Robbie S. Wilson

Tradeoffs and the evolution of thermal reaction norms

Tradeoffs have played a prominent role in the development of theories describing the evolution of reaction norms. Different classes of tradeoffs are known to constrain the evolution of phenotypes, but current theories incorporate only a subset of these tradeoffs. Consequently, these theories cannot account for some of the variation in reaction norms that has been observed within and among species. Empirical studies of thermal reaction norms for physiological and life historical traits have shown that different proximate mechanisms can produce similar reaction norms. As a consequence, certain tradeoffs can be circumvented when the fitness costs imposed by these tradeoffs are severe. We argue that a unified theory that includes all classes of tradeoffs would provide a better understanding of the mechanisms that drive the evolution of reaction norms.

Testing the beneficial acclimation hypothesis

Recent developments in evolutionary physiology have seen many of the long-held assumptions within comparative physiology receive rigorous experimental analysis. Studies of the adaptive significance of physiological acclimation exemplify this new evolutionary approach. The beneficial acclimation hypothesis (BAH) was proposed to describe the assumption that all acclimation changes enhance the physiological performance or fitness of an individual organism. To the surprise of most physiologists, all empirical examinations of the BAH have rejected its generality. However, we suggest that these examinations are neither direct nor complete tests of the functional benefit of acclimation. We consider them to be elegant analyses of the adaptive significance of developmental plasticity, a type of phenotypic plasticity that is very different from the traditional concept of acclimation that is used by comparative physiologists.

Coadaptation: A Unifying Principle in Evolutionary Thermal Biology*

Over the last 50 yr, thermal biology has shifted from a largely physiological science to a more integrated science of behavior, physiology, ecology, and evolution. Today, the mechanisms that underlie responses to environmental temperature are being scrutinized at levels ranging from genes to organisms. From these investigations, a theory of thermal adaptation has emerged that describes the evolution of thermoregulation, thermal sensitivity, and thermal acclimation. We review and integrate current models to form a conceptual model of coadaptation. We argue that major advances will require a quantitative theory of coadaptation that predicts which strategies should evolve in specific thermal environments. Simply combining current models, however, is insufficient to understand the responses of organisms to thermal heterogeneity; a theory of coadaptation must also consider the biotic interactions that influence the net benefits of behavioral and physiological strategies. Such a theory will be challenging to develop because each organism’s perception of and response to thermal heterogeneity depends on its size, mobility, and life span. Despite the challenges facing thermal biologists, we have never been more pressed to explain the diversity of strategies that organisms use to cope with thermal heterogeneity and to predict the consequences of thermal change for the diversity of communities.

Predicting the Physiological Performance of Ectotherms in Fluctuating Thermal Environments

SUMMARY Physiological ecologists have long sought to understand the plasticity of organisms in environments that vary widely among years, seasons and even hours. This is now even more important because human-induced climate change is predicted to affect both the mean and variability of the thermal environment. Although environmental change occurs ubiquitously, relatively few researchers have studied the effects of fluctuating environments on the performance of developing organisms. Even fewer have tried to validate a framework for predicting performance in fluctuating environments. Here, we determined whether reaction norms based on performance at constant temperatures (18, 22, 26, 30 and 34°C) could be used to predict embryonic and larval performance of anurans at fluctuating temperatures (18–28°C and 18–34°C). Based on existing theory, we generated hypotheses about the effects of stress and acclimation on the predictability of performance in variable environments. Our empirical models poorly predicted the performance of striped marsh frogs (Limnodynastes peronii) at fluctuating temperatures, suggesting that extrapolation from studies conducted under artificial thermal conditions would lead to erroneous conclusions. During the majority of ontogenetic stages, growth and development in variable environments proceeded more rapidly than expected, suggesting that acute exposures to extreme temperatures enable greater performance than do chronic exposures. Consistent with theory, we predicted performance more accurately for the less variable thermal environment. Our results underscore the need to measure physiological performance under naturalistic thermal conditions when testing hypotheses about thermal plasticity or when parameterizing models of life-history evolution.

Thermal acclimation of locomotor performance in tadpoles of the frog Limnodynastes peronii.

Abstract Previous analyses of thermal acclimation of locomotor performance in amphibians have only examined the adult life history stage and indicate that the locomotor system is unable to undergo acclimatory changes to temperature. In this study, we examined the ability of tadpoles of the striped marsh frog (Limnodynastes peronii) to acclimate their locomotor system by exposing them to either 10 °C or 24 °C for 6 weeks and testing their burst swimming performance at 10, 24, and 34 °C. At the test temperature of 10 °C, maximum velocity (Umax) of the 10 °C-acclimated tadpoles was 47% greater and maximum acceleration (Amax) 53% greater than the 24 °C-acclimated animals. At 24 °C, Umax was 16% greater in the 10 °C-acclimation group, while there was no significant difference in Amax or the time taken to reach Umax (T-Umax). At 34 °C, there was no difference between the acclimation groups in either Umax or Amax, however T-Umax was 36% faster in the 24 °C-acclimation group. This is the first study to report an amphibian (larva or adult) possessing the capacity to compensate for cool temperatures by thermal acclimation of locomotor performance. To determine whether acclimation period affected the magnitude of the acclimatory response, we also acclimated tadpoles of L. peronii to 10 °C for 8 months and compared their swimming performance with tadpoles acclimated to 10 °C for 6 weeks. At the test temperatures of 24 °C and 34 °C, Umax and Amax were significantly slower in the tadpoles acclimated to 10 °C for 8 months. At 10 °C, T-Umax was 40% faster in the 8-month group, while there were no differences in either Umax or Amax. Although locomotor performance was enhanced at 10 °C by a longer acclimation period, this was at the expense of performance at higher temperatures.

