Avoidance behavior of Hyalella azteca in response to three common-use insecticides.

Non-target organisms in aquatic environments may experience lethal or sublethal effects following exposure to contaminants. Most protocols and regulations, however, are designed to provide protection from lethal effects and are thus based on conventional estimates of population lethality. The relative lack of reliable behavioral endpoints makes it challenging to implement regulations that are similarly protective against sublethal toxicity. The objective of this study was to quantify the avoidance behavior of Hyalella azteca when exposed to three insecticides-bifenthrin (B), chlorpyrifos (C), and permethrin (P)-at a range of estimated lethal concentrations. A two-choice behavioral arena was used for each chemical to quantify H. azteca activity and time spent in either uncontaminated sediment or sediment spiked at concentrations reflecting estimated 48-h lethal concentrations (LC50, LC25, and LC10). For all three insecticides, naïve H. azteca demonstrated a preference for the uncontaminated sediment over the contaminated sediment at the LC50 (B: 312 ng/gOC; C: 1265 ng/gOC; P: 5042 ng/gOC) and LC25 (B: 230 ng/gOC; C: 859 ng/gOC; P: 3817 ng/gOC), spending significantly more time in the uncontaminated side of the arena. H. azteca did not avoid sediment at LC10 (B: 204 ng/gOC; C: 609 ng/gOC; P: 1515 ng/gOC) levels, indicating the existence of a potential threshold of detection. Despite the lack of substrate preference at this exposure level, H. azteca were nevertheless more active (i.e., increased zone-switching) when exposed to bifenthrin at the LC10, suggesting a possible irritation response (e.g., movement after exposure) to this chemical. Our results provide evidence that H. azteca exhibit innate avoidance responses to sediments contaminated with common insecticides at concentrations below those represented by traditional toxicological endpoints (e.g., LC50). The sensitivity and ease with which this behavioral endpoint can be assayed demonstrates the potential utility of behavioral endpoints in toxicological assessments using model organisms.

Eavesdropping Micropredators as Dynamic Limiters of Sexual Signal Elaboration and Intrasexual Competition.

AbstractTo thoroughly understand the drivers of dynamic signal elaboration requires assessing the direct and indirect effects of naturally interacting factors. Here, we use structural equation modeling to test multivariate data from in situ observations of sexual signal production against a model of causal processes hypothesized to drive signal elaboration. We assess direct and indirect effects, and relative impacts, of male-male competition and attacks by eavesdropping frog-biting midges (Diptera: Corethrellidae) on call elaboration of male túngara frogs (Engystomops pustulosus). We find that the intensity of attacks by these micropredator flies drives the extent to which frogs elaborate their calls, likely due to a temporal trade-off between signaling and antimicropredator defense. Micropredator attacks appear to dynamically limit a male's call rate and complexity and consequently dampen the effects of intrasexual competition. In accounting for naturally interacting drivers of signal elaboration, this study presents a counterpoint to the mechanisms traditionally thought to drive sexual selection in this system. Moreover, the results shed light on the relatively unexamined and potentially influential role of eavesdropping micropredators in the evolution of sexual communication systems.

Integrating theoretical and empirical approaches for a robust understanding of endocrine flexibility.

There is growing interest in studying hormones beyond single 'snapshot' measurements, as recognition that individual variation in the endocrine response to environmental change may underlie many rapid, coordinated phenotypic changes. Repeated measures of hormone levels in individuals provide additional insight into individual variation in endocrine flexibility - that is, how individuals modulate hormone levels in response to the environment. The ability to quickly and appropriately modify phenotype is predicted to be favored by selection, especially in unpredictable environments. The need for repeated samples from individuals can make empirical studies of endocrine flexibility logistically challenging, but methods based in mathematical modeling can provide insights that circumvent these challenges. Our Review introduces and defines endocrine flexibility, reviews existing studies, makes suggestions for future empirical work, and recommends mathematical modeling approaches to complement empirical work and significantly advance our understanding. Mathematical modeling is not yet widely employed in endocrinology, but can be used to identify innovative areas for future research and generate novel predictions for empirical testing.

Mathematical modeling reveals how the speed of endocrine regulation should affect baseline and stress-induced glucocorticoid levels.

