Avian thermoregulation in the heat: phylogenetic variation among avian orders in evaporative cooling capacity and heat tolerance

Little is known about the phylogenetic variation of avian evaporative cooling efficiency and heat tolerance in hot environments. We quantified thermoregulatory responses to high air temperature (Ta) in ~100-g representatives of three orders: African cuckoo (Cuculus gularis, Cuculiformes), lilac-breasted roller (Coracias caudatus, Coraciiformes), and Burchell’s starling (Lamprotornis australis, Passeriformes). All three species initiated respiratory mechanisms to increase evaporative heat dissipation when body temperature (Tb) approached 41.5°C in response to increasing Ta, with gular flutter observed in cuckoos and panting in rollers and starlings. Resting metabolic rate (RMR) and evaporative water loss (EWL) increased by quantitatively similar magnitudes in all three species, although maximum rates of EWL were proportionately lower in starlings. Evaporative cooling efficiency [defined as the ratio of evaporative heat loss (EHL) to metabolic heat production (MHP)] generally remained below 2.0 in cuckoos and starlings, but reached a maximum of ~3.5 in rollers. The high value for rollers reveals a very efficient evaporative cooling mechanism, and is similar to EHL/MHP maxima for similarly sized columbids which very effectively dissipate heat via cutaneous evaporation. This unexpected phylogenetic variation among the orders tested in the physiological mechanisms of heat dissipation is an important step toward determining the evolution of heat tolerance traits in desert birds.Summary statementWe show that avian evaporative cooling efficiency and heat tolerance display substantial taxonomic variation that are, unexpectedly, not systematically related to the use of panting versus gular flutter processes.

Thermoregulatory responses of sindhi and guzerat heifers under shade in a tropical environment

This study characterized the thermal environment and assessed the physiological aspects of acclimatization of Sindhi and Guzerat heifers in a tropical environment (Brazil) under shade. Eight Sindhi and eight Guzerat purebred heifers (Bos indicus) had their physiological traits measured twice a day (9:00 a.m. and 2:00 p.m.). Environmental data during the experimental period were collected at two-hour intervals between 5:00 a.m. and 5:00 p.m. The temperature-humidity (THI) and the black globe temperature-humidity (BGHI) indices were calculated, and surface temperature (St), respiratory rate (Rr), and rectal temperature (Rt) were collected, being used to estimate heat loss by cutaneous (Ec) and respiratory (Er) evaporation. In the warmer parts of the day (1:00 and 3:00 p.m.), the THI and BGHI reached values of 80.26 and 81.25, respectively. There was no significant difference in rectal temperatures between the breeds, but higher values were observed in the afternoon. Heat transfer by cutaneous evaporation reached 118.71±12.91 W.m-2 and 103.43±6.82 W.m-2 at 2:00 p.m. for the Sindhi and Guzerat heifers, respectively. Under these conditions (air temperature was between 29 and 30°C), 84% of the total latent heat loss in Sindhi and Guzerat heifers was represented by Ec. It can be concluded that Sindhi and Guzerat heifers can maintain homeothermy with minimal thermoregulatory effort under shade conditions in a tropical environment.

Evaporative cooling and cutaneous surface temperature of Holstein cows in tropical conditions

The effects of skin temperature (T S) on the rate of heat loss by cutaneous evaporation (E S) in Holstein cows chronically exposed to sun, considering hair coat colour were studied. Sixteen purebred cows were measured for E S and T S at 01:00 p.m. after 6 hours of exposure to sun, on three body regions (flank, neck and gluteus) and considering dark and white spots separately. Sweating rate (S) and E S were measured by means of a ventilated capsule. Black skin areas presented mean S (138.9 ± 8.5 gm-2 h-1), E S (93.3 ± 5.7 Wm-2), and T S (33.1 ± 0.2°C) higher than those in the white areas (109.5 ± 9.7 gm-2h-1), 73.6 ± 6.5 Wm-2 and 32.6 ± 0.2°C, respectively). There is an exponential relationship among cutaneous temperature and cutaneous evaporation, which can be represented by the equation: E S = 31.5 + exp{(T S - 27.9)/2.19115}, with coefficient of determination r² = 0.68. Cutaneous evaporative heat loss remains almost constant around 48 Wm-2 until T S reaches nearly 31°C.

