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“A gentle immunological balance thus has to be
maintained in the decidua, where immunological activity
operates to eliminate a pathogen without damaging
the fetus”Markel et al. (2002)Journal of Clinical Investigation 110: 943
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“The border zone … is not a sharp line,
for it is in truth the fighting line where
the conflict between the maternal cells
and the invading trophoderm takes
place, and it is strewn with such of the
dead on both sides as have not already
been carried off the field or otherwise
disposed of.”
Johnstone (May 1914)Journal of Obstetrics and Gynaecology of the British Empire 25: 231
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Maternal provisioning of a fetus is associated
with an opportunity cost
The opportunity cost translates into lower
expected fitness through other offspring
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If extra resources are transferred to an
embryothe embryo’s expected
fitness increasesthe mother’s expected
fitness from other offspring decreases
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maternal investment in fetus
benefit
ben
efi
t to
fetu
s cost cost to
sib
ling
s
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X minimizes cost to siblings
benefit
ben
efi
t to
fetu
s cost cost to
sib
ling
s
X
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Z maximizes benefit to fetus
benefit
ben
efi
t to
fetu
s cost cost to
sib
ling
s
X Z
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Y maximizes (benefit — cost)
benefit
ben
efi
t to
fetu
s cost cost to
sib
ling
s
X ZY
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maternal(non-inherited)
maternal(inherited)
paternal(inherited)
mother
fetus
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Relative shares
p = probability of shared paternity
geneBenefit
(to fetus)Cost
(to sibs)
maternal(non-inherited) 0 1/2
maternal(inherited) 1 1/2
paternal(inherited)
1 p/2
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A non-inherited maternal gene gains no benefit from the
survival and reproduction of a
fetus
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worse than that!
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Non-inherited maternal genes will
benefit from the early demise of the fetus
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How is pregnancy possible?
rarity of genetic self-recognition
“the parliament of the genes”(mutual policing)
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Paternally-derived genes in fetuses favor greater demands on
mothers than maternally-derived
genes
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PRONUCLEAR SUBSTITUTIONS
egg nucleus sperm nucleus
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QuickTime™ and aPhoto - JPEG decompressor
are needed to see this picture.
fetus
yolk sac
trophoblast
mum+
dad
mum+
mum
dad+
dad
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46,XX paternal origin
massively proliferating placental tissues
1,000-fold increased risk of choriocarcinoma
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46,XX maternal origin
ovarian teratomas; benignproduce most tissues (but not
placenta)
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maternal(non-inherited)
maternal(inherited)
paternal(inherited)
mother
fetus
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incomplete information
p = probability of shared paternity
Benefit(to fetus)
Cost(to sibs)
mother 1/2 1/2
fetus 1 (1+p)/4
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umbilical cord
spiral artery
uterine vein
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Conflict can exist over
whether or not to miscarry
the nutrient quality of maternal blood
the volume of blood reaching the placenta
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ovulation(day 0)
hCG (day 7)
onset of menstruation(day 14)
CL regresses(days 8-10)
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women attempting to conceive
number of cycles
chemical pregnancies
clinical pregnancies
term pregnancies
707
198
155
136
data from Wilcox et al. (1988)
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anterior pituitary
corpus luteum
uterus
luteinizinghormone
progesterone
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anterior pituitary
corpus luteum
uterus
placenta
luteinizinghormone
progesterone
chorionicgonadotropin
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anterior pituitary
corpus luteum
uterus
placenta
luteinizinghormone
progesterone
chorionicgonadotropin
progesterone
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CONCENTRATIONS IN MATERNAL SERUM
non-pregnant pregnant
hLH/hCG
hGH/hPL
progesterone
estradiol
100 mIU/ml
5 ng/ml
10 ng/ml
0.4 ng/ml
50,000 mIU/ml
10,000 ng/ml
200 ng/ml
20 ng/ml
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Placental hormonesWhy shout?
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Placental hormones originate as fetal
attempts to manipulate maternal physiology for
fetal benefit
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Placental hormones may evolve to become
little more than endocrine SPAM
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maternal carbohydrate metabolism
• fasting blood glucose falls in first trimester
• maternal sensitivity to insulin decreases as pregnancy progresses
• maternal insulin production increases in parallel with reduced sensitivity
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MATERNAL CIRCULATION DURING PREGNANCYplacentamaternaltissues
RpRm
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maternal blood pressure in pregnancy
• blood pressure reduced during most pregnancies; rises toward term
• ≈ 10% women develop hypertension= pregnancy-induced hypertension
(PIH)
• preeclampsia (PIH + proteinuria) affects ≈ 3% pregnancies
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Placental factors
Maternal factors
Uteroplacentalresistance
decrease
increase
Non-placentalresistance
increase
decrease
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MATERNAL CIRCULATION DURING PREGNANCYplacentamaternaltissues
RpRm
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Maternal-fetal relations lack important
feedback controls because signals are not evolutionarily credible
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non-pregnant mothers of sons
time since birthof last son
XY cellsin blood
data from Bianchi et al. (1996)
6 months10 months12 months
2 yrs3 yrs6 yrs7 yrs27 yrs
nonoyesyesyesyesyesyes
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PRONUCLEAR SUBSTITUTIONS
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data of E. B. Keverne
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data of E. B. Keverne
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Androgenetic/normal chimeras have large
bodies with relatively small brains
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Gynogenetic/normal chimeras have small bodies with relatively
large brains
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photos from E. B. Keverne
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Contribution to brains of chimeric mice
hypothalamusneocortex
“two dads” + + + —
“two mums” + + +—
Keverne et al. (1996)Developmental Brain Research 92: 91
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Genomic imprinting concerns differences between genomes of
maternal and paternal origin, not differences
between males and females
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