陆地棉种子品质性状遗传和蛋白质QTL定位研究
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摘要
棉籽是棉花生产中的最重要的副产品,应用于食用油、饲料和燃料等领域,并且是潜在的食用蛋白质源。随着世界人口的迅速增长,棉籽作为人类食物资源的潜能将越大。因此,棉籽品质性状的改良将成为棉花育种改良的重要目标之一。全面深入地研究棉籽品质性状的遗传特性及QTL定位将对于棉籽品质的遗传改良具有重要的理论和指导意义。本研究以具有氨基酸含量差异的445份棉籽为材料,构建了17种氨基酸的近红外定标模型。以188份陆地棉重组近交系为材料,按照不完全双列杂交方式,创建了永久F2群体(IF2群体)。采用包括基因型和基因型×环境互作的双子叶植物二倍体种子数量性状遗传模型,基于胚、母体、细胞质三套遗传体系,研究了陆地棉种子品质性状的遗传特性。同时,采用新发展的包括环境互作效应在内的多遗传体系QTL作图方法和基因定位软件,对棉籽蛋白质等性状进行了胚和母体植株不同遗传体系的QTL定位分析。主要研究结果如下：
1棉籽氨基酸含量的NIRS定标模型构建：本研究采用一阶导数的数学处理(1,4,4,1)、标准正态变换和去趋势(SNV+D)最佳组合的预处理方法,结合改良的偏最小二乘法(MPLS),创建了棉仁粉中17种氨基酸含量的近红外反射光谱(NIRS)定标模型。天冬氨酸、苏氨酸等12种氨基酸含量的定标模型较好,其相对分析误差(RPDc)为3.735～7.132,外部检验的决定系数(r2)为0.910～0.979,完全可以替代化学测定；而丝氨酸、蛋氨酸、酪氨酸和脯氨酸等4种氨基酸的定标模型预测效果略差,其RPDc为2.205～2.814,外部检验r2为0.800～0.830,但仍可用于棉仁粉中氨基酸含量的测定。半胱氨酸定标方程的RPDc较小(RPDc=1.358).因此,棉仁粉中半胱氨酸含量不能用近红外分析方法进行检测。氨基酸含量的NIRS定标模型为陆地棉种仁氨基酸含量测定提供一种快速、低廉、准确可靠的分析方法。
2IF2群体的构建：基于来自陆地棉品种间杂交组合(HS46xMARCABUCAG8US-1-88的188个重组近交系,在重组近交系间进行随机交配,创建了一套包括了376个组合的陆地棉品种间“永久F2”群体。该群体中的重要棉籽品质性状均表现出典型的数量性状特点,呈正态分布,并且各性状出现超亲分离。因此,该群体可作为优良的遗传研究和育种资源。
3棉籽品质性状的遗传特性分析：基于上述IF2群体的种子品质性状表型数据,对陆地棉7个重要种子品质性状进行了遗传方差、遗传率和遗传相关分析。
(1)遗传方差分析表明,子指、仁／壳的基因表达主要受胚加性和母体加性主效应控制,其次受细胞质×环境互作效应控制；仁指的基因表达主要受胚加性和母体加性效应控制；种仁率的基因表达主要受胚加性效应控制,其次受胚显性×环境互作效应和母体加性×环境互作效应；油分含量的基因表达主要受细胞质主效应控制,其次受胚加性×环境互作效应控制。以上说明子指、仁／壳、种仁率、油分含量和仁指的变异主要来自基因型本身的作用,但环境因素不可忽视。蛋白质含量的基因表达主要受种子胚加性×环境互作效应和种子胚显性×环境互作效应控制,其次受母体加性和母体显性主效应控制；棉酚含量的基因表达主要受胚显性x环境互作效应和母体加性x环境互作效应控制,其次受母体加性主效应控制。以上结果说明这两个性状的变异易遭受很大的环境影响,因此,在性状改良时,必需考虑环境因素。
(2)遗传率分析表明,子指、仁／壳、仁指和蛋白质含量总狭义遗传率较低,这些性状在中高代进行选择可以取得更好的选择效果；棉酚、油分含量总狭义遗传率较高,在早世代进行性状选择能取得一定效果。子指、种仁率、仁／壳、仁指、棉酚含量、油分含量以普通狭义遗传率为主,其中,子指、仁指、棉酚含量以普通母体遗传率为主；种仁率、仁／壳和油分含量以普通细胞质遗传率为主；蛋白质含量以互作细胞质遗传率为主。在育种改良中,子指、仁指、棉酚含量、仁／壳和油分含量可以根据母体植株的总体表现和单粒选择相结合的方法在低世代进行选择。由于棉籽蛋白质含量具有互作遗传率高的特点,其在育种改良中虽也可根据母体植株的总体表现在低世代进行单株选择,但一般与特定环境有密切关系。
(3)遗传相关性分析表明,种仁率与棉酚、仁／壳以胚加性负相关为主；子指与仁／壳、棉酚含量、种仁率、仁指、蛋白质含量,种仁率与蛋白质含量,仁／壳与蛋白质含量,油分与蛋白质含量,仁指与蛋白质、棉酚含量以母体加性负相关为主；仁／壳与仁指、棉酚含量、种仁率以胚加性、细胞质、母体加性负相关为主；棉酚与油分含量以胚加性正相关为主；棉酚与蛋白质含量以细胞质正相关为主；种仁率与仁指以胚加性和母体加性正相关为主。因此,上述成对性状可以在早世代进行单粒或单株间接选择。油分含量与子指、仁／壳、种仁率和仁指未发现明显的相关性,但仍有可能通过这些物理品质性状间接改良油分含量。
4棉籽蛋白质及氨基酸含量的QTL定位：以来自陆地棉品种间杂交组合(HS46×MARCABUCAG8US-1-88的188个重组近交系随机交配获得的“永久F2"群体为材料,采用本实验室构建的重组自交系分子连锁图谱,应用新发展的程序QTL Network-CL-2.0-Seed对“永久F2"群体在两个环境下测得的蛋白质及氨基酸含量进行多遗传体系QTL定位及其遗传分析。共检测到6个控制蛋白质含量的QTL,解释了58.5%的表型变异,其中,2个QTL主要在胚核基因组上表达,其余4个QTL同时在胚和母体核基因组上表达,3个QTL具有明显的环境互作效应。在3个胚加性效应贡献率大于5%的QTL中,2个QTL已定位在具体的染色体上,为蛋白质含量的分子育种提供重要基础。
