Pattern formation from homogeneity is well studied, but less is known concerning symmetry-breaking instabilities in heterogeneous media. It is non-trivial to separate observed spatial patterning due to inherent spatial heterogeneity from emergent patterning due to nonlinear instability. We employ WKBJ asymptotics to investigate Turing instabilities for a spatially heterogeneous reaction-diffusion system, and derive conditions for instability which are local versions of the classical Turing conditions. We find that the structure of unstable modes differs substantially from the typical trigonometric functions seen in the spatially homogeneous setting. Modes of different growth rates are localized to different spatial regions. This localization helps explain common amplitude modulations observed in simulations of Turing systems in heterogeneous settings. We numerically demonstrate this theory, giving an illustrative example of the emergent instabilities and the striking complexity arising from spatially heterogeneous reaction-diffusion systems. Our results give insight both into systems driven by exogenous heterogeneity, as well as successive pattern forming processes, noting that most scenarios in biology do not involve symmetry breaking from homogeneity, but instead consist of sequential evolutions of heterogeneous states. The instability mechanism reported here precisely captures such evolution, and extends Turing's original thesis to a far wider and more realistic class of systems.

Cardiff School of Mathematics Cardiff University Senghennydd Road Cardiff CF24 4AG UK

Department of Mathematics FNSPE Czech Technical University Prague Trojanova 13 120 00 Prague Czech Republic

Wolfson Centre for Mathematical Biology Mathematical Institute University of Oxford Andrew Wiles Building Radcliffe Observatory Quarter Woodstock Road Oxford OX2 6GG UK

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