RNAi转基因作物安全评价研究进展
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摘要
在秀丽隐杆线虫中首次发现双链RNA(dsRNA)能特异性地导致基因沉默(RNAi)现象后,人们开始大量地研究RNAi技术,并将其应用于功能基因的研究,来提高作物的抗性和改良遗传育种等。本文详细介绍了RNAi的技术原理,并且对RNAi技术与传统转基因技术的区别进行分析,阐述了该技术具有重要的生物学意义,以及在农作物害虫防治领域的占据独特优势。基于RNAi技术存在的潜在脱靶效应,从改良植物、靶标生物和生态环境的3个方面具体分析该技术可能存在的风险,为RNAi技术的风险评估提供参考。由于RNAi技术仍存在风险,为了维护生态多样性和保障人们的人身安全,应尽快建立起符合实际需求的安全性评价方法,本文针对RNAi转基因作物的环境安全和食用安全2个方面的评估方案进行概述。RNAi技术对减少害虫数量、提高水稻产量、降低种植成本以及减少化学农药污染、促进农业可持续发展来说具有重要意义,但该技术仍存在风险,需要进一步监管和研究,建立完善的生态评价系统,让RNAi技术在农业生产上发挥作用。
With the discovery that double-stranded RNA(dsRNA) could result in gene silencing(RNA interference, RNAi) in Caenorhabditis elegans, scientists began to exploit this RNAi technology for the study of gene functions, in order to improve crop resistances and genetic breeding. In this paper, the principle of RNAi technology is introduced, and the differences between the RNAi and traditional transgenic technology are discussed. Based on the potential miss-target effect of RNAi technology, we analyzed the potential risks of RNAi technology from improving plants, target organisms and ecological environment, which provided reference for risk assessment of RNAi technology. Because there are still risks in RNAi technology, in order to maintain ecological diversity and safeguard people′s personal safety, we should establish a safety evaluation method. Therefore, we summarize the evaluation schemes of environmental safety and food safety of RNAi genetically modified crops. RNAi technology may play an important role in reducing pest densities, increasing rice yield, reducing planting costs, reducing chemical pesticide pollution. However, there are still risks in this technology. Further supervision and research are needed to establish a sound ecological evaluation system so that RNAi technology can play a role in agricultural production.
With the discovery that double-stranded RNA(dsRNA) could result in gene silencing(RNA interference, RNAi) in Caenorhabditis elegans, scientists began to exploit this RNAi technology for the study of gene functions, in order to improve crop resistances and genetic breeding. In this paper, the principle of RNAi technology is introduced, and the differences between the RNAi and traditional transgenic technology are discussed. Based on the potential miss-target effect of RNAi technology, we analyzed the potential risks of RNAi technology from improving plants, target organisms and ecological environment, which provided reference for risk assessment of RNAi technology. Because there are still risks in RNAi technology, in order to maintain ecological diversity and safeguard people′s personal safety, we should establish a safety evaluation method. Therefore, we summarize the evaluation schemes of environmental safety and food safety of RNAi genetically modified crops. RNAi technology may play an important role in reducing pest densities, increasing rice yield, reducing planting costs, reducing chemical pesticide pollution. However, there are still risks in this technology. Further supervision and research are needed to establish a sound ecological evaluation system so that RNAi technology can play a role in agricultural production.
引文
高梦烛,赵亚林,闫青地,张海丽,王凤茹,董金皋,2016.拟南芥PMRP基因在叶绿体淀粉粒积累和抗寒性中的作用.中国农业科学,49(5):832-839.
郭强,范仲学,褚栋,李晓晴,2012.RNA干扰技术在农作物害虫防治中的应用.山东农业科学,44(12):16-18.
贺炜华,曾富华,陈信波,2008.转基因植物遗传传递稳定性.湛江师范学院学报,29(6):57-61.