本试验还共检测了控制16种氨基酸含量性状表现的QTL在胚和母体植株2套核基因组上的分布情况：包括亮氨酸(Leu)、苯丙氨酸(Phe)、苏氨酸(Thr)、缬氨酸(Val)、蛋氨酸(Met)、异亮氨酸(Ile)、赖氨酸(Lys)、天冬氨酸(Asp)、丝氨酸(Ser)、谷氨酸(Glu)、甘氨酸(Gly)、丙氨酸(Ala)、酪氨酸(Tyr)、组氨酸(His)、精氨酸(Arg)、脯氨酸(Pro)含量。研究结果发现控制这些性状的QTL数目分别为2-8个,可以解释表型变异为23.37%-64.32%。所检测到的每种氨基酸QTL中,在胚核基因组上表达的QTL分别有0-2个,其中,苯丙氨酸、缬氨酸、蛋氨酸、酪氨酸和组氨酸各为2个,天冬氨酸和甘氨酸未发现仅在胚核基因组中表达的QTL；在母体植株核基因组中表达的分别有0-2个,其中,异亮氨酸为2个,缬氨酸、天冬氨酸和组氨酸各为1个,其余的12种氨基酸未发现仅在母体基因组中表达的QTL。在胚核基因组和母体植株核基因组中同时表达的分别有0-6个,其中苯丙氨酸为6个,组氨酸为0个；有显著的环境互作效应的有1-5个,其中苏氨酸、天冬氨酸、谷氨酸、甘氨酸和丙氨酸各为1个,苯丙氨酸为6个。在七种必需氨基酸中,具有良好的定位效果的为苯丙氨酸(8个QTL,共解释的表型变异为54.5%)、缬氨酸(6个QTL,共解释的表型变异为64.32%)、异亮氨酸(6个QTL,共解释的表型变异为60.95%),这些QTL在相应的性状改良中将可发挥重要的作用。
Cotton (Gossypium spp.) is one of important economic crops in the world. The fiber which is its main product is used in the textile industry while the cottonseed, which is a byproduct is widely used in making food, animal feed, fuel and other products. Cottonseed contains many kinds of nutrients, making it a potential food-source for humans as the world population continues to rapidly grow. Cottonseed improvement has therefore become one of the major objectives in cotton breeding requiring a more detailed understanding of the genetic mechanisms in cottonseed for effective manipulation. The aims of the present study are the following:(1) Studying the inheritance of upland cotton seed traits, which include determination of genetic effects, heritability and correlation based on the embryo genome, the maternal plant genome and the plasmon. It was conducted with genetic models that were developed purposely for quantitative traits in diploid plant seed and their corresponding statistical approaches using two years data from the188inbred lines and their hybrids, all of which were cultivated within a partial diallel mating design;(2) Identifying and analyzing the QTL for protein-related traits based on embryo genome and maternal plant genome. An "immortalized F2"(IF2) population including376crosses was adopted, for this study which was conducted using the newly developed QTL mapping method that includes embryo and maternal main effects as well as their genetype×environment (GE) interaction effects on quantitative traits of dicotyledonous seeds. The main results were summarized as follows:
1Calibration equations were developed with near infrared spectroscopy (NIRS) for amino acid content in cottonseeds using a total of445samples with a high variability in amino acid contents were used for developing calibration equations of amino acid content of cottonseeds. The spectral data of cotton kernel powder was processed using the first derivative mathematical treatment combined with SNV and de-trend, as well as Modified Partial Least Squares (MPLS) regression method. Chemometric models for17amino acids present in cottonseed were developed, and12of them were found to be excellent for determination of related amino acids including