Structure of the ovotestis and evidence for heterosynthetic incorporation of yolk precursors in the oocytes of the nudibranch mollusc,Spurilla neapolitana

The ovotestis of Spurilla neapolitana consists of a series of spherical lobes, each of which is composed of radially arranged, sac-like acini or follicles. The male and female portions of each acinus are separated by ovarian follicle cells and testicular accessory cells. A thick basal lamina serves as a barrier between adjacent acini. The surface of each ovotestis lobe is covered by several layers of myoepithelial cells resting on a connective tissue layer. Developing oocytes are intimately associated with follicle cells except in the last stages of vitellogenesis. Follicle cells are characterized by the presence of extensive arrays of rough endoplasmic reticulum (RER) and Golgi complexes and may play a role in vitellogenesis. An ultrastructural analysis of vitellogenesis suggests that oocytes utilize both auto- and heterosynthetic mechanisms of yolk formation. Autosynthetsis is suggested by the activity of the Golgi complex and RER, while heterosynthesis is indicated by high levels of endocytotic activity by the oocyte. Follicle cell development and high endocytotic activity in the oocytes may be a reproductive adaptation to accelerate yolk synthesis, resulting in more rapid egg production.     

Ovarian maturation and oogenesis in the blue swimmer crab,Portunus pelagicus (Decapoda: Portunidae)

The study was aimed at understanding the process of reproduction and the changes happening in the ovary of Portunus pelagicus during maturation, which would be useful for its broodstock development for hatchery purposes. For that, tissue samples from different regions of the ovary at various stages of maturation were subjected to light and electron microscopy, and based on the changes revealed and the differences in ovarian morphology, the ovary was divided into five stages such as immature (previtellogenic oocytes), early maturing (early vitellogenic oocytes), late maturing (late vitellogenic oocytes), mature (vitellogenic oocytes), and spent (resorbing oocytes). The ovarian wall comprised of an outermost thin pavement epithelium, a middle layer of connective tissue, and an innermost layer of germinal epithelium. The oocytes matured as they moved from the centrally placed germinal zone toward the ovarian wall. The peripheral arrangement of nucleolar materials and the high incidence of cell organelles during the initial stages indicated vitellogenesis I. Movement of follicle cells toward oocytes in the early maturing stage and low incidence of mitochondria and endoplasmic reticulum in the ooplasm during late vitellogenic stage marked the commencement and end of vitellogenesis II, respectively. Yolk granules at various stages of development were seen in the ooplasm from late vitellogenic stage onwards. The spent ovary had an area with resorbing oocytes and empty follicle cells denoting the end of one reproductive cycle and another area with oogonial cells and previtellogenic oocytes indicating the beginning of the next.     

Reproductive biology of the polychaete Kefersteinia cirrata Keferstein (Hesionidae). I. Ovary structure and oogenesis

The ultrastructure of the ovary and the developing oocytes of the polychaete Kefersteinia cirrata have been described. The paired ovaries occur in all segments from the 11th to the posterior. Each consists of several finger-like lobes around an axial genital blood vessel. Oogenesis is well synchronised, young oocytes start to develop in September and vitellogenesis begins in January and is completed by May.

The young oocytes are embedded among the peritoneal cells of the blood vessel wall which have accumulations of glycogen and other storage products. Each oocyte becomes associated with a follicle cell with abundant rough endoplasmic reticulum. Yolk synthesis involves the accumulation of electron dense granules along the cisternae of the abundant rough endoplasmic reticulum. Active Golgi complexes are present and are involved in the production of cortical alveoli. The oocyte has branched microvilli, which contact the follicle cells or blood sinuses between the follicle cells and peritoneal cells. In post-spawning individuals the lysosome system of the follicle cells is hypertrophied and the cells play a role in oocyte breakdown and resorption.     

