Evolution and spread of glyphosate resistance in Conyza bonariensis in California and a comparison with closely related Conyza canadensis

Glyphosate‐resistant weeds are an increasing problem in perennial cropping systems in the Central Valley of California, USA. To elucidate the evolutionary origins and spatial spread of resistance, we investigated the geographical distribution of glyphosate resistance and the population genetic diversity and structure of Conyza bonariensis and compared the results with previously studied C. canadensis. Thirty‐five populations from orchards and vineyards across the Central Valley were sampled. Population genetic structure was assessed using microsatellite markers. Population‐level resistance was assessed in glasshouse screening of plants grown from field‐collected seed. Bayesian clustering and analyses of multilocus genotypes indicated multiple origins of resistance, as observed in C. canadensis. Pairwise F_ST analysis detected spatial spread of resistance in the south of the Central Valley, also similar to C. canadensis. The results strongly indicate that the southern valley was an environment markedly more suitable than the northern valley for resistance spread and that spread in Conyza species was driven by increased uniformity of strong selection in the southern valley, due to recent regulation on herbicides other than glyphosate. Accordingly, resistant C. canadensis individuals occurred at high frequencies only in the southern valley, but interestingly resistant C. bonariensis occurred at high frequencies throughout the valley. Expression of resistance showed varying degrees of plasticity in C. bonariensis. The lower selfing rate and substantially greater genotypic diversity in C. bonariensis, relative to C. canadensis, indicate greater evolutionary potential over shorter time periods. Interspecific hybridisation was detected, but its role in resistance evolution remains unclear.     

Enhanced weed‐crop competition effects on growth and seed production of herbicide‐resistant and herbicide‐susceptible annual sowthistle (Sonchus oleraceus)

Enhanced crop competition could aid in the management of annual sowthistle (Sonchus oleraceus L.), a dominant weed of Australian cropping systems. A two‐year pot study was conducted to evaluate the effect of wheat (Triticum aestivum L.) planting densities (0, 82, and 164 wheat plants/m²) on growth and seed production of glyphosate‐resistant (GR) and glyphosate‐susceptible (GS) biotypes of annual sowthistle. Without competition, both biotypes produced a similar number of leaves and biomass, but the GS biotype produced 80% more seeds (46,050 per plant) than the GR biotype. In competition with 164 wheat plants/m², the number of leaves in the GR and GS biotypes was reduced by 62 and 61%, respectively, in comparison with the no‐competition treatment, and similarly, weed biomass was reduced by 78 and 77%, respectively. Compared to no‐competition treatment, the seed production of GR and GS biotypes was reduced by 33 and 69%, respectively, when grown with 82 wheat plants/m², but increasing wheat density from 82 to 164 plants/m² reduced the number of seeds only in the GS biotype (81%). Both biotypes produced greater than 6,000 seeds per plant when grown in competition with 164 plants/m², suggesting that increased crop density should be integrated with other weed management strategies for efficient control of annual sowthistle.     

Reduced translocation is involved in resistance to glyphosate and paraquat in Conyza bonariensis and Conyza canadensis from California

Resistance to glyphosate and paraquat has evolved in some populations of Conyza spp. from California, USA. This study evaluated whether herbicide absorption and translocation were involved in the mechanism of resistance to both herbicides. Three lines of each species were used: glyphosate‐paraquat‐susceptible (GPS), glyphosate‐resistant (GR) and glyphosate‐paraquat‐resistant (GPR). Radiolabelled herbicide was applied to a fully expanded leaf, and absorption and movement out of the treated leaf were monitored for up to 24 h for paraquat and 72 h for glyphosate. Plants treated with paraquat were incubated in darkness for the first 16 h and then subjected to light conditions. More glyphosate was absorbed in C. bonariensis (52.9–58.3%) compared with C. canadensis (28.5–37.6%), but no differences in absorption were observed among lines within a species. However, in both species, the GR and GPR lines translocated less glyphosate out of the treated leaf when compared with their respective GPS lines. Paraquat absorption was similar among lines and across species (71.3–77.6%). Only a fraction of paraquat was translocated in the GPR lines (3% or less) when compared with their respective GPS or GR lines (20% or more) in both species. Taken together, these results indicate that reduced translocation is involved in the mechanism of resistance to glyphosate and paraquat in C. bonariensis and C. canadensis.     

