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Herbicide resistance has the potential to impose fitness costs in plants. In Mexico, Phalaris minor Retz. has evolved resistance to acetyl-coa carboxylase (ACCase) inhibiting herbicides. However, it is unknown whether these changes may involve other ecological costs for resistant individuals under non-selective conditions. The aim of this study was to determine whether the evolution of resistance has reduced the ability of P. minor to compete for resources at the seedling stage. De Wit's replacement series experiments (susceptible:resistant 100:0, 50:50, 0:100) were performed under two initial conditions. In the first experiment, we examined the effect of differences in germination time between resistant and susceptible genotypes on canopy cover. In the second experiment, germination was synchronized, and all the replacement series began at the same time. Delayed germination of resistant accessions reduced their ability to compete for canopy space, competition intensity index, and relative productivity in relation to susceptible accessions. When germination was synchronized, the resistant accessions had the same canopy cover and productivity as susceptible accessions. The delayed germination may cause displacement of resistant individuals in the absence of the selective factor (herbicide).

Littleseed canarygrass (∼canarygrass) evolved populations that are resistant to isoproturon during the early 1990s in north-western India. Clodinafop-propargyl (∼clodinafop) was recommended for controlling these populations. It has been used extensively in wheat for the last several years. Recently, poor or no control of canarygrass by clodinafop has been observed in large areas, which could be related to cross-resistance or multiple resistance. This study was designed to test whether resistance has evolved in canarygrass populations against clodinafop and to explore control of the resistant populations with sulfosulfuron and pinoxaden. Among the 311 canarygrass populations that were tested, 86, 55 and 34 showed variable phytotoxicity (0–99%) due to 0.030, 0.060 and 0.120 kg ha−1 clodinafop, respectively. Based on the resistance index, 11 populations were “highly resistant”, 60 were “resistant” and the rest (240) were “susceptible” to clodinafop. Five and six clodinafop-resistant populations showed slight resistance to 0.0125 kg ha−1 sulfosulfuron and 0.025 kg ha−1 pinoxaden, respectively. But, sulfosulfuron at 0.025 and 0.050 kg ha−1 and pinoxaden at 0.050 and 0.100 kg ha−1 controlled all the canarygrass populations. Clodinafop used for 4 years increased the chance of resistance evolving, whereas its rotation with sulfosulfuron reduced the chance of resistance evolving. This study showed that considerable canarygrass populations have evolved a low-to-high degree of resistance against clodinafop. The further use of clodinafop would lead to the spread of resistance in larger areas through the dispersal of resistant seeds. Clodinafop should be replaced with 0.025 kg ha−1 sulfosulfuron or 0.050 kg ha−1 pinoxaden. Besides, where canarygrass has not evolved resistance, the yearly rotation of sulfosulfuron with clodanafop or pinoxaden might delay the evolution of resistance.

Gherekhloo J, Osuna MD & De Prado R (2012). Biochemical and molecular basis of resistance to ACCase-inhibiting herbicides in Iranian Phalaris minor populations. Weed Research52, 367–372. Summary Phalaris minor is a common weed in wheat and barley fields of Iran. Repeated use of ACCase inhibiting herbicides during the last two decades to control this weed has resulted in the appearance of populations of P. minor that are resistant to the aryloxyphenoxy propionates (APP) in some wheat fields of the country. Dose–response assays were conducted to investigate the level of resistance in two P. minor populations (AR and MR4) which have developed resistance to three APP herbicides, including diclofop-methyl, fenoxaprop-P ethyl and clodinafop propargyl. A high level of resistance in these populations may be due to the presence of an altered ACCase enzyme in these plants, and we hypothesised that one or more mutations in the gene encoding ACCase enzyme are responsible for insensitivity of the enzyme. Results confirmed that resistance in both populations was target site-based, and molecular studies revealed that substitutions of Trp-2027-Cys and Asp-2078-Gly, respectively in AR and MR4, are responsible for insensitivity of the enzyme in these populations. This is the first report to show that these substitutions endow resistance to APP herbicides in P. minor, though other resistant biotypes are reported from elsewhere. These mutations may result in resistance of P. minor to some DIM and DEN herbicides. It seems that lack of adequate herbicide and crop rotation has selected plants with different target site mutations, which cause differential responses of the ACCase enzyme to ACCase inhibitors. Additional resistance management practices may be necessary to prevent ACCase-inhibiting herbicides from becoming ineffective over wide areas.

