Amphibian and reptile adaptations to the environment: interplay between physiology and behavior

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This book provides a comprehensive and integrative view of the interplay between physiology and behavior in amphibians and reptiles, leading to a better understanding of the subject. The book covers topics that have recently been in the spotlight for scientific research on the physiology, behavior, and conservation of amphibians and reptiles. It brings together recent information from a range of disciplines that address critical topics for understanding their biology. As these studies are scattered across articles in specialized journals, this book provides a single and expanded source summarizing such advancements.

Amphibian and Reptile Adaptations to the Environment: Interplay Between Physiology and Behavior maintains a solid scientific basis for the biological topics covered. However, it presents the material in a clear and direct manner so that it is accessible even to non-biologists interested in the basic biology, behavior, and ecology of these animals as well as how these elements are connected to their conservation.

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We moved your item s to Saved for Later. There was a problem with saving your item s for later. You can go to cart and save for later there. Amphibian and Reptile Adaptations to the Environment - eBook. Average rating: 0 out of 5 stars, based on 0 reviews Write a review. Comparative physiologists are well aware of the joint effects of physical environmental conditions e. At the same time, ecologists emphasize that environmental constraints on water, temperature, and energy budget may vary nonindependently in time and space.

Ecological studies often reveal spatial covariations in, for example, air temperature, atmospheric moisture, and soil humidity that result from changes in landscape structure and microhabitats, such as the degree of shading or the vegetation type e. Concurrent changes in rainfall, water availability, and ambient humidity might accentuate or buffer the ecological consequences of climate warming depending on patterns of change in hydration state at the organism level Cahill et al.

This emphasizes the need to carefully investigate and account for dual changes in temperature and water availability in the environment. Body temperature influences the speed of enzymatic reactions and the structure of cellular membranes Angilletta, , and water is the solvent of biochemical reactions and a fluid for nutritional provisioning of cells Chaplin, In tandem, temperature and hydration conditions are therefore crucial for biochemical reactions and cell metabolism. Appropriate regulation of body temperature and water balance movement of water out and into the organism is therefore critical for organism performance and involves the modulation of a wide range of physiological and behavioral mechanisms through multiple hormone secretions Bradshaw, Recent studies have indeed emphasized that the vulnerability of animals to climate change may be the consequence of abnormally high costs resulting from the regulation of their body temperature thermoregulation, see Box 1 and of their water balance hydroregulation, see Box 2 and Cahill et al.

Body temperature has immediate effects on physiological performances, such as maximal locomotor capacities or energy assimilation, and on survival due to the existence of critical thermal limits. Spatial variation in operative temperatures depends on topography and habitat complexity, and temporal variation is often strong but predictable because of daily and seasonal cycles Paaijmans et al. Heat and water exchange rules in ectotherms and endotherms.

Body temperature and water balance are jointly influenced by heat and water exchanges within the organism and between the organism and its environment. These exchanges are modulated by i the biophysical and physiological properties of the organism and by ii behavioral strategies. Biophysical properties include morphology, surface properties skin ultrastructure, fur, feathers, etc.

For instance, skin color, thickness, and ultrastructure in reptiles and amphibians determine heating capacity and resistance to water loss. Heat and water exchanges can influence each other; for example, evaporative water loss EWL induces heat loss evaporative cooling whereas metabolic reactions are a source of heat and water, for example, through fat catabolism.

The regulation of body temperature and water balance involves neuronal integration and feedback reactions from the central nervous system that orchestrate hormone secretions involved in the maintenance of temperature and water balance homeostasis. Performance curves. Thermal performance curves are typically characterized by a linear to geometric growth with increasing body temperature until a maximum is reached at an optimal range of temperatures.

This is then followed by a rapid performance decrease at body temperatures above the optimum, meaning that a small increase above the optimum can be lethal Dowd et al. The hydric performance curves can be calculated from studies of dehydration and describe the relationship between an animal performance and hydration state. The common pattern is that performances are maximized when water is provided at libitum i. The optimal body temperature for performance remains the same whatever the hydration state.

