异质生境对二型花柱植物雪地报春花性状分化的影响
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摘要
该研究以生长在新疆北部的二型花柱植物雪地报春(Primula nivalis)为研究材料,对7个低海拔(1 657~2 013m)森林居群和7个高海拔(2 400~2 700m)草原居群的花型比例进行调查,对花部特征进行检测,并对长、短花柱植株在不同海拔居群的花性状分化进行了比较研究,以揭示该类群植物的演化与环境的关系。结果表明:(1)在高海拔草原居群中长花柱植株比例高于短花柱植株。(2)比较草原和森林生态系统中的雪地报春居群,长花柱花形态特征间存在显著差异,但短花柱花形态特征间的差异不显著。(3)高海拔草原居群中长花柱花的花冠筒长度、雄蕊长度、雌雄蕊距离、雌雄异位程度以及雌雄蕊空间位置是导致花性状分化的主要因素。(4)在草原居群中,长花柱花的雄蕊、雌雄异位程度及花冠口至雄蕊距离的变化减少了长短花柱花间的互补式对应性。研究认为,在异质生境中长花柱花性状的变化是导致该物种长短花柱花性状分化的主要因素。
Distylous plant is a special group of angiosperm which has reciprocal stigma and anthers within a species.Comparative investigations of breeding system of distylous species on different elevations can be revealing the relationship between distylous plant evolution and their environments.Primula nivalis is a native primrose in the northern Xinjiang,NW China.In order to investigate the comparative effect of habitat heterogeneity on frequency of morphs in different sexual morph plants and differentiation of floral characters were quantified at seven low-elevation forest(1 657-2 013 m)and seven high-elevation grassland populations in this research.The study results will clarify the reproductive strategies and adaptation mechanisms of P.nivalis at different elevational environments.(1)The investigations on effect of heterogeneous habitats on P.nivalis floral phenotype suggested that frequency of LS(long style)morph individuals in high elevation grassland population was higher than that of SS(short style)morph individuals;(2)Thefloral morphological characters of LS among populations in different ecological conditions were significant different,but no significant difference for SS morphs with different ecological conditions;(3)The length of corolla tube and stamen,and stamen-stigma spatial distance,herkogemy and relative reciprocity of LS flowers in high elevation grassland population are the main factors for floral trait differentiation in P.nivalis at heterogeneous environments;(4)This revealed that variation of LS floral characters at different population decreased the herkogamy level of both morph flower sexual organs in high elevation grassland populations.Therefore,changes of floral trait of LS plants are the main factors for floral trait differentiation and relative reciprocity of both morph flowers in heterogeneous environments.
Distylous plant is a special group of angiosperm which has reciprocal stigma and anthers within a species.Comparative investigations of breeding system of distylous species on different elevations can be revealing the relationship between distylous plant evolution and their environments.Primula nivalis is a native primrose in the northern Xinjiang,NW China.In order to investigate the comparative effect of habitat heterogeneity on frequency of morphs in different sexual morph plants and differentiation of floral characters were quantified at seven low-elevation forest(1 657-2 013 m)and seven high-elevation grassland populations in this research.The study results will clarify the reproductive strategies and adaptation mechanisms of P.nivalis at different elevational environments.(1)The investigations on effect of heterogeneous habitats on P.nivalis floral phenotype suggested that frequency of LS(long style)morph individuals in high elevation grassland population was higher than that of SS(short style)morph individuals;(2)Thefloral morphological characters of LS among populations in different ecological conditions were significant different,but no significant difference for SS morphs with different ecological conditions;(3)The length of corolla tube and stamen,and stamen-stigma spatial distance,herkogemy and relative reciprocity of LS flowers in high elevation grassland population are the main factors for floral trait differentiation in P.nivalis at heterogeneous environments;(4)This revealed that variation of LS floral characters at different population decreased the herkogamy level of both morph flower sexual organs in high elevation grassland populations.Therefore,changes of floral trait of LS plants are the main factors for floral trait differentiation and relative reciprocity of both morph flowers in heterogeneous environments.
