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

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.

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Evolution of target and non-target based multiple herbicide resistance in a single Palmer amaranth (Amaranthus palmeri) population from Kansas

Weed Technology ◽

10.1017/wet.2020.32 ◽

2020 ◽

Vol 34 (3) ◽

pp. 447-453

Author(s):

Sushila Chaudhari ◽

Vijay K. Varanasi ◽

Sridevi Nakka ◽

Prasanta C. Bhowmik ◽

Curtis R. Thompson ◽

...

Keyword(s):

Resistance Mechanism ◽

Acetolactate Synthase ◽

Palmer Amaranth ◽

Target Site ◽

Epsp Synthase ◽

High Incidence ◽

Evolution Of Resistance ◽

Target Site Resistance ◽

Herbicide Atrazine ◽

Sites Of Action

AbstractThe evolution of resistance to multiple herbicides in Palmer amaranth is a major challenge for its management. In this study, a Palmer amaranth population from Hutchinson, Kansas (HMR), was characterized for resistance to inhibitors of photosystem II (PSII) (e.g., atrazine), acetolactate synthase (ALS) (e.g., chlorsulfuron), and EPSP synthase (EPSPS) (e.g., glyphosate), and this resistance was investigated. About 100 HMR plants were treated with field-recommended doses (1×) of atrazine, chlorsulfuron, and glyphosate, separately along with Hutchinson multiple-herbicide (atrazine, chlorsulfuron, and glyphosate)–susceptible (HMS) Palmer amaranth as control. The mechanism of resistance to these herbicides was investigated by sequencing or amplifying the psbA, ALS, and EPSPS genes, the molecular targets of atrazine, chlorsulfuron, and glyphosate, respectively. Fifty-two percent of plants survived a 1× (2,240 g ai ha−1) atrazine application with no known psbA gene mutation, indicating the predominance of a non–target site resistance mechanism to this herbicide. Forty-two percent of plants survived a 1× (18 g ai ha−1) dose of chlorsulfuron with proline197serine, proline197threonine, proline197alanine, and proline197asparagine, or tryptophan574leucine mutations in the ALS gene. About 40% of the plants survived a 1× (840 g ae ha−1) dose of glyphosate with no known mutations in the EPSPS gene. Quantitative PCR results revealed increased EPSPS copy number (50 to 140) as the mechanism of glyphosate resistance in the survivors. The most important finding of this study was the evolution of resistance to at least two sites of action (SOAs) (~50% of plants) and to all three herbicides due to target site as well as non–target site mechanisms. The high incidence of individual plants with resistance to multiple SOAs poses a challenge for effective management of this weed.

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

Weed Science ◽

10.1017/wsc.2019.1 ◽

2019 ◽

Vol 67 (3) ◽

pp. 273-280 ◽

Cited By ~ 4

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.

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Molecular confirmation of resistance to PPO inhibitors in Amaranthus tuberculatus and Amaranthus palmeri, and isolation of the G399A PPO2 substitution in A. palmeri

Weed Technology ◽

10.1017/wet.2020.86 ◽

2020 ◽

pp. 1-7

Author(s):

Jacob S. Montgomery ◽

Darci A. Giacomini ◽

Patrick J. Tranel

Keyword(s):

Goodness Of Fit ◽

Resistance Mechanism ◽

Amino Acid Position ◽

Resistance Mechanisms ◽

Palmer Amaranth ◽

Target Site ◽

Coding Region ◽

Common Waterhemp ◽

Growing Seasons ◽

Target Site Resistance

Abstract During the 2017 to 2019 growing seasons, samples of waterhemp and Palmer amaranth that had reportedly survived field-rate applications of protoporphyrinogen oxidase (PPO)–inhibiting herbicides were collected from the American Midwest and tested for target-site mutations known at the time to confer resistance. Target-site resistance was identified in nearly all (135 of 145) tested common waterhemp populations but in only 8 of 13 Palmer amaranth populations. Follow-up research on one population of Palmer amaranth (W-8), which tested negative for all such mutations, confirmed it was resistant to lactofen, with a magnitude of resistance comparable to that conferred by the ΔG210 PPO2 mutation. Gene sequences from both isoforms of PPO (PPO1 and PPO2) were compared between W-8 and known PPO inhibitor–sensitive sequence. A glycine-to-alanine substitution at the 399th amino acid position (G399A) of PPO2, recently identified to reduce target-site herbicide sensitivity, was observed in a subset of resistant W-8 plants. Because no missense mutation completely delimited resistant and sensitive sequences, we initially suspected the presence of a secondary, non-target-site resistance mechanism in this population. To isolate G399A, a segregating F2 population was produced and screened with a delimiting rate of lactofen. χ2 goodness-of-fit analysis of dead/alive ratings indicated single-locus inheritance of resistance in the F2 population, and molecular markers for the W-8 parental PPO2 coding region co-segregated tightly, but not perfectly, with resistance. More research is needed to fully characterize Palmer amaranth PPO inhibitor–resistance mechanisms, which appear to be more diverse than those found in common waterhemp.

