Fitness costs associated with acetyl‐coenzyme A carboxylase mutations endowing herbicide resistance in American sloughgrass (Beckmannia syzigachne Steud.)

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高温对抗性和敏感小菜蛾生命表参数及后代杀虫剂敏感性的影响

为研究高温对抗性和敏感小菜蛾Plutella xylostella生物学适合度及抗药性进化的影响,本研究测定了温度对杀虫剂抗性和敏感小菜蛾实验种群生命表参数及后代对阿维菌素敏感性的影响。 结果表明：与25℃相比,33.5℃条件下抗性和敏感种群的平均世代历期（D）、净生殖率（R₀）、内禀增长率（r_m）、周限增长率（λ）值、总产卵量、雌蛾平均寿命、日产卵量和孵化率均显著下降。 在25℃下,抗性和敏感种群的r_m,λ,D和日产卵量无显著差异,但抗性种群的R₀略低于敏感种群（为敏感种群的87%）。 在33.5℃下,抗性种群的r_m和R₀显著低于敏感种群（分别为敏感种群的75%和64%）,抗性种群雌成虫寿命、总产卵量和孵化率显著也低于敏感种群。 抗性种群在33.5℃下饲养一代后,后代对阿维菌素的抗性水平大幅下降,而相同处理下敏感种群对阿维菌素的抗性水平没有显著下降。 上述结果表明,无论在常温还是高温条件下,抗性种群生物学适合度均低于敏感种群,尤其是高温下抗性种群适合度远低于敏感种群。 抗性种群经高温饲养一代后对阿维菌素的抗性水平大幅下降可能与高温下抗性种群适合度显著较低有关。     

EMS诱变甘蓝型油菜M_2代群体的表型突变研究

利用4种浓度EMS处理甘蓝型油菜NJ7982种子,选取其中诱变效果好的处理(0.4%EMS),研究其后代突变型和突变频率。对4.8万株的M2代群体鉴定结果表明,6种器官性状在群体中均出现了突变。子叶突变性状包括3子叶、子叶黄化等,占群体的0.22%;叶片突变性状包括黄化叶、白化叶、紫色叶、上卷叶、下卷叶等,占0.74%;花器突变性状包括紫色花蕾、死蕾、3花瓣、6花瓣、白色瓣、花瓣黄白镶嵌、花瓣皱缩、完全不育、部分可育等,占9.38%;株型突变性状包括矮秆、紫茎等,占4.98%;角果突变性状包括粗角、长角、紫角等,占2.79%;种皮黄色,占0.40%;总的表型突变频率为18.51%。这些遗传多样性的突变材料,为甘蓝型油菜种质创新及品种遗传改良提供基础性材料。     

Phenotypic mechanisms of host resistance against greenbug (Homoptera: Aphididae) revealed by near isogenic lines of wheat

Interactions between biotype E greenbug, Schizaphis graminum (Rondani), and wheat, Triticum aestivum L., were investigated using resistant and susceptible near isogenic lines of the greenbug resistance gene Gb3. In an antixenosis test, the greenbugs preferred susceptible plants to resistant ones when free choice of hosts was allowed. Aphid feeding resulted in quick and severe damage to susceptible plants, which seemed to follow a general pattern spatially and was affected by the position where the greenbugs were initially placed. Symptom of damage in resistant plants resembled senescence. Within-plant distribution of aphids after infestation was clearly different between the two genotypes. Significantly more greenbugs fed on the first (oldest) leaf than on the stem in resistant plants, but this preference was reversed in the susceptible one. After reaching its peak, aphid population on the susceptible plants dropped quickly. All susceptible plants were dead in 10-14 d after infestation due to greenbug feeding. Aphid population dynamics on resistant plants exhibited a multipeak curve. After the first peak, the greenbug population declined slowly. More than 70% of resistant plants were killed 47 d after infestation. Performance of both biotype E and I greenbugs on several Gb3-related wheat germplasm lines were also examined. It seems that the preference-on-stem that was characteristic of biotype E greenbugs on the susceptible plants was aphid biotype- and host genotype-dependent. Results from this study suggested that antixenosis, antibiosis, and tolerance in the resistant plants of wheat might all contribute to resistance against greenbug feeding.     

