Evolvability in Artificial Development of Large, Complex Structures and the Principle of Terminal Addition

Epigenetic tracking (ET) is a model of development that is capable of generating diverse, arbitrary, complex three-dimensional cellular structures starting from a single cell. The generated structures have a level of complexity (in terms of the number of cells) comparable to multicellular biological organisms. In this article, we investigate the evolvability of the development of a complex structure inspired by the "French flag" problem: an "Italian Anubis" (a three-dimensional, doglike figure patterned in three colors). Genes during development are triggered in ET at specific developmental stages, and the fitness of individuals during simulated evolution is calculated after a certain stage. When this evaluation stage was allowed to evolve, genes that were triggered at later stages of development tended to be incorporated into the genome later during evolutionary runs. This suggests the emergence of the property of terminal addition in this system. When the principle of terminal addition was explicitly incorporated into ET, and was the sole mechanism for introducing morphological innovation, evolvability improved markedly, leading to the development of structures much more closely approximating the target at a much lower computational cost.