丹红杨×通辽1号杨杂交子代苗期抗旱性初步评价
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摘要
杨树是重要的用材林和防护林树种,土壤干旱不仅影响杨树正常生长发育,甚至降低木材产量与品质。评价杨树全同胞家系抗旱性,筛选抗旱指标,对培育抗旱新品种、干旱土地的利用及生态环境建设具有重要意义。本研究以123个丹红杨×通辽1号杨杂交子代无性系为材料,采用盆栽控水试验,在中度干旱胁迫下,测定植株的生长、光合、叶形和水分生理等12个指标;建立田间试验林,测定一年生田间生长量。结果表明,干旱胁迫对杂交子代各指标均有极显著影响(P<0.01),其中气孔导度(Ga)变异系数最大。相关性分析发现,各抗旱性状之间呈现不同程度的相关性。利用主成分分析法和隶属函数法进行苗期抗旱性综合评价,将杂交群体划分为干旱敏感型、中等抗旱型和高抗旱型3种类型,分别包含30个、73个和22个无性系,DX-27、DX-4、DX-3等10个无性系抗旱性强于父本通辽1号杨,其中无性系DX-55和DX-81抗旱性D值(均为0.60)最大,抗旱能力最强。逐步线性回归分析和相关性分析表明基径(D)、净光合速率(Pn)、气孔导度(Ga)和蒸腾速率(Tr)对抗旱性影响显著,可作为苗期抗旱性评价指标,其中Ga对抗旱性直接效应最大。采用布雷金多性状综合评价法对杂交子代田间生长性状综合评价,将杂交群体划分为4级,亲本丹红杨和通辽1号杨分别隶属优级和差级,6个抗旱性强于父本通辽1号杨的无性系(DX-76、DX-78、DX-53、DX-55、DX-3、DX-20)隶属优级和良级,初步筛选为抗旱速生材料。本研究为杨树抗旱遗传改良及苗期抗旱性鉴定提供理论依据。
Poplar is an important timber and shelterbelt tree species.The soil drought severely limits its growth,development and quality.Deciphering the theory and production for drought resistance is of interest on the poplar varieties cultivation,arid land use,and ecological environment construction to evaluate poplar full sib family drought resistance and select drought index.In this study,123 progenies derived from a cross of Populus deltoides‘Danhong'×P. simonii ‘Tongliao1' were subjected for moderate pot water control experiment via determining 12 growth parameters such as photosynthesis,leaf and water indexes.In addition,we established field experimental forest and measured one year growth increment.The results showed that the drought stress had a significant effect on all the tested traits of hybrid progeny(P < 0.01),among which stomatal conductance(Ga)had the largest variation coefficient.Correlation analysis indicated that responsive traits exhibited different degrees of correlation.Drought resistances of the progenies at seedling stage were evaluated with principle component analysis and subordinate function.The hybrid population were classified into three types as drought sensitive type,moderate drought resistance type and high drought resistance type with each contained 30,73,and 22 individuals.DX-27,DX-4,DX-3 and other seven lines adopt to drought better than male parent P.simonii‘Tongliao1'.Line DX-55 and DX-81 got the maximum drought resistance D value(all 0.60),and showed the strongest drought resistance.Stepwise linear regression and correlation analysis indicated diameter(D),net photosynthesis(Pn),stomatal conductance(Ga)and transpiration rate(Tr)could be used as drought resistance indexes,of which Ga had the largest direct effect on drought resistance.The comprehensive evaluation was used to analyze field growth characters,and the progenies were divided into four classes. P.deltoides ‘Danhong' and P.simonii ‘Tongliao1' belong to the first and fourth class,respectively.Six lines(DX-76,DX-78,DX-53,DX-55,DX-3,DX-20)with better drought resistance ability than male parent P.simonii ‘Tongliao1' belong to the first and second class,and preliminarily selected as drought-resistant and fast-growing materials.Thus,this study provides theoretical basis in breeding for drought resistance in poplar and test for drought resistance at the seedlingstage.
