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.     

Competitive ability of ALS‐inhibitor herbicide‐resistant Fimbristylis miliacea

Fimbristylis miliacea, a weed in rice, has evolved resistance to acetolactate synthase (ALS) inhibitors. This study aimed to investigate the competitive abilities of ALS‐resistant (R) and ALS‐susceptible (S) F. miliacea with rice. A replacement series experiment was conducted in the glasshouse at the Federal University of Pelotas, Brazil. The proportions of rice to F. miliacea were 100:0, 75:25, 50:50, 25:75 and 0:100, with 1060 plants m^?2. The experimental units were arranged in a completely randomised design with four replications. A follow‐up study was conducted at the University of Arkansas, Fayetteville, USA, in a split‐plot design with four replications. The main plot was species mixture (rice × R, rice × S, R × S). The subplot was competition partitioning (below‐ and above‐ground, below‐ground only, above‐ground only and no interspecific competition). Leaf area, plant height and shoot dry mass were recorded. Rice was more competitive than the R or S F. miliacea. In equal proportions of rice and F. miliacea, regardless of ecotype, the relative leaf area, height and dry mass of rice were greater than that of F. miliacea. The ALS‐resistant ecotype was less competitive with rice than the S ecotype. Intraspecific competition among rice plants was stronger than rice competition with F. miliacea. Competition for below‐ground resources was the most critical aspect of interference among rice and F. miliacea. In production fields, high infestation levels of F. miliacea results in significant yield losses; thus, resistance to ALS inhibitors needs to be curtailed.     

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.     

Our top 10 herbicide‐resistant weed management practices

Although proactive or reactive herbicide‐resistant weed management (HRWM) practices have been recommended to growers in different agroecoregions globally, there is a need to identify and prioritise those having the most impact in mitigating or managing herbicide selection pressure in the northern Great Plains of North America. Our perspective on this issue is based on collaborative research, extension activities and dialogue with growers or farming experience (cereal, oilseed and pulse crop production) during the past 30 years. We list our top 10 HRWM practices, concluding with the number 1 practice which is the foundation of the other nine practices: crop diversity. Although our top 10 HRWM practices have broad applicability across agroecoregions, their ranking may vary widely. © 2017 Society of Chemical Industry     

Characterization of acetolactate synthase from sulfonylurea herbicide‐resistant Schoenoplectus juncoides

Ten accessions of sulfonylurea‐resistant Schoenoplectus juncoides were collected from paddy fields in Japan. In order to characterize acetolactate synthase from sulfonylurea‐resistant S. juncoides, acetolactate synthase amino acid substitutions, whole‐plant growth inhibition and acetolactate synthase enzyme inhibition were examined. Schoenoplectus juncoides has two acetolactate synthase genes (ALS1 and ALS2). The sulfonylurea‐resistant accessions harbored amino acid substitutions at Pro197 or Trp574 in either ALS1 or ALS2 (the amino acid number is standardized to the Arabidopsis thaliana sequence). The whole plants of all the sulfonylurea‐resistant accessions showed resistance to imazosulfuron. The resistance level depended on the altered amino acid residues in acetolactate synthase. The acetolactate synthase enzyme that was partially purified from all the sulfonylurea‐resistant accessions was less sensitive to imazosulfuron, compared to the susceptible accession, suggesting that the resistance is related to the altered acetolactate synthase enzyme. In addition, the concentration–response inhibition of acetolactate synthase activity by imazosulfuron in the sulfonylurea‐resistant accessions was remarkably different with the presence of an amino acid substitution in either ALS1 or ALS2. Furthermore, the concentration–response inhibition of acetolactate synthase activity in the sulfonylurea‐resistant accessions with a P197S, P197T or W574L mutation showed a double‐sigmoid curve. The regression analysis of enzyme inhibition suggested that the abundance ratio of ALS1 to ALS2 enzymes was approximately 70:30%, with a range of ±15%. Taken together, these results suggest that the resistance of sulfonylurea‐resistant accessions of S. juncoides is related to altered acetolactate synthase in either ALS1 or ALS2, although the abundance of the altered acetolactate synthase in the plants is different among the sulfonylurea‐resistant accessions.     

An approach to investigate the costs of herbicide‐resistant Alopecurus myosuroides

Herbicide resistance in Alopecurus myosuroides causes severe problems in Western European cropping systems. Costs of herbicide resistance were investigated in this study by analysing variable production costs and sales revenues. Three farms were selected for this study, with winter wheat as the dominating crop in all farms. Resistance in A. myosuroides populations was verified at all locations. Four farming approaches were simulated over a period of 20 years: (i) continuing the actual cropping system without increase of resistance, (ii) continuing the actual cropping system with increase of resistance, (iii) changing cropping practice to overcome resistance and (iv) changing cropping practice to prevent resistance. Contribution margins representing the proportion of sales revenue that is not consumed by variable costs were calculated for all approaches. Comparative static simulations showed that average contribution margins in a cropping system with more than 60% winter cereals and reduced tillage practice dropped from 807 € ha^?1 a^?1 without herbicide resistance to 307 € ha^?1 a^?1 with herbicide resistance. Alopecurus myosuroides population densities increased to more than 1000 plants m^?2. Diverse crop rotations, including spring crops, clover–grass leys and intensive tillage, suppressed A. myosuroides populations, and average contribution margin was 630 € ha^?1 a^?1. Preventive methods with rotations of winter cereals and spring crops with less clover–grass leys resulted in an average contribution margin of 691 € ha^?1 a^?1. In conclusion, rotations of winter cereals and spring crops combined with inversion tillage and herbicides provide stable yields and can prevent weed population increase.     

