A redescription of the rictulariid nematode Pterygodermatites (Pterygodermatites) plagiostoma Wed... more A redescription of the rictulariid nematode Pterygodermatites (Pterygodermatites) plagiostoma Wedl, 1861, is made by means of light and scanning electron microscopy, including the first data on male specimens. The morphologic study was based on specimens recovered from two Algerian hedgehogs (Atelerix algirus) from Tenerife and Gran Canaria islands (Canary Archipelago, Spain). The main characteristics of P. (P.) plagiostoma males are the presence of 49–53 pairs of cuticular projections, the presence of one or two midventral precloacal fans (generally one), and the size of two unequal spicules, measuring 98–123 µm (right spicule) and 185–236 µm (left spicule). The cloacal papillae are arranged according to the Ascaridida type. They include two precloacal pairs, an unpaired precloacal papilla, one pair lateral to the cloaca, six postcloacal pairs, and a pair of phasmids near the tail tip. Females are mainly characterized by the presence of 71–77 pairs of cuticular projections, with 43...
The present work constitutes the first ultrastructural analysis of the spermatozoon in the Pleuro... more The present work constitutes the first ultrastructural analysis of the spermatozoon in the Pleurogenidae, with the study of three species belonging to three of the 16 genera included in this family, namely Pleurogenes claviger, Pleurogenoides medians and Prosotocus confusus. The mature spermatozoa of these pleurogenids present two axonemes of the 9+“1” trepaxonematan pattern, a nucleus, two mitochondria, two bundles of parallel cortical microtubules, external ornamentation, spine-like bodies and granules of glycogen. The organization of these characters in the sperm cell is similar in the three species. Thus, the anterior spermatozoon extremity is filiform and a continuous and submembranous layer of parallel cortical microtubules surrounds the axonemes at their anterior end. The posterior spermatozoon extremity exhibits the second axoneme and corresponds to the Cryptogonimidean type of Quilichini et al. (2010). Slight differences were noted between the spermatozoon of P. confusus an...
A new microsporidian Toguebayea baccigeri n. gen., n. sp., hyperparasite of Bacciger israelensis ... more A new microsporidian Toguebayea baccigeri n. gen., n. sp., hyperparasite of Bacciger israelensis (Digenea, Faustulidae), parasite of Boops boops (Teleostei, Sparidae) is described by means of transmission electron microscopy. The phylogenetic analysis, based on the SSU rDNA gene, places the new species in the clade containing mainly crustacean-infecting microsporidia of the genus Cucumispora, within superclade V (Marinosporidia) sensu Vossbrinck et al., 2014. Mature spores of T. baccigeri are ovoid, uninucleated and measure 2.5 × 1.4 μm. The number of coils of the polar tube is 8–10. The polaroplast is composed of an external lamellar part and an internal vesicular or granular part. The main differences that distinguish the new genus and new species from the closely related microsporidia include hyperparasitism in a digenean host infecting a marine fish, the geographic distribution (coast of Tunisia), presence of one nucleus at all developmental stages, disporoblastic sporogony, and...
The present study describes the ultrastructural organization of the spermatozoa of the brachylaim... more The present study describes the ultrastructural organization of the spermatozoa of the brachylaimid digenean Ityogonimus ocreatus (Ityogoniminae) by means of transmission electron microscopy (TEM). Live digeneans were collected from the digestive tract of an Iberian mole Talpa occidentalis (Eulipotyphla, Talpidae) captured accidentally during a vole pest control campaign in Priesca (Asturias, Spain). The TEM study reveals that the I. ocreatus sperm are filiform, tapered at both extremities, and have two 9+'1' trepaxonematan axonemes, external ornamentation of the plasma membrane associated with cortical microtubules, spine-like bodies, two bundles of parallel cortical microtubules and one mitochondrion overlapping the anterior part of the nucleus. The external ornamentation of the plasma membrane is located in the posterior part of the anterior region. The maximum number of parallel cortical microtubules (45) is located in the anterior part of the sperm cells. Our results are compared with the available data on the family Brachylaimidae, especially on the other Ityogoniminae studied to date (Scaphiostomum palaearcticum).
