杂交–渐渗进化理论在转基因逃逸及其环境风险评价和研究中的意义

doi:10.3724/SP.J.1003.2009.09132

摘要/Abstract

摘要：

转基因作物的商品化生产和大规模环境释放, 引起了全球对生物安全问题的广泛关注和争议, 其中转基因通过花粉介导的基因漂移逃逸到非转基因作物及其野生近缘种, 进而带来不同类型的环境风险就是备受争议的生物安全问题之一。有效的生物安全评价和研究能够为转基因作物的安全持久利用保驾护航。按照风险评价的原则, 对于转基因逃逸及其潜在环境风险的评价应包括两个重要步骤: (1)检测转基因向野生近缘种(包括杂草类型)群体逃逸的频率; (2)确定逃逸后的转基因能否通过遗传渐渗在野生近缘种群体中存留和扩散。杂交–渐渗是进化生物学中非常重要的科学命题和普遍的自然现象, 杂交–渐渗的进化理论与转基因逃逸及其潜在环境风险的研究和评价有密切的关系。杂交–渐渗过程往往导致物种形成、适应性进化和自然群体的濒危与灭绝, 这是因为在杂交–渐渗过程中, 不同的机制如遗传同化作用、群体湮没效应以及群体的选择性剔除效应等都会在很大程度上影响群体的进化过程。转基因通过杂交–渐渗进入野生群体, 使这一过程更加复杂化。如果转基因能提高群体的适合度, 则更有利于其渐渗速率, 从而在群体中迅速扩散并带来一定的生态后果。杂交–渐渗的进化理论和思想将有益于指导转基因逃逸及其潜在环境风险的研究和评价。

关键词: 生物安全, 转基因逃逸, 生态风险, 杂交, 渐渗, 进化, 适合度, 风险评价

Abstract

The commercial production and extensive environmental release of genetically modified (GM) crops have aroused worldwide concerns and debates over the biosafety of these crops. Transgene escape and its potential environmental risks are among the most debated biosafety issues. Transgene(s) can move from a GM crop to its non-GM counterparts and wild relatives via pollen-mediated gene flow, potentially causing various types of environmental problems. Effective biosafety assessment and research can facilitate the safe and sustainable application of GM crops. Following the framework of risk assessment, there are two critical steps for assessing environmental risks caused by transgene escape: (1) to measure frequencies of transgene escape from a GM crop to its non-GM counterpart or wild relative species (including weedy forms) via pollen-mediated gene flow; and (2) to determine the persist and spread of escaped transgene(s) in wild or weedy populations through introgression. The study of hybridization-introgression represents one of the most important and common phenomena in plant evolutionary, the study of which also includes the two important steps: to estimate frequencies of hybridization and to know the introgression of gene(s) in question within or among populations. The evolutionary theory of hybridization-introgression has a very close relationship with research and assessment of transgene escape and its potential environmental risks. The process of hybridization-introgression usually results in speciation, endangered status, extinction, or adaptive evolution of plant species. This is because important effects such as genetic assimilation, demographic swamping, and selective sweeps during the hybridization-introgression process, can considerably affect the evolutionary process of plant populations, into which the incorporation of transgenes may complicate the evolutionary process. If transgenes in question can significantly increase the fitness of individuals, they will quickly spread in the populations through fast introgression and significantly influence population dynamics bring certain evolutionary consequences. Therefore, we recommend applying the evolutionary theory of hybridization-introgression to guide the research and assessment of potential environmental risks caused by transgene escape.

Key words: biosafety, transgene escape, ecological consequence, hybridization, introgression, evolution, fitness, risk assessment

卢宝荣, 夏辉, 杨箫, 金鑫, 刘苹, 汪魏 (2009) 杂交–渐渗进化理论在转基因逃逸及其环境风险评价和研究中的意义. 生物多样性, 17, 362-377. DOI: 10.3724/SP.J.1003.2009.09132.

Bao-Rong Lu, Hui Xia, Xiao Yang, Xin Jin, Ping Liu, Wei Wang (2009) . Biodiversity Science, 17, 362-377. DOI: 10.3724/SP.J.1003.2009.09132.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2009.09132

