The dive response redefined: underwater behavior influences cardiac variability in freely diving dolphins

A hallmark of the dive response, bradycardia, promotes the conservation of onboard oxygen stores and enables marine mammals to submerge for prolonged periods. A paradox exists when marine mammals are foraging underwater because activity should promote an elevation in heart rate (f(H)) to support increased metabolic demands. To assess the effect of the interaction between the diving response and underwater activity on f(H), we integrated interbeat f(H) with behavioral observations of adult bottlenose dolphins diving and swimming along the coast of the Bahamas. As expected for the dive response, f(H) while resting during submergence (40±6 beats min(-1)) was significantly lower than f(H) while resting at the water surface (105±8 beats min(-1)). The maximum recorded f(H) (f(H,max)) was 128±7 beats min(-1), and occurred during post-dive surface intervals. During submergence, the level of bradycardia was modified by activity. Behaviors such as simple head bobbing at depth increased f(H) by 40% from submerged resting levels. Higher heart rates were observed for horizontal swimming at depth. Indeed, the dolphins operated at 37-58% of their f(H,max) while active at depth and approached 57-79% of their f(H,max) during anticipatory tachycardia as the animals glided to the surface. f(H) was significantly correlated with stroke frequency (range=0-2.5 strokes s(-1), r=0.88, N=25 dives) and calculated swim speed (range=0-5.4 m s(-1), r=0.88, N=25 dives). We find that rather than a static reflex, the dive response is modulated by behavior and exercise in a predictable manner.