scholarly journals Diversified Resistance Mechanisms in Multi-Resistant Lolium spp. in Three European Countries

2020 ◽  
Vol 11 ◽  
Author(s):  
Laura Scarabel ◽  
Silvia Panozzo ◽  
Donato Loddo ◽  
Solvejg K. Mathiassen ◽  
Michael Kristensen ◽  
...  
Keyword(s):  
Management Strategies ◽  
Acetolactate Synthase ◽  
Resistance Mechanisms ◽  
Target Site ◽  
European Countries ◽  
Cereal Crops ◽  
Resistance Pattern ◽  
Italian Population ◽  
Als Inhibitors ◽  
Moderate Resistance

Annual ryegrass species (Lolium spp.) infest cereal crops worldwide. Ryegrass populations with multiple resistance to the acetyl coenzyme A carboxylase (ACCase) and acetolactate synthase (ALS) inhibitors are an increasing problem in several European countries. We investigated the resistance pattern and level of resistance in ryegrass populations collected in Denmark, Greece and Italy and studied the diversity of mechanisms endowing resistance, both target-site and metabolism based. All populations showed high resistance indexes (RI) to the ALS inhibitors, iodosufuron-methyl-sodium + mesosulfuron-methyl (RI from 8 to 70), whereas the responses to the two ACCase inhibitors, clodinafop-propargyl and pinoxaden, differed. The Greek and Italian populations were moderately to highly resistant to clodinafop (RI > 8) and showed low to moderate resistance to pinoxaden (RI ranged from 3 to 13) except for one Italian population. In contrast, the Danish Lolium populations showed low to moderate resistance to clodinafop (RI ranged from 2 to 7) and only one population was resistant to pinoxaden. Different mutant ACCase alleles (Leu1781, Cys2027, Asn2041, Val2041, Gly2078, Arg2088, Ala2096) and ALS alleles (Gly122, Ala197, Gln197, Leu197, Ser197, Thr197, Val205, Asn376, Glu376, Leu574) endowing resistance were detected in the Greek and Italian populations. In several plants, no mutated ALS and ACCase alleles were found showing a great heterogeneity within and among the Greek and Italian populations. Conversely, no mutant ACCase alleles were identified in the four Danish populations and only one mutant ALS allele (Leu574) was detected in two Danish populations. The expression level of nitronate monooxygenase (NMO), glutathione S-transferase (GST) and cytochrome P450s (CYP72A1 and CYP72A2) varied broadly among populations and individual plants within the populations. Constitutive up-regulation of GST, CYP72A1 and CYP72A2 was detected in resistant plants respect to susceptible plants in one Danish and one Italian population. It appears that the mechanisms underlying resistance are rather complex and diversified among Lolium spp. populations from the three countries, coevolution of both target-site resistance and metabolic based herbicide resistance appears to be a common feature in Denmark and Italy. This must be considered and carefully evaluated in adopting resistance management strategies to control Lolium spp. in cereal crops.

scholarly journals Multiple mutations in the EPSPS and ALS genes of Amaranthus hybridus underlie resistance to glyphosate and ALS inhibitors

2020 ◽  
Vol 10 (1) ◽  
Author(s):  
Maria J. García ◽  
Candelario Palma-Bautista ◽  
José G. Vazquez-Garcia ◽  
Antonia M. Rojano-Delgado ◽  
María D. Osuna ◽  
...  
Keyword(s):  
Amino Acid ◽  
Target Genes ◽  
Acetolactate Synthase ◽  
Resistance Mechanisms ◽  
Activity Level ◽  
Cross Resistance ◽  
Weed Species ◽  
Amaranthus Hybridus ◽  
Als Inhibitors ◽  
Moderate Resistance

