玉米抗旱性指标的筛选及其遗传特性研究
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摘要
本文以抗旱性不同玉米自交系和杂交种为试材,采用盆栽、田间和实验室模拟水分胁迫等手段,分别从形态学、生理学对玉米育种抗旱性鉴定指标进行了系统研究,对不同杂交种和自交系的抗旱性进行了分析。在研究结果的数量分析过程中,借助灰色关联度分析法、隶属函数值法等科学统计分析方法,并在此基础上建立了玉米育种抗旱性鉴定指标体系。同时用Duncan氏新复极差测验法进行差异显著性测验；根据Griffing双列杂交配合力统计原理,选用仅包含正交组合,共P(P-1)/2个试验材料的试验方法获取试验数据,用DPS数据处理软件进行数据处理,计算自交系主要农艺性状的一般配合力(GCA)和特殊配合力(SCA)及遗传参数,并进行统计分析；用Mather的A、B、C尺度检验方法及方差分析方法分析控制玉米籽粒含水量的基因作用；分析了玉米抗旱性的遗传规律。主要研究结果如下：
1.以三个玉米杂交种及亲本自交系为试验材料,苗期和中后期进行中度水分胁迫,测定了一系列形态和生理生化指标。结果表明各杂交种间及各自交系间抗旱性差异明显。以抗旱系数进行排序,杂交种：铁单17)农大108)铁单10；亲本自交系：沈136>P178>丹340>铁9010>黄C>C8605-2。以抗旱指数进行排序,杂交种：铁单17)农大108>铁单10；亲本自交系：沈136>P178>铁9010>丹340>黄C>C8605-2。抗旱指数和抗旱系数具有相同的趋势。亲本及F1代间抗旱性在某些性状上有一定相关性,抗旱性强的自交系,其杂交种的抗旱性也较好,抗旱性弱的自交系,其杂交种抗旱性也相对较弱。利用隶属函数值法进行分析结果,其中自交系沈136苗期、花期平均抗旱隶属值最高,具有较强的抗旱性,P178、铁9010、铁单17、农大108、340属中等抗旱类型,C8605-2、黄C属弱抗旱类型,而铁单10属不抗旱类型。
2.中度水分胁迫处理玉米自交系和杂交种后,测定了一些农艺性状指标和生理生化指标,通过综合分析,结果表明：中度水分胁迫条件下,玉米植株叶片相对含水量、丙二醛含量和ASI与玉米子粒产量密切相关,能够比较准确地评价玉米自交系和杂交种抗旱性的强弱。
在中度水分胁迫条件下,利用上述几个相关指标对10个骨干自交系进行抗旱性测定,结果显示相对含水量、电导率、丙二醛与子粒产量呈极显著的正相关,ASI与子粒产量呈极显著负相关。说明在鉴定筛选玉米种植资源时,玉米植株叶片相对含水量、丙二醛含量、电导率和ASI与玉米子粒产量密切相关,能够比较准确地评价玉米自交系抗旱性的强弱,和抗旱系数、抗旱指数一样可以作为玉米育种耐旱自交系和杂交种筛选的鉴定指标。
3.进行玉米抗旱育种,首先要筛选抗旱资源,对种质资源进行抗旱性鉴定,选择抗旱性较好的材料作为玉米抗旱性育种的基础材料,鉴定有利的抗旱基因并加以利用。本文通过对一些骨干玉米自交系生理、形态指标和产量的测定,结果表明,玉米自交系间的耐旱性存在较大差异。其中,自交系沈136、齐319、丹598和沈137耐旱性较好,可以作为玉米耐旱育种的基础材料,进行抗旱自交系和杂交种的选育。
4.采用PEG处理模拟水分胁迫,能够达到与田间水分胁迫相似的效果。因此可以利用PEG在芽苗期对玉米幼苗进行模拟干旱处理,结合一些抗旱指标的测定,能够比较准确地评价玉米种质的抗旱性。本文采用PEG处理幼苗模拟水分胁迫,通过叶片丙二醛含量、电导率和叶片水势等几个相关生理指标测定,分析玉米自交系的抗旱性,同时结合盆栽试验进行干旱胁迫处理测定玉米自交系的产量,通过抗旱指数评价玉米自交系的抗旱性,并对苗期指标与抗旱指数进行了相关性分析,结果PEG处理幼苗模拟干旱胁迫鉴定玉米自交系的抗旱性与抗旱指数具有显著的相关性,因此苗期采用PEG处理模拟水分胁迫进行大量玉米种质的抗旱性筛选,结合田间干旱鉴定,可以更好地评价玉米种质的抗旱性。采用PEG模拟水分胁迫处理玉米幼苗,方法简单、有效、低成本、短周期就能够分析大量样品,解决了盆栽试验、田间试验进行水分干旱胁迫处理样品少,胁迫时间、胁迫强度难以控制等问题,是快速鉴定玉米种质抗旱性的比较准确的评价方法。
5.本研究利用耐旱性不同的丹598、辽618、K12、沈136、沈137、丹340、铁9010、P178,8个自交系按Griffing方法4组配的28个不完全双列杂交组合及自交系沈136和铁9010为亲本材料,配制的P1 P2 F1 F2四个世代群体,进行了抗旱性性状的遗传分析。研究结果表明,水分胁迫下穗长、秃尖、百粒重、行粒数这些性状受加性效应影响大,产量、株高和穗位性状受非加性基因作用较大。行粒数、穗长和秃尖的广义遗传力和狭义遗传力均较低,性状受环境影响大,百粒重性状的广义遗传力和狭义遗传力均较高,而且相差较小,性状受环境影响小,且为加性遗传,产量、株高和穗位性状广义遗传力较高,性状受环境影响较大。其中穗位主要受非加性效应控制,产量和株高加性效应和非加性效应均有影响。抗旱指数性状的最适遗传模型为E-1模型,抗旱性指数主要受2对主基因+多基因控制。其中主基因遗传力大于多基因遗传力,分别为61.59%和17.72%,主基因对抗旱指数的变异度的影响远远大于多基因对抗旱指数的变异度的影响,而且主基因+多基因决定了抗旱指数表型变异的79.32%,另外有20.68%是由环境因素决定的。
