Genesis of cilia and microvilli of rat nasal epithelia during pre-natal development I. Olfactory epithelium, qualitative studies *

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Abstract
Rat foetuses from intra-uterine days E13 through E22 (day before parturition) and adults were used for a qualitative electron-microscopic investigation of the development of ciliated/microvilldus surfaces of the olfactory epithelium. In the E13 and most of the E14 embryos the epithelial surface is not yet characteristically olfactory. Apical cell profiles show primary cilia. These can arise at the epithelial surface or below. From E14 onwards the epithelial surface acquires olfactory characteristics. Dendritic endings of the olfactory receptor cells can be found amidst microvillous profiles of supporting cells. Either cell type may bear primary cilia. F rom E16 onwards the receptor cells sprout multiple olfactory cilia, but cells with primary cilia are found throughout pre-natal development. These primary cilia are, at least for a while, retained during the formation of the secondary cilia. Primary cilia always have distinct necklaces at their base. Otherwise, especially with respect to their tips, their morphology can vary. Originally they have expanded tips (up to E14) ; later on such wide tips are no longer encountered (E16 and E17). Primary cilia of receptor cells never have wide tips. Appreciable numbers of endings with tapering olfactory cilia are discerned around E18 and especially E19. Throughout pre-natal development posterior/superior parts of the septal olfactory epithelium are more precocious than anterior/inferior parts, in particular in the region of transition with the respiratory epithelium. This advance in development includes total densities of dendritic endings of olfactory receptor cells, densities of multiciliated endings alone and lengths of supporting cell microvilli. This difference is discussed with respect to the topography of the olfactory epithelial surface in adult animals. In addition to the systematic topographic variation, a number of more local, apparently not-systematically distributed, topographic variations present during development are described. Most of these also occur in adult animals and they include heterogeneity in length of supporting cell microvilli and the presence of patches of supporting cells with rounded apical protuberances, of patches displaying dendrites with polyaxonemes rather than individual cilia and of scattered atypical cells (neither typical olfactory receptor nor olfactory supporting cells). At their surfaces such atypical cells can resemble inner-ear hair cells. Relative to olfactory receptor and supporting cells there are only very few atypical cells. Finally, we describe some differences in the appearance of membrane surfaces of olfactory cilia and supporting cell microvilli as a function of development.