Can poisons stimulate bees? Appreciating the potential of hormesis in bee–pesticide research

Hormesis, a biphasic dose response whereby exposure to low doses of a stressor can stimulate biological processes, has been reported in many organisms, including pest insects when they are exposed to low doses of a pesticide. However, awareness of the hormesis phenomenon seems to be limited among bee researchers, in spite of the increased emphasis of late on pollinator toxicology and risk assessment. In this commentary, we show that there are several examples in the literature of substances that are toxic to bees at high doses but stimulatory at low doses. Appreciation of the hormetic dose response by bee researchers will improve our fundamental understanding of how bees respond to low doses of chemical stressors, and may be useful in pollinator risk assessment. © 2015 Society of Chemical Industry     

Low doses of glyphosate change the responses of soyabean to subsequent glyphosate treatments

Many herbicides promote plant growth at doses well below the recommended application rate (hormesis). The objectives of this study were to evaluate glyphosate‐induced hormesis in soyabean (Glycine max) and determine whether pre‐treating soyabean seedlings with low doses of glyphosate would affect their response to subsequent glyphosate treatments. Seven doses (1.8–720 g a.e. ha^?1) of glyphosate were applied to 3‐week‐old seedlings, and the effects on the electron transport rate (ETR), metabolite (shikimate, benzoate, salicylate, AMPA, phenylalanine, tyrosine and tryptophan) levels and dry weight were determined. The lowest dose stimulated ETR and increased biomass the most. Benzoate levels increased 203% with 3.6 g a.e. ha^?1 glyphosate. Salicylate content and tyrosine content were unaffected, whereas phenylalanine and tryptophan levels were increased by 60 and 80%, respectively, at 7.2 g a.e. ha^?1. Dose–response curves for these three amino acids were typical for hormesis. In another experiment that was replicated twice, soyabean plants were pre‐treated with low doses of glyphosate (1.8, 3.6 or 7.2 g a.e. ha^?1) and treated with a second application of glyphosate (1.8, 3.6, 7.2, 36, 180 or 720 g a.e. ha^?1) 14 days later. For total seedling dry weight, a 3.6 and 7.2 g a.e. ha^?1 glyphosate dose preconditioned the soyabean seedlings to have greater growth stimulation by a later glyphosate treatment than plants with no preconditioning glyphosate exposure. Optimal hormetic doses were generally higher with pre‐treated plants than plants that had not been exposed to glyphosate. Thus, pre‐exposure to low doses of glyphosate can change the hormetic response to later low‐dose exposures.     

Herbicides and plant hormesis

Herbicide hormesis is commonly observed at subtoxic doses of herbicides and other phytotoxins. The occurrence and magnitude of this phenomenon are influenced by plant growth stage and physiological status, environmental factors, the endpoint measured and the timing between treatment and endpoint measurement. The mechanism in some cases of herbicide hormesis appears to be related to the target site of the herbicide, whereas in other examples hormesis may be by overcompensation to moderate stress induced by the herbicides or a response to disturbed homeostasis. Theoretically, herbicide hormesis could be used in crop production, but this has been practical only in the case of the use of herbicides as sugar cane ‘ripeners’ to enhance sucrose accumulation. The many factors that can influence the occurrence, the magnitude and the dose range of hormetic increases in yield for most crops make it too unpredictable and risky as a production practice with the currently available knowledge. Herbicide hormesis can cause undesired effects in situations in which weeds are unintentionally exposed to hormetic doses (e.g. in adjacent fields, when shielded by crop vegetation). Some weeds that have evolved herbicide resistance may have hormetic responses to recommended herbicide application rates. Little is known about such effects under field conditions. A more complete understanding of herbicide hormesis is needed to exploit its potential benefits and to minimize its potential harmful effects in crop production. © 2014 Society of Chemical Industry     

