Downy brome (Bromus tectorum L.) is a difficult species to control in the dryland wheat (Triticum aestivum L.) production areas of northeastern Oregon. The selection of herbicide-resistant B. tectorum populations has further complicated B. tectorum management. A survey of wheat growers was conducted in 2021 and 2022 to understand B. tectorum management practices. The survey included four questions based on the growing seasons from 2017 to 2022 related to crop rotation, tillage versus no-tillage, irrigation versus dryland, and herbicide programs. To determine the extent of herbicide resistance, seeds were collected from 49 B. tectorum populations in wheat fields in northeastern Oregon and tested for resistance. Herbicides tested were clethodim, glyphosate, imazamox, mesosulfuron, metribuzin, propoxycarbazone, pyroxasulfone, pyroxsulam, quizalofop, and sulfosulfuron. Winter wheat–summer fallow rotation was the most predominant cropping system in the region. Most of the fields were in no-tillage systems, and none were irrigated. Pyroxasulfone applied preemergence and acetolactate synthase (ALS) inhibitors applied postemergence were the most often used herbicides for B. tectorum control in winter wheat. Glyphosate was the most frequently used herbicide for B. tectorum control in summer fallow. Resistance screenings confirmed that 46 of the 49 B. tectorum populations were resistant to ALS inhibitors with different cross-resistance patterns. Two populations were resistant to metribuzin and exhibited multiple resistance to ALS inhibitors. All populations were susceptible to clethodim, glyphosate, pyroxasulfone, and quizalofop. The widespread occurrence of ALS inhibitor–resistant B. tectorum populations limits effective postemergence herbicide options in winter wheat.

Due to increased food demand, the use of herbicides is both necessary and on the rise. Several herbicide classes target photosynthetic electron transport: Herbicide Resistance Action Committee (HRAC) Groups 5, 6, and 22. These herbicides are used in large amounts in many different cropping systems to control several species of broadleaf and grass weeds. This article provides a comprehensive review of what these photosynthesis inhibitors are, how they are used, and their modes of action. Presently, commercial herbicides only inhibit electron flow at two different sites: photosystem II (PSII) and photosystem I (PSI). Herbicides that inhibit electron flow at PSII block the movement of electrons down the electron transport chain, while those that inhibit electron flow at PSI accept electrons. Necrosis developing on the leaves of plants treated with PSII and PSI inhibitors is due to the accumulation of reactive oxygen species. Evolution of resistance, toxicity concerns, and other limitations of these herbicides call for the exploration of new chemistries that can be used to target this pathway.

Protoporphyrinogen oxidase (PPO)-inhibiting herbicides remain an important and useful chemistry 60 yr after their first introduction. In this review, based on topics introduced at the Weed Science Society of America 2021 symposium titled “A History, Overview, and Plan of Action on PPO Inhibiting Herbicides,” we discuss the current state of PPO-inhibiting herbicides. Renewed interest in the PPO-inhibiting herbicides in recent years, due to increased use and increased cases of resistance, has led to refinements in knowledge regarding the mechanism of action of PPO inhibitors. Herein we discuss the importance of the two isoforms of PPO in plants, compile a current knowledge of target-site resistance mechanisms, examine non–target site resistance cases, and review crop selectivity mechanisms. Consistent and reproducible greenhouse screening and target-site mutation assays are necessary to effectively study and compare PPO-inhibitor resistance cases. To this end, we cover best practices in screening to accurately identify resistance ratios and properly interpret common screens for point mutations. The future of effective and sustainable PPO-inhibitor use relies on development of new chemistries that maintain activity on resistant biotypes and the promotion of responsible stewardship of PPO inhibitors both new and old. We present the biorational design of the new PPO inhibitor trifludimoxazin to highlight the future of PPO-inhibitor development and discuss the elements of sustainable weed control programs using PPO inhibitors, as well as how responsible stewardship can be incentivized. The sustained use of PPO inhibitors in future agriculture relies on the effective and timely communication from mode of action and resistance research to agronomists, Extension workers, and farmers.

