Evaluating thermoregulation in reptiles: an appropriate null model

Established indexes of thermoregulation in ectotherms compare body temperatures of real animals with a null distribution of operative temperatures from a physical or mathematical model with the same size, shape, and color as the actual animal but without mass. These indexes, however, do not account for thermal inertia or the effects of inertia when animals move through thermally heterogeneous environments. Some recent models have incorporated body mass, to account for thermal inertia and the physiological control of warming and cooling rates seen in most reptiles, and other models have incorporated movement through the environment, but none includes all pertinent variables explaining body temperature. We present a new technique for calculating the distribution of body temperatures available to ectotherms that have thermal inertia, random movements, and different rates of warming and cooling. The approach uses a biophysical model of heat exchange in ectotherms and a model of random interaction with thermal environments over the course of a day to create a null distribution of body temperatures that can be used with conventional thermoregulation indexes. This new technique provides an unbiased method for evaluating thermoregulation in large ectotherms that store heat while moving through complex environments, but it can also generate null models for ectotherms of all sizes.     

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.     

Human males and females body thermoregulation: Perfusion effect analysis

Skin temperature is a common physiological parameter that reflects thermal responses. Blood perfusion is an important part of the physiological processes that the human body undergoes in order to maintain homeostasis. This study focuses on the effect of perfusion on the temperature distribution in human males and females body in different thermal environment. The study has been carried out for one dimensional steady cases using finite element method. The input parameter of the model is the blood perfusion or volumetric flow rate within the tissue. The appropriate physical and physiological parameters together with suitable boundary conditions that affect the heat regulations have been incorporated in the model. The study is to have a better understanding that how does thermoregulation change in human males and females skin layered due to perfusion.     

Physiological evolution during adaptive radiation: A test of the island effect in Anolis lizards

Phenotypic evolution is often exceptionally rapid on islands, resulting in numerous, ecologically diverse species. Although adaptive radiation proceeds along various phenotypic axes, the island effect of faster evolution has been mostly tested with regard to morphology. Here, we leveraged the physiological diversity and species richness of Anolis lizards to examine the evolutionary dynamics of three key traits: heat tolerance, body temperature, and cold tolerance. Contrary to expectation, we discovered slower heat tolerance evolution on islands. Additionally, island species evolve toward higher optimal body temperatures than mainland species. Higher optima and slower evolution in upper physiological limits are consistent with the Bogert effect, or evolutionary inertia due to thermoregulation. Correspondingly, body temperature is higher and more stable on islands than on the American mainland, despite similarity in thermal environments. Greater thermoregulation on islands may occur due to ecological release from competitors and predators compared to mainland environments. By reducing the costs of thermoregulation, ecological opportunity on islands may actually stymie, rather than hasten, physiological evolution. Our results emphasize that physiological diversity is an important axis of ecological differentiation in the adaptive radiation of anoles, and that behavior can impart distinct macroevolutionary footprints on physiological diversity on islands and continents.     

Thermoregulation in the comfort temperature zone

Under constant adequate temperature conditions (comfortable for a man) the retention of temperature homeostasis was shown to require a continuous functioning of the physiological thermoregulation system to prevent short-term and whole day deviations of the temperature from the physiological level. Under adequate temperature conditions the thermoregulation system was shown to attain its highest sensitivity and accuracy. It is possible that this occurs owing to physiological control over the total body heat content being included into the process of thermoregulation. The data are given on the existence and "structure" of the physiological mechanisms of such a control.     

Physiological thermoregulation in bigeye tuna,Thunnus obesus

Synopsis Although a growing body of evidence has indicated that tuna can thermoregulate and have body temperatures that are decoupled from immediate changes in ambient temperature, demonstrating the extent and time-course of body temperature changes in tuna moving through their natural environments has proved to be elusive. Here we use body temperature data telemetered from free-ranging fish to demonstrate short-latency physiological thermoregulation in bigeye tuna. We used a recently developed modeling system to determine the magnitude and time-course of the whole-body thermal conductivity changes that would result in the body temperature changes observed in fish in the wild. The results indicate rapid, 100 to 1000-fold changes in whole-body thermal conductivity that occur in response to quickly changing ambient temperatures. Coupling this physiological response with behavioral thermoregulation expands the foraging space of these animals by permitting activity in wide ranges of water temperatures and depths.Paper from the International Union of Biological Societies symposium The biology of tunas and billfishes: an examination of life on the knife edge, organized by Richard W. Brill and Kim N. Holland.     

