水稻紫叶性状遗传和基因定位的研究
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摘要
水稻紫叶性状是一种在水稻整个生育期都能识别的明显标记,是一类特殊的稻种资源,由于受环境影响小,性状表现稳定,因而在水稻分类、遗传研究和育种实践上具有重要价值。利用易于识别的紫叶标记不育系是解决杂交水稻繁制种纯度降低和快速简便鉴定杂种F_1种子纯度的最有效途径。本研究利用来自国际水稻研究所的紫叶稻特异种质资源进行遗传分析和基因定位,弄清其遗传规律,探索紫叶基因及紫叶抑制基因的数量及其在染色体上的位置和相互关系,探讨紫叶性状导入不育系后对不育系生育后期叶片光合和衰老生理以及杂种F_1植株生长发育、籽粒灌浆和产量的影响,明确该紫叶稻在杂交水稻育种中利用的可能性,从而为紫叶标记性状的育种应用提供理论依据和实践基础。主要研究结果如下:
1.以紫叶稻IR1552与22个不同遗传背景和恢保关系的绿叶稻品种杂交,通过对杂种F_1、F_2、BCF_1、BCF_2和F_3群体植株叶色的分离进行调查和遗传分析,结果表明,紫叶稻IR1552的紫叶性状由核内一对隐性紫叶基因(pl-pl)和一对显性紫叶抑制基因(Ipl-Ipl)控制,同时也受紫叶调节基因(C_i-C_i)的作用,而紫叶调节基因数目的多少与双亲之间存在差异的基因数目有关,即与亲本的遗传背景有关。并推断,紫叶稻IR1552的紫叶色素遗传模式为:plplIplIplC_iC_i。
2.以先红A与广恢128 BCF_2为定位群体,利用SSR分子标记对紫叶基因及紫叶抑制基因进行定位的结果表明,紫叶基因位于第6染色体短臂端距SSR标记RM587和RM584分别为3.1 cM和2.2 cM的区域内,暂命名为pl_1。紫叶抑制基因位于第1染色体长臂端距离SSR标记RM9和RM488分别1.2 cM和0.8 cM的区域内,暂命名为Ipl_1。
3.通过对紫叶标记不育系及其杂种F_1生育后期叶片光合及衰老生理指标进行研究,发现水稻紫叶标记不育系先红A、知红A和桂紫-1S生育后期剑叶中叶绿素含量、光合速率、可溶性蛋白含量和SOD活性均高于对照不育系博A和GD-1S,而MDA含量和POD活性却低于对照;紫叶不育系所配杂种F_1生育后期剑叶中叶绿素含量、光合速率、可溶性蛋白含量、MDA含量、SOD活性和POD活性则与对照品种博优253相当,无明显差异。表明水稻紫叶标记不育系先红A、知红A、桂紫-1S及其杂种F_1叶片功能正常,具有良好的光合性能,后期不早衰。紫叶性状对水稻紫叶标记不育系及杂种F_1的农艺性状和产量无不良影响,是一理想的紫叶标记供体,在育种实践上具有重要的应用价值。
4.用不同类型的籼型三系不育保持系、两系不育系与紫叶稻IR1552杂交,导入这一优异紫叶标记性状,育成带紫叶标记的籼型三系不育系和两系不育系。实践证明,该紫叶性状不仅能导入籼型的三系不育系、两系不育系中,还能有效地选育出符合育种目标的紫叶标记三系不育系和两系不育系,转育效果良好。现已育成5个带紫叶标记的籼型三系不育系和两系不育系,其中籼型紫叶标记三系不育系先红A和知红A均已于2007年8月通过了广西农作物品种审定委员会组织的专家鉴定,可以在生产上应用。
5.籼型紫叶标记三系不育系先红A和知红A是广西最早育成的紫叶标记的野败型水稻三系不育系,紫叶标记特征明显,繁茂性好,柱头外露率高,易于恢复,一般配合力强,所配组合株叶形态好,优势强,应用前景广阔。
6.用育成的水稻紫叶三系不育系先红A、紫叶两系不育系桂紫-1S分别与不同类型恢复系测交配组的结果表明,所有组合杂种F_1育性均能恢复,但不同组合之间主要农艺性状和产量差异较大,部分组合主要农艺和产量性状优于对照特优63,最大增产幅度达8.61%,表现出明显的杂种优势。这表明通过渗入广亲和基因和粳稻血缘,聚合抗病性和广适性性状,扩大亲本遗传距离,强化恢复系改良,经广泛测交筛选,是可以选育出强优势杂交稻组合供生产应用的。
Purple leaf of rice is a significant mark and could be recognized in its whole life. Purple leaf rice is a special type of rice germplasm. The purple leaf character has important value in genetic study and breeding practice as well as taxonomy due to its stability and not greatly affected by environment. By use of purple leafed male sterile line is the most effective way to solve seed purity problems in hybrid rice seed production and parental line multiplication, and the fast and simple way to identify the purity of F_1 seed. This study is to conduct genetic analysis and gene locating, to understand the rule of inheritance, and to know the number of expression and control gene (s), the gene location on chromosome, and interaction of the genes of purple leaf character and its effects on the photosynthetic and senescence physiology of male sterile lines during the late stages of growth and development, and its effects on the consequence of growth, development, seed filling and yield of the hybrid F_1 plants by use of purple leafed rice germplasm introduced from the International Rice Research Institute (IRRI) to explore the possibility of using purple leafed rice in hybrid rice breeding, and provide theoretical and practical bases for purple leaf character of rice in breeding program. The major results are shown as follows:
