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Since the commercialization of herbicide-resistant (HR) crops, primarily glyphosate-resistant crops, their adoption has increased rapidly. Multiple herbicide resistance traits in crops such as canola (Brassica napus L.), corn (Zea mays L.), cotton (Gossypium hirsutum L.), and soybean [Glycine max (L.) Merr.] have become available in recent years, and management of their volunteers needs attention to prevent interference and yield loss in rotational crops. The objectives of this review were to summarize HR crop traits in barley (Hordeum vulgare L.), canola, corn, cotton, rice (Oryza sativa L.), soybean, sugarbeet (Beta vulgaris L.), and wheat (Triticum aestivum L.); assess their potential for volunteerism; and review existing literature on the interference of HR crop volunteers, yield loss, and their management in rotational crops. HR crop volunteers are problem weeds in agronomic cropping systems, and the impact of volunteerism depends on several factors, such as crop grown in rotation, the density of volunteers, management practices, and microclimate. Interference of imidazolinone-resistant (IR) barley or wheat volunteers can be a problem in rotational crops, particularly when IR crops such as canola or wheat are grown. HR canola volunteers are abundant in the Northern Great Plains due to high fecundity, seed loss before or during harvest, and secondary seed dormancy, and they can interfere in crops grown in rotation such as flax (Linum usitatissimum L.), field peas (Pisum sativum L.), and soybean. HR corn volunteers are competitive in crops grown in rotation such as corn, cotton, soybean, and sugarbeet, with yield loss depending on the density of HR corn volunteers. Volunteers of HR cotton, rice, soybean, and sugarbeet are not major concerns and can be controlled with existing herbicides. Herbicide options would be limited if the crop volunteers are multiple HR; therefore, recording the cultivar planted the previous year and selecting the appropriate herbicide are important. The increasing use of 2,4-D, dicamba, glufosinate, and glyphosate in North American cropping systems requires research on herbicide interactions and alternative herbicides or methods for controlling multiple HR crop volunteers.
Weed resistance surveys that monitor the spread of resistant weeds have mainly been conducted through time-consuming, labor-intensive, and destructive greenhouse herbicide screens. As an alternative, we introduce here a nondestructive leaf-disk assay based on chlorophyll fluorescence (Fv/Fm values that measure photosynthetic efficiency) that allows the detection of resistance to both systemic and contact herbicides within ∼48 h. The current study validated the assay for detecting resistance to fomesafen, glyphosate, and dicamba in Palmer amaranth (Amaranthus palmeri S. Watson), waterhemp [Amaranthus tuberculatus (Moq.) Sauer], kochia [Bassia scoparia (L.) A.J. Scott], and goosegrass [Eleusine indica (L.) Gaertn.]. Negative correlation between Fv/Fm values and spray injury levels was observed in all herbicide–weed combinations at the discriminating doses, except for glyphosate in Amaranthus. The correlation coefficients were –0.41 for fomesafen (10 µM, P < 0.0001) in Amaranthus, –0.92 for glyphosate in E. indica (250 µM, P < 0.0001), and –0.44 for dicamba in B. scoparia (800 µM, P = 0.0023). At the population level, the assay clearly separated susceptible from highly resistant populations. However, the assay showed lower sensitivity in distinguishing populations of different resistance levels or separating populations with low resistance from susceptible populations. At the individual plant level, results from the leaf-disk assay and whole-plant spray tests were concordant in 85.5%, 92.3%, and 71.7% of the plants tested for fomesafen–Amaranthus, glyphosate–Eleusine, and dicamba–Bassia, respectively. The assay yielded 1% to 15% false-positive and 6% to 13% false-negative results across herbicides. The current study demonstrated that the leaf-disk assay is a useful tool to identify weed resistance. Optimization is needed to improve its sensitivities and expand its usage to more diverse herbicide–weed species combinations.
