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Respiration and heat exchange in relation to brain temperature (Th) and body temperature (Tb) were investigated in four heat stressed camels subjected daily to high temperature (47 degrees C) in a climate chamber while resting when hydrated and dehydrated by approximately 10%, 15% and 20% of initial weight. Diurnally Tb followed patterns described previously. Th was usually 0.2-0.5 degrees C greater than Tb: occasional reversals with brain cooling were observed. Minute ventilation increased with Tb: above 37.5 degrees C it was approximately half as much in dehydrated as in hydrated animals. Respiratory frequency increased with Tb up to 60/min. Tidal volume fell with increasing frequency; above 25 breaths/min, tidal volume approximated dead space volume. Exhaled air was almost always unsaturated with no systematic effect of dehydration. Metabolic rate fell on dehydration reducing ventilatory demand. Th and Tb were measured in two of the animals walking outdoors: then Th fell below Tb if exercise exceeded 30 min. The data indicate that heat stressed camels pant, but turbinate vasoconstriction in a hot environment prevents cooling of the brain by carotid rete heat exchange.

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The avian lung has been considered to be rigid and to remain isovolumetric during the respiratory cycle. We tested this hypothesis by implanting radiopaque markers of tantalum on the dorsal pulmonary surfaces and ventral pulmonary aponeuroses of Pekin ducks (Anas platyrhynchos) and measuring changes in lung thickness during the respiratory cycle using high speed cineradiography. We found small but regular changes in lung thickness that were synchronous with respiratory phase. Lung thickness was greatest at mid-inspiration (0.6% greater than mean) and least at mid-expiration (0.8% less than mean). Measurements made on ostrich (Struthio camelus) respiratory structures suggest that the maximal force that could be generated by the muscles (Mm. costopulmonales) at the margins of the ventral pulmonary aponeurosis is more than two orders of magnitude greater than would be required to resist pressure-induced changes in lung volume during respiration at rest. The action of these muscles could account for the very small magnitude of the volume changes measured during the respiratory cycle.

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The purpose of this study was to determine the effect of heavy thermal panting on arterial oxygen (PaO2) and carbon dioxide (PaCO2) tension in emus. The birds showed no significant change in body temperature during a 3-4 h heat stress caused by increasing ambient air temperature from 21 to 46 degrees C. However, the emus increased their respiratory frequency 10-fold (from 5.3 to 52.9 breaths X min-1). The high respiratory frequency resulted in a slight but significant decrease in PaCO2 (from 33.5 to 29.8 mm Hg), coupled with a slight increase in pH (from 7.449 to 7.469). Paradoxically, these changes were accompanied by a significant decrease in the arterial oxygen tension (from 99.7 to 84.6 mm Hg). The arterial hypoxia suggests hypoventilation while the hypocapnia suggests hyperventilation of the lungs. This could result from various spatial and/or temporal changes in ventilation/perfusion ratios.

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The complex pattern of air flow in the respiratory system of birds suggests that certain sites function as valves. To examine the possibility of mechanical valving, rather than aerodynamic valving, we recorded radiographic images of the orifices where the medioventral secondary bronchi branch from the primary bronchus in resting Pekin ducks. Analysis of the images indicated that the orifices do not change size or shape during the respiratory cycle, suggesting that they function as aerodynamic rather than mechanical valves in directing air flow through the lung.

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Rates of oxygen consumption and respiratory water loss were studied in camels that were exposed to desert heat and water deprivation. We found that changes in body temperature are accompanied by considerable changes in respiratory water loss. Body temperature fluctuations are greatest in dehydrated camels (up to 7 degrees C), and in these the respiratory water loss might vary from abut 0.06 to 1.2 g/min. The respiratory frequency varied from about 4 to 28 min-1, while the metabolic rate varied less than twofold. The lowest values for respiratory water loss can be explained by the exhalation of air at temperatures far below body temperature, and, in addition, removal of water vapour from the exhaled air, resulting in exhalation of air at less than 100% relative humidity.

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We have found that camels can reduce the water loss due to evaporation from the respiratory tract in two ways: (1) by decreasing the temperature of the exhaled air and (2) by removal of water vapour from this air, resulting in the exhalation of air at less than 100% relative humidity (r.h.). Camels were kept under desert conditions and deprived of drinking water. In the daytime the exhaled air was at or near body core temperature, while in the cooler night exhaled air wat at or near ambient air temperature. In the daytime the exhaled air was fully saturated, but at night its humidity might fall to approximately 75% r.h. The combination of cooling and desaturation can provide a saving of water of 60% relative to exhalation of saturated air at body temperature. The mechanism responsible for cooling of the exhaled air is a simple heat exchange between the respiratory air and the surfaces of the nasal passageways. On inhalation these surfaces are cooled by the air passing over them, and on exhalation heat from the exhaled air is given off to these cooler surfaces. The mechanism responsible for desaturation of the air appears to depend on the hygroscopic properties of the nasal surfaces when the camel is dehydrated. The surfaces give off water vapour during inhalation and take up water from the respiratory air during exhalation. We have used a simple mechanical model to demonstrate the effectiveness of this mechanism.

