Ultrastructure of the Protonephridia in the Dorvilleid Polychaete Apodotrocha progenerans (Annelida)

The achaetous dorvilleid polychaete Apodotrocha progenerans Westheide & Riser, 1983, possesses several pairs of segmentally arranged protonephridia. They consist of a blindly ending terminal cell and three tubule (emission) cells. The terminal cell is a flame bulb with unusual filtration area consisting of interdigitated pedicel-like projections. Each cell has its own tuft of flagella; and the emission channel is intracellular. The tube-shaped 'seamless' cells are slotted into each other like water-pipes.     

The nephridia of the interstitial polychaete Hesionides arenaria and their phylogenetic significance (Polychaeta,Hesionidae)

Summary The small hesionid polychaete Hesionides arenaria possesses paired segmental excretory organs that closely resemble solenocytic protonephridia. The nephridium consists of one terminal cell and four tubule cells which form the emission channel. From the terminal cell, up to six flagella arise each surrounded by a weir of ten regularly arranged cytoplasmic rods. The structure of the cytoplasm of three of the following cells suggests that they function in active transport and storage. Because all of the larger, more primitive species of this family are equipped with metanephridia, the possibility is discussed that these organs have been developed out of metanephridia. The Hesionides arenaria nephridium may be a morphological stage in the evolutionary pathway from metanephridia to solenocytes. This would mean that solenocytes can no longer be considered to be homologous in every case with other protonephridial organs in polychaetes and may well be derived several times independently out of metanephridia or true protonephridia.     

The Nephridium of the Bonellia viridis Male (Echiura)

The nephridium of the dwarf male of Bonellia viridis was investigated by means of transmission electron microscopy. The nephridium proved to be of a distinct protonephridial type and not a metanephridium as maintained in the older literature. The nephridium is composed of a ciliated duct that projects into the coelom. Five crown cells at the end of the duct function as terminal filtration cells. Each crown cell has a bundle of about 20 cilia, surrounded by a labyrinthic weir of cell processes that are presumably involved in filtration. The ciliary bundles enter the nephridial duct through perforations of the adjacent tubule cells. This finding of a protonephridium in a minute, coelomate animal that lacks a circulatory system corroborates a recently formulated functional theory on the distribution of nephridial types.     

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     

A new interpretation of plasmodesmatal ultrastructure

Summary It is shown that simple, unbranched, plasmodesmata between young xylem ray cells of willow have no direct intercellular continuity apart from the plasmalemma which limits the cytoplasm and lines the plasmodesmatal canal. Each plasmodesma is traversed by a 200 Å diameter tubule (the desmotubule) which has a wall with probably 11 subunits arranged around a central cavity through which runs a 40 Å diameter rod. This rod is connected to the inside of the tubule wall, by fine filaments. At the ends of each plasmodesma the plasmalemma and cell wall are closely appressed to the tubule, thus precluding direct continuity between the cytoplasm of adjacent cells. Through the central part of the plasmodesmata the tubule is separated from the plasmalemma by a 90–100 Å wide gap. Cytoplasmic microtubules in the same tissue have a diameter of approximately 250 Å and a wall probably composed of 13 subunits: both desmotubules and cytoplasmic microtubules therefore have a centre-to-centre subunit spacing of about 47 Å. It is suggested that the desmotubules are not microtubules but may be nuclear spindle fibres which become trapped in the wall during cell plate formation. The endoplasmic reticulum, while closely approaching the plasmodesmata, is not continuous across them. It is thought most unlikely that the endoplasmic reticulum traverses plasmodesmata, as the dimensions of the central tubule — found here as well as by other workers — are smaller than those which would be expected to allow a stable molecular configuration in a unit membrane. The plasmalemma, where it lines the plasmodesmatal canal, appears to have particulate subunits in the outer opaque layers and the presence of these subunits may be attributable to the need for stability in membranes arranged about so small a radius.     

