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Germination and emergence assays represent the most notable examples of time-to-event data in agriculture and related disciplines. In spite of the peculiar characteristics of this type of data, there has been little effort to establish a specific and comprehensive framework for their analyses. Indeed, a brief survey of the literature shows that germination and emergence data, along with other phenological measurements such as flowering time, have been analyzed through myriad approaches, giving rise to confusion and uncertainty among scientists and practitioners as to what may represent the best statistical practice. This lack of coherence in statistical approach may reduce the efficiency of research, while making the communication of results and the cross-study comparisons extremely challenging. Here, we attempt to provide a coherent framework and protocol for the analyses of germination/emergence and other time-to-event data in weed science and related disciplines, together with a software implementation in the form of a new R package. We propose a similar approach to biological assays in ecotoxicology, based on: (1) fitting a time-to-event model to describe the whole time course of events; (2) comparing time-to-event curves across experimental treatments, and (3) deriving further information from the fitted model to better focus on some traits of interest. The most appropriate methods to accomplish this procedure were carefully selected from the framework of survival analysis and related sources and were modified to comply with the specific needs of weed, seed, and plant sciences. Finally, they were implemented in the new R package drcte. In this article, we describe the procedure and its limitations by way of providing examples of several types of germination/emergence assays. We highlight that our proposed procedure can also serve as the first step of data analyses, with its output subsequently submitted to traditional or meta-analytic approaches.
Herbicide-resistant (HR) crops are widely grown throughout the United States and Canada. These crop-trait technologies can enhance weed management and therefore can be an important component of integrated weed management (IWM) programs. Concomitantly, evolution of HR weed populations has become ubiquitous in agricultural areas where HR crops are grown. Nevertheless, crop cultivars with new or combined (stacked) HR traits continue to be developed and commercialized. This review, based on a symposium held at the Western Society of Weed Science annual meeting in 2021, examines the impact of HR crops on HR weed management in the U.S. Great Plains, U.S. Pacific Northwest, and the Canadian Prairies over the past 25 yr and their past and future contributions to IWM. We also provide an industry perspective on the future of HR crop development and the role of HR crops in resistance management. Expanded options for HR traits in both major and minor crops are expected. With proper stewardship, HR crops can reduce herbicide-use intensity and help reduce selection pressure on weed populations. However, their proper deployment in cropping systems must be carefully planned by considering a diverse crop rotation sequence with multiple HR and non-HR crops and maximizing crop competition to effectively manage HR weed populations. Based on past experiences in the cultivation of HR crops and associated herbicide use in the western United States and Canada, HR crops have been important determinants of both the selection and management of HR weeds.
Acetolactate synthase (ALS) inhibitors provide postemergence control of green kyllinga (Kyllinga brevifolia Rottb.) in turfgrass and other cropping systems. A suspected resistant (R) biotype of K. brevifolia was collected from a golf course and evaluated for resistance to ALS inhibitors. In greenhouse experiments, the sulfosulfuron rates required to cause 50% shoot biomass reduction from the nontreated at 4 wk after treatment (WAT) were 10 and 792 g ai ha–1 for the susceptible (S) and R biotypes, respectively. The rates required to cause 50% injury at 4 WAT were 189 and >3,360 g ai ha–1, respectively. In other experiments, shoot mass of the R biotype was not reduced by imazaquin, trifloxysulfuron-sodium, pyrimisulfan, thiencarbazone + foramsulfuron + halosulfuron, florasulam + halauxifen-methyl, and bentazon compared with the nontreated, while sulfentrazone reduced biomass similarly for both R and S biotypes. Gene sequencing of the R biotype revealed a mutation at Asp-376-Glu that has previously conferred resistance to five families of ALS inhibitors. This is the first report of ALS-inhibitor resistance in K. brevifolia.