Herbicides can stimulate plant growth

Low dose stimulations by toxicants have long been observed. Great controversies exist concerning the interpretation of these observations, spanning from believing that they are a general stress response occurring for all chemicals, to simply being an experimental artefact resulting from poorly growing control plants or from biomass allocation between plant parts. This study investigates the growth response and biomass allocation pattern of barley exposed to 10–15 doses of eight different herbicides. The results show that the globally most widely used herbicide, glyphosate, together with the sulfonylurea, metsulfuron‐methyl, can induce a real stimulation in biomass growth of approximately 25% when applied at doses corresponding to 5–10% field rate. The other six herbicides tested did not induce consistent hormesis, thereby undermining the theory of hormesis being a general stress response. Biomass allocations between plant parts did take place, but were not the cause of the hormetic growth stimulations. The results demonstrate that plant physiological responses to low herbicide doses cannot be extrapolated from our knowledge of effects of higher, commercially used, doses. Other physiological mechanisms seem to be triggered in the low dose‐range, and the investigation of these mechanisms poses new challenges for agronomists, environmentalists and plant physiologists.     

Insecticide‐induced hormesis and arthropod pest management

Ecological backlashes such as insecticide resistance, resurgence and secondary pest outbreaks are frequent problems associated with insecticide use against arthropod pest species. The last two have been particularly important in sparking interest in the phenomenon of insecticide‐induced hormesis within entomology and acarology. Hormesis describes a biphasic dose–response relationship that is characterized by a reversal of response between low and high doses of a stressor (e.g. insecticides). Although the concept of insecticide‐induced hormesis often does not receive sufficient attention, or has been subject to semantic confusion, it has been reported in many arthropod pest species and natural enemies, and has been linked to pest outbreaks and potential problems with insecticide resistance. The study of hormesis remains largely neglected in entomology and acarology. Here, we examined the concept of insecticide‐induced hormesis in arthropods, its functional basis and potential fitness consequences, and its importance in arthropod pest management and other areas. © 2013 Society of Chemical Industry     

Pesticide-mediated homeostatic modulation in arthropods

The term hormesis was coined to describe a phenomenon where exposure to high levels of stressors is inhibitory whereas low (mild, sublethal, and subtoxic) doses are stimulatory. The stimulatory effects are believed to be the result of compensatory biochemical processes following a destabilization of normal homeostasis. Exposure of arthropods to mild levels of chemical stressors (i.e., pesticides) may result in enhanced reproduction that has been associated along with other factors with pest outbreaks and resurgences. Hormesis, however, cannot be claimed for cases in which the observed stimulatory effects were due to exposure of non-target pests (i.e., mites) to pesticides (DDT, carbaryl, insecticidal pyrethroids or imidacloprid). Pesticides applied to non-target pests cannot be regarded as stressors since inhibition or mortality at very high doses can hardly be observed and measured. Pesticide-induced homeostatic modulation (PIHM) is suggested as a broader term to include both hormesis and stimulatory effects of pesticides on non-target pests. The specific role played by PIHM in inducing pest outbreaks in agroecosystems is difficult to evaluate as other complex environmental factors are most likely involved. The time factor is significant where applied pesticides undergo physical dissipation as well as biological, chemical, and/or photochemical modifications. A delay in outbreaks may be anticipated as arthropod pests exposed to effective residues and degradation products will be subjected to PIHM resulting in enhanced reproduction. Knowledge about hormesis and PIHM has practical aspects for designing pest control strategies and pest resistance management practices.     