Developmental origin of segmental differences in the leech ectoderm: survival and differentiation of the distal tubule cell is determined by the host segment

The body plan of the adult leech is metameric, with each hemisegmental complement of ectodermal and mesodermal tissues being produced from a set of seven serially repeated embryonic blast cells. Previous studies have shown that homologous o blast cells give rise to an almost identical complement of descendant cells in each of the 21 abdominal segments, but that one o blast cell derivative--the distalmost cell of the nephridial tubule--is only present in 15 abdominal segments in the mature leech. Here we show that all o blast cells generate a presumptive distal tubule cell and that this cell migrates to its normal position in all abdominal segments. However, in segments which normally do not contain the mesodermal portion of the nephridium, the distal tubule cell dies before undergoing its terminal morphological differentiation. To ascertain whether the fate of the distal tubule cell is determined by its lineage history or by the segmental environment into which it is born, we utilized a previously described procedure for altering the segmental register between different embryonic cell lines. This procedure allowed us to effectively transplant o blast cells into more posterior segments prior to the cell divisions which generate their descendant clones. The results indicate that the survival or death of the distal tubule cell is determined by the identity of the host segment and that a given distal tubule cell could be effectively murdered or rescued by slipping its blast cell precursor into an appropriate segment. These findings suggest that the segment-specific pattern of distal tubule cell survival is not inherent to the O cell line, but arises from interactions with surrounding tissues.     

Regional cytology and cytochemistry of the crayfish kidney tubule

Cytological and cytochemical methods were used to identify and characterzie six distinct regions of the crayfish kidney: coelomosac, labyrinth I and II, and nephridial canal I, II, and III. Cells of the coelomosac possess cytoplasm which is strongly PAS-positive, but poor in RNA and protein. Their nuclei possess unusual projections which extend to the basal plasmalemma. Labyrinth I contains columnar cells rich in glycogen. Labyrinth II is characterized by a distended lumen and by shorter cells with high cytoplasmic RNA, many possessing a large intracellular vacuole. A PAS-positive brush border is unique to the two portions of the labyrinth. Cells in the nephridial canal show strong reactions for RNA and Mg⁺⁺-dependent ATPase. In nephridial canal I and II, cells are flattened to cuboidal with the lumen being more distended in nephridial canal I than anywhere else in the tubule. In nephridial canal III, the cells are large and columnar, and the cytoplasmic RNA concentration is greatest apically. Nuclei in all regions of the tubule epithelium, except coelomosac, are large and react strongly for protein. Coelomosac nuclei and those in blood cells are condensed and contain little protein. The cytoplasm of blood cells displays a significant amount of RNA, and traces of polysaccharide material. These observations demonstrate the presence of highly specialized morphological and histochemical alterations along the length of the kidney tubule and indicate sequential modification of urine in the lumen. Evident morphological and cytochemical likenesses between analogous regions of the mammlian nephron and the crayfish kidney tubule suggest that basic physiological similarities may also exist.     

Ultrastructure of the tubular nephron of the lizard Podarcis (= Lacerta) Taurica

The proximal, intermediate, and distal convoluted tubules of the neprhon of Podarcis (= Lacerta) taurica were examined by electron microscopy. Proximal tubule cells have large, apical cytoplasmic protrusions and microvilli interpreted to function in urate secretion. Adjacent cells are bound apically by tight junctions and desmosomes but interdigitate in their basal region. This situation is repeated in the other tubules with significant differences in intercellular space width. The basal surfaces bear numerous cytoplasmic processes. The intermediate tubule has proximal and distal segments each with dark, ciliated, and light cells, the cuboidal dark cells with dense cytoplasm constituting the main bulk of the wall. As the cells of the proximal and distal segments resemble those of the proximal and distal convoluted tubules, respectively, the intermediate tubule is considered as a transition region. The ciliated cell body has two broad processes extending from the lumen, one to the basement membrane and one to a foot process of a light cell. The light cell is surrounded by dark and ciliated cells. It does not reach the lumen, but contacts the basement membrane through a process running below a ciliated cell to form a mushroom-shaped structure in tubule cross-section, the light cell process forming the stalk and a ciliated cell the cap. The cilia probably propel the glomerular filtrate towards the distal convoluted tubule. This latter tubule has initial, middle, and terminal zones, all nonciliated but with different lumen widths and cell shapes.     

