转基因作物的生物安全:基因漂移及其潜在生态风险的研究和管控
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摘要
转基因作物品种在商业化推广的20多年间发展迅猛,在保障食品供应、拓展农业功能、缓解资源约束、保护生态环境等方面做出了卓越贡献。在创造巨大经济、环境和社会效益的同时,转基因作物的生物安全问题也引起了全球的广泛关注和讨论。其中,难以准确预见的外源基因通过基因漂移逃逸至非转基因作物及其野生近缘种,进而导致潜在的生态风险就是国内外学者的研究热点。围绕基因漂移的机制及其生态风险、风险评估、控制措施等问题进行介绍和讨论,并展望转基因生物技术的发展趋势。
Genetically modified(GM) crop is developing fast in the past 20 years on a large-scale commercialization and extension, which makes a great contribution to the food supply, extension of agriculture function, alleviation of resources constraints and protection of ecological environment. Global concerns and debates have been raised by the biosafety of GM crop while great economic, environmental and social benefits are brought by their planting. Transgene can escape to non-GM crops and wild relatives via gene flow resulting in ecological risk, which is mostly studied by domestic and foreign researchers. This paper mainly compassed the introduction and discussion about the mechanism,ecological risk, risk assessment, and the control measures of gene flow, furthermore, the future utilization of GM technology was proposed.
Genetically modified(GM) crop is developing fast in the past 20 years on a large-scale commercialization and extension, which makes a great contribution to the food supply, extension of agriculture function, alleviation of resources constraints and protection of ecological environment. Global concerns and debates have been raised by the biosafety of GM crop while great economic, environmental and social benefits are brought by their planting. Transgene can escape to non-GM crops and wild relatives via gene flow resulting in ecological risk, which is mostly studied by domestic and foreign researchers. This paper mainly compassed the introduction and discussion about the mechanism,ecological risk, risk assessment, and the control measures of gene flow, furthermore, the future utilization of GM technology was proposed.
引文
[1]Lavigne C,Klein E K,Vallée P,et al. A pollen-dispersal experiment with transgenic oilseed rape. Estimation of the average pollen dispersal of an individual plant within a field. Theoretical and Applied Genetics,1998,96(9):886-896.
[2]Nap J P,Metz P L J,Escaler M,et al. The release of genetically modified crops into environment:Part I overview of current status and regulations. The Plant Journal,2003,33(1):1-18.
[3]李瑞峰,王莹,王宇,等.转基因作物及其生物安全性.东北农业大学学报,2007,38(3):405-410.
[4]O’Callaghan M,Glare T R,Burgess E P J,et al. Effects of plants genetically modified for insect resistance on nontarget organisms.Annual Review of Entomology,2005,50(4):271-292.
[5]朱彦涛,徐虹,郭蔼光,等.植物转基因技术与当代社会发展.中国农学通报,2008,24(4):509-522.
[6]Hermannsson J,Kristjansdottir T A,Stefansson T S,et al. Measuring gene flow in barley fields under Icelandic sub-arctic conditions using closed-flowering varieties. Icelandic Agricultural Sciences,2010,23(64):51-59.
[7]James C. 2015年全球生物技术/转基因作物商业化发展态势.中国生物工程杂志,2016,36(4):1-11.
[8]James C. 2017年全球生物技术/转基因作物商业化发展态势.中国生物工程杂志,2018,38(6):1-8.
[9]李静,李红芳,张换样,等.全球转基因作物的产业化发展.山西农业科学,2009,37(1):3-8.
[10]James C. 2014年全球生物技术/转基因作物商业化发展态势.中国生物工程杂志,2015,35(1):1-14.
[11]卢宝荣,张文驹,李博.转基因的逃逸及生态风险.应用生态学报,2003,14(6):989-994.
[12]卢宝荣,夏辉.转基因植物的环境生物安全:转基因逃逸及其潜在生态风险的研究和评价.生命科学,2011,23(2):186-194.
[13]韦祖生,田益农,马崇熙.作物基因漂移研究综述.现代农业科技,2011(13):13-15.
[14]Bertolla F,Simonet P. Horizontal gene transfers in the environment:natural transformation as a putative process for gene transfers between transgenic plants and microorganisms. Research in Microbiology,1999,150(6):375-384.
