARRETT S C H.Ecological and evolutionary consequences of sexual and clonal reproduction in aquatic plants[J].Aquatic Botany,2016,135:46-61.
[15]ZIPPERLE A M,COYER J A,REISE K,et al.An evaluation of small-scale genetic diversity and the mating system in Zostera noltii on an intertidal sandflat in the Wadden Sea[J].Annals of Botany,2011,107:127-134.
[16]VAN DRUNEN W E,VAN KLEUNEN M,DORKEN M E.Consequences of clonality for sexual fitness:clonal expansion enhances fitness under spatially restricted dispersal[J].Proceedings of the National Academy of Sciences,2015,112(29):8929-8936.
[17]HU Y,BARRETT S C H,ZHANG D Y,et al.Experimental analysis of mating patterns in a clonal plant reveals contrasting modes of self-pollination[J].Ecology&Evolution,2016,5(22):5423-5431.
[18]SINCLAIR E A,GECAN I,KRAUSS S L,et al.Against the odds:complete outcrossing in a monoecious clonal seagrass Posidonia australis(Posidoniaceae)[J].Annals of Botany,2014,113(7):1185-1196.
[19]FRIEDMAN J,BARRETT S C H.Wind of change:new insights on the ecology and evolution of pollination and mating in wind-pollinated plants[J].Annals of Botany,2009,103(9):1515-1527.
[20]FRIEDMAN J,BARRETT S.A phylogenetic analysis of the evolution of wind pollination in the angiosperms[J].International Journal of Plant Sciences,2008,169(1):49-58.
[21]CRUDEN R W.Temporal dioecism:systematic breadth,associated traits,and temporal patterns[J].Botanical Gazette,1988,149(1):1-15.
[22]WATANABE S,NOMA N,NISHIDA T.Flowering phenology and mating success of the heterodichogamous tree Machilus thunbergii Sieb.et Zucc(Lauraceae)[J].Plant Species Biology,2016,31(1):29-37.
[23]KUBITZKI K,KURZ H.Synchronized dichogamy and dioecy in neotropical lauraceae[J].Plant Systematics and Evolution,1984,147(3-4):253-266.
[24]CERINO M C,RICHARD G A,TORRETTA J P,et al.Reproductive biology of Ziziphus mistol Griseb.(Rhamnaceae),a wild fruit tree of saline environments[J].Flora-Morphology,Distribution,Functional Ecology of Plants,2015,211:18-25.
[25]PANG C,SAUNDERS R M K.Floral Biology and Pollination Ecology of Desmos chinensis(Annonaceae):assessing the efficacy of floral synchrony for promoting xenogamy[J].International Journal of Plant Sciences,2015,176(4):333-345.
[26]PANG C,SAUNDERS R M K.The evolution of alternative mechanisms that promote outcrossing in annonaceae,a selfcompatible family of early-divergent angiosperms[J].Botanical Journal of the Linnean Society,2014,174(1):93-109.
[27]KHANDURI V P,SHARMA C M,KUMAR K S,et al.Annual variation in flowering phenology,pollination,mating system,and pollen yield in two natural populations of Schima wallichii(DC.)Korth[J].The Scientific World Journal,2013,2013:11.
[28]NARBONA E,ORTIZ P L,ARISTA M.Linking self-incompatibility,dichogamy,and flowering synchrony in two Euphorbiaspecies:alternative mechanisms for avoiding self-fertilization?[J].PLoS ONE,2011,6(6):e20668.
[29]PEI N,LUO Z,SCHLESSMAN M,et al.Synchronized protandry and hermaphroditism in a tropical secondary forest tree,Schefflera heptaphylla(Araliaceae)[J].Plant Systematics and Evolution,2011,296(1-2):29-39.
[30]王翠翠,刘文哲,张莹.喜树开花特性及繁育系统的研究[J].热带亚热带植物学报,2009,17(3):275-282.WANG C C,LIU W Z,ZHANG Y.Flowering characteristics and breeding system in Camptotheca acuminata Decne[J].Journal of Tropical and Subtropical Botany,2009,17(3):275-282.(in Chinese)
[31]刘宇,周桂玲.大赖草的交配系统及生殖对策[J].中国沙漠.2010,30(1):92-96.LIU Y,ZHOU G L Mating system and reproductive strategy of Leymus racemosus[J].Journal of Desert Research,2010,30(1):92-96.(in Chinese)
[32]赵元杰.芒繁育系统的研究[D].长沙:湖南农业大学,2010.
