Non‐target site resistance to flucarbazone,imazamethabenz and pinoxaden is controlled by three linked genes in Avena fatua

Extensive herbicide usage has led to the evolution of resistant weed populations that cause substantial crop yield losses and increase production costs. The multiple herbicide‐resistant (MHR) Avena fatua populations utilised in this study are resistant to members of all selective herbicide families, across five modes of action, available for A. fatua control in US small grain production, and thus pose significant agronomic and economic threats. Resistance to acetolactate synthase and acetyl‐CoA carboxylase inhibitors is not conferred by known target site mutations, indicating that non‐target site resistance (NTSR) mechanisms are involved. Understanding the inheritance of NTS MHR is of upmost importance for continued agricultural productivity in the face of the rapid increase in resistant weed populations worldwide. As few studies have examined the inheritance of NTSR in autogamous weeds, we investigated the inheritance and genetic control of NTSR in the highly autogamous, allohexaploid species A. fatua. We found that NTSR in MHRA. fatua is controlled by three separate, closely‐linked nuclear genes for flucarbazone‐sodium, imazamethabenz‐methyl and pinoxaden. The single‐gene NTSR inheritance patterns reported here contrast with other examples in allogamous species and illustrate the diversity of evolutionary responses to strong selection.     

抗药性杂草与治理

抗药性杂草对农田杂草治理和农业生产构成严重威胁,成为备受全球关注的严重问题。随着长期、大量使用相对有限的化学除草剂,全球抗药性杂草发展迅猛,目前已有217种杂草对化学除草剂产生了抗药性,我国抗药性杂草发展也十分迅猛。本文在介绍杂草抗药性基本概念、抗药性杂草发展过程、抗药性杂草现状的基础上,重点描述了抗药性杂草治理策略,以期为我国抗药性杂草研究和治理提供参考。     

Developmental changes in glutathione S‐transferase activity in herbicide‐resistant populations of Alopecurus myosuroides Huds (black‐grass) in the field

Herbicide‐resistant populations of Alopecurus myosuroides Huds (black‐grass) have become widespread throughout the UK since the early 1980s. Clear evidence suggests that more than one resistance mechanism exists, and glutathione S‐transferases (GSTs) have been implicated in resistance due to enhanced metabolism. This study reports the determination of GST activity in four UK black‐grass populations from field sites situated in the East Midlands. Data demonstrate that, as untreated plants in the field mature, there is an accompanying natural elevation of GST activity with natural environmental changes from winter to spring. We speculate that this endogenous change in enzyme activity with plant development in the field contributes to reduced efficacy of some graminicides applied in the spring. These observations are discussed in relation to predicting herbicide efficacy to achieve maximum control of this important grass weed. © 2001 Society of Chemical Industry     

Simulating evolution of glyphosate resistance in Lolium rigidum I: population biology of a rare resistance trait

Despite frequent use for the past 25 years, resistance to glyphosate has evolved in few weed biotypes. The propensity for evolution of resistance is not the same for all herbicides, and glyphosate has a relatively low resistance risk. The reasons for these differences are not entirely understood. A previously published two‐herbicide resistance model has been modified to explore biological and management factors that account for observed rates of evolution of glyphosate resistance. Resistance to a post‐emergence herbicide was predicted to evolve more rapidly than it did to glyphosate, even when both were applied every year and had the same control efficacy. Glyphosate is applied earlier in the growing season when fewer weeds have emerged and hence exerts less selection pressure on populations. The evolution of glyphosate resistance was predicted to arise more rapidly when glyphosate applications were later in the growing season. In simulations that assumed resistance to the post‐emergence herbicide did not evolve, the evolution of glyphosate resistance was less rapid, because post‐emergence herbicides were effectively controlling rare glyphosate‐resistant individuals. On their own, these management‐related factors could not entirely account for rates of evolution of resistance to glyphosate observed in the field. In subsequent analyses, population genetic parameter values (initial allele frequency, dominance and fitness) were selected on the basis of empirical data from a glyphosate‐resistant Lolium rigidum population. Predicted rates of evolution of resistance were similar to those observed in the field. Together, the timing of glyphosate applications, the rarity of glyphosate‐resistant mutants, the incomplete dominance of glyphosate‐resistant alleles and pleiotropic fitness costs associated with glyphosate resistance, all contribute to its relatively slow evolution in the field.     

