Origins of differentiation via phenotypic plasticity.

How cell types of multicellular organisms came to be differentiated is still an open issue. Here I offer a model that posits that the origins of some cell differentiation patterns were originally passive outcomes of environmental effects. As cells' contact with the external environment was diminished, their patterns of gene expression were altered, due to changes in concentrations of externally supplied substances. Later, as multicellular growth continued, the relationships of cell layers to each other shifted, producing concentration gradients of signaling molecules. These gradients emanated both from the external cell layer toward the inside and from internal cell layers to adjacent layers. In this scenario then, differentiation arose initially as a by-product of the changing patterns of gene expression and of the complex mixtures and changing concentrations of substances passing among layers. Subsequent selection would operate to stabilize the expression patterns in those cell layers whose phenotypes provide a fitness advantage to the organism.