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First published online March 28, 2008
Journal of Experimental Biology 211, 1169-1179 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.013235
Heart rate regulation and extreme bradycardia in diving emperor penguins
Center for Marine Biotechnology and Biomedicine, Scripps Institution of Oceanography, University of California San Diego, La Jolla, CA 92093-0204, USA
* Author for correspondence (e-mail: jmeir{at}ucsd.edu)
Accepted 3 February 2008
To investigate the diving heart rate (fH) response of
the emperor penguin (Aptenodytes forsteri), the consummate avian
diver, birds diving at an isolated dive hole in McMurdo Sound, Antarctica were
outfitted with digital electrocardiogram recorders, two-axis accelerometers
and time depth recorders (TDRs). In contrast to any other freely diving bird,
a true bradycardia (fH significantly
<fH at rest) occurred during diving [dive
fH (total beats/duration)=57±2 beats
min–1, fH at rest=73±2 beats
min–1 (mean ± s.e.m.)]. For dives less than the
aerobic dive limit (ADL; duration beyond which [blood lactate] increases above
resting levels), dive fH=85±3 beats
min–1, whereas fH in dives greater than
the ADL was significantly lower (41±1 beats min–1). In
dives greater than the ADL, fH reached extremely low
values: fH during the last 5 mins of an 18 min dive was 6
beats min–1. Dive fH and minimum
instantaneous fH during dives declined significantly with
increasing dive duration. Dive fH was independent of swim
stroke frequency. This suggests that progressive bradycardia and peripheral
vasoconstriction (including isolation of muscle) are primary determinants of
blood oxygen depletion in diving emperor penguins. Maximum instantaneous
surface interval fH in this study is the highest ever
recorded for emperor penguins (256 beats min–1), equivalent
to fH at
O2 max.,
presumably facilitating oxygen loading and post-dive metabolism. The classic
Scholander–Irving dive response in these emperor penguins contrasts with
the absence of true bradycardia in diving ducks, cormorants, and other penguin
species.
Key words: diving, electrocardiogram (ECG), emperor penguin, heart rate, respiratory sinus arrhythmia, stroke frequency