Phalaris minor (littleseed canary grass) is a major weed in wheat fields in some parts of Iran. Diclofop-methyl, fenoxaprop-P-ethyl, and clodinafop-propargyl are three acetyl coenzyme A carboxylase (ACCase)-inhibiting herbicides that are commonly used to control this grass in wheat fields.Thirty-four P. minor populations with suspected resistance to ACCase-inhibiting herbicides were sampled from wheat fields in the provinces of Fars and Golestan in Iran.The dose‐response assays that were conducted under controlled greenhouse conditions indicated that 14 populations were resistant to fenoxaprop-P-ethyl, seven populations were resistant to both fenoxaprop-P-ethyl and diclofop-methyl, and three populations were resistant to fenoxaprop-P-ethyl, diclofop-methyl, and clodinafop-propargyl. These populations showed different levels of resistance to the applied herbicides,compared to the susceptible population. These results suggest that different mechanisms of resistance could be involved.The enzyme assay revealed that the existence of modified ACCase in the three most-resistant populations (AR, MR4, and SR3) is responsible for the resistance of these populations.

BACKGROUND: The increasing use of ACCase-inhibiting herbicides has resulted in evolved resistance in key grass weeds infesting cereal cropping systems worldwide. Here, a thorough and systematic approach is proposed to elucidate the basis of resistance to three ACCase herbicides in a Lolium multiflorum Lam. (Italian rye grass) population from the United Kingdom (UK24). RESULTS: Resistance to sethoxydim and pinoxaden was always associated with a dominant D2078G (Alopecurus myosuroides Huds. equivalent) target-site mutation in UK24. Conversely, whole-plant herbicide assays on predetermined ACCase genotypes showed very high levels of resistance to diclofop-methyl for all three wild DD2078 and mutant DG2078 and GG2078 ACCase genotypes from the mixed resistant population UK24. This indicates the presence of other diclofop-methyl-specific resistance mechanism(s) yet to be determined in this population. The D2078G mutation could be detected using an unambiguous DNA-based dCAPS procedure that proved very transferable to A. myosuroides, Avena fatua L., Setaria viridis (L.) Beauv. and Phalaris minor Retz. CONCLUSION: This study provides further understanding of the molecular basis of resistance to ACCase inhibitor herbicides in a Lolium population and a widely applicable PCR-based method for monitoring the D2078G target-site resistance mutation in five major grass weed species. Copyright © 2010 Society of Chemical Industry

BACKGROUND: To reduce herbicide use by 75%, integrated use of sorghum and sunflower extracts each at 18 L ha−1 combined with 1/4th (75% less) of label rates of four herbicides (mesosulfuron + idosulfuron, metribuzin, phenoxaprop-p-ethyl and isoproturon) were investigated for the management of wild oat and canary grass, the two pernicious weeds in wheat fields worldwide. RESULTS: The results revealed that sorghum + sunflower extracts combined with 1/4th (75% less) of label rates of herbicides inhibited dry matter production of wild oat by up to 89% and canary grass by up to 92%. The wild oat and canary grass persistence index in sorghum + sunflower extracts combined with 1/4th (75% less) of label rates of herbicides was either lower or equal to respective label rates of herbicides, except sorghum + sunflower extract + 1/4th phenoxaprop-p-ethyl. Lower herbicide rates + water extracts also produced wheat grain yield statistically equal with label rates of respective herbicides. Two treatments having water extracts + lower herbicides rates were economical and sorghum + sunflower + 1/4th mesosulfuron + idosulfuron produced the highest (4404%) marginal rate of return. CONCLUSION: Herbicides use can be reduced by 75% through integration with sorghum + sunflower extracts without compromising yield and net benefits for cost-effective and eco-friendly management of wild oat and canary grass in wheat. Copyright © 2010 Society of Chemical Industry

