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.     

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.     

Effects of Bt transgenic rice line on stripe stem borer,Chilo suppressalis and its consequences on egg parasitoid,Trichogramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) in the laboratory

Interactions among Chilo suppressalis (Walker), its larval parasitoid Trichogramma brassicae Bezdenko and insect-resistant transgenic rice lines were evaluated under laboratory conditions. The results showed that all neonates and second larvae of stem borer were dead regardless of being fed rotationally or permanently on Bt rice, but 18% and 28% of the third and fourth instar larvae could complete development and turned to adults, respectively, when fed rotationally. In the second year of project, third instar larvae of stem borer were used to evaluate Bt rice impact on T. brassicae. The result showed that emergence rate was lower, the abnormal female rate increased and adult female longevity of the wasp decreased significantly in comparison with the conventional variety. There was no significant difference in gender ratio and parasitism rate under Bt rice treatment in comparison with conventional variety.     

Occurrence and field densities of Coleoptera in the maize herb layer: implications for Environmental Risk Assessment of genetically modified <Emphasis Type="Italic">Bt</Emphasis>-maize

Beetles (Coleoptera) are a diverse and ecologically important group of insects in agricultural systems. The Environmental Risk Assessment (ERA) of genetically modified Bt-crop varieties with insect resistances thus needs to consider and assess the potential negative impacts on non-target organisms belonging to this group. We analysed data gathered during 6 years of field-release experiments on the impact of two genetically modified Bt-maize varieties (Ostrinia-resistant MON810 and Diabrotica-resistant MON88017) on the occurrence and field densities of Coleoptera, especially the two families Coccinellidae and Chrysomelidae. Based on a statistical analysis aimed at establishing whether Bt-maize varieties are equivalent to their near-isogenic counterparts, we discuss the limitations of using field experiments to assess the effects of Bt-maize on these two beetle families. The densities of most of the beetle families recorded in the herb layer were very low in all growing seasons. Coccinellidae and Chrysomelidae were comparatively abundant and diverse, but still low in numbers. Based on their role as biological control agents, Coccinellidae should be a focus in the ERA of Bt-plants, but given the large natural variability in ladybird densities in the field, most questions need to be addressed in low-tier laboratory tests. Chrysomelidae should play a negligible role in the ERA of Bt-plants, since they occur on-crop as secondary pests only. Species occurring off-crop, however, can be addressed in a similar fashion as non-target Lepidoptera in Cry1Ab expressing Bt-maize.     

Development of insect-resistant transgenic indica rice with a synthetic cry1C* gene

Stemborers and leaffolders are two groups of lepidopteran pests that cause severe damage to rice in many areas of the world. In this study, a cry1C* gene encoding Bacillus thuringiensis (Bt) δ-endotoxin was synthesized by codon optimization as the first step towards gene stacking in our resistance management strategy of transgenic rice. Agrobacterium-mediated transformation of this gene into Minghui 63 (Oryza sativa L.), an elite indica CMS restorer line, produced 120 independently transformed plants, 19 of which had a single-copy transgene. Preliminary screening of T₁ families of these 19 transformants in the field identified five lines showing a high level of resistance to leaffolders (Cnaphalocrocis medinalis) and stemborers. Hybrids were produced by crossing these five lines with Zhenshan 97A, the male-sterile line for Shanyou 63, the most widely cultivated hybrid in China. These five lines and their hybrids were highly resistant to yellow stemborer (Tryporyza incertulas) as revealed by an insect bioassay. The content of Cry1C* protein varied considerably among the five lines as well as among the corresponding hybrids. T1c-19, a line showing the highest content of Cry1C* protein, and its hybrid were tested in the field for insect resistance and agronomic performance and found to be highly resistant to stemborers and leaffolders throughout the growth period, resulting in a significantly increased grain yield compared with the respective controls. These results indicate that T1c-19 can be used for production of insect-resistant hybrid rice and as a germplasm for gene stacking to produce rice plants with two toxins.     