[33]SNYDER J M,RICHARDS J H.Floral phenology and compatibility of sawgrass,Cladium jamaicense(Cyperaceae)[J].American Journal of Botany,2005,92(4):736-743.
[34]MICHALSKI S G,DURKA W.Synchronous pulsed flowering:analysis of the flowering phenology in Juncus(Juncaceae)[J].Annals of Botany,2007,100(6):1271-1285.
[35]BHARDWAJ M,ECKERT C G.Functional analysis of synchronous dichogamy in flowering rush,Butomus umbellatus(Butomaceae)[J].American Journal of Botany,2001,88(12):2204-2213.
[36]HARDER L D,AIZEN M A.The functional significance of synchronous protandry in Alstroemeria aurea[J].Functional Ecology,2004,18(3):467-474.
[37]AUGSPURGER C K.Phenology,flowering synchrony,and fruit set of six neotropical shrubs[J].Biotropica,1983,15(4):257-267.
[38]MARQUIS R J.Phenological variation in the neotropical understory shrub Piper arielanum:causes and consequences[J].Ecology,1988,69(5):1552-1565.
[39]THOMSON J D,BARRETT S C.Temporal variation of gender in Aralia hispida Vent.(Araliaceae)[J].Evolution,1981:1094-1107.
[40]BRIGGS B G,TINKER A.Synchronous monoecy in ecdeiocoleaceae(Poales),in Western Australia[J].Australian Journal of Botany,2014,62(5):391-402.
[41]LIAO W,HARDER L D.Consequences of multiple inflorescences and clonality for pollinator behavior and plant mating[J].The American Naturalist,2014,184(5):580-592.
[42]ISAGI Y,SHIMADA K,KUSHIMA H,et al.Clonal structure and flowering traits of a bamboo[Phyllostachys pubescens(Mazel)Ohwi].stand grown from a simultaneous flowering as revealed by AFLP analysis[J].Molecular Ecology,2017,13(7):2017-2021.
[43]邱月群,董文渊,王逸之,等.筇竹天然居群遗传多样性研究[J].西北林学院学报,2017,32(2):155-160.QIU Y Q,DONG W Y,WANG Y Z,et al.Genetic diversity in natural populations of Qiongzhuea tumidinoda[J].Journal of Northwest Forestry University,2017,32(2):155-160.(in Chinese)
[44]BERTIN R I.Incidence of monoecy and dichogamy in relation to self-fertilization in angiosperms[J].American Journal of Botany,1993,80(5):557-560.
[45]CRUDEN R W.Temporal dioecism:systematic breadth,associated traits,and temporal patterns[J].Botanical Gazette,1988,149:1-15.
[46]ECKERT C G.Contributions of autogamy and geitonogamy to self-fertilization in a mass-flowering,clonal plant[J].Ecology,2000,81(2):532-542.
[2]DONG M,YU F,ALPERT P.Ecological consequences of plant clonality[J].Annals of Botany,2014,114(2):367-379.
[3]李甜江,李根前,韦宇,等.克隆植物中国沙棘人工林生长动态研究[J].西北林学院学报,2007,22(4):30-34.LI T J,LI G Q,WEI Y,et al.Growth dynamics of plantation of clonal tree species Hippophae rhamniodes L.subsp sinensis[J].Journal of Northwest Forestry University,2007,22(4):30-34.(in Chinese)
[4]VALLEJO-MARIN M,DORKEN M E,BARRETT S C.The ecological and evolutionary consequencesof clonality for plant mating[J].The Annual Review of Ecology,Evolution,and Systematics,2010,41(3):193-213.
[5]王宁,李卫芳,周兵,等.中国入侵克隆植物入侵性、克隆方式及地理起源[J].生物多样性,2016,24(1):12-19.WANG N,LI W F,ZHOU B,et al.Invasiveness,clonal form and geographical origin of invasive clonal plant species in China[J].Biodiversity,2016,24(1):12-19.(in Chinese)
[6]安锋,阚丽艳,谢贵水,等.海南外来植物入侵的现状与对策[J].西北林学院学报,2007,22(5):193-197.AN F,KAN L Y,XIE G S,et al.Alien invasion plants in Hainan Island and contral countermeasures[J]Journal of Northwest Forestry University,2007,22(5):193-197.