The functional ultrastructure of eggs and cellular organization of hexacanths from gravid proglot... more The functional ultrastructure of eggs and cellular organization of hexacanths from gravid proglottids of Thysanotaenia congolensis, from black rats from Cape Verde, were examined by transmission electron microscopy. Mature eggs with fully formed hexacanths are grouped within parenchymatous capsules of gravid proglottids. Oncospheral envelopes surrounding mature hexacanths are reduced to a very thin membranous embryophore as their protective function is taken over by the parenchymatous capsules originating from the medullary parenchyma of immature proglottids and composed of three layers. Six major cell types are present: a bi-nucleate medullary centre; a six-nucleate U-shaped penetration gland; a second type of penetration gland; two neurosecretory-type nerve cells; about 30 somatic cells; and about 12 germinative cells. Present results on the functional ultrastructure of eggs and cellular organization of hexacanths support the phylogenetic distinction between T. congolensis and ces...
The ultrastructural characters of the mature spermatozoon of Taenia taeniaeformis are described b... more The ultrastructural characters of the mature spermatozoon of Taenia taeniaeformis are described by means of transmission electron microscopy. Materials were obtained from a naturally infected road-killed cat (Felis catus) from La Palma (Canary Islands, Spain). The mature spermatozoon of T. taeniaeformis is a filiform cell, which is tapered at both extremities and lacks mitochondria. It is characterised by the presence of (1) a single spirallised crested body about 140 nm thick, (2) a single axoneme of the 9+'1' pattern of trepaxonematan Platyhelminthes, (3) a twisted (40 degrees ) layer of submembranous cortical microtubules, (4) a periaxonemal sheath surrounding the axoneme, (5) transverse intracytoplasmic walls and (6) a spirallised nucleus encircling the axoneme. The mature spermatozoon of T. taeniaeformis is also characterised by the presence of an apical cone in its anterior extremity and by the disorganisation of the axoneme in its posterior extremity. The ultrastructural characters of the mature spermatozoon of T. taeniaeformis are compared with those of other cestodes studied to date, with particular emphasis on other representatives of the family Taeniidae.
The ultrastructure of the infective oncosphere of the cestode Mosgovoyia ctenoides (Anoplocephali... more The ultrastructure of the infective oncosphere of the cestode Mosgovoyia ctenoides (Anoplocephalidae) is described. The surface of the infective oncosphere is covered by a thin cytoplasmic layer of tegument connected by a narrow cytoplasmic process with the binucleate subtegumental cell, situated deeper in the body. Below the basal matrix of the cytoplasmic layer of the tegument are situated wide bands of the peripheral, somatic musculature responsible for body movements. The 3 pairs of hooks and their muscles form a complex hook muscle system, responsible for coordinated hook action. Five major types of cells have been distinguished: (1) a binucleate subtegumental cell, (2) a binucleate penetration gland, (3) 2 nerve cells, (4) numerous somatic cells, and (5) about 6 germinative cells. The approximate number of cells is 24 (26 nuclei, including 2 syncytial structures). The results of this study, when compared with other published reports from other cestode taxa, support previous hypotheses that the progressive reduction of oncosphere cells is an adaptive feature in cestode evolution.
This study presents the first comprehensive helminthological data on three sympatric riparian mus... more This study presents the first comprehensive helminthological data on three sympatric riparian mustelids (the European mink Mustela lutreola, the polecat M. putorius and the American mink M. vison) in south-western France. One hundred and twenty-four specimens (45 M. lutreola, 37 M. putorius and 42 M. vison) from eight French departments were analysed. Globally, 15 helminth species were detected: Troglotrema acutum, Pseudamphistomum truncatum, Euryhelmis squamula, Euparyphium melis and Ascocotyle sp. (Trematoda), Taenia tenuicollis (Cestoda), Eucoleus aerophilus, Pearsonema plica, Aonchotheca putorii, Strongyloides mustelorum, Molineus patens, Crenosoma melesi, Filaroides martis and Skrjabingylus nasicola (Nematoda) and larval stages of Centrorhynchus species (Acanthocephala). The autochthonous European mink harboured the highest species richness (13 species) followed by the polecat with 11 species. The introduced American mink presented the most depauperate helminth community (nine ...