Investigating the mechanisms of glyphosate resistance in <Emphasis Type="Italic">Lolium multiflorum</Emphasis>

Evolved resistance to the herbicide glyphosate has been reported in eleven weed species, including Lolium multiflorum. Two glyphosate-resistant L. multiflorum populations were collected, one from Chile (SF) and one from Oregon, USA (OR), and the mechanisms conferring glyphosate resistance were studied. Based on a Petri dish dose–response bioassay, the OR and the SF populations were two and fivefold more resistant to glyphosate when compared to the susceptible (S) population, respectively; however, based on a whole-plant dose–response bioassay, both OR and SF populations were fivefold more resistant to glyphosate than the S population, implying that different resistance mechanisms might be involved. The S population accumulated two and three times more shikimic acid in leaf tissue 96 h after glyphosate application than the resistant OR and SF populations, respectively. There were no differences between the S and the glyphosate-resistant OR and SF populations in ¹⁴C-glyphosate leaf uptake; however, the patterns of ¹⁴C-glyphosate translocation were significantly different. In the OR population, a greater percentage of ¹⁴C-glyphosate absorbed by the plant moved distal to the treated section and accumulated in the tip of the treated leaf. In contrast, in the S and in the SF populations, a greater percentage of ¹⁴C-glyphosate moved to non-treated leaves and the stem. cDNA sequence analysis of the EPSP synthase gene indicated that the glyphosate-resistant SF population has a proline 106 to serine amino acid substitution. Here, we report that glyphosate resistance in L. multiflorum is conferred by two different mechanisms, limited translocation (nontarget site-based) and mutation of the EPSP synthase gene (target site-based).     

Selection for Resistance to Leptosphaerulina Leaf Spot in Alfalfa

Significant differences were observed among 11 alfalfa cultivars for the percentage of plants resitant to Leptospbaerulina briosiana after one inoculation. Screening for resistance was done by inoculating 18 day old plants (cv. Europe), held in a moist chamber, with L. briosiana. The self- and cross-progenies of plants resistant to L. briosiana after two inoculations showed slightly higher resistance to Leptosphaerulina leaf spot than the unselected original population but the difference was significant for the cross-progeny only. Larger variations were observed among self-progenies than cross-progenies. No correlation was found between self - and cross-progenies. At least three inoculations may be necessary to eliminate most susceptible plants. Mas selection was as effective as the polycross progeny test to improve the level of resistance of the selected populations as compared with the unselected one but the difference was agronomically unimportant. Under field conditions, the plants selected as resistant after two inoculations in the laboratory showed better resistance to L. briosiana than the susceptible ones but only on the first of three notations. The population derived from plants selected from cv. Europe as resistant to L. briosiana in the laboratory was not significantly more resitant than cv. Europe in the filed but generally showed less foliar infection.     

Rearing history affects behaviour and performance of two virulent Nasonovia ribisnigri populations on two lettuce cultivars

Many aphid species have become virulent to host‐plant resistance, which limits the sustainability of insect resistance breeding. However, when this adaptation to resistant plants is associated with fitness costs for the aphids, virulence can be lost in the absence of resistant plants. For two populations of the lettuce aphid, Nasonovia ribisnigri (Mosely) (Hemiptera: Aphididae), we evaluated whether virulence to Nr‐gene‐based resistance was lost on a susceptible lettuce, Lactuca sativa L. (Asteraceae), and assessed possible costs of virulence. The feeding behaviour and performance of these aphids, reared and tested on susceptible and resistant lettuce, were investigated. The rearing plant affected feeding behaviour and performance of the aphids. Temporary reduction and long‐term loss of virulence were found. The total duration of phloem intake was shorter after being reared on susceptible lettuce and tested on resistant lettuce. In addition, one population had a lower survival on resistant lettuce after being reared on susceptible lettuce. There were also indications of fitness costs of the virulence in both populations.     

Microsatellite mutation directed by an external stimulus

Microsatellites are regions of DNA containing tandem repeats of a core 2-6 bp nucleotide sequence. To test the hypothesis that microsatellite mutation can be directed by exposure to specific external cues, control and treatment groups of resistant and susceptible wheat varieties were grown under controlled conditions and genotyped at a number of microsatellite loci that map to chromosomes known to contain Fusarium head blight (FHB) resistance/susceptibility loci. Genotyping was undertaken both prior to and following exposure to Fusarium graminearum, the FHB pathogen. Within a month of inoculation of inflorescences, 58% of experimental plants, and no control plants, had acquired a novel allele at the locus Xgwm112.1. This allele was detected only in head blight affected tissue. Uninoculated control plants, and leaf samples from inoculated plants, showed no mutation. Cloning and sequencing of PCR products indicates that the new allele was generated by contraction of the (CT)(n) repeat motif. Observation of the same deletion-based mutation in all varieties, its absence in control plants not exposed to the head blight pathogen, and the detection of no similar mutational events in a control panel of loci not expected to show mutation, indicates that this example of microsatellite mutation is induced and/or caused by FHB infection.     

