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Why Maternal Favouritism Leads
to the Battle of the Sexes?
Weihao Zhong
Nicholas Priest
Talk Outline
• Sexual Conflict
• Maternal Effects & Sex Allocation
• An Intralocus Population Genetic Model
• Implications for Sexual Conflict
Photo credits, clockwise from top left: Richard Naylor, Andrew Syred , Kamimura (2007) & Rezác (2009).
Mendelian Inheritance
Phenotype,
Z
Genetic
Component, G
Environmental
Component, E
Maternal effects
Phenotype,
ZO
Genetic
Component, GO
Environmental
Component, EO
Phenotype,
ZM
Offspring Generation
Cross-Generational Effects on Inheritance
Maternal Generation Offspring Generation
Maternal effects
Phenotype,
ZO
Genetic
Component, GO
Environmental
Component, EO
Phenotype,
ZM
Genetic
Component, GM
Environmental
Component, EM
Cross-Generational Effects on Inheritance
Maternal Generation Offspring Generation
Maternal effects
Phenotype,
ZO
Genetic
Component, GO
Environmental
Component, EO
Phenotype,
ZM
Genetic
Component, GM
Environmental
Component, EM
Cross-Generational Effects on Inheritance
Sexual conflict theory needs to consider the role of
maternal effects.
Sex-Specific Maternal Effects
• Maternal effects often depend on offspring sex
– Egg-laying order & offspring growth in birds (Badyaev
2006)
– High early fertility of daughters from promiscuous mothers
in Drosophila (Priest et al. 2008)
Sex-Specific Maternal Effects
• Maternal effects often depend on offspring sex
– Egg-laying order & offspring growth in birds (Badyaev
2006)
– High early fertility of daughters from promiscuous mothers
in Drosophila (Priest et al. 2008)
• Sex allocation theories predict sex-specific maternal
favouritism (Trivers & Willard 1973; Charnov 1982)
– Sex ratio adjustment
– Offspring quality adjustment
Can sex-specific maternal
favouritism drive the evolution of
sexual conflict?
Population Genetic Model of Sex Allocation
• Assumptions:
– One locus with 2 alleles: A1 A2, with frequencies of p1 & p2
– Either directly or maternally expressed
– Sex-specific fitness with random mating in infinite
population
Population Genetic Model of Sex Allocation
A1A1 A1A2 A2A1 A2A2
A1A1 p13 p12p2 0 0
A1A2 p12p2
2
p1p22
2
p12p2
2
p1p22
2
A2A1 p12p2
2
p1p22
2
p12p2
2
p1p22
2
A2A2 0 0 p1p22 p23
Offspring Genotype
Mother
Genotype
Population Genetic Model of Sex Allocation
A1A1 A1A2 A2A1 A2A2
A1A1 p13 p12p2 0 0
A1A2 p12p2
2
p1p22
2
p12p2
2
p1p22
2
A2A1 p12p2
2
p1p22
2
p12p2
2
p1p22
2
A2A2 0 0 p1p22 p23
Offspring Genotype
Mother
Genotype
+SAdf
/
+SAdm
+SDdf
/
+SDdm
+SDdf
/
+SDdm
- SAdf
/
- SAdm
Offspring
Fitness Effects
Population Genetic Model of Sex Allocation
A1A1 A1A2 A2A1 A2A2
A1A1 p13 p12p2 0 0
A1A2 p12p2
2
p1p22
2
p12p2
2
p1p22
2
A2A1 p12p2
2
p1p22
2
p12p2
2
p1p22
2
A2A2 0 0 p1p22 p23
Offspring Genotype
Mother
Genotype
+SDmf /+SDmm
+SDmf /+SDmm
- SAmf /- SAmm
Maternal Fitness
Effects
+SAmf /+SAmm
Population Genetic Model of Sex Allocation
A1A1 A1A2 A2A1 A2A2
A1A1 p13 p12p2 0 0
A1A2 p12p2
2
p1p22
2
p12p2
2
p1p22
2
A2A1 p12p2
2
p1p22
2
p12p2
2
p1p22
2
A2A2 0 0 p1p22 p23
Offspring Genotype
Mother
Genotype
+SAmf
+SAdf
wA1A1f =1 +SAdf + SAmf
Population Genetic Model of Sex Allocation
• Fitness
Population Genetic Model of Sex Allocation
• Fitness
Population Genetic Model of Sex Allocation
• Fitness
• Evolution
Population Genetic Model of Sex Allocation
• Fitness
• Evolution
Maternal and direct effects contribute equally to fitness.
