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.     

Integrated pest management and weed management in the United States and Canada

There is interest in more diverse weed management tactics because of evolved herbicide resistance in important weeds in many US and Canadian crop systems. While herbicide resistance in weeds is not new, the issue has become critical because of the adoption of simple, convenient and inexpensive crop systems based on genetically engineered glyphosate‐tolerant crop cultivars. Importantly, genetic engineering has not been a factor in rice and wheat, two globally important food crops. There are many tactics that help to mitigate herbicide resistance in weeds and should be widely adopted. Evolved herbicide resistance in key weeds has influenced a limited number of growers to include a more diverse suite of tactics to supplement existing herbicidal tactics. Most growers still emphasize herbicides, often to the exclusion of alternative tactics. Application of integrated pest management for weeds is better characterized as integrated weed management, and more typically integrated herbicide management. However, adoption of diverse weed management tactics is limited. Modifying herbicide use will not solve herbicide resistance in weeds, and the relief provided by different herbicide use practices is generally short‐lived at best. More diversity of tactics for weed management must be incorporated in crop systems. © 2014 Society of Chemical Industry     

Metabolism and selectivity of rice herbicides in plants

Chemical weed control with effective and highly active herbicides has been very useful and convenient means. It has contributed to stable crop production and is labor saving. Recent herbicides have had characteristics such as high effectiveness without causing environmental pollution or harmful effects, and appropriate herbicides having high activity, low toxicity, high selectivity and being non-persistent have been developed. The metabolism of rice herbicides used mainly in Japan, such as sulfonylurea, chloroacetamide, acylamide, urea, thiocarbamate, pyrazole, triazine, diphenyl ether, phthalimide, phenoxy, aryloxyphenoxypropionate, etc., is reviewed, and its involvement in selectivity is also discussed. The metabolism of herbicides is closely related to their activity and selectivity. Differential herbicide metabolism in plants is a contributing factor of selectivity between crops and weeds. Chemicals that are more detoxified in crops and/or more activated or less detoxified in weeds are considered as being effective and selective herbicides. The metabolism of various types of rice herbicides includes: oxidative reaction (ring and chain hydroxylation, O - and N -dealkylation), hydrolysis and subsequent glucose conjugation, and glutathione conjugation in rice. These detoxicative activities are much higher in rice than weeds in paddies, and this leads to the selectivity of herbicides. Enzymes, oxidase, P-450 mono-oxygenase, esterase, acylamidase, glucosyl transferase, glutathione transferase, etc., play important roles in herbicide metabolism and selectivity.     

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.     

Herbicides and plant hormesis

Herbicide hormesis is commonly observed at subtoxic doses of herbicides and other phytotoxins. The occurrence and magnitude of this phenomenon are influenced by plant growth stage and physiological status, environmental factors, the endpoint measured and the timing between treatment and endpoint measurement. The mechanism in some cases of herbicide hormesis appears to be related to the target site of the herbicide, whereas in other examples hormesis may be by overcompensation to moderate stress induced by the herbicides or a response to disturbed homeostasis. Theoretically, herbicide hormesis could be used in crop production, but this has been practical only in the case of the use of herbicides as sugar cane ‘ripeners’ to enhance sucrose accumulation. The many factors that can influence the occurrence, the magnitude and the dose range of hormetic increases in yield for most crops make it too unpredictable and risky as a production practice with the currently available knowledge. Herbicide hormesis can cause undesired effects in situations in which weeds are unintentionally exposed to hormetic doses (e.g. in adjacent fields, when shielded by crop vegetation). Some weeds that have evolved herbicide resistance may have hormetic responses to recommended herbicide application rates. Little is known about such effects under field conditions. A more complete understanding of herbicide hormesis is needed to exploit its potential benefits and to minimize its potential harmful effects in crop production. © 2014 Society of Chemical Industry     

Perspectives on transgenic,herbicide‐resistant crops in the United States almost 20 years after introduction

