Behavioural Processes 63 (2003) 53–61
Timing of presentation of an audience: aggressive priming
and audience effects in male displays of Siamese
fighting fish (Betta splendens)
Ricardo J. Matos
∗
, Tom M. Peake, Peter K. McGregor
Department of Animal Behaviour, Zoological Institute, Copenhagen University, Tagensvej 16, DK-2200 Copenhagen N, Denmark
Received 22 July 2002; received in revised form 31 January 2003; accepted 5 February 2003
Abstract
Studies of animal communication often underestimate the presence of individuals other than the signaller–receiver dyad.
Signalling interactions often occur in the presence of non-participating individuals (audiences); the effect of these individuals
upon the dynamics of interactions has been called the audience effect. Recent studies of fighting fish Betta splendens have shown
that the presence of a male audience can increase aggression during interactions. However, in many of these studies males were
allowed to see the audience prior to the interaction, thus such pre-exposure may have facilitated aggressive behaviour (aggressive
priming). Here we present results of two experiments designed to examine the relative importance of priming and audience effects
on the dynamics of aggressive interactions. Males that were pre-exposed showed higher levels of aggression during subsequent
interactions regardless of the presence or absence of an audience. When only one of the interactants had been pre-exposed to the
audience, the non-exposed male showed similar increases in aggressive behaviour, i.e. matching the level of aggression showed
by his opponent. Taken together these results suggest that aggressive priming may have resulted in an over-estimation of the
audience effect in previous studies. The results still highlight the importance of social environment in determining the dynamics
and outcomes of aggressive contests.
© 2003 Elsevier Science B.V. All rights reserved.
Keywords: Aggressive behaviour; Audience effect; Betta splendens; Communication network; Priming
1. Introduction
Most studies of animal communication have as-
sumed that individuals exchange signals in social iso-
lation. Consequently, models of communication and
empirical work have addressed primarily the evolution
of signals in a dyadic context. However, an increasing
number of studies have considered the influence of
the social environment on signalling behaviour (e.g.
∗
Corresponding author. Tel.: +45-35-32-13-46;
fax: +45-35-32-12-99.
E-mail address: rmatos@zi.ku.dk (R.J. Matos).
McGregor and Peake, 2000; McGregor et al., 2000;
Johnstone, 2001; Dugatkin, 2001). In these studies
the natural environment where communication occurs
is considered to be a network of several potential re-
ceivers and signallers. The dynamics created by such
an environment have marked effects on the costs and
benefits of signalling, and thus on the behavioural
strategies used by signallers during interactions (e.g.
McGregor and Peake, 2000; McGregor et al., 2000;
Johnstone, 2001). One of the major costs of signalling
in a network environment is imposed by “unwanted”
conspecific receivers. For example, in several species
the individuals (eavesdroppers or audiences) that
0376-6357/03/$ – see front matter © 2003 Elsevier Science B.V. All rights reserved.
doi:10.1016/S0376-6357(03)00029-9