Thermal acclimation of locomotor performance in tadpoles and adults of the aquatic frog Xenopus laevis

Abstract Among amphibians, the ability to compensate for the effects of temperature on the locomotor system by thermal acclimation has only been reported in larvae of a single species of anuran. All other analyses have examined predominantly terrestrial adult life stages of amphibians and found no evidence of thermal acclimatory capacity. We examined the ability of both tadpoles and adults of the fully aquatic amphibian Xenopus laevis to acclimate their locomotor system to different temperatures. Tadpoles were acclimated to either 12 °C or 30 °C for 4 weeks and their burst swimming performance was assessed at four temperatures between 5 °C and 30 °C. Adult X. laevis were acclimated to either 10 °C or 25 °C for 6 weeks and their burst swimming performance and isolated muscle performance was determined at six temperatures between 5 °C and 30 °C. Maximum swimming performance of cold-acclimated X. laevis tadpoles was greater at cool temperatures and lower at the highest temperature in comparison with the warm-acclimated animals. At the test temperature of 12 °C, maximum swimming velocity of tadpoles acclimated to 12 °C was 38% higher than the 30 °C-acclimation group, while at 30 °C, maximum swimming velocity of the 30 °C-acclimation group was 41% faster than the 12 °C-acclimation group. Maximum swimming performance of adult X. laevis acclimated to 10 °C was also higher at the lower temperatures than the 25 °C acclimated animals, but there was no difference between the treatment groups at higher temperatures. When tested at 10 °C, maximum swimming velocity of the 10 °C-acclimation group was 67% faster than the 25 °C group. Isolated gastrocnemius muscle fibres from adult X. laevis acclimated to 10 °C produced higher relative tetanic tensions and decreased relaxation times at 10 °C in comparison with animals acclimated to 25 °C. This is only the second species of amphibian, and the first adult life stage, reported to have the capacity to thermally acclimate locomotor performance.

Urban Physiology: City Ants Possess High Heat Tolerance

Urbanization has caused regional increases in temperature that exceed those measured on a global scale, leading to urban heat islands as much as 12°C hotter than their surroundings. Optimality models predict ectotherms in urban areas should tolerate heat better and cold worse than ectotherms in rural areas. We tested these predications by measuring heat and cold tolerances of leaf-cutter ants from South Americas largest city (São Paulo, Brazil). Specifically, we compared thermal tolerances of ants from inside and outside of the city. Knock-down resistance and chill-coma recovery were used as indicators of heat and cold tolerances, respectively. Ants from within the city took 20% longer to lose mobility at 42°C than ants from outside the city. Interestingly, greater heat tolerance came at no obvious expense of cold tolerance; hence, our observations only partially support current theory. Our results indicate that thermal tolerances of some organisms can respond to rapid changes in climate. Predictive models should account for acclimatory and evolutionary responses during climate change.

Morphological and physiological specialization for digging in amphisbaenians, an ancient lineage of fossorial vertebrates.

SUMMARY Amphisbaenians are legless reptiles that differ significantly from other vertebrate lineages. Most species dig underground galleries of similar diameter to that of the animal. We studied the muscle physiology and morphological attributes of digging effort in the Brazilian amphisbaenid Leposternon microcephalum (Squamata; Amphisbaenia), which burrows by compressing soil against the upper wall of the tunnel by means of upward strokes of the head. The individuals tested (<72 g) exerted forces on the soil of up to 24 N. These forces were possible because the fibres of the longissimus dorsi, the main muscle associated with burrowing, are highly pennated, thus increasing effective muscle cross-sectional area. The muscle is characterized by a metabolic transition along its length: proximal, medial and distal fibres are fast contracting and moderately oxidative, but fibres closer to the head are richer in citrate synthase and more aerobic in nature. Distal fibres, then, might be active mainly at the final step of the compression stroke, which requires more power. For animals greater than a given diameter, the work required to compress soil increases exponentially with body diameter. Leposternon microcephalum, and probably some other highly specialized amphisbaenids, are most likely constrained to small diameters and can increase muscle mass and effective muscle cross-sectional area by increasing body length, not body diameter.

Dishonest Signals of Strength in Male Slender Crayfish (Cherax dispar) during Agonistic Encounters

Many animals resolve disputes without combat by displaying signals of potential strength during threatening displays. Presumably, competitors use each others displays to assess their relative strengths, and current theory predicts that these signals of strength should generally be honest. We tested this prediction by investigating the relationships among morphology, performance, and social dominance in males of the slender crayfish Cherax dispar. Crayfish routinely use their enlarged front claws (chelae) for both intimidation and fighting, making this species ideal for studying the honesty of weapon size. We evaluated five competing models relating morphological and physiological traits to dominance during paired competitive bouts. Based on the best model, larger chelae clearly resulted in greater dominance; however, chela strength had no bearing on dominance. Thus, displays of chela size were dishonest signals of strength, and the enlarged chelae of males seemingly function more for intimidation than for fighting. In addition, an analysis of the performance of isolated chela muscle showed that muscle from male crayfish produced only half the force that muscle from female crayfish produced ( \documentclass{aastex} \usepackage{amsbsy} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{bm} \usepackage{mathrsfs} \usepackage{pifont} \usepackage{stmaryrd} \usepackage{textcomp} \usepackage{portland,xspace} \usepackage{amsmath,amsxtra} \usepackage[OT2,OT1]{fontenc} \newcommand\cyr{ \renewcommand\rmdefault{wncyr} \renewcommand\sfdefault{wncyss} \renewcommand\encodingdefault{OT2} \normalfont \selectfont} \DeclareTextFontCommand{\textcyr}{\cyr} \pagestyle{empty} \DeclareMathSizes{10}{9}{7}{6} \begin{document} \landscape