Morphology and physiological indexes were studied to identify drought resistance of maize according to pot experiment, filed experiment and lab simulated water stress using different maize inbred lines and hybrids with different drought resistance. The grey correlation analysis method and membership function value method were used to establish the maize drought resistance identification system. Duncan's new multiple range test method was used to significant test. According to Griffing double row hybrid general combining ability statistical principle, the test samples were P(P-1)/2 orthogonal combinations and DPS were used for data analysis to calculate the GCA, SCA and genetic parameter of maize inbred lines. The gene effect controlled maize grain water content and genetic law of maize drought resistance of maize were analyzed using Mather's A、B、C scale inspection and analysis of variance method. The main results were as follows:
1. Morphology and physiological and biochemical indexes of three maize hybrids and their inbred lines were studied during moderate water stress at seedling stage and anthesis. The results indicated that the drought resistance of maize hybrids and inbred lines had significant differences. The drought resistance coefficient ordering of maize hybrids were as follows: Tiedan17> Nongda108> Tiedan 10; while the drought resistance coefficient ordering of inbred lines were as follows:Shen136> P178> Dan340> Tie 9010> Huang C> C8605-2. The drought resistance index and coefficient had same trend. The drought resistance of parents and F1 had some correlations on some traits. The drought resistance of inbred line was higher, the corresponding hybrid was better. According to membership function value method, the result indicated that Shen 136 which had the highest membership value at seedling stage and anthesis had higher drought resistance; P178, Tie9010, Tiedan17, Nongda108 and 340 were medium drought resistance type; C8605-2 and Huang C were weak drought resistance type; while Tiedan10 was sensitive drought resistance type.
2. Agronomic characters and physiological and biochemical indexes of maize hybrids and inbred lines were measured under moderate water stress. The results showed that the relative water content, MDA and ASI closely related with maize yield. These indexes could be accurately to evaluate the drought resistance of maize hybrids and inbred lines.
The drought resistance of ten ordinary maize inbred lines was studied using above related indexes. The results indicated that the relative water content, MDA content, electric conductivity of maize leaves were significantly correlated to the yield of maize under moderate drought stress. While there was a significant negative correlation between ASI and yield. It showed that these physiological indexes of leaves could accurately evaluate the drought resistance of maize and could be selection and identification index of maize inbred lines in maize drought resistance breeding as well as drought resistance coefficient and drought resistance index.