RAPD and SCAR Markers Linked to a Gene Conferring Resistance to Angular Leaf Spot in Common Bean

Angular leaf spot, caused by the fungus Phaeoisariopsis griseola , is one of the most important bean diseases in Brazil. The objectives of this study were to determine the inheritance of angular leaf spot resistance and to identify random amplified polymorphic DNA (RAPD) and sequence-characterized amplified region (SCAR) markers linked to the resistance gene present in cv. Cornell 49-242, in a cross between this cultivar and susceptible cv. Rudá. The parents, F₁, F₂ and backcross-derived plants were inoculated with P. griseola pathotype 31-17 under environmentally controlled greenhouse conditions. The results indicate that one single dominant gene controls the resistance in Cornell 49–242. Two RAPD markers, OPN 02_890c and OPE 04_650c, were found to be linked in the coupling phase, at 3.2 and 12.5 c m of the resistance gene, respectively. To increase the reproducibility of the detection of marker OPN02_890c it was converted into a SCAR marker. It is proposed that the designation of Phg-2 be used for the angular leaf spot resistant gene present in Cornell 49-242.     

A novel cost of R gene resistance in the presence of disease

Resistance responses can impose fitness costs when pests are absent. Here, we test whether the induction of resistance can decrease fitness even in plants under attack; we call this potential outcome a net cost with attack. Using lines in which genetic background was controlled, we investigated whether susceptible Arabidopsis thaliana plants can outperform R gene resistant plants when infected with pathogens. For the R gene RPS2, there was a fitness benefit of resistance in the presence of intraspecific competition, but there was a net cost in the absence of competition: resistant plants produced less seed than susceptible plants even though infected with Pseudomonas syringae. This net cost was primarily due to overcompensation by susceptible plants, which occurred because of a developmental response to infection. For the R gene RPP5, there was no fitness effect of resistance without competition but a net cost when plants were infected with Peronospora parasitica in the presence of competition. This net cost was due to a reduction in the fitness of infected, resistant plants and complete compensation in susceptible plants. A spatially variable model suggests that a trade-off between net benefits and net costs with attack may help explain the persistence of individuals lacking R gene resistance to disease.     

Necrotic Lesions as Unusual Symptoms of Ring Rot in the Potato Leaves

Test-tube plants and suspension cell cultures of two cultivars of the potato (Solanum tuberosum L.) differing in their resistance to ring rot caused by Clavibacter michiganensis subsp. sepedonicus and six strains of this bacterium were used to test the relationship between the virulence, the leaf ability to adsorb bacteria, and the symptoms of the disease. In addition to chlorosis and drying, heavy inoculation with virulent strains caused unusual symptoms, such as leaf necrotic lesions. In the resistant cultivar, the necrotic lesions were predominantly local, whereas in the susceptible cultivar, they expanded. Unlike the susceptible cultivar, suspension cells of the resistant cultivar weakly adhered bacteria of the tested strains. Bacteria entered the plants through the leaf stomata. The sorption and penetration were much more pronounced in the susceptible cultivar. It was concluded that strain virulence varies depending on the conditions of inoculation, and uncharacteristic symptoms (necrotic lesions) arise. The local necrotic lesions are considered a hypersensitive response, and exopolysaccharides of the pathogen as the factors of virulence.     

