The topology of robustness and evolvability in evolutionary systems with genotype-phenotype map

In this paper we formulate a topological definition of the concepts of robustness and evolvability. We start our investigation by formulating a multiscale model of the evolutionary dynamics of a population of cells. Our cells are characterised by a genotype-phenotype map: their chances of survival under selective pressure are determined by their phenotypes, whereas the latter are determined their genotypes. According to our multiscale dynamics, the population dynamics generates the evolution of a genotype-phenotype network. Our representation of the genotype-phenotype network is similar to previously described ones, but has a novel element, namely, our network contains two types of nodes: genotype and phenotype nodes. This network representation allows us to characterise robustness and evolvability in terms of its topological properties: phenotypic robustness by means of the clustering coefficient of the phenotype nodes, and evolvability as the emergence of giant connected component which allows navigation between phenotypes. This topological definition of evolvability allows us to characterise the so-called robustness of evolvability, which is defined in terms of the robustness against attack (i.e. edge removal) of the giant connected component. An investigation of the factors that affect the robustness of evolvability shows that phenotypic robustness and the cryptic genetic variation are key to the integrity of the ability to innovate. These results fit within the framework of a number of models which point out that robustness favours rather than hindering evolvability. We further show that the corresponding phenotype network, defined as the one-component projection of the whole genotype-phenotype network, exhibits the small-world phenomenon, which implies that in this type of evolutionary system the rate of adaptability is enhanced.