Performance decreases when temperature and hydration state departs from their optimal values. Hydroregulation, defined as the set of behavioral and physiological mechanisms to control water balance and remain hydrated, is one component of osmoregulation, that is, the regulation of ionic concentration such as salts and minerals in body fluids in which water is the solvent. Water balance determines the hydration state of the organism, defined as the volumetric quantity of water or percentage of body water.

There are numerous markers of water balance depending on the model species such as direct measures of water content, or indirect measures, for example, measures of body mass changes or plasma osmolality. Hydroregulation only applies to semiterrestrial organisms living at the transition zone between aquatic and terrestrial environments, like amphibians, and terrestrial animals. Desiccation risks depend on water vapor pressure deficit and skin permeability, such that species differ tremendously in water loss rates through evaporation and therefore vulnerability to dehydration.

Species also differ importantly in tolerance to dehydration, and many organisms can go for extent period of time in places without permanent access to water e. Water constraints are not universal and are restricted to desiccating environments, such as terrestrial habitats and salt water, and thermoregulation is therefore virtually free from water limitation in freshwater or very humid environments. Desiccation risk depends on spatiotemporal patterns of air moisture, which is a nondepreciable resource that covary with environmental temperatures.

Studies of endotherms have focused on the consequences of this coupling for the thermal physiology of large mammals Mitchell et al. On the other hand, it has been suggested that ectotherms must adjust their body temperature within a preferred range that depends on species' resistance to water loss and water availability in the environment Angilletta, Comparative physiologists have shown that thermoregulation and hydroregulation influence each other through several pathways with strong interspecific variation between the two dominant modes of thermoregulation endothermy vs.

Here, we focus on three different hierarchical levels of functional integration between thermoregulation and hydroregulation relevant to ectotherms in general: cutaneous and respiratory heat and water exchanges, behavioral regulation of thermal and water balance, and the coupling between the mass and water budgets of ectotherms. Comparative studies have shown that thermal and water budgets are partly determined by biophysical properties of the body e.

Ectotherms have evolved diverse indirect means to control rates of water loss such as mucus secretion in anurans Lillywhite, , changes in the lipid barrier to water in nonavian reptiles and amphibians reviewed in Lillywhite, , or changes in cuticular properties in insects reviewed in Chown et al. On the other hand, water balance is variably influenced by respiratory water loss in terrestrial ectotherms. The high diversity of respiratory modes in ectotherms makes it hard to generalize patterns of respiratory water loss responses to changes in thermal and water conditions e.

Those may also apply in some ectotherm species Tattersall et al. For example, Pirtle et al. As a consequence, water deprivation can cause thermal depression i. This is true for some insects' species with unusually strong resistance to evaporative water loss Chown et al. Another set of mechanisms linking water budget and temperature regulation involves behavioral choices. On the one hand, in ectotherms, habitat humidity and individual water balance can influence daily and annual activity patterns, habitat selection or movement, leading to behavioral hydroregulation, whereby individuals flexibly adjust their behavior according to external and internal conditions to regulate their hydration state Guillon et al.

On the other hand, behavioral thermoregulation also involves changes in habitat choice decisions or activity patterns, and is the dominant mode of thermoregulation in ectotherms.

Amphibian and Reptile Adaptations to the Environment: Interplay Between Physiology and Behavior

Unfortunately, most behavioral studies of thermoregulation have not considered effects of nonenergetic mechanisms related to water balance. Yet, recent evidence suggests that water availability in the environment can modify the costs and benefits of behavioral thermoregulation in ectotherms such as basking, foraging, or resting Caillon et al.

Basking decisions apparently related to the regulation of temperature and the energy budget could prioritize water conservation or be highly constrained by water conditions under some circumstances e. The mass balance equation for water is mechanistically linked with those for nutrients and energy because food can provide both nutrients and water, energy catabolism produces water, and feces production combines both nutrient and water loss. The original model was designed to understand patterns of variation in thermoregulation primarily in lizards. We suggest that the cost—benefit model of thermoregulation can easily accommodate the functional integration of thermoregulation with hydroregulation.