Abstract Amaranthus hybridus is one of the main weed species in Córdoba, Argentina. Until recently, this weed was effectively controlled with recurrent use of glyphosate. However, a population exhibiting multiple resistance (MR2) to glyphosate and imazamox appeared in a glyphosate resistant (GR) soybean field, with levels of resistance up to 93 and 38-fold higher to glyphosate and imazamox, respectively compared to the susceptible (S) population. In addition to imidazolinones, MR2 plants showed high resistance levels to sulfonylamino-carbonyl (thio) benzoates and moderate resistance to sulfonylureas and triazolopyrimidines. Multiple amino acid substitutions were found in both target genes, acetolactate synthase (ALS) and 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS), responsible for conferring high herbicides resistance levels in this A. hybridus population. In the case of EPSPS, the triple amino acid substitution TAP-IVS was found. In addition, MR2 plants also showed increased EPSPS gene expression compared to susceptible plants. A Ser653Asn substitution was found in the ALS sequence of MR2, explaining the pattern of cross-resistance to the ALS-inhibitor herbicide families found at the ALS enzyme activity level. No other mutations were found in other conserved domains of the ALS gene. This is the first report worldwide of the target site resistance mechanisms to glyphosate and ALS inhibitors in multiple herbicide resistance Amaranthus hybridus.

scholarly journals A Statewide Survey of PPO-Inhibitor Resistance and the Prevalent Target-Site Mechanisms in Palmer amaranth (Amaranthus palmeri) Accessions from Arkansas

Weed Science ◽  
2017 ◽  
Vol 66 (2) ◽  
pp. 149-158 ◽  
Author(s):  
Vijay K. Varanasi ◽  
Chad Brabham ◽  
Jason K. Norsworthy ◽  
Haozhen Nie ◽  
Bryan G. Young ◽  
...  
Keyword(s):  
Resistance Mechanism ◽  
Acetolactate Synthase ◽  
Resistance Mechanisms ◽  
Palmer Amaranth ◽  
Target Site ◽  
Evolution Of Resistance ◽  
Target Site Resistance ◽  
Novel Target ◽  
Als Inhibitors ◽  
Inhibitor Resistance

Palmer amaranth is one of the most problematic weeds in the midsouthern United States, and the evolution of resistance to protoporphyrinogen oxidase (PPO) inhibitors in biotypes already resistant to glyphosate and acetolactate synthase (ALS) inhibitors is a major cause of concern to soybean and cotton growers in these states. A late-season weed-escape survey was conducted in the major row crop–producing counties (29 counties) to determine the severity of PPO-inhibitor resistance in Arkansas. A total of 227 Palmer amaranth accessions were sprayed with fomesafen at 395 g ha−1to identify putative resistant plants. A TaqMan qPCR assay was used to confirm the presence of the ΔG210 codon deletion or the R128G/M (homologous to R98 mutation in common ragweed) target-site resistance mechanisms in thePPX2gene. Out of the 227 accessions screened, 44 were completely controlled with fomesafen, and 16 had only one or two severely injured plants (≥98% mortality) when compared with the 1986 susceptible check (100% mortality). The remaining 167 accessions were genotypically screened, and 82 (49%) accessions were found to harbor the ΔG210 deletion in thePPX2gene. The R128G was observed in 47 (28%) out of the 167 accessions screened. The mutation R128M, on the other hand was rare, found in only three accessions. About 13% of the accessions were segregating for both the ΔG210 and R128G mutations. Sixteen percent of the tested accessions had mortality ratings <90% and did not test positive for the ΔG210 or the R128G/M resistance mechanisms, indicating that a novel target or non–target site resistance mechanism is likely. Overall, PPO inhibitor–resistant Palmer amaranth is widespread in Arkansas, and the ΔG210 resistance mechanism is especially dominant in the northeast corridor, while the R128G mutation is more prevalent in counties near Memphis, TN.

Target site–based resistance to penoxsulam in late watergrass (Echinochloa phyllopogon) from China

Weed Science ◽  
2019 ◽  
Vol 67 (4) ◽  
pp. 380-388 ◽  
Author(s):  
Jian Liu ◽  
Jiapeng Fang ◽  
Zongzhe He ◽  
Jun Li ◽  
Liyao Dong
Keyword(s):  
Binding Affinity ◽  
Acetolactate Synthase ◽  
Resistance Mechanisms ◽  
Target Site ◽  
Future Research ◽  
Target Site Resistance ◽  
Target Gene Sequence ◽  
Als Inhibitors ◽  
Echinochloa Phyllopogon