[15]EllstrandNC,PrentieeHC,HancockJF.Geneflowand introgression from domesticated plants into their wild relatives.Annual Review of Ecology and Systematics,1999,30(1):539-563.
[16]Ellstrand N C. Current knowledge of gene flow in plants:implications for transgene flow. Philosophical Transactions of the Royal Society B:Biological Science,2003,358(1434):1163-1170.
[17]Ochman H,Lawrenee J G,Groisman E A. Lateral gene transfer and the nature of bacterial innovation. Nature,2000,405(6784):299-304.
[18]Gogarten J P,Townsend J P. Horizontal gene transfer,genome innovation and evolution. Nature Reviews Microbiology,2005,3(9):679-687.
[19]Pasquet R S,Peltier A,Hufford M B,et al. Long-distance pollen flow assessment through evaluation of pollinator foraging range suggests transgene escape distances. Proceedings of the National Academy of Sciences of the United States of America,2008,105(36):13456-13461.
[20]MarceauA,LoubetB,AndrieuB,etal.Modelingdiurnal and seasonal patterns of maize pollen emission in relation to meteorological factors. Agricultural and Forest Meteorology,2011,151(1):11-21.
[21]Zhang K,Li Y,Lian L. Pollen-mediated transgene flow in maize grown in the Huang-huai-hai region in China. Journal of Agricultural Science,2011,149(2):205-216.
[22]贺娟,朱威龙,朱家林,等.风、蜜蜂因素对转Cry1Ac基因棉花花粉介导的基因漂移的影响.棉花学报,2013,25(5):453-458.
[23]朱家林,贺娟,牛建群,等.风向因素对转基因抗虫棉花基因漂移效率的影响.生态学报,2013,33(21):6803-6812.
[24]Yan S,Zhu J L,Zhu W L,et al. Pollen-mediated gene flow from transgenic cotton under greenhouse conditions is dependent on different pollinators. Scientific Reports,2015,5:15917.
[25]闫硕,朱家林,朱威龙,等.风速对转基因棉花基因漂移的影响.生态学杂志,2017,36(8):2217-2223.
[26]Yoshimura Y,Beckie H J,Matsuo K. Transgenic oilseed rape along transportation routes and port of Vancouver in western Canada.Environmental Biosafety Research,2006,5(2):67-75.
[27]GarnierA,PivardS,LecomteJ.Measuringandmodeling anthropogenic secondary seed dispersal along roadverges for feral oilseed rape. Basic and Applied Ecology,2008,9(5):533-541.
[28]Amsellem L,Noyer J L,Hossaert-McKey M. Evidence for a switch in the reproductive biology of Rubus alceifolius(Rosaceae)towards apomixis,between its native range and its area of introduction.American Journal of Botany,2001,88(12):2243-2251.
[29]Cureton A N,Newbury H J,Raybould A F,et al. Genetic structure and gene flow in wild beet populations:the potential influence of habitat on transgene spread and risk assessment. Journal of Applied Ecology,2006,43(6):1203-1212.
[30]Chandler S,Dunwell J M. Gene flow,risk assessment and the environmental release of transgenic plants. Critical Reviews in Plant Sciences,2008,27(1):25-49.
[31]Lu B R,Yang C. Gene flow from genetically modified rice to its wild relatives:Assessing potential ecological consequences.Biotechnology Advance,2009,27(6):1083-1091.
[32]Friesen L F,Nelson A G,Van Acker R C. Evidence of contamination of pedigreed canola(Brassica napus)seedlots in western Canada with genetically-engineered herbicide-resistance traits. Agronomy Journal,2003,95(5):1342-1347.
[33]Andow D A,Zwahlen C. Assessing environmental risks of transgenic plants. Ecology Letters,2006,9(2):196-214.
[34]Haxel G R. Rapid displacement of native species by invasive species:effects of hybridization. Critical Care Medicine,1999,39(4):879-880.
[35]Wolf D E,Takebayashi N,Rieseberg L H. Predicting the risk of extinction through hybridization. Conservation Biology,2001,15(4):1039-1053.
