On the cilium cell relationship in the adenohypophysis of the mouse

Summary The 8+1 cilia previously reported in the adenohypophysis have been reinvestigated with special emphasis on their relation to the parent cell. In contrast to the fibril pattern which is remarkably constant, this relation shows great variation, supporting the hypothesis that the cilia are rudimentary.This study was supported in part by Grant NB 02215 of The National Institute of Neurological Diseases and Blindness, U. S. Public Health Service. This aid is gratefully acknowledged. The author wishes to thank Dr. Th. Blackstad for valuable advices and Mrs. J. L. Vaaland for skillful technical assistance.     

Some aspects of the fine structure of the statocysts of the molluscsPecten andPterotrachea

Summary The statocyst ofPecten is composed of hair cells and supporting cells. The hair cells bear kinocilia and microvilli at their distal ends and the supporting cells bear microvilli. The cilia have a 9+2 internal filament content, and arise from basal bodies that have roots, basal feet and microtubular connections. Two different ciliary arrangements are described, one with a small number of cilia arranged in a ring, and another with many more cilia arranged in rows. Below the hair cells are probable synapses. A ciliated duct connects to the lumen of the static sac and passes through the centre of the static nerve. The hair cells in the statocyst ofPterotrachea bear kinocilia and microvilli. The possible importance of cilia and microvilli in the transduction process is discussed.We would like to thank ProfessorJ. Z. Young for bringing specimens ofPterotrachea from Naples and also the staff of the Stazione Zoologica for the provision of specimens, Dr.M. Land for providing specimens ofPecten, the Science Research Council (U.K.) for providing the electron microscope used in much of the study and also for a grant to one of us (V.C.B.), and Mrs.J. Parkers and Mr.R. Moss and Mrs.J. Hamilton for much photographic and technical assistance.     

Fine structure of the Paraventricular organ of Xenopus laevis tadpoles

Summary The ultrastructure of the Paraventricular organ in the hypothalamus of Xenopus laevis tadpoles is described. It appeares that the Paraventricular organ of this anuran species is homologous with the Organon vasculosum hypothalami or the Paraventricular organ of other vertebrates.The Paraventricular organ of Xenopus laevis is characterized by an ependymal lining with only few cilia and by two types of nerve cells. Both types of nerve cells have ventricular processes, protruding into the lumen of the third ventricle and forming a network. The protrusions bear cilia of the 8+1 pattern. It has been possible to distinguish both types of nerve cells on account of their dense-core vesicles. A secretory function of both cell types is suggested.In a region close to the Paraventricular organ, another granulated type of nerve cell has been observed. A relationship between these cells and the preoptic nucleus is discussed.The author thanks Prof. Dr. P. G. W. J. van Oordt for his helpful comments and criticism, Mr. H. van Kooten for photographic assistance and Mr. F. Dijk for technical assistance.     

Changes in the fine structure of the human testicular interstitial cells after treatment with human gonadotrophins

Summary Study of the fine structure of the human interstitial cells after prolonged stimulation with human gonadotrophin reveals a striking increase in the quantity of the agranular endoplasmic reticulum. This is accompanied by an increase in the number of mitochondria which exhibit more extensive cristae, collections of intramitochondrial lipid and aggregations of electron-dense granular deposits. A rise is also evident in the number of lipofuscin pigment deposits and granular membrane-bounded bodies, both of which exhibit acid phosphatase activity. These changes after gonadotrophic stimulation are discussed in relation to steroid biosynthesis.In the pretreatment biopsies of these patients aged between 25–35 years, some interstitial cells contain intranuclear crystals which exhibit a hexagonal structure. The relationship of these intranuclear crystals to the cytoplasmic crystals of Reinke is discussed.The author is indebted to Dr. J. W. Johnstone and Dr. A. Long for the human material used in this study. Thanks are also due to Dr. H. P. Taft for helpful suggestions in the management of these patients, to Professor B. Hudson for the estimations of plasma testosterone and to Dr. J. B. Brown for the supply of human pituitary gonadotrophin and the estimations of urinary oestrogens. The technical help of Mr. T. Mezciems and the photographic assistance of Mr. J. S. Simmons F. R. P. S. and Miss S. Flett is gratefully acknowledged.     