Herbicide‐resistant crops have had a profound impact on weed management. Most of the impact has been by glyphosate‐resistant maize, cotton, soybean and canola. Significant economic savings, yield increases and more efficacious and simplified weed management have resulted in widespread adoption of the technology. Initially, glyphosate‐resistant crops enabled significantly reduced tillage and reduced the environmental impact of weed management. Continuous use of glyphosate with glyphosate‐resistant crops over broad areas facilitated the evolution of glyphosate‐resistant weeds, which have resulted in increases in the use of tillage and other herbicides with glyphosate, reducing some of the initial environmental benefits of glyphosate‐resistant crops. Transgenic crops with resistance to auxinic herbicides, as well as to herbicides that inhibit acetolactate synthase, acetyl‐CoA carboxylase and hydroxyphenylpyruvate dioxygenase, stacked with glyphosate and/or glufosinate resistance, will become available in the next few years. These technologies will provide additional weed management options for farmers, but will not have all of the positive effects (reduced cost, simplified weed management, lowered environmental impact and reduced tillage) that glyphosate‐resistant crops had initially. In the more distant future, other herbicide‐resistant crops (including non‐transgenic ones), herbicides with new modes of action and technologies that are currently in their infancy (e.g. bioherbicides, sprayable herbicidal RNAi and/or robotic weeding) may affect the role of transgenic, herbicide‐resistant crops in weed management. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.     

Contamination of internationally traded wheat by herbicide‐resistant Lolium rigidum

As herbicide‐resistant weeds have spread in the agricultural fields of grain‐exporting countries, their seeds could be introduced into other countries as contaminants in imported grain. The spread of resistance genes through seed and pollen can cause significant economic loss. In order to assess the extent of the problem, we investigated the contamination by herbicide‐resistant annual ryegrass (Lolium rigidum) of wheat imported from Western Australia into Japan. Annual ryegrass seeds were recovered from wheat shipments and seed bioassays were conducted to identify resistance to the herbicides that are commonly used in Australia: diclofop‐methyl, sethoxydim, chlorsulfuron, and glyphosate. Nearly 4500 ryegrass seeds were detected in 20 kg of wheat that was imported in both 2006 and 2007. About 35% and 15% of the seeds were resistant to diclofop‐methyl, 5% and 6% were resistant to sethoxydim, and 56% and 60% were resistant to chlorsulfuron in 2006 and 2007, respectively. None was resistant to glyphosate in either year. As the contamination of crops by herbicide‐resistant weeds is probably a common phenomenon, the monitoring of incoming grain shipments is necessary to stem the further spread of herbicide‐resistant weeds into importing countries.     

除草剂安全剂作用机理研究进展

除草剂安全剂是一类可在不影响除草剂对靶标杂草活性的前提下,有选择性地保护作物免受除草剂伤害的特殊用途化合物,有关安全剂作用机理的研究对新安全剂的开发具有重要意义。目前关于除草剂安全剂的作用机理主要有4种观点:1）影响除草剂在作物体内的吸收和转运;2）与除草剂竞争靶标位点;3）影响靶标酶的活性;4）增强作物对除草剂的代谢。文章对近年来安全剂作用机理及安全剂对杂草的影响等研究进展进行了综述,并分析了当前存在的问题及未来的研究方向,旨在为深入研究安全剂的作用机理及新安全剂开发提供参考。     

Why have no new herbicide modes of action appeared in recent years?

Herbicides with new modes of action are badly needed to manage the evolution of resistance of weeds to existing herbicides. Yet no major new mode of action has been introduced to the market place for about 20 years. There are probably several reasons for this. New potential products may have remained dormant owing to concerns that glyphosate-resistant (GR) crops have reduced the market for a new herbicide. The capture of a large fraction of the herbicide market by glyphosate with GR crops led to significantly diminished herbicide discovery efforts. Some of the reduced herbicide discovery research was also due to company consolidations and the availability of more generic herbicides. Another problem might be that the best herbicide molecular target sites may have already been discovered. However, target sites that are not utilized, for which there are inhibitors that are highly effective at killing plants, suggests that this is not true. Results of modern methods of target site discovery (e.g. gene knockout methods) are mostly not public, but there is no evidence of good herbicides with new target sites coming from these approaches. In summary, there are several reasons for a long dry period for new herbicide target sites; however, the relative magnitude of each is unclear. The economic stimulus to the herbicide industry caused by the evolution of herbicide-resistant weeds, especially GR weeds, may result in one or more new modes of action becoming available in the not too distant future.     

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.     

