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ORIGINAL PAPER
Received: 25 November 2010 / Revised: 14 June 2011 / Accepted: 30 June 2011 / Published online: 16 July 2011
# Springer-Verlag 2011
Communicated by A. Schulte-Hostedde
A. Duek (*) : L. Barto
Department of Ethology, Institute of Animal Science,
Ptelstv 815,
CZ-10400 Praha, Czech Republic
e-mail: adamdusek@yahoo.com
A. Duek
Biodiversity Research Group, Department of Zoology,
Faculty of Science, Charles University,
Vinin 7,
CZ-12844 Praha, Czech Republic
F. Sedlek
Department of Zoology, Faculty of Science,
University of South Bohemia,
Braniovsk 31,
CZ-37005 esk Budjovice, Czech Republic
Introduction
The sex allocation theory (Charnov 1982) predicts that natural
selection will favor parental ability to vary offspring sex
whenever fitness returns from investing in sons and daughters
differ. To predict offspring sex ratio (sons to daughters)
variation in mammals, several adaptive hypotheses have been
proposed (reviewed in Clutton-Brock and Iason 1986; Frank
1990; Cockburn et al. 2002). The hypothesis proposed by
Trivers and Willard (1973) has become one of the most
influential.
Trivers and Willard (1973) pointed out that in polygynous
or highly promiscuous mammals males can achieve higher
reproductive success than females and extra parental
investment can thus have a larger effect on the fitness return
from sons. Taking this aspect into account, Trivers and
Willard (1973) hypothesized that mothers in good condition
should tend to produce sons whereas mothers in poor
condition should tend to produce daughters. In looking for
the mechanism whereby females could adjust the offspring
sex ratio, Trivers and Willard (1973) suggested that In
species such as mammals, in which males determine sex of
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Table 1 The predicted effects of food restriction-decreased maternal condition on the litter characteristics at birth
Hypothesis
Myers
Williams
TriversWillards
Decreasing
Unchanged/increasing
Decreasing
Decreasing
Decreasing
Unchanged/decreasing
The table shows the predictions relevant for a particular hypothesis. Although the plausibility of the TriversWillard hypothesis for polytocous
mammals is controversial, for an illustration of the differences between the discussed hypotheses (TriversWillards, Myers, Williams), the table
also shows the predictions derived from the TriversWillard hypothesis. The predictions on the secondary sex ratio and the birth litter size concern
both the feeding treatment and the maternal weight. By contrast, to assess gradual change in the total production costs, the predictions on the total
production costs concern only the maternal weight.
Results
In total, we examined 865 pups of 65 food-restricted and
961 pups of 72 control mothers. Consistent with our
assumptions, (1) producing a male was, regardless of the
feeding treatment, 1.04 times costlier than producing a
female, and (2) the pre-mating food restriction significantly
decreased (a) the proportion of fertile females, (b) the
maternal weight (both W2 and W), and (c) delayed the
delivery date (Table 2), indicating a general decline in the
maternal condition. The food-restricted and control mothers
did not differ (t135 =0.46, p=0.65) in their weight before the
period of food restriction (W1).
Table 2 The effect of feeding
treatment on female fertility,
maternal weight at mating (W2),
maternal weight change (W),
and mothers delivery date
Feeding treatment
Control
n
75
n
72
72
72
Food-restricted
Proportion
96.00
MeanSE
27.550.33
2.050.43
21.900.14
n
77
n
65
65
65
Proportion
84.42
MeanSE
24.510.30
9.810.71
22.800.22
2 value
5.73
t value
6.80
14.61
3.41
p value
<0.02
p value
<0.0001
<0.0001
<0.001
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the secondary sex ratio and the birth litter size (F1,132 =66.33,
p<0.0001; Fig. 4c).
Discussion
Regardless of the Offspring sex, the pre-weaning offspring mortality was significantly lower in the foodrestricted than control mothers (21 =4.68, p=0.03; Fig. 6).
mating are nested within the feeding treatment of the mother. Each
data point represents an observation of one litter per group (control,
n=72; food-restricted, n=65). The best-fit regression lines are shown
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