Ammannia multiflora Roxb. is a dominant broadleaf weed that is a serious problem in southern China rice fields, and acetolactate synthase (ALS)-inhibiting herbicides have been used for its control for more than 20 years. Excessive reliance on ALS-inhibiting herbicides has led to herbicide resistance in A. multiflora. In this study, 10 A. multiflora populations from the Jiangsu Province of China were collected, and the resistance levels and target site–resistance mechanisms to ALS-inhibiting herbicides bensulfuron-methyl and penoxsulam were investigated. The dose–response assays showed that eight populations evolved resistance to bensulfuron-methyl (9.1- to 90.9-fold) and penoxsulam (5.0- to 103.1-fold). Amplification of ALS genes indicated that there were three ALS genes (AmALS1, AmALS2, and AmALS3) in A. multiflora. Sequence analysis revealed amino acid mutations at Pro-197 in either AmALS1 (Pro-197-Ala, Pro-197-Ser, and Pro-197-His) or AmALS2 (Pro-197-Ser and Pro-197-Arg) in resistant populations, and no mutations were found in AmALS3. Moreover, two independent mutations (Pro-197-Ala in AmALS1 and Pro-197-Ser in AmALS2 or Pro-197-Ala in AmALS1 and Pro-197-Arg in AmALS2) coexisted in two resistant populations, respectively. In addition, the auxin mimic herbicides MCPA and florpyrauxifen-benzyl, the photosystem II inhibitor bentazon, and the protoporphyrinogen oxidase inhibitor carfentrazone-ethyl can effectively control the resistant A. multiflora populations. Our study demonstrates the wide prevalence of ALS inhibitor–resistant A. multiflora populations in Jiangsu Province and the diversity of Pro-197 mutations in ALS genes and provides alternative herbicide options for controlling resistant A. multiflora populations.
Jéssica F. L. Leal, Amanda dos S. Souza, Junior Borella, André Lucas S. Araujo, Ana Claudia Langaro, Monique M. Alves, Luana Jéssica S. Ferreira, Sarah Morran, Luiz H. S. Zobiole, Felipe R. Lucio, Aroldo F. L. Machado, Todd A. Gaines, Camila F. de Pinho
In this work, we evaluated the short time-induced oxidative stress–mediated rapid metabolic and physiological responses of resistant and susceptible Sumatran fleabane [Conyza sumatrensis (Retz.) E. Walker; syn.: Erigeron sumatrensis Retz.] to 2,4-D herbicide. Under fixed conditions (25 C and 65 ± 5% relative humidity), we assayed injury symptoms, chlorophyll a fluorescence, and antioxidative systems of biotypes both resistant and susceptible to 2,4-D (1,005 g ae ha–1). Under 15 versus 25 C temperatures and light and dark conditions, oxidative stress–mediated damage was assayed on plants that received 2,4-D herbicide applications. The injury symptoms observed in the 2,4-D–resistant biotype were rapid necrosis in leaves within 30 min, with the reestablishment of normal growth within 1 to 2 wk after 2,4-D treatment. The basal antioxidant enzyme activities of superoxide dismutase, catalase, and ascorbate peroxidase were greater in the resistant than in the susceptible biotype, although the activities of all enzymes generally did not differ between untreated and treated in the resistant biotype. The resistant biotype showed great reduction (at 1 and 4 h after application) in the photosynthetic electron transport chain performance index, while these metabolic changes were only detected after 4 h in the susceptible biotype. The resistant biotype recovered from the foliar damage 1 to 2 wk after 2,4-D application, while the susceptible biotype was controlled. The production of H2O2 was responsive to temperature and increased more rapidly in the 2,4-D–resistant biotype than in the susceptible one at both 15 and 25 C; however, there was a greater increase at 25 C in the resistant biotype. H2O2 production was not light dependent in 2,4-D–resistant C. sumatrensis, with increases even under dark conditions. The 2,4-D–resistant biotype showed rapid photosynthetic damage, possibly due to the rapid necrosis and leaf disruption, and increased H2O2 content compared with the susceptible biotype.
