转基因栽培稻基因漂移是否会带来环境生物安全影响?

转基因通过基因漂移可以渐渗到作物的野生近缘种,由此而导致的环境风险是全球广泛关注的生物安全问题.有3个关键因素可以决定环境风险的程度:特定空间距离的转基因漂移频率,转基因在野生近缘种中的表达水平,以及转基因为野生近缘种群体带来的适合度效应.本文将根据现有研究结果,从上述3方面对转基因漂移到非转基因栽培稻、杂草稻和野生稻造成的潜在环境影响进行回顾.栽培稻品种之间的基因漂移频率很低,可以通过空间隔离或其他方法使其降低到可忽略的水平.在共同分布的环境中,栽培稻基因(包括转基因)向杂草稻和野生稻的漂移不可避免.尽管抗虫转基因(Bt或Bt/CpTI)在栽培稻和野生近缘种杂交后代中可以正常表达,但由于在低虫压环境中,抗虫转基因不会明显改变野生近缘种的适合度,抗虫转基因漂移所造成的环境影响十分有限.因此对基因漂移而言,抗虫转基因栽培稻的商品化种植应该比较安全.然而,抗除草剂转基因渐渗到杂草稻或野生稻会改变群体的适合度,可能会引起不可预测的环境后果.     

适合度分析对转基因逃逸潜在环境风险评价的意义北大核心CSCD

转基因作物的全球大规模种植引起了全世界的广泛关注甚至争议。经过遗传改良并具有自然选择优势的转基因作物进入商品化种植,可能将带来环境生物安全的顾虑。在这些生物安全的顾虑中,转基因通过花粉介导的基因漂移向栽培作物的野生近缘种逃逸及其导致的潜在环境风险,就是世人最为关注的环境生物安全问题之一。包括中国在内的许多国家,在转基因作物进行商品化生产之前都必须对转基因逃逸及其带来的潜在环境风险进行严格评价。按照风险评价的框架,转基因向野生近缘种逃逸及其带来潜在环境风险的评价包括3个连续的步骤:1)检测转基因漂移到作物野生近缘种的频率;2)分析转基因在野生近缘种中的表达;3)确定转基因对野生近缘种群体适合度和进化潜力的影响。大量基因漂移的研究结果已表明,转基因通过基因漂移向栽培作物邻近的野生近缘种群体逃逸难以避免,而转基因也会在作物的野生近缘种群体中正常表达。因此分析和评价转基因为野生近缘种带来的适合度效应,对于转基因逃逸及其环境风险的评价至关重要。对适合度的概念及其进化意义进行介绍,并对如何利用转基因的适合度效应分析转基因逃逸的环境风险,以及对此类环境风险进行研究和评价的具体方法予以介绍。上述知识和方法的掌握将有助于人们对转基因作物环境生物安全及其评价的全面理解。     

适合度分析对转基因逃逸潜在环境风险评价的意义

转基因作物的全球大规模种植引起了全世界的广泛关注甚至争议。经过遗传改良并具有自然选择优势的转基因作物进入商品化种植,可能将带来环境生物安全的顾虑。在这些生物安全的顾虑中,转基因通过花粉介导的基因漂移向栽培作物的野生近缘种逃逸及其导致的潜在环境风险,就是世人最为关注的环境生物安全问题之一。包括中国在内的许多国家,在转基因作物进行商品化生产之前都必须对转基因逃逸及其带来的潜在环境风险进行严格评价。按照风险评价的框架,转基因向野生近缘种逃逸及其带来潜在环境风险的评价包括3个连续的步骤:1)检测转基因漂移到作物野生近缘种的频率;2)分析转基因在野生近缘种中的表达;3)确定转基因对野生近缘种群体适合度和进化潜力的影响。大量基因漂移的研究结果已表明,转基因通过基因漂移向栽培作物邻近的野生近缘种群体逃逸难以避免,而转基因也会在作物的野生近缘种群体中正常表达。因此分析和评价转基因为野生近缘种带来的适合度效应,对于转基因逃逸及其环境风险的评价至关重要。对适合度的概念及其进化意义进行介绍,并对如何利用转基因的适合度效应分析转基因逃逸的环境风险,以及对此类环境风险进行研究和评价的具体方法予以介绍。上述知识和方法的掌握将有助于人们对转基因作物环境生物安全及其评价的全面理解。     

