Amaranthus hybridus populations resistant to triazine and acetolactate synthase-inhibiting herbicides

Amaranthus hybridus L. populations (A, B and C) obtained from escapes in Massac County and Pope County fields in southern Illinois, USA were subjected to greenhouse and laboratory experiments to measure multiple resistance to triazine and acetolactate synthase (ALS)‐inhibiting herbicides and cross‐resistance between sulfonylurea and imidazolinone herbicides. Phytotoxicity responses of the three populations revealed that only population B exhibited multiple resistances to triazine and ALS‐inhibiting herbicides. This population was >167‐, >152‐ and >189‐fold resistant to atrazine, imazamox and thifensulfuron, respectively, at the whole plant level compared with the susceptible population. Population A was only resistant to triazines and population C was only resistant to ALS‐inhibiting herbicides. Results from in vivo ALS enzyme and chlorophyll fluorescence assays confirmed these findings and indicated that an altered site‐of‐action mediated resistance to both triazine and ALS‐inhibiting herbicides. Gene sequencing revealed that a glycine for serine substitution at residue 264 of the D1 protein, and a leucine for tryptophan substitution at residue 574 of ALS were the causes of resistance for the three populations.     

Triazine herbicide resistance in Chenopodium album L.: Occurrence and characteristics of an intermediate biotype

Intermediate (I) biotypes for triazine herbicide resistance in Chenopodium album (as defined by a peculiar fluorescence curve), had the same ID₅₀ values as resistant(R) plants for chloroplast response to atrazine, but proved to be more susceptible at lower doses. Furthermore, the lethal dose in seedling treatments was lower than that of the R plants, but six times higher than for susceptible (S) plants. These I characteristics of I biotypes were maternally inherited in crosses. I biotypes were isolated from various progenies of susceptible precursor (Sp) plants in two garden populations. This could be the first step in the occurrence of triazine herbicide resistance. However, Sp plants have not been observed in field populations. The significance of the presence of a single isozyme pattern for all Sp plants is discussed. The results suggest an evolutionary pathway from S to R plants via I biotypes.     

Resistance of barnyardgrass (Echinochloa crus-galli) to atrazine and quinclorac

Two populations of Echinochloa crus-galli (R and I) exhibited resistance to quinclorac. Another population (X) exhibited resistance to quinclorac and atrazine. The R and I populations were collected from monocultures of rice in southern Spain. The X population was collected from maize fields subjected to the application of atrazine over several years. The susceptible (S) population of the same genus was collected from locations which had never been treated with herbicides. The quinclorac ED₅₀ value (dose causing 50% reduction in shoot fresh weight) for the R and I biotypes were 26- and 6-fold greater than for the S biotype. The X biotype was 10 times more tolerant to quinclorac than the S biotype and also showed cross-resistance to atrazine, being 82-fold more resistant to atrazine than the R, I and S biotypes. Chlorophyll fluorescence and Hill reaction analysis supported the view that the mechanism of resistance to atrazine in the X biotype was modification of the target site, the DI protein. Quinclorac at 20 mg litre^-1 did not inhibit photosynthetic electron transport in any of the test biotypes. The quinclorac I₅₀ values (herbicide dose needed for 50% Hill reaction reduction) of the S population was over 50000-fold higher than the atrazine I₅₀ value for the same S population, indicating that quinclorac is not a PS II inhibiting herbicide. Propanil at doses greater than 0·5 kg ha^-1 controlled all the biotypes. © 1997 SCI     

靶标基因突变导致入侵性杂草长芒苋对咪唑乙烟酸产生抗性

长芒苋Amaranthus palmeri生长迅速,适应性广,繁殖系数高,具有很强的竞争性,已在我国多地定植,对作物产量及生态环境构成潜在威胁.一旦其对除草剂产生抗性,将大大增加治理难度.本试验研究了采自不同地点的长芒苋种群对除草剂咪唑乙烟酸的抗性水平和抗性机理.整株生物测定得出,长芒苋疑似抗性种群和敏感种群对咪唑乙烟...     