Summary Photocontrol of weeds requires a better understanding of the effect of light on seed germination. Therefore, the effect of light intensity and exposure duration was tested on seed germination of two winter annual weeds, Phalaris minor and Poa annua. Photo-sensitised seeds of both species were subjected to four light intensities, including 130, 250, 500 and 1000 lmol m )2 s )1 , each at six exposure durations of 0.1, 1, 10, 100, 1000 and 10 000 s. Light stimulated the germination of both weeds. A modelling approach was used to describe the relationship between light intensity and exposure duration with the germination of weed seeds. A combination of logistic and Gaussian models provided good fits to P. minor percentage germination data, whereas in the case of P. annua, the incorporation of logistic and quadratic models better described the relationships. According to the model, increasing light intensity from 0 (full darkness) to 1000 lmol m )2 s )1 increased the germination of P. minor and P. annua from 28% to 57% and from 0% to 35% respectively. The seed germination of both weed species increased with exposure duration, but declined at duration periods longer than 100 s in P. minor or 1000 s in P. annua. The model predicted a trade-off between light intensity and light duration for both weeds. That is, a high light intensity irradiated for a short time exposure may result in the same percentage germination as one with a low light intensity, but at longer exposure duration.

Optimizing the herbicide dose by the addition of adjuvants is an acceptable way to reduce the risk of side-effects from herbicides. Therefore, to detect a suitable adjuvant for diclofop-methyl, cycloxydim, and clodinafop-propargyl against littleseed canarygrass (Phalaris minor) and wild oat (Avena ludoviciana), six dose-response experiments were conducted. The treatments consisted of diclofop-methyl at 0, 112, 225, 450, 675, and 900 g ai ha⁻¹, cycloxydim at 0, 15, 30, 60, 90, and 120 g ai ha⁻¹, and clodinafop-propargyl at 0, 8, 16, 32, 48, and 64 g ai ha⁻¹ with and without the adjuvants of Frigate, olive oil, and castor oil at 0.2% (v/v) in order to control both littleseed canarygrass and wild oat. Tested herbicides performance was enhanced by all adjuvants against littleseed canarygrass and wild oat. The addition of Frigate and the vegetable oils had the lowest and the highest effect on the performance of all of the herbicides on both littleseed canarygrass and wild oat, respectively, which confirms the solubilizing nature of the cuticular waxes by vegetable oils. A comparison between the two vegetable oils revealed that olive oil exerted a greater control of littleseed canarygrass than did the castor oil. In contrast, castor oil exerted a greater control of wild oat than did the olive oil, which can be related to differences in the leaf surface micromorphology of the weeds.

