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Oogenesis,Vitellogenesis and Cocoon Secretion in Two Freshwater Leeches from Assiut, Egypt

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Oogenesis,Vitellogenesis and Cocoon Secretion in Two Freshwater Leeches from Assiut, Egypt Research in Zoology 2012, 2(5): 47-59 DOI: 10.5923/j.zoology.20120205.02 Oogenesis,Vitellogenesis and Cocoon Secretion in Two Freshwater Leeches from Assiut, Egypt Hanaa A. Gouda Zoology Department, Faculty of Science, Assiut University, Assiut, Egypt Abstract The organization of the ovisacs, fine structure of the oogenesis, vitellogenesis and formation of cocoons were studied in two freshwater leeches; Alboglossiphonia levis (Glossiphoniidae) and Barbronia assiuti (Barbronidae). Each of them was composed of two elongated ovisacs of germ cells' c lusters. The ovisacs, in the studied species, were me roistic (nutrimental), without regionalization and mitotically dividing oogonia, germ cells entering the meiosis prophase as well as trophocytes, prominent cytophores and growing oocytes were occurred. During late previtellogenesis/early vitellogenesis, the oocytes detached from the cytophore and floated in the coelom; they are usually enveloped by the peritoneal epithelium and associated with blood vessels. Meantime, the activity of clitellar gland cells in both studied leeches could be distinguished. Both light and electron microscopy revealed two types of clitellar gland cells. These exocrine gland cells were responsible for the formation of cocoons which enclosed the fertilized ova. The organization of ovisacs and the course of oogenesis in the two genera were studied ultrastructurally and compared with other related leeches. Keywo rds Alboglossiphonia, Barbronia, Oogenesis, Vite llogenesis, Ultrastructure, Egypt prominence of the clitellum is variable in leeches (usually 1. Introduction somites X to XIII; that is, the eighth through eleventh true segments) and often is not evident externally in mos t Ooogenesis in leeches begins with the release of epithelial species[22]. Various studies of oogenesis have been cells from the walls of specialized coelomic sacs. The restricted mainly to glossiphoniids[8-11,15,17-19], little in released oogonia float freely in the lumen of the coelomic hirudinids[1], p iscicolids[23]. So far, no ultrastructural sacs wherein they undergo mitotic divisions followed by studies investigated the gametogenesis in Barbroniida. The meiosis and the haploid nuclei eventually are found arranged present study is aimed to study the oogenesis, vitellogenesis around the periphery of an anucleate cytophore to which the and secretion of cocoons in Barbronia assiuti gametes remain connected[1]. Multiple oogonia, many (Arhynchobdellida: Barbronidae) as we ll as, trophocytes and a number of growing oocytes are Alboglossiphonia levis (Rhynchobdellida: Glossiphoniidae). for med[2,3]. Trophocytes (nurse cells) are connected by inter-cellular bridges and form clusters. Only one within each cluster differentiates into an oocyte and continues the 2. Materials and Methods me iosis. Trophocytes are mostly responsible for rRNA synthesis and its transport to oocytes[4]. Early studies[5-7] 2.1. Sampling and other modern[8-20] on leech oogenesis have shown that Sexually mature leeches of Alboglossiphonia levis Gouda, the course of this process and the spatial re lationship 2010 and Barbronia assiuti Hussein and El-Shimy, 1982 between different kinds of cells (oocytes, trophocytes and ° ° were collected from Al-Sont canal (27 11' N / 31 11' E) follic le cells) inside the ovary vary considerably among , various families. Moreover, only glossiphoniid leeches near Assiut governorate. produce many large yolky eggs[15,21], whereas leeches in the remaining families lay s mall and yolk-poor eggs[15]. 2.2. Techniques Like other clitellates, leeches possess a clitellum, the Nine leeches from each studied species were relaxed in % s addle-like glandular region associated with cocoon 8 ethanol, specimens were examined at the clitellu m and deposition in the anterior portion of the body. The gonopores to detect their maturity. Two specimens of both spp., were processed and stuck onto a holder in ventral and * Corresponding author: dorsal surfaces using a suitable glue then and [email protected] (Han aa A. Gouda) photomicrographed with Jeol SM (JSM-5400 LV scanning Published online at http://journal.sapub.org/zoology Copyright © 2012 Scientific & Academic Publishing. All Rights Reserved electron microscope), at the EM Unit. The other seven 48 Hanaa A. Gouda: Oogenesis,Vitellogenesis and Cocoon Secretion in Two Freshwater Leeches from Assiut, Egypt specimens were cut into two halves at a level below the i.e. no zones with consecutive stages of oogenesis could be ovisacs. The anterior halves of both Alboglossiphonia levis found within ovary cords. During oogenesis three types of and Barbronia assiuti were fixed in formol saline (1 g cells; enveloping follicle cells, numerous trophocytes and calcium chloride + 100 ml neutral formalin) for 48 hrs. Four few developing oocytes, could be detected clearly. During specimens were used for paraffin sections and three for the previtellogenesis, each trophocyte is