https://www.biodiversity-science.net/CN/Y2009/V17/I4/362

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作物 Crop	逃逸对象 Recipient	实验方法 Method	转基因 Transgene	鉴定方法 Method of identification	转基因逃逸频率* Frequency of transgene escape	参考文献 References
转基因油菜 GM oil seed rape (Brassica napus)	白菜型油菜 B. rapa	田间调查 Field survey	EPSPS	除草剂筛选 Herbicide selection	1.1-17.5%	Simard et al., 2006
转基因油菜 GM oil seed rape (Brassica napus)	白菜型油菜 B. rapa	田间实验与调查 Field survey and designed experiment	EPSPS、 GFP	除草剂筛选、荧光检测、AFLP分子标记 Herbicide selection, GFP protein detection and AFLP markers	7.0-13.6%	Warwick et al., 2003
转基因油菜 GM oil seed rape (Brassica napus)	非转基因油菜 Non-GM B. napus	田间实验 Designed experiment	EPSPS	除草剂筛选 Herbicide selection	0.16% (~1 m) 0.05% (~5 m)	Du et al., 2009
匍匐翦股颖 GM creeping bentgrass (Agrostis gigantea)	匍匐翦股颖的自生株 Creeping bentgrass volunteer	田间调查 Field survey	EPSPS	除草剂筛选 Herbicide selection	34.0-93.0%	Zapiola et al., 2008
转基因棉花 GM cotton (Gossypium hirsutum)	非转基因棉花 Non-GM cotton	田间实验 Designed experiment	Bt	Bt基因特异性PCR Bt gene specific PCR	19.57% (~1 m)	Wang et al., 2007
转基因莴苣 GM lettuce (Lactuca sativa)	非转基因莴苣 Non-GM lettuce	田间实验 Designed experiment	KNAT、NPTII	卡那徽素筛选结合转基因特异性PCR Kanamycin selection and NPTII gene specific PCR	0.49% (~0.5 m) 0.071% (~11 m) 0.035% (22 m)	Giannino et al., 2008
转基因栽培稻 GM rice (Oryza sativa)	杂草稻 O. sativa f. spontanea	田间实验 Designed experiment	Bar	除草剂筛选 Herbicide selection	< 0.1%	Chen et al., 2004; Messeguer et al., 2004
转基因栽培稻 GM rice (Oryza sativa)	普通多年生野生稻 O. rufipogon	田间实验 Designed experiment	Bar	除草剂筛选 Herbicide selection	11-18% (~1 m) 0.01% (~250 m)	Wang et al., 2006
转基因栽培稻 GM rice (Oryza sativa)	非转基因栽培稻 Non-GM rice	田间实验 Designed experiment	Bt、CpTI、hpt	潮霉素筛选 Hygromycin selection	0.05-0.79% (< 1 m)	Rong et al., 2005
转基因栽培稻 GM rice (Oryza sativa)	非转基因的雄性不育系 Non-GM male sterile rice line	田间实验 Designed experiment	Bar	除草剂筛选 Herbicide selection	3.145-36.12%	Jia et al., 2007
转基因栽培稻 GM rice (Oryza sativa)	非转基因杂交稻 Non-GM hybrid rice	田间实验 Designed experiment	Bar	除草剂筛选 Herbicide selection	0.037-0.045%	Jia et al., 2007

作物 Crop	逃逸对象 Recipient	实验方法 Method	转基因 Transgene	鉴定方法 Method of identification	转基因逃逸频率* Frequency of transgene escape	参考文献 References
转基因油菜 GM oil seed rape (Brassica napus)	白菜型油菜 B. rapa	田间调查 Field survey	EPSPS	除草剂筛选 Herbicide selection	1.1-17.5%	Simard et al., 2006
转基因油菜 GM oil seed rape (Brassica napus)	白菜型油菜 B. rapa	田间实验与调查 Field survey and designed experiment	EPSPS、 GFP	除草剂筛选、荧光检测、AFLP分子标记 Herbicide selection, GFP protein detection and AFLP markers	7.0-13.6%	Warwick et al., 2003
转基因油菜 GM oil seed rape (Brassica napus)	非转基因油菜 Non-GM B. napus	田间实验 Designed experiment	EPSPS	除草剂筛选 Herbicide selection	0.16% (~1 m) 0.05% (~5 m)	Du et al., 2009
匍匐翦股颖 GM creeping bentgrass (Agrostis gigantea)	匍匐翦股颖的自生株 Creeping bentgrass volunteer	田间调查 Field survey	EPSPS	除草剂筛选 Herbicide selection	34.0-93.0%	Zapiola et al., 2008
转基因棉花 GM cotton (Gossypium hirsutum)	非转基因棉花 Non-GM cotton	田间实验 Designed experiment	Bt	Bt基因特异性PCR Bt gene specific PCR	19.57% (~1 m)	Wang et al., 2007
转基因莴苣 GM lettuce (Lactuca sativa)	非转基因莴苣 Non-GM lettuce	田间实验 Designed experiment	KNAT、NPTII	卡那徽素筛选结合转基因特异性PCR Kanamycin selection and NPTII gene specific PCR	0.49% (~0.5 m) 0.071% (~11 m) 0.035% (22 m)	Giannino et al., 2008
转基因栽培稻 GM rice (Oryza sativa)	杂草稻 O. sativa f. spontanea	田间实验 Designed experiment	Bar	除草剂筛选 Herbicide selection	< 0.1%	Chen et al., 2004; Messeguer et al., 2004
转基因栽培稻 GM rice (Oryza sativa)	普通多年生野生稻 O. rufipogon	田间实验 Designed experiment	Bar	除草剂筛选 Herbicide selection	11-18% (~1 m) 0.01% (~250 m)	Wang et al., 2006
转基因栽培稻 GM rice (Oryza sativa)	非转基因栽培稻 Non-GM rice	田间实验 Designed experiment	Bt、CpTI、hpt	潮霉素筛选 Hygromycin selection	0.05-0.79% (< 1 m)	Rong et al., 2005
转基因栽培稻 GM rice (Oryza sativa)	非转基因的雄性不育系 Non-GM male sterile rice line	田间实验 Designed experiment	Bar	除草剂筛选 Herbicide selection	3.145-36.12%	Jia et al., 2007
转基因栽培稻 GM rice (Oryza sativa)	非转基因杂交稻 Non-GM hybrid rice	田间实验 Designed experiment	Bar	除草剂筛选 Herbicide selection	0.037-0.045%	Jia et al., 2007