The lamina propria of the bovine seminiferous tubule

Summary The boundary tissue of bovine testicular seminiferous tubules exhibits remarkable regional differences at the level of the seminiferous tubule proper, as compared with its terminal segment. The basal lamina of the seminiferous tubule proper is multilayered and possesses knob-like protrusions. At the level of the terminal segment the basal lamina is highly specialized; in the region of the terminal plug candelabrum-like projections of the tubular basal lamina invade the bases of the modified supporting cells up to a depth of 3.5 m. The adjoining surface of these supporting cells is densely studded with hemidesmosomes. The elongated peritubular cells are arranged in 3–5 concentric layers around the tubulus seminiferus proper but form a loose association at the level of the terminal segment. Where the terminal segment joins the testicular straight tubule, peritubular cells may assemble to constitute a contractile spiral. Elastic tissue is situated mainly subjacent to the tubular basal lamina and to a lesser degree between the peritubular cell layers. A peritubular space lined by endothelium-like cells may surround the seminiferous tubule proper and also the transitional zone of the terminal segment.Supported by a grant from the Deutsche Forschungsgemeinschaft     

Ultrastructure of the metanephridial system in Aeolosoma bengalense (Annelida)

Summary The excretory system of Aeolosoma bengalense has been examined by light and electron microscopy. The system consists of seven serially arranged paris of metanephridia and six pairs of podocytes (referring to the first zoid of an animal chain). The podocytes surround blood spaces of the alimentary canal forming dorsoventrally running loops that emerge on both sides of it. The two elements of the system have a correlative position, each podocyte extending in close proximity to the funnel of a metanephridium. Only in the region of the first metanephridia are podocytes lacking. The nephrostome of the metanephridia consists of two cells, an inner one, the terminal duct cell, and an outer one enwrapping it, called the mantle cell. Nephrostomal cilia that extend into the coelomic space arise exclusively from the rim of the mantle cell whereas those of the terminal duct cell arranged on its luminal surface protrude into the canal forming a flame. The nephridial canal is ciliated throughout and is either intra- or extracellular. Its initial loops aggregate to form a compact organ, the nephridial body. The middle part of the duct constitutes a loop that ascends at each side of the alimentary canal where it is in intimate contact with its blood spaces. Ultrastructural features of the duct cells suggest a reabsorptive function in two regions, the nephridial body and the uppermost part of the loop. The terminal part of the duct passes through the nephridial body and opens ventrolaterally. Generally, the transverse vascular loops at the gut consist of one podocyte each. In the oesophageal region, where only one pair of podocytes is present, the loops connect the dorsal with the ventral longitudinal vessel. Three pairs of podocytes are present in the dilated region of the intestine emerging from its lateral wall and joining the median ventral vessel or blood spaces near by. In the hind gut, where two pairs of podocytes occur, the loops arise from the dorsolateral part and enter directly the ventral vessel. Cytological features of podocytes resemble those of other animals. The results are discussed on the basis of current theories on the function and the phylogenetic significance of excretory systems in the Annelida.Abbreviations bl basal lamina - bs blood space - bv blood vessel - cf ciliary flame - ci cilia - co connection of the vascular loop with the intestinal blood space - cu cuticle - db dense body - dc duct cell - di dictyosome - za zonula adhearens - dv dorsal vessel - ecb epicuticular body - ev endocytotic vesicle - ic intestinal cell - ici inner cilia - iv intestinal vessel - lm longitudinal muscle - mc mantle cell - mg midgut - mi mitochondrion - mv microvilli - nu nucleus - oci outer cilia - oe oesophagus - pc podocyte - pe pedicel - pel primary elongation of the podocyte - sm slit membrane - tc terminal duct cell - ve vesicle with heterogeneous contents - vv ventral vessel     

Ultrastructure of the Protonephridial Terminal Organ in the Terrestrial Nemertean Geonemertes pelaensis (Rhynchocoela,Enopla, Hoplonemertini)

The protonephridial terminal organ of Geonemertes pelaensis consists of two cells that are equal in both size and shape and form mirror images of each other. From the perinucleate lump-shaped region basally in each cell arises a cytoplasmic column which branches at regular intervals to form 3–4 nearly circular bars. The opposed columns and alternating bars of the two cells are arranged in such a way that they form an obviously rigid, cylindrical structure which both supports and gives rise to the thin-walled weir. The fenestration of the weir consists of a single, enormously extended and sinuous cleft which represents part of the boundary between the two terminal cells. The ciliary flame comprises 92–118 cilia that originate basally in the two cells and is surrounded by a palisade of long, straight microvilli, positioned immediately within the weir's wall. The structure is very similar to that of the terminal organ of the land nemertean Pantinonemertes, except that the cytoskeleton which supports columns and bars consists of a fibro-granular substance, not, as in Pantinonemertes, of an abundance of oriented microtubules.     