Poplar is an important timber and shelterbelt tree species.The soil drought severely limits its growth,development and quality.Deciphering the theory and production for drought resistance is of interest on the poplar varieties cultivation,arid land use,and ecological environment construction to evaluate poplar full sib family drought resistance and select drought index.In this study,123 progenies derived from a cross of Populus deltoides‘Danhong'×P. simonii ‘Tongliao1' were subjected for moderate pot water control experiment via determining 12 growth parameters such as photosynthesis,leaf and water indexes.In addition,we established field experimental forest and measured one year growth increment.The results showed that the drought stress had a significant effect on all the tested traits of hybrid progeny(P < 0.01),among which stomatal conductance(Ga)had the largest variation coefficient.Correlation analysis indicated that responsive traits exhibited different degrees of correlation.Drought resistances of the progenies at seedling stage were evaluated with principle component analysis and subordinate function.The hybrid population were classified into three types as drought sensitive type,moderate drought resistance type and high drought resistance type with each contained 30,73,and 22 individuals.DX-27,DX-4,DX-3 and other seven lines adopt to drought better than male parent P.simonii‘Tongliao1'.Line DX-55 and DX-81 got the maximum drought resistance D value(all 0.60),and showed the strongest drought resistance.Stepwise linear regression and correlation analysis indicated diameter(D),net photosynthesis(Pn),stomatal conductance(Ga)and transpiration rate(Tr)could be used as drought resistance indexes,of which Ga had the largest direct effect on drought resistance.The comprehensive evaluation was used to analyze field growth characters,and the progenies were divided into four classes. P.deltoides ‘Danhong' and P.simonii ‘Tongliao1' belong to the first and fourth class,respectively.Six lines(DX-76,DX-78,DX-53,DX-55,DX-3,DX-20)with better drought resistance ability than male parent P.simonii ‘Tongliao1' belong to the first and second class,and preliminarily selected as drought-resistant and fast-growing materials.Thus,this study provides theoretical basis in breeding for drought resistance in poplar and test for drought resistance at the seedlingstage.
引文
[1]贾黎明,刘诗琦,祝令辉,胡建军,王小平.我国杨树林的碳储量和碳密度.南京林业大学学报:自然科学版,2013,37(2):1-7Jia L M,Liu S Q,Zhu L H,Hu J J,Wang X P.Carbon storage and density of poplars in China.Journal of Nanjing Forestry University:Natural Sciences Edition,2013,37(2):1-7
[2]王宇超,王得祥,彭少兵,何帆.干旱胁迫对木本滨藜生理特性的影响.林业科学,2010,46(1):61-67Wang Y C,Wang D X,Peng S B,He F.Effects of drought stress on physiological characteristics of woody saltbush.Scientia Silvae Sinicae,2010,46(1):61-67
[3]Dai A.Increasing drought under global warming in observations and models.Nature Climate Change,2012,3:52-58
[4]Allen C D,Macalady A K,Chenchouni H,Bachelet D,Mcdowell N G,Vennetier M,Kitzberger T,Rigling A,Breshears D D,Hogg E H,Gonzalez P,Fensham R J,Zhang Z,Castro J,Demidova N,Lim J H,Allard G,Running S W.Aglobal overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests.Forest Ecology Management,2010,259:660-684