But, maternal effects contribute only half as much to evolution.
Scenarios of Conflict Evolution
• Assuming conflicting alleles, can sex-specific
maternal effects increase the intensity of sexual
conflict?
1. Additive Genetic Effects
2. Dominance Genetic Effects
• Assuming no allelic conflict, can sex-specific maternal
effects generate sexual conflict?
3. Additive + Dominance Genetic Effects
Additive Intralocus Sexual Conflict
A1A1 A1A2 A2A1 A2A2
Male (Direct) +SAdm 1 1 - SAdm
Female (Direct) - SAdf 1 1 +SAdf
A1A1 A1A2 A2A1 A2A2
Male (Direct) +SAdm 1 1 - SAdm
Female (Maternal) - SAmf 1 1 +SAmf
Additive Intralocus Sexual Conflict
♀
♂
Additive Intralocus Sexual Conflict
♀
♂
Additive Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♀
♂
Additive Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♀
♂
♀
♂
♂+♀
Additive Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
No net change under
Mendelian inheritance.
No selection for conflict
Additive Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♀
♂ No net change under
Mendelian inheritance.
No selection for conflict
Additive Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♀
♂
♂+♀
No net change under
Mendelian inheritance.
No selection for conflict
But, evolution towards
maximum sexual
conflict with maternal
effects.
Dominance Intralocus Sexual Conflict
A1A1 A1A2 A2A1 A2A2
Male (Direct) 1 +SDdm +SDdm 1
Female (Direct) 1 -SDdf -SDdf 1
A1A1 A1A2 A2A1 A2A2
Male (Direct) 1 +SDdm +SDdm 1
Female (Maternal) 1 -SDmf -SDmF 1
Dominance Intralocus Sexual Conflict
♀
♂
Dominance Intralocus Sexual Conflict
♀
♂
Dominance Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♀
♂
Dominance Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♀
♂
Dominance Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♂+♀
♀
♂
No net change under
Mendelian inheritance.
No selection for conflict
Dominance Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♀
♂
No net change under
Mendelian inheritance.
No selection for conflict
Dominance Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=25 generations
♂+♀
♀
♂
No net change under
Mendelian inheritance.
No selection for conflict
But, evolution towards
maximum sexual
conflict with maternal
effects.
Add + Dom Intralocus Sexual Conflict
A1A1 A1A2 A2A1 A2A2
Male (Direct) +SAdm +SDdm +SDdm -SAdm
Female (Direct) +SAdf +SDdf +SDdf -SAdf
A1A1 A1A2 A2A1 A2A2
Male (Direct) +SAdm +SDdm +SDdm -SAdm
Female (Direct +
Maternal)
+ SAmf +SDdf +SDdF -SAmf
Add + Dom Intralocus Sexual Conflict
♀
♂
Add + Dom Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=75 generations
♀
♂
*
Identical equilibria for
the sexes under
Mendelian inheritance.
No conflict evolves.
Add + Dom Intralocus Sexual Conflict
Arrow=25 generations
Space between arrows=75 generations
♀
♂
* *
Identical equilibria for
the sexes under
Mendelian inheritance.
No conflict evolves.
But, maternal effects
drives lower equilibrium
for females.
Add + Dom Intralocus Sexual Conflict
♂+♀
♀
♂
* * *
Arrow=25 generations
Space between arrows=75 generations
Identical equilibria for
the sexes under
Mendelian inheritance.
No conflict evolves.
But, maternal effects
drives lower equilibrium
for females.
Population equilibrium
is sub-optimal for either
sex. Conflict evolves.
Implications
• Sex-specific maternal allocation can drive the
evolution of sexual conflict without invoking
adaptation
Implications
• Sex-specific maternal allocation can drive the
evolution of sexual conflict without invoking
adaptation
• Conflict results whenever slight imbalance in
maternal allocation exists between the sexes
Implications
• Sex-specific maternal allocation can drive the
evolution of sexual conflict without invoking
adaptation
• Conflict results whenever slight imbalance in
maternal allocation exists between the sexes
• A simple and general mechanism of conflict
evolution with no assumptions of sex linkage
Acknowledgements
Jason WolfNick Priest
Maternal Effect Gene Evolution
Maternally expressed gene
Directly expressed gene
Both genes have additive
genetic effect of 0.1

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Zhong Liverpool Jan 2010 Talk [FINAL]

  • 1. Why Maternal Favouritism Leads to the Battle of the Sexes? Weihao Zhong Nicholas Priest
  • 2. Talk Outline • Sexual Conflict • Maternal Effects & Sex Allocation • An Intralocus Population Genetic Model • Implications for Sexual Conflict
  • 3. Photo credits, clockwise from top left: Richard Naylor, Andrew Syred , Kamimura (2007) & Rezác (2009).
