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.     

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 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.     

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     

On the Larval Protonephridial System of Gyrocotyle and the Evolution of Cercomeromorphae (Platyhelminthes)

Eggs of Gyrocotyle urna specimens were hatched. Certain results from the hatching work are given. The protonephridial system of the larvae/lycophores was investigated. The system was found to be bilaterally symmetrical with two nephridiopores in the anterior body-half. The system is very similar to the posterior protonephridial system in the oncomiracidium (the monogenoidean larva). Unlike the oncomiracidium the Gyrocotyle lycophore has no anterior system. The hooks of amphilinidean and gyrocotylidean lycophores were found to be of the same basic type as those of other cercomero-morphaean larvae. A preliminary investigation of the hook ontogeny of Amphilina foliacea was made: all ten hooks appear to begin developing at the same time, the six ordinary hooks early assumed a similar shape and arrangement to those in the oncosphaera, the four aberrant hooks seem to be less aberrant in their earlier developmental stages. On a basis of the number of hooks and other facts relating to the larvae, cercomero-morphaean evolution at the larval level is claimed. Because there are no traces of a mouth, a pharynx and an intestine in the ontogeny of Gyrocotyle , the lack of these body parts in the lycophore is assumed to be a primitive condition. It is claimed that the monogenoidean protonephridial system originated from a system of the Gyrocotyle lycophore type and that the mouth, pharynx and intestine are secondarily evolved in the monogenoideans. It is assumed that mouth-, pharynx- and intestine-less creatures with an ingestive body-covering gave rise to the digenean and the cercomeromorphaean evolutionary lines. On a basis of the ontogeny and the life cycles in the different cercomeromorphaean groups it is claimed that the adult monogenoideans never reached a certain evolutionary level which is repeated in the ontogeny of the ces-todes, the amphilinideans and the gyrocotylideans.     

Ultrastructure of the Flame Bulbs and Protonephridial Capillaries of Artioposthia sp. (Platyhelminthes,Tricladida, Geoplanidae)

The protonephridial terminal complex of Artioposthia is formed by one or two terminal cells, each with a nucleus located in the lateral wall of the flame bulb, and probably two proximal canal cells forming the wall of the protonephridial capillary. The weir is restricted to the proximal parts of the flame bulbs and consists of convoluted slits separated by thick cytoplasmic columns. Cross-striated ciliary rootlets running parallel with and obliquely or transversely to the longitudinal axis of the flame bulbs strengthen the walls of the flame bulbs and, to a lesser degree, that of the capillary. Numerous cristate mitochondria are present in the terminal and proximal canal cells. Cytoplasmic processes extend from the terminal cells into the adjacent tissue, and narrow internal leptotriches extend from the cytoplasm of the terminal cells into the lumen of the flame bulbs. The wall of the capillary contains many interconnected, liquid filled spaces that communicate with the lumen of the capillary, and two septate junctions. Phylogenetic implications of the findings are discussed.     

Ultrastructure of the ciliary pits in the Geocentrophora group (Platyhelminthes,Lecithoepitheliata)

The ultrastructure of the paired lateral ciliary pits in several endemic species of Geocentrophora from Lake Baikal and in one cosmopolitan species, G. baltica, has been compared and the possible functional significance is discussed. The pit is composed of two distinctive parts; the bottom of the pit is an extensive sensitive area, filled with uni-and biciliary sensory receptors with reduced rootlets and numerous neurotubules. The walls of the pit are formed by several large dark cells, characterized by a dark cytoplasm with numerous mitochondria, a large nucleus, intracellular canaliculi, basal infoldings of the cell membrane, glycogen granules and a varying number of cilia. A protruding, densely ciliated ridge occurs along the anterior wall of the pit. The cilia have a strengthened rootlet system and seem to provide a strong water current into the pit. Dark cell processes penetrate the basement membrane of the pit and come into the vicinity of large cells with a cytoplasm similar to that of the dark cells of the pit. These large cells in their turn come close to the terminal parts of the protonephridial canals, containing a weir. Smaller protonephridial capillaries without a weir seem to open directly into the pit lumen. The morphological data obtained suggest that the ciliary pit in not only a sensory structure, but plays a part in osmoregulation and ion exchange as well.     

Ultrastructure of the flame bulbs and protonephridial capillaries of Prorhynchus (Lecithoepitheliata, Prorhynchidae, Turbellaria)

The flame bulb of Prorhynchus is formed by a single cell. Its nucleus is not located in the cytoplasm at the base of the flame. Cilia of the flame have cross-striated hollow ciliary rootlets converging towards their tips. The maximum number of cilia counted was 13. The weir consists of a single row of longitudinal ribs that contain longitudinal filaments and possess regularly arranged protrusions along their surface appearing as transverse bands in horizontal section. A 'membrane' of extracellular material extends between the ribs. and loose material fills the places between the ribs, with a denser layer midway between adjacent ribs. Distally, the ribs fuse to form a continuous tube without a junction. Small protonephridial capillaries lack junctions, larger ducts have lateral flames and patches of long microvilli. Large excretory ducts open into a ciliated and lamellated cavity which is connected by a narrow canal to the excretory pore. The terminal part of the canal close to the pore possesses many cilia and microvilli. Phylogenetic implications of the findings are discussed.     