3. Selecting and identification of drought resistance germplasm resources was the primary problem to maize drought resistance breeding. Physiology and morphological indexes and yield of some ordinary maize inbred lines were measured in this study. The results indicated that the drought resistance of maize inbred lines had significant differences. The drought resistance of Shen136, Qi319, Dan598 and Shen137 were better and these inbred lines could be basic material as maize drought resistance breeding.
4. The maize using PEG simulated drought stress could reach the same effect with the field water stress. Hence the maize which treated using PEG simulated drought stress at seedling stage and combined with drought resistance indexes could be accurately evaluate the drought resistance of maize. MDA, electric conductivity and water potential were measured to analysis the drought resistance of maize inbred lines using PEG simulated drought stress. Besides, the pot experiment was performed to measure yield under water stress. The drought resistance was evaluated according to drought resistance index and the relationship between indexes at seedling stage and drought resistance index were also analyzed. The results indicated that the drought resistance and drought resistance index had significant relationship using PEG simulated water stress. It can be used to select the drought resistance of maize. Combined with identification in the field, it would be good to evaluate the drought resistance of maize. It is an ordinary, effective, low cost and short time method to analyze plenty samples using PEG simulated water stress to treat maize seedling and also be an accurate and quick method to evaluate maize drought resistance.
5. Eight inbred lines including Dan598, Liao618, K12, Shen136, Shen137, Dan340, Tie9010, P178 which had different drought resistance were studied. P1 P2 F1 F2 generations were builded according to 28 incomplete diallel crosses of 8 inbred lines using Griffing 4 methods, Shen136and Tie9010, and genetic analysis of drought resistance was studied. The result indicated that, additive effect had great effect on ear length, bare tip length,100 grain weight and kernels in ear row under water stress. While yield, plant height and ear position were determined by non-additive genes. Both of the general and narrow genetic ability of kernels in ear row, ear length and bare tip length were low which influnced by environment greatly. While both of the general and narrow genetic ability of 100 grain weight were high which less influnced by environment and were controlled by additive gene. The general genetic ability of yield, plant height and ear position were higher which influnced by environment greatly. Ear position was mainly affected by non-additive genes. While both of additive and non-additive genes had effect on yield and plant height. The optimum genetic model of drought resistance index was E-1. Drought resistance index was maily affected by two pairs of major gene and polygenes. The heritability of major gene was higher than polygenes which were 61.59%and 17.7%, respectively. Variability effection of major gene to drought resistance was larger than polygenes. Phenotypic variation of drought resistance controlled by major gene and polygenes was 79.32%, while the other 20.68% was controlled by environment.
1.以三个玉米杂交种及亲本自交系为试验材料,苗期和中后期进行中度水分胁迫,测定了一系列形态和生理生化指标。结果表明各杂交种间及各自交系间抗旱性差异明显。以抗旱系数进行排序,杂交种：铁单17)农大108)铁单10；亲本自交系：沈136>P178>丹340>铁9010>黄C>C8605-2。以抗旱指数进行排序,杂交种：铁单17)农大108>铁单10；亲本自交系：沈136>P178>铁9010>丹340>黄C>C8605-2。抗旱指数和抗旱系数具有相同的趋势。亲本及F1代间抗旱性在某些性状上有一定相关性,抗旱性强的自交系,其杂交种的抗旱性也较好,抗旱性弱的自交系,其杂交种抗旱性也相对较弱。利用隶属函数值法进行分析结果,其中自交系沈136苗期、花期平均抗旱隶属值最高,具有较强的抗旱性,P178、铁9010、铁单17、农大108、340属中等抗旱类型,C8605-2、黄C属弱抗旱类型,而铁单10属不抗旱类型。
