Development of herbicide resistance in weeds in a crop rotation with acetolactate synthase‐tolerant sugar beets under varying selection pressure

The development of acetolactate synthase (ALS) tolerant sugar beet provides new opportunities for weed control in sugar beet cultivation. The system consists of an ALS?inhibiting herbicide (foramsulfuron + thiencarbazone‐methyl) and a herbicide‐tolerant sugar beet variety. Previously, the use of ALS‐inhibitors in sugar beet was limited due to the susceptibility of the crop to active ingredients from this mode of action. The postulated benefits of cultivation of the ALS‐tolerant sugar beet are associated with potential risks. Up to now, with no relevant proportion of herbicide‐tolerant crops in Germany, ALS‐inhibitors are used in many different crops. An additional use in sugar beet cultivation could increase the selection pressure for ALS‐resistant weeds. To evaluate the impact of varying intensity of ALS‐inhibitor use on two weed species (Alopecurus myosuroides and Tripleurospermum perforatum) in a crop rotation, field trials were conducted in Germany in two locations from 2014 to 2017. Weed densities, genetic resistance background and crop yields were annually assessed. The results indicate that it is possible to control ALS‐resistant weeds with an adapted herbicide strategy in a crop rotation including herbicide‐tolerant sugar beet. According to the weed density and species, the herbicide strategy must be extended to graminicide treatment in sugar beet, and a residual herbicide must be used in winter wheat. The spread of resistant biotypes in our experiments could not be attributed to the integration of herbicide‐tolerant cultivars, although the application of ALS‐inhibitors promoted the development of resistant weed populations. Annual use of ALS‐inhibitors resulted in significant high weed densities and caused seriously yield losses. Genetic analysis of surviving weed plants confirmed the selection of ALS‐resistant biotypes.     

Establishment of Lolium species resistant to acetolactate synthase‐inhibiting herbicide in and around grain‐importation ports in Japan

Compared with natural seed dispersal, human‐mediated seed dispersal could spread herbicide resistance genes on a much larger scale. Herbicide‐resistant weed seeds have been reported as contaminants in commercial grain. We investigated the contamination of seeds of Lolium species with target‐site mutations conferring resistance to acetolactate synthase (ALS)‐inhibiting herbicides in wheat imported from the USA, Canada and Australia into Japan. We also investigated the establishment of ALS‐inhibiting herbicide‐resistant Lolium species in 12 seaports in Japan that are major entry points for international commodities. We found herbicide‐resistant Lolium spp. seeds from all classes of wheat samples. Resistant individuals became established at six of eight ports where more than 50 kt of imported wheat is unloaded every year. The establishment of resistant Lolium spp. individuals was common at major grain landing ports. Monitoring over 3 years at one port revealed that the frequency of resistant individuals did not fluctuate between years. Many resistant individuals were distributed in front of the entrance of a fodder company, but a few resistant individuals were found in areas 2 km away from the port. The results indicate that gene flow is rare through pollen or seed movement from resistant plants to peripheral populations. Further extensive and long‐term monitoring is necessary to perform a comprehensive risk assessment of herbicide‐resistant plants entering Japan through major commercial ports.     

Selectivity of SMY 1500 (4-amino-6-tert-butyl-3-ethylthio-l,2,4,triazin-5(4H)-one) in Triticum durum

Susceptibility to SMY 1500 (4-amino-6-tert-butyl-3-ethylthio-1.2.4-triazin-5(4H)-one) was studied in durum wheat (Triticum durum Desf.) cultivars in growth-chamber assays. Weight reduction, photosynthesis inhibition and herbicide levels were determined at various times after a 24-h herbicide treatment in two cultivars of durum wheat, one cultivar of spring wheat (Triticum aestivum L). and in the weed Bromus diandrus (Roth). Measurements of weight reduction distinguished between plants tolerant and plants susceptible to SMY 1500. Tolerant cultivars showed a lower herbicide content and photosynthesis inhibition than susceptible plants after the 24-h treatment. Thereafter, herbicide levels and photosystem II inhibition decreased faster in tolerant plants. Comparison with previous work showed that plants tolerated doses of SMY 1500 about four times higher than of its analogue, metribuzin. Herbicide content in leaves just after treatment, and rate of detoxification may explain the selectivity of SMY 1500 in these cereals.     

