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Competition between genotypes within a plant population can result in the displacement of the least competitive by more competitive genotypes. Although evolutionary processes in plants may occur over thousands and millions of years, it has been suggested that changes in key fitness traits could occur in as little as decades, with herbicide resistance being a common example. However, the rapid evolution of complex traits has not been proven in weeds. We hypothesized that changes in weed growth and competitive ability can occur in just a few years because of selection in agroecosystems. Seed of multiple generations of a single natural population of the grassy weed giant foxtail (Setaria faberi Herrm.) were collected during 34 yr (i.e., 1983 to 2017). Using a “resurrection” approach, we characterized life-history traits of the different year-lines under noncompetitive and competitive conditions. Replacement-series experiments comparing the growth of the oldest year-line (1983) versus newer year-lines (1991, 1996, 1998, 2009, and 2017) showed that plant competitive ability decreased and then increased progressively in accordance with oscillating selection. The adaptations in competitive ability were reflected in dynamic changes in leaf area and biomass when plants were in competition. The onset of increased competitive ability coincided with the introduction of herbicide-resistant crops in the landscape in 1996. We also conducted a genome-wide association study and identified four loci that were associated with increased competitive ability over time, confirming that this trait changed in response to directional selection. Putative transcription factors and cell wall–associated enzymes were linked to those loci. This is the first study providing direct in situ evidence of rapid directional evolution of competitive ability in a plant species. The results suggest that agricultural systems can exert enough pressure to cause evolutionary adaptations of complex life-history traits, potentially increasing weediness and invasiveness.

The triazines are one of the most widely used herbicide classes ever developed and are critical for managing weed populations that have developed herbicide resistance. These herbicides are traditionally valued for their residual weed control in more than 50 crops. Scientific literature suggests that atrazine, and perhaps others-triazines, may no longer remain persistent in soils due to enhanced microbial degradation. Experiments examined the rate of degradation of atrazine and two other triazine herbicides, simazine and metribuzin, in both atrazine-adapted and non-history Corn Belt soils, with similar soils being used from each state as a comparison of potential triazine degradation. In three soils with no history of atrazine use, thet1/2of atrazine was at least four times greater than in three soils with a history of atrazine use. Simazine degradation in the same three sets of soils was 2.4 to 15 times more rapid in history soils than non-history soils. Metribuzin in history soils degraded at 0.6, 0.9, and 1.9 times the rate seen in the same three non-history soils. These results indicate enhanced degradation of the symmetrical triazine simazine, but not of the asymmetrical triazine metribuzin.

Herbicides have been the principal means of weed control in developed countries for approximately 50 yr because they are the most cost-effective method. Such general use of herbicides has resulted in weed resistance to herbicides, which continues to be a growing problem. Within the past decade, the evolution of resistance to the once-dominant herbicide glyphosate has resulted in major concerns about the future ability to control weeds in many crop systems. Moreover, many weed species have evolved resistance to multiple mechanisms of herbicide action. Given the dearth of new herbicides with novel mechanisms of action, it appears inevitable that weed management programs will need to be supplemented by the use of tactics other than herbicides. However, the inclusion of more diversity for weed management also introduces complexity, cost, and time constraints to current crop production systems. This paper describes broadly the considerations, opportunities, and constraints of diverse weed management tactics to address the burgeoning problems with herbicide resistance.

Field experiments were conducted near Boone, IA, to quantify postdispersal seed predation of common lambsquarters and common waterhemp in corn (2003) and soybean (2004) managed with conventional, reduced, and zero-tillage systems. Seed predation in each tillage regime was quantified using selective exclusion treatments during July through September 2003 and June through October 2004. In addition, the activity density of ground-dwelling invertebrates was estimated with pitfall traps. Choice and no-choice feeding trials were conducted in the laboratory using the most abundant weed seed predators found in the field to determine seed preferences of the potential predator organisms. The greatest seed loss occurred during July and August. In 2003, seed predation was lower in zero tillage than in conventional and reduced tillages, but no differences in seed predation between tillage regimes were observed in 2004. Maximum seed predation for common lambsquarters was 53% in 2003 and 64% in 2004. Common waterhemp seed predation reached 80% in 2003 and 85% in 2004. The majority of seed predation was by invertebrate organisms. The most common invertebrate species captured with pitfall traps were field crickets (Gryllus pennsylvanicusDe Geer [Gryllidae, Orthoptera]) and ground beetles (Harpalus pensylvanicusBurmeister [Coleoptera, Carabidae]). In 2003, field crickets were relatively more abundant in conventional and reduced tillage than in zero-tillage plots. In 2004, field crickets were more abundant in the reduced tillage than in the other two tillage regimes. No differences were detected for ground beetles among tillage regimes (P = 0.57). Choice and no-choice feeding experiments confirmed the preferences of field crickets and ground beetles for common lambsquarters and common waterhemp seeds over the larger seeds of giant foxtail and velvetleaf. Under field conditions, the activity density of field crickets was a significant predictor of common lambsquarters (r2= 0.47) and common waterhemp (r2= 0.53) seed predation. Positive relationships were also detected between the activity density of ground beetles and common lambsquarters (r2= 0.30) and common waterhemp (r2= 0.30) seed predation. This research demonstrated that weed seed predation is an important component affecting weed seedbanks and that crop management practices that favor the activity of predators such as field crickets or ground beetles could influence weed populations. Also, the results suggested that tillage is more important in determining the number of weed seeds available on the soil surface to predators than directly affecting predator activity density.