Establishment of Lolium species resistant to acetolactate synthase‐inhibiting herbicide in and around grain‐importation ports in Japan

Compared with natural seed dispersal, human‐mediated seed dispersal could spread herbicide resistance genes on a much larger scale. Herbicide‐resistant weed seeds have been reported as contaminants in commercial grain. We investigated the contamination of seeds of Lolium species with target‐site mutations conferring resistance to acetolactate synthase (ALS)‐inhibiting herbicides in wheat imported from the USA, Canada and Australia into Japan. We also investigated the establishment of ALS‐inhibiting herbicide‐resistant Lolium species in 12 seaports in Japan that are major entry points for international commodities. We found herbicide‐resistant Lolium spp. seeds from all classes of wheat samples. Resistant individuals became established at six of eight ports where more than 50 kt of imported wheat is unloaded every year. The establishment of resistant Lolium spp. individuals was common at major grain landing ports. Monitoring over 3 years at one port revealed that the frequency of resistant individuals did not fluctuate between years. Many resistant individuals were distributed in front of the entrance of a fodder company, but a few resistant individuals were found in areas 2 km away from the port. The results indicate that gene flow is rare through pollen or seed movement from resistant plants to peripheral populations. Further extensive and long‐term monitoring is necessary to perform a comprehensive risk assessment of herbicide‐resistant plants entering Japan through major commercial ports.     

Frequency and distribution of herbicide resistance in Raphanus raphanistrum populations randomly collected across the Western Australian wheatbelt

In 2003, a random survey was conducted across the Western Australian wheatbelt to establish the extent and frequency of herbicide resistance in Raphanus raphanistrum populations infesting crop fields. Five hundred cropping fields were visited, with 90 R. raphanistrum populations collected, representative of populations present in crop fields throughout the Western Australian wheatbelt. Collected populations were screened with four herbicides of various modes of action that are commonly used for the control of this weed. The majority of Western Australian R. raphanistrum populations were found to contain plants resistant to the acetolactate synthase (ALS)‐inhibiting herbicide chlorsulfuron (54%) and auxin analogue herbicide, 2,4‐D amine (60%). This survey also determined that over half (58%) of these populations were multiple resistant across at least two of the four herbicide modes of action used in the screening. Only 17% of R. raphanistrum populations have retained their initial status of susceptibility to all four herbicides. The distribution patterns of the herbicide‐resistant populations identified that there were higher frequencies of resistant and developing resistance populations occurring in the intensively cropped northern regions of the wheatbelt. These results clearly indicate that the reliance on herbicidal weed control in cropping systems based on reduced tillage and stubble retention will lead to higher frequencies of herbicide‐resistant weed populations. Therefore, within intensive crop production systems, there is a need to diversify weed management strategies and not rely entirely on too few herbicide control options.     

抗性杂草的鉴定与治理

杂草抗性的日益严重已成为化学药剂除草的一大障碍。本文综述了最新杂草抗性的发展，抗性机理，抗性杂草的识别、鉴定及评价等，分析其发生的原因并探讨治理措施，试图为解决杂草抗性提供借鉴。     

Prevalence of herbicide‐resistant weed species in Malaysian rice fields: A review

The management of weeds in Malaysian rice fields is very much herbicide‐based. The heavy reliance on herbicide for weed control by many rice‐growers arguably eventually has led to the development and evolution of herbicide‐resistant biotypes in Malaysian rice fields over the years. The continuous use of synthetic auxin (phenoxy group) herbicides and acetohydroxyacid synthase‐inhibiting herbicides to control rice weeds was consequential in leading to the emergence and prevalence of resistant weed biotypes. This review discusses the history and confirmed cases and incidence of herbicide‐resistant weeds in Malaysian rice fields. It also reviews the Clearfield Production System and its impact on the evolution of herbicide resistance among rice weed species and biotypes. This review also emphasizes the strategies and management options for herbicide‐resistant rice field weeds within the framework of herbicide‐based integrated weed management. These include the use of optimum tillage practices, certified clean seeds, increased crop competition through high seeding rates, crop rotation, the application of multiple modes of action of herbicides in annual rotations, tank mixtures and sequential applications to enable a broad spectrum of weed control, increase the selective control of noxious weed species in a field and help to delay the resistance evolution by reducing the selection pressure that is forced on those weed populations by a specific herbicidal mode of action.     