[36]Levin D A,Franciseo-Ortega J,Jansen R K. Hybridization and the extinction of rare plants pecies. Conservation Biology,1996,10(1):10-16.
[37]Gealy D R,Mitten D H,Rutger J N. Gene flow between red rice(Oryza sativa)and herbicide-resistant rice(O. sativa):implications for weed management. Weed Technology,2003,17(3):627-645.
[38]Xia H,Lu B R,Su J,et al. Normal expression of insect-resistant transgene in progenies of common wild rice crossed with genetically modified rice:its implication in ecological biosafety assessment.Theoretical and Applied Genetics,2009,119(4):635-644.
[39]Van Deynze A E,Sundstrom F J,Bradford K J. Pollen-mediated gene flow in California cotton depends on pollinator activity. Crop Science,2005,45(4):1565-1570.
[40]Johnson P G,Larson S R,Anderton A L,et al. Pollen-mediated gene flow from Kentucky bluegrass under cultivated field conditions. Crop Science,2006,46(5):1990-1997.
[41]Chapman M A,Burke J M. Letting the gene out of the bottle:the population genetics of genetically modified crops. New Phytologist,2006,170(3):429-443.
[42]Yan S,Zhu W L,Zhang B Y,et al. Pollen-mediated gene flow from transgenic cotton is constrained by physical isolation measures.Scientific Reports,2018,8:2862.
[43]Heuberger S,Ellers-Kirk C,Tabashnik B E,et al. Pollen-and seedmediated transgene flow in commercial cotton seed production fields.PLoS ONE,2010,5(11):e14128.
[44]Scorza R,Kriss A B,Callahan A M,et al. Spatial and temporal assessment of pollen-and seed-mediated gene flow from genetically engineered plum Prunus domestica. PLoS ONE,2013,8(10):e75291.
[45]Llewellyn,D,Tyson C,Constable G,et al. Containment of regulated genetically modified cotton in the field. Agriculture Ecosystems and Environment,2007,121(4):419-429.
[46]Daniell H,Kumar S,Dufourmantel N. Breakthrough in chloroplast genetic engineering of agronomically important crops. Trends in Biotechnology,2005,23(5):238-245.
[47]Grevich J J,Daniell H. Chloroplast genetic engineering:recent advances and future perspectives. Critical Reviews in Plant Sciences,2005,24(2):83-107.
[48]Quesada-Vargas T,Ruiz O N,Daniell H. Characterization of heterologous multigene operons in transgenic chloroplasts. Plant Physiology,2005,138(3):1746-1762.
[49]Daniell H,Datta R,Varma S,et al. Containment of herbicide resistance through genetic engineering of chloroplast genome. Nature Biotechnology,1998,16(4):345-348.
[50]Ruf S,Hermann M,Berger I J,et al. Stable genetic transformation of tomato plastids and expression of a foreign protein in fruit. Nature Biotechnology,2001,19(9):870-875.
[51]Haygood R A,Ives A R,Andow D A. Population genetics of transgene containment. Ecology Letters,2004,7(3):213-220.
[52]Wang T,Li Y,Shi Y,et al. Low frequency transmission of a plastidencoded trait in setaria italic. Theoretical and Applied Genetics,2004,108(2):315-320.
[53]Azhagiri A K,Maliga P. Exceptional paternal inheritance of plastids in Arabidopsis suggests that low-frequency leakage of plastids via pollen may be universal in plants. Plant Journal for Cell and Molecular Biology,2007,52(5):817-823.
[54]Ruf S,Karcher D,Bock R. Determining the transgene containment level provided by chloroplast transformation. Proceedings of the National Academy of Sciences of the United States of America,2007,104(17):6998-7002.
[55]Budar F,Touzet P,De Paepe R. The nucleomitochondrial conflict in cytoplasmic male sterilities revisited. Genetica,2003,117(1):3-16.
[56]Chase C D. Cytoplasmic male sterility:a window to the world of plantmitochon-drial-nuclear interaction. Trends in Genetics,2007,23(2):81-90.
[57]Weider C,Stamp P,Christov N,et al. Stability of cytoplasmic male sterility in maize under different environmental conditions. Crop Science,2009,49(1):77-84.