Herbicides used in combination can reduce the probability of herbicide resistance in finite weed populations

A simulation study was conducted to examine the effect of pattern of herbicide use on development of resistance to two herbicides with different modes of action in finite weed populations. The effects of the size of the treatment area (analogous to initial weed population), germination fraction and degree of self‐pollination in the weed were investigated. The results indicate that the probability of developing resistance to one or both herbicides decreases as the size of the area/initial population decreases. For treatment areas of 100 ha or less with an initial weed seedbank of 100 seeds m^?2 and initial frequencies of the resistance genes of 10^?6, development of resistance to both herbicides (double‐resistance) is uncommon within 50 years for all types of weeds if both herbicides are used in all years (used in combination). If herbicides are used in alternate years (rotated) double‐resistance almost always occurs in 100 ha areas but is uncommon in areas of 1 ha or less. The results suggest that adoption of practices that limit movement of weeds in conjunction with using herbicides in combination rather than in rotation can substantially delay development of herbicide resistance.     

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。     

Effects of rye cover crop residue and herbicides on weed control in narrow and wide row soybean planting systems

Α three‐year, non‐irrigated field study was conducted in 1998, 1999, and 2000 at the Southern Weed Science Research Unit farm, Stoneville, MS to study the effects of rye cover crop residue, soybean planting systems, and herbicide application programs on the control, density and biomass of several weed species and soybean yield. The soybean planting systems comprised 19 cm rows with high plant density, 57 cm rows with medium plant density, and 95 cm rows with low plant density. The herbicide programs evaluated were pre‐emergence, postemergence, pre‐emergence followed by postemergence, and no herbicide. Flumetsulam and metolachlor were applied pre‐emergence, and acifluorfen, bentazon, and clethodim were applied postemergence. The presence or absence of rye cover crop residue and a soybean planting system did not affect weed control of the species evaluated (browntop millet, barnyard grass, broadleaf signal grass, pitted morningglory, yellow nutsedge, Palmer amaranth and hyssop spurge), when herbicides were applied, regardless of the application program. In addition, rye cover crop residue was not an effective weed management tool when no herbicide was applied, because density and biomass of most weeds evaluated were higher than a no cover crop residue system. Among soybean planting systems, narrow with high plant density soybeans reduced density of grasses, broadleaf weeds and yellow nutsedge by 24–83% and total weed biomass by 38%, compared to wide with low plant density soybeans. Although weed pressure was reduced by narrow with high plant density soybeans, herbicide applications had the most impact on weed control, weed density and biomass. All herbicide programs controlled all weed species 81–100% at two weeks after postemergence herbicide applications, in comparison to no‐herbicide. Density of grasses and all broadleaf weeds as well as total weed biomass was lower with the pre‐emergence followed by postemergence program than these programs alone. Soybean yields were higher in the pre‐emergence followed by postemergence, and postemergence only programs than the pre‐emergence alone program. Planting crops in narrow rows is one cultural method of reducing weed pressure. However, even with the use of this cultural practice, prevalent weed pressure often requires management with herbicides.     

Herbicide‐resistant weed management: focus on glyphosate

This review focuses on proactive and reactive management of glyphosate‐resistant (GR) weeds. Glyphosate resistance in weeds has evolved under recurrent glyphosate usage, with little or no diversity in weed management practices. The main herbicide strategy for proactively or reactively managing GR weeds is to supplement glyphosate with herbicides of alternative modes of action and with soil‐residual activity. These herbicides can be applied in sequences or mixtures. Proactive or reactive GR weed management can be aided by crop cultivars with alternative single or stacked herbicide‐resistance traits, which will become increasingly available to growers in the future. Many growers with GR weeds continue to use glyphosate because of its economical broad‐spectrum weed control. Government farm policies, pesticide regulatory policies and industry actions should encourage growers to adopt a more proactive approach to GR weed management by providing the best information and training on management practices, information on the benefits of proactive management and voluntary incentives, as appropriate. Results from recent surveys in the United States indicate that such a change in grower attitudes may be occurring because of enhanced awareness of the benefits of proactive management and the relative cost of the reactive management of GR weeds. Copyright © 2011 Society of Chemical Industry     

Benchmark study on glyphosate-resistant cropping systems in the United States. Part 4: Weed management practices and effects on weed populations and soil seedbanks