AbstractLate watergrass [Echinochloa phyllopogon (Stapf) Koso-Pol.] is one of the most persistent weeds in rice fields and shows resistance to some acetolactate synthase (ALS)-inhibiting herbicides, such as penoxsulam. Previous studies of E. phyllopogon’s herbicide resistance have focused on non–target site resistance mechanisms. In this study, E. phyllopogon populations from Heilong Jiang Province, China, that were possibly resistant to penoxsulam were used to identify the target site–based mechanisms of resistance. Population HSRH-520 showed a 25.4-fold higher resistance to penoxsulam than the sensitive population, HSRH-538. HSRH-520 was resistant to other ALS inhibitors, with resistance indexes ranging from 17.1 to 166. Target-gene sequence analysis revealed two different ALS genes in E. phyllopogon; a Pro-197-Ser substitution occurred in the ALS-2 gene of HSRH-520. In vitro activity assays revealed that the penoxsulam concentrations required to inhibit 50% of the ALS activity were 13.7 times higher in HSRH-520 than in HSRH-538. Molecular-docking tests showed that the Pro-197-Ser mutation reduced the binding affinity between ALS and ALS inhibitors belonging to the triazolopyrimidine, sulfonylaminocarbonyltriazolinone, and sulfonylurea families, and there were almost no effects on binding affinity when the ALS inhibitors were of the pyrimidinylthiobenzoate and imidazolinone families. Overall, the results indicated and verified that the Pro-197-Ser mutation leads to increased ALS activity by reducing the binding affinity of the inhibitor and ALS. This is the first report on the Pro-197-Ser mutation in the complete ALS gene of E. phyllopogon and will aid future research of target site–based resistance mechanisms of E. phyllopogon to ALS inhibitors.

Controlling Herbicide-Resistant Annual Bluegrass (Poa annua) Phenotypes with Methiozolin

2017 ◽  
Vol 31 (3) ◽  
pp. 470-476 ◽  
Author(s):  
James T. Brosnan ◽  
Jose J. Vargas ◽  
Gregory K. Breeden ◽  
Sarah L. Boggess ◽  
Margaret A. Staton ◽  
...  
Keyword(s):  
Acetolactate Synthase ◽  
Target Site ◽  
Multiple Resistance ◽  
Herbicide Resistant ◽  
Annual Bluegrass ◽  
Microtubule Inhibitors ◽  
Effective Option ◽  
Target Site Resistance ◽  
Susceptible Control ◽  
Als Inhibitors

Methiozolin is an isoxazoline herbicide being investigated for selective POST annual bluegrass control in managed turfgrass. Research was conducted to evaluate methiozolin efficacy for controlling two annual bluegrass phenotypes with target-site resistance to photosystem II (PSII) or enolpyruvylshikimate-3-phosphate synthase (EPSPS)-inhibiting herbicides (i.e., glyphosate), as well as phenotypes with multiple resistance to microtubule and EPSPS or PSII and acetolactate synthase (ALS)-inhibiting herbicides. All resistant phenotypes were established in glasshouse culture along with a known herbicide-susceptible control and treated with methiozolin at 0, 125, 250, 500, 1000, 2000, 4000, or 8000 g ai ha−1. Methiozolin effectively controlled annual bluegrass with target-site resistance to inhibitors of EPSPS, PSII, as well as multiple resistance to EPSPS and microtubule inhibitors. Methiozolin rates required to reduce aboveground biomass of these resistant phenotypes 50% (GR50 values) were not significantly different from the susceptible control, ranging from 159 to 421 g ha−1. A phenotype with target-site resistance to PSII and ALS inhibitors was less sensitive to methiozolin (GR50=862 g ha−1) than a susceptible phenotype (GR50=423 g ha−1). Our findings indicate that methiozolin is an effective option for controlling select annual bluegrass phenotypes with target-site resistance to several herbicides.

Target site–based penoxsulam resistance in barnyardgrass (Echinochloa crus-galli) from China

Weed Science ◽  
2019 ◽  
Vol 67 (3) ◽  
pp. 281-287 ◽  
Author(s):  
Jiapeng Fang ◽  
Tingting Liu ◽  
Yuhua Zhang ◽  
Jun Li ◽  
Liyao Dong
Keyword(s):  
Amino Acid ◽  
Amino Acid Sequence ◽  
Acetolactate Synthase ◽  
Target Site ◽  
Cross Resistance ◽  
Predicted Amino Acid Sequence ◽  
Sequence Analyses ◽  
Als Inhibitors ◽  
Higher Sensitivity

