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Surface ultrastructure of the gills of the mullets Mugil curema, M. liza and M. platanus (Mugilidae, Pisces).

Mugil curema, M. liza, and M. platanus were collected from the southeastern and southern coast of Brazil. The second gill arches were analyzed by scanning electron microscopy and histology. The highest density of chloride and mucus-secreting cells was observed in the gill filaments of M. liza and M. platanus. Spines are scarce and were found only in the pharyngeal region of M. curema. The dorsal angle of curvature of the simple projections is most reduced in the rakers of M. liza and M. platanus. The raker borderline on the internal side of the arches of M. curema has grooves that do not occur in the other two species. On the external side of the branchial arches, the borders of the rakers of M. liza and M. platanus are smooth. The shape of the rakers is characteristic for each species: in M. curema, it resembles the letter "D"; in M. liza, it is trapezoidal, and in M. platanus, it is triangular. Thus there is a morphologic similarity between M. liza and M. platanus, and both differ from M. curema. All three species show elongated and extremely elaborated rakers that are placed next to each other and turned toward the opercular cavity. There are few taste buds and only several mucus-secreting cells along the whole pharyngeal region. These characteristics suggest that these species do not select food chemically but obtain it mechanically with the rakers and aggregate it with mucus.

Animals↗

Cytokeratin expression in human tongue epithelium.

The epithelium of the human tongue shows diverse morphological variations from one site to another and even within the epithelium of the same papilla. This complexity has led to confusion regarding tongue epithelium as being orthokeratinized, parakeratinized, or nonkeratinized. Cytokeratins have been shown to characterize different epithelia. The present paper describes cytokeratin expression by adult tongue epithelia and relates their distribution to morphology. Six healthy human tongue specimens were obtained after plastic surgery and cytokeratin expression was investigated immunohistochemically, using a panel of 15 antibodies for cytoskeletal proteins, and biochemically using two-dimensional gel electrophoresis. The results showed that the ventral and lateral surfaces of the tongue are related to the nonkeratinizing stratified squamous epithelia, esophageal type, whereas the dorsal surface showed mixed expression of cytokeratins. In the tip of filiform and on the surface of fungiform papillae, cytokeratins of terminal differentiation are expressed as skin type; and in the rest of the papillae as well as in interpapillary areas, the epithelium expresses esophageal type cytokeratins. Certain simple epithelial cytokeratins were found in taste buds. Cytokeratin 19 was also detected in the basal cell layer of all esophageal type epithelia in the tongue. The present results provide basis for studies on the biological events in epithelial differentiation during development and in pathology.

Cytoskeletal Proteins↗

Expression of blood group-related glycoconjugates in the junctional and other oral epithelia of rodents.

BACKGROUND: The junctional epithelium (JE) attaches the gingiva to the non-vital tooth surface and has other unusual properties which protect the underlying periodontal tissues. The JE differs from other gingival and oral epithelia in its unusual expression of cytokeratins typical of both stratifying and of simple epithelia, a phenotypic pattern possibly related to its specialized functions. METHODS: The patterns of differentiation of rodent gingival and other epithelia were examined using monoclonal antibodies against various glycoconjugates which are expressed on epithelial cell surfaces and provide an alternative marker system for regionally-differing patterns of cell maturation. RESULTS: Markers that are typical of basal cells in other stratifying epithelia were expressed by all cell strata of JE. JE lacked differentiation markers typical of other stratifying oral epithelial but showed suprabasal expression of markers typically expressed by simple epithelia and specialized epithelia, such as taste buds. CONCLUSIONS: The phenotype of rodent JE differs from that of other oral epithelia and the pattern of differentiation assessed by its expression of glycoconjugates parallels that for other phenotypic markers, such as cytokeratins. Differentiation of rodent JE is similar to that of human JE. The functional significance of these patterns of expression is not yet clear but the markers characterizing this unusual epithelium in rodents may be associated with its behavior in periodontal disease and of value to experimental studies of its development.

ABO Blood-Group System↗

Maternal folate deficiency results in selective upregulation of folate receptors and heterogeneous nuclear ribonucleoprotein-E1 associated with multiple subtle aberrations in fetal tissues.