[58]Hvarleva T,Hristova M,Bakalova A,et al. CMS lines for evaluation of pollen flow in sunflower relevance for transgene flow mitigation.Biotechnology and Biotechnological Equipment,2009,23(3):1309-1315.
[59]Latha R,Thiyagarajan K,Senthilvel S. Genetics,fertility behaviour and molecular marker analysis of a new TGMS line,TS6,in rice.Plant Breeding,2004,123(3):235-240.
[60]Sawhney V K. Photoperiod-sensitive male sterile mutant in tomato and its potential use in hybrid seed production. Journal of Horticultural Science and Biotechnology,2004,79(1):138-141.
[61]MlynárováL,Conner A,Nap J P. Directed microspore-specific recombination of transgenic alleles to prevent pollen-mediated transmission of transgenes. Plant Biotechnology Journal,2006,4(4):445-452.
[62]GidoniD,SrivastavaV,CarmiN.Site-specificexcisional recombination strategies for elimination of undesirable transgenes from crop plants. In Vitro Cellular and Developmental BiologyPlant,2008,44(6):457-467.
[63]Metz P L J,Jacobsen E,Nap J P,et al. The impact on biosafety of the phosphinothricin-tolerance transgene in inter-specific B. rapa×B.napus hybrids and their successive backcrosses. Theoretical and Applied Genetics,1997,95(3):442-450.
[64]Turuspekov Y,Honda I,Watanabe Y,et al. An inverted and microcolinear genomic regions of rice and barley carrying the cly1 gene for cleistogamy. Breeding Science,2009,59(5):657-663.
[65]Benitez E R,Khan N A,Matsumura H,et al. Varietal differences and morphology of cleistogamy in soybean. Crop Science,2010,50(1):185-190.
[66]Diaz A,MacNair M R. The effect of plant size on the expression of cleistogamy in Mimulus nasutus. Functional Ecology,1998,12(1):92-98.
[67]Lu B R. Transgene containment by molecular means-is it possible and cost effective? Environmental Biosafety Research,2003,2(1):3-8.
[68]Yoshida G,Itoh J I,Ohmori S,et al. Sperwoman-1-cleistogamy,a hopeful allele for gene containment in GM-rice. Plant Biotechnology Journal,2007,5(6):835-846.
[69]Gressel J. Tandem construct:preventing the rise of superweeds.Trends in Biotechnology,1999,17(9):361-366.
[70]Al-Ahmad H,Galili S,Gressel J. Tandem constructs to mitigate transgene persistence:tobacco as a model. Molecular Ecology,2004,13(3):697-710.
[71]姚万军,吴晗.我国应否将主粮转基因技术产业化?——基于一般均衡框架的经济学分析.南开学报(哲学社会科学版),2014(5):85-94.
[72]孙卓婧,张安红,叶纪明.转基因作物研发现状及展望.中国农业科技导报,2018,20(7):11-18.
[73]中国农村技术开发中心.依托“七大农作物育种”专项实施,推进基因编辑技术发展和应用.中国农业科技导报,2018,20(8):155.
[74]何晓丹,陈琦琦,展进涛.欧美等国基因组编辑生物安全管理政策及对中国的启示.中国科技论坛,2018(8):183-188.
[2]Nap J P,Metz P L J,Escaler M,et al. The release of genetically modified crops into environment:Part I overview of current status and regulations. The Plant Journal,2003,33(1):1-18.
[3]李瑞峰,王莹,王宇,等.转基因作物及其生物安全性.东北农业大学学报,2007,38(3):405-410.
[4]O’Callaghan M,Glare T R,Burgess E P J,et al. Effects of plants genetically modified for insect resistance on nontarget organisms.Annual Review of Entomology,2005,50(4):271-292.
[5]朱彦涛,徐虹,郭蔼光,等.植物转基因技术与当代社会发展.中国农学通报,2008,24(4):509-522.
[6]Hermannsson J,Kristjansdottir T A,Stefansson T S,et al. Measuring gene flow in barley fields under Icelandic sub-arctic conditions using closed-flowering varieties. Icelandic Agricultural Sciences,2010,23(64):51-59.
[7]James C. 2015年全球生物技术/转基因作物商业化发展态势.中国生物工程杂志,2016,36(4):1-11.