AbstractBarnyardgrass [Echinochloa crus-galli (L.) P. Beauv.] is acknowledged to be the most troublesome weed in rice fields in Anhui and Jiangsu provinces of China. It cannot be effectively controlled using certain acetolactate synthase (ALS)-inhibiting herbicides, including penoxsulam. Echinochloa crus-galli samples with suspected resistance to penoxsulam were collected to identify the target site–based mechanism underlying this resistance. Populations AXXZ-2 and JNRG-2 showed 33- and 7.3-fold resistance to penoxsulam, respectively, compared with the susceptible JLGY-3 population. Cross-resistance to other ALS inhibitors was reported in AXXZ-2 but not in JNRG-2, and occasionally showed higher sensitivity than JLGY-3. In vitro ALS activity assays revealed that penoxsulam concentrations required to inhibit 50% of ALS activity were 11 and 5.2 times greater in AXXZ-2 and JNRG-2, respectively, than in JLGY-3. DNA and predicted amino acid sequence analyses of ALS revealed Ala-205-Val and Ala-122-Gly substitutions in AXXZ-2 and JNRG-2, respectively. Our results indicate that these substitutions in ALS are at least partially responsible for resistance to penoxsulam.

scholarly journals Target Site Resistance to Acetolactate Synthase Inhibitors in Diplotaxis erucoides and Erucaria hispanica–Mechanism of Resistance and Response to Alternative Herbicides

Agronomy ◽  
2020 ◽  
Vol 10 (4) ◽  
pp. 471 ◽  
Author(s):  
Maor Matzrafi ◽  
Ofri Gerson ◽  
Moshe Sibony ◽  
Baruch Rubin
Keyword(s):  
Management Strategies ◽  
Acetolactate Synthase ◽  
Poor Control ◽  
First Case ◽  
Evolution Of Resistance ◽  
Target Site Resistance ◽  
Excellent Control ◽  
Diplotaxis Erucoides ◽  
Als Inhibitors ◽  
Two Populations

Diplotaxis erucoides and Erucaria hispanica are common weeds of the Mediterranean region; they infest various habitats including cultivated fields and roadsides. In several fields across Israel, farmers have reported on poor control of D. erucoides and E. hispanica plants using acetolactate synthase (ALS) inhibitors. Greenhouse experiments were conducted to determine the effect of various ALS inhibitors on plants from two potentially resistant D. erucoides and E. hispanica populations. Additionally, alternative management strategies using auxinic herbicides were studied. Plants from both populations exhibited resistance to all tested ALS inhibitors, up to 20-fold the label field rate, as compared with ALS sensitive populations of D. erucoides and E. hispanica. Sequencing of the ALS gene revealed Trp574 to Leu substitution in ALS-resistant D. erucoides plants, whereas a Pro197 to Ser substitution was detected in ALS-resistant E. hispanica plants. Although high levels of resistance were observed in individuals from both putative resistant populations, sensitive individuals were also detected, suggesting the evolution of resistance in these two populations is still in progress. Auxinic herbicides, 2,4-D, and mecoprop-P, provided excellent control of plants from both ALS-resistant populations. This study documents and confirms the first case of evolution of resistance to ALS inhibitors in D. erucoides and E. hispanica populations.

Multiple herbicide resistance in California Italian ryegrass (Lolium perenne ssp. multiflorum): characterization of ALS-inhibiting herbicide resistance

Weed Science ◽  
2019 ◽  
Vol 67 (3) ◽  
pp. 273-280 ◽  
Author(s):  
Parsa Tehranchian ◽  
Vijay K. Nandula ◽  
Maor Matzrafi ◽  
Marie Jasieniuk
Keyword(s):  
Lolium Perenne ◽  
Herbicide Resistance ◽  
Resistance Mechanism ◽  
Acetolactate Synthase ◽  
Target Site ◽  
Italian Ryegrass ◽  
Resistance Level ◽  
Als Inhibitors ◽  
Inhibitor Resistance ◽  
Als Inhibitor