Herbicide resistance, documented in many economically damaging weed species, is a major threat to global crop production. The injudicious use of herbicides, often in the absence of diverse weed control strategies, poses an immense selection pressure on weed communities for resistance evolution and weed adaptive traits such as high seed dormancy. This study evaluates the interaction among developing herbicide resistance, seed size, and seed dormancy of ripgut brome (Bromus diandrus Roth), wild oat (Avena fatua L.), and hare barley [Hordeum leporinum Link; syn. Hordeum murinum L. ssp. leporinum (Link) Arcang.] collected from within intensively managed fields (in-crop) in comparison with populations in surrounding ruderal (non-crop disturbed) areas with no history of exposure to herbicides within the Western Australian grainbelt. Seed size of the three species varied by farming system (continuous cereal-intensive annual crops, diverse annual crops, pasture based) and habitat (in-crop, ruderal). Field populations of H. leporinum and B. diandrus tended to have greater seed size compared with ruderal populations. Larger seeds had significantly more dormancy in all three weed species. Field-collected populations that were exposed to herbicide applications for at least the past 5 yr exhibited significantly greater seed dormancy compared with their counterparts present in ruderal areas within the same geographic area. The association between increased seed dormancy and developing multiple herbicide resistance further complicates effective weed management.
Glyphosate-resistant (GR) horseweed [Conyza canadensis (L.) Cronquist; syn.: Erigeron canadensis L.] interference can substantially reduce corn (Zea mays L.) yield. The complementary activity of 4-hydroxyphenylpyruvate dioxygenase (HPPD) and photosystem II (PSII) inhibitors has been investigated for the control of several weed species, and in many cases has been synergistic; however, there is little information on the interaction of HPPD- and PSII-inhibiting herbicides for postemergence control of GR C. canadensis in corn. Four field trials were studied over 2 yr (2019, 2020) in Ontario, Canada, in commercial corn fields with natural infestations of GR C. canadensis to evaluate the level of GR C. canadensis control with three HPPD-inhibiting herbicides (mesotrione, tolpyralate, and topramezone) and three PSII-inhibiting herbicides (atrazine, bromoxynil, and bentazon) applied individually and in tank-mix combinations, and to document the interaction of the three HPPD inhibitors tank mixed with the three PSII inhibitors. Mesotrione, tolpyralate, and topramezone controlled GR C. canadensis 83%, 84%, and 72%, respectively, at 8 wk after application (WAA). Bromoxynil and bentazon controlled GR C. canadensis 71% and 79%, respectively, while atrazine provided only 31% control at 8 WAA. The joint application of atrazine, bromoxynil, or bentazon with mesotrione increased GR C. canadensis control from 83% to 100% at 8 WAA. Tolpyralate tank mixed with atrazine, bromoxynil, or bentazon controlled GR C. canadensis 96%, 98%, and 98%, respectively, which was comparable to the mesotrione tank mixes at 8 WAA. Topramezone plus atrazine, bromoxynil, or bentazon controlled GR C. canadensis 91%, 93%, and 95%, respectively, at 8 WAA. Interactions between HPPD and PSII inhibitors were synergistic for all combinations of mesotrione or tolpyralate with atrazine, bromoxynil, or bentazon. The interaction between topramezone and PSII inhibitors was additive. All nine tank mixes controlled GR C. canadensis >90%. This study concludes that bromoxynil or bentazon, instead of atrazine, can be co-applied with mesotrione, tolpyralate, or topramezone without compromising GR C. canadensis control in corn.