Littleseed canarygrass (Phalaris minor Retz.), a troublesome weed of wheat in India, has evolved multiple herbicide resistance across three modes of action: photosynthesis at the photosystem II site A, acetyl-coA carboxylase (ACCase), and acetolactate synthase inhibition. The multiple herbicide-resistant (MHR) populations had a low level of sulfosulfuron resistance but a high level of resistance to clodinafop and fenoxaprop (ACCase inhibitors). Some of the populations had GR50 (50% growth reduction) values for clodinafop that were 11.7-fold greater than that of the most susceptible population. The clodinafop-resistant populations also showed a higher level of cross-resistance to fenoxaprop (fop group) but a low level of cross-resistance to pinoxaden (den group). Although clodinafop and pinoxaden are from two different chemical families (fop and den groups), their same site of action is responsible for cross-resistance behavior. The populations that were resistant to four groups of herbicides (phenylureas, sulfonylurea, aryloxyphenoxypropionate, and phenylpyrazolin) were susceptible to the triazine (metribuzin and terbutryn) and dinitroaniline (pendimethalin) herbicides. The P. minor populations that were resistant to the aryloxyphenoxypropionate and phenylurea herbicides were effectively controlled by the sulfonylurea herbicide, sulfosulfuron. In the fields infested with P. minor that was resistant to clodinafop, a sulfosulfuron application (25 g ha−1) increased the wheat yield by 99.2% over that achieved using the recommended rate of clodinafop (60 g ha−1). However, the evolution of multiple resistance against the four groups is a threat to wheat production. To prevent the spread of MHR P. minor populations, as well as the extension of multiple resistance to new chemicals, concerted efforts in developing and implementing a sound, integrated weed management program are needed. The integrated approach, consisting of crop and herbicide rotation with cultural and mechanical weed control tactics, should be considered as a long-term resistance management strategy that will help to sustain wheat productivity and farmers' income.

Summary Studies were conducted to investigate seed longevity, seedling emergence and seed production of the weed Phalaris minor in wheat in northern India. The longevity of P. minor seeds buried in bags in the field was often limited to less than 1 year, although many seeds buried at 30 cm depth in a rice–wheat rotation remained viable for longer. The application of direct seeding in wheat reduced the seedling emergence rate of P. minor, when compared with conventional ploughing and sowing. However, a larger P. minor seedbank in the upper soil layer in plots under direct seeding partly reversed this positive effect in one of the two studies. Besides differences in relative distribution of weed seeds through the soil profile, it was likely that other factors such as reduced soil disturbance and soil characteristics associated with the application of direct seeding were also involved in regulating the emergence rate. Mature P. minor plants in wheat were found to invest a stable part (27%) of their aboveground biomass in seed, so that total seed weight was strongly linearly correlated with the aboveground biomass of the mother plant. Individual seed weight, however, was little affected by the weight of the mother plant.

Phalaris minor, the most serious weed in wheat in north-western India, has developed extensive isoproturon resistance due to continuous isoproturon use. For its control, alternative herbicides (flufenacet, metribuzin and sulfosulfuron) at different application rates, and timing were evaluated in wheat. In addition, herbicide carryover risk onto rotational crops (sorghum; maize and green gram, Vigina radiata) was also assessed. Isoproturon at 1 and 2 kg a.i. ha -1 provided only 10.5% and 51.8% P. minor control respectively. Of the other herbicides, early post-emergent [15-21 days after sowing (DAS)] flufenacet at 180-480 g a.i. ha -1 provided acceptable control of P. minor, but failed to control broad-leaved weeds and was phytotoxic to the wheat crop. Metribuzin at 210 g a.i. ha -1 was effective in controlling both Phalaris and dicotyledonous weeds. Mixtures of both flufenacet and metribuzin at reduced rates were better than flufenacet for weed control and grain yield. The efficacy of flufenacet and metribuzin was drastically reduced with later growth stages of P. minor (four to five leaf). Whereas sulfosulfuron at 25-30 g a.i. ha -1 , applied either early post-emergence (19 DAS) or post-emergence (30-42 DAS), was quite effective. Overall, sulfosulfuron was the most effective treatment with regard to weed control and crop yield. However, maize and sorghum grown in rotation after harvest of sulfosulfuron-treated wheat plots showed 65-73% crop biomass inhibition. The residual effect of sulfosulfuron was also noticed on Trianthema portulacastrum (Horse purslane), causing 73.5% dry matter reduction. By contrast, no carryover damage with flufenacet was observed on maize, sorghum and green gram. Glasshouse pot experiments and field trials investigating crop sensitivity to pre-plant applications of sulfosulfuron found the decreasing order: sorghum > maize > green gram. The risk of carryover onto rotational crops should be considered when choosing alternative herbicides for P. minor control in wheat.

