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PubMed2006

Loss of primary cilia results in deregulated and unabated apical calcium entry in ARPKD collecting duct cells.

Recent genetic analysis has identified a pivotal role of primary cilia in the pathogenesis of polycystic kidney disease (PKD). However, little is known regarding how cilia loss/dysfunction contributes to cyst development. In epithelial cells, changes in apical fluid flow induce cilia-mediated Ca2+ entry via polycystin-2 (PC2), a cation channel. The Oak Ridge Polycystic Kidney (orpk) mouse contains a mutated Tg737 gene that disrupts expression of polaris, a protein required for ciliogenesis. These studies examine the effect of cilia malformation on Ca2+ entry in orpk cilia(-) collecting duct principal cells, and in orpk cells in which wild-type Tg737 was reintroduced, orpk cilia(+). [Ca2+]i was monitored in confluent cell monolayers using fluorescence microscopy. Intrinsic apical Ca2+ entry was measured by Mn2+ quenching and Ca2+ depletion/readdition under flow conditions below the threshold for stimulation. We found that unstimulated apical Ca2+ entry was markedly increased in cilia(-) cells and was sensitive to Gd3+, an inhibitor of PC2. Electrophysiological measurements demonstrate increased abundance of an apical channel, consistent with PC2, in cilia(-) cells. Immunofluorescence studies revealed that PC2, normally expressed on and at the base of cilia in orpk cilia(+) cells, was observed throughout the apical membrane in cilia(-) cells. Furthermore, cilia(-) cells displayed elevated subapical Ca2+ levels measured with the near-membrane Ca2+ indicator FFP-18. We propose that cilia exert a tonic regulatory influence on apical Ca2+ entry, and absence of cilia results in loss of spatial organization of PC2, causing unregulated Ca2+ entry and elevations in subapical [Ca2+], a factor which may contribute to cyst formation.

Animals↗

PubMed1986

Mechanical stimulation activates beating in calcium-arrested lateral cilia of Mytilus edulis gill.

Lateral cilia of Mytilus edulis gill arrest upon mechanical stimulation, the result of calcium influx. A mechanical stimulus that deflects these cilia toward the effective stroke, and is normally sufficient to cause transient arrest in beating lateral cilia or transient movement into the recovery stroke in quiescent cilia, initiates beating in Ca2+ ionophore-arrested cilia at 9-15 Hz, for periods as long as 30 s. This movement is restricted to the stimulated cilia and the beat pattern appears constrained in the first half of the beat cycle. Application of dopamine causes ciliary arrest in the presence (but not absence) of Ca2+ and mechanical stimulation will also activate such cilia to beat. In the presence of ATP, mechanical stimulation of detergent-permeabilized lateral cell models arrested in the presence of 50 microM Ca2+ will also cause activation comparable in frequency, duration, and beat pattern to that seen in Ca2+-arrested cells, but the initiation is more difficult. Upon application of ionophore in Ca2+-free (EGTA) seawater, the cilia become quiescent, stopped at the end of the recovery stroke. Mechanical stimulation will cause activation of beat, with a similar range of frequencies and duration as in Ca2+-arrested lateral cilia, but the beat pattern is normal and cilia of adjacent cells may also beat, presumably initiated by mechanical coupling. Gently lifting cilia at their basal ends, using small, slow movements of a mechanical probe, will initiate several beat cycles in quiescent lateral cilia but will cause Ca2+-arrested cilia to 'snap' into the effective stroke and back.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1980

Biochemical studies of olfaction: isolation, characterization, and odorant binding activity of cilia from rainbow trout olfactory rosettes.

The role of cilia in recognition of olfactory stimuli has been controversial. Cilia from the intact olfactory rosettes of the rainbow trout Salmo gairdneri were isolated, characterized biochemically, and examined by electron microscopy. The markers studied are those associated with cilia in other organisms. Dynein arms contain Mg2+-AtPase; this enzyme was enriched in the isolated cilia preparation. Guanine nucleotides are associated with the outer microtubule doublets of cilia but adenine nucleotides are not; a substantial enrichment in guanine, relative to adenine, was found in the cilia preparation. Tubulin, the structural protein component of microtubules, occurs in large amounts in cilia. Disc gel electrophoresis indicated tubulin in the cilia preparation. Electron microscopy confirmed the presence of cilia in the isolated preparation. Rainbow trout have an acute sense of smell and many amino acids are odorants to this species. Functional activity of the cilia preparation relevant to odorant recognition was assessed by using binding of radioactively labeled odorant amino acids. L-Alanine, L-serine, L-threonine, L-lysine, and D-alanine bound to the cilia preparation. This study provides direct biochemical evidence that olfactory cilia bind odorant molecules and supports the hypothesis that odorant recognition sites are integral parts of the cilia.