Identification of a target‐site mutation conferring resistance to triazine herbicides in oriental mustard (Sisymbrium orientale L.) from Australia

In southern Australia, oriental mustard (Sisymbrium orientale) has been controlled successfully by triazine herbicides for several decades. The screening of 40 populations that were collected from the southern grain belt of Australia during 2010 and 2013 for resistance to six different herbicides (glyphosate, diflufenican, imazamox, chlorsulfuron, atrazine and 2,4‐dichlorophenoxyacetic acid) identified two oriental mustard populations as highly resistant to atrazine. Compared to the known oriental mustard‐susceptible populations (S1 and S2), these two resistant populations (P17 and P18) from near Horsham, Victoria, Australia, were 311‐ and 315‐fold resistant to atrazine, as determined by a comparison of the LD₅₀ values. However, there was no resistance to diuron detected in these populations. Sequencing of the chloroplast psbA gene identified a missense mutation of serine 264 to glycine in both herbicide‐resistant oriental mustard populations, which is known to confer high‐level atrazine resistance in other species.     

Increased herbicidal activity of triazine herbicides by piperonyl butoxide

Piperonyl butoxide (PB) is a known Synergist which enhances the activity of insecticides by inhibiting their biotransformation to less active products. We have evaluated the possible use of PB as a herbicide synergist using triazine herbicides in sensitive, tolerant, and resistant plants. The effects of PB, triazine herbicides, and their combinations were examined in whole plants as well as in chloroplasts isolated from triazine-sensitive (S) and -resistant (R) weed biotypes. PB itself, applied postemergence (0.1–0.5%, v/v), was slightly toxic to the plants tested. However, foliar application of PB combined with atrazine, terbutryn or prometryn to maize seedlings significantly increased the phytotoxicity of the herbicides. Low rates of atrazine, prometryn, and terbutryn in a tank-mixture with PB, effectively controlled Solatium nigrum L. and Abutilon theophrasli Medik. PB enhanced atrazine efficacy in both S and R biotypes of Lolium rigidum Gaud. The synergistic effect of PB was evident also in vitro when atrazine and methabenzthiazuron were used to inhibit photosystem II electron transport in chloroplasts isolated from resistant weeds. These data demonstrate the potential of PB as a herbicide synergist and its possible utilization as an aid for improving the activity of triazine herbicides in sensitive, tolerant and resistant plants.     

Differential translocation of paraquat in paraquat-resistant populations of Hordeum leporinum

Two populations of Hordeum leporinum have evolved resistance to paraquat within a small area in central Tasmania, Australia. One population (THL1) was more than 80-fold resistant to paraquat when treated in winter, compared with a susceptible population (THL4) collected nearby, whereas the other population (THL2) was only 19-fold resistant. Translocation of paraquat was examined in all three populations at warm and cool temperature regimes. Herbicide was applied to a basal section of the second leaf of plants kept in the dark and translocation measured after 16 h of dark and during a subsequent light period. Paraquat absorption into the treated leaf was uniformly high in susceptible and resistant populations, with >93% of the applied herbicide absorbed within 16 h in the dark at both temperatures. Translocation of paraquat out of the treated leaf was low in the dark, with <4% of the herbicide translocated to the remainder of the plant. More herbicide was translocated out of the treated leaves in susceptible plants in the dark, compared with resistant plants at both temperature regimes and more paraquat was translocated at warmer temperatures. Extensive basipetal translocation of paraquat to the rest of the plant occurred in susceptible plants following exposure of the treated plants to light. However, basipetal translocation was much reduced in resistant plants in the light and corresponded to the degree of resistance. Resistance to paraquat in H. leporinum is the result of reduced translocation of paraquat out of the treated leaves.     