Laboratory assessment of the impacts of transgenic Bt rice on the ecological fitness of the soil non-target arthropod, Folsomia candida (Collembola: Isotomidae)

Transgenic rice expressing Bacillus thuringiensis (Bt) endotoxins (Bt rice) for pest control is considered an important solution to food security in China. However, tests for potential effects on non-target soil organisms are required for environmental risk assessment. The soil collembolan Folsomia candida L. (Collembola: Isotomidae) is a potential non-target arthropod that is often used as a biological indicator in bio-safety assessments of transgenic crops. In the present study, the roots, stems, and leaves of Bt rice were exposed to F. candida under laboratory conditions, with survival, reproduction and growth of the collembolan as ecological fitness parameters. Significant differences in ecological fitness were found among the different treatments, including differences in the plant parts and varieties of non-Bt rice, presumably as the result of three factors: gene modification, plant parts and rice varieties. The fitness of F. candida was less affected by the different diets than by the exposure to the same materials mixed with soil. Our results clearly showed that there was no negative effect of different Bt rice varieties on the fitness of F. candida through either diet or soil exposure.     

转基因逃逸及其环境生物安全评价研究进展——抗虫水稻案例分析

转基因技术研发为提高我国水稻产量和减少劳动力投入提供了巨大机遇。我国对转基因水稻研发进行了大量的投入,目前已培育了具有不同新性状的转基因水稻品系,许多品系已进入生物安全评价阶段。风险评价对转基因水稻的安全生产至关重要,是其商品化生产之前必须解决的问题,其中包括转基因逃逸及其潜在环境影响。对水稻抗虫转基因逃逸及其潜在环境风险的评价包括3个重要环节：（1）通过田间试验和模型模拟检测转基因漂移到非转基因栽培稻及其野生近缘种的频率;（2）检测转基因在栽培稻和野生近缘种后代中的表达;（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。大量研究表明,在近距离的空间范围内栽培稻品种之间的基因漂移频率很低（〉0.1%）,但栽培稻与其野生近缘种的基因漂移频率变异很大。进一步研究还表明,Bt抗虫转基因在栽培稻与普通野生稻后代中均能正常表达,但在其不同生长阶段,表达量有很大变异。在有较高水平的害虫虫压下,含有抗虫转基因的栽培稻及野生近缘种杂交后代与不含转基因的对照相比,抗虫性显著提高且适合度利益明显;但是,在虫害发生水平较低时,含有抗虫转基因的群体与不含抗虫转基因的群体相比没有显著的适合度优势。综上,转基因逃逸到非转基因水稻的频率极低,并且可以通过空间隔离阻断其逃逸。虽然抗虫转基因向杂草稻以及与栽培稻距离较近的野生稻群体的逃逸无法避免,但是野生稻和杂草稻群体周围环境中的总体虫压较低,所以基因漂移带来的环境影响应十分有限。     

Bt rice could provide ecological resistance against nontarget planthoppers

Genetically engineered (GE) rice lines expressing Lepidoptera‐active insecticidal cry genes from the bacterium Bacillus thuringiensis (Bt) have been developed in China. Field surveys indicated that Bt rice harbours fewer rice planthoppers than non‐Bt rice although planthoppers are not sensitive to the produced Bt Cry proteins. The mechanisms underlying this phenomenon remain unknown. Here, we show that the low numbers of planthoppers on Bt rice are associated with reduced caterpillar damage. In laboratory and field‐cage experiments, the rice planthopper Nilapavata lugens had no feeding preference for undamaged Bt or non‐Bt plants but exhibited a strong preference for caterpillar‐damaged plants whether Bt or non‐Bt. Under open‐field conditions, rice planthoppers were more abundant on caterpillar‐damaged non‐Bt rice than on neighbouring healthy Bt rice. GC–MS analyses showed that caterpillar damage induced the release of rice plant volatiles known to be attractive to planthoppers, and metabolome analyses revealed increased amino acid contents and reduced sterol contents known to benefit planthopper development. That Lepidoptera‐resistant Bt rice is less attractive to this important nontarget pest in the field is therefore a first example of ecological resistance of Bt plants to nontarget pests. Our findings suggest that non‐Bt rice refuges established for delaying the development of Bt resistance may also act as a trap crop for N. lugens and possibly other planthoppers.     