Amino Acids↗

PubMed2004

Motor or sensor: a new aspect of primary cilia function.

Cilia are hair-like structures that protrude from the surface of the cell and are evolutionary well conserved. The characteristic feature of cilia is their motility and, in ciliated epithelia such as the trachea, their principal function is to transport materials along the cell surface. Each epithelial cell has many cilia on its surface. As well as this multiple form of motile cilia seen in the epithelium, there are primary cilia, also known as a monocilium because each cell has only one cilium. These types of cilia are thought to be non-motile because they lack a central pair of microtubules, are anomalous and have no function. However, recent studies have shown that primary cilia are involved in both developmental and pathological processes, including the establishment of left-right asymmetry and polycystic kidney disease. During development, cells in the node rotate their primary cilia to produce an extracellular current that is essential for the determination of left-right asymmetry of the body. In the kidney, primary cilia act as mechanosensors to detect fluid flow. Without such cilia, the kidney develops multiple cysts that eventually destroy kidney function. Furthermore, studies have identified a variety of proteins that are localized in the cilia and their diverse roles in various ciliary functions. These studies suggest the diversity of primary cilia. To elucidate how ciliary proteins interact and perform their functions in primary cilia will help us understand both their function and their diversity.

Animals↗

PubMed1991

Acetaldehyde-mediated cilia dysfunction in bovine bronchial epithelial cells.

Acetaldehyde, which is present in significant concentrations in cigarette smoke and is elevated during alcohol ingestion, has been demonstrated to impair mucociliary clearance of the lung. Acetaldehyde is also known to impair protein function through the formation of acetaldehyde-protein adducts. We hypothesized that acetaldehyde impairs bronchial epithelial cilia motion by inhibiting cilia dynein adenosinetriphosphatase (ATPase) activity through the formation of acetaldehyde adducts with cilia proteins. Acetaldehyde induced concentration- and time-dependent slowing of cilia beating and cilia-derived dynein ATPase activity in primary cultures and isolated axonemes of bovine airway epithelial cells. Cilia slowing and ATPase inhibitory effects were also observed with related aldehydes but not with ethanol. Acetaldehyde binding, assessed by gel electrophoresis using [14C] acetaldehyde, was demonstrated to occur with the dynein heavy chains and with tubulin and closely paralleled ATPase inhibition. We conclude that acetaldehyde directly impairs bronchial cilia function causing slowing of cilia beating, inhibits cilia dynein ATPase activity, and binds to cilia proteins critical for motion including dynein and tubulin. These data suggest that acetaldehyde-induced cilia dysfunction may be related to direct cilia ATPase inactivation and adduct formation with cilia dynein and tubulin. This may be an important mechanism by which airway host defenses are impaired in clinical settings where acetaldehyde exposure occurs, e.g., with cigarette smoking and alcohol ingestion.

Acetaldehyde↗

PubMed1988

Pre-natal development of rat nasal epithelia. V. Freeze-fracturing on necklaces of primary and secondary cilia of olfactory and respiratory epithelial cells.

