Behavioural Processes 68 (2005) 237–240
Commentary
Do all parasites manipulate their hosts?
Sam P. Brown
a,b
a
Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK
b
Section of Integrative Biology, University of Texas, 1 University Station, Austin, TX 78712, USA
Received 31 August 2004; accepted 31 August 2004
Thomas et al. (2005) offer a very stimulating review
of both the achievements of and prospects for research
into parasitic manipulation. Perhaps in keeping with the
maturity of this research field, the notion of parasitic
manipulation received only a brief passing definition
in their review, before current controversies took cen-
tre stage. At the risk of pedantry, my aim in this com-
mentary is to focus closely on definitions of the related
concepts of host manipulation and virulence. I argue
that an inspection of these concepts and their biolog-
ical underpinnings leads to the conclusion that while
parasites are not necessarily manipulative, a majority
of parasites can be seen to be manipulators. Perhaps
the only unambiguous cases of non-manipulation are
restricted to socially-defective microparasites.
To begin, consider the opening definition offered by
Thomas et al. (2005).
... host manipulation, which occurs when a parasite en-
hances its own transmission by altering host behaviour
[p. 2, lines 4–5]
Throughout their review, and indeed throughout the
host-manipulation literature, we see numerous cases
that fit exactly with this definition—typically the par-
E-mail address: sam@biosci.utexas.edu.
asite is a helminth worm, and the enhanced transmis-
sion is between an intermediate and definitive host in
a complex life cycle. However, the review also drops
hints towards a larger scope for the host manipulation
concept. At times, the authors talk of parasites altering
the host phenotype—a more inclusive term than be-
haviour. Elsewhere, they allude to parasites enhancing
aspects of fitness other than transmission (for instance
their persistence in the face of a host immune response).
Putting these two generalisations together, we can ex-
pand the above definition to
... host manipulation, which occurs when a parasite
enhances its fitness by altering its host’s phenotype
This broader definition is consistent with the full
range of examples cited in Thomas et al. (2005),
and mirrors more inclusive previous definitions (e.g.
Brown, 1999). It is also remains consistent with the
interpretation of host-manipulation as an example of
Dawkins’ notion of the extended phenotype (Dawkins,
1982, 1990; Thomas et al., 2005).
By broadening the definition of host-manipulation
beyond its historical roots in macroparasites, do we
run the risk of creating a definition that is merely
tautological with parasitism? Consider two mutually-
exclusive classes of parasite: manipulators and non-
0376-6357/$ – see front matter © 2004 Published by Elsevier B.V.
doi:10.1016/j.beproc.2004.08.015