[5]杨建伟,梁宗锁,韩蕊莲,孙群,崔浪军.不同干旱土壤条件下杨树的耗水规律及水分利用效率研究.植物生态学报,2004,28(5):657-664Yang J W,Liang Z S,Han R L,Sun Q,Cui L J.Water use efficiency and water consumption characterics of poplar under soil drought conditions.Acta Phytoecological Sinica,2004,28(5):657-664
[6]Mazzoleni S,Dickmann D I.Differential physiological and morphological responses of two hybrid Populus clones to water stress.Tree Physiology,1988,4:61-70
[7]Souch C A,Stephens W.Growth,productivity and water use in three hybrid poplar clones.Tree Physiology,1998,18:829-835
[8]付士磊,周永斌,何兴元,陈玮.干旱胁迫对杨树光合生理指标的影响.应用生态学报,2006,17(11):2016-2019Fu S L,Zhou Y B,He X Y,Chen W.Effects of drought stress on photosynthesis physiology of Populus pseudo-simonii.Chinese Journal of Applied Ecology,2006,17(11):2016-2019
[9]Viger M,Smith H K,Cohen D,Dewoody J,Trewin H,Steenackers M,Bastien C,Taylor G.Adaptive mechanisms and genomic plasticity for drought tolerance identified in European black poplar(Populus nigra L.).Tree Physiology,2016,36:909
[10]杨淑红,宋德才,刘艳萍,杨清淮,朱延林.土壤干旱胁迫和复水后3个杨树品种叶片部分生理指标变化及抗旱性评价.植物资源与环境学报,2014,23(3):65-73Yang S H,Song D C,Liu Y P,Yang Q H,Zhu Y L.Changes of some physiological indexes in leaf of three cultivars of Populus after drought stress in soil and re-watering and evaluation on their drought resistance.Journal of Plant Resources and Environment,2014,23(3):65-73
[11]邱兴,樊军锋,李晓东,吕小锋.6个杨树新无性系叶片旱生结构研究.西北林学院学报,2015,30(1):96-101Qiu X,Fan J F,Li X D,Lv X F.Research on drought resistance on leaf anatomical structure of six new poplar clones.Journal of Northwest Forestry University,2015,30(1):96-101
[12]Loutfy N,Eltayeb M A,Hassanen A M,Moustafa M FM,Sakuma Y,Inouhe M.Changes in the water status and osmotic solute contents in response to drought and salicylic acid treatments in four different cultivars of wheat(Triticum aestivum).Journal of Plant Research,2012,125:173-184
[13]Gower S T,Kucharik C J,Norman J M.Direct and indirect estimation of leaf area index,f APAR,and net primary production of Terrestrial Ecosystems.Remote Sensing Environment,1999,70:29-51
[14]Nicolas M,Erwin D,Boudouresque E,Delay D,Petit J,Delmotte F,Brignolas F.Impact of successive drought and re-watering cycles on growth and specific leaf area of two Populus×canadensis(Moench)clones,‘Dorskamp’and‘Luisa-Avanzo’.Tree Physiology,2003,23:1225-1235
[15]Yang Y,Yao Y,Xu G,Li C Y.Growth and physiological responses to drought and elevated ultraviolet-b in two contrasting populations of Hippophae rhamnoides.Physiologia Plantarum,2010,124:431-440
[16]李忠武,蔡强国,唐政洪,吴淑安,王宏.作物生产力模型及其应用研究.应用生态学报,2002,13(9):1174-1178L i Z W,C a i G Q,Ta n g Z H,Wu S A,Wa n g H.C r o p productivity model and its application.Chinese Journal of Applied Ecology,2002,13(9):1174-1178
[17]姬慧娟,贾会霞,章小铃,胡建军.干旱胁迫对红皮柳光合特性日变化及生长的影响.南京林业大学学报:自然科学版,2016,40(6):41-46Ji H J,Jia H X,Zhang X L,Hu J J.Effect of drought stress on photosynthetic diurnal course and growth of Salix purpurea.Journal of Nanjing Forestry University:Natural Sciences Edition,2016,40(6):41-46