Many cilium types have at their proximal base a particulated membrane structure, the so-called ciliary necklace. Necklaces of primary and secondary cilia of olfactory receptor cells and ciliated respiratory cells, and of primary cilia of olfactory supporting cells were studied as a function of embryonic age. Strand numbers in necklaces of primary cilia of these cell types do not differ, but they differ significantly from those of necklaces of secondary cilia. Primary cilia have 2 to 4, but most commonly 3, necklace strands. This is true for necklaces of primary cilia of 8 different nasal cell types: olfactory epithelial basal and glandular cells, vomeronasal receptor and supporting cells, and microvillous respiratory epithelial cells, in addition to the 3 cell types mentioned above. Comparison with other systems suggests that primary cilia resemble flagella of eukaryotic flagellates and spermatozoa of some invertebrates with respect to their number of necklace strands. Average numbers of necklace strands in secondary olfactory cilia increase from 3-4 at the 16th and 17th gestational days to 6-7 in adults. Those in secondary respiratory cilia increase from 2-3 at the 18th and 19th gestational days to 5-6 in adults. Longer cilia have more strands than shorter ones. Necklaces often have free strand endings, also in primary cilia, suggesting that they spiral. Comparing the present data with those in the literature suggests that necklace features occurring during reciliation differ from those of de novo ciliogenesis. Primary and secondary cilia share the following qualities: 1) Membrane regions above necklace strands can differ quite drastically from those below the strands.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1986

Mutant sensory cilia in the nematode Caenorhabditis elegans.

Eight classes of chemosensory neurons in C. elegans fill with fluorescein when living animals are placed in a dye solution. Fluorescein enters the neurons through their exposed sensory cilia. Mutations in 14 genes prevent dye uptake and disrupt chemosensory behaviors. Each of these genes affects the ultrastructure of the chemosensory cilia or their accessory cells. In each case, the cilia are shorter or less exposed than normal, suggesting that dye contact is the principal factor under selection. Ten genes affect many or all of the sensory cilia in the head. The daf-19 (m86) mutation eliminates all cilia, leaving only occasional centrioles in the dendrites. The cilia in che-13 (e1805), osm-1 (p808), osm-5 (p813), and osm-6 (p811) mutants have normal transition zones and severely shortened axonemes. Doublet-microtubules, attached to the membrane by Y links, assemble ectopically proximal to the cilia in these mutants. The amphid cilia in che-11 (e1810) are irregular in diameter and contain dark ground material in the middle of the axonemes. Certain mechanocilia are also affected. The amphid cilia in che-10 (e1809) apparently degenerate, leaving dendrites with bulb-shaped endings filled with dark ground material. The mechanocilia lack striated rootlets. Cilia defects have also been found in che-2, che-3, and daf-10 mutants. The osm-3 (p802) mutation specifically eliminates the distal segment of the amphid cilia. Mutations in three genes affect sensillar support cells. The che-12 (e1812) mutation eliminates matrix material normally secreted by the amphid sheath cell. The che-14 (e1960) mutation disrupts the joining of the amphid sheath and socket cells to form the receptor channel. A similar defect has been observed in daf-6 mutants. Four additional genes affect specific classes of ciliated sensory neurons. The mec-1 and mec-8 (e398) mutations disrupt the fasciculation of the amphid cilia. The cat-6 (e1861) mutation disrupts the tubular bodies of the CEP mechanocilia. A cryophilic thermotaxis mutant, ttx-1 (p767), lacks fingers on the AFD dendrite, suggesting this neuron is thermosensory.

Animals↗

PubMed1984

Observations on axonemes and membranes of olfactory and respiratory cilia in frogs and rats using tannic acid-supplemented fixation and photographic rotation.

With tannic acid-supplemented fixation and a photographic rotation technique, ultrastructural features of axonemes of frog olfactory cilia resemble those of respiratory cilia in virtually all respects. Different types of ciliary axonemes corresponding to motile and immotile olfactory cilia are not discernible. In rats, however, axonemes of olfactory cilia are quite different from those of respiratory cilia and always lack microtubule-attached structures in proximal parts. Their distal parts usually have only two microtubules which terminate in a cap-like structure. In either species, tannic acid-supplemented fixation reveals that outer leaflets of membranes of olfactory cilia are thicker than inner leaflets. This is not the case for the respiratory cilia, and the overall thickness of ciliary membranes is smaller in respiratory cilia. From our observations and literature data on vertebrates and invertebrates, it is inferred that the ultrastructure of axonemes of olfactory cilia is not evolutionarily stable. This implies that this structure does not play any specific role in the olfactory transduction process. However, the motility associated with microtubule-attached arms of those cilia which have complete axonemes may be involved in the efficacy of the olfactory process. The consistent differences between membranes of olfactory and respiratory cilia suggest that membranes of olfactory cilia may have specific properties important to the initial events of the olfactory transduction process.

