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Mixed sex allocation strategies in a

polytocous mammal, the house mouse


(Mus musculus)
Adam Duek, Ludk Barto & Frantiek
Sedlek

Behavioral Ecology and Sociobiology


ISSN 0340-5443
Volume 65
Number 12
Behav Ecol Sociobiol (2011)
65:2209-2217
DOI 10.1007/s00265-011-1229-8

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Behav Ecol Sociobiol (2011) 65:22092217
DOI 10.1007/s00265-011-1229-8

ORIGINAL PAPER

Mixed sex allocation strategies in a polytocous mammal,


the house mouse (Mus musculus)
Adam Duek & Ludk Barto & Frantiek Sedlek

Received: 25 November 2010 / Revised: 14 June 2011 / Accepted: 30 June 2011 / Published online: 16 July 2011
# Springer-Verlag 2011

Abstract The issue of adaptive adjustment of offspring sex


ratio (proportion of male births) in polytocous mammals,
producing several offspring per litter, is controversial because
females of these species can maximize their fitness mainly by
adjusting offspring number. To address this issue, we
examined the effect of maternal condition at mating,
experimentally decreased by pre-mating food restriction, on
the sex ratio variation in 137 female mice. We tested two basic
sex allocation hypotheses plausible for polytocous mammals:
(1) the Myers hypothesis, predicting that cheaper sex should
be favored in poor environmental conditions to maximize
offspring number; and (2) the Williams hypothesis, predicting
maximum fitness returns by adjusting size- and sex-specific
composition of the litter according to the maternal condition.
The food-restricted mothers produced larger litters with a
higher proportion of cheaper daughters than the control
mothers. By contrast, the control mothers optimized size and
sex composition of the litter according to their weight at
mating. In addition, the offspring of the food-restricted
mothers suffered less from pre-weaning mortality than those

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

of the control mothers. Therefore, when comparing the


groups, the Myers hypothesis had a general significance
while the Williams hypothesis was plausible only for the
control mothers. Furthermore, some of the food-restricted
mothers partly coped with the pre-mating food restriction and
increased the proportion of sons in the litter with the
increasing maternal weight loss (during the period of food
restriction). The sex ratio variation was thus a result of three
sex allocation strategies depending on the maternal condition
at mating.
Keywords Sex allocation . Maternal condition . Food
restriction . Sex ratio . Litter size . Mouse

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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offspring, female control of the sex ratio must involve


differential mortality by sex, either of sperm cells or of
growing young during parental investment.
Although polytocous mammals (i.e., species producing
several offspring per litter) were not explicitly excluded
from the original hypothesis of Trivers and Willard (1973)
other authors (e.g., Frank 1990; Hardy 1997; Krackow
1997; Carranza 2004) later suggested that the Trivers
Willard hypothesis may not be plausible for this type of
mammal. The reason for this suggestion is that the mothers
of polytocous mammals can optimize their reproductive
fitness not only by adjusting offspring sex ratio but mainly
by adjusting offspring number. This key feature has been
considered in two alternative hypotheses, the first proposed
by Myers (1978) and the second by Williams (1979).
Myers (1978) critically reviewed the TriversWillard
hypothesis (Trivers and Willard 1973) and proposed another
hypothesis predicting that offspring sex ratio should be
adjusted to maximize the number of successful offspring
rather than the average reproductive potential of each
offspring. This prediction is based on the argument
originally pointed out by Fisher (1930) that if the
production of one sex is more costly than that of the other,
parents can increase their reproductive success by producing
the cheaper sex. Myers (1978) related this argument to the
situation when the food resources are limited, and proposed
that the cheaper sex should be favored over the more
expensive sex in poor environmental conditions since it is
less vulnerable to inadequate parental investment. In most
mammals, males are the more expensive sex (Clutton-Brock
et al. 1985; Cockburn et al. 2002; Lindenfors et al. 2007)
and the offspring sex ratio should, therefore, decrease with a
decreasing maternal condition. In contrast to Trivers and
Willard (1973), Myers (1978) supposed a mechanism
operating before or at conception.
Williams (1979), on the other hand, pointed out that if
more than one offspring is produced, mothers should adjust
the offspring sex ratio along with the size of the litter to
maximize fitness returns from sons and daughters. In
polytocous mammals, maternal condition should, therefore,
be related to a specific sequence of increasing production
costs based on size and sex composition of the litters (Frank
1990; Servanty et al. 2007). Suppose, for example, that
producing a male (m) is 1.4 times costlier than producing a
female (f), then the specific sequence of production costs
should increase as follows: 1f<1m<2f<1m1f<2m<3f<
1m2f<2m1f<4f<3m<1m3f, etc. (see Fig. 1 for a diagram).
To test the Williams hypothesis, using the wild boar (Sus
scrofa scrofa), Servanty et al. (2007) recently proposed a
new computational method. They quantified the relative
costs of producing a male versus a female and related the
sequence of increasing production costs to maternal
condition. To assess the production costs of the whole