Ultrastructure of the ovary and oogenesis in six species of patellid limpets (Gastropoda: Patellogastropoda) from South Africa

Abstract. The ultrastructural features of the ovary and oogenesis have been described in 6 species of patellid limpets from South Africa. The ovary is a complex organ that is divided radially into numerous compartments or lacunae by plate-like, blind-ended, hollow trabeculae that extend from the outer wall of the ovary to its central lumen. Trabeculae are composed of outer epithelial cells, intermittent smooth muscle bands, and extensive connective tissue. Oocytes arise within the walls of the trabeculae and progressively bulge outward into the ovarian lumen during growth while partially surrounded by squamous follicle cells. During early vitellogenesis, the follicle cells lift from the surface of the underlying oocytes and microvilli appear in the perivitelline space. Follicle cells restrict their contact with the oocytes to digitate foot processes that form desmosomes with the oolamina. When vitellogenesis is initiated, the trabecular epithelial cells hypertrophy and become proteosynthetically active. Yolk synthesis involves the direct incorporation of extraoocytic precursors from the lumen of the trabeculae (hemocoel) into yolk granules via receptor-mediated endocytosis. Lipid droplets arise de novo and Golgi complexes synthesize cortical granules that form a thin band beneath the oolamina. A fibrous jelly coat forms between the vitelline envelope and the overlying follicle cells in all species.     

Cytochemical and fine structural analysis of oogenesis in the gastropod, Ilyanassa obsoleta

An analysis of differentiating oocytes of the gastropod, Ilyanassa obsoleta, has been made by techniques of light and electron microscopy. Early previtellogenic oocytes are limited by a smooth surfaced oolemma and are associated with each other by maculae adhaerentes. Previtellogenic oocytes are also distinguished by a large nucleus containing randomly dispersed aggregates of chromatin. Within the ooplasm are Golgi complexes, mitochondria and a few cisternae of the rough endoplasmic reticulum. When vitellogenesis begins, the oolemma becomes morphologically specialized by the formation of microvilli. One also notices an increase in the number of organelles and inclusions such as lipid droplets. During vitellogenesis there is a dilation of the saccules of the Golgi complexes and cisternae of the endoplasmic reticulum. Associated with the Golgi complexes are small protein-carbohydrate yolk precursors encompassed by a membrane. These increase in size by fusing with each other. The “mature” yolk body is a membrane-bounded structure with a central striated core and a granular periphery. At maturity a major portion of the ooplasmic constituents such as as mitochondria and lipid droplets occupy the animal region while the bulk of the population of yolk bodies are situated in the vegetal hemisphere. The follicle cells incompletely encompass the developing oocyte. In addition to the regularly occurring organelles, follicle cells are characterized by the presence of large quantities of rough endoplasmic reticulum and Golgi complexes whose saccules are filled with a dense substance. Associated with the Golgi saccules are secretory droplets of varied size. Amongst the differentiating oocytes and follicle cells are Leydig cells. These cells are characterized by a large vacuole containing glycogen. A possible function for the follicle and Leydig cells is discussed.     

Ultrastructure of the ovary and oogenesis in the polychaete Capitella jonesi (Hartman, 1959)

Ultrastructural features of the ovary and oogenesis in the polychaete Capitella jonesi (Hartman, '59) have been described. The ovaries are paired, sac-like follicles suspended by mesenteries in the ventral coelom throughout the midbody region of the mature worm. Oogenesis is unsynchronized and occurs entirely within the ovary, where developing gametogenic stages are segregated spatially within a germinal and a growth zone. Multiplication of oogonia and differentiation of oocytes into the late stages of vitellogenesis occur in the germinal region of the ovary, whereas late-stage vitellogenic oocytes and mature eggs are located in a growth zone. Follicle cells envelop the oocytes in the germinal zone of the ovary and undergo hypertrophy and ultrastructural changes that correlate with the onset of vitellogenesis. These changes include the development of extensive arrays of rough ER and numerous Golgi complexes, formation of microvilli along the surface of the ovary, and the initiation of extensive endocytotic activity. Oocytes undergo similar, concomitant changes such as the differentiation of surface microvilli, the formation of abundant endocytotic pits and vesicles along the oolemma, and the appearance of numerous Golgi complexes, cisternae of rough ER, and yolk bodies. Yolk synthesis appears to occur by both autosynthetic and heterosynthetic processes involving the conjoined efforts of the Golgi complex and rough ER of the oocyte and the probable addition of extraovarian (heterosynthetic) yolk precursors. Evidence is presented that implicates the follicle cells in the synthesis of yolk precursors for transport to the oocytes. At ovulation, mature oocytes are released from the overy after the overlying follicle cells apparently withdraw. Bundles of microfilaments within the follicle cells may play a role in this withdrawal process.     