Structure and development of the nephridia of Tomopteris helgolandica (Annelida)

 Nephridial diversity is high in Phyllodocida (Annelida) and ranges from protonephridia to metanephridia. The nephridia of Tomopteris helgolandica (Tomopteridae) can be characterized as metanephridia which bear a multiciliated solenocyte. This cell is medially apposed to the proximal part of the nephridial duct and bears several cilia, each of which is surrounded by a ring of 13 microvilli. An extracellular matrix connects the microvilli and thus leads to the impression of a tube surrounding the central cilium. Each tube separately enters a subjacent duct cell and the cilia extend into a cup-shaped compartment within the duct cell. This compartment is not connected to the duct. The funnel consists of eight multiciliated cells and is connected to the nephridial duct, which initially runs intercellularly and later percellularly. The last duct cell bears a neck-like process which pierces the subepidermal basal membrane and is connected to epidermal cells forming a small invagination, the nephropore. The nephridia of T. helgolandica develop from a band of cells and all structural components are differentiated at an early developmental stage. Further development is characterized by enlargment of the funnel, ciliogenesis in the solenocyte, merging of different sections of the duct and, finally, the formation of the nephropore. An evaluation of the nephridia of T. helgolandica leads to the hypothesis that the nephridial diversity in Phyllodocida can be explained by the retainment of different stages in the transition of protonephridia into metanephridia; this is caused by the formation of a ciliated funnel at different ontogenetic stages. Although the protonephridia in Phyllodocida are regarded as primary nephridial organs, protonephridia are also presumed to have evolved secondarily in progenetic interstitial species of the Annelida by an incomplete differentiation of the nephridial anlage. Accepted: 18 December 1996     

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     

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     

Immunoelectron microscopy of fodrin in the rat uriniferous and collecting tubular epithelium

We examined the localization of fodrin in epithelial cells of rat uriniferous and collecting tubules by immunofluorescence and immunoelectron microscopy of frozen sections. In the uriniferous tubule, fodrin was found along the cell membrane and in the well-developed terminal web, as previously reported in other epithelial cells: in the terminal web and along the basolateral cell membrane in the proximal tubule; all around the cell surface in the thin limb of Henle; along the basolateral surface in the thick limb of Henle's thick segment and the distal tubule. In the intercalated cells of the collecting tubule, fodrin was found not only along the basolateral cell membrane but also in the apical cytoplasm. The most peculiar labeling was obtained in the principal cells of the collecting tubule. In addition to labeling in the basolateral cell membrane, fodrin was found diffusely in the cytoplasmic matrix. Association of fodrin with any particular structure could not be identified, but the Golgi area was apparently free of labeling. Cytoplasmic labeling was more conspicuous in the principal cells of the medulla than in those of the cortex. The present results show that fodrin need not always exist in association with the cell membrane or the cytoskeleton but can occur in the cytoplasmic matrix, at least in epithelial cells. We discuss the possible physiological significance of the latter distribution.     

A neurofibrillar body in the perikaryon of nervus terminalis ganglion cells in teleosts

Summary In three species of Teleosts (Tinea tinea L., Leuciscus cephalus cabeda Risso, Epinephelus guaza L.) a round strongly argentophilic body of considerable size occurs in the cytoplasm of the nervus terminalis ganglion cells. In Tinea, surgical interruption of functional connections of the ganglion cells does not produce any apparent change either in the number or in the size of these cytoplasmic bodies.Electron microscopical observations show that the neurofibrillar body is made up of densely packed and irregularly arranged bundles. In cross section, each of these bundles appears to be composed of neurofilaments (100 Å in thickness) and neurotubules (diameter: 300 Å). Each tubule is surrounded by 9–10 filaments equi-distant from one another, and at a distance of 30–40 Å from the central tubule.The authors are indebted to Prof. G. Palladini for helpful histochemical advice, to Prof. B. Bertolini for electron micrographs and to Mr. D. Scorsini for skilful technical assistance.     

Ultrastructure of the Flame Bulbs of Urastoma cyprinae (Platyhelminthes, ‘Prolecithophora’, Urastomidae)

The ultrastructure of the flame bulbs of the turbellarian Urastoma cyprinae from Mytilus galloprovincialis in the Mediterranean is described. The nucleus of the terminal cell is located some distance basal to the rootlets of the cilia forming the flame; the cytoplasm contains numerous tubules approximately 54–66 nm in diameter, and vesicles. Thick walled, densely packed rod-like structures coil around each other with a tendency towards longitudinal orientation close to the flame. The rod-like structures tightly surround the basal part of the flame and the distal cytoplasmic tube in the apical part of the flame. Some of them, including the inner predominantly longitudinally directed ones, are continuous with the cytoplasm of the terminal cell, others are continuous with the cytoplasm of the distal cytoplasmic tube. Internal leptotriches arise from the cytoplasm of the terminal cell and intrude between the basal parts of the cilia of the flame. The distal cytoplasmic tube possesses a septate junction. The flame bulb of Urastoma differs distinctly from those known from other Platyhelminthes; implications for the phylogeny of Platyhelminthes are discussed.     