[8]James C. 2017年全球生物技术/转基因作物商业化发展态势.中国生物工程杂志,2018,38(6):1-8.
[9]李静,李红芳,张换样,等.全球转基因作物的产业化发展.山西农业科学,2009,37(1):3-8.
[10]James C. 2014年全球生物技术/转基因作物商业化发展态势.中国生物工程杂志,2015,35(1):1-14.
[11]卢宝荣,张文驹,李博.转基因的逃逸及生态风险.应用生态学报,2003,14(6):989-994.
[12]卢宝荣,夏辉.转基因植物的环境生物安全:转基因逃逸及其潜在生态风险的研究和评价.生命科学,2011,23(2):186-194.
[13]韦祖生,田益农,马崇熙.作物基因漂移研究综述.现代农业科技,2011(13):13-15.
[14]Bertolla F,Simonet P. Horizontal gene transfers in the environment:natural transformation as a putative process for gene transfers between transgenic plants and microorganisms. Research in Microbiology,1999,150(6):375-384.
[15]EllstrandNC,PrentieeHC,HancockJF.Geneflowand introgression from domesticated plants into their wild relatives.Annual Review of Ecology and Systematics,1999,30(1):539-563.
[16]Ellstrand N C. Current knowledge of gene flow in plants:implications for transgene flow. Philosophical Transactions of the Royal Society B:Biological Science,2003,358(1434):1163-1170.
[17]Ochman H,Lawrenee J G,Groisman E A. Lateral gene transfer and the nature of bacterial innovation. Nature,2000,405(6784):299-304.
[18]Gogarten J P,Townsend J P. Horizontal gene transfer,genome innovation and evolution. Nature Reviews Microbiology,2005,3(9):679-687.
[19]Pasquet R S,Peltier A,Hufford M B,et al. Long-distance pollen flow assessment through evaluation of pollinator foraging range suggests transgene escape distances. Proceedings of the National Academy of Sciences of the United States of America,2008,105(36):13456-13461.
[20]MarceauA,LoubetB,AndrieuB,etal.Modelingdiurnal and seasonal patterns of maize pollen emission in relation to meteorological factors. Agricultural and Forest Meteorology,2011,151(1):11-21.
[21]Zhang K,Li Y,Lian L. Pollen-mediated transgene flow in maize grown in the Huang-huai-hai region in China. Journal of Agricultural Science,2011,149(2):205-216.
[22]贺娟,朱威龙,朱家林,等.风、蜜蜂因素对转Cry1Ac基因棉花花粉介导的基因漂移的影响.棉花学报,2013,25(5):453-458.
[23]朱家林,贺娟,牛建群,等.风向因素对转基因抗虫棉花基因漂移效率的影响.生态学报,2013,33(21):6803-6812.
[24]Yan S,Zhu J L,Zhu W L,et al. Pollen-mediated gene flow from transgenic cotton under greenhouse conditions is dependent on different pollinators. Scientific Reports,2015,5:15917.
[25]闫硕,朱家林,朱威龙,等.风速对转基因棉花基因漂移的影响.生态学杂志,2017,36(8):2217-2223.
[26]Yoshimura Y,Beckie H J,Matsuo K. Transgenic oilseed rape along transportation routes and port of Vancouver in western Canada.Environmental Biosafety Research,2006,5(2):67-75.
[27]GarnierA,PivardS,LecomteJ.Measuringandmodeling anthropogenic secondary seed dispersal along roadverges for feral oilseed rape. Basic and Applied Ecology,2008,9(5):533-541.
[28]Amsellem L,Noyer J L,Hossaert-McKey M. Evidence for a switch in the reproductive biology of Rubus alceifolius(Rosaceae)towards apomixis,between its native range and its area of introduction.American Journal of Botany,2001,88(12):2243-2251.
[29]Cureton A N,Newbury H J,Raybould A F,et al. Genetic structure and gene flow in wild beet populations:the potential influence of habitat on transgene spread and risk assessment. Journal of Applied Ecology,2006,43(6):1203-1212.
[30]Chandler S,Dunwell J M. Gene flow,risk assessment and the environmental release of transgenic plants. Critical Reviews in Plant Sciences,2008,27(1):25-49.