2.中度水分胁迫处理玉米自交系和杂交种后,测定了一些农艺性状指标和生理生化指标,通过综合分析,结果表明：中度水分胁迫条件下,玉米植株叶片相对含水量、丙二醛含量和ASI与玉米子粒产量密切相关,能够比较准确地评价玉米自交系和杂交种抗旱性的强弱。
在中度水分胁迫条件下,利用上述几个相关指标对10个骨干自交系进行抗旱性测定,结果显示相对含水量、电导率、丙二醛与子粒产量呈极显著的正相关,ASI与子粒产量呈极显著负相关。说明在鉴定筛选玉米种植资源时,玉米植株叶片相对含水量、丙二醛含量、电导率和ASI与玉米子粒产量密切相关,能够比较准确地评价玉米自交系抗旱性的强弱,和抗旱系数、抗旱指数一样可以作为玉米育种耐旱自交系和杂交种筛选的鉴定指标。
3.进行玉米抗旱育种,首先要筛选抗旱资源,对种质资源进行抗旱性鉴定,选择抗旱性较好的材料作为玉米抗旱性育种的基础材料,鉴定有利的抗旱基因并加以利用。本文通过对一些骨干玉米自交系生理、形态指标和产量的测定,结果表明,玉米自交系间的耐旱性存在较大差异。其中,自交系沈136、齐319、丹598和沈137耐旱性较好,可以作为玉米耐旱育种的基础材料,进行抗旱自交系和杂交种的选育。
4.采用PEG处理模拟水分胁迫,能够达到与田间水分胁迫相似的效果。因此可以利用PEG在芽苗期对玉米幼苗进行模拟干旱处理,结合一些抗旱指标的测定,能够比较准确地评价玉米种质的抗旱性。本文采用PEG处理幼苗模拟水分胁迫,通过叶片丙二醛含量、电导率和叶片水势等几个相关生理指标测定,分析玉米自交系的抗旱性,同时结合盆栽试验进行干旱胁迫处理测定玉米自交系的产量,通过抗旱指数评价玉米自交系的抗旱性,并对苗期指标与抗旱指数进行了相关性分析,结果PEG处理幼苗模拟干旱胁迫鉴定玉米自交系的抗旱性与抗旱指数具有显著的相关性,因此苗期采用PEG处理模拟水分胁迫进行大量玉米种质的抗旱性筛选,结合田间干旱鉴定,可以更好地评价玉米种质的抗旱性。采用PEG模拟水分胁迫处理玉米幼苗,方法简单、有效、低成本、短周期就能够分析大量样品,解决了盆栽试验、田间试验进行水分干旱胁迫处理样品少,胁迫时间、胁迫强度难以控制等问题,是快速鉴定玉米种质抗旱性的比较准确的评价方法。
5.本研究利用耐旱性不同的丹598、辽618、K12、沈136、沈137、丹340、铁9010、P178,8个自交系按Griffing方法4组配的28个不完全双列杂交组合及自交系沈136和铁9010为亲本材料,配制的P1 P2 F1 F2四个世代群体,进行了抗旱性性状的遗传分析。研究结果表明,水分胁迫下穗长、秃尖、百粒重、行粒数这些性状受加性效应影响大,产量、株高和穗位性状受非加性基因作用较大。行粒数、穗长和秃尖的广义遗传力和狭义遗传力均较低,性状受环境影响大,百粒重性状的广义遗传力和狭义遗传力均较高,而且相差较小,性状受环境影响小,且为加性遗传,产量、株高和穗位性状广义遗传力较高,性状受环境影响较大。其中穗位主要受非加性效应控制,产量和株高加性效应和非加性效应均有影响。抗旱指数性状的最适遗传模型为E-1模型,抗旱性指数主要受2对主基因+多基因控制。其中主基因遗传力大于多基因遗传力,分别为61.59%和17.72%,主基因对抗旱指数的变异度的影响远远大于多基因对抗旱指数的变异度的影响,而且主基因+多基因决定了抗旱指数表型变异的79.32%,另外有20.68%是由环境因素决定的。
Morphology and physiological indexes were studied to identify drought resistance of maize according to pot experiment, filed experiment and lab simulated water stress using different maize inbred lines and hybrids with different drought resistance. The grey correlation analysis method and membership function value method were used to establish the maize drought resistance identification system. Duncan's new multiple range test method was used to significant test. According to Griffing double row hybrid general combining ability statistical principle, the test samples were P(P-1)/2 orthogonal combinations and DPS were used for data analysis to calculate the GCA, SCA and genetic parameter of maize inbred lines. The gene effect controlled maize grain water content and genetic law of maize drought resistance of maize were analyzed using Mather's A、B、C scale inspection and analysis of variance method. The main results were as follows:
1. Morphology and physiological and biochemical indexes of three maize hybrids and their inbred lines were studied during moderate water stress at seedling stage and anthesis. The results indicated that the drought resistance of maize hybrids and inbred lines had significant differences. The drought resistance coefficient ordering of maize hybrids were as follows: Tiedan17> Nongda108> Tiedan 10; while the drought resistance coefficient ordering of inbred lines were as follows:Shen136> P178> Dan340> Tie 9010> Huang C> C8605-2. The drought resistance index and coefficient had same trend. The drought resistance of parents and F1 had some correlations on some traits. The drought resistance of inbred line was higher, the corresponding hybrid was better. According to membership function value method, the result indicated that Shen 136 which had the highest membership value at seedling stage and anthesis had higher drought resistance; P178, Tie9010, Tiedan17, Nongda108 and 340 were medium drought resistance type; C8605-2 and Huang C were weak drought resistance type; while Tiedan10 was sensitive drought resistance type.
