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Quantitative Genetics
• Up until now, we have dealt with characters (actually genotypes) controlled by a single locus, with only two alleles:
Discrete Variation
Many Traits are Polygenic
• continuous and quantitative refer to variation, polygenic refers to the mode of inheritance, ("many genes")
Quantitative Variationor Continuous variation
Studying Quantitative Traits
• It would be impossibly difficult to use the same approach as population genetics to consider inheritance at many many loci, especially if the number of loci is unknown
• We need to look at DISTRIBUTIONS of characters rather than frequencies of alleles
Characterizing a Quantitative Trait#
of i
nd
ivid
ua
ls
Z
Mean (average)
( )2
iz z
N
Σ −=
z=
Variance in Phenotype (VP) (mean squared deviation from mean)
(Phenotype)
What Causes Phenotypic Variation Among Individuals
Z
# o
f in
div
idu
als Genetics?
Environment?
Both?
P G E G x EV V V V= + +
Partitioning Variance
Total Phenotypic Variance (VP)
VG VE VG x E
VADDVEPI
Genetic Variance can be subdivided:
VADD= phenotypic variation due to the additive effects of alleles
VDOM = phenotypic variation due to dominance effects (when the effect of the allele depends on the identity of the other allele at that locus)
VEPI = phenotypic variation due to epistatic effects (when the effect of the allele depends on the identity of alleles at different loci)
VDOM
Additive Genetic Variation
VADD= phenotypic variation due to the additive effects of alleles
Consider a gene with 2 alleles, A1 and A2:
A1 A1 A1 A2 A2 A2 a d
Additive effects only
10 8 6
W/ dominance 10 10 6
How much does each additional copy of A1 add to phenotype?
Which of these can be passed on to offspring?
Partitioning Variance
Total Phenotypic Variance (VP)
VG VE VG x E
Environmental Variance can be subdivided:
VEN V= phenotypic variation due to random environmental influences
VCOM = phenotypic variation due to common family influences
VMAT = phenotypic variation due to maternal influences
VENV VCOM VMAT
Maternal Environment Effect in Guppy Offspring Size
Food stressed mothers produce larger offspring
0.9
1
1.1
1.2
1.3
1.4
Low High
Maternal Food Level
Offspring Size (mg)
Reznick and Yang 1993
Breaking the Stick of Variation
• By breaking the stick of variation, we can determine how much of the phenotypic variation is due to each component.
• Selection acts on phenotypic variation, but can only cause evolution if the variation is heritable
• Broad-sense heritability: H2 = VG/VP
• Narrow-sense heritability: h2 = VA/VP
Partitioning Variance
Total Phenotypic Variance (VP)
VG VE VG x E
VDOM VEPIVADDVENV VCOM VMAT VG x E
heritability (h2) = the proportion of phenotypic variation that is due to the additive effects of alleles [how much of VP is made up by VADD]
Total Phenotypic Variance (VP)VADD
Why only Additive Genetic Variance?The additive effects of alleles are responsible for the
degree of similarity between parents and offspring
Additive effects
a = the effect of substituting an A1 or A2 allele
Why is there spread around the phenotypic values of 6, 8, and 10 for each genotype?
VE
Dominant A2
A2A2 A1A2 A1A1 a dADD only 10 8 6 2 0w/ DOM 10 10 6 2 2
Why only Additive Genetic Variance?The additive effects of alleles are responsible for the
degree of similarity between parents and offspring
Additive effects Dominant A2
A1A2 x A1A2
Parents = 8 Parents = 10
Offspring = .25(6)+.5(8)+.25(10) = 8 Offspring = .25(6)+.5(10)+.25(10) = 9
Dominance causes offspring phenotype to deviate from parental phenotype!
So, What is Heritability?Heritability describes the proportion of variation in trait
that can respond to selection
Broad-sense Heritability (H2 = h2B = VG/VP)
– could include dominance and epistatic variation
Narrow-sense Heritability (h2= VA/VP)– proportion of phenotypic variance that is due to
additive genetic causes
Measuring Heritability
Heritability is the slope of the regression between offspring and mid-parent phenotype
Mid-parent phenotypic trait value
Off
sprin
g ph
enot
ypic
tra
it va
lue
Slope = 0.89
h2= 0.89
covariance between parent & offspringvariance of parentsslope =
Can look at other relatives too!
Slope(mom,daughter) = ½ h2
Slope(half-sibs) = ¼ h2
Meaning of Heritability
• Evolution by natural selection can only occur in pops A & B• h2=0 in pop C--> none of the variation is due to VA
• h2 is undefined, there is no variation
Notes about h2
1) Heritability is NOT THE PROBABILITY A TRAIT IS INHERITED OR THE PROBABILITY A TRAIT HAS A GENETIC BASIS
2) Estimates of heritability are specific to the population in which they are measured.
3) heritabilities are statements about variance, not means (e.g., the number of eyes in humans has a 0 heritability, but this doesn't mean that eye number is not under genetic control)
4) high heritability doesn't mean environment doesn't matter or, vice versa, low heritability doesn't mean genes aren't important.
Total Phenotypic Variance (VP)VADD
Total Phenotypic Variance (VP)VADD