Comparing after-ripening response and germination requirements of Conyza canadensis and C. bonariensis (Asteraceae) through logistic functions

Germination requirements and after‐ripening effects during one year of dry storage at 15/5 and 25/15°C (day/night) were compared for Conyza bonariensis and C. canadensis (Asteraceae). A logistic function was fitted to the results from tests over time in various incubation conditions, using three populations of each species as replicates. Time required for response to dry storage was measured by using a new method, the third derivative of the logistic function. Therefore, a point when major germination was achieved could be detected, without having to rely on maximum germination (which is uncertain), individual data points or any subjectively chosen limit. Fresh seeds of both species were dependent on light for germination and after‐ripening was mainly manifested by increasing germination in darkness. Low dormancy status and light requirement might indicate that soil cultivations should rapidly reduce the seedbanks of these species, although fecundity and wind dispersal will affect population levels. The species differed in their germination response, with C. bonariensis germinating at lower temperatures than C. canadensis. This seemingly counter‐intuitive result may explain the prevention of fatal germination of C. canadensis in cold conditions and its behaviour as a summer annual in northern climates, while C. bonariensis is restricted to warmer parts of the world.     

Seed germination response to temperature for a range of international populations of Conyza canadensis

Conyza canadensis is a surface‐germinating ruderal facultative winter annual with recruitment that is highly susceptible to changes in microsite conditions. A key adaptive characteristic for a facultative winter annual species, like C. canadensis, is germination response to temperature. The objective of this study was to determine the germination response to temperature for C. canadensis seed sourced from regions around the world with differing climates and, by doing so, gain insight into the role that seed germination biology plays in the adaptiveness and weediness of facultative winter annual weeds. Seed was sourced from populations in Málaga, Spain, Hertfordshire, UK, Shiraz, Iran and southern Ontario, Canada, and grown out in a common garden under controlled conditions to produce seed for this study. These seeds were then subjected to temperatures from 6.5 to 20°C at 1.5°C increments using a thermogradient plate. Cumulative daily germination counts for 30 days were recorded. Results indicated that temperature and source location had a significant effect on germination response. Estimated base germination temperature ranges were significantly different among the populations [Ontario (8–9.5°C), Iran (9.5–11°C), Spain (12.5–14°C), UK (11–12.5°C)], as were accumulated growing degree days (GDDs; d°C) required to reach 50% germination. For three of the four populations, estimated base germination temperature range values were below those previously reported in the literature. These differences are most likely rapid evolutionary adaptations to local climate and highlight the potential C. canadensis has to be problematic as a native and invasive species.     

Seedbank persistence,germination and early growth of glyphosate‐resistant Kochia scoparia

This study describes the seedbank persistence of glyphosate‐resistant (GR) Kochia scoparia at two sites in western Canada and examines if GRK. scoparia from western Canada and mid‐western United States (USA) differ from their susceptible counterparts in seed germination and early growth characteristics at low‐temperature regimes. Site or depth of seed burial (surface, 2.5 cm, 10 cm) did not affect seed viability over time and time to 50% and 90% loss of viability averaged 210 and 232 days respectively. Glyphosate‐resistant K. scoparia generally germinated later and had lower cumulative germination than glyphosate‐susceptible (GS) K. scoparia from Saskatchewan, Canada; and Kansas, USA; but not Colorado, USA. Similarly, time to 10% first leaf of GSK. scoparia from Saskatchewan and Kansas tended to be sooner than that of GRK. scoparia, with a greater percentage of GS vs. GR seedlings of populations from all regions having attained first leaf by the end of the experiment. The short seedbank longevity and delayed and reduced germination and time to first leaf of GRK. scoparia may potentially be exploited to maximise management efficacy through delayed preseeding weed control or alternatively by early seeding date to enhance crop competitiveness.     

