“金冠霞帔”文冠果种实性状可塑性
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摘要
表型可塑性是有机体适应异质环境,增强生存能力的重要机制。以文冠果新品种"金冠霞帔"为研究对象,于2016年在不同纬度以及同一纬度两种不同土壤类型3个区域4个试验地,选取无性嫁接扩繁、树体结构相近的植株,开展控制授粉与开放授粉对比实验,并通过无性嫁接植株之间的控制授粉实现了文冠果的自交。果实成熟后采收并风干,测量果实质量、果壳质量、结果数、座果率、种子粒数、种子质量、出籽率、单粒重等12个性状,筛选了3个表型可塑性指标,并以表型可塑性指数PPI进行了综合评价。结果表明:随北京、辽宁、内蒙古的纬度变化以及褐土到风沙土的生境变化,种实性状均表现出可塑性。随种植纬度的增加,植株增加对果实种子的资源性投入,果实体积变大且数量增加,果实质量、果壳质量、种子质量、种子粒数均显著增加。从相对肥沃褐土到相对贫瘠风沙土的生境变化,种实性状表现出"趋贫特化,优先后代"的响应机制,即资源相对匮乏时,植株将有限资源分配给少量果实,减少果壳资源分配,相对多的资源分配给少量种子,种子大粒化的方式适应贫瘠生境,这种"繁殖策略"支持了文冠果具有分布广泛、适应性强、抗逆性强等特性。本研究为文冠果适应与进化研究提供基础资料,并对其栽培管理和保育利用提供重要参考。
Phenotypic plasticity is an important mechanism for the adaptation of organisms to heterogeneous environments and for the enhancement of their survival ability. Trees of new variety"Jinguanxiapei"of Xanthoceras sorbifolium Bunge with similar structure were selected for this experiment in 2016,which consisted of a replicated diallel mating of controlled pollination and open pollination in four sites of three regions. Self-pollination was achieved by grafting individuals pollinated each other. Four experimental sites located in three regions of Beijing,Liaoning and Inner Mongolia,consisting of three different latitudes and two different soil types in one of the three regions. After harvesting and drying the fruits,we measured 12 traits including fruit quality,shell quality,number of fruits,fruit rate,seed number,seed quality,seed yield,and single grain weight. Three phenotypic plasticity indicators were evaluated and finally the phenotypic plasticity index( PPI) was used to estimate the plasticity of each trait. The results showed that seed and fruit traits exhibited phenotypic plasticity across different latitudes and different soil conditions. With increasing latitude,the seed and fruit traits showed a positive response by increasing resource input to the fruits and seeds,showing increases in fruit volume,fruit number,fruit mass,fruit shell mass,seed mass and seed number. With the change of habitat from cinnamon soil to aeolian sandy soil,seed and fruit traits exhibited the response mechanism of specialization for scarce resource and priority of resource allocation to progeny. When the resources are relatively scarce,plants allocate the limited resources to a small amount of fruit,reduce the allocation of fruit shell resources,and allocate more resources to a small amount of seeds,so that the seeds are bigger to adapt to the environment of aeolian sandy soil. Such a reproductive strategy makes the species has wide distribution,strong adaptability,and strong resistance. Our results provide basic data for study of adaptation and evolution and scientific reference for breeding and cultivation of X. sorbifolium.
Phenotypic plasticity is an important mechanism for the adaptation of organisms to heterogeneous environments and for the enhancement of their survival ability. Trees of new variety"Jinguanxiapei"of Xanthoceras sorbifolium Bunge with similar structure were selected for this experiment in 2016,which consisted of a replicated diallel mating of controlled pollination and open pollination in four sites of three regions. Self-pollination was achieved by grafting individuals pollinated each other. Four experimental sites located in three regions of Beijing,Liaoning and Inner Mongolia,consisting of three different latitudes and two different soil types in one of the three regions. After harvesting and drying the fruits,we measured 12 traits including fruit quality,shell quality,number of fruits,fruit rate,seed number,seed quality,seed yield,and single grain weight. Three phenotypic plasticity indicators were evaluated and finally the phenotypic plasticity index( PPI) was used to estimate the plasticity of each trait. The results showed that seed and fruit traits exhibited phenotypic plasticity across different latitudes and different soil conditions. With increasing latitude,the seed and fruit traits showed a positive response by increasing resource input to the fruits and seeds,showing increases in fruit volume,fruit number,fruit mass,fruit shell mass,seed mass and seed number. With the change of habitat from cinnamon soil to aeolian sandy soil,seed and fruit traits exhibited the response mechanism of specialization for scarce resource and priority of resource allocation to progeny. When the resources are relatively scarce,plants allocate the limited resources to a small amount of fruit,reduce the allocation of fruit shell resources,and allocate more resources to a small amount of seeds,so that the seeds are bigger to adapt to the environment of aeolian sandy soil. Such a reproductive strategy makes the species has wide distribution,strong adaptability,and strong resistance. Our results provide basic data for study of adaptation and evolution and scientific reference for breeding and cultivation of X. sorbifolium.