Intra- and inter-sample similarities for four populations of the annual grass weed Phalaris minor from Haryana state, India, were examined using inter-simple sequence repeat (ISSR) DNA markers. Levels of polymorphism within and between populations were low in comparison with values reported from other grassy weed species. Analysis of inter-population similarities allowed a partial differentiation of the four populations and of pairs of populations classified by cropping system. Analysis of the intra-population similarity data showed a weak but consistent and statistically significant negative correlation between the molecular similarity of seedlings and the physical distance between their mother plants over distances up to 40 m (the maximum separation tested) in all four populations. The consistency of the observed relationship between molecular similarity and physical separation, and the differences in cultivation practices at the four sites, suggested that the relationship may be a result of localized out-crossing, rather than an effect of localized seed rain. The results of the analyses are discussed in relation to the potential for evolution of multiple traits in the weed in response to changes in the wheat production system in the region.

Summary Seed placement, soil temperature and soil moisture content influenced the process of after-ripening in Phalaris minor seeds. Seeds of P. minor collected from the soil just after wheat harvesting exhibited higher germination than seeds from P. minor threshed directly. There was a pronounced impact of periodic inhabitation of seed into the soil on germination after its dispersal. Germination was strongly inhibited when the seed was kept in soil at more than field capacity (FC) or in water. Maximum germination of seed incubated in soil at FC occurred at 30°C while a temperature of 40°C favoured after-ripening of seed when mixed with dry soil or kept dry without any medium. Release from conditional dormancy was quicker in the seed retrieved from the soil kept at 20°C than at 10°C. Seed release from conditional dormancy and germination increased with a rise in temperature from 30 to 40°C when the seed was retrieved from incubation in soil at FC for 70 days. The seed kept immersed in water was least responsive to a rise in temperature. Seed recovered from dry soil, or kept without any medium, responded quickly at both temperatures. Light enhanced the germination of Phalaris minor seed. The seedbank subjected to rice (Oryza sativa) field management conditions lost vigour in comparison with the seed stored in laboratory. There was significant variability in seed viability when exposed to differential water management conditions in rice.

Summary Germination of Phalaris minor declined with the increase in duration of imbibition in water from 30 min to 72 h at temperatures above 22 °C. Germination was reduced down to 10 cm and 2 cm soil depth by wheat straw burning in puddled and non-puddled soil, respectively, with maximum reduction near the soil surface. The dormancy of P. minor seed was not more than 60 days under field conditions. In puddled soil, 38–60% of the viable seeds of P. minor remained concentrated in the upper 5-cm layer. Germination decreased with an increase in soil depth. In total, 15% of seeds stored in the laboratory emerged from 10-cm depth, whereas seeds did not germinate below 4.2-cm depth under field conditions. Depth of emergence of P. minor was shallower in zero tillage compared with the conventional method of wheat sowing. The seeds retrieved from rice soils kept under continuous submergence for 60 days exhibited 26% and 57% loss of germination over semi-submergence and semi-wet conditions respectively. There was 100% loss of germination in 10-month-old seeds retrieved from the soil under rice-growing conditions. Plant density of P. minor was lower in zero tillage than with the conventional method of wheat sowing. Cross-ploughing in the upper 2–5 cm of soil (shallow tillage) and drill-sowing of wheat 1 week after shallow tillage reduced germination of P. minor by 44% and 37% and increased grain yield by 21% and 47% over zero-tillage and conventional methods respectively.