Waterhemp is an increasingly problematic weed in the U.S. Midwest, having now evolved resistances to herbicides from six different site-of-action groups. Glyphosate-resistant waterhemp in the Midwest is especially concerning given the economic importance of glyphosate in corn and soybean production. Amplification of the target-site gene, 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS) was found to be the mechanism of glyphosate resistance in Palmer amaranth, a species closely related to waterhemp. Here, the relationship between glyphosate resistance and EPSPS gene amplification in waterhemp was investigated. Glyphosate dose response studies were performed at field sites with glyphosate-resistant waterhemp in Illinois, Kansas, Kentucky, Missouri, and Nebraska, and relative EPSPS copy number of survivors was determined via quantitative real-time polymerase chain reaction (qPCR). Waterhemp control increased with increasing glyphosate rate at all locations, but no population was completely controlled even at the highest rate (3,360 g ae ha−1). EPSPS gene amplification was present in plants from four of five locations (Illinois, Kansas, Missouri, and Nebraska) and the proportion of plants with elevated copy number was generally higher in survivors from glyphosate-treated plots than in plants from the untreated control plots. Copy number magnitude varied by site, but an overall trend of increasing copy number with increasing rate was observed in populations with gene amplification, suggesting that waterhemp plants with more EPSPS copies are more resistant. Survivors from the Kentucky population did not have elevated EPSPS copy number. Instead, resistance in this population was attributed to the EPSPS Pro106Ser mutation. Results herein show a quantitative relationship between glyphosate resistance and EPSPS gene amplification in some waterhemp populations, while highlighting that other mechanisms also confer glyphosate resistance in waterhemp.

A segment of the debate surrounding the commercialization and use of glyphosate-resistant (GR) crops focuses on the theory that the implementation of these traits is an extension of the intensification of agriculture that will further erode the biodiversity of agricultural landscapes. A large field-scale study was initiated in 2006 in the United States on 156 different field sites with a minimum 3-yr history of GR-corn, -cotton or -soybean in the cropping system. The impact of cropping system, crop rotation, frequency of using the GR crop trait, and several categorical variables on seedbank weed population density and diversity was analyzed. The parameters of total weed population density of all species in the seedbank, species richness, Shannon's H′ and evenness were not affected by any management treatment. The similarity between the seedbank and aboveground weed community was more strongly related to location than management; previous year's crops and cropping systems were also important while GR trait rotation was not. The composition of the weed flora was more strongly related to location (geography) than any other parameter. The diversity of weed flora in agricultural sites with a history of GR crop production can be influenced by several factors relating to the specific method in which the GR trait is integrated (cropping system, crop rotation, GR trait rotation), the specific weed species, and the geographical location. Continuous GR crop, compared to fields with other cropping systems, only had greater species diversity (species richness) of some life forms, i.e., biennials, winter annuals, and prostrate weeds. Overall diversity was related to geography and not cropping system. These results justify further research to clarify the complexities of crops grown with herbicide-resistance traits to provide a more complete characterization of their culture and local adaptation to the weed seedbank.

Understanding abundance and distribution of weed species within the landscape of an agroecosystem is an important goal for weed science. Abundance is a measure of the number or frequency of individuals in an area. Distribution is a measure of the geographical range of a weed species. The study of weed population's abundance and distribution is helpful in determining how a population changes over time in response to selective pressures applied by our agronomic practices. Accurate estimates, however, of these two key variables are very important if we are to manage agricultural land both for productivity and for biodiversity.

Although the prolonged emergence pattern of common waterhemp is well known, its effect on flowering phenology and success is less well understood. The ecological significance of later common waterhemp cohorts could have been underestimated. Ecological knowledge of common waterhemp, such as reproductive phenology and success, would help us better understand the invasiveness of this weed and thus facilitate the development of more targeted control methods. Field studies were conducted at Ames, IA, to evaluate temporal variation in flowering phenology (e.g., date of flower initiation, number of flowering plants per day) and reproductive success (e.g., seed production) of natural common waterhemp cohorts. Later-emerging common waterhemp cohorts flowered quicker and had a relatively shorter flowering period than early cohorts. Common waterhemp cohorts showed very large variations for the duration of their vegetative phases, indicating different photoperiod sensitivity among cohorts. Furthermore, common waterhemp cohorts exhibited a pulsed flowering pattern, which was probably influenced by temporal distribution of rain events; up to seven distinct flowering pulses within 40 d were observed in 2009, and about eight flowering pulses scattered over a 60 d period were observed in 2010. Common waterhemp maintained high seed production throughout the growing season. Seed production for the entire cohort was influenced more by emergence timing, whereas individual plant fecundity was more affected by plant population densities. Common waterhemp demonstrates strong phenological plasticity by evolving a pulsed flowering pattern, which rendered it plastic enough to tailor flowering to variable environmental conditions thus facilitating effective pollination. Common waterhemp later cohorts have a strong seed production potential, which may be ecologically significant with regard to herbicide resistance evolution.