AbstractMultiple resistance to glyphosate, sethoxydim, and paraquat was previously confirmed in two Italian ryegrass [Lolium perenne L. ssp. multiflorum (Lam.) Husnot] populations, MR1 and MR2, in northern California. Preliminary greenhouse studies revealed that both populations were also resistant to imazamox and mesosulfuron, both of which are acetolactate synthase (ALS)-inhibiting herbicides. In this study, three subpopulations, MR1-A (from seed of MR1 plants that survived a 16X rate of sethoxydim), MR1-P (from seed of MR1 plants that survived a 2X rate of paraquat), and MR2 (from seed of MR2 plants that survived a 16X rate of sethoxydim), were investigated to determine the resistance level to imazamox and mesosulfuron, evaluate other herbicide options for the control of these multiple resistant L. perenne ssp. multiflorum, and characterize the underlying ALS-inhibitor resistance mechanism(s). Based on LD50 values, the MR1-A, MR1-P, and MR2 subpopulations were 38-, 29-, 8-fold and 36-, 64-, and 3-fold less sensitive to imazamox and mesosulfuron, respectively, relative to the susceptible (Sus) population. Only MR1-P and MR2 plants were cross-resistant to rimsulfuron, whereas both MR1 subpopulations were cross-resistant to imazethapyr. Pinoxaden (ACCase inhibitor [phenylpyrazoline 'DEN']) only controlled MR2 and Sus plants at the labeled field rate. However, all plants were effectively controlled (>99%) with the labeled field rate of glufosinate. Based on I50 values, MR1-A, MR-P, and MR2 plants were 712-, 1,104-, and 3-fold and 10-, 18-, and 5-fold less responsive to mesosulfuron and imazamox, respectively, than the Sus plants. Sequence alignment of the ALS gene of resistant plants revealed a missense single-nucleotide polymorphism resulting in a Trp-574-Leu substitution in MR1-A and MR1-P plants, heterozygous in both, but not in the MR2 plants. An additional homozygous substitution, Asp-376-Glu, was identified in the MR1-A plants. Addition of malathion or piperonyl butoxide did not alter the efficacy of mesosulfuron on MR2 plants. In addition, the presence of 2,4-D had no effect on the response of mesosulfuron on the MR2 and Sus. These results suggest an altered target site is the mechanism of resistance to ALS inhibitors in MR1-A and MR1-P plants, whereas a non–target site based resistance apparatus is present in the MR2 plants.

scholarly journals Nontarget-Site Resistance to ALS Inhibitors in Waterhemp (Amaranthus tuberculatus)

Weed Science ◽  
2015 ◽  
Vol 63 (2) ◽  
pp. 399-407 ◽  
Author(s):  
Jiaqi Guo ◽  
Chance W. Riggins ◽  
Nicholas E. Hausman ◽  
Aaron G. Hager ◽  
Dean E. Riechers ◽  
...  
Keyword(s):  
Acetolactate Synthase ◽  
Enzyme Assays ◽  
Whole Plant ◽  
Site Of Action ◽  
Amaranthus Tuberculatus ◽  
Resistance Trait ◽  
Als Inhibitors ◽  
Moderate Resistance ◽  
Inhibitor Resistance ◽  
Als Inhibitor

A waterhemp population (MCR) previously characterized as resistant to 4-hydroxyphenylpyruvate dioxygenase and photosystem II inhibitors demonstrated both moderate and high levels of resistance to acetolactate synthase (ALS) inhibitors. Plants from the MCR population exhibiting high resistance to ALS inhibitors contained the commonly found Trp574Leu ALS amino acid substitution, whereas plants with only moderate resistance did not have this substitution. A subpopulation (JG11) was derived from the MCR population in which the moderate-resistance trait was isolated from the Trp574Leu mutation. Results from DNA sequencing and ALS enzyme assays demonstrated that resistance to ALS inhibitors in the JG11 population was not due to an altered site of action. This nontarget-site ALS-inhibitor resistance was characterized with whole-plant dose–response experiments using herbicides from each of the five commercialized families of ALS-inhibiting herbicides. Resistance ratios ranging from 3 to 90 were obtained from the seven herbicides evaluated. Nontarget-site resistance to ALS has been rarely documented in eudicot weeds, and adds to the growing list of resistance traits evolved in waterhemp.

scholarly journals Target-Site Resistances to ALS and PPO Inhibitors Are Linked in Waterhemp (Amaranthus tuberculatus)