转基因水稻基因流的发生与生态学后果

中国是水稻的起源中心之一,分布着丰富的野生种质资源.自转基因水稻获得安全证书以来,转基因水稻与其近缘野生种间的基因流受到广泛关注.本文对转基因水稻基因流的发生及其可能引起的生态学后果进行了综述和展望.认为转基因水稻能够与栽培稻、野生稻、杂草稻、稗草等成功杂交,但基因流发生频率较低且变化较大.基因流成功发生后,由于转基因水稻具转基因新性状而有适合度优势,转基因可能只通过少数几代就进入野生种群.当转基因植株进入野生种群并在自然条件下长期存在时,转基因植株与近缘野生种间的竞争关系和相对适合度将决定混合种群的动态变化.研究转基因水稻基因流的影响及其长期生态学后果对合理保护与利用野生种质资源具有重要意义.     

转基因水稻外源基因向近缘种群漂流和渐渗的研究进展

水稻是我国最重要的粮食作物之一,我国有8亿以上的人口以稻米作为主食。但在水稻生产中,由于病、虫、草害及不良气候等逆境因子的影响,严重制约了水稻的高产、稳产。转基因生物技术的迅速发展,为水稻抗性育种提供了新途径。自20世纪80年代以来,我国全方位地开展了转基因水稻的研发,目前已经培育出大量的抗病、抗虫、抗除草剂和抗逆的转基因水稻品种,这将为提高我国水稻的生产力和确保粮食安全做出重要的贡献。但转基因水稻的基因漂流及其可能带来的生物安全问题备受关注。已有报道证明,外源转基因可以通过异交向非转基因品种和野生近缘种漂流。在不同的试验条件下,抗除草剂基因有0．05％-0．53％逃逸的可能,其向不育系的最大漂移频率可达4．518％。抗虫基因向相邻非转基因水稻的平均漂移频率最高为0．875％。因此,本文对水稻与其近缘野生种的杂交情况,转基因水稻外源基因向非转基因品种、野生近缘种以及野生非近缘种的漂流和渐渗及其潜在的生态环境风险等方面进行了简要分析,并对转基因水稻的发展进行了展望,以期为转基因水稻的安全应用提供参考。     

Bt/CpTI转基因稻及其非转基因亲本对照在间隔种植条件下的转基因漂移

随着转基因技术的迅速发展,越来越多的转基因作物被培育出来。转基因作物的外源转基因通过花粉传播向非转基因作物的漂移,会影响非转基因作物品种的种子纯度,从而可能导致一系列生物安全问题。为了研究转基因栽培水稻(Oryzasativa)中的外源转基因通过花粉介导向非转基因水稻品种逃逸的可能性及其频率,我们选用3个含双价抗虫基因(Bt/CpTI)的转基因水稻品系及其相对应的非转基因水稻亲本品种(近等基因系)进行了转基因漂移的实验。为了获取在近距离状况下转基因水稻与非转基因水稻品种之间的转基因漂移频率,采用了转基因与非转基因水稻品种间隔种植的栽培方式,分别在福建省福州市和海南省三亚市的转基因环境安全实验地进行实验,并利用潮霉素抗性筛选标记基因来鉴定转基因和非转基因稻的杂种。共检测了从非转基因水稻品种随机收获的70,056颗种子,以此计算转基因漂移频率。结果表明,在相邻种植的情况下,由这3个转基因水稻向对应的非转基因水稻品种的转基因漂移的频率比较低(0.275–0.832%)。如此近距离条件下获得的低转基因漂移频率表明,对于严格自花授粉的水稻而言,通过一定的隔离措施,能有效地降低由花粉介导的转基因漂移导致的非转基因种子混杂。     

转基因稻及其非转基因亲本对照在间隔种植条件下的转基因漂移

随着转基因技术的迅速发展, 越来越多的转基因作物被培育出来。转基因作物的外源转基因通过花粉传播向非转基因作物的漂移, 会影响非转基因作物品种的种子纯度, 从而可能导致一系列生物安全问题。为了研究转基因栽培水稻(Oryza sativa)中的外源转基因通过花粉介导向非转基因水稻品种逃逸的可能性及其频率, 我们选用3个含双价抗虫基因(Bt/CpTI)的转基因水稻品系及其相对应的非转基因水稻亲本品种(近等基因系)进行了转基因漂移的实验。为了获取在近距离状况下转基因水稻与非转基因水稻品种之间的转基因漂移频率, 采用了转基因与非转基因水稻品种间隔种植的栽培方式, 分别在福建省福州市和海南省三亚市的转基因环境安全实验地进行实验, 并利用潮霉素抗性筛选标记基因来鉴定转基因和非转基因稻的杂种。共检测了从非转基因水稻品种随机收获的70,056颗种子, 以此计算转基因漂移频率。结果表明, 在相邻种植的情况下, 由这3个转基因水稻向对应的非转基因水稻品种的转基因漂移的频率比较低(0.275–0.832％)。如此近距离条件下获得的低转基因漂移频率表明, 对于严格自花授粉的水稻而言, 通过一定的隔离措施, 能有效地降低由花粉介导的转基因漂移导致的非转基因种子混杂。     