Review on modeling heat transfer and thermoregulatory responses in human body

Several mathematical models of human thermoregulation have been developed, contributing to a deep understanding of thermal responses in different thermal conditions and applications. In these models, the human body is represented by two interacting systems of thermoregulation: the controlling active system and the controlled passive system. This paper reviews the recent research of human thermoregulation models. The accuracy and scope of the thermal models are improved, for the consideration of individual differences, integration to clothing models, exposure to cold and hot conditions, and the changes of physiological responses for the elders. The experimental validated methods for human subjects and manikin are compared. The coupled method is provided for the manikin, controlled by the thermal model as an active system. Computational Fluid Dynamics (CFD) is also used along with the manikin or/and the thermal model, to evaluate the thermal responses of human body in various applications, such as evaluation of thermal comfort to increase the energy efficiency, prediction of tolerance limits and thermal acceptability exposed to hostile environments, indoor air quality assessment in the car and aerospace industry, and design protective equipment to improve function of the human activities.     

Bombesian alters behavioral thermoregulation in fish

Bombesin was microinjected into the ventricular system of a teleost fish, $\text{Catostomus commersoni}$, in order to evaluate its effects on behavioral thermoregulation in a horizontal thermal gradient. General locomotor activity was also examined. Results demonstrated that bombesin decreased the behaviorally selected internal body temperatures by 5.0–7.0°C relative to controls and that the latency of the response was dose-dependent. The peptide did not change locomotor activity. Results are discussed in relation to body temperature setpoint(s) and possible implications for endotherm thermoregulation.     

Incorporating neurophysiological concepts in mathematical thermoregulation models

Skin blood flow (SBF) is a key player in human thermoregulation during mild thermal challenges. Various numerical models of SBF regulation exist. However, none explicitly incorporates the neurophysiology of thermal reception. This study tested a new SBF model that is in line with experimental data on thermal reception and the neurophysiological pathways involved in thermoregulatory SBF control. Additionally, a numerical thermoregulation model was used as a platform to test the function of the neurophysiological SBF model for skin temperature simulation. The prediction-error of the SBF-model was quantified by root-mean-squared-residual (RMSR) between simulations and experimental measurement data. Measurement data consisted of SBF (abdomen, forearm, hand), core and skin temperature recordings of young males during three transient thermal challenges (1 development and 2 validation). Additionally, ThermoSEM, a thermoregulation model, was used to simulate body temperatures using the new neurophysiological SBF-model. The RMSR between simulated and measured mean skin temperature was used to validate the model. The neurophysiological model predicted SBF with an accuracy of RMSR?<?0.27. Tskin simulation results were within 0.37 °C of the measured mean skin temperature. This study shows that (1) thermal reception and neurophysiological pathways involved in thermoregulatory SBF control can be captured in a mathematical model, and (2) human thermoregulation models can be equipped with SBF control functions that are based on neurophysiology without loss of performance. The neurophysiological approach in modelling thermoregulation is favourable over engineering approaches because it is more in line with the underlying physiology.     

Regulate or tolerate: Thermal strategy of a coral reef flat resident,the epaulette shark,Hemiscyllium ocellatum

Highly variable thermal environments, such as coral reef flats, are challenging for marine ectotherms and are thought to invoke the use of behavioural strategies to avoid extreme temperatures and seek out thermal environments close to their preferred temperatures. Common to coral reef flats, the epaulette shark (Hemiscyllium ocellatum) possesses physiological adaptations to hypoxic and hypercapnic conditions, such as those experienced on reef flats, but little is known regarding the thermal strategies used by these sharks. We investigated whether H. ocellatum uses behavioural thermoregulation (i.e., movement to occupy thermally favourable microhabitats) or tolerates the broad range of temperatures experienced on the reef flat. Using an automated shuttlebox system, we determined the preferred temperature of H. ocellatum under controlled laboratory conditions and then compared this preferred temperature to 6 months of in situ environmental and body temperatures of individual H. ocellatum across the Heron Island reef flat. The preferred temperature of H. ocellatum under controlled conditions was 20.7 ± 1.5°C, but the body temperatures of individual H. ocellatum on the Heron Island reef flat mirrored environmental temperatures regardless of season or month. Despite substantial temporal variation in temperature on the Heron Island reef flat (15–34°C during 2017), there was a lack of spatial variation in temperature across the reef flat between sites or microhabitats. This limited spatial variation in temperature creates a low-quality thermal habitat limiting the ability of H. ocellatum to behaviourally thermoregulate. Behavioural thermoregulation is assumed in many shark species, but it appears that H. ocellatum may utilize other physiological strategies to cope with extreme temperature fluctuations on coral reef flats. While H. ocellatum appears to be able to tolerate acute exposure to temperatures well outside of their preferred temperature, it is unclear how this, and other, species will cope as temperatures continue to rise and approach their critical thermal limits. Understanding how species will respond to continued warming and the strategies they may use will be key to predicting future populations and assemblages.     