1. Hybridization was made between purple leafed line IR1552 and 22 green leaf varieties with different genetic backgrounds and restoring-maintaining relationship, and investigation and genetic analysis were done for leaf color segregation on F_1, F_2, BCF_1 and BCF_2 populations. The results showed that the purple leaf character from IR1552 was controlled by a pair of recessive gene (pl-pl) and a pair of dominant gene (Ipl-Ipl), while also affected by the purple regulation gene (Ci-Ci). The amount of regulation gene is related to the gene number difference between the two parents, i.e. related to their genetic backgrounds. It was concluded that the inheritance model of the purple pigment was plplIplIplCiCi.
2. The purple gene and control gene were mapped in the BCF_2 population of Xianhong A and Guanghui 128 found by using SSR molecular markers. The result showed as follows: the purple gene was flanked by SSR markers RM587 and RM584 on the short arm of chromosome 6 with a genetic distance of 3.1 cM and 2.2 cM, respectively, while the purple control gene was flanked by RM9 and RM488 on the long arm of chromosome 1 with a distance of 1.2 cM and 0.8 cM, respectively. The two pairs of genes were tentatively named pl_1 and Ipl_1.
3. Studies on physiological indexes of leaf photosynthesis and aging in the late stage of growth of purple leafed male sterile line and its F_1 hybrids were done and showed that the chlorophylls contents, photosynthetic rate, soluble protein and activity of SOD of flag leaf in the late growth stage of purple leafed male sterile lines Xiahong A, Zhihong A and Guizhi-1S were higher than the controls Bo A and GD-1S, while MDA content and activity of POD were lower than the controls. The chlorophyll contents, photosynthetic rate, soluble protein content, MDA content, activity of SOD and POD in the flag leaf during the late growth stage of F_1 hybrids by use of purple leafed male sterile lines were similar to those of CK Boyou 253, not significantly different. All the results showed that the leaf function of purple leafed male sterile lines Xianhong A, Zhihong A, Guizhi-1S and their F_1 hybrids is normal with good photosynthesis, no early aging, and there is no negative effect on the agronomic traits and yield of IR1552 and derived male sterile lines and F_1 hybrids. It is a good donor of purple leaf mark for rice breeding practice and has important use value.
4. Through hybridization between purple leafed IR1552 and different indica three-line sterile maintainers, two-line sterile lines, the ideal purple leaf mark was introduced into three-line and two-line male sterile lines. The results showed that the purple leaf character not only could be introduced into indica type of three and two lines sterile lines, but also could be used to develop breeding goals' purple leafed three and two lines sterile lines with good transferring results. So far five purple leafed three and two line sterile lines have been developed, among which both Xianhong A and Zhihong A have passed the experts' evaluation organized by Guangxi Crop Variety Registration Committee (GCVRC) in August, 2007, and ready for commercial use.
5. Indica type purple leafed three-line male sterile lines Xianhong A and Zhihong A are the first wild abortion (WA) type three-line male sterile lines bred in Guangxi with significant purple leaf mark, vigorous plant, high stigma exertion ratio, easy to be restored, strong general combining ability (GCA), good plant type and strong heterosis. It is promising for use of their hybrids.