黄静,贾瑞宗,孔华,郭静远,王绪朋,郭安平,2017.基于RNAi抗PRSV番木瓜株系的培育及其分子特征验证.热带作物学报,38(3):513-520.
焦悦,付伟,翟勇,2018.RNAi技术在作物中的应用及安全评价研究.作物杂志 (1):9-15.
李翠萍,吴民耀,王宏元,2012.3种半数致死浓度计算方法之比较.动物医学进展,33(9):89-92.
李娇,2016.利用RNA干扰技术沉默稻纵卷叶螟几丁质合成酶A基因.硕士学位论文.贵阳:贵州大学.
李丽文,朱延明,李杰,柏锡,才华,2007.RNAi技术在植物功能基因组学中的研究进展.东北农业大学学报,38(1):119-124.
李媛媛,2017.探究农业转基因作物及其食用安全评价对策研究进展.农业与技术,37(18):247.
林珠凤,陈海燕,吉训聪,2015.害虫对转Bt基因作物的抗性及治理.热带生物学报,6(4):497-503.
刘华清,李胜清,陈浩,2010.转基因作物安全评价及检测技术.华中农业大学学报(社会科学版) (6):14-19.
陆宴辉,梁革梅,2016.Bt作物系统害虫发生演替研究进展.植物保护,42(1):7-11.
尚仁福,吴立刚,2016.RNA干扰的机制及其应用.生命科学,28(5):576-583.
周红建,黄松,王雄伟,2010.RNAi技术研究新进展.生物技术通报 (12):84-87.
ALLEN M L,RD W W,2012.Saliva of Lygus lineolaris digests double stranded ribonucleic acids.Journal of Insect Physiology,58:391-396.
ANDRADE E C,HUNTER W B,2017.RNAi feeding bioassay:development of a non-transgenic approach to control Asian citrus psyllid and other hemipterans.Entomologia Experimentalis et Applicata,162(3):389-396.
BACHMAN P M,BOLOGNESI R,MOAR W J,MUELLER G M,PARADISE M S,RAMASESHADRI P,TAN J,UFFMANJ P,WARREN J A,WIGGINS B E,2013.Characterization of the spectrum of insecticidal activity of a double-stranded RNA with targeted activity against western corn rootworm (Diabrotica virgifera virgifera LeConte).Transgenic Research,22:1207-1222.
BACHMAN P M,HUIZINGA K M,JENSEN P D,MUELLER G,TAN J,UFFMAN J P,LEVINE S L,2016.Ecological risk assessment for DvSnf7 RNA:a plant-incorporated protectant with targeted activity against western corn rootworm.Regulatory Toxicology and Pharmacology:RTP,81:77-88.
BAUM J A,BOGAERT T,CLINTON W,HECK G R,FELDMANN P,ILAGAN O,JOHNSON S,PLAETINCK G,MUNYIKWA T,PLEAU M,2007.Control of coleopteran insect pests through RNA interference.Nature Biotechnology,25:1322-1326.
BELDEN J B,LYDY M J,2006.Joint toxicity of chlorpyrifos and esfenvalerate to fathead minnows and midge larvae.Environmental Toxicology and Chemistry,25:623-629.
BRYAN W C,2005.Environmental fate and effects of Bacillus thuringiensis (Bt) proteins from transgenic crops:a review.Journal of Agricultural and Food Chemistry,53:4643-4695.
CAMARGO C,WU K,FISHILEVICH E,NARVA K E,SIEGFRIED B D,2018.Knockdown of RNA interference pathway genes in western corn rootworm,Diabrotica virgifera virgifera,identifies no fitness costs associated with Argonaute 2 or Dicer-2.Pesticide Biochemistry & Physiology,148:103-110.
CHEN X,TIAN H,ZOU L,TANG B,HU J,ZHANG W,2008.Disruption of Spodoptera exigua larval development by silencing chitin synthase gene A with RNA interference.Bulletin of Entomological Research,98:613-619.