[31]Lu B R,Yang C. Gene flow from genetically modified rice to its wild relatives:Assessing potential ecological consequences.Biotechnology Advance,2009,27(6):1083-1091.
[32]Friesen L F,Nelson A G,Van Acker R C. Evidence of contamination of pedigreed canola(Brassica napus)seedlots in western Canada with genetically-engineered herbicide-resistance traits. Agronomy Journal,2003,95(5):1342-1347.
[33]Andow D A,Zwahlen C. Assessing environmental risks of transgenic plants. Ecology Letters,2006,9(2):196-214.
[34]Haxel G R. Rapid displacement of native species by invasive species:effects of hybridization. Critical Care Medicine,1999,39(4):879-880.
[35]Wolf D E,Takebayashi N,Rieseberg L H. Predicting the risk of extinction through hybridization. Conservation Biology,2001,15(4):1039-1053.
[36]Levin D A,Franciseo-Ortega J,Jansen R K. Hybridization and the extinction of rare plants pecies. Conservation Biology,1996,10(1):10-16.
[37]Gealy D R,Mitten D H,Rutger J N. Gene flow between red rice(Oryza sativa)and herbicide-resistant rice(O. sativa):implications for weed management. Weed Technology,2003,17(3):627-645.
[38]Xia H,Lu B R,Su J,et al. Normal expression of insect-resistant transgene in progenies of common wild rice crossed with genetically modified rice:its implication in ecological biosafety assessment.Theoretical and Applied Genetics,2009,119(4):635-644.
[39]Van Deynze A E,Sundstrom F J,Bradford K J. Pollen-mediated gene flow in California cotton depends on pollinator activity. Crop Science,2005,45(4):1565-1570.
[40]Johnson P G,Larson S R,Anderton A L,et al. Pollen-mediated gene flow from Kentucky bluegrass under cultivated field conditions. Crop Science,2006,46(5):1990-1997.
[41]Chapman M A,Burke J M. Letting the gene out of the bottle:the population genetics of genetically modified crops. New Phytologist,2006,170(3):429-443.
[42]Yan S,Zhu W L,Zhang B Y,et al. Pollen-mediated gene flow from transgenic cotton is constrained by physical isolation measures.Scientific Reports,2018,8:2862.
[43]Heuberger S,Ellers-Kirk C,Tabashnik B E,et al. Pollen-and seedmediated transgene flow in commercial cotton seed production fields.PLoS ONE,2010,5(11):e14128.
[44]Scorza R,Kriss A B,Callahan A M,et al. Spatial and temporal assessment of pollen-and seed-mediated gene flow from genetically engineered plum Prunus domestica. PLoS ONE,2013,8(10):e75291.
[45]Llewellyn,D,Tyson C,Constable G,et al. Containment of regulated genetically modified cotton in the field. Agriculture Ecosystems and Environment,2007,121(4):419-429.
[46]Daniell H,Kumar S,Dufourmantel N. Breakthrough in chloroplast genetic engineering of agronomically important crops. Trends in Biotechnology,2005,23(5):238-245.
[47]Grevich J J,Daniell H. Chloroplast genetic engineering:recent advances and future perspectives. Critical Reviews in Plant Sciences,2005,24(2):83-107.
[48]Quesada-Vargas T,Ruiz O N,Daniell H. Characterization of heterologous multigene operons in transgenic chloroplasts. Plant Physiology,2005,138(3):1746-1762.
[49]Daniell H,Datta R,Varma S,et al. Containment of herbicide resistance through genetic engineering of chloroplast genome. Nature Biotechnology,1998,16(4):345-348.
[50]Ruf S,Hermann M,Berger I J,et al. Stable genetic transformation of tomato plastids and expression of a foreign protein in fruit. Nature Biotechnology,2001,19(9):870-875.
[51]Haygood R A,Ives A R,Andow D A. Population genetics of transgene containment. Ecology Letters,2004,7(3):213-220.
[52]Wang T,Li Y,Shi Y,et al. Low frequency transmission of a plastidencoded trait in setaria italic. Theoretical and Applied Genetics,2004,108(2):315-320.
