The evolution of accessory air-breathing structures is typically associated with reduction of the gills, although branchial ion transport remains pivotal for acid-base and ion regulation. Therefore, airbreathing fishes are believed to have a low capacity for extracellular pH regulation during a respiratory acidosis. In the present study, we investigated acid-base regulation during hypercapnia in the airbreathing fish Pangasianodon hypophthalmus in normoxic and hypoxic water at 28-30°C. Contrary to previous studies, we show that this air-breathing fish has a pronounced ability to regulate extracellular pH (pH e ) during hypercapnia, with complete metabolic compensation of pH e within 72 h of exposure to hypoxic hypercapnia with CO 2 levels above 34 mmHg. The high capacity for pH e regulation relies on a pronounced ability to increase levels of HCO 3 − in the plasma. Our study illustrates the diversity in the physiology of air-breathing fishes, such that generalizations across phylogenies may be difficult.
The aim of the present study was to determine the roles of externally versus internally oriented CO/H-sensitive chemoreceptors in promoting cardiorespiratory responses to environmental hypercarbia in the facultative air-breathing fish, Chitala ornata (the clown knifefish). Fish were exposed to environmental acidosis (pH ~ 6.0) or hypercarbia (≈ 30 torr PCO) that produced changes in water pH equal to the pH levels of the acidotic water to distinguish the relative roles of CO versus H. We also injected acetazolamide to elevate arterial levels of PCO and [H] in fish in normocarbic water to distinguish between internal and external stimuli. We measured changes in gill ventilation frequency, air breathing frequency, heart rate and arterial blood pressure in response to each treatment as well as the changes produced in arterial PCO and pH. Exposure to normocarbic water of pH 6.0 for 1 h did not produce significant changes in any measured variable. Exposure to hypercarbic water dramatically increased air breathing frequency, but had no effect on gill ventilation. Hypercarbia also produced a modest bradycardia and fall in arterial blood pressure. Injection of acetazolamide produced similar effects. Both hypercarbia and acetazolamide led to increases in arterial PCO and falls in arterial pH although the changes in arterial PCO/pH were more modest following acetazolamide injection as were the increases in air breathing frequency. The acetazolamide results suggest that the stimulation of air breathing was due, at least in part, to stimulation of internally oriented CO/H chemoreceptors monitoring blood gas changes.
Chitala ornata is a facultative air-breathing fish, which at low temperatures shows an arterial P CO2 (Pa CO2 ) level only slightly elevated above that of water breathers. By holding fish with indwelling catheters at temperatures from 25 to 36°C and measuring blood gasses, we show that this animal follows the ubiquitous poikilotherm pattern of reducing arterial pH with increasing temperature. Surprisingly, the temperature increase caused an elevation of Pa CO2 from 5 to 12 mmHg while the plasma bicarbonate concentration remained constant at around 8 mmol l −1 . The temperature increase also gave rise to a larger fractional increase in air breathing than in gill ventilation frequency. These findings suggest that air breathing, and hence the partitioning of gas exchange, is to some extent regulated by acid-base status in airbreathing fish and that these bimodal breathers will be increasingly likely to adopt respiratory pH control as temperature rises, providing an interesting avenue for future research.
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