农作物抗除草剂遗传工程研究进展

控制杂草提高农作物产量是农业生产中共同面临的问题,发展抗除草剂农作物将是最经济最方便控制杂草的技术。由于对除草剂的作用模式和除草剂代谢途径的了解,弄清了除草剂的关键靶酶及其基因,因此分离除草剂靶酶基因,克隆能解毒除草剂的酶基因,通过转化技术可获得抗除草剂农作物,大量的抗除草剂转基因农作物大田试验表明,将最有希望在２０００年进入市场。     

Current trends in the global market of transgenic plants and environmental safety issues

The world market for the first generation of transgenic crops (insecticidal and herbicide-resistant plants) has been expanding since 2012, mostly owing to developing countries. The cautious attitude in the majority of economically developed countries to the first-generation transgenic agricultural crops is due to several objective circumstances: the negative impact of insecticidal Bt-crops on useful and endangered invertebrate species, the allergenic properties of Bt-toxin for humans, toxicity of glyphosate to humans and animals, the widely spreading resistance of weeds to glyphosate, the increasing resistance of–harmful–insects to insecticidal Bt-plants, the danger of–genetic pollution–of aboriginal plant varieties, and the flow of herbicide resistance traits to weed plants.     

Non-target-site herbicide resistance: a family business

We have witnessed a dramatic increase in the frequency and diversity of herbicide-resistant weed biotypes over the past two decades, which poses a threat to the sustainability of agriculture at both local and global levels. In addition, non-target-site mechanisms of herbicide resistance seem to be increasingly implicated. Non-target-site herbicide resistance normally involves the biochemical modification of the herbicide and/or the compartmentation of the herbicide (and its metabolites). In contrast to herbicide target site mutations, fewer non-target mechanisms have been elucidated at the molecular level because of the inherently complicated biochemical processes and the limited genomic information available for weedy species. To further understand the mechanisms of non-target-site resistance, we propose an integrated genomics approach to dissect systematically the functional genomics of four gene families in economically important weed species.     

Using empirical data to model transgene dispersal

One element of the current public debate about genetically modified crops is that gene flow from transgenic cultivars into surrounding weed populations will lead to more problematic weeds, particularly for traits such as herbicide resistance. Evolutionary biologists can inform this debate by providing accurate estimates of gene flow potential and subsequent ecological performance of resulting hybrids. We develop a model for gene flow incorporating exponential distance and directional effects to be applied to windpollinated species. This model is applied to previously published data on gene flow in experimental plots of Agrostis stolonifera L. (creeping bentgrass), which assessed gene flow from transgenic plants resistant to the herbicide glufosinate to surrounding non-transgenic plants. Our results show that although pollen dispersal can be limited in some sites, it may be extensive in others, depending on local conditions such as exposure to wind. Thus, hybridization under field conditions is likely to occur. Given the nature of the herbicide resistance trait, we regard this trait as unlikely to persist in the absence of herbicide, and suggest that the ecological consequences of such gene flow are likely to be minimal.     

Integrated weed management systems with herbicide-tolerant crops in the European Union: lessons learnt from home and abroad

Conventionally bred (CHT) and genetically modified herbicide-tolerant (GMHT) crops have changed weed management practices and made an important contribution to the global production of some commodity crops. However, a concern is that farm management practices associated with the cultivation of herbicide-tolerant (HT) crops further deplete farmland biodiversity and accelerate the evolution of herbicide-resistant (HR) weeds. Diversification in crop systems and weed management practices can enhance farmland biodiversity, and reduce the risk of weeds evolving herbicide resistance. Therefore, HT crops are most effective and sustainable as a component of an integrated weed management (IWM) system. IWM advocates the use of multiple effective strategies or tactics to manage weed populations in a manner that is economically and environmentally sound. In practice, however, the potential benefits of IWM with HT crops are seldom realized because a wide range of technical and socio-economic factors hamper the transition to IWM. Here, we discuss the major factors that limit the integration of HT crops and their associated farm management practices in IWM systems. Based on the experience gained in countries where CHT or GMHT crops are widely grown and the increased familiarity with their management, we propose five actions to facilitate the integration of HT crops in IWM systems within the European Union.     