Differential response of Amaranthus tuberculatus (Moq ex DC) JD Sauer to glyphosate

Midwest USA farmers have reported inconsistent control of Amaranthus tuberculatus (= rudis) (Moq ex DC) JD Sauer by glyphosate in glyphosate-resistant crops. The potential of selection for decreased A tuberculatus sensitivity to glyphosate was therefore investigated in a reportedly resistant Everly, IA population (P0-EV). Evaluation of six A tuberculatus populations from the Midwest USA estimated a seedling baseline sensitivity of 2.15 mM glyphosate. Based on these results, three generations of divergent recurrent selection were implemented on P0-EV to isolate resistant and susceptible populations. A seedling assay was developed to screen large amounts of seeds and thus expedite the selection process. Whole-plant and seedling rate responses of P0-EV and a known pristine A tuberculatus population from Paint Creek, OH (P0-WT) identified no significant difference in response to glyphosate; however, greater phenotypic variance was ostensibly evident in P0-EV. The first recurrent generation selected for resistance at 3.2 mM glyphosate (RS1-R) had a 5.9- and 1.7-fold resistance increase at the seedling and whole-plant levels, respectively, compared with the susceptible generation selected at 32 microM glyphosate. After three cycles of recurrent selection, 14.6-fold difference in resistance at the seedling level and 3.1-fold difference at the whole-plant level were observed when comparing the populations selected for resistance (RS3-R) and susceptibility (RS3-S). Overall, recurrent selection increased the frequency of resistant individuals and decreased the variability to glyphosate at the population level. Nevertheless, variability for glyphosate resistance was still evident in RS3-R. Results herein suggested that A tuberculatus is inherently variable to glyphosate and that selection decreased the sensitivity to glyphosate. We purport that evolved glyphosate resistance in A tuberculatus may require multiple cycles of selection under field conditions. Historic estimated use of glyphosate alludes to the evolution of tolerant weed populations.     

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.     

Fitness effects of the triazine resistance mutation in Amaranthus hybridus: relative fitness in maize and soyabean crops

Fitness costs of the triazine resistance mutation were estimated in experimental populations of Amaranthus hybridus L. from Maryland and Virginia (USA). Estimated costs were attributed directly to effects of the resistance mutation, through comparisons of lines bearing similar nuclear genomes, but either resistant or susceptible cytoplasm. Experimental weed plants were established from seed in the field at a wide range of A. hybridus and crop (maize or soyabean) densities, and experiments were repeated over 3 years. Resistant cytoplasm caused large reductions in several components of fitness, but substantial differences were observed among years in these fitness costs of resistance. For the fitness component of total biomass above-ground biomass production, relative fitnesses of resistant genotypes (resistant/susceptible ratios) ranged between 0.42 and 0.70. Costs of resistance were similar in weeds growing with either maize or soyabean crops. Certain costs of resistance differed between populations: impairment of seedling establishment and early growth by the resistance mutation was evident in the Virginia population, but not in the Maryland population. This functional difference between populations was associated with greater costs of resistance in the Virginia population with respect to mid-season leaf number and total above-ground biomass production; however, only the former population difference in fitness cost was statistically significant.     

Resistance of Amaranthus hybridus to atrazine

Resistance of weeds to triazine herbicides has been recorded in many countries. The extent of the problem in South Africa is uncertain. In a pilot study, the atrazine resistance of Amaranthus hybridus L. (smooth pigweed) was investigated. Suspected resistant (R) and susceptible (S) biotypes were treated with commercially formulated atrazine. After post-and pre-emergence applications under tunnel conditions, it was found that the suspected R biotype plants were not affected at herbicide dosages of belween 1.25 and 25.0 kg a.i. ha^-1, i.e. up to 20 times gieater than the lowest recommended dosage. However, the S biotype plants were killed by the lowest dosage. In the fieid, mortalities in the R biolype were not observed after post-emergence applications of 1.25-10.00 kg a.i. ha^-1. In contrast, all S biotype plants were killed. In tunnel experiments, the R biotype was also found to be resistant to cyanazine and cyanazine+atrazine, while slight tolerance to linuron was observed. All these treatments resulted in 100% mortality of the S biotype. Although S biotype seeds oi A. hybridus were found to germinate slightly sooner under controlled conditions than R biotype seeds, preliminary results suggest that there are no major differences. Indications are that, although the growth of the S biotype may be greater than that of the R biotype, the competitive effect of the two biotypes on crop seedlings may well be similar.     

