Sex ratio distortion

Joe Ironside
 Strategies to increase
cytoplasmic transmission
• Increase fitness of female offspring

X
– Enhance metabolism
– Reduce sibling rivalry by killing males
• Increase proportion of female
offspring
– Convert males or hermaphrodites into
females
• Kill offspring of unrelated females
– Cytoplasmic incompatibility

X
 Parasitic sex determination

• Parthenogenesis
– Caused by Wolbachia
in hymenopterans and
mites
• Feminization
– Caused by
• Wolbachia and
Microsporidia in
crustaceans
• Male-sterility
– Caused by
mitochondria in plants
 Thelygeny in crustaceans
• Thelygenic females
– Produce only (or mainly) female offspring
– Associated with parasitic infection
• Gammarus duebeni
– aquatic amphipod
– Infected with two microsporidian parasites
• Orchestia gammarellus
– semi-terrestrial amphipod
– Infected with one paramyxid parasites
• Armadillidium vulgare
– terrestrial isopod
– Infected with one bacterial parasite
– No reduction in brood size
• Feminisation rather than male-killing
 Why feminise?
• Feminising parasites are
vertically transmitted
– From females to offspring
– Never from males to
offspring
• Infected males are a dead
end
• Increasing proportion of
infected hosts that are
female
– Increases vertical parasite
transmission
 Why feminise?
• Vertically transmitted
parasites
– Number of infected female
offspring produced by
infected females
• Must be greater than
– Number of uninfected female
offspring produced by all
females
• Vertical transmission
– Can fail
• Feminisation
– Can fail
 Question
• What is the impact of feminising parasites
on sex ratio evolution?
– What is the optimal sex ratio for uninfected
females to produce?
 Impact on sex ratio evolution
• Assuming additive genetic variation
for sex ratio
– E.g. G. duebeni
• As feminising parasite increases in
prevalence
– Males become less common than
females
• Total reproductive output of males remains
equal to that of females
– Male offspring are fitter than female offspring

• Natural selection
– Alleles for male sex determination
• increase in frequency
– Alleles for female sex determination
• decrease in frequency
Hatcher and Dunn 1995
• Predicted outcome
– All uninfected individuals are male
– All infected individuals are female
• Not observed in nature
 Question
• What happens to sex chromosomes in the
presence of a feminising parasite?
 Impact on X and
Y chromosomes
• Male-heterogametic (XY) sex
determination
– E.g. Orchestia gammarellus
• Parasite Paramarteilia orchestiae
– overrides Y chromosome
• Results in XY females
• XY♀ x XY♂ = XX, XY and YY offspring
– Uninfected XX♀, XY♂, YY♂
– Infected XX♀, XY♀, YY♀
• YY females
– All uninfected offspring are male
• Fittest genotype at high parasite prevalence
• Predicted outcome
– X chromosome should go extinct
• All females are infected YY
• All males are uninfected YY
– Supported by data from O. gammarellus
populations
 Impact on Z and W
chromosomes
• Female-heterogametic (ZW) sex determination
– E.g. Armadillidium vulgare
• Wolbachia parasite
– overrides chromosomal sex determination
• Results in ZZ females
• ZZ♀ x ZZ♂ = only ZZ offspring
– Uninfected ZZ♀, ZZ♂
– Infected ZZ♀
• ZZ females
– All uninfected offspring are male
• Fitter than ZW at high parasite prevalence
• Predicted outcome
– W chromosome should go extinct
• All females are infected ZZ
• All males are uninfected ZZ
– Supported by data from A. vulgare populations
 Horizontal gene transfer
• f-lineages of A. vulgare
– ZZ genotype
– Demonstrate thelygeny
• few male offspring
– Not infected with
Wolbachia
• Wolbachia genome
– Integrated into host
chromosome
• Forms new sex chromosome Leclercq et al. 2016
 Question
• What is the impact of the f-element on sex
ratio evolution?
– What is the optimal sex ratio for females to
produce?
 The M gene
• Dominant autosomal gene
• Overrides f-element
– Restores male sex determination
 Male-killing
• Males killed to
prevent competition
with infected female
siblings
– Caused by
Wolbachia in
butterflies
– Caused by
• Wolbachia, Rickettsia
and Spiroplasma in
ladybirds
 Cytoplasmic male sterility

• Increase proportion
of female offspring
– Male-sterility
• Converts
hermaphrodites into
females
– Less pollen enables
more seed to be
produced
 Cytoplasmic incompatibility

• Sperm of infected males

is sabotaged
– If female is uninfected
• Paternal genome self-
destructs after fertilisation
– If female is infected
• Paternal genome is rescued
• Caused by Wolbachia in
insects