Exotic wildland weeds serve as reservoirs for a newly introduced cole crop pest,Bagrada hilaris (Hemiptera: Pentatomidae)

The role of wildland weeds that serve as alternate hosts for insect pests has not been directly examined for the potential to sustain pest populations or facilitate pest outbreaks. The development of weed biological control programmes is also complicated by weed species that are closely related to economically important or native plants, especially rare or special status species. In recent surveys in southern California, USA, we found a newly introduced insect pest of cole crops, Bagrada hilaris Burmeister (Bagrada bug), feeding on Brassicaceae weeds in riparian areas adjacent to agricultural fields where cole crops are routinely grown. Insect populations grew to levels well over action thresholds and caused severe damage to populations of the invasive weed, Lepidium latifolium (perennial pepperweed). The numerical response of B. hilaris on L. latifolium and other Brassicaceae weeds in natural areas may pose a significant challenge to effectively managing pest populations in crops. However, the accidental introduction of this insect provides the opportunity to examine plant–insect interactions with important implications for development of biologically based control methods for weeds.     

An individual‐based model of seed‐ and rhizome‐propagated perennial plant species and sustainable management of Sorghum halepense in soybean production systems in Argentina

Perennial plants which propagate through both seeds and rhizomes are common in agricultural and nonagricultural systems. Due to their multifaceted life cycle, few population models are available for studying such species. We constructed a novel individual‐based model to examine the effects of ecological, evolutionary, and anthropogenic factors on the population dynamics of perennial species. To exemplify the application of the model, we presented a case study of an important weed, Sorghum halepense (L.) Pers. (Johnsongrass), in soybean productions in Argentina. The model encompasses a full perennial weed life cycle with both sexual (seeds) and asexual (rhizomes) propagations. The evolution of herbicide resistance was modeled based on either single genes or quantitative effects. Field experiments were conducted in the species' native environment in Argentina to parameterize the model. Simulation results showed that resistance conferred by single‐gene mutations was predominantly affected by the initial frequency of resistance alleles and the associated fitness cost. Population dynamics were influenced by evolved resistance, soil tillage, and rhizome fecundity. Despite the pivotal role of rhizomes in driving the population dynamics of Johnsongrass, most herbicides target the aboveground biomass, and chemical solutions to control rhizomes are still very limited. To maintain effective (short‐term) and sustainable (long‐term) weed management, it is recommended to combine soil tillage with herbicide applications for suppressing the rhizomes and delaying the evolution of resistance. This novel model of seed‐ and rhizome‐propagated plants will also be a useful tool for studying the evolutionary processes of other perennial weeds, cash crops, and invasive species.     

Can the growing of transgenic maize threaten protected Lepidoptera in Europe?

We evaluated whether protected European butterflies can potentially be at risk if transgenic maize is extensively grown in Central Europe. We explored potential consequences of both insect resistant (IR) and herbicide resistant (HR) transgenic maize. IR maize can produce pollen that is toxic to lepidopteran larvae, and this puts butterfly species at possible risk if the presence of young larvae coincides with maize flowering, during which large quantities of maize pollen can be deposited on vegetation. By considering the timing of maize flowering in Europe and the phenology of the protected Lepidoptera species, we found that 31 species had at least one generation where 50% of the larval stage overlapped with maize flowering, and 69 species for which first instar larvae were present during maize pollen shedding. HR maize allows high concentration herbicide treatments on fields without seasonal limitation, which can drastically reduce weed densities. In cases where such weed species are host plants for protected butterflies, reduced host plant/food availability can result, causing population decreases. By using published information, we first identified the important weed species in major maize-growing European countries. Subsequently, we checked whether the host plants of protected Lepidoptera included species that are common maize weeds. We identified 140 protected species having food plants that are common weeds in one or more of the major European maize-growing countries. If HR maize is grown in Europe, there is a potential hazard that their food plants will seriously decline, causing a subsequent decline of these protected species.     

Herbicide resistance-endowing ACCase gene mutations in hexaploid wild oat (Avena fatua): insights into resistance evolution in a hexaploid species