Post-embryonic development and fully-formed polycephalic larvae of Taenia parva Baer, 1926 were e... more Post-embryonic development and fully-formed polycephalic larvae of Taenia parva Baer, 1926 were examined by light (LM) and transmission electron microscopy (TEM). Three developmental stages were recognised: (1) an early stage of exogenous budding at the surface of the central vesicle; (2) a stage of polycephalic cyst development accompanied by segmentation of the growing larval strobile and an obvious decrease in the size of the central vesicle; (3) fully-formed larval strobile and invaginated scoleces. In fully-developed encysted polycephalic larvae, there are usually 14–24 segmented larval strobilae, each terminating with an invaginated scolex; larval strobilae arise from a common central vesicle and remain attached posterior to it during the entire development. The number of segments varies between 109 and 120 per larval strobila. The polycephalic larvae examined closely resemble the strobilocercus type of taeniid larvae. The structure of developing and fully-formed larvae was ex...
We describe the spermiogenesis process and the ultrastructural characters of the spermatozoon of ... more We describe the spermiogenesis process and the ultrastructural characters of the spermatozoon of Acanthobothrium crassicolle by means of transmission electron microscopy, including cytochemical analysis for glycogen. Spermiogenesis in A. crassicolle begins with the formation of the differentiation zone that contains two centrioles associated with striated rootlets and an intercentriolar body. The latter is formed by one electron-dense layer. The centrioles develop into two free flagella that first grow orthogonally to a median cytoplasmic process and then undergo flagellar rotation becoming parallel to that median cytoplasmic process. After flagellar rotation only one of the flagella completes its growth and both short and long flagella undergo proximodistal fusion with the median cytoplasmic process. In the final stages of spermiogenesis, the nucleus becomes filiform and migrates into the spermatid body. Later, the ring of arched membranes constricts and the spermatozoon is liberat...
The present study describes the ultrastructure of mature vitellocytes of the trypanorhynch cestod... more The present study describes the ultrastructure of mature vitellocytes of the trypanorhynch cestode Progrillotia pastinacae Dollfus, 1946 (Progrillotiidae), a parasite of the common stingray Dasyatis pastinaca (Linnaeus, 1758) (Dasyatidae). The vitelline cells of this species measure about 24 μm in length and about 20 μm in width. They have small, elongated, slightly lobulated nuclei, about 4–5 μm in length, with large dense elongated nucleoli and numerous irregularly-shaped dense clumps of heterochromatin. The extensive cytoplasm is rich in numerous cell organelles and cell inclusions. While the perinuclear cytoplasm contains numerous long parallel cisternae of GER, ribo-and polyribosomes, several Golgi complexes and mitochondria, the peripheral cytoplasm contains predominantly three types of cell inclusions: a great number of large lipid droplets, several shell globule clusters, and a very small amount of glycogen-like particles. The most characteristic features of vitellocytes in ...
The first description of vitellogenesis in the Trypanorhyncha is presented in this paper. Though ... more The first description of vitellogenesis in the Trypanorhyncha is presented in this paper. Though the type of vitellogenesis and mature vitellocyte in Dollfusiella spinulifera appear to be unique among the Eucestoda, to some extent they resemble that observed in the lower cestodes, namely the Tetraphyllidea and Pseudophyllidea. Maturation is characterized by: (1) an increase in cell volume; (2) extensive development of large, parallel, frequently concentric cisternae of GER that produce proteinaceous granules; (3) development of Golgi complexes engaged in packaging this material; (4) continuous enlargement of proteinaceous granules within vesicles and their transformation into shell globule clusters; and (5) progressive fusion of all vesicles, with flocculent material containing the proteinaceous granules and shell globule clusters, into a single very large vesicle that characterises mature vitellocytes of this tapeworm. Cell inclusions in and around the large vesicle consist of floc...
The intrauterine, mature and fully embryonated eggs of the prosthogonimid trematode Mediogonimus ... more The intrauterine, mature and fully embryonated eggs of the prosthogonimid trematode Mediogonimus jourdanei Mas-Coma et Rocamora, 1978 were examined by means of transmission electron microscopy (TEM), using high pressure freezing, freeze substitution and infiltration with resin techniques. Each embryonated egg is composed of a miracidium surrounded by three envelopes: (1) the egg shell, (2) the outer and (3) inner envelopes. Egg envelopes play an important role in the protection, metabolism, storage of nutritive reserves and the general biology of the M. jourdanei egg. The inner envelope is characterised by large, flattened nuclei, and its syncytial cytoplasm contains a heavy accumulation of glycogen, lipid droplets, mitochondria and large vesicles. These traits indicate that this layer has the features of a metabolically-active syncytial layer with an energy storage capability. In the infective eggs observed before the hatching of the miracidium, areas of so-called “focal cytoplasmi...
