Ultrastructure of the protonephridial system of regenerating Stenostomum sp. (Plathelminthes,Catenulida)

Summary The ultrastructure of the flame bulbs, protonephridial capillaries and duct of fully developed and regenerating Stenostomum sp. is described. Flame bulbs are formed by a single cell whose nucleus is located basally or laterally to the weir. The weir is formed by a single row of transverse ribs connected by a thin membrane, apparently of extracellular matrix. Internal leptotriches arise from the proximal cytoplasm and extend in a (usually) single row along the weir and into the lumen of the distal cytoplasmic tube. Many or all leptotriches do not fuse with the distal cytoplasm. Two cilia are each anchored in the proximal cytoplasm by a cross-striated vertical and lateral rootlet, the latter bent forward and extending for some distance into one of the two cytoplasmic cords along the weir. Each cord contains the lateral rootlet in its proximal part, as well as many microtubules. The distal cytoplasmic tube contains two (longitudinal) junctions, i.e. lines of contact between cell processes of the same, terminal cell. Occasionally, more than two junctions were seen, apparently due to branches of the terminal cell in contact with each other. Flame bulbs join capillaries lined by several canal cells type I, containing few or no microvilli but lateral flames. Such capillaries join a duct (or ducts?) lined by canal cells type II with many long microvilli. The large protonephridial duct is lined by numerous cells with lateral flames and many long microvilli. In regenerating tissue (10.5 hours after cutting) some flame bulbs were free, i.e. not connected to capillaries, and some capillaries openly communicated with the surrounding intercellular space. In the presence of a single row of ribs in the weir, of internal leptotriches, and of vertical and lateral ciliary rootlets, the flame bulb of Stenostomum sp. resembles that of other Plathelminthes much more closely than hitherto thought. The species differs from non-catenulid plathelminths mainly in the large number of glandular cells lining the large protonephridial ducts, in the transverse orientation of the ribs in the weir and in the presence of only two cilia in the flame.     

Investigations on the protonephridial system of post-larval Gyrocotyle urna and Amphilina foliacea (Cestoda).

The gross morphology and ultrastructure of the different parts of the protonephridial system of the monozoic tapeworms Gyrocotyle urna and Amphilina foliacea are described. The terminal cell in both species has numerous cilia which are interconnected and extend into the lumen of the first canal cell. The filtration area is built up from projections of two cells, the terminal cell and the first canal cell. The first canal cell forms a solid hollow cylinder without a cell gap and a desmosome as found in Neodermata other than cestodes and Udonella. In Gyroctyle the nucleus of the first canal cell is located in the wall cytoplasma whereas more distally located ductules of both species have subepithelial cell bodies containing the nuclei. In both taxa the protonephridial canal system is reticulate. In Amphilina the distal canals lack non-terminal ciliary flames, such ciliary tufts can be found in the larger capillaries of Gyrocotyle. The capillary cilia have rootlets and the ultrastructure of the duct wall cytoplasm containing large numbers of vesicles indicates highly active transport processes. The morphology of the protonephridial systems is discussed with regard to the evolution of Neodermata (especially of the Cestoda) and the function of the protonephridial system in cestodes as a probable organ of nutrient distribution.     

Ultrastructure of the Protonephridium in Acteonemertes bathamae Pantin (Rhynchocoela: Enopla: Hoplonemertini)

The protonephridial system consists of terminal cell, protonephridial capillary, protonephridial tubule and efferent duct. The terminal cell is an elongated, thin-walled, fenestrated basket containing a ciliary flame circumscribed by a palisade of straight microvilli. The filtration area is confined to the terminal cell and consists of slits bridged by a filtration membrane. The cilia, as well as the microvilli, projects into the proximal bell-shaped part of the thin-walled protonephridial capillary. The terminal cells are often found in pairs connected to the same capillary, which has a very narrow lumen. The proximal part of the thick-walled, convoluted protonephridial tubule is ciliated and shows characteristic foldings of the luminal plasma membrane and numerous small vesicles in the cytoplasm. The cells of the following, non-ciliated part of the tubule have interdigitating lateral surfaces and the bases deeply invaginated to form compartments with numerous mitochondria; in the cytoplasm are many large vesicles, possibly containing lipid droplets, and small amounts of glycogen. The distal protonephridial tubule resembles various epithelia with an osmoregulatory function, including the vertebrate nephron.     