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 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.     

The ultrastructure of the protonephridial system of Götte's larva of Stylochus mediterraneus (Polycladida, Platyhelminthes)

The protonephridial system of Götte's larva of Stylochus mediterraneus was studied by electron microscopy. There is one protonephridium on each side of the body, formed by one terminal and one canal cell. The terminal filtration apparatus is formed by a single cell (the terminal cell) with several globular processes, the largest of which includes the nucleus. Fingers of cytoplasm (leptotriches) from each process penetrate the lumen surrounding the bundle of cilia and fingers from adjacent processes interdigitate to form a pattern of convoluted slits which constitute the weir. The single canal cell is inserted internally to the terminal cell at the top of the weir and encloses the lumen without a junction. Septate junctions are present between the terminal and canal cells. The lumen of the canal cell is smooth-walled for most of its length and cilia arise and terminate at all levels of the terminal and canal cells. Posterior to the larval mouth opening, the canal cell crosses the epithelium and the lumen ramifies to form the excretory opening. The terminal apparatus closely resembles that found in the freshwater planarian Bdellocephala brunnea .     

A review of protonephridial excretory systems in Acanthocephala

Our present understanding of the excretory system of Acanthocephala is largely the result of work done by 5 German scholars: Kaiser, Schepotieff, Meyer, Kilian, and von Haffner. Present studies indicate that a protonephridial system is restricted to the family Oligacanthorhynchidae. However, many members of this family have not had a protonephridial system described. Three nephridial designs have been described: 1) dendritic type, organized as branches of a tree where each final branch terminates in a ciliated bulb; 2) capsular type, in which all ciliated bulbs empty directly into a common chamber; and 3) rudimentary type, consisting of a single cell with a patent ciliary pouch but no ducts to the outside. The first 2 types are a syncytia with 3 nuclei located in the capsule or stem wall and none in the flame bulbs. These excretory systems consist of 2 clusters of flame bulbs that empty separately into an expandable excretory bladder which in turn empties into ducts of the reproductive system. This urogenital system empties to the outside through a gonopore located at the tip of the penis in males and the posterior terminus of the vagina in females. Cilia occur in certain excretory tubes, depending on sex and species, but are unknown in the excretory bladder or ducts leading into it. The rudimentary type consists of a cell whose posterior extension terminates near the bursal lumen, but it is not known if this is significant for the discharge of material. There is no information on the physiology or biochemistry of the excretory system or its products.     

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 lycophora larva ofGyrocotyle urna (Cestoda,Gyrocotylidea)

Summary The fine structure of the glands of the lycophora larva ofGyrocotyle urna is described. There are four pairs of different glands which can be differentiated by the ultrastructure of their vesicles, their location and their gland pores. Their cell bodies are located at the transition from the median to the posterior third of the larva. Gland ducts formed by extensions of the gland cells run anteriorly and terminate at the anterior tip of the larva. In the cytoplasm of the terminal duct regions peripheral microtubules are found. The structure and function of the glands inGyrocotyle is discussed with regard to the evolution of cestodes and other Neodermata.     

Ultrastructural observations of the protonephridia of Polymerurus nodicaudus (Gastrotricha: Paucitubulatina)

Kieneke, A. and Hochberg, R. 2012. Ultrastructural observations of the protonephridia of Polymerurus nodicaudus (Gastrotricha: Paucitubulatina). —Acta Zoologica (Stockholm) 93 : 115–124. We studied different regions of the protonephridia of the limnic gastrotrich Polymerurus nodicaudus by means of light and electron microscopy to determine how freshwater species might differ from their marine relatives. Microscopic and ultrastructural characters are in accordance with another limnic species of Paucitubulatina, Chaetonotus maximus, whose protonephridial system has been previously reconstructed. Shared protonephridial characters of both species include the presence of highly elongate terminal organ cilia, microvilli, and the canal cell lumen as well as the presence of a conspicuous anterior loop of the protonephridial lumen. These features are not present in representatives of earlier, marine, paucitubulatan lineages (i.e., Xenotrichulidae) and so are assessed as evolutionary novelties that were likely important for the successful colonization of the freshwater environment.     