Behav Ecol Sociobiol (2011) 65:22092217

Fig. 1 Diagram illustrating an increase in the sequence of production


costs with the increasing maternal condition. In this example, producing a
male () is 1.4 costlier than producing a female (), i.e., production
costs of 1.4 for a male and 1 for a female. Note that not only the
secondary sex ratio (solid line), but also the birth litter size (dashed line)
can change non-linearly with the increasing maternal condition

litter, we propose to use the term Total production costs,


summarizing the production costs of all male and female
pups. This term will be used further throughout the text.
In order to determine the significance of litter size for
offspring sex ratio variation, we tested the Myers and
Williams hypotheses in a typical polytocous mammal, the
house mouse (Mus musculus). We investigated the effect of
maternal condition at mating, experimentally decreased by
pre-mating food restriction, on (1) the secondary sex ratio
(proportion of male births), (2) the birth litter size, and (3)
the total production costs. First, to assess the effect of food
restriction on the maternal condition, we hypothesized that
the food-restricted females will show (a) a lower fertility (a
lower proportion of females littering), (b) a lower maternal
weight, and (c) a delayed delivery date. Assuming that (1)
producing a male will be costlier than producing a female,
and (2) the maternal condition at mating will decrease due
to the pre-mating food restriction, we then tested the
predictions on the litter characteristics at birth (13),
derived from the Myers and Williams hypotheses (see
Table 1). In addition, to fully evaluate the Myers hypothesis,
we also investigated the effect of pre-mating food restriction
on the pre-weaning offspring mortality (proportion of
offspring deaths from day 1 to 21). Assuming that the
food-restricted mothers will optimize the sex composition of
the litter to maximize the number of successful offspring (as
predicted by Myers 1978), we hypothesized that, regardless
of the sex, the offspring of the food-restricted mothers will
be more viable than those of the control mothers.

Materials and methods


Animals and data collection
We used outbred CD-1 mice (Faculty of Medicine MU,
Czech Republic). The animals were housed individually in

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Table 1 The predicted effects of food restriction-decreased maternal condition on the litter characteristics at birth
Hypothesis

Secondary sex ratio (%)


Birth litter size
Total production costs

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.

plastic cages (223614 cm) with a standard grain-based


pellet (ST-1) diet (Velas, a. s., Hrabanov, Lys nad Labem,
Czech Republic) and water provided ad libitum. The colony
was maintained in a temperature controlled room (241C)
with a 14:10-h light/dark cycle.
Reproduction was studied in three independent, consecutive
experimental trials consisted of a total of 152 adult (6080 days
old) virgin females. In 77 randomly selected females, the
maternal condition at mating was experimentally modified by
the intermittent feeding (four 24-h food deprivation
periods every other day) during the week before pairing
(as implemented by Meikle and Drickamer 1986; Meikle
and Thornton 1995). To assess the effect of food restriction
on the maternal condition at mating, we also used an absolute
weight at pairing, i.e., on the day after the period of food
restriction (W2), and a relative weight change during the
period of food restriction (W), computed as follows:


