After 1952: The later development of Alan Turing's ideas on the mathematics of pattern formation

The paper 'The chemical basis of morphogenesis' [Phil. Trans. R. Soc. Lond. B 237, 37-72 (1952)] by Alan Turing remains hugely influential in the development of mathematical biology as a field of research and was his only published work in the area. In this paper I discuss the later development of his ideas as revealed by lesser-known archive material, in particular the draft notes for a paper with the title 'Outline of development of the Daisy'. These notes show that, in his mathematical work on pattern formation, Turing developed substantial insights that go far beyond Turing (1952). The model differential equations discussed in his notes are substantially different from those that are the subject of Turing (1952) and present a much more complex mathematical challenge. In taking on this challenge, Turing's work anticipates (i) the description of patterns in terms of modes in Fourier space and their nonlinear interactions, (ii) the construction of the well-known model equation usually ascribed to Swift and Hohenberg, published 23 years after Turing's death, and (iii) the use of symmetry to organise computations of the stability of symmetrical equilibria corresponding to spatial patterns. This paper focuses on Turing's mathematics rather than his intended applications of his theories to phyllotaxis, gastrulation, or the unicellular marine organisms Radiolaria. The paper argues that this archive material shows that Turing encountered and wrestled with many issues that became key mathematical research questions in subsequent decades, showing a level of technical skill that was clearly both ahead of contemporary work, and also independent of it. His legacy in recognising that the formation of patterns can be understood through mathematical models, and that this mathematics could have wide application, could have been far greater than just the single paper of 1952. A revised and substantially extended draft of 'Outline of development of the Daisy' is included in the Supplementary material. (C) 2015 Elsevier Inc. All rights reserved.