Toxicity of glyphosate to animals: A meta-analytical approach.

Glyphosate (GLY)-based herbicides (GBHs) are the most commonly applied pesticide worldwide, and non-target organisms (e.g., animals) are now regularly exposed to GLY and GBHs due to the accumulation of these chemicals in many environments. Although GLY/GBH was previously considered to be non-toxic, growing evidence indicates that GLY/GBH negatively affects some animal taxa. However, there has been no systematic analysis quantifying its toxicity to animals. Therefore, we used a meta-analytical approach to determine whether there is a demonstrable effect of GLY/GBH toxicity across animals. We further addressed whether the effects of GLY/GBH vary due to (1) taxon (invertebrate vs. vertebrate), (2) habitat (aquatic vs. terrestrial), (3) type of biological response (behavior vs. physiology vs. survival), and (4) dosage or concentration of GLY/GBH. Using this approach, we also determined whether adjuvants (e.g., surfactants) in commercial formulations of GBHs increased toxicity for animals relative to exposure to GLY alone. We analyzed 1282 observations from 121 articles. We conclude that GLY is generally sub-lethally toxic for animals, particularly for animals in aquatic or marine habitats, and that toxicity did not exhibit dose-dependency. Yet, our analyses detected evidence for widespread publication bias so we encourage continued experimental investigations to better understand factors influencing GLY/GBH toxicity to animals.

A simulated heat wave-but not herbicide exposure-alters resource investment strategy in an insect.

Animals are increasingly exposed to potential stressors related to environmental change, and multiple stressors may alter the dynamics by which animals acquire resources and invest those resources into important life-history traits. Stress may lead to the prioritization of current reproduction to maximize lifetime reproduction (i.e., terminal investment [TI]) or, in contrast, prioritize somatic investment over current reproduction to facilitate future reproductive opportunities (i.e., reproductive restraint [RR]). Tests of the TI and RR hypotheses typically use immune challenges as stressors, and have not been explicitly tested in the context of environmental change even though warming influences resource allocation patterns across taxa. Further, the multiple-stressor framework has been a useful construct to clarify the costs of complex environmental shifts to animals, but it has not been leveraged to understand such effects on investment strategy. Thus, we tested the TI and RR hypotheses by manipulating widespread features of environmental change-glyphosate-based herbicide (GBH; Roundup®) exposure and a simulated heat wave-in the variable field cricket (Gryllus lineaticeps). A simulated heat wave affected the life-history tradeoff between investment into reproduction and soma. Specifically, heat wave prioritized investment into ovary mass over non-reproductive tissue, even after accounting for food consumption, in support of the TI hypothesis. In contrast, GBH exposure did not affect any measured trait, and crickets did not discriminate between tap water and GBH solution during drinking. Therefore, some-but not all-aspects of environmental change may alter resource investment strategies in animals. We encourage continued integration of the multiple-stressor framework and life-history theory to better understand how animals respond to their rapidly changing environments.

Pesticides in a warmer world: Effects of glyphosate and warming across insect life stages.

Glyphosate (GLY) is a broad-spectrum herbicide that is the most commonly applied pesticide in terrestrial ecosystems in the U.S. and, potentially, worldwide. However, the combined effects of warming associated with climate change and exposure to GLY and GLY-based formulations (GBFs) on terrestrial animals are poorly understood. Animals progress through several life stages (e.g., embryonic, larval, and juvenile stages) that may exhibit different sensitivities to stressors. Therefore, we factorially manipulated temperature and GLY/GBF exposure in the variable field cricket (Gryllus lineaticeps) during two life stages-nymphal development and adulthood-and examined key animal traits, such as developmental rate, body size, food consumption, reproductive investment, and lifespan. A thermal environment simulating future climate warming obligated several costs to fitness-related traits. For example, warming experienced during nymphal development reduced survival, adult body mass and size, and investment into flight capacity and reproduction. Warming experienced by adults reduced lifespan and growth rate. Alternatively, the effects of GBF exposure were more subtle, often context-dependent (e.g., effects were only detected in one sex or temperature regime), and were stronger during adult exposure relative to exposure during development. There was evidence of additive costs of warming and GBF exposure to rates of feeding and growth in adults. Yet, the negative effect of GBF exposure to adult lifespan did not occur in warming conditions, suggesting that ongoing climate change may obscure some of the costs of GBFs to non-target organisms. The effects of GLY alone (i.e., in the absence of proprietary surfactants found in commercial formulations) were non-existent. Animals will be increasingly exposed to warming and GBFs, and our results indicate that GBF exposure and warming can entail additive costs for an animal taxon (insects) that plays critical roles in terrestrial ecosystems.