Benzobicyclon tolerance in rice (Oryza sativa L.) is dependent on the presence of a functional HIS1 gene, but the level of sensitivity might vary among different cultivars. Greenhouse, laboratory, and field experiments were conducted to further explore the role of HIS1 in cultivated rice tolerance and to exploit findings toward optimizing benzobicyclon activity on weedy rice (unwanted rice; Oryza sativa L.). In a heredity experiment, benzobicyclon tolerance was confirmed to be a semidominant trait conferred by HIS1 based on the intermediate response (ED50 values) of HIS1 heterozygous F1 plants. The spatial–temporal expression of HIS1 was next studied in tissue types (blades, sheaths, and whorls) across tolerant cultivars (‘Roy J’, ‘Diamond’, ‘LaKast’, ‘CLXL745’, and ‘XL753’) and growth stages (2- to 3- compared with 5- to 6-leaf). The relative expression of HIS1 was tissue specific and highest in whorls, followed by blades and then sheaths. Minimal differences in expression across cultivars and growth stages were observed. Furthermore, HIS1 was not largely upregulated at 6 h after benzobicyclon treatment. In the same experiment, cultivar tolerance to benzobicyclon at the label rate of 371 g ha–1 was found to be growth stage dependent. Plant growth was reduced by ∼35% when rice plants were at the 2- to 3- compared with 5- to 6-leaf growth stages. These results show that differences in benzobicyclon tolerance among HIS1 homozygous cultivars is likely not directly correlated with the expression of HIS1. In this research a model was proposed and supported by a field proof of concept study, indicating benzobicyclon efficacy on weedy rice is a function of HIS1 zygosity by growth stage at application. Prior research indicates HIS1 is the dominant allele in weedy rice accessions in Arkansas, and thus, based on our model, benzobicyclon should be applied to weedy rice with ≤2 leaves for suppression.
Leucaena [Leucaena leucocephala (Lam.) de Wit] is a perennial weed in more than 25 countries, including Australia. Knowledge regarding the seed biology of L. leucocephala could help in making weed management decisions. Experiments were conducted to study the effect of hot water (scarification), alternating temperatures, heat stress, salt stress, water stress, and burial depth on seed germination of two populations of L. leucocephala collected from Toowoomba and Gatton, Australia. The optimum duration of hot water treatment to break the hard seed coat dormancy was 2 min for both populations. The highest germination (92% to 98%) was recorded at 35/25 C for both populations, and similar germination occurred at 30/20 C. The Toowoomba population recorded greater germination at low temperature (15/5 to 25/15 C) than the Gatton population. Additionally, the Gatton population had higher germination than the Toowoomba population after 5 min of exposure to temperatures of up to 100 C, suggesting that the Gatton population may be more tolerant to heat stress. Germination was completely inhibited at pretreatment (5 min) temperatures of 150 to 250 C. The Toowoomba population recorded 17% greater germination than the Gatton population at a high salt concentration (160 mM NaCl), indicating its greater salt tolerance. At low moisture stress (–0.1 and –0.2 MPa), higher germination was observed in the Toowoomba population than in the Gatton population, whereas germination was similar for both populations at higher water stress levels (–0.4 MPa or lower). Germination was similar for both populations at shallow depths (0 and 1 cm) but higher emergence was recorded for the Toowoomba population at 2 to 8 cm than the Gatton population. Differential germination behaviors of both populations suggest that they adapted differently in their respective local environments. Knowledge gained from this study will help in formulating integrated management practices for L. leucocephala.
Automated guidance systems have advanced precise interrow hoeing in narrowly spaced cereals. Compared with other direct mechanical strategies, hoeing provides superior weed control and improved yields. However, weeds in the uncultivated intrarow zone may survive and compete intensely with the crop, causing yield loss. Therefore, improved intrarow weed management strategies in hoed cereals must be investigated. In spring barley (Hordeum vulgare L.), the effect of crop density was assessed at four levels (200, 300, 400, and 500 plants m–2); interrow spacing at two levels (15 and 20 cm), relevant to the abilities of current automated equipment to hoe between narrowly spaced rows; and weed management treatment at three levels (no additional controls, herbicide, and preemergence tine harrowing). All treatments received interrow hoeing, and a surrogate weed (white mustard, Sinapis alba L.) was sown and monitored throughout experiments. The manipulation of crop density was a reliable method for suppressing the growth of intrarow weeds. As barley density increased from the target 200 to 500 plants m–2, percent reduction in intrarow surrogate and ambient weed biomass (g m–2) increased from 49% to 82% and 53% to 99%, respectively. Increasing crop density caused a decrease in grain bulk density (kg hl–1) both years, and grain protein (%) and 1,000-kernel weight (g) in one year; whether these changes constitute a loss in grain quality depends upon end use. While row spacing had no effect on intrarow weeds, crop yields were 7% to 8% lower at 20 cm compared with 15 cm, incentivizing narrow row sowing. Barley yields were unaffected by increasing crop density, and the effect of preemergence tine harrowing was inconsistent. In one year, harrowing reduced surrogate and ambient weed biomass and increased barley yield; however, in another year, ambient weed biomass increased, and harrowing did not affect yield or surrogate weed biomass.
