  Much of the current theory of adaptation is based on Gillespie's mutational
landscape model (MLM), which assumes that the fitness values of genotypes
linked by single mutational steps are independent random variables. On the
other hand, a growing body of empirical evidence shows that real fitness
landscapes, while possessing a considerable amount of ruggedness, are smoother
than predicted by the MLM. In the present article we propose and analyse a
simple fitness landscape model with tunable ruggedness based on the Rough Mount
Fuji (RMF) model originally introduced by Aita et al. [Biopolymers 54:64-79
(2000)] in the context of protein evolution. We provide a comprehensive
collection of results pertaining to the topographical structure of RMF
landscapes, including explicit formulae for the expected number of local
fitness maxima, the location of the global peak, and the fitness correlation
function. The statistics of single and multiple adaptive steps on the RMF
landscape are explored mainly through simulations, and the results are compared
to the known behavior in the MLM model. Finally, we show that the RMF model can
explain the large number of second-step mutations observed on a highly-fit
first step backgound in a recent evolution experiment with a microvirid
bacteriophage [Miller et al., Genetics 187:185-202 (2011)].