嘉庚蛸雌性生殖系统组织学观察

对象山港自然海区中的嘉庚蛸(Octopus tankahkeei)雌性生殖系统的组织学结构进行了研究.结果表明,雌性生殖系统由卵巢、输卵管、输卵管腺组成.卵巢单个、球形,内包裹滤泡细胞围成的卵子,输卵管1对,开口于外套腔中部,每条输卵管中部膨大形成圆球状的输卵管腺.近端输卵管内具两瓣蘑菇状突起,上有不规则短指状分枝,突...     

泥螺卵子发生的超微结构研究

利用透射电镜观察了泥螺卵子发生过程。结果表明 ,泥螺的卵子发生可划分为卵原细胞、卵黄发生早期、卵黄发生中期及卵黄发生后期卵母细胞 4个时期。卵原细胞核大而圆 ,胞质内分布有少量的线粒体和高尔基囊泡 ,细胞表面具微绒毛。卵黄发生早期的卵母细胞 ,胞质中各类细胞器发达 ,并出现数量较多的类朦胧子。卵黄发生中期的卵母细胞胞体迅速增大 ,核伸出伪足状突起 ,卵质中各种细胞器活动活跃 ,并参与形成卵黄粒和脂滴。此期还可观察到卵母细胞与滤泡细胞间的物质交换现象。卵黄发生后期的卵母细胞体积增至最大 ,细胞器数量减少。本文就卵黄发生前后卵母细胞内部构造的变化、意义及滤泡细胞与卵母细胞蛋白来源间的关系作了探讨     

The development of the hexagonally structured egg envelope of the C-O sole (Pleuronichthys coenosus)

The surface of a mature, pelagic C-O sole egg is composed of polygonal chambers having four to eight sides, most of which are hexagonally shaped. This honeycomb pattern initially appears on primary oocytes as a thin layer of compact, electron-dense material. Discrete thickenings begin to develop on the envelope of perinuclear stage oocytes. The thickenings lengthen and thin to form the hexagonal walls of the envelope in oocytes undergoing yolk vesicle formation. The walls of each hexagonal chamber occur in an area corresponding to the lateral margins of the adjacent follicle cell, suggesting that the hexagonal walls are produced by the follicle cells. The hexagonal layer is nearly complete at the beginning of vitellogenesis, and as vitellogenesis continues, a striated envelope layer composed of fibrillar lamellae develops between the oocyte and the hexagonal layer. The striated layer appears to be secreted by the oocyte. After vitellogenesis is completed, oocytes are ovulated and double in size during a period of maturation. Concurrently, the striated primary envelope stretches and thins into eight to nine horizontal lamellae. On the mature egg surface, the polygonal chambers are about 24–31 μm in diameter. Within each chamber there is a subpattern of polygonal areas; each polygon is 1.5–2.0 μm in diameter, and circumscribes a pore canal opening. This exceptional envelope may furnish the egg with some degree of protection, resiliency, and buoyancy, but its specific functions are not known.     

Muco-follicle cells of the jelly coat in the oocyte envelope of the sheatfish (Silurus glanis L.)