asparagic acid, threonine, glutamic acid, etc, with-the ratio of SD to SECV in the calibration set (RPDc) of3.735～7.132and adetermination coefficient (r2) of0.910～0.979in external validation. For these12amino acids, their values obtained by the chemical method were accurately predicted using NIRS. The RPDc of serine, methionine, tyrosine and proline ranged between2.205and2.814, and they determination coefficient (r2) were from0.800to0.830in external validation. For these4amino acids, the NIRS values were not as accurate as those from the chemical analysis, but could be used for sample screening in cotton breeding program. The equation for cystine however was not reliable as its RPDc was only1.358, which was not suitable to estimate its content in cottonseeds.
2An immortal F2population (IF2) comprising of376crosses was constructed using a set of recombinant inbred lines (RIL) derived from the hybrid of HS46XMARCABUCAG8US-1-88. In this population, most cottonseed quality traits including seed index (SI), kernel index (KI), kernel percentage (KP), kernel/hull (K/H), protein content (PC), oil content (OC), gossypol content (GC) and17amino acids contents have characters of quantitative traits with normal distribution. In addition, means of most traits in some crosses of the IF2population were beyond those of the parents (HS46and MARCABUCAG8US-1-88). This population thus is suitable resource for genetic studies and breeding.
3Based on phenotypic data of cottonseed in188recombinant inbred lines and their IF2population, genetic variances, heritability and correlation analysis of7important cottonseed quality traits were conducted.
(1) The results of variance analysis showed:SI, KI and K/H were mainly affected by embryo additive and maternal additive effects. Secondly SI and K/H were affected by cytoplasmatic interaction; KP was mainly affected by embryo additive effect, and on a lesser note by embryo dominance and maternal additive interaction effects. OC was predominantly affected by cytoplasmic effects, and secondly by embryo additive interaction effects; The phenotypic variance of the above mentioned traits was mainly atributed to their genetypes, although environmental factors were not neglected. PC was mainly controlled by embryo dominance and maternal additive interaction effects, secondly by maternal additive and dominance main effects; GC was mainly affected by embryo dominance×environment interaction effects, also by maternal additive effect. The phenotypic variance of these two traits could be more affected by environment. The environmental factor needed to be considered in the improvement of these traits.
