玉米耐盐碱鉴定技术体系构建与耐盐碱种质筛选
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摘要
全球盐碱化进程的加快加重了盐碱对人类的威胁,它给农业生产造成的损失仅次于干旱。吉林省是我国粮食主产区,其西部地区土地盐碱化严重,并已对该区的生态环境与经济发展构成严重威胁。快速开发和有效利用西部地区盐碱化土壤,不仅直接关系到西部生态环境的恢复和改善,更牵系如何更好地确保吉林省保障国家粮食安全的重大战略问题。
本研究以玉米杂交种郑单958及其双亲自交系为材料,在4个浓度水平的Na_2CO_3溶液(12.5、25、37.5、50 mmol/L)胁迫下和4个浓度水平的NaCl溶液(50、100、150、200 mmol/L)胁迫下,进行种子萌发和幼苗胁迫试验,分别测定发芽率、相对电导率、脯氨酸(Pro)含量、超氧化物歧化酶(SOD)活性、丙二醛(MDA)含量等生理生化指标,比较杂交种与双亲自交系之间的耐盐碱差异,在此基础上构建玉米耐盐碱鉴定技术体系。然后利用已构建的玉米耐盐碱鉴定技术体系对吉林省农科院主推的29份玉米杂交种进行苗期耐盐碱性分析,将部分杂交种种植在院洮南综合试验站和院公主岭院区试验地进行大田产比分析,验证已构建的玉米耐盐碱鉴定技术体系的有效性。同时,利用玉米回交群体BC_2F_2进行苗期胁迫和大田胁迫,进行耐盐碱一致性分析,对苗期胁迫后的部分回交群体幼苗进行SSR标记分析,进而筛选获得耐盐碱玉米回交导入系。研究结果将为揭示玉米耐盐碱生理机制、挖掘玉米耐盐碱种质奠定实验基础,为玉米杂交种的区域布局提供参考依据。主要研究结果如下:
(1)玉米耐盐碱筛选的适宜浓度为25 mmol/L的Na_2CO_3溶液与100 mmol/L的NaCl溶液,种子发芽率、相对电导率、SOD活性、MDA含量以及Pro含量是玉米耐盐碱筛选的适宜生理生化指标;
(2)通过对29份吉林省农科院主推玉米杂交种进行苗期耐盐碱性分析筛到极耐盐碱玉米杂交种3份;通过14份玉米杂交种公主岭(非胁迫对照)和洮南(胁迫区)两地大田减产率的比较分析表明,苗期筛选的结果简单有效,能够比较客观地反映玉米杂交种的耐盐碱性;
(3)通过对6个玉米回交群体BC_2F_2实验室苗期胁迫和洮南大田胁迫的比较分析,结果表明苗期筛选结果与大田筛选结果一致,以郑58为背景的回交群体整体耐盐碱性较好;通过对玉米回交群体BC_2F_2【(郑58×吉4112)×郑58】进行SSR标记分析,针对找到导入供体亲本片段较多且耐盐等级在1、2级的玉米12株自交授粉,获得耐盐碱回交导入系BC_2F_3。
Salt soil was widely distributed in the world and could affect normal growth of crops. As the global salinity accelerated, it increased the threat to humanity and became one of the ecological crisis to humanity. Salinity was inferior to drought in causing the loss of agricultural production. Jilin Province was one of corn main production areas in China. The western region of Jilin province is the salinization severity area. The salinization poses a grave threat to the development of ecological environment and economic in this region. So rapid development and effective use of saline soil is important and have practical significance to the economic benefits, environmental protection and sustainable development in Jilin Province.
