278 CHAPTER 11
Costs of reproduction
Is there evidence for a cost of reproduction? This concept plays a large role in the
theory of life history evolution. Genotypes that allocate more to reproduction and
less to themselves may display decreased survival or growth. This allocation trade-off
would be manifested as a negative genetic correlation (see Chapter 6) between repro-
duction and survival. If there were also genetic variation in the amount of resources
individuals acquired from the environment, however, that variation could give rise to
a positive genetic correlation between reproduction and survival (FIGURE 11.3) [7, 59].
Both kinds of correlation were found in a seed beetle (Callosobruchus maculatus) that
develops as a larva within a bean [37]. Reproductive adults may continue to feed, but
females lay eggs even if they are deprived of food. Variation among families showed a
positive genetic correlation if females were given food (hence, variation in acquisition
of resource), but a negative genetic correlation between fecundity and survival—evi-
dence of a cost of reproduction—when females were deprived of food.
Costs of reproduction have been detected in many kinds of organisms. Genetic
correlations found in a wild population of Drosophila melanogaster showed strong
trade-offs between the number of eggs females laid when young and both their
longevity and their fecundity later in life. However, longer-lived genotypes showed
higher fecundity late in life—a rather significant observation, as we will soon see
[58]. In a study of brown anoles (Anolis sagrei), Robert Cox and Ryan Calsbeek sur-
gically removed the ovaries from wild females and then released them [14]. Even
though this species lays only one egg per clutch, these females, prevented from
allocating energy and nutrients to reproduction, showed higher growth and sur-
vival than did sham-operated females with intact ovaries (FIGURE 11.4).
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Survival A B
Resources
1 – x
x
Reproduction
Growth
Maintenance,
survival
Reproduction
(A)
(B)
Resource
acquisition locus
A
Resource
allocation locus
B
FIGURE 11.3 actors giving rise to genetic correlations between life history F
traits such as survival (or growth) and reproduction. (A) Alleles at locus A affect
the amount of energy or other resources that individuals acquire from the
environment. Alleles at locus B affect allocation of resources to functions
such as growth or survival versus reproduction, in proportions x and 1 – x.
(B) Genotypes that differ in their ability to acquire resources (for example, be-
cause of variation at locus A) are represented by blue circles. Genotypes that
differ in how resources are allocated between survival and reproduction (for
example, because of variation at locus B) are shown by red circles. The overall
genetic correlation between survival and reproduction depends on the rela-
tive magnitude of variation in resource acquisition versus resource allocation.
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Change in size (SVL in mm)
Change in mass (g)
SVL Mass
0
1.0
0.5
1.5
2.0
3.0
2.5
3.5
0
0.10
0.05
0.15
0.20
0.30
0.25
0.35
(A)
Measures of growth
Survival
2007 2008 Both
0
0.2
0.1
0.3
(B)
Year of study
SHAM
OVX
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Evolution4e_11.03.ai Date 11-20-2016
Survival A B
Resources
1 – x
x
Reproduction
Growth
Maintenance,
survival
Reproduction
(A)
(B)
Resource
acquisition locus
A
Resource
allocation locus
B
FIGURE 11.4 Evidence of the cost of
reproduction. (A) Female Anolis sagrei
from which ovaries were removed (OVX)
grew larger and gained more weight
than sham-operated (SHAM) females. SVL
is the snout-to-vent length. (B) Over the
2-year study, the proportion of females that
survived to the following year was higher
for ovariectomized females (blue columns)
than for sham-operated females (red
columns), which produced eggs. Allocation
to reproduction reduced females’ growth
and survival. (From [14].)
11_EVOL4E_CH11.indd 278 3/22/17 1:11 PM