Latent heat loss of Holstein cows in a tropical environment: a prediction model

Nine lactating Holstein cows with average 526 ± 5 kg of BW, five predominantly black and four predominantly white, bred in a tropical region and managed in open pasture were observed to measure cutaneous and respiratory evaporation rates under different environmental conditions. Cows were separated in three weight class: 1 (<450 kg), 2 (450-500 kg) and 3 (>500 kg). Latent heat loss from cutaneous surface was measured using a ventilated capsule; evaporation in the respiratory system was measured using a facial mask. The results showed that heaviest cows (2 and 3 classes) presented the least evaporation rates. When air temperature increased from 10 to 36ºC the relative humidity decreased from 90 to 30%. In these conditions the heat loss by respiratory evaporation increased from 5 to 57 Wm-2, while the heat loss by cutaneous evaporation increased from 30 to 350 Wm-2. The results confirm that latent heat loss was the main way of thermal energy elimination under high air temperatures (>30ºC); cutaneous evaporation was the main mechanism of heat loss, responding for about 85% of the heat loss. A model was presented for the prediction of the latent heat loss that was based on physiological and environmental variables and could be used to estimate the contribution of evaporation to thermoregulation; a second, based on air temperature only, should be used to make a simple characterization of the evaporation process.

Prevalence of cutaneous evaporation in Merriam's kangaroo rat and its adaptive variation at the subspecific level

Previous estimates suggested that ventilatory evaporation constitutes the major source of water loss in kangaroo rats (Dipodomys spp.). We quantified rates of water loss in Merriam's kangaroo rat (Dipodomys merriami) and demonstrate the degree to which acclimation to a particular thermal and hydric environment plays a role in the intraspecific variation in water loss evident in this species. We draw the following conclusions: (1) that water loss varies intraspecifically in Merriam's kangaroo rat, in association with habitats of contrasting aridity and temperature; (2) that animals from more xeric locations have lower water loss rates than those from more mesic sites; (3) that most water loss is cutaneous, with ventilatory evaporative water loss contributing, at most, only 44% to total evaporative water loss; and (4) that intraspecific differences in rates of water loss are not acclimatory, but fixed. After acclimating under the same conditions, xeric-site animals still show a 33% lower rate of evaporative water loss than mesic-site animals.

Inhibiting ventilatory evaporation produces an adaptive increase in cutaneous evaporation in mourning doves Zenaida macroura

We tested the hypothesis that birds can rapidly change the conductance of water vapor at the skin surface in response to a changing need for evaporative heat loss. Mourning doves (Zenaida macroura) were placed in a two-compartment chamber separating the head from the rest of the body. The rate of cutaneous evaporation was measured in response to dry ventilatory inflow at three ambient temperatures and in response to vapor-saturated ventilatory inflow at two ambient temperatures. At 35 degrees C, cutaneous evaporation increased by 72 % when evaporative water loss from the mouth was prevented, but no increase was observed at 45 degrees C. For both dry and vapor-saturated treatments, cutaneous evaporation increased significantly with increased ambient temperature. Changes in skin temperature made only a minor contribution to any observed increase in cutaneous evaporation. This indicates that Z. macroura can effect rapid adjustment of evaporative conductance at the skin in response to acute change in thermoregulatory demand.

Heat stress induces ultrastructural changes in cutaneous capillary wall of heat-acclimated rock pigeon

In heat-acclimated rock pigeons, cutaneous water evaporation is the major cooling mechanism when exposed at rest to an extremely hot environment of 50–60°C. This evaporative pathway is also activated in room temperature by a β-adrenergic antagonist (propranolol) or an α-adrenergic agonist (clonidine) and inhibited by a β-adrenergic agonist (isoproterenol). In contrast, neither heat exposure nor drug administration activates cutaneous evaporation in cold-acclimated pigeons. To elucidate the mechanisms underlying this phenomenon, we studied the role of the ultrastructure and permeability of the cutaneous vasculature. During both heat stress and the administration of propranolol and clonidine, we observed increased capillary fenestration and endothelial gaps. Similarly, propranolol increased the extravasation of Evans blue-labeled albumin in the skin tissue. We concluded that heat acclimation reinforces a mechanism by which the activation of adrenergic signal transduction pathways alters microvessel permeability during heat stress. Consequently the flux of plasma proteins and water into the interstitial space is accelerated, providing an interstitial source of water for sustained cutaneous evaporative cooling.