引文
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[14]DE VOS J M,KELLER B,ISHAM S T,et al.Reproductive implications of herkogamy in homostylous primroses:variation during anthesis and reproductive assurance in alpine environments[J].Functional Ecology,2012,26:854-865.
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[17]HERRERA C M,CASTELLANOS M C,MEDRANO M.Geographical context of floral evolution:Towards an improved research programme in floral diversification[M]//HARDER L D,BARRETT S C H.Ecology and Evolution of Flowers.UK.Oxford University Press.2006:278-294.
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[19]PAILLER T,THOMPSON J D.Distyly and variation in heteromorphic incompatibility in Gaertnera vaginata(Rubiaceae)endemic to La Réunion Island[J].American Journal of Botany,1997,84:315-327.
[20]ZHAO Z G,WANG Y K.Selection by pollinators on floral traits in generalized Trollius ranunculoides(Ranunculaceae)along altitudinal gradients[J].PLoS One.2015,10:e0118299.doi:10.1371/journal.pone.0118299,
[21]BRYS R,JACQUEMYN H.Disruption of the distylous syndrome in Primula veris[J].Annals of Botany,2015,115:27-39.
[22]DE VOS J M,HUGHES C E,SCHNEEWEISS G M,et al.Heterostyly accelerates diversification via reduced extinction in primroses[J].Proceedings of the Royal Society B:Biological Sciences,2014,281:20140075.
[23]KELLER B,DE VOS J M,CONTI E.Decrease of sexual organ reciprocity between heterostylous primrose species,with possible functional and evolutionary implications[J].Annals of Botany,2012,110:1 233-1 244.
[24]CONTRERAS P S,ORMELAS J F.Reproductive conflicts of Palicourea padifolia(Rubiaceae)a distylous shrub of a tropical cloud forest in Mexico[J].Plant Systematics and Evolution,1999,219:225-241.
[25]LIU S J,WU L Y,HUANG S Q.Shortened anther-stigma distance reduces compatible pollination in two distylous Primulaspecies[J].Journal of Plant Ecology,2015,doi:10.1093/jpe/rtv049
[26]HARGREAVES A L,WEINER J L,ECKERT C G.Highelevation range limit of an annual herb is neither caused nor reinforced by declining pollinator service[J].Journal of Ecology,2015,103:572-584.
[27]MCKEE J,RICHARDS A J.The effect of temperature on reproduction in five Primulaspecies[J].Annals of Botany,1998,82:359-374.
[2]BARRETT S C H.Evolution and Function of Heterostyly[M].Berlin:Springer-Verlag,1992.
[3]BARRETT S C H,JESSON L,BAKER A M.The evolution and function of stylar polymorphisms in flowering plants[J].Annals of Botany,2000,85:253-265.
[4]SANCHEZ J M,FERRERO V,ARROYO J,et al.Patterns of style polymorphism in five species of the south African genus Nivenia(Iridaceae)[J].Annals of Botany,2010,106:321-331.
[5]RODRIGUES E B,CONSOLARO H.Atypical distyly in Psychotria goyazensis Mull.Arg.(Rubiaceae),an intramorph self-compatible species[J].Acta Botanica Brasilica,2013,27:155-161.
[6]ARROYO J,BARRETT S C H.Discovery of distyly in Narcissus(Amaryllidaceae)[J].American Journal of Botany,2000,87:748-751.
[7]BARRETT S C H.Mating strategies in flowering plants:the outcrossing-selfing paradigm and beyond[J].The Royal Society of London Series B Biological Sciences,2002,358:991-1 004.
[8]ARROYO M T K,PACHECO D A,AGUILERA P.Floral allocation at different altitudes in highly autogamous alpine Chaetanthera euphrasioides(Asteraceae)in the central Chilean Andes[J].Alpine Botany,2013,123:7-12.