Many herbicide-resistant weed species are polyploids, but far too little about the evolution of resistance mutations in polyploids is understood. Hexaploid wild oat (Avena fatua) is a global crop weed and many populations have evolved herbicide resistance. We studied plastidic acetyl-coenzyme A carboxylase (ACCase)-inhibiting herbicide resistance in hexaploid wild oat and revealed that resistant individuals can express one, two or three different plastidic ACCase gene resistance mutations (Ile-1781-Leu, Asp-2078-Gly and Cys-2088-Arg). Using ACCase resistance mutations as molecular markers, combined with genetic, molecular and biochemical approaches, we found in individual resistant wild-oat plants that (1) up to three unlinked ACCase gene loci assort independently following Mendelian laws for disomic inheritance, (2) all three of these homoeologous ACCase genes were transcribed, with each able to carry its own mutation and (3) in a hexaploid background, each individual ACCase resistance mutation confers relatively low-level herbicide resistance, in contrast to high-level resistance conferred by the same mutations in unrelated diploid weed species of the Poaceae (grass) family. Low resistance conferred by individual ACCase resistance mutations is likely due to a dilution effect by susceptible ACCase expressed by homoeologs in hexaploid wild oat and/or differential expression of homoeologous ACCase gene copies. Thus, polyploidy in hexaploid wild oat may slow resistance evolution. Evidence of coexisting non-target-site resistance mechanisms among wild-oat populations was also revealed. In all, these results demonstrate that herbicide resistance and its evolution can be more complex in hexaploid wild oat than in unrelated diploid grass weeds. Our data provide a starting point for the daunting task of understanding resistance evolution in polyploids.     

Drought exposure leads to rapid acquisition and inheritance of herbicide resistance in the weed Alopecurus myosuroides

Globally, herbicide resistance in weeds poses a threat to food security. Resistance evolves rapidly through the co‐option of a suite of physiological mechanisms that evolved to allow plants to survive environmental stress. Consequently, we hypothesize that stress tolerance and herbicide resistance are functionally linked. We address two questions: (i) does exposure to stress in a parental generation promote the evolution of resistance in the offspring? (ii) Is such evolution mediated through non‐genetic mechanisms? We exposed individuals of a grass weed to drought, and tested whether this resulted in herbicide resistance in the first generation. In terms of both survival and dry mass, we find enhanced resistance to herbicide in the offspring of parents that had been exposed to drought. Our results suggest that exposure of weeds to drought can confer herbicide resistance in subsequent generations, and that the mechanism conferring heritability of herbicide resistance is non‐genetic.     

Herbicidal potential of catechol as an allelochemical

Catechol is an allelochemical which belongs to phenolic compounds synthesized in plants. Its herbicidal effects on weed species; field poppy (Papaver rhoeas), creeping thistle (Cirsium arvense), henbit (Lamium amplexicaule) and wild mustard (Sinapis arvensis) were investigated using wheat (Triticum vulgare) and barley (Hordeum vulgare) species as control plants. In comparison to 2,4-D (a common synthetic herbicide), 13.64 mM of catechol have been found to have a strong herbicidal effect, as effective as 2,4-D on field poppy weed by killing it, and a suppressive herbicidal effect on the other weeds by inhibiting their growth significantly. Concerning all the weeds, in general, elongation of the shoot was affected more negatively than that of the root. Fresh weights of the weeds were decreased by catechol significantly only in field poppy but not in other weeds. The study reveals that catechol is a potent inhibitor of growth of the weeds and therefore it can be evaluated as a herbicide for future weed management strategies.     

Herbicide-resistance conferred by expression of a catalytic antibody in Arabidopsis thaliana

Engineering herbicide resistance in crops facilitates control of weed species, particularly those that are closely related to the crop, and may be useful in selecting lines that have undergone multiple transformation events. Here we show that herbicide-resistant plants can be engineered by designing an herbicide and expressing a catalytic antibody that destroys the herbicide in planta. First, we developed a carbamate herbicide that can be catalytically destroyed by the aldolase antibody 38C2. This compound has herbicidal activity on all three plant species tested. Second, the light chain and half of the heavy chain (Fab) of the catalytic antibody were targeted to the endoplasmic reticulum in two classes of Arabidopsis thaliana transformants. Third, the two transgenic plants were crossed to produce an herbicide-resistant F1 hybrid. The in vitro catalytic activity of the protein from F1 hybrids corroborates that catalytic antibodies can be constitutively expressed in transgenic plants, and that they can confer a unique trait.     