扩张莫尼茨绦虫(绦虫纲:圆叶目)的原肾管及原肾管概念的评述

本研究应用透射电子显微镜研究了扩张莫尼茨绦虫原肾管的细胞学特征 ,莫尼茨绦虫原肾管的焰茎球为一个过滤器结构 ,类似于“挡河坝”样构造 ,此构造由端细胞和近管细胞外突形成的肋条 (或称杆 )相互交错排列而成。肋条之间由细胞外物质构成的“膜”结构连接 ,过滤作用通过该“膜”发生。焰细胞与近管细胞交界处有裂缝或孔与细胞外的结缔组织 (实质组织 )相通 ;原肾管的毛细排泄管细胞质索之间没有隔状联结 ;毛细排泄管及排泄管的管腔内有大量珠状微绒毛突起以增加表面积。从扩张莫尼茨绦虫及其它一些无脊椎动物原肾管的研究结果表明 ,原原肾管概念将焰细胞作为封闭的盲端已不再合适 ,需要进行修订 ,建议修订为 :原肾管是一种焰细胞系统 ,通常由焰细胞、管细胞和肾孔细胞组成 ,焰茎球作为过滤装置与周围的结缔组织 (实质组织 )有或没有裂缝 (孔 )相通     

Ultrastructure of excretory system in <Emphasis Type="Italic">Bothrioplana semperi</Emphasis> (Platyhelminthes,Turbellaria)

The ultrastructure of flame bulbs and epithelium of excretory canals in Bothrioplana semperi (Turbellaria, Seriata) have been studied. The flame bulbs consist of two cells, the terminal cell and the proximal canal cell. The weir is formed by two rows of longitudinal ribs. The ribs of the internal row originate from the flame cell, external ribs are formed by the proximal canal cell. Each external rib has a remarkable bundle of microfilaments, originating in the cytoplasm of the first canal cell distally to the bases of external ribs. Membrane of internal ribs is marked by small electrondense granules, separate or fused to an electron-dense layer, continuous to dense “membrane,” connecting both external and internal ribs. Sparse internal leptotrichs originate from the bottom of the flame bulb cavity. External leptotrichs are lacking. Septate junction is present only in proximal canal cell at the level of tips of cilia. The apical surface of the canal cell bears rare short microvilli. The basal membrane of canal cells forms long invaginations that may reach nearly the apical membrane. The epithelium of excretory canals lacks the cilia. The ultrastructure of flame bulbs and epithelium of the excretory canals in B. semperi shares representatives of suborder Proseriata (Seriata). The contradiction exists in interpretation of the structure of flame bulbs in Proseriata. Ehlers and Sopott-Ehlers assumed that the external ribs are derivatives of the proximal canal cell and internal ones are outgrowths of the terminal cell, while Rohde has found conversely: the external ribs are outgrowths of the terminal cell, the internal ones are outgrowths of the proximal canal cell. However, the illustrations provided by Rohde do not enable to ascertain what cells the internal and external ribs derive from, while illustrations provided by Ehlers justify his interpretation. The order of weir formation in B. semperi confirms the viewpoint of Ehlers. The implication of ultrastructure of flame bulbs in Proseriata, especially of the order of flame bulb formation, in the Platyhelminthes phylogeny has been discussed.     

Ultrastructure of the lycophora larva of Gyrocotyle urna (Cestoda,Gyrocotylidea)