BACKGROUND: Homocysteine, which increases in folate deficiency, can upregulate folate receptors (FR) at the translational level by increasing the interaction between a short cis-element in the 5'-untranslated region of FR-alpha mRNA and heterogeneous nuclear ribonucleoprotein-E1 (hnRNP-E1). Perturbation of this RNA-protein interaction on GD8.5 induces neural tube defects and neurocristopathies in mice. FR upregulation can also reduce cell proliferation independently of folate deficiency in some human cells. Accordingly, we tested the hypothesis that sustained murine maternal folate deficiency would negatively impact pregnancy outcomes, upregulate FR, and selectively reduce fetal cell proliferation. METHODS: Dams were fed chow with various levels of folic acid added for eight weeks before and throughout pregnancy. Following sacrifice on GD17, dams were compared for folate and homocysteine status as well as pregnancy outcomes. Fetuses from some groups were evaluated by specific biochemical, molecular, and immunohistochemical studies for FR, hnRNP-E1, and apoptosis. RESULTS: When compared to dams fed a folate-replete diet, those dams on a folate-depleted diet developed reduced red cell folates and hyperhomocysteinemia and an inverse dose-dependent upregulation of FR and hnRNP-E1 on GD17 without alterations in cell number in the majority of tissues. However, FR overexpression was accompanied by a significant reduction in the net number of cells in the midgut, lung, pons, tongue, and olfactory epithelium, and with premature differentiation in dorsal root ganglion cells and dysplasia of taste buds. By contrast, in the brain, spinal cord, diaphragm, and primordium of follicles of vibrissae, there was less FR expression, which accompanied a net reduction in number of cells and architectural anomalies. Subtle "immunohistochemical footprints" of apoptosis on GD17 fetuses corresponded with net cell loss in the lung and olfactory epithelium. Upregulation of FR could be explained by a homocysteine-induced RNA-protein interaction in folate-depleted fetuses that led to a proportionate increase in murine FR biosynthesis. CONCLUSIONS: Maternal folate deficiency results in selective upregulation of FR and hnRNP-E1 associated with multiple aberrations in fetal tissues that include increased cell loss, architectural anomalies, and premature differentiation. The potential significance of these findings to explain the wide spectrum of folate-responsive birth defects in humans is discussed.

Animals↗

Zinc deficiency and taste dysfunction; contribution of carbonic anhydrase, a zinc-metalloenzyme, to normal taste sensation.

The present study was designed to clarify the effect of zinc deficiency on sodium chloride preference, the lingual trigeminal and taste nerves transduction, and carbonic anhydrase (CA) activity of the tongue surface and salivary gland. Male SD rats, 4 weeks old, were divided into four groups, and fed zinc-deficient (Zn-Def), low-zinc (Low-Zn), and zinc-sufficient diets with free access (Zn-Suf) and pair-feeding (Pair-fed). After taking part in the preference tests for 42 days, the rats were provided for the chorda tympani and lingual trigeminal nerves recordings, then finally sacrificed and the tongue and submandibular gland excised to measure CA activity. Sodium chloride preference increased only after 4 days of the feeding of zinc-deficient and low-zinc diets, which means that the taste abnormality appears abruptly in zinc deficieny and even though in marginal zinc deficiency. Reduced CA activities of the taste-related tissues in zinc-deficient group paralleled well with the decreased taste and lingual trigeminal nerves sensitivities.

Animals↗

Localization of P2X3 receptors and coexpression with P2X2 receptors during rat embryonic neurogenesis.

It is well known that extracellular ATP mediates rapid excitatory signaling by means of the ionotropic P2X receptors. One of its subunits, the P2X(3) receptor, is well documented to be associated with sensory innervation in adult animals. It is speculated that the P2X(3) receptor may have already been present in the early sensory system. The aim of this study was to investigate the distribution of the P2X(3) receptor during neurogenesis by using immunohistochemistry on rat embryos from embryonic day (E)9.5-18.5. The P2X(3) receptor was first identified in the hindbrain neural tube and the sensory ganglia in E11-11.5 embryos. At E14.5, the optic tract and retina, nucleus tractus solitarius, mesencephalic trigeminal nucleus, and sensory nerves in both respiratory and digestive tract showed positive staining. The facial nucleus, the prepositus hypoglossal nucleus, and the sympathetic ganglia also showed P2X(3) immunoreactivity, even though these are not sensory associated. P2X(3) immunoreactivity was detected in the vestibular nucleus, the nerves in mesentery, bladder, and kidney in E16.5 and in nerves in vibrissae in E18.5. P2X(3) immunoreactivity in the facial nucleus, spinal trigeminal tract, the mesencephalic trigeminal nucleus, and the vestibular nucleus were undetectable in postnatal day 16 rat brainstem. The P2X(3) receptor was coexpressed with the P2X(2) receptor in nucleus tractus solitarius, dorsal root ganglion, nodose ganglion, and the taste bud in E16.5 embryo, which was 5 days later than the first appearance of the native P2X(3) receptor. In summary, we present a detailed expression pattern of the P2X(3) receptor during neurogenesis and report that P2X(3) immunoreactivity is down-regulated in early postnatal brainstems.