BACKGROUND: Weed management in glyphosate‐resistant (GR) maize, cotton and soybean in the United States relies almost exclusively on glyphosate, which raises criticism for facilitating shifts in weed populations. In 2006, the benchmark study, a field‐scale investigation, was initiated in three different GR cropping systems to characterize academic recommendations for weed management and to determine the level to which these recommendations would reduce weed population shifts. RESULTS: A majority of growers used glyphosate as the only herbicide for weed management, as opposed to 98% of the academic recommendations implementing at least two herbicide active ingredients and modes of action. The additional herbicides were applied with glyphosate and as soil residual treatments. The greater herbicide diversity with academic recommendations reduced weed population densities before and after post‐emergence herbicide applications in 2006 and 2007, particularly in continuous GR crops. CONCLUSION: Diversifying herbicides reduces weed population densities and lowers the risk of weed population shifts and the associated potential for the evolution of glyphosate‐resistant weeds in continuous GR crops. Altered weed management practices (e.g. herbicides or tillage) enabled by rotating crops, whether GR or non‐GR, improves weed management and thus minimizes the effectiveness of only using chemical tactics to mitigate weed population shifts. Copyright © 2011 Society of Chemical Industry     

Evolving understanding of the evolution of herbicide resistance

A greater number of, and more varied, modes of resistance have evolved in weeds than in other pests because the usage of herbicides is far more extensive than the usage of other pesticides, and because weed seed output is so great. The discovery and development of selective herbicides are more problematic than those of insecticides and fungicides, as these must only differentiate between plant and insect or pathogen. Herbicides are typically selective between plants, meaning that before deployment there are already some crops possessing natural herbicide resistance that weeds could evolve. The concepts of the evolution of resistance and the mechanisms of delaying resistance have evolved as nature has continually evolved new types of resistance. Major gene target‐site mutations were the first types to evolve, with initial consideration devoted mainly to them, but slowly ‘creeping’ resistance, gradually accruing increasing levels of resistance, has become a major force owing to an incremental accumulation of genetic changes in weed populations. Weeds have evolved mechanisms unknown even in antibiotic as well as other drug and pesticide resistances. It is even possible that cases of epigenetic ‘remembered’ resistances may have appeared. Copyright © 2009 Society of Chemical Industry     

抗除草剂转基因水稻的研究进展

水稻是我国乃至世界上最重要的粮食作物之一,杂草是影响水稻产量和品质的主要因素。近年来,抗除草剂转基因水稻的研究与应用取得了较大进展。本文概述了稻田的主要杂草及防除方法,简述了稻田常用的几种除草剂及其作用机理,介绍了全球抗除草剂转基因作物的现状,着重阐述了抗除草剂转基因水稻的研发进展、应用及未来发展方向。     

Cross‐resistance of Echinochloa species to acetolactate synthase inhibitor herbicides

This study was conducted to evaluate the cross‐resistance of acetolactate synthase (ALS) inhibitors with different chemistries, specifically azimsulfuron (sulfonylurea), penoxsulam (triazolopyrimidine sulfonanilide) and bispyribac‐sodium (pyrimidinyl thio benzoate), in Echinochloa oryzicola and Echinochloa crus‐galli that had been collected in South Korea and to investigate their herbicide resistance mechanism. Both Echinochloa spp. showed cross‐resistance to the ALS inhibitors belonging to the above three different chemistries. In a whole plant assay with herbicides alone, the resistant/susceptible ratios for azimsulfuron, penoxsulam and bispyribac‐sodium were 12.6, 28.1 and 1.9 in E. oryzicola and 21.1, 13.7 and 1.8 in E. crus‐galli, respectively. An in vitro ALS enzyme assay with herbicides showed that the I ₅₀‐values of the resistant accessions were approximately two‐to‐three times higher than the susceptible accessions, with no statistical difference, suggesting that the difference in ALS sensitivity cannot explain ALS inhibitor resistance in Echinochloa spp. for azimsulfuron, penoxsulam and bispyribac‐sodium. A whole plant assay with fenitrothion showed that the GR ₅₀‐values significantly decreased in both the resistant E. oryzicola and E. crus‐galli accessions when azimsulfuron, penoxsulam and bispyribac‐sodium were applied with the P450 inhibitor, while no significant decrease was observed in the susceptible accessions when the P450 inhibitor was used. Thus, these results suggest that ALS inhibitor cross‐resistance for azimsulfuron, penoxsulam and bispyribac‐sodium is related to enhanced herbicide metabolism.