Three resistant (R) rigid ryegrass (Lolium rigidum Gaudin) populations from southern Australia (EP162, 375-14, and 198-15) with cross-resistance to thiocarbamate, chloroacetamide, and sulfonylisoxazoline herbicides displayed reduced sensitivity to the isoxazolidinone herbicides bixlozone and clomazone. Each of these R populations was exposed to two cycles of recurrent selection (RS) in which plants were treated with the field rate of bixlozone, survivors were bulk crossed, and seed was collected. After the first cycle of recurrent selection (RS1), the LD50 to bixlozone in population 198-15 increased to 17.5-fold compared with the S population and increased further to 26.9-fold after a second cycle of recurrent selection (RS2). The recurrent selection process also increased the level of resistance to bixlozone in populations EP162 and 375-14 (7.8- to 18.4-fold) compared with the S population. Phorate antagonized bixlozone and clomazone in SLR4 (34.6- and 28.1-fold increase in LD50) and both herbicides in populations EP162 (36.5- to 46.6-fold), 375-14 (71.4- to 73.9-fold), and 198-15 (86.4- to 91.5-fold) compared with the absence of phorate. The increase in LD50 of all L. rigidum RS populations when treated with phorate suggests a lack of herbicide activation is not the likely resistance mechanism to these herbicides. This research highlights the elevated risk of thiocarbamate-resistant L. rigidum populations to rapidly evolve resistance to the isoxazolidinone herbicides bixlozone and clomazone.
Redroot pigweed (Amaranthus retroflexus L.) is a dominant weed in soybean [Glycine max (L.) Merr.] fields in Heilongjiang Province, China. High selective pressure caused by the extensive application of the protoporphyrinogen oxidase (PPO)-inhibiting herbicide fomesafen has caused A. retroflexus to evolve resistance to this herbicide. Two susceptible and two resistant populations (S1, S2, R1, and R2) were selected in this study to illustrate the target-site resistance mechanism in resistant A. retroflexus. Whole-plant bioassays indicated that R1 and R2 had evolved high-level resistance to fomesafen, with resistance factors of 27.0 to 27.9. Sequence alignment of the PPO gene showed an Arg-128-Gly substitution in PPX2. The basal expression differences of PPX1 and PPX2 between the S1 and R1 plants were essentially nonsignificant, whereas the basal expression of PPX2 in R2 plants was slightly lower than in S1 plants. Compared with the PPX1 gene, the PPX2 gene maintained higher expression in the resistant plants after treatment with fomesafen. An enzyme-linked immunosorbent assay showed a similar basal PPO content between the susceptible and resistant plants without treatment. After fomesafen treatment, the PPO content decreased sharply in the susceptible plants compared with the resistant plants. Furthermore, after 24 h of treatment, the resistant plants showed increased PPO content, whereas the susceptible plants had died. The PPO2 mutation resulted in high extractable PPO activity and low sensitivity to fomesafen along with changes in PPO enzyme kinetics. Although the mutant PPO2 exhibited increased Km values in the resistant plants, the Vmax values in these plants were also increased. Changes in the properties of the PPO enzyme due to an Arg-128-Gly substitution in PPX2, including changes in enzyme sensitivity and enzyme kinetics, are the target-site mechanism of resistance in A. retroflexus.
