Electron microscope studies on developing oocytes of a coelenterate medusa with special reference to vitellogenesis

In a hydrozoan jellyfish, the female gonad is differentiated from a specialized region of the epidermis near the manubrium. Changes in the oocytes during growth and vitellogenesis are described as observed with electron microscopic and cytochemical techniques. Three major types of yolk are formed; these include lipid, glycogen, and membrane-bound granules consisting of both protein and carbohydrate. The latter first appear evident within vesicular and cisternal elements of the numerous Golgi complexes. The orientation and structural variations noted between the endoplasmic reticulum and forming face of the Golgi complexes suggest that the protein component of the yolk granules may be transferred from the cisternae of the endoplasmic reticulum to the Golgi complex where it is joined to carbohydrate perhaps synthesized by the Golgi complexes. Stages in the release of the precursor yolk material sequestered in cisternal elements of the Golgi complexes are illustrated. The presence of coated and uncoated vesicles in the Golgi regions and their possible role in intracellular transport are described and discussed. The presence and possible method of morphogenesis of vesiculate yolk bodies are also described. What appear to represent invaginations of the oolemma extend into the ooplasm and display a special orientation with respect to lamellae of the rough-surfaced endoplasmic reticulum. Intraooplasmic synthesis appears to constitute the major pathway for protein-carbohydrate yolk deposition.     

Oogenesis in the annelid Enchytraeus albidus with special reference to the origin and cytochemistry of yolk

In the annelid Enchytraeus albidus the ovary is composed of packets containing eight synchronously developing oocytes. Each oocyte in the packet is connected, via a bridge, to a common cytoplasmic mass. Developmental synchrony of oocytes within individual packets is probably related to the ooplasmic continuity. The young previtellogenic oocyte contains many polysomes, a few cisternae of smooth and rough endoplasmic reticulum, small Golgi complexes, and mitochondria. Many of the mitochondria are dumbbell-shaped and may thus represent division stages. Vitellogenesis is marked by the appearance of peripherally located lipid yolk and small, densely staining granules scattered throughout the ooplasm. There is an increase of smooth endoplasmic reticulum, mitochondria, and enlarged Golgi elements. Small multivesicular-like bodies, the early stages of developing yolk, are derived from the Golgi complex. The mature yolk sphere is bipartite and consists of (a) a variable number of dense spheres, the core bodies, which are produced in the ooplasm by the Golgi complex and which become embedded in (b) a dense matrix. The electron opaque tracer, horseradish peroxidase is incorporated into the oocyte and deposited in the matrix suggesting that this component of the yolk sphere is obtained by micropinocytosis. Enzyme digestions and various cytochemical techniques suggest that the core bodies are rich in carbohydrate, probably as glyco- or mucoproteins, and that the matrix is rich in lipid.     

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.     

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.     

Fine Structures of Oocytes and Accessory Cells of the Ovary in the Starfish Patiria (Asterina) pectinifera at Different Stages of Oogenesis and After 1-Methyladenine-Induced Maturation

Morphological changes in the growing and maturing oocytes of Patiria ( Asterina ) pectinifero were studied by electron microscopy. Oogenesis is of the solitary type. An extensive system of rough endoplasmic reticulum (ER) and Golgi complex (GC) develops in the ooplasm forming the cortical, yolk and secretory granules in its peripheral regions. The contents of the latter granules are released from the oocyte and form the vitelline membrane. At early stages of oogenesis, extensive multiplication of mitochondria results in formation of a large aggregate of these organelles in the perinuclear cytoplasm ("yolk nucleus"). After maturation of full grown oocytes has been induced by 1-methyladenine, the membranous cell structures are rapidly rearranged: vast aggregates of ER cisternae in the surface cytoplasm layer and single ER cisternae among yolk granules are disintegrated to small vesicles; the GC is reduced. These processes are suggested to be somehow related to changes in hydration of the cytoplasm and in rigidity of its surface layer. In maturing oocytes, the yolk granules form characteristic linear rows, trabeculae, traversing the cytoplasm and their boundary membranes fuse in zones of contact. Some granules are converted to multivesicular bodies, thus suggesting the activation of hydrolytic enzymes that form part of the yolk in echinoderms.     