(2) The results of heritability analysis showed:The total narrow-sense heritabilities for GC and OC were relatively high, it was thus feasible to select these traits in early generation; those for SI, K/H, KI and PC were relatively low, then these traits were not suitable for selection in early generation. Maternal general heritability was the most important for SI, KI and GC. Cytoplasmic heritability was the main parts for K/H, KP and OC; Selection advance for SI, K/H, OC, KI and GC could therefore be expected in the early generation through the combination of the performance of maternal plant and single seed selection. While for PC, it was cytoplasmic interaction heritability, and improving PC would be more efficient in the early generation by single plant selection based on the performance of maternal plant under specific environment.
(3) The analysis of genetic correlation showed:The relationship between KI and GC, KI and K/H were mainly controlled by embryo additive effect, which showed a significantly negative correlation; Significantly negative maternal additive correlation were the main part between SI and K/H, SI and GC, SI and KI, SI and/KP, SI and PC, KP and PC, K/H and PC, OC and PC, KI and PC, KI and GC. The relationship between K/H and KI, K/H and GC, K/H and KP were mainly controlled by embryo additive, cytoplasmic and maternal additive effects, which showed significantly negative correlation; Significantly positive embryo additive correlation was main part between GC and OC; Significantly positive cytoplasmic correlation was main part between GC and PC. Significantly positive embryo and maternal additive correlation were main part between KP and KI. Therefore, direct selection for above pair traits could be conduceted in the early generation by single seed or plant selected. The relationship was not significant between OC and SI, OC and K/H, OC andKP, OC and KI, and it was not reliable to get high oil content in cottonseed through physical quality trait selected
4QTL mapping of cottonseed protein and amino acid contents were conducted on the immortal F2population obtained from the random mating among the188recombinant inbred lines derived from the cross of two upland cotton cultivars (HS46×MARCABUCAG8US-1-88). A molecular linkage map of recombinant inbred lines constructed in our laboratory and a newly developed program called the QTL Network-CL-2.0-Seed was used for QTL analysis for protein and amino acid contents based on multiple genetic systems in two different environments. Six QTL were detected for protein content explaining58.5%of the phenotypic variation. Among them, two QTL were expressed in the embryo, While the remaining four QTL were expressed in the embryo and maternal genomes. Three of the six QTL had environmental interaction effects. Among three QTL with more than5%embryo additive heritability, two QTL were mapped on particular chromosome. This indicated these two QTL were major for improving protein content in cottonseed.
QTL for sixteen amino acid including phenylalanine (phe), lysine (Lys), leucine (Leu), isoleucine (ILe), methionine (Met), threnine (Thr), valine (Val), aspartic acid (Asp), alanine (Ala), proline(Pro), serine(Ser), tyrosine(Try), histidine(His), argnine (Arg), glutamic acid (Glu) and glycine (Gly) contents were identified. The number of QTL for these traits ranged from2to8, explaining23.37%～64.32%of the phenotypic variance. Among these QTL, the range between0and2QTL were expressed in embryo genome. No QTL for Asp and Gly contents was expressed in embryo genome.Two QTL for Phe, Val, Met, Tyr and His contents were expressed in the embryo genome, respectively. Among these QTL for the sixteen aimno acid contents, the range between0and2QTL were merely expressed in maternal genome. Two for ILe and one QTL for Val, Asp and His contents were merely expressed in maternal genome, respectively. No QTL for the remaining twelve amino acid contens was merely expressed in maternal genome. Among the QTL for the sixteen amino acid contents, the range between0and6QTL were simultaneouly expressed in embryo and maternal genomes. No QTL for His content was simultaneouly expressed in embryo and maternal genomes. Six QTL for Phe content were simultaneouly expressed in the embryo and maternal genomes. The number of QTL with significant environmental interaction effect varied from1to5QTL for sixteen amino acid contents. One for Thr, Asp, Glu and Gly and Six QTL for Phe content had environmental interaction effects, respectively. The results of mapping QTL for Phe, Val and Ile content were excellent. A total of eight QTL for Phe content, six QTL for Val and Ile contents explained54.5%,64.32%and60.95%of phenotypic variance, respectively. These QTL could play an important role for the improvement of their corresponding traits.