Resistance to glyphosate in Lolium rigidum

Annual ryegrass (Lolium rigidum) is a widespread and important weed of Australia and populations of this weed have developed resistance to most major herbicides, including glyphosate. The possible mechanisms of resistance have been examined in one glyphosate-resistant Lolium population. No major differences were observed between resistant and susceptible biotypes in respect of (i) the target enzyme (EPSP synthase), (ii) DAHP synthase, the first enzyme of the target (shikimate) pathway, (iii) absorption of glyphosate, or (iv) translocation. Following treatment with glyphosate, there was greater accumulation of shikimate (derived from shikimate-3-Pi) in susceptible than in resistant plants. In addition, the resistant population exhibited cross-resistance to 2-hydroxy-3-(1,2,4-triazol-1-yl)propyl phosphonate, a herbicide which, although structurally similar to glyphosate, acts at an unrelated target site. On the basis of these observations we speculate that movement of glyphosate to its site of action in the plastid is involved in the resistance mechanism. © 1999 Society of Chemical Industry     

Multiple herbicide resistance in littleseed canarygrass (Phalaris minor): A threat to wheat production in India

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.     

You cannot fight fire with fire: a demographic model suggests alternative approaches to manage multiple herbicide‐resistant Avena fatua

Multiple herbicide‐resistant (MHR ) weed populations pose significant agronomic and economic threats and demand the development and implementation of ecologically based tactics for sustainable management. We investigated the influence of nitrogen fertiliser rate (56, 112, 168, or 224 kg N ha^?1) and spring wheat seeding density (67.3 kg ha^?1 or 101 kg ha^?1) on the demography of one herbicide susceptible and two MHR Avena fatua populations under two cropping systems (continuous cropping and crop‐fallow rotation). To represent a wide range of environmental conditions, data were obtained in field conditions over 3 years (2013–2015). A stochastic density‐dependent population dynamics model was constructed using the demographic data to project A. fatua populations. Elasticity analysis was used to identify demographic processes with negative impacts on population growth. In both cropping systems, MHR seedbank densities were negatively impacted by increasing nitrogen fertilisation rate and wheat density. Overall, MHR seedbank densities were larger in the wheat‐ fallow compared with the continuous wheat cropping system and seedbank densities stabilised near zero in the high nitrogen and high spring wheat seeding rate treatment. In both cropping systems, density‐dependent seed production was the most influential parameter impacting population growth rate. This study demonstrated that while the short‐term impact of weed management tactics can be investigated by field experiments, evaluation of long‐term consequences requires the use of population dynamics models. Demographic models, such as the one constructed here, will aid in selecting ecologically based weed management tactics, such as appropriate resource availability and modification to crop competitive ability to reduce the impact of MHR .     

Observations on the physiological basis for the selectivity of morfamquat to graminaceous plants

The bipyridylium herbicide morfamquat damages dicotyledon plants in a similar way to paraquat but unlike paraquat, has little activity on Gramineae such as cocksfoot, wheat and maize. Results are presented which show that this selective action is not due to differences in the amounts of chemical taken up by treated leaves of susceptible and resistant plants or to differences in the degree of movement in plants. Freeze-thaw experiments show that the plasmalemma and tonoplast membranes of sensitive and resistant weeds are less permeable to morfamquat than to paraquat. Whilst this could account for differences in activity of the two herbicides on cocksfoot, further evidence is given which shows that selectivity is not due to differences in morfamquat movement from free space to cytoplasm. Differential movement from cytoplasm into the chloroplasts of resistant and susceptible species may be the basis of the selective action, but several other mechanisms of selectivity are also mentioned.     