Amino Acid Sequence↗

Developmental expression pattern of monoamine oxidases in sensory organs and neural crest derivatives.

Serotonin (5-HT) has been shown to act as a morphogen in craniofacial and heart development and in the migration of neural crest derivatives. Some of these structures are capable of capturing 5-HT during development, but nothing is known about the localization of the main monoamine degradation enzymes, monoamine oxidase (MAO) A and B, in these developing tissues. We generated a highly specific antibody to MAOB; immunoreactivity is entirely abolished in brain extracts or brain sections of mice lacking MAOB. From the use of this antibody and specific riboprobes, we report that MAOB is expressed early in a variety of neural crest derivatives, in facial sensory organs, and in the heart. From E11.5 to P0, MAOB was found to be strongly expressed in the following neural crest derivatives: the aorta, cranial mesenchyme (developing bones, sensory neurons of the cranial ganglia, cartilages, thyroid, and striate muscles), dental mesenchyme, several soft palate derivatives, and boundary cap cells (E11.5-P4). Boundary cap cells contribute to the formation of nerve exit-entry points between the central and the peripheral nervous systems. Several facial sensory organs also contained MAOB mRNA, protein, and activity. High MAOB expression was noted in the olfactory placode, the dorsal part of the olfactory epithelium, the olfactory nerve layer (probably the ensheathing glia), the cochlear ganglionic cells, the taste buds, and the Merkel cells in the vibrissae follicles. Finally, we found that MAOB is massively expressed in the pharyngeal organ, heart, liver, and mast cells. In contrast, MAOA expression was restricted to the sympathetic ganglia and to the meningeal and capillary blood vessels. The pattern of MAOB expression generally matched the previously reported patterns of expression of the plasma 5-HT transporter expression or of the histamine biosynthetic enzyme L-histidine decarboxylase, suggesting a role for MAOB in fine regulation of the levels of 5-HT and histamine in the developing embryo.

Aging↗

Medullary efferent and afferent neurons of the facial nerve of the pit viper Gloydius brevicaudus.

For the purposes of comparative anatomy, we used tracer techniques and immunohistochemistry to study the facial nerve in the pit viper Gloydius brevicaudus and obtained much new data applicable to the function of this nerve in snakes and, in particular, pit vipers. We were able to identify the superior salivatory nucleus in these snakes. Preganglionic fibers from this nucleus pass along the palatine nerve and an anterior communicating branch to reach the pterygopalatine ganglion attached to the deep branch of the trigeminal maxillary nerve. The palatine nerve also contains general somatic afferents and a very few special visceral afferents from some taste buds on the palate. In the mandibular direction, preganglionic fibers from the superior salivatory nucleus join special visceral efferents from the motor nucleus in the hyomandibular nerve, from which they pass into the chorda tympani to course together for a short distance. The special visceral efferents branch off outside the cranium, and the preganglionic fibers continue on to join the trigeminal mandibular nerve to project to small ganglia within the mandible. The chorda tympani also contains general somatic afferents from the mandibular region but no special visceral afferents. This is the first time that the superior salivatory nucleus and its adjuncts have been identified in a snake. The chorda tympani of these snakes is also distinguished from the mammalian condition by lacking any special visceral afferents and by branching outside the cranium.

Animals↗

Ontogeny of the somatostatin variant [Pro2,Met13]somatostatin-14 in the brain, pituitary, and sensory organs of the frog Rana esculenta.

