‘Central place forager’ is a term used to describe the
behaviour of a foraging animal that repeatedly returns to the
same place with food (Orians and Pearson, 1979; Lessells and
Stephens, 1983). The close analogy between a bird returning
to its nest with food (Kacelnik, 1984) and a diving animal
returning to the surface to breathe means that the concept of
central place foraging can be applied to surfacing divers
(Houston and McNamara, 1985). In this case, the surface of
the water is the ‘central place’. Thus, models that examine the
behaviour of central place foragers can be transposed to
examine the behaviour of a diving animal.
During foraging bouts, assuming that a diver wants to be
under the water acquiring food for as much time as possible,
repeatedly returning to the surface for oxygen is (generally) in
direct competition with food gain (Ydenberg and Clark, 1989).
Optimal foraging models, which are based on the marginal
value theorem (Charnov, 1976), have been developed to
determine the optimal surface time for oxygen loading that
maximises the proportion of time spent foraging (Kramer,
1988; Houston and Carbone, 1992; Carbone and Houston,
1996). These models are based on the physiological gains and
losses of oxygen over the dive cycle and therefore the post-
dive loading curve is an integral, but as yet unquantified,
part of many theoretical studies of optimal time allocation
(Thompson et al., 1993; Carbone and Houston, 1994, 1996;
Lea et al., 1996; Walton et al., 1998; Mori, 1998, 1999).
Because the mammalian lung system rapidly collapses on
immersion (Kooyman and Ponganis, 1998), the majority of the
oxygen stores of marine mammals consist of haemoglobin and
myoglobin. Thus it is likely that the generic shape for the
oxygen loading curve proposed and used by Kramer (1988) in
his optimal breathing model (Fig. 1) may be correct for most
diving mammals. However, Walton et al. (1998) have argued
that this shape is not accurate for diving birds.
The respiratory system of birds differs from that of
mammals and consists mainly of large distended air sacs and
a lung, which is comparatively small and rigid (Scheid, 1979).
However, the estimated contribution of the air sacs to the total
body oxygen stores of diving birds ranges from 23% to 64%,
e.g. tufted duck (Keijer and Butler, 1982), thick-billed murre
(Croll et al., 1992), lesser scaup (Stephenson, 1995), Adélie
penguin (Kooyman and Ponganis, 1998), king penguin
(Ponganis et al., 1999). Therefore, it has been suggested that
the respiratory system will have a profound influence on the
dynamics of post-dive oxygen loading. The oxygen stored
within the air sacs of tufted ducks during a dive is made
available for consumption through locomotion, and associated
abdominal activity. This produces pressure differentials
3945 The Journal of Experimental Biology 205, 3945–3954 (2002)
Printed in Great Britain © The Company of Biologists Limited
JEB4342
The rate of oxygen uptake at the surface between dives
was measured for four tufted ducks, Aythya fuligula,
during bouts of foraging dives to a depth of 1.8 m. The
ducks surfaced into a respirometer box after each dive so
that the rate of oxygen uptake (V
.
O
∑) could be measured.
V
.
O
∑ decreased over time at the surface and there was a
particularly rapid phase of oxygen uptake for
approximately the first 3 s. The specific shape of the
oxygen uptake curve is dependent upon the duration of
the preceding dive. The uptake curve after longer dives
was significantly steeper during the first 3 s at the surface
than after shorter dives, although V
.
O
∑ after the first 3 s
was not significantly different between these two dive
duration bins. Thus, the mean total oxygen uptake (V
O
∑)
was higher after surface periods following longer dives.
Due to the high V
.
O
∑ during the initial part of the surface
period, the curve associated with longer dives was
statistically biphasic, with the point of inflection at 3.3 s.
The curve for shorter dives was not statistically biphasic.
The birds may increase their respiratory frequency during
the first 3 s after longer dives, producing the increased
V
.
O
∑, which would enable the birds to resaturate their
oxygen stores more rapidly in response to the increased
oxygen depletion of the longer submergence time.
Key words: tufted duck, Aythya fuligula, diving, oxygen uptake,
optimal foraging, model.
Summary
Introduction
Oxygen uptake during post dive recovery in a diving bird Aythya fuligula:
implications for optimal foraging models
Roland Parkes
1
, Lewis G. Halsey
1,
*, Anthony J. Woakes
1
, Roger L. Holder
2
and Patrick J. Butler
1
1
School of Biosciences, and
2
School of Mathematics and Statistics, University of Birmingham, Edgbaston,
Birmingham, B15 2TT, UK
*Author for correspondence (e-mail:lgh013@bham.ac.uk)
Accepted 16 September 2002