Impact of Conyza bonariensis density and establishment period on soyabean grain yield,yield components and economic threshold

Conyza spp. have become a major weed around the world, mainly because of weed resistance issues. The objective of this work was to test the hypothesis that the soyabean crop yield is dependent on the density of Conyza bonariensis and on the timing of weed establishment in relation to the crop sowing date. It was also theorised that these variables affect soyabean crop yield components and the economic threshold of C. bonariensis on soyabean. Field experiments were conducted during 2010 and 2011 using a randomised complete block design. In each experiment, several densities (0, 3, 6, 12, 24, 48, 96 and 192 plants m^?2) of C. bonariensis were established in soyabean fields. Conyza bonariensis establishment dates varied considerably between the experiments [81, 38 and 0 days before soyabean sowing (DBSS)]. Conyza bonariensis plants were first cultivated in a glasshouse and then transplanted to the field at the three‐leaf growth stage. At the lower densities, each C. bonariensis plant decreased soyabean yield by 36%, 12% and 1.0%, when established at 81, 38 and 0 DBSS respectively. The economic thresholds based on sensitivity analysis were below 0.5 plant m^?2 when C. bonariensis was introduced at 81 and 38 DBSS; in contrast, they were between 2 and 4 plants m^?2 when the weed was established at the crop sowing time. The results emphasise the importance of proper C. bonariensis management prior to soyabean sowing and highlight the need for residual herbicides to avoid grain yield losses.     

A hydrothermal seedling emergence model for Conyza bonariensis

Conyza bonariensis is a South American native annual Asteraceae that has been introduced to the Mediterranean, where it behaves as a ruderal plant and a weed that is difficult to control in several crops. The development of predictive models can contribute to control measures at early growth stages, but currently there are no studies to predict seedling emergence of Conyza species. Our objectives were to develop and evaluate a model for predicting emergence response of C. bonariensis to the soil hydrothermal environment. A hydrothermal seed germination model was fitted to time course germination data from germination tests carried out at different constant temperatures and water potentials with the aim of establishing the hydrothermal parameters characterising C. bonariensis seed germination. The relationship between cumulative seedling emergence and cumulative hydrothermal time under field conditions was analysed using the Gompertz function. Model development was based on 2 years' data from a field experiment. Base temperature and base water potential for seed germination were estimated at 10.6°C and ?0.70 ± 0.151 MPa, respectively. The emergence model showed a very good fit to the experimental data. According to this model, seedling emergence starts at 15 accumulated hydrothermal time (HTT) after sowing, and 50 and 95% emergence is completed at 53 HTT and 105 HTT, respectively. For model evaluation, independent field experiments were carried out in two localities. Cumulative seedling emergence was accurately predicted by the model. Results indicate that this model can be useful as a predictive tool contributing to effective control of C. bonariensis populations.     

Glyphosate resistance in common ragweed ( A mbrosia artemisiifolia L.) from Mississippi,USA

Glyphosate is one of the most commonly used broad‐spectrum herbicides over the last 40 years. Due to the widespread adoption of glyphosate‐resistant (GR) crop technology, especially corn, cotton and soybean, several weed species have evolved resistance to this herbicide. Research was conducted to confirm and characterize the magnitude and mechanism of glyphosate resistance in two GR common ragweed ( A mbrosia artemisiifolia L.) biotypes from Mississippi, USA. A glyphosate‐susceptible (GS) biotype was included for comparison. The effective glyphosate dose to reduce the growth of the treated plants by 50% for the GR1, GR2 and GS biotypes was 0.58, 0.46 and 0.11 kg ae ha^?1, respectively, indicating that the level of resistance was five and fourfold that of the GS biotype for GR1 and GR2, respectively. Studies using ¹⁴ C‐glyphosate have not indicated any difference in its absorption between the biotypes, but the GR1 and GR2 biotypes translocated more ¹⁴ C‐glyphosate, compared to the GS biotype. This difference in translocation within resistant biotypes is unique. There was no amino acid substitution at codon 106 that was detected by the 5‐enolpyruvylshikimate‐3‐phosphate synthase gene sequence analysis of the resistant and susceptible biotypes. Therefore, the mechanism of resistance to glyphosate in common ragweed biotypes from Mississippi is not related to a target site mutation or reduced absorption and/or translocation of glyphosate.     

Fenugreek root exudates show species-specific stimulation of Orobanche seed germination

Various Orobanche species are weedy and cause severe reduction in the yields of many important crops. The seeds of these parasitic weeds may remain dormant in the soil for many years until germination is stimulated by the release of a chemical signal from a host plant. In order to determine the effects of fenugreek root exudate on the induction of Orobanche crenata, Orobanche ramosa and Orobanche foetida seed germination, root exudate was collected from hydroponically grown fenugreek seedlings. Fractionation patterns obtained from column and thin layer chromatography of the fenugreek root exudate showed a set of metabolites differing in their polarity with stimulatory activity on Orobanche seed germination. The crude root exudate stimulated both O. ramosa and O. crenata seed germination to the same level caused by the synthetic germination stimulant GR24 at 10 mg L^?1. It also stimulated O. foetida seed germination which did not respond to GR24. Active fractions of root exudate stimulated the germination of Orobanche species differentially.     