[9]BARRETT S C H,RICHARDS J H.Heterostyly in tropical plants[M]//PRANCE G,GOTTSBERGER G.Reproductive Systems and Speciation in Tropical Woody Plants.Memoirs of the New York Botanical Garden,1990,55:35-61.
[10]HADDADCHI A.Stylar polymorphism,reciprocity and incompatibility systems in Nymphoides montana(Menyanthaceae)endemic to southeastern Australia[J].Plant Systematics and Evolution,2013,299:389-401.
[11]LLOYD D G.Evolution towards dioecy in heterostylous populations[J].Plant Systematics and Evolution,1979,31:71-80.
[12]PAILLER T,HUMEAU L,THOMESON J D.Distyly and heteromorphic incompatibility in oceanic island species of Erythroxylum(Erythroxylaceae)[J].Plant Systematics and Evolution,1998,213:187-198.
[13]BARRETT S C H,HARDER L D.The evolution of polymorphic sexual systems in daffodils(Narcissus)[J].New Phytologist,2004,165:45-53.
[14]DE VOS J M,KELLER B,ISHAM S T,et al.Reproductive implications of herkogamy in homostylous primroses:variation during anthesis and reproductive assurance in alpine environments[J].Functional Ecology,2012,26:854-865.
[15]GROSSENBACHER D,RUNQUIST R B,GOLDBERG E E,et al.Geographic range size is predicted by plant mating system[J].Ecology Letters,2015,18:706-713.
[16]SAMPSON D A,KREBS R A.Quantitative evaluation of reciprocal herkogamy in the distylous species,Hedyotis caerulea(Rubiaceae)[J].Plant Systematics and Evolution,2012,298:1 361-1 370.
[17]HERRERA C M,CASTELLANOS M C,MEDRANO M.Geographical context of floral evolution:Towards an improved research programme in floral diversification[M]//HARDER L D,BARRETT S C H.Ecology and Evolution of Flowers.UK.Oxford University Press.2006:278-294.
[18]THOMPSON J N.The Geographic Mosaic of Coevolution[M].Chicago University of Chicago Press,2005.
[19]PAILLER T,THOMPSON J D.Distyly and variation in heteromorphic incompatibility in Gaertnera vaginata(Rubiaceae)endemic to La Réunion Island[J].American Journal of Botany,1997,84:315-327.
[20]ZHAO Z G,WANG Y K.Selection by pollinators on floral traits in generalized Trollius ranunculoides(Ranunculaceae)along altitudinal gradients[J].PLoS One.2015,10:e0118299.doi:10.1371/journal.pone.0118299,
[21]BRYS R,JACQUEMYN H.Disruption of the distylous syndrome in Primula veris[J].Annals of Botany,2015,115:27-39.
[22]DE VOS J M,HUGHES C E,SCHNEEWEISS G M,et al.Heterostyly accelerates diversification via reduced extinction in primroses[J].Proceedings of the Royal Society B:Biological Sciences,2014,281:20140075.
[23]KELLER B,DE VOS J M,CONTI E.Decrease of sexual organ reciprocity between heterostylous primrose species,with possible functional and evolutionary implications[J].Annals of Botany,2012,110:1 233-1 244.
[24]CONTRERAS P S,ORMELAS J F.Reproductive conflicts of Palicourea padifolia(Rubiaceae)a distylous shrub of a tropical cloud forest in Mexico[J].Plant Systematics and Evolution,1999,219:225-241.
[25]LIU S J,WU L Y,HUANG S Q.Shortened anther-stigma distance reduces compatible pollination in two distylous Primulaspecies[J].Journal of Plant Ecology,2015,doi:10.1093/jpe/rtv049
[26]HARGREAVES A L,WEINER J L,ECKERT C G.Highelevation range limit of an annual herb is neither caused nor reinforced by declining pollinator service[J].Journal of Ecology,2015,103:572-584.
[27]MCKEE J,RICHARDS A J.The effect of temperature on reproduction in five Primulaspecies[J].Annals of Botany,1998,82:359-374.