6. The results of test crosses of different R lines with three-line Xianhong A and two-line Guizhi-1S, respectively, showed that the sterility could be restored in all F_1 hybrids, but there were differences in the agronomic traits and yield among the hybrids. The agronomic traits and yield components of some hybrids are better than the CK Teyou 63, and the highest yield increased by 8.61%, showing significant heterosis. This suggested that, through adding WC gene (s) and japonica rice germplasm, combining disease resistances and wide adaptability, widening genetic distance of the parental lines, enhancing R line improvement, widely test crossing and screening, it is possible to develop hybrid rice with very strong heterosis for commercial production.
1.以紫叶稻IR1552与22个不同遗传背景和恢保关系的绿叶稻品种杂交,通过对杂种F_1、F_2、BCF_1、BCF_2和F_3群体植株叶色的分离进行调查和遗传分析,结果表明,紫叶稻IR1552的紫叶性状由核内一对隐性紫叶基因(pl-pl)和一对显性紫叶抑制基因(Ipl-Ipl)控制,同时也受紫叶调节基因(C_i-C_i)的作用,而紫叶调节基因数目的多少与双亲之间存在差异的基因数目有关,即与亲本的遗传背景有关。并推断,紫叶稻IR1552的紫叶色素遗传模式为:plplIplIplC_iC_i。
2.以先红A与广恢128 BCF_2为定位群体,利用SSR分子标记对紫叶基因及紫叶抑制基因进行定位的结果表明,紫叶基因位于第6染色体短臂端距SSR标记RM587和RM584分别为3.1 cM和2.2 cM的区域内,暂命名为pl_1。紫叶抑制基因位于第1染色体长臂端距离SSR标记RM9和RM488分别1.2 cM和0.8 cM的区域内,暂命名为Ipl_1。
3.通过对紫叶标记不育系及其杂种F_1生育后期叶片光合及衰老生理指标进行研究,发现水稻紫叶标记不育系先红A、知红A和桂紫-1S生育后期剑叶中叶绿素含量、光合速率、可溶性蛋白含量和SOD活性均高于对照不育系博A和GD-1S,而MDA含量和POD活性却低于对照;紫叶不育系所配杂种F_1生育后期剑叶中叶绿素含量、光合速率、可溶性蛋白含量、MDA含量、SOD活性和POD活性则与对照品种博优253相当,无明显差异。表明水稻紫叶标记不育系先红A、知红A、桂紫-1S及其杂种F_1叶片功能正常,具有良好的光合性能,后期不早衰。紫叶性状对水稻紫叶标记不育系及杂种F_1的农艺性状和产量无不良影响,是一理想的紫叶标记供体,在育种实践上具有重要的应用价值。
4.用不同类型的籼型三系不育保持系、两系不育系与紫叶稻IR1552杂交,导入这一优异紫叶标记性状,育成带紫叶标记的籼型三系不育系和两系不育系。实践证明,该紫叶性状不仅能导入籼型的三系不育系、两系不育系中,还能有效地选育出符合育种目标的紫叶标记三系不育系和两系不育系,转育效果良好。现已育成5个带紫叶标记的籼型三系不育系和两系不育系,其中籼型紫叶标记三系不育系先红A和知红A均已于2007年8月通过了广西农作物品种审定委员会组织的专家鉴定,可以在生产上应用。
5.籼型紫叶标记三系不育系先红A和知红A是广西最早育成的紫叶标记的野败型水稻三系不育系,紫叶标记特征明显,繁茂性好,柱头外露率高,易于恢复,一般配合力强,所配组合株叶形态好,优势强,应用前景广阔。
6.用育成的水稻紫叶三系不育系先红A、紫叶两系不育系桂紫-1S分别与不同类型恢复系测交配组的结果表明,所有组合杂种F_1育性均能恢复,但不同组合之间主要农艺性状和产量差异较大,部分组合主要农艺和产量性状优于对照特优63,最大增产幅度达8.61%,表现出明显的杂种优势。这表明通过渗入广亲和基因和粳稻血缘,聚合抗病性和广适性性状,扩大亲本遗传距离,强化恢复系改良,经广泛测交筛选,是可以选育出强优势杂交稻组合供生产应用的。
Purple leaf of rice is a significant mark and could be recognized in its whole life. Purple leaf rice is a special type of rice germplasm. The purple leaf character has important value in genetic study and breeding practice as well as taxonomy due to its stability and not greatly affected by environment. By use of purple leafed male sterile line is the most effective way to solve seed purity problems in hybrid rice seed production and parental line multiplication, and the fast and simple way to identify the purity of F_1 seed. This study is to conduct genetic analysis and gene locating, to understand the rule of inheritance, and to know the number of expression and control gene (s), the gene location on chromosome, and interaction of the genes of purple leaf character and its effects on the photosynthetic and senescence physiology of male sterile lines during the late stages of growth and development, and its effects on the consequence of growth, development, seed filling and yield of the hybrid F_1 plants by use of purple leafed rice germplasm introduced from the International Rice Research Institute (IRRI) to explore the possibility of using purple leafed rice in hybrid rice breeding, and provide theoretical and practical bases for purple leaf character of rice in breeding program. The major results are shown as follows:
1. Hybridization was made between purple leafed line IR1552 and 22 green leaf varieties with different genetic backgrounds and restoring-maintaining relationship, and investigation and genetic analysis were done for leaf color segregation on F_1, F_2, BCF_1 and BCF_2 populations. The results showed that the purple leaf character from IR1552 was controlled by a pair of recessive gene (pl-pl) and a pair of dominant gene (Ipl-Ipl), while also affected by the purple regulation gene (Ci-Ci). The amount of regulation gene is related to the gene number difference between the two parents, i.e. related to their genetic backgrounds. It was concluded that the inheritance model of the purple pigment was plplIplIplCiCi.