A segment of the debate surrounding the commercialization of genetically engineered (GE) crops, such as glyphosate-resistant (GR) crops, focuses on the theory that implementation of these traits is an extension of the intensification of agriculture that will further erode the biodiversity of agricultural landscapes. A large field-scale study was conducted in 2006 in the United States on 156 different field sites with a minimum 3-yr history of GR corn, cotton, or soybean in the cropping system. The impact of cropping system, crop rotation, frequency of using the GR crop trait, and several categorical variables on emerged weed density and diversity was analyzed. Species richness, evenness, Shannon's H′, proportion of forbs, erect growth habit, and C3species diversity were all greater in agricultural sites that lacked crop rotation or were in a continuous GR crop system. Rotating between two GR crops (e.g., corn and soybean) or rotating to a non-GR crop resulted in less weed diversity than a continuous GR crop. The composition of the weed flora was more strongly related to location (geography) than any other parameter. The diversity of weed flora in agricultural sites with a history of GR crop production can be influenced by several factors relating to the specific method in which the GR trait is integrated (cropping system, crop rotation, GR trait rotation), the specific weed species, and the geographical location. The finding that fields with continuous GR crops demonstrated greater weed diversity is contrary to arguments opposing the use of GE crops. These results justify further research to clarify the complexities of crops grown with herbicide-resistance traits, or more broadly, GE crops, to provide a more complete characterization of their culture and local adaptation.

Controlled environment experiments showed that velvetleaf plants grown under drought stress or low temperature (LT) treatments had greater leaf epicuticular wax (ECW) deposition compared to plants grown in soil with moisture at field capacity (FC) or a high temperature (HT) regime. Light intensity did not affect ECW deposition; however, increasing light intensity decreased the leaf ECW ester content and increased the secondary alcohol content. Plants grown at an LT regime or under FC had leaf ECW with fewer hydrocarbons and more esters than those grown at an HT or drought stress regime. Velvetleaf absorption of acifluorfen increased as light intensity decreased for plants grown in adequate soil water content, while the opposite was true for drought-stressed plants. Velvetleaf absorption of acifluorfen was approximately 3 and 10 times greater, respectively, with the addition of 28% urea ammonium nitrate (UAN) in comparison to crop oil concentrate (COC) or no adjuvant, regardless of the environmental treatments. Plants absorbed more acifluorfen when subjected to the LT regime in comparison to the HT regime when UAN was the adjuvant, while the opposite was true when COC was the adjuvant. Velvetleaf absorption of acifluorfen was not affected by drought stress when COC or no adjuvant was used and varied between studies when UAN was used. Velvetleaf absorption of bentazon was greatest for plants grown under HT/FC or high light/FC treatments and least with plants grown under HT/drought stress or low light/drought stress treatments, regardless of the adjuvant. However, bentazon absorption was higher with the addition of an adjuvant and for plants grown at a high light intensity or FC condition compared with medium to low light intensity or drought stress treatments.

Asiatic dayflower has recently become a troublesome weed in eastern Iowa. This weed demonstrates an extended emergence period and there is anecdotal evidence of glyphosate tolerance. Thus, Asiatic dayflower is difficult to manage in glyphosate-resistant (GR) corn and soybean. Greenhouse experiments were conducted to evaluate the response of Asiatic dayflower to glyphosate applied at different rates and growth stages. Field research was conducted in 2005 and 2006 to evaluate different herbicides for Asiatic dayflower control in soybean. PRE herbicides were applied at planting and POST herbicides were applied 21 and 42 d after planting (DAP). In addition, shikimate accumulation in response to glyphosate was compared among Asiatic dayflower and GR and non-GR corn and soybean. Under greenhouse conditions, a single application of glyphosate (0.84 kg ae ha−1) did not control Asiatic dayflower. Only the highest rate evaluated, 13.44 kg ae ha−1 (16X), was lethal to Asiatic dayflower. Even when applied at an early growth stage (two leaves) and using high rates (3.36 kg ae ha−1), glyphosate controlled Asiatic dayflower just 28%. In the field, metribuzin and KIH-485 controlled Asiatic dayflower 80 and 73%, respectively. Early POST applications (21 DAP) of cloransulam or lactofen controlled Asiatic dayflower 80 and 67%, respectively. A single glyphosate application of 0.86 kg ae ha−1 controlled Asiatic dayflower approximately 50%. Glyphosate-treated Asiatic dayflower and non-GR corn and soybeans accumulated shikimate after application. GR corn and soybeans did not accumulate shikimate in response to glyphosate. Twenty-one days after treatment, all the non-GR soybean and corn plants died; however, Asiatic dayflower plants survived.

Understanding weed seedling emergence patterns is important for successful implementation of many weed management strategies. Identifying the sources of variation of emergence patterns could greatly improve our ability to predict emergence timing. Differences in seed dormancy levels between populations or biotypes are usually not considered when studying seedling emergence of many weed species despite evidence that dormancy levels can affect weed seedling emergence patterns. We studied the importance of seed dormancy on seedling emergence patterns of common waterhemp using three biotypes (Ames, Everly, and Ohio) that differed in dormancy regulation and level (5, 26, and 87% germination, respectively) and three tillage systems (no-tillage, chisel plow, and moldboard plow) in 2004 and 2005. Seedling emergence was at least four times greater under no-tillage than under chisel or moldboard plow conditions. Fewer seedlings emerged in moldboard plow than in other tillage systems. Furthermore, seedling ...