Rapid vacuolar sequestration: the horseweed glyphosate resistance mechanism

BACKGROUND: Glyphosate‐resistant (GR) weed species are now found with increasing frequency and threaten the critically important GR weed management system. RESULTS: The reported ³¹P NMR experiments on glyphosate‐sensitive (S) and glyphosate‐resistant (R) horseweed, Conyza canadensis (L.) Cronq., show significantly more accumulation of glyphosate within the R biotype vacuole. CONCLUSIONS: Selective sequestration of glyphosate into the vacuole confers the observed horseweed resistance to glyphosate. This observation represents the first clear evidence for the glyphosate resistance mechanism in C. canadensis. Copyright © 2010 Society of Chemical Industry     

AM真菌对加拿大一枝黄花与菌根植物和非菌根植物种间作用的影响

为明确丛枝菌根(arbuscular mycorrhizal,AM)真菌对加拿大一枝黄花Solidago canadensis与本地菌根植物和非菌根植物种间竞争格局的调控作用,采用温室盆栽试验,通过接种摩西球囊霉Glomus mosseae(GM)、根内球囊霉G. intraradices(GI)及其混合菌种(GM+GI)3种处理,分析AM真菌对加拿大一枝黄花与本地菌根植物玉米Zea mays和非菌根植物油菜Brassica campestris种间作用的影响。结果表明:与对照相比,接种AM真菌均显著提高了加拿大一枝黄花和玉米的菌根侵染率,菌根侵染率为13.720%～50.015%,且前者的菌根侵染率明显高于后者。单独种植时,与对照相比,接种AM真菌尤其是接种混合菌种显著提高了加拿大一枝黄花的株高、叶片数和总干重。在加拿大一枝黄花与玉米混合种植时,与单独种植相比,加拿大一枝黄花的株高、叶片数、根长和总干重均较低;同时,与对照相比,接种AM真菌显著提高了玉米的相对竞争强度而对加拿大一枝黄花的相对竞争强度没有显著影响。在加拿大一枝黄花和油菜混合种植时,与对照相比,接种AM真菌则显著提高了加拿大一枝黄花的株高、叶片数、净光合速率和总干重;同时,接种AM真菌促进了入侵种的竞争优势而抑制了非菌根植物油菜的生长。说明加拿大一枝黄花与本地种的竞争格局受到与之混生物种的菌根依赖性强度以及AM真菌的种类差异影响。     

Rising CO2 can alter fodder–weed interactions and suppression of Parthenium hysterophorus

Three C₄ grass (Setaria incrassata, Astrebla squarrosa and Bothriochloa decipiens) and one C₃ legume (Clitoria ternatea) suppressive fodder species, were re‐evaluated against the growth of the C₃ Parthenium hysterophorus under an ambient (390 μmol mol^?1) and an elevated atmospheric CO₂ concentration (550 μmol mol^?1). Under the elevated atmospheric CO₂, shoot dry biomass and suppression index (SI) value of the C₄ S. incrassata were both reduced by 32% and 0.7 respectively, while those for A. squarrosa were reduced by 23% and 0.3. In contrast and under the same elevated atmospheric CO₂ concentration, the shoot dry biomass and SI of the C₄ B. decipiens were increased by 8% and 0.1 respectively, while those for the C₃ C. ternatea were increased by 38% and 0.8. Our results suggest that C₃ fodder plants along with certain C₄ species could be utilised for the effective management of P. hysterophorus under the future elevated atmospheric CO₂ conditions. However, this system needs more fodder species to be investigated. Our results suggest that rising CO₂ per se may alter the efficacy of suppressive fodder management of an invasive C₃ species, P. hysterophorus.     