Scale effect on rice pollen‐mediated gene flow: implications in assessing transgene flow from genetically engineered plants

Transgene flow from genetically engineered (GE) crops to non‐GE varieties and wild relatives via pollen‐mediated gene flow (PMGF) may create food and environmental biosafety concerns. Assessing the level of PMGF from GE crops is required before commercialization. Whether the level of PMGF estimated at relatively small scales can sufficiently represent the actual scenario at large production scales remains unresolved. Here, we estimated average PMGF frequencies from three insect‐resistant GE rice lines to their non‐GE counterparts at four scales ranging from 9 to 576 m², having the number of GE to non‐GE plants constantly at the ratio of 8:1. Based on nearly 1.3 million examined seedlings from non‐GE rice plots, very low frequencies (<0.1%) of transgene flow were detected. The highest frequencies were found in plots at the smallest scales. Scale had a significantly negative effect on the frequency of PMGF in rice, with decreased gene flow at increased scale. An extended PMGF model could well represent the experimental data. Field experiments at relatively small scales combined with mathematical modelling could provide useful prediction on the level of rice transgene flow at large production scales. This is probably applicable for other crop species with wind‐ and self‐pollination.     

Bt rice expressing Cry1Ab does not stimulate an outbreak of its non-target herbivore, Nilaparvata lugens

In this study, the non-target effects of Bt rice “KMD2” expressing a Cry1Ab protein on the performance of the brown planthopper (BPH), Nilaparvata lugens, over multiple generations were evaluated under laboratory and field conditions. In the laboratory, BPH was reared to observe the impact of the Bt rice as compared to its parental non-Bt cultivar Xiushui 11, while the population dynamics and oviposition performance of BPH were investigated in the field. The survival of BPH nymphs fed Bt and non-Bt rice did not differ significantly. The nymph developmental duration of BPH was significantly delayed by the Bt rice by comparison with the non-Bt rice for the 1st and 2nd but not the 4th generation. Most importantly, the fecundity of BPH on the Bt rice was significantly decreased in every generation when compared with the non-Bt rice. In the field investigations, the population density of BPH nymphs was significantly lower in the Bt rice field. However, the temporal pattern of population dynamics of BPH adults was similar between the Bt and non-Bt rice, presumably due to migratory interference of the adults. In the Bt rice field, the percentage of tillers with eggs and the number of eggs per tiller were also significantly lower from tillering to mature stage. Additionally, Cry1Ab protein could not be detected in guts from single BPH adults. In general, our results suggest that the Bt rice “KMD2” could not stimulate an outbreak of BPH.     

No effect of transgene and strong wild parent effects on seed dormancy in crop–wild hybrids of rice: implications for transgene persistence in wild populations

Soil seed banks act as a gene pool for local plant species and, as such, can buffer local populations, especially those experiencing challenging environmental conditions. Seed dormancy has important implications to dynamics of soil seed banks. Therefore, estimating the seed dormancy of transgenic crop–wild hybrids could shed light on the persistence of transgenes in wild‐plant soil seed banks. Individuals from eight populations of wild rice Oryza rufipogon were crossed with those of three insect‐resistant transgenic rice lines. Selfed (F2–F4) and backcrossed populations (BC1, BC1F2 and BC1F3) were then made from the hybrids. Seed germination was tested under three treatments: (a) normal; (b) overwintering in soil; and (c) one‐week heat‐shocking. The effects of transgene, wild parent and hybrid generation on hybrid seed germination were examined. No significant effect of insect‐resistant transgenes (Bt and CpTI) was detected on the seed dormancy of crop–wild hybrids, while a significant wild parent effect was found. The seeds of advanced generation hybrids have higher germination percentages and lower dormancy than do those of F1 and BC1 generations. The study showed that the dormancy of hybrid seeds was determined mainly by their genetic backgrounds. All hybrid seeds have higher germination percentages and lower dormancy (and, consequently, a poorer overwintering ability), compared with wild seeds, and reduce dormancy would contribute to a fitness disadvantage, compared with wild types. Therefore, such seeds might form part of naturally occurring soil seed banks, through which crop genes would persist in wild populations.     