引文
敖妍,韩墨,赵磊磊,等.2015.主要分布区文冠果类型的划分.西北林学院学报,30(3):100-106.
毕泉鑫,蔡龙,马兴华,等.2011.中国特有能源植物文冠果的遗传学及产业化.中国野生植物资源,30(5):37-41.
毕泉鑫.2015.基于形态性状和SSR标记的文冠果交配组合优化研究(博士学位论文).北京:北京林业大学.
柴春山,芦娟,蔡国军,等.2013.文冠果人工种群的果实表型多样性及其变异.林业科学研究,26(2):181-191.
杜乐山.2014.加拿大一枝黄花的表型可塑性与适应性研究(硕士学位论文).北京:北京林业大学.
耿宇鹏,张文驹,李博,等.2004.表型可塑性与外来植物的入侵能力.生物多样性,12(4):447-455.
耿占礼.2012.红花文冠果嫁接繁育技术.水土保持应用技术,21(1):25-26.
侯元凯,黄琳,高巍,等.2013.不同种源文冠果果实及种子表型性状变异的研究.中南林业科技大学学报,33(7):20-24.
侯元凯,黄琳,杨超伟,等.2011.文冠果种子性状与果实性状的相关性研究.中南林业科技大学学报,31(9):24-27.
牟洪香,侯新村,刘巧哲.2007.木本能源植物文冠果的表型多样性研究.林业科学研究,20(3):350-355.
汪子洋,黄炎子,王俊杰,等.2017.文冠果不同果型种实性状的概率分布与选择.北京林业大学学报,39(9):17-31.
王姝,周道玮.2017.植物表型可塑性研究进展.生态学报,37(24):8161-8169.
王涛.2012.中国能源植物文冠果的研究.北京:中国科学技术出版社.
魏胜利,王文全,王继永,等.2011.甘草野生种群果实与种子形态特征的地理变异研究.现代生物医学进展,11(23):4460-4464.
吴征镒,孙航,周浙昆,等.2005.中国植物区系中的特有性及其起源和分化.云南植物研究,27(6):577-601.
徐东翔.2010.文冠果生物学.北京:科学出版社.
杨贺雨.2016.华中五味子表型可塑性研究(硕士学位论文).西安:陕西师范大学.
杨雨春,赵佳宁,张忠辉,等.2011.文冠果不同群体果实和种子性状综合评价研究.中国农学通报,27(16):36-40.
杨韫嘉.2014.文冠果种实性状变异与根孽幼苗可塑性分析(硕士学位论文).北京:北京林业大学.
张萍,王连军,彭华.2012.文冠果优良类型选择.中国林副特产,117(2):10-12.
张维,巴雅尔塔,杨晓绒,等.2014.新疆野核桃种子的生物量可塑性及其生长模型.生态科学,33(5):991-997.
周浩然,徐明,于秀波,等.2014.生物适应性和表型可塑性理论研究进展.西北农林科技大学学报:自然科学版,42(4):215-220.
Abraham EM,Kyriazopoulos AP,Parissi ZM,et al.2014.Growth,dry matter production,phenotypic plasticity,and nutritive value of three natural populations of Dactylis glomerata L.under various shading treatments.Agroforestry Systems,88:287-299.
Bloom AJ,Chapin FS,Mooney HA.1985.Resource limitation in plants:An economic analogy.Annual Review of Ecology&Systematics,16:363-392.
Bonser SP,Aarssen LW.2001.Allometry and plasticity of meristem allocation throughout development in Arabidopsis thaliana.Journal of Ecology,89:72-79.
Bradshaw AD.1965.Evolutionary significance of phenotypic plasticity in plants.Advances in Genetics,13:115-155.
Canli FA,Pektas M.2015.Improving fruit size and quality of low yielding and small fruited pear cultivars with benzyladenine and gibberellin applications.European Journal of Horticultural Science,80:103-108.
El-Kassaby YA,Cook C.1994.Female reproductive energy and reproductive success in a Douglas-fir seed orchard and its impact on genetic diversity.Silvae Genetica,43:243-246.