During early vitellogenesis of the oocytes of Silurus glanis, the follicular cells proliferate, their epithelial organization becomes disrupted, and they transform into an irregularly structured large mass of cells engaged in intensive secretory activity. They contain nuclei, rough endoplasmic reticulum, Golgi bodies, and secretory inclusions termed “acorn bodies,” which are synthesized in the cytoplasm. The acorn bodies have two components: an electron-dense cap and a moderately electron-dense body. As development proceeds, the acorn bodies become modified into spherules of mucous material, the mucosomes. The electron-dense part persists as a small calotte or crescent often irregularly structured at the periphery of the mucosome, and fragments of it are dispersed into the interior of the mucosomal body. The mucosomes are membrane-bound and contain small granules, 55 nm in diameter. At the end of vitellogenesis, the follicle cells are filled with mucosomes, and cytoplasmic residua can only sparingly be observed among them. Oocytic microvilli extend through the zona radiata and intermingle with follicular cell processes in the cleft between the zona radiata and the belt of mucosomes during growth of the oocyte. Capillaries develop in connective tissue of the theca layer as vitellogenesis proceeds. © 1993 Wiley-Liss, Inc.     

Ovarian 3-hydroxy-3-methylglutaryl-CoA reductase in Blattella germanica (L.): pattern of expression and critical role in embryogenesis

In the ovary of adult Blattella germanica, the enzyme 3-hydroxy-3-methylglutaryl-CoA reductase (HMG-CoA reductase) is highly expressed in mid-late vitellogenesis, suggesting a functional link of the mevalonate pathway with choriogenesis. The inhibitor of HMG-CoA reductase, fluvastatin, applied in females in late vitellogenesis, inhibits the activity of the enzyme in the ovary and in the developing embryos within the ootheca. This does not affect choriogenesis or ootheca formation but reduces the number of larvae per ootheca. Our results suggest that fluvastatin is incorporated into the oocytes and has delayed inhibitory effects on the oviposited eggs. HMG-CoA reductase is essential for embryogenesis, but not for chorion formation.     

Allatostatin in ovaries, oviducts, and young embryos in the cockroach Diploptera punctata

The quantity and localization of -Phe-Gly-Leu-amide allatostatins (-F-G-L-amide AST) was determined by ELISA and immunohistochemistry in ovaries and oviducts and in pre-dorsal closure embryos. AST in the cytoplasm of basal oocytes gradually increased from 4 to 35 fmol/ovary pair from the start (day 2) to the completion of vitellogenesis (day 6), then rapidly increased to 121 fmol/ovary pair during choriogenesis. In oviducts, AST-immunoreactivity was found in nerves to the muscle layer and in epithelial cells. AST-immunoreactivity in oviduct epithelial cells increased during vitellogenesis. A marked increase in quantity of AST in oviduct tissue between completion of chorion formation and immediately after ovulation appears to result from AST released from oocytes as they travel down the oviducts because AST content of newly ovulated eggs was 40% lower than late stage chorionated oocytes, and these oocytes released AST when incubated in saline. AST in embryos, localized in yolk cells, decreased as embryos approached dorsal closure. That this material in ovaries and embryos is AST was confirmed by its ability to inhibit JH synthesis in vitro and identification by MALDI-TOF mass spectrometry of a peptide with a mass corresponding to that of a Diploptera punctata AST. These findings indicate likely novel functions for ASTs: facilitation of ovulation and utilization of yolk.     

Morphology and ultrastructure of the somatic cells in Astacus leptodactylus ovary