Chlorophyll a fluorescence is often used to estimate various types of damage in lichens. In order to optimize the output and improve interpretations of such measurements the protocol for pretreatment and measuring is important. To study the effects of measurement conditions, the lichens Lobaria pulmonaria, L. scrobiculata, Xanthoria parietina and Parmelia sulcata were first stressed by high light intensities at 600 or 1000 µmol photons m−2 s−1 for 4 h. Then various conditions during recovery or pretreatment were used to optimize the detection of more lasting damage. Recovery from photoinhibition was incomplete in darkness, whereas light as low as 0·2 or 1·0 µmol m−2 s−1 resulted in complete recovery if the recovery period was long enough. Additionally, low intensity light given for1·5 h after one day in darkness caused rapid and complete recovery. In conclusion, before measuring maximal PSII efficiency (Fv/Fm) with chlorophyll fluorescence, it is important to let lichens recover in low intensity light and not in darkness, to optimize recovery from photoinhibition; dark adaptation can only be recommended if the photoinhibition status of the lichens is of interest.

A comparison of the structure of the α-methylbenzyl analogue (MBAT) of atrazine with the Photosystem II herbicides atrazine [6-chloro-N-ethyl-N′-(1-methylethyl)-1,3,5-triazine-2,4-diamine] and diuron [N′-(3,4-dichlorophenyl)-N,N-dimethylurea] suggested that N-alkylbenzyltriazines may be a structural bridge between the triazines and the phenylureas. In the phenylureas, the addition of chlorines at the meta and/or para positions produces a marked increase in activity. Chloro-substituted derivatives of MBAT were synthesized to determine whether this structure-activity relationship also applies to the alkylaryltriazines. Addition of a chlorine to MBAT at the 4-position (CMBAT) caused a substantial increase in intrinsic activity, and a second chlorine at the 3-position (DCMBAT) caused a further increase. In direct comparisons, DCMBAT was more active in vitro than terbuthylazine [6-chloro-N-ethyl-N′-(1,1-dimethylethyl)-1,3,5-triazine-2,4-diamine], the most active chlorotriazine, and was also more active than diruon. The effects of DCMBAT were also measured on triazine-resistant pigweed (Amaranthus hybridus L. # AMACH) both in vivo and in vitro. The activity of this compound in triazine-resistant chloroplasts was intermediate between that of atrazine and diruon both in inhibition of photosynthetic electron transport and in competition for diuron binding sites, with half-maximal values falling in the micromolar range. Whole plant phytotoxicity of DCMBAT on triazine-resistant pigweed was also intermediate between that of diuron and atrazine. Since DCMBAT is a triazine with biological properties similar to that of a urea, we conclude that in a functional as well as structural sense, DCMBAT is a herbicide that is a hybrid between a triazine and a urea.

Repeated use of propanil to control barnyardgrass (BYG) and other weeds in rice has led to the development of propanil-resistant barnyardgrass (R-BYG). R-BYG possesses elevated aryl acylamidase activity levels, which cause rapid metabolism of propanil analogous to propanil degradation in rice. The current screening method for determining propanil resistance in BYG requires about 10 mo. The present study examined the use of chlorophyll fluorescence as a more rapid method to identify propanil resistance in BYG soon after it is suspected. Chlorophyll fluorescence data from excised BYG leaf tissue (R-BYG and susceptible-BYG [S-BYG]; 13- to 41-d-old) exposed to 100 μM propanil for 2 h indicated a 95 to 100% inhibition of electron transport (photosynthesis inhibition) in both R- and S-BYG. However, when incubated in water in the dark for 22 h after the initial 2-h treatment, metabolism in R-BYG was sufficient to reduce levels of absorbed propanil and facilitate recovery. Lack of metabolism of propanil prevented recovery in S-BYG, thus allowing the two biotypes to be distinguished easily by the chlorophyll fluorescence assay. Further studies using this 2-h exposure to 100 μM propanil followed by a 22-h recovery period evaluated several assay parameters. A longer recovery time and the effects of various propanil concentrations were also evaluated. A herbicide dose-response curve showed the greatest difference in photosynthesis inhibition for both biotypes at about 100 μM propanil, but both biotypes were inhibited > 95% when treated with 400 μM propanil. Inhibition of photosynthesis in both biotypes was greatest when the recovery incubation temperature was 35 C compared to 20, 25, and 30 C. Fluorescence data from harvested tissue stored moist in plastic bags at 23 C (to simulate shipment) showed that biotypes could be differentiated even when received as late as 4 d after harvest. Thus, samples can be harvested from the field soon after propanil failure and resistance or susceptibility to propanil determined after only a few days. This technique can greatly reduce the time, space, and labor currently required to determine propanil resistance in BYG.

