Modern herbicides make major contributions to global food production by easily removing weeds and substituting for destructive soil cultivation. However, persistent herbicide selection of huge weed numbers across vast areas can result in the rapid evolution of herbicide resistance. Herbicides target specific enzymes, and mutations are selected that confer resistance-endowing amino acid substitutions, decreasing herbicide binding. Where herbicides bind within an enzyme catalytic site very few mutations give resistance while conserving enzyme functionality. Where herbicides bind away from a catalytic site many resistance-endowing mutations may evolve. Increasingly, resistance evolves due to mechanisms limiting herbicide reaching target sites. Especially threatening are herbicide-degrading cytochrome P450 enzymes able to detoxify existing, new, and even herbicides yet to be discovered. Global weed species are accumulating resistance mechanisms, displaying multiple resistance across many herbicides and posing a great challenge to herbicide sustainability in world agriculture. Fascinating genetic issues associated with resistance evolution remain to be investigated, especially the possibility of herbicide stress unleashing epigenetic gene expression. Understanding resistance and building sustainable solutions to herbicide resistance evolution are necessary and worthy challenges.

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Evolution in Action: Plants Resistant to Herbicides

Glyphosate is the world's most important herbicide, with many uses that deliver effective and sustained control of a wide spectrum of unwanted (weedy) plant species. Until recently there were relatively few reports of weedy plant species evolving resistance to glyphosate. Since 1996, the advent and subsequent high adoption of transgenic glyphosate-resistant crops in the Americas has meant unprecedented and often exclusive use of glyphosate forweedcontroloververylarge areas.Consequently, in regionsoftheUSA wheretransgenic glyphosate- resistant crops dominate, there are now evolved glyphosate-resistant populations of the economically damaging weed species Ambrosia artemissifolia L., Ambrosia trifida L., Amaranthus palmeri SW atson,Amaranthus rudis JD Sauer, Amaranthus tuberculatus (Moq) JD Sauer and various Conyza and Lolium spp. Likewise, in areas of transgenic glyphosate-resistant crops in Argentina and Brazil, there are now evolved glyphosate-resistant populations ofSorghum halepense (L.) Pers andEuphorbia heterophylla L. respectively. As transgenic glyphosate- resistant crops will remain very popular with producers, it is anticipated that glyphosate-resistant biotypes of other prominent weed species will evolve over the next few years. Therefore, evolved glyphosate-resistant weeds are a major risk for the continued success of glyphosate and transgenic glyphosate-resistant crops. However, glyphosate-resistant weeds are not yet a problem in many parts of the world, and lessons can be learnt and actions taken to achieve glyphosate sustainability. A major lesson is that maintenance of diversity in weed management systems is crucial for glyphosate to be sustainable. Glyphosate is essential for present and future world food production, and action to secure its sustainability for future gene rations is ag lobal imperative.  2008 Society of Chemical Industry

Evolved glyphosate-resistant weeds around the world: lessons to be learnt

Studies of mechanisms of resistance to glyphosate have increased current understanding of herbicide resistance mechanisms. Thus far, single-codon non-synonymous mutations of EPSPS (5-enolypyruvylshikimate-3-phosphate synthase) have been rare and, relative to other herbicide mode of action target-site mutations, unconventionally weak in magnitude for resistance to glyphosate. However, it is possible that weeds will emerge with non-synonymous mutations of two codons of EPSPS to produce an enzyme endowing greater resistance to glyphosate. Today, target-gene duplication is a common glyphosate resistance mechanism and could become a fundamental process for developing any resistance trait. Based on competition and substrate selectivity studies in several species, rapid vacuole sequestration of glyphosate occurs via a transporter mechanism. Conversely, as the chloroplast requires transporters for uptake of important metabolites, transporters associated with the two plastid membranes may separately, or together, successfully block glyphosate delivery. A model based on finite glyphosate dose and limiting time required for chloroplast loading sets the stage for understanding how uniquely different mechanisms can contribute to overall glyphosate resistance.

https://onlinelibrary.wiley.com/doi/pdf/10.1002/ps.3743

Glyphosate resistance: state of knowledge.