  • 5. Maternal effects Phenotype, ZO Genetic Component, GO Environmental Component, EO Phenotype, ZM Offspring Generation Cross-Generational Effects on Inheritance
  • 6. Maternal Generation Offspring Generation Maternal effects Phenotype, ZO Genetic Component, GO Environmental Component, EO Phenotype, ZM Genetic Component, GM Environmental Component, EM Cross-Generational Effects on Inheritance
  • 7. Maternal Generation Offspring Generation Maternal effects Phenotype, ZO Genetic Component, GO Environmental Component, EO Phenotype, ZM Genetic Component, GM Environmental Component, EM Cross-Generational Effects on Inheritance Sexual conflict theory needs to consider the role of maternal effects.
  • 8. Sex-Specific Maternal Effects • Maternal effects often depend on offspring sex – Egg-laying order & offspring growth in birds (Badyaev 2006) – High early fertility of daughters from promiscuous mothers in Drosophila (Priest et al. 2008)
  • 9. Sex-Specific Maternal Effects • Maternal effects often depend on offspring sex – Egg-laying order & offspring growth in birds (Badyaev 2006) – High early fertility of daughters from promiscuous mothers in Drosophila (Priest et al. 2008) • Sex allocation theories predict sex-specific maternal favouritism (Trivers & Willard 1973; Charnov 1982) – Sex ratio adjustment – Offspring quality adjustment
  • 10. Can sex-specific maternal favouritism drive the evolution of sexual conflict?
  • 11. Population Genetic Model of Sex Allocation • Assumptions: – One locus with 2 alleles: A1 A2, with frequencies of p1 & p2 – Either directly or maternally expressed – Sex-specific fitness with random mating in infinite population
  • 12. Population Genetic Model of Sex Allocation A1A1 A1A2 A2A1 A2A2 A1A1 p13 p12p2 0 0 A1A2 p12p2 2 p1p22 2 p12p2 2 p1p22 2 A2A1 p12p2 2 p1p22 2 p12p2 2 p1p22 2 A2A2 0 0 p1p22 p23 Offspring Genotype Mother Genotype
  • 13. Population Genetic Model of Sex Allocation A1A1 A1A2 A2A1 A2A2 A1A1 p13 p12p2 0 0 A1A2 p12p2 2 p1p22 2 p12p2 2 p1p22 2 A2A1 p12p2 2 p1p22 2 p12p2 2 p1p22 2 A2A2 0 0 p1p22 p23 Offspring Genotype Mother Genotype +SAdf / +SAdm +SDdf / +SDdm +SDdf / +SDdm - SAdf / - SAdm Offspring Fitness Effects
  • 14. Population Genetic Model of Sex Allocation A1A1 A1A2 A2A1 A2A2 A1A1 p13 p12p2 0 0 A1A2 p12p2 2 p1p22 2 p12p2 2 p1p22 2 A2A1 p12p2 2 p1p22 2 p12p2 2 p1p22 2 A2A2 0 0 p1p22 p23 Offspring Genotype Mother Genotype +SDmf /+SDmm +SDmf /+SDmm - SAmf /- SAmm Maternal Fitness Effects +SAmf /+SAmm
  • 15. Population Genetic Model of Sex Allocation A1A1 A1A2 A2A1 A2A2 A1A1 p13 p12p2 0 0 A1A2 p12p2 2 p1p22 2 p12p2 2 p1p22 2 A2A1 p12p2 2 p1p22 2 p12p2 2 p1p22 2 A2A2 0 0 p1p22 p23 Offspring Genotype Mother Genotype +SAmf +SAdf wA1A1f =1 +SAdf + SAmf
  • 16. Population Genetic Model of Sex Allocation • Fitness
  • 17. Population Genetic Model of Sex Allocation • Fitness
  • 18. Population Genetic Model of Sex Allocation • Fitness • Evolution
  • 19. Population Genetic Model of Sex Allocation • Fitness • Evolution Maternal and direct effects contribute equally to fitness. But, maternal effects contribute only half as much to evolution.