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

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。     

海南万宁普通野生稻居群开花习性和生殖特性研究

对目前海南最大的普通野生稻居群——万宁普通野生稻居群的花期、花时、结实率和花粉育性进行了调查研究。结果发现,该普通野生稻居群内东部和西部群体花期相差15d;东部群体单穗平均结实率为8．67％,西部群体单穗平均结实率达62．1％,呈极显著差异;而东部和西部群体的花时基本相同,花粉育性与栽培稻均无显著差异。该居群东部群体与毗邻再生栽培稻的花期相遇,花时一致,满足基因漂移发生的部分必须条件。虽然目前尚无充分证据证明普通野生稻与其毗邻的栽培稻有基因漂移发生,但出于野生稻保护的需要,转基因水稻种植应设置安全距离,避开该野生稻分布区域。     

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

转基因作物的商品化生产和大规模环境释放在带来巨大利益的同时,也引起了全球对其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,进而导致的潜在环境和生态风险就是备受争议的生物安全问题之一。转基因植物的环境生物安全涉及两方面关键问题：如何科学评价转基因植物商品化种植以后带来的环境和生态影响;如何利用环境生物安全的研究成果来制定科学有效的风险监测和管理措施。对转基因逃逸及其潜在生态风险的科学评价应包括三个重要环节：（1）检测转基因的逃逸的频率;（2）检测转基因逃逸后的表达和遗传规律;（3）确定逃逸后的转基因对野生近缘种群体适合度的影响及其进化潜力,本文将围绕对转基因逃逸及其潜在环境风险的科学评价,以转基因水稻为案例来对转基因逃逸带来生态影响的研究好评价的进展进行简要介绍,并对目前依据风险评价研究成果制定的各种管理策略进行了讨论。只有提高对转基因生物环境安全研究和评价的水平,并制定有效的风险监测和管理措施,才能为我国转基因技术的发展和转基因产品的商品化应用保驾护航。     

Limited fitness advantages of crop-weed hybrid progeny containing insect-resistant transgenes (Bt/CpTI) in transgenic rice field

Background

The spread of insect-resistance transgenes from genetically engineered (GE) rice to its coexisting weedy rice (O. sativa f. spontanea) populations via gene flow creates a major concern for commercial GE rice cultivation. Transgene flow to weedy rice seems unavoidable. Therefore, characterization of potential fitness effect brought by the transgenes is essential to assess environmental consequences caused by crop-weed transgene flow.

Methodology/Principal Findings

Field performance of fitness-related traits was assessed in advanced hybrid progeny of F₄ generation derived from a cross between an insect-resistant transgenic (Bt/CpTI) rice line and a weedy strain. The performance of transgene-positive hybrid progeny was compared with the transgene-negative progeny and weedy parent in pure and mixed planting of transgenic and nontransgenic plants under environmental conditions with natural vs. low insect pressure. Results showed that under natural insect pressure the insect-resistant transgenes could effectively suppress target insects and bring significantly increased fitness to transgenic plants in pure planting, compared with nontransgenic plants (including weedy parent). In contrast, no significant differences in fitness were detected under low insect pressure. However, such increase in fitness was not detected in the mixed planting of transgenic and nontransgenic plants due to significantly reduced insect pressure.

Conclusions/Significance

Insect-resistance transgenes may have limited fitness advantages to hybrid progeny resulted from crop-weed transgene flow owning to the significantly reduced ambient target insect pressure when an insect-resistant GE crop is grown. Given that the extensive cultivation of an insect-resistant GE crop will ultimately reduce the target insect pressure, the rapid spread of insect-resistance transgenes in weedy populations in commercial GE crop fields may be not likely to happen.     