[7]BARRETT S C.Influences of clonality on plant sexual reproduction[J].Proceedings of the National Academy of Sciences,2015,112(29):8859-8866.
[8]BARRETT S C H,HARDER L D.The ecology of mating and its evolutionary consequences in seed plants[J].Annual Review of Ecology Evolution&Systematics,2016,48(1):1-23.
[9]LLOYD D G,WEBB C J.The avoidance of interference between the presentation of pollen and stigmas in angiosperms I.dichogamy[J].New Zealand Journal of Botany,1986,24:135-162.
[10]CETINBAS A,UNAL M.An overview of dichogamy in angiosperms[J].Research in Plant Biology,2014,4(5):9-27.
[11]予茜,张彦文,郭友好.传粉生物学常用术语释译[J].植物分类学报,2008,46(1):96-102.YU Q,ZHANG Y W,GUO Y H.Translation and elucidation of common terms in pollination biology[J].Journal of Systematics and Evolution,2008,46(1):96-102.(in Chinese)
[12]DE KROON H,HUBER H,STUEFER J F,et al.A modular concept of phenotypic plasticity in plants[J].New Phytologist,2005,166(1):73-82.
[13]CHARPENTIER A.Consequences of clonal growth for plant mating[J].Evolutionary Ecology,2002,15(3):521-530.
[14]ECKERT C G,DORKEN M E,BARRETT S C H.Ecological and evolutionary consequences of sexual and clonal reproduction in aquatic plants[J].Aquatic Botany,2016,135:46-61.
[15]ZIPPERLE A M,COYER J A,REISE K,et al.An evaluation of small-scale genetic diversity and the mating system in Zostera noltii on an intertidal sandflat in the Wadden Sea[J].Annals of Botany,2011,107:127-134.
[16]VAN DRUNEN W E,VAN KLEUNEN M,DORKEN M E.Consequences of clonality for sexual fitness:clonal expansion enhances fitness under spatially restricted dispersal[J].Proceedings of the National Academy of Sciences,2015,112(29):8929-8936.
[17]HU Y,BARRETT S C H,ZHANG D Y,et al.Experimental analysis of mating patterns in a clonal plant reveals contrasting modes of self-pollination[J].Ecology&Evolution,2016,5(22):5423-5431.
[18]SINCLAIR E A,GECAN I,KRAUSS S L,et al.Against the odds:complete outcrossing in a monoecious clonal seagrass Posidonia australis(Posidoniaceae)[J].Annals of Botany,2014,113(7):1185-1196.
[19]FRIEDMAN J,BARRETT S C H.Wind of change:new insights on the ecology and evolution of pollination and mating in wind-pollinated plants[J].Annals of Botany,2009,103(9):1515-1527.
[20]FRIEDMAN J,BARRETT S.A phylogenetic analysis of the evolution of wind pollination in the angiosperms[J].International Journal of Plant Sciences,2008,169(1):49-58.
[21]CRUDEN R W.Temporal dioecism:systematic breadth,associated traits,and temporal patterns[J].Botanical Gazette,1988,149(1):1-15.
[22]WATANABE S,NOMA N,NISHIDA T.Flowering phenology and mating success of the heterodichogamous tree Machilus thunbergii Sieb.et Zucc(Lauraceae)[J].Plant Species Biology,2016,31(1):29-37.
[23]KUBITZKI K,KURZ H.Synchronized dichogamy and dioecy in neotropical lauraceae[J].Plant Systematics and Evolution,1984,147(3-4):253-266.
[24]CERINO M C,RICHARD G A,TORRETTA J P,et al.Reproductive biology of Ziziphus mistol Griseb.(Rhamnaceae),a wild fruit tree of saline environments[J].Flora-Morphology,Distribution,Functional Ecology of Plants,2015,211:18-25.
[25]PANG C,SAUNDERS R M K.Floral Biology and Pollination Ecology of Desmos chinensis(Annonaceae):assessing the efficacy of floral synchrony for promoting xenogamy[J].International Journal of Plant Sciences,2015,176(4):333-345.
[26]PANG C,SAUNDERS R M K.The evolution of alternative mechanisms that promote outcrossing in annonaceae,a selfcompatible family of early-divergent angiosperms[J].Botanical Journal of the Linnean Society,2014,174(1):93-109.
