title: A living mesoscopic cellular automaton made of skin scales.
abstract: In vertebrates, skin colour patterns emerge from nonlinear dynamical microscopic systems of cell interactions. Here we show that in ocellated lizards a quasi-hexagonal lattice of skin scales, rather than individual chromatophore cells, establishes a green and black labyrinthine pattern of skin colour. We analysed time series of lizard scale colour dynamics over four years of their development and demonstrate that this pattern is produced by a cellular automaton (a grid of elements whose states are iterated according to a set of rules based on the states of neighbouring elements) that dynamically computes the colour states of individual mesoscopic skin scales to produce the corresponding macroscopic colour pattern. Using numerical simulations and mathematical derivation, we identify how a discrete von Neumann cellular automaton emerges from a continuous Turing reaction-diffusion system. Skin thickness variation generated by three-dimensional morphogenesis of skin scales causes the underlying reaction-diffusion dynamics to separate into microscopic and mesoscopic spatial scales, the latter generating a cellular automaton. Our study indicates that cellular automata are not merely abstract computational systems, but can directly correspond to processes generated by biological evolution.
authors.affiliation: Laboratory of Artificial and Natural Evolution (LANE), Department of Genetics and Evolution, University of Geneva, Geneva, Switzerland.|SIB Swiss Institute of Bioinformatics, Geneva, Switzerland.
authors.forename: Liana
authors.initials: L
authors.lastname: Manukyan
authors.affiliation: Laboratory of Artificial and Natural Evolution (LANE), Department of Genetics and Evolution, University of Geneva, Geneva, Switzerland.
authors.forename: Sophie A
authors.initials: SA
authors.lastname: Montandon
authors.affiliation: Laboratory of Artificial and Natural Evolution (LANE), Department of Genetics and Evolution, University of Geneva, Geneva, Switzerland.|SIB Swiss Institute of Bioinformatics, Geneva, Switzerland.
authors.forename: Anamarija
authors.initials: A
authors.lastname: Fofonjka
authors.affiliation: Department of Mathematics, University of Geneva, Geneva, Switzerland.|Skolkovo Institute of Science and Technology, Skolkovo, Russia.|Chebyshev Laboratory, Department of Mathematics and Mechanics, St Petersburg State University, St Petersburg, Russia.
authors.forename: Stanislav
authors.initials: S
authors.lastname: Smirnov
authors.affiliation: Laboratory of Artificial and Natural Evolution (LANE), Department of Genetics and Evolution, University of Geneva, Geneva, Switzerland.|SIB Swiss Institute of Bioinformatics, Geneva, Switzerland.
authors.forename: Michel C
authors.initials: MC
authors.lastname: Milinkovitch
mesh_terms: D000818:Animals; D005075:Biological Evolution; D002450:Cell Communication; D003116:Color; D004058:Diffusion; D008116:Lizards / Q000033:anatomy & histology / Q000254:growth & development*; D008297:Male; D009024:Morphogenesis; D012867:Skin / Q000033:anatomy & histology / Q000166:cytology* / Q000254:growth & development*; D012880:Skin Pigmentation / Q000502:physiology*