The fine structure of the maxillary gland of the brine shrimp,Artemia salina: The efferent duct

Summary The efferent duct of the maxillary gland of adult brine shrimp, Artemia salina, is specialized into two morphologically distinct regions: an efferent tubule and a terminal duct. The wall of the efferent tubule is composed of epithelial cells which possess an apical microvillous border and, more basally, membranous configurations with which large numbers of mitochondria are closely associated. These membranous configurations are of two types: 1) infoldings of the plasma membrane of a single cell, and 2) interdigitations of lateral processes from adjoining cells. In contrast to the efferent tubule, the cells of the terminal duct possess a secreted cuticle and lack modifications which markedly increase the area of the plasma membrane. Mitochondria of the terminal duct are smaller and less numerous than those of the efferent tubule and typically are not found in close association with the plasma membrane. The ultrastructure of the efferent tubule and terminal duct suggests that the former region plays an active role in modifying the luminal contents and the latter region functions primarily as a conduit for the final urine.     

PROTONEPHRIDIA

(1) The flame cell of platyhelminths is a composite organ formed from two cells. One cell contains a large nucleus and bears the flagella which form the flame. The other cell which contributes the barrel is the first tubule cell. There is a region of interdigitation between the two cells at the top of the barrel and the interdigitations are joined along their length by desmosomes. The tubule lumina are extracellular, the smaller tubules, at least, being formed by encircling projections from a single cell, joined at their tips by desmosomes. Cestodes apparently have no desmosomes in their tubule walls so that the tubule lumina may be intracellular. The tubules of most platyhelminths are lined by folds or microvilli, and flagella may be present in the lumen. The protonephridia of nemertines, entoprocts and priapulids appear to be of this type. (2) Direct evidence of the physiological role of flame cell systems is limited. There is, for example, no proven instance of the production of urine hypo-osmotic to body fluids by any fresh-water platyhelminth or nemertine. Endoparasitic platyhelminths are apparently unable to osmoregulate. The relative permeability of the body surface of marine, fresh-water, terrestrial and parasitic platyhelminths and nemertines may be related to protonephridial function. It seems highly likely that the function of the flame cell is to filter interstitial fluid, separating water and crystalloids from macromolecules. The ultrafiltrate produced then flows down the tubules as a result of the hydrostatic pressure generated by the beating of flame flagella, or as a result of peristaltic waves of the whole body generated by the musculature of the worm. The fluid may be modified in the canal lumen by both active and passive resorption of solutes or the secretion of material from the walls into the lumen. Experiments with the larger platyhelminths suggest that the main function of the system is to remove organic metabolites from the interstitial spaces of the deeper tissues of the worm by a mechanism more efficient than simple diffusion. (3) The flame bulbs of rotifers are fan-shaped and the nucleus is in the tubule not the cap. The barrel of the flame bulb is composed of a series of columns in scalloped formation, each arc of the scallop being supported by a cytoplasmic pillar. A membrane interconnects the columns and each column is linked to its neighbouring central pillar by fibrils. The tubules leading from the flame bulbs are a complex system of three to four multinucleate cells. They empty into a contractile bladder. The protonephridia of acanthocephalans and gastrotrichs may be of this type. (4) The mode of action of flame bulbs is probably to filter the pseudocoelomic fluid which is then modified by selective reabsorption in the tubule system. Rotifers are able to osmoregulate and this may be the chief function of their protonephridia. (5) Solenocytes are morphologically diverse, usually with a cytoplasmic cap containing a nucleus, and a long tubule, in the lumen of which lie one or two flagella. The walls of the tubule are pierced by fenestrations, probably the site of fluid ‘filtration’. (6) Kümmel (1962) has suggested that the many types of terminal organ are the result of divergent evolution from a single ancestral type. We suggest that protone-phridial terminal organs can be divided on structural grounds into three or four different groups which are probably not inter-related. This would mean that the apparent structural similarities which do appear would be the result of convergent evohtion imposing a conformity based on functional requirements.