Resistance to propanil by a barnyardgrass (BYG) biotype has been reported, and its occurrence is becoming widespread in U.S. rice-producing regions. Interactions between propanil and the herbicides anilofos, molinate, pendimethalin, piperophos, quinclorac, and thiobencarb and the insecticide carbaryl were evaluated in laboratory studies for improved control of propanil-resistant barnyardgrass (R-BYG). Chlorophyll fluorescence measurements of BYG leaf segments were used to ascertain electron transport inhibition and to assess the synergy/antagonism of propanil (100 μM) with the various additives (50 μM). Synergistic effects on photosynthetic inhibition were found with anilofos, carbaryl, pendimethalin, and piperophos in combination with propanil. Such synergistic actions of chemicals with propanil may lead to chemical combinations useful for R-BYG control.

The effects of buthidazole {3-[5-(1,1-dimethylethyl)-1,3,4-thiadiazol-2-yl]-4-hydroxy-1-methyl-2-imidazolidinone} on the photochemical reactions of isolated pea (Pisum sativum L.) chloroplast thylakoids were analyzed. Buthidazole was found to inhibit electron transport at two distinct sites in the photosynthetic electron transport chain. The major site of inhibition was on the reducing side of photosystem II at the site of diuron [3-(3,4-dichlorophenyl)-1,1-dimethylurea] and atrazine [2-chloro-4-(ethylamino)-6-(isopropylamino)-s-triazine] inhibition. Buthidazole also had a secondary site of electron transport inhibition on the oxidizing side of photosystem II. No evidence was found for buthidazole to act as an uncoupler or as an energy transfer inhibitor.

Propanil is an acylanilide herbicide introduced in the early 1960s to control dicotyledonous weeds and grasses, including Echinochloa species in cultivated rice. Since then, propanil has been used extensively in rice production in the United States and in several other countries. Propanil is an inhibitor of photosystem II, but rice is tolerant to propanil because of the presence of a high level of aryl acylamidase that catalytically degrades the compound to nonphytotoxic products, i.e., 3,4-dichloroaniline and propionic acid. About 10 yr ago, biotypes of barnyardgrass and junglerice were discovered to be resistant to propanil. The resistance mechanism of these two biotypes has been shown to be elevated levels of aryl acylamidase activity. Various strategies to combat propanil resistance and to more fully understand the biochemistry involved in this resistance have been investigated. These include studies on the interactions of herbicides and other chemicals with propanil, rotation of rice with other crops (consequently the use of other herbicide modes of action), and use of alternative herbicides in rice. Certain compounds, including some organophosphate insecticides, are potent inhibitors of aryl acylamidase, which can act as synergists with propanil to increase phytotoxicity. Another compound that lacks insecticidal or herbicidal activity, PPG-124, has been commercialized as a herbicide synergist for propanil. These chemical and biochemical interactions and other factors involved in propanil-resistant Echinochloa weeds are presented and discussed.

Herbicides inhibit biochemical and physiological processes or both with lethal consequences. The target sites of these small molecules are usually enzymes involved in primary metabolic pathways or proteins carrying out essential physiological functions. Herbicides tend to be highly specific for their respective target sites and have served as tools to study these physiological and biochemical processes in plants (Dayan et al. 2010b).