Herbicide cross resistance in weeds

Non-target-site-based resistance (NTSR) can confer unpredictable cross-resistance to herbicides. However, the genetic determinants of NTSR remain poorly known. The current, urgent challenge for weed scientists is thus to elucidate the bases of NTSR so that detection tools are developed, the evolution of NTSR is understood, the efficacy of the shrinking herbicide portfolio is maintained and integrated weed management strategies, including fully effective herbicide applications, are designed and implemented. In this paper, the importance of NTSR in resistance to herbicides is underlined. The most likely way in which NTSR evolves-by accumulation of different mechanisms within individual plants-is described. The NTSR mechanisms, which can interfere with herbicide penetration, translocation and accumulation at the target site, and/or protect the plant against the consequences of herbicide action, are then reviewed. NTSR is a part of the plant stress response. As such, NTSR is a dynamic process unrolling over time that involves 'protectors' directly interfering with herbicide action, and also regulators controlling 'protector' expression. NTSR is thus a quantitative trait. On this basis, a three-step procedure is proposed, based on the use of the 'omics' (genomics, transcriptomics, proteomics or metabolomics), to unravel the genetic bases of NTSR.

Unravelling the genetic bases of non-target-site-based resistance (NTSR) to herbicides: a major challenge for weed science in the forthcoming decade

Glyphosate behavior was examined in Italian ryegrass plants from Chile that were sensitive (S) and resistant (R) to this herbicide. In order to explain the resistance to glyphosate, contact angles, spray retention, foliar uptake, herbicide translocation, and target enzyme activity were studied. Contact angles of glyphosate solutions at a field concentration were 40° to 45° on the abaxial surface of R leaves as compared to 70° on S. Glyphosate spray retention by R plants was 35% lower than by S plants. Glyphosate uptake by the abaxial leaf surface of R plants was about 40% lower than that of S plants. In addition, in the R plants more glyphosate migrated to the tip of the treated leaves. The target enzyme in R and S plants was sensitive to the herbicide. Based on these and previous results, it is concluded that resistance in this Italian ryegrass biotype results from lower spray retention, lower foliar uptake from the abaxial leaf surface, and altered translocation pattern. The decreases in spray ...

Mechanisms of Resistance to Glyphosate in a Ryegrass (Lolium multiflorum) Biotype from Chile

The evolution of glyphosate-resistant weeds has recently increased dramatically. Six suspected glyphosate-resistant Amaranthus tuberculatus populations were studied to confirm resistance and determine the resistance mechanism. Resistance was confirmed in greenhouse for all six populations with glyphosate resistance factors (R/S) between 5.2 and 7.5. No difference in glyphosate absorption or translocation was observed between resistant and susceptible individuals. No mutation at amino acid positions G101, T102, or P106 was detected in the EPSPS gene coding sequence, the target enzyme of glyphosate. Analysis of EPSPS gene copy number revealed that all glyphosate-resistant populations possessed increased EPSPS gene copy number, and this correlated with increased expression at both RNA and protein levels. EPSPS Vmax and Kcat values were more than doubled in resistant plants, indicating higher levels of catalytically active expressed EPSPS protein. EPSPS gene amplification is the main mechanism contributing to glyphosate resistance in the A. tuberculatus populations analyzed.

Characterization of glyphosate resistance in Amaranthus tuberculatus populations.

