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.     

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.     

Ultrastructural aspects of the germovitellarium of two prorhynchids (Platyhelminthes,Lecithoepitheliata)

Summary

The female gonad of two fresh-water prorhynchids, Geocentrophora baltica and Prorhynchus stagnalis, has been investigated by means of conventional electron microscopy and cytochemical techniques. Both species have an unpaired germovitellarium located under the gut; accessory cells surround the germovitellarium of G. baltica. The germovitellarium consists of a restricted germinative area where early differentiating oocytes and vitellocytes are randomly associated, and an extensive growth area with follicular organization. Each follicle consists of a single alecithal oocyte surrounded by numerous vitellocytes. The main features of oocyte differentiation are the accumulation of lipid droplets and the appearance of Golgi complexes and small bodies possibly representing secondary lysosomes. Vitellocytes show features typical of secretory cells, including well-developed rough endoplasmic reticulum (RER) and Golgi complexes which are involved in the production of type A and type B inclusions, hi both species, type A inclusions appear first, have a glycoprotein content, do not contain polyphenols, and become localized in the peripheral cytoplasm of mature vitellocytes; they have been interpreted as eggshell forming granules. Type B inclusions are larger, have a proteinaceous content with a different structure in the two species examined, and remain scattered in the cytoplasm of mature vitellocytes; they are considered to be yolk. The finding of eggshell forming granules without polyphenols in prorhynchids contrasts with the condition in most platyhelminths that have a sclerotized eggshell formed through a tanning process of polyphenolic substances. The small bodies in the oocytes and the eggshell granules in the vitellocytes of Lecithoepitheliata differ from those observed in prolecithophorans, which have oocyte and vitellocyte inclusions similar to those of the Rhabdocoela.     

Ultrastructural and cytochemical studies of the female gonad of Prorhynchus sp. (Platyhelminthes,Lecithoepitheliata)

The female gonad of Prorhynchus is heterocellular (neoophoran organization) and consists of an unpaired, elongate germovitellarium enveloped by a finely granular extracellular lamina. It is composed of a posterior germinative area where early oocytes are randomly associated with differentiating vitellocytes and a growth area with follicular organization. In each follicle a single oocyte is surrounded by a layer of vitellocytes. By electron microscopy, the oocytes showed features typical of non-vitellogenic germ cells; they had chromatoid bodies, annulate lamellae, lipid droplets and R.E.R. and Golgi complexes producing small granules with a multilamellar pattern. Vitellocytes showed features typical of secretory cells with the R.E.R. and Golgi complex developed to a great extent and involved in the production of type A and type B globules, respectively. We speculate that type A globules are shell-globules and type B globules are yolk. The structure, composition and role of vitellocyte globules of Prorhynchus are compared with those of homologous inclusions from other Platyhelminthes.Abbreviations A type A globule - B type B globule - ECL extracellular lamina - GC Golgi complex - L lipid - RER rough endoplasmic reticulum - O oocyte - V vitellocyte     

Ultrastructure of the female gonad of two temnocephalids (Platyhelminthes,Rhabdocoela)

The female gonad of Temnocephala dendyi and T. minor consists of a single germarium and two rows of vitellaria. It is enveloped by an extracellular lamina and accessory cells. Accessory cells are only peripherally located in the germarium while their cytoplasmic projections also fill the spaces between vitellocytes in the vitellarium. The main feature of oocyte maturation is the appearance of chromatoid bodies and the development of rough endoplasmic reticulum (R.E.R.) and Golgi complexes which appear to be correlated with the production of double-structured egg granules. The egg granules, which are localized in the cortical cytoplasm of mature oocytes, contain glycoproteins, are devoid of polyphenols and are similar in structure and composition to the cortical granules observed in some Digenea and Monogenea. Vitellocytes are typical secretory cells with well-developed R.E.R. and Golgi complexes which are involved in the production of shell globules and yolk. The multigranular pattern and the polyphenolic composition of the shell globules of the temnocephalids investigated are similar to those observed in other rhabdocoels, and in some Prolecithophora and Neodermata. This feature may represent a synapomorphy shared by these taxa. This revised version was published online in July 2006 with corrections to the Cover Date.     

