普通玉米4个籽粒性状的遗传分析
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摘要
本试验以籽粒长度、籽粒容重、出籽率等籽粒性状差异明显的6个自交系为供试材料，按6世代遗传交配设计，利用6世代均数分析、6世代遗传方差分析及6世代联合分析等数量性状分析方法，对籽粒长度、籽粒比重、HKWI和出籽率等与产量和商品品质有关的籽粒性状进行了遗传机制研究，取得以下主要研究结果：
(1)根据6世代均数分析和遗传方差分析结果，4个籽粒性状在多数组合中不符合加性—显性模型，存在上位性效应，其加性效应和显性效应均显著存在，加性效应除籽粒长度在多数组合中表现为减效，其余性状均表现为增效；4个籽粒性状的显性效应均为增效，且显性效应大于加性效应；上位性效应则因材料而异。上述4个籽粒性状的广义遗传率在多数组合中表现较高，狭义遗传率则表现出较大的差异，且出籽率在多数组合中表现为超显性遗传，而其余3个籽粒性状则表现为部分显性，但籽粒比重和HKWI已接近完全显性。
(2)根据6世代联合分析结果，除籽粒比重外的3个籽粒性状的最适模型为主基因+多基因混合遗传模型，且在多数组合中表现为2对主基因+多基因混合遗传。其中籽粒长度和HKWI检测到的2对主效基因的加性效应为减效，显性效应为增效，且数值相当，并加性效应大于显性效应，互作效应中以1效应和i效应较为明显，作用方向为负向，j_(ab)效应和j_(ba)效应作用较小，作用方向为正向。多基因的加性效和显性效应均为减效，且显性效应大于加性效应，3种上位性效应也较为明显，且[i]效应为减效，[j]效应和[1]效应为增效。出籽率虽然在3个组合中差异较大，但总的来看，该性状检测到的2对主效基因的显性效应均为增效，且加性效应大于显性效应，上位性效应以1效应较为明显，且作用方向为负，多基因的加性效应和显性效应均为减效，且显性效应大于加性效应。此外，3个籽粒性状(籽粒长度、HKWI和出籽率)的主基因遗传率在不同组合的不同世代中存在明显差异，高者可达80％以上，低者仅为10％左右。但总的看来，籽粒长度的主基因遗传率中等偏低，而HKWI和出籽率则表现较高。多基因遗传率同样在不同组合的不同世代中同样存在明显差异，似乎籽粒长度的多基因比主基因所起的作用要大一些，而HKWI和出籽率则多基因比主基因所起的作用要小一些。
(3)比较4个籽粒性状在不同组合中的遗传表现，结果发现，4个籽粒性状在不同遗传背景下表现出基因型间的差异。而且也存在一定的规律性。从6世代均数分析和遗传方差分析的结果看，其差异主要体现在加性效应的作用程度和作用方
向、上位性效应是否存在、上位性效应存在时的作用程度和作用方向以及遗传率的
大小等方面。而从6世代联合分析的结果看，其差异主要体现在主效基因是否存在、
主效基因的数量、主效基因加性效应和上位性效应的作用程度和作用方向以及多基
因上位性效应的作用程度和作用方向等方面。
(4)根据4个籽粒性状的遗传特性，可为制定育种方案时提供参考。对于4
个籽粒性状的选育，通过亲本的选择是有效的，但用于选育亲本的基础群体籽粒性
状必须表现优良;选育二环系是选育籽粒性状优良自交系的比较有效的方法。籽粒
长度、HKWI、出籽率等籽粒性状，可在较低世代进行单株选择，但籽粒长度的选
择标准应适当放宽一些;而籽粒比重，应采用混合选择的方法或在较高世代进行单
株选择，且选择标准应适当放宽。对于自交系的改良，籽粒长度、HKWI和出籽率
等籽粒性状，可通过回交转育将外源有利基因导入而予以改良。籽粒比重可通过聚
合回交法加以改良。
在亲本选择的同时，还应注意亲本间的组配，以利用上述籽粒性状的显性效应
和上位性效应，充分发挥杂种优势的增产作用，且必须始终坚持以高配合力为核心，
以血缘关系和杂种优势为主线，注重杂种优势群和杂种优势模式的挖掘和利用。
(5)比较本文结果与前人研究的结果，发现4个籽粒性状的遗传比较复杂，
并且还可能存在本试验无法检测到的其它遗传效应，因此尚需作进一步研究。一方
面可采用不同的遗传交配设计和试验方法，探索在不同交配方式下籽粒性状的遗传
机制。另一方面，可利用分子标记对4个籽粒性状进行QTL定位，对遗传试验结
果作进一步的解释和验证。此外，对于主效基因遗传机制的探讨，本试验采用的遗
传试验检测方法(6世代联合方法)与分子标记检测方法(QTL定位)各有优缺点，
可否将两种分析方法相互结合，共同发展，也有待作进一步的探讨。
In the study the material were 6 inbred lines which were obvious diversity in kernel traits, such as kernel length, kernle desity, helling percentage etc. According to 6 generation genetic mating design, we had made genetic mechanism researches on four characters, such as kernel length, kernel desity, HKWI, and helling percentage, with combing quantitative-character genetic analyzing methods such as 6 generation mean analysis, 6 generation genetic variance analysis, and 6 generation joint analysis. The main results are followed.
