菜用大豆黄化新突变体的主要农艺性状和生理特性
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摘要
对菜用大豆自然黄化突变体VS-5Y及其亲本VS-5,以及上海市崇明岛菜用大豆地方品种‘乌大青’的主要农艺性状、叶绿素含量、光合参数,以及抗氧化物质等进行了比较。结果显示,与VS-5相比,VS-5Y的株高降低了1/3,单株鲜荚产量降低了1/2,荚皮和籽粒均突变为黄色,种子百粒重也显著降低。与VS-5相比,VS-5Y和‘乌大青’的叶绿素a、叶绿素b和类胡萝卜素含量在苗期显著低于VS-5,其净光合速率(Pn)、气孔导度(Gs)和蒸腾速率(Tr)在生长前期也明显降低。随着生长至成株期,VS-5Y的叶绿素含量和净光合速率与VS-5相当,但是到了开花期和结荚期,VS-5Y的叶片、茎秆和豆荚均迅速转黄,表现为整株黄化。VS-5Y在黄化期间的叶绿素a/叶绿素b明显高于VS-5,其叶绿素b的缺失程度高于叶绿素a。‘乌大青’则仅在苗期表现黄化,随着生长,其叶绿素含量和Pn等逐渐转为正常。此外,VS-5Y在黄化期间的超氧化物歧化酶(SOD)活性和丙二醛(MDA)含量相对较高,而脯氨酸和可溶性蛋白含量相对较低。本研究为探究大豆黄化机理和利用叶色突变作为标记性状用于菜用大豆遗传改良和种质创新提供了种质和理论研究基础。
The main agronomic characters, chlorophyll contents, photosynthetic parameters and antioxidant substances of vegetable soybean mutant VS-5 Y and its parent VS-5, as well as the local vegetable soybean cultivar'Wudaqing' from Chongming Island of Shanghai, were comparatively studied. The results showed that, compared with VS-5, the plant height of VS-5 Y was reduced by 1/3, and the yield of fresh pods per plant was reduced by 1/2.The pod skin and grain of VS-5 Y were changed to yellow, and its 100-seed weight was also significantly reduced.The contents of chlorophyll a, chlorophyll b and carotenoids in VS-5 Y and 'Wudaqing' were significantly lower than those in VS-5 at seedling stage. The net photosynthetic rate(Pn), stomatal conductance(Gs) and transpiration rate(Tr) of VS-5 Y and 'Wudaqing' were also significantly decreased at the early growth stage. At adult stage, the chlorophyll contents and net photosynthetic rate of VS-5 Y were similar to that of VS-5. However, the whole plant of VS-5 Y includes the leaves, po ds and beans were turned to yellow rapidly at the flowering and pod-setting stages, and during yellowish stage, the chlorophyll a/chlorophyll b ratio(Chl a/Chl b) of VS-5 Y was significantly higher than that of VS-5, and the Chl b deficiency of VS-5 Y was higher than that of Chl a. The chlorophyll contents and Pn of 'Wudaqing' became normal gradually with the growth of plants. In addition, the SOD activity and malondialdehyde(MDA) content of VS-5 Y during yellowish stage was relatively higher, but the contents of proline and soluble protein were relatively lower. This study might provide a significant yellowish germplasm and theoretical basis for soybean yellowish mechanism research and for the use of xantha mutants as marker trait for genetic improvement and germplasms innovation of vegetable soybean.