The parasitic plant purple witchweed [Striga hermonthica (Delile) Benth.] poses a serious threat to cereal production in sub-Saharan Africa. Under natural infestation, the wild pearl millet [Pennisetum glaucum (L.) R. Br.] line 29Aw demonstrates resistance against the parasite, but the mechanism of its resistance is unknown. Striga resistance can be due to: (1) low induction of Striga germination (pre-attachment resistance) or (2) inhibition of parasite attachment and development (post-attachment resistance). Germination bioassays and root chamber (rhizotron) resistance-screening assays were used to determine the extent of pre- and post-attachment Striga resistance in 29Aw compared with the Striga-susceptible ‘SOSAT-C88-P10’ variety. Regarding pre-attachment resistance, 29Aw stimulated 10-fold less Striga seed germination at a maximum germination distance of 7.96 ± 2.75 mm from the host root compared with 35.94 ± 2.88 mm in SOSAT-C88-P10. Post-attachment resistance revealed 10- to 19-fold fewer and 2.5-fold shorter Striga seedlings with 28-fold less biomass growing on 29Aw compared with SOSAT-C88-P10. Microscopic analysis showed that Striga penetration in 29Aw was blocked at the endodermal and cortical levels. Post-attachment resistance in 29Aw was further supported by fewer (22%) Striga–host vascular connections in 29Aw compared with 79% in SOSAT-C88-P10. Together, these findings demonstrate that 29Aw harbors both pre- and post-attachment resistance mechanisms against S. hermonthica.
Invasive species face new selective pressures and low genetic variation caused by genetic bottlenecks and founder effects when they are introduced into novel environments. Epigenetic variation may help them to cope with these problems. Mile-a-minute (Mikania micrantha Kunth) is a highly invasive exotic weed that has seriously damaged biodiversity and agricultural ecosystems. We first adopted methylation-sensitive amplified polymorphism (MSAP) markers to investigate epigenetic variation of 21 M. micrantha populations in southern China, and further explored the effects of environmental factors on population epigenetic differentiation by correlating epigenetic and climate and soil data. Adaptive epiloci positively correlated with climate/soil variables were identified. Minimum temperature of the coldest month and mean temperature of the coldest quarter were considered as decisive factors for its distribution. Climate is presumed to play a relatively more important role than soil in shaping the adaptive epigenetic differentiation in M. micrantha. Under ongoing global warming, populations of M. micrantha are predicted to expand northward. In addition, the weed also presented higher epigenetic variation compared with genetic variation. Leaf shape variation was detected related to methylation-state change at the population level.
Redroot pigweed (Amaranthus retroflexus L.) and slender amaranth (Amaranthus viridis L.) are considered emerging problematic weeds in summer crops in Australia. An outdoor pot experiment was conducted to examine the effects of planting time on two populations of A. retroflexus and A. viridis at the research farm of the University of Queensland, Australia. Both species were planted every month from October to January (2017 to 2018 and 2018 to 2019), and their growth and seed production were recorded. Although both weeds matured at a similar number of growing degree days (GDD), they required a different number of days to complete their life cycles depending on planting date. The growth period was reduced and flowering occurred sooner as both species experienced cooler temperatures and shorter daylight hours. Both species exhibited increased height, biomass, and seed production for the October-sown plants compared with other planting times, and these parameters were reduced by delaying the planting time. The shoot and root biomass of A. retroflexus and A. viridis (averaged over both populations) was reduced by more than 70% and 65%, respectively, when planted in January, in comparison to planting in October. When planted in October, A. retroflexus and A. viridis produced 11,350 and 5,780 seeds plant–1, but these were reduced to 770 and 365 seeds plant–1 for the January planting date, respectively. Although the growth and fecundity of these species were dependent on planting time, these weeds could emerge throughout the late spring to summer growing season (October to March) in southeast Australia and could produce a significant number of seeds. The results showed that when these species emerged in the late spring (October), they grew vigorously and produced more biomass in comparison with the other planting dates. Therefore, any early weed management practice for these species could be beneficial for minimizing the subsequent cost and energy inputs toward their control.