CHOI M Y,VANDER MEER R K,COY M,SCHARF M E,2012.Phenotypic impacts of PBAN RNA interference in an ant,Solenopsis invicta,and a moth,Helicoverpa zea.Journal of Insect Physiology,58:1159-1165.
DUBELMAN S,FISCHER J,ZAPATA F,HUIZINGA K,JIANG C,UFFMAN J,LEVINE S,CARSON D,2014.Environmental fate of double-stranded RNA in agricultural soils.PLoS ONE,9:e93155.
DUTTA T K,BANAKAR P,RAO U,2014.The status of RNAi-based transgenic research in plant nematology.Frontiers in Microbiology,5:760.
FIGUEIRAMANSUR J,FERREIRAPEREIRA A,MANSUR J F,FRANCO T A,ALVARENGA E S,SORGINE M H,NEVES B C,MELO A C,LEAL W S,MASUDA H,2013.Silencing of P-glycoprotein increases mortality in temephos-treated Aedes aegypti larvae.Insect Molecular Biology,22:648-658.
FIRE A,XU S,MONTGOMERY M K,KOSTAS S A,DRIVER S E,MELLO C C,1998.Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans.Nature,391:806-811.
FISHILEVICH E,VELEZ A M,STORER N P,LI H,BOWLING A J,RANGASAMY M,WORDEN S E,NARVA K E,SIEGFRIED B D,2016.RNAi as a management tool for the western corn rootworm,Diabrotica virgifera virgifera.Pest Management Science,72:1652-1663.
FITT G P,MARES C L,LLEWELLYN D J,1994.Field evaluation and potential ecological impact of transgenic cottons (Gossypium hirsutum) in Australia.Biocontrol Science and Technology,4:535-548.
GOULD F,1998.Sustainability of transgenic insecticidal cultivars:integrating pest genetics and ecology.Annual Review of Entomology,43:701-726.
GUO Z,KANG S,ZHU X,XIA J,WU Q,WANG S,XIE W,ZHANG Y,2015.The novel ABC transporter ABCH1 is a potential target for RNAi-based insect pest control and resistance management.Scientific Reports,5:13728.
GUO S,KEMPHUES K J,1995.Par-1,a gene required for establishing polarity in C.elegans embryos,encodes a putative Ser/Thr kinase that is asymmetrically distributed.Cell,81:611-620.
HE Z B,CAO Y Q,YIN Y P,WANG Z K,CHEN B,PENG G X,XIA Y X,2010.Role of hunchback in segment patterning of Locusta migratoria manilensis revealed by parental RNAi.Development Growth & Differentiation,48:439-445.
HU J,XIA Y,2016.F1-ATP synthase α-subunit:a potential target for RNAi-mediated pest management of Locusta migratoria manilensis.Pest Management Science,72:1433-1439.
HUVENNE H,SMAGGHE G,2010.Mechanisms of dsRNA uptake in insects and potential of RNAi for pest control:a review.Journal of Insect Physiology,56:227-235.
KENNERDELL J R,CARTHEW R W,1998.Use of dsRNA-mediated genetic interference to demonstrate that frizzled and frizzled 2 act in the wingless pathway.Cell,95:1017-1026.
KNORR E,FISHILEVICH E,TENBUSCH L,FREY M L F,RANGASAMY M,BILLION A,WORDEN S E,GANDRA P,ARORA K,LO W,2018.Gene silencing in Tribolium castaneum as a tool for the targeted identification of candidate RNAi targets in crop pests.Scientific Reports,8:206.
KOUGH J L,EDELSTEIN R,2012.EPA regulatory requirements for plant-incorporated protectants//CHRIS W,MCHUGHEN A.Regulation of agricultural biotechnology:the United States and Canada:163-174.
LEE Y,KIM M,HAN J,YEOM K H,LEE S,BAEK S H,KIM V N,2004.MicroRNA genes are transcribed by RNA polymerase II.The EMBO Journal,23:4051-4060.