We defined the somatic environment in which female germinal cells develop, and performed ultrastructural analyses of various somatic cell types, with particular reference to muscle cells and follicle cells, that reside within the ovary at different stages of oogenesis. Our findings show that ovarian wall of the crayfish is composed of long muscle cells, blood cells, blood vessels and hemal sinuses. The follicle and germinal cells lie within a common compartment of ovarian follicles that is defined by a continuous basal matrix. The follicle cells form branching cords and migrate to surround the developing oocytes. A thick basal matrix separates the ovarian interstitium from ovarian follicles compartment. Transmission electron microscopy shows that inner layer of basal matrix invaginates deeply into the ovarian compartment. Our results suggest that before being surrounded by follicle cells to form follicles, oogonia and early previtellogenic oocytes reside within a niche surrounded by a basal matrix that separates them from ovarian interstitium. We found coated pits and coated vesicles in the cortical cytoplasm of previtellogenic and vitellogenic oocytes, suggesting the receptor mediated endocytosis for transfer of material from the outside of the oocytes, via follicle cells. The interstitial compartment between the inner muscular layer of the ovarian wall and the basal matrix of the ovarian follicle compartment contains muscle cells, hemal sinuses, blood vessels and blood cells. Granular hemocytes, within and outside the vessels, were the most abundant cell population in the ovarian interstitium of crayfish after spawning and in the immature ovary. J. Morphol. 277:118–127, 2016. © 2015 Wiley Periodicals, Inc.     

Anatomy of the ovaries of the starfish Asterias rubens (Echinodermata)

Summary The ovaries of the starfish Asterias rubens were studied histologically and ultrastructurally. The reproductive system in female specimens consists of ten separate ovaries, two in each ray. Each ovary is made up of a rachis with lateral primary and secondary folds: the acini maiores and acini minores. The ovarian wall is composed of an outer and an inner part, separated by the genital coelomic sinus. The ovarian lumen contains oocytes in various phases of oogenesis, follicle cells, nurse cells, phagocytosing cells and steroid-synthesizing cells.Oogenesis is divided into four phases: (i) multiplication phase of oogonia, (ii) initial growth phase of oocytes I, (iii) growth phase proper of oocytes I, and (iv) post-growth phase of oocytes I. The granular endoplasmic reticulum and the Golgi complex of the oocytes appear to be involved in yolk formation, while the haemal system, haemal fluid and nurse cells may also be important for vitellogenesis. The haemal system is discussed as most likely being involved in synchronizing the development of the ovaries during the annual reproductive cycle and in inducing, stimulating and regulating the function of the ovaries.Steroid-synthesizing cells are present during vitellogenesis; a correlation between the presence of these cells and vitellogenesis is discussed.     

Differentiation and function of the ovarian somatic cells in the pseudoscorpion,Chelifer cancroides (Linnaeus, 1761) (Chelicerata: Arachnida: Pseudoscorpionida)

Pseudoscorpion females carry fertilized eggs and embryos in specialized brood sacs, where embryos are fed with a nutritive fluid produced and secreted by somatic ovarian cells. We used various microscopic techniques to analyze the organization of the somatic cells in the ovary of a pseudoscorpion, Chelifer cancroides. In young specimens, the ovary is a cylindrical mass of internally located germline cells (oogonia and early previtellogenic oocytes) and two types of somatic cells: the epithelial cells of the ovarian wall and the internal interstitial cells. In subsequent stages of the ovary development, the oocytes grow and protrude from the ovary into the hemocoel (opisthosomal cavity). At the same time the interstitial cells differentiate into the follicular cells that directly cover the oocyte surface, whereas some epithelial cells of the ovarian wall form the oocyte stalks – tubular structures that connect the oocytes with the ovarian tube. The follicular cells do not seem to participate in oogenesis. In contrast, the cells of the stalk presumably have a dual function. During ovulation the stalk cells appear to contribute to the formation of the external egg envelope (chorion), while in the post-ovulatory phase of ovary function they cooperate with the other cells of the ovarian wall in the production of the nutritive fluid for the developing embryos.     