[18]张春玲,李淑梅,赵自成,胡建军,韩一凡.杨树新品种‘丹红杨’.林业科学,2008,44(1):169Zhang C L,Li S M,Zhao Z C,Hu J J,Han Y F.A new poplar variety Populus deltoides CL.‘Dahong’.Scientia Silvae Sinicae,2008,44(1):169
[19]Chen J H,Song Y P,Zhang H,Zhang D Q.Genome wide analysis of gene expression in response to drought stress in Populus simonii.Plant Molecular Biology Reporter,2013,31:946-962
[20]杜森,高祥照.土壤分析技术规范.北京:中国农业出版社,2006:47-49Du S,Gao X Z.Technical specification for soil analysis.Beijing:China Agricultural Publishers,2006:47-49
[21]曾辉,曹苑南,王述民,王兰芬,董海鸿,陈吉宝.30份普通菜豆苗期抗旱性鉴定及抗旱指标的确定.植物遗传资源学报,2016,17(6):1014-1021Zeng H,Cao Y N,Wang S M,Wang L F,Dong H H,Chen J B.Evaluation of drought tolerance of 30 common bean in seedling stage and screening of drought-tolerance assessment index.Journal of Plant Genetic Resource,2016,17(6):1014-1021
[22]胡标林,杨平,万勇,李霞,罗世友,罗向东,谢建坤.东乡野生稻BILs群体苗期抗旱性综合评价及其遗传分析.植物遗传资源学报,2013,14(2):249-256Hu B L,Yang P,Wan Y,Li X,Luo S Y,Luo X D,Xie JK.Comprehensive assessment of drought resistance of BILs population derived from Dongxiang wild rice(Oryza rufipogon Griff)at seedling stage and its genetic analysis.Journal of Plant Genetic Resources,2013,14(2):249-256
[23]谢季坚.农业科学中的模糊数学方法.武汉:华中理工大学出版社,1993:99-193Xie J J.Fuzzy mathematics in agricultural science.Wuhan:Huazhong university of science and technology publishing house,1993:99-193
[24]周永学,苏晓华,樊军锋,刘永红,高建社.引种欧洲黑杨无性系苗期生长测定与选择.西北农林科技大学学报:自然科学版,2004,32(10):102-106Zhou Y X,Su X H,Fan J F,Liu Y H,Gao J S.Growth determination and selection of introduced Populus nigra clones in seedling stage.Journal of Northwest Sci-tech University of Agricultural and Forestry:Natural Science Edition,2004,32(10):102-106
[25]Wang W,Altman A.Plant responses to drought,salinity and extreme temperatures:towards genetic engineering for stress tolerance.Planta,2003,218:1-14
[26]Zhang X,Zang R,Li C.Population differences in physiological and morphological adaptations of Populus davidiana seedlings in response to progressive drought stress.Plant Science,2004,166:791-797
[27]Yin C Y,Wang X,Duan B L,Luo J X,Li C Y.Early growth,dry matter allocation and water use efficiency of two sympatric Populus,species as affected by water stress.Environmental Experimental Botany,2005,53:315-322
[28]Anjum S A,Xie X Y,Wang L C,Saleem M,Man C,Lei W.Morphological,physiological and biochemical responses of plants to drought stress.African Journal of Agricultural Research,2011,6:2026-2032
[29]Lee T Y,Je S M,Kwak M J,Akhmadi K,Tumurbaatar E,Khaine I,Lee H,Jang J Y,Kim H N,Ahn H J,Woo SY.Physiological responses of Populus sibirica to different irrigation regimes or reforestation in arid area.South African Journal of Botany,2017,112:329-335
[30]Vanlerberghe G C,Martyn G D,Dahal K.Alternative oxidase:a respiratory electron transport chain pathway essential for maintaining photosynthetic performance during drought stress.Physiologia Plantarum,2016,157:322-337
[31]Yin C Y,Peng Y H,Zang R G,Zhu Y P,Li C Y.Adaptive responses of Populus kangdingensis to drought stress.Physiologia Plantarum,2005,123:445-451