W2  W1
W
 100%
W1
where W1 is the maternal weight before and W2 after the
period of food restriction. For control females (the rest of the
mice), fed ad libitum at all times, W from an equivalent
period was computed. For simplification, W2 will be further
referred as Maternal weight at mating and W as
Maternal weight change.
Each female was paired with a randomly selected adult
male (n=152; removed after 10 days) and checked for
delivery daily. On the day of birth, the pups were counted,
individually marked and sexed. Mothers and pups were
then weighed and the total production costs (Ct) for all
litters were computed with the formula:
Ct Cm  Nm Cf  Nf
where Cm and Cf represent the production costs for a single
male and a single female, while Nm and Nf indicate the
numbers of male and female pups in the litter, respectively.
Cm and Cf were derived from the average birth weights of

all male (Wm) and female (Wf) pups (as proposed by


Servanty et al. 2007). Assuming Cf <Cm, when Cf =1, then
Cm can be written as:


Wm  Wf
Cm Cf
Wm
The body weight was measured with a digital balance
(model EMB 200-2; Kern & Sohn GmbH, Germany;
weighing accuracy of 0.01 g).
During the lactation period (day 1 to 21), mothers and
pups were checked every fourth day (1, 5, 9, 13, 17, and
21 days) for health status and mortality. On the day of
weaning (day 21), the number and sex of the surviving
pups was recorded.
Statistical analysis
All analyses were performed using the SAS System (V 9.2,
20022008, SAS Institute, Cary). A chi-square contingency
table analysis was used to assess (1) the effect of Feeding
treatment on Female fertility (proportion of females
littering), and (2) the effect of Feeding treatment and
Offspring sex on Pre-weaning offspring mortality
(proportion of offspring deaths from day 1 to 21). The
effect of Feeding treatment on Maternal weight (both
W2 and W) and Mothers delivery date (days from
pairing to delivery) was assessed by a t test. In addition, a t
test was also used to compare the food-restricted and
control mothers for their weight before the period of food
restriction (W1).
A General Linear Mixed Model (GLMM) using the PROC
MIXED with restricted maximum likelihood estimation was
applied to test factors affecting (1) Secondary sex ratio
(proportion of male births); (2) Birth litter size; and (3)
Total production costs, derived from (4) Offspring birth
weight. In particular GLMM models, with random effects
Experimental trial (14) and Maternal identity (4), the
tested fixed effects were the class variables Feeding

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Behav Ecol Sociobiol (2011) 65:22092217

treatment (14) and Offspring sex (4), and the continuous


variables Maternal weight at mating (W2: 14), Maternal
weight change (W: 14), Maternal delivery weight (4),
Mothers delivery date (14), Birth litter size (1, 3, and 4)
and Secondary sex ratio (2, 3, and 4). The assumption of
normality was met in all continuous variables.
All interaction terms were tested but were not reported
unless statistically significant. With respect to the Williams
hypothesis-predicted specific sequence of increasing
production costs (see Introduction), the dependent variable
Total production costs was explicitly tested for the
interaction between the Secondary sex ratio and the Birth
litter size. The predicted values of the dependent variable
were computed and plotted against the continuous fixed
effect with the predicted regression line (as recommended by
Tao et al. 2002). In the case that the predicted values were
distributed on the fitted regression line, the raw data points
were presented. The within-class differences were indicated
by modeladjusted least-square means with standard errors
(SE) to account for an unbalanced design with unequal
numbers of control and food-restricted mothers. Differences
within the class were tested by a t test. When a significant
difference was detected, the least-square means were
compared by the TukeyKramer adjustment for multiple
comparisons. All cited p values were two-tailed, with a
significance level () set at 0.05.

