SUMMARYThe aim of the present study was to determine the roles that externally versus internally oriented CO 2 /H + -sensitive chemoreceptors might play in promoting cardiorespiratory responses to environmental hypercarbia in the air-breathing fish, Hoplerythrinus unitaeniatus (jeju). Fish were exposed to graded hypercarbia (1, 2.5, 5, 10 and 20% CO 2 ) and also to graded levels of environmental acidosis (pH ~7.0, 6.0, 5.8, 5.6, 5.3 and 4.7) equal to the pH levels of the hypercarbic water to distinguish the relative roles of CO 2 versus H + . We also injected boluses of CO 2 -equilibrated solutions (5, 10 and 20% CO 2 ) and acid solutions equilibrated to the same pH as the CO 2 boluses into the caudal vein (internal) and buccal cavity (external) to distinguish between internal and external stimuli. The putative location of the chemoreceptors was determined by bilateral denervation of branches of cranial nerves IX (glossopharyngeal) and X (vagus) to the gills. The data indicate that the chemoreceptors eliciting bradycardia, hypertension and gill ventilatory responses (increased frequency and amplitude) to hypercarbia are exclusively branchial, externally oriented and respond specifically to changes in CO 2 and not H + . Those involved in producing the cardiovascular responses appeared to be distributed across all gill arches while those involved in the gill ventilatory responses were located primarily on the first gill arch. Higher levels of aquatic CO 2 depressed gill ventilation and stimulated air breathing. The chemoreceptors involved in producing air breathing in response to hypercarbia also appeared to be branchial, distributed across all gill arches and responded specifically to changes in aquatic CO 2 . This would suggest that chemoreceptor groups with different orientations (blood versus water) are involved in eliciting air-breathing responses to hypercarbia in jeju.Key words: gills, cardiorespiratory control, hypercarbia, CO 2 and H + chemoreceptors, air breathing, jeju, Hoplerythrinus unitaeniatus.
THE JOURNAL OF EXPERIMENTAL BIOLOGY
2798and Baird, 1982; Graham, 1997;Sanchez et al., 2005). In other species, however, increases in environmental CO 2 are without effect on gill ventilation (Johansen, 1966;Todd, 1972;McMahon and Burggren, 1987) or air breathing (Johansen et al., 1968;Lomholt and Johansen, 1974). Just as with aquatic CO 2 , some air-breathing fish show no response to increasing levels of CO 2 in inspired air Perry et al., 2008) while others show an increase (Smith, 1930; Delaney et al., 1974; Delaney et al., 1976; Delaney et al., 1977; Babiker, 1979) or even a decrease (Jesse et al., 1967) in ventilation of the air-breathing organ (ABO).The equivocal nature of the data raises questions about the existence and role of internally oriented CO 2 /H + -sensitive chemoreceptors in driving gill ventilation in exclusively waterbreathing fish, as well as in driving gill ventilation or air breathing in facultative and obligate air-breathing fish. Clearly much remains to be done to resolve this issue...