A test of the Australian Weed Risk Assessment system in China

The Australian Weed Risk Assessment system (AWRA) is an effective pre-border weed-screening tool that has played an active role in preventing the introduction of alien weeds into Australia and has been utilized in several other countries worldwide. Here, we selected 131 species of naturalized exotic plants (including 76 species of given non-weeds and 55 species of given weeds) to evaluate the AWRA in China for the first time. The AWRA performed better for discriminating major weeds than non-weeds and minor weeds, as it correctly rejected 84% of major weeds and did not wrongly accept a major weed. Among non-weeds, 76% were correctly classified with the final outcome of “accept” and 7.9% were wrongly rejected by the AWRA. This system correctly rejected 56% of minor weeds but accepted only 2.8% of minor weeds. The remaining 23% of all alien plants tested were classified as “evaluate further” by the AWRA. The area under the ROC curve (AUC) was 0.944, suggesting that the AWRA would be highly efficient at discriminating alien plants in China. In addition, we compared the scores of seven attributes of the AWRA between prior plant categories and analyzed their correlation with weed status. The average score for each attribute differed significantly between the two prior categories (weed and non-weed), but the average scores of the attribute “undesirable traits” did not significantly differ between any two of the three categories (non-weeds, minor weeds, and major weeds). There was a significant positive correlation between the scores of each attribute of the AWRA and weed status. The correlation coefficient for “dispersal mechanisms” and weed status was the highest and that for “undesirable traits” was the lowest. We believe that the AWRA can serve as an important weed-screening tool for plant introduction management in China.     

A plant‐traits approach to assessing the success of alien weed species in irrigated Mediterranean orchards

In addition to their impact on natural habitats, invasive alien plants can have a significant negative effect on agricultural systems and cause economic losses. Flood‐irrigated orchards in the Mediterranean Basin are vulnerable to the invasion of alien weeds, primarily because of the traditional management practices used in the orchards, which are characterized by high soil moisture during the dry summer period, nutrient availability and high levels of disturbance. This study sought to determine whether their biological traits can explain the success of alien weed species. To answer this question, 408 floristic relevés were conducted in 136 flood‐irrigated orchards on the Plains of Lleida (Catalonia, NE of Spain). Richness and cover of native and alien weeds were compared. Furthermore, a set of biological traits were compared between successful and non‐successful weeds for the whole data and separately between native and alien weeds using logistic regression and classification trees. In flood‐irrigated orchards, alien species covered most of their area, even though the richness of alien species was lower than that of the native species. The most important species were C4 species with seeds dispersed by water, and on the other hand, rosulate and caespitose‐reptant hemicryptophytes with long flowering period. Most of these traits fitted with those of the invasive alien weeds, which were mostly C4 species with seeds dispersed by water. Perennial life form characterized successful native weeds. In this study, we discuss how the traditional management of flood irrigation in fruit‐tree orchards favours invasive alien weeds that have specific traits, acting as a reservoir for the spread of alien weeds into other crops and surrounding riparian habitats. We also propose changing management practices in order to avoid the selection of alien weeds and to promote native species.     

Italian weedy rice—A case of de‐domestication?

Weedy rice is a representative of the extensive group of feral weeds that derive from crops, but has returned to the lifestyle of a wild species. These weeds develop either from a hybridization of crops with wild relatives (exoferality), or by mutation of crops to weedy forms (endoferality). Due to the close relation of weed and crop, the methods for weed‐targeted containment are limited to date. A deeper understanding of the development of such weeds might help to design more efficient and sustainable approaches for weed management. Weedy rice poses a serious threat to rice yields worldwide. It is widely accepted that weedy rice has originated independently in different regions all over the world. However, details of its evolution have remained elusive. In the current study, we investigated the history of weedy rice in northern Italy, the most important rice‐growing area in Europe. Our approach was to analyze genes related to weedy traits (SD1, sh4, Rc) in weedy rice accessions compared to cultivars, and to integrate these results with phenotypic and physiological data, as well as historical information about rice farming in Italy. We arrive at a working model for the timeline of evolution of weedy rice in Italy indicating that both exoferality and endoferality acted as forces driving the development of the diverse weedy rice populations found in the region today. Models of weed evolution can help to predict the direction which weed development might take and to develop new, sustainable methods to control feral weeds.     

Terbuthylazine interferes with iron nutrition in maize (<Emphasis Type="Italic">Zea mays</Emphasis>) plants

Many chemicals, including herbicides, are routinely applied to crops for weed management and food production improvement. However, the intensive use of herbicides could lead eventually to a great environmental threat due to their persistence and accumulation in the ecosystems and contamination of soils. Furthermore, the possible effect of these chemicals on nutrient uptake and assimilation in crops has only recently been discussed. The present study aimed at understanding the effect of the herbicide terbuthylazine (TBA), a herbicide commonly applied to control weeds in leguminous species and triazine tolerant crops, on the capability of maize plants to cope with iron (Fe) shortage. The application of 2 and 5 mg L^?1 TBA caused a significant reduction of root Fe concentration. This reduction might be attributed to a decreased release of phytosiderophores, which in turn could be ascribed to a reduced sulfur assimilation. Results provide evidence that TBA impairs Fe uptake and accumulation in non-target plants most likely interacting with sulfur-assimilating enzymes [ATP sulfurylase and O-acetylserine(thiol)lyase].     