Under 4 alkali-stress conditions (Na2CO3 solution) and 4 salt-stress conditions (NaCl solution), maize hybrid Zhengdan958 and their parents were cultivated by sand culture. The concentration of Na2CO3 is 12.5, 25, 37.5 and 50 mmol/L, respectively. The concentration of NaCl is 50, 100, 150 and 200 mmol/L, respectively. Some physiological and biochemical indicies were tested and used to evaluate salt-alkali tolerances of hybrids and their parents, which included germination percentage, relative conductivity, SOD activity, MDA content and proline content, etc. Based on the differences between salt-alkali tolerances of hybrids and their parents the screening system of salt or alkali-stress tolerance in maize seedling were established. Salt-alkali tolerance in seedling stage among twenty-nine maize hybrids used predominantly in Jilin Province was analysised and part of these hybrids were planted in Taonan Research Center and the other in Gongzhuling Research Center. Reduction percentage analysis were carried out among fourteen hybrids so as to terrify the screening system of maize salinity we building previously. Based on the consensus analysis of Salt-alkali tolerance in lab and field for maize BC_2F_2 population, the survival seedling was marked by SSR makers and elite maize germplasm was further acquired.
The main results were summarized as followed: (1) The appropriate concentration applied in germplasm screening were 25 mmol/L Na_2CO_3 and 100 mmol/L NaCl and germination percentage, relative conductivity, SOD activity, MDA content and proline content were the appropriate physiological and biochemical indices; (2) Three elite hybrids were screened ou of twenty-nine maize hybrids used predominantly in Jilin Province. The evaluation system of salt or alkali-stress tolerance in maize seedling was verified simply and effectively and could reflect the salinity of maize hybrids correctly. (3) Through agronomic investigation of six BC_2F_2 populations of maize, the results indicated that the analysis between seedling and field were consistent. Twelve BC_2F_2 lines were self-pollinated, whose recurrent parent was zheng58. Based on SSR marker, We screened the elite maize germplasm successfully.
The results would lay a foundation for revealing the physiological mechanism of maize salt-alkali tolerant and screening elite maize germplasm, and provide reference for regional distribution of maize hybrids.
本研究以玉米杂交种郑单958及其双亲自交系为材料,在4个浓度水平的Na_2CO_3溶液(12.5、25、37.5、50 mmol/L)胁迫下和4个浓度水平的NaCl溶液(50、100、150、200 mmol/L)胁迫下,进行种子萌发和幼苗胁迫试验,分别测定发芽率、相对电导率、脯氨酸(Pro)含量、超氧化物歧化酶(SOD)活性、丙二醛(MDA)含量等生理生化指标,比较杂交种与双亲自交系之间的耐盐碱差异,在此基础上构建玉米耐盐碱鉴定技术体系。然后利用已构建的玉米耐盐碱鉴定技术体系对吉林省农科院主推的29份玉米杂交种进行苗期耐盐碱性分析,将部分杂交种种植在院洮南综合试验站和院公主岭院区试验地进行大田产比分析,验证已构建的玉米耐盐碱鉴定技术体系的有效性。同时,利用玉米回交群体BC_2F_2进行苗期胁迫和大田胁迫,进行耐盐碱一致性分析,对苗期胁迫后的部分回交群体幼苗进行SSR标记分析,进而筛选获得耐盐碱玉米回交导入系。研究结果将为揭示玉米耐盐碱生理机制、挖掘玉米耐盐碱种质奠定实验基础,为玉米杂交种的区域布局提供参考依据。主要研究结果如下:
(1)玉米耐盐碱筛选的适宜浓度为25 mmol/L的Na_2CO_3溶液与100 mmol/L的NaCl溶液,种子发芽率、相对电导率、SOD活性、MDA含量以及Pro含量是玉米耐盐碱筛选的适宜生理生化指标;
(2)通过对29份吉林省农科院主推玉米杂交种进行苗期耐盐碱性分析筛到极耐盐碱玉米杂交种3份;通过14份玉米杂交种公主岭(非胁迫对照)和洮南(胁迫区)两地大田减产率的比较分析表明,苗期筛选的结果简单有效,能够比较客观地反映玉米杂交种的耐盐碱性;
(3)通过对6个玉米回交群体BC_2F_2实验室苗期胁迫和洮南大田胁迫的比较分析,结果表明苗期筛选结果与大田筛选结果一致,以郑58为背景的回交群体整体耐盐碱性较好;通过对玉米回交群体BC_2F_2【(郑58×吉4112)×郑58】进行SSR标记分析,针对找到导入供体亲本片段较多且耐盐等级在1、2级的玉米12株自交授粉,获得耐盐碱回交导入系BC_2F_3。
Salt soil was widely distributed in the world and could affect normal growth of crops. As the global salinity accelerated, it increased the threat to humanity and became one of the ecological crisis to humanity. Salinity was inferior to drought in causing the loss of agricultural production. Jilin Province was one of corn main production areas in China. The western region of Jilin province is the salinization severity area. The salinization poses a grave threat to the development of ecological environment and economic in this region. So rapid development and effective use of saline soil is important and have practical significance to the economic benefits, environmental protection and sustainable development in Jilin Province.