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

Weed Science ◽

10.1017/wsc.2019.14 ◽

2019 ◽

Vol 67 (4) ◽

pp. 380-388 ◽

Cited By ~ 2

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.

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

Weed Technology ◽

10.1017/wet.2017.13 ◽

2017 ◽

Vol 31 (3) ◽

pp. 470-476 ◽

Cited By ~ 3

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.

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Distribution and validation of genotypic and phenotypic glyphosate and PPO-nhibitor resistance in Palmer amaranth (Amaranthus palmeri) from southwestern Nebraska

Weed Technology ◽

10.1017/wet.2020.74 ◽

2020 ◽

pp. 1-12 ◽

Cited By ~ 1

Author(s):

Maxwel C Oliveira ◽

Darci A Giacomini ◽

Nikola Arsenijevic ◽

Gustavo Vieira ◽

Patrick J Tranel ◽

...

Keyword(s):

Cluster Analysis ◽

Gene Amplification ◽

Cropping Systems ◽

Resistance Mechanism ◽

Geographic Area ◽

Glyphosate Resistance ◽

Resistance Mechanisms ◽

Palmer Amaranth ◽

Resistance Evolution ◽

Inhibitor Resistance

Abstract Failure to control Palmer amaranth with glyphosate and protoporphyrinogen IX oxidase (PPO)-inhibitor herbicides was reported across southwestern Nebraska in 2017. The objectives of this study were to 1) confirm and 2) validate glyphosate and PPO-inhibitor (fomesafen and lactofen) resistance in 51 Palmer amaranth accessions from southwestern Nebraska using genotypic and whole-plant phenotypic assay correlations and cluster analysis, and 3) determine which agronomic practices might be influencing glyphosate resistance in Palmer amaranth accessions in that location. Based on genotypic assay, 88% of 51 accessions contained at least one individual with amplification (>2 copies) of the 5-enolypyruvyl-shikimate-3-phosphate synthase (EPSPS) gene, which confers glyphosate resistance; and/or a mutation in the PPX2 gene, either ΔG210 or R128G, which endows PPO-inhibitor resistance in Palmer amaranth. Cluster analysis and high correlation (0.83) between genotypic and phenotypic assays demonstrated that EPSPS gene amplification is the main glyphosate resistance mechanism in Palmer amaranth accessions from southwestern Nebraska. In contrast, there was poor association between genotypic and phenotypic responses for PPO-inhibitor resistance, which was attributed to segregation for PPO-inhibitor resistance within these accessions and/or the methodology that was adopted herein. Genotypic assays can expedite the process of confirming known glyphosate and PPO-inhibitor resistance mechanisms in Palmer amaranth from southwestern Nebraska and other locations. Phenotypic assays are also a robust method for confirming glyphosate resistance but not necessarily PPO-inhibitor resistance in Palmer amaranth. Moreover, random forest analysis of glyphosate resistance in Palmer amaranth indicated that EPSPS gene amplification, county, and current and previous crops are the main factors influencing glyphosate resistance within that geographic area. Most glyphosate-susceptible Palmer amaranth accessions were found in a few counties in areas with high crop diversity. Results presented here confirm the spread of glyphosate resistance and PPO-inhibitor resistance in Palmer amaranth accessions from southwestern Nebraska and demonstrate that less diverse cropping systems are an important driver of herbicide resistance evolution in Palmer amaranth.

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

Agronomy ◽

10.3390/agronomy10040471 ◽

2020 ◽

Vol 10 (4) ◽

pp. 471 ◽

Cited By ~ 2

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.

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Point Mutations as Main Resistance Mechanism Together With P450-Based Metabolism Confer Broad Resistance to Different ALS-Inhibiting Herbicides in Glebionis coronaria From Tunisia

Frontiers in Plant Science ◽

10.3389/fpls.2021.626702 ◽

2021 ◽

Vol 12 ◽

Author(s):

Zeineb Hada ◽

Yosra Menchari ◽

Antonia M. Rojano-Delgado ◽

Joel Torra ◽

Julio Menéndez ◽

...

