小麦品种(系)延绿性的遗传变异及其特征
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摘要
为了解高湿环境下小麦品种(系)延绿性的遗传变异和延绿特点,2012-2014年以我国主要麦区近年来育成的47个品种(系)为材料,以品种(系)开花后不同日龄下旗叶的绿色叶面积百分比(%GLA)为指标,采用模糊聚类法对品种(系)的延绿性进行分类,用Gompertz模型拟合品种(系)开花后旗叶的衰老过程并解析其衰老特性。结果表明,3年试验中,小麦品种(系)均可分为延绿、中等延绿、中等早衰和早衰4种类型,其中,CP02-8-5-6-2-1、天民668、川农16、川麦60、川麦107和泰山046219为延绿稳定表达品种(系);小麦旗叶衰老特征参数达到最大衰老时间(TMRS)、绿色叶面积持续期(GLAD)和平均衰老速度(ARS)与灌浆后期旗叶的%GLA均呈极显著正相关;同一年度不同延绿型品种(系)间旗叶的TMRS、GLAD和ARS存在显著差异,MRS无显著差异,TMRS和GLAD表现为延绿型>中等延绿型>中等早衰型>早衰型,ARS表现为早衰型>中等早衰型>中等延绿型>延绿型;不同年度间小麦旗叶TMRS和GLAD表现为2014年>2013年>2012年,MRS表现为2013年>2014年>2012年,ARS以2012年>2013年>2014年。这说明旗叶衰老特征参数可作为品种延绿的评价指标。
To understand the extent and features of genetic variation of stay green traits in wheat varieties under water-logged environment,forty-seven varieties released from major wheat growing regions were evaluated during 2012-2014 wheat growing seasons,and the percentage green leaf area(%GLA)of flag leaf was used to group the tested varieties by fuzzy cluster method,and to fit flag leaf senescence process by Gompertz model the tested varieties.The derived parameters of leaf senescence curve were used to analyze stay green features of the tested varieties.The results indicated that forty-seven varieties tested in years 2012-2014 could be clustered into four different types,stay green(SG),moderately stay green(MSG),moderately non-stay green(MNSG)and non-stay green(NSG),and CP02-8-5-6-2-1,Tianmin 668,Chuannong 16,Chuanmai 60,Chuanmai 107 and Taishan 046219 could be as-signed as SG type based on their stay green performances in the three consecutive years;the time of maximum rate of leaf senescence(TMRS),green leaf area duration(GLAD)and average rate of leaf senescence(ARS)were significantly and positively correlated with %GLA at late filling stage;in the same growing season except for MRS,the other three parameters TMRS,GLAD and ARS were significantly different among the four different types,and the orders of the mean values were as following:TMRS and GLAD ranked as SG>MSG>MNSG>NSG,ARS ranked as NSG>MNSG>MSG>SG;in different growing seasons,the mean values for the flag leaf senescence parameters of the four different types ranked as 2014>2013>2012 for TMRS and GLAD,2013>2014>2012 for MRS,and 2012>2013>2014 for ARS.The characteristic parameters of leaf senescence process could be used as an indicator to distinguish the extent of the stay green performance of wheat variety.
To understand the extent and features of genetic variation of stay green traits in wheat varieties under water-logged environment,forty-seven varieties released from major wheat growing regions were evaluated during 2012-2014 wheat growing seasons,and the percentage green leaf area(%GLA)of flag leaf was used to group the tested varieties by fuzzy cluster method,and to fit flag leaf senescence process by Gompertz model the tested varieties.The derived parameters of leaf senescence curve were used to analyze stay green features of the tested varieties.The results indicated that forty-seven varieties tested in years 2012-2014 could be clustered into four different types,stay green(SG),moderately stay green(MSG),moderately non-stay green(MNSG)and non-stay green(NSG),and CP02-8-5-6-2-1,Tianmin 668,Chuannong 16,Chuanmai 60,Chuanmai 107 and Taishan 046219 could be as-signed as SG type based on their stay green performances in the three consecutive years;the time of maximum rate of leaf senescence(TMRS),green leaf area duration(GLAD)and average rate of leaf senescence(ARS)were significantly and positively correlated with %GLA at late filling stage;in the same growing season except for MRS,the other three parameters TMRS,GLAD and ARS were significantly different among the four different types,and the orders of the mean values were as following:TMRS and GLAD ranked as SG>MSG>MNSG>NSG,ARS ranked as NSG>MNSG>MSG>SG;in different growing seasons,the mean values for the flag leaf senescence parameters of the four different types ranked as 2014>2013>2012 for TMRS and GLAD,2013>2014>2012 for MRS,and 2012>2013>2014 for ARS.The characteristic parameters of leaf senescence process could be used as an indicator to distinguish the extent of the stay green performance of wheat variety.