Ultrastructural characteristics of the protonephridial terminal organ and associated ducts of adult specimens of the Aspidogastrea, Digenea and Monogenea, with comments on the relationships between these groups

Transmission electron microscopical observations were made on the protonephridial terminal organ and associated ducts of three adult trematodes, the aspidogastrean Aspidogaster limacoides Diesing, 1835 and the digeneans Azygia lucii (Müller, 1776) and Phyllodistomum angulatum Linstow, 1907, and the monogenean Ancyrocephalus paradoxus Creplin, 1839. Previously unreported ultrastructural details of the terminal organ of adult trematodes include multiple contact sites (septate junctions and zonulae adherentes) between the membranes of the terminal and adjacent canal cells. Septate junctions traverse the epithelial cytoplasm of the canal wall, and the same type of septate junctions are observed within the cytoplasmic cord at the level of the tip of the flame tuft in both longitudinal and oblique sections of all three trematode species studied. In the monopisthocotylean Ancyrocephalus paradoxus, the absence of any junctions in the cytoplasmic cord and the presence of septate junction within all of the protonephridial ducts are reported. On the basis of the small number of monogenean species in which these features have been studied, in relation to the size of the group, there seems to be a high diversity in some characters of the protonephridial terminal organ. The study confirms that the Aspidogastrea and Digenea possess the same morphology of their protonephridial terminal organ and, although this differs slightly from that of most members of the Monogenea so far studied, it supports previous views on the close relationship of these groups.     

The evolution of the protonephridial terminal organ across Rotifera with particular emphasis on Dicranophorus forcipatus,Encentrum mucronatum and Erignatha clastopis (Rotifera: Dicranophoridae)

Riemann, O. and Ahlrichs, W.H. 2009. The evolution of the protonephridial terminal organ across Rotifera with particular emphasis on Dicranophorus forcipatus, Encentrum mucronatum and Erignatha clastopis (Rotifera: Dicranophoridae). —Acta Zoologica (Stockholm) 91 : 199–211 We report on the ultrastructure of the protonephridial terminal organ in three species of dicranophorid rotifers (Dicranophorus forcipatus, Encentrum mucronatum and Erignatha clastopis). Differences between the three species relate to shape and size, the morphology of the filter region and the number of microvilli and cilia inside the terminal organ. A comparison across Rotifera indicates that the terminal organs in D. forcipatus display a number of plesiomorphic characters, but are modified in E. mucronatum and Er. clastopis. This is in accordance with the results of phylogenetic analyses suggesting a basal position of D. forcipatus compared with the more derived species E. mucronatum and Er. clastopis. Moreover, we survey available data on the terminal organ in Rotifera and discuss its evolutionary transformations. The protonephridial terminal organ in the common ancestor of Rotifera consisted of a cytoplasmic cylinder with cilia united into a vibratile flame and a single circle of circumciliary microvilli. Depending on the topology on which characters are optimized, the site of ultrafiltration was formed by longitudinal cytoplasmic columns spanned by a fine filter diaphragm or by pores in the wall of the terminal organ. In several taxa of Rotifera, the terminal organ – probably independently – lost its circumciliary microvilli.     

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)     

The fine structure of protonephridia in Gnathostomulida and their comparison within Bilateria

Summary The fine structure of the protonephridia of Haplognathia rosea (Filospermoidea) and Gnathostomula paradoxa (Bursovaginoidea) is described. Each protonephridium consists of three different cells: (1) a monociliated terminal cell which constitutes the filtration area, (2) a nonciliated canal cell showing a special protonephridial outlet system, and (3) an intraepidermal cell — the nephroporus cell — constituting the nephroporus. The protonephridia are arranged serially. There is no canal system connecting the protonephridial units.Protonephridial characters in other Bilateria are considered. The pattern of characters in the protonephridia in the last common gnathostomulid stem species and presumed apomorphies in the protonephridia of the Gnathostomulida investigated are discussed.Abbreviations used in figures ac acessory centriole - AC additional epidermal cell - bb basal body - bl basal lamina - bm bundle of microvilli - c cilium - cc cilium duct cell - cd cilium duct - cr ciliary rootlet - crs structures resembling ciliary rootlets - di diplosome - ds desmosome - dy dictyosome - f filtration area - g granules - m mitochondrium - mv microvillus - n nucleus - NC nephroporus cell - np nephroporus - oc outlet canal - TC terminal cell - tl tubules of lacunar system     

The Terminal Protonephridial Complex of Haplopharynx rostratus (Platyhelminthes,Haplopharyngida)

The terminal protonephridial complex of Haplopharynx rostratus consists of three terminal cells. There are no weirs consisting of ribs connected by a filtration “membrane”, but some cytoplasmic outgrowths into the lumen of the terminal cells. Excretion is by exocytotic vesicles. The terminal cells also contain Golgi complexes and large membrane-bound vacuoles containing electron-dense material. The ciliary bundles (flames) of terminal cells 2 and 3 protrude into the lumen of the centrally located terminal cell I. The complex is surrounded by a sheath containing numerous filaments. The terminal complex of H. rostratus resembles that of the macrostomid Paromalostomum proceracauda, lending support to the view that the two taxa are closely related. © 1998 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd. All rights reserved