Kurashige NS,Agrawal AA.2005.Phenotypic plasticity to light competition and herbivory in Chenopodium album(Chenopodiaceae).American Journal of Botany,92:21-26.
Leishman MR,Westoby M.1992.Classifying plants into groups on the basis of associations of individual traits evidence from Australian semi-arid woodlands.Journal of Ecology,80:417-424.
Quero JL,Villar R,Mara1ón T,et al.2006.Interactions of drought and shade effects on seedlings of four Quercus species:Physiological and structural leaf responses.New Phytologist,170:819-833.
Rapport DJ,Turner JE.1977.Economic Models in Ecology.Science,195:367-373.
Rejmanek M,Richardson DM.1996.What attributes make some plant species more invasive?Ecology,77:1655-1661.
Scheiner SM.1993.Genetics and evolution of phenotypic plasticity.Annual Review of Ecology and Systematics,24:35-68.
Schlichting CD,Levin DA.1986.Phenotypic plasticity:An evolving plant character.Biological Journal of the Linnean Society,29:37-47.
Stajnko D,Rakun J,Blanke M.2009.Modelling apple fruit yield using image analysis for fruit colour,shape and texture.European Journal of Horticultural Science,74:260-267.
Sultan SE.2000.Phenotypic plasticity for plant development,function and life history.Trends in Plant Science,5:537-542.
Sultan SE.2010.Plant developmental responses to the environment:Eco-devo insights.Current Opinion in Plant Biology,13:96-101.
Sultan SE.2013.Phenotypic plasticity and plant adaptation.Plant Biology,44:363-383.
Valladares F,Gianoli E,Gómez JM.2007.Ecological limits to plant phenotypic plasticity.New Phytologist,176:749-763.
Valladares F,Sanchez-Gomez D,Zavala MA.2006.Quantitative estimation of phenotypic plasticity:Bridging the gap between the evolutionary concept and its ecological applications.Journal of Ecology,94:1103-1116.
Valladares F,Wright SJ,Lasso E,et al.2000.Plastic phenotypic response to light of 16 congeneric shrubs from a Panamanian rainforest.Ecology,81:1925-1936.
Wang Q,Huang Y,Wang Z,et al.2017.Fruit shape and reproductive self and cross compatibility for the performance of fruit set in an andromonoecious species:Xanthoceras sorbifolium Bunge.Tree Genetics&Genomes,13:116.
Westeberhard MJ.1989.Phenotypic plasticity and the origins of diversity.Annual Review of Ecology&Systematics,20:249-278.
毕泉鑫,蔡龙,马兴华,等.2011.中国特有能源植物文冠果的遗传学及产业化.中国野生植物资源,30(5):37-41.
毕泉鑫.2015.基于形态性状和SSR标记的文冠果交配组合优化研究(博士学位论文).北京:北京林业大学.
柴春山,芦娟,蔡国军,等.2013.文冠果人工种群的果实表型多样性及其变异.林业科学研究,26(2):181-191.
杜乐山.2014.加拿大一枝黄花的表型可塑性与适应性研究(硕士学位论文).北京:北京林业大学.
耿宇鹏,张文驹,李博,等.2004.表型可塑性与外来植物的入侵能力.生物多样性,12(4):447-455.
耿占礼.2012.红花文冠果嫁接繁育技术.水土保持应用技术,21(1):25-26.
侯元凯,黄琳,高巍,等.2013.不同种源文冠果果实及种子表型性状变异的研究.中南林业科技大学学报,33(7):20-24.
侯元凯,黄琳,杨超伟,等.2011.文冠果种子性状与果实性状的相关性研究.中南林业科技大学学报,31(9):24-27.
牟洪香,侯新村,刘巧哲.2007.木本能源植物文冠果的表型多样性研究.林业科学研究,20(3):350-355.
汪子洋,黄炎子,王俊杰,等.2017.文冠果不同果型种实性状的概率分布与选择.北京林业大学学报,39(9):17-31.
王姝,周道玮.2017.植物表型可塑性研究进展.生态学报,37(24):8161-8169.
王涛.2012.中国能源植物文冠果的研究.北京:中国科学技术出版社.
魏胜利,王文全,王继永,等.2011.甘草野生种群果实与种子形态特征的地理变异研究.现代生物医学进展,11(23):4460-4464.
吴征镒,孙航,周浙昆,等.2005.中国植物区系中的特有性及其起源和分化.云南植物研究,27(6):577-601.