(1)According to the result of 6 generation mean analysis and 6 generation genetic variance analysis, we found that 4 kernel traits didn't accord with additive-domiance genetic model, and exited epistasis effect. It was obvious exit additive effect and domiance effect. The values of additive effect of kernel length were negative in most generation, other kernel traits were positive in most generation. The values of domiance effect of 4 kernel traits were positive, and larger than additive effect. The values of epistasis effect were diversity because of different materials. The heritability in broad sense of 4 kernel traits was high, but the heritability in narrow sense of 4 kernel traits was diversity in different materials. Helling percentage was super-dominant, other kernel traits were additive, but kernel desity and HKWI were near to daominant.
(2)According to 6 generation joint analysis result,. The most suitable genetic model of 4 kernel traits was two-major gene inheritance model in most generation execpt of kernel desity. The values of additive effect were negative ,but the domiance effect were positive of the two major gene in kernel length and HKWI. The values of 1 effect and i effect were negative, and more important than jab effect and jba effect., positive. In polygene, the values of additive effect and domiance effect were negative, and larger than additive. It was important to be 3 epistasis effects. The values of [i] effect was negative, but [j] effect and [1] effect were positive. The values of additive effect and domiance effect were negative, and domiance effect was larger than additive effect. In addition, it was diversity about major gene heritability of 3 kernel traits in different generations of different combinations, from 80% to 10%, but looking totally, the major
gene heritability of kernel length was lower than HKW1 and helling percentage. The polygene heritability of 4 kernel traits were diversity in different generation of different combinations, too. The function of the polygene heritability was more important than the major heritability in kernel length, on the contrary, the function of the major heritability was more important than polygene heritability in other 2 kernel traits.
(3)Comparing the genetic characters of 4 kernel traits in different combinations. We found that the genotype of 4 kernel traits was diversity in different heredity backgrounds, and follow any regulation. From the 6 generation mean analysis and the 6 generation genetic variance analysis, the mainly diversity was function and degree of additive effect, exiting epistasis effect or not, function and degree of epistasis effect, and size of gene heritability, etc. From the 6 generation joint analysis, the mainly diversity was exiting major gene, quantity of the major gene, function and degree of additive effect and domiance effect in major gene, and function and degree of epistasis effect in polygene, etc.