The effect of temperature on glutathione S-transferase activity and glutathione content in Alopecurus myosuroides (black grass) biotypes susceptible and resistant to herbicides

Herbicide‐resistant populations of Alopecurus myosuroides (black grass) have become widespread throughout the UK since the early 1980s. Previous observations in this laboratory have demonstrated that natural climatic fluctuations caused increases in endogenous glutathione S‐transferase (GST) enzyme activity in A. myosuroides plants as they mature, which is thought to be linked to herbicide resistance in this species. The present study has investigated the effects of plant growth at 10°C and 25°C, and reports GST specific activity and glutathione (GSH) pool size in resistant and susceptible A. myosuroides biotypes. Findings demonstrate differences in GST activity between resistant and susceptible populations, which are transient at lower growth temperatures. The GSH pool size was elevated at lower growth temperature in both biotypes. We speculate that these endogenous responses are part of a natural mechanism of acclimation to environmental change in this species and suggest that resistant plants are more able to adapt to environmental stress, as indicated in this instance by temperature change. These observations imply that the control of resistant A. myosuroides by graminicides may be more effective when applied at lower temperatures and at earlier growth stages.     

Relationship between weed dormancy and herbicide rotations: implications in resistance evolution

It is suggested that selection for late germinating seed cohorts is significantly associated with herbicide resistance in some cropping systems. In turn, it is conceivable that rotating herbicide modes of action selects for populations with mutations for increased secondary dormancy, thus partially overcoming the delaying effect of rotation on resistance evolution. Modified seed dormancy could affect management strategies – like herbicide rotation – that are used to prevent or control herbicide resistance. Here, we review the literature for data on seed dormancy and germination dynamics of herbicide‐resistant versus susceptible plants. Few studies use plant material with similar genetic backgrounds, so there are few really comparative data. Increased dormancy and delayed germination may co‐occur with resistance to ACCase inhibitors, but there is no clear‐cut link with resistance to other herbicide classes. Population shifts are due in part to pleiotropic effects of the resistance genes, but interaction with the cropping system is also possible. We provide an example of a model simulation that accounts for genetic diversity in the dormancy trait, and subsequent consequences for various cropping systems. We strongly recommend adding more accurate and detailed mechanistic modelling to the current tools used today to predict the efficiency of prevention and management of herbicide resistance. These models should be validated through long‐term experimental designs including mono‐herbicide versus chemical rotation in the field. © 2017 Society of Chemical Industry     

Control by glyphosate and its alternatives of glyphosate‐susceptible and glyphosate‐resistant Echinochloa colona in the fallow phase of crop rotations in subtropical Australia

Echinochloa colona is the most common grass weed of summer fallows in the grain‐cropping systems of the subtropical region of Australia. Glyphosate is the most commonly used herbicide for summer grass control in fallows in this region. The world's first population of glyphosate‐resistant E. colona was confirmed in Australia in 2007 and, since then, >70 populations have been confirmed to be resistant in the subtropical region. The efficacy of alternative herbicides on glyphosate‐susceptible populations was evaluated in three field experiments and on both glyphosate‐susceptible and glyphosate‐resistant populations in two pot experiments. The treatments were knockdown and pre‐emergence herbicides that were applied as a single application (alone or in a mixture) or as part of a sequential application to weeds at different growth stages. Glyphosate at 720 g ai ha^?1 provided good control of small glyphosate‐susceptible plants (pre‐ to early tillering), but was not always effective on larger susceptible plants. Paraquat was effective and the most reliable when applied at 500 g ai ha^?1 on small plants, irrespective of the glyphosate resistance status. The sequential application of glyphosate followed by paraquat provided 96–100% control across all experiments, irrespective of the growth stage, and the addition of metolachlor and metolachlor + atrazine to glyphosate or paraquat significantly reduced subsequent emergence. Herbicide treatments have been identified that provide excellent control of small E. colona plants, irrespective of their glyphosate resistance status. These tactics of knockdown herbicides, sequential applications and pre‐emergence herbicides should be incorporated into an integrated weed management strategy in order to greatly improve E. colona control, reduce seed production by the sprayed survivors and to minimize the risk of the further development of glyphosate resistance.     