In these species, modest to severe dehydration usually leads to a decrease in the maximal locomotor performance capacities as expected, but also a decrease in the optimal body temperature for performances as well as a reduction of the performance breadth and the thermal tolerance range.

One proximate explanation for nonadditive effects of water balance and body temperature on performance traits is that dehydration modifies the thermal sensitivity of cell and tissue metabolism as well as protection against thermal stress. Not only can the benefits of thermoregulation and hydroregulation be nonadditive, but the costs of hydroregulation must also interact with the costs of thermoregulation in ectotherms. Whereas body temperature fluctuates quickly in ectotherms, especially in the smallest species, some of the costs of hydroregulation are likely delayed because the water balance changes more gradually as a function of water intakes and losses.

Behavioral ecologists often envision three different kinds of costs opportunity, energy, and risks of thermoregulation. Unfortunately, we still know extremely little about the costs of behavioral hydroregulation in ectotherms and can only speculate on their interactions with the costs of thermoregulation. Increased basking effort allows ectotherms to reach faster their optimal temperature but can compromise their water balance on the long run, possibly reducing performance and thus growth and survival.

For example, in lizards, the net effect of an increased behavioral activity in full sun on water balance is negative and not offset by potential positive effects of thermoregulation on metabolic water production and dietary water intake Pirtle et al. Studies of movements of ectotherms between water sources in arid or semiarid environments provide an excellent opportunity to address behavioral constraints induced by thermal and water availability, as seen with research done in ungulates Cain et al. Areas near water sources could have more or less vegetation cover. This will affect the ability of individuals to find both basking spots, shade, and water easily.

Individuals do not necessarily have to endure environmental conditions in their local habitat, but they can also disperse to more suitable environmental conditions. Global climate change could reduce complementation at the landscape scale when local warming is associated with higher drought frequencies and increase it when local warming is associated with higher rainfall, such as in temperate areas Dore, Unfortunately, empirical studies of climate niche shifts and dispersal plasticity of ectotherms in response to joint needs for temperature and water are exceedingly rare.

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Thus, whether rising temperature will increase or reduce dispersal is not easy to predict in this species given a regional context of drier climate conditions in the coming years. For example, parental care to the eggs and embryos has emerged repeatedly in invertebrates and vertebrates, and it is often stated that benefits of parental care are primarily derived from enhanced regulation of thermal conditions in ectotherms Farmer, ; Shine, Therefore, the need of jointly regulating thermal and water balance during periods of high parental investment may influence the evolution of parental behaviors and brooding in ectotherms.

This situation may apply in other viviparous species of ectotherms as well. We suggest that incorporating water requirements and interactions with parental thermoregulation is a crucial facet that has been neglected so far. Guidelines for future studies. With this paper, we aim at proposing guidelines for future studies trying to understand the responses of ectotherms to changes in their temperature and water environment. The first task would be to better disentangle the environmental temperature and water implications on microclimate properties.

Peter Funch

In the environment, a wet habitat is also often cooler than a dry one. Future studies should pay attention to be able to have all combinations of water and thermal environments or focus on other variables such as the biophysics of the microhabitat evaporative water losses, operative temperature. A second task is to better understand the functional responses of ectotherms to these habitats. We should now ask what mechanisms are common to thermoregulation and hydroregulation, and also investigate their plasticity and flexibility that could be critical in understanding organism responses to global changes.

Eventually, such studies should try to disentangle the physiological and ecological effects of thermal and hydric conditions including additive and interactive effects of multiple environmental factors. In particular, we need more investigations of the influence of environmental temperatures on water balance and of the influence of hydric conditions on the thermal biology. At least three different solutions are available. First, empirical studies should include detailed descriptions of both thermal and hydric conditions, including potential evaporative water loss rates and water availability in the environment.

For example, biophysical models currently make it possible to produce maps of operative temperature and evaporative water loss for ectotherms across a wide range of spatial scales Bartelt et al. Second, comparisons across environmental gradients in target species should focus on independent geographic variation in water and thermal conditions to reduce colinearity between these environmental factors.