Normal expression of insect-resistant transgene in progeny of common wild rice crossed with genetically modified rice: its implication in ecological biosafety assessment

Transgene outflow from genetically modified (GM) rice to its wild relatives may cause undesirable ecological consequences. Understanding the level of transgene expression in wild rice following gene flow is important for assessing such consequences, providing that transgene escape from GM rice cannot be prevented. To determine the expression of a transgene in common wild rice (Oryza rufipogon), we analyzed the content of Cry1Ac protein in three GM rice lines containing a Bt transgene, their F₁ hybrids with common wild rice and F₂ progeny at different growth stages, using the sandwich enzyme-linked immunosorbent assay. The average content of Cry1Ac protein in leaf samples of the wild rice lines ranged between 0.016 and 0.069% during the entire growth period, whereas that in stems varied between 0.12 and 0.39%. A great variation in Cry1Ac protein content was detected among individuals of F₁ hybrids and F₂ progeny, with some wild individuals showing higher level of Bt toxin than the cultivated GM rice. The results suggest that the Bt transgene can express normally in the interspecific hybrids between insect-resistant GM rice and common wild rice, and may have similar effects on the target insects as in GM rice.     

Non-target organism effects tests on Vip3A and their application to the ecological risk assessment for cultivation of MIR162 maize

Transgenic crops producing insecticidal proteins from Bacillus thuringiensis (Bt) provide economic, environmental and health benefits by maintaining or increasing crop yields with fewer applications of insecticide. To sustain these benefits, it is important to delay the evolution of insect resistance to the proteins, and to ensure that the proteins do not harm non-target organisms, particularly those that may control secondary pests that would otherwise flourish because of reduced insecticide applications. Vip3A is a Bt vegetative insecticidal protein that is active against lepidopterous pests. It has a different mode of action from other proteins for control of Lepidoptera in current Bt crops, and when combined with these proteins, it should help to delay the evolution of pest resistance to Bt crops. This paper presents data on the effects of Vip3A on non-target organisms, and an ecological risk assessment of MIR162 maize, which expresses Vip3Aa20. Laboratory studies indicate few adverse effects of Vip3A to non-target organisms: 11 of 12 species tested showed no adverse effects when exposed to high concentrations of Vip3A relative to estimated exposures resulting from cultivation of MIR162 maize. Daphnia magna exposed to Vip3Aa20 were unaffected in terms of survival or fecundity, but grew slightly more slowly than unexposed controls. The data indicate that cultivation of MIR162 maize poses negligible risk to non-target organisms, and that crops producing Vip3A are unlikely to adversely affect biological control organisms such that benefits from reduced insecticide applications are lost.     

Field evaluation and risk assessment of transgenic indica basmati rice

We report the first field trial of different transgenic lines of Indica Basmati rice (B-370) expressing cry1Ac and cry2A genes. Different transgenic lines were grown under field conditions for two consecutive years, according to RCBD and Split Plot Design respectively. All the biosafety measures were taken into consideration. Sixty neonate larvae of yellow stem borer were artificially infested into each plant in three installments. Data was recorded in terms of dead hearts and white heads at vegetative and flowering stage respectively. Transgenic lines exhibited inherent ability to protect rice plants from target insects (p<0.01). Natural infestations of rice skipper and rice leaf folder were also observed and transgenic plants were statistically superior to their untransformed counterparts. Green house whole plant bioassays were done by infesting two 2^nd instar larvae of rice leaf folder per tiller. Transgenics were 96% more resistant than untransformed control plants. The presence of cry genes was observed with Dot blot, PCR and Southern blot analysis, while ELISA and Western blot analysis confirmed the expression of Cry proteins. All lines expressed higher level of Cry proteins when compared with commercially released cultivars of Bt cotton, maize and potato. It was also observed that although toxin titer substantially decreased with increasing age of the plants, it remained well within the limits to kill the target insects. Morphological studies showed significant variation for days to maturity, plant height and panicle length. Cooking qualities of seeds harvested from these lines were compared with the untransformed control. The transgenic lines had no effect on non-target insects (insects belonging to orders other than diptera and lepidoptera) and germination of three local varieties of wheat. Chances of gene spread were calculated at a level of 0.18% cross pollination in experimental lines.     