引文
[1]GONG Y H,ZHANG J,GAO J F,et al.Slow export of photoassimilate from stay-green leaves during late grain-filling stage in hybrid winter wheat(Triticum aestivum L.)[J].Journal of Agronomy&Crop Science,2005,191:292.
[2]赵会杰,李有,邹琦.两个不同穗型小麦品种(系)的冠层辐射和光合特征的比较研究[J].作物学报,2002,28(5):654.ZHAO H J,LI Y,ZHOU Q.A comparative study on characteristics of radiation and photosynthesis in canopy of two different spike-type cultivars of wheat[J].Acta Agronomica Sinica,2002,28(5):654.
[3]龚月桦,刘迎洲,高俊风.K型杂交小麦901及亲本籽粒灌浆的生长分析[J].中国农业科学,2004,37(9):1288.GONG Y H,LIU Y Z,GAO J F.Growth analysis on the process of grain filling in hybrid wheat 901and its parent[J].Scientia Agricultura Sinica,2004,37(9):1288.
[4]武永胜,薛晖,刘洋,等.持绿型小麦叶片衰老和叶绿素荧光特征的研究[J].干旱地区农业研究,2010,28(4):117.WU Y S,XUE H,LIU Y,et al.Study on senescence and chlorophyll fluorescence traits of stay-green leaf in wheat[J].Agricultural Research in the Arid Areas,2010,28(4):117.
[5]杨东清,李玉玲,倪英丽,等.外源ABA和6-BA对不同持绿型小麦籽粒灌浆进程及蛋白质含量影响[J].作物学报,2014,40(2):301.YANG D Q,LI Y L,NI Y L,et al.Effects of exogenous ABA and 6-BA on protein content and grain filling process in different types of stay-green wheat[J].Acta Agronomica Sinica,2014,40(2):301.
[6]KUMARI M,PUDAKE R N,SINGH V P.Identification of microsatellite markers associated with stay green trait in wheat RILs[J].Indian Journal of Genetics&Plant Breeding,2012,74(4):415.
[7]LUCHE H S,SILVA J A G,NORNBERG R,et al.Stay-green effects on adaptability and stability in wheat[J].African Journal of Agricultural Research,2015,10(11):1142.
[8]CHRISTOPHER J T,CHRISTOPHER M J,BORRELL AK,et al.Stay-green traits to improve wheat adaptation in wellwatered and water-limited environments[J].Journal of Experimental Botany,2016,67(17):5159.
[9]李永攀,罗培高,任正隆.小麦持绿性与产量关系研究[J].西南农业学报,2008,21(5):1221.LI Y P,LUO P G,REN Z H.Studies on the relation between the yield and trait of green-keeping wheat[J].Southwest China Journal of Agricultural Sciences,2008,21(5):1221.
[10]李涛,龚月桦,吕金印,等.不同基因型小麦14 C-储备物在花后的转运与分配规律[J].核农学报,2010,24(1):149.LI T,GONG Y H,LU J Y,et al.Transportation and distribution of 14C-reserves on different genotype wheat varieties after anthesis[J].Acta Agriculturae Nucleatae Sinica,2010,24(1):149.
[11]石慧清,龚月桦,张东武.花后高温对持绿型小麦叶片衰老及籽粒淀粉合成相关酶的影响[J].植物生态学报,2011,35(7):769.SHI H Q,GONG Y H,ZHANG D W,et al.Effect of high temperature on leaf senescence and related enzymes of grain starch synthesis in stay-green wheat after anthesis[J].Acta Phytoecologica Sinic,2011,35(7):769.