The fine structure of sensory receptor processes in the auricular epithelium of the planarian,Dugesia tigrina

Summary The marginal epithelium of the lateral auricles of the planarian, Dugesia tigrina, includes a cell type with surface cilia and microvilli, a basal nucleus, and dense cytoplasm containing secretory vacuoles, Golgi elements, mitochondria and ribosomes. Through channels within the epithelial cytoplasm, cellular processes, interpreted as extensions of neurosensory receptor cells located in the subepidermis, project to the surface. The receptor processes, containing microtubules, mitochondria, vesicles and an agranular tubular reticulum, project beyond the epithelial cell surface; one or two cilia each emerge from a basal body in the apex of the projection. Close to the point of emergence to the epithelial surface, each cylindrical receptor process is surrounded by a collar-like septate junction between adjacent plasma membranes. The cilia of the projections differ from those of the epithelial cells in diameter, density of matrix and in the banding patterns of the rootlets. A few projections appear with the apex and basal body retracted below the epithelial surface. The possible function of these ciliated processes in sensory reception is discussed.This work was supported by Grant No. SO 1 FR 5369 from the U.S. Public Health Service to the University of Illinois at the Medical Center.I thank Dr. J. P. Marbarger, Director of the Research Resources Laboratory, for use of the electron microscope facilities, Miss Irena Kairys for technical help, Miss Marie Jaeger for assistance with photography, and Mr. Robert Parshall for the drawing.To Professor Arthur Wagg Pollister, I respectfully dedicate this article on the occasion of his retirement from Columbia University.     

The ultrastructure of primary cilia in the endocrine and excretory duct cells of the pancreas of mice and rats

A primary cilium was frequently observed in the endocrine alpha, beta and delta cells, as well as in the excretory duct cells of the pancreas of normal mice and rats. The characteristic components of the cilium including the basal body, axoneme (shaft), and terminal part were clearly recognizable. The basal body or distal centriole surrounded by Golgi vesicles was perpendicularly oriented to the proximal centriole, and a dense striated band was seen filling the gap between them. The microtubules of the basal body consisted of nine peripheral triplets exhibiting a 9 + 0 pattern, an appearance similar to that of the proximal centriole. Rootlets, basal feet and alar sheets associated with the basal body were occasionally seen. The axoneme usually consisted of a 9 + 0 pattern of microtubule doublets, but other irregular patterns of 7 + 2, 7 + 3, and 8 + 1 were also seen. The microtubules in the terminal part of the cilium became fewer in number and had no peculiar arrangement. The cilium of the endocrine cells always projected into the intercellular canaliculus and was covered by the ciliary sheath, and occasionally, double cilia were visualized in the vicinity of beta cells. In the excretory duct cells, the cilium showed similar features, but it was slightly longer and always projected into the dense secretory content of duct lumen. On the other hand, no primary cilium was ever observed in the acinar cells of mouse and rat pancreas. In conclusion, the present study describes the morphology of primary cilia and its associated components in the endocrine and excretory duct cells of the pancreas of mice and rats. The findings suggest that the primary cilium should be considered as a constant intracellular organelle though its function and significance remain speculative.     