An ultrastructural study of oogenesis in the planarian Schmidtea mediterranea (Platyhelminthe, Paludicola)

The ovary of the freshwater planarian Schmidtea mediterranea has been studied for the first time using both light and electron microscopy methods. The ultrastructure of the ovary revealed two types of cells: accessory cells and germinal cells at various stages of differentiation, distributed along a maturation axis. Initially, oogonia underwent cytoplasm growth due to the development of organelles, such as endoplasmic reticulum, Golgi complex, and mitochondria, which are all involved in the production of cytoplasmic inclusions or yolk globules. It is shown that the chromatoid body and fibrogranular aggregates may participate in the synthesis of vitelline inclusions. When completely mature, the oocytes have become larger, due to the accumulation of nutritive inclusions, which are round in shape and have a paracrystalline structure. These inclusions are interpreted as being yolk globules and may represent a kind of nutritive material for the developing embryo. These ultrastructural features of the ovary agree with the available phylogenetic tree, based on morphological and karyological characters that considers Schmidtea group as a genus and not a subgenus. The presence of sperm between the oocytes suggests that fertilization may occur within the ovary, representing an uncommon condition within the Triclads, in which fertilization usually takes places outside of the ovaries.     

Ultrastructural and cytochemical aspects of oogenesis in Castrada viridis (Platyhelminthes, Rhabdocoela).

The heterocellular female gonad of the typhloplanoid Castrada viridis consists of a single germarium and two rows of vitellaria. The germarium, composed of a germinative zone and a growth zone, is surrounded externally by a layer of accessory cells the function of which is hypothesized in this study. The main feature of oocyte differentiation is the synthesis of small electron-dense inclusions produced by the rough endoplasmic reticulum (R.E.R.) and Golgi complex. The electron-dense content of the egg inclusions reacts positively to the cytochemical test used to detect polyphenols and is only partially extracted following incubation in protease. The genesis, composition, and peripheral location of egg inclusions in mature oocytes suggest that they could represent residual eggshell granules. The presumed function of eggshell granules is discussed and their fine morphology is compared with that observed in other neoophoran Platyhelminthes.     

Ultrastructural and cytochemical studies on the germarium of Dicrocoelium dendriticum (Plathelminthes,Digenea)

Summary The unpaired germarium of Dicrocoelium dendriticum contains many female germ cells at different stages of maturation and is enveloped by a fibrous basal lamina-like structure and a multilayered cytoplasmic sheath whose origins and functions are discussed. The maturation process of primary oocytes occurs completely within the prophase of the first meiotic division. It has been divided into three stages, as previously suggested for monogeneans. Stage I corresponds to oogonia and early oocytes which are located in the distal germinative area of the gonad. These cells are characterized by a high nucleo/cytoplasmic ratio and a poorly differentiated cytoplasm. Stage II corresponds to maturing oocytes grouped in the central area of the gonad and exhibiting long synaptonemal complexes and a prominent nucleolus. The main feature of cytoplasmic differentiation is the increase in the number of RER and Golgi complex which are involved in the production of small electron-dense granules. Stage III corresponds to mature oocytes located in the proximal area of the germarium near the origin of the oviduct. In this stage, the granules become regularly distributed in a monolayer in the peripheral ooplasm and make contact with the oolemma. They show a distinctive complex structure, are composed of proteins and glycoproteins and do not contain polyphenols. Their possible role as cortical granules is discussed in relation to chemical composition and previous studies on other Plathelminthes. Neither yolk globules nor glycogen are present in the oocytes.Abbreviations I oogonium and early oocyte - II growing oocyte - III mature oocyte - cg cortical granule - cs cytoplasmic sheath - db dense body - ecm extra cellular matrix - ER endoplasmic reticulum - fl fibrous extracellular layer - gc Golgi complex - m mitochondria - N nucleus - nu nucleolus - RER rough endoplasmic reticulum - sc synaptonemal complex     

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.     

The evolution of the female gonad in Platyhelminthes-Turbellaria: ultrastructural investigations

Summary

The structure of the female gonad has undergone important evolutionary modifications in Platyhelminthes-Turbellaria. The primitive condition consists of single oocytes freely distributed in the parenchyma autosynthetically producing both yolk and eggshell forming granules (e.f.g.) (archoophoran level of organization). A derived condition is the presence of a compact, hetero-cellular gonad enveloped by a cellular tunica and/or an extracellular lamina-like layer and composed of germaria with alecithal oocytes and vitellaria with vitelline or yolk cells producing and accumulating both yolk and e.f.g. (neoophoran level of organization). In the last three decades the female gonad of a number of turbellarian taxa has been investigated by means of electron microscopy and cytochemistry, and several characters of oocytes and vitellocytes have been hypothesized to have a potential phylogenetic value. Some of these characters and their possible phylogenetic implications are briefly reviewed.     