[32]李禄军,蒋志荣,李正平,邵玲玲.3树种抗旱性的综合评价及其抗旱指标的选取.水土保持研究,2006,13(6):253-254Li L J,Jiang Z R,Li Z P,Shao L L.Comprehensive evaluation on drought resistance of three tree species and the choice of drought resistance indexes.Research of Soil and Water Conservation,2006,13(6):253-254
[33]王兴荣,张彦军,李玥,刘天鹏,张金福,祁旭升.干旱胁迫对大豆生长的影响及抗旱性评价方法与指标筛选,植物遗传资源学报,2018,19(1):49-56Wang X R,Zhang Y J,Li Y,Liu T P,Zhang J F,Qi X S.Effect of drought stress on growth and screening methods and indexes for drought-resistance in soybean.Journal of Plant Genetic Resource,2018,19(1):49-56
[34]张文英,智慧,柳斌辉,彭海成,李伟,王永芳,李海权,栗雨勤,刁现民.谷子全生育期抗旱性鉴定及抗旱指标筛选.植物遗传资源学报,2010,11(5):560-565Zhang W Y,Zhi H,Liu B H,Peng H C,Li W,Wang Y F,Li HQ,Li Y Q,Diao X M.Indexes screening for drought resistance test of foxtail millet.Journal of Plant Genetic Resources,2010,11(5):560-565
[35]董孔军,刘天鹏,何继红,任瑞玉,张磊,杨天育.糜子育成品种抗旱性评价与鉴定指标筛选.植物遗传资源学报,2015,16(5):968-975Dong K J,Liu T P,He J H,Ren R Y,Zhang L,Yang TY.Evaluation and identification index selection on the drought resistance of broomcorn millet elite cultivars at seeding stage.Journal of Plant Genetic Resource,2015,16(5):968-975
[36]Hoecker N,Keller B,Piepho H,Hochholdinger F.Manifestation of heterosis during early maize(Zea mays L.)root development.Theoretical Applied Genetics,2006,112:421-429
[37]Bruce A B.The Mendelian theory of heredity and the augmentation of vigor.Science,1910,32:627-628
[38]Crow J F.Alternative hypothesis of hybrid vigor.Genetics,1948,33:477-487
[39]Semel Y,Nissenbaum J,Menda N,Zinder M,Krieger U,I s s m a n N,P l e b a n T,L i p p m a n Z B,G u r A,Z a m i r D.Overdominant quantitative trait loci for yield and fitness in tomato.Proceedings of the National Academy of Science,2006,103:12981-12986
[40]East E M.Heterosis.Genetics,1936,21:375
[41]Grassi G,Magnani F.Stomatal,mesophyll conductance and biochemical limitations to photosynthesis as affected by drought and leaf ontogeny in ash and oak trees.Plant,Cell and Environment,2005,28:834-849
[42]Susiluoto S,Berninger F.Interactions between morphological and physiological drought responses in Eucalyptus microtheca.Silva Fennica,2007,41:221-233
[43]He F,Wang H L,Li H G,Su Y Y,Li S,Yang Y L,Feng CH,Yin W L,Xia X L.PeCHYR1,a ubiquitin E3 ligase from Populus euphratica,enhances drought tolerance via ABA-induced stomatal closure by ROS production in Populus.Plant Biotechnology Journal,DOI:10.1111/pbi.12893
[44]Alexieva V,Sergiev I,Mapelli S,Karanov E.The effect of drought and ultraviolet radiation on growth and stress markers in pea and wheat.Plant,Cell and Environment,2001,24:1337-1344
[45]尹春英,李春阳.杨树抗旱性研究进展.应用与环境生物学报,2003,9(6):662-668Yin C Y,Li C Y.Advance in research on drought resistance of Populus.Chinese Journal of Applied and Environment Biology,2003,9(6):662-668
[46]Xu X,Yang F,Xiao X,Sheng Z,Helen K,Chun Y L.Sexspecific response of Populus cathayana to drought and elevated temperatures.Plant,Cell and Environment,2008,31:850-860
[2]王宇超,王得祥,彭少兵,何帆.干旱胁迫对木本滨藜生理特性的影响.林业科学,2010,46(1):61-67Wang Y C,Wang D X,Peng S B,He F.Effects of drought stress on physiological characteristics of woody saltbush.Scientia Silvae Sinicae,2010,46(1):61-67
