Morphogenesis of the Retinal Rods

The morphogenesis of the retinal rods has been studied with the electron microscope in white mice from birth up to the 16th day of age. Observations have been mainly concentrated on specimens of the 8th and 12th days and on the differentiation of the inner and outer segments of the retinal rods. In the morphogenesis of the outer segment three main stages have been considered. The first stage consists in the development of a primitive cilium projecting from a bulge of protoplasm which constitutes the primordium of the inner segment. A basal body, nine pairs of peripheral filaments, a surface membrane, and a matrix filled with a fine vesicular material have been recognized as components of the primitive cilium. The vesicles are called "morphogenetic material" because it is believed that they represent the macromolecular primordium of the rod sacs of the future outer segment. The second stage corresponds to the great enlargment of the apical region of the primitive cilium due to the rapid building up of the lamellar material of the rod sacs. The primitive rod sacs appear to be connected with the ciliary filaments. The basal portion of the primitive cilium remains undifferentiated and constitutes the connecting cilium of the adult rod (1). The third stage consists in the remodelling and reorientation of the rod sacs into their permanent transverse disposition. This process starts in the middle portion of the outer segments and proceeds towards both extremities which can be considered as zones of growth of the outer segment. The inner segment is at the beginning a bulge of protoplasm containing unoriented mitochondria, a basal body, a small Golgi zone, and numerous dense particles. Then this region becomes elongated and the different components assume the stratified disposition found in the adult (1). The demonstration that the entire outer segment of the rod cell is the result of the differentiation of a primitive cilium is discussed in view of the conflicting interpretations found in the literature. The possible macromolecular mechanisms that may be involved in the submicroscopic morphogenesis of the rod sacs are discussed and the possible role of the morphogenetic material is considered. The results described in this paper confirm and extend the interpretation of the submicroscopic morphology of the adult rod cell as presented in a previous paper (1).