connected with an transmission electron microscopy (TEM). Pa raffin tissues anuclear cytoplasmic mass, the cytophore, by a cytoplasmic were processed routinely for light microscopy based on[24]. bridge. Organelles such as mitochondria, cisternae of rER, Tissues were sectioned at 6 um at the clitellar region. So me Golgi complexes and ribosomes are transported via histological procedures were performed to distinguish the intercellular bridges and the cytophore towards the growing different types of clitellar cells in both studied species. oocytes (Figs. 3a-h). Neutral mucosubstances were demonstrated by the periodic acid-Schiff (PAS) reaction. Neutral and acidic 3.2.1. Follicle Ce lls mucosubstances were revealed in the same section by In A. levis there were outer follicular cells surrounding the staining with a combination of the PAS reaction and Alcian trophocytes and oocytes (Figs. 1d,f,g) as well as inner blue. The section was first stained with Alcian blue at pH 2.5 follicular ce lls between the developing oocytes (Figs. 1e,f). and Alcian blue-positive acid mucosubstances that were also Outer follicular ce lls differed considerably from inner ones. PAS positive will not react with PAS. Only neutral The outer follicular ce lls were thin and elongated except for compounds were PAS positive. Also, toluidine blue could the nuclear region, which was much broader, oval nucleus, stain metachromatically the acidic mucus and sulphated rounded prominent nucleolus and rich in mitochondria and compounds. After examination, photomicrographs were rough endoplasmic ret iculu m especially at the extremities taken by a digital camera (Canon G6) fixed on a binocular (Fig. 1f,g). No evident signs of synthetic activity by these Zeiss light microscope. For transmission electron cells were observed in A. levis. Inner fo llicula r cells were microscopy, three anterior halves of A. levis and B. assiuti found in groups (2-3), between the developing oocytes (Figs. were placed in 2.5 % glutaraldehyde in 0.1 M phosphate 1e,f). These cells appeared oval in shape, with oval nucleus buffer at pH 7.2 for 3 hrs. at roo m temperature. The and rounded prominent nucleolus. These cells showed great specimens were post fixed in 1% buffered osmium tetroxide synthetic activity and their cytoplasm were filled with huge at pH 7.2 for 3 hours at 4 °c. They were then dehydrated and amounts of electron dense cortical granules. These granules embedded in Epon. Semithin sections of 0.5 um were cut and occurred between the microvilli and periphery of oocytes via stained with toluidine blue for examination under a light exocytosis (Figs. 1e,4b). Furthermore, cortical granules microscope. Ultrathin sections from selected areas of the constituted a condensed layer around large oocytes (Fig. 4b). trimmed blocks were made and collected in copper grids. As the oocytes enlarged, the inner follicular cells became The ultrathin sections were contrasted in uranyl acetate for much thinner (Fig. 4b). At the end of oogenesis, when the 10 minutes, lead citrate for 5 minutes and examined under cortical granules are formed, follicle cells detached fro m the transmission electron microscope JEM, 100 Cx11 TEM, at oocytes and degenerated. In B. assiuti there were only outer Assiut University. follicular cells surrounding the oocytes (Figs. 2b -d,5a) and the inner follicular cells could not be detected. Follicular ce ll in B. assiuti was greatly simila r, in structure and shape, to the 3. Results inner follicular ce ll of A. levis. The cells were rich in rough endoplasmic reticulum and golgi complex (Figs. 2c,d). 3.1. Gross Mor phology of the Ovisac Various sized vesicles of both clear and electron dense Both Alboglossiphonia levis (Fig. 1a) and Barbronia secretions were found in follicle cells of B. assiuti (Figs. assiuti (Fig. 2a ) have similar pair of elongated ovisacs . 2c,d). At the onset of vitellogenesis, the follic le cells tightly Inside the ovisacs, numerous germ cells were arranged in enveloped the oocytes (Figs. 3h,4b,5a). The plas ma solid cords (Figs. 1a-c, 2a). me mb rane of fo llic le ce lls appeared smooth towards the ovisac wall but slightly protruded towards the oocyte (Fig. 3.2. Oogenesis 2c), which seemed to form gaps filled with he mocoelo mic Histology showed ovisac of both current species was flu id. At late vitellogenesis, some various sized yolk beside composed of cords of germ-cell line, cords are non-polarized; few cortical granules were found. Research in Zoology 2012, 2(5): 47-59 49 Fi gures 1. Mat ure Alboglossiphonia levis; (a) gross anat omy of ovisacs (ov), germ cells (gc), t rophocyt es (t), cytophore (asterisk), degenerating t rophocyt es (arrows); (b) semithin section through germ cells (gc); (c) ult rat hin sect ion, germ cells (gc), nucleus (n), scale bar = 2um; (d) semithin section, outer follicular cells (of), t rophocyt es (t); (e-g) T EM o f outer follicular cells (of), nucleus (n), rounded nucleolus (ne), mitochondria (m), rough endoplasmic ret iculum (rr), cort ical granules (asterisk) in front of and behind (arrows) the surface of developing oocyte (do), (a,b,d) stained with toluidine blue, scale bar = 25 um, (c,e-g) = 2 um 50 Hanaa A.
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