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Exercise
places demands on the respiratory, circulatory and
muscular systems, and limitations in any one of
these could, in turn, limit the exercise
capabilities of the whole organism. One of the most
extreme forms of exercise is the long migrations of
some species of birds and yet very little is known
about the detailed behaviour and physiology of
migrating birds (see review by Butler & Bishop,
2000). Using the latest lightweight satellite
transmitters (PTTs) and implantable data loggers, we
have obtained unique physiological and behavioural
data (route taken, stopover areas, duration of
non-stop flights, etc) on adult barnacle geese
during their autumn migration from a breeding colony
(at Ny-Ĺlesund) in the NW of the island of
Spitsbergen in the high Arctic to Caerlaverock in
the Solway Firth, southern Scotland (Butler et al.,
1998; Butler & Woakes, 1998, 2001).
Accurate estimates of the rates of energy
expenditure of animals are essential components of
many behavioural and ecological studies and yet
there are very few means by which field metabolic (FMR)
rate can be determined. We have developed (Woakes et
al., 1995) and deployed a data logging system which
enables us to record heart rate (fH) and body
temperature (Tb) over
long periods of time. With proper calibration and
validation experiments, it has been possible to
demonstrate that, in a number of species of
terrestrial and aquatic birds, there is a linear
relationship between fH and rate of oxygen
consumption and that the former can be used to
estimate the latter (Nolet et al., 1992; Bevan et
al., 1994, 1995a, 1995b; Hawkins et al., 2000; Green
et al., 2001; Froget et al., 2001). However, it has
become clear that various factors, such as body
condition associated with periods of fasting during
breeding and moulting in penguins, and different
forms of exercise, such as running and flying, can
affect the relationship between fH and the oxygen
consumption rate (Green et al., 2001; Froget et al.,
2001; Butler et al., 2000; Ward et al., 2002; Clarke
et al., 2006). Thus
it is important to perform calibration experiments
under conditions which, as far as is possible,
simulate those to which the animals will be exposed
in the field. One advantage of the heart rate method
for estimating of animals, is that it can be used to
estimate the energy cost of specific activities,
such as breeding, foraging, moulting, migration,
etc. (Bevan et al., 1995a, 2002; Butler et al.,
1998; Green et al., 2002), as well as give an
indication of the overall energy expenditure, and
hence food requirements, of animals in the field. We
have completed a study on year-round energetics of
macaroni penguins and are now studying king penguins
and black-browed albatrosses. A combined laboratory
and field study of foraging behaviour and physiology
of cormorants is also underway.
Recent
Publications
(click
HERE
for full publication list):
Halsey, L.G., Butler, P.J. and Woakes, A.J. 2005.
Breathing hypoxic gas affects the physiology as well
and the diving behaviour of tufted ducks. Physiol.
Biochem. Zool. 78, 273-284.
Guillemette, M., Woakes, A. J., Henaux, V.,
Grandbois, J-M. and Butler, P. J. 2005. The effect
of depth on the diving behaviour of common eiders.
Can. J. Zool. 82, 1818-1826
Clark, T.D, Ryan, T., Ingram, B.A., Woakes, A.J.,
Butler, P.J. and Frappell, P.B. 2005. Factorial
aerobic scope is independent of temperature and
primarily modulated by heart rate in exercising
Murray cod (Maccullochella peellii peellii). Physiol.
Biochem. Zool. 78, 347-355.
Clark, T.D., Wang, T., Butler, P.J. and Frappell,
P.B. 2005. Factorial scopes of cardio-metabolic
variables remain constant with changes in body
temperature in the varanid lizard, Varanus
rosenbergi. Am. J. Physiol. 288, R992-R997.
Green, J., Woakes, A., Boyd, I. and Butler, P J.
2005. Cardiovascular adjustments during locomotion
in penguins. Can. J. Physiol. 83, 445-454.
Halsey, L.G., Wallace, S.E, Woakes, A.J., Winkler,
H. and Butler P.J. 2005. Tufted ducks Aythya
fuligula do not control buoyancy during diving. J.
av. Biol. 36, 261-267.
Clark, T.D., Butler, P.J. and Frappell, P.B. 2005.
Digestive state influences the heart rate hysteresis
and rates of heat exchange in the varanid lizard,
Varanus rosenbergi. J. exp. Biol. 208, 2269-2276.
Green, J.A., Boyd, I.L., Woakes, A.J., Green, C.J.
and Butler, P.J. 2005. Do seasonal changes in
metabolic rate facilitate changes in diving
behaviour? J. exp. Biol. 208, 2581-2593
Day,
N. and Butler, P.J. 2005. The effects of acclimation
to reversed seasonal temperatures on the swimming
performance of adult brown trout Salmo trutta. J.
exp. Biol. 208, 2683-2692.
Winter, M.J. Verweij, F., Garofalo, E., Ceradini,
S., McKenzie, D. J., Williams, M. A., Taylor, E. W.,
Butler, P. J., van der Oost, R. and Chipman, J. K.
2005. Tissue levels and biomarkers of organic
contaminants in feral and caged chub (Leuciscus
cephalus) from rivers in the West Midlands, UK. Aq
Toxicol. 73, 394-405.
Green, J. A., Boyd, I. L., Woakes, A. J. and Butler,
P. J. 2005. Behavioural flexibility during
year-round foraging in macaroni penguins. Mar. Ecol.
Prog. Ser. 296, 183-196.
Fahlman, A., Schmidt, A., Handrich, Y., Woakes, A.J.
and Butler, P.J. 2005. Metabolism and
thermoregulation during fasting in king penguins,
Aptenodytes patagonicus, in air and water. Am. J.
Physiol. 289, R670-679.
Grémillet, D., Kuntz, G., Gilbert, C., Woakes, A.J.,
Butler, P.J and Le Maho, Y. 2005. Cormorants dive
through the Polar night. Biol Letts 1, 469-471.
Grémillet, D., Kuntz, G., Woakes, A.J., Gilbert, C.,
Robin, J-P., Le Maho Y. and Butler. P.J. 2005.
Year-round recordings of behavioural and
physiological parameters reveal the survival
strategy of a poorly insulated diving endotherm
during the Arctic winter. J exp Biol 208, 4231-4241.
Green, J.A., White, C.R. and Butler, P.J. 2005.
Allometric estimation of metabolic rate from heart
rate in penguins. Comp Biochem Physiol A 142,
478-484.
Halsey, L., Brand, O.J., Woakes, A.J. and Butler,
P.J. 2006. Experiments on single diving animals in
the laboratory often measure dives of decreased
effort. Ibis 148, 164-166
Watkins, E. J., Hall, A. D., Wiseman, J. and Butler,
P. J. 2006. Feed metabolism and resting metabolic
rate in mallards and domestic ducks. J Zool. (in
press).
Clark, T.D., Butler, P.J. and Frappell, P.B. 2006.
Factors influencing the prediction of metabolic rate
in a reptile. Func. Ecol. (in press).
Halsey, L.G., Butler, P.J. and Blackburn, T.M. 2006.
A phylogenetic analysis of the allometry of diving.
Am Nat (in press).
McKenzie, D.J., Garofalo, E., Winter, M.J., Ceradini,
S., Verweij, F., Day, N., Hayes, R., van der Oost,
R., Butler, P.J., Chipman, J.K. and Taylor, E.W.
2006. Swimming performance in fish: a physiological
biomarker of aquatic pollution? Phil Trans Roy Soc
(in press). |