Temporal Effects of a Begomovirus Infection and Host Plant Resistance on the Preference and Development of an Insect Vector,Bemisia tabaci,and Implications for Epidemics

Persistent plant viruses, by altering phenotypic and physiological traits of their hosts, could modulate the host preference and fitness of hemipteran vectors. A majority of such modulations increase vector preference for virus-infected plants and improve vector fitness, ultimately favouring virus spread. Nevertheless, it remains unclear how these virus-induced modulations on vectors vary temporally, and whether host resistance to the pathogen influences such effects. This study addressed the two questions using a Begomovirus-whitefly-tomato model pathosystem. Tomato yellow leaf curl virus (TYLCV) -susceptible and TYLCV-resistant tomato genotypes were evaluated by whitefly-mediated transmission assays. Quantitative PCR revealed that virus accumulation decreased after an initial spike in all genotypes. TYLCV accumulation was less in resistant than in susceptible genotypes at 3, 6, and 12 weeks post inoculation (WPI). TYLCV acquisition by whiteflies over time from resistant and susceptible genotypes was also consistent with virus accumulation in the host plant. Furthermore, preference assays indicated that non-viruliferous whiteflies preferred virus-infected plants, whereas viruliferous whiteflies preferred non-infected plants. However, this effect was prominent only with the susceptible genotype at 6 WPI. The development of whiteflies on non-infected susceptible and resistant genotypes was not significantly different. However, developmental time was reduced when a susceptible genotype was infected with TYLCV. Together, these results suggest that vector preference and development could be affected by the timing of infection and by host resistance. These effects could play a crucial role in TYLCV epidemics.     

Characteristics of resistance in rice to rice hoja blanca virus (RHBV) and its vector, Tagosodes orizicolus (Muir)

This paper characterises resistance to rice hoja blanca virus (RHBV) which is transmitted by the planthopper Tagosodes orizicolus (Muir). Resistance is expressed as decreased proportion of plants infected compared to susceptible lines, although severity of symptom expression is similar in both types. This resistance is not due to differences in vector feeding behaviour. Vectors reared eight generations on resistant plants showed no increased ability to transmit to resistant lines or decreased ability to transmit to susceptible ones. Longevity of vectors was similar when reared either on virus-resistant or susceptible plants. Incubation period of the virus in resistant plants are significantly longer than in susceptible plants. Resistance increased with plant age in resistant and susceptible cultivars. Increased virus dosage, as determined by increased number of viru-liferous vectors per inoculated plant, caused an increase in transmission to both resistant and susceptible cultivars. However, the ranking of resistant and susceptible remained the same across the experimental range of dosage and plant age. It is concluded that the resistance studied is to virus infection and there is little risk of “breakdown” occurring as a result of genetic or behavioural changes in the vector population. This will permit the use of economic thresholds to planthopper feeding damage with little risk of RHBV epidemic outbreak.     

Differential Light Responses of Photosynthesis by Triazine-resistant and Triazine-susceptible Senecio vulgaris Biotypes

Studies were conducted to determine a physiological basis for competitive differences between Senecio vulgaris L. biotypes which are either resistant or susceptible to triazine herbicides. Net carbon fixation of intact leaves of mature plants was higher at all light intensities in the susceptible biotype than in the resistant biotype. Quantum yields measured under identical conditions for each biotype were 20% lower in the resistant than in the susceptible biotype. Oxygen evolution in continuous light measured in stroma-free chloroplasts was also higher at all light intensities in the susceptible biotype than in the resistant biotype. Oxygen evolution in response to flashing light was measured in stroma-free chloroplasts of both biotypes. The steady-state yield per flash of resistant chloroplasts was less than 20% that of susceptible chloroplasts. Susceptible chloroplasts displayed oscillations in oxygen yield per flash typically observed in normal chloroplasts, whereas the pattern of oscillations in resistant chloroplasts was noticeably damped. It is suggested that modification of the herbicide binding site which confers s-triazine resistance may also affect the oxidizing side of photosystem II, making photochemical electron transport much less efficient. This alteration has resulted in a lowered capacity for net carbon fixation and lower quantum yields in whole plants of the resistant type.     

Consequences of the overproduction of methyl jasmonate on seed production, tolerance to defoliation and competitive effect and response of Arabidopsis thaliana

* Accumulation of methyl jasmonate (MeJA) after herbivore attack in plants is associated with the induction of defenses that can benefit fitness, but are costly to express; effects often explored using exogenous application of jasmonates. * Here I explored the consequences of the overexpression of MeJA on seed production, tolerance to defoliation and competitive effect and response, using a genotype of Arabidopsis thaliana that overexpresses jasmonic acid carboxyl methyltransferase (JMT) and contains threefold higher levels of MeJA than wild-type plants. * Without competition, JMT plants produced 37-40% less total seed mass than vector controls or wild-type plants, and had reduced seed germination. Defoliation reduced height more strongly in wild-type than in JMT plants, but reduced total seed production equally. In a competition experiment, the presence of a neighbor reduced fitness more strongly in wild-type than in JMT plants, but JMT plants exhibited dampened opportunity costs and benefits of induction with jasmonic acid of itself or its neighbor. This may have related to the higher constitutive expression but reduced inducibility of jasmonate-mediated defenses, including trypsin inhibitors, exhibited by JMT plants. * In natural plant populations, overexpression of MeJA-mediated responses should be beneficial to resistance to herbivores, pathogens and competitors, but is directly costly to fitness and probably constrains plasticity in response to changing environmental conditions.     