Under 4 alkali-stress conditions (Na2CO3 solution) and 4 salt-stress conditions (NaCl solution), maize hybrid Zhengdan958 and their parents were cultivated by sand culture. The concentration of Na2CO3 is 12.5, 25, 37.5 and 50 mmol/L, respectively. The concentration of NaCl is 50, 100, 150 and 200 mmol/L, respectively. Some physiological and biochemical indicies were tested and used to evaluate salt-alkali tolerances of hybrids and their parents, which included germination percentage, relative conductivity, SOD activity, MDA content and proline content, etc. Based on the differences between salt-alkali tolerances of hybrids and their parents the screening system of salt or alkali-stress tolerance in maize seedling were established. Salt-alkali tolerance in seedling stage among twenty-nine maize hybrids used predominantly in Jilin Province was analysised and part of these hybrids were planted in Taonan Research Center and the other in Gongzhuling Research Center. Reduction percentage analysis were carried out among fourteen hybrids so as to terrify the screening system of maize salinity we building previously. Based on the consensus analysis of Salt-alkali tolerance in lab and field for maize BC_2F_2 population, the survival seedling was marked by SSR makers and elite maize germplasm was further acquired.
The main results were summarized as followed: (1) The appropriate concentration applied in germplasm screening were 25 mmol/L Na_2CO_3 and 100 mmol/L NaCl and germination percentage, relative conductivity, SOD activity, MDA content and proline content were the appropriate physiological and biochemical indices; (2) Three elite hybrids were screened ou of twenty-nine maize hybrids used predominantly in Jilin Province. The evaluation system of salt or alkali-stress tolerance in maize seedling was verified simply and effectively and could reflect the salinity of maize hybrids correctly. (3) Through agronomic investigation of six BC_2F_2 populations of maize, the results indicated that the analysis between seedling and field were consistent. Twelve BC_2F_2 lines were self-pollinated, whose recurrent parent was zheng58. Based on SSR marker, We screened the elite maize germplasm successfully.
The results would lay a foundation for revealing the physiological mechanism of maize salt-alkali tolerant and screening elite maize germplasm, and provide reference for regional distribution of maize hybrids.
引文
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郑国琦,许兴,徐兆祯. 2002.耐盐分胁迫的生物学机理及其基因工程研究进展.生命科学研究学报. 23(1):22- 23.
Ali A.J., Xu J.L., Ismail A.M., Fu B.Y., Vijaykumar C.H.M., Gao Y.M., Domingo J., Maghirang R., Yu S.B., Gregorio G., Yanagihara S., Cohen M., Mackill D., and Li Z.K. 2006.
Hidden diversity for abiotic and biotic stress tolerances in the primary gene pool of rice revealed by a large backcross breeding program. Field Crop Res. 97(1): 66-76.
Almodares A, Hadi M R, Dosti B. 2007. Effects of Salt Stress on Germination Percentage and Seedling Growth in Sweet Sorghum Cultivars. Journal of Biological Sciences. 7(8): 1492-1495.
Bernstein L. 1991. Osmotic adjustment of plants to saline media. Amer J Bot. 48:909-918.
Bohnert H J, Ayoubi P, Borchert C. 2001. A genomics approach towards salt stress tolerance [J]. Plant Physiol Biochem. 39:295-311.
Burdon R H, O Kane D, Fadzillah N. 1996. Oxidative stress and responses in Arabidopsis thaliana and Oryza sativa subjected to chilling and salinity stress. Biochem Soc Trans. 24:469-472.
Chen Yao, Zheng Hai lei, Xiao Qiang. 2005. Effects of Salinity on Oxidative and Antioxidative System of Spartina alterni flora. Journal of Xiamen University (Natural Science). 44(4): 576-579.