Animals↗

PubMed2002

Claudin localization in cilia of the retinal pigment epithelium.

Using immunocytochemistry and confocal microscopy we demonstrate that claudin-immunoreactivity is a novel marker for retinal pigment epithelial cilia. Claudin-immunoreactivity obtained by polyclonal anti-claudin 1 antibody, which could crossreact with claudin 3, was colocalized with acetylated tubulin-immunoreactivity in cultured human retinal pigment epithelial cells. Claudin-immunoreactivity associated with the retinal pigment epithelium (RPE) cilia was more intense than was claudin-immunoreactivity in the junctional complex. Approximately two-thirds of the RPE cells in the rat contain cilia that are immunoreactive with acetylated tubulin on postnatal day 1, and a significant portion of these cilia label with the anti-claudin 1 antibody. Cilia decrease in frequency over subsequent postnatal days, and are absent by postnatal day 30. As RPE cilia decrease in number during postnatal rat development, claudin-immunoreactivity is lost earlier than acetylated tubulin, suggesting that the loss of claudin may initiate RPE cilium degeneration. Claudin-immunoreactivity was not evident in cilia of photoreceptor cells, epithelia of nasal mucosa, small intestine, or colon, suggesting that claudin may be a unique molecule in RPE cilia. These data suggest that cilia of the RPE, unlike cilia on other cell types, contain claudin, and that this molecule may play an important and specific role in the function and/or maintenance of RPE cilia.

Animals↗

PubMed1981

Microtubule sliding in cilia of the rabbit trachea and oviduct.

Evidence for active sliding of microtubules during ciliary activity has been demonstrated in a number of organisms: sea urchin sperm flagella, protozoan cilia, and mollusc gill cilia. Although there is evidence that active sliding also occurs in mammalian sperm flagella, there is little or no information on whether active sliding of microtubules also occurs in the short (5-micron) cilia of the mammalian trachea or oviduct. Since these cilia are important in tracheobronchial clearance and ovum transport, respectively, it has been important to demonstrate that microtubule sliding is also involved in the activity of somatic cilia. Ciliated apical portions (cortices) and cilia were isolated from rabbit trachea and oviduct, using Triton X-100 to demembranate the cilia. Most of the ciliated cortices reactivated upon addition of ATP, whereas isolated cilia reactivated to a lesser extent. When preparations of cilia were digested with trypsin before or after ATP addition, disintegration of axonemal doublets occurred with about the same frequency as reactivation. These events were recorded using Nomarski optics and dark-field microscopy. When isolated cilia which had been digested by trypsin and exposed to ATP were also prepared for electron microscopy by negative staining, telescoping of doublet microtubules from axonemes could be shown. These results demonstrate that mammalian somatic ciliary doublet microtubules actively slide in a manner similar to that described for invertebrate cilia.

Adenosine Triphosphate↗

PubMed1986

Abnormal cilia in a fourth ventricular ependymoma.

Although the luminal surface of the ependymal cells of the human ventricular system is characterized by the presence of cilia with a typical 9 + 2 arrangement of microtubules, cilia are usually described as sparse or absent in ependymal cells of ependymomas. The incidence of aberrant cilia and structurally abnormal cilia in normal ependymal cells and ependymomal cells has not been documented. This report describes detailed ultrastructural and quantitative studies of cilia of the ependymal cells of a fourth ventricular ependymoma in a 5-year-old boy. Of a total of 267 cilia studied, 50% were located in the extracellular microrosettes, 30% in the intracytoplasmic lumina and 20% in the cytoplasm. Thirty precent of all cilia studied showed structural abnormalities. Those present in the aberrant locations, e.g., intracytoplasmic lumina and in the cytoplasm, had a higher incidence of abnormality. Ciliary abnormalities fell into four majors categories: (1) cilia with abnormal axial microtubules, (2) compound cilia, (3) swollen cilia, and (4) cilia with dynein arm defect. Of these, addition, deletion and disorganization of axonemal microtubules were most common. The present study suggested that ciliary abnormality is common in ependymoma and may represent another example of oncogenetic effect on the ciliogenesis of ciliated epithelium.