From phenoloxidase to fecundity: food availability does not influence the costs of oxidative challenge in a wing-dimorphic cricket.

Stressed animals often struggle to maintain optimal investment into a number of fitness-related traits, which can result in some traits being more adversely affected than others. Variation in stress-related costs may also depend on the environment-costs can be facultative and only occur when resources are limited, or they may be obligate and occur regardless of resource availability. Dynamics of oxidative stress may be important in life-history evolution given their role in a range of biological processes-from reproduction to immunity to locomotion. Thus, we examined how resource (food) availability influences the costs of oxidative challenge to fitness-related traits spanning several levels of biological organization. We manipulated food availability and oxidative status in females of the wing-dimorphic sand field cricket (Gryllus firmus) during early adulthood. We then determined investment into several traits: reproduction (ovary mass), soma (body mass and flight musculature), and immune function (total phenoloxidase activity). Oxidative challenge (paraquat exposure) obligated costs to somatic tissue and a parameter of immune function regardless of food availability, but it did not affect reproduction. We show that the costs of oxidative challenge are trait-specific, but we did not detect a facultative (food-dependent) cost of oxidative challenge to any trait measured. Although the dynamics of oxidative stress are complex, our study is an important step toward a more complete understanding of the roles that resource availability and redox systems play in mediating life histories.

A Simulated Heat Wave Has Diverse Effects on Immune Function and Oxidative Physiology in the Corn Snake (Pantherophis guttatus).

Animals will continue to encounter increasingly warm environments, including more frequent and intense heat waves. Yet the physiological consequences of heat waves remain equivocal, potentially because of variation in adaptive plasticity (reversible acclimation) and/or aspects of experimental design. Thus, we measured a suite of physiological variables in the corn snake (Pantherophis guttatus) after exposure to field-parameterized, fluctuating temperature regimes (moderate temperature and heat wave treatments) to address two hypotheses: (1) a heat wave causes physiological stress, and (2) thermal performance of immune function exhibits adaptive plasticity in response to a heat wave. We found little support for our first hypothesis because a simulated heat wave had a negative effect on body mass, but it also reduced oxidative damage and did not affect peak performance of three immune metrics. Likewise, we found only partial support for our second hypothesis. After exposure to a simulated heat wave, P. guttatus exhibited greater performance breadth and reduced temperature specialization (the standardized difference between peak performance and performance breadth) for only one of three immune metrics and did so in a sex-dependent manner. Further, a simulated heat wave did not elicit greater performance of any immune metric at higher temperatures. Yet a heat wave likely reduced innate immune function in P. guttatus because each metric of innate immune performance in this species (as in most vertebrates) was lower at elevated temperatures. Together with previous research, our study indicates that a heat wave may have complex, modest, and even positive physiological effects in some taxa.

Context dependency and generality of fever in insects.

Fever can reduce mortality in infected animals. Yet, despite its fitness-enhancing qualities, fever often varies among animals. We used several approaches to examine this variation in insects. Texas field crickets (Gryllus texensis) exhibited a modest fever (1 °C increase in preferred body temperature, T pref) after injection of prostaglandin, which putatively mediates fever in both vertebrates and invertebrates, but they did not exhibit fever during chronic exposure to heat-killed bacteria. Further, chronic food limitation and mating status did not affect T pref or the expression of behavioural fever, suggesting limited context dependency of fever in G. texensis. Our meta-analysis of behavioural fever studies indicated that behavioural fever occurs in many insects, but it is not ubiquitous. Thus, both empirical and meta-analytical results suggest that the fever response in insects 'is widespread, although certainly not inevitable' (Moore 2002). We highlight the need for future work focusing on standardizing an experimental protocol to measure behavioural fever, understanding the specific mechanism(s) underlying fever in insects, and examining whether ecological or physiological costs often outweigh the benefits of fever and can explain the sporadic nature of fever in insects.