  We consider the evolution of populations under the joint action of mutation
and differential reproduction, or selection. The population is modelled as a
finite-type Markov branching process in continuous time, and the associated
genealogical tree is viewed both in the forward and the backward direction of
time. The stationary type distribution of the reversed process, the so-called
ancestral distribution, turns out as a key for the study of mutation-selection
balance. This balance can be expressed in the form of a variational principle
that quantifies the respective roles of reproduction and mutation for any
possible type distribution. It shows that the mean growth rate of the
population results from a competition for a maximal long-term growth rate, as
given by the difference between the current mean reproduction rate, and an
asymptotic decay rate related to the mutation process; this tradeoff is won by
the ancestral distribution.
  Our main application is the quasispecies model of sequence evolution with
mutation coupled to reproduction but independent across sites, and a fitness
function that is invariant under permutation of sites. Here, the variational
principle is worked out in detail and yields a simple, explicit result.