psbA mutation (Asn266 to Thr) in Senecio vulgaris L. confers resistance to several PS II-inhibiting herbicides

DNA sequence analysis of the psbA gene encoding the D1 protein of photosystem II (PS II), the target site of PS II-inhibiting herbicides, identified a point mutation (Asn266 to Thr) in a bromoxynil-resistant Senecio vulgaris L. population collected from peppermint fields in Oregon. Although this mutation has been previously reported in Synechocystis, this is the first report of this particular point mutation in a higher plant exhibiting resistance to PS II-inhibiting herbicides. The resistant population displayed high-level resistance to bromoxynil and terbacil (R/S ratio 10.1 and 9.3, respectively) and low-level resistance to metribuzin and hexazinone (R/S ratio 4.2 and 2.6, respectively) when compared with the susceptible population. However, the population was not resistant to the triazine herbicides atrazine and simazine or to the urea herbicide diuron. A chlorophyll fluorescence assay confirmed the resistance levels and patterns of cross-resistance of the whole-plant studies. The resistant S. vulgaris plants produced fewer seeds. Differences in cross-resistance patterns to PS II-inhibiting herbicides and the difference in fitness cost could be exploited in a weed management program.     

Status of black grass (Alopecurus myosuroides) resistance to acetyl-coenzyme A carboxylase inhibitors in France

We assessed the contributions of target site‐ and non‐target site‐based resistance to herbicides inhibiting acetyl‐coenzyme A carboxylase (ACC) in Alopecurus myosuroides (black grass). A total of 243 A. myosuroides populations collected across France were analysed using herbicide sensitivity bioassay (24 300 seedlings analysed) and ACC genotyping (13 188 seedlings analysed). Seedlings resistant to at least one ACC‐inhibiting herbicide were detected in 99.2% of the populations. Mutant, resistant ACC allele(s) were detected in 56.8% of the populations. Among the five resistant ACC alleles known in A. myosuroides, alleles containing an isoleucine‐to‐leucine substitution at codon 1781 were predominant (59.5% of the plants containing resistant ACC alleles). Comparison of the results from herbicide sensitivity bioassays with genotyping indicated that more than 75% of the plants resistant to ACC‐inhibiting herbicides in France would be resistant via increased herbicide metabolism. Analysis of herbicide application records suggested that in 15.9% of the populations studied, metabolism‐based resistance to ACC‐inhibiting herbicides was mostly selected for by herbicides with other modes of action. Our study revealed the importance of non‐target site‐based resistance in A. myosuroides. Using herbicides with alternative modes of action to control populations resistant to ACC‐inhibiting herbicides, the recommended management approach, may thus be jeopardised by the widespread occurrence of metabolism‐based resistance mechanisms conferring broad‐spectrum cross‐resistance.     

Mutations of the ALS gene endowing resistance to ALS-inhibiting herbicides in Lolium rigidum populations

BACKGROUND: In the important grass weed Lolium rigidum (Gaud.), resistance to ALS‐inhibiting herbicides has evolved widely in Australia. The authors have previously characterised the biochemical basis of ALS herbicide resistance in a number of L. rigidum biotypes and established that resistance can be due to a resistant ALS and/or enhanced herbicide metabolism. The purpose of this study was to identify specific resistance‐endowing ALS gene mutation(s) in four resistant populations and to develop PCR‐based molecular markers. RESULTS: Six resistance‐conferring ALS mutations were identified: Pro‐197‐Ala, Pro‐197‐Arg, Pro‐197‐Gln, Pro‐197‐Leu, Pro‐197‐Ser and Trp‐574‐Leu. All six mutations were found in one population (WLR1). Each Pro‐197 mutation conferred resistance to the sulfonylurea (SU) herbicide sulfometuron, whereas the Trp‐574‐Leu mutation conferred resistance to both sulfometuron and the imidazolinone (IMS) herbicide imazapyr. A derived cleaved amplified polymorphic sequences (dCAPS) marker was developed for detecting resistance mutations at Pro‐197. Furthermore, cleaved amplified polymorphic sequences (CAPS) markers were developed for detecting each of the six mutant resistant alleles. Using these markers, the authors revealed diverse ALS‐resistant alleles and genotypes in these populations and related them directly to phenotypic resistance to ALS‐inhibiting herbicides. CONCLUSION: This study established the existence of a diversity of ALS gene mutations endowing resistance in L. rigidum populations: 1–6 different mutations were found within single populations. At field herbicide rates, resistance profiles were determined more by the specific mutation than by whether plants were homo‐ or heterozygous for the mutation. Copyright © 2008 Society of Chemical Industry     