Thermal ecology and physiology of an elongate and semi-fossorial arthropod,the bark centipede

Organismal performance is strongly linked to temperature because of the fundamental thermal dependence of chemical reaction rates. However, the relationship between the environment and body temperature can be altered by morphology and ecology. In particular, body size and body shape can impact thermal inertia, as high surface area to volume ratios will possess low thermal mass. Habitat type can also influence thermal physiology by altering the opportunity for thermoregulation. We studied the thermal ecology and physiology of an elongate invertebrate, the bark centipede (Scolopocryptops sexspinosus). We characterized field body temperature and environmental temperature distributions, measured thermal tolerance limits, and constructed thermal performance curves for a population in southern Georgia. We found evidence that bark centipedes behaviorally thermoregulate, despite living in sheltered microhabitats, and that performance was maintained over a broad range of temperatures (over 20 °C). However, both the thermal optimum for performance and upper thermal tolerance were much higher than mean body temperature in the field. Together, these results suggest that centipedes can thermoregulate and maintain performance over a broad range of temperatures but are sensitive to extreme temperatures. More broadly, our results suggest that wide performance breadth could be an adaptation to thermal heterogeneity in space and time for a species with low thermal inertia.     

A study on body temperature regulation and residential thermal environments of the elderly—I. whole country survey of residential thermal environment and evaluation methods: RTE-index

1. 1. As part of “research on environmental comfort,” that is, research which aims to make people's living environment more comfortable, we attempted to clarify the relation between the living environment of elderly people and physiological and psychological factors. We carried out a comprehensive study with a view to establishing comfort standards for the residential thermal environment, and for creating evaluation and control systems.

2. 2. The comfort of the living environment is closely related to the thermal environment and the temperature-regulation ability of the human body. This ability of the body to regulate temperature develops during childhood and recedes as the adult ages.

3. 3. We therefore carried out: (a) experiments on body-temperature regulation and on special characteristics of the body-temperature regulation of elderly people, (b) a nationwide survey of the actual residential thermal environments of elderly people and (c) a survey of the daily activities of elderly people, and how elderly people's physiological and psychological conditions change when they are engaged in these activities.

4. 4. As a result of these experiments and surveys, we were able to: (d) formulate standards for evaluation of the residential thermal environment and (e) numerically express the results of systematic evaluation of residential thermal environments of elderly people, by means of a RTE-index.

An application of randomization for detecting evidence of thermoregulation in timber rattlesnakes (Crotalus horridus) from northwest Arkansas

Most reptiles maintain their body temperatures within normal functional ranges through behavioral thermoregulation. Under some circumstances, thermoregulation may be a time-consuming activity, and thermoregulatory needs may impose significant constraints on the activities of ectotherms. A necessary (but not sufficient) condition for demonstrating thermoregulation is a difference between observed body temperature distributions and available operative temperature distributions. We examined operative and body temperature distributions of the timber rattlesnake (Crotalus horridus) for evidence of thermoregulation. Specifically, we compared the distribution of available operative temperatures in the environment to snake body temperatures during August and September. Operative temperatures were measured using 48 physical models that were randomly deployed in the environment and connected to a Campbell CR-21X data logger. Body temperatures (n=1,803) were recorded from 12 radiotagged snakes using temperature-sensitive telemetry. Separate randomization tests were conducted for each hour of day within each month. Actual body temperature distributions differed significantly from operative temperature distributions at most time points considered. Thus, C. horridus exhibits a necessary (but not sufficient) condition for demonstrating thermoregulation. However, unlike some desert ectotherms, we found no compelling evidence for thermal constraints on surface activity. Randomization may prove to be a powerful technique for drawing inferences about thermoregulation without reliance on studies of laboratory thermal preference.     

