小麦黄绿突变体特性研究与遗传分析
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摘要
小麦是全世界最重要的粮食作物之一，为全球三分之一的人提供稳定的粮食来源，其产量的提高对解决未来全球粮食问题具有重要的战略意义。叶色突变是一类具有明显表型特点的突变体类型，它不仅在高等植物叶绿素的生物代谢、叶绿体分化与发育、植物生理、基因功能等研究中具有重要意义，而且在实际的生产中也具有潜在的应用价值。
本研究分别对来源于四倍体小麦Cappelli和六倍体小麦偃展4110的黄绿叶色突变体的农艺性状、叶绿体超微结构、温敏特性、遗传特性等方面进行了研究，并对控制四倍体小麦黄绿变体的基因ygld进行了定位。主要研究结果如下：
1.黄绿叶色突变体的表型分析：两个小麦叶色突变体植株与野生型相比，叶片颜色表现出明显的黄绿色，而且在全生育期稳定地表达。色突变体植株的生长发育受到阻碍，但是可以正常生长和结实。此外，突变体的株高，有效分蘖，小穗数，穗长，穗粒数，单株重量，千粒重等农艺性状都受到了较大程度的影响，而且四倍体黄绿突变体的表型还会在一定程度上受到外界环境的影响。
2.叶绿体超微结构的观察：透射电子显微镜（TEM）的分析表明，与野生型的叶绿体超微结构相比，两个突变体的叶肉细胞中叶绿体数量和大小并没有明显的变化，但四倍体小麦ygld突变体内叶片中的叶绿体形状已变为近圆形，基粒类囊体的数目变异很大，基粒类囊体与基质类囊体在基质中的排列较为散乱；六倍体小麦ygla突变体内，许多叶绿体外层膜结构破裂严重，叶绿体内部结构暴露在外，但是类囊体和基粒等内部结构相对完好。
3.叶片中色素含量分析：通过对植株叶片中色素含量的测量，发现与野生型相比，四倍体小麦ygld突变体的叶绿素a、叶绿素b、类胡萝卜素的含量都发生不同程度的下降，但叶绿素a与叶绿素b含量之间的比值升高。六倍体小麦ygla突变体的叶绿素含量和类胡萝卜素含量同样较野生型中有较大幅度的下降。
4.叶绿素荧光特性研究：四倍体小麦ygld突变体中光合系统（PS）最大光化学效率Fv/Fm在突变体与野生型间无明显差异，但突变体的初始荧光值（Fo）、最大荧光值（Fm）、光化学猝灭系数（Qp）、非光化学猝灭系数（NPQ）和电子传递速率（ETR）和野生型相比均有一定程度的减少。该结果表明ygld突变体光合系统（PS）受到了抑制，使得其光合能力减弱。六倍体小麦ygla突变体的各数值较野生型都有所下降，这些荧光特性也说明其光合能力和效率的减弱。
5.四倍体小麦突变体ygld中基因的遗传及定位分析：遗传分析表明四倍体小麦ygld突变体的黄绿叶色表型是由两对隐性核基因控制的（χ~2(15:1)=0.0056, p﹥0.9），分别命名为ygld1和ygld2，其中ygld1被定位在染色体5AL上，与SSR分子标记wmc110共分离；ygld2定位在染色体5BL上，与SSR标记wmc28共分离。该研究结果为小麦黄绿叶色突变基因的克隆及其分子机制的解析奠定了基础。
Wheat is one of the most important cereal crops in the world, and it is the main food for one thirdpeople in the world. It is essential to meet the demands of increasing wheat yield to ensure globalfood safety. Chlorophyll-deficient mutation is a kind of mutations with significant phenotypes. For itsadvantages, chlorophyll-deficient mutant has been used in the research of chlorophyll metabolism,chloroplast differentiation and development, plant physiology and gene function, it also has thepotential to be used in application.
In this study, two yellow green leaf mutants were studied, one of the yellow green leaf mutantygld was derived from Triticum durum var. Cappelli treated by gamma radiation, the other yellowgreen leaf mutant ygla identified from population of Triticum aestivum var. YZ4110treated by EMS.The agronomic traits, chloroplast ultrastructure, environment-sensitivity and genetic characteristics ofthe mutants were studied. And the genes controlled yellow green leaf trait from Triticum durum var.Cappelli were mapped. The results are as follows:
1. Phenotype characteristics of two mutants: All mutant leaves displayed yellow green leaf color,which is obviously different from the leaves on wild type, and expressed in the whole growthstage. Plant height, number of spikes per plant, number of grains of spike, grain yield per plant,1000-grain weight in the two mutants were significantly decreased than those in their wild types.Moreover, the phenotype of ygld is influenced by environment stresses.