The fine structure of the eye of the mollusc Pecten maximus

Summary The retina of Pecten maximus is divided into two light sensitive layers forming the distal and proximal retinae. The cells from these layers have different electrophysiological responses, the distal cells giving primary off responses, and the proximal cells giving on responses. The receptor surfaces of the distal retinal cells are formed from lamellae produced by the outer membranes of flattened cilia. These cilia have a basal body, basal foot, no root system and a 9 + 0 internal filament content. Each cell gives rise to an axon from its distal side, and this process goes up to the basement membrane, which is present below the cellular lens, passes along beneath it, and joins the distal optic nerve. The receptor part of the proximal retinal cells is formed from a vast array of microvilli. Each of these cells also bears one or two cilia with a probable 9 + 0 internal filament complement and no roots. The proximal cells give rise to axons, forming the proximal optic nerve. Below the proximal retina is a reflecting layer, the argentea, and below this is a pigment cell layer.We would like to acknowledge the advice and encouragement of Professor A. F. Huxley, Professor J. Z. Young and Dr. E. G. Gray. — We would like to thank Mrs. J. I. Astafiev for drawing Fig. 1, Mr. S. Waterman for photographic help and Miss C. Martin for clerical assistance.     

The fine structure of the olfactory mucosa and nerve in the teleost Carassius carassius L.

Summary Morphological studies on teleost olfactory mucosa confirm the findings of previous authors regarding the general arrangement of conventional cell types, viz. receptor, sustentacular, mucous and basal, but teleosts show certain distinct differences. The receptor cells have the general mammalian bipolar shape but their peripheral dendrite does not project beyond the epithelial surface. In addition to numerous typical cilia, an exceptional ciliary formation was observed in which the filaments, instead of forming individual cilia, are grouped together in clusters and are enveloped in a single limiting membrane.At the junction between the finger-like process and the mucosal fold myelinated nerve fibres are observed within the subepithelial stroma.Within the postero-medial zone of the mucosa is a conspicuous well-differentiated new cell type. A thick rim of electron-dense cytoplasm, bounded by an outer trilaminar membrane, encloses prominent foliate (leaf-like) organelles, a basal nucleus, numerous mitochondria and vacuolar spaces. These foliaceous cells communicate with the external environment through a small stoma, their close association with epithelial components suggesting a possible secretory or absorptive function. Their intricate morphology, however, suggests that they may be receptors, but their role and neural connections still require definition.Supported by Grant 5 RO 5 TWOO 154-02 from the National Institutes of Health, United States Public Health Service.The authors are indebted to Dr. A. S. Wilson for his helpful criticism and gratefully acknowledge the photographic technical assistance of Mr. J. Simmons and Mr. S. Frank.     

Cilia in the accessory hyperstriatum of the domestic fowl

Summary Numerous neurons and glia in the accessory hyperstriatum of the domestic fowl contain a cilium that is attached to a basal body. The accessory centriole is in the vicinity of the basal body and in some instances a connection between the two centrioles is noted. Cross-striated rootlets are associated with the basal body and the accessory centriole, however, some rootlets are found distant to centrioles. Cross sections of cilia show that most accessory hyperstriatal cilia have an 8+1 fiber pattern. Several proposed functional roles of neuronal cilia are discussed.This investigation was supported by a research grant from the National Institute of Neurological Diseases and Stroke (5 RO 1 NSO 7557-02) awarded to Norma Jean Adamo.     

Cilia and other surface structures of the trochophore of Harmothoë imbricata (Polychaeta)

Summary Ciliary aggregations occur as the prototroch, neurotroch, apical system and as tufts associated with the eyes and superficial glands. The major collection of cilia is the locomotory organ or prototroch that runs around the equatorial plane of the larva. This band is composed of four contiguous rows of cells, the two medial rows bearing the long locomotory cilia. The cilia occur in clumps, with several clumps arising from each prototroch cell while both the main cells contribute to each clump. The central filaments of these cilia are orientated at right angles to the long axis of the clump, the direction of ciliary beat being at right angles to the progression of the metachronal wave along the prototroch. The neurotroch, extending from the mouth to the posterior pole of the larva, beats away from the mouth. The rate of beating is rapid, and the cilia are short. The apical area of the larva is bordered by five single lines of compound cilia that surround a few stiff cilia. All the cilia beat occasionally. A further line of cilia, the akrotroch, exists at a position halfway between the apical area and the prototroch on the same side as the mouth. These cilia beat towards the prototroch. Some of these cilia are associated with sets of glandular openings. The fine structure of the glands and cuticle is described. The glands are small mucous glands that open via a projecting pore which is encircled by rings of microvilli. They often occur in groups of four or in pairs. The cuticle is similar to that described previously for adult polychaetes.This work was started under a Science Research Council (U.K.) grant (B/SR/1871) for a Research Assistantship to Dr. M. S. Laverack and grateful acknowledgement is made for this. We would like to thank Dr. A. Boyde for all his advice and use of apparatus. The scanning electron microscope used in this study was provided by a Science Research Council (U. K.) grant to Dr. Boyde. We would like to thank Mrs. J. Parkes for photographic assistance.     