Summary The ultrastructure of the protonephridial system of the lycophore larva of Gyrocotyle urna Grube and Wagener, 1852, is described. It consists of six terminal cells, at least two proximal canal cells, two distal canal cells and two nephridiopore cells. The terminal cells and the proximal canal cell build up the filtration weir with its two circles of weir rods. The proximal canal cell constitutes a solid, hollow cylinder without a cell gap and desmosome. The distal canal cell is characterized by a strong reduction of the canal lumen by irregularly shaped microvilli. The nephridiopore region is formed by a nephridiopore cell; its cell body is located at some distance proximally within the larva. The connection among different canal cells is brought about by septate desmosomes. Morphological, evolutionary and functional aspects of the protonephridial system within Platyhelminthes are discussed. The structure of the proximal canal cells without a desmosome is considered an autapomorphy of Cestoda.Abbreviations ci cilia of the terminal cell - Co distal canal cell - col lumen of the distal canal cell - Ep epidermis - er outer rods of the filtration weir - il inner leptotriches - ir inner rods of the filtration weir - ld lipid droplets - mt microtubule - mv microvilli - Nc nephridiopore cell - Ne neodermis anlage cells - nu nucleus - pC proximal canal cell - ro ciliary rootlets - sd septate desmosome - Tc terminal cell     

Ultrastructure of flame cells and protonephridial capillaries of Craspedella and Didymorchis (Plathelminthes,Rhabdocoela)

Summary The ultrastructure of the flame cells and protonephridial capillaries of the Rhabdocoela Craspedella sp. and Didymorchis sp., ectocommensals on the freshwater crayfish Cherax destructor in eastern Australia is described. The flame cells of both species have variable numbers of cilia without distinct rootlets and with decreasing numbers of axonemal tubules towards the ciliary tips. Bundles of microtubules extend from the cytoplasm adjacent to the ciliary rootlets through the ribs of the weir apparatus into the distal cytoplasmic tube, where the numbers of microtubules gradually decrease. The weir apparatus is formed by a single row of longitudinal ribs connected by a membrane. In Craspedella, but not in Didymorchis, the ribs have external branched leptotriches. Mitochondria are common in the wall of the flame cell of both species. The protonephridial capillary just above the end of the ciliary tuft narrows in both species and bends sharply in Craspedella. The lumen of the flame cell and the capillary is lined by a dark layer of cytoplasm; there is no enlargement of the surface area by microvilli or lamellae. Centrioles were seen in the capillary wall of Craspedella, and in Didymorchis the cytoplasm around the capillaries has a very loose and light appearance. The ultrastructure of the flame cells and capillaries of both species corresponds closely to that of Temnocephala sp.Abbreviations in the figures BB basal body - CE centriole - L leptotrich - M microtubules - ME membrane of weir apparatus - MI mitochondrion - PC protonephridial capillary - R rib (rod) of weir apparatus     

Ultrastructure of the protonephridium in Prostoma graecense (Bohmig) (Rhynchocoela, Enopla, Hoplonemertini)

Fine structure studies show that (1) the terminal cell is an elongated thin-walled and fenestrated basket with a multiciliary flame. Many short curved microvilli are confined to the cell lumen, while longer straight microvilli project from the cell's apical end into the proximal part of the protonephri-dial capillary, forming a sheath around the flame. The filtration area consists of slits between narrow cytoplasmic bars and is entirely confined to the terminal cell, which consequently is defined as a flame bulb, not closely similar to those of other phyla. (2) The protonephridial capillary is short and narrow, with few scattered cilia and luminal microvilli. (3) The coiled tubule is thick-walled, with several ciliated cells very rich in glycogen. The luminal border shows specializations probably concerned with modifying the ultrafiltrate.     

Ultrastructure of the protonephridia of larval Rugiloricus cf. cauliculus, male Armorloricus elegans, and female Nanaloricus mysticus (Loricifera)

The protonephridial system of several Loricifera was studied by transmission electron microscopy. A larval specimen of Rugiloricus cf. cauliculus possesses two protonephridia, which are "capped" frontally by a compact mass of still undifferentiated gonadal cells. Each protonephridium consists of four monociliary terminal cells and four canal cells with a diplosome but no cilia. Because of incomplete series of sections and unsatisfactory fixation, the outleading cell(s) could not be detected. In a male specimen of Armorloricus elegans, each gonad contains two protonephridia that open into the gonadal lumen. Each protonephridium consists of two monociliary terminal cells, each forming a filter, two nonciliated canal cells, and two nephroporus cells. The protonephridial lumina of the latter cells fuse to one common lumen, which unites with the gonadal lumen. Preliminary observations on the protonephridia of a female Nanaloricus mysticus reveal a more complicated arrangement of interdigitating terminal and canal cells. One or two terminal cells form their own individual filter or four cells form a common compound filter. The cilium of the terminal cells of all species investigated are surrounded by a palisade of nine microvilli that support the filter barrier made of an extracellular matrix. An additional filter diaphragm could be traced between the pores in the cell wall of each terminal cell of A. elegans. The urogenital system of the Loricifera differs from that of the Priapulida in that the protonephridia of the former are completely integrated into the gonad, whereas the excretory organs of the latter open into the urogenital duct caudally of the gonads.     