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The respiratory air of the giraffe is exhaled at temperatures substantially below body core temperature. As a consequence, the water content of the exhaled air is reduced to levels below that in pulmonary air, resulting in substantial reductions in respiratory water loss. Measurements under outdoor conditions showed that at an ambient air temperature of 24 degrees C, the exhaled air was 7 degrees C below body core temperature, and at ambient air temperature of 17 degrees C, the exhaled air was 13 degrees C below core temperature. The observations were extended to two additional species of wild and four species of domestic ungulates. All these animals exhaled air at temperatures below body core temperature. The average amount of water recovered due to cooling of the air during exhalation, calculated as per cent of the water loss that would occur if air were exhaled at body core temperature, amounted to between 24 and 58%, the average value for the giraffe being 56%.

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To clarify the problems of altitude tolerance in birds, we studied the combined effect of hypocapnia and hypoxia on cerebral blood flow (CBF) in ducks. CBF was measured by the xenon clearance method. Normocapnic hypoxia causes CBF to increase when the arterial O2 tension (PaO2) falls below 60--70 mmHg. Hypocapnic hypoxia significantly shifts the blood flow curve so that blood flow does not increase until a lower PaO2 (50--60 mmHg) is reached. This gives the appearance that hypocapnia suppresses the hypoxia-induced increase in CBF. However, due to the Bohr effect, the hypocapnic blood contains significantly more O2 than does the normocapnic blood at the same PaO2. Therefore, when CBF is expressed as a function of O2 content, rather than PO2, CBF in the hypocapnic group does not differ significantly from the CBF in the normocapnic group. We interpret this to mean that because of the significantly greater oxygen content of the hypocapnic blood at a given PaO2, the degree of hypoxia experienced by these brains is not as severe as that experienced by the normocapnic brains.

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The effect of hypoxia on cerebral blood flow in ducks was investigated by the rate at which arterially injected xenon-133 was cleared from the duck's brain. A two-component clearance curve resulted, which we have attributed to flow through the grey and white matter. Decreasing the arterial oxygen tension (PaO2) to 75 mmHg had no effect on cerebral blood flow. However, decreasing the PaO2 below 75 mmHg significantly increased blood flow to the fast-clearing compartment. The greatest increase in blood flow was seen when the arterial PO2 was below 50 mmHg. At an arterial PO2 of 30 mmHg, the cerebral blood flow to the fast-clearing compartment was increased more than 600% above the normoxic level. The magnitude of this increase is much greater in the duck than has been reported for mammals at roughly equivalent arterial oxygen tensions. The ability of avian cerebral blood flow to increase at moderate levels of hypoxia, plus the magnitude of the increase, may play a role in the exceptional tolerance of birds to hypoxia.

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Unacclimated pigeons were exposed to various simulated altitudes. Arterial and mixed venous blood gas levels were measured. At the highest altitude 9150m (Pb = 235 mm Hg) a remarkable degree of alkalosis was tolerated for several hours without ill effecrtant factor in the bird's ability to survive such altitudes.

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The purpose of this study was to determine the effect of arterial PCO2 on blood flow to the avian brain. Cerebral blood flow was measured on curarized, artificially ventilated Pekin ducks by the rate at which intra-arterially injected xenon-133 was cleared from the duck's brain. A two-component clearance curve resulted: the blood flow calculated from the fast and slow components was similar to the blood flow to mammalian grey and white matter, respectively. Hypercapnia markedly increased the fast component of blood flow, whereas hypocapnia had no effect on this component. These effects were not due to changes in blood pressure, which was independent of arterial PCO2. Blood flow calculated from the slow component was independent of arterial PCO2. We conclude that the lack of response to hypocapnia may contribute to the exceptional tolerance of birds to high altitude by maintaining normal cerebral blood flow.

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Domestic ducks were exposed to simulated altitudes of 0, 3000, 6000, and 9000 m in order to study the respiratory changes that take place. We found that the respiratory minute volume (VE,BTPS) increased with altitude, the increase being due to increased respiratory frequency while tidal volume (VT, BTPS) showed only minor changes. The quantity of air moved (VE, STPD), however, remained nearly unchanged with increasing altitude. The oxygen extraction, calculated as 1--(FIN2FEO2)/(FEN2FIO2), remained constant at about 0.28 up to 6000 m and declined to 0.17 at 9000 m. The fractional gas concentrations (FO2 and FCO2) in exhaled air and in the interclavicular and posterior thoracic air sacs changed only little up to 6000 m, but at 9000 m FO2 increased and FCO2 decreased. The relative constancy of expired and air sac gas up to 6000 m seems remarkable. However, when applied to current models of air flow in the avian respiratory system the results seem fully explainable and permit a detailed analysis of the functioning of the avian respiratory system.

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The energetic cost for walking is relatively higher for penguins than for other birds or for quadrupeds of similar body mass. The morphology of penguins seems to represent a compromise between aquatic and terrestrial locomotion wherein both energy economy and speed suffer when the birds move on land.

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