Imidacloprid-induced hormesis on the fecundity and juvenile hormone levels of the green peach aphid Myzus persicae (Sulzer)

Pesticide-induced hormesis may be an alternative mechanism for pest resurgence which is a serious problem in agriculture. Confirmation of the general phenomenon of hormesis may have significant implications for the design of pest control strategies and pest resistance management practices, although this has proved difficult due to the lack of appropriate methodology and the absence of well-defined mechanisms to support the experimental observations. In this study, a model-based approach to describe a dose-response relationship incorporating the hormetic effect was applied to the detection and estimation of imidacloprid-induced hormesis in the green peach aphid, Myzus persicae (Sulzer). The results indicated that imidacloprid at low concentrations induced stimulation of fecundity, however, high concentrations inhibition. This was reflected in an inverted U-shaped curve and related to the change of juvenile hormone III (JH III) levels in M. persicae. Fitting the data with quadratic and Weibull functions, which included a parameter for hormesis, showed that the magnitude of the hormetic effect was 31.31% for fecundity and 32.21% for JH III levels. The presence of hormesis in fecundity induced by imidacloprid may be related to the change in JH III levels in M. persicae.     

Green peach aphid,Myzus persicae (Hemiptera: Aphididae), reproduction during exposure to sublethal concentrations of imidacloprid and azadirachtin

BACKGROUND: Resurgence of insect pests following insecticide applications is often attributed to natural enemy disturbance, but hormesis could be an alternative or additional mechanism. Green peach aphid, Myzus persicae (Sulzer), is an important insect pest of many crops worldwide that may be exposed to sublethal insecticide concentrations over time. Here, the hypothesis that exposure to low concentrations of imidacloprid and azadirachtin can induce hormetic responses in M. persicae is tested in the laboratory. RESULTS: When insects were exposed to potato leaf discs dipped in sublethal concentrations of insecticide, almost all measured endpoints—adult longevity, F1 production, F1 survival and F2 production—were affected, and a statistically significant (P < 0.05) stimulatory response was recorded for F2 production following exposure to imidacloprid. No other measures for hormesis were statistically significant, but other trends of hormetic response were consistently observed. CONCLUSIONS: Given that variable distribution and degradation of insecticides in the field would result in a wide range of concentrations over time and space, these laboratory experiments suggest that exposure to sublethal concentrations of imidacloprid and azadirachtin could stimulate reproduction in M. persicae. Copyright © 2008 Society of Chemical Industry     

Response of a red clone of Myzus persicae (Hemiptera: Aphididae) to sublethal concentrations of imidacloprid in the laboratory and greenhouse

BACKGROUND: Pest resurgence following a pesticide application may occur owing to a stimulatory (hormetic) response to sublethal insecticide concentrations. The objective of the present study was to examine the potential for a greenhouse‐derived red clone of Myzus persicae to exhibit resurgence owing to a hormetic response following a systemic imidacloprid treatment in a bell pepper greenhouse. RESULTS: No differences in mortality and fecundity were observed among apterous adults exposed to sublethal imidacloprid concentrations on excised pepper leaves fed aqueous solutions of imidacloprid. Survival of first‐generation progeny was negatively affected by imidacloprid exposure, yet surviving progeny exhibited no differences in development rates or fecundity from progeny of adults unexposed to imidacloprid. Aphid mortality declined most rapidly in clip cages on pepper leaves at the top of the pepper canopy as compared with leaves present at the middle or bottom of the pepper canopy. CONCLUSION: Imidacloprid decays rapidly in mature pepper plants, resulting in sublethal concentrations in the upper canopy in as little as 4 weeks. Sublethal insecticide concentrations have been implicated in the resurgence of pest populations; however, exposure to sublethal doses of imidacloprid are unlikely to result in pesticide‐induced resurgence of the M. persicae aphid clone examined in this study. Copyright © 2011 Society of Chemical Industry     

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

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

农药对害虫天敌的Hormesis效应研究进展

低剂量有毒物质引起的Hormesis效应是毒理学研究的热点问题。近年来,农药对害虫天敌的Hormesis效应引起了研究人员的广泛关注。本文就低剂量农药对天敌发育历期、体重、雌虫生殖率及对天敌控害能力的Hormesis效应及其机制进行了简要总结,并讨论了今后的研究方向。旨在为协调害虫综合防治中生物防治与化学防治的矛盾提供新的思路。     

Impact of Brassica juncea biofumigation on viability of propagules of pernicious weed species