[12]CHRISTOPHER J T,MANSCHADI A M,HAMMER G L,et al.Developmental and physiological traits associated with high yield and stay-green phenotype in wheat[J].Australian Journal of Agricultural Research,2008,59(4):354.
[13]隋娜,李萌,田纪春,等.超高产小麦品种(系)生育后期光合特性的研究[J].作物学报,2005,31(6):808.SUI N,LI M,TIAN J Cet al.Photosynthetic characteristics of super high yield wheat cultivars at late growth period[J].Acta Agronomica Sinica,2005,31(6):808.
[14]ZHANG C J,CHEN G X,GAO X X,et al.Photosynthetic decline in flag leaves of two field-grown spring wheat cultivars with different senescence properties[J].South African Journal of Botany,2006,72(1):15.
[15]DERKX A P,ORFORD S,GRIFFITHS S,et al.Identification of differentially senescing mutants of wheat and impacts on yield,biomass and nitrogen partitioning[J].Journal of Integrative Plant Biology,2012,54(8):555.
[16]TIAN F X,GONG J F,WANG G P,et al.Improved drought resistance in a wheat stay-green mutant Tasg1under field conditions[J].Biologia Plantarum,2012,56(3):509.
[17]NARUOKA Y,SHERMAN J D,LANNING S P,et al.Genetic analysis of green leaf duration in spring wheat[J].Crop Science,2012,52:99.
[18]JOSHI A K,KUMARI M,SINGH V P,et al.Stay green trait:variation,inheritance and its association with spot blotch resistance in spring wheat(Triticum aestivum L.)[J].Euphytica,2007,153(1-2):59.
[19]VERMA V,FOULKES M J,WORLAND A J,et al.Mapping quantitative trait loci for flag leaf senescence as a yield determinant in winter wheat under optimal and droughtstressed environments[J].Euphytica,2004,135:255.
[20]VIJAYALAKSHMI K,FRITZ A K,PAULSEN G M,et al.Modeling and mapping QTL for senescence-related traits in winter wheat under high temperature[J].Molecular Breeding,2010,26(2):163.
[21]陈朝儒,奚亚军,王竹林,等.冬小麦持绿和灌浆特征及其抗早衰特性评价[J].西北植物学报,2011,31(4):715.CHEN C R,XI Y J,WANG Z L,et al.Characteristics of stay-green and grain-filling and the evaluation of anti-senescence properties of winter wheat[J].Acta Botanica BorealiOccidentalia Sinica,2011,31(4):715.
[22]AHLAWAT S,CHHABRA A K,BEHL R K,et al.Genotypic divergence analysis for stay green characters in wheat(Triticum aestivum L.em.Thell)[J].The South Pacific Journal of Natural Sciences,2009,26(1):73.
[23]KUMARI M,PUDAKE R N,SINGH V P,et al.Association of staygreen trait with canopy temperature depression and yield traits under terminal heat stress in wheat(Triticum aestivum L.)[J].Euphytica,2013,190(1):87.
[24]PASK A,PIETRAGALLA J,MULLAN D,et al.Physiological breedingⅡ:A field guide to wheat phenotyping[M].Chapter 12:Leaf area,green crop area and senescence.Mexico,CIMMYT,2012:58.
[25]SEBER G A F,WILD C J.Nonlinear regression[M].Chapter 7:Growth model.New York,Wiley,1989:325-365.
[26]WU X Y,KUAI B K,JIA J Z,et al.Regulation of leaf senescence and crop genetic improvement[J].Journal of Integrative Plant Biology,2012,54(12):936.
[27]THOMAS H,HOWARTH C J.Five ways to stay green[J].Journal of Experimental Botany,2000,51:329.
[28]LUO P G,REN Z L.A single recessive gene controls leaf chlorosis in wheat[J].Journal of Plant Physiology Molecular Biology,2006,32:330.
[29]任天恒,陈放,张怀琼,等.1RS.1BL易位在川农号系列小麦新品种(系)选育中的作用[J].麦类作物学报,2011,31(3):430.REN T H,CHEN F,ZHANG H Q,et al.Application of1RS.1BL translocation in the breeding of“Chuannong”series wheat cultivars[J].Journal of Triticeae Crops,2011,31(3):430.