LEVINE S L,TAN J,MUELLER G M,BACHMAN P M,JENSEN P D,UFFMANJ P,2015.Independent action between DvSnf7 RNA and Cry3Bb1 protein in southern corn rootworm,Diabrotica undecimpunctata howardi and Colorado potato beetle,Leptinotarsa decemlineata.PLoS ONE,10:e0118622.
LI T,CHEN J,FAN X,CHEN W,ZHANG W,2017.MicroRNA and dsRNA targeting chitin synthase a reveal a great potential for pest management of the hemipteran insect Nilaparvata lugens.Pest Management Science,73:1529-1537.
LIN Y H,HUANG J H,LIU Y,BELLES X,LEE H J,2016.Oral delivery of dsRNA lipoplexes to German cockroach protects dsRNA from degradation and induces RNAi response.Pest Management Science,73:960-966.
MAO J,ZENG F,2013.Plant-mediated RNAi of a gap gene-enhanced tobacco tolerance against the Myzus persicae.Transgenic Research,23:145-152.
MEISTER G,TUSCHL T,2004.Mechanisms of gene silencing by double-stranded RNA.Nature,431:343-349.
OBBARD D J,GORDON K H J,BUCK A H,JIGGINS F M,2009.The evolution of RNAi as a defence against viruses and transposable elements.Philosophical Transactions Biological Sciences,364:99-115.
PAIM R M,ARAUJO R N,LEHANE M J,GONTIJO N F,PEREIRA M H,2013.Long-term effects and parental RNAi in the blood feeder Rhodnius prolixus (Hemiptera;Reduviidae).Insect Biochemistry & Molecular Biology,43:1015-1020.
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YANG M,ZHAO L,SHEN Q,XIE G,WANG S,TANG B,2017.Knockdown of two trehalose-6-phosphate synthases severely affects chitin metabolism gene expression in the brown planthopper Nilaparvata lugens.Pest Management Science,73:206-216.
YOGINDRAN S,RAJAM M V,2016.Artificial miRNA-mediated silencing of ecdysone receptor (EcR) affects larval development and oogenesis in Helicoverpa armigera.Insect Biochemistry and Molecular Biology,77:21-30.
ZHU F,XU J,PALLI R,FERGUSON J,PALLI S R,2011.Ingested RNA interference for managing the populations of the Colorado potato beetle,Leptinotarsa decemlineata.Pest Management Science,67:175-182.
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郭强,范仲学,褚栋,李晓晴,2012.RNA干扰技术在农作物害虫防治中的应用.山东农业科学,44(12):16-18.
贺炜华,曾富华,陈信波,2008.转基因植物遗传传递稳定性.湛江师范学院学报,29(6):57-61.
黄静,贾瑞宗,孔华,郭静远,王绪朋,郭安平,2017.基于RNAi抗PRSV番木瓜株系的培育及其分子特征验证.热带作物学报,38(3):513-520.
焦悦,付伟,翟勇,2018.RNAi技术在作物中的应用及安全评价研究.作物杂志 (1):9-15.
李翠萍,吴民耀,王宏元,2012.3种半数致死浓度计算方法之比较.动物医学进展,33(9):89-92.
李娇,2016.利用RNA干扰技术沉默稻纵卷叶螟几丁质合成酶A基因.硕士学位论文.贵阳:贵州大学.
李丽文,朱延明,李杰,柏锡,才华,2007.RNAi技术在植物功能基因组学中的研究进展.东北农业大学学报,38(1):119-124.
李媛媛,2017.探究农业转基因作物及其食用安全评价对策研究进展.农业与技术,37(18):247.
林珠凤,陈海燕,吉训聪,2015.害虫对转Bt基因作物的抗性及治理.热带生物学报,6(4):497-503.
刘华清,李胜清,陈浩,2010.转基因作物安全评价及检测技术.华中农业大学学报(社会科学版) (6):14-19.