Biofumigation may be a promising tool for depletion of persistent weed seedbanks/bud banks. This technique is based on the incorporation of chopped glucosinolate‐rich plant biomass into the soil, upon which isothiocyanates with herbicidal properties are released. To gain acceptance by farmers and foster its implementation, the biofumigation process should be further optimised. This study elucidated the impact of biological (species), technical (burial depth, ground cover) and pedohydrological (temperature and moisture content) factors on efficacy of Brassica juncea biofumigation under (semi‐)natural conditions. In a first experiment (field experiment), seeds and vegetative propagules of various weed species were buried at different depths and exposed to different doses of fresh fine‐chopped B. juncea biomass in the presence or absence of a plastic ground cover. In a second experiment (container experiment), buried seeds of ten species were subjected to biofumigation at diverging soil organic matter content, soil moisture content and soil temperature. In a third experiment (dose–response Petri dish bioassay), unburied seeds of eight species were subjected to various doses of rehydrated B. juncea powder. Biofumigation efficacy was determined by analysing viability of treated and untreated propagules. In general, efficacy of biofumigation increased with decreasing burial depth and increasing B. juncea dose. Biofumigation was highly effective (mortality >85%) against small‐seeded species but less effective (mortality 0%–20%) against hard‐seeded and large‐seeded species at 200 t ha^?1. Vegetative propagules of Sonchus arvensis, Equisetum arvense and Convolvulus sepium were highly sensitive (mortality >90%) to biofumigation. Efficacy was most pronounced under moist warm incubation conditions, in the presence of a plastic ground cover.     

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

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

Sensitivity of Echinochloa crus‐galli populations to maize herbicides: a comparison between cropping systems

Echinochloa crus‐galli is an important maize weed with significant variation in herbicide sensitivity. This differential response may reflect differences in selection pressure caused by years of cropping system‐related herbicide usage. The herbicide sensitivity of E. crus‐galli populations from three divergent cropping systems was evaluated in dose–response pot experiments. Populations were collected from sandy fields with (i) a long‐term organic cropping system, (ii) a conventional cropping system with maize in the crop rotation or (iii) a conventional cropping system with long‐term monocropping of maize. Each cropping system was represented by six E. crus‐galli populations. The effectiveness of three foliar‐applied maize herbicides (nicosulfuron, cycloxydim and topramezone) and two soil‐applied maize herbicides (S‐metolachlor and dimethenamid‐P) was tested at three doses and two runs. Foliar‐applied herbicides were applied at the three true leaves stage. Soil‐applied herbicides were applied immediately after sowing. The foliage dry weight per pot was determined 4 weeks after treatment. Plant responses were expressed as biomass reduction. Herbicide sensitivity was consistently lowest for populations from maize monocropping systems. Compared with populations from organic cropping systems, populations from monocropping systems showed 6.9%, 9.8% and 29.3% lower sensitivity to cycloxydim, topramezone and nicosulfuron respectively. Populations from the conventional crop rotation system showed intermediate sensitivity levels, which did not significantly differ from sensitivity levels of populations from the other cropping systems. Sensitivity to dimethenamid‐P and S‐metolachlor was not affected by cropping system. Environmental conditions influenced herbicidal response . This study indicated that integrated weed management may be necessary to preserve herbicide efficacy over the long term.     

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

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

Synthesis and herbicidal activities of novel 3-(substituted benzyloxy or phenoxy)-6-methyl-4-(3-trifluoromethylphenyl)pyridazine derivatives

BACKGROUND: 4‐(3‐Trifluoromethylphenyl)pyridazine represents a new series of compounds with bleaching and herbicidal activities. RESULTS: A total of 43 novel 3‐(substituted benzyloxy or phenoxy)‐6‐methyl‐4‐(3‐trifluoromethylphenyl)pyridazine derivatives were synthesised, and their bleaching and herbicidal activities were evaluated through Spirodela polyrrhiza and greenhouse tests. Some compounds exhibited excellent herbicidal activities, even at a dose of 7.5 g ha^?1. CONCLUSION: The results showed that a substituted phenoxy group at the 3‐position of the pyridazine ring and the electron‐withdrawing group at the para ‐position on the benzene ring were essential for high herbicidal activity. Copyright © 2011 Society of Chemical Industry     

