棉花细胞质雄性不育的研究与利用
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摘要
棉花具有十分明显的杂种优势。杂交棉通常比常规棉增产15%左右,而且在纤维品质、抗病、抗虫、抗逆境和光合效率等性状上也有明显改良。在棉花杂种优势的利用中,最重要的环节之一是杂交种子的生产(制种)。目前,杂交棉制种常有四条途径,人工去雄授粉法制种、化学杀雄法制种、利用核雄性不育的"两系法"制种和利用细胞质雄性不育的"三系法"制种。生产实践表明,利用棉花雄性不育既可简化制种又可节省成本,特别是利用棉花细胞质雄性不育系、保持系和恢复系的"三系法"制种,可较有效克服其他制种方法的一些缺点,是最有效的途径。为此,文章在阐述棉花杂种优势利用途径的基础上,重点综述棉花细胞质雄性不育的遗传学、细胞学和生理生化的特点;深入阐述不育细胞质对杂种F_1的正/负效应,并就如何培育强恢复系的问题,以培育转GST的强恢复系为例,探讨克服不育细胞质对杂种F_1负效应的可能机制;根据棉花为常异花授粉作物和花器具有虫媒花特征的特点,详细介绍三系杂交棉制种的亲本(不育系和恢复系)选配、地点选择和环境优化等条件,以及如何综合优化这些条件提高制种产量的关键技术。利用棉花细胞质雄性不育的"三系法"制种,与其他作物比较,在杂种优势利用中具有4个突出的优点:(1)不育系为无花粉不育类型,育性不受气候等环境的影响,可保证杂种的纯度;(2)棉花开花期长达3个月,不存在制种时花期不遇的现象,制种产量有保证;(3)棉花生态适应性广,育成的组合可在各地种植,种子产业化效益明显;(4)可利用种间(海岛棉与陆地棉间)杂种优势。可以预言,基于细胞质雄性不育的三系杂交棉是大有前途的,将是棉花杂种优势利用的主要途径。最后,就本领域的发展趋势,特别是在利用现代生物技术培育新的不育系和恢复系方面进行了初步探讨。
Cotton has significant heterosis. Hybrid cotton usually can increase production in lint yield by about 15% compared with conventional self-pollinated cultivars, and also can get obvious improvement in fiber quality, disease resistance, insect resistance, adversity resistance and photosynthetic efficiency. Among some links of cotton heterosis use, the most important link is the castration in the production of hybrid seeds. At present, there are four ways for the castration, such as hand emasculation,chemical male gametocide, nuclear male sterility and cytoplasmic male sterility(CMS). The production practice showed that use of cotton male sterility could not only simplify the hybrid seed production but also save the production cost on a commercial economic scale. In particular, the way of hybrid seed production by use of cotton CMS line, maintainer line and restorer line were the most effective way since it could overcome some disadvantages in the other ways. Therefore, in this paper, the study and application of the cotton CMS system in hybrid seed production were overviewed and some of problems currently limiting application were also addressed. At first, the genetic, cytological and biochemical characteristics of the cotton CMS were reviewed. Secondly, the positive/negative effects of sterile cytoplasm in hybrid F_1 were analyzed, and how to overcome these negative effects, such as pollen temperature sensitive and F_1 not expressing complete fertility, by developing strong restorer lines with a stronger ability for F_1 fertility restoration, was discussed in detail. For an example, transgenic strong restorer line could be developed by introducing the exogenous GST gene, which was assumed to have the function of enhancing pollen vitality, into some conventional restorer lines,and so that hybrids with higher heterosis could be produced by crossing this strong restorer with sterile lines. According to the characteristics of cotton as an often cross-pollination crop, this paper recommended in detail the key techniques of hybrid cotton seed production, such as rules of parent(sterile line and restorer line) selection, location selection and environment optimization for enriching native pollinators to produce more hybrid seeds. Then, the paper pointed out that compared with other crops, cotton hybrid seed production based on CMS system has four advantages in the cotton heterosis use:(1) The purity of hybrid seeds can be guaranteed because there is no pollen in anthers of cotton CMS line and its sterility is very stable and not affected by the climate and other environments;(2) The high yield of hybrid seed can be obtained since the long flowering period(about 3 months) of cotton does not result in the flowering asynchronism between sterile line and restorer line;(3) The wide ecological adaptability of cotton and the possibility of large-scale hybrid seed production will be benefited to popularize hybrid cotton; and(4) Interspecific heterosis between upland cotton(Gossypium hirsutum L.) and sea-island cotton(G. barbadense L.) can be used. It is predicted that the hybrid cotton production based on CMS system will be the main approach to utilize heterosis of cotton. Finally, the future works in study and application of CMS in cotton heterosis, especially in development of new sterile lines and restorer lines by use of modern biotechnology, was also discussed.