Secondary sex ratio


The food-restricted mothers produced litters with a lower
proportion of sons than the control mothers (F1,133 = 4.21,
p = 0.04; t133 = 2.05, p = 0.04; TukeyKramer adjustment
p = 0.04; Fig. 2a). In addition, the mothers adjusted the
proportion of sons in the litter according to their weight
change during the period of food restriction (F2,133 = 3.22,
p = 0.04; Fig. 3a). This effect was, however, apparent only
in the food-restricted mothers where the proportion of sons
increased with the increasing maternal weight loss
(t133 =2.01, p=0.05). In the control mothers, the same but
non-significant (t133 =1.55, p=0.12) trend was apparent.
Birth litter size
The food-restricted mothers tended to produce larger litters
than the control mothers (F1,132 =5.76, p=0.02; t132 =2.40,
p=0.02; TukeyKramer adjustment p=0.06; Fig. 2b). In
addition, the number of pups in the litter was affected by
the maternal weight at mating (F2,97.4 =9.75, p=0.0001;
Fig. 3b). The birth litter size of the control mothers
increased with the increasing maternal weight (t112 =4.38,
p<0.0001). On the contrary, the birth litter size of the foodrestricted mothers was independent of the maternal weight
(t108 =0.89, p=0.38).
Total production costs

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

The total production costs of the food-restricted mothers


tended to be higher than those of the control mothers
(F1,132 =4.36, p=0.04; t132 =2.09, p=0.04; TukeyKramer
adjustment p=0.07; Fig. 4a). But more importantly, the total
production costs were affected by the maternal weight at
mating (F2,132 =10.89, p<0.0001; Fig. 4b). In the control
mothers, the total production costs increased with the
increasing maternal weight (t132 =4.49, p<0.0001). By
contrast, the total production costs of the food-restricted
mothers were maternal weight-independent (t132 = 1.24,
p = 0.22). Further, regardless of the feeding treatment, the
total production costs increased with the interaction between

Feeding treatment
Control

Female fertility (%)


Maternal weight at mating (g)
Maternal weight change (%)
Mothers delivery date (days)

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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Fig. 2 The secondary sex ratio


a and the birth litter size
b depending on the feeding
treatment of the mother
(control, n=72; food-restricted,
n=65). Bars indicate
least-square means (SE)

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).

The sex ratio variation was a result of three sex allocation


strategies depending on the maternal condition at mating. In
accordance with earlier studies (Meikle and Drickamer
1986; Meikle and Thornton 1995), we have observed that
the food-restricted mothers tended to produce a lower
proportion of sons than the control mothers (Fig. 2a). In
addition, we have also observed that the more weight the
food-restricted mothers lost (during the period of food
restriction) the more sons they tended to produce (Fig. 3a).
By contrast, the control mothers optimized the total
production costs (based on the size and sex composition
of the litter) according to their weight at mating (Fig. 4b).
Presuming that the maternal weight loss reflects a
decline in the maternal condition, the increase in the
proportion of sons with the increasing maternal weight loss
seems to be in contrast to recent studies (e.g., Alexenko et
al. 2007; Cameron et al. 2008) reporting a decrease in the
proportion of sons due to a condition decline. However, it is
possible that some of the food-restricted females had
actually recovered from the pre-mating food restriction
and their condition in part increased (Meikle and Thornton

Fig. 3 The secondary sex ratio depending on the maternal weight


change a, and the birth litter size depending on the maternal weight at
mating b. Both the maternal weight change and the maternal weight at

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

Offspring birth weight


The offspring birth weight was not affected either by the
Feeding treatment or by the Maternal weight at
mating and/or the Maternal weight change. The
female offspring were lighter than the male offspring
(F1,1710 = 157.69, p < 0.0001; t1710 = 12.56, p < 0.0001;
TukeyKramer adjustment p<0.0001; Fig. 5a). In addition,
the birth weight of the offspring decreased with the
increasing birth litter size (F 1,135 = 58.61, p < 0.0001;
Fig. 5b), while increased with the increasing maternal
delivery weight (F1,131 =9.14, p<0.01; Fig. 5c) and the
increasing mothers delivery date (F1,133 =8.89, p<0.01;
Fig. 5d).
Pre-weaning offspring mortality

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Fig. 4 The total production


costs depending on the feeding
treatment of the mother a, the
maternal weight at mating
nested within the feeding
treatment of the mother b,
and the interaction between the
secondary sex ratio and the birth
litter size c. Bars indicate leastsquare means (SE), and each
data point represents a predicted
value of one litter per group
(control, n=72; food-restricted,
n=65). The best-fit regression
lines (b) and regression surface
(c) are shown