Two forms of somatostatin are expressed in the frog brain, i.e., somatostatin-14 (SS1) and the [Pro(2), Met(13)]somatostatin-14 variant (SS2). We have previously described the ontogeny of SS1-immunoreactive cells in the brain of Rana esculenta. In the present study, we have investigated the distribution of prepro-SS2 (PSS2)-expressing neurons in the brain of the same species during development by using antibodies directed against the N-flanking region of SS2 (PSS2(54-66)). Immunoreactive perikarya first appeared in the ventral hypothalamus at stages IV-VII. Subsequently, positive neurons were seen in the nucleus of the diagonal band of Broca, the anterior preoptic area, the posterior tuberculum (stages VIII-XII), as well as the dorsal (stages XIII-XV) and medial (stages XIX-XX) periventricular preoptic nucleus. At metamorphic climax and in newly metamorphosed frogs, positive perikarya were found in the striatum and in the interpeduncular nucleus. PSS2(54-66)-immunoreactive fibers were already widely distributed during the first stages of development, indicating that SS2 may act as a neuromodulator and/or neurotransmitter during ontogeny. The presence of PSS2(54-66)-positive nerve fibers in olfactory structures suggests that, in tadpoles, SS2 may be involved in the processing of olfactory information. The occurrence of PSS2(54-66)-like immunoreactivity in taste buds, and in the olfactory and vomeronasal organs indicates that SS2 may mediate the unconditioned and reinforcing properties of natural chemicals. Finally, the intenseexpression of PSS2(54-66)-like immunoreactivity in melanotrope cells of the pituitary suggests that SS2 may diffuse toward the pars distalis to regulate the activity of adenohypophysial cells during tadpole development.

Animals↗

Central projections and motor nuclei of the facial, glossopharyngeal, and vagus nerves in the mormyrid fish Gnathonemus petersii.

Most of the information about the anatomy of the fish's cranial nerves was collected in the first two decades of this century. Experimental analysis of the VIIth, IXth, and Xth cranial nerves by modern tract tracing techniques started about 20 years ago. Several species have been investigated to date, including one species of Agnatha (Myxinoidea), two species of elasmobranchs, and species of some orders of Teleostei like Cyprinidae, Siluriformes, Perciformes, and Gadidae. The sensory and motor nuclei of the VIIth, IXth, and Xth cranial nerves of Gnathonemus petersii were studied by anterograde and retrograde axoplasmatic transport of horseradish peroxidase and cobaltous lysine complex. The sensory nuclei form a continuous column of cells in the brain stem extending caudal to the obex. The rostral one-fourth of this column is occupied by the overlapping terminals of the VIIth and IXth nerves. The vagus nerve has 5 roots. The first 4 of these innervate the gills and the fifth supplies viscera. Afferents from the gills terminate ipsilaterally rostral to the obex in topographic order and their terminal fields overlap. Viscerosensory fibers terminate ipsilaterally in the obex region and bilaterally in the commissural nucleus of Cajal. The facial motor nucleus is located rostral to the sensory nucleus. Facial motoneurons have pear-shaped and multipolar perikarya. Their axons form a rostrally directed knee before leaving the brain. The motoneurons of the IXth and Xth nerves have a common cell column. The vagal motoneurons form a periventricular, a medial, and an intermediate cell group rostral to the obex. In the obex region and also caudal to it, a lateral and a caudal group can be distinguished. Vagal motoneurons show a topographic arrangement that is similar to that of the sensory vagal projections. The majority of motoneurons have pear-shaped perikary and ventrolaterally oriented dendrites. In the caudal nucleus the dendrites extend dorsally and overlap the terminals of sensory fibers. The axons form a dorsolaterally directed arch before joining the sensory roots. Since G. petersii uses its electrosensory system primarily for detection of food, its gustatory system is less developed than in other fishes, which possess a large number of taste buds.

Animals↗

[Psychophysics of sweet taste. II. Statistical theory of the stimulus-response behavior of sweet taste receptors].

Considering various conditions of adaptation, the authors determined the Weber discrimination thresholds for the sweet taste of saccharose. When the adaptation to the basic stimulus is maintained and use is made of the differential coefficient behaviour, the discrimination threshold is an exponential function of the concentration. The discrimination threshold is by definition that amount of stimulant per unit of volume which will produce an effect of the magnitude of 1. Consequently, the numbers of moles being substituted, its reciprocal, the probability of obtaining a stimulus response, is greater than 1. A differential equation, namely (formula: see text), can be formulated as the basis of the stimulus-response behaviour of saccharose, the integration of which yields a discrimination characteristic expressed as (formula: see text). S = concentration of the solution as the stimulus intensity; R = stimulus response expressed in steps. The parameter b is the maximum probability of the stimulus response and can be termed substance-specific concentration coefficient. Rm is the maximum step number when the stimulus intensity tends to infinity; it is also substance-specific. As this characteristic involves both the proportional factors and the differential coefficients, it is called P-D characteristic. From this invariant relationship, the static P characteristic for a very slow increase in stimulus intensity can be derived. When the adaptation to the basic stimulus is abolished by mouth rinsing between tests in paired comparisons, the dependence of the discrimination threshold on the concentration is more than exponential. The stimulus-response characteristics obtained with different time-courses of stimulation are approximated by both the Weber-Fechner and the Stevens law.