徐东翔.2010.文冠果生物学.北京:科学出版社.
杨贺雨.2016.华中五味子表型可塑性研究(硕士学位论文).西安:陕西师范大学.
杨雨春,赵佳宁,张忠辉,等.2011.文冠果不同群体果实和种子性状综合评价研究.中国农学通报,27(16):36-40.
杨韫嘉.2014.文冠果种实性状变异与根孽幼苗可塑性分析(硕士学位论文).北京:北京林业大学.
张萍,王连军,彭华.2012.文冠果优良类型选择.中国林副特产,117(2):10-12.
张维,巴雅尔塔,杨晓绒,等.2014.新疆野核桃种子的生物量可塑性及其生长模型.生态科学,33(5):991-997.
周浩然,徐明,于秀波,等.2014.生物适应性和表型可塑性理论研究进展.西北农林科技大学学报:自然科学版,42(4):215-220.
Abraham EM,Kyriazopoulos AP,Parissi ZM,et al.2014.Growth,dry matter production,phenotypic plasticity,and nutritive value of three natural populations of Dactylis glomerata L.under various shading treatments.Agroforestry Systems,88:287-299.
Bloom AJ,Chapin FS,Mooney HA.1985.Resource limitation in plants:An economic analogy.Annual Review of Ecology&Systematics,16:363-392.
Bonser SP,Aarssen LW.2001.Allometry and plasticity of meristem allocation throughout development in Arabidopsis thaliana.Journal of Ecology,89:72-79.
Bradshaw AD.1965.Evolutionary significance of phenotypic plasticity in plants.Advances in Genetics,13:115-155.
Canli FA,Pektas M.2015.Improving fruit size and quality of low yielding and small fruited pear cultivars with benzyladenine and gibberellin applications.European Journal of Horticultural Science,80:103-108.
El-Kassaby YA,Cook C.1994.Female reproductive energy and reproductive success in a Douglas-fir seed orchard and its impact on genetic diversity.Silvae Genetica,43:243-246.
Kurashige NS,Agrawal AA.2005.Phenotypic plasticity to light competition and herbivory in Chenopodium album(Chenopodiaceae).American Journal of Botany,92:21-26.
Leishman MR,Westoby M.1992.Classifying plants into groups on the basis of associations of individual traits evidence from Australian semi-arid woodlands.Journal of Ecology,80:417-424.
Quero JL,Villar R,Mara1ón T,et al.2006.Interactions of drought and shade effects on seedlings of four Quercus species:Physiological and structural leaf responses.New Phytologist,170:819-833.
Rapport DJ,Turner JE.1977.Economic Models in Ecology.Science,195:367-373.
Rejmanek M,Richardson DM.1996.What attributes make some plant species more invasive?Ecology,77:1655-1661.
Scheiner SM.1993.Genetics and evolution of phenotypic plasticity.Annual Review of Ecology and Systematics,24:35-68.
Schlichting CD,Levin DA.1986.Phenotypic plasticity:An evolving plant character.Biological Journal of the Linnean Society,29:37-47.
Stajnko D,Rakun J,Blanke M.2009.Modelling apple fruit yield using image analysis for fruit colour,shape and texture.European Journal of Horticultural Science,74:260-267.
Sultan SE.2000.Phenotypic plasticity for plant development,function and life history.Trends in Plant Science,5:537-542.
Sultan SE.2010.Plant developmental responses to the environment:Eco-devo insights.Current Opinion in Plant Biology,13:96-101.
Sultan SE.2013.Phenotypic plasticity and plant adaptation.Plant Biology,44:363-383.
Valladares F,Gianoli E,Gómez JM.2007.Ecological limits to plant phenotypic plasticity.New Phytologist,176:749-763.
Valladares F,Sanchez-Gomez D,Zavala MA.2006.Quantitative estimation of phenotypic plasticity:Bridging the gap between the evolutionary concept and its ecological applications.Journal of Ecology,94:1103-1116.
Valladares F,Wright SJ,Lasso E,et al.2000.Plastic phenotypic response to light of 16 congeneric shrubs from a Panamanian rainforest.Ecology,81:1925-1936.
Wang Q,Huang Y,Wang Z,et al.2017.Fruit shape and reproductive self and cross compatibility for the performance of fruit set in an andromonoecious species:Xanthoceras sorbifolium Bunge.Tree Genetics&Genomes,13:116.
Westeberhard MJ.1989.Phenotypic plasticity and the origins of diversity.Annual Review of Ecology&Systematics,20:249-278.