Field experiments were carried out in Northern Greece from 1994 to 1997 to study interference between Avena sterilis L. or Phalaris minor Retz. and five autumn-sown barley cultivars. Weed:crop interference began in early April. Avena sterilis at 120 plants m−2 showed greater interference against barley than P. minor at 400 plants m−2. The greatest grain yield and ear number reduction due to interference by either weed was recorded for cvs Klipper and Plaisant, and the least for cv. Athinaida; with cvs Carina and Thermi intermediate. Yield reduction due to A. sterilis for cvs Athinaida, Carina, Thermi, Klipper and Plaisant was 8, 16, 27, 61 and 67%, respectively, while corresponding losses to P. minor were 1, 8, 14, 45 and 55%. These results clearly indicate that growth and consequently yield components of cv. Athinaida were unaffected by the presence of either weed species, while those of cv. Carina were affected by A. sterilis, but not by P. minor. However, dry weight and panicle number of both weed species were severely reduced by interference with cvs Carina, Athinaida and Thermi compared with cvs Klipper and Plaisant. The order of interference of the five barley cultivars tested against A. sterilis and P. minor was Athinaida > Carina > Thermi > Klipper ≥ Plaisant.

Accessions of annual Phalaris species in germplasm collections lack adequate genetic characterization for utilization. The objective of this study was to assess the level of isozyme variability of five annual Phalaris species, including 49 annual canarygrass (P. canariensis L.; 2n = 2x = 12), 24 P. brachystachys Link. (2n = 2x = 12), 48 P. minor Retz. (2n = 4x = 28), 19 P. paradoxa L. (2n = 2x = 14), and P. angusta Nees ex Tris (2n = 2x = 14) accessions from North American germplasm collections. Eight enzymes were used to detect polymorphism within and among accessions by scoring the presence or absence of 62 bands. Polymorphism was detected in 6 of 19 bands in P. canariensis, 12 of 22 bands in P. brachystachys, 34 of 40 bands in P. minor, 19 of 29 bands in P. paradoxa, and 6 of 21 bands in P. angusta. Polymorphism was detected in 23 accessions (47%) P. canariensis, 15 accessions (63%) of P. brachystachys, 35 accessions (73%) of P. minor, four accessions (19%) of P. paradoxa and three accessions of P. angusta (100%). Within-accession polymorphic indices ranged from 0 to 0.03 in P. canariensis, 0 to 0.02 in P. brachystachys, 0 to 0.06 in P. minor, 0 to 0.04 in P. paradoxa, and 0.01 to 0.04 in P. angusta. Jaccard's similarity coefficients ranged from 0.83 to 1.0 in P. canariensis, 0.78 to 1.0 in P. brachystachys, 0.47 to 1.0 in P. minor, 0.44 to 1.0 in P. paradoxa, and 0.81 to 0.86 in P. angusta. Little isozyme variation existed either within or among accessions of the four diploid species. Phalaris minor, an autotetraploid, possessed greater variation among accessions rather than within accessions. In summary, limited variation was detected by means of isozymes in these five annual Phalaris species obtained from North American germplasm collections.

The effect of the monooxygenase inhibitor, 1-aminobenzotriazole (ABT) on isoproturon phytotoxicity and metabolism was studied in resistant (R) and susceptible (S) biotypes of Phalaris minor and in wheat (Triticum aestivum). Addition of ABT (2·5, 5 and 10 mg litre-1) to isoproturon (0·25, 0·5, 1, 2 and 4 mg litre-1) in the nutrient solution significantly enhanced the phytotoxicity of isoproturon against the R biotype. Isoproturon at 0·25 mg litre-1 reduced the dry weight (DW) of the S biotype by 77%, whereas the R biotype required 4·0 mg litre-1 for similar reduction. Addition of 10 mg litre-1 of ABT to the 0·25 mg litre-1 isoproturon caused 71 and 82% reduction in DW of R and S biotypes, respectively. Wheat was more sensitive to the mixture of isoproturon and ABT than the R biotype of P. minor. Reduced concentrations of ABT in the mixture from 10 to 2·5 mg litre-1 increased the DW of the R biotype more than that of the S biotype. The R biotype metabolised [14C]isoproturon at a faster rate than the S biotype. ABT (5 mg litre-1) inhibited the degradation of [14C]isoproturon in both biotypes of P. minor and in wheat. In the presence of ABT, about half of the applied [14C]isoproturon remained as parent herbicide in all the three species after two days. The metabolites were similar in the R and S biotypes and wheat as determined by co-chromatography with reference standards and mass spectroscopy (MS). ABT inhibited the appearance of the hydroxy and monomethyl metabolites and their conjugates in all the test plants. These results suggest that the activity of the enzymes responsible for the degradation of isoproturon is greater in the R than in the S biotype of P. minor, resulting in its rapid detoxification. Incorporation of the monooxygenase inhibitor ABT into the nutrient solution greatly inhibited the degradation of [14C]isoproturon in the R biotype and increased its phytotoxicity. Both hydroxylation and N-dealkylation reactions were found to be sensitive to ABT; inhibition of hydroxylation was greater than that of demethylation. Since ABT could not completely suppress isoproturon degradation, it is possible that more than one monooxygenase is involved. © 1998 SCI

