Biological Journal of the Linnean Society, 2003, 78, 43–49. With 3 figures
© 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 78, 43–49 43
Blackwell Science, LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066The Linnean Society of London, 2003
78
Original Article
EPIGENETIC REGULATION OF TROPHIC MORPHOLOGYC. E. ADAMS
ET AL.
*Corresponding author. E-mail: C.Adams@bio.gla.ac.uk
Epigenetic regulation of trophic morphology through
feeding behaviour in Arctic charr, Salvelinus alpinus
COLIN E. ADAMS*, CHRIS WOLTERING and GAVIN ALEXANDER
Fish Biology Group, University Field Station, Institute of Biomedical and Life Sciences, University of
Glasgow, Rowardennan, Loch Lomond, Glasgow G63 0AW, Scotland, UK
Received 11 December 2001; accepted for publication 13 August 2002
Several models of speciation suggest that in species that are phenotypically plastic, selection can act on phenotypic
variation that is environmentally induced in the earliest stages of divergence. One trait that could be subject to this
process is foraging behaviour, where discrete foraging strategies are common. One species which is highly plastic in
the expression of phenotype, the Arctic charr, Salvelinus alpinus (L.), is characterized by discrete variation in the
anatomy of the head and mouthparts. These traits have been shown to have a functional significance, but the expres-
sion of which is thought to be at least partly phenotypically plastic. Here we test the hypothesis that foraging behav-
iour may regulate the anatomy of the head and mouthparts in Arctic charr. In a dyad experiment, size-matched pairs
of fish from a mixed family group were fed a diet of either Mysis (a hard-bodied shrimp) or Chironomid larvae. Nine
morphometric measures of head dimensions that describe wild trophic morphs were measured at the start of the
experiment and 24 weeks later. Principal component scores of size-corrected morphometric measures showed highly
significant differences between fish exposed to the two diets. Univariate ANOVA analysis of the head morphometric
variables showed that fish fed on Chironomids developed longer, wider jaws, longer heads and a larger eye for a given
body length than did those fish fed upon Mysis. We conclude that foraging anatomy in Arctic charr is phenotypically
plastic and that variation in foraging behaviour that results in feeding specialization in the wild could induce vari-
ation in head anatomy. This in turn could reinforce foraging specialization. Very rapid epigenetic divergence into
distinct feeding morphs (as demonstrated here) would allow selection to act at more than one mode and thus
could promote rapid evolutionary divergence, initially prior to genetic segregation, in species which are highly
plastic. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 78, 43–49.
ADDITIONAL KEYWORDS: environmental regulation – evolution – phenotypic plasticity – speciation.
INTRODUCTION
Several models have shown that diversifying evolu-
tionary forces operating at the very earliest stages of
speciation could act on phenotypic variation that is
environmentally induced in a single gene pool, i.e.
epigenetic variation (West-Eberhard, 1989, 1998;
Wimberger, 1994, Skúlason, Snorrason & Jónsson,
1999). This occurs when the gene pool has the ability
to produce more than one discrete, alternative pheno-
type for a given characteristic; that is, where the pop-
ulation is phenotypically plastic. When this occurs,
selection may be able to operate on two or more dis-
crete phenotypes simultaneously resulting in diver-
gence (West-Eberhard, 1989). An important element of
these models is that phenotypic divergence may occur
prior to any genetic segregation (West-Eberhard,
1986, 1989, 1998; Wimberger, 1994; Skúlason et al.,
1999). The type of phenotypic variation on which these
epigenetic evolutionary mechanisms are most likely to
act initially is variation in behaviour. A likely candi-
date for selection to act upon is foraging behaviour,
because of the possibility of alternative strategies for
successful foraging and its potential effects on fitness
(see West-Eberhard, 1989; Wimberger, 1994; Skúlason
et al., 1999).
In nature, the existence of more than one discrete
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