# Quantitative Genetics (16 pages)

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1999 2000 Gregory Carey Chapter 19 Advanced Topics 1 Quantitative Genetics II Advanced Topics In this section mathematical models are developed for the computation of different types of genetic variance Several substantive points about genetic variance components and their effect on the analysis of behavioral data are also made The reader uninterested in the mathematics can read the two text boxes to gain the substantive conclusions Quantitative geneticists partition total genetic variance into three types additive dominance and epistatic variance Additive genetic variance measures the extent to which phenotypic individual differences are predictable from the additive effects of allelic substitutions Dominance genetic variance is variance associated with dominant gene action the fact that the genetic value for a heterozygote is not exactly the average of the genetic value of the two homozygotes Epistatic genetic variance is the variance associated with the statistical interaction among loci gene by gene interaction as it is often called Additive and dominance variance may be illustrated by examining a single locus designated here as M locus with two alleles M1 and M2 The additive effect of allele M2 is the average change in genotypic values seen by substituting an M2 allele for an M1 allele To find this effect simply construct a new variable called X1 here that equals the number of M2 alleles for the individual s genotype For genotype M1M1 X1 0 for M1M2 X1 1 and for M2M2 X1 2 To account for dominance construct another new variable X2 with values of 0 1 and 0 for respectively genotypes M1M1 M1M2 and M2M2 Table 18A 1 provides hypothetical data set up with the new variables1 It is assumed that the phenotype is scaled so that the population mean is 100 and the population standard deviation is 15 Insert Table 18A 1 here 1 This coding system can be used to account for any number of alleles at a locus For example to model additive effects of allele M3 construct another variable giving the number of M3 alleles in a genotype There would then be three dominance variables one for the heterozygote M1M2 a second for the 1999 2000 Gregory Carey Chapter 19 Advanced Topics 2 If we had actual data on individuals we would calculate the additive and dominance effects and their variance components by performing two regression In the first we would regress the phenotypic score noted as Y herein on X1 The squared multiple correlation 2 from this regression equals the additive heritability hA the proportion of phenotypic variance associated with additive gene action at this locus For the data in Table 18A 1 R2 2 137 hA so 13 7 of phenotypic variance is predicted from additive gene effects at this locus The regression line for this equation will be the strait line of best fit that through the three genotypic values for the three genotypes It is illustrated in Figure 18A 1 The second regression model would be of the form Y b0 b1 X 1 b2 X 2 The intercept b0 will equal the genotypic value for M1M1 The regression coefficient b1 equals the average effect of substituting allele M2 for M1 in any genotype Finally the coefficient b2 equals the genotypic value of the heterozygote less the average of the genotypic values of the two homozygotes This measures dominant gene action For the present example b0 94 b1 7 and b2 5 The value of coefficient b1 informs us that on average one M2 allele increases phenotypic values by 7 units Because the value of b2 is not 0 we can conclude that there is some degree of dominance Because b2 5 we can conclude that there is partial dominance for allele M1 so that the genotypic value of the heterozygote is moved 5 units away from the midpoint of the two homozygotes and toward genotype M1M1 The multiple correlation from this model equals the additive heritability plus the 2 2 dominance heritability hA hD For the present example R2 162 Dominance heritability can be found by subtracting the R2 from the first regression model from this 2 value hD 162 137 025 heterozygote M1M3 and the third for the heterozygote M2M3 For each of the three dominance variables the 1999 2000 Gregory Carey Chapter 19 Advanced Topics 3 The regression coefficients can also be used to calculate additive and dominance heritability Let p1 denote the frequency of allele M1 and p2 the frequency of M2 and let V denote the variance of the phenotype which would be 225 for this example Then hA2 2 p1 p2 b1 p1 p2 b2 2 V and hD2 2 p1 p2 b2 2 V You should verify that these equations give the same heritability estimates and the regression procedure To examine epistasis consider the N locus with alleles N1 and N2 Just as we created two new variables for the M locus we could also create two new variables to model the additive effect and the dominance effect at this locus Call these variables X3 and X4 For genotypes N1N1 N1N2 and N2N2 the respective values for X3 will be 0 1 and 2 the respective values for X4 would be 0 1 and 0 The coding for the additive and the dominance effects at both the M and the N loci are given in Table 18A 2 Insert Table 18A 2 here In regression an interaction between two predictor variables is modeled by creating a new variable that is the product of the two predictor variables and entering this variable Genetic epistasis is modeled in the same way Multiplying the additive variable for the M locus X1 by the additive variable for the N locus X3 gives a new variable X5 in Table 18A 2 that geneticists call additive by additive epistasis The variance associated with this is termed additive by additive epistatic variance There are two different ways to model the interaction between an additive effect at one locus and a dominance effect at a second locus First we could multiply the additive variable for the M locus by the dominance variable for the N locus This new variable is appropriate heterozygote would have a value of 1 and all other genotypes would be assigned a value of 0 1999 2000 Gregory Carey Chapter 19 Advanced Topics 4 given as X6 in Table 18A 2 The second way is to multiply the dominance variable for M by the additive variable for N giving variable X7 in Table 18A 2 Together variables X6 and X7 model additive by dominance epistasis and the variance associated with these two variables is called additive by dominance epistatic variance The final interactive term is the product of the two dominance variables for the M and N loci It is given as X8 in Table 18A 2 This is dominance by dominance