Summary Three pot experiments have investigated the effects of nitrogen (N) supply on interspecific competition between three weed species (Phalaris minor Retz., Chenopodium album L, and Sinapis arvensis L.) and spring wheat (Triticum aestivum L.) cv, Alexandria. The treatments tested included monocultures of each species and a mix-ture containing them in equal proportions that were combined factorially with two levels of N supply (20, 120 kg N ha-1). Low N supply decreased net photosynthetic rate (Pn), leaf N percentage, plant dry weight and N uptake of both wheat and weed species and gram dry weight of wheat. The effects of low N on Pn and dry weight of weeds were greater than the effects on wheat. In most cases the decrease in Pn at low N was due to non-stomatal factors. The relative competitive abilities of wheat and weeds were influenced by N supply. At high N, S.arvensis was more competitive than wheat, whereas P. minor was less competitive than wheat. C. album was more competitive than wheat at both N levels. The rank order of competitive ability of the weed species was C. album > P. minor > S. arvensis. The effects of interspecific competition on Pn were smaller than the effects of N supply and were not associated with corresponding effects on leaf N percentage and plant dry weight of both wheat and weed species and grain dry weight of wheat.

The 1995/6 International Survey of Herbicide-Resistant Weeds recorded 183 herbicide-resistant weed biotypes (124 different species) in 42 countries. The increase in the number of new herbicide-resistant weeds has remained relatively constant since 1978, at an average of nine new cases per year worldwide. Whilst 61 weed species have evolved resistance to triazine herbicides, this figure now only accounts for one-third of all documented herbicide-resistant biotypes. Triazine-resistant weeds have been controlled successfully in many countries by the use of alternative herbicides. Due to the economic importance of ALS and ACCase inhibitor herbicides worldwide, and the ease with which weeds have evolved resistance to them, it is likely that ALS and ACCase inhibitor-resistant weeds will present farmers with greater problems in the next five years than triazine-resistant weeds have caused in the past 25 years. Thirty-three weed species have evolved resistance to ALS-inhibitor herbicides in 11 countries. ALS-inhibitor-resistant weeds are most problematic in cereal, corn/soybean and rice production. Thirteen weed species have evolved resistance to ACCase inhibitors, also in 11 countries. ACCase inhibitor resistance in Lolium and Avena spp. threatens cereal production in Australia, Canada, Chile, France, South Africa, Spain, the United Kingdom and the USA. Fourteen weed species have evolved resistance to urea herbicides. Isoproturon-resistant Phalaris minor infesting wheat fields in North West India and chlorotoluron-resistant Alopecurus myosuroides in Europe are of significant economic importance. Although 27 weed species have evolved resistance to bipyridilium herbicides, and 14 weed species have evolved resistance to synthetic auxins, the area infested and the availability of alternative herbicides have kept their impact minimal. The lack of alternative herbicides to control weeds with multiple herbicide resistance, such as Lolium rigidum and Alopecurus myosuroides, makes these the most challenging resistance problems. The recent discovery of glyphosate-resistant Lolium rigidum in Australia is a timely reminder that sound herbicide-resistant management strategies will remain important after the widespread adoption of glyphosate-resistant crops.