Gene flow from genetically modified rice to its wild relatives: Assessing potential ecological consequences

Pollen-mediated gene flow is the major pathway for transgene escape from GM rice to its wild relatives. Transgene escape to wild Oryza species having AA-genome will occur if GM rice is released to environments with these wild Oryza species. Transgenes may persist to and spread in wild populations after gene flow, resulting unwanted ecological consequences. For assessing the potential consequences caused by transgene escape, it is important to understand the actual gene flow frequencies from GM rice to wild relatives, transgene expression and inheritance in the wild relatives, as well as fitness changes that brought to wild relatives by the transgenes. This article reviews studies on transgene escape from rice to its wild relatives via gene flow and its ecological consequences. A framework for assessing potential ecological consequences caused by transgene escape from GM rice to its wild relatives is discussed based on studies of gene flow and fitness changes.     

Performance of Hybrids between Weedy Rice and Insect-resistant Transgenic Rice under Field Experiments： Implication for Environmental Biosafety Assessment

Transgene escape from genetically modified (GM) rice Into weedy rice via gene flow may cause undesired environmental consequences. Estimating the field performance of crop-weed hybrids will facilitate our understanding of potential introgression of crop genes (including transgenes) into weedy rice populations, allowing for effective biosafety assessment. Comparative studies of three weedy rice strains and their hybrids with two GM rice lines containing different insect-resistance transgenes (CpTl or BtlCpTI) indicated an enhanced relative performance of the crop-weed hybrids, with taller plants, more tillers, panicles, and spikelets per plant, as well as higher 1000-seed weight, compared with the weedy rice parents, although the hybrids produced less filled seeds per plant than their weedy parents. Seeds from the F1 hybrids had higher germination rates and produced more seedlings than the weedy parents, which correlated positively with 1000-seed weight. The crop-weed hybrids demonstrated a generally enhanced relative performance than their weedy rice parents in our field experiments. These findings indicate that transgenes from GM rice can persist to and introgress into weedy rice populations through recurrent crop-to-weed gene flow with the aid of slightly increased relative fitness in F1 hybrids.     

Genes invading new populations: a risk assessment perspective

One of the concerns raised over the introduction of genetically modified crops is that transgenes will invade populations of wild relatives, causing ecologically significant changes in fitness. In recent years, this has given rise to several studies estimating hybridization rates and the fitness of crop-wild relative hybrids. These studies have established that transgenes are likely to move to F1 hybrids, albeit at low frequency. Hybridization, however, is not synonymous with introgression, and questions remain as to whether particular transgenes will cause ecologically significant changes in recipient plant populations. Research effort should now focus on estimating any changes in the fitness of a population as a consequence of having a transgene, understanding genotype x environment interactions, and deducing the extent to which pathogens and herbivores (transgene targets) regulate wild relative populations. This will involve a combination of manipulative experiments and empirically motivated mathematical models.     

Persistence of transgenes in wild rice populations depends on the interaction between genetic background of recipients and environmental conditions

The persistence of transgenes in wild populations may cause unintended ecological consequences, and the possibility of transgenes' persistence and introgression is dependent on fitness performance of transgenic crop–wild hybrids. To investigate the effects of transgene and genotype × environment on the fitness of crop–wild rice hybrids, a total of 11 cross‐combination progenies between insect‐resistant transgene (CpTI and Bt/CpTI) rice lines and wild rice (Oryza rufipogon) were evaluated at different sites with contrasting insect treatments. The results showed that fitness performance varied between transgenic hybrids having different wild parents and under different environmental conditions, indicating that fitness effects of transgenes on hybrid progenies depend heavily on the genetic background of recipient plants and growing environment. Significant fitness advantages conferred by transgenes were found only in some hybrids under high insect pressure condition, demonstrating that the level of target insects in the field environment influences the persistence and spread of insect‐resistant transgenes in wild rice populations. These findings suggest that evolutionary fate of escaped transgenes is different in wild populations with diverse genetic backgrounds under various environmental conditions.     

Gene flow from cultivated rice (Oryza sativa) to its weedy and wild relatives

BACKGROUND AND AIMS: Transgene escape through gene flow from genetically modified (GM) crops to their wild relative species may potentially cause environmental biosafety problems. The aim of this study was to assess the extent of gene flow between cultivated rice and two of its close relatives under field conditions. METHODS: Experiments were conducted at two sites in Korea and China to determine gene flow from cultivated rice (Oryza sativa L.) to weedy rice (O. sativa f. spontanea) and common wild rice (O. rufipogon Griff.), respectively, under special field conditions mimicking the natural occurrence of the wild relatives in Asia. Herbicide resistance (bar) and SSR molecular finger printing were used as markers to accurately determine gene flow frequencies from cultivated rice varieties to their wild relatives. KEY RESULTS: Gene flow frequency from cultivated rice was detected as between approx. 0.011 and 0.046 % to weedy rice and between approx. 1.21 and 2.19 % to wild rice under the field conditions. CONCLUSIONS: Gene flow occurs with a noticeable frequency from cultivated rice to its weedy and wild relatives, and this might cause potential ecological consequences. It is recommended that isolation zones should be established with sufficient distances between GM rice varieties and wild rice populations to avoid potential outcrosses. Also, GM rice should not be released when it has inserted genes that can significantly enhance the ecological fitness of weedy rice in regions where weedy rice is already abundant and causing great problems.     