Cerebral Ventricle Neoplasms↗

PubMed2005

Cilium-generated signaling and cilia-related disorders.

Biologists have long known that humans experience their environment through cilia. Light, odorant, and sound perception depend on these microtubule-filled, complex organelles present on cells in primary sensory tissues. Recently, discoveries on the mechanism of assembly of cilia (flagella) in the lowly, biflagellated, eucaryotic green alga Chlamydomonas have triggered a renaissance of interest in the organelles along with a recognition of their key sensory roles in nonsensory tissues. Chlamydomonas researchers uncovered an entirely new set of cellular machinery essential for transporting the protein components of cilia and flagella in all ciliated/flagellated eukaryotic cells between their site of synthesis in the cell body and their site of assembly at the tip of the flagellum (intraflagellar transport: IFT). Prompted by the surprising observations that disruption of IFT genes in mice led to polycystic kidney disease (PKD) and that PKD proteins are present on the sensory cilia of Caenorhabditis elegans, researchers have made a direct connection between PKD and cilia. At least five (and possibly all) of the seven identified human genes disrupted in PKD and a related disorder nephronophthisis encode proteins expressed in the primary cilia that project into the lumen from the epithelial cells that line renal tubules. Moreover, the renal cilia are flow sensors and at least two of the PKD genes encode ciliary transmembrane proteins essential for mechanosensation. Although their roles have not yet been as clearly identified, cilia also are at the center of a rare human disorder, Bardet-Biedl syndrome (BBS), in which patients exhibit phenotypes of common human diseases, including obesity and increased incidence of hypertension and diabetes. Five of the eight known BBS genes encode basal body or cilia proteins in mice or humans, and homologues of two of the remaining genes are present in basal bodies/cilia of model organisms. Here we briefly describe the biology of cilia and flagella, we outline how studies on model organisms have led to our current understanding of the roles of these organelles and their proteins in health and disease, and we highlight the notion that the primary cilia present on cells throughout the body, even those on brain neurons, may be essential for as yet undiscovered cilium-generated signaling functions.

Animals↗

PubMed1980

Motile statocyst cilia transmit rather than directly transduce mechanical stimuli.

We have investigated the role of motile cilia in mechanotransduction by statocysts of the nudibranch mollusk Hermissenda crassicornis. Movement of the cilia that experience the weight of statoconia causes increased variance of voltage noise and membrane depolarization of the statocyst hair cell. Two complementary approaches were used to immobilize the cilia. Vanadate anion was iontophoretically injected into hair cells. This reversible inhibitor of vibratile form and to assume a more classic, pliable beat pattern. Voltage noise decreased as the cilia slowed and bent more extremely, nearly disappearing as motility was lost. When the intracellular vanadate concentration approached 10(-5) M, the cilia were arrested in an effective stroke against the cell membrane. The cell no longer depolarized upon gravitational or local mechanical stimulation. Rapid reversal of ciliary inhibition by norepinephrine or slow reversal with time restored both the voltage noise and depolarization response. Cilia were rendered rigid and upright by covalent cross-linkage of their membrane "sleeve" to the 9 + 2 axoneme, using the photoactivated, lipophilic, bifunctional agent 4,4'-dithiobisphenyl azide. In the initial stages of cross-linkage, the cilia remained vibratile but slowed and moved through wider excursions. Voltage noise decreased in frequency but increased in amplitude. When the cilia were fully arrested, voltage noise was minimized while the resting potential and membrane resistance remained essentially constant. Mechanical stimulation of the rigid cilia, normal to the cell membrane, elicited a generator potential of the same amplitude but of greater duration than before treatment. Because cilia that are partially arrested by vanadate undergo increased bending, although the hair cell shows decreased noise, neither the axoneme nor the ciliary membrane proper would appear to be sites of direct transduction. In cells with beating but stiffened cilia, however, the voltage noise becomes amplified, implying an increased efficiency of transduction. We suggest that active but rigid flexure of the axoneme is involved in amplification and continuous signal detection. The basal insertion area is the most likely transduction site, being the terminal leverage point through which force is applied to the plasma membrane via the flexing ciliary shaft.

Animals↗

PubMed2005

Elucidating the function of primary cilia by conditional gene inactivation.