[30]姜朋,陈小霖,张平平,等.宁麦9号对其衍生品种(系)的遗传贡献[J].作物学报,2014,40(5):830.JIANG P,CHEN X L,ZHANG P P,et al.Genetic contribution of wheat variety Ningmai 9to its derivates[J].Acta Agronomica Sinica,2014,40(5):830.
[31]GREGERSEN P L,CULETIC A,BOSCHIAN L,et al.Plant senescence and crop productivity[J].Plant Molecular Biology,2013,82:603.
[2]赵会杰,李有,邹琦.两个不同穗型小麦品种(系)的冠层辐射和光合特征的比较研究[J].作物学报,2002,28(5):654.ZHAO H J,LI Y,ZHOU Q.A comparative study on characteristics of radiation and photosynthesis in canopy of two different spike-type cultivars of wheat[J].Acta Agronomica Sinica,2002,28(5):654.
[3]龚月桦,刘迎洲,高俊风.K型杂交小麦901及亲本籽粒灌浆的生长分析[J].中国农业科学,2004,37(9):1288.GONG Y H,LIU Y Z,GAO J F.Growth analysis on the process of grain filling in hybrid wheat 901and its parent[J].Scientia Agricultura Sinica,2004,37(9):1288.
[4]武永胜,薛晖,刘洋,等.持绿型小麦叶片衰老和叶绿素荧光特征的研究[J].干旱地区农业研究,2010,28(4):117.WU Y S,XUE H,LIU Y,et al.Study on senescence and chlorophyll fluorescence traits of stay-green leaf in wheat[J].Agricultural Research in the Arid Areas,2010,28(4):117.
[5]杨东清,李玉玲,倪英丽,等.外源ABA和6-BA对不同持绿型小麦籽粒灌浆进程及蛋白质含量影响[J].作物学报,2014,40(2):301.YANG D Q,LI Y L,NI Y L,et al.Effects of exogenous ABA and 6-BA on protein content and grain filling process in different types of stay-green wheat[J].Acta Agronomica Sinica,2014,40(2):301.
[6]KUMARI M,PUDAKE R N,SINGH V P.Identification of microsatellite markers associated with stay green trait in wheat RILs[J].Indian Journal of Genetics&Plant Breeding,2012,74(4):415.
[7]LUCHE H S,SILVA J A G,NORNBERG R,et al.Stay-green effects on adaptability and stability in wheat[J].African Journal of Agricultural Research,2015,10(11):1142.
[8]CHRISTOPHER J T,CHRISTOPHER M J,BORRELL AK,et al.Stay-green traits to improve wheat adaptation in wellwatered and water-limited environments[J].Journal of Experimental Botany,2016,67(17):5159.
[9]李永攀,罗培高,任正隆.小麦持绿性与产量关系研究[J].西南农业学报,2008,21(5):1221.LI Y P,LUO P G,REN Z H.Studies on the relation between the yield and trait of green-keeping wheat[J].Southwest China Journal of Agricultural Sciences,2008,21(5):1221.
[10]李涛,龚月桦,吕金印,等.不同基因型小麦14 C-储备物在花后的转运与分配规律[J].核农学报,2010,24(1):149.LI T,GONG Y H,LU J Y,et al.Transportation and distribution of 14C-reserves on different genotype wheat varieties after anthesis[J].Acta Agriculturae Nucleatae Sinica,2010,24(1):149.
[11]石慧清,龚月桦,张东武.花后高温对持绿型小麦叶片衰老及籽粒淀粉合成相关酶的影响[J].植物生态学报,2011,35(7):769.SHI H Q,GONG Y H,ZHANG D W,et al.Effect of high temperature on leaf senescence and related enzymes of grain starch synthesis in stay-green wheat after anthesis[J].Acta Phytoecologica Sinic,2011,35(7):769.
[12]CHRISTOPHER J T,MANSCHADI A M,HAMMER G L,et al.Developmental and physiological traits associated with high yield and stay-green phenotype in wheat[J].Australian Journal of Agricultural Research,2008,59(4):354.
[13]隋娜,李萌,田纪春,等.超高产小麦品种(系)生育后期光合特性的研究[J].作物学报,2005,31(6):808.SUI N,LI M,TIAN J Cet al.Photosynthetic characteristics of super high yield wheat cultivars at late growth period[J].Acta Agronomica Sinica,2005,31(6):808.
