Fitness Landscape Models
Kauffman (1993, 1995; Kauffman and Levin 1987; Kauffman and Johnsen 1991) has proposed and studied in depth a class of models referred to as NK models, which are models of random fitness landscapes on which one can implement a variety of types of evolutionary dynamics and study the development and interaction of species. (The letters N and K do not stand for anything; they are the names of parameters in the model.) Based on the results of extensive simulations of NK models, Kauffman and co-workers have suggested a number of possible connections between the dynamics of evolution and the extinction rate. To a large extent it is this work which has sparked recent interest in biotic mechanisms for mass extinction. In this chapter we review Kauffman's work in detail. An NK model is a model of a single rugged landscape, which is similar in construction to the spin-glass models of statistical physics (Fischer and Hertz 1991), particularly p-spin models (Derrida 1980) and random energy models (Derrida 1981). Used as a model of species fitness the NK model maps the states of a model genome onto a scalar fitness W. This is a simplification of what happens in real life, where the genotype is first mapped onto phenotype and only then onto fitness. However, it is a useful simplification which makes simulation of the model for large systems tractable. As long as we bear in mind that this simplification has been made, the model can still teach us many useful things. The NK model is a model of a genome with N genes. Each gene has A alleles. In most of Kauffman's studies of the model he used A = 2, a binary genetic code, but his results are not limited to this case. The model also includes epistatic interactions between genes—interactions whereby the state of one gene affects the contribution of another to the overall fitness of the species. In fact, it is these epistatic interactions which are responsible for the ruggedness of the fitness landscape.