In Kathleen Keeler's Cost-Benefit Model of Mutualism, the Condition for Persistence

In Kathleen Keeler's cost-benefit model of mutualism, the condition for persistence of mutualists is pw_mw + qw_mu > w_nm. The parameters p and q represent the

A) cost and benefit of the mutualism.
B) fitness of successful and unsuccessful mutualists.
C) frequency of "mutualist" and "nonmutualist" alleles in the population.
D) proportion of successful and unsuccessful mutualists in the population.
E) amount of herbivore protection provided by ant and non-ant defenses.