Abstract Refuge use provides a good model for the
study of trade-offs between the benefits of predator
avoidance and the costs of lost feeding opportunities. We
manipulated the latter costs by subjecting similar-sized
three-spine sticklebacks to 2 days of food deprivation
followed by a 2-day re-feeding period and recorded asso-
ciated changes in body weight and refuge use. Food dep-
rivation resulted in a decrease and re-feeding in an in-
crease in the duration of refuge use by fish. Emergence
times of fish from the refuge were extremely variable
(with a ratio of 1:127 between the shortest and the long-
est ones) but individual ranks were highly consistent be-
tween different days of testing, suggesting that emer-
gence times were individually characteristic. Percentage
weight change of fish in response to the experimental
treatments also showed a high level of inter-individual
variation ranging from 0–17%. A significant positive
correlation was found between the percentage weight
lost and the percentage decrease in emergence time from
a refuge after food deprivation and similarly between the
percentage weight gained and the percentage increase in
refuge use after re-feeding. The relationship between en-
ergy turnover and behavioural strategies is discussed.
Key words Gasterosteus aculeatus · Foraging
behaviour · Refuge use · Weight change
Introduction
Two of the most important factors that determine refuge
use in animals are the intensity of a predation risk and
the nutritional state (Dill and Gillett 1991; Sih 1992).
Staying in a refuge precludes (or severely limits) forag-
ing activity; thus refuge use results in a trade-off be-
tween the benefits of predator avoidance and the costs of
lost feeding opportunities (Sih 1997). Some recent stud-
ies have focused on the importance of the nutritional
state as a predictor of emergence time in a number of
different taxa (Dill and Gillett 1991: barnacles; Koivula
et al. 1995: birds; Dill and Fraser 1997: polychaete
worm; Krause et al. 1998: fish). Many of these studies
investigated (among other factors) whether and to what
degree hiding behaviour was reduced as a function of the
duration of food deprivation but little attention was paid
to inter-individual variation in the effects of food depri-
vation periods on energy reserves and corresponding be-
havioural strategies. Krause et al. (1998) reported that
food deprivation periods of the same length caused
body-length-dependent weight losses in three-spine
sticklebacks (Gasterosteus aculeatus) with larger indi-
viduals losing a smaller proportion of their initial weight
than small ones. This difference in relative weight loss
was explained by the difference in mass-specific meta-
bolic efficiency, which is greater in larger individuals
(Weatherley and Gill 1987; Wootton 1994). As a result
of higher relative metabolic costs, smaller fish generally
showed shorter hiding times and decreased their hiding
behaviour more strongly in response to food deprivation
periods than large fish, suggesting a key role of weight
loss in influencing refuging behaviour (Krause et al.
1998).
Previous experiments have shown that three-spine
sticklebacks respond to the introduction to a new envi-
ronment by seeking shelter from which they subsequent-
ly start to investigate the surrounding area for food
(Krause et al. 1998). In this study, we used this proce-
dure to examine the effects of food deprivation and re-
feeding periods on refuge behaviour in fish of similar
J. Krause (
✉
) · S.P. Loader · E. Kirkman
Ecology & Evolution Group,
Centre of Biodiversity and Conservation, School of Biology,
University of Leeds, Leeds LS2 9JT, UK
e-mail: J.Krause@leeds.ac.uk;
Tel.: +44-113-2332840; Fax: +44-114-2332835
G.D. Ruxton
Division of Environmental and Evolutionary Biology,
Graham Kerr Building, University of Glasgow,
Glasgow G12 8QQ, UK
acta ethol (1999) 2:29–34 © Springer-Verlag and ISPA 1999
ORIGINAL ARTICLE
Jens Krause · Simon P. Loader · Emma Kirkman
Graeme D. Ruxton
Refuge use by fish as a function of body weight changes
Received: 13 December 1998 / Received in revised form: 2 May 1999 / Accepted: 7 June 1999