Ultrastructural studies of differentiation in the oocyte of the polyclad turbellarian,Prostheceraeus floridanus

Oocyte differentiation in the polyclad turbellarian Prostheceraeus floridanus has been examined to determine the nature of oogenesis in a primitive spiralian. The process has been divided into five stages. (1) The early oocyte: This stage is characterized by a large germinal vesicle surrounded by dense granular material associated with the nuclear pores and with mitochondria. (2) The vesicle stage: The endoplasmic reticulum is organized into sheets which often contain dense particles. Vesicles are found in clusters in the cytoplasm, some of which are revealed to be lysosomes by treatment with the Gomori acid phosphatase medium. (3) Cortical granule formation: Cortical granules are formed by the fusion of filled Golgi vasuoles which have been released from the Golgi saccules. The association between the endoplasmic reticulum and Golgi suggests that protein is synthesized in the ER and transferred to the Golgi where polysaccharides are added to form nascent cortical granules. (4) Yolk synthesis: After a large number of cortical granules are synthesized, yolk bodies appear. They originate as small membrane-bound vesicles containing flocculent material which subsequently increase in size and become more compact. Connections between the forming yolk bodies and the endoplasmic reticulum indicate that yolk synthesis occurs in the ER. (5) Mature egg: In the final stage, the cortical granules move to the periphery and yolk platelets and glycogen fill the egg. At no time is there any evidence of uptake of macromolecules at the oocyte surface. Except for occasional desmosomes between early oocytes, no membrane specialization or cell associations are seen throughout oogenesis. Each oocyte develops as an independent entity, a conclusion supported by the lack of an organized ovary.     

东方杯叶吸虫卵黄腺和卵巢的超微结构研究

本文应用透射电镜观察了东方杯叶吸虫卵黄腺和卵巢的超微结构,并与体外培养成虫进行比较。根据形态特征和内含物的存在情况,将卵黄细胞和卵母细胞的发育均分为不同时期,详细描述了各期的形态特征,探讨了卵黄球和皮质颗粒等内含物的生理功能。体外培养成虫成熟卵黄细胞中有散在的卵黄物质,成熟卵母细胞中线粒体囊泡化,这些可作为体外培养的评价指标。     

An ultrastructural study of oogenesis in a marine triclad

The ultrastructural features of oocyte differentiation were studied in the marine triclad Cercyra hastata. Oocytes at several stages of maturation, each surrounded by follicle cell projections, are present within each of the two ovaries. A pre-vitellogenic and a vitellogenic stage have been detected in the oogenesis of C. hastata. The pre-vitellogenic stage is mainly characterized by an increase in the nuclear and nucleolar volume and activity, and the appearance and development of cortical granule precursors which are elaborated by the Golgi complex. In early phases of the vitellogenic stage, intense delamination and blebbing of the nuclear envelope occurs which probably contributes to an increase in number of cytoplasmic membranes and to transfer of nuclear material to the cytoplasm. The rough endoplasmic reticulum is extensively developed and often assumes a ‘whorl’ array. Several areas of yolk precursor formation appear in the whorls. Numerous 2–5 μm protein yolk globules are subsequently formed which appear surrounded by a double membrane (cisternae of the smooth endoplasmic reticulum) and become randomly distributed throughout the cytoplasm of mature oocytes. The peripheral ooplasm is occupied by a monolayer of electron-dense cortical granules. Finally, the evolutionary significance of the autosynthetic mechanism of yolk production is discussed.     

Differentiation of cytoplasmic organelles and storage of yolk during vitellogenesis in Hemidiaptomus ingens and Mixodiaptomus kupelwieseri (Copepoda,Calanoida)