Cramer G R, Epstein E,Lauchli. 1991. Effects of sodium,potassium and calcium on salt-stressed barley. Physiologia Plantarum. (81): 197-202.
Foolad M R, Jones R A. 1993. Mapping salt-tolerance genes in tomato using trait-based marker analysis. Theor Appl Genet. 87: 184-192.
Godfrey W N, John C O, Erwin B. 2004. Gas Exchange and Chlorophyll Fluorescence of Sorghum under Salt Stress. Crop Science. 44(3): 806-811.
Godfrey W N, John C O, Erwin B. 2004. Response of Growth, Water Relations, and Ion Accumulation to NaCl Salinity. Crop Science. 44(3): 797-805.
Grieve C M, Lesch S M, Maas E V, Francois L E. 1993. Leaf and spikelet primordial initiation in salt-stressed wheat. Crop Science. 33: 1286-1292.
Guo Fangqing, Bang zhangcheng. 1999. Enhanced H+transport activity of tonplast vesicles isolated from roots of salt-tolerant multanto. Chinese Science Bulltein. 4(13): 1198.
Haro R, Baneulosma, Quintero F J, et al. 1999.Genetec basis of sodium exclusion and sodium tolerance in yeast A model for plants. Physilol Plant. 89: 868- 874.
Kasuga M, Liu Q, Miura S,et al. 1999. Improving plant drought,salt and freezing tolerance by gene transfer of a single stress-inducible transcription factor. Nat Biotechnol. 17(3): 287-291.
Li Z K, Fu BY, Gao Y M, Xu J L, Ali J, Lafitte R, Jiang Y Z, Domingo R J, Vijayakumar C.H.M., Maghirang R, Zheng T Q, Zhu L H. 2005. Genome-wide introgression lines and a forward genetics strategy for functional genomic research of complex phenotypes in rice(Oryza sativa L.).Plant Molecular Biology. 59: 33-52.
Lin Z-F(林植芳), Li S-Q(李双全), Lin G-Z(林桂珠), et al. 1984. Super oxide dismutase activity and lipid peroxidetion in relation to senescence of rice leaves.Acta Botanica Sinica(植物学报). 26(6): 605-615.
Maslenkora L T. 1993. Adaptation to salinity as monitored by PSⅡoxygen evolving reactions in barley thylakoids. Plant Physiol. 142: 629-634.
Michelet B, Boutr Y M. 1995. The plasma membrane H+-ATPase. Ahighly regulated enzyme with multiple physiological functions. Plant Physiol. 108: 1-6.
Morabito D, Jolivet Y, Prat D. 1996. Differences in the physiological responses of two clones of Eucalyptus Microtheca selected for their salt tolerance.Plant Science. 114(2): 129-139.
Muller M, Santarius K A. 1978. Changes in chloroplast membrane lipids during adaptation of barley to extreme salinity. Plant Physiol. 62: 326-333.
Mumtaz S S, Maqvi S M, Shereen A, et al. 1995. Proline accumulation in wheat seedlings subjected to various stress. Acta Plant. 17: 17-20.
Neumman P M. 1995. The of the sell wall adjustment in plant resistance to water deficit [J]. Crop. 35: 1258-1266.
Overtli T J. 1968. Extrcellular salt accumulation, a possible mechanism of salt injury in plants. Agrochimica. 12: 461-469.
Petrusa L M, W inicolL. 1997. Proline status in salt tolerant and salt sensitive alfalfa cell lines and plants in response to NaCl[ J]. Plant Physiol Biochem. 35: 303-310.
Rana Munns, Mark Tester, 2008. Mechanisms of Salinity Toleranc. Plant Boil. 59: 651-681.
Richard D, Bliss K A, Platt-Aloia,Thomson W W. 1984. Changes in plasma lemma organization in cow pea radicle during imbition in water and NaCl solutions.Plant Cell and Environment. 7: 601-606.
Rygol J, Zimmermann U. 1990. Radial and axial turgor pressure measure ments in individual root cells of Msembryanthemun crystalliuum grown under various saline conditions.Plant Cell Environ. 13: 15-26.
