In contrast to mammals in which the pineal gland is a discrete structure situated dorsally in the brain, the "pineal gland" in teleost fishes is composed of a number of separate but connected constituent parts, collectively described as the "pineal complex." In this paper, we have described the pineal complex in a common Indian carp, Catla catla, which exhibits an annual reproductive cycle. Attempts have been made to (a) provide an in-depth description of the structure of the pineal complex; and (b) identify the photoreceptor cells of the pineal, by exposing the animals to constant light (LL) and constant darkness (DD). Furthermore, we examined any possible influence of the reproductive status of the fish on the responsiveness of the pineal photoreceptor cells in C. catla following exposure to LL and DD. To this end, a total of four experiments were carried out during the four different phases of the annual reproductive cycle that is characteristic of this species. Each of these four experiments was carried out for a period of 30 days after which the fishes were sacrificed, different parts of the pineal complex were dissected out, and processed for histological and karyometric studies. Our results showed that the pineal complex in this species is composed of three separate but connected parts, (a) an end vesicle (EV); (b) a dorsal sac (DS); and (c) a long and thin pineal stalk (PS) that attaches the EV to the DS. Detailed karyometric and histo-morphologic studies following exposure of the animals to DD and LL showed that constant darkness led to a stimulatory effect on the pineal photoreceptor cells of the EV as evident from a significant increase in the nuclear diameter. In contrast, the nuclear diameter of the photoreceptor cells in animals subjected to constant light showed a significant reduction. Furthermore, the observed cellular changes in the EV of fish exposed either to LL or DD were independent of the stage of the gonadal cycle. The apparent lack of any cellular responses either in the PS, or in the DS, following exposure to LL and DD, suggests that in C. catla the photoreceptor cells are located only within the epithelial lining of the EV and that these cells respond in a manner similar to mammalian pinealocytes when subjected to comparable photoperiod-induced experimental conditions.
The present study attempted for the first time to explore the importance of photoperiod in the regulation of seasonal ovarian functions in any subtropical major carp. Adult Indian major carp Catla catla were transferred to a long photoperiod (LP; LD 16:8) or a short photoperiod (SP; LD 8:16) for 30 days on 4 dates corresponding to the beginnings of 4 reproductive phases in an annual cycle, and responsiveness of the ovary was evaluated by comparison with the gonadal weight (I(G)), relative number of developing oocytes, serum levels of vitellogenin, and the activity of 2 important steroidogenic enzymes, that is, Delta(5)3beta-hydroxysteroid dehydrogenase and 17.beta-hydroxysteroid dehydrogenase, in the ovary of fish in a natural photoperiod. Exposure of fish to LP during the preparatory phase (February-March) resulted in a significant (p < 0.001) increase in the values of vitellogenin and in the activity of both the steroidogenic enzymes but not in the ovarian weight and in the relative number of different stages of oocytes. A more stimulatory influence of LP was noted during the prespawning phase (April-May), when precocious maturation of ovary was evident from a significant (p < 0.001) rise in the values of each studied features of ovarian functions. However, no ovarian response was found when the fish were transferred to LP during the spawning (July-August) and the postspawning (September-October) phases. On the other hand, the SP was found to have an inhibitory influence on ovarian growth and maturation during the prespawning and the spawning phases or to have no influences on ovarian functions during the preparatory and the postspawning phases of an annual cycle. The results of our study provide the first evidence that photoperiod per se plays an important role in the seasonal maturation of ovary in a subtropical freshwater major carp.
The structure of the pineal complex and the annual reproductive cycle in a major Indian carp, Catla catla, were investigated in the present study. Additionally, given the well-known inhibitory effects of the pineal on reproductive function in mammals, attempts were made to investigate whether or not the pineal exerts an inhibitory influence on reproductive function in this piscine species as well. Sexually adult animals were utilized in all experiments. The cytomorphology of the pineal complex and a number of parameters for testicular function--such as testicular cytology, serum testosterone levels, and testicular activities of two steroidogenic enzymes, 17beta-hydroxysteroid dehydrogenase (17beta-HSD) and delta5-3beta-hydroxysteroid dehydrogenase (delta5-3beta-HSD) were examined over a period of two years. Our studies showed that the pineal complex in this species consists of three separate but distinctly connected components: (a) an end vesicle (EV); (b) a long pineal stalk (PS); and (c) a dorsal sac (DS). Of these, the epithelial lining of the EV consists of cells that have rounded vesicular nuclei and long apical cytoplasmic processes that reach the lumen, features suggestive of photoreceptor cells. The cells of the PS have some similarity with those of the EV, while DS cells appear columnar and ciliated. With regard to gonadal activity, germ cell profiles revealed that this species has four distinct phases during the annual reproductive cycle: (a) preparatory (January-April); (b) pre-spawning (May-June); (c) spawning (July); and (d) post-spawning (August-December). During the spawning phase (July), seminiferous tubular diameter, percentage of late spermatids within seminiferous tubules, and serum testosterone levels showed the highest values compared to those obtained in most of the other phases of the reproductive cycle. Also in July, along with peak serum testosterone levels, the activities of 17beta-HSD and delta5-3beta-HSD were at their highest levels. In a correlation between the pineal cytology and testicular functional status, it was noted that both the nuclear diameter and the apical cytoplasmic projections of the EV photoreceptor cells showed a significant reduction, thus suggesting a reduced synthetic activity, during the month of July, the spawning phase of the reproductive cycle. In contrast, the same features of the EV cells during the other phases of the reproductive cycle showed an increased cellular and metabolic activity--a time when the gonads were less active and in a quiescent stage. These data suggest an inhibitory role of the pineal on gonadal function and thus provide additional credence to the concept that, as in higher mammals, there exists an inverse relationship between the pineal activity and gonadal function in teleost fishes as well.
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