Efficient Evaporative Cooling and Pronounced Heat Tolerance in an Eagle-Owl, a Thick-Knee and a Sandgrouse

Avian evaporative cooling and the maintenance of body temperature (Tb) below lethal limits during heat exposure has received more attention in small species compared to larger-bodied taxa. Here, we examined thermoregulation at air temperatures (Tair) approaching and exceeding normothermic Tb in three larger birds that use gular flutter, thought to provide the basis for pronounced evaporative cooling capacity and heat tolerance. We quantified Tb, evaporative water loss (EWL) and resting metabolic rate (RMR) in the ∼170-g Namaqua sandgrouse (Pterocles namaqua), ∼430-g spotted thick-knee (Burhinus capensis) and ∼670-g spotted eagle-owl (Bubo africanus), using flow-through respirometry and a stepped Tair profile with very low chamber humidities. All three species tolerated Tair of 56–60°C before the onset of severe hyperthermia, with maximum Tb of 43.2°C, 44.3°C, and 44.2°C in sandgrouse, thick-knees and eagle-owls, respectively. Evaporative scope (i.e., maximum EWL/minimum thermoneutral EWL) was 7.4 in sandgrouse, 12.9 in thick-knees and 7.8 in eagle-owls. The relationship between RMR and Tair varied substantially among species: whereas thick-knees and eagle-owls showed clear upper critical limits of thermoneutrality above which RMR increased rapidly and linearly, sandgrouse did not. Maximum evaporative heat loss/metabolic heat production ranged from 2.8 (eagle-owls) to 5.5 (sandgrouse), the latter the highest avian value yet reported. Our data reveal some larger species with gular flutter possess pronounced evaporative cooling capacity and heat tolerance and, when taken together with published data, show thermoregulatory performance varies widely among species larger than 250 g. Our data for Namaqua sandgrouse reveal unexpectedly pronounced variation in the metabolic costs of evaporative cooling within the genus Pterocles.

Keeping Cool

Measurement of giraffe body temperature has shown that it is ~38.5oC but it can vary by ~5oC over the course of a day. Body heat is derived from fermentation of browse, other metabolic processes and radiant heat. Heat loss mechanisms partly depend on body surface area. Despite their unusual shape the body surface area of giraffes is similar to that in other equivalent body mass mammals: a shorter trunk is offset by a longer neck and legs. Heat loss by radiation is constant, by conduction rare and minimal. Their long, slender legs and neck are an advantage for convective and evaporative heat loss from the skin: heat transfer is inversely proportional to the square root of diameter. Evaporation from the respiratory system occurs through the nasal mucosa, the surface area of which in giraffes is large. Cooling of the nasal mucosa and blood follows and cool blood drains in to the jugular vein and contributes to whole body cooling and cooling of the blood supplying the brain by heat exchange in the carotid rete. Similar heat exchange may occur across the surface of the ossicones. Behavior changes when ambient temperature exceeds skin temperature. Giraffes re-orientate their bodies to minimize radiant heat gain and seek shade. A unique arrangement of blood vessels supplying blood to skin patches allows patches to act as thermal windows through which heat can be lost an arrangement enhanced by evaporation: sweat gland density in the skin of patches is greater than it is elsewhere.

Effects of environmental temperature and humidity on evaporative heat loss through firefighter suit materials made with semi-permeable and microporous moisture barriers