Ultrastructure of growing oocytes and accessory cells in Austrognathia (Gnathostomulida, bursovaginoidea)

The ovary of Austrognathia cf. riedli consists of 4-6 oocytes which are located in the mid-body region between the epidermis and the gut epithelium. The ovary is not enveloped by a tunica and each growing oocyte is surrounded by one or more accessory cells, the function of which is hypothesized in this study. Oogenesis is not synchronous and can be subdivided into a previtellogenic phase and a vitellogenic phase. Previtellogenic oocytes undergo a number of cell differentiations which consist mainly of an increase in size of the nucleus and nucleolus and the appearance in the cytoplasm of chromatoid bodies, annulate lamellac and short cisternae of rough endoplasmic reticulum (RER). Vitellogenic oocytes are characterized by the increase of RER, the appearance of numerous Golgi complexes and the accumulation of electron-dense globules, glycogen and lipid droplets. The electron-dense globules have been interpreted as yolk on the basis of both their localization and composition. Yolk synthesis occurs mainly by an endogenous mechanism and, to a lesser extent, by micropinocytosis. No shell-granules have been identified in the oocytes. The present ultrastructural findings are discussed in comparison with those from other lower metazoans.     

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     

Ultrastructural features of oogenesis in a free-living marine Platyhelminth, Vorticeros luteum

The female gonad of the prolecithophoran turbellarian Voticeros luteum has a heterocellular organization as is typical in the neoophoran Platyhelminthes. It is composed of an ovarian and a vitellarian area that are not well separated from one another and from the surrounding tissues by a tunica. Each oocyte during maturation becomes completely surrounded by a single accessory cell whose function has been hypothesized on the basis of the present investigation. The main feature in oocyte differentiation appears to be the synthesis and accumulation of small electron-dense inclusions; these are about 1-2 mum in size, are elaborated by the Golgi apparatus and have a content which reacts partially to the cytochemical test for polyphenols. The genesis, composition and cortical localization of the inclusions suggest that they could be homologous to the egg-shell granules present in the oocytes of 'archoophoran' species. A comparative analysis and discussion of the oocyte inclusions observed to date in other turbellarians is undertaken and the hypothetical use of these structures as characters providing some new insights in clarifying the phylogenetic relationships among free-living Platyhelminthes is finally discussed.     

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.     

Oogenesis in Actinoposthia beklemischevi (Platyhelminthes, Acoela): an ultrastructural and cytochemical study

The oogenesis of the acoel Actinoposthia beklemischevi can be divided into a previtellogenic and a vitellogenic stage. Maturing oocytes are surrounded by accessory cells (a.c.) that produce electrondense granules, the content of which is released into the space between the oocyte and a.c. and gives rise to a thin primary egg envelope. The a.c. may also contribute to yolk synthesis by transferring low molecular weight precursors to the oocyte. Two types of inclusion are produced in maturing oocytes. Type I inclusions are small, roundish granules produced by the Golgi complex. They have a proteinaceous non-polyphenolic content which is discharged in the intercellular space and produce a thicker secondary egg envelope. Type I inclusions represent eggshell-forming granules (EFGs). Type II inclusions are variably sized globules progressively changing their shape from round to crescent. They appear to be produced by the ER, contain glycoproteins and remain scattered throughout the cytoplasm in large oocytes. Type II inclusions represent yolk. The main features of oogenesis in Actinoposthia are: (a) EFGs have a non-polyphenolic composition; (b) the egg envelope has a double origin and is not sclerotinized; (c) yolk production appears to be autosynthetic. The present ultrastructural findings are compared with those from other Acoelomorpha and Turbellaria.     

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.     

Oogenesis in pig ovaries during the prenatal period: ultrastructure and morphometry

Oogenesis in fetal pig ovaries comprises the successive changes from the primordial germ cells to the dictyotene oocytes in primordial ovarian follicles. In this study the observations were carried out with an electron microscope and stereological analysis was performed. At the ultrastructural level there are no differences between the primordial germ cells and oogonia, but oogonia are connected with the intercellular bridges. The onset of the dictyotene phase was accompanied by the changes in the cytoplasm of oocytes. Near the nucleus, the yolk nucleus is formed containing numerous Golgi bodies, endoplasmic reticulum (ER), mitochondria and granules. ER proliferates in contact with the external leaflet of the nuclear envelope forming the narrow ER cisterns. Between the nuclear envelope and ER cisterns, the vesicles with grey content are visible. The proliferating ER forms numerous concentric cisterns around the nucleus. Next, the most external cisterns fragment, detach, and then form the cup-like structures. These structures separate the distinct areas of cytoplasm-compartments, which contain mitochondria, ribosomes and lipid droplets. The cells of cortical sex cords of the ovary, which encloses the oocyte, form the follicles. The volume of oocytes in forming follicle increases due to the increase in the number of the cell inclusions: lipid droplets, vacuoles and yolk globules. In the oocytes of primordial ovarian follicles, the compartments are transformed into the yolk globules, which are encountered by a sheath of ER cisterns and the grey vesicles; they contain the mitochondria, lipid droplets and light vacuoles. The role of the compartments and yolk globules as metabolic units is discussed in comparison with similar structures of the mature eggs of pigs and other mammal species.     