Ultrastructure of the nephridio-circulatory connections in Tubulanus annulatus (Nemertini,Anopla)

Summary The protonephridial terminal organs in the nemertean Tubulanus annulatus form an integral part of the blood vessel wall. Both endothelial and muscle-cell layers of the vessel's wall are discontinued at the site of each terminal organ. The terminal organs are usually composed of from one to three terminal cells enclosing a central lumen provided with many microvilli and separated from the blood vessel's lumen by a membranous filtration area. The latter is perforated by numerous winding clefts formed by interdigitation of minute cytoplasmic pedicels arising from processes issued by each of the involved terminal cells. Ultrafiltration of blood plasma takes place across a filtration membrane which spans the cleft system and the basal lamina of the terminal cells. Fluid is propelled into the lumen of the terminal organs through the activity of ciliary bundles, one for each terminal cell involved, perhaps supplemented by vascular turgor. All efferent conduits of the protonephridium have profuse infoldings of the luminal cell surfaces and/or numerous pinocytotic pits suggestive of reabsorption of substances from the primary urine.Abbreviations BL basal lamina - C cilium - CP coated pit - CT collecting tubule - CV inzcoated vesicle - D dictyosome - E endothelial cell - F fenestration of endothelial cell - FA filtration area - FM filtration membrane - G glycogen granule - LV lateral vessel - M mitochondrion - MC muscle cell - MV microvillus - N nucleus of terminal cell - NE nucleus of endothelial cell - NP nephridiopore - PC protonephridial capillary cell - PT protonephridial tubule - R rootlet - TC terminal cell     

Ultrastructure of protonephridia in Xenotrichula carolinensis syltensis and Chaetonotus maximus (Gastrotricha: Chaetonotida): comparative evaluation of the gastrotrich excretory organs

In an attempt to obtain detailed information on the entire protonephridial system in Gastrotricha, we have studied the protonephridial ultrastructure of two paucitubulatan species, Xenotrichula carolinensis syltensis and Chaetonotus maximus by means of complete sets of ultrathin sections. In spite of some differences in detail, the morphology of protonephridia in both examined species shows a common pattern: Both species have one pair of protonephridia that consist of a bicellular terminal organ, a voluminous, aciliar canal cell and an adjacent, aciliar nephridiopore cell. The terminal organ consists of two monociliar terminal cells each with a distal cytoplasmic lobe. These lobes interdigitate and surround cilia and microvilli of the terminal cells. Where both lobes interdigitate, a meandering cleft is formed that is covered by the filtration barrier. We here term the entire structure composite filter. The elongated, in some regions convoluted protonephridial lumen opens distally to the outside via a permanent nephridiopore. A comparison with the protonephridia of other species of the Gastrotricha allows hypothesising the following autapomorphies of the Paucitubulata: The bicellular terminal organ with a composite filter, the convoluted distal canal cell lumen and the absence of cilia, ciliary basal structures and microvilli within the canal cell. Moreover, this comparative survey could confirm important characteristics of the protonephridial system assumed for the ground pattern of Gastrotricha like, for example, the single terminal cell with one cilium surrounded by eight microvilli.     

