A population evolves due to changes in its gene frequencies arising due to
mutations, natural selection, random genetic drifts and migrations, Svirezh
ev introduced an integral variational principle, in analogy with the least-
action principle of classical mechanics, by defining a Lagrangian which rem
ained stationary on the trajectory followed by the population undergoing se
lection, This principle can also be extended to multiple loci in some simpl
e cases. However, in a two-locus model or more general models, there is not
straightforward extension of this principle if linkage and epistasis are p
resent. The local optimality principle can be geometrically formulated in a
Riemannian metric space of gene frequencies: so that, under evolutionary p
ressures, the population trajectory moves in that direction along which the
increment of the mean fitness is maximum.