[14]ZHANG C J,CHEN G X,GAO X X,et al.Photosynthetic decline in flag leaves of two field-grown spring wheat cultivars with different senescence properties[J].South African Journal of Botany,2006,72(1):15.
[15]DERKX A P,ORFORD S,GRIFFITHS S,et al.Identification of differentially senescing mutants of wheat and impacts on yield,biomass and nitrogen partitioning[J].Journal of Integrative Plant Biology,2012,54(8):555.
[16]TIAN F X,GONG J F,WANG G P,et al.Improved drought resistance in a wheat stay-green mutant Tasg1under field conditions[J].Biologia Plantarum,2012,56(3):509.
[17]NARUOKA Y,SHERMAN J D,LANNING S P,et al.Genetic analysis of green leaf duration in spring wheat[J].Crop Science,2012,52:99.
[18]JOSHI A K,KUMARI M,SINGH V P,et al.Stay green trait:variation,inheritance and its association with spot blotch resistance in spring wheat(Triticum aestivum L.)[J].Euphytica,2007,153(1-2):59.
[19]VERMA V,FOULKES M J,WORLAND A J,et al.Mapping quantitative trait loci for flag leaf senescence as a yield determinant in winter wheat under optimal and droughtstressed environments[J].Euphytica,2004,135:255.
[20]VIJAYALAKSHMI K,FRITZ A K,PAULSEN G M,et al.Modeling and mapping QTL for senescence-related traits in winter wheat under high temperature[J].Molecular Breeding,2010,26(2):163.
[21]陈朝儒,奚亚军,王竹林,等.冬小麦持绿和灌浆特征及其抗早衰特性评价[J].西北植物学报,2011,31(4):715.CHEN C R,XI Y J,WANG Z L,et al.Characteristics of stay-green and grain-filling and the evaluation of anti-senescence properties of winter wheat[J].Acta Botanica BorealiOccidentalia Sinica,2011,31(4):715.
[22]AHLAWAT S,CHHABRA A K,BEHL R K,et al.Genotypic divergence analysis for stay green characters in wheat(Triticum aestivum L.em.Thell)[J].The South Pacific Journal of Natural Sciences,2009,26(1):73.
[23]KUMARI M,PUDAKE R N,SINGH V P,et al.Association of staygreen trait with canopy temperature depression and yield traits under terminal heat stress in wheat(Triticum aestivum L.)[J].Euphytica,2013,190(1):87.
[24]PASK A,PIETRAGALLA J,MULLAN D,et al.Physiological breedingⅡ:A field guide to wheat phenotyping[M].Chapter 12:Leaf area,green crop area and senescence.Mexico,CIMMYT,2012:58.
[25]SEBER G A F,WILD C J.Nonlinear regression[M].Chapter 7:Growth model.New York,Wiley,1989:325-365.
[26]WU X Y,KUAI B K,JIA J Z,et al.Regulation of leaf senescence and crop genetic improvement[J].Journal of Integrative Plant Biology,2012,54(12):936.
[27]THOMAS H,HOWARTH C J.Five ways to stay green[J].Journal of Experimental Botany,2000,51:329.
[28]LUO P G,REN Z L.A single recessive gene controls leaf chlorosis in wheat[J].Journal of Plant Physiology Molecular Biology,2006,32:330.
[29]任天恒,陈放,张怀琼,等.1RS.1BL易位在川农号系列小麦新品种(系)选育中的作用[J].麦类作物学报,2011,31(3):430.REN T H,CHEN F,ZHANG H Q,et al.Application of1RS.1BL translocation in the breeding of“Chuannong”series wheat cultivars[J].Journal of Triticeae Crops,2011,31(3):430.
[30]姜朋,陈小霖,张平平,等.宁麦9号对其衍生品种(系)的遗传贡献[J].作物学报,2014,40(5):830.JIANG P,CHEN X L,ZHANG P P,et al.Genetic contribution of wheat variety Ningmai 9to its derivates[J].Acta Agronomica Sinica,2014,40(5):830.
[31]GREGERSEN P L,CULETIC A,BOSCHIAN L,et al.Plant senescence and crop productivity[J].Plant Molecular Biology,2013,82:603.