Outdoor thermal comfort characteristics in the hot and humid region from a gender perspective

Thermal comfort is a subjective psychological perception of people based also on physiological thermoregulation mechanisms when the human body is exposed to a combination of various environmental factors including air temperature, air humidity, wind speed, and radiation conditions. Due to the importance of gender in the issue of outdoor thermal comfort, this study compared and examined the thermal comfort-related differences between male and female subjects using previous data from Taiwanese questionnaire survey. Compared with males, the results indicated that females in Taiwan are less tolerant to hot conditions and intensely protect themselves from sun exposure. Our analytical results are inconsistent with the findings of previous physiological studies concerning thermal comfort indicating that females have superior thermal physiological tolerance than males. On the contrary, our findings can be interpreted on psychological level. Environmental behavioral learning theory was adopted in this study to elucidate this observed contradiction between the autonomic thermal physiological and psychological–behavioral aspects. Women might desire for a light skin tone through social learning processes, such as observation and education, which is subsequently reflected in their psychological perceptions (fears of heat and sun exposure) and behavioral adjustments (carrying umbrellas or searching for shade). Hence, these unique psychological and behavioral phenomena cannot be directly explained by autonomic physiological thermoregulation mechanisms. The findings of this study serve as a reference for designing spaces that accommodates gender-specific thermal comfort characteristics. Recommendations include providing additional suitable sheltered areas in open areas, such as city squares and parks, to satisfy the thermal comfort needs of females.     

Responses to Rapid Temperature Change in Vertebrate Ectotherms

Vertebrate ectotherms often encounter rapid, large scale changesin body temperature. In this paper, I discuss the direct effectsof changing body temperature on physiological parameters, aswell as corrective responses initiated by the animal. For manybiological functions, mean body temperature provides a usefulmeasure of the thermal effects produced by an altered environmentaltemperature. Under most conditions, the fins and body surfaceof fish are more important avenues of heat exchange than thegills. The local thermal sensitivity of peripheral blood vesselsresults in vasomotor adjustments which can alter thermal conductivity.Acid-base balance is challenged by changes in body temperature.Shifts in body temperature also alter metabolic demands, enzymeconformation, ionic and osmotic relationships, spontaneous activitylevels and nervous system function. Compensatory mechanismsinclude behavioral thermoregulation, by which animals seek toavoid stressful thermal environments, and autonomic restorativeresponses such as high temperature panting in reptiles. Waterbreathers may initiate anticipatory responses to minimize arterialoxygen fluctuations during termperature change. The organizationof the central neuronal network underlying the above regulatoryresponses is unclear. Both air and water breathers are ableto initiate compensatory acid-base responses, but the strategiesutilized by the two groups are quite different. Altered bodytemperature initiates long-term acclimation responses, and ifrapid, can also trigger stress responses.     

Drivers of Daily Routines in an Ectothermic Marine Predator: Hunt Warm,Rest Warmer?

Animal daily routines represent a compromise between maximizing foraging success and optimizing physiological performance, while minimizing the risk of predation. For ectothermic predators, ambient temperature may also influence daily routines through its effects on physiological performance. Temperatures can fluctuate significantly over the diel cycle and ectotherms may synchronize behaviour to match thermal regimes in order to optimize fitness. We used bio-logging to quantify activity and body temperature of blacktip reef sharks (Carcharhinus melanopterus) at a tropical atoll. Behavioural observations were used to concurrently measure bite rates in herbivorous reef fishes, as an index of activity for potential diurnal prey. Sharks showed early evening peaks in activity, particularly during ebbing high tides, while body temperatures peaked several hours prior to the period of maximal activity. Herbivores also displayed peaks in activity several hours earlier than the peaks in shark activity. Sharks appeared to be least active while their body temperatures were highest and most active while temperatures were cooling, although we hypothesize that due to thermal inertia they were still warmer than their smaller prey during this period. Sharks may be most active during early evening periods as they have a sensory advantage under low light conditions and/or a thermal advantage over cooler prey. Sharks swam into shallow water during daytime low tide periods potentially to warm up and increase rates of digestion before the nocturnal activity period, which may be a strategy to maximize ingestion rates. “Hunt warm, rest warmer” may help explain the early evening activity seen in other ectothermic predators.     