Nitrogen (N) management markedly affects weed competition with crops. The effect of N availability on plant competition varies with a plant's abilities to capture and use N. Accordingly, we expected the N effect on plant competition to change with the relative emergence time of competing individuals. This hypothesis was tested by growing corn and velvetleaf plants in target and neighbor roles and comparing their N responses. Sowing times were varied, so that target plant emergent dates were constant, whereas neighbor plants were sown to emerge 7 d before, with, or 7 d after targets. Seedlings were fertilized daily with 2.5 mmol (“low N”) or 10.0 mmol (“high N”) ammonium nitrate (NH4NO3). Corn had greater total weight, leaf area, and root-to-shoot ratio (RSR) than velvetleaf. Different dry weights may have reflected seed weights; corn seed weight was greater than velvetleaf. Regardless of role, corn and velvetleaf dry weight and leaf area were greater with high N than low N; in contrast, RSR was lower with high N than low N. The RSR response to N availability suggested plant resources were shifted from N foraging, toward competition for photosynthetically active radiation (PAR). In target plants of each species, dry weight and leaf area increased linearly with time between target and neighbor emergence. Conversely, dry weight and leaf area of neighbor plants decreased with time between target and neighbor emergence. The N response of neighbor plants increased with time of emergence between target and neighbor emergence. Larger plants likely required more N to sustain growth than smaller plants; plants that emerged earlier likely had greater daily N requirements than those that emerged later. These results support factoring in emergence when predicting N effects on plant competition. Optimal N provisioning for integrated weed management may differ with emergence times of competing plants.

Controlled-environment experiments were conducted to determine giant foxtail epicuticular wax (ECW) deposition and fluazifop-P absorption under different environmental conditions and with two adjuvants. Drought stress and low temperature increased leaf ECW content, whereas low light intensity decreased ECW content compared with medium light intensity. Drought stress conditions decreased the fatty acid and primary alcohol content of ECW and increased the hydrocarbon content compared with field capacity. Compositional changes would make the ECW more hydrophobic and reduce leaf wetting by herbicide spray. Increasing air temperature decreased the aldehyde content of ECW, whereas decreasing light intensity increased ECW fatty acid and aldehyde content while decreasing primary alcohols and esters. Compositional changes under low light intensity would make the ECW more hydrophilic and increase leaf wetting by a herbicide spray. Drought stress reduced fluazifop-P absorption regardless of the temperature but could not further reduce fluazifop-P absorption under low light intensity. Fluazifop-P absorption by plants under low light and drought stress conditions was similar to plants under low or medium light intensity and field capacity conditions. Similarly, the rate of fluazifop-P absorption was less under drought stress and low light conditions. Fluazifop-P absorption was greater when crop oil concentrate was added compared with 28% urea ammonium nitrate or no additive. Crop oil concentrate, added to the herbicide solution, overcame reduced fluazifop-P absorption under the low light conditions and in one of the two drought stress regimes but could not overcome reduced fluazifop-P absorption with the high temperature regime.

Common waterhemp is an obligate outcrosser that has high genetic variability. However, under selection pressure, this weed shows population differentiation for adaptive traits. Intraspecific variation for herbicide resistance has been studied, but no studies have been conducted to determine the existence of variation for other adaptive traits that could influence weed management. The objective of this study was to examine the existence of different seed dormancy regulatory mechanisms in common waterhemp. Seed dormancy regulation, in response to different temperature and moisture regimes, was studied through germination experiments and proteome analysis using two common waterhemp biotypes (Ames and Everly) collected from agricultural fields in Iowa, and one biotype (Ohio) collected from a pristine area in Ohio. Without stratification, germination percentage among the different biotypes was 9, 29 and 88% for Ames, Everly, and Ohio respectively. The germination rate of seeds from Ames was dramatically increased after incubation at either 4 or 25 C under wet conditions, whereas germination of seeds from Everly was only increased at 25 C under wet conditions. The Ohio biotype showed no change in germination response to any of the incubation treatments. Germination studies indicated that the rate of seed dormancy alleviation differed between biotypes. Seed protein profiles obtained from the three biotypes differed in protein abundance, number, and type. A putative small heat-shock protein (sHSP) of 17.6 kDa and isoelectric point (pI) 6.1 increased whereas a putative glyceraldehyde-3-phosphate dehydrogenase (G3PDH) of 30.9 kDa and pI 6.4 decreased in abundance in the Ames biotype as seed dormancy was reduced in response to incubation at 4 C and wet conditions. These two proteins did not change in the Everly and Ohio biotypes, suggesting that these proteins changed their abundance in response to seed dormancy alleviation. The results of this study suggest that differences in seed dormancy levels between the biotypes were due to different physiological regulatory mechanisms.

