Heart rate and energetics of free-ranging king penguins (Aptenodytes patagonicus).

The main objective of this study was to determine heart rate (fh) and the energetic costs of specific behaviours of king penguins while ashore and while foraging at sea during their breeding period. In particular, an estimate was made of the energetic cost of diving in order to determine the proportion of dives that may exceed the calculated aerobic dive limit (cADL; estimated usable O2 stores/estimated rate of oxygen consumption during diving). An implanted data logger enabled fh and diving behaviour to be monitored from 10 free-ranging king penguins during their breeding period. Using previously determined calibration equations, it was possible to estimate rate of oxygen consumption (VO2) when the birds were ashore and during various phases of their foraging trips. Diving behaviour showed a clear diurnal pattern, with a mixture of deep (>40 m), long (>3 min) and shallow (<40 m), short (<3 min) dives from dawn to dusk and shallow, short dives at night. Heart rate during dive bouts and dive cycles (dive + post-dive interval) was 42% greater than that when the birds were ashore. During diving, fh was similar to the 'ashore' value (87+/-4 beats min(-1)), but it did decline to 76% of the value recorded from king penguins resting in water. During the first hour after a diving bout, fh was significantly higher than the average value during diving (101+/-4 beats min(-1)) and for the remainder of the dive bout. Rates of oxygen consumption estimated from these (and other) values of fh indicate that when at sea, metabolic rate (MR) was 83% greater than that when the birds were ashore [3.15 W kg(-1) (-0.71, +0.93), where the values in parentheses are the computed standard errors of the estimate], while during diving bouts and dive cycles, it was 73% greater than the 'ashore' value. Although estimated MR during the total period between dive bouts was not significantly different from that during dive bouts [5.44 W kg(-1) (-0.30, +0.32)], MR during the first hour following a dive bout was 52% greater than that during a diving bout. It is suggested that this large increase following diving (foraging) activity is, at least in part, the result of rewarming the body, which occurs at the end of a diving bout. From the measured behaviour and estimated values of VO2, it was evident that approximately 35% of the dives were in excess of the cADL. Even if VO2 during diving was assumed to be the same as when the birds were resting on water, approximately 20% of dives would exceed the cADL. As VO2 during diving is, in fact, that estimated for a complete dive cycle, it is quite feasible that VO2 during diving itself is less than that measured for birds resting in water. It is suggested that the regional hypothermia that has been recorded in this species during diving bouts may be at least a contributing factor to such hypometabolism.

The heart rate/oxygen consumption relationship during cold exposure of the king penguin: a comparison with that during exercise.

This study investigated whether exposure to low ambient temperature could be used as an alternative to exercise for calibrating heart rate (fH) against rate of oxygen consumption ((O(2))) for subsequent use of fH to estimate (O(2)) in free-ranging animals. Using the relationship between the oxygen pulse (OP, the amount of oxygen used per heart beat) and an index of body condition (or nutritional index, NI), a relationship between fH and (O(2)) was established for resting king penguins exposed to a variety of environmental temperatures. Although there was a small but significant increase in the OP above and below the lower critical temperature (-4.9 degrees C), there was no difference in the relationship obtained between the OP and body condition (NI) obtained above or below the lower critical temperature. These results were then compared with those obtained in a previous study in which the relationship between fH and (O(2)) had been established for king penguins during steady-state exercise. The relationship between OP and NI in the present study was not significantly different from the relationship between resting OP and NI in the previous study. However, the relationship was different from that between active OP and NI. We conclude that, at least for king penguins, although thermoregulation does not affect the relationship between resting OP and NI, temperature cannot be used as an alternative to exercise for calibrating fH against (O(2)) for subsequent use of fH to estimate (O(2)) in free-ranging animals.

Heart rate as an indicator of oxygen consumption: influence of body condition in the king penguin.

The use of heart rate to estimate field metabolic rate has become a more widely used technique. However, this method also has some limitations, among which is the possible impact that several variables such as sex, body condition (i.e. body fat stores) and/or inactivity might have on the relationship between heart rate and rate of oxygen consumption. In the present study, we investigate the extent to which body condition can affect the use of heart rate as an indicator of the rate of oxygen consumption. Twenty-two breeding king penguins (Aptenodytes patagonicus) were exercised on a variable-speed treadmill. These birds were allocated to four groups according to their sex and whether or not they had been fasting. Linear regression equations were used to describe the relationship between heart rate and the rate of oxygen consumption for each group. There were significant differences between the regression equations for the four groups. Good relationships were obtained between resting and active oxygen pulses and an index of the body condition of the birds. Validation experiments on six courting king penguins showed that the use of a combination of resting oxygen pulse and active oxygen pulse gave the best estimate of the rate of oxygen consumption V(O2). The mean percentage error between predicted and measured V(O2) was only +0.81% for the six birds. We conclude that heart rate can be used to estimate rate of oxygen consumption in free-ranging king penguins even over a small time scale (30 min). However, (i) the type of activity of the bird must be known and (ii) the body condition of the bird must be accurately determined. More investigations on the impact of fasting and/or inactivity on this relationship are required to refine these estimates further.