Cotton has significant heterosis. Hybrid cotton usually can increase production in lint yield by about 15% compared with conventional self-pollinated cultivars, and also can get obvious improvement in fiber quality, disease resistance, insect resistance, adversity resistance and photosynthetic efficiency. Among some links of cotton heterosis use, the most important link is the castration in the production of hybrid seeds. At present, there are four ways for the castration, such as hand emasculation,chemical male gametocide, nuclear male sterility and cytoplasmic male sterility(CMS). The production practice showed that use of cotton male sterility could not only simplify the hybrid seed production but also save the production cost on a commercial economic scale. In particular, the way of hybrid seed production by use of cotton CMS line, maintainer line and restorer line were the most effective way since it could overcome some disadvantages in the other ways. Therefore, in this paper, the study and application of the cotton CMS system in hybrid seed production were overviewed and some of problems currently limiting application were also addressed. At first, the genetic, cytological and biochemical characteristics of the cotton CMS were reviewed. Secondly, the positive/negative effects of sterile cytoplasm in hybrid F_1 were analyzed, and how to overcome these negative effects, such as pollen temperature sensitive and F_1 not expressing complete fertility, by developing strong restorer lines with a stronger ability for F_1 fertility restoration, was discussed in detail. For an example, transgenic strong restorer line could be developed by introducing the exogenous GST gene, which was assumed to have the function of enhancing pollen vitality, into some conventional restorer lines,and so that hybrids with higher heterosis could be produced by crossing this strong restorer with sterile lines. According to the characteristics of cotton as an often cross-pollination crop, this paper recommended in detail the key techniques of hybrid cotton seed production, such as rules of parent(sterile line and restorer line) selection, location selection and environment optimization for enriching native pollinators to produce more hybrid seeds. Then, the paper pointed out that compared with other crops, cotton hybrid seed production based on CMS system has four advantages in the cotton heterosis use:(1) The purity of hybrid seeds can be guaranteed because there is no pollen in anthers of cotton CMS line and its sterility is very stable and not affected by the climate and other environments;(2) The high yield of hybrid seed can be obtained since the long flowering period(about 3 months) of cotton does not result in the flowering asynchronism between sterile line and restorer line;(3) The wide ecological adaptability of cotton and the possibility of large-scale hybrid seed production will be benefited to popularize hybrid cotton; and(4) Interspecific heterosis between upland cotton(Gossypium hirsutum L.) and sea-island cotton(G. barbadense L.) can be used. It is predicted that the hybrid cotton production based on CMS system will be the main approach to utilize heterosis of cotton. Finally, the future works in study and application of CMS in cotton heterosis, especially in development of new sterile lines and restorer lines by use of modern biotechnology, was also discussed.
引文
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[2]王学德,李悦有.细胞质雄性不育棉花的转基因恢复系的选育.中国农业科学,2002,35(2):137-141.WANG X D,LI Y Y.Development of transgenic restorer of cytoplasmic male sterility in upland cotton.Scientia Agricultura Sinica,2002,35(2):137-141.(in Chinese)
[3]王学德.三系杂交棉--棉花细胞质雄性不育的研究与利用.北京:科学出版社,2011.WANG X D.Hybrid Cotton Based on System of Cytoplasmic Male Sterility.Beijing:Science Press,2011.(in Chinese)
[4]王学德,张天真,潘家驹.细胞质雄性不育棉花小孢子发生的细胞学观察和线粒体DNA的RAPD分析.中国农业科学,1998,31(2):70-75.WANG X D,ZHANG T Z,PAN J J.Cytological observation of microsporogenesis and RAPD analysis of mitochondrial DNAs for cytoplasmic male sterile cotton lines.Scientia Agricultura Sinica,1998,31(2):70-75.(in Chinese)
[5]王学德,张天真,潘家驹.棉花细胞质雄性不育系育性恢复的遗传基础:I.恢复基因及其遗传效应.中国农业科学,1996,29(5):32-40.WANG X D,ZHANG T Z,PAN J J.Genetic basis of fertility restoration to cytoplasmic male sterile lines available in China:Ⅰ.Restorer genes and their effects.Scientia Agricultura Sinica,1996,29(5):32-40.(in Chinese)
[6]王学德,潘家驹.我国棉花细胞质雄性不育系育性恢复的遗传基础:Ⅱ.恢复基因与育性增强基因间的互作效应.遗传学报,1997,24(3):271-277.WANG X D,PAN J J.Genetic basis of fertility restoration to cytoplasmic male sterile lines available in China:Ⅱ.Interation effects between restorer genes and fertility enhancer gene.Chinese Journal of Genetics,1997,24(3):271-277.(in Chinese)
[7]WEAVER J B,WESVER J B Jr.Inheritance of pollen fertility restoration in cytoplasmic male-sterile upland cotton.Crop Science,1977,17:497-499.