Carlos Sousa-Ortega, Aritz Royo-Esnal, Iñigo Loureiro, Ana I. Marí, Juan A. Lezáun, Fernando Cordero, Milagros Saavedra, José A. Paramio, José L. Fernández, Joel Torra, José M. Urbano
Winter wild oat [Avena sterilis ssp. ludoviciana (Durieu) Gillet & Magne; referred to as A. sterilis here] is one of the major weed species of the Avena genus, given its high competitive ability to infest cereal crops worldwide, with special concern in Spain. A nine-location field experiment was established across Spain where a total of 400 A. sterilis seeds per location were sowed in four replicates in autumn 2016 to monitor the emergence during two growing seasons in dryland conditions. The data were used to test the prediction ability of previously published thermal (TT) and hydrothermal time (HTT) models and to develop new models, if required. Overall, the average percentage of emergence was 30% during the first season and 21% during the second season. In both seasons, the main emergence flush occurred between November and February. According to the phenological stage, A. sterilis achieved the tillering earlier in southern sites, between November 25 and the end of December, compared with northern sites, where this stage was reached at the end of January. The newly developed model described the emergence with precision, using three cardinal temperatures to estimate the TT. The three cardinal points were established at –1.0, 5.8, and 18.0 C for base (Tb), optimum (To), and ceiling temperature (Tc), while the base water potential (Ψb) was established at –0.2 MPa for the HTT estimation. This study contributes to improving prediction of the emergence of A. sterilis and provides knowledge for decision support systems (DSS) for the control of this weed.
In central Europe, barnyardgrass [Echinochloa crus-galli (L.) P. Beauv.], has commonly been found in humid lowland areas. As a result of the introduction of new crops and farming practices, in the northwest Carpathians, E. crus-galli has spread from lowland (<200 m altitude) to highland (>400 m altitude) areas. We collected seed samples from local populations lying at a distance of approximately 5 km from each other and lined up along transects following the flows of two rivers. The rivers first flow through the valleys separated by mountain ridges and eventually flow into a common lowland. After ripening, the seeds of all populations were germinated at 25 C under long-day conditions. Only the seeds of some lowland populations germinated up to 75%. The frequency of germinated seeds decreased as the altitude where the population was collected increased, and above 200 m above sea level, germination was mostly zero. We then studied the phenological and morphological differentiation of plants from the original (lowland) and recently occupied (highland) areas. Seeds of the lowest and the highest localities lying on the transect of each river were sown in a common garden experiment. In plants from the highland localities, heading and seed dispersal were earlier, while tiller height and tiller mass were lower than in plants from the lowland localities. Seed mass produced per tiller in the lowland and highland plants was similar, and as a result, highland plants allocated a larger proportion of body mass to seed production than did lowland plants. Echinochloa crus-galli populations from highland localities thus produce their progeny earlier and at a lower energy cost than populations from lowland localities. The plasticity of phenological characters likely facilitated adaptation during E. crus-galli spread from lowlands to highlands. Similar adaptations in plant phenology may contribute to the spread of E. crus-galli in other geographic areas.
Weed emergence time and the longevity of weed seeds within the soil play an important role in implementing a timely and effective weed control program. In this study, the seed longevity and emergence pattern of wild oat (Avena fatua L.) and sterile oat [Avena sterilis ssp. ludoviciana (Durieu) Gillet & Magne] were monitored in field conditions. Fresh seeds of A. fatua and A. ludoviciana were placed into nylon bags (50 seeds per bag in three replications for three locations in eastern Australia: Gatton, Narrabri, and St George) and buried at depths of 0, 2, and 10 cm in November 2017. Bags were exhumed at 6-mo intervals over 30 mo to evaluate seed germination, viability, and decay components. The seed decay component of A. fatua and A. ludoviciana followed an exponential pattern. On both the surface and at the 10-cm burial depth, 50% of the seeds of A. fatua and A. ludoviciana had decayed by 6 mo. The seeds of A. fatua persisted longer at 2-cm depth than at other depths, particularly at St George, where 90% of the seeds decayed after the 30-mo study. However, at Gatton and Narrabri, 90% of the seeds of A. fatua at this depth had decayed after 18 mo of burial in the soil. In the emergence pattern experiment (2017 to 2019), the emergence of A. fatua and A. ludoviciana from different burial depths was also studied. The emergence of A. fatua and A. ludoviciana was greater from 2-cm (29% to 36%) and 5-cm (18% to 43%) soil depths compared with the surface (5% to 10%) and 10-cm (3-9%) soil depth. Avena ludoviciana emerged earlier (2,253 growing degree days [GDD]; March 14, 2018) than A. fatua (3,364 GDD; May 23, 2018). Both species exhibited high emergence between May to June 2018, and the last cohort of each species was observed in October 2018. The highest seedling emergence occurred at the start of the winter season (May), which emphasizes the need for early PRE weed control such as tillage, herbicide application, and cover crops to ensure crops are planted in a clean seedbed. The continued emergence of these weeds into the spring season (October) emphasizes the need for extended periods of A. fatua and A. ludoviciana management. The results also suggest that management strategies that can control all emerged seedlings over 2 yr and restrict seed rain in the field could lead to complete control of Avena spp. in the field.