陆宴辉,梁革梅,2016.Bt作物系统害虫发生演替研究进展.植物保护,42(1):7-11.
尚仁福,吴立刚,2016.RNA干扰的机制及其应用.生命科学,28(5):576-583.
周红建,黄松,王雄伟,2010.RNAi技术研究新进展.生物技术通报 (12):84-87.
ALLEN M L,RD W W,2012.Saliva of Lygus lineolaris digests double stranded ribonucleic acids.Journal of Insect Physiology,58:391-396.
ANDRADE E C,HUNTER W B,2017.RNAi feeding bioassay:development of a non-transgenic approach to control Asian citrus psyllid and other hemipterans.Entomologia Experimentalis et Applicata,162(3):389-396.
BACHMAN P M,BOLOGNESI R,MOAR W J,MUELLER G M,PARADISE M S,RAMASESHADRI P,TAN J,UFFMANJ P,WARREN J A,WIGGINS B E,2013.Characterization of the spectrum of insecticidal activity of a double-stranded RNA with targeted activity against western corn rootworm (Diabrotica virgifera virgifera LeConte).Transgenic Research,22:1207-1222.
BACHMAN P M,HUIZINGA K M,JENSEN P D,MUELLER G,TAN J,UFFMAN J P,LEVINE S L,2016.Ecological risk assessment for DvSnf7 RNA:a plant-incorporated protectant with targeted activity against western corn rootworm.Regulatory Toxicology and Pharmacology:RTP,81:77-88.
BAUM J A,BOGAERT T,CLINTON W,HECK G R,FELDMANN P,ILAGAN O,JOHNSON S,PLAETINCK G,MUNYIKWA T,PLEAU M,2007.Control of coleopteran insect pests through RNA interference.Nature Biotechnology,25:1322-1326.
BELDEN J B,LYDY M J,2006.Joint toxicity of chlorpyrifos and esfenvalerate to fathead minnows and midge larvae.Environmental Toxicology and Chemistry,25:623-629.
BRYAN W C,2005.Environmental fate and effects of Bacillus thuringiensis (Bt) proteins from transgenic crops:a review.Journal of Agricultural and Food Chemistry,53:4643-4695.
CAMARGO C,WU K,FISHILEVICH E,NARVA K E,SIEGFRIED B D,2018.Knockdown of RNA interference pathway genes in western corn rootworm,Diabrotica virgifera virgifera,identifies no fitness costs associated with Argonaute 2 or Dicer-2.Pesticide Biochemistry & Physiology,148:103-110.
CHEN X,TIAN H,ZOU L,TANG B,HU J,ZHANG W,2008.Disruption of Spodoptera exigua larval development by silencing chitin synthase gene A with RNA interference.Bulletin of Entomological Research,98:613-619.
CHOI M Y,VANDER MEER R K,COY M,SCHARF M E,2012.Phenotypic impacts of PBAN RNA interference in an ant,Solenopsis invicta,and a moth,Helicoverpa zea.Journal of Insect Physiology,58:1159-1165.
DUBELMAN S,FISCHER J,ZAPATA F,HUIZINGA K,JIANG C,UFFMAN J,LEVINE S,CARSON D,2014.Environmental fate of double-stranded RNA in agricultural soils.PLoS ONE,9:e93155.
DUTTA T K,BANAKAR P,RAO U,2014.The status of RNAi-based transgenic research in plant nematology.Frontiers in Microbiology,5:760.
FIGUEIRAMANSUR J,FERREIRAPEREIRA A,MANSUR J F,FRANCO T A,ALVARENGA E S,SORGINE M H,NEVES B C,MELO A C,LEAL W S,MASUDA H,2013.Silencing of P-glycoprotein increases mortality in temephos-treated Aedes aegypti larvae.Insect Molecular Biology,22:648-658.
FIRE A,XU S,MONTGOMERY M K,KOSTAS S A,DRIVER S E,MELLO C C,1998.Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans.Nature,391:806-811.