Animals↗

Worldwide haplotype diversity and coding sequence variation at human bitter taste receptor loci.

Bitter taste perception in humans is mediated by receptors encoded by 25 genes that together comprise the TAS2R (or T2R) gene family. The ability to identify the ligand(s) for each of these receptors is dependent on understanding allelic variation in TAS2R genes, which may have a significant effect on ligand recognition. To investigate the extent of coding variation among TAS2R alleles, we performed a comprehensive evaluation of sequence and haplotype variation in the human bitter taste receptor gene repertoire. We found that these genes exhibit substantial coding sequence diversity. In a worldwide population sample of 55 individuals, we found an average of 4.2 variant amino acid positions per gene. In aggregate, the 24 genes analyzed here, along with the phenylthiocarbamide (PTC) receptor gene analyzed previously, specify 151 different protein coding haplotypes. Analyses of the ratio of synonymous and nonsynonymous nucleotide substitutions using the Ka/Ks ratio revealed an excess of amino acid substitutions relative to most other genes examined to date (Ka/Ks = 0.94). In addition, comparisons with more than 1,500 other genes revealed that levels of diversity in the TAS2R genes were significantly greater than expected (pi = 0.11%; p < 0.01), as were levels of differentiation among continental populations (FST = 0.22; p < 0.05). These diversity patterns indicate that unusually high levels of allelic variation are found within TAS2R loci and that human populations differ appreciably with respect to TAS2R allele frequencies. Diversity in the TAS2R genes may be accounted for by natural selection, which may have favored alleles responsive to toxic, bitter compounds found in plants. These findings are consistent with the view that different alleles of the TAS2R genes encode receptors that recognize different ligands, and suggest that the haplotypes we have identified will be important in studies of receptor-ligand recognition.

Alleles↗

Odontode morphology and skin surface features of Andean astroblepid catfishes (Siluriformes, Astroblepidae).

Two types of odontodes, or dermal teeth, occur in the neotropical Andean astroblepid catfishes. Both odontode types conform in structure to dermal teeth of gnathostomes in having dentine surrounding a central pulp cavity covered by a superficial layer of enameloid, but differ from one another in terms of attachment and association with other epidermis features. Type I odontodes in astroblepids, also found in all representatives of the superfamily Loricarioidea, are larger (40-50 microm base diameter), generally conical and sharply pointed, occur on the fin rays, and are associated with dermal bone. Type I odontodes attach to an elevated pediment of dermal bone of the fin lepidotrich, and to dermal bone generally in loricarioids, via a ring of connective tissue. Type II odontodes of astroblepids are smaller (15-20 microm base diameter) and blunt, occur in the skin of the head, maxillary barbels, nasal flap, and lip margins, and are not associated with dermal bone. Observations based on histology and scanning electron microscopy indicate that Type II odontodes are associated with other epithelial structures to form a putative mechanosensory organ. The odontode base lies deep in the dermis. The shaft is surrounded by a dense patch of microvillous epithelium and projects from within a pit formed by an elevated ring of laminar epithelial cells bearing several columnar, knob-like putative mechanosensory structures. Type II odontode organs have thus far been observed in only three astroblepid species, Astroblepus longifilis, A. chotae, A. rosei, where they occur in especially dense arrays on the maxillary barbels, surrounded by discrete patches of microvilli and separate mechanoreceptors. Type II odontode organs are less dense elsewhere on the body, but also occur in the skin of the snout, head, and lips. Typical taste buds are absent from the barbels of these species, but present in other astroblepids. The presence of Type II odontodes and their association with specialized epithelial pit organs are unique to astroblepids among siluriforms and may be potentially important adaptations to life in torrential mountain streams.

Animals↗

Comparative morphology and cytology of the male sperm-transmission organs in viviparous species of clinid fishes (Clinidae: Teleostei, Perciformes).