The fine structure of the protonephridial system in the land nemertean Pantinonemertes californiensis (Rhynchocoela,Enopla, Hoplonemertini)

Summary The protonephridial terminal organ in the nemertean Pantinonemertes californiensis is composed of two cells that are similar in size and shape and are mirror images of each other. Basally in the organ the two cells combine to form a binucleate cytoplasmic mass. Apically they are intimately joined to form a subcylindrical thin-walled weir apparatus; this part is supported by two opposed cytoplasmic columns running the length of the weir region, one originating from each of the two cells, and by a number of regularly spaced circular bars that arise from the two columns. The ciliary flame consists of 94–114 cilia that originate in the bases of the two cells, and it is surrounded by a palisade of incomplete circlets of long, straight microvilli. The convoluted protonephridial tubule is rich in structures that indicate intensive reabsorption from the primary urine. It is argued that the terminal organs in Pantinonemertes and Geonemertes are fundamentally similar and differ only in the amount of microtubules present in the longitudinal supports.Abbreviations BL basal lamina - CF ciliary flame - CT connective tissue - CV coated vesicle - E endocytotic pit - FM filtration membrane - G Golgi complex - LC longitudinal cytoplasmic column - M mitochondrion - MT microtubules - MV microvilli - N nucleus - NPC nucleus of protonephridial capillary cell - PC protonephridial capillary cell - R rootlets - TB transverse bar - TC terminal cell - WE weir, exterior of fenestrated wall - WI weir, interior of same     

Ultrastructure of the protonephridial system of Dactylogyrus sp. and an unidentified ancyrocephaline (Monogenea: Dactylogyridae)

The ultrastructure of the flame bulbs and capillaries of the protonephridia of Dactylogyrus (probably anchoratus) from Carassius auratus in southeastern Australia, and of an unidentified ancyrocephaline from the marine teleost Priacanthus macracanthus in southern Queensland is described. The cilia of the flame are anchored in the terminal cell by means of basal bodies without distinct rootlets. The nucleus of the terminal cell is basal, and (in Dactylogyrus) partly lateral to the basal bodies. The weir consists of a row of internal and a row of external ribs (rods) connected by a ‘membrane’. The external ribs are continuations of the cytoplasm of a thick-walled ‘cytoplasmic cylinder’ (proximal canal cell) which tightly surrounds most of the flame and contains a septate junction; the internal ribs are continuous with the terminal cell. Internal leptotriches arise from the perikaryon of the terminal cell, and, in the ancyrocephaline, also from the internal ribs. The wall of the protonephridial capillaries contains a septate junction, a reticulum of interconnected spaces and, in the ancyrocephaline, also lamellae. Lateral flames are common in the capillaries of Dactylogyrus.     

Ultrastructure of the ampullae of Lorenzini of <Emphasis Type="Italic">Aptychotrema rostrata</Emphasis> (Rhinobatidae)

Small epidermal pores of the electrosensory ampullae of Lorenzini located both ventrally and dorsally on the disk of Aptychotrema rostrata (Shaw and Nodder, 1794) open to jelly-filled canals, the distal end of which widens forming an ampulla that contains 6 ± 0.7 alveolar bulbs (n = 13). The sensory epithelium is restricted to the alveolar bulbs and consists of receptor cells and supportive cells. The receptor cells are ellipsoid and their apical surfaces are exposed to the alveolar lumen with each bearing a single central kinocilium. Presynaptic bodies occur in the basal region of the receptor cell immediately proximal to the synaptic terminals. The supportive cells that surround receptor cells vary in shape. Microvilli originate from their apical surface and extend into the alveolar lumen. Tight junctions and desmosomes connect the supportive cells with adjacent supportive and receptor cells in the apical region. The canal wall consists of two cell layers, of which the luminal cells are squamous and interconnect via desmosomes and tight junctions, whereas the cells of the deeper layer are heavily interdigitated, presumably mechanically strengthening the canal wall. Columnar epithelial cells form folds that separate adjacent alveoli. The same cells separate the ampulla and canal wall. An afferent sensory nerve composed of up to nine myelinated nerve axons is surrounded by several layers of collagen fibers and extends from the ampulla. Each single afferent neuron can make contacts with multiple receptor cells. The ultrastructural characteristics of the ampullae of Lorenzini in Aptychotrema rostrata are very similar to those of other elasmobranch species that use electroreception for foraging.     