The main agronomic characters, chlorophyll contents, photosynthetic parameters and antioxidant substances of vegetable soybean mutant VS-5 Y and its parent VS-5, as well as the local vegetable soybean cultivar'Wudaqing' from Chongming Island of Shanghai, were comparatively studied. The results showed that, compared with VS-5, the plant height of VS-5 Y was reduced by 1/3, and the yield of fresh pods per plant was reduced by 1/2.The pod skin and grain of VS-5 Y were changed to yellow, and its 100-seed weight was also significantly reduced.The contents of chlorophyll a, chlorophyll b and carotenoids in VS-5 Y and 'Wudaqing' were significantly lower than those in VS-5 at seedling stage. The net photosynthetic rate(Pn), stomatal conductance(Gs) and transpiration rate(Tr) of VS-5 Y and 'Wudaqing' were also significantly decreased at the early growth stage. At adult stage, the chlorophyll contents and net photosynthetic rate of VS-5 Y were similar to that of VS-5. However, the whole plant of VS-5 Y includes the leaves, po ds and beans were turned to yellow rapidly at the flowering and pod-setting stages, and during yellowish stage, the chlorophyll a/chlorophyll b ratio(Chl a/Chl b) of VS-5 Y was significantly higher than that of VS-5, and the Chl b deficiency of VS-5 Y was higher than that of Chl a. The chlorophyll contents and Pn of 'Wudaqing' became normal gradually with the growth of plants. In addition, the SOD activity and malondialdehyde(MDA) content of VS-5 Y during yellowish stage was relatively higher, but the contents of proline and soluble protein were relatively lower. This study might provide a significant yellowish germplasm and theoretical basis for soybean yellowish mechanism research and for the use of xantha mutants as marker trait for genetic improvement and germplasms innovation of vegetable soybean.
引文
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Cao L.,2007,Characterization and genetics of a novel aurea mutant in wheat,Dissertation for Ph.D.,Northwest A&F University,Supervosor:Wang H.,pp.1-113(曹莉,2007,一个新的小麦黄化突变体研究,博士学位论文,西北农林科技大学,导师:王辉,pp.1-113)
Chen W.,2014,Performance and genetic analysis of the new yellow leaf mutants in soybean,Thesis for M.S.,Nanjing Agriculture University,Supervisor:Zhao T.J.,pp.21-32(陈薇,2014,大豆叶片黄化新突变体的表现特点与遗传研究,硕士学位论文,南京农业大学,导师:赵团结,pp.21-32)
Cheng T.S.,and Chandlee J.M.,1999,The structural,biochemical,and genetic characterization of a new radiation-induced,variegated leaf mutant of soybean(Glycine max(L.)Merr),Proc.Natl.Sci.Counc.Repub.China B.,23(1):27-37
Dai N.,Yuan G.X.,Liu X.M.,and Gao H.B.,2017,Genetic mapping of a chlorosis mutant k60 in Arabidopsis thaliana,Beijing Linye Daxue Xuebao(Journal of Beijing Forestry University),39(7):40-45(戴楠,袁光孝,刘小敏,高宏波,2017,拟南芥黄化突变体k60的基因作图定位,北京林业大学学报,39(7):40-45)
