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10. Molecular Phylogeny
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                                   this revelation, then use formula (10.17) to find the equilibrium dis-
                                   tribution of the nucleotide substitution chain of Section 10.5 when
                                   detailed balance is not assumed. Your answer should match the dis-
                                   tribution appearing in Problem 15.
                                17. In his method of evolutionary parsimony, Lake [14] has highlighted
                                   the balanced transversion assumption. This assumption implies the
                                   constraints λ AC = λ AT , λ GC = λ GT , λ CA = λ CG , and λ TA = λ TG
                                   in the nucleotide substitution model with general transition rates.
                                   Without further restrictions, infinitesimal balanced transversions do
                                   not imply finite-time balanced transversions. For example, the iden-
                                   tity p AC (t)= p AT (t) may not hold. Prove that finite-time balanced
                                   transversions follow if the additional closure assumptions
                                                      λ AG − λ GA  = λ G − λ A
                                                      λ CT − λ TC  = λ T − λ C
                                                                                      k
                                   are made [1]. (Hint: Show by induction that the matrices Λ have the
                                   balanced transversion pattern for equality of entries.)
                                18. In Lake’s balanced transversion model of the last problem, show that

                                                       λ AG λ GT  = λ AT λ GA
                                                       λ CT λ TA  = λ TC λ CA
                                   are necessary and sufficient conditions for the corresponding Markov
                                   chain to be reversible.
                                19. In the Ising model, one can explicitly calculate the partition function
                                        H(c)
                                       e    of Section 10.10. To simplify matters, we impose circular
                                      c
                                   symmetry and write
                                                             m         m

                                                 H(c)  = θ 0    c i + θ 1  1 {c i=c i+1 } ,
                                                             i=1      i=1
                                   where c m+1 = c 1 . Show that

                                                           1      1  m
                                             	   H(c)
                                                                            e
                                                e     =       ···       e θ 0c i θ 1 1 {c i =c i+1 }
                                              c           c 1 =0  c m =0 i=1
                                                                    m
                                                          	     	      t
                                                      =      ···      u Zu i+1 .         (10.18)
                                                                       i
                                                          u 1   u m i=1
                                                                                 t
                                                                                          t
                                   Here each u i ranges over the set of two vectors (1, 0) and (0, 1) , and
                                   Z is the 2 × 2 matrix

                                                               e θ 1  e θ 0
                                                      Z   =                 .
                                                               e θ 0  e 2θ 0 +θ 1
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