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Three general methods are used to estimate heritability. First, as discussed earlier, we can measure heritability by the response of a population to selection. Second, we can directly estimate the components of variance by minimizing one component; the remaining variance can then be attributed to other causes. For example, by minimizing environmental causes of variance, we can estimate the genetic component directly. Or, by eliminating the genetic causes of variance, we can estimate the environmental component directly. Third, we can measure the similarity between relatives. We look now at the latter two methods. Variance components can be minimized in several different ways. If we use genetically identical organisms, then the additive, dominance, and epistatic variances are zero, and all that is left is the environmental variance. For example, F. Robertson determined the variance components for the length of the thorax in Drosophila. The total variance (VPh) in a genetically heterogeneous population was 0.366 (measured directly from the distribution of the trait, as in tables 18.2 and 18.3). He then looked at the variance in ies that were genetically homogeneous. These were from isolated lines inbred in the laboratory over many generations to become virtually homozygous. Robertson studied the F1 in several different matings of inbred lines and found the variance in thorax length to be 0.186 (VE). By subtraction (0.366 0.186), we know that the total genetic variance (VG) was 0.180. From this, we can calculate heritability in the broad sense as HB VG/VPh 0.180/0.366 0.49

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One of the important features of the Sallen and Key lter prototype is that we can choose the values for the resistors that set the circuit gain independently of those of the resistors that set the cutoff frequency (the converse is not true) Thus, we can separately select K = 10 for both stages by requiring that 1 + RA /RB = 10, for example, RA = 100 k , RA = 111 k Next, we compute the component values using equations 1520 Since we have only two equations and four unknowns, two values will have to be selected arbitrarily We can write: C = and 1 = Q R2 R1 C2 + C1 R1 R2 C2 R1 + (1 K) C1 R2 C1 C2 1 1 C C1 C2 = R 1 R2 C1 C2 R1 R 2 R1 R 2 = 1 C C1 C2

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Data for air at 15 C Pa = 147 Ibf in2 Po Pa = 50 lbf in2 8 jets per row 2 rows of jets at station or 1 row of jets at station L/D = 1

and operate a change of variables to obtain: x= R2 R1 c x c= C2 + (1 K) C1 C1 C2 a= C2 C1 b= 1 Q

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Genetic variance can be measured directly by minimizing the in uence of the environment. This is most easily done with plants grown in a greenhouse. Under that circumstance, environmental variables, such as soil quality, water, and sunlight, can be controlled to a very high degree. Hence, the variance among individuals grown under these circumstances is almost all genetic variance. The total phenotypic variance can be obtained from the plants grown under natural circumstances. This allows us to calculate heritability in the broad sense. Several methods exist to sort out the additive from the dominant and epistatic portions of the genetic variance. The methods rely mostly on correlations between relatives. That is, the expected amount of genetic similarity between certain relatives can be compared with the actual similarity. The expected amount of genetic similarity is the proportion of genes shared; this is a known quantity for any form of relatedness. For example, parents and offspring have half their genes in common. The relation of observed and expected correlations between relatives is a direct measure of heritability in the narrow sense. We can thus de ne HN robs rexp

ax 2 bx + c = 0 There is always a positive root in the preceding equation, as a and b are both positive One can easily show that there is another positive root if 1 +K 1< 4Q2 C2 < K 1 C1

(18.13)

and in that case there are two solutions for R1 , R2 ; also, there are real solutions only if C2 < (K 1)C1 Solving the preceding equations gives: x= b+ R2 = R1 b2 4 ac 2a

To calculate a heritability in the narrow sense, it is necessary to extract the components of the genetic variance, VG.

80 (Ibf/in2)

Now, we have the new system: 1 C2 1 + 4 + 4(K 1) 2 R Q Q C1 2 = R1 C2 2 C1 1 R1 R2 = wC C1 C2 that can by easily be solved by substitution, as follows: 1 + Q C2 1 4 + 4(K 1) 2 Q C1 C2 2 C1 1 + Q C2 1 4 + 4(K 1) 2 Q C1 2 That is, 2 R1 = C C1 C2 1 + Q 1 + Q C2 C1 C2 C1

R2 =

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