Reduced paraquat translocation in paraquat resistant Arctotheca calendula (L.) Levyns is a consequence of the primary resistance mechanism, not the cause

Resistance to paraquat has been studied in detail in many weed species for more than a decade, with the precise mechanism of resistance still unclear. Several studies have indicated that reduced movement of the herbicide to the site of action in the chloroplast is at least partly responsible for endowing resistance. Although paraquat translocation studies have been performed in the past it has been rare for these studies to have been conducted on whole plants in the light, despite early observations which clearly showed that paraquat translocation is minimal unless treated plants are exposed to light. This study has addressed this issue in Arctotheca calendula by tracing the movement of ¹⁴C-paraquat in resistant and susceptible plants in both the dark and light. Differences in paraquat translocation between the resistant and susceptible biotypes of A. calendula were only observed when treated plants were exposed to light. It was observed that paraquat translocation was significantly reduced in the resistant compared to the susceptible biotype when plants were exposed to light but not in the dark. It is postulated that paraquat translocation is dependent on light mediated damage. As paraquat-induced damage is less severe in paraquat resistant plants, overall paraquat translocation is reduced in the resistant biotype.     

Modelling effects of vector acquisition threshold on disease progression in a perennial crop following deployment of a partially resistant variety

Deployment of resistant varieties is a key strategy to mitigating economic losses due to arthropod‐transmitted plant pathogens of perennial crops. In many cases, the best available resistant traits for introgression confer only partial resistance. Plants displaying partial resistance have lower pathogen titres than susceptible counterparts, but remain hosts for the pathogen. As partially resistant varieties maintain yield after infection, infected plants are unlikely to be rogued (i.e. removed). Accordingly, there is a risk that partially resistant plants could serve as a source of inoculum for pathogen spread to susceptible plants. Here, a mathematical model that tracked spread of an arthropod‐transmitted pathogen in a plant population consisting of susceptible and partially resistant plants was used to identify a threshold acquisition rate from partially resistant plants that resulted in limited spread of the pathogen from partially resistant plants to susceptible plants. The acquisition threshold from partially resistant plants varied with parameters influenced by disease management decisions such as number of vectors per plant, vector turnover, replacement of susceptible plants, and proportion of plants that were partially resistant. In model simulations, effects of deploying a partially resistant variety on disease incidence in a susceptible variety depended on the extent to which pathogen spread among susceptible plants was suppressed and acquisition rates from partially resistant plants. Collectively, the results indicate that risk of partially resistant plants serving as inoculum sources could be assessed prior to deployment, thereby enabling design of complementary disease management tactics to minimize economic losses in susceptible varieties following deployment.     

Glyphosate resistance in four different populations of Lolium rigidum is associated with reduced translocation of glyphosate to meristematic zones

Weed populations with resistance to glyphosate have evolved over the last 7 years, since the discovery of the first glyphosate‐resistant populations of Lolium rigidum in Australia. Four populations of L. rigidum from cropping, horticultural and viticultural areas in New South Wales and South Australia were tested for resistance to glyphosate by dose–response experiments. All populations required considerably more glyphosate to achieve 50% control compared with a known susceptible population, indicating they were resistant to glyphosate. Translocation of glyphosate within these resistant populations was examined by following the movement of radiolabelled glyphosate applied to a mature leaf. All resistant plants translocated significantly more herbicide to the tip of the treated leaf than did susceptible plants. Susceptible plants translocated twice as much herbicide to the stem meristematic portion of the plant compared with resistant plants. These different translocation patterns suggest an association between glyphosate resistance in L. rigidum and the ability of glyphosate to accumulate in the shoot meristem.     

Modelling herbicide dose and weed density effects on crop:weed competition

The effects of a range of herbicide doses on crop:weed competition were investigated by measuring crop yield and weed seed production. Weed competitivity of wheat was greater in cv. Spark than in cv. Avalon, and decreased with increasing herbicide dose, being well described by the standard dose–response curve. A combined model was then developed by incorporating the standard dose–response curve into the rectangular hyperbola competition model to describe the effects of plant density of a model weed, Brassica napus L., and a herbicide, metsulfuron‐methyl, on crop yield and weed seed production. The model developed in this study was used to describe crop yield and weed seed production, and to estimate the herbicide dose required to restrict crop yield loss caused by weeds and weed seed production to an acceptable level. At the acceptable yield loss of 5% and the weed density of 200 B. napus plants m^–2, the model recommends 0.9 g a.i. metsulfuron‐methyl ha^–1 in Avalon and 2.0 g a.i. in Spark.     