FISHILEVICH E,VELEZ A M,STORER N P,LI H,BOWLING A J,RANGASAMY M,WORDEN S E,NARVA K E,SIEGFRIED B D,2016.RNAi as a management tool for the western corn rootworm,Diabrotica virgifera virgifera.Pest Management Science,72:1652-1663.
FITT G P,MARES C L,LLEWELLYN D J,1994.Field evaluation and potential ecological impact of transgenic cottons (Gossypium hirsutum) in Australia.Biocontrol Science and Technology,4:535-548.
GOULD F,1998.Sustainability of transgenic insecticidal cultivars:integrating pest genetics and ecology.Annual Review of Entomology,43:701-726.
GUO Z,KANG S,ZHU X,XIA J,WU Q,WANG S,XIE W,ZHANG Y,2015.The novel ABC transporter ABCH1 is a potential target for RNAi-based insect pest control and resistance management.Scientific Reports,5:13728.
GUO S,KEMPHUES K J,1995.Par-1,a gene required for establishing polarity in C.elegans embryos,encodes a putative Ser/Thr kinase that is asymmetrically distributed.Cell,81:611-620.
HE Z B,CAO Y Q,YIN Y P,WANG Z K,CHEN B,PENG G X,XIA Y X,2010.Role of hunchback in segment patterning of Locusta migratoria manilensis revealed by parental RNAi.Development Growth & Differentiation,48:439-445.
HU J,XIA Y,2016.F1-ATP synthase α-subunit:a potential target for RNAi-mediated pest management of Locusta migratoria manilensis.Pest Management Science,72:1433-1439.
HUVENNE H,SMAGGHE G,2010.Mechanisms of dsRNA uptake in insects and potential of RNAi for pest control:a review.Journal of Insect Physiology,56:227-235.
KENNERDELL J R,CARTHEW R W,1998.Use of dsRNA-mediated genetic interference to demonstrate that frizzled and frizzled 2 act in the wingless pathway.Cell,95:1017-1026.
KNORR E,FISHILEVICH E,TENBUSCH L,FREY M L F,RANGASAMY M,BILLION A,WORDEN S E,GANDRA P,ARORA K,LO W,2018.Gene silencing in Tribolium castaneum as a tool for the targeted identification of candidate RNAi targets in crop pests.Scientific Reports,8:206.
KOUGH J L,EDELSTEIN R,2012.EPA regulatory requirements for plant-incorporated protectants//CHRIS W,MCHUGHEN A.Regulation of agricultural biotechnology:the United States and Canada:163-174.
LEE Y,KIM M,HAN J,YEOM K H,LEE S,BAEK S H,KIM V N,2004.MicroRNA genes are transcribed by RNA polymerase II.The EMBO Journal,23:4051-4060.
LEVINE S L,TAN J,MUELLER G M,BACHMAN P M,JENSEN P D,UFFMANJ P,2015.Independent action between DvSnf7 RNA and Cry3Bb1 protein in southern corn rootworm,Diabrotica undecimpunctata howardi and Colorado potato beetle,Leptinotarsa decemlineata.PLoS ONE,10:e0118622.
LI T,CHEN J,FAN X,CHEN W,ZHANG W,2017.MicroRNA and dsRNA targeting chitin synthase a reveal a great potential for pest management of the hemipteran insect Nilaparvata lugens.Pest Management Science,73:1529-1537.
LIN Y H,HUANG J H,LIU Y,BELLES X,LEE H J,2016.Oral delivery of dsRNA lipoplexes to German cockroach protects dsRNA from degradation and induces RNAi response.Pest Management Science,73:960-966.
MAO J,ZENG F,2013.Plant-mediated RNAi of a gap gene-enhanced tobacco tolerance against the Myzus persicae.Transgenic Research,23:145-152.
MEISTER G,TUSCHL T,2004.Mechanisms of gene silencing by double-stranded RNA.Nature,431:343-349.
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