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The notion that evolution can be viewed as a hill-climbing process in an adaptive landscape was introduced in 1932 by Wright [1], and remains one of the most powerful images in evolutionary biology [2]. Since the discovery of the molecular structure of genes, it has been clear that the substrate over which the adaptive landscape should be properly defined is the space of genetic sequences [3]. Nevertheless, apart from a few landmark papers [4, 5], adaptation has not been in the focus of the theory of molecular evolution, which instead has concentrated on the effects of stochastic drift in a neutral (flat) fitness landscape [6]. This situation is presently changing [7, 8]. Long-term evolution experiments on microbial populations [9] are beginning to produce a wealth of data, on the phenotypic as well as on the genotypic level, which make it meaningful to ask precise questions about the timing and size of adaptive events, and what they can tell us about the structure of the underlying adaptive landscape.

Adaptation in Simple and Complex Fitness Landscapes