1棉籽氨基酸含量的NIRS定标模型构建：本研究采用一阶导数的数学处理(1,4,4,1)、标准正态变换和去趋势(SNV+D)最佳组合的预处理方法,结合改良的偏最小二乘法(MPLS),创建了棉仁粉中17种氨基酸含量的近红外反射光谱(NIRS)定标模型。天冬氨酸、苏氨酸等12种氨基酸含量的定标模型较好,其相对分析误差(RPDc)为3.735～7.132,外部检验的决定系数(r2)为0.910～0.979,完全可以替代化学测定；而丝氨酸、蛋氨酸、酪氨酸和脯氨酸等4种氨基酸的定标模型预测效果略差,其RPDc为2.205～2.814,外部检验r2为0.800～0.830,但仍可用于棉仁粉中氨基酸含量的测定。半胱氨酸定标方程的RPDc较小(RPDc=1.358).因此,棉仁粉中半胱氨酸含量不能用近红外分析方法进行检测。氨基酸含量的NIRS定标模型为陆地棉种仁氨基酸含量测定提供一种快速、低廉、准确可靠的分析方法。
2IF2群体的构建：基于来自陆地棉品种间杂交组合(HS46xMARCABUCAG8US-1-88的188个重组近交系,在重组近交系间进行随机交配,创建了一套包括了376个组合的陆地棉品种间“永久F2”群体。该群体中的重要棉籽品质性状均表现出典型的数量性状特点,呈正态分布,并且各性状出现超亲分离。因此,该群体可作为优良的遗传研究和育种资源。
3棉籽品质性状的遗传特性分析：基于上述IF2群体的种子品质性状表型数据,对陆地棉7个重要种子品质性状进行了遗传方差、遗传率和遗传相关分析。
(1)遗传方差分析表明,子指、仁／壳的基因表达主要受胚加性和母体加性主效应控制,其次受细胞质×环境互作效应控制；仁指的基因表达主要受胚加性和母体加性效应控制；种仁率的基因表达主要受胚加性效应控制,其次受胚显性×环境互作效应和母体加性×环境互作效应；油分含量的基因表达主要受细胞质主效应控制,其次受胚加性×环境互作效应控制。以上说明子指、仁／壳、种仁率、油分含量和仁指的变异主要来自基因型本身的作用,但环境因素不可忽视。蛋白质含量的基因表达主要受种子胚加性×环境互作效应和种子胚显性×环境互作效应控制,其次受母体加性和母体显性主效应控制；棉酚含量的基因表达主要受胚显性x环境互作效应和母体加性x环境互作效应控制,其次受母体加性主效应控制。以上结果说明这两个性状的变异易遭受很大的环境影响,因此,在性状改良时,必需考虑环境因素。
(2)遗传率分析表明,子指、仁／壳、仁指和蛋白质含量总狭义遗传率较低,这些性状在中高代进行选择可以取得更好的选择效果；棉酚、油分含量总狭义遗传率较高,在早世代进行性状选择能取得一定效果。子指、种仁率、仁／壳、仁指、棉酚含量、油分含量以普通狭义遗传率为主,其中,子指、仁指、棉酚含量以普通母体遗传率为主；种仁率、仁／壳和油分含量以普通细胞质遗传率为主；蛋白质含量以互作细胞质遗传率为主。在育种改良中,子指、仁指、棉酚含量、仁／壳和油分含量可以根据母体植株的总体表现和单粒选择相结合的方法在低世代进行选择。由于棉籽蛋白质含量具有互作遗传率高的特点,其在育种改良中虽也可根据母体植株的总体表现在低世代进行单株选择,但一般与特定环境有密切关系。
(3)遗传相关性分析表明,种仁率与棉酚、仁／壳以胚加性负相关为主；子指与仁／壳、棉酚含量、种仁率、仁指、蛋白质含量,种仁率与蛋白质含量,仁／壳与蛋白质含量,油分与蛋白质含量,仁指与蛋白质、棉酚含量以母体加性负相关为主；仁／壳与仁指、棉酚含量、种仁率以胚加性、细胞质、母体加性负相关为主；棉酚与油分含量以胚加性正相关为主；棉酚与蛋白质含量以细胞质正相关为主；种仁率与仁指以胚加性和母体加性正相关为主。因此,上述成对性状可以在早世代进行单粒或单株间接选择。油分含量与子指、仁／壳、种仁率和仁指未发现明显的相关性,但仍有可能通过这些物理品质性状间接改良油分含量。
4棉籽蛋白质及氨基酸含量的QTL定位：以来自陆地棉品种间杂交组合(HS46×MARCABUCAG8US-1-88的188个重组近交系随机交配获得的“永久F2"群体为材料,采用本实验室构建的重组自交系分子连锁图谱,应用新发展的程序QTL Network-CL-2.0-Seed对“永久F2"群体在两个环境下测得的蛋白质及氨基酸含量进行多遗传体系QTL定位及其遗传分析。共检测到6个控制蛋白质含量的QTL,解释了58.5%的表型变异,其中,2个QTL主要在胚核基因组上表达,其余4个QTL同时在胚和母体核基因组上表达,3个QTL具有明显的环境互作效应。在3个胚加性效应贡献率大于5%的QTL中,2个QTL已定位在具体的染色体上,为蛋白质含量的分子育种提供重要基础。
本试验还共检测了控制16种氨基酸含量性状表现的QTL在胚和母体植株2套核基因组上的分布情况：包括亮氨酸(Leu)、苯丙氨酸(Phe)、苏氨酸(Thr)、缬氨酸(Val)、蛋氨酸(Met)、异亮氨酸(Ile)、赖氨酸(Lys)、天冬氨酸(Asp)、丝氨酸(Ser)、谷氨酸(Glu)、甘氨酸(Gly)、丙氨酸(Ala)、酪氨酸(Tyr)、组氨酸(His)、精氨酸(Arg)、脯氨酸(Pro)含量。研究结果发现控制这些性状的QTL数目分别为2-8个,可以解释表型变异为23.37%-64.32%。所检测到的每种氨基酸QTL中,在胚核基因组上表达的QTL分别有0-2个,其中,苯丙氨酸、缬氨酸、蛋氨酸、酪氨酸和组氨酸各为2个,天冬氨酸和甘氨酸未发现仅在胚核基因组中表达的QTL；在母体植株核基因组中表达的分别有0-2个,其中,异亮氨酸为2个,缬氨酸、天冬氨酸和组氨酸各为1个,其余的12种氨基酸未发现仅在母体基因组中表达的QTL。在胚核基因组和母体植株核基因组中同时表达的分别有0-6个,其中苯丙氨酸为6个,组氨酸为0个；有显著的环境互作效应的有1-5个,其中苏氨酸、天冬氨酸、谷氨酸、甘氨酸和丙氨酸各为1个,苯丙氨酸为6个。在七种必需氨基酸中,具有良好的定位效果的为苯丙氨酸(8个QTL,共解释的表型变异为54.5%)、缬氨酸(6个QTL,共解释的表型变异为64.32%)、异亮氨酸(6个QTL,共解释的表型变异为60.95%),这些QTL在相应的性状改良中将可发挥重要的作用。