The ecological and evolutionary aspects of thermoregulation behavior on fish

The ecological and evolutionary aspects of the thermoregulation behavior in fish are reviewed. The values of the Final Preference Temperature (“Final Preferendum” or FPT) and their relations with the temperature of species origin are analyzed. The analogy in the thermal niche of the poikilotherm animals and of the FPT zone is shown. The similarity and difference in thermal acclimation and thermoregulation behavior are considered. The seasonal and age-related peculiarities of fish thermoregulation behavior and of its manifestations upon impact of a variety of external factors (food, parasites, etc.) are analyzed. The characteristic is given for relation of thermoregulation behavior with schooling, territorial, and feeding behaviors, as well as a species range. The FPT values for some fish families are given.     

Modern theoretical and practical problems of homoiothermia and thermoregulation

The adjuncts to the existing determinations of homoiothermia are made on the basis of new data, the principles of temperature adaptation of humans and homoiothermal animals are presented. The main purposes of the thermoregulation system of homoiothermal animals and humans in various temperature excesses are formulated. The arguments are advanced in favor of the fact that in the thermoneutral zone the thermoregulation system goes from the principles of regulating the temperature homeostasis by one or several temperature points of a body to the regulation by the fluctuations of the total heat content of an organism, which increases the sensitivity and the accuracy of the thermoregulation system operation. The physiological mechanisms are described of determining (measuring) the total heat content of a body.     

Components and mechanisms of thermal hyperpnea.

The pattern of breathing during a hyperthermia-induced hyperventilation varies across different species. Thermal tachypnea is a first phase panting response adopted during hyperthermia when tidal volume is minimized and the frequency of breathing is maximized. Blood-gas tensions and pH are maintained during this hyperventilation, and the associated heat loss helps the animal regulate its body temperature. A second pattern of breathing adopted in hyperthermia is thermal hyperpnea; this response is the focus of this review. This form of hyperventilation is evident after an increase in core temperature and it is apparent in humans. Increases of tidal volume as well as frequency of breathing are evident during this response that results in a respiratory alkalosis. The cause of thermal hyperpnea is not resolved; evidence of the potential mechanisms underlying this response support that modulators of the response act in either a multiplicative or additive manner with body temperatures. The details of the designs and methodologies of the studies supporting or refuting these two views are discussed. A physiological rationale for thermal hyperpnea is presented in which it is suggested this response serves a heat-loss role and contributes to selective brain cooling in hyperthermic humans. Ongoing research in this area is focused on resolving the mechanisms underlying thermal hyperpnea and its contribution to cranial thermoregulation. The direct application of this research is for the care of febrile and hyperthermic patients.     

Body temperature null distributions in reptiles with nonzero heat capacity: seasonal thermoregulation in the American alligator (Alligator mississippiensis)

Regulation of body temperature may increase fitness of animals by ensuring that biochemical and physiological processes proceed at an optimal rate. The validity of current methods of testing whether or not thermoregulation in reptiles occurs is often limited to very small species that have near zero heat capacity. The aim of this study was to develop a method that allows estimation of body temperature null distributions of large reptiles and to investigate seasonal thermoregulation in the American alligator (Alligator mississippiensis). Continuous body temperature records of wild alligators were obtained from implanted dataloggers in winter (n=7, mass range: 1.6-53.6 kg) and summer (n=7, mass range: 1.9-54.5 kg). Body temperature null distributions were calculated by randomising behavioural postures, thereby randomly altering relative animal surface areas exposed to different avenues of heat transfer. Core body temperatures were predicted by calculations of transient heat transfer by conduction and blood flow. Alligator body temperatures follow regular oscillations during the day. Occasionally, body temperature steadied during the day to fall within a relatively narrow range. Rather than indicating shuttling thermoregulation, however, this pattern could be predicted from random movements. Average daily body temperature increases with body mass in winter but not in summer. Daily amplitudes of body temperature decrease with increasing body mass in summer but not in winter. These patterns result from differential exposure to heat transfer mechanisms at different seasons. In summer, alligators are significantly cooler than predictions for a randomly moving animal, and the reverse is the case in winter. Theoretical predictions show, however, that alligators can be warmer in winter if they maximised their sun exposure. We concluded that alligators may not rely exclusively on regulation of body temperature but that they may also acclimatise biochemically to seasonally changing environmental conditions.