Multiple resistance of Papaver rhoeas L. to 2,4‐D and acetolactate synthase inhibitors in four European countries

The issue of cross‐ or multiple resistance to acetolactate synthase (ALS) inhibitors and the auxinic herbicide 2,4‐D was investigated in Papaver rhoeas L., a common and troublesome weed in winter cereals, in a broad‐scale study across four European countries. A combination of herbicide sensitivity bioassays and molecular assays targeting mutations involved in resistance was conducted on 27 populations of P. rhoeas originating from Greece (9), Italy (5), France (10) and Spain (3). Plants resistant to the field rate of 2,4‐D were observed in 25 of the 27 populations assayed, in frequencies ranging from 5% to 85%. Plants resistant to ALS‐inhibiting herbicides (sulfonylureas) were present in 24 of the 27 populations, in frequencies ranging from 4% to 100%. Plants resistant to 2,4‐D co‐occurred with plants resistant to sulfonylureas in 23 populations. In four of these, the probability of presence of plants with cross‐ or multiple resistance to 2,4‐D and sulfonylureas was higher than 0.5. ALS genotyping of plants from the field populations or of their progenies, identified ALS alleles carrying a mutation at codon Pro197 or Trp574 in 2,4‐D‐sensitive and in 2,4‐D‐resistant plants. The latter case confirmed multiple resistance to 2,4‐D and ALS inhibitors at the level of individual plants in all four countries investigated. This study is the first to identify individual plants with multiple resistance in P. rhoeas, an attribute rarely assessed in other weed species, but one with significant implications in designing chemical control strategies.     

Growth and yield responses of Italian ryegrass (Lolium multiflorum) to diclofop-methyl and ozone

We evaluated the combined effects of diclofop‐methyl herbicide application and the air pollutant ozone (O₃) on diclofop‐methyl‐resistant and ‐susceptible biotypes of Italian ryegrass (Lolium multiflorum). We conducted two experiments, one with a long vegetative growth period and the other with a short vegetative growth in late spring with seed production in summer. As expected, because of its phytotoxicity, the herbicide alone reduced total vegetative biomass, leaf area, tiller number and seed production at most sampling periods in susceptible plants for both experiments. However, it had variable effects on resistant plants, including a positive effect on seed production. Ozone alone delayed vegetative biomass accumulation and reduced leaf area and seed biomass in both experiments. However, the effects of O₃ on some parameters were altered by herbicide rate and/or biotype. Especially notable was a greater reduction in seed biomass because of O₃ in resistant than in susceptible plants with no herbicide. If these apparent differential responses to herbicide and O₃ stress of susceptible and resistant plants are confirmed and persist over time, evolutionary tradeoffs may occur. For example, the frequency of resistant plants in a population may be altered in response to interactions between herbicides and other anthropogenic stresses.     

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.     

Competition between Imperata cylindrica and maize in the forest savannah transition zone of Nigeria

Imperata cylindrica is a noxious weed that infests annual and perennial crops in most tropical regions. High crop densities may offer opportunities to reduce I. cylindrica competition in small‐scale farming systems. The competitive ability of maize relative to I. cylindrica was evaluated in an addition series experiment in the forest savannah transition zone in 2006 and 2007 at Ibadan, Nigeria. Maize and I. cylindrica were planted in eight monoculture densities (4, 8, 12, 16, 20, 32, 48 and 64 plants m^?2) and in a 1:1 mixture at eight total densities (2:2, 4:4, 6:6, 8:8, 10:10, 16:16, 24:24 and 32:32 maize: I. cylindrica plants m^?2) as in monoculture. Non‐linear regression models were used to relate crop and weed shoot biomass to their densities and total grain yield to maize density. In maize, intraspecific competition was more than interspecific competition; in I. cylindrica, interspecific competition was higher than intraspecific. As expected, total grain yield was lower in the mixture than in maize monoculture at all total densities. Average maize grain yield in maize monoculture differed from that in mixtures by 0.77 t ha^?1 in 2006 and 0.57 t ha^?1 in 2007. Niche differentiation indices were <1 in 2006 and >1 in 2007, indicating that both species competed for similar resources in 2006, but not in 2007. The greater competitive ability of maize over I. cylindrica may be associated with rapid growth and canopy development observed in the field.     

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.     