A redescription of the rictulariid nematode Pterygodermatites (Pterygodermatites) plagiostoma Wed... more A redescription of the rictulariid nematode Pterygodermatites (Pterygodermatites) plagiostoma Wedl, 1861, is made by means of light and scanning electron microscopy, including the first data on male specimens. The morphologic study was based on specimens recovered from two Algerian hedgehogs (Atelerix algirus) from Tenerife and Gran Canaria islands (Canary Archipelago, Spain). The main characteristics of P. (P.) plagiostoma males are the presence of 49–53 pairs of cuticular projections, the presence of one or two midventral precloacal fans (generally one), and the size of two unequal spicules, measuring 98–123 µm (right spicule) and 185–236 µm (left spicule). The cloacal papillae are arranged according to the Ascaridida type. They include two precloacal pairs, an unpaired precloacal papilla, one pair lateral to the cloaca, six postcloacal pairs, and a pair of phasmids near the tail tip. Females are mainly characterized by the presence of 71–77 pairs of cuticular projections, with 43...
The present work constitutes the first ultrastructural analysis of the spermatozoon in the Pleuro... more The present work constitutes the first ultrastructural analysis of the spermatozoon in the Pleurogenidae, with the study of three species belonging to three of the 16 genera included in this family, namely Pleurogenes claviger, Pleurogenoides medians and Prosotocus confusus. The mature spermatozoa of these pleurogenids present two axonemes of the 9+“1” trepaxonematan pattern, a nucleus, two mitochondria, two bundles of parallel cortical microtubules, external ornamentation, spine-like bodies and granules of glycogen. The organization of these characters in the sperm cell is similar in the three species. Thus, the anterior spermatozoon extremity is filiform and a continuous and submembranous layer of parallel cortical microtubules surrounds the axonemes at their anterior end. The posterior spermatozoon extremity exhibits the second axoneme and corresponds to the Cryptogonimidean type of Quilichini et al. (2010). Slight differences were noted between the spermatozoon of P. confusus an...
A new microsporidian Toguebayea baccigeri n. gen., n. sp., hyperparasite of Bacciger israelensis ... more A new microsporidian Toguebayea baccigeri n. gen., n. sp., hyperparasite of Bacciger israelensis (Digenea, Faustulidae), parasite of Boops boops (Teleostei, Sparidae) is described by means of transmission electron microscopy. The phylogenetic analysis, based on the SSU rDNA gene, places the new species in the clade containing mainly crustacean-infecting microsporidia of the genus Cucumispora, within superclade V (Marinosporidia) sensu Vossbrinck et al., 2014. Mature spores of T. baccigeri are ovoid, uninucleated and measure 2.5 × 1.4 μm. The number of coils of the polar tube is 8–10. The polaroplast is composed of an external lamellar part and an internal vesicular or granular part. The main differences that distinguish the new genus and new species from the closely related microsporidia include hyperparasitism in a digenean host infecting a marine fish, the geographic distribution (coast of Tunisia), presence of one nucleus at all developmental stages, disporoblastic sporogony, and...
The present study describes the ultrastructural organization of the spermatozoa of the brachylaim... more The present study describes the ultrastructural organization of the spermatozoa of the brachylaimid digenean Ityogonimus ocreatus (Ityogoniminae) by means of transmission electron microscopy (TEM). Live digeneans were collected from the digestive tract of an Iberian mole Talpa occidentalis (Eulipotyphla, Talpidae) captured accidentally during a vole pest control campaign in Priesca (Asturias, Spain). The TEM study reveals that the I. ocreatus sperm are filiform, tapered at both extremities, and have two 9+'1' trepaxonematan axonemes, external ornamentation of the plasma membrane associated with cortical microtubules, spine-like bodies, two bundles of parallel cortical microtubules and one mitochondrion overlapping the anterior part of the nucleus. The external ornamentation of the plasma membrane is located in the posterior part of the anterior region. The maximum number of parallel cortical microtubules (45) is located in the anterior part of the sperm cells. Our results are compared with the available data on the family Brachylaimidae, especially on the other Ityogoniminae studied to date (Scaphiostomum palaearcticum).