These investigations concern two freshwater calanoid copepods Hemidiaptomus ingens and Mixodiaptomus kupelwieseri. The first aspect of the research relates to the processes involved in the formation and the differentiation of the ooplasmic organelles at the time of primary vitellogenesis. During this phase, a number of complex associations develop in the ooplasm. They consist chiefly of nuage-like structures, corresponding to extruded nuclear material, and vesicular formations, some arising from the nuclear envelope and the others neoformed in the ooplasm. These associations represent centers of maturation for ribosomes and synthesis for reticulum membranes. Annulate lamellae may be observed near these associations. Biogenesis of the reticulum always precedes the differentiation of the Golgi apparatus. Indeed, the dictyo-somes develop in characteristic complexes including endoplasmic reticulum cisternae and numerous vesicles resulting from intensive blebbing from cisternae. The second aspect of this research concerns yolk synthesis and accumulation of hyaloplasmic inclusions. A preliminary synthesis of yolk occurs early in these complexes and becomes more important after achievement of Golgi apparatus biogenesis. However, the most important yolk storage results from exogenous molecules and consists of complex globules, which develop into the ooplasm during secondary vitellogenesis. Formation of these globules is associated with the accumulation of two categories of inclusions in the hyaloplasm, i.e., lipid droplets and clusters of glycogen particles. At the end of vitellogenesis, a new type of endogenous material develops into small cisternae localized in the cortical ooplasm. © 1993 Wiley-Liss, Inc.     

Electron microscopic and autoradiographic studies on vitellogenesis in Necturus maculosus

Electron microscope studies on Necturus maculosus oocytes ranging in size from 1.1–1.5 mm in diameter indicate the primary proteinaceous yolk to arise within structures referred to in other amphibian oocytes as yolk precursor sacs or bodies. The origin of these yolk precursor sacs appears to result from the activity of the Golgi complexes which form multivesicular and granular-vesicular bodies, the limiting membrane of which is at times incomplete. During differentiation, the yolk precursor sacs contain small vesicles similar in size to Golgi vesicles, larger vesicles similar to vesicular elements of the agranular endoplasmic reticulum and, on occasion, a portion of a mitochondrion. The interior of these sacs becomes granular, perhaps by a dissolution of the components just described, and soon becomes organized into a crystalline configuration. In oocytes 2.0–2.5 mm in diameter, an extensive micropinocytotic activity begins, continues throughout vitellogenesis, and constitutes the primary mechanism for the formation of secondary yolk protein. Numerous coated and smooth-surfaced vesicles, as well as electron-dense and electronlucent ones, fuse in the cortical ooplasm to form progressively larger yolk platelets.     

Ultrastructural studies on the formation of yolk granules in the statoblast of a fresh-water bryozoan,Pectinatella gelatinosa

The formation of protein-carbohydrate yolk in the statoblast of a fresh-water bryozoan, Pectinatella gelatinosa, was studied by electron microscopy. Two types (I and II) of yolk cells were distinguished. The type I yolk cells are mononucleate and comprise a large majority of the yolk cells. The type II yolk cells are small in number; they become multinucleate by fusion of cells at an early stage of vitellogenesis. In both types of yolk cells, electron-dense granules (dense bodies) are formed in Golgi or condensing vacuoles, which are then called yolk granules. For the formation of yolk granules, the following processes are considered: 1. Yolk protein is synthesized in the rough-surfaced endoplasmic reticulum (RER) of the yolk cells. 2. The synthesized protein condenses in the cisternal space of the RER and is packaged into small oval swellings, which are then released from the RER as small vesicles (Golgi vesicles, 300-600 A in diameter). 3. The small vesicles fuse with one another to form condensing vacuoles, or with pre-existing growing yolk granules. 4. In the matrix of the condensing vacuoles or growing yolk granules, electron-dense fibers are fabricated and then arranged in a paracrystalline pattern to form the dense body. 5. After the dense body reaches its full size, excess membrane is removed and eventually the yolk granules come to mature. Toward the end of vitellogenesis of the yolk cells, the cytoplasmic organelles are ingested by autophagosomes derived from multivesicular bodies and disappear.     