Weed Science ◽  
2016 ◽  
Vol 65 (1) ◽  
pp. 4-8 ◽  
Author(s):  
Patrick J. Tranel ◽  
Chenxi Wu ◽  
Ahmed Sadeque
Keyword(s):  
Resistance Mechanism ◽  
Acetolactate Synthase ◽  
Resistance Mechanisms ◽  
Target Site ◽  
Recurrent Parent ◽  
Common Waterhemp ◽  
Independent Assortment ◽  
Resistance Evolution ◽  
Amaranthus Tuberculatus ◽  
Molecular Marker Analysis

It is generally expected that, in the case of multiple herbicide resistance, different resistance mechanisms within a weed will follow Mendel’s law of independent assortment. Research was conducted to investigate anecdotal observations suggesting that target site–based resistances to inhibitors of acetolactate synthase (ALS) and protoporphyrinogen oxidase (PPO) did not follow independent assortment in common waterhemp. Cosegregation of the two resistances was observed in backcross lines (population sensitive to both herbicides as recurrent parent). Specifically, whereas 52% of backcross plants were resistant to a PPO inhibitor, this percentage increased to 92% when the backcross plants were preselected for resistance to an ALS inhibitor. Molecular marker analysis confirmed that the corresponding genes (ALSandPPX2) were genetically linked. When data from all plants analyzed were pooled, the genetic distance between the two genes was calculated to be 7.5 cM. The two genes were found to be about 195 kb apart in the recently published grain amaranth genome, explaining the observed genetic linkage. There is likely enough recombination that occurs between the linked genes to prevent the linkage from having significant implications in terms of resistance evolution. Nevertheless, documentation of the happenstance linkage between target-site genes for resistance to ALS and PPO inhibitors in waterhemp is a reminder that one should not assume distinct resistance mechanism will independently assort.

scholarly journals Multiple-Herbicide Resistance in a 2,4-D–Resistant Waterhemp (Amaranthus tuberculatus) Population from Nebraska

Weed Science ◽  
2017 ◽  
Vol 65 (6) ◽  
pp. 743-754 ◽  
Author(s):  
Roberto J. Crespo ◽  
Ana B. Wingeyer ◽  
Greg R. Kruger ◽  
Chance W. Riggins ◽  
Patrick J. Tranel ◽  
...  
Keyword(s):  
D1 Protein ◽  
Acetolactate Synthase ◽  
Mechanisms Of Action ◽  
Target Site ◽  
Site Mutation ◽  
Auxin Receptor ◽  
Application Rates ◽  
Chlorimuron Ethyl ◽  
Als Inhibitors ◽  
Susceptible Populations

A 2,4-D-resistant tall waterhemp population (FS) from Nebraska was evaluated for resistance to other TIR1 auxin receptor herbicides and to herbicides having alternative mechanisms of action using greenhouse bioassays and genetic markers. Atrazine, imazethapyr, lactofen, mesotrione, glufosinate, and glyphosate were applied in a single-dose bioassay, and tissue was collected from marked plants for genetic analysis. The FS population was not injured by atrazine or by imazethapyr. Approximately 50% of the plants survived lactofen and were actively growing 28 d after treatment. The population was susceptible to mesotrione, glufosinate, and glyphosate. Ametryn, chlorimuron-ethyl, 2,4-D, aminocyclopyraclor, aminopyralid, and picloram were applied in dose–response studies. The FS population was sensitive to ametryn, and the Ser-264-Gly substitution in the D1 protein was not detected, suggesting the lack of response to atrazine is not due to a target-site mutation. The FS population exhibited less than 50% injury to chlorimuron-ethyl at application rates 20 times the labeled use rate. The Ser-653-Asn acetolactate synthase (ALS) substitution, which confers resistance to imidazolinone herbicides, was present in the FS population. However, this does not explain the lack of response to the sulfonylurea herbicide, chlorimuron-ethyl. Sequencing of a portion of thePPX2Lgene did not show the ΔG210 mutation that confers resistance to protoporphyrinogen oxidase–inhibiting herbicides, suggesting that other factors were responsible for waterhemp survival after lactofen application. The FS population was confirmed to be at least 30-fold resistant to 2,4-D relative to the susceptible populations. In addition, it was at least 3-fold less sensitive to aminopyralid and picloram, two other TIR1 auxin receptor herbicides, than the 2,4-D-susceptible populations were. These data indicated that the FS population contains both target and non–target site mechanisms conferring resistance to herbicides spanning at least three mechanisms of action: TIR1 auxin receptors, ALS inhibitors, and photosystem II inhibitors.

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