Atmospheric pollen dispersion from herbicide-resistant horseweed (<Emphasis Type="Italic">Conyza canadensis</Emphasis> L.)

Horseweed (Conyza canadensis (L.) Cronq.) with evolved herbicide resistance has become an especially problematic weed in crop production across the USA and on four continents (North America, South America, Asia, and Europe). Spread of herbicide resistance can occur through pollen-mediated gene flow between resistant and susceptible horseweed populations. However, there are little knowledge, preventive guidelines, and mechanism modeling for pollen transport in this system. We need to better understand pollen dispersion and deposition in the context of atmospheric conditions, herbicide-resistant horseweed patch size, and buffer crop type, height, and field size. A mechanistic model is needed to account for these. A pollen dispersion and deposition model was calibrated and validated using 2013 experimental field data. The validated model was run for various combinations of atmospheric conditions, horseweed characteristics (source strength), and buffer species and size (pollen can be intercepted by crop plants). Large fields with crops with a high leaf area density and tall plants can effectively prevent pollen dispersion. The information will help provide guidelines for preventing herbicide resistance spread from herbicide-resistant weeds and genetically modified plants in general.     

Assessing broomrape risk due to weeds in cropping systems with an indicator linked to a simulation model

Integrated crop protection tolerates residual weed floras if they are not harmful for crop production. These weeds can host harmful crop pests, among which parasitic plants such as branched broomrape (Phelipanche ramosa). This holoparasite is responsible for large yield losses in French crops such as oilseed rape. To date, there are no herbicides available to control it. To evaluate ex ante the impact of crop management practices on weed-mediated parasite infection of crops, we developed an indicator calculated from outputs of the weed dynamics model FlorSys. It consists of three components assessing weed impact on (1) stimulation of parasite germination during the whole cropping season, i.e. the potential risk reduction for future crops via a reduction of the parasite seed bank, (2) the stimulation of parasite germination in host crops, i.e. the potential risk increase for the current crop, (3) parasite reproduction on weed plants, i.e. the potential risk increase for future crops. This indicator was then used to predict weed-mediated broomrape risk in cropping systems from six regions from France and one from Spain. Antagonisms and synergies with other indicators of weed-harmfulness for crop production and weed contribution to plant and functional biodiversity were investigated with Pearson correlation analyses. For instance, cropping systems with a high parasite risk also had a high functional biodiversity (e.g. weed-based food offer for bees). Effects of crop management practices on the weed-mediated parasite risk indicator were identified with linear models; regression trees were used to identify the combinations of management practices that maximised or minimised weed-mediated broomrape risk. Parasite risk depended on crop rotation, sowing and harvest dates, tillage, herbicides and mechanical weeding. The lowest risk was observed in fields that were last tilled less than 21 days before sowing, with more than 0.6 herbicides per year (i.e. 3 applications in 5 years) with multiple entry modes into the weeds (e.g. leaves and roots) and the last herbicide sprayed no later than 127 days before harvest. RLQ analyses were used to identify correlations between weed species traits (Q matrix) and simulated parasite risk (R matrix), via simulated weed densities (L matrix). Early summer-emerging weed species increased parasite risk. No other notable correlations were found, indicating that parasite risk results from a weed community of interacting species, and not simply from individual weed species. An advice table was built to summarize and explain the effects of crop management practices on weed-mediated parasite risk.     

Weed and invertebrate community compositions in arable farmland

A large proportion of the land surface area of Great Britain (GB) is used for arable agriculture. Due to changes in farm management practices over the last 50 years, there have been marked declines in the abundance of arable wildlife groups of conservation importance, including weeds, invertebrates and birds. Here we ask whether changes in weed species composition, driven by changes in management, such as a change in crop or a modification in herbicide regime, might be expected to lead to changes in the species compositions of other wildlife groups, including invertebrates. Using multivariate analyses, on data from eight crop and herbicide management groups sampled across 266 arable fields, we show that the weed composition changes with the crop and herbicide management adopted and the invertebrate composition changes with the crop grown. We conclude that each conventional crop sampled had a unique composition of weeds and invertebrates, and expect this to be true for all conventional arable crops. Changes in weed species composition, driven by changes in crop or herbicide management, will lead to changes in the compositions of invertebrates, and possibly other wildlife groups. However, these changes will probably be buffered by the effect of functional redundancy, the crop and dispersal. Handling editor: Gimme Walter.