Weed management in potato and mint rely heavily on two Group 5 photosystem II–inhibiting herbicides, metribuzin and terbacil, respectively. Seed from weed escapes was collected in 2010 from 69 potato and mint production fields in Washington State and tested for resistance to metribuzin and terbacil. Of the seed screened, 26 of 51 Powell amaranth and redroot pigweed accessions and 8 of 23 common lambsquarters accessions tested expressed at least some degree of resistance to one or both herbicides. A Ser264Gly mutation in the chloroplast psbA gene was present in all but three resistant accessions. Herbicides with other mechanisms of action registered in potato and mint controlled all identified photosystem II–resistant accessions when applied at typical labeled rates.

Amicarbazone is a new triazolinone herbicide with a broad spectrum of weed control. The phenotypic responses of sensitive plants exposed to amicarbazone include chlorosis, stunted growth, tissue necrosis, and death. Its efficacy as both a foliar- and root-applied herbicide suggests that absorption and translocation of this compound is very rapid. This new herbicide is a potent inhibitor of photosynthetic electron transport, inducing chlorophyll fluorescence and interrupting oxygen evolution ostensibly via binding to the QB domain of photosystem II (PSII) in a manner similar to the triazines and the triazinones classes of herbicides. As a result, its efficacy is susceptible to the most common form of resistance to PSII inhibitors. Nonetheless, amicarbazone has a good selectivity profile and is a more potent herbicide than atrazine, which enables its use at lower rates than those of traditional photosynthetic inhibitors.

A population of waterhemp was identified in Adams County, Illinois, that survived treatment of several acetolactate synthase (ALS) inhibitors and a postemergence (POST) application of lactofen, a protoporphyrinogen oxidase (PPO)–inhibiting herbicide. Greenhouse studies were conducted to quantify the responses of this waterhemp population, designated ACR, to multiple PPO inhibitors and various other herbicides with different sites of action. Resistance ratios were obtained by comparing herbicide dose–response curves between the ACR population and a herbicide-susceptible waterhemp population. The ACR population was resistant to lactofen (23-fold) and to five other PPO-inhibiting herbicides (ranging from 2.2- to 6.2-fold). Furthermore, the ACR waterhemp population was 17,000-fold and 18,000-fold resistant to imazamox and thifensulfuron, respectively, two ALS-inhibiting herbicides. In response to atrazine, a Photosystem II inhibitor, the ACR population was 38-fold resistant. Plants within the ACR waterhemp population survived treatment of a herbicide mixture containing lactofen at 175 g ai ha−1, imazamox at 44 g ae ha−1, and atrazine at 1,000 g ai ha−1. Thus, individual plants—not just the population as a whole—displayed multiple herbicide resistance. The ACR population was not resistant to glyphosate or paraquat. This is the first reported weed population from the United States with resistances to herbicides inhibiting three unique sites of action. Furthermore, this research identifies a significant reduction in the number of POST herbicide options available for waterhemp control in soybean production.

Turfgrass injury from triazines has limited the use of photosystem II (PS II) inhibitors for weed control in seashore paspalum. Amicarbazone is a new PS II inhibitor with potential safety in seashore paspalum, but the effects of application timing on turf tolerance has received limited investigation. Field experiments were conducted in Griffin, GA to evaluate the tolerance of ‘Sea Isle 1’ seashore paspalum to amicarbazone applications in winter, spring, and summer. Seashore paspalum had minimal injury (< 5%) from amicarbazone treatments (98, 196, and 392 g ai ha−1) applied for annual bluegrass control in winter and spring. By 6 wk after treatment (WAT), amicarbazone at 392 g ha−1 provided 78 and 90% annual bluegrass control in 2013 and 2014, respectively, and was similar to pronamide at 1,680 g ai ha−1. Amicarbazone at 196 g ha−1 provided 71% control of annual bluegrass in 2014, but control was poor (< 70%) in 2013. Sequential amicarbazone applications at 98 g ha−1 provided poor control in both years by 6 WAT. From six amicarbazone rates (up to 984 g ha−1) applied in summer, seashore paspalum required 510 and < 123 g ha−1 for 20% turfgrass injury (I20) and 20% clipping reduction (CR20), respectively, whereas I20 and CR20 measured > 984 g ha−1 for ‘Tifway’ bermudagrass. Overall, amicarbazone may be safely applied to seashore paspalum in winter, spring, and summer at rates and regimens evaluated. However, seashore paspalum may exhibit shoot growth inhibition up to 4 WAT, suggesting that end users should be cautious when using amicarbazone during active growth in summer.