The ciliated epidermis of Xenoturbella bocki (Platyhelminthes, Xenoturbellida) with some phylogenetic considerations

The epidermis of Xenoturbella bocki Westblad was studied by scanning and transmission electron microscopy. Two cell types predominate in the epidermis: multiciliated epidermal cells and non-ciliated or monociliated gland cells. A conspicuous feature is the dense ciliary coverage and the numerous gland cell openings. Xenoturbella has a characteristic pattern of axonemal filament termination in the distal tips of their cilia. Each epidermal cilium has the typical 9 + 2 patten through the major part of its shaft. Near the tip there is a shelf at which doublets 4–7 terminate. Doublets 1, 2, 3, 8 and 9 continue into the thinner distal part of the cilium. A similar shelf in cilia is known only from the turbellarian orders Nemertodermatida and Acoela, and hence may be an apomorphic feature which indicates a close relationship between Xenoturbellida, Nemertoder-matida and Acoela. The basal body is provided with a so-called basal foot which has a cross-striated appearance and an expanded distal plate that seems to act as a microtubule organizing center. Approximately 15–25 microtubuli radiate from the endplate of the basal foot to the basal bodies caudally. The arrangement of basal foot and ciliary rootlets in Xenoturbella differs from that of Acoela and related orders in that there are two striated rootlets only (an anterior and a posterior one), rather than one main rootlet and two lateral rootlets.     

Scanning electron microscopic studies of cilia

Summary A simple method for the preparation of ciliated epithelia for study with the scanning electron microscope is described. Ciliary groups are well preserved and it is possible to discern individual cilia and work out their numbers and orientation. Following scanning electron microscopical study some of the material was prepared for transmission electron microscopy and the ultrastructure of the tissue was found to be surprisingly well preserved. The tracheal epithelium of the rabbit, the olfactory epithelia of the goldfish and the rabbit, and the sensory epithelia in the statocyst of a cephalopod mollusc were examined with the scanning electron microscope to demonstrate the possibilities of the method. Acknowledgements. We would like to thank Professor J. Z. Young for his continued interest and support. The scanning electron microscope was purchased with a grant provided by the Science Research Council to Dr. Boyde, Mr. R. Willis helped in the initial stages of the study, Mr. G. Savage provided help with the goldfish material, Mr. S. Waterman provided much photographic assistance, and Mrs. N. Finney the secretarial assistance.     

Development of the ciliary complex and microtubules in the cells of rat subcommissural organ

Summary The fine structure of the rat subcommissural organs from the late stages of gestation through the postnatal to the adult stages was studied with the electron microscope. Emphasis in this report is placed on the development of the cilium with its affiliated structures. With the progress of cytodifferentiation centrioles originally located in the Golgi region migrate to the cell apex, where each then serves as a basal body to form a cilium which has a 9+2 organization of substructures. Thus, each of the mature subcommissural cells is provided with two cilia of motile type. Satellites first appear on one side of the basal body at about 17 fetal days, rapidly increase in number with age, and finally surround the basal body, forming an elaborate latticework. In the perinatal period microtubules progressively increase in number in the distal cytoplasm, which concurrently elongates and forms a prominent projection in the brain ventricle. Closely associated with the microtubules are large clusters of dense granular masses with an undefined border, which bear a close resemblance to the dense masses appearing in the differentiating cells of respiratory epithelium and having been generally assumed to be the precursor substance for centriole replication. However, the mature subcommissural cells contain no centrioles other than the preexisting pair, each of which has organized a cilium. The dense masses in the subcommissural cells are presumed to be involved in the formation of the cytoplasmic microtubules instead of new centrioles.Work supported by the National Science Council, the Republic of China, and by the China Medical Board of New York, Inc. A preliminary report was presented at the 6th International Congress for Electron Microscopy, Kyoto, 1966 (Lin, H.-S., andI-1. Chen: Satellites of the ciliary basal body and microtubules in the cells of the rat subcommissural organ. In: Electron Microscopy (R. Ueda, ed.) Vol. II, 461–462. Tokyo: Maruzen Co., Ltd. 1966).     