(4)According to the genetic characters of 4 kernel traits,. Selecting the parents is availabe, but the base population must to be eminent. Selecting second cycle lines is a good way to select eminent inbred lines about kernel traits. Some kernel traits such as kernel length, HKWI, and helling percentage, should be individual selection in lower generations, but the stand of selection must be looseness adequately. Kernel desity should be bulk selection in higher generations, but the stand of selection must be looseness. For improved inbred lines, some kernel traits such as kernel length,
(1)根据6世代均数分析和遗传方差分析结果，4个籽粒性状在多数组合中不符合加性—显性模型，存在上位性效应，其加性效应和显性效应均显著存在，加性效应除籽粒长度在多数组合中表现为减效，其余性状均表现为增效；4个籽粒性状的显性效应均为增效，且显性效应大于加性效应；上位性效应则因材料而异。上述4个籽粒性状的广义遗传率在多数组合中表现较高，狭义遗传率则表现出较大的差异，且出籽率在多数组合中表现为超显性遗传，而其余3个籽粒性状则表现为部分显性，但籽粒比重和HKWI已接近完全显性。
(2)根据6世代联合分析结果，除籽粒比重外的3个籽粒性状的最适模型为主基因+多基因混合遗传模型，且在多数组合中表现为2对主基因+多基因混合遗传。其中籽粒长度和HKWI检测到的2对主效基因的加性效应为减效，显性效应为增效，且数值相当，并加性效应大于显性效应，互作效应中以1效应和i效应较为明显，作用方向为负向，j_(ab)效应和j_(ba)效应作用较小，作用方向为正向。多基因的加性效和显性效应均为减效，且显性效应大于加性效应，3种上位性效应也较为明显，且[i]效应为减效，[j]效应和[1]效应为增效。出籽率虽然在3个组合中差异较大，但总的来看，该性状检测到的2对主效基因的显性效应均为增效，且加性效应大于显性效应，上位性效应以1效应较为明显，且作用方向为负，多基因的加性效应和显性效应均为减效，且显性效应大于加性效应。此外，3个籽粒性状(籽粒长度、HKWI和出籽率)的主基因遗传率在不同组合的不同世代中存在明显差异，高者可达80％以上，低者仅为10％左右。但总的看来，籽粒长度的主基因遗传率中等偏低，而HKWI和出籽率则表现较高。多基因遗传率同样在不同组合的不同世代中同样存在明显差异，似乎籽粒长度的多基因比主基因所起的作用要大一些，而HKWI和出籽率则多基因比主基因所起的作用要小一些。
(3)比较4个籽粒性状在不同组合中的遗传表现，结果发现，4个籽粒性状在不同遗传背景下表现出基因型间的差异。而且也存在一定的规律性。从6世代均数分析和遗传方差分析的结果看，其差异主要体现在加性效应的作用程度和作用方
向、上位性效应是否存在、上位性效应存在时的作用程度和作用方向以及遗传率的
大小等方面。而从6世代联合分析的结果看，其差异主要体现在主效基因是否存在、
主效基因的数量、主效基因加性效应和上位性效应的作用程度和作用方向以及多基
因上位性效应的作用程度和作用方向等方面。
(4)根据4个籽粒性状的遗传特性，可为制定育种方案时提供参考。对于4
个籽粒性状的选育，通过亲本的选择是有效的，但用于选育亲本的基础群体籽粒性
状必须表现优良;选育二环系是选育籽粒性状优良自交系的比较有效的方法。籽粒
长度、HKWI、出籽率等籽粒性状，可在较低世代进行单株选择，但籽粒长度的选
择标准应适当放宽一些;而籽粒比重，应采用混合选择的方法或在较高世代进行单
株选择，且选择标准应适当放宽。对于自交系的改良，籽粒长度、HKWI和出籽率
等籽粒性状，可通过回交转育将外源有利基因导入而予以改良。籽粒比重可通过聚
合回交法加以改良。
在亲本选择的同时，还应注意亲本间的组配，以利用上述籽粒性状的显性效应
和上位性效应，充分发挥杂种优势的增产作用，且必须始终坚持以高配合力为核心，
以血缘关系和杂种优势为主线，注重杂种优势群和杂种优势模式的挖掘和利用。
(5)比较本文结果与前人研究的结果，发现4个籽粒性状的遗传比较复杂，
并且还可能存在本试验无法检测到的其它遗传效应，因此尚需作进一步研究。一方
面可采用不同的遗传交配设计和试验方法，探索在不同交配方式下籽粒性状的遗传
机制。另一方面，可利用分子标记对4个籽粒性状进行QTL定位，对遗传试验结
果作进一步的解释和验证。此外，对于主效基因遗传机制的探讨，本试验采用的遗
传试验检测方法(6世代联合方法)与分子标记检测方法(QTL定位)各有优缺点，
可否将两种分析方法相互结合，共同发展，也有待作进一步的探讨。
In the study the material were 6 inbred lines which were obvious diversity in kernel traits, such as kernel length, kernle desity, helling percentage etc. According to 6 generation genetic mating design, we had made genetic mechanism researches on four characters, such as kernel length, kernel desity, HKWI, and helling percentage, with combing quantitative-character genetic analyzing methods such as 6 generation mean analysis, 6 generation genetic variance analysis, and 6 generation joint analysis. The main results are followed.
