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Physical principles underlying biological pattern formation are discussed. In particular, the combination of local self-enhancement and long-range (“lateral”) inhibition (Gierer and Meinhardt, 1972) accounts for de-novo pattern formation, and for striking features of developmental regulation such as induction, spacing and proportion regulation of centers of activation in tissues and cells. Part I explains physical principles of spatial organisation in biological development. Part II demonstrates in mathematical terms that and how short-range activation and long-range inhibition are conditions for the generation of spatial concentration patterns. The conditions can be expressed in terms of ranges, rates and orders of reactions. These conditions, in turn, can also be derived by analysis of dynamic instabilities by means of Fourier waves, showing the neither obvious nor trivial relation between the latter approach and the theory based primarily on autocatalysis and lateral inhibition.
The paper addresses the formation of striking patterns within originally near-homogenous tissue, the process prototypical for embryology, and represented in particularly puristic form by cut sections of hydra regenerating a complete animal with head and foot. Essential requirements are autocatalytic, self-enhancing activation, combined with inhibitory or depletion effects of wider range - “lateral inhibition”. Not only de-novo-pattern formation, but also well known, striking features of developmental regulation such as induction, inhibition, and proportion regulation can be explained on this basis. The theory provides a mathematical recipe for the construction of molecular models with criteria for the necessary non-linear interactions. It has since been widely applied to different developmental processes.