Open Access

Alfred Gierer

• 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.