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the puparium). To minimize inbreeding, a population size of at least
100 was maintained in all lines. With the exception of exposure to
parasitoids, control and selection lines were cultured using identical
procedures. Larvae were exposed to parasitoids in the selection
treatment in conditions that lead to high rates of parasitism to avoid
selection on traits other than parasitoid resistance. The encapsulation ability of the flies was assessed by exposing 200 second-instar
larvae to parasitoid females for 2 h, after which the larvae were kept
at 20 8C for five days and then dissected to score capsule frequency.
The response to selection was rapid and similar across the four
lines (Fig. 1). The probability of encapsulation rose from 5% to a
maximum of ,60% in five generations. The maximum encapsulation ability is similar to that found in Mediterranean populations,
which are heavily attacked by A. tabida13. After selection, four pairs
of F1 lines were created by crossing each of the selected lines with a
different control line. In one cross of the pair, males derived from
selected lines and females from control lines, whereas in the other
cross the sexes were reversed. The mean encapsulation rate of the
offspring was 34% (standard error 4%), and was not influenced by
the origin of the female parent (t-test, P 0:9), excluding maternal
or cytoplasmic effects, and strongly suggesting that the genes
influencing resistance are not sex-linked.
We searched for correlated responses to selection by comparing a
range of fitness parameters in the control and selected lines. In a first
set of experiments we compared the performance of flies reared
under conditions of abundant food and very low competition. The
full details of the experiments will be reported elsewhere, but in
summary we found no significant differences in larval and pupal
survival, larval and pupal development time, adult longevity in the
absence of food, early female fecundity, adult size or fluctuating
asymmetry. Working with only eight lines in total we have limited
statistical power to detect subtle differences, but in no cases were
strong but insignificant trends detected.
In a second set of experiments we compared the performance of
control and selected flies under conditions of weak to strong
intraspecific competition (Fig. 2). Competitive ability was assessed
by rearing larvae with flies from a genetically marked tester
stock14,15 at four different levels of competition for food (see
Methods for experimental and statistical details). When competition was weak (0.4 ml and 0.2 ml larval food), survival was high
(,80%) and there was no difference between the two sets of lines
(P . 0:1). However, for more severe competition (0.1 ml), survival
was reduced and there was significantly greater mortality among
selected than control flies (P 0:011). This difference was also seen
at the most severe level of competition (0.05 ml), where survival was
much lower (,50%), although the statistical significance was less
(P 0:061) owing to the much greater within-line variability in
survival when resources were very scarce. At higher levels of
competition, flies are smaller, take longer to develop, and show
more fluctuating asymmetry, but we found no significant differences between the performances of the control and selected lines
(although there was a trend for selected flies to be smaller at the two
lower levels of competition).
Thus reduced survival in a high-competition environment is a
correlated response to selection for improved defence against
endoparasitoid attack in D. melanogaster. This suggests there is a
trade-off between defence against parasitoids and other components of fitness. We suspect that selected larvae allocate more
resources to the machinery of cellular encapsulation or to counteracting the wasps attempts to disable encapsulation, and hence are
less able to withstand very stressful conditions. Several other tradeoffs in the life-history strategies of D. melanogaster have been
identified only in circumstances when the fly is under stress2.
D. melanogaster populations in the wild feed in rotting fruit and
other fermenting substrates, and often experience a level of competition in the field comparable with those at which survival of the
selected larvae was reduced16.
We believe this to be the first demonstration of a cost of resistance
against parasitoids. For many species of hosts, parasitoid attack
rates vary both temporally and spatially, and the resulting fluctuating selection pressures, combined with costs to parasitoid defences,
may explain the additive genetic variance in defensive ability
observed in several host species810. The population dynamics of
hosts and their parasitoids are often coupled, and our findings
suggest that there may be complex joint evolutionary and population dynamic interactions. More generally, evidence for costs of
means and standard errors of the four selected and control lines. For logistical
symbols; selected lines, filled symbols) relative to a tester strain, showing means
and standard errors of the four selected or control lines at four levels of
competition for larval food. The competition index and the statistical analysis are
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resistance against pathogens and parasites in animals is scarce, and
comes largely from studies on populations of farm or laboratory
animals with highly modified genetic backgrounds (reviewed in
ref. 1). Finally, because of the enormous amount known about
Drosophila genetics, we suggest that the interactions between
D. melanogaster and A. tabida may be a valuable model system for
the study of the evolution of resistance and of the genetic basis of
M
adaptation.
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Methods
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