Unpredictable environmental changes displace individuals from homeostasis and elicit a stress response. In vertebrates, the stress response is mediated mainly by glucocorticoids (GCs) which initiate physiological changes while minimizing allostatic overload. Individuals and species vary consistently in baseline and stress-induced GC levels and the speed with which GC levels can be upregulated or downregulated, but the extent to which variation in hormone regulation influences baseline and stress-induced GC levels is unclear. Using mathematical modeling, we tested how GC regulation rate, frequencies and durations of acute stressors, fitness functions, and allostatic overload affect GC levels during control and acute stress periods. As GC regulation rate slows, baseline and acute stress-induced GC levels become more similar. When the speed of up- and downregulation decreased, hormone levels became more linked to anticipated future conditions to avoid fitness costs of mismatching a new environmental state. More frequent acute stressors caused baseline and acute stress-induced GC levels to converge. When fitness was more tightly linked to hormone levels during acute stress periods than during control states, the speed of upregulation influenced optimal hormone levels more than the downregulation rate. With allostatic overload costs included, predicted GC levels were lower and more dependent on the frequency of past acute stressors. Our results show the value of optimality modeling to study the hormonal response to stressors and suggest GC levels depend on past and anticipated future environmental states as well as individual differences in hormone regulation.

Mixed Sex Effects on the Second-to-Fourth Digit Ratio of Túngara Frogs (Engystomops pustulosus) and Cane Toads (Rhinella marina).

Sexual dimorphism in the ratio of digit lengths has been correlated to behavioral, physiological, and morphological traits in a variety of taxa. While sexual dimorphism in the second-to-fourth digit length ratio (2D:4D) is a well-established indicator of prenatal androgen exposure in mammals, investigations into the patterns of 2D:4D and the drivers of such variation in other taxa are lacking. We used linear mixed effects models to gain a mechanistic understanding of the factors that drive variation in the scaling relationship between the lengths of the second and fourth digits in two species of anurans: túngara frogs (Engystomops pustulosus) and cane toads (Rhinella marina). We found evidence for sexual dimorphism of the 2D:4D scaling relationship on the front feet of túngara frogs, with female frogs having a larger ratio than males resulting from a relatively longer second digit on females. To our knowledge, this mammal-like pattern of sex differences in digit ratio has not yet been reported for anurans. However, given the reduced number of digits on the front feet of anurans, and uncertainty about which digit was lost during evolutionary history, this apparent sexual dimorphism in the front feet of túngara frogs should be treated with caution. In contrast, we found no evidence of sexual dimorphism in 2D:4D on either the front or rear feet of cane toads. This study highlights ambiguities in 2D:4D across taxa and suggests that further research is needed to evaluate the effect of androgens on 2D:4D in animals other than placental mammals.

Size matters: insights from an allometric approach to evaluate control methods for invasive Australian Rhinella marina.

Invasive species are costly and difficult to control. In order to gain a mechanistic understanding of potential control measures, individual-based models uniquely parameterized to reflect the salient life-history characteristics of invasive species are useful. Using invasive Australian Rhinella marina as a case study, we constructed a cohort- and individual-based population simulation that incorporates growth and body size of terrestrial stages. We used this allometric approach to examine the efficacy of nontraditional control methods (i.e., tadpole alarm chemicals and native meat ants) that may have indirect effects on population dynamics mediated by effects on body size. We compared population estimates resulting from these control methods with traditional hand removal. We also conducted a sensitivity analysis to investigate the effect that model parameters, specifically those associated with growth and body size, had on adult population estimates. Incremental increases in hand removal of adults and juveniles caused nonlinear decreases in adult population estimates, suggesting less return with increased investment in hand-removal efforts. Applying tadpole alarm chemicals or meat ants decreased adult population estimates on the same level as removing 15-25% of adults and juveniles by hand. The combined application of tadpole alarm chemicals and meat ants resulted in approximately 80% decrease in adult abundance, the largest of any applied control method. In further support of the nontraditional control methods, which greatly affected the metamorph stage, our model was most sensitive to changes in metamorph survival, juvenile survival, metamorph growth rate, and adult survival. Our results highlight the use and insights that can be gained from individual-based models that incorporate growth and body size and the potential success that nontraditional control methods could have in controlling established, invasive Rhinella marina populations.