Mechanisms of protein yolk synthesis and deposition in crustacean oocytes

Summary Electron microscope studies on the oocytes of several crustacean species demonstrate that the protein yolk arises within vesicular and lamellar forms of the rough-surfaced endoplasmic reticulum. The vesicular form of the endoplasmic reticulum may have its origin from a blebbing process of the outer layer of the nuclear envelope. Disc-shaped granules, representing precursor elements of the yolk granules, appear within the vesicular and lamellar profiles of endoplasmic reticulum. Autoradiographic results suggest that the ribosomes attached to the endoplasmic reticulum take part in the biosynthesis of yolk proteins. Numerous disc-shaped granules accumulate within the cisternae of the endoplasmic reticulum, but eventually they undergo a transformation into a finely granular yolk granule. Thus, both the origin and growth of protein yolk granules occur within membranes constituting the endoplasmic reticulum. The results provide evidence that intra-ooplasmic synthesis of yolk protein occurs in these oocytes.This investigation was supported by research grants (HD-00699; GM-09229) and a Career Development Award (GM-11,524) from the National Institutes of Health, U.S. Public Health Service.     

Oogenesls in the terrestrial hermit crab,coenobita clypeatus (decapoda,anomura): II. Vitellogenesis

Oocytes from the land hermit crab, Coenobita clypeatus, in various stages of vitellogenesis were examined by light and electron microscopy. Early vitellogenic oocytes are characterized by accumulations of discrete vesicles of endoplasmic reticulum in the perinuclear cytoplasm. As oocytes develop, the endoplasmic reticulum becomes abundant, and numerous Golgi complexes are seen. There is a well developed Golgi-endoplasmic reticulum interaction. Within the confines of the reticulum are discrete intracisternal granules, which can be seen coalescing into electron-dense yolk bodies. Lipid accumulation is seen throughout the cytoplasm. Coincident with the burst of intra-oocytic metabolism are oolemma modifications and micropinocytosis, which provide ultrastructural evidence for extra-oocytic yolk production. The mature oocyte contains numerous yolk and lipid vesicles of varying electron density that comprise both intra- and extra-oocytic substrates.     

Ultrastructural and cytochemical aspects of the germarium and the vitellarium in Syndesmis patagonica (Platyhelminthes,Rhabdocoela, Umagillidae)

The cytoarchitecture of the female gonad of the endosymbiont umagillid Syndesmis patagonica has been investigated using electron microscopy and cytochemical techniques. The female gonad consists of paired germaria and vitellaria located behind the pharynx in the mid‐posterior region of the body. Both the germaria and the vitellaria are enveloped by an outer extracellular lamina and an inner sheath of accessory cells which contribute to the extracellular lamina. Oocyte maturation occurs completely during the prophase of the first meiotic division. Oocyte differentiation is characterized by the appearance of chromatoid bodies and the development of endoplasmic reticulum and Golgi complexes. These organelles appear to be involved in the production of round granules, about 2–2.5 μm in diameter, with a homogeneous electron‐dense core surrounded by a granular component and a translucent halo delimited by a membrane. These egg granules migrate to the periphery of mature oocytes, are positive to the cytochemical test for polyphenol detection, are unaffected by protease and have been interpreted as eggshell granules. The mature oocytes also contain a small number of yolk granules, lipid droplets, and glycogen particles scattered throughout the ooplasm. The vitellaria are branched organs composed of vitelline follicles with vitellocytes at different stages of maturation. Developing vitellocytes contain well‐developed rough endoplasmic reticulum and small Golgi complexes involved in the production of eggshell and yolk globules. Eggshell globules are round, measure 4–5 μm in diameter, and have a mosaic‐like patterned content which contains polyphenols. The yolk globules, 2–3 μm in diameter, show a homogeneous protein content of medium electron density, devoid of polyphenols, and completely digested by protease. The mature vitellocytes also contain glycogen as further reserve material. The presence of polyphenolic eggshell granules in the oocytes and of polyphenolic eggshell globules with a mosaic‐like pattern in the vitellocytes have been considered apomorphic features of the Rhabdocoela + Prolecithophora. J. Morphol. 275:703–719, 2014. © 2014 Wiley Periodicals, Inc.     