The Olfactory System in the Hagfish Myxine glutinosa

The apical part of the olfactory epithelium in Myxine glutinosa was investigated by optical and electron microscopy. This part of the epithelium consists of supporting cells and two types of olfactory receptor cells, i.e., ciliated receptor cells and microvillous receptor cells. The olfactory cilia have a 9 + 0 pattern of the microtubules, occasionally with one pair of the doublets dislocated towards the center of the cilium. Giant cilia were observed. The supporting cells bear microvilli and are rich in tonofilaments. The supporting cells also have a secretory function, their secretion consisting mainly of acid mucopolysaccharides. An asymmetrical type of desmosome was found between the olfactory receptor cells and the supporting cells.     

Morphology of the proboscis of Hubrechtella Juliae (nemertea,pilidiophora): Implications for pilidiophoran monophyly

The proboscis of Hubrechtella juliae was examined using transmission electron microscopy, scanning electron microscopy, and confocal laser scanning microscopy to reveal more features of basal pilidiophoran nemerteans for morphological and phylogenetic analysis. The proboscis glandular epithelium consists of sensory cells and four types of gland cells (granular, bacillary, mucoid, and pseudocnidae‐containing cells) that are not associated with any glandular systems; rod‐shaped pseudocnidae are 15–25 μm in length; the central cilium of the sensory cells is enclosed by two rings of microvilli. The nervous plexus lies in the basal part of glandular epithelium and includes 26–33 (11–12 in juvenile) irregularly anastomosing nerve trunks. The proboscis musculature includes four layers: endothelial circular, inner diagonal, longitudinal, and outer diagonal; inner and outer diagonal muscles consist of noncrossing fibers; in juvenile specimen, the proboscis longitudinal musculature is divided into 7–8 bands. The endothelium consists of apically situated support cells with rudimentary cilia and subapical myocytes. Unique features of Hubrechtella's proboscis include: acentric filaments of the pseudocnidae; absence of tonofilament‐containing support cells; two rings of microvilli around the central cilium of sensory cells; the occurrence of subendothelial diagonal muscles and the lack of an outer diagonal musculature (both states were known only in Baseodiscus species). The significance of these characters for nemertean taxonomy and phylogeny is discussed. The proboscis musculature in H. juliae and most heteronemerteans is bilaterally arranged, which can be considered a possible synapomorphy of Hubrechtellidae + Heteronemertea (= Pilidiophora). J. Morphol. 274:1397–1414, 2013. © 2013 Wiley Periodicals, Inc.     

Fine structural observations on the ciliary receptors in the epidermis of three otoplanid species (Turbellaria proseriata).

In Notocaryoturbella bigermaria, Otoplana truncaspina and Paroto-planella heterorhabditica three types of epidermal receptors are recognized. Type I: with a single cilium running in a duct, piercing the distal dendrite process of the receptor. The internal wall of the dendrite process has eight ridges with longitudinal filaments lying inside them. The ciliary basal body lacks a longitudinal rootlet but is encircled by a thin annular formation. Type II: with a single (A) or several (B) cilia which protrude from the outer epithelial surface and are provided with a large and striped rootlet. Both types are considered as mechanoreceptors. Type III: with two or more short and stumpy cilia devoid of rootlets and displaying the usual 9 + 2 pattern in the proximal part only. They are considered as chemoreceptors.     