The limited understanding of soil nitrogen (N) effects on crop yield loss to weeds prevents the refinement of integrated weed management to consider soil fertility. A 3-yr field experiment compared ammonium nitrate applications before corn emergence (PRE N) or in late June (POST N) on the growth of giant foxtail, velvetleaf, common waterhemp, and grain corn. The N responses were studied in corn population densities of 5.4 and 7.9 plants m−2 to understand how crop density might influence N effects on yield loss. In late June of each year, leaf areas were 25% (corn), 64% (velvetleaf), and 41% (giant foxtail) lower with POST N than with PRE N. Common waterhemp leaf area in late June was unaffected by N timing. Corn population density did not affect weed shoot dry weights evaluated in late June. Corn and velvetleaf heights in late July were up to 19 and 48% lower, respectively, with POST N compared with PRE N. Common waterhemp and giant foxtail heights in late July were unaffected by N timing. Corn y...

Weed biotypes putatively resistant to acetolactate synthase (ALS)–inhibiting herbicides were reported by Iowa farmers from 1997 to 2001. Greenhouse studies confirmed cross-resistance to triazolopyrimidine sulfonanilide and sulfonylurea (SU) herbicides in giant ragweed from Scott County, IA (Werner Farm), which corresponded to resistance to susceptibility (R:S) GR50(50% growth reduction) ratios of 21 and 28 to cloransulam and primisulfuron + prosulfuron, respectively. At the enzyme level, this represented a 49- and 20-foldI50(50% enzyme inhibition) increase. Cross-resistance to imidazolinone (IMI) and SU herbicides was also observed in common sunflower from Cherokee, IA. Compared with a susceptible biotype, the resistant common sunflower biotype demonstrated GR50R:S ratios of 36 and 43 to imazethapyr and chlorimuron, respectively. Shattercane from Malvern, IA, was susceptible to nicosulfuron but was resistant to imazethapyr (GR50R:S ratio = 29). The woolly cupgrass biotypes from Union County, IA (Pettit Farm and Travis Farm), were reportedly resistant but were identified susceptible to both IMI and SU herbicides. Using an in vivo ALS assay, extractable endogenous 2,3-diketone concentrations ranged from 25 to 71 nmol g−1fresh weight for all species. Compared with susceptible biotypes, 2,3-diketone levels accumulated to at least twofold higher levels in treated resistant plants 120 h after herbicide application. Field history data suggested that resistance evolved independently in three environments where ALS-inhibiting herbicides represented an important component of the selection pressure.

The nitrogen (N) response of competing plants may be affected by photosynthetically active radiation (PAR) availability and maximum potential growth rate, which determine N requirements. The responses of two crop (corn and soybean) and six weed species (common lambsquarters, common waterhemp, giant foxtail, velvetleaf, wild mustard, and woolly cupgrass) in low and high (150 and 450 μmol m−2s−1) PAR levels to daily fertilization with either low or high (0.2 or 7.5 mM) NH4NO3levels were studied. Leaf area of all species responded positively to N by 8 d after emergence (DAE) when grown in high PAR; in low PAR, most species did not respond until 11 DAE. Dry weight and leaf area of all species at 18 DAE were greater with high than with low N. These responses to high N were also greater in high than in low PAR for all species. Dry weights with high N were up to 100% greater in low PAR and up to 700% greater in high PAR than dry weights with low N. These responses suggest that low PAR reduced the benefit of N to the plants. The regression of relative growth rate (RGR) with high N to RGR with low N had a slope that was less than unity (β = 0.79), indicating that species with a higher RGR with high N experienced greater decreases in RGR with low N. Similarly, the sensitivity (change in RGR) of plants grown with high and low N was positively related to RGR with high N. RGR differences among crop and weed species may be related to differences in N requirement that could be exploited for weed management. RGR and seed size were negatively correlated, which may explain previous observations that small-seeded weeds were more sensitive to environmental stress.

Common waterhemp, giant foxtail, and velvetleaf seed germination in response to temperature was studied with a two-way thermogradient plate. Seeds were maintained under dark and wet conditions at 4 C for 12 wk, and velvetleaf seeds were scarified before the experiments were conducted. The seeds were germinated at 25 different temperature treatments. Minimum and optimum temperatures for velvetleaf germination were approximately 8 and 24 C, respectively. Temperature alternation did not affect the germination of this species. The minimum germination temperature was 10 C for common waterhemp and 14 C for giant foxtail. The optimum germination of giant foxtail occurred at approximately 24 C, but common waterhemp optimum germination was variable depending on temperature alternation. Increased amplitude of the diurnal temperature alternation increased percent germination of these two species, and this was more evident at lower temperatures. In the case of common waterhemp, the temperature required to reach specific germination percentages was reduced by increasing the amplitude of the temperature alternation.

A common sunflower population from Howard, SD (HSD) was previously determined to be cross-resistant to imazethapyr and chlorimuron-ethyl, both acetolactate synthase–inhibiting (ALS) herbicides. Experiments were conducted to determine if target-site polymorphisms could act as a mechanism of ALS-inhibitor herbicide resistance in the HSD common sunflower. Approximately 1,600 nucleotides were amplified by polymerase chain reaction and sequenced from putativeALSgene(s) in common sunflower and Jerusalem artichoke. In sunflower, two different amplification products were detected that differed by a nine-basepair deletion. This suggested the presence of at least two ALS genes in common sunflower that could contribute to the herbicide resistance phenotype. In addition, an Ala205to Val205substitution was observed in several clones from resistant common sunflower (amino acid position is relative to the full-length mouse-ear cress ALS protein). Previously documented mutations at this position in other species indicated that it might play a vital role in conferring resistance to one or more ALS-inhibitor herbicides.