The functional ultrastructure of eggs and cellular organization of hexacanths from gravid proglot... more The functional ultrastructure of eggs and cellular organization of hexacanths from gravid proglottids of Thysanotaenia congolensis, from black rats from Cape Verde, were examined by transmission electron microscopy. Mature eggs with fully formed hexacanths are grouped within parenchymatous capsules of gravid proglottids. Oncospheral envelopes surrounding mature hexacanths are reduced to a very thin membranous embryophore as their protective function is taken over by the parenchymatous capsules originating from the medullary parenchyma of immature proglottids and composed of three layers. Six major cell types are present: a bi-nucleate medullary centre; a six-nucleate U-shaped penetration gland; a second type of penetration gland; two neurosecretory-type nerve cells; about 30 somatic cells; and about 12 germinative cells. Present results on the functional ultrastructure of eggs and cellular organization of hexacanths support the phylogenetic distinction between T. congolensis and ces...
The ultrastructural characters of the mature spermatozoon of Taenia taeniaeformis are described b... more The ultrastructural characters of the mature spermatozoon of Taenia taeniaeformis are described by means of transmission electron microscopy. Materials were obtained from a naturally infected road-killed cat (Felis catus) from La Palma (Canary Islands, Spain). The mature spermatozoon of T. taeniaeformis is a filiform cell, which is tapered at both extremities and lacks mitochondria. It is characterised by the presence of (1) a single spirallised crested body about 140 nm thick, (2) a single axoneme of the 9+'1' pattern of trepaxonematan Platyhelminthes, (3) a twisted (40 degrees ) layer of submembranous cortical microtubules, (4) a periaxonemal sheath surrounding the axoneme, (5) transverse intracytoplasmic walls and (6) a spirallised nucleus encircling the axoneme. The mature spermatozoon of T. taeniaeformis is also characterised by the presence of an apical cone in its anterior extremity and by the disorganisation of the axoneme in its posterior extremity. The ultrastructural characters of the mature spermatozoon of T. taeniaeformis are compared with those of other cestodes studied to date, with particular emphasis on other representatives of the family Taeniidae.
The ultrastructure of the infective oncosphere of the cestode Mosgovoyia ctenoides (Anoplocephali... more The ultrastructure of the infective oncosphere of the cestode Mosgovoyia ctenoides (Anoplocephalidae) is described. The surface of the infective oncosphere is covered by a thin cytoplasmic layer of tegument connected by a narrow cytoplasmic process with the binucleate subtegumental cell, situated deeper in the body. Below the basal matrix of the cytoplasmic layer of the tegument are situated wide bands of the peripheral, somatic musculature responsible for body movements. The 3 pairs of hooks and their muscles form a complex hook muscle system, responsible for coordinated hook action. Five major types of cells have been distinguished: (1) a binucleate subtegumental cell, (2) a binucleate penetration gland, (3) 2 nerve cells, (4) numerous somatic cells, and (5) about 6 germinative cells. The approximate number of cells is 24 (26 nuclei, including 2 syncytial structures). The results of this study, when compared with other published reports from other cestode taxa, support previous hypotheses that the progressive reduction of oncosphere cells is an adaptive feature in cestode evolution.
This study presents the first comprehensive helminthological data on three sympatric riparian mus... more This study presents the first comprehensive helminthological data on three sympatric riparian mustelids (the European mink Mustela lutreola, the polecat M. putorius and the American mink M. vison) in south-western France. One hundred and twenty-four specimens (45 M. lutreola, 37 M. putorius and 42 M. vison) from eight French departments were analysed. Globally, 15 helminth species were detected: Troglotrema acutum, Pseudamphistomum truncatum, Euryhelmis squamula, Euparyphium melis and Ascocotyle sp. (Trematoda), Taenia tenuicollis (Cestoda), Eucoleus aerophilus, Pearsonema plica, Aonchotheca putorii, Strongyloides mustelorum, Molineus patens, Crenosoma melesi, Filaroides martis and Skrjabingylus nasicola (Nematoda) and larval stages of Centrorhynchus species (Acanthocephala). The autochthonous European mink harboured the highest species richness (13 species) followed by the polecat with 11 species. The introduced American mink presented the most depauperate helminth community (nine ...