[53]Azhagiri A K,Maliga P. Exceptional paternal inheritance of plastids in Arabidopsis suggests that low-frequency leakage of plastids via pollen may be universal in plants. Plant Journal for Cell and Molecular Biology,2007,52(5):817-823.
[54]Ruf S,Karcher D,Bock R. Determining the transgene containment level provided by chloroplast transformation. Proceedings of the National Academy of Sciences of the United States of America,2007,104(17):6998-7002.
[55]Budar F,Touzet P,De Paepe R. The nucleomitochondrial conflict in cytoplasmic male sterilities revisited. Genetica,2003,117(1):3-16.
[56]Chase C D. Cytoplasmic male sterility:a window to the world of plantmitochon-drial-nuclear interaction. Trends in Genetics,2007,23(2):81-90.
[57]Weider C,Stamp P,Christov N,et al. Stability of cytoplasmic male sterility in maize under different environmental conditions. Crop Science,2009,49(1):77-84.
[58]Hvarleva T,Hristova M,Bakalova A,et al. CMS lines for evaluation of pollen flow in sunflower relevance for transgene flow mitigation.Biotechnology and Biotechnological Equipment,2009,23(3):1309-1315.
[59]Latha R,Thiyagarajan K,Senthilvel S. Genetics,fertility behaviour and molecular marker analysis of a new TGMS line,TS6,in rice.Plant Breeding,2004,123(3):235-240.
[60]Sawhney V K. Photoperiod-sensitive male sterile mutant in tomato and its potential use in hybrid seed production. Journal of Horticultural Science and Biotechnology,2004,79(1):138-141.
[61]MlynárováL,Conner A,Nap J P. Directed microspore-specific recombination of transgenic alleles to prevent pollen-mediated transmission of transgenes. Plant Biotechnology Journal,2006,4(4):445-452.
[62]GidoniD,SrivastavaV,CarmiN.Site-specificexcisional recombination strategies for elimination of undesirable transgenes from crop plants. In Vitro Cellular and Developmental BiologyPlant,2008,44(6):457-467.
[63]Metz P L J,Jacobsen E,Nap J P,et al. The impact on biosafety of the phosphinothricin-tolerance transgene in inter-specific B. rapa×B.napus hybrids and their successive backcrosses. Theoretical and Applied Genetics,1997,95(3):442-450.
[64]Turuspekov Y,Honda I,Watanabe Y,et al. An inverted and microcolinear genomic regions of rice and barley carrying the cly1 gene for cleistogamy. Breeding Science,2009,59(5):657-663.
[65]Benitez E R,Khan N A,Matsumura H,et al. Varietal differences and morphology of cleistogamy in soybean. Crop Science,2010,50(1):185-190.
[66]Diaz A,MacNair M R. The effect of plant size on the expression of cleistogamy in Mimulus nasutus. Functional Ecology,1998,12(1):92-98.
[67]Lu B R. Transgene containment by molecular means-is it possible and cost effective? Environmental Biosafety Research,2003,2(1):3-8.
[68]Yoshida G,Itoh J I,Ohmori S,et al. Sperwoman-1-cleistogamy,a hopeful allele for gene containment in GM-rice. Plant Biotechnology Journal,2007,5(6):835-846.
[69]Gressel J. Tandem construct:preventing the rise of superweeds.Trends in Biotechnology,1999,17(9):361-366.
[70]Al-Ahmad H,Galili S,Gressel J. Tandem constructs to mitigate transgene persistence:tobacco as a model. Molecular Ecology,2004,13(3):697-710.
[71]姚万军,吴晗.我国应否将主粮转基因技术产业化?——基于一般均衡框架的经济学分析.南开学报(哲学社会科学版),2014(5):85-94.
[72]孙卓婧,张安红,叶纪明.转基因作物研发现状及展望.中国农业科技导报,2018,20(7):11-18.
[73]中国农村技术开发中心.依托“七大农作物育种”专项实施,推进基因编辑技术发展和应用.中国农业科技导报,2018,20(8):155.
[74]何晓丹,陈琦琦,展进涛.欧美等国基因组编辑生物安全管理政策及对中国的启示.中国科技论坛,2018(8):183-188.