Structure-activity relationship of herbicides inhibiting ACh-induced contractions in a molluscan smooth muscle

Certain herbicides are known to influence the muscle function of molluscs. The penis-retractor muscle complex (PRM complex) of the edible snail, Helix pomatia, is a suitable test object for studying these side effects in smooth muscle, because sequential contractions can be induced in vitro by adding and removing acetylcholine (ACh). In the presence of herbicidal amides, carbamates, and ureas (concentration, 10^?4 mol/liter) the muscle tension was found to be reduced to a variable extent. In contrast, the relaxing effect of herbicidal phenoxycarbonic acids was weak. The inhibitory properties of the herbicides tested were correlated with the electron-withdrawing properties of certain substituents. These substituents are bound to an amino function. Their structure is either aromatic (amides, biscarbamates, carbamates, ureas), aliphatic (thiocarbamates), or cycloaliphatic (cycloat). The structure-activity relationship is comparable with that found for herbicidal activity in plants.     

Olive processing wastes for weed control

The herbicidal effect of olive processing wastes (OPW) on some weed species in wheat, maize and sunflower was investigated in the Aegean region of Turkey. In trials with maize and sunflower, OPW was applied as an air‐dried solid form at 3 and 4.5 kg m^?2. It provided an effectiveness level on Portulaca oleracea of 63–98%. In trials with wheat, OPW was applied as solid and liquid forms, each at two different doses, namely 4.5 and 6 kg m^?2 (solid), and 5 and 10 L m^?2 (liquid). Solid OPW provided a reduction in total weed coverage of 75% and 81% at doses of 4.5 and 6 kg m^?2, respectively. The weed coverage reduction by liquid OPW was 39% and 62% with 5 and 10 L m^?2, respectively. Apart from 12–26% reduction of the number of germinating seeds, OPW showed no toxic effects on maize and sunflower. Wheat was affected in the initial stages but no adverse effect was detected at harvest. It can be concluded that the herbicidal effect of OPW may be considered as an alternative to chemical weed control in some important summer crops (maize and sunflower) and for most of the weeds in winter wheat.     

Modelling the effects of herbicide dose and weed density on rice‐weed competition

The effects of herbicide dose on rice‐weed competition were investigated to develop a combined model, which can be utilised to estimate an optimum herbicide dose for a given weed density in paddy rice cultivation. Field studies were conducted in Suwon for rice‐Echinochloa crus‐galli competition and Iksan for rice‐Eleocharis kuroguwai during 2007. The competitive effect of the weeds E. crus‐galli and E. kuroguwai decreased with increasing doses of flucetosulfuron and azimsulfuron, respectively, in the same manner as the standard dose–response curve. The combination of the rectangular hyperbolic model and the standard dose–response curve adequately described the complex effects of herbicide dose and weed competition on rice yield. Parameter estimates were used with the model to predict rice yield and estimate the doses of flucetosulfuron and azimsulfuron required to restrict rice yield loss caused by E. crus‐galli and E. kuroguwai, respectively, to an acceptable level. For a rice yield of 5.0 t ha^?1, the model recommended flucetosulfuron doses of 8.7, 13.4 and 20.1 g a.i. ha^?1 when infested with E. crus‐galli at 12, 24 and 48 plants m^?2 respectively. For a rice yield of 5.2 t ha^?1, the model recommended azimsulfuron doses of 3.9, 7.5 and 12.6 g a.i. ha^?1 when infested with E. kuroguwai at 24, 48 and 96 plants m^?2 respectively. The theoretical outputs of the combined model appear robust and indicate there are opportunities for reduced herbicide use in the field. These now require evaluation under field conditions.