[8]SHEETZ R H,WESVER J B.Inheritance of a fertility enhancer factor from Pima cotton when transferred into upland cotton with Gosypium harknessii Brandegree cytoplasm.Crop Science,1980,20:272-275.
[9]张小全,王学德.细胞质雄性不育陆地棉与海岛棉间杂种优势的初步研究.棉花学报,2005,17(2):79-83.ZHANG X Q,WANG X D.Preliminary study on heterosis of interspecific hybrid cotton(Gossypium hirsutum×G.barbadense)based on cytoplasmic male-sterility system.Cotton Science,2005,17(2):79-83.(in Chinese)
[10]ZHANG X Q,WANG X D,JIANG P D,ZHU W.Inheritance of fertility restoration for cytoplasmic male sterility in a new Gossypium barbadense restorer.Agricultural Sciences in China,2010,9(4):101-105.
[11]倪密,王学德,张昭伟,朱云国,张海平,邵明彦,袁淑娜,刘英新,文国吉.三系杂交棉花粉育性对高温和低温胁迫的反应.作物学报,2009,35(11):2085-2090.NI M,WANG X D,ZHANG Z W,ZHU Y G,ZHANG H P,SHAO MY,YUAN S N,LIU Y X,WEN G J.Reaction of pollen fertility on extreme temperature stress in CMS-based hybrid cotton.Acta Agronomica Sinica,2009,35(11):2085-2090.(in Chinese)
[12]王学德.细胞质雄性不育棉花线粒体蛋白质和DNA的分析.作物学报,2000,26(1):35-39.WANG X D.Analyses of mitochondrial protein and DNA from cytoplasmic male sterile cotton.Acta Agronomica Sinica,2000,26(1):35-39.(in Chinese)
[13]蒋培东.棉花细胞质雄性不育机理的研究[D].杭州:浙江大学,2007.JIANG P D.Studies on the mechanism of cotton cytoplasmic male sterility[D].Hangzhou:Zhejiang University,2007.(in Chinese)
[14]巩养仓,张雪林,吴建勇,张兴平,彭凡嘉,张志刚,贺云新,梅正鼎,周德桂,邢朝柱.哈克尼西棉细胞质雄性不育相关线粒体基因多态性分析.棉花学报,2017,29(4):327-335.GONG Y C,ZHANG X L,WU J Y,ZHANG X P,PENG F J,ZHANG Z G,HE Y X,MEI Z D,ZHOU D G,XING C Z.Polymorphism analysis of mitochondrial genes associated cytoplasmic male sterility in cotton(Gossypium harknessii Brandegee).Cotton Science,2017,29(4):327-335.(in Chinese)
[15]王学德,潘家驹.细胞质雄性不育陆地棉的细胞质效应.作物学报,1997,23(4):393-399.WANG X D,PAN J J.Cytoplasmic effects of cytoplasmic male sterile upland cotton.Acta Agronomica Sinica,1997,23(4):393-399.(in Chinese)
[16]赵存鹏,王兆晓,王凯辉,刘素恩,耿军义,郭宝生.胞质雄性不育系冀2658A细胞质对陆地棉主要性状的影响.植物学报,2017,52(5):560-567.ZHAO C P,WANG Z X,WANG K H,LIU S E,GENG J Y,GUO B S.Effect of cytoplasmic male sterility on main characters of cotton.Chinese Bulletin of Botany,2017,52(5):560-567.(in Chinese)
[17]VIRGINA P R,ROGER K S,ERIC R A,RANDY D A.Overexpression of glutathione S-transferase/glutathione peroxidase enhances the growth of transgenic tobacco seedlings during stress.Nature Biotechnology,1997,15:988-991.
[18]VIRGINA P R,SUNDUS A L,DANIEL K G,JAMES R M,RANGYD A.Stress tolerance in transgenic tobacco seedlings that overexpress glutathione S-transferase/glutathione peroxidase.Plant Cell Physiology,2000,41(11):1229-1234.
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