Effect of phosphate‐based seed priming on strigolactone production and Striga hermonthica infection in cereals

Strigolactones, plant‐secreted underground signalling molecules, play an important role in agricultural ecosystems, because they mediate the interaction of crops with symbiotic AM fungi and parasitic weeds like Striga hermonthica. Cereal host plants secret these signalling molecules particularly under nutrient‐deficient conditions and especially when phosphate (P) is limiting. The objective of the present study was to see the potential of P seed priming for S. hermonthica management in cereals in relation to strigolactone production. It has been demonstrated that P fertiliser application down‐regulates the production of these signalling molecules in the rhizosphere, which results in lower S. hermonthica infection of cereals. The laboratory study showed maximum production of strigolactones from dry and water‐soaked seeds, while seed soaking in P solution reduced their production. Similarly, maximum S. hermonthica infection was observed under control treatments with dry sowing or water soaking, while P seed soaking decreased S. hermonthica germination, emergence and dry biomass in all cereal crops. Our study shows that P seed priming resulted in lower exudation of strigolactones, which induced less S. hermonthica seeds germination and hence may lead to lower S. hermonthica infection. P‐based seed priming could prove to be an effective and affordable strategy to reduce S. hermonthica infection in cereals. Further research for practical field application is needed.     

Anoda cristata control with glyphosate in narrow- and wide-row soyabean

Experiments evaluated the effect of glyphosate rate and Anoda cristata density, on crop and weed biomass and weed seed production in wide (70 cm) and narrow rows (35 cm) glyphosate‐resistant soyabean (Glycine max). Soyabean density was higher at 35 cm row spacing as an increase in planting rate in narrow‐row soyabean is recommended for producers in Argentina. Soyabean biomass at growth stage V4 (four nodes on the main stem with fully developed leaves beginning with the unifoliate leaves) was higher when grown on narrow than in wide‐rows but was not affected by the presence of A. cristata. At growth stage R5 (seed initiation – seed 3 mm long in a pod at one of the four uppermost nodes on the main stem, with a fully developed leaf and full canopy development), crop biomass was greater in narrow rows compared with wide rows with 12 plants m^?2 of A. cristata. In narrow‐row soyabean, a single application of a reduced rate of glyphosate maintained soyabean biomass at R5 and provided excellent weed control regardless of weed density. In wide‐row soyabean control was reduced at the high weed density. Regardless of row spacing, A. cristata biomass and seed production were severely reduced by half of the recommended dose rate of glyphosate but the relationship between biomass and seed production was not altered. Glyphosate rates as low as 67.5 g a.e. ha^?1 in narrow rows or 540 g a.e. ha^?1 in wide rows provided excellent control of A. cristata. To minimize glyphosate use, planting narrow‐row soyabean are effective where A. cristata density is low.     

Improved chemical control of Conyza bonariensis in wheat limits problems in the following fallow

Conyza bonariensis is a major weed infesting zero‐tilled cropping systems in subtropical Australia, particularly in wheat and winter fallows. Uncontrolled C. bonariensis survives to become a problem weed in the following crops or fallows. As no herbicide has been registered for C. bonariensis in wheat, the effectiveness of 11 herbicides, currently registered for other broad‐leaved weeds in wheat, was evaluated in two pot and two field experiments. As previous research showed that the age of C. bonariensis, and to a lesser extent, the soil moisture at spraying affected herbicide efficacy, these factors also were investigated. The efficacy of the majority of herbicide treatments was reduced when large rosettes (5–15 cm diameter) were treated, compared with small rosettes (<5 cm diameter). However, for the majority of herbicide treatments, the soil moisture did not affect the herbicide efficacy in the pot experiments. In the field, a delay in herbicide treatment of 2 weeks reduced the herbicide efficacy consistently across herbicide treatments, which was related to weed age but not to soil moisture differences. Across all the experiments, four herbicides controlled C. bonariensis in wheat consistently (83–100%): 2,4‐D; aminopyralid + fluroxypyr; picloram + MCPA + metsulfuron; and picloram + high rates of 2,4‐D. Thus, this problem weed can be effectively and consistently controlled in wheat, particularly when small rosettes are treated, and therefore C. bonariensis will have a less adverse impact on the following fallow or crop.     