The goal of this research was to understand how firefighter protective suits perform in different operational environments. This study used a sweating guarded hotplate to examine the effect of environmental temperature (20–45°C) and relative humidity (25–85% RH) on evaporative heat loss through firefighter turnout materials. Four firefighter turnout composites containing three different bi-component (semi-permeable) and one microporous moisture barriers were selected. The results showed that the evaporative resistance of microporous moisture barrier systems was independent of environmental testing conditions. However, absorbed moisture strongly affected evaporative heat loss through semi-permeable moisture barriers coated with a layer of nonporous hydrophilic polymer. Moisture absorption in mild environment (20–25°C) tests, or when testing at high humidity (>85% RH), significantly increased water vapor transmission in semi-permeable turnout systems. It was also found that environmental conditions used in the total heat loss (THL) test (25°C and 65% RH) produced moisture condensation in bi-component barrier systems, making them appear more breathable than could be expected when worn in hotter environments. Regression models successfully qualified the relationships between moisture uptake levels in semi-permeable barrier systems and evaporative resistance and THL. These findings reveal the limitations in relying on THL, the heat strain index currently called for by the NFPA 1971 Standard for Structural Firefighter personal protective equipment, and supports the need to measure turnout evaporative resistance at 35°C (Ret), in addition to THL at 25°C.

Thermoregulation in desert birds: scaling and phylogenetic variation in heat tolerance and evaporative cooling

ABSTRACTEvaporative heat dissipation is a key aspect of avian thermoregulation in hot environments. We quantified variation in avian thermoregulatory performance at high air temperatures (Ta) using published data on body temperature (Tb), evaporative water loss (EWL) and resting metabolic rate (RMR) measured under standardized conditions of very low humidity in 56 arid-zone species. Maximum Tb during acute heat exposure varied from 42.5±1.3°C in caprimulgids to 44.5±0.5°C in passerines. Among passerines, both maximum Tb and the difference between maximum and normothermic Tb decreased significantly with body mass (Mb). Scaling exponents for minimum thermoneutral EWL and maximum EWL were 0.825 and 0.801, respectively, even though evaporative scope (ratio of maximum to minimum EWL) varied widely among species. Upper critical limits of thermoneutrality (Tuc) varied by >20°C and maximum RMR during acute heat exposure scaled to Mb0.75 in both the overall data set and among passerines. The slope of RMR at Ta>Tuc increased significantly with Mb but was substantially higher among passerines, which rely on panting, compared with columbids, in which cutaneous evaporation predominates. Our analysis supports recent arguments that interspecific within-taxon variation in heat tolerance is functionally linked to evaporative scope and maximum ratios of evaporative heat loss (EHL) to metabolic heat production (MHP). We provide predictive equations for most variables related to avian heat tolerance. Metabolic costs of heat dissipation pathways, rather than capacity to increase EWL above baseline levels, appear to represent the major constraint on the upper limits of avian heat tolerance.

Do endotherms have thermal performance curves?

ABSTRACTTemperature is an important environmental factor governing the ability of organisms to grow, survive and reproduce. Thermal performance curves (TPCs), with some caveats, are useful for charting the relationship between body temperature and some measure of performance in ectotherms, and provide a standardized set of characteristics for interspecific comparisons. Endotherms, however, have a more complicated relationship with environmental temperature, as endothermy leads to a decoupling of body temperature from external temperature through use of metabolic heat production, large changes in insulation and variable rates of evaporative heat loss. This has impeded our ability to model endothermic performance in relation to environmental temperature as well as to readily compare performance between species. In this Commentary, we compare the strengths and weaknesses of potential TPC analogues (including other useful proxies for linking performance to temperature) in endotherms and suggest several ways forward in the comparative ecophysiology of endotherms. Our goal is to provide a common language with which ecologists and physiologists can evaluate the effects of temperature on performance. Key directions for improving our understanding of endotherm thermoregulatory physiology include a comparative approach to the study of the level and precision of body temperature, measuring performance directly over a range of body temperatures and building comprehensive mechanistic models of endotherm responses to environmental temperatures. We believe the answer to the question posed in the title could be ‘yes’, but only if ‘performance’ is well defined and understood in relation to body temperature variation, and the costs and benefits of endothermy are specifically modelled.