On the Early Development of the Protonephridial Systems in Some Species Belonging to the Genera Diphyllobothrium, Triaenophorus and Schistocephalus (Cestoda, Pseudophyllidea)

Malmberg, G. {Department of Zoology, University of Stockholm, Stockholm, Sweden.) On the early development of the protonephridial systems in some species belonging to the genera Diphyllobothrium, Triaenophorus and Schistocephalus {Cestoda, Pseudophyllidea). Zool. Scripta 1 (5): 227–228, 1972.–The protonephridial systems of coracidia (oncospheres) and procercoids of four Diphyllobothrium species, of Triaenophorus nodulosus (Pallas) and of Schistocephalus solidus (Muller) were studied. In the oncospheres of Diphyllobothrium and Schistocephalus the two flame bulbs of the primary protonephridial system were present, but not in the oncospheres of Triaenophorus. In Schistocephalus the two flame bulbs were found to be inactive in the oncosphere (studied inside the coracidium), but very active in the youngest procercoids, which may imply that the primary protonephridial system does not start its function until the oncosphere has entered the copepod body cavity. The primary protonephridial system of the Triaenophorus procercoids was totally (most specimens) or partly reduced. The secondary protonephridial system, however, began developing more or less simultaneously with the integumental hooklets, the cercomer and the first calcareous bodies, which is in accordance with what is described concerning Diphyllobothrium. The ciliated, excretory bladder described by Rosen in 1919 was found to be a posteriorly open invagination, surrounding the “cercomer shaft”. Reverse bends of the posterior main canals of the secondary system are located closely around the wall of, though very likely not emptying into this invagination.     

Ultrastructure of the Flame Bulbs and Protonephridial Capillaries of Microstomum sp. (Platyhelminthes,Macrostomida)

The terminal part of the protonephridia of Microstomum is formed by a branching proximal canal cell and (at least?) two terminal cells. Each weir consists of longitudinal (sometimes convoluted) ribs continuous with the cytoplasm of the terminal cell. Internal leptotriches arise from the terminal and proximal canal cells. Near the tip of the flame, the proximal canal cell tube is surrounded by the more external terminal cell and connected to it by a septate junction. Large cristate mitochondria are densely packed in the terminal and canal cells. The flame bulb of Microstomum differs markedly from that of other macrostomids (Macrostomum, Paramalostomum) examined. Phylogenetic implications are discussed.     

The excretory organs in Sphaerodorum flavum (Phyllodocida, Sphaerodoridae): a rare case of co-occurrence of protonephridia, coelom and blood vascular system in Annelida

The excretory organs of Sphaerodorum flavum (Sphaerodoridae) were investigated by TEM and reconstructed from serial ultrathin sections. These organs are segmentally arranged paired protonephridia, which are in close association with a well-developed blood vascular system. Each protonephridium consists of a terminal part made up of two monociliary terminal cells (solenocytes), and a nephridioduct, formed by two cells. The two solenocytes lie close together. Each cilium is surrounded by 12 microvillar rods projecting from the perikaryon of each solenocyte. These rods form a weir-like structure in the coelomic space. The distal part of the weir is embedded in the proximal nephridioduct. The largest part of the cell bodies of the solenocytes, containing the nucleus, is lateral or basal to the weir-like structures. The lumen of the nephridioduct is formed by two multiciliated cells, which enclose the extracellular nephridial canal one behind the other. The canal opens through the nephropore beneath the cuticle without penetrating the cuticle. Both nephridioduct cells are surrounded by a blood vessel, which is partially folded into several layers. The significance of a simultaneous occurrence of protonephridial excretory organs and a well-developed blood vascular system as well as coelomic cavities is discussed. The results of this investigation indicate a close relationship of Sphaerodoridae to Phyllodocidae instead of to Syllidae within the Phyllodocida. Accepted: 27 November 2000     

Ultrastructure of the protonephridia of Seison annulatus (Rotifera)

Summary Each of the two protonephridial systems of Seison annulatus consists of three sections which are separated by cell borders with septate junctions: (a) a terminal syncytium with eight terminal organs and a capillary canal, (b) a canal syncytium which is divided into a multiciliary canal region and a main canal region, and (c) a nephroporus cell. The terminal syncytium is branched and linked twice to the canal syncytium. The supporting structure of each filtration barrier is a hollow cylinder which is perforated by pores and lacks microvilli (pillars). A protonephridial spine is situated in the multiciliary canal region and stabilizes the neck region. The ored, hollow cylinder and the protonephridial spine are new characteristics for the Rotifera.     