Modelling the effects of sub-lethal doses of herbicide and nitrogen fertilizer on crop–weed competition

The effects of sub‐lethal dose of herbicide and nitrogen fertilizer on crop–weed competition were investigated. Biomass increases of winter wheat and a model weed, Brassica napus, at no‐herbicide treatment with increasing nitrogen were successfully described by the inverse quadratic model and the linear model respectively. Increases in weed competitivity (β₀) of the rectangular hyperbola and parameter B in the dose–response curve for weed biomass, with increasing nitrogen were also successfully described by the exponential model. New models were developed by incorporating inverse quadratic and exponential models into the combined rectangular hyperbola with the standard dose–response curve for winter wheat biomass yield and the combined standard dose—response model with the rectangular hyperbola for weed biomass, to describe the complex effects of herbicide and nitrogen on crop–weed competition. The models developed were used to predict crop yield and weed biomass and to estimate the herbicide doses required to restrict crop yield loss caused by weeds and weed biomass production to an acceptable level at a range of nitrogen levels. The model for crop yield was further modified to estimate the herbicide dose and nitrogen level to achieve a target crop biomass yield. For the target crop biomass yield of 1200 g m^?2 with an infestation of 100 B. napus plants m^?2, the model recommended various options for nitrogen and herbicide combinations: 140 and 2.9, 180 and 0.9 and 360 kg ha^?1 and 1.7 g a.i. ha^?1 of nitrogen and metsulfuron‐methyl respectively.     

Frequency and distribution of herbicide resistance in Raphanus raphanistrum populations randomly collected across the Western Australian wheatbelt

In 2003, a random survey was conducted across the Western Australian wheatbelt to establish the extent and frequency of herbicide resistance in Raphanus raphanistrum populations infesting crop fields. Five hundred cropping fields were visited, with 90 R. raphanistrum populations collected, representative of populations present in crop fields throughout the Western Australian wheatbelt. Collected populations were screened with four herbicides of various modes of action that are commonly used for the control of this weed. The majority of Western Australian R. raphanistrum populations were found to contain plants resistant to the acetolactate synthase (ALS)‐inhibiting herbicide chlorsulfuron (54%) and auxin analogue herbicide, 2,4‐D amine (60%). This survey also determined that over half (58%) of these populations were multiple resistant across at least two of the four herbicide modes of action used in the screening. Only 17% of R. raphanistrum populations have retained their initial status of susceptibility to all four herbicides. The distribution patterns of the herbicide‐resistant populations identified that there were higher frequencies of resistant and developing resistance populations occurring in the intensively cropped northern regions of the wheatbelt. These results clearly indicate that the reliance on herbicidal weed control in cropping systems based on reduced tillage and stubble retention will lead to higher frequencies of herbicide‐resistant weed populations. Therefore, within intensive crop production systems, there is a need to diversify weed management strategies and not rely entirely on too few herbicide control options.     

Response of Sorghum halepense demographic processes to plant density and rimsulfuron dose in maize

In spatially heterogeneous weed infestations, variable dose technologies could be used to minimise herbicide use; high doses could be applied to reduce high‐density patches and low doses to maintain weed populations in low‐density portions of a field. To assess the potential short‐ and long‐term effects of variable herbicide dose and site‐specific management, the major weed demographic processes were described and parameterised in this study. Various doses of rimsulfuron (from 0 to 12.5 g a.i. ha^?1) were applied to different densities of Sorghum halepense (0–100 plants m^?2). Contrary to similar studies with other weed species, higher herbicide efficacy was not observed at low densities, suggesting that the same rimsulfuron dose should be applied regardless of the S. halepense density. The highest percentage of control was obtained with the full rimsulfuron dose. However, it did not guarantee a decrease of the infestation in the following season in the field areas where the initial S. halepense density was lower than 60 plants m^?2. Reduced doses of rimsulfuron to control S. halepense cannot be recommended based on our results.     

Inheritance of resistance to paraquat in barley grass Hordeum glaucum Steud.

Two biotypes of the grass weed barley grass (Hordeum glaucum), one resistant and the other susceptible to the herbicide paraquat, were studied along with their F₁, F₂ and F₃ progeny to determine the inheritance of paraquat resistance. The plants were sprayed with 50–200 g a.i. paraquat ha^?1. These concentrations killed the susceptible type. The data obtained from segregating populations indicated that paraquat resistance in H. glaucum is controlled by a single nuclear gene with incomplete dominance.