The effects of temperature and light on the dormancy of velvetleaf, common waterhemp, and giant foxtail seeds were studied under controlled growth chamber conditions. Seeds were either kept chilled at 4 C for 12 wk under wet conditions or nonchilled at 4 C in dry storage. Then, seeds were germinated under increasing and decreasing temperatures and under continuous red light (R) and far-red light (FR). In addition, chilled and nonchilled seeds were germinated in the dark after being exposed to alternating R and FR flashes. Velvetleaf germination was increased by exposure to high temperatures (36 C) immediately after exposure to low temperatures (4 C), but light had no effect. Chilling increased common waterhemp seed germination and sensitivity to light and temperature. R promoted common waterhemp seed germination, whereas FR inhibited germination and maintained dormancy. In addition, the effect of light was reversible. Therefore, common waterhemp dormancy was phytochrome regulated. However, high t...

Five common cocklebur biotypes from southern Minnesota, central Iowa, southern Iowa, and Ohio were grown at the Iowa State University Curtiss Research Farm in 1995 and 1996 to examine intraspecific variations. Maximum plant heights were measured, and anthesis dates and day of bur set were recorded. At the end of the growing season, all plants were excised at the soil surface and weighed. Common cocklebur biotypes did not differ significantly in height. Flowering date was associated strongly with photoperiod and varied little between years within a biotype. But flowering date and bur set date differed among biotypes. The highest dry matter yields occurred in later flowering biotypes.

Two experiments were conducted to examine woolly cupgrass seed reserve utilization and the effect of germination depth on seed reserve utilization, tiller number, and timing of tiller emergence. The first experiment included germination in light with water, in light with Hoagland's solution, and in darkness with water; the second included five planting depths of 1, 3, 5, 7, and 9 cm. Endosperm utilization and shoot and root growth over time were recorded. Endosperms lost weight rapidly until 6 d after germination (DAG), when weight loss slowed down. Endosperm weight loss ceased 8 or 10 DAG. Endosperm weight loss was fastest for germination in light with Hoagland's solution. Deep planting significantly decreased emergence rate and increased the time to emergence. Differences in residual endosperms may have affected the early development of woolly cupgrass seedlings. Germination from a depth of 9 cm depleted the endosperm by the time of emergence. Deep planting also decreased shoot dry weight and t...

In 1996 a common sunflower population near Howard, SD, was suspected to be cross-resistant to imazethapyr and chlorimuron. Whole-plant acetolactate synthase (ALS) assays confirmed ALS-inhibitor resistance in the Howard biotype. The I50 values (inhibition of 50% of the enzyme activity) indicated that the resistant population required 39 and 9 times more imazethapyr and chlorimuron, respectively, to obtain the same level of enzyme inhibition compared with the sensitive biotype. Herbicide dose response data supported the whole-plant enzyme assay data; control (> 90%) was not achieved with less than a four-times application rate of chlorimuron. Control with imazethapyr was not achieved even with a 16-times rate. Chlorimuron and imazethapyr controlled 70 and 95% of the population, respectively, when a four-times rate of each herbicide was applied separately. Differences in 14C-herbicide absorption were observed, suggesting that there may be physical or chemical differences in leaf surface composition ...

The influence of tillage, crop rotation, and weed management regimes on the weed seedbank in land previously under the conservation reserve program (CRP) for 8 yr was determined from 1994 through 1997. The study was a split-plot design with four replications, two tillage systems, two crop rotations, and three weed management treatments. Eleven weed species were recorded in 1994 and 1995, and 13 in 1996 and 1997. The weed seedbank was dominated by broadleaf species. In 1994, the first year after CRP, the seed population density in the top 15 cm of the soil profile was 51,480 seeds m−2, of which 60 and 20% were pigweed and common lambsquarters. The population density of pigweed seeds in the seedbank increased over time and reached 51,670 seeds m−2 in 1996. In contrast, the seed population density for foxtail species was only 417 seeds m−2 in 1994, but it increased to 7,820 seeds m−2 in 1997. The large increase in foxtail species seed population density in the 4-yr period was mainly in the no-herbic...

Failure to control Xanthium strumarium with acetolactate synthase (ALS) inhibitor herbicides has been reported in Iowa and surrounding states. Single-seed descent techniques were used to isolate three X. strumarium biotypes: CAM-10 from near Cambridge, Iowa; Colo-25 from near Colo, Iowa; and Ohio-1 from Fulton County, Ohio. Ohio-1 and Colo-25 were selected because of apparent resistance to imazethapyr, whereas CAM-10 was selected for observed sensitivity to imazethapyr. The biotypes were assayed in vitro with three different ALS inhibitor herbicides, and ALS activity was measured. The 50% inhibition values (I50) of ALS for imazethapyr were determined to be ninefold or higher for the Ohio-1 and Colo-25 biotypes compared to the CAM-10 biotype. The I50 for imazaquin was determined to be about ninefold higher for the Colo-25 biotype and sixfold higher for the Ohio-1 biotype when compared to the CAM-10 biotype. All biotypes were equally sensitive to chlorimuron ethyl. The resistance was due to a singl...