Guo J.J.,2009,Study on photosynthesis characteristics in chlorophyll-deficient soybean mutant during the development of leaves,Thesis for M.S.,Nanjing Agriculture University,Supervisor:Xu X.M.,pp.21-34(郭晶晶,2009,大豆叶绿素缺乏突变体发育过程中光合特性的研究,硕士学位论文,南京农业大学,导师:许晓明,pp.21-34)
Hu Y.H.,Wang M.L.,Zhang N.H.,Liao W.T.,and Zhao Y.,2004,Molecular cloning and sequence analysis of the Toc33cDNA in Brassica napus line Cr3529,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),37(8):1198-1202(胡远辉,王茂林,张年辉,廖问陶,赵云,2004,幼叶黄化油菜突变体Cr3529中Toc33 cDNA的克隆和序列分析,中国农业科学,37(8):1198-1202)
Jarvis P.,Chen L.J.,Li H.,Peto C.A.,Fankhauser C.,and Chory J.,1998,An Arabidopsis mutant defective in the plastid general protein import apparatus,Science,282(5386):100-103
Jiang S.H.,Zhou H.,Lin D.Z.,Dong Y.J.,Ye S.H.,and Zhang X.M.,2013,Identification and gene mapping of a thermo-sensitive leaf-color mutant at seedling stage in rice,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),27(4):359-364(江少华,周华,林冬枝,董彦君,叶胜海,张小明,2013,水稻幼苗叶色温敏感突变体的鉴定及其基因定位,中国水稻科学,27(4):359-364)
Konovsky J.,Lumpkin T.A.,and McClary D.,1994,Edamame:the vegetable soybean,In:O'Rourke A.D.(ed.),Understanding the Japanese Food and Agrimarket:A Multifaceted Opportunity,Haworth Press,Binghamton,USA,pp.173-181
Kusumi K.,Komori H.,Satoh H.,and Iba K.,2000,Characterization of a zebra mutant of rice with increased susceptibility to light stress,Plant Cell Physiol.,41(2):158-164
Li H.S.,Sun Q.,Zhao S.J.,and Zhang W.H.,eds.,2000,Principles and techniques of plant physiology and biochemistry experiments,Higher Education Press,Beijing,China,pp.258-261(李合生,孙群,赵世杰,章文华,编著,2000,植物生理生化实验原理和技术,高等教育出版社,中国,北京,pp.258-261)
Li W.,Yu C.Y.,and Hu S.W.,2007,Primary investigation on a chlorsis mutant in Brassica juncea L.,Xibei Nonglin Keji Daxue Xuebao(Journal of Northwest A&F University(Natural Science Edition)),35(9):79-82(李玮,于澄宇,胡胜武,2007,芥菜型油菜叶片黄化突变体的初步研究,西北农林科技大学学报(自然科学版),35(9):79-82)
Liu H.Y.,Zhou F.,Li J.,Yang M.M.,Zhou T.,Hao G.C.,and Zhao Y.Z.,2017,Anatomical structure and photosynthetic characteristics of a yellow leaf mutant YL1 in sesame(Sesamum indicum L.),Zuowu Xuebao(Acta Agronomica Sinica),43(12):1856-1863(刘红艳,周芳,李俊,杨敏敏,周婷,郝国存,赵应忠,2017,芝麻黄化突变体YL1的叶片解剖学及光合特性,作物学报,43(12):1856-1863)
Ma Z.H.,Hou X.L.,Song C.,and Shen X.X.,2005,Breeding of CMS lines with yellow bud mutant marker in chili pepper,Lajiao Zazhi(Journal of Pepper),(4):19-23(马志虎,侯喜林,宋春,沈晓昆,2005,辣椒黄绿苗胞质雄性不育系的选育,辣椒杂志,(4):19-23)
Ma Z.H.,Shen X.K.,and Song C.,2003,Utilization of leaf color mark of yellow-green in hot pepper purity identification and male sterility breeding,Changjiang Shucai(Journal of Changjiang Vegetables),(1):36-38(马志虎,沈晓昆,宋春,2003,黄绿苗叶色标记技术在辣椒纯度鉴定及雄性不育中的应用,长江蔬菜,(1):36-38)
Qiu L.J.,Chang R.Z.,and Liu Z.X.,eds.,2006,Specification and data standard for description of soybean germplasm resources,China Agriculture Press,Beijing,China,pp.49-61(邱丽娟,常汝镇,刘章雄,编著,2006,大豆种质资源描述规范和数据标准,中国农业出版社,中国,北京,pp.49-61)