Fenoxaprop-p-ethyl (FE), 2-{4-[(6-chloro-2-benzoxazolyl)oxy]phenoxy} propanoate, ethyl ester (R), is an aryloxyphenoxypropionate herbicide for postemergence control of annual and perennial grasses in paddy fields; its site of action is acetyl-coenzyme A carboxylase (ACCase), an enzyme in fatty acids biosynthesis. The possible mechanism(s) of resistance to FE in a resistant biotype of Echinochloa phyllopogon was examined, namely, absorption, translocation, and metabolism of FE and ACCase susceptibility to fenoxaprop acid (FA). Studies of the in vitro inhibition of ACCase discounted any differential active site sensitivity as the basis of resistance to FE. There were differences in absorption rates between biotypes from 3 to 48 h after application (HAA). Biotypes did not differ in either the amounts or the rates of FE translocated; 98% of applied [14C]FE remaining in the treated leaf. However, there was a good correlation between the rate of herbicide metabolism and the plant resistance. The R biotype produ...

Late watergrass (Echinochloa phyllopogon): mechanisms involved in the resistance to fenoxaprop-p-ethyl.

The continuous use of a single herbicide for weed control can result in selection of biotypes resistant to that compound. Greenhouse experiments were conducted to assess the occurrence of wild poinsettia (Euphorbia heterophylla, EPHHL) resistant biotypes to glyphosate. Two suspected glyphosateresistant biotypes from the northern part of Rio Grande do Sul, Brazil, were compared to known glyphosate-susceptible biotypes. Dose-response curves were used to compare the biotypes, with rates ranging from 0 to 450 g ha -1 in one experiment, and from 0 to 1200 g ha -1 in another. The resistance factor, calculated with the I 50 data, indicated the resistant biotypes were about three times less sensitive to glyphosate than the susceptible biotypes. This is the first report of a glyphosate-resistant biotype in a weed species of major importance and distribution in Brazil. A risk analysis is discussed for the occurrence of glyphosate-resistant wild poinsettia in glyphosate-tolerant soybeans.

Glyphosate resistant biotypes of wild poinsettia (Euphorbia heterophylla L.) and its risk analysis on glyphosate-tolerant soybeans

Cyhalofop-butyl (CB), 2-[4-(4-cyano-2-fluorophenoxy)phenoxy]propanoic acid, butyl ester (R), is an aryloxyphenoxypropionate (AOPP) herbicide for postemergence use in rice to control grasses, mainly Echinochloa spp. Similar to other AOPP and cyclohexanedione herbicides, the site of action of CB is acetyl-coenzyme A carboxylase (ACCase), an enzyme in fatty acid biosynthesis. The mechanisms involved in the selectivity of CB in rice (Oryza sativa L.)—absorption, translocation, metabolism, and ACCase susceptibility—were studied. Studies of in vitro inhibition of ACCase in E. oryzoides and O. sativa L. species discounted any differential active site sensitivity as the basis of tolerance to CB. The O. sativa L. cuticle was uniformly covered by waxes, with predominantly unshaped large waxes randomly distributed, obtaining absorption values of under 30%, 24 h after application (HAA). The E. oryzoides cuticle formed a non-uniform covered reticule, with less wax density and areas lacking in waxes reaching maximum values of absorption rising to 73%, 24 HAA. Translocation studies revealed no significant differences, either between species, or between times, remaining in the treated leaf. There was a good correlation between the rate of metabolism and plant tolerance. Plant metabolism studies demonstrated that tolerant rice inactivated the esterases producing a lack of functionality thus reducing the conversion of CB to cyhalofop acid, which is the active form of the herbicide. Moreover, it increased the metabolism of the herbicide forming non toxic metabolites much faster than E. oryzoides. It was concluded that the basis of rice tolerance to CB was a lack of esterase functionality, a reduced absorption through the cuticle and an increase in cyhalofop acid metabolism.

J. P. Ruiz-Santaella

Papers

Mechanisms of Resistance to Glyphosate in a Ryegrass (Lolium multiflorum) Biotype from Chile

Characterization of glyphosate resistance in Amaranthus tuberculatus populations.

Late watergrass (Echinochloa phyllopogon): mechanisms involved in the resistance to fenoxaprop-p-ethyl.

Glyphosate resistant biotypes of wild poinsettia (Euphorbia heterophylla L.) and its risk analysis on glyphosate-tolerant soybeans

Basis of selectivity of cyhalofop-butyl in Oryza sativa L.