Micromorphology of ovaries and oogenesis in Grania postclitellochaeta (Clitellata: Enchytraeidae).

The genus Grania comprises over 70 species of exclusively marine clitellate annelids belonging to the family Enchytraeidae. Morphologically, this genus is well separated from other enchytraeids, with thick cuticles, anterior segments I-IV fused into a "head", chaetal bundles consisting only of one stout chaeta, and reduction of circular musculature. The aim of the present study is to describe the ovary organization and the course of oogenesis in Grania postclitellochaeta, and to compare it with other known systems of ovary organization and oogenesis in clitellate annelids, especially in enchytraeids. Generally, oogenesis in G. postclitellochaeta can be divided into two phases: (i) early stages of oogenesis, occurring within the paired ovaries - each ovary is similar to a bunch of grapes, where each 'lobe' is a germ-line cyst enveloped by flat somatic cells, and (ii) oogenesis proper, which takes place within the body lumen where each growing oocyte is accompanied by its own group of nurse cells. Germ cells are interconnected by cytoplasmic channels (intercellular bridges, ring canals) and form syncytial cysts. As in other clitellate annelids, the cyst center contains a common cytoplasm (cytophore) to which each cell is connected by one ring canal only. Initially, within the ovary, all interconnected cells develop synchronously and are morphologically similar. At the time when the cysts detach from the ovary, one of the interconnected cells begins to gather nutrients, grows and becomes an oocyte, whereas the rest of the cells (nurse cells) do not continue meiosis and instead seem to provide the oocyte with macromolecules and cell organelles. Analysis of serial sections reveals that cysts are always composed of 16 cells - one oocyte and fifteen nurse cells. A comparative analysis showed that almost all features of oogenesis in G. postclitellochaeta are similar to that in other representatives of Enchytraeidae (mainly Enchytraeus albidus), suggesting evolutionary conservation of the process across this family.