[27]KHANDURI V P,SHARMA C M,KUMAR K S,et al.Annual variation in flowering phenology,pollination,mating system,and pollen yield in two natural populations of Schima wallichii(DC.)Korth[J].The Scientific World Journal,2013,2013:11.
[28]NARBONA E,ORTIZ P L,ARISTA M.Linking self-incompatibility,dichogamy,and flowering synchrony in two Euphorbiaspecies:alternative mechanisms for avoiding self-fertilization?[J].PLoS ONE,2011,6(6):e20668.
[29]PEI N,LUO Z,SCHLESSMAN M,et al.Synchronized protandry and hermaphroditism in a tropical secondary forest tree,Schefflera heptaphylla(Araliaceae)[J].Plant Systematics and Evolution,2011,296(1-2):29-39.
[30]王翠翠,刘文哲,张莹.喜树开花特性及繁育系统的研究[J].热带亚热带植物学报,2009,17(3):275-282.WANG C C,LIU W Z,ZHANG Y.Flowering characteristics and breeding system in Camptotheca acuminata Decne[J].Journal of Tropical and Subtropical Botany,2009,17(3):275-282.(in Chinese)
[31]刘宇,周桂玲.大赖草的交配系统及生殖对策[J].中国沙漠.2010,30(1):92-96.LIU Y,ZHOU G L Mating system and reproductive strategy of Leymus racemosus[J].Journal of Desert Research,2010,30(1):92-96.(in Chinese)
[32]赵元杰.芒繁育系统的研究[D].长沙:湖南农业大学,2010.
[33]SNYDER J M,RICHARDS J H.Floral phenology and compatibility of sawgrass,Cladium jamaicense(Cyperaceae)[J].American Journal of Botany,2005,92(4):736-743.
[34]MICHALSKI S G,DURKA W.Synchronous pulsed flowering:analysis of the flowering phenology in Juncus(Juncaceae)[J].Annals of Botany,2007,100(6):1271-1285.
[35]BHARDWAJ M,ECKERT C G.Functional analysis of synchronous dichogamy in flowering rush,Butomus umbellatus(Butomaceae)[J].American Journal of Botany,2001,88(12):2204-2213.
[36]HARDER L D,AIZEN M A.The functional significance of synchronous protandry in Alstroemeria aurea[J].Functional Ecology,2004,18(3):467-474.
[37]AUGSPURGER C K.Phenology,flowering synchrony,and fruit set of six neotropical shrubs[J].Biotropica,1983,15(4):257-267.
[38]MARQUIS R J.Phenological variation in the neotropical understory shrub Piper arielanum:causes and consequences[J].Ecology,1988,69(5):1552-1565.
[39]THOMSON J D,BARRETT S C.Temporal variation of gender in Aralia hispida Vent.(Araliaceae)[J].Evolution,1981:1094-1107.
[40]BRIGGS B G,TINKER A.Synchronous monoecy in ecdeiocoleaceae(Poales),in Western Australia[J].Australian Journal of Botany,2014,62(5):391-402.
[41]LIAO W,HARDER L D.Consequences of multiple inflorescences and clonality for pollinator behavior and plant mating[J].The American Naturalist,2014,184(5):580-592.
[42]ISAGI Y,SHIMADA K,KUSHIMA H,et al.Clonal structure and flowering traits of a bamboo[Phyllostachys pubescens(Mazel)Ohwi].stand grown from a simultaneous flowering as revealed by AFLP analysis[J].Molecular Ecology,2017,13(7):2017-2021.
[43]邱月群,董文渊,王逸之,等.筇竹天然居群遗传多样性研究[J].西北林学院学报,2017,32(2):155-160.QIU Y Q,DONG W Y,WANG Y Z,et al.Genetic diversity in natural populations of Qiongzhuea tumidinoda[J].Journal of Northwest Forestry University,2017,32(2):155-160.(in Chinese)
[44]BERTIN R I.Incidence of monoecy and dichogamy in relation to self-fertilization in angiosperms[J].American Journal of Botany,1993,80(5):557-560.
[45]CRUDEN R W.Temporal dioecism:systematic breadth,associated traits,and temporal patterns[J].Botanical Gazette,1988,149:1-15.
[46]ECKERT C G.Contributions of autogamy and geitonogamy to self-fertilization in a mass-flowering,clonal plant[J].Ecology,2000,81(2):532-542.