A survey of weed flora and basic weed management practices was conducted in commercial strawberry [Fragaria × ananassa (Weston) Duchesne ex Rozier ssp. ananassa] fields in central Florida during the 2019 to 2020 field season. Forty-one fields on 14 farms were surveyed, which represents a total of 907 ha or 23% of the acreage planted to strawberries in Florida. All conventional fields were fumigated, and combinations of chloropicrin and 1,3-dichloropropene were used on 72% of the fumigated acreage. Preemergence or postemergence herbicides were used in row middles on 100% of the conventional acreage, but preemergence herbicides applied under the plastic mulch were only used on 12% of the acreage. A total of 47 weed species were identified during the survey. The five weed species with the highest frequency were goosegrass [Eleusine indica (L.) Gaertn.], cutleaf evening primrose (Oenothera laciniata Hill), Carolina geranium (Geranium carolinianum L.), common ragweed (Ambrosia artemisiifolia L.), and eclipta [Eclipta prostrata (L.) L.] which occurred in 83%, 63%, 58%, 58%, and 46% of all fields surveyed, respectively. The five species with the highest overall relative abundance were E. indica, O. lanciniata, G. carolinium, A. artemisiifolia, and Brazil pusley (Richardia brasiliensis Gomes). The frequency and distribution of E. indica is of particular concern, because populations in strawberry fields are thought to be resistant to paraquat, which is the primary herbicide used for crop termination. Weed density tended to be low overall, which suggests the integrated weed management (IWM) programs adopted by most growers are effective. Mean weed density tended to be similar on conventional and organic fields and unaffected by fumigant applied the year of the survey or the number of years strawberries had been grown consecutively in a field. Weed management research moving forward should focus on the development of IWM programs for E. indica, O. lanciniata, G. carolinium, and A. artemisiifolia.
Almonds [Prunus dulcis (Mill.) D.A. Webb] are grown on nearly 650,000 ha in California and generate nearly $4.9 billion in export revenue annually, primarily to the European Union (EU). To facilitate harvest operations, broad-spectrum herbicides such as glyphosate and/or glufosinate are commonly used to control vegetation before harvest. The current minimum preharvest intervals (PHIs) for glyphosate and glufosinate herbicides registered in the United States are 3 and 14 d, respectively. The maximum residue limit (MRL) for glyphosate and glufosinate in almonds in the EU is 0.1 mg kg–1; however, a recent study recommended the glyphosate MRL be reduced to 0.05 mg kg–1. Laboratory and field experiments were conducted to evaluate herbicide transfer from soil to almonds and the effect of longer PHIs on glyphosate and glufosinate residues in harvested almonds. After harvest operations, almonds were dissected into hulls, shells, and kernels for analysis of glyphosate, glufosinate, and their metabolites using liquid chromatography–tandem mass spectroscopy. In the field experiment, glyphosate and glufosinate were detected at 0.121 to 0.291 mg kg–1 in almond hulls and shells. Glyphosate and primary metabolites were below the limit of detection (LOD) in almond kernels at all PHIs. Glufosinate was below the LOD, but the metabolite 3-(methylphosphinico)propionic acid was detected at 0.03 to 0.075 mg kg–1 in kernels from some replicate plots. There were no significant differences in either herbicide or any metabolite among PHI treatments. The lab experiment showed decreasing residue levels from hull to shell to kernel; furthermore, rinsing kernels resulted in 71% and 46% reduction in [14C]glyphosate and [14C]glufosinate, respectively, which suggests much of the herbicide residue may be associated with dust on the kernel surfaces. The results of these experiments indicate very low levels of herbicide transfer from soil to almonds, and increasing the PHI within the tested range did not reduce the already low amounts of herbicide or metabolites in almonds.
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