1995). This possibility can be supported: (1) by the food


restriction-modified effect of maternal condition on the
birth litter size (Figs. 2b and 3b) and the total production
costs (Figs. 4a and b); and (2) by the positive effect of
food restriction-delayed delivery date on the offspring
birth weight (Fig. 5d), indicating a coping strategy (i.e.,
the ability to cope with adverse conditions, in this case, the
pre-mating food restriction; for more information see, e.g.,
Mendl and Deag 1995).
The above-mentioned food restriction-modified effect of
maternal condition on the birth litter size and the total
production costs is one of the most interesting findings of
this study. While in the control mothers both these litter
characteristics increased with the increasing maternal
weight (as predicted by Williams 1979), in the foodrestricted mothers, these characteristics were higher and
weight-independent (Figs. 3b and 4b). Meikle and Thornton
(1995) obtained similar results and suggested that the
positive effect of pre-mating food restriction on female
reproduction, often used for the reproductive enhancement
in some domestic animals (e.g., Beltranena et al. 1991),
might result from an abundant food intake after food
restriction. The ultimate explanation of this effect may be
that the food-restricted mothers increased the production of
cheaper daughters (1) to maximize the number of successful
offspring (as predicted by Myers 1978), (2) to compensate

potential reproductive costs (as predicted by Schaffer 1974;


Hirshfield and Tinkle 1975), or (3) possibly even to
maximize terminal investment (as predicted by CluttonBrock 1984). Moreover, these effects could be strengthened
by using laboratory mice selected for generally high
reproductive output (Rauw et al. 1999).
Since male mouse embryos are more vulnerable than
females to maternal nutritional stress (Krackow and Hoeck
1989), one could expect that the food-restricted mothers
deliver lighter sons than the control mothers. We have not
confirmed this assumption. By contrast, the sons were
heavier than the daughters irrespective of the feeding
treatment (Fig. 5a), indicating that males were the more
expensive sex (as observed by Krackow et al. 2003). As a
consequence of the difference in producing sons and
daughters, the mothers tended to balance between the
secondary sex ratio and the birth litter size, indicating an
adaptive strategy. The control mothers tended to produce
smaller litters with a higher proportion of sons whereas the
food-restricted mothers tended to produce larger litters with
a higher proportion of daughters. The maternal trade-off
between the offspring size and number (as predicted by
Lack 1948; Smith and Fretwell 1974) was not affected by
the feeding treatment (Fig. 5b).
We do not know what mechanism had caused the
female-biased sex ratio of the food-restricted mothers. As

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Fig. 5 The offspring birth


weight depending on the offspring sex a, the birth litter size
b, the maternal delivery weight
c, and the mothers delivery date
d. Bars indicate least-square
means (SE), and each data
point represents a predicted
value of one offspring per group
(control, black circle, n=961;
food-restricted, white circle,
n=865). The best-fit regression
lines are shown

presumed by Meikle and Thornton (1995), the sex-specific


embryonic mortality seems unlikely because the pre-mating
food restriction did not reduce the litter size. A potential
mechanism may be the timing of insemination within the
estrous cycle (Krackow 1995a). Taking into account a
potential adaptive significance of the food restrictiondelayed delivery date, it is possible that the food-restricted
mothers deferred their insemination to favor the implantation
of female blastocysts (Krackow 1995b; Krackow and
Burgoyne 1998). An alternative explanation may be that
the pre-mating food restriction decreased the glucose level