作物 Crop	野生近缘种 Wild relative	性状 Trait	目标转基因 Transgene	世代 Generation	对适合度影响 Impact to fitness	资料来源 References
向日葵 Helianthus annus	野生向日葵 H. annus (wild type)	抗虫 Insect resistance	Bt (cry1Ac)	BC₁	提高 Increase	Snow et al., 2003
甘蓝型油菜 Brassica napus	白菜型野油菜 B. rapa	抗虫 Insect resistance	Bt (cry1Ac)	F₁hybrids	提高 Increase	Vacher et al., 2004
甘蓝型油菜 Brassica napus	白菜型野油菜 B. rapa	抗虫 Insect resistance	Bt (cry1Ac)	F₁, BC₁F1, BC₂F₁, BC₂F₂	不显著 No significance	Halfhill et al., 2005
甘蓝型油菜 Brassica napus	白菜型野油菜 B. rapa	抗除草剂 Herbicide tolerance	Bar	BC₃	不显著 No significance	Snow et al., 1999
玉米 Zea mays	墨西哥杂草玉米 Z. mays ssp. mexicana	抗除草剂 Herbicide tolerance	EPSPS	F₁hybrids	不显著 No significance	Guadagnuolo et al., 2006
水稻 Oryza sativa	一年生野生稻 O. nivara	抗虫 Insect resistance	CpTI	F₃, F₄	不显著 No significance	Ye et al., 2008
水稻 Oryza sativa	杂草稻 O. sativa f. spontanea	抗除草剂 Herbicide tolerance	Bar	F₁, F₂	不显著 No significance	Zhang et al., 2003
向日葵 Helianthus annus	野生向日葵 H. annus (wild type)	抗病 Disease resistance	OxOx	BC₃	不显著 No significance	Burke & Rieseberg, 2003
甘蓝型油菜 Brassica napus	野芥菜 Sinapis arvensis	抗除草剂 Herbicide tolerance	Bar	BC₁-BC₇	降低 Decrease	Gueritaine et al., 2002

作物 Crop	野生近缘种 Wild relative	性状 Trait	目标转基因 Transgene	世代 Generation	对适合度影响 Impact to fitness	资料来源 References
向日葵 Helianthus annus	野生向日葵 H. annus (wild type)	抗虫 Insect resistance	Bt (cry1Ac)	BC₁	提高 Increase	Snow et al., 2003
甘蓝型油菜 Brassica napus	白菜型野油菜 B. rapa	抗虫 Insect resistance	Bt (cry1Ac)	F₁hybrids	提高 Increase	Vacher et al., 2004
甘蓝型油菜 Brassica napus	白菜型野油菜 B. rapa	抗虫 Insect resistance	Bt (cry1Ac)	F₁, BC₁F1, BC₂F₁, BC₂F₂	不显著 No significance	Halfhill et al., 2005
甘蓝型油菜 Brassica napus	白菜型野油菜 B. rapa	抗除草剂 Herbicide tolerance	Bar	BC₃	不显著 No significance	Snow et al., 1999
玉米 Zea mays	墨西哥杂草玉米 Z. mays ssp. mexicana	抗除草剂 Herbicide tolerance	EPSPS	F₁hybrids	不显著 No significance	Guadagnuolo et al., 2006
水稻 Oryza sativa	一年生野生稻 O. nivara	抗虫 Insect resistance	CpTI	F₃, F₄	不显著 No significance	Ye et al., 2008
水稻 Oryza sativa	杂草稻 O. sativa f. spontanea	抗除草剂 Herbicide tolerance	Bar	F₁, F₂	不显著 No significance	Zhang et al., 2003
向日葵 Helianthus annus	野生向日葵 H. annus (wild type)	抗病 Disease resistance	OxOx	BC₃	不显著 No significance	Burke & Rieseberg, 2003
甘蓝型油菜 Brassica napus	野芥菜 Sinapis arvensis	抗除草剂 Herbicide tolerance	Bar	BC₁-BC₇	降低 Decrease	Gueritaine et al., 2002