Cotton (Gossypium spp.) is one of important economic crops in the world. The fiber which is its main product is used in the textile industry while the cottonseed, which is a byproduct is widely used in making food, animal feed, fuel and other products. Cottonseed contains many kinds of nutrients, making it a potential food-source for humans as the world population continues to rapidly grow. Cottonseed improvement has therefore become one of the major objectives in cotton breeding requiring a more detailed understanding of the genetic mechanisms in cottonseed for effective manipulation. The aims of the present study are the following:(1) Studying the inheritance of upland cotton seed traits, which include determination of genetic effects, heritability and correlation based on the embryo genome, the maternal plant genome and the plasmon. It was conducted with genetic models that were developed purposely for quantitative traits in diploid plant seed and their corresponding statistical approaches using two years data from the188inbred lines and their hybrids, all of which were cultivated within a partial diallel mating design;(2) Identifying and analyzing the QTL for protein-related traits based on embryo genome and maternal plant genome. An "immortalized F2"(IF2) population including376crosses was adopted, for this study which was conducted using the newly developed QTL mapping method that includes embryo and maternal main effects as well as their genetype×environment (GE) interaction effects on quantitative traits of dicotyledonous seeds. The main results were summarized as follows:
1Calibration equations were developed with near infrared spectroscopy (NIRS) for amino acid content in cottonseeds using a total of445samples with a high variability in amino acid contents were used for developing calibration equations of amino acid content of cottonseeds. The spectral data of cotton kernel powder was processed using the first derivative mathematical treatment combined with SNV and de-trend, as well as Modified Partial Least Squares (MPLS) regression method. Chemometric models for17amino acids present in cottonseed were developed, and12of them were found to be excellent for determination of related amino acids including asparagic acid, threonine, glutamic acid, etc, with-the ratio of SD to SECV in the calibration set (RPDc) of3.735～7.132and adetermination coefficient (r2) of0.910～0.979in external validation. For these12amino acids, their values obtained by the chemical method were accurately predicted using NIRS. The RPDc of serine, methionine, tyrosine and proline ranged between2.205and2.814, and they determination coefficient (r2) were from0.800to0.830in external validation. For these4amino acids, the NIRS values were not as accurate as those from the chemical analysis, but could be used for sample screening in cotton breeding program. The equation for cystine however was not reliable as its RPDc was only1.358, which was not suitable to estimate its content in cottonseeds.
2An immortal F2population (IF2) comprising of376crosses was constructed using a set of recombinant inbred lines (RIL) derived from the hybrid of HS46XMARCABUCAG8US-1-88. In this population, most cottonseed quality traits including seed index (SI), kernel index (KI), kernel percentage (KP), kernel/hull (K/H), protein content (PC), oil content (OC), gossypol content (GC) and17amino acids contents have characters of quantitative traits with normal distribution. In addition, means of most traits in some crosses of the IF2population were beyond those of the parents (HS46and MARCABUCAG8US-1-88). This population thus is suitable resource for genetic studies and breeding.
3Based on phenotypic data of cottonseed in188recombinant inbred lines and their IF2population, genetic variances, heritability and correlation analysis of7important cottonseed quality traits were conducted.
(1) The results of variance analysis showed:SI, KI and K/H were mainly affected by embryo additive and maternal additive effects. Secondly SI and K/H were affected by cytoplasmatic interaction; KP was mainly affected by embryo additive effect, and on a lesser note by embryo dominance and maternal additive interaction effects. OC was predominantly affected by cytoplasmic effects, and secondly by embryo additive interaction effects; The phenotypic variance of the above mentioned traits was mainly atributed to their genetypes, although environmental factors were not neglected. PC was mainly controlled by embryo dominance and maternal additive interaction effects, secondly by maternal additive and dominance main effects; GC was mainly affected by embryo dominance×environment interaction effects, also by maternal additive effect. The phenotypic variance of these two traits could be more affected by environment. The environmental factor needed to be considered in the improvement of these traits.