Susceptibility and tolerance in hybrid and pure‐line rice varieties to herbivore attack: biomass partitioning and resource‐based compensation in response to damage

Hybrid rice has been noted for its susceptibility to insects and diseases compared to pure‐line (conventional) rice varieties. We investigated herbivory by Nilaparvata lugens, Sogatella furcifera and Scirpophaga incertulas on replicated three‐line hybrid sets (parental and hybrid lines) in field and greenhouse experiments. In a field experiment, caterpillar densities and stemborer damage was similar among hybrid and parental lines. In field and greenhouse experiments, the cytoplasmic male sterile (CMS)‐lines and maintainer lines had higher densities of planthoppers (including N. lugens and S. furcifera) than restorer or hybrid lines likely because of their wild abortive CMS‐lineage. High nitrogen levels increased plant mortality due to N. lugens, but often reduced mortality from S. furcifera and S. incertulas: this was similar between hybrid and pure‐line varieties. The hybrids were generally more tolerant of herbivory (lower biomass reductions per unit weight of insect) than the inbred parental lines. The addition of nitrogen to both the hybrid and pure‐line varieties had contrasting effects on tolerance depending on the nature of the attacking insect: fertiliser increased tolerance to S. furcifera (lower losses of yield and shoot biomass per mg insect) and S. incertulas (lower yield, shoot and root biomass loss) but fertiliser reduced tolerance to N. lugens (higher loss of root biomass and no effects on yield and shoot biomass loss). Our results indicate that hybrid rice is not physiologically more susceptible to herbivores than are pure‐line varieties even under high nitrogen conditions, but does have higher tolerance to insect damage.     

Field evaluation of effects of transgenic <Emphasis Type="Italic">cry1Ab/cry1Ac,cry1C</Emphasis> and <Emphasis Type="Italic">cry2A</Emphasis> rice on <Emphasis Type="Italic">Cnaphalocrocis medinalis</Emphasis> and its arthropod predators

The impacts of transgenic Bt rice on target pests and their predators need to be clarified prior to the commercialization of Bt rice. In this study, the percentages of folded leaves of three transgenic Bt rice lines and non-transgenic parental rice line caused by Cnaphalocrocis medinalis were studied over two successive growing seasons. In addition, the population densities, relative abundance and population dynamics of C. medinalis and four species of its natural arthropod predators were investigated at three sites in China. The results showed that rice line significantly affected the percentages of folded leaves and population densities of C. medinalis larvae. Significantly higher percentages of folded leaves were observed on the non-transgenic rice compared with the three transgenic Bt rice on most sampling dates. Significantly higher densities of C. medinalis larvae and higher relative abundance of C. medinalis within phytophages were found on non-transgenic rice compared with three transgenic Bt rice at different sites across the study period. The population dynamics of C. medinalis larvae were significantly affected by rice line, rice line×sampling date, rice line×year, rice line×sampling date×year. However, there was little, if any, significant difference in the relative abundance, population density and population dynamics of the four arthropod predators between the three Bt rice lines and non-transgenic rice. The results of this study indicate that the Bt toxin in transgenic Bt rice can effectively suppress the occurrence of C. medinalis, but has no significant effects on the occurrence of the four predatory arthropod species.     

Impact of insect-resistant transgenic rice on target insect pests and non-target arthropods in China

Progress on the research and development of insect-resistant transgenic rice, especially expressing insecticidal proteins from Bacillus thuringiensis （Bt）, in China has been rapid in recent years. A number of insect-resistant transgenic rice lines/varieties have passed restricted and enlarged field testing, and several have been approved for productive testing since 2002 in China, although none was approved for commercial use until 2006. Extensive laboratory and field trials have been conducted for evaluation of the efficiency of transgenic rice on target lepidoteran pests and potential ecological risks on non-target arthropods. The efficacy of a number of transgenic rice lines currently tested in China was excellent for control of the major target insect pests, the rice stem borers （Chilo suppressalis, Scirpophaga incertulas, Sesamia inferens） and leaffolder （ Cnaphalocrocis medinalis）, and was better than most insecticides extensively used by millions of farmers at present in China. No significantly negative or unintended effects of transgenic rice on non-target arthropods were found compared with non-transgenic rice. In contrast, most of the current insecticides used for the control of rice stem borers and leaffolders proved harmful to natural enemies, and some insecticides may directly induce resurgence of rice planthoppers. Studies for developing a proactive insect resistance management of transgenic rice in the future are discussed to ensure the sustainable use of transgenic rice.     