Fig. 6 The pre-weaning offspring mortality depending on the feeding


treatment of the mother (control, n=961; food-restricted, n=865).
Bars indicate proportions of offspring deaths from day 1 to 21

which could favor the survival of female blastocysts


(Cameron 2004; Cameron et al. 2008). On the other hand,
the increase in the proportion of sons with the increasing
maternal weight loss might result from an increase of the
maternal testosterone level in response to the nutritional
stress (Grant 2007) or from an increase of the glucose level
due to the food repletion after the period of food restriction
(Ahima et al. 1996; Cameron 2004; Pecoraro et al. 2004;
Cameron et al. 2008). The above-mentioned timing of
insemination within the estrous cycle, differentially influencing
the survival of male and female blastocysts (Krackow 1995a,
b), could also be a mechanism whereby the control mothers
might adjust the size- and sex-specific composition of the
litter according to their weight. All these mechanisms are not
mutually exclusive and may interact with each other.
It is important to point out that, regardless of the sex, the
offspring of the food-restricted mothers suffered less from
pre-weaning mortality than those of the control mothers
(Fig. 6). These results are all the more interesting because
the effect of pre-mating food restriction persisted until the
end of the lactation period during which both control and
food-restricted mothers were fed ad libitum. Considering
this aspect, it indeed seems that the food-restricted mothers
aimed to produce cheaper daughters to maximize the
number of successful (surviving) offspring (as predicted
by Myers 1978). By contrast, the food restriction-modified
effect of maternal condition on the total production costs
(Fig. 4b) shows that only the control mothers might adjust
the offspring sex ratio along with the size of the litter to

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maximize fitness returns from sons and daughters (as


predicted by Williams 1979). Therefore, when comparing
the groups, the Myers hypothesis (Myers 1978) had
a general significance while the Williams hypothesis
(Williams 1979) was plausible only for the control mothers.
In addition, the results indicate that some of the foodrestricted mothers partly coped with the pre-mating food
restriction and increased the proportion of sons in the litter
with the increasing maternal weight loss (during the period
of food restriction).
To date, a combination of the sex allocation strategies
(conditional sex allocation: Charnov et al. 1981; Jones 1982;
local mate competition: Hamilton 1967) has been reported in
the parasitoid wasps (Lariophagus distinguendus: Werren
and Simbolotti 1989; Heterospilus prosopidis: Ikawa et al.
1993; Laelius pedatus: Mayhew and Godfray 1997).
Another combination of the sex allocation strategies (sensu
Wild and West 2007), conditional sex allocation (Trivers and
Willard 1973) and local resource competition (Clark 1978;
Silk 1983), has been reported in the mouflon (Ovis musimon:
Landete-Castillejos et al. 2001) and the common brushtail
possum (Trichosurus vulpecula: Isaac et al. 2005). In
addition, Allaine et al. (2000) has reported a combination
of the Williams hypothesis (Williams 1979) and the helper
repayment hypothesis (Gowaty and Lennartz 1985; Emlen
et al. 1986; Lessells and Avery 1987) in the Alpine marmot.
To our knowledge, our study is the first to show a
combination of three sex allocation strategies (Myers,
Williams, and maternal coping strategy) in a typical
polytocous mammal, the house mouse. Taking into account
multiple selective forces acting on parental and offspring
fitness (Krackow 2002), we assume that mixed sex
allocation strategies may be widely dispersed and could
potentially explain part of the sex ratio variation across
various taxa.
The house mouse is a typical r-selected species
producing many offspring in an unpredictable environment
(MacArthur and Wilson 1967; Pianka 1970). Therefore, the
optimization of litter composition under stable conditions
and the production of cheaper daughters under changeable
conditions may be envisaged as a part of a more general
strategythe maximization of offspring number. Considering
this aspect, the Myers hypothesis (Myers 1978) and the
Williams hypothesis (Williams 1979), both emphasizing the
offspring quantity, should be implemented to investigate the
variation in offspring sex ratio in polytocous mammals.
Acknowledgments We are grateful to Petr imeek for helpful
comments on the statistical analysis, George A. Bubenik for correcting
the English text, Sven Krackow for a stimulating discussion, and two
anonymous referees for valuable comments improving the final
version of the article. This research was supported by the Czech
Science Foundation (523/03/H076) and by the Ministry of Agriculture
of the Czech Republic (MZE0002701404).

Behav Ecol Sociobiol (2011) 65:22092217


Ethical standards The authors declare that the present study complies
with the current laws of the Czech Republic.

Conflict of interest The authors declare that they have no conflict of


interest.

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