Studies on extant bimodally breathing vertebrates offer us a chance to
gain insight into the changes in respiratory control during the evolu
tionary transition from water to air breathing. In primitive Actinopte
rygian air-breathing fishes (Lepisosteus and Amia), gill ventilation i
s driven by an endogenously active central rhythm generator that is po
werfully modulated by afferent input from internally and externally or
iented branchial chemoreceptors, as it is in water-breathing Actinopte
rygians. The effects of internal or external chemoreceptor stimulation
on water and air breathing vary substantially in these aquatic air br
eathers, suggesting that their roles are evolutionarily malleable. Air
breathing in these bimodal breathers usually occurs as single breaths
taken at irregular intervals and is an on-demand phenomenon activated
primarily by afferent input from the branchial chemoreceptors. There
is no evidence for central CO2/pH sensitive chemoreceptors and air-bre
athing organ mechanoreceptors have little influence over branchial- or
air-breathing patterns in Actinopterygian air breathers. In the Sarco
pterygian lungfish Lepidosiren and Protopterus, ventilation of the hig
hly reduced gills is relatively unresponsive to chemoreceptor or mecha
noreceptor input. The branchial chemoreceptors of the anterior arches
appear to monitor arterialized blood, while chemoreceptors in the post
erior arches may monitor venous blood. Lungfish respond vigorously to
hypercapnia, but it is not known whether these responses are mediated
by central or peripheral chemoreceptors. A major difference between th
e Sarcopterygian and Actinopterygian bimodal breathers is that lungfis
h can inflate their lungs using rhythmic bouts of air breathing, and l
ung mechanoreceptors influence the onset and termination of these lung
inflation cycles. The control of breathing in amphibians appears simi
lar to that of lungfish. Branchial ventilation may persist as rhythmic
buccal oscillations in most adults, and stimulation of peripheral che
moreceptors in the aortic arch or carotid labyrinths initiates short b
outs of breathing. Ventilation is much more responsive to hypercapnia
in adult amphibians than in Actinopterygian fishes because of central
CO2/pH sensitive chemoreceptors that act to convert periodic to more c
ontinuous breathing patterns when stimulated.