Ultrastructure of the preseptal part of metanephridia in Nais variabilis and Dero digitata (Annelida, Clitellata)

The composition and arrangement of cells in the preseptal region of metanephridia have been examined by ultrastructural methods in two naidid species, Nais variabilis and Dero digitata. Within this region special attention has been paid to the portion around the orifice and the region where the metanephridium penetrates the septum. In N. variabilis, the preseptal region is composed of four cells and, in D. digitata, three cells are present. In both species three cells correspond in position and ultrastructural details and, hence, are interpreted as homologous. These are the mantle cell, the flame cell, and the canal cell. The mantle cell covers the preseptal region and surrounds the opening. The margin around the orifice is endowed with cilia, which extend into the coelomic space and beat irregularly. They do not enter the orifice and, thus, are not part of the internal ciliary flame. Posteriorly, in D. digitata, the mantle cell originates from the septal wall, i.e., its extensions spread in the plane of the frontal coelothelium of the septum. In N. variabilis, the mantle cell is continued by a further cell, enwrapping the posterior region of the preseptal part. This cell, called the septal cell, is anchored in the septal wall like the mantle cell in D. digitata. Both cells are interpreted as mesodermal components of the metanephridium. The flame cell lies beneath the mantle cell. In front, on its dorsal wall, many cilia are inserted which extend posteriorly into the nephridial canal forming a flame. In D. digitata, the caudal extension of this cell was examined in more detail; it originates from an intraseptal position. The canal cell lines the anterior lumen of the nephridial duct. While the mantle cell and flame cell enclose the organ from a dorsal position, the canal cell lies opposite embracing the lumen from a ventromedial position. Behind, it extends into the postseptal region for a certain distance. It is concluded that metanephridia in the Clitellata have a coelothelial component and, probably, are not just descendants of a single cell, the nephridioblast. The results further indicate that a flame cell and a mantle cell or some corresponding coelothelial cells may be constitutive elements of the ground plan of the clitellate metanephridium. Phylogenetic consequences for non-clitellate Annelida are discussed. Accepted: 21 December 1999     

Ultrastructure of the protonephridial system, of Anoplodiscus cirrusspiralis (Monogenea Monopisthocotylea).

The flame bulb is formed by a terminal cell and a proximal canal cell. The weir consists of interdigitating ribs all of which form one circle, i.e. alternating ribs do not have distinctly 'internal' or 'external' positions. Cytoplasmic cords are absent and all ribs, i.e. those continuous with the proximal canal cell and those continuous with the terminal cell, form external leptotriches. At least some external leptotriches have interconnected branches extending along the flame bulb. Internal leptotriches are not branched and arise from the basal perikaryon of the terminal cell. In the cytoplasmic cylinder at the tip of the flame bulb, structures resembling incomplete septate junctions were seen. However, neither the cytoplasmic cylinder nor the small protonephridial capillaries contain complete septate junctions as found in all other Monogenea Polyopisthocotylea, Monogenea Monopisthocotylea, Trematoda Aspidogastrea and Trematoda Digenea examined to date. In the lack of a septate junction, Anoplodiscus resembles Udonella, Amphilinidea, Gyrocotylidea and Eucestoda. However, the presence in this species of rudimentary septate junctions in the small capillaries and of complete junctions in larger ones indicates that complete junctions have been secondarily lost. Anoplodiscus resembles the Monogenea and Trematoda in the presence of lamellae in the larger protonephridial ducts. For the first time in a monogenean, the ultrastructure of the excretory bladder is described. A nucleated convoluted duct opens through a narrow connecting duct into the bladder, which in turn opens through a narrow connecting duct into the excretory pore lined by tegument. Convoluted duct, connecting ducts and bladder are lined by a lamellated epitheliu.(ABSTRACT TRUNCATED AT 250 WORDS)