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Shen B, Jensen R G, Bohnert H. 1997. Increased resistance to oxidative stress in transgenic plants by targeting mannitol biosynthesis to chloroplasts.Plant Physiol. 113:1177-1183.
Shinozaki K,Yamaguchi-Shinozaki K. 1997. Gene expression and signal transduction in water-stress response. Plant Physiol. 115: 327-334.
Szabolcs I. 1989.Salt-Affected soils. Florida: CRC Press Inc Boca Raton.
Walker R R, Blackmore D H, Sung Qing. 1993. Carbon Dioside Assimilation and Foliar Ion Concentrations in Leaves of Lemon(Citrus Limon L)Trees Irrigated with NaCl or Na2SO4. Aust JPlant Physiol. 20: 73-85.
ZhangJ-F, FangY-F, MakeschinF, LiX-F, QinG-H. 2002. A-groforestry and its application in amelioration of saline soils in eastern China coastal region. In the Proceedings of the Second International Conference on Sustainable Agriculture for Food,Energy and Industry(inpress).
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赵可夫,范海. 1995.盐胁迫下外源ABA对玉米幼苗耐盐性的影响.植物学报. 37(4): 295-300.
郑国琦,许兴,徐兆祯. 2002.耐盐分胁迫的生物学机理及其基因工程研究进展.生命科学研究学报. 23(1):22- 23.
Ali A.J., Xu J.L., Ismail A.M., Fu B.Y., Vijaykumar C.H.M., Gao Y.M., Domingo J., Maghirang R., Yu S.B., Gregorio G., Yanagihara S., Cohen M., Mackill D., and Li Z.K. 2006.
Hidden diversity for abiotic and biotic stress tolerances in the primary gene pool of rice revealed by a large backcross breeding program. Field Crop Res. 97(1): 66-76.
Almodares A, Hadi M R, Dosti B. 2007. Effects of Salt Stress on Germination Percentage and Seedling Growth in Sweet Sorghum Cultivars. Journal of Biological Sciences. 7(8): 1492-1495.
Bernstein L. 1991. Osmotic adjustment of plants to saline media. Amer J Bot. 48:909-918.
Bohnert H J, Ayoubi P, Borchert C. 2001. A genomics approach towards salt stress tolerance [J]. Plant Physiol Biochem. 39:295-311.
Burdon R H, O Kane D, Fadzillah N. 1996. Oxidative stress and responses in Arabidopsis thaliana and Oryza sativa subjected to chilling and salinity stress. Biochem Soc Trans. 24:469-472.
Chen Yao, Zheng Hai lei, Xiao Qiang. 2005. Effects of Salinity on Oxidative and Antioxidative System of Spartina alterni flora. Journal of Xiamen University (Natural Science). 44(4): 576-579.
Cramer G R, Epstein E,Lauchli. 1991. Effects of sodium,potassium and calcium on salt-stressed barley. Physiologia Plantarum. (81): 197-202.
Foolad M R, Jones R A. 1993. Mapping salt-tolerance genes in tomato using trait-based marker analysis. Theor Appl Genet. 87: 184-192.
Godfrey W N, John C O, Erwin B. 2004. Gas Exchange and Chlorophyll Fluorescence of Sorghum under Salt Stress. Crop Science. 44(3): 806-811.
Godfrey W N, John C O, Erwin B. 2004. Response of Growth, Water Relations, and Ion Accumulation to NaCl Salinity. Crop Science. 44(3): 797-805.
Grieve C M, Lesch S M, Maas E V, Francois L E. 1993. Leaf and spikelet primordial initiation in salt-stressed wheat. Crop Science. 33: 1286-1292.
Guo Fangqing, Bang zhangcheng. 1999. Enhanced H+transport activity of tonplast vesicles isolated from roots of salt-tolerant multanto. Chinese Science Bulltein. 4(13): 1198.
Haro R, Baneulosma, Quintero F J, et al. 1999.Genetec basis of sodium exclusion and sodium tolerance in yeast A model for plants. Physilol Plant. 89: 868- 874.
Kasuga M, Liu Q, Miura S,et al. 1999. Improving plant drought,salt and freezing tolerance by gene transfer of a single stress-inducible transcription factor. Nat Biotechnol. 17(3): 287-291.
