Dear All
My co-authors and I would like to share our recent publication measuring 
cardiovascular responses in the bottlenose dolphin during breath-holds at rest. 

In this paper we measure heart rate and stroke volume and calculate cardiac 
output. We correct the values for the cofounding changes in cardiac function 
associated with each breath, the Respiratory Sinus Arrhythmia, and conclude 
that the dive response is minimal. We propose that in the dolphin, the dive 
response (reduced heart rate and cardiac output) is an extension of the 
Respiratory Sinus Arrhythmia and a more generalized physiological response and 
not a specialized trait for diving.

We also provide results that suggest that dolphins can voluntarily alter heart 
rate and blood flow. This mechanism would allow them to adjust blood flow based 
on the dive they are planning, and adjust flow during the dive to alter changes 
due to metabolic demand. Voluntary adjustment of heart rate has been shown 
previously in other species, such as the harbor porpoise or sea lion, and a 
limited ability for such an adjustment have also been shown in human elite 
athletes. We propose that this ability to voluntary alter blood flow as needed 
during the dive is the key adaptation how they match demand and supply, and may 
explain how failure of this mechanism during stress can cause diving related 
issues such as the bends. 

While controversial, this new article questions the importance of the dive 
response that has for long been thought to be one of the main mechanisms for 
how diving animals can perform extended breath-holds, and argues that evolution 
has enabled dolphins and marine mammals to become efficient at voluntary 
adjustment of heart rate and blood flow to improve use of the available oxygen.

For anyone interested in this Open Access publication, the details and a link 
are below, or send me an email at [email protected] <mailto:[email protected]> 
for a pdf, or for questions of comments.

Sincerely,
Andreas

Title:Re-evaluating the significance of the dive response during voluntary 
surface apneas in the bottlenose dolphin, Tursiops truncatus
Authors:Fahlman, A.,Miedler, S., Rocho-Levine, J., Jabois, A., Arenarez, J., 
Marti-Bonmati, L., García-Párraga, D., Cauture, F.
Journal: Scientific Reports, 9, 
DOI: 10.1038/s41598-019-45064-8
URL: www.nature.com/articles/s41598-019-45064-8

Abstracts: The dive response is well documented for marine mammals, and 
includes a significant reduction in heart rate (fH) 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 fH while breathing. Using transthoracic echocardiography we measured 
fH, 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.) fH, 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 fH, 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 fH, SV, and CO rapidly increased by as much as 117%, 34%, 
and 190%, respectively. Next, fH, 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 fH is estimated at the surface, and 
separating it from the RSA associated with each breath to evaluate the 
significance of cardiorespiratory matching during diving.

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