The evolutionary origins of glia are lost in time, as soft tissues rarely leave behind fossil footprints, and any molecular footprints they might have been left we have yet to decipher. Nevertheless, because of the growing realization of the importance glia plays in the development and functioning of the nervous system, lessons we can draw about commonalities among different taxa (including vertebrates) brought about either from a common origin, or from common adaptational pressures, shed light on the roles glia play in all nervous systems. The Acoelomorpha, primitive interstitial flatworms with very simple cellular organization and currently at the base of the bilaterian phylogeny, possess glia-like cells. If they indeed represent the ancestors of all other Bilateria, then it is possible that all glias derive from a common ancestor. However, basal taxa lacking convincing glia are found in most major phyletic lines: urochordates, hemichordates, bryozoans, rotifers, and basal platyhelminths. With deep phylogenies currently in flux, it is equally possible that glia in several lines had different origins. If developmental patterns are any indication, glia evolved from ectodermal cells, possibly from a mobile lineage, and even possibly independently in different regions of the body. As to what functions might have brought about the evolution of glia, by-product removal, structural support, phagocytic needs, developmental programming, and circuit modulation may be the more likely. Explaining possible cases of glial loss is more difficult, as once evolved, glia appears to keep inventing new functions, giving it continued value even after the original generative need becomes obsolete. Among all the uncertainties regarding the origin of glia, one thing is certain: that our ideas about those origins will change with every rearrangement in deep phylogeny and with continued advances in invertebrate molecular and developmental areas. (C) 2011 Wiley-Liss, Inc.