PURPOSE OF REVIEW: This review discusses recent experimental approaches to determine the function of primary cilia by conditional inactivation of genes crucial for cilia formation. RECENT FINDINGS: A functional role in the sensing of fluid flow was recently assigned to the primary cilia. This discovery shed light onto how cells sense dynamic fluid movements. Conditional inactivation of primary cilia formation in later ontogenic stages demonstrated the crucial role renal primary cilia play in the control of cell proliferation. SUMMARY: Primary cilia can act as flow sensors, transmitting signals by means of calcium influx into the cells. Structures based on primary cilia are also crucial for the function of photoreceptor cells and it can be expected that additional functions of these organelles will be determined in the future. An important experimental approach to elucidate the involvement of primary cilia in other physiological processes is to specifically inactivate genes crucial for formation of primary cilia. Morphological and physiological changes induced by the loss of primary cilia will help determine additional roles primary cilia play in physiology and organ development.

Animals↗

PubMed1983

The distribution and assessment of electron-microscopic abnormalities of human cilia.

The aim of this study was to overcome difficulties of assessing the true incidence of electromicroscopic abnormalities of microtubular structure of cilia by examining large numbers of cilia from each case. The effects of different fixatives on the appearances of cilia were also studied. Bronchial biopsies were examined from 35 subjects who were being investigated for various lung diseases and nasal biopsies from 12 subjects (7 with retinitis pigmentosa (R.P.), and 5 healthy controls). Numerous pieces of normal looking bronchial wall from a lobectomy specimen were used to examine the effect of six different fixatives. 2.9% of bronchial cilia (mean of 890 cilia examined) and 2.4% of nasal cilia (mean of 808 cilia examined) showed microtubular abnormalities. Examining large numbers of cilia established that increased microtubular abnormalities were associated with smoking, chronic pulmonary infection and carcinoma of the lung. There was a significant increase (p less than 0.001) in microtubular abnormalities in nasal cilia in R.P. The appearances of cilia varied considerably with different fixatives. The numbers of dynein arms seen and the ease of recognising radial spokes and microtubules was particularly effected by fixation. The true incidence of microtubular abnormalities can only be ascertained by examining large numbers of cilia.

Adult↗

PubMed1985

The antagonistic effects of 5-hydroxytryptamine and methylxanthine on the gill cilia of Mytilus edulis.

The laterofrontal (LF) cirri on isolated gill filaments of Mytilus edulis, prepared in natural seawater, are active and initially beat with an average frequency of about 8 Hz (with a range of 6-14 Hz). However, the lateral (L) cilia on these filaments are arrested in a position at the end of their recovery stroke. Perfusion of the filament with artificial seawater (ASW), with or without 1% ethanol, has little or no biological effect on the activity of the LF cirri, although a transitory decrease in frequency often accompanies the perfusion process. The L cilia remain arrested during perfusion with ASW. The exposure of the gill to low levels of 5-hydroxytryptamine (5HT) (10(-8) less than 5HT less than 10(-7) M) has no effect on the activity of the LF cirri but stimulates the L cilia to beat. Exposure to higher concentrations of 5HT (greater than 10(-7) M) elevates the beat frequency of the L cilia and simultaneously inhibits the activity of the LF cirri, leading to their arrest in a position at the end of the effective stroke. This arrest of the LF cirri occurs as the L cilia attain a 5HT-induced beat frequency between 12 to 14 Hz. The influence of 5HT on the L cilia and the LF cirri can be reversibly mimicked or enhanced by the phosphodiesterase inhibitor 3-isobutyl-1-methylxanthine (IBMX). A concentration of 0.5 mM IBMX mimics low 5HT concentrations (about 10(-7) M) by stimulating the L cilia to beat without affecting the beat frequency of the LF cirri. A combination of 10(-7) M 5HT and 0.5 mM IBMX in ASW mimics high (greater than 10(-6) M) 5HT concentrations by arresting the LF cirri and increasing the beat frequency of the L cilia. Under these conditions, the threshold of the LF cirri arrest response is again found to occur as the L cilia attain a beat frequency of 12-14 Hz. These results suggest that the mechanisms of LF cirri arrest and L cilia activation are mediated by 5HT-induced changes in intracellular cyclic AMP levels.

