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The Felsenstein pruning algorithm computes a site likelihood function by sum-product belief propagation from leaves to root. For a finite-state Markov kernel $K$, the subtree likelihood function obeys $L_v(a)={\prod }_{w\text{ child of }v}{\sum }_{b}K(a,b)L_w(b)$, with observed-state indicators at leaves. Summing against the root probability distribution gives the site likelihood function. An outward pass gives edge posterior probabilities and expected transition counts for the expectation-maximization algorithm.

Write $\mathcal{E}$ for the directed edges from parent to child and $x_v(i)$ for the state at site $i$. The global Markov property for an undirected graph on this tree, together with the specified parent transitions, gives the complete-data likelihood functionOne can obtain the factorization by successively removing terminal subtrees: after conditioning on their parent, each subtree is independent of the rest. The root factor and transition products also show that different sites are independent. Only ancestral states are latent variables.

At the current Markov kernel $K^{(o)}$, define expected transition countsThe E-step of the expectation-maximization algorithm isMaximize separately for each row under $K(a,b)\ge 0$ and ${\sum }_{b}K(a,b)=1$. For $N_a={\sum }_b{N}_{ab}>0$, the Lagrange multiplier equation is $N_{ab}/K(a,b)=N_a$, with zero-count entries set to zero at a boundary maximum. Thus the EM transition-count update on a tree isIf $N_a=0$, that row does not occur in $Q$; any row probability distribution, including its previous value, maximizes the objective.

The expected counts can be computed exactly by the Felsenstein pruning algorithm and an outward belief propagation pass. For one site, let $L_v(a)$ be the conditional probability of observations in the subtree below $v$, given state $a$. For a leaf $v$ with observed state $d_v$, $L_v(a)=1_{\{a=d_v\}}$. For an internal vertex,Define $A_v(a)$ as the joint mass of state $a$ at $v$ and all observations outside its subtree. The root has $A_{v_0}(a)=1/4$. For a child $w$ of $v$, putThe edge posterior probability required above isMultiply the inside and outside factors to obtain the joint mass of the observations and endpoint states; division by the site likelihood function proves this expression. Sum $\xi$ over edges and sites to form $N_{ab}$. The two passes take $O(k|\mathcal{E}|4^2)=O(kn)$ operations on a binary tree. Scaling messages or computing them logarithmically avoids underflow.

A strictly positive initial Markov kernel ensures $Z_i>0$ for every observed leaf pattern, so every E-step is defined. At a later boundary iterate, require positive likelihood for the actual data and restrict the conditional distribution to its support. EM likelihood monotonicity proves that the update cannot decrease observed-data likelihood function. It is a maximum-likelihood iteration, not a guarantee of finding the global maximum-likelihood estimate; different initial kernels may lead to different stationary points.