Electron microscopical observations on vitellocytes and germocytes inNematoplana coelogynoporoides (Plathelminthes,Proseriata)

Summary Fine-structural features of vitellaria and germaria inNematoplana coelogynoporoides are documented and compared with those of other free-living plathelminths with ectolecithal eggs. Emphasis is mainly put on the pattern of eggshell material, yolk bodies deposited in vitellocytes, and marginal granules of the female germ cells. In this species, encompassed in the taxon Proseriata Unguiphora, the eggshell granules show a meandering pattern also known from species of the taxon Proseriata Lithophora. In contrast, the yolk globules exhibit crystalline components unknown from the Lithophora. The marginal granules in the ooplasm have an extremely large diameter. They consist of a flocculent core and a crescent-shaped cortex. Marginal granules of this appearance have not been found in any other taxon of free-living Neoophora.Abbreviations cc crystalline component - co cortex - gER granular endoplasmic reticulum - go Golgi complex - gl glycogen - lp lipid droplet - mg marginal granule - n nucleus - nl nucleolus - sg eggshell granule - sp spermatozoa - yg yolk globule     

Previtellogenic and vitellogenic oocytes in ovarian follicles of cultured siberian sturgeon Acipenser baerii (Chondrostei,Acipenseriformes)

Previtellogenic and vitellogenic oocytes in ovarian follicles from cultured Siberian sturgeon Acipenser baerii were examined. In previtellogenic oocytes, granular and homogeneous zones in the cytoplasm (the ooplasm) are distinguished. Material of nuclear origin, rough endoplasmic reticulum, Golgi complexes, complexes of mitochondria with cement and round bodies are numerous in the granular ooplasm. In vitellogenic oocytes, the ooplasm comprises three zones: perinuclear area, endoplasm and periplasm. The endoplasm contains yolk platelets, lipid droplets, and aggregations of mitochondria and granules immersed in amorphous material. In the nucleoplasm, lampbrush chromosomes, nucleoli, and two types of nuclear bodies are present. The first type of nuclear bodies is initially composed of fibrillar threads only. Their ultrastructure subsequently changes and they contain threads and medium electron dense material. The second type of nuclear bodies is only composed of electron dense particles. All nuclear bodies impregnate with silver, stain with propidium iodide, and are DAPI‐negative. Their possible role is discussed. All oocytes are surrounded by follicular cells and a basal lamina which is covered by thecal cells. Egg envelopes are not present in previtellogenic oocytes. In vitellogenic oocytes, the plasma membrane (the oolemma) is covered by three envelopes: vitelline envelope, chorion, and extrachorion. Vitelline envelope comprises four sublayers: filamentous layer, trabecular layer 2 (t2), homogeneous layer, and trabecular layer 1 (t1). In the chorion, porous layer 1 and porous layer 2 are distinguished in most voluminous examined oocytes. Three micropylar cells that are necessary for the formation of micropyles are present between follicular cells at the animal hemisphere. J. Morphol. 278:50–61, 2017. ©© 2016 Wiley Periodicals,Inc.     

The female gonad in two species of Microplana (Platyhelminthes,Tricladida, Rhynchodemidae): Ultrastructural and cytochemical investigations

The female gonad of the land planarians Microplana scharffi and Microplana terrestris consists of two small germaria located ventrally in the anterior third of the body and of two ventro‐lateral rows of oblong vitelline follicles distributed between the intestinal pouches. Both these structures are enveloped by a tunica composed of an outer extracellular lamina and an inner sheath of accessory cells. Oocyte maturation is characterized by the appearance of chromatoid bodies and the development of endoplasmic reticulum and Golgi complexes. These organelles appear to be correlated with the production of egg granules with a fenestrated/granular content of medium electron density, about 4–5 μm in diameter, which remain dispersed in the ooplasm of mature oocytes. On the basis of cytochemical tests showing their glycoprotein composition, and their localization in mature oocytes, these egg granules have been interpreted as yolk. In the vitelline follicles, vitellocytes show the typical features of secretory cells with well‐developed rough endoplasmic reticulum and Golgi complexes involved in the production of eggshell globules and yolk. The eggshell globules, which appear to arise from repeated coalescences of two types of Golgi‐derived vesicles, contain polyphenols and, when completely mature, they measure about 1–1,2 μm in diameter and show a meandering/concentric content pattern as is typical of the situation observed in most Proseriata and Tricladida. Mature vitellocytes also contain a large amount of glycogen and lipids as further reserve material. On the basis of the ultrastructural features of the female gonad and in relation to the current literature the two species of rhynchodemids investigated appear to be closely related to the freshwater planarians belonging to the family Dugesiidae. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Electron microscope studies on the oocyte of the fresh-water mussel (Anodonta), with special reference to the stalk and mechanism of yolk deposition

