The chemical basis of morphogenesis
 A. M. Turing
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It is suggested that a system of chemical substances, called morphogens, reacting together and diffusing through a tissue, is adequate to account for the main phenomena of morphogenesis. Such a system, although it may originally be quite homogeneous, may later develop a pattern or structure due to an instability of the homogeneous equilibrium, which is triggered off by random disturbances. Such reactiondiffusion systems are considered in some detail in the case of an isolated ring of cells, a mathematically convenient, though biologically unusual system. The investigation is chiefly concerned with the onset of instability. It is found that there are six essentially different forms which this may take. In the most interesting form stationary waves appear on the ring. It is suggested that this might account, for instance, for the tentacle patterns onHydra and for whorled leaves. A system of reactions and diffusion on a sphere is also considered. Such a system appears to account for gastrulation. Another reaction system in two dimensions gives rise to patterns reminiscent of dappling. It is also suggested that stationary waves in two dimensions could account for the phenomena of phyllotaxis.
The purpose of this paper is to discuss a possible mechanism by which the genes of a zygote may determine the anatomical structure of the resulting organism. The theory does not make any new hypotheses; it merely suggests that certain wellknown physical laws are sufficient to account for many of the facts. The full understanding of the paper requires a good knowledge of mathematics, some biology, and some elementary chemistry. Since readers cannot be expected to be experts in all of these subjects, a number of elementary facts are explained, which can be found in textbooks, but whose omission would make the paper difficult reading.
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 Title
 The chemical basis of morphogenesis
 Journal

Bulletin of Mathematical Biology
Volume 52, Issue 12 , pp 153197
 Cover Date
 19900101
 DOI
 10.1007/BF02459572
 Print ISSN
 00928240
 Online ISSN
 15229602
 Publisher
 Kluwer Academic Publishers
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 Authors

 A. M. Turing ^{(1)}
 Author Affiliations

 1. University of Manchester, Manchester, UK