How hornbills handle heat: sex-specific thermoregulation in the southern yellow-billed hornbill

ABSTRACTAt a global scale, thermal physiology is correlated with climatic variables such as temperature and aridity. There is also evidence that thermoregulatory traits vary with fine-scale microclimate, but this has received less attention in endotherms. Here, we test the hypothesis that avian thermoregulation varies with microclimate and behavioural constraints in a non-passerine bird. Male and female southern yellow-billed hornbills (Tockus leucomelas) experience markedly different microclimates while breeding, with the female sealing herself into a tree cavity and moulting all her flight feathers during the breeding attempt, becoming entirely reliant on the male for provisioning. We examined interactions between resting metabolic rate (RMR), evaporative water loss (EWL) and core body temperature (Tb) at air temperatures (Ta) between 30°C and 52°C in male and female hornbills, and quantified evaporative cooling efficiencies and heat tolerance limits. At thermoneutral Ta, neither RMR, EWL nor Tb differed between sexes. At Ta >40°C, however, RMR and EWL of females were significantly lower than those of males, by ∼13% and ∼17%, respectively, despite similar relationships between Tb and Ta, maximum ratio of evaporative heat loss to metabolic heat production and heat tolerance limits (∼50°C). These sex-specific differences in hornbill thermoregulation support the hypothesis that avian thermal physiology can vary within species in response to fine-scale microclimatic factors. In addition, Q10 for RMR varied substantially, with Q10 ≤2 in some individuals, supporting recent arguments that active metabolic suppression may be an underappreciated aspect of endotherm thermoregulation in the heat.

A review of intervention methods used to reduce flying-fox mortalities in heat stress events

Heat stress events in Australian flying-fox camps have resulted in significant numbers of flying-fox deaths. The frequency and intensity of such events have increased in recent decades, attributed to anthropogenic climate change. Evidence-based interventions are required to address this growing threat. Responders currently use different combinations of a range of intervention methods. We undertook a systematic review of heat stress interventions, which we classified as either ‘camp-scale’ or ‘individual-scale’. Camp-scale interventions included manual and automated misting of roost vegetation, whereas individual-scale interventions included spraying individual animals or removing them for intensive cooling and rehydration procedures. Our study showed that to date, evaluation of the efficacy of heat stress interventions has been largely anecdotal rather than empirical. This highlights the need for dedicated rigorous studies to evaluate the effectiveness of all the intervention methods described here. It will be especially important to understand the relationship between camp temperature and humidity levels and their influence on flying-foxes’ ability to regulate their body temperature, because high relative humidity reduces the ability of mammals to cool themselves using evaporative heat loss. The development of biophysiological measures such as temperature and humidity indices for different flying-fox species would enable meaningful interpretations of intervention trials under controlled conditions.

Phenotypic flexibility in heat production and heat loss in response to thermal and hydric acclimation in the zebra finch, a small arid-zone passerine

To maintain constant body temperature (Tb) over a wide range of ambient temperatures (Ta) endothermic animals require large amounts of energy and water. In hot environments, the main threat to endothermic homeotherms is insufficient water to supply that necessary for thermoregulation. We investigated flexible adjustment of traits related to thermoregulation and water conservation during acclimation to hot conditions or restricted water availability, or both, in the zebra finch, Taeniopygia guttata a small arid-zone passerine. Using indirect calorimetry, we measured changes in whole animal metabolic rate (MR), evaporative heat loss (EHL) and Tb before and after acclimation to 23 or 40 °C, with different availability of water. Additionally, we quantified changes in partitioning of EHL into respiratory and cutaneous avenues in birds exposed to 25 and 40 °C. In response to heat and water restriction zebra finches decreased MR, which together with unchanged EHL resulted in increased efficiency of evaporative heat loss. This facilitated more precise Tb regulation in heat-acclimated birds. Acclimation temperature and water availability had no effect on the partitioning of EHL into cutaneous or respiratory avenues. At 25 °C, cutaneous EHL accounted for ~ 60% of total EHL, while at 40 °C, its contribution decreased to ~ 20%. Consistent among-individual differences in MR and EHL suggest that these traits, provided that they are heritable, may be a subject to natural selection. We conclude that phenotypic flexibility in metabolic heat production associated with acclimation to hot, water-scarce conditions is crucial in response to changing environmental conditions, especially in the face of current and predicted climate change.