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Recent literature on the evolution and interrelationships of the Caryophyllidea based on molecular and morphological criteria is reviewed. Molecular analyses with SSU rDNA, LSU rDNA and ef-1 alpha reaffirms the basal or near basal position of these nonozoic cestodes. Major emphasis is on an evaluation of the scoring in morphological character matrices used in cladistic studies. Suggested changes to present scoring are: uterus is dorsal; scolex is afossate, fossate or difossate with little support for monofossate; ciliated coracidium is absent; vitellaria are circum-cortical and circum-medullary; testes are cortical and medullary; metacercoid stage is absent; and the spermatozoan lacks a crested body, flagellar rotation and proximodistal fusion. Of the 41 recognized genera of the Caryophyllidea, 59% have an afossate scolex and the remainders are fossate. The use of a new character, “nuclear vacuole” in the nucleus of mature vitellocytes, is suggested. To aid in identifying cestode body types in an evolutionary context, they are designated as monopleuroid, polypleuroid and strobila. Tabulated differences between the monozoic Caryophyllidea and polyzoic eucestodes suggest that the two groups may warrant separate taxonomic status. The question of whether or not the monozoic state is primary or secondarily derived is not resolved. Using the life cycle characterstics of the Pseudophyllidea and of Archigetes as models, it is hypothesized that progenesis may have played a major role in the evolution of the Caryophyllidea. If the role of progenesis can be substantiated through total evidence incorporating cytohistological data, then the monozoic condition becomes coincidental and the hypothesis is not supported that the Caryophyllidea are ancestral and preceded polyzoic eucestodes.
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.
Spermatogenesis of Glaridacris catostomi from Catostomus commersoni (Catostomidae) from Albany, New York (USA) was studied by means of TEM, SEM and squashes. Mature testes of G. catostomi contain all consecutive stages of spermatogenesis; primary spermatogonia are usually situated at the periphery and mature spermatozoa in the centre of testes. The primary spermatogonium divides mitotically, but the two daughter cells, secondary spermatogonia, remain connected with each other by a cytoplasmic bridge. Spermatogenesis in G. catostomi is of a rosette type. Six incomplete, synchronic cytokineses, five mitotic and meiotic divisions occur simultaneously, resulting in a cluster of four tertiary spermatogonia, then eight quaternary spermatogonia, and subsequently sixteen primary spermatocytes are formed. These enlarge, their nuclei move to the periphery and the cluster of cells takes on the form of rosette. After the first meiotic division, a rosette of thirty-two secondary spermatocytes is formed. The haploid nuclei of these are smaller and the cell membranes near the centre of the rosette become indistinct as the displacement of nuclei toward the periphery continues. The second maturation division results in sixty-four spermatids. During spermiogenesis, their nuclei subsequently elongate, migrate, and are transformed into electron-dense, filiform nuclei of spermatozoa. Each spermatid forms at the surface a so-called "zone of differentiation". From this conical zone arise initially three elongating processes: cytoplasmic extension and two lateral flagella. The spermiogenesis type observed in G. catostomi is characterised by an early abortion of the second axoneme. The remaining single axoneme, forming the sperm flagellum, elongates in parallel with cytoplasmic process accommodating the sperm nucleus; the two parts are initially separated. The migration of the sperm nucleus induces their lateral fusion, which is, however, very superficial; the flagellar axoneme and sperm nucleus are never incorporated completely into a common sperm body as observed in pseudophyllideans or cyclophyllideans. The spermatozoon of G. catostomi consists of two elongate parts: nucleated sperm body with a row of cortical microtubules and flagellum connected by a narrow, longitudinal bridge throughout nearly the entire length. The flagellum consists of a single axoneme of the 9 + '1' Platyhelminthes type. The value of spermiogenesis type and sperm ultrastructure as taxonomie tools in platyhelminth phylogeny is 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.
Vitellogenesis in Khaxvia armeniaca was examined by means of transmission electron microscopy (TEM) and cytochemical staining with periodic acid-thiosemicarbazide-silver proteinate (PA-TSC-SP) for specific detection of glycogen at the ultrastructural level. Mature vitelline follicles consist of cells in various stages of development, progressing from immature cells of gonial type near the periphery to mature vitellocytes towards the centre. Maturation of vitelline cells is characterized by: (1) increase in cell volume; (2) increase in nuclear surface area restoring the N/C ratio; (3) nucleolar transformation; (4) extensive development of large parallel cisternae of GER, the shell-protein producing units; (5) development of Golgi complexes engaged in shell-granule/shell-globule vitelline material formation and package; (6) formation 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 that fuse into large shell-globule clusters with a progressive increase in the number and size of the latter; and (9) degeneration of GER in the medial layer of vitellocyte cytoplasm with degenerative changes and accumulation of glycogen and shell-globule clusters within the cytoplasm, associated with a massive accumulation of glycogen in the nucleus. The functional significance of the large amount of nuclear and cytoplasmic glycogen and numerous shell-globule clusters is analysed. The ultrastructural aspect of vitellogenesis is compared with that in other monozoic and polyzoic cestodes. Conclusions concerning the interrelationships of vitellogenesis patterns and ultrastructural cytochemistry of mature vitellocytes to the various types of embryogenesis, are drawn and discussed.
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.
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.
In Lytocestus indicus putative neurosecretory cells (pNSC) are recognized on the basis of phloxinophilic and fuchsinophilic nature of their cytoplasm; pNSC in L. indicus are dimensionally small. Morphologically, there are four types of pNSC: a-, uni-, bi-, and multipolar cells. They are found both in the cortical and medullary parenchyma of the neck and strobilar regions; in the latter the pNSC occur in close vicinity with the longitudinal muscles and components of the reproductive system.
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