Cytodifferentiation and vitellogenesis during oogenesis in arachnida: Cytological studies on developing oocytes of a harvestman

Light and electron microscope studies were made on harvestman oocytes during the course of their origin, differentiation, and vitellogenesis. The germ cells appear to originate from the ovarian epithelium. They subsequently migrate to the outer surface of the epithelium, where they remain attached often by means of stalk cells which suspend them in the hemocoel during oogenesis. The “Balbiani bodies,” “yolk nuclei,” or “nuage” constitute a prominent feature of young, previtellogenic oocytes, and take the form of large, but variable sizes of electron-dense cytoplasmic aggregates with small fibrogranular components. The cytoplasmic aggregates fragment and disperse, and cannot be detected in vitellogenic oocytes. The young oocytes become surrounded by a vitelline envelope that appears to represent a secretory product of the oocyte. The previtellogenic oocytes are impermeable to horseradish peroxidase under both in vivo and in vitro conditions. In addition to mitochondria, dictyosomes, and abundant ribosomes, the ooplasm of the previtellogenic oocyte acquires both vesicular and lamellar forms of the rough-surfaced endoplasmic reticulum. In many areas, a dense homogeneous product appears within the cisternae of the endoplasmic reticulum and represents nascent yolk protein synthesized by the oocyte during early stages of vitellogenesis. Later in vitellogenesis, the oocyte becomes permeable to horseradish peroxidase under both in vivo and in vitro conditions. This change is associated with a massive process of micropinocytosis which is reflected in the presence of large numbers of vesicles of variable form and structure in the cortical ooplasm. Both spherical and tubular vesicles are present, as are coated and uncoated vesicles. Stages in the fusion of the vesicles with each other and with developing yolk platelets are illustrated. In the harvester oocytes, vitellogenesis is a process that involves both autosynthetic and heterosynthetic mechanisms.     

The ultrastructure of oogenesis and yolk formation in Labidocera aestiva (Copepoda: Calanoida)

Yolk formation in the oocytes of the free-living, marine copepod, Labidocera aestiva (order Calanoida) involves both autosynthetic and heterosynthetic processes. Three morphologically distinct forms of endogenous yolk are produced in the early vitellogenic stages. Type 1 yolk spheres are formed by the accumulation and fusion of dense granules within vesicular and lamellar cisternae of endoplasmic reticulum. A granular form of type 1 yolk, in which the dense granules within the cisternae of endoplasmic reticulum do not fuse, appears to be synthesized by the combined activity of endoplasmic reticulum and Golgi complexes. Type 2 yolk bodies subsequently appear in the ooplasm but their formation could not be attributed to any particular oocytic organelle. In the advanced stages of vitellogenesis, a single narrow layer of follicle cells becomes more developed and forms extensive interdigitations with the oocytes. Extra-oocytic yolk precursors appear to pass from the hemolymph into the follicle cells and subsequently into the oocytes via micropinocytosis. Pinocytotic vesicles fuse in the cortical ooplasm to form heterosynthetically derived type 3 yolk bodies.     

An ultrastructural study of relationships between the ovarian haemal system,follicle cells,and primary oocytes in the sea star,Asterias rubens

Summary The genital haemal sinus, present throughout the gonad wall of sea stars, is supposed to be the site of ultimate accumulation of nutrients for the germinal epithelium. Early vitellogenic pear-shaped oocytes are attached to this sinus by stalk-like processes. The ultrastructure of this association and of the oocyte-follicle cell complex is described with emphasis on mechanisms involved in oocyte nutrition.The genital haemal sinus, and sometimes portions of the surrounding genital coelomic sinus, contain a fine granular ground substance and amoeboid cells. Material similar to the haemal ground substance also fills vacuities in the inner basal laminae of the haemal sinus and intervenes between this layer and adjacent germinal and follicle cells in the ovarian lumen.Vitellogenesis is first detectable as numerous vacuoles accumulate within the oocyte-stalk near the haemal sinus; they contain flocculent material and often fuse with adjacent lysosome-like vacuoles. As vitellogenesis proceeds, oocytes develop complex and tenuous connections with the haemal sinus. These consist of a network of pseudopodia that interdigitate with thin sheet-like extensions of follicle cells. These cells are attached to the oolemma by microfilamentous processes and contain regularly arranged concentrations of glycogen granules and well developed rough endoplasmic reticulum.It is concluded, (1) that follicle cells provide each oocyte with a compartmentalized microenvironment within the ovarian lumen, (2) that such compartments are intimately associated with the nutrient laden haemal sinus, and (3) that nutritive and vitellogenic substances, derived extragonadally and stored temporarily in the ovarian wall, can pass through the oocyte-stalk.     

