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Vitellogenesis in Wenyonia virilis was examined by transmission electron microscopy (TEM), including the cytochemical detection of glycogen at the ultrastructural level with the periodic acid-thiosemicarbazide-silver proteinate (PA-TSC-SP) technique. Mature vitelline follicles have cells in various stages of development, progressing from immature cells of gonial type near the periphery of the follicle to maturing and mature vitellocytes towards the centre. Maturation is characterized by: (1) increase in cell volume; (2) increase in nuclear surface area restoring the N/C (nucleo-cytoplasmic) ratio; (3) nucleolar transformation; (4) extensive development of parallel cisternae of GER, the shell-protein producing units; (5) development of Golgi complexes, engaged in shell-granule/shell-globule formation and packaging; (6) synthesis and storage of glycogen in the cytoplasm; (7) simultaneous, independent formation and storage of intranuclear glycogen; (8) continuous fusion of small shell-granules into larger shell-globules and fusion of these into large shell-globule clusters with a progressive increase in the number and size of the latter; and (9) disintegration of GER in the medial layer of vitellocyte cytoplasm, degenerative changes and accumulation of glycogen and shell-globule clusters within the cytoplasm. The functional significance of numerous shell-globule clusters and the relatively small amount of nuclear and cytoplasmic glycogen is analysed. Unlike vitellogenesis of other caryophyllids, the nuclear glycogen of mature vitellocytes in W. virilis is randomly dispersed in the nucleoplasm and never forms a high central accumulation, the so-called “nuclear vacuole”. The nutritive function of vitellocytes appears greatly reduced in W. virilis, a fact perhaps related to the intrauterine development of the early embryos. The ultrastructure of vitellogenesis in W. virilis is compared with that in other lower cestodes, both monozoic and polyzoic. Conclusions concerning interrelationships of the vitellogenesis pattern of the ultrastructural cytochemistry of mature vitellocytes of W. virilis to intrauterine embryonation, absence of uterine glands and an extensive uterus characteristic for this species, are drawn and discussed.
Development and morphology of the scolex and mode of attachment of Wenyonia virilis Woodland, 1923, a caryophyllaeid cestode from the silurid Nile fish Synodontis schall (Bloch et Schneider, 1801), were studied by means of light and scanning electron microscopy (SEM). Scolex and genital primordia changes through four stages of juvenile development are described. Longitudinal ridges do not appear on the scolex until the cestode has well defined genital primordia. This is in stark contrast to other caryophyllidean genera in which the basic morphology of the adult scolex becomes evident at the procercoid stage in the oligochaete intermediate host. The scolex of the adult has 13 to 19 prominent longitudinal ridges and deep furrows that come together at the apex to form an apical ring, a protrusible terminal introvert within the apical ring that forms a deep apical pouch when fully retracted, and a central group of Faserzellen. The scolex of W. virilis appears similar to the rugomonobothriate scolex of another African caryophyllid, Monobothrioides chalmersius (Woodland, 1924). Comparisons are made with other caryophyllideans having a scolex with a terminal structure: Monobothrium Diesing, 1863, Djombangia Bovien, 1926 and Caryoaustralus Mackiewicz et Blair, 1980. The terminal introvert may be responsible for attachment in early juvenile stages, but may be supplemented by the longitudinal ridges and furrows later in development. Host tissue appears to be drawn into these furrows that function as weak organs of attachment. We could not determine how the introvert of adult worms functions in attachment. At the site of attachment, the mucosa showed necrosis and degeneration and the submucosa exhibited vacuolization and infiltration with lymphocytes and leucocytes.
Ultrastructural and cytochemical characteristics of GER-bodies observed in the vitellocyte cytoplasm of the intrauterine eggs of the caryophyllidean cestode Wenyonia virilis are described. In this species GER-bodies were observed only in the cytoplasm of vitellocytes, surrounded by a newly formed egg-shell. They are composed of spherical areas of condensed, electron-dense cytoplasm which contains concentrically arranged parallel lamellae of granular endoplasmic reticulum (GER), forming characteristic balls of different sizes. Each GER-body is surrounded by numerous free ribosomes, polyribosomes, α-glycogen rosettes and large mitochondria. Results of cytochemical analysis by means of PATSC-SP test for polysaccharides indicated that glycogen is absent within the GER-bodies, however, a strongly positive reaction was observed only in large aggregations of α-glycogen rosettes and β-glycogen particles, localised usually near GER-bodies.
Ultrastructural evidence for early intraurerine embryonic development of Wenyonia virilis is presented. At the initial stage of egg formation, the fertilized oocyte or ovum is surrounded by numerous vitellocytes and newly formed eggshell. Individual vitellocytes undergo progressive fusion into a vitelline syncytium. During cleavage divisions, three types of blastomeres are formed: macromeres, mesomeres and micromeres. Two large macromeres contain a large nucleus with spherical nucleolus and numerous small heterochromatin islands dispersed in moderately electron-dense nucleoplasm. The granular cytoplasm shows a few large mitochondria. Medium-sized mesomeres contain a spherical nucleus with numerous heterochromatin islands, adjacent to the nuclear envelope, and a prominent electron-dense nucleolus. Their nuclei are embedded in granular cytoplasm with a few large and numerous small mitochondria and Golgi complexes. The small micromeres are characterized by presence of spherical nucleoli with large areas of highly condensed heterochromatin and a few islands of granular electron-lucent nucleoplasm. Their granular cytoplasm shows a few small lipid droplets and several spherical mitochondria. Majority of micromeres give rise to the hexacanth but many of them also undergo degeneration or apoptosis. Both mesomeres and macromeres are engaged in the formation of the oncospheral envelopes. The outer envelope is formed by a fusion of two macromeres whereas the inner envelope originates from a fusion of mesomeres. The intrauterine eggs of W. virilis usually contain an embryo at the early preoncopheral phase of development and possesses three primary envelopes: (1) thick eggshell; (2) thin cytoplasmic layer of the outer envelope and (3) inner envelope. Based on embryonic development, egg type and life-cycle characteristics, caryophyllideans tend to show closer affinities to spathebothriideans than to the former pseudophyllideans.
The ultrastructure of spermiogenesis in Wenyonia virilis Woodland, 1923, a caryophyllaeid cestode from the silurid Nile fish Synodontis schall (Bloch et Schneider, 1801), is described by means of transmission electron microscopy (TEM) for the first time. Spermiogenesis follows the characteristic caryophyllidean type and is initiated by the formation of a differentiation zone. This area, delimited at its base by a ring of arching membranes and bordered by cortical microtubules, contains two centrioles associated with typical striated rootlets with a reduced intercentriolar body between them. The apical area of the differentiation zone exhibits electron-dense material that is present only during the early stages of spermiogenesis. Only one of the centrioles develops into a free flagellum that grows at an angle of >90° in relation to the cytoplasmic extension. Spermiogenesis is also characterized by a flagellar rotation and a proximodistal fusion of the flagellum with the cytoplasmic extension. The most interesting features observed in W virilis are the presence of a reduced, very narrow intercentriolar body and the unique type of flagellar rotation >90°. Results are compared with those described in two caryophyllideans, Glaridacris catostomi Cooper, 1920 and Khawia armeniaca (Cholodkovski, 1915). Contrary to the original report of Świderski and Mackiewicz (2002), that flagellar rotation has never been observed in spermiogenesis of G. catostomi, re-assessment of their description and illustrations leads us to conclude that flagellar rotation must logically occur in that species. The value of various morphological features of sperm in phylogenetic inference is discussed.
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