Evaluating genetic containment strategies for transgenic plants

One of the primary concerns about genetically engineered crop plants is that they will hybridize with wild relatives, permitting the transgene to escape into the environment. The likelihood that a transgene will spread in the environment depends on its potential fitness impact. The fitness conferred by various transgenes to crop and/or wild-type hybrids has been evaluated in several species. Different strategies have been developed for reducing the probability and impact of gene flow, including physical separation from wild relatives and genetic engineering. Mathematical models and empirical experimental evidence suggest that genetic approaches have the potential to effectively prevent transgenes from incorporating into wild relatives and becoming established in wild populations that are not reproductively isolated from genetically engineered crops.     

The establishment of genetically engineered canola populations in the U.S

Concerns regarding the commercial release of genetically engineered (GE) crops include naturalization, introgression to sexually compatible relatives and the transfer of beneficial traits to native and weedy species through hybridization. To date there have been few documented reports of escape leading some researchers to question the environmental risks of biotech products. In this study we conducted a systematic roadside survey of canola (Brassica napus) populations growing outside of cultivation in North Dakota, USA, the dominant canola growing region in the U.S. We document the presence of two escaped, transgenic genotypes, as well as non-GE canola, and provide evidence of novel combinations of transgenic forms in the wild. Our results demonstrate that feral populations are large and widespread. Moreover, flowering times of escaped populations, as well as the fertile condition of the majority of collections suggest that these populations are established and persistent outside of cultivation.     

Glyphosate-drift but not herbivory alters the rate of transgene flow from single and stacked trait transgenic canola (Brassica napus) to nontransgenic B. napus and B. rapa

? Transgenic plants can offer agricultural benefits, but the escape of transgenes is an environmental concern. In this study we tested the hypothesis that glyphosate drift and herbivory selective pressures can change the rate of transgene flow between the crop Brassica napus (canola), and weedy species and contribute to the potential for increased transgene escape risk and persistence outside of cultivation. ? We constructed plant communities containing single transgenic B. napus genotypes expressing glyphosate herbicide resistance (CP4 EPSPS), lepidopteran insect resistance (Cry1Ac), or both traits ('stacked'), plus nontransgenic B. napus, Brassica rapa and Brassica nigra. Two different selective pressures, a sublethal glyphosate dose and lepidopteran herbivores (Plutella xylostella), were applied and rates of transgene flow and transgenic seed production were measured. ? Selective treatments differed in the degree in which they affected gene flow and production of transgenic hybrid seed. Most notably, glyphosate-drift increased the incidence of transgenic seeds on nontransgenic B. napus by altering flowering phenology and reproductive function. ? The findings of this study indicate that transgenic traits may be transmitted to wild populations and may increase in frequency in weedy populations through the direct and indirect effects of selection pressures on gene flow.     

The persistence of cultivar alleles in wild populations of sunflowers five generations after hybridization

 The development of transgenic plants has heightened concern about the possible escape of genetically engineered material into the wild. Hybridization between crops and their wild relatives provides a mechanism by which this could occur. While hybridization has been documented between several crops and wild or weedy relatives, little is known about the persistence of cultivar genes in wild populations in the generations following hybridization. Wild and weedy sunflowers occur sympatrically with cultivated sunflowers throughout much of the cultivation range, and hybridization is known to occur. We surveyed two cultivar-specific RAPD markers in 2700 progeny in a naturally occurring population of wild Helianthus annuus over five generations following a single generation of hybridization with the cultivar. Moderate levels of gene flow were detected in the first generation (42% hybrids at the crop margin) and cultivar allele frequencies did not significantly decline over four subsequent generations. These results indicate that gene flow from cultivated into wild populations of sunflowers can result in the long-term establishment of cultivar alleles in wild populations. Furthermore, we conclude that neutral or favorable transgenes have the potential to escape and persist in wild sunflower populations. Received: 1 November 1996/Accepted: 17 January 1997