Somerville C.R.,1986,Analysis of photosynthesis with mutants of higher plants and algae,Annu.Rev.Plant Biol.,37(37):467-506
Stern D.B.,Hanson M.R.,and Barkan A.,2004,Genetics and genomics of chloroplast biogenesis:maize as a model system,Trends in Plant Science,9(6):293-301
Viswanathan C.,Jagendorf A.,and Zhu J.K.,2005,Understanding and improving salt tolerance in plants,Crop Sci.,45(2):437-448
Wang C.L.,1990,Analysis of artificially induced chlorophyll mutants in plants,Henongxue Tongbao(Journal of Nuclear A-gricultural Science),11(6):261-265(王彩莲,1990,人工诱发植物叶绿素突变分析,核农学通报,11(6):261-265)
Xiao H.G.,Yang H.W.,Rao Y.,Yang B.,Zhu Y.,and Zhang W.L.,2013,Analysis of chloroplast ultrastructure,stomatal characteristic parameters and photosynthetic characteristics of chlorophyll-reduced mutant in Brassica napus L.,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),46(4):715-727(肖华贵,杨焕文,饶勇,杨斌,朱英,张文龙,2013,甘蓝型油菜黄化突变体的叶绿体超微结构、气孔特征参数及光合特性,中国农业科学,46(4):715-727)
Yang C.,2014,Morphology,physiology,heredity and molecular markers of yellow leaf mutant YL-1 in cabbage(Brassica oleracea L.car.capitata),Thesis for M.S.,Chinese Academy of Agricultural Science,Supervisor:Fang Z.Y.,pp.32-39(杨冲,2014,甘蓝叶色黄化突变体YL-1的形态、生理、遗传和分子标记的研究,硕士学位论文,中国农业科学院,导师:方智远,pp.32-39)
Yang X.M.,Wu X.L.,Liu Y.F.,Li T.L.,and Qi M.F.,2018,Analysis of chlorophyll and photosynthesis of tomato chlorophyll-deficient mutant induced by EMS,Yingyong Shengtai Xuebao(Chinese Journal of Applied Ecology),29(6):1983-1989(杨小苗,吴新亮,刘玉凤,李天来,齐明芳,2018,一个番茄EMS叶色黄化突变体的叶绿素含量及光合作用,应用生态学报,29(6):1983-1989)
Yu B.S.,and Wang P.Y.,1990,Identification and utilization of soybean chlorophyll deficient mutants,Henongxue Tongbao(Journal of Nuclear Agricultural Science),11(3):121-122(于佰双,王培英,1990,大豆叶绿素缺失突变体的鉴定和利用,核农学通报,11(3):121-122)
Zhang T.Y,Zhou C.L.,Liu X.,Sun A.L.,Cao P.H.,Thanhliem N.,Tian Y.L.,Zhai H.Q.,and Jiang L.,2017,Phenotypes and gene mapping of a thermo-sensitive yellow leaf mutant of rice,Zuowu Xuebao(Acta Agronomica Sinica),43(10):1426-1433(张天雨,周春雷,刘喜,孙爱伶,曹鹏辉,Thanhliem N.,田云录,翟虎渠,江玲,2017,一个水稻温敏黄化突变体的表型分析和基因定位,作物学报,43(10):1426-1433)
Zhang T.Z.,Pan J.J.,and Feng F.Z.,1989,Genetic identification of a male sterile line of cotton with yellow-bud marker,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),22(4):17-21(张天真,潘家驹,冯福祯,1989,一个有芽黄标记性状的棉花雄性不育系的遗传鉴定,中国农业科学,22(4):17-21)
Zhao Y.,Wang M.L.,Zhang Y.Z.,Du L.F.,and Pan T.,2000,Achlorophyll-reduced seedling mutant in oilseed rape,Brassica napus,for utilization in F1hybrid production,Plant Breeding,119(2):131-135
Zhu J.K.,2000,Genetic analysis of plant salt tolerance using Arabidopsis,Plant Physiol.,124(3):941-948
Cang J.,Yu L.F.,Wang Y.Y.,Zhang D.,Hao Z.B.,and Hu B.Z.,2007,Agronomic and biochemical characters of chlorophyll deficient mutant HS 821 of soybean,Henong Xuebao(Journal of Nuclear Agricultural Sciences),21(1):9-12(苍晶,于龙凤,王豫颖,张达,郝再彬,胡宝忠,2007,大豆叶绿素缺失突变体HS 821的农艺性状和生化特性,核农学报,21(1):9-12)