2. Agronomic characters and physiological and biochemical indexes of maize hybrids and inbred lines were measured under moderate water stress. The results showed that the relative water content, MDA and ASI closely related with maize yield. These indexes could be accurately to evaluate the drought resistance of maize hybrids and inbred lines.
The drought resistance of ten ordinary maize inbred lines was studied using above related indexes. The results indicated that the relative water content, MDA content, electric conductivity of maize leaves were significantly correlated to the yield of maize under moderate drought stress. While there was a significant negative correlation between ASI and yield. It showed that these physiological indexes of leaves could accurately evaluate the drought resistance of maize and could be selection and identification index of maize inbred lines in maize drought resistance breeding as well as drought resistance coefficient and drought resistance index.
3. Selecting and identification of drought resistance germplasm resources was the primary problem to maize drought resistance breeding. Physiology and morphological indexes and yield of some ordinary maize inbred lines were measured in this study. The results indicated that the drought resistance of maize inbred lines had significant differences. The drought resistance of Shen136, Qi319, Dan598 and Shen137 were better and these inbred lines could be basic material as maize drought resistance breeding.
4. The maize using PEG simulated drought stress could reach the same effect with the field water stress. Hence the maize which treated using PEG simulated drought stress at seedling stage and combined with drought resistance indexes could be accurately evaluate the drought resistance of maize. MDA, electric conductivity and water potential were measured to analysis the drought resistance of maize inbred lines using PEG simulated drought stress. Besides, the pot experiment was performed to measure yield under water stress. The drought resistance was evaluated according to drought resistance index and the relationship between indexes at seedling stage and drought resistance index were also analyzed. The results indicated that the drought resistance and drought resistance index had significant relationship using PEG simulated water stress. It can be used to select the drought resistance of maize. Combined with identification in the field, it would be good to evaluate the drought resistance of maize. It is an ordinary, effective, low cost and short time method to analyze plenty samples using PEG simulated water stress to treat maize seedling and also be an accurate and quick method to evaluate maize drought resistance.
5. Eight inbred lines including Dan598, Liao618, K12, Shen136, Shen137, Dan340, Tie9010, P178 which had different drought resistance were studied. P1 P2 F1 F2 generations were builded according to 28 incomplete diallel crosses of 8 inbred lines using Griffing 4 methods, Shen136and Tie9010, and genetic analysis of drought resistance was studied. The result indicated that, additive effect had great effect on ear length, bare tip length,100 grain weight and kernels in ear row under water stress. While yield, plant height and ear position were determined by non-additive genes. Both of the general and narrow genetic ability of kernels in ear row, ear length and bare tip length were low which influnced by environment greatly. While both of the general and narrow genetic ability of 100 grain weight were high which less influnced by environment and were controlled by additive gene. The general genetic ability of yield, plant height and ear position were higher which influnced by environment greatly. Ear position was mainly affected by non-additive genes. While both of additive and non-additive genes had effect on yield and plant height. The optimum genetic model of drought resistance index was E-1. Drought resistance index was maily affected by two pairs of major gene and polygenes. The heritability of major gene was higher than polygenes which were 61.59%and 17.7%, respectively. Variability effection of major gene to drought resistance was larger than polygenes. Phenotypic variation of drought resistance controlled by major gene and polygenes was 79.32%, while the other 20.68% was controlled by environment.
引文
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