Post-embryonic development and fully-formed polycephalic larvae of Taenia parva Baer, 1926 were e... more Post-embryonic development and fully-formed polycephalic larvae of Taenia parva Baer, 1926 were examined by light (LM) and transmission electron microscopy (TEM). Three developmental stages were recognised: (1) an early stage of exogenous budding at the surface of the central vesicle; (2) a stage of polycephalic cyst development accompanied by segmentation of the growing larval strobile and an obvious decrease in the size of the central vesicle; (3) fully-formed larval strobile and invaginated scoleces. In fully-developed encysted polycephalic larvae, there are usually 14–24 segmented larval strobilae, each terminating with an invaginated scolex; larval strobilae arise from a common central vesicle and remain attached posterior to it during the entire development. The number of segments varies between 109 and 120 per larval strobila. The polycephalic larvae examined closely resemble the strobilocercus type of taeniid larvae. The structure of developing and fully-formed larvae was ex...
We describe the spermiogenesis process and the ultrastructural characters of the spermatozoon of ... more We describe the spermiogenesis process and the ultrastructural characters of the spermatozoon of Acanthobothrium crassicolle by means of transmission electron microscopy, including cytochemical analysis for glycogen. Spermiogenesis in A. crassicolle begins with the formation of the differentiation zone that contains two centrioles associated with striated rootlets and an intercentriolar body. The latter is formed by one electron-dense layer. The centrioles develop into two free flagella that first grow orthogonally to a median cytoplasmic process and then undergo flagellar rotation becoming parallel to that median cytoplasmic process. After flagellar rotation only one of the flagella completes its growth and both short and long flagella undergo proximodistal fusion with the median cytoplasmic process. In the final stages of spermiogenesis, the nucleus becomes filiform and migrates into the spermatid body. Later, the ring of arched membranes constricts and the spermatozoon is liberat...
The present study describes the ultrastructure of mature vitellocytes of the trypanorhynch cestod... more The present study describes the ultrastructure of mature vitellocytes of the trypanorhynch cestode Progrillotia pastinacae Dollfus, 1946 (Progrillotiidae), a parasite of the common stingray Dasyatis pastinaca (Linnaeus, 1758) (Dasyatidae). The vitelline cells of this species measure about 24 μm in length and about 20 μm in width. They have small, elongated, slightly lobulated nuclei, about 4–5 μm in length, with large dense elongated nucleoli and numerous irregularly-shaped dense clumps of heterochromatin. The extensive cytoplasm is rich in numerous cell organelles and cell inclusions. While the perinuclear cytoplasm contains numerous long parallel cisternae of GER, ribo-and polyribosomes, several Golgi complexes and mitochondria, the peripheral cytoplasm contains predominantly three types of cell inclusions: a great number of large lipid droplets, several shell globule clusters, and a very small amount of glycogen-like particles. The most characteristic features of vitellocytes in ...
The first description of vitellogenesis in the Trypanorhyncha is presented in this paper. Though ... more The first description of vitellogenesis in the Trypanorhyncha is presented in this paper. Though the type of vitellogenesis and mature vitellocyte in Dollfusiella spinulifera appear to be unique among the Eucestoda, to some extent they resemble that observed in the lower cestodes, namely the Tetraphyllidea and Pseudophyllidea. Maturation is characterized by: (1) an increase in cell volume; (2) extensive development of large, parallel, frequently concentric cisternae of GER that produce proteinaceous granules; (3) development of Golgi complexes engaged in packaging this material; (4) continuous enlargement of proteinaceous granules within vesicles and their transformation into shell globule clusters; and (5) progressive fusion of all vesicles, with flocculent material containing the proteinaceous granules and shell globule clusters, into a single very large vesicle that characterises mature vitellocytes of this tapeworm. Cell inclusions in and around the large vesicle consist of floc...
The intrauterine, mature and fully embryonated eggs of the prosthogonimid trematode Mediogonimus ... more The intrauterine, mature and fully embryonated eggs of the prosthogonimid trematode Mediogonimus jourdanei Mas-Coma et Rocamora, 1978 were examined by means of transmission electron microscopy (TEM), using high pressure freezing, freeze substitution and infiltration with resin techniques. Each embryonated egg is composed of a miracidium surrounded by three envelopes: (1) the egg shell, (2) the outer and (3) inner envelopes. Egg envelopes play an important role in the protection, metabolism, storage of nutritive reserves and the general biology of the M. jourdanei egg. The inner envelope is characterised by large, flattened nuclei, and its syncytial cytoplasm contains a heavy accumulation of glycogen, lipid droplets, mitochondria and large vesicles. These traits indicate that this layer has the features of a metabolically-active syncytial layer with an energy storage capability. In the infective eggs observed before the hatching of the miracidium, areas of so-called “focal cytoplasmi...
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