Activity of mesotrione on resistant weeds in maize

Mesotrione is a new callistemone herbicide that inhibits the HPPD enzyme (p-hydroxyphenylpyruvate dioxygenase) and introduces a new naturally selective tool into weed-management programmes for use in maize. Mesotrione provides control of the major broad-leaved weeds, and it can be used in integrated weed-management programmes depending on the grower's preferred weed-control strategy. At post-emergence rates of 150 g AI ha-1 or less, mesotrione provides naturally selective control of key species that may show triazine resistance (TR), e.g. Chenopodium album L, Amaranthus species, Solanum nigrum L, as well as species of weed that show resistance to acetolactase synthase (ALS) inhibitors e.g. Xanthium strumarium L, Amaranthus spp and Sonchus spp. The data presented show that resistant and susceptible biotypes of these species with resistance to triazine herbicides, such as atrazine, simazine, terbutylazine and metribuzin, or ALS-inhibitor herbicides, such as imazethepyr, remain susceptible to mesotrione. These results confirm that there is no cross-resistance in biotypes with target site resistance to triazine or ALS-inhibiting herbicides. It is important that herbicide choice and rotation becomes an integral part of planning weed management, so as to minimise the risks of crop losses from weed competition, build-up of weed seed in the soil and the further development of weed resistance across a range of herbicide modes of action.     

Characterisation of a triazine-resistant biotype of Bromus tectorum found in Spain

A population of Bromus tectorum infesting an olive grove at Córdoba (Spain) survived simazine use rates of 3.0 kg a.i. ha⁻¹ over two consecutive years. Non‐tillage olive monoculture and two annual simazine applications had been carried out for 10 years. The resistant biotype showed a higher ED₅₀ value (7.3 kg a.i. ha⁻¹) than that of the susceptible control (0.1 kg a.i. ha⁻¹), a 73‐fold increase in herbicide tolerance. The use of fluorescence, Hill reaction, absorption, translocation and metabolism assays showed that simazine resistance in this biotype was caused by a modification of the herbicide target site, since chloroplasts from the resistant biotype of B. tectorum were more than 300 times less sensitive to simazine than those from the susceptible biotype. In addition, non‐treated resistant plants of B. tectorum displayed a significant reduction in the Q_A to Q_B electron transfer rate when compared with the susceptible biotype, a characteristic that has been linked to several mutations in the protein D1 conferring resistance to PS II inhibiting herbicides. Resistant plants showed cross‐resistance to other groups of triazine herbicides with the hierarchy of resistance level being methoxy‐s‐triazines ≥chloro‐s‐triazines > methylthio‐s‐triazines > cis‐triazines. The results indicate a naturally occurring target‐site point mutation is responsible for conferring resistance to triazine herbicides. This represents the first documented report of target site triazine resistance in this downy brome biotype.     

A molecular genetic assessment of herbicide-resistant Sinapis arvensis

The acquisition of resistance to both the auxinic herbicide dicamba and the sulfonylurea herbicide chlorsulfuron has been recorded in Canadian populations of the weed species Sinapis arvensis L. (charlock, wild mustard) To study the effect of this selection for herbicide resistance on levels of genetic variation, polymerase chain reaction-based DNA fingerprinting techniques were used to characterize two herbicide-resistant and one susceptible population of S. arvensis . Analysis of the resultant DNA marker profiles revealed extensive polymorphism between individuals. However, segregation of the three biotypes was detectable despite high levels of intrabiotype polymorphism. No reduction in the levels of heterozygosity within the resistant populations were found compared with the susceptible population.     