An interesting relationship exists between the ovary and the developing oocyte in the fresh-water mussel. As the oocytes grow, they elongate and bulge into the ovarian cavity. In the early stages, the nucleus migrates from the attached region (“foot”) to the distal region of the cell. With continued growth and maturation the connection between the proximal “foot” and distal nucleated portion becomes reduced to a narrow stalk. Microtubules appear in the young oocytes as they start to elongate and become packed in the stalks of older oocytes. It is suggested that the microtubules function as supporting structures and possibly also as channels for the transfer of materials from one portion of the oocyte to the other. The fine structure of the oocyte reveals evidence that the developing yolk bodies or spheres are formed, in part at least, by the incorporation of many smaller “precursor yolk vesicles.” These appear in the region of the Golgi complex and are presumed to be derived from the Golgi saccules. The oocyte contains an unusually well developed endoplasmic reticulum whose cisternae are filled with a rather conspicuous material.     

Cytochemical characterization of the endomembranous system during the oocyte primary growth in Serrasalmus spilopleura (Teleostei, Characiformes, Characidae)

The morphophysiological changes that occur during oocyte primary growth in Serrasalmus spilopleura were studied using ultrastructural cytochemical techniques. In the previtellogenic oocytes endoplasmic reticulum components, Golgi complex cisternae and vesicles, lysosomes, multivesicular bodies and some electron-dense vesicles react to acid phosphatase (AcPase) detection. The endoplasmic reticulum components, Golgi complex cisternae and vesicles also react to osmium tetroxide and potassium iodide impregnation (KI). These structures, except for the Golgi complex cisternae, are strongly contrasted by osmium tetroxide and zinc iodide impregnation (ZIO). Some electron-dense vesicles are ZIO-stained, while microvesicles in the multivesicular bodies and other large isolated cytoplasmic vesicles are contrasted by KI. At primary oocyte growth, the activity of the endomembranous system and the proliferation of membranous organelles are intense. The biosynthetic pathway of the lysosomal proteins such as acid phosphatase, involves the endoplasmic reticulum, Golgi complex, vesicles with inactive hydrolytic enzymes and, finally, the lysosomes. The oocyte endomembranous system have reduction capacity and are involved in the metabolism of rich in SH groups.     

Oogenesis in the leech Glossiphonia heteroclita (Annelida, Hirudinea, Glossiphoniidae) II. Vitellogenesis, follicle cell structure and egg shell formation

By the end of previtellogenesis, the oocytes of Glossiphonia heteroclita gradually protrude into the ovary cavity. As a result they lose contact with the ovary cord (which begins to degenerate) and float freely within the hemocoelomic fluid. The oocyte's ooplasm is rich in numerous well-developed Golgi complexes showing high secretory activity, normal and transforming mitochondria, cisternae of rER and vast amounts of ribosomes. The transforming mitochondria become small lipid droplets as vitellogenesis progresses. The oolemma forms microvilli, numerous coated pits and vesicles occur at the base of the microvilli, and the first yolk spheres appear in the peripheral ooplasm. A mixed mechanism of vitellogenesis is suggested. The eggs are covered by a thin vitelline envelope with microvilli projecting through it. The envelope is formed by the oocyte. The vitelline envelope is produced by exocytosis of vesicles containing two kinds of material, one of which is electron-dense and seems not to participate in envelope formation. The cortical ooplasm of fully grown oocytes contains many cytoskeletal elements (F-actin) and numerous membrane-bound vesicles filled with stratified content. Those vesicles probably are cortical granules. The follicle cells surrounding growing oocytes have the following features: (1) they do not lie on a basal lamina; (2) their plasma membrane folds deeply, forming invaginations which eventually seem to form channels throughout their cytoplasm; (3) the plasma membrane facing the ovary lumen is lined with a layer of dense material; and (4) the plasma membrane facing the oocyte forms thin projections which intermingle with the oocyte microvilli. In late oogenesis, the follicle cells detach from the oocytes and degenerate in the ovary lumen.