1-Methyl-3-isobutylxanthine↗

PubMed1974

Ciliary membrane differentiations in Tetrahymena pyriformis. Tetrahymena has four types of cilia.

We have examined thin sections and replicas of freeze-fractured cilia of Tetrahymena pyriformis. The ciliary necklace located at the base of all freeze-fractured oral and somatic cilia has been studied in thin sections. Since electron-dense linkers have been found to connect both microtubule doublets and triplets to the ciliary membrane at the level of the necklace, the linkers and the associated necklace seem to be related to the transition region between the doublets and triplets of a cilium. Plaque structures, consisting of small rectangular patches of particles located distal to the ciliary necklace, are found in strain GL, but are absent in other strains examined in this study. In freeze-cleaved material, additional structural differentiations are observed in the distal region of the ciliary membranes of somatic and oral cilia. Somatic cilia contain many randomly distributed particles within their membrane. Oral cilia can be divided into three categories on the basis of the morphology of their freeze-fractured membranes: (a) undifferentiated cilia with very few randomly distributed particles: (b) cilia with particles arranged in parallel longitudinal rows spaced at intervals of 810-1080 A that are located on one side of the cilium; and (c) cilia with patches of particles arranged in short rows oriented obliquely to the main axis of the cilium. The latter particles, found on one side of the cilium, seem to serve as attachment sites for bristles 375-750 A long and 100 A wide which extend into the surrounding medium. The particles with bristles are located at the tips of cilia in the outermost membranelle and may be used to detect food particles and/or to modify currents in the oral region so that food particles are propelled more efficiently into the buccal cavity. Examination of thin-sectioned material indicates that the particles in oral cilia which form the longitudinal rows could be linked to microtubule doublets. Linkage between microtubule doublets and adjacent membrane areas on one side of the cilium could modify the form of ciliary beat by restricting the sliding of the microtubules. It is suggested that membrane-microtubule interactions may form the basis for the various forms of ciliary beat observed in different organisms.

Animals↗

PubMed1985

Genesis of cilia and microvilli of rat nasal epithelia during pre-natal development. I. Olfactory epithelium, qualitative studies.

Rat foetuses from intra-uterine days E13 through E22 (day before parturition) and adults were used for a qualitative electron-microscopic investigation of the development of ciliated/microvillous surfaces of the olfactory epithelium. In the E13 and most of the E14 embryos the epithelial surface is not yet characteristically olfactory. Apical cell profiles show primary cilia. These can arise at the epithelial surface or below. From E14 onwards the epithelial surface acquires olfactory characteristics. Dendritic endings of the olfactory receptor cells can be found amidst microvillous profiles of supporting cells. Either cell type may bear primary cilia. From E16 onwards the receptor cells sprout multiple olfactory cilia, but cells with primary cilia are found throughout pre-natal development. These primary cilia are, at least for a while, retained during the formation of the secondary cilia. Primary cilia always have distinct necklaces at their base. Otherwise, especially with respect to their tips, their morphology can vary. Originally they have expanded tips (up to E14); later on such wide tips are no longer encountered (E16 and E17). Primary cilia of receptor cells never have wide tips. Appreciable numbers of endings with tapering olfactory cilia are discerned around E18 and especially E19. Throughout pre-natal development posterior/superior parts of the septal olfactory epithelium are more precocious than anterior/inferior parts, in particular in the region of transition with the respiratory epithelium. This advance in development includes total densities of dendritic endings of olfactory receptor cells, densities of multiciliated endings alone and lengths of supporting cell microvilli. This difference is discussed with respect to the topography of the olfactory epithelial surface in adult animals. In addition to the systematic topographic variation, a number of more local, apparently not-systematically distributed, topographic variations present during development are described. Most of these also occur in adult animals and they include heterogeneity in length of supporting cell microvilli and the presence of patches of supporting cells with rounded apical protuberances, of patches displaying dendrites with polyaxonemes rather than individual cilia and of scattered atypical cells (neither typical olfactory receptor nor olfactory supporting cells). At their surfaces such atypical cells can resemble inner-ear hair cells. Relative to olfactory receptor and supporting cells there are only very few atypical cells.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