Multiple herbicide‐resistant Lolium rigidum (annual ryegrass) now dominates across the Western Australian grain belt

Lolium rigidum (annual or rigid ryegrass) is a widespread annual weed in cropping systems of southern Australia, and herbicide resistance in L. rigidum is a common problem in this region. In 2010, a random survey was conducted across the grain belt of Western Australia to determine the frequency of herbicide‐resistant L. rigidum populations and to compare this with the results of previous surveys in 1998 and 2003. During the survey, 466 cropping fields were visited, with a total of 362 L. rigidum populations collected. Screening of these populations with the herbicides commonly used for control of L. rigidum revealed that resistance to the ACCase‐ and ALS‐inhibiting herbicides was common, with 96% of populations having plants resistant to the ACCase herbicide diclofop‐methyl and 98% having plants resistant to the ALS herbicide sulfometuron. Resistance to another ACCase herbicide, clethodim, is increasing, with 65% of populations now containing resistant plants. Resistance to other herbicide modes of action was significantly lower, with 27% of populations containing plants with resistance to the pre‐emergent herbicide trifluralin, and glyphosate, atrazine and paraquat providing good control of most of the populations screened in this survey. Ninety five per cent of L. rigidum populations contained plants with resistance to at least two herbicide modes of action. These results demonstrate that resistance levels have increased dramatically for the ACCase‐ and ALS‐inhibiting herbicides since the last survey in 2003 (>95% vs. 70–90%); therefore, the use of a wide range of integrated weed management options are required to sustain these cropping systems in the future.     

Modelling binary mixtures of herbicides in populations resistant to one of the components: evaluation for resistance management

BACKGROUND: Herbicide mixtures are commonly proposed to delay the selection of herbicide resistance in susceptible populations (called the SM strategy). However, in practice, herbicide mixtures are often used when resistance to one of the two active ingredients has already been detected in the targeted population (called the RM strategy). It is doubtful whether such a practice can select against resistance, as the corresponding selection pressure is still exerted. As a consequence, the effect of mixtures on the evolution of an already detected resistance to one of the herbicides in the combination remains largely unexplored. In the present work, a simple model was developed to explore further the necessary and sufficient conditions under which a binary RM strategy might stabilise or even reduce resistance frequency. RESULTS: Covering the hypothetical largest range of parameters, 39% of 9000 random simulations attest that the RM strategy might theoretically reduce resistance frequency. When strong enough, high genetic cost of resistance, negative cross‐resistance between the herbicides associated in the mixture and reduced selection differential between resistant and susceptible plants can counterbalance the resistance advantage to one of the two applied herbicides. However, the required conditions for an RM strategy to ensure resistance containment in natural conditions seldom overlap with experimental parameter estimates given in the literature. CONCLUSION: It is concluded that the sufficient conditions for an RM strategy to be effective would rarely be encountered. As a consequence, the strategy of formulating mixtures with herbicides for which resistance has already been detected should be avoided. Copyright © 2008 Society of Chemical Industry     

Chlorophyll fluorescence imaging: a new method for rapid detection of herbicide resistance in Alopecurus myosuroides