Effects of seed mixture sowing with transgenic Bt rice and its parental line on the population dynamics of target stemborers and leafrollers,and non‐target planthoppers

The widespread planting of insect‐resistant crops has caused a dramatic shift in agricultural landscapes, thus raising concerns about the potential impacts on both target and non‐target pests. In this study, we examined the potential effects of intra‐specific seed mixture sowing with transgenic Bt rice (Bt) and its parental non‐transgenic line (Nt) (100% Bt rice [Bt₁₀₀], 5% Nt+95% Bt [Nt₀₅Bt₉₅], 10% Nt+90% Bt [Nt₁₀Bt₉₀], 20% Nt+80% Bt [Nt₂₀Bt₈₀], 40% Nt+60% Bt [Nt₄₀Bt₆₀] and 100% Nt rice [Nt₁₀₀]) on target and non‐target pests in a 2‐year field trial in southern China. The occurrence of target pests, Sesamia inferens, Chilo suppressalis and Cnaphalocrocis medinalis, decreased with the increased ratio of Bt rice, and the mixture ratios with more than 90% Bt rice (Bt₁₀₀ and Nt₀₅Bt₉₅) significantly increased the pest suppression efficiency, with the lowest occurrences of non‐target planthoppers, Nilaparvata lugens and Sogatella furcifera in Nt₁₀₀ and Nt₀₅Bt₉₅. Furthermore, there were no significant differences in 1000‐grain dry weight and grain dry weight per 100 plants between Bt₁₀₀ and Nt₀₅Bt₉₅. Seed mixture sowing of Bt rice with ≤10% (especially 5%) of its parent line was sufficient to overcome potential compliance issues that exist with the use of block or structured refuge to provide most effective control of both target and non‐target pests without compromising the grain yield. It is also expected that the strategy of seed mixture sowing with transgenic Bt rice and the non‐transgenic parental line would provide rice yield stability while decreasing the insecticide use frequency in rice production.     

The development and status of Bt rice in China

Multiple lines of transgenic rice expressing insecticidal genes from the bacterium Bacillus thuringiensis (Bt) have been developed in China, posing the prospect of increases in production with decreased application of pesticides. We explore the issues facing adoption of Bt rice for commercial production in China. A body of safety assessment work on Bt rice has shown that Bt rice poses a negligible risk to the environment and that Bt rice products are as safe as non‐Bt control rice products as food. China has a relatively well‐developed regulatory system for risk assessment and management of genetically modified (GM) plants; however, decision‐making regarding approval of commercial production has become politicized, and two Bt rice lines that otherwise were ready have not been allowed to enter the Chinese agricultural system. We predict that Chinese farmers would value the prospect of increased yield with decreased use of pesticide and would readily adopt production of Bt rice. That Bt rice lines may not be commercialized in the near future we attribute to social pressures, largely due to the low level of understanding and acceptance of GM crops by Chinese consumers. Hence, enhancing communication of GM crop science‐related issues to the public is an important, unmet need. While the dynamics of each issue are particular to China, they typify those in many countries where adoption of GM crops has been not been rapid; hence, the assessment of these dynamics might inform resolution of these issues in other countries.     

Transgenic fertile japonica rice plants expressing a modified cryIA(b) gene resistant to yellow stem borer

The japonica rice variety Taipei 309 was cotransformed by particle bombardment of immature embryo-derived embryogenic calli with a modified δ-endotoxin gene cryIA(b) of Bacillus thuringiensis (Bt) under the control of the rice Actin1 promoter, and the hygromycin resistance gene, hph driven by the CaMV35S promoter. Selected transgenic rice plants showed enhanced insecticidal activity against yellow stem borer (Scirpophaga incertulas), with mortality rates reaching up to 100% in a bioassay with cut stems. Introduction and expression of the Actin1 promoter-Bt gene into rice provides japonica rice germplasm resistant to insect attack. Received: 21 March 1997 / Revision received: 23 June 1997 / Accepted: 5 July 1997