[3]Dai A.Increasing drought under global warming in observations and models.Nature Climate Change,2012,3:52-58
[4]Allen C D,Macalady A K,Chenchouni H,Bachelet D,Mcdowell N G,Vennetier M,Kitzberger T,Rigling A,Breshears D D,Hogg E H,Gonzalez P,Fensham R J,Zhang Z,Castro J,Demidova N,Lim J H,Allard G,Running S W.Aglobal overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests.Forest Ecology Management,2010,259:660-684
[5]杨建伟,梁宗锁,韩蕊莲,孙群,崔浪军.不同干旱土壤条件下杨树的耗水规律及水分利用效率研究.植物生态学报,2004,28(5):657-664Yang J W,Liang Z S,Han R L,Sun Q,Cui L J.Water use efficiency and water consumption characterics of poplar under soil drought conditions.Acta Phytoecological Sinica,2004,28(5):657-664
[6]Mazzoleni S,Dickmann D I.Differential physiological and morphological responses of two hybrid Populus clones to water stress.Tree Physiology,1988,4:61-70
[7]Souch C A,Stephens W.Growth,productivity and water use in three hybrid poplar clones.Tree Physiology,1998,18:829-835
[8]付士磊,周永斌,何兴元,陈玮.干旱胁迫对杨树光合生理指标的影响.应用生态学报,2006,17(11):2016-2019Fu S L,Zhou Y B,He X Y,Chen W.Effects of drought stress on photosynthesis physiology of Populus pseudo-simonii.Chinese Journal of Applied Ecology,2006,17(11):2016-2019
[9]Viger M,Smith H K,Cohen D,Dewoody J,Trewin H,Steenackers M,Bastien C,Taylor G.Adaptive mechanisms and genomic plasticity for drought tolerance identified in European black poplar(Populus nigra L.).Tree Physiology,2016,36:909
[10]杨淑红,宋德才,刘艳萍,杨清淮,朱延林.土壤干旱胁迫和复水后3个杨树品种叶片部分生理指标变化及抗旱性评价.植物资源与环境学报,2014,23(3):65-73Yang S H,Song D C,Liu Y P,Yang Q H,Zhu Y L.Changes of some physiological indexes in leaf of three cultivars of Populus after drought stress in soil and re-watering and evaluation on their drought resistance.Journal of Plant Resources and Environment,2014,23(3):65-73
[11]邱兴,樊军锋,李晓东,吕小锋.6个杨树新无性系叶片旱生结构研究.西北林学院学报,2015,30(1):96-101Qiu X,Fan J F,Li X D,Lv X F.Research on drought resistance on leaf anatomical structure of six new poplar clones.Journal of Northwest Forestry University,2015,30(1):96-101
[12]Loutfy N,Eltayeb M A,Hassanen A M,Moustafa M FM,Sakuma Y,Inouhe M.Changes in the water status and osmotic solute contents in response to drought and salicylic acid treatments in four different cultivars of wheat(Triticum aestivum).Journal of Plant Research,2012,125:173-184
[13]Gower S T,Kucharik C J,Norman J M.Direct and indirect estimation of leaf area index,f APAR,and net primary production of Terrestrial Ecosystems.Remote Sensing Environment,1999,70:29-51
[14]Nicolas M,Erwin D,Boudouresque E,Delay D,Petit J,Delmotte F,Brignolas F.Impact of successive drought and re-watering cycles on growth and specific leaf area of two Populus×canadensis(Moench)clones,‘Dorskamp’and‘Luisa-Avanzo’.Tree Physiology,2003,23:1225-1235
[15]Yang Y,Yao Y,Xu G,Li C Y.Growth and physiological responses to drought and elevated ultraviolet-b in two contrasting populations of Hippophae rhamnoides.Physiologia Plantarum,2010,124:431-440
[16]李忠武,蔡强国,唐政洪,吴淑安,王宏.作物生产力模型及其应用研究.应用生态学报,2002,13(9):1174-1178L i Z W,C a i G Q,Ta n g Z H,Wu S A,Wa n g H.C r o p productivity model and its application.Chinese Journal of Applied Ecology,2002,13(9):1174-1178