Amicarbazone is a photosystem II (PSII) inhibiting herbicide of the triazolinone herbicide family. Greenhouse experiments were conducted to compare the effects of amicarbazone and atrazine on annual bluegrass control and quantum yield (ΦPSII) when applied at three treatment placements (soil-only, foliage-only, and foliage + soil). Herbicide rates for amicarbazone and atrazine were 0.53 and 2.25 kg ha−1, respectively. Amicarbazone applied soil-only and foliage + soil controlled annual bluegrass 57 and 59%, respectively, 1 wk after treatment (WAT). Atrazine applied to foliage + soil controlled annual bluegrass 48% at 1 WAT. All soil-only and foliage + soil treatments were similar 2 WAT. Foliage-only application of amicarbazone provided significantly less control than other amicarbazone treatments throughout the study. Amicarbazone applied soil-only and foliage + soil controlled annual bluegrass 100% at 3 WAT. Soil-only and foliage + soil applications of atrazine and amicarbazone had similar reductions in quantum yield (ΦPSII) at 1 to 3 WAT. Foliar-applied amicarbazone reduced ΦPSII 78, 84, and 86% at 1, 2, and 3 WAT, respectively. The rapid reduction in annual bluegrass ΦPSII and the increase in control resulting from soil contact of amicarbazone indicate root exposure of amicarbazone is beneficial for annual bluegrass control.

Laboratory and greenhouse studies were conducted to determine if the reported red morningglory control failures with atrazine in sugarcane are caused by triazine-resistant mutants. Plants were grown from seeds collected at 24 locations in Louisiana and Arkansas where atrazine had never been used, and where it had been used in sugarcane with poor results. Terminal fluorescence of leaf material from all locations increased after treatment with 10−3 M atrazine, indicating electron transport inhibition and, hence, triazine susceptibility. However, small differences in the magnitude of fluorescence increase were observed among populations, possibly indicating the existence of biotypes with slightly different inherent tolerances to atrazine. Some phenotypic differences were observed among the red morningglory populations. In a separate study, postemergence application of atrazine at 1.1 kg ai/ha plus nonionic surfactant controlled greenhouse-grown plants from all populations at least 99%, which supports the findings of the fluorescence assay. This research was unable to verify that reduced red morningglory control with atrazine was the result of an altered binding site mutation, even in populations exposed to atrazine annually for more than 10 yr. Other factors should be evaluated to determine their impact on atrazine performance.

Greenhouse studies were conducted in 2003 at the Stine–Haskell Research Center to determine whether herbicide inhibitors of six specific sites in the carotenoid biosynthesis pathway would elicit synergistic responses when applied postemergence (POST) in combination with the photosystem II (PSII) inhibitor atrazine. Based on data analysis with the Isobole method, synergistic responses were observed on red morningglory, common cocklebur, and giant foxtail when atrazine was applied in mixtures with the deoxy-D-xylulose-5-phosphate reductoisomerase (DOXP reductoisomerase) inhibitor fosmidomycin, the p-hydroxyphenylpyruvate dioxygenase (HPPD) inhibitor mesotrione, and the DuPont proprietary zeta-carotene desaturase (ZDS) inhibitor DFPC. Clomazone (its metabolite ketoclomazone is the actual enzyme inhibitor), an inhibitor of 1-deoxy-D-xylulose-5-phosphate synthatase (DOXP synthase), provided synergistic responses on red morningglory, but antagonistic responses on both common cocklebur and giant foxtail when applied in mixtures with atrazine. Combinations of the lycopene cyclase (LC) inhibitor, CPTA, with atrazine produced synergistic responses on both common cocklebur and giant foxtail but were antagonistic on red morningglory. Norflurazon, a phytoene desaturase (PDS) inhibitor, applied in mixtures with atrazine provided synergistic responses on red morningglory, antagonistic responses on giant foxtail, and independent responses on common cocklebur. Because carotenoids have been determined to play a key role in quenching singlet oxygen species in the chloroplast and also assist in the maintenance of the D1 protein in PSII, this might help explain the synergistic responses with atrazine observed in our studies.