No fitness cost for wheat's H gene-mediated resistance to Hessian fly (Diptera: Cecidomyiidae)

Resistance (R) genes have a proven record for protecting plants against biotic stress. A problem is parasite adaptation via Avirulence (Avr) mutations, which allows the parasite to colonize the R gene plant. Scientists hope to make R genes more durable by stacking them in a single cultivar. However, stacking assumes that R gene-mediated resistance has no fitness cost for the plant. We tested this assumption for wheat's resistance to Hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae). Our study included ten plant fitness measures and four wheat genotypes, one susceptible, and three expressing either the H6, H9, or H13 resistance gene. Because R gene-mediated resistance has two components, we measured two types of costs: the cost of the constitutively-expressed H gene, which functions in plant surveillance, and the cost of the downstream induced responses, which were triggered by Hessian fly larvae rather than a chemical elicitor. For the constitutively expressed Hgene, some measures indicated costs, but a greater number of measures indicated benefits of simply expressing the H gene. For the induced resistance, instead of costs, resistant plants showed benefits of being attacked. Resistant plants were more likely to survive attack than susceptible plants, and surviving resistant plants produced higher yield and quality. We discuss why resistance to the Hessian fly has little or no cost and propose that tolerance is important, with compensatory growth occurring after H gene-mediated resistance kills the larva. We end with a caution: Given that plants were given good growing conditions, fitness costs may be found under conditions of greater biotic or abiotic stress.     

Effects of gossypol on fitness costs associated with resistance to Bt cotton in pink bollworm

Fitness costs associated with insect resistance to Bacillus thuringiensis (Bt) crops may help to delay or prevent the spread of resistance alleles, especially when refuges of non-Bt host plants are present. The potential for such delays increases as the magnitude and dominance of fitness costs increase. Here, we examined the idea that plant secondary chemicals affect expression of fitness costs associated with resistance to Bt cotton in Pectinophora gossypiella (Saunders). Specifically, we tested the hypotheses that gossypol affects the magnitude or dominance of fitness costs, by measuring performance of three independent sets of pink bollworm populations fed artificial diet with and without gossypol. Each set had an unselected susceptible population, a resistant population derived by selection from the susceptible population, and the F1 progeny of the susceptible and resistant populations. No individuals completed development on diets with gossypol in one set, suggesting that these individuals partially lost the ability to detoxify this chemical. In the other two sets, costs affecting survival did not support the hypotheses, but costs affecting pupal weight did. Adding gossypol to diet increased the magnitude and dominance of costs affecting pupal weight. In one of the two sets with survivors on diet with gossypol, costs affecting development time were less recessive when gossypol was present in diet. These results indicate that gossypol increased the magnitude and dominance of some fitness costs. Better understanding of the effects of natural plant defenses on fitness costs could improve our ability to design refuges for managing insect resistance to Bt crops.     

Isofemale Line Analysis of Meloidogyne incognita Virulence to Cowpea Resistance Gene Rk

Isofemale lines (IFL) from single egg masses were studied for genetic variation in Meloidogyne incognita isolates avirulent and virulent to the resistance gene Rk in cowpea (Vigna unguiculata). In parental isolates cultured on susceptible and resistant cowpea, the virulent isolate contained 100% and the avirulent isolate 7% virulent lineages. Virulence was selected from the avirulent isolate within eight generations on resistant cowpea (lineage selection). In addition, virulence was selected from avirulent females (individual selection). Virulence differed (P ≤ 0.05) both within and between cohorts of IFL cultured for up to 27 generations on susceptible or resistant cowpea. Distinct virulence profiles were observed among IFL. Some remained avirulent on susceptible plants and became extinct on resistant plants; some remained virulent on resistant and susceptible plants; some changed from avirulent to virulent on resistant plants; and others changed from virulent to avirulent on susceptible plants. Also, some IFL increased in virulence on susceptible plants. Single descent lines from IFL showed similar patterns of virulence for up to six generations. These results revealed considerable genetic variation in virulence in a mitotic parthenogenetic nematode population. The frequencies of lineages with stable or changeable virulence and avirulence phenotypes determined the overall virulence potential of the population.     