2. Chloroplast ultrastructure observation: The TEM analysis showed that there were no differencesin the size or the number of the chloroplasts between the mutants and their wild types. However,the shape and structure of the chloroplast were different between the ygld mutant and its wildtype. The wild-type plant had ellipsoidal chloroplast, while in the ygld mutant, the shape ofchloroplast changed from ellipsoidal to circular and the grana stacks in the stroma were highlyvariable in size and exhibited a disorganized appearance. In ygla mutant, compared with the wildtype, the envelop of chloroplast was ruptrued, but thylakoid and grana remain intact, nosignificant change in number were observed.
3. Pigment content analysis: Compared with the wild types, the pigment content of the mutants,including chlorophyll a, chlorophyll b, total chlorophyll, and carotene, was decreased in themutants. In contrast, the chla/chlb ratio of the mutant was increased in comparison with thenormal green leaves.
4. Photosynthetic efficiency analysis: Chl fluorescence kinetic parameters indicated that there waslittle different from maximal quantum yield of PS Ⅱ (Fv/Fm) between ygld and wild-type.However, the minimal fluorescence (Fo), maximal fluorescence (Fm), photochemical quenching(qP), non-photochemical quenching (NPQ) and the electron transport (ETR) were largelydecreased in the mutant. These results indicated that the PS in ygld mutant were severelyinhibited, and photosynthetic capacity was reduced. In ygla mutant, the same decreases in all the above index were observed. These results indicated that photosynthesis efficiency was decreased inthe mutant.
5. Genetic analysis and mapping of the mutants: The phenotype of yellow green leaf in the ygldmutant was controlled by two recessive nuclear loci. The genes were designated ygld1and ygld2.Two molecular markers co-segregated with these genes. ygld1co-segregated with the SSRmarker wmc110on chromosome5AL, and ygld2co-segregated with the SSR marker wmc28onchromosome5BL. These results will contribute to the field of gene cloning and theunderstanding of the mechanisms underlying chlorophyll metabolism and chloroplastdevelopment in wheat.
本研究分别对来源于四倍体小麦Cappelli和六倍体小麦偃展4110的黄绿叶色突变体的农艺性状、叶绿体超微结构、温敏特性、遗传特性等方面进行了研究，并对控制四倍体小麦黄绿变体的基因ygld进行了定位。主要研究结果如下：
1.黄绿叶色突变体的表型分析：两个小麦叶色突变体植株与野生型相比，叶片颜色表现出明显的黄绿色，而且在全生育期稳定地表达。色突变体植株的生长发育受到阻碍，但是可以正常生长和结实。此外，突变体的株高，有效分蘖，小穗数，穗长，穗粒数，单株重量，千粒重等农艺性状都受到了较大程度的影响，而且四倍体黄绿突变体的表型还会在一定程度上受到外界环境的影响。
2.叶绿体超微结构的观察：透射电子显微镜（TEM）的分析表明，与野生型的叶绿体超微结构相比，两个突变体的叶肉细胞中叶绿体数量和大小并没有明显的变化，但四倍体小麦ygld突变体内叶片中的叶绿体形状已变为近圆形，基粒类囊体的数目变异很大，基粒类囊体与基质类囊体在基质中的排列较为散乱；六倍体小麦ygla突变体内，许多叶绿体外层膜结构破裂严重，叶绿体内部结构暴露在外，但是类囊体和基粒等内部结构相对完好。
3.叶片中色素含量分析：通过对植株叶片中色素含量的测量，发现与野生型相比，四倍体小麦ygld突变体的叶绿素a、叶绿素b、类胡萝卜素的含量都发生不同程度的下降，但叶绿素a与叶绿素b含量之间的比值升高。六倍体小麦ygla突变体的叶绿素含量和类胡萝卜素含量同样较野生型中有较大幅度的下降。
4.叶绿素荧光特性研究：四倍体小麦ygld突变体中光合系统（PS）最大光化学效率Fv/Fm在突变体与野生型间无明显差异，但突变体的初始荧光值（Fo）、最大荧光值（Fm）、光化学猝灭系数（Qp）、非光化学猝灭系数（NPQ）和电子传递速率（ETR）和野生型相比均有一定程度的减少。该结果表明ygld突变体光合系统（PS）受到了抑制，使得其光合能力减弱。六倍体小麦ygla突变体的各数值较野生型都有所下降，这些荧光特性也说明其光合能力和效率的减弱。
5.四倍体小麦突变体ygld中基因的遗传及定位分析：遗传分析表明四倍体小麦ygld突变体的黄绿叶色表型是由两对隐性核基因控制的（χ~2(15:1)=0.0056, p﹥0.9），分别命名为ygld1和ygld2，其中ygld1被定位在染色体5AL上，与SSR分子标记wmc110共分离；ygld2定位在染色体5BL上，与SSR标记wmc28共分离。该研究结果为小麦黄绿叶色突变基因的克隆及其分子机制的解析奠定了基础。