Wheat (cv. WH-147) and five biotypes of Phalaris minor Retz. (KR-1, H-4, K-2, H-2 and J-1) were treated with isoproturon in controlled environmental conditions to assess their level of resistance. Resistance of P. minor to isoproturon was found in the order of KR-1>H-4> K-2 > H-2 = J-1. Compared with the susceptible (S) biotype (H-2), the resistant (R) biotypes (KR-1, H-4 and K-2) of P. minor required 13.0, 4.5 and 2.7 times higher doses of isoproturon for a 50% reduction in growth (GR 50 ) and 2.4 times that of the S biotype (H-2) by wheat. The corresponding figures for KR-1, H-4, K-2 biotypes and wheat were 18, 4.1, 2.4 and 4.6 times based on dry weight reduction. The effect of isoproturon on photosynthesis was studied in vitro using five biotypes of P. minor and in vivo with wheat, KR-I (R) and H-2 (S) biotypes of P. minor. Under in vitro treatment conditions isoproturon inhibited the photosynthesis of all five P. minor biotypes, whereas in vivo the recovery was greater in the R biotype than in the wheat and the S biotype. Effects on chlorophyll fluorescence were also measured in wheat and the KR-1 (R) and H-2 (S) biotypes of P. minor. A 4-h treatment of excised leaves incubated in isoproturon solution (0.025 and 0.05mM concentration) resulted in a decreased fluorescence coefficient (F v /F m ratio, in which F v = variable fluorescence (F m - F o ); F m = the maximum fluorescence and F o = initial fluorescence) in wheat (Triticum aestivum L.) and both biotypes of P. minor. The recovery was, however, greater in the R biotype than in wheat and it was completely recovered within 24 h. No recovery was recorded in the case of the S biotype of P. minor and a greater recovery time was required for wheat than the R biotype. The higher dose required for growth inhibition in the R biotype and rapid recovery of oxygen evolution and fluorescence coefficient under in vivo conditions together with the absence of selectivity in vitro suggests that the target site was unaffected. It can be conjectured that resistance to isoproturon is most probably because of enhanced metabolism or sequestration of isoproturon, resulting in decreased target site delivery.

We report here the discovery and characterization of a fibrous mineral contaminant of the diet in that area of north-east Iran where oesophageal cancer has a very high incidence. This contaminant has a smoothly tapering shape and is between 50 and 150 micrometers long. The greatest diameter is between 1 and 10 micrometers and this decreases to a sharply pointed tip with a radius of curvature of between 0.25 and 0.60 micrometers. Electron microscope X-ray analysis shows that this fibre consists almost entirely of silica. It is free from alkali metals, aluminium and iron, and therefore differs from other known natural or manmade mineral fibres. Examination of the seeds of more than sixty different species of weed know to contaminate the wheat in this area of the Middle East shows that the fibre originates from the seeds of the common Mediterranean grass Phalaris minor. This seed bears fibres of the same dimensions, composition and birefringence, borne upon the inflorescence bracts which envelop the pericarp of the seeds of this and other members of the phalaris genus. They are broken off from the seed when the wheat is milled but persist in the flour, where up to 3,000 are found in each gram. Similar fibres can be isolated in quantity from the seeds of related species which are grown commercially, and they have a similar size and composition. When cells of the 3T3 mouse fibroblast line are exposed to these fibres in semi-solid suspension culture, their proliferation is stimulated more than 100-fold. We present an hypothesis for the involvement of these plant mineral fibres in the aetiology of oesophageal cancer in Iran and in other areas of high incidence.