Substructure of solitary cilia in mouse kidney

Summary Recent scanning electron microscopic studies confirm the presence of solitary cilia on most epithelial cells along the mammalian nephron and collecting ducts.By transmission electron microscopy we have found that the axonemata of such cilia consist of a maximal number of 9 doublet and no singlet filaments. 10% of the cross-sectioned cilia contain 9 doublets arranged in a peripheral ring (9+0 pattern). 30 % of the cross-sections contain 8 or 7 doublets in peripheral ring and 1 or 2 doublets in the central region (8+1 and 7+2 patterns). Serial sections and goniometer tilt reveal the central doublets to originate as dislodged peripheral doublets. 60% of the sectioned cilia contain filament numbers between 8 and 4. In patterns of 5 and 4 filaments single microtubules predominate.The functional significance of these atypical cilia is discussed.We are indebted to Prof. B. Afzelius and Prof. Th. Brun for valuable information and discussions during this work. The technical assistance of Miss K. Weltzin, Mr. E. Erichsen, Mr. R. Jensen and Mr. J. Røli is greatly appreciated     

The fine structure of atypical ciliated cells in the human gastric epithelium

Gastric mucosa obtained from the body and pyloric portions of the human stomach were observed by light and transmission electron microscopy. Ciliated cells were found in two of 18 subjects examined, one patient with gastric ulcer and the other one with gastric adenocarcinoma. The ciliated cells were found in epithelia at sites away from the main lesions. The tissues containing ciliated cells showed intestinal metaplasia combined with mild chronic gastritis in both cases. The epithelial layer facing the gastric lumen was composed of columnar cells with numerous uniform microvilli and goblet cells. This epithelium extended to the superficial parts of the tubules surrounded by the lamina propria. The deeper portions of the tubules were composed of mucous secretory, endocrine, and rarely ciliated cells. These ciliated cells were provided with numerous cilia the numbers of which varied considerably from cell to cell. This was in contrast to the primary cilium which is usually single. The central part of the apical cell membrane was sometimes concave in the area from where cilia tended to arise. It was also observed that numerous basal bodies as well as mucus-like granules were contained in the same cell. The axonemal pattern was different from that of ordinary cilia and showed 9 + 0 and 8 + 1 patterns. In longitudinal sections it was found that one peripheral doublet was displaced to the center of the axoneme as it left the basal body.     

Die Feinstruktur der Cerebraldrüse von Planorbarius corneus L. (Basommatophora)

Summary The fine structure of the cerebral gland of the fresh water snail Planorbarius corneus has been described. The apical cell surfaces of the epithelium bordering the lumen are covered with short microvilli and in some parts with cilia. There is a remarkable enlargement of the basal surface of those cells underlying the perineurium by long branched cell processes. The fine structure and density of the substances in the lumen differ; inclusions of paracrystallin structure are found. The possible functional significance of these organs in Pulmonates is discussed.

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Frau Prof. Dr. B. Scharrer zum 60. Geburtstag gewidmet.

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Mit dankenswerter Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Distinctive features of cilia in metazoans and their significance for systematics.

A comparative study of epidermal cilia in the Turbellaria and Nemertea has revealed features in these organelles that are specific to certain taxonomic groups. Turbellarians of the order Acoela, in particular, have a characteristic pattern of axonemal filament termination in the distal tips of their cilia and a characteristic ciliary rootlet system that is not seen in other turbellarian orders nor in other metazoans. Each epidermal cilium in acoels has a typical 9 + 2 axonemal pattern through the main part of its length, but near its distal tip there is an abrupt shelf-life narrowing at which filaments 4-7 terminate; filaments 1, 2, 8 and 9 continue into the thinner distal-most part of the shaft along with singlet microtubules from the axonemal center. The rootlet system in acoel cilia involves an interconnecting pattern with lateral connectives. The unique structure of these cilia has systematic and phylogenetic significance for the Acoela, and it is argued that ultrastructural characters in general, including characters of organelles, can be validly applied to the phylogeny and systematics of the Metazoa.