Wheat is one of the most important cereal crops in the world, and it is the main food for one thirdpeople in the world. It is essential to meet the demands of increasing wheat yield to ensure globalfood safety. Chlorophyll-deficient mutation is a kind of mutations with significant phenotypes. For itsadvantages, chlorophyll-deficient mutant has been used in the research of chlorophyll metabolism,chloroplast differentiation and development, plant physiology and gene function, it also has thepotential to be used in application.
In this study, two yellow green leaf mutants were studied, one of the yellow green leaf mutantygld was derived from Triticum durum var. Cappelli treated by gamma radiation, the other yellowgreen leaf mutant ygla identified from population of Triticum aestivum var. YZ4110treated by EMS.The agronomic traits, chloroplast ultrastructure, environment-sensitivity and genetic characteristics ofthe mutants were studied. And the genes controlled yellow green leaf trait from Triticum durum var.Cappelli were mapped. The results are as follows:
1. Phenotype characteristics of two mutants: All mutant leaves displayed yellow green leaf color,which is obviously different from the leaves on wild type, and expressed in the whole growthstage. Plant height, number of spikes per plant, number of grains of spike, grain yield per plant,1000-grain weight in the two mutants were significantly decreased than those in their wild types.Moreover, the phenotype of ygld is influenced by environment stresses.
2. Chloroplast ultrastructure observation: The TEM analysis showed that there were no differencesin the size or the number of the chloroplasts between the mutants and their wild types. However,the shape and structure of the chloroplast were different between the ygld mutant and its wildtype. The wild-type plant had ellipsoidal chloroplast, while in the ygld mutant, the shape ofchloroplast changed from ellipsoidal to circular and the grana stacks in the stroma were highlyvariable in size and exhibited a disorganized appearance. In ygla mutant, compared with the wildtype, the envelop of chloroplast was ruptrued, but thylakoid and grana remain intact, nosignificant change in number were observed.
3. Pigment content analysis: Compared with the wild types, the pigment content of the mutants,including chlorophyll a, chlorophyll b, total chlorophyll, and carotene, was decreased in themutants. In contrast, the chla/chlb ratio of the mutant was increased in comparison with thenormal green leaves.
4. Photosynthetic efficiency analysis: Chl fluorescence kinetic parameters indicated that there waslittle different from maximal quantum yield of PS Ⅱ (Fv/Fm) between ygld and wild-type.However, the minimal fluorescence (Fo), maximal fluorescence (Fm), photochemical quenching(qP), non-photochemical quenching (NPQ) and the electron transport (ETR) were largelydecreased in the mutant. These results indicated that the PS in ygld mutant were severelyinhibited, and photosynthetic capacity was reduced. In ygla mutant, the same decreases in all the above index were observed. These results indicated that photosynthesis efficiency was decreased inthe mutant.
5. Genetic analysis and mapping of the mutants: The phenotype of yellow green leaf in the ygldmutant was controlled by two recessive nuclear loci. The genes were designated ygld1and ygld2.Two molecular markers co-segregated with these genes. ygld1co-segregated with the SSRmarker wmc110on chromosome5AL, and ygld2co-segregated with the SSR marker wmc28onchromosome5BL. These results will contribute to the field of gene cloning and theunderstanding of the mechanisms underlying chlorophyll metabolism and chloroplastdevelopment in wheat.
引文
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