Genetic diversity and phylogeny of strains of Clavibacter nebraskensis associated with recent and historic Goss's wilt epidemics in the north Central USA

Goss's wilt and blight of maize, caused by Clavibacter nebraskensis, is an important disease in the USA and Canada. The re-emergence of Goss's wilt in the mid-2000s and the subsequent spread of this disease probably resulted from changes in crop management practices and possibly changes in the pathogen population. The objectives of this study were to determine the genetic diversity and phylogeny of strains of C. nebraskensis associated with recent and historic Goss's wilt epidemics in the north Central USA, to measure the effects of collection period on population structure, and to identify the phylogenetic status of avirulent strains of C. nebraskensis. Multilocus sequence analysis and typing were used to address these objectives. Bayesian analysis of the sequences of atpD, dnaK, and kdpA separated the type strain of C. nebraskensis from type or reference strains of three other Clavibacter spp. The 125 strains of C. nebraskensis used in this study formed a monophyletic taxon. Four lineages were identified. Clades I, II, and IV contained a high proportion of highly virulent strains from the Upper Midwest. Clade III contained a high proportion of strains from Nebraska, Colorado, and South Dakota collected between 1969 and 1998. A significant substructure was detected between subpopulations from historic outbreaks in Nebraska and Colorado and more recent outbreaks in Minnesota. Virulence or avirulence in maize was not correlated with a particular MLSA clade. The results support the hypothesis that genetic changes in the population of C. nebraskensis were correlated with recent area expansions of Goss's wilt.     

Physiological and Cytological Studies on the Inhibition of Striga Seed Germination by the Plant Growth-promoting Bacterium Azospirillum brasilense

Striga hermonthica (Del.) Benth. is an obligate parasitic weed of tropical cereals whose rhizosphere can also be colonised by bacteria of the genus Azospirillum. A previous study demonstrated that the two organisms (Azospirillum and striga) interacted during cerealroot colonisation. Two strains of A. brasilense isolated from an African sorghum rhizosphere prevented the germination of striga seeds although they were stimulated to germinate by the presence of sorghum roots. Azospirillum cells suspended in a synthetic germination stimulant (GR24) did not inhibit striga seed germination, but did block radicle elongation. Those radicles had an abnormal morphology, and contained no vacuolated cells in the root elongation zone. Lipophilic compounds extracted from the medium of bacteria in the log and stationary growth phases prevented the germination of striga seeds.     

Characteristics of seeds and seedlings from weeds treated with sublethal herbicide doses

Size, germination and viability of seeds as well as growth of seedlings derived from three weed species were studied in a pot trial. Fallopin convolvulus (L.) A. Loeve, Galium. spurium L. and Thlaspi arvense L. were treated with MCPA or tribenuron-methyl at four doses and at five growth stages, from seedling stage to flowering, In G. spurium subnormal doses of tribenuron-methyl, applied at intermediate growth stages, greatly reduced seed weight, gennination, viability, seedling shoot biomass and root biomass. Germination and viability, as well as the shoot biomass and root biomass of seedlings, were highly correlated with seed weight. In addition, but to a smaller extent, seed weight was reduced in F. convolvidus and T. arvense by tribenuron-methyl and in G. spurium by MCPA. Germination was reduced in F. convolvulus by MCPA and in T. arvense by tribenuron-methyl. However, the effects varied greatly depending on the growth stage at application.     

Predicting the competitive effects of weed and crop density on weed biomass, weed seed production and crop yield in wheat

Competition between winter-sown wheat and Viola arvensis Murray or Papaver rhoeas L. was studied in two experiments in two successive years. The effects of varying crop and weed density were modelled in terms of weed biomass over time, weed seed production and crop yield. Biomass model parameters, representing maximum weed biomass and intra- and interspecific competition, were obtained for different assessment dates, enabling biomass levels to be predicted during the two growing seasons. Weed biomass declined, and its maximum level was reached earlier, with increasing crop density. Intraspecific competition was higher in the absence than in the presence of crop, increasing with time and with weed density. Halving the wheat population increased June biomass of V. arvensis by 74% and of P. rhoeas by 63%. Crop yield losses with increasing weed density were greater with low than with medium and high crop populations. P. rhoeas was significantly more competitive than V. arvensis in both years. Weed biomass in 1989 responded more to reductions in crop density following the milder winter of 1988/89 than in the previous year; however crop yields were less affected in 1989 due to summer drought, restricting late weed growth and competition. Weed seed production was related to weed biomass; the progressive lowering of crop density increased seed production, and both species were very prolific in the absence of crop. By combining models, seed production could be derived for a given competitive effect on the crop. Threshold weed populations, based on low weed levels that are not economic to control, could then be equated with the accompanying weed seed production.