This work comprises the first comparative study of the morphology and cytology of the sperm transmission organs in males of 14 species of viviparous clinid fishes (Clinidae, Blennioidei, Teleostei). The form and dimensions of these organs differ among the various species studied. The organs are composed of intra-abdominal ampullae, into which the sperm ducts and urinary bladder anchor, and an external protruding intromittent papilla used for insemination. The form of the ampullae differs among the various species, from pear-shaped to horseshoe-shaped. It increases in dimensions with increasing length of the male. In all the species this organ is covered by a connective-tissue tunic that encompasses both circular and longitudinal striated muscle bundles. The lumina of the ampullae harbor the epididymis, a strongly convoluted and plicated duct, which becomes filled with spermatozeugmata during reproduction. From here, the epididymis continues into the protruding intromittent papillae, where its folds gradually straighten at the apical part of the intromittent organ. The form and dimensions of this copulatory organ also differ in the various species. Papillae bearing taste buds are found on the apical parts of the intromittent organ, and it is probable that these, together with the difference in forms of the organ, help to prevent interspecific copulation.

Animals↗

Structure and carbohydrate histochemistry of the esophagus in ten teleostean species.

The histology and carbohydrate histochemistry of ten teleostean esophagi were compared. Structurally, the four layers of atypical vertebrate digestive tract were consistently present. The epithelium was always stratified and in all but one species (Ictalurus nebulosus) contained taste buds. Esophageal mucous cells were not the typical goblet cells seen in other vertebrates but appeared to be of six different types, pairs of which were associated with particular families. In esocids, poorly developed mucous acini and serous monogranular cells were present. In all species, the subepithelial connective tissue was not divided into definitive lamina propriae submucosae due to the absence of muscularis mucosae. Variably present in this connective tissue region were argentophilic fibers and in esocids only, randomly dispersed striated muscle fibers. The arrangement of the muscularis was reverse to that of the general vertebrate plan. In mucous cells, three general types of epithelial mucosubstances were identified and in broad terms were recognized as sulfomucins, sialomucins and neutral mucosubstances. Morphological differences were accompained by differences in carbohydrate localization, each esophageal epithelium containing at least two different mucosubstances. However, the mucosubstances identified in each mucous cell had a profile of characteristics different in some respects from any other. Thus teleostean esophagi appear to perform an integrated diversity of functions as reflected by their complex morphology and carbohydrate histochemistry.

Animals↗

Papillary projections at the lingual margin in the striped dolphin, Stenella coeruleoalba, with special reference to their development and regression.

Papillary projections along the anterolateral margin of the tongue were observed in fetal and young stages of the striped dolphin, Stenella coeruleoalba. Papillary projections appear in the prenatal period and attain maximum development in the early postnatal period. They almost disappear by weaning. Vestigial eminences remain at the corresponding region, but they completely disappear in the adult. The papillary projections observed differ markedly from the lingual papillae of the general mammalian tongue as they are temporary, localized at the anterolateral margin and large in size. The projections were also present in young individuals of some other dolphins. No taste buds could be seen on the projections in any of the stages of all specimens observed. Such projections may have important mechanical functions during suckling in these mammals.

Animals↗

Ultrastructure of a complex epithelial system: the pharyngeal lining of the larval lamprey Petromyzon marinus.

Electron microscopy shows that the pharyngeal lining of the larval lamprey Petromyzon marinus is a structurally complex epithelial system that can be separated into eight epithelial types: gill lamellar, gill interlamellar, goblet cell, protective, terminal (taste) bud, preciliated, ciliated in tracts, and ciliated in grooves. Furthermore, these epithelial types encompass at least sixteen different cell types based on ultrastructure and, in some cases, correlative histochemistry (PAS, Alcian blue). Common to nearly all the epithelial types are basal cells and intermediate cells. These two cell types are seen as undifferentiated. Among mature cells, structural specialization as proceeded in three directions: 1) elaboration of mitochondria, probably related to molecular transport (ion-uptake cells, chloride cells); 2) ciliogenesis (preciliated and ciliated cell types); and 3) production of mucous secretory granules (mucous-platelet cells, goblet cells, superficial protective cells, columnar mucous cells, "cobblestone" cells, and marginal and dark cells in the terminal buds). Many of the functions of the cell types relate to the process of suspension feeding in this animal.

Animals↗