Due to the steadily increasing number of putative herbicide‐resistant weed populations, the demand for rapid in‐season tests is rising. In this study, we introduce a new quantitative herbicide‐resistance test system based on chlorophyll fluorescence imaging analysis of photosynthesis‐related parameters. Susceptible and herbicide‐resistant populations of Alopecurus myosuroides (black‐grass) were cultivated in multiwell tissue culture plates containing nutrient agar and different dosages of fenoxaprop‐P‐ethyl and mesosulfuron+iodosulfuron. The maximum quantum efficiency of the PSII was measured 3 h after transplanting (HAT) and then for seven days every 24 h. Data of maximum quantum efficiency of the PSII were compared with standard whole‐plant pot tests and molecular tests for target‐site mutations. It was possible to fit dose‐response curves and calculate corresponding resistance factors for ED90 for all populations tested using the chlorophyll fluorescence imaging. It was possible to distinguish between resistant and susceptible populations. The results of the chlorophyll fluorescence imaging corresponded well with the standard whole‐plant pot tests in the glasshouse. However, populations with proved target‐site mutations did not differ from other herbicide‐resistant populations in the maximum quantum efficiency values of the PSII. We conclude that the chlorophyll fluorescence imaging provides reliable data on herbicide resistance for both modes of action tested in a shorter time and using less space, compared with standard whole‐plant pot tests in the glasshouse.     

Herbicide‐resistant genetically‐modified crop: its risks with an emphasis on gene flow

Herbicide‐resistant genetically‐modified (GM) crops are the most widely cultivated worldwide, representing 78% of GM crops in 1999, followed by insect‐resistant GM crops with Bt gene. Gene flow is the most touching risk arising from GM crops, and is categorized as three types: within species, between species and between GM crop and other organisms. This review shows that gene flow is a reality in the plant kingdom with evolutionary change. Herbicide resistance evolves naturally and spreads dynamically in weeds. One of the most concerning crop in relation to gene flow is Brassica napus, which has a high outcrossing rate and many relative species. In contrast, frequency of gene flow via outcrossing is relatively low in inbreeding cereal crops such as rice, wheat and barley, but published reports have shown that substantial gene flow is possible. Another possible and immediate risk is herbicide‐resistant GM crops becoming volunteer weeds. Dry direct‐seeded rice is one of the most likely crops in this respect. Stacking different resistance genes in a crop would accelerate multiple resistance evolution in weeds. Multiple resistance to three major herbicides has already been observed in oilseed rape cultivation. More efforts must be made for long‐term risk assessment on GM crops in the natural ecosystem. More studies on weed biology and ecology, particularly reproductive processes in weeds, are essential for better understanding of gene flow and systematic management strategy. We hope that this review motivates researchers to analyze data available now, to collect fundamental information on crops and weeds in agro‐ecosystem, and to lead to better risk assessment and management.     

Potential roles for microbial endophytes in herbicide tolerance in plants

Herbicide tolerance in crops and weeds is considered to be monotrophic, i.e. determined by the relative susceptibility of the physiological process targeted and the plant's ability to metabolise and detoxify the agrochemical. A growing body of evidence now suggests that endophytes, microbes that inhabit plant tissues and provide a range of growth, health and defence enhancements, can contribute to other types of abiotic and biotic stress tolerance. The current evidence for herbicide tolerance being bitrophic, with both free‐living and plant‐associated endophytes contributing to tolerance in the host plant, has been reviewed. We propose that endophytes can directly contribute to herbicide detoxification through their ability to metabolise xenobiotics. In addition, we explore the paradigm that microbes can ‘prime’ resistance mechanisms in plants such that they enhance herbicide tolerance by inducing the host's stress responses to withstand the downstream toxicity caused by herbicides. This latter mechanism has the potential to contribute to the growth of non‐target‐site‐based herbicide resistance in weeds. Microbial endophytes already contribute to herbicide detoxification in planta, and there is now significant scope to extend these interactions using synthetic biology approaches to engineer new chemical tolerance traits into crops via microbial engineering. © 2015 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.     

Differential translocation of paraquat in paraquat-resistant populations of Hordeum leporinum