A field study was established in southern Iowa in 1994 to study seasonal and long-term weed population dynamics on land being brought back into production after 8 yr as part of the conservation reserve program (CRP). The study was a split-plot design with four replications; two tillage regimes, two crop rotations, and three herbicide application methods were used. Even though the tillage regime did not influence individual weed population density throughout the study, the no-till (NT) regime had more weeds compared to conventional tillage (CT). However, when weeds were grouped into categories, tillage influenced broadleaf weeds in 1994 and 1996 and total weeds in 1995. Plots under the NT regime had an average of 46 broadleaf weeds m−2compared to 27 in CT in 1994, withAmaranthus rudisSauer (common waterhemp) being the most prevalent. NT had a total of 186 weeds m−2compared to 125 m−2weeds in CT in 1995; however, in 1996, CT plots had 184 weeds m−2compared to 121 m−2in the NT regime. Except for broadleaf weeds in 1994, crop rotation did not influence the number of weeds, and herbicide application methods had the greatest effect on weed populations. Overall, weed populations were greater in 1997, 1996, and 1995 than in 1994 for all herbicide application methods. The no-herbicide treatment had the highest number of weeds throughout the study. The total number of weeds in band and broadcast treatments averaged 41 and 26 m−2in 1994; 96 and 24 m−2in 1995; 96 and 12 m−2in 1996; and 109 and 95 m−2in 1997. The use of broadcast herbicides in NT should be recommended for land coming out of CRP. Regardless of the herbicide application method or crop rotation, CT plots had better yields for bothZea maysL. (corn) andGlycine maxL. (soybean).Glycine maxhad a better stand compared toZ. maysin the first year, indicating that a rotation starting withG. maxmight be preferred in the land coming out of CRP.

Micheal D. K. Owen Corresponding author. Agronomy Department, Iowa State University, Ames, IA 5001 1; mdowen@iastate.edu Laboratory and field experiments were conducted to determine the effects of temperature, oxygen, and seed burial depth on woolly cupgrass seed dormancy. Woolly cupgrass seeds at physiological maturity are innately dormant. Stratifying these seeds at 5 C for 8 wk increased germination to greater than 90%, regardless of the germination temperature. Alternating the germination temperature hastened germination by 2 wk, while seedcoat removal hastened germination by another 4 wk. Low oxygen concentration was more detrimental to seed germination when seeds were subjected to constant temperatures in comparison to alternating temperatures. Oxygen concentration did not affect seed germination when seeds were placed in an alternating 15/25 C regime. Seed germination was approximately 15% less when seeds were stratified for 2 wk and subjected to oxygen concentrations below the ambient oxygen concentration, whereas this decrease was present only at the 8% oxygen concentration when seeds were stratified 4 wk. In the field, the stratification requirement was satisfied by early December for the study year. Seeds remaining on the soil surface overcame dormancy sooner than seeds buried 2 or 4 cm.

The effect of herbicides applied to V3 soybeans on race 3 soybean cyst nematode (SCN) reproduction and glyceollin production in roots was measured. Soybeans were treated postemergence with 1 × and 2 × rates of herbicides plus adjuvants or with adjuvants alone, and SCN development was measured. Acifluorfen, bentazon, lactofen, crop oil concentrate (COC), and nonionic surfactant (NIS) applications reduced SCN egg population densities 50 to 60% compared with the untreated control 4 and 8 wk after application. The SCN reproduction on plants treated with fluazifop-P, sethoxydim, and imazethapyr was similar to the untreated control. Crop oil concentrate or NIS applications alone were as effective as acifluorfen, bentazon, or lactofen applications for reducing SCN reproduction. However, no additive effect of adjuvant-herbicide combinations was observed, nor did herbicide rate affect SCN reproduction. Treatments reduced SCN reproduction only when applied to soybeans and had no effect on SCN reproduction when applied directly to the soil. No treatment stimulated SCN reproduction relative to the untreated control. Soybeans treated with COC, NIS, acifluorfen, and bentazon also had more glyceollin detected than the untreated control. Herbicide-induced glyceollin production may have increased the resistance of soybean to SCN.