Cao L.,2007,Characterization and genetics of a novel aurea mutant in wheat,Dissertation for Ph.D.,Northwest A&F University,Supervosor:Wang H.,pp.1-113(曹莉,2007,一个新的小麦黄化突变体研究,博士学位论文,西北农林科技大学,导师:王辉,pp.1-113)
Chen W.,2014,Performance and genetic analysis of the new yellow leaf mutants in soybean,Thesis for M.S.,Nanjing Agriculture University,Supervisor:Zhao T.J.,pp.21-32(陈薇,2014,大豆叶片黄化新突变体的表现特点与遗传研究,硕士学位论文,南京农业大学,导师:赵团结,pp.21-32)
Cheng T.S.,and Chandlee J.M.,1999,The structural,biochemical,and genetic characterization of a new radiation-induced,variegated leaf mutant of soybean(Glycine max(L.)Merr),Proc.Natl.Sci.Counc.Repub.China B.,23(1):27-37
Dai N.,Yuan G.X.,Liu X.M.,and Gao H.B.,2017,Genetic mapping of a chlorosis mutant k60 in Arabidopsis thaliana,Beijing Linye Daxue Xuebao(Journal of Beijing Forestry University),39(7):40-45(戴楠,袁光孝,刘小敏,高宏波,2017,拟南芥黄化突变体k60的基因作图定位,北京林业大学学报,39(7):40-45)
Guo J.J.,2009,Study on photosynthesis characteristics in chlorophyll-deficient soybean mutant during the development of leaves,Thesis for M.S.,Nanjing Agriculture University,Supervisor:Xu X.M.,pp.21-34(郭晶晶,2009,大豆叶绿素缺乏突变体发育过程中光合特性的研究,硕士学位论文,南京农业大学,导师:许晓明,pp.21-34)
Hu Y.H.,Wang M.L.,Zhang N.H.,Liao W.T.,and Zhao Y.,2004,Molecular cloning and sequence analysis of the Toc33cDNA in Brassica napus line Cr3529,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),37(8):1198-1202(胡远辉,王茂林,张年辉,廖问陶,赵云,2004,幼叶黄化油菜突变体Cr3529中Toc33 cDNA的克隆和序列分析,中国农业科学,37(8):1198-1202)
Jarvis P.,Chen L.J.,Li H.,Peto C.A.,Fankhauser C.,and Chory J.,1998,An Arabidopsis mutant defective in the plastid general protein import apparatus,Science,282(5386):100-103
Jiang S.H.,Zhou H.,Lin D.Z.,Dong Y.J.,Ye S.H.,and Zhang X.M.,2013,Identification and gene mapping of a thermo-sensitive leaf-color mutant at seedling stage in rice,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),27(4):359-364(江少华,周华,林冬枝,董彦君,叶胜海,张小明,2013,水稻幼苗叶色温敏感突变体的鉴定及其基因定位,中国水稻科学,27(4):359-364)
Konovsky J.,Lumpkin T.A.,and McClary D.,1994,Edamame:the vegetable soybean,In:O'Rourke A.D.(ed.),Understanding the Japanese Food and Agrimarket:A Multifaceted Opportunity,Haworth Press,Binghamton,USA,pp.173-181
Kusumi K.,Komori H.,Satoh H.,and Iba K.,2000,Characterization of a zebra mutant of rice with increased susceptibility to light stress,Plant Cell Physiol.,41(2):158-164
Li H.S.,Sun Q.,Zhao S.J.,and Zhang W.H.,eds.,2000,Principles and techniques of plant physiology and biochemistry experiments,Higher Education Press,Beijing,China,pp.258-261(李合生,孙群,赵世杰,章文华,编著,2000,植物生理生化实验原理和技术,高等教育出版社,中国,北京,pp.258-261)
Li W.,Yu C.Y.,and Hu S.W.,2007,Primary investigation on a chlorsis mutant in Brassica juncea L.,Xibei Nonglin Keji Daxue Xuebao(Journal of Northwest A&F University(Natural Science Edition)),35(9):79-82(李玮,于澄宇,胡胜武,2007,芥菜型油菜叶片黄化突变体的初步研究,西北农林科技大学学报(自然科学版),35(9):79-82)
