The International Journal of Developmental Biology

Int. J. Dev. Biol. 50: 289 - 299 (2006)

Vol 50, Issue 2-3

Special Issue: Developmental Morphodynamics

Before programs: The physical origination of multicellular forms

Open Access | Published: 15 February 2006

Stuart A. Newman1,*, Gabor Forgacs2 and Gerd B. Müller3

1New York Medical College, Valhalla, NY, USA, 2University of Missouri, Columbia, MO, USA and 3University of Vienna, Wien, Austria


By examining the formative role of physical processes in modern-day developmental systems, we infer that although such determinants are subject to constraints and rarely act in a "pure" fashion, they are identical to processes generic to all viscoelastic, chemically excitable media, non-living as well as living. The processes considered are free diffusion, immiscible liquid behavior, oscillation and multistability of chemical state, reaction-diffusion coupling and mechanochemical responsivity. We suggest that such processes had freer reign at early stages in the history of multicellular life, when less evolution had occurred of genetic mechanisms for stabilization and entrenchment of functionally successful morphologies. From this we devise a hypothetical scenario for pattern formation and morphogenesis in the earliest metazoa. We show that the expected morphologies that would arise during this relatively unconstrained "physical" stage of evolution correspond to the hollow, multilayered and segmented morphotypes seen in the gastrulation stage embryos of modern-day metazoa as well as in Ediacaran fossil deposits of ~600 Ma. We suggest several ways in which organisms that were originally formed by predominantly physical mechanisms could have evolved genetic mechanisms to perpetuate their morphologies.


self-organization, canalization, differential adhesion, epigenetic determinant, generative entrenchment

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