Microsomes (100,000 g pellet containing mixed membrane fractions but primarily endoplasmic reticulum) were isolated from shoots of corn, shattercane, and woolly cupgrass grown from naphthalic anhydride treated or untreated seed to determine if metabolism of bentazon, nicosulfuron, and primisulfuron could be demonstrated in the preparations. Corn is tolerant of all three herbicides, shattercane is tolerant of bentazon, and woolly cupgrass is tolerant of bentazon and primisulfuron. Naphthalic anhydride treatment was required for detectable bentazon, nicosulfuron, and primisulfuron hydroxylation in corn microsomes and for bentazon hydroxylation in woolly cupgrass microsomes. Bentazon hydroxylation was low, but detectable, in microsomes from shattercane shoots without naphthalic anhydride treatment. Naphthalic anhydride-treated corn microsomes hydroxylated 292, 120, and 52 pmol mg−1protein min−1of bentazon, nicosulfuron, and primisulfuron, respectively. Primisulfuron (19 pmol mg−1protein min−1), but not nicosulfuron, was hydroxylated in woolly cupgrass microsomes. Neither nicosulfuron nor primisulfuron was hydroxylated in shattercane microsomes. Bentazon and primisulfuron inhibited nicosulfuron hydroxylation in corn microsomes. Bentazon, but not nicosulfuron, also inhibited primisulfuron hydroxylation in the corn microsomes. This indicates that the three herbicides can interact at the same cytochrome P-450(s) in corn. Primisulfuron hydroxylation was not inhibited by either bentazon or nicosulfuron in woolly cupgrass microsomes. This suggests that the cytochrome P-450(s) for primisulfuron hydroxylation are different between corn and woolly cupgrass. Also, bentazon hydroxylation in corn and shattercane microsomes was inhibited by the cytochrome P-450 inhibitor tetcyclasis, while that in woolly cupgrass was not. Again, this suggests a difference in the cytochrome P-450(s) responsible for bentazon metabolism among the species. Although absolute conclusions comparing in vitro microsomal activities to whole plant herbicide tolerance cannot be made because it is unknown whether the same cytochrome P-450(s) are studied in microsomes from naphthalic anhydride-treated tissue as are responsible for in vivo herbicide metabolism, there was a broad correlation between metabolism of a particular herbicide in microsomes of a species and the species' tolerance of that herbicide.

Micheal D. K. Owen Department of Agronomy, Iowa State University, Ames, IA 50011 Horseweed emergence and survival were evaluated in no-tillage soybean and corn at Rosemount, MN, and Ames, IA, from the fall of 1992 through the summer of 1995. Most of the horseweed at both locations emerged in the fall. Winter survival of fall-emerged seedlings ranged from 59 to 91%. Timing and extent of horseweed emergence in spring varied by year and location. Spring emergence ranged from 5 to 32% of total emergence, with greater spring emergence at Rosemount than at Ames. Emergence occurred as late as early June at Rosemount and late May at Ames. Results indicated that horseweed may emerge well into the growing season, and spring emergence should be considered in weed management systems for no-tillage crop production.

Research was conducted to elucidate mechanism(s) of nicosulfuron and primisulfuron selectivity in corn, woolly cupgrass, and shattercane. Corn absorbed less than one half the nicosulfuron and primisulfuron that woolly cupgrass and shattercane absorbed. Acetolactate synthase (ALS) isolated from three species responded similarly to nicosulfuron and primisulfuron. Values for 50% (I50) inhibition of ALS for nicosulfuron were 36.9, 37.2 and 29.5 nM for corn, woolly cupgrass and shattercane, respectively. Primisulfuron I50values were 13.8, 11.5 and 10.0 nM for corn, woolly cupgrass and shattercane, respectively. Shattercane's I50was different from corn's but not from that of woolly cupgrass. Corn rapidly metabolized nicosulfuron and primisulfuron, with a half-life of less than 4 h. Shattercane metabolized the herbicides more slowly, with a half-life greater than 72 h for nicosulfuron and 36 h for primisulfuron. Nicosulfuron and primisulfuron half-lives were greater than 72 h and less than 4 h, respectively, in woolly cupgrass. Therefore, selectivity with nicosulfuron and primisulfuron is likely based on metabolism to nonphytotoxic compounds. Corn tolerance to nicosulfuron and primisulfuron was also attributed to reduced herbicide penetration and translocation below the treated leaf.

Laboratory studies were conducted to determine the effect of seed coat on woolly cupgrass seed moisture and oxygen uptake, and to determine if water-soluble growth inhibitors are present in the seed. Intact dormant seeds did not respond to any temperature regime or to oxygen concentrations above atmospheric level. Dehulling increased germination of dormant seed to about 85%. Oxygen concentrations above atmospheric level increased germination of partially dehulled (1/4 distal end of the seed dehulled) seeds an additional 10%. Oxygen uptake by dehulled dormant and intact nondormant seeds was greater than intact dormant seeds. Leaching intact dormant seeds did not promote germination, nor did the leachate inhibit germination of nondormant seeds. Embryos excised from dormant seeds germinated under laboratory conditions. Results from this study suggest that the woolly cupgrass seed coat may inhibit germination by controlling oxygen availability to the embryo.

More than 70% of all14C-bentazon absorption occurred within 4 h after herbicide application regardless of adjuvant Moisture stress reduced14C-bentazon absorption by common cocklebur and velvetleaf. Mature (second true leaf) and moisture-stressed leaves of velvetleaf had 50 and 17 μg cm−1more epicuticular wax (ECW) than did juvenile and unstressed leaves, respectively. Common cocklebur had less14C in the ECW and lower total14C in treated mature leaves compared to juvenile leaves. The use of 28% urea ammonium nitrate (UAN) or crop oil concentrate (COC) increased14C in ECW samples of both plant species, regardless of leaf age or moisture stress. More14C in the ECW did not always correlate with more14C in the leaf tissue. Adjuvants increased14C-bentazon absorption into leaves of plants that had been stressed.