Knowing how much energy an individual requires is fundamental to resolving a number of questions about marine mammals. These include determining how they have solved complex bioenergetic problems to successfully exploit their environment, and whether or not they compete with fisheries for prey (Markussen and Oritsland, 1991;Olesiuk, 1993;Rosen and Trites, 1997;Stenson et al., 1997;Boyd et al., 1999). Unfortunately, determining energy expenditure of free-ranging individuals is difficult. Measuring doubly labelled water (DLW) turnover rates and recording heart rate are two common techniques that have been used.The DLW method suffers from two major limitations. The first is that it provides only a mean estimate of metabolism over the entire period between blood samplings. The second limitation is that there is a finite time over which the measurement can be made, due to the biological half-life of the chemical agents. In large vertebrates, this period is usually 5-10 days, after which the animals must be recaptured for blood sampling. Many assumptions, estimates and logistics needed to use DLW also compromise the applicability of this technique (see Costa, 1987;Speakman, 1993).Recording heart rate is a technique for estimating energy expenditure that offers the possibility of monitoring metabolism for a year or longer, with a fine time resolution of hours or minutes that can be related to specific activities (Butler, 1993;Bevan et al., 1995b;Woakes et al., 1995;Andrews, 1998). When coupled with dive profiles from time-depth recorders, for example, heart rate may be used to estimate the energy expenditure of specific dives. Several comparative studies have shown that this technique is as robust as using the DLW method (Nolet et al., 1992;Bevan et al., 1994Bevan et al., , 1995aBoyd et al., 1995).The heart rate method is based on Fick's relationship of oxygen consumption (V . O∑) and heart rate (fH) (Fick, 1870):where VS is stroke volume of the heart, and CaO∑-Cv -O∑ is the difference in oxygen concentration between arterial and mixed venous blood, respectively (also referred to as tissue oxygen The validity of using heart rate to estimate energy expenditure in free-ranging Steller sea lions Eumetopias jubatus was investigated by establishing whether there is a relationship between heart rate (fH) and oxygen consumption rate (V . O∑) in captive sea lions while swimming and resting. Four trained Steller sea lions (2 males and 2 females; mass 87.4-194.4·kg; age 16 months-3 years) were each equipped with a datalogger and two dorsal subcutaneous electrodes to record electrocardiograms from which fH was calculated. V .
O∑(measured using open-circuit respirometry) was simultaneously recorded while the previously fasted animals were at rest within an enclosed dry metabolic chamber or while they swam in an enclosed swim mill against water currents of various speeds (0-1.5·m·s -1 ). O∑ while fasting suggests that heart rate can potentially be used to monitor energy consumption in free-ranging Steller sea lions. However, a shor...