Two populations of Hordeum leporinum have evolved resistance to paraquat within a small area in central Tasmania, Australia. One population (THL1) was more than 80-fold resistant to paraquat when treated in winter, compared with a susceptible population (THL4) collected nearby, whereas the other population (THL2) was only 19-fold resistant. Translocation of paraquat was examined in all three populations at warm and cool temperature regimes. Herbicide was applied to a basal section of the second leaf of plants kept in the dark and translocation measured after 16 h of dark and during a subsequent light period. Paraquat absorption into the treated leaf was uniformly high in susceptible and resistant populations, with >93% of the applied herbicide absorbed within 16 h in the dark at both temperatures. Translocation of paraquat out of the treated leaf was low in the dark, with <4% of the herbicide translocated to the remainder of the plant. More herbicide was translocated out of the treated leaves in susceptible plants in the dark, compared with resistant plants at both temperature regimes and more paraquat was translocated at warmer temperatures. Extensive basipetal translocation of paraquat to the rest of the plant occurred in susceptible plants following exposure of the treated plants to light. However, basipetal translocation was much reduced in resistant plants in the light and corresponded to the degree of resistance. Resistance to paraquat in H. leporinum is the result of reduced translocation of paraquat out of the treated leaves.     

An aspartate to glycine change in the carboxyl transferase domain of acetyl CoA carboxylase and non‐target‐site mechanism(s) confer resistance to ACCase inhibitor herbicides in a Lolium multiflorum population

BACKGROUND: The increasing use of ACCase‐inhibiting herbicides has resulted in evolved resistance in key grass weeds infesting cereal cropping systems worldwide. Here, a thorough and systematic approach is proposed to elucidate the basis of resistance to three ACCase herbicides in a Lolium multiflorum Lam. (Italian rye grass) population from the United Kingdom (UK24). RESULTS: Resistance to sethoxydim and pinoxaden was always associated with a dominant D2078G (Alopecurus myosuroides Huds. equivalent) target‐site mutation in UK24. Conversely, whole‐plant herbicide assays on predetermined ACCase genotypes showed very high levels of resistance to diclofop‐methyl for all three wild DD2078 and mutant DG2078 and GG2078 ACCase genotypes from the mixed resistant population UK24. This indicates the presence of other diclofop‐methyl‐specific resistance mechanism(s) yet to be determined in this population. The D2078G mutation could be detected using an unambiguous DNA‐based dCAPS procedure that proved very transferable to A. myosuroides, Avena fatua L., Setaria viridis (L.) Beauv. and Phalaris minor Retz. CONCLUSION: This study provides further understanding of the molecular basis of resistance to ACCase inhibitor herbicides in a Lolium population and a widely applicable PCR‐based method for monitoring the D2078G target‐site resistance mutation in five major grass weed species. Copyright © 2010 Society of Chemical Industry     

A novel amino acid substitution Ala-122-Tyr in ALS confers high-level and broad resistance across ALS-inhibiting herbicides

BACKGROUND: Wild radish, a problem weed worldwide, is a severe dicotyledonous weed in crops. In Australia, sustained reliance on ALS‐inhibiting herbicides to control this species has led to the evolution of many resistant populations endowed by any of several ALS mutations. The molecular basis of ALS‐inhibiting herbicide resistance in a novel resistant population was studied. RESULTS: ALS gene sequencing revealed a previously unreported substitution of Tyr for Ala at amino acid position 122 in resistant individuals of a wild radish population (WARR30). A purified subpopulation individually homozygous for the Ala‐122‐Tyr mutation was generated and characterised in terms of its response to the different chemical classes of ALS‐inhibiting herbicides. Whole‐plant dose‐response studies showed that the purified subpopulation was highly resistant to chlorsulfuron, metosulam and imazamox, with LD₅₀ or GR₅₀ R/S ratio of > 1024, > 512 and > 137 respectively. The resistance to imazypyr was found to be relatively moderate (but still substantial), with LD₅₀ and GR₅₀ R/S ratios of > 16 and > 7.8 respectively. In vitro ALS activity assays showed that Ala‐122‐Tyr ALS was highly resistant to all tested ALS‐inhibiting herbicides. CONCLUSION: The molecular basis of ALS‐inhibiting herbicide resistance in wild radish population WARR30 was identified to be due to an Ala‐122‐Tyr mutation in the ALS gene. This is the first report of an amino acid substitution at Ala‐122 in the plant ALS that confers high‐level and broad‐spectrum resistance to ALS‐inhibiting herbicides, a remarkable contrast to the known mutation Ala‐122‐Thr endowing resistance to imidazolinone herbicide. Copyright © 2012 Society of Chemical Industry