Li Z K, Fu BY, Gao Y M, Xu J L, Ali J, Lafitte R, Jiang Y Z, Domingo R J, Vijayakumar C.H.M., Maghirang R, Zheng T Q, Zhu L H. 2005. Genome-wide introgression lines and a forward genetics strategy for functional genomic research of complex phenotypes in rice(Oryza sativa L.).Plant Molecular Biology. 59: 33-52.
Lin Z-F(林植芳), Li S-Q(李双全), Lin G-Z(林桂珠), et al. 1984. Super oxide dismutase activity and lipid peroxidetion in relation to senescence of rice leaves.Acta Botanica Sinica(植物学报). 26(6): 605-615.
Maslenkora L T. 1993. Adaptation to salinity as monitored by PSⅡoxygen evolving reactions in barley thylakoids. Plant Physiol. 142: 629-634.
Michelet B, Boutr Y M. 1995. The plasma membrane H+-ATPase. Ahighly regulated enzyme with multiple physiological functions. Plant Physiol. 108: 1-6.
Morabito D, Jolivet Y, Prat D. 1996. Differences in the physiological responses of two clones of Eucalyptus Microtheca selected for their salt tolerance.Plant Science. 114(2): 129-139.
Muller M, Santarius K A. 1978. Changes in chloroplast membrane lipids during adaptation of barley to extreme salinity. Plant Physiol. 62: 326-333.
Mumtaz S S, Maqvi S M, Shereen A, et al. 1995. Proline accumulation in wheat seedlings subjected to various stress. Acta Plant. 17: 17-20.
Neumman P M. 1995. The of the sell wall adjustment in plant resistance to water deficit [J]. Crop. 35: 1258-1266.
Overtli T J. 1968. Extrcellular salt accumulation, a possible mechanism of salt injury in plants. Agrochimica. 12: 461-469.
Petrusa L M, W inicolL. 1997. Proline status in salt tolerant and salt sensitive alfalfa cell lines and plants in response to NaCl[ J]. Plant Physiol Biochem. 35: 303-310.
Rana Munns, Mark Tester, 2008. Mechanisms of Salinity Toleranc. Plant Boil. 59: 651-681.
Richard D, Bliss K A, Platt-Aloia,Thomson W W. 1984. Changes in plasma lemma organization in cow pea radicle during imbition in water and NaCl solutions.Plant Cell and Environment. 7: 601-606.
Rygol J, Zimmermann U. 1990. Radial and axial turgor pressure measure ments in individual root cells of Msembryanthemun crystalliuum grown under various saline conditions.Plant Cell Environ. 13: 15-26.
Saghai-Maroof M A, Soliman K M, Jorgensen R A, Allard R W. 1984. Ribosomal DNA spacer-length polymorphisms in barley: mendelian inheritance, chromosomal location and population dynamics. Proc. Natl. Acad. Sci. 81: 8014-8018.
Shen B, Jensen R G, Bohnert H. 1997. Increased resistance to oxidative stress in transgenic plants by targeting mannitol biosynthesis to chloroplasts.Plant Physiol. 113:1177-1183.
Shinozaki K,Yamaguchi-Shinozaki K. 1997. Gene expression and signal transduction in water-stress response. Plant Physiol. 115: 327-334.
Szabolcs I. 1989.Salt-Affected soils. Florida: CRC Press Inc Boca Raton.
Walker R R, Blackmore D H, Sung Qing. 1993. Carbon Dioside Assimilation and Foliar Ion Concentrations in Leaves of Lemon(Citrus Limon L)Trees Irrigated with NaCl or Na2SO4. Aust JPlant Physiol. 20: 73-85.
ZhangJ-F, FangY-F, MakeschinF, LiX-F, QinG-H. 2002. A-groforestry and its application in amelioration of saline soils in eastern China coastal region. In the Proceedings of the Second International Conference on Sustainable Agriculture for Food,Energy and Industry(inpress).
Zhu G-L(朱广廉), Zhong H-W(钟诲文), Zhang A-Q(张爱琴). 1990. Experiments of Plant Physiology(植物生理学实验). Beijing: Peking University Press.