Two-year Investigations on Herbicide-Resistant Silky Bent Grass (Apera spica-venti L. Beauv.) Populations in Winter Wheat��Population Dynamics, Yield Losses, Control Efficacy and Introgression into Sensitive Population

In this study, the results of two-year investigations on herbicide resistance in silky bent grass (Apera spica-venti) populations are presented. Two populations of A. spica-venti were sown in a winter wheat field at the Ihinger Hof Research Station near Stuttgart in Germany in October 2008. Whole-plant bioassays conducted with both populations in the greenhouse before the field trial was set up revealed that population A was strongly resistant to acetolactate synthase (ALS)-inhibitors, whereas population B was sensitive to this group of herbicides. Each block was treated with isoproturon, fenoxaprop-P-ethyl, sulfosulfuron and meso-iodosulfuron at the recommended field dose. One treatment remained unsprayed. It was found that the average number of silky bent grass panicles was significantly higher in the second year (2010) with 343 panicles m^{? 2} in the untreated control compared to 44 panicles m^{? 2} in the first year (2009). Efficacy of both ALS-inhibitors was significantly reduced in the resistant population compared to the sensitive population. Grass-weed plants surviving treatments with ALS-inhibitors produced the same number of seeds as the untreated plants. It was found that germination rate of seeds from the resistant population was more than threefold higher than from the sensitive population. Grain yield was equal in all treatments and populations in 2009. In 2010, applications of isoproturon and fenoxaprop-P-ethyl resulted in higher grain yields, with a mean of 4.6 t ha^{? 1} compared to an average of 3.9 t ha^{? 1} in the plots treated with sulfosulfuron and meso-iodosulfuron and 4.3 t ha^{? 1} in the untreated plots. However, these differences were not statistically significant. The sensitive population of the second generation (2009/2010) was approximately 20% more tolerant to ALS-inhibitors than the sensitive population of the first generation (2008/2009) which indicates introgression of herbicide resistance traits already after one year. These results clearly show that herbicide resistance to ALS-inhibitors in silky bent grass is likely to spread rapidly causing significant economic losses. Therefore, management strategies need to be developed and tested to prevent and overcome herbicide resistance in European cereal production systems.     

Molecular characterisation of resistance to ALS-inhibiting herbicides in Hordeum leporinum biotypes

BACKGROUND: The acetolactate synthase (ALS)-inhibiting herbicide sulfosulfuron is registered in Australia for the selective control of Hordeum leporinum Link. in wheat crops. This herbicide failed to control H. leporinum on two farms in Western Australia on its first use. This study aimed to determine the level of resistance of three H. leporinum biotypes, identify the biochemical and molecular basis and develop molecular markers for diagnostic analysis of the resistance. RESULTS: Dose-response studies revealed very high level (>340-fold) resistance to the sulfonylurea herbicides sulfosulfuron and sulfometuron. In vitro ALS assays revealed that resistance was due to reduced sensitivity of the ALS enzyme to herbicide inhibition. This altered ALS sensitivity in the resistant biotypes was found to be due to a mutation in the ALS gene resulting in amino acid proline to serine substitution at position 197. In addition, two- to threefold higher ALS activities were consistently found in the resistant biotypes, compared with the known susceptible biotype. Two cleaved amplified polymorphic sequence (CAPS) markers were developed for diagnostic testing of the resistant populations. CONCLUSION: This study established the first documented case of evolved ALS inhibitor resistance in H. leporinum and revealed that the molecular basis of resistance is due to a Pro to Ser mutation in the ALS gene.     

An association between triazine resistance and powdery mildew resistance in Epilobium ciliatum and Senecio vulgaris

The response of four naturally occurring biotypes of Epilobium ciliatum and four sources of Senecio vulgaris to the herbicide atrazine were compared with their susceptibility to the powdery mildews Sphaerotheca epilobii and Erysiphe cichoracearum. Biotypes that were resistant to atrazine were also resistant to mildew. Mechanisms that might explain the association between atrazine resistance and mildew resistance are discussed, along with possible implications of these findings for farmland ecology and for the production of herbicide- and mildew-resistant crop plants.