(2) The results of heritability analysis showed:The total narrow-sense heritabilities for GC and OC were relatively high, it was thus feasible to select these traits in early generation; those for SI, K/H, KI and PC were relatively low, then these traits were not suitable for selection in early generation. Maternal general heritability was the most important for SI, KI and GC. Cytoplasmic heritability was the main parts for K/H, KP and OC; Selection advance for SI, K/H, OC, KI and GC could therefore be expected in the early generation through the combination of the performance of maternal plant and single seed selection. While for PC, it was cytoplasmic interaction heritability, and improving PC would be more efficient in the early generation by single plant selection based on the performance of maternal plant under specific environment.
(3) The analysis of genetic correlation showed:The relationship between KI and GC, KI and K/H were mainly controlled by embryo additive effect, which showed a significantly negative correlation; Significantly negative maternal additive correlation were the main part between SI and K/H, SI and GC, SI and KI, SI and/KP, SI and PC, KP and PC, K/H and PC, OC and PC, KI and PC, KI and GC. The relationship between K/H and KI, K/H and GC, K/H and KP were mainly controlled by embryo additive, cytoplasmic and maternal additive effects, which showed significantly negative correlation; Significantly positive embryo additive correlation was main part between GC and OC; Significantly positive cytoplasmic correlation was main part between GC and PC. Significantly positive embryo and maternal additive correlation were main part between KP and KI. Therefore, direct selection for above pair traits could be conduceted in the early generation by single seed or plant selected. The relationship was not significant between OC and SI, OC and K/H, OC andKP, OC and KI, and it was not reliable to get high oil content in cottonseed through physical quality trait selected
4QTL mapping of cottonseed protein and amino acid contents were conducted on the immortal F2population obtained from the random mating among the188recombinant inbred lines derived from the cross of two upland cotton cultivars (HS46×MARCABUCAG8US-1-88). A molecular linkage map of recombinant inbred lines constructed in our laboratory and a newly developed program called the QTL Network-CL-2.0-Seed was used for QTL analysis for protein and amino acid contents based on multiple genetic systems in two different environments. Six QTL were detected for protein content explaining58.5%of the phenotypic variation. Among them, two QTL were expressed in the embryo, While the remaining four QTL were expressed in the embryo and maternal genomes. Three of the six QTL had environmental interaction effects. Among three QTL with more than5%embryo additive heritability, two QTL were mapped on particular chromosome. This indicated these two QTL were major for improving protein content in cottonseed.
QTL for sixteen amino acid including phenylalanine (phe), lysine (Lys), leucine (Leu), isoleucine (ILe), methionine (Met), threnine (Thr), valine (Val), aspartic acid (Asp), alanine (Ala), proline(Pro), serine(Ser), tyrosine(Try), histidine(His), argnine (Arg), glutamic acid (Glu) and glycine (Gly) contents were identified. The number of QTL for these traits ranged from2to8, explaining23.37%～64.32%of the phenotypic variance. Among these QTL, the range between0and2QTL were expressed in embryo genome. No QTL for Asp and Gly contents was expressed in embryo genome.Two QTL for Phe, Val, Met, Tyr and His contents were expressed in the embryo genome, respectively. Among these QTL for the sixteen aimno acid contents, the range between0and2QTL were merely expressed in maternal genome. Two for ILe and one QTL for Val, Asp and His contents were merely expressed in maternal genome, respectively. No QTL for the remaining twelve amino acid contens was merely expressed in maternal genome. Among the QTL for the sixteen amino acid contents, the range between0and6QTL were simultaneouly expressed in embryo and maternal genomes. No QTL for His content was simultaneouly expressed in embryo and maternal genomes. Six QTL for Phe content were simultaneouly expressed in the embryo and maternal genomes. The number of QTL with significant environmental interaction effect varied from1to5QTL for sixteen amino acid contents. One for Thr, Asp, Glu and Gly and Six QTL for Phe content had environmental interaction effects, respectively. The results of mapping QTL for Phe, Val and Ile content were excellent. A total of eight QTL for Phe content, six QTL for Val and Ile contents explained54.5%,64.32%and60.95%of phenotypic variance, respectively. These QTL could play an important role for the improvement of their corresponding traits.
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