Liu H.Y.,Zhou F.,Li J.,Yang M.M.,Zhou T.,Hao G.C.,and Zhao Y.Z.,2017,Anatomical structure and photosynthetic characteristics of a yellow leaf mutant YL1 in sesame(Sesamum indicum L.),Zuowu Xuebao(Acta Agronomica Sinica),43(12):1856-1863(刘红艳,周芳,李俊,杨敏敏,周婷,郝国存,赵应忠,2017,芝麻黄化突变体YL1的叶片解剖学及光合特性,作物学报,43(12):1856-1863)
Ma Z.H.,Hou X.L.,Song C.,and Shen X.X.,2005,Breeding of CMS lines with yellow bud mutant marker in chili pepper,Lajiao Zazhi(Journal of Pepper),(4):19-23(马志虎,侯喜林,宋春,沈晓昆,2005,辣椒黄绿苗胞质雄性不育系的选育,辣椒杂志,(4):19-23)
Ma Z.H.,Shen X.K.,and Song C.,2003,Utilization of leaf color mark of yellow-green in hot pepper purity identification and male sterility breeding,Changjiang Shucai(Journal of Changjiang Vegetables),(1):36-38(马志虎,沈晓昆,宋春,2003,黄绿苗叶色标记技术在辣椒纯度鉴定及雄性不育中的应用,长江蔬菜,(1):36-38)
Qiu L.J.,Chang R.Z.,and Liu Z.X.,eds.,2006,Specification and data standard for description of soybean germplasm resources,China Agriculture Press,Beijing,China,pp.49-61(邱丽娟,常汝镇,刘章雄,编著,2006,大豆种质资源描述规范和数据标准,中国农业出版社,中国,北京,pp.49-61)
Somerville C.R.,1986,Analysis of photosynthesis with mutants of higher plants and algae,Annu.Rev.Plant Biol.,37(37):467-506
Stern D.B.,Hanson M.R.,and Barkan A.,2004,Genetics and genomics of chloroplast biogenesis:maize as a model system,Trends in Plant Science,9(6):293-301
Viswanathan C.,Jagendorf A.,and Zhu J.K.,2005,Understanding and improving salt tolerance in plants,Crop Sci.,45(2):437-448
Wang C.L.,1990,Analysis of artificially induced chlorophyll mutants in plants,Henongxue Tongbao(Journal of Nuclear A-gricultural Science),11(6):261-265(王彩莲,1990,人工诱发植物叶绿素突变分析,核农学通报,11(6):261-265)
Xiao H.G.,Yang H.W.,Rao Y.,Yang B.,Zhu Y.,and Zhang W.L.,2013,Analysis of chloroplast ultrastructure,stomatal characteristic parameters and photosynthetic characteristics of chlorophyll-reduced mutant in Brassica napus L.,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),46(4):715-727(肖华贵,杨焕文,饶勇,杨斌,朱英,张文龙,2013,甘蓝型油菜黄化突变体的叶绿体超微结构、气孔特征参数及光合特性,中国农业科学,46(4):715-727)
Yang C.,2014,Morphology,physiology,heredity and molecular markers of yellow leaf mutant YL-1 in cabbage(Brassica oleracea L.car.capitata),Thesis for M.S.,Chinese Academy of Agricultural Science,Supervisor:Fang Z.Y.,pp.32-39(杨冲,2014,甘蓝叶色黄化突变体YL-1的形态、生理、遗传和分子标记的研究,硕士学位论文,中国农业科学院,导师:方智远,pp.32-39)
Yang X.M.,Wu X.L.,Liu Y.F.,Li T.L.,and Qi M.F.,2018,Analysis of chlorophyll and photosynthesis of tomato chlorophyll-deficient mutant induced by EMS,Yingyong Shengtai Xuebao(Chinese Journal of Applied Ecology),29(6):1983-1989(杨小苗,吴新亮,刘玉凤,李天来,齐明芳,2018,一个番茄EMS叶色黄化突变体的叶绿素含量及光合作用,应用生态学报,29(6):1983-1989)
Yu B.S.,and Wang P.Y.,1990,Identification and utilization of soybean chlorophyll deficient mutants,Henongxue Tongbao(Journal of Nuclear Agricultural Science),11(3):121-122(于佰双,王培英,1990,大豆叶绿素缺失突变体的鉴定和利用,核农学通报,11(3):121-122)
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