[17]姬慧娟,贾会霞,章小铃,胡建军.干旱胁迫对红皮柳光合特性日变化及生长的影响.南京林业大学学报:自然科学版,2016,40(6):41-46Ji H J,Jia H X,Zhang X L,Hu J J.Effect of drought stress on photosynthetic diurnal course and growth of Salix purpurea.Journal of Nanjing Forestry University:Natural Sciences Edition,2016,40(6):41-46
[18]张春玲,李淑梅,赵自成,胡建军,韩一凡.杨树新品种‘丹红杨’.林业科学,2008,44(1):169Zhang C L,Li S M,Zhao Z C,Hu J J,Han Y F.A new poplar variety Populus deltoides CL.‘Dahong’.Scientia Silvae Sinicae,2008,44(1):169
[19]Chen J H,Song Y P,Zhang H,Zhang D Q.Genome wide analysis of gene expression in response to drought stress in Populus simonii.Plant Molecular Biology Reporter,2013,31:946-962
[20]杜森,高祥照.土壤分析技术规范.北京:中国农业出版社,2006:47-49Du S,Gao X Z.Technical specification for soil analysis.Beijing:China Agricultural Publishers,2006:47-49
[21]曾辉,曹苑南,王述民,王兰芬,董海鸿,陈吉宝.30份普通菜豆苗期抗旱性鉴定及抗旱指标的确定.植物遗传资源学报,2016,17(6):1014-1021Zeng H,Cao Y N,Wang S M,Wang L F,Dong H H,Chen J B.Evaluation of drought tolerance of 30 common bean in seedling stage and screening of drought-tolerance assessment index.Journal of Plant Genetic Resource,2016,17(6):1014-1021
[22]胡标林,杨平,万勇,李霞,罗世友,罗向东,谢建坤.东乡野生稻BILs群体苗期抗旱性综合评价及其遗传分析.植物遗传资源学报,2013,14(2):249-256Hu B L,Yang P,Wan Y,Li X,Luo S Y,Luo X D,Xie JK.Comprehensive assessment of drought resistance of BILs population derived from Dongxiang wild rice(Oryza rufipogon Griff)at seedling stage and its genetic analysis.Journal of Plant Genetic Resources,2013,14(2):249-256
[23]谢季坚.农业科学中的模糊数学方法.武汉:华中理工大学出版社,1993:99-193Xie J J.Fuzzy mathematics in agricultural science.Wuhan:Huazhong university of science and technology publishing house,1993:99-193
[24]周永学,苏晓华,樊军锋,刘永红,高建社.引种欧洲黑杨无性系苗期生长测定与选择.西北农林科技大学学报:自然科学版,2004,32(10):102-106Zhou Y X,Su X H,Fan J F,Liu Y H,Gao J S.Growth determination and selection of introduced Populus nigra clones in seedling stage.Journal of Northwest Sci-tech University of Agricultural and Forestry:Natural Science Edition,2004,32(10):102-106
[25]Wang W,Altman A.Plant responses to drought,salinity and extreme temperatures:towards genetic engineering for stress tolerance.Planta,2003,218:1-14
[26]Zhang X,Zang R,Li C.Population differences in physiological and morphological adaptations of Populus davidiana seedlings in response to progressive drought stress.Plant Science,2004,166:791-797
[27]Yin C Y,Wang X,Duan B L,Luo J X,Li C Y.Early growth,dry matter allocation and water use efficiency of two sympatric Populus,species as affected by water stress.Environmental Experimental Botany,2005,53:315-322
[28]Anjum S A,Xie X Y,Wang L C,Saleem M,Man C,Lei W.Morphological,physiological and biochemical responses of plants to drought stress.African Journal of Agricultural Research,2011,6:2026-2032
[29]Lee T Y,Je S M,Kwak M J,Akhmadi K,Tumurbaatar E,Khaine I,Lee H,Jang J Y,Kim H N,Ahn H J,Woo SY.Physiological responses of Populus sibirica to different irrigation regimes or reforestation in arid area.South African Journal of Botany,2017,112:329-335
[30]Vanlerberghe G C,Martyn G D,Dahal K.Alternative oxidase:a respiratory electron transport chain pathway essential for maintaining photosynthetic performance during drought stress.Physiologia Plantarum,2016,157:322-337