Crop–weed interactions are affected by environmental alterations resulting from a crop's presence, such as modifications in temperature, light quality and quantity, and moisture conditions that could modify weed performance. The objectives of this work were to study (1) how soybean [Glycine max (L.) Merr.] crop structure modifies the environment under the canopy and large crabgrass [Digitaria sanguinalis (L.) Scop.] plant structure, biomass, and seed production and dormancy; and (2) the relative importance of these environmental changes on the weed's characteristics. A field experiment in a completely randomized block design with five replicates was performed to evaluate narrow and wide interrow spacing and soybean maturity groups 3 and 4. Measured variables were intercepted solar radiation (RAD); red–far red (R-FR) ratio; humidity; minimum, maximum, and alternating temperatures; and weed biomass, tillers per plant, height, and seed dormancy. Crop canopy reduced solar radiation, R-FR ratio, and daily average maximum and alternating temperatures. Soybean presence reduced the weed biomass, tillers and seeds per plant, and seed dormancy. High solar radiation intercepted by the crop during the reproductive phase was the main environmental variable related to reductions in weed biomass, tillers per plant, and fecundity. The combination of low temperature and solar radiation received by developing seeds was more related to seed dormancy than the rest of the variables. Crop management decisions focused on the fact that keeping the crop canopy alive for a longer time at the end of the season would not only reduce the weed growth but also seed dormancy.
Ryegrass (Lolium spp.) is a troublesome weed in major wheat (Triticum aestivum L.) production regions in the United States. High diversity and adaptive potential are known to contribute to its success as a weed species and also create difficulties in correct species identification in fields. The objective of this research was to characterize diversity for 16 different morphological traits among 56 Lolium populations collected from wheat production fields across the Texas Blackland Prairies region and identify Lolium species based on taxonomic characteristics. Populations were highly diverse (both at inter- and intrapopulation levels) for the traits studied, and a taxonomic comparison with USDA-GRIN reference samples revealed that all the populations were variants of Italian ryegrass [Lolium perenne L. ssp. multiflorum (Lam.) Husnot] with a few offtypes of perennial ryegrass (Lolium perenne L.) or probable hybrids between the two species. Hierarchical clustering grouped the populations into six clusters based on their similarities for the morphological traits investigated. Principal component analysis showed that the variability for yield traits greatly contributed to the total diversity. Pre-flowering plant height (stage 10 on Feekes scale) was positively correlated with tiller count, shoot biomass, and spike count, but not with total seed count per plant, whereas plant height at maturity (stage 11.3 to 11.4 on Feekes scale) was highly correlated with total seeds per plant. Further, basal node color was positively correlated with plant growth habit, regrowth rate, and leaf color. Leaf blade width was positively correlated with survival to pinoxaden and multiple herbicides, whereas, spike count was negatively correlated with survival to mesosulfuron. The high levels of intra- as well as interpopulation variability documented in this study indicate the potential of this species to rapidly adapt to herbicides and emphasize the need for implementing diverse management tactics, including the integration of harvest weed seed control.
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