  • 20. Scenarios of Conflict Evolution • Assuming conflicting alleles, can sex-specific maternal effects increase the intensity of sexual conflict? 1. Additive Genetic Effects 2. Dominance Genetic Effects • Assuming no allelic conflict, can sex-specific maternal effects generate sexual conflict? 3. Additive + Dominance Genetic Effects
  • 21. Additive Intralocus Sexual Conflict A1A1 A1A2 A2A1 A2A2 Male (Direct) +SAdm 1 1 - SAdm Female (Direct) - SAdf 1 1 +SAdf A1A1 A1A2 A2A1 A2A2 Male (Direct) +SAdm 1 1 - SAdm Female (Maternal) - SAmf 1 1 +SAmf
  • 22. Additive Intralocus Sexual Conflict ♀ ♂
  • 23. Additive Intralocus Sexual Conflict ♀ ♂
  • 24. Additive Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♀ ♂
  • 25. Additive Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♀ ♂
  • 26. ♀ ♂ ♂+♀ Additive Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations No net change under Mendelian inheritance. No selection for conflict
  • 27. Additive Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♀ ♂ No net change under Mendelian inheritance. No selection for conflict
  • 28. Additive Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♀ ♂ ♂+♀ No net change under Mendelian inheritance. No selection for conflict But, evolution towards maximum sexual conflict with maternal effects.
  • 29. Dominance Intralocus Sexual Conflict A1A1 A1A2 A2A1 A2A2 Male (Direct) 1 +SDdm +SDdm 1 Female (Direct) 1 -SDdf -SDdf 1 A1A1 A1A2 A2A1 A2A2 Male (Direct) 1 +SDdm +SDdm 1 Female (Maternal) 1 -SDmf -SDmF 1
  • 30. Dominance Intralocus Sexual Conflict ♀ ♂
  • 31. Dominance Intralocus Sexual Conflict ♀ ♂
  • 32. Dominance Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♀ ♂
  • 33. Dominance Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♀ ♂
  • 34. Dominance Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♂+♀ ♀ ♂ No net change under Mendelian inheritance. No selection for conflict
  • 35. Dominance Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♀ ♂ No net change under Mendelian inheritance. No selection for conflict
  • 36. Dominance Intralocus Sexual Conflict Arrow=25 generations Space between arrows=25 generations ♂+♀ ♀ ♂ No net change under Mendelian inheritance. No selection for conflict But, evolution towards maximum sexual conflict with maternal effects.
  • 37. Add + Dom Intralocus Sexual Conflict A1A1 A1A2 A2A1 A2A2 Male (Direct) +SAdm +SDdm +SDdm -SAdm Female (Direct) +SAdf +SDdf +SDdf -SAdf A1A1 A1A2 A2A1 A2A2 Male (Direct) +SAdm +SDdm +SDdm -SAdm Female (Direct + Maternal) + SAmf +SDdf +SDdF -SAmf
  • 38. Add + Dom Intralocus Sexual Conflict ♀ ♂
  • 39. Add + Dom Intralocus Sexual Conflict Arrow=25 generations Space between arrows=75 generations ♀ ♂ * Identical equilibria for the sexes under Mendelian inheritance. No conflict evolves.
  • 40. Add + Dom Intralocus Sexual Conflict Arrow=25 generations Space between arrows=75 generations ♀ ♂ * * Identical equilibria for the sexes under Mendelian inheritance. No conflict evolves. But, maternal effects drives lower equilibrium for females.
  • 41. Add + Dom Intralocus Sexual Conflict ♂+♀ ♀ ♂ * * * Arrow=25 generations Space between arrows=75 generations Identical equilibria for the sexes under Mendelian inheritance. No conflict evolves. But, maternal effects drives lower equilibrium for females. Population equilibrium is sub-optimal for either sex. Conflict evolves.
  • 42. Implications • Sex-specific maternal allocation can drive the evolution of sexual conflict without invoking adaptation
  • 43. Implications • Sex-specific maternal allocation can drive the evolution of sexual conflict without invoking adaptation • Conflict results whenever slight imbalance in maternal allocation exists between the sexes
  • 44. Implications • Sex-specific maternal allocation can drive the evolution of sexual conflict without invoking adaptation • Conflict results whenever slight imbalance in maternal allocation exists between the sexes • A simple and general mechanism of conflict evolution with no assumptions of sex linkage
  • 46. Maternal Effect Gene Evolution Maternally expressed gene Directly expressed gene Both genes have additive genetic effect of 0.1