[31]Yin C Y,Peng Y H,Zang R G,Zhu Y P,Li C Y.Adaptive responses of Populus kangdingensis to drought stress.Physiologia Plantarum,2005,123:445-451
[32]李禄军,蒋志荣,李正平,邵玲玲.3树种抗旱性的综合评价及其抗旱指标的选取.水土保持研究,2006,13(6):253-254Li L J,Jiang Z R,Li Z P,Shao L L.Comprehensive evaluation on drought resistance of three tree species and the choice of drought resistance indexes.Research of Soil and Water Conservation,2006,13(6):253-254
[33]王兴荣,张彦军,李玥,刘天鹏,张金福,祁旭升.干旱胁迫对大豆生长的影响及抗旱性评价方法与指标筛选,植物遗传资源学报,2018,19(1):49-56Wang X R,Zhang Y J,Li Y,Liu T P,Zhang J F,Qi X S.Effect of drought stress on growth and screening methods and indexes for drought-resistance in soybean.Journal of Plant Genetic Resource,2018,19(1):49-56
[34]张文英,智慧,柳斌辉,彭海成,李伟,王永芳,李海权,栗雨勤,刁现民.谷子全生育期抗旱性鉴定及抗旱指标筛选.植物遗传资源学报,2010,11(5):560-565Zhang W Y,Zhi H,Liu B H,Peng H C,Li W,Wang Y F,Li HQ,Li Y Q,Diao X M.Indexes screening for drought resistance test of foxtail millet.Journal of Plant Genetic Resources,2010,11(5):560-565
[35]董孔军,刘天鹏,何继红,任瑞玉,张磊,杨天育.糜子育成品种抗旱性评价与鉴定指标筛选.植物遗传资源学报,2015,16(5):968-975Dong K J,Liu T P,He J H,Ren R Y,Zhang L,Yang TY.Evaluation and identification index selection on the drought resistance of broomcorn millet elite cultivars at seeding stage.Journal of Plant Genetic Resource,2015,16(5):968-975
[36]Hoecker N,Keller B,Piepho H,Hochholdinger F.Manifestation of heterosis during early maize(Zea mays L.)root development.Theoretical Applied Genetics,2006,112:421-429
[37]Bruce A B.The Mendelian theory of heredity and the augmentation of vigor.Science,1910,32:627-628
[38]Crow J F.Alternative hypothesis of hybrid vigor.Genetics,1948,33:477-487
[39]Semel Y,Nissenbaum J,Menda N,Zinder M,Krieger U,I s s m a n N,P l e b a n T,L i p p m a n Z B,G u r A,Z a m i r D.Overdominant quantitative trait loci for yield and fitness in tomato.Proceedings of the National Academy of Science,2006,103:12981-12986
[40]East E M.Heterosis.Genetics,1936,21:375
[41]Grassi G,Magnani F.Stomatal,mesophyll conductance and biochemical limitations to photosynthesis as affected by drought and leaf ontogeny in ash and oak trees.Plant,Cell and Environment,2005,28:834-849
[42]Susiluoto S,Berninger F.Interactions between morphological and physiological drought responses in Eucalyptus microtheca.Silva Fennica,2007,41:221-233
[43]He F,Wang H L,Li H G,Su Y Y,Li S,Yang Y L,Feng CH,Yin W L,Xia X L.PeCHYR1,a ubiquitin E3 ligase from Populus euphratica,enhances drought tolerance via ABA-induced stomatal closure by ROS production in Populus.Plant Biotechnology Journal,DOI:10.1111/pbi.12893
[44]Alexieva V,Sergiev I,Mapelli S,Karanov E.The effect of drought and ultraviolet radiation on growth and stress markers in pea and wheat.Plant,Cell and Environment,2001,24:1337-1344
[45]尹春英,李春阳.杨树抗旱性研究进展.应用与环境生物学报,2003,9(6):662-668Yin C Y,Li C Y.Advance in research on drought resistance of Populus.Chinese Journal of Applied and Environment Biology,2003,9(6):662-668
[46]Xu X,Yang F,Xiao X,Sheng Z,Helen K,Chun Y L.Sexspecific response of Populus cathayana to drought and elevated temperatures.Plant,Cell and Environment,2008,31:850-860