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     

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.     

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.     

Enhanced weed‐crop competition effects on growth and seed production of herbicide‐resistant and herbicide‐susceptible annual sowthistle (Sonchus oleraceus)

Enhanced crop competition could aid in the management of annual sowthistle (Sonchus oleraceus L.), a dominant weed of Australian cropping systems. A two‐year pot study was conducted to evaluate the effect of wheat (Triticum aestivum L.) planting densities (0, 82, and 164 wheat plants/m²) on growth and seed production of glyphosate‐resistant (GR) and glyphosate‐susceptible (GS) biotypes of annual sowthistle. Without competition, both biotypes produced a similar number of leaves and biomass, but the GS biotype produced 80% more seeds (46,050 per plant) than the GR biotype. In competition with 164 wheat plants/m², the number of leaves in the GR and GS biotypes was reduced by 62 and 61%, respectively, in comparison with the no‐competition treatment, and similarly, weed biomass was reduced by 78 and 77%, respectively. Compared to no‐competition treatment, the seed production of GR and GS biotypes was reduced by 33 and 69%, respectively, when grown with 82 wheat plants/m², but increasing wheat density from 82 to 164 plants/m² reduced the number of seeds only in the GS biotype (81%). Both biotypes produced greater than 6,000 seeds per plant when grown in competition with 164 plants/m², suggesting that increased crop density should be integrated with other weed management strategies for efficient control of annual sowthistle.     

Evaluation of six commonly used reference genes for gene expression studies in herbicide‐resistant Avena fatua biotypes

Avena fatua of the family Poaceae is one of the most common and economically damaging grass weeds. Resistance to herbicides that inhibit acetyl‐coenzyme A carboxylase and acetolactate synthase activities has recently been detected in A. fatua. The resistance may be due to mutations in the herbicide targets and/or enhanced herbicide metabolism resulting from changes in gene expression, including in genes involved in detoxifying herbicide active ingredients. To analyse gene expression, stable housekeeping genes must be experimentally determined and used for data normalisation. In this study, A. fatua plants were treated with different herbicide types and plant materials were harvested at three time points following treatment. Six candidate reference genes (18S rRNA, ACT, EF1α, GAPDH, TBP, and TUB) were selected, sequenced and analysed by RT‐qPCR. The resulting data were assessed using four algorithms from the RefFinder software to determine gene expression stability. We identified TBP and GAPDH as the most stably expressed A. fatua reference genes following herbicide treatment.     

Penoxsulam‐resistant barnyardgrass (Echinochloa crus‐galli) in rice fields in China

Barnyardgrass (Echinochloa crus‐galli) proliferation seriously threatens rice production worldwide. Whole‐plant bioassays were conducted in order to test the sensitivity to penoxsulam of 52 barnyardgrass populations and the resistance of six penoxsulam‐resistant populations to 12 other herbicides that are commonly used in rice fields. Among the 48 populations that had escaped penoxsulam control in the rice fields, 8.3% showed a very high level of resistance, 58.3% showed a high level of resistance and 10.4% showed a moderate level of resistance. Multiple resistance was confirmed in all six penoxsulam‐resistant populations that were tested further. They exhibited at least a moderate level of resistance; that is, to 6–10 of the total of 13 herbicides that was tested. Most of the six penoxsulam‐resistant populations showed at least a moderate level of resistance to bispyribac‐sodium, quinclorac, metamifop, cyhalofop‐butyl and oxadiazon, three populations held at least a moderate level of resistance to oxyfluorfen and pretilachlor, two populations also held at least a moderate level of resistance to pyrazosulfuron‐ethyl, pyribenzoxim and fenoxaprop‐P‐ethyl, but the resistance indices of the six populations to pendimethalin were all low. This study has confirmed resistance to pretilachlor and oxadiazon in weeds for the first time.     

Distribution of herbicide‐resistant acetyl‐coenzyme A carboxylase alleles in Lolium rigidum across grain cropping areas of South Australia

Resistance to the acetyl‐coenzyme A carboxylase (ACCase)‐inhibiting herbicides in Lolium rigidum is widespread in grain cropping areas of South Australia. To better understand the occurrence and spread of resistance to these herbicides and how it has changed with time, the carboxyl transferase (CT) domain of the ACCase gene from resistant L. rigidum plants, collected from both random surveys of the mid‐north of Southern Australia over 10 years as well as stratified surveys in individual fields, was sequenced and target site mutations characterised. Amino acid substitutions occurring as a consequence of these target site mutations, at seven positions in the ACCase gene previously correlated with herbicide resistance, were identified in c. 80% of resistant individuals, indicating target site mutation is a common mechanism of resistance in L. rigidum to this herbicide mode of action. Individuals containing multiple amino acid substitutions (two, and in two cases, three substitutions) were also found. Substitutions at position 2041 occurred at the highest frequency in all years of the large area survey, while substitutions at position 2078 were most common in the single farm analysis. This study has shown that target site mutations leading to amino acid substitutions in ACCase of L. rigidum are widespread across South Australia and that these mutations have likely evolved independently in different locations. The results indicate that seed movement, both within and between fields, may contribute to the spread of resistance in a single field. However, over a large area, the independent appearance and selection of target site mutations conferring resistance through herbicide use is the most important factor.