Antioxidative enzymes and isozymes analysis of taro genotypes and their implications in <Emphasis Type="Italic">Phytophthora</Emphasis> blight disease resistance

Assessment of the differential expression of antioxidative enzymes and their isozymes, was done in 30 day-old ex vitro raised plants of three highly resistant (DP-25, Jhankri and Duradim) and one highly susceptible (N-118) genotypes of taro [Colocasia esculenta (L.) Schott]. Antioxidative enzymes were assayed in the ex vitro plants, 7 days after inoculation with the spores (15,000 spores ml⁻¹ water) of Phytophthora colocasiae Raciborski to induce taro leaf blight disease. Uninoculated ex vitro plants in each genotype were used as control. The activity of superoxide dismutase (SOD) and guaiacol peroxidase (GPX) increased under induced blight condition when compared with control. Increase in antioxidative enzymes was more (67–92%) in the resistant genotypes than that (21–29%) of the susceptible genotype. The zymograms of SOD and GPX in the resistant genotypes, with pathogenic infection, showed increased activity for anodal isoform of SOD and increased expression and/or induction of either POX 1 or POX 2 isoforms of GPX. In susceptible genotype, expression of the above isoforms was faint for SOD and nearly absent for GPX under both blight free and induced blight conditions. Induction and/or increased activity of particular isoform of SOD and GPX against infection of Phytophthora colocasiae in the resistant genotypes studied led to the apparent conclusion of linkage of isozyme expression with blight resistance in taro. This might be an important criterion in breeding of taro for Phytophthora leaf blight resistance.     

A mechanism of chlorotoluron resistance in Lolium rigidum

Many biotypes of Lolium rigidum Gaud, (annual ryegrass) have developed resistance to herbicides; however, few have developed resistance to phenylurea herbicides. Two biotypes with different histories of herbicide selection pressure were six to eight times less sensitive to the phenylurea herbicide, chlorotoluron, than a susceptible biotype. Resistance was not due to differences in the herbicide target site as oxygen evolution by thylakoids isolated from resistant and susceptible biotypes was similarly inhibited by diuron and chlorotoluron. There was no difference in the uptake and distribution of chlorotoluron into resistant and susceptible plants. There was a twofold greater rate of chlorotoluron detoxification in resistant plants with N-demethylation being a major detoxification reaction. Resistant plants treated with a 3-h pulse of 120 M chlorotoluron recovered net carbon fixation after 42 h, half the time taken by susceptible plants. The mixed-function oxidase inhibitor 1-aminobenzotriazole (70 M) intensified the effects of chlorotoluron in resistant plants when applied in combination with the herbicide for 7 d. 1-Aminobenzotriazole also inhibited the metabolism of chlorotoluron in both resistant and susceptible plants. The cytochrome P-₄₅₀ inhibitor, piperonyl butoxide piperonyl butoxide, interacted with chlorotoluron when applied to plants growing in soil. Chlorotoluron applied with reduced plant dry weight to a greater extent than chlorotoluron alone. It appears, therefore, that enhanced detoxification is the major mechanism of resistance to chlorotoluron in the resistant biotypes studied.Abbreviations ABT 1-aminobenzotriazole - VLR1 Victorian L. rigidum biotype 1 — herbicide susceptible - VLR69 Victorian L. rigidum biotype 69 — herbicide resistant - WLR2 Western Australian L. rigidum biotype 2 — herbicide resistant M.W.M.B, was supported by an Australian Postgraduate Research Award and a supplementary scholarship from the Grains Research and Development Corporation. We are very grateful to Dr. E. Ebert, Ciba Geigy, Basal, Switzerland for providing [¹⁴C]chlorotoluron and standards of chlorotoluron metabolites. We express our gratitude to Dr. John Huppatz of the CSIRO Division of Plant Industry for providing ABT. We also thank Ciba Geigy Australia for providing technical-grade chlorotoluron and formulated phenylurea herbicides.