2. The purple gene and control gene were mapped in the BCF_2 population of Xianhong A and Guanghui 128 found by using SSR molecular markers. The result showed as follows: the purple gene was flanked by SSR markers RM587 and RM584 on the short arm of chromosome 6 with a genetic distance of 3.1 cM and 2.2 cM, respectively, while the purple control gene was flanked by RM9 and RM488 on the long arm of chromosome 1 with a distance of 1.2 cM and 0.8 cM, respectively. The two pairs of genes were tentatively named pl_1 and Ipl_1.
3. Studies on physiological indexes of leaf photosynthesis and aging in the late stage of growth of purple leafed male sterile line and its F_1 hybrids were done and showed that the chlorophylls contents, photosynthetic rate, soluble protein and activity of SOD of flag leaf in the late growth stage of purple leafed male sterile lines Xiahong A, Zhihong A and Guizhi-1S were higher than the controls Bo A and GD-1S, while MDA content and activity of POD were lower than the controls. The chlorophyll contents, photosynthetic rate, soluble protein content, MDA content, activity of SOD and POD in the flag leaf during the late growth stage of F_1 hybrids by use of purple leafed male sterile lines were similar to those of CK Boyou 253, not significantly different. All the results showed that the leaf function of purple leafed male sterile lines Xianhong A, Zhihong A, Guizhi-1S and their F_1 hybrids is normal with good photosynthesis, no early aging, and there is no negative effect on the agronomic traits and yield of IR1552 and derived male sterile lines and F_1 hybrids. It is a good donor of purple leaf mark for rice breeding practice and has important use value.
4. Through hybridization between purple leafed IR1552 and different indica three-line sterile maintainers, two-line sterile lines, the ideal purple leaf mark was introduced into three-line and two-line male sterile lines. The results showed that the purple leaf character not only could be introduced into indica type of three and two lines sterile lines, but also could be used to develop breeding goals' purple leafed three and two lines sterile lines with good transferring results. So far five purple leafed three and two line sterile lines have been developed, among which both Xianhong A and Zhihong A have passed the experts' evaluation organized by Guangxi Crop Variety Registration Committee (GCVRC) in August, 2007, and ready for commercial use.
5. Indica type purple leafed three-line male sterile lines Xianhong A and Zhihong A are the first wild abortion (WA) type three-line male sterile lines bred in Guangxi with significant purple leaf mark, vigorous plant, high stigma exertion ratio, easy to be restored, strong general combining ability (GCA), good plant type and strong heterosis. It is promising for use of their hybrids.
6. The results of test crosses of different R lines with three-line Xianhong A and two-line Guizhi-1S, respectively, showed that the sterility could be restored in all F_1 hybrids, but there were differences in the agronomic traits and yield among the hybrids. The agronomic traits and yield components of some hybrids are better than the CK Teyou 63, and the highest yield increased by 8.61%, showing significant heterosis. This suggested that, through adding WC gene (s) and japonica rice germplasm, combining disease resistances and wide adaptability, widening genetic distance of the parental lines, enhancing R line improvement, widely test crossing and screening, it is possible to develop hybrid rice with very strong heterosis for commercial production.
引文
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