What causes sexual ornaments and behavior (secondary sexual
characteristics): primary are the gonads and reproductive
equipment themselves)?
Peacock’s tail, bird song repertoires, conspicuous color patterns,
sexual displays
This is a conspicuous problem: occurred to Darwin
Two possibilities for sexual selection are (1) male-male competition
(horns etc.) or (2) female choice. (Competition is usually among males,
and choice usually by females, because females usually invest more in
offspring.) We will focus on female choice.
Fisherian “runaway” model (“sexy son”)
- trait initially advantageous
- preference and trait become genetically correlated (trait expressed
in males, preference in females)
- Requirements: heritability of trait and preference. trait
not too expensive (no requirement that the trait should be
expensive)
- Tests/predictions:
- negative or zero correlation between male quality and trait
- genetic correlation between trait and preference (Houde and Endler
compared guppy populations in different streams)
- high variability among populations in ornaments?
Zahavi “handicap” principle
The handicap principle says that traits are not
chosen despite their being costly, but because they are costly.
Good genes model. Ornaments are a signal of good genes:
they must be costly, otherwise you can’t rely on the honesty of the
signal.
- revealing traits: indicate male
condition/quality
- conditional traits: only expressed by high-quality
males
- viability (“Zahavi”) traits: expressed by everyone, but kill off
low-quality males (may not work??)
Penn & Számadó (2020):
Rather than being wasteful over-investments, honest signals evolve in
this scenario because selection favours efficient and optimal investment
into signal expression and minimizes signalling costs … This model is
better understood within a Darwinian framework of adaptive signalling
trade-offs, without the added burden and confusing logic of the Handicap
Principle.
Problem with handicap principle: what maintains genetic variation for
“goodness”? Under normal conditions (directional selection), one would
expect that soon everyone would have good genes and there would be no
variability (nothing for females to choose from except luck). Constant
mutation could lead to a mutation-selection balance, or a
variable environment could keep changing what genes are “good”. Of
course (as we know), parasites represent a strong driver of variability
in the biotic environment.
Hamilton-Zuk hypothesis
(the Red Queen returns)
a subset of ‘good-genes’ models
requirements (Combes
(2005) p. 185)
- females prefer resistant (showy) males (also required by
Fisher)
- correlation between quality (resistance) and trait (not
required by Fisher), or negative correlation between parasite load and
trait
- (genetic) heritable variation in resistance (the hardest one to
prove)
- Parasite load decreases host viability (otherwise there is no point
in selecting resistant males)
- (genetic) heritable variation in trait
Evidence for H-Z: between-species
If H-Z is operating, we expect a positive
correlation between parasite load and showiness across species. This
correlation has been found, for example, in Hamilton and Zuk’s original
study which correlated human judgements of species brightness with
information on ectoparasite load. Other studies have been more
equivocal.
There are some problems with between-species comparisons, though:
- reverse causality: showiness could cause parasitism
(e.g. brighter species could attract more ectoparasites, or species that
invest more in showiness could have less to invest in parasite
resistance)
- ecological correlates: showiness and parasitism
could both be driven by other factors (hole-nesting, polygyny)
- falsifiability: when a correlation is not found, how do you decide
whether the data are really good enough to reject?
- phylogenetic inertia
- dynamics: how do we know that parasite-driven
sexual selection isn’t so effective that it drives parasite loads to
lower levels in showy species?
Evidence for H-Z: within species
Guppies (Martin &
Johnsen, 2007):
- Gyrodactylus turbulli infestation lowers brightness of
parasitized males
- it also reduces the number of sexual displays
- females prefer brighter males
- we don’t know whether resistance is heritable
- we can’t rule out transmission avoidance (see below)
Barn swallows (Møller,
1990):
- females prefer males with longer tails (manipulative experiments,
shortening and lengthening male tails)
- males with longer tails have fewer mites (field observation)
- chicks with heavy mite loads are smaller at fledging, leading to
lower survivorship/fecundity/fitness
- heritable variation in resistance: cross-fostering
offspring of long-tailed males in the nests of short-tailed males (and
vice versa) shows that offspring of long-tailed fathers inherit their
fathers’ low mite loads. This rules out the hypotheses that
- long-tailed males have just avoided parasitism by luck
- females are selecting males for paternal care (direct
benefits) [and offspring are less parasitized, e.g., because
they’re better nourished]
- offspring benefit by avoiding direct transmission of mites from
fathers (also direct benefits)
Alternatives
Alternatives to H-Z: both can be tested (with some difficulty) by
manipulative cross-fostering experiments
- selection for parental care (direct benefits) [can be ruled out in
species without parental care]
- selection for parasite avoidance.
- e.g.: sage grouse & red ink hematomas (Spurrier et al. in Clayton (1991)) [can be ruled out for
non-directly transmitted parasites; stronger evidence for ectoparasites
suggests parasite avoidance]
- selection for female fecundity (pipefish, Combes (2005) p. 180)
Immunocompetence handicap hypothesis
Secondary sexual traits seem to be negatively correlated with
immunosuppression
Folstad & Karter (1992): turning
off your immune system is the ultimate handicap. Thus, it is not a
coincidence that females prefer traits that are linked to (temporary)
immunosuppression; males that can afford to turn off their immune
systems during the mating season must have good genes …
(note switch from original H-Z!)
Good links between androgens and secondary sex characteristics. Some
support for the link between testosterone (and other androgens) and
immunosuppression, but variable results in different experiments.
Immunoredistribution rather than immunosuppression? [Evans et al. (2000);Roberts et al. (2004);
Other axes of signalling and immune function: carotenoid, melanin
systems (McGraw & Ardia, 2003; Simons et al.,
2012)
Stress-linked immunocompetence handicap (Buchanan, 2000) “testosterone has a dual
effect: it leads to immunosuppression through a mechanism involving
corticosterone but, conversely, leads to increased immunocompetence
probably via dominance influencing access to resources”.
possibly explains the mess … (Bortolotti et al., 2009)
Where are we? Balenger & Zuk
(2014)
Recent meta-analyses: (Dougherty et al.,
2023)
Humans: do females prefer male faces associated with lower
cortisol/higher testosterone/better immunity (Rantala et al., 2012)? across countries, do
females prefer ‘masculine’ men more in countries with more parasite
exposure (DeBruine et al., 2012)?
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Last updated: 2023-11-27 12:05:16.287138