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semi-definite but can be singular if two individuals have identical genotypes or the
         number of markers (m) is less than genotyped individuals (n). If number of markers are
         limited (m < n), an improved non-singular matrix G  can be obtained as wtG + (1 – wt)A.
                                                  wt
         VanRaden (2008) indicated that wt = 0.90, 0.95 and 0.98 gave good results.
            Another method for computing G involves scaling ZZ′ by the reciprocals of
         the expected variance of marker loci (VanRaden, 2008). Thus G = ZDZ′, where
         D is diagonal with:
            d =      1
                  21-
             ii  mp  (  p )]
                  [
                     j   j
            The MME for Eqn 11.8 are:
               ′
                 −1
                           −1
                                                −1
                                              ′
            ⎛ XR X     X ′R W        ⎞ ⎛ ⎞ ˆ b  ⎛ ⎛ XR y ⎞
            ⎜    −1        −1     −1  ⎟ ⎜ ⎟  =  ⎜  −1  ⎟                    (11.9)
                                               ′
               ′
            ⎝ WR X     W ′R W +  G a ⎠ ⎝ ⎠ a ˆ  ⎝ WR y⎠
                               2
         where a now equals s /s .
                            2
                            e  a
            This approach for genomic evaluation has the advantage that existing software
         for genetic evaluation can be used by replacing A with G and the systems of equations
         are of the size of animals, which tend to be fewer than the number of SNPs. In pedi-
         gree populations, G discriminates among sibs, and other relatives, allowing us to say
         whether these sibs are more or less alike than expected, so we can capture informa-
         tion on Mendelian sampling. Also, the method is attractive for populations without
         good pedigree, as G will capture this information among the genotyped individuals
         (Hayes and Daetwyler, 2013).
                                                                   2
            Note that Eqn 11.9 assumes all the additive genetic variance (s ) is captured by
                                                                   a
         the SNP, but this may not be the case if the linkage disequilibrium between SNP and
         QTL is not perfect. Later, in Section 11.6, a model is discussed that might capture any
         residual polygenic variance not captured by the SNPs. Another possible limitation is
                                                −1
         that there are no direct rules for computing G  and in large populations the compu-
         tation may not be feasible.
         Example 11.3
         The data in Example 11.1 is analysed using Eqns 11.8 and 11.9 and the same genetic
         parameters to compute DGVs for both the reference and validation animals without
         using weights.
            The matrix X in Eqn 11.9 is the same X as in Example 11.1, W is a diagonal
         matrix for the eight reference animals with records and a = 245/35.25 = 6.950.
            The G matrix constructed from Z for the ten SNPs as:
               ZZ′
            2Σp ( 1 p )−
                j   j
         with 2Σp (1 − p ) = 3.5383 is:
                j     j
         13  1.472
         14 −0.446  0.746
         15  0.988 −0.930  1.634
         16  0.059 −0.446  0.422  0.907
         17  0.685 −0.950  1.048  0.402  1.593
         18 −0.163  0.180 −0.365 −0.163 −0.102  0.746
         19 −0.708  0.201 −0.627  0.423 −0.365  0.201 0.786
         20 −0.547  0.079 −0.183  0.301 −0.203  0.079 0.382  0.826


          186                                                            Chapter 11
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