The cnidarian premises of metazoan evolution: from triploblasty, to coelom formation, to metamery

The hydromedusan subumbrellar muscle tissues originate from a mass of endo- and ectoderm derived cells proliferating inwardly. This mass of cells, called entocodon, is separated by the ecto- and endoderm through a layer of extracellular matrix, thus forming a locally triploblastic arrangement of tissues. By cavitation and differentiation, the entocodon gives rise to the striated and smooth muscle layers of the subumbrella. The structure of the striated muscle is histologically identical to that described for triploblasts, where striated muscle is mainly mesodermic. Together with the mode of development, this suggests that not all Cnidaria are diploblastic and that the subumbrellar cavity is a coelom-like structure. The subumbrellar cavity is formed late in ontogeny, whereas the coelom in higher animals is normally formed during embryonic development. Instead of remaining closed, the subumbrella becomes open, with the muscular mesothelium in contact with the environment. This view of cnidarian structures can be pushed further while considering the regulation of modularity and metamery. The occurrence of homeotic genes in cnidarians and other 'lower' animals shows that the specification of body architecture has a common molecular ground throughout the animal kingdom. It is suggested that the expression of homeotic genes while specifying a given architecture might be regulated by iteration of their activity, leading to 'low' modular organisms and to 'higher' metameric ones. Replicated parts as modules and metameres might be analogous, but the affinity in their molecular specification (and possibly regulation) might indicate homology. The number of resemblances between cnidarians and 'higher' metazoa suggests that this phylum might be a centre of metazoan radiation, having structures which represent the living premises of the main steps of animal evolution.