Reproductive System in the Male Phase of a Parasitic Isopod (Crustacea) - Morphological, Histological and Ultrastructural Evidence for Sequential Protandrous Hermaphroditic Changes.

This paper reports the protandric hermaphroditic changes in the reproductive system of the male-phased Norileca indica, a cymothoid that parasitizes the scombrid fish Rastrelliger kanagurta. Each part of N. indica's paired reproductive system lies on either side of the gut. This study considers the three successive size classes of the male phase - designated as M1, M2 and M3 - using light microscopy and ultrastructural methods. The testis comprises of three bulged sac-like lobes labelled t1, t2 and t3, all of which open into the ovary of their respective side. The vas deferens, which emerges as a posterior extension of the ovary, opens into the penis and the distal end of each oviduct leads to a sealed gonopore on their respective sides. Each testis lobe (t1/t2/t3) displays clusters of germ cells undergoing stage-specific differentiation. Spermatids undergoing sequential changes associated with spermiogenesis keep close proximity to somatic accessory cells. The characteristic histological changes associated with protandric hermaphroditism are visible in the ovaries of sequential size classes (M1, M2 and M3). In early M1, besides spermatophores, the ovary has abundant polymorphic nuclei; in the mid/late M1, the posterior ovary has abundant spermatophores, anterior displayed oogonia, previtellogenic oocytes and two distinct forms of follicle cells. In M2, the anterior ovary shows compactly arranged oocytes while the posterior region accommodates spermatophores - fewer, however, than during M1. The entire ovary during M3 is crowded with previtellogenic oocytes, which marginalize the spermatophore passage. The vas deferens of the smallest M1 lack spermatophores. As the size class progresses through late M1 into M2 and M3, the posterior vas deferens is filled with spermatophores, which closely associate with the glandular epithelial lining.