(1)According to the result of 6 generation mean analysis and 6 generation genetic variance analysis, we found that 4 kernel traits didn't accord with additive-domiance genetic model, and exited epistasis effect. It was obvious exit additive effect and domiance effect. The values of additive effect of kernel length were negative in most generation, other kernel traits were positive in most generation. The values of domiance effect of 4 kernel traits were positive, and larger than additive effect. The values of epistasis effect were diversity because of different materials. The heritability in broad sense of 4 kernel traits was high, but the heritability in narrow sense of 4 kernel traits was diversity in different materials. Helling percentage was super-dominant, other kernel traits were additive, but kernel desity and HKWI were near to daominant.
(2)According to 6 generation joint analysis result,. The most suitable genetic model of 4 kernel traits was two-major gene inheritance model in most generation execpt of kernel desity. The values of additive effect were negative ,but the domiance effect were positive of the two major gene in kernel length and HKWI. The values of 1 effect and i effect were negative, and more important than jab effect and jba effect., positive. In polygene, the values of additive effect and domiance effect were negative, and larger than additive. It was important to be 3 epistasis effects. The values of [i] effect was negative, but [j] effect and [1] effect were positive. The values of additive effect and domiance effect were negative, and domiance effect was larger than additive effect. In addition, it was diversity about major gene heritability of 3 kernel traits in different generations of different combinations, from 80% to 10%, but looking totally, the major
gene heritability of kernel length was lower than HKW1 and helling percentage. The polygene heritability of 4 kernel traits were diversity in different generation of different combinations, too. The function of the polygene heritability was more important than the major heritability in kernel length, on the contrary, the function of the major heritability was more important than polygene heritability in other 2 kernel traits.
(3)Comparing the genetic characters of 4 kernel traits in different combinations. We found that the genotype of 4 kernel traits was diversity in different heredity backgrounds, and follow any regulation. From the 6 generation mean analysis and the 6 generation genetic variance analysis, the mainly diversity was function and degree of additive effect, exiting epistasis effect or not, function and degree of epistasis effect, and size of gene heritability, etc. From the 6 generation joint analysis, the mainly diversity was exiting major gene, quantity of the major gene, function and degree of additive effect and domiance effect in major gene, and function and degree of epistasis effect in polygene, etc.
(4)According to the genetic characters of 4 kernel traits,. Selecting the parents is availabe, but the base population must to be eminent. Selecting second cycle lines is a good way to select eminent inbred lines about kernel traits. Some kernel traits such as kernel length, HKWI, and helling percentage, should be individual selection in lower generations, but the stand of selection must be looseness adequately. Kernel desity should be bulk selection in higher generations, but the stand of selection must be looseness. For improved inbred lines, some kernel traits such as kernel length,
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