Selective annual bluegrass (ABG) control with mesotrione is often inconsistent, and sequential applications might be required for complete control. The complementary nature of p-hydroxyphenylpyruvate dioxygenase (HPPD)- and photosystem II (PSII)-inhibiting herbicides is well documented. The HPPD-inhibiting herbicide mesotrione and the PSII-inhibiting herbicide amicarbazone both have efficacy against annual bluegrass and safety on certain cool-season turfgrasses. Topramezone is a HPPD-inhibiting herbicide being investigated for use in turfgrass. Field and greenhouse experiments were conducted to examine single applications of topramezone and mesotrione alone or in combination with amicarbazone for POST ABG control in spring. In greenhouse experiments, the combination of mesotrione (280 g ai ha−1) and amicarbazone (75 g ai ha−1) controlled ABG 70% by 21 d after treatment, > 29% more than either herbicide applied alone; these combinations were determined to be synergistic. Amicarbazone combined with topramezone (14.5 g ai ha−1) provided < 10% ABG control and was not synergistic. When combined with mesotrione, increasing amicarbazone rate to 150 or 255 g ha−1 did not increase ABG control compared to 75 g ha−1in field experiments. Combining mesotrione with amicarbazone resulted in a synergistic increase in POST ABG control at 1 and 2 wk after treatment (WAT). When applied alone or in combination with amicarbazone, increasing the mesotrione rate from 90 to 280 g ha−1 increased efficacy on ABG in field experiments. The combination of mesotrione at 280 g ha−1 and amicarbazone at 75 g ha−1 provided > 90% ABG control in field experiments. Future research should focus on sequential applications of mesotrione–amicarbazone combinations for ABG control in locations where ABG is historically more difficult to control.

Lobaria pulmonaria is an epiphytic lichen that, in south-eastern Canada, inhabits deciduous forests where it must acclimate to large seasonal changes in temperature and in light caused by closing and opening of the leaf canopy. On a seasonal timescale, this acclimation occurs via large shifts in the macromolecular complexes of the photosynthetic system, within a photobiont population that shows no seasonal change in cell numbers. In this study, samples of L. pulmonaria were harvested in February and in May from a natural population near Sackville, New Brunswick, and subjected to two simulated intense seasonal changes: (1) early spring warming, simulated by a shift from high light at 5°C to high light at 16°C (February shift), and (2) late spring canopy closure, simulated by a shift from high light at 16°C to low light at 16°C (May shift). Thallus samples were collected daily throughout each week-long shift. There were no significant changes in photobiont cell population size or in the fraction of cells dividing during either shift. During the first day of the February temperature shift, there were, however, large changes in the pools of chlorophyll, the major light capture molecule in the photobionts, the PsbA (D1) core protein of photosystem II whose turnover is highly responsive to changing light and temperature, and the RbcL major subunit of the carbon-fixing RUBISCO enzyme whose levels correlate strongly with achieved photosynthesis in lichens. A static population of photobionts was therefore able to perform large and rapid macromolecular reallocations to cope with rapid environmental change. No significant changes were seen in the chlorophyll, photosystem II or RUBISCO pools across the May light shift, although seasonal-scale macromolecular reallocation does occur in response to decreased light in the summer.