Vitellogenesis of basal trematode Aspidogaster limacoides(Aspidogastrea: Aspidogastridae)

The vitellogenesis of the trematode Aspidogaster limacoides (Aspidogastrea: Aspidogastridae), a parasite of cyprinid fishes, is described here using transmission electron microscopy. Four different stages of vitellocytes are differentiated: immature vitellocytes, early maturing vitellocytes, advanced maturing vitellocytes and mature vitellocytes. The process follows the same general pattern already described in other free-living neoophorans and parasitic flatworms (i.e. Trematoda, Monogenea and Cestoda): differentiation into mature vitelline cells involves the development of mitochondria, granular endoplasmic reticulum, Golgi complexes, lipid droplets and shell-globules. Mature vitellocytes of A. limacoides are composed of numerous shell-globule clusters, few lipid droplets and glycogen granules. They differ from those of another aspidogastrean Rugogaster hydrolagi in that they possess numerous globules tightly packed and by the presence of only one type of vitelline material. The interstitial tissue of vitelline follicles of A. limacoides contains a peripheral nucleus and long cytoplasmic projections extending between vitelline cells. Since aspidogastreans are considered as an archaic group of parasitic flatworms and thus have a strategic phylogenetic position, future works needs to pay special attention to the ultrastructural and chemical composition of mature vitellocytes within this basal group of trematodes.     

Reproduction and sexual development in a freshwater gastrotrich

Summary Post-embryonic development of parthenogenic eggs of Lepidodermella squammata was studied by light and electron microscopy in animals of known age and reproductive history. Each bilateral gonad initially contains eight cells. No mitotic proliferation occurs during parthenogenic egg development. Germ cells are tightly clustered, have smooth plasma membranes with no interconnections, and are uninucleate. There is no surrounding ovary or oviduct. At hatching, two cells in each gonad are identifiable as parthenogenic eggs. The enlarged nucleolus of the most mature egg has already attained the morphology that persists throughout vitellogenesis, with intertwined granular and fibrillar threads. Less mature eggs have earlier stages of nucleolar development, and lack indications of meiotic events. Parthenogenic eggs enter vitellogenesis singly, with formation of RER and active Golgi complexes, and the accumulation of lipid, yolk, and various granules. The shell is formed in situ, whereas the spines elongate after egg deposition. Most animals produce four parthenogenic eggs, which undergo immediate development (tachyblastic eggs). Resting (opsiblastic) eggs are rare in isolation culture. Both types of eggs are produced only prior to the formation of sperm and primary oocytes. The absence of synaptonemal complexes, which would indicate synapsis of homologous chromosomes in prophase of meiosis I, implies that parthenogenesis is by apomixis in L. squammata.     

Lysosomes in the cessation of vitellogenin secretion by the mosquito fat body; selective degradation of Golgi complexes and secretory granules

A massive and selective degradation of Golgi complexes, secretory granules, and RER is the mechanism responsible for the rapid termination of Vg secretion by trophocytes of the mosquito fat body. These cells are involved in an intensive synthesis of a glycoprotein, vitellogenin (Vg), which is accumulated by developing oocytes as yolk protein. Previously, assays for lysosomal enzymes have demonstrated that the cessation of Vg synthesis is characterized by a sharp increase in lysosomal activity; and fluorescent microscopy has shown that, during this intense lysosomal activity, Vg concentrates in lysosomes. In this report, electron microscopy combined with cytochemistry for lysosomal enzymes and localization of Vg with colloidal gold immunocytochemistry has shown that this lysosomal activity is directed towards selective degradation of Vg and organelles associated with its synthesis and secretion. Three organelles undergo lysosomal breakdown: the Golgi complex, Vg-containing secretory granules, and RER. The degradation of Golgi complexes occurs in two steps similar to that for RER: first, the organelle is sequestered by double isolation membranes, and the resulting pre-lysosome then fuses with a primary or secondary lysosome. In contrast, mature Vg-containing secretory granules fuse with lysosomes directly. This combination of crino- and autophagy is a specific, highly intense, and precisely timed event.