The dive response is well documented for marine mammals, and includes a significant reduction in heart rate ( f H ) during submersion as compared while breathing at the surface. In the current study we assessed the influence of the Respiratory Sinus Arrhythmia (RSA) while estimating the resting f H while breathing. Using transthoracic echocardiography we measured f H , and stroke volume (SV) during voluntary surface apneas at rest up to 255 s, and during recovery from apnea in 11 adult bottlenose dolphins ( Tursiops truncatus , 9 males and 2 females, body mass range: 140–235 kg). The dolphins exhibited a significant post-respiratory tachycardia and increased SV. Therefore, only data after this RSA had stabilized were used for analysis and comparison. The average (±s.d.) f H , SV, and cardiac output (CO) after spontaneous breaths while resting at the surface were 44 ± 6 beats min −1 , 179 ± 31 ml, and 7909 ± 1814 l min −1 , respectively. During the apnea the f H , SV, and CO decreased proportionally with the breath-hold duration, and after 255 s they, respectively, had decreased by an average of 18%, 1–21%, and 12–37%. During recovery, the f H , SV, and CO rapidly increased by as much as 117%, 34%, and 190%, respectively. Next, f H , SV and CO rapidly decreased to resting values between 90–110 s following the surface apnea. These data highlight the necessity to define how the resting f H is estimated at the surface, and separating it from the RSA associated with each breath to evaluate the significance of cardiorespiratory matching during diving.

Heart-rate ( f H ) changes during diving and exercise are well documented for marine mammals, but changes in stroke volume (SV) and cardiac output (CO) are much less known. We hypothesized that both SV and CO are also modified following intense exercise. Using transthoracic ultrasound Doppler at the level of the aortic valve, we compared blood flow velocities in the left ventricle and cardiac frequencies during rest and at 1, 3 and 4 min after a bout of exercise in 13 adult bottlenose dolphins (Tursiops truncatus, six male and seven female, body mass range 143-212 kg). Aortic cross-sectional area and ventricle blood velocity at the aortic valve were used to calculate SV, which together with f H provided estimates of left CO at rest and following exercise. f H and SV stabilized approximately 4-7 s following the post-respiratory tachycardia, so only data after the f H had stabilized were used for analysis and comparison. There were significant increases in f H , SV and CO associated with each breath. At rest, f H , SV and CO were uncorrelated with body mass, and averaged 41±9 beats min , respectively. One minute following high intensity exercise, the cardiac variables had increased by 104±43%, 63±11% and 234±84%, respectively. All variables remained significantly elevated in all animals for at least 4 min after the exercise. These baseline values provide the first data on SV and CO in awake and unrestrained cetaceans in water.

In the current study we used transthoracic echocardiography to measure stroke volume (SV), heart rate (fH) and cardiac output (CO) in adult bottlenose dolphins (Tursiops truncatus), a male beluga whale calf [Delphinapterus leucas, body mass (Mb) range: 151–175 kg] and an adult female false killer whale (Pseudorca crassidens, estimated Mb: 500–550 kg) housed in managed care. We also recorded continuous electrocardiogram (ECG) in the beluga whale, bottlenose dolphin, false killer whale, killer whale (Orcinus orca) and pilot whale (Globicephala macrorhynchus) to evaluate cardiorespiratory coupling while breathing spontaneously under voluntary control. The results show that cetaceans have a strong respiratory sinus arrythmia (RSA), during which both fH and SV vary within the interbreath interval, making average values dependent on the breathing frequency (fR). The RSA-corrected fH was lower for all cetaceans compared with that of similarly sized terrestrial mammals breathing continuously. As compared with terrestrial mammals, the RSA-corrected SV and CO were either lower or the same for the dolphin and false killer whale, while both were elevated in the beluga whale. When plotting fR against fH for an inactive mammal, cetaceans had a greater cardiac response to changes in fR as compared with terrestrial mammals. We propose that these data indicate an important coupling between respiration and cardiac function that enhances gas exchange, and that this RSA is important to maximize gas exchange during surface intervals, similar to that reported in the elephant seal.

Man-made environmental change may have significant impact on apex predators, like marine mammals. Thus, it is important to assess the physiological boundaries for survival in these species, and assess how climate change may affect foraging efficiency and the limits for survival. In the current study, we investigated whether the respiratory sinus arrhythmia (RSA) could estimate tidal volume ( V T ) in resting bottlenose dolphins ( Tursiops truncatus ). For this purpose, we measured respiratory flow and electrocardiogram (ECG) in five adult bottlenose dolphins at rest while breathing voluntarily. Initially, an exponential decay function, using three parameters (baseline heart rate, the change in heart rate following a breath, and an exponential decay constant) was used to describe the temporal change in instantaneous heart rate following a breath. The three descriptors, in addition to body mass, were used to develop a Generalized Additive Model (GAM) to predict the inspired tidal volume ( V Tinsp ). The GAM allowed us to predict V Tinsp with an average ( ± SD) overestimate of 3 ± 2%. A jackknife sensitivity analysis, where 4 of the five dolphins were used to fit the GAM and the 5th dolphin used to make predictions resulted in an average overestimate of 2 ± 10%. Future studies should be used to assess whether similar relationships exist in active animals, allowing V T to be studied in free-ranging animals provided that heart rate can be measured.

We measured respiratory flow (V̇), breathing frequency ( f R ), tidal volume (V T ), breath duration and end-expired O 2 content in bottlenose dolphins (Tursiops truncatus) before and after static surface breathholds ranging from 34 to 292 s. There was considerable variation in the end-expired O 2 , V T and f R following a breath-hold. The analysis suggests that the dolphins attempt to minimize recovery following a dive by altering V T and f R to rapidly replenish the O 2 stores. For the first breath following a surface breath-hold, the end-expired O 2 decreased with dive duration, while V T and f R increased. Throughout the recovery period, end-expired O 2 increased while the respiratory effort (V T , f R ) decreased. We propose that the dolphins alter respiratory effort following a breath-hold according to the reduction in end-expired O 2 levels, allowing almost complete recovery after 1.2 min.

No abstract