This is likely to be feasible at a regional scale by combining climate gradients e. Third, one can also rely on classical factorial experimental design in the field e. To elucidate the joint physiological and behavioral mechanisms of body temperature and water balance in ectotherms, we need a stronger emphasis on hydroregulation strategies than in current climate change research Sinclair et al. For example, such models applied to terrestrial lizards predict that changes in the skin properties and behavioral tactics are a much more important contribution for hydroregulation in response to changes in water availability than metabolic changes Pirtle et al.

In addition, bioenergetic models require a good understanding of all water balance mechanisms including dietary and metabolic water inputs. There is indeed great scope to improve our understanding of the behavioral responses of ectotherms to variation in water availability and hydric conditions relative to thermal conditions, and to disentangle temperature and water effects on behavior Kearney et al. This is particularly pressing because habitat selection mechanisms are of great importance to both temperature and water balance regulation Pintor et al.

Laboratory experiments with shuttle boxes or contrasted microhabitats Pintor et al. An explanation for this is that direct manipulations and quantitative measures of hydration state are more difficult to perform than those of thermal conditions and body temperature in ectotherms, in which thousands of thermal performance curves have been quantified Angilletta, ; Angilletta et al. The physiological and behavioral regulation of body temperature and water balance should be considered as an integrated functional property of terrestrial and semiterrestrial ectotherms.

The content and structure of the paper was conceived collectively by all authors. We thank the anonymous reviewers for their helpful comments that helped improving this manuscript. We thank Dale DeNardo for his critical advices in the elaboration of this article. Ecol Evol. National Center for Biotechnology Information , U. Journal List Ecol Evol v. Published online Aug 2. Author information Article notes Copyright and License information Disclaimer.

Corresponding author. Email: gro. Received May 15; Accepted Jun Abstract The regulation of body temperature thermoregulation and of water balance defined here as hydroregulation are key processes underlying ecological and evolutionary responses to climate fluctuations in wild animal populations. Keywords: behavioral decisions, body temperature, performance curves, physiological adjustments, water balance. Box 1 Thermoregulation. Open in a separate window. Figure 1. Figure 2. Box 2 Hydroregulation. If water is limiting, optimal body temperatures for heat exchanges and energy metabolism may not be reached Posture changes Change in the posture according to the daytime in frogs Pough et al.

Figure 3. Heat stress and dehydration in adapting for performance: Good, bad, both, or neither? Temperature , 3 , — Trading heat and hops for water: Dehydration effects on locomotor performance, thermal limits, and thermoregulatory behavior of a terrestrial toad.

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American Zoologist , 32 , — Ecological Modelling , , — Behavioural thermoregulation and the relative roles of convection and radiation in a basking butterfly. Journal of Thermal Biology , 41 , 65— The cost—benefit model of thermoregulation does not predict lizard thermoregulatory behavior. Ecology , 86 , — Vertebrate ecophysiology: An introduction to its principles and applications.

Environmental endocrinology. General and Comparative Endocrinology , , — Muscles provide an internal water reserve for reproduction. Temperature extremes: Geographic patterns, recent changes, and implications for organismal vulnerabilities. Global Change Biology , 22 , — How does climate change cause extinction? Warming decreases thermal heterogeneity of leaf surfaces: Implications for behavioural thermoregulation by arthropods. Functional Ecology , 28 , — III, Krausman P.

Mechanisms of thermoregulation and water balance in desert ungulates. Wildlife Society Bulletin , 34 , — Do we underestimate the importance of water in cell biology? Nature Reviews Molecular Cell Biology , 7 , — Dehydration: Physiology, assessment, and performance effects. Comprehensive Physiology , 4 , — Respiratory water loss in insects. Insect physiological ecology: Mechanisms and patterns.

Water loss in insects: An environmental change perspective. Journal of Insect Physiology , 57 , — Australian Journal of Zoology , 47 , Scaling of body temperature in mammals and birds. Functional Ecology , 22 , 58— Evolutionary shifts in habitat aridity predict evaporative water loss across squamate reptiles.

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