Oogenesis in the leech Glossiphonia heteroclita (Annelida, Hirudinea, Glossiphonidae). I. Ovary structure and previtellogenic growth of oocytes

Glossiphonia heteroclita has paired ovaries whose shape and dimensions change as oogenesis proceeds: during early previtellogenesis they are small and club-shaped, whereas during vitellogenesis they broaden and elongate considerably. During early oogenesis (previtellogenesis), each ovary is composed of an outer envelope (ovisac) that surrounds the ovary cavity and is filled with hemocoelomic fluid, in which a single and very convoluted ovary cord is bathed. The ovary cord consists of germline cells, including nurse cells and young oocytes surrounded by a layer of elongated follicle cells. Additionally, follicle cells with long cytoplasmic projections occur inside the ovary cord, where they separate germ cells from each other. The ovary cord contains thousands of nurse cells. Each nurse cell has one intercellular bridge, connecting it to a central anucleate cytoplasmic mass, the cytophore (rachis); it in turn is connected by one intercellular bridge with each growing oocyte. Numerous mitochondria, RER cisternae, ribosomes, and Golgi complexes are transported from the nurse cells, via the intercellular bridge and cytophore, to the growing oocytes. Oogenesis in G. heteroclita is synchronous with all oocytes in the ovary in the same stage of oogenesis. The youngest observed oocytes are slightly larger than nurse cells, and usually occupy the periphery of the ovary cord. As previtellogenesis proceeds, the oocytes gather a vast amount of cell organelles and become more voluminous. As a result, in late previtellogenesis the oocytes gradually protrude into the ovary cavity. Simultaneously with oocyte growth, the follicle cells differentiate into two subpopulations. The morphology of the follicle cells surrounding the nurse cells and penetrating the ovary cord does not change, whereas those enveloping the growing oocytes become more voluminous. Their plasma membrane invaginates deeply, forming numerous broad vesicles that eventually seem to form channels or conducts through which the hemocoelomic fluid can easily access the growing oocytes.     

Ultrastructural and histochemical study of previtellogenic oogenesis in the desert lizard Scincus mitranus (Squamata,Sauropsida)

The structure of the granulosa in reptilian sauropsids varies between groups. We investigated the follicle development in the desert lizard Scincus mitranus. In the germinal bed, oogonia, and primary oocytes were identified and found to be interspersed between the epithelial cells. Previtellogenesis was divided into three stages: early, transitional, and late previtellogenic stages. During the early previtellogenic stage (diplotene), the oocyte is invested by small epithelia cells that formed a complete single layer, which may be considered as a young follicle. The transitional previtellogenic stage was marked by proliferation and differentiation of the granulosa layer from a homogenous layer consisting of only small cells to a heterogeneous layer containing three cell types: small, intermediate, and large cells. The late previtellogenic stage was marked by high-synthetic activity of large cells and the initiation of cytoplasmic bridges between large granulosa cells and the oocyte. Small cells were the only type of granulosa cells that underwent division. Thus, these cells may be stem cells for the granulosa cell population and may develop into intermediate and subsequently large cells. The intermediate cells may be precursors of large cells, as suggested by their ultrastructure. The ultrastructure of the large granulosa was indicative of their high synthetic activity. Histochemical analysis indicated the presence of cholesterol and phospholipids in the cytoplasm of large cells, the zona pellucida, among the microvilli, in the bridges region, and in the cortical region of the oocyte cytoplasm. These materials may be transferred from large cells into the oocyte through cytoplasmic bridges and provide nutritive function to large cells rather than functioning in steroidogenesis or vitellogenesis.