Craig E. Franklin

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.

What is conservation physiology? Perspectives on an increasingly integrated and essential science †

The definition of ‘conservation physiology’ is refined to be more inclusive, with an emphasis on characterizing diversity, understanding and predicting responses to environmental change and stressors, and generating solutions. The integrative discipline is focused on mechanisms and uses physiological tools, concepts, and knowledge to advance conservation and resource management.

Determining environmental causes of biological effects: the need for a mechanistic physiological dimension in conservation biology

The emerging field of Conservation Physiology links environmental change and ecological success by the application of physiological theory, approaches and tools to elucidate and address conservation problems. Human activity has changed the natural environment to a point where the viability of many ecosystems is now under threat. There are already many descriptions of how changes in biological patterns are correlated with environmental changes. The next important step is to determine the causative relationship between environmental variability and biological systems. Physiology provides the mechanistic link between environmental change and ecological patterns. Physiological research, therefore, should be integrated into conservation to predict the biological consequences of human activity, and to identify those species or populations that are most vulnerable.

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.

A falsification of the thermal specialization paradigm: compensation for elevated temperatures in Antarctic fishes.

Specialization to a particular environment is one of the main factors used to explain species distributions. Antarctic fishes are often cited as a classic example to illustrate the specialization process and are regarded as the archetypal stenotherms. Here we show that the Antarctic fish Pagothenia borchgrevinki has retained the capacity to compensate for chronic temperature change. By displaying astounding plasticity in cardiovascular response and metabolic control, the fishes maintained locomotory performance at elevated temperatures. Our falsification of the specialization paradigm indicates that the effect of climate change on species distribution and extinction may be overestimated by current models of global warming.

Physiological mechanisms of thermoregulation in reptiles: a review

The thermal dependence of biochemical reaction rates means that many animals regulate their body temperature so that fluctuations in body temperature are small compared to environmental temperature fluctuations. Thermoregulation is a complex process that involves sensing of the environment, and subsequent processing of the environmental information. We suggest that the physiological mechanisms that facilitate thermoregulation transcend phylogenetic boundaries. Reptiles are primarily used as model organisms for ecological and evolutionary research and, unlike in mammals, the physiological basis of many aspects in thermoregulation remains obscure. Here, we review recent research on regulation of body temperature, thermoreception, body temperature set-points, and cardiovascular control of heating and cooling in reptiles. The aim of this review is to place physiological thermoregulation of reptiles in a wider phylogenetic context. Future research on reptilian thermoregulation should focus on the pathways that connect peripheral sensing to central processing which will ultimately lead to the thermoregulatory response.

Antarctic fish can compensate for rising temperatures: Thermal acclimation of cardiac performance in Pagothenia borchgrevinki

SUMMARY Antarctic fish Pagothenia borchgrevinki in McMurdo Sound, Antarctica, inhabit one of the coldest and most thermally stable of all environments. Sea temperatures under the sea ice in this region remain a fairly constant –1.86°C year round. This study examined the thermal plasticity of cardiac function in P. borchgrevinki to determine whether specialisation to stable low temperatures has led to the loss of the ability to acclimate physiological function. Fish were acclimated to– 1°C and 4°C for 4–5 weeks and cardiac output was measured at rest and after exhaustive exercise in fish acutely transferred from their acclimation temperature to –1, 2, 4, 6 and 8°C. In the– 1°C acclimated fish, the factorial scope for cardiac output was greatest at –1°C and decreased with increasing temperature. Increases in cardiac output with exercise in the –1°C acclimated fish was achieved by increases in both heart rate and stroke volume. With acclimation to 4°C, resting cardiac output was thermally independent across the test temperatures; furthermore, factorial scope for cardiac output was maintained at 4, 6 and 8°C, demonstrating thermal compensation of cardiac function at the higher temperatures. This was at the expense of cardiac function at –1°C, where there was a significant decrease in factorial scope for cardiac output in the 4°C acclimated fish. Increases in cardiac output with exercise in the 4°C acclimated fish at the higher temperatures was achieved by changes in heart rate alone, with stroke volume not varying between rest and exercise. The thermal compensation of cardiac function in P. borchgrevinki at higher temperatures was the result of a change in pumping strategy from a mixed inotropic/chronotropic modulated heart in –1°C acclimated fish at low temperatures to a purely chronotropic modulated heart in the 4°C acclimated fish at higher temperatures. In spite of living in a highly stenothermal cold environment, P. borchgrevinki demonstrated the capacity to thermally acclimate cardiac function to elevated temperatures, thereby allowing the maintenance of factorial scope and the support of aerobic swimming at higher temperatures.

Adaptation of rainbow fish to lake and stream habitats

Abstract Fish occupy a range of hydrological habitats that exert different demands on locomotor performance. We examined replicate natural populations of the rainbow fishes Melanotaenia eachamensis and M. duboulayi to determine if colonization of low-velocity (lake) habitats by fish from high-velocity (stream) habitats resulted in adaptation of locomotor morphology and performance. Relative to stream conspecifics, lake fish had more posteriorly positioned first dorsal and pelvic fins, and shorter second dorsal fin bases. Habitat dimorphism observed between wild-caught fish was determined to be heritable as it was retained in M. eachamensis offspring raised in a common garden. Repeated evolution of the same heritable phenotype in independently derived populations indicated body shape divergence was a consequence of natural selection. Morphological divergence between hydrological habitats did not support a priori expectations of deeper bodies and caudal peduncles in lake fish. However, observed divergence in fin positioning was consistent with a family-wide association between habitat and morphology, and with empirical studies on other fish species. As predicted, decreased demand for sustained swimming in lakes resulted in a reduction in caudal red muscle area of lake fish relative to their stream counterparts. Melanotaenia duboulayi lake fish also had slower sustained swimming speeds (Ucrit) than stream conspecifics. In M. eachamensis, habitat affected Ucrit of males and females differently. Specifically, females exhibited the pattern observed in M. duboulayi (lake fish had faster Ucrit than stream fish), but the opposite association was observed in males (stream males had slower Ucrit than lake males). Stream M. eachamensis also exhibited a reversed pattern of sexual dimorphism in Ucrit (males slower than females) relative to all other groups (males faster than females). We suggest that M. eachamensis males from streams responded to factors other than water velocity. Although replication of muscle and Ucrit phenotypes across same habitat populations within and/or among species was suggestive of adaptation, the common garden experiment did not confirm a genetic basis to these associations. Kinematic studies should consider the effect of the position and base length of dorsal fins.

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.

The aerobic scope of an antarctic fish, Pagothenia borchgrevinki and its significance for metabolic cold adaptation

SummaryResting weight-specific oxygen consumption of the cryopelagic Antarctic nototheniid Pagothenia borchgrevinki at 0°C was 39.6 ml kg-1 · h-1 for a 50 g fish, with oxygen consumption being described by the regression equation: log10 VO2(ml/h)=−1.104+0.825 log10 Mb (g). These values are considerably below those raported by Wohlschlag (1964a,b). VO2 max. in forced swimming was described by the regression equation: log10 VO2 max = −0.507+0.823 log10 Mb. Despite low basal metabolism, factorial aerobic scope is similar to that reported for most other teleost fish, as is the cost of net transport. Myotomal muscles were used only at the highest swimming speeds and once they were recruited the fish fatigued rapidly. After swimming, oxygen debt was repaid rapidly, with a half-time of 20 min.