Keyword(s):

Enzyme Activity ◽

Dose Response ◽

Resistance Mechanism ◽

Point Mutations ◽

Acetolactate Synthase ◽

Target Site ◽

Cross Resistance ◽

P450 Monooxygenase ◽

Target Site Resistance ◽

Sites Of Action

Resistance to acetolactate synthase (ALS) inhibiting herbicides has recently been reported in Glebionis coronaria from wheat fields in northern Tunisia, where the weed is widespread. However, potential resistance mechanisms conferring resistance in these populations are unknown. The aim of this research was to study target-site resistance (TSR) and non-target-site resistance (NTSR) mechanisms present in two putative resistant (R) populations. Dose–response experiments, ALS enzyme activity assays, ALS gene sequencing, absorption and translocation experiments with radiolabeled herbicides, and metabolism experiments were carried out for this purpose. Whole plant trials confirmed high resistance levels to tribenuron and cross-resistance to florasulam and imazamox. ALS enzyme activity further confirmed cross-resistance to these three herbicides and also to bispyribac, but not to flucarbazone. Sequence analysis revealed the presence of amino acid substitutions in positions 197, 376, and 574 of the target enzyme. Among the NTSR mechanisms investigated, absorption or translocation did not contribute to resistance, while evidences of the presence of enhanced metabolism were provided. A pretreatment with the cytochrome P450 monooxygenase (P450) inhibitor malathion partially synergized with imazamox in post-emergence but not with tribenuron in dose–response experiments. Additionally, an imazamox hydroxyl metabolite was detected in both R populations in metabolism experiments, which disappeared with the pretreatment with malathion. This study confirms the evolution of cross-resistance to ALS inhibiting herbicides in G. coronaria from Tunisia through TSR and NTSR mechanisms. The presence of enhanced metabolism involving P450 is threatening the chemical management of this weed in Tunisian wheat fields, since it might confer cross-resistance to other sites of action.

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Confirmation and Characterization of Non–target site Resistance to Fomesafen in Palmer amaranth (Amaranthus palmeri)

Weed Science ◽

10.1017/wsc.2018.60 ◽

2018 ◽

Vol 66 (6) ◽

pp. 702-709 ◽

Cited By ~ 23

Author(s):

Vijay K. Varanasi ◽

Chad Brabham ◽

Jason K. Norsworthy

Keyword(s):

Cytochrome P450 ◽

Cropping Systems ◽

Palmer Amaranth ◽

Target Site ◽

Amaranthus Palmeri ◽

Allelic Discrimination Assay ◽

Allelic Discrimination ◽

First Case ◽

Target Site Resistance ◽

Novel Target

AbstractPalmer amaranth (Amaranthus palmeri S. Watson), a dioecious summer annual species, is one of the most troublesome weeds in U.S. cropping systems. The evolution of resistance to protoporphyrinogen oxidase inhibitors in A. palmeri biotypes is a major cause of concern to soybean [Glycine max (L.) Merr.] and cotton (Gossypium hirsutum L.) growers in the midsouthern United States. The objective of this study was to confirm and characterize the non–target site mechanism in a fomesafen-resistant accession from Randolph County, AR (RCA). A dose–response assay was conducted to assess the level of fomesafen resistance, and based on the GR50 values, the RCA accession was 18-fold more resistant to fomesafen than a susceptible (S) biotype. A TaqMan allelic discrimination assay and sequencing of the target-site genes PPX2 and PPX1 revealed no known or novel target-site mutations. An SYBR Green assay indicated no difference in PPX2 gene expression between the RCA and S biotypes. To test whether fomesafen resistance is metabolic in nature, the RCA and the S biotypes were treated with different cytochrome P450 (amitrole, piperonyl butoxide [PBO], malathion) and glutathione S-transferase (GST) (4-chloro-7-nitrobenzofurazan [NBD-Cl]) inhibitors, either alone or in combination with fomesafen. Malathion followed by (fb) fomesafen in RCA showed the greatest reduction in survival (67%) and biomass (86%) compared with fomesafen alone (45% and 66%, respectively) at 2 wk after treatment. Interestingly, NBD-Cl fb fomesafen also resulted in low survival (35%) compared with the fomesafen-only treatment (55%). Applications of malathion or NBD-Cl preceding fomesafen treatment resulted in reversal of fomesafen resistance, indicating the existence of cytochrome P450– and GST-based non–target site mechanisms in the RCA accession. This study confirms the first case of non–target site resistance to fomesafen in A. palmeri.

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

Weed Science ◽

10.1614/ws-d-16-00090.1 ◽

2016 ◽

Vol 65 (1) ◽

pp. 4-8 ◽

Cited By ~ 4

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.

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