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Epithelial overexpression of BDNF and NT4 produces distinct gustatory axon morphologies that disrupt initial targeting.

Most fungiform taste buds fail to become innervated when BDNF or NT4 is overexpressed in the basal layer of tongue epithelium. Here, we examined when and how overexpression of BDNF and NT4 disrupt innervation to fungiform papillae. Overexpression of either factor disrupted chorda tympani innervation patterns either before or during the initial innervation of fungiform papillae. NT4 and BDNF overexpression each disrupted initial innervation by producing different gustatory axon morphologies that emerge at distinct times (E12.5 and E14.5, respectively). Chorda tympani nerve branching was reduced in NT4 overexpressing mice, and neuronal fibers in these mice were fasciculated and remained below the epithelial surface, as if repelled by NT4 overexpression. In contrast, many chorda tympani nerve branches were observed near the epithelial surface in mice overexpressing BDNF, and most were attracted to and invaded non-taste filiform papillae instead of gustatory papillae. These results suggest that BDNF, but not NT4, normally functions as a chemoattractant that allows chorda tympani fibers to distinguish their fungiform papillae targets from non-gustatory epithelium. Since BDNF and NT4 both signal through the p75 and TrkB receptors, trophin-specific activation of different internal signaling pathways must regulate the development of the distinct gustatory axon morphologies in neurotrophin-overexpressing mice.

Animals↗

Perireceptor events in taste.

The microenvironment at chemical receptor sites is important for ligand-receptor interaction as it can influence the entry, residence time or exit of odorant and sapid molecules. The perireceptor milieu at apical taste cell microvilli consists of taste pore mucus and secretions from salivary glands. The majority of taste buds are sheltered in epithelial folds of the foliate and circumvallate papillae where saliva is provided predominantly by the lingual von Ebner's glands (VEGs). To investigate possible saliva-tastant interactions, we have characterized a prominent 18 kDa secretory protein expressed in human, rat and pig VEGs. The human and rat VEG proteins share 60% sequence identity and, by virtue of their protein and gene structure, can be assigned to the lipocalin superfamily of lipophilic ligand carrier proteins. VEG proteins might function as transporters of hydrophobic molecules, for example bitter substances, like the nasal odorant-binding proteins that belong to the same protein family. Because binding experiments using various bitter substances have so far failed, and in light of the species-specific expression, other functions for VEG proteins must be considered. These include the protection of taste epithelia, pheromone transport and lipid binding.

Amino Acid Sequence↗

Role of taste in the microstructure of quinine ingestion by rats.

The microstructure of the licking behavior of water-deprived rats presented with either water or quinine during 45-min single-bottle tests was analyzed. The chorda tympani (CT) and glossopharyngeal (GL) nerves, which innervate the taste buds of the tongue, were transected in deeply anesthetized rats to discern their contribution to the behavioral pattern of quinine drinking. Rats were presurgically habituated to the testing protocol and postsurgically tested first with water and then novel 0.2 mM quinine-HCl in a subsequent session. The substantial decrease in intake observed in sham-operated controls (n = 16) when quinine was the stimulus was entirely a function of a decrease in lick volume and burst size (a run of licks with interlick intervals <1 s). Contrary to the intake-suppressing effects of quinine, pause duration decreased and burst number increased. Combined transection of the CT and GL (n = 6) strikingly opposed all of these quinine-induced behavioral changes, whereas CT transection (n = 7) was without effect and GL transection (n = 8) had an intermediate influence. These results suggest that taste acts more on neural circuits governing burst termination as opposed to burst initiation, which, in turn, appears to be more sensitive to signals related to physiological state. These findings are discussed in terms of other known nerve transection effects on quinine responsiveness, and the implications of the microstructural results are considered with respect to probabilistic as opposed to deterministic control of licking behavior.

Animals↗

Chorda tympani responses under lingual voltage clamp: implications for NH4 salt taste transduction.

Rat chorda tympani (CT) responses to NH4Cl, ammonium acetate (NH4Ac), and ammonium hippurate (NH4Hp) were obtained during simultaneous current and voltage clamping of the lingual field potential. Although functional and developmental similarities for gustation have been reported for NH4+ and K+ salts, we report here that significant differences are discernible in the CT responses to both salts. Unlike neural responses to KCl, those to NH4Cl are voltage sensitive, enhanced by submucosa negative and suppressed by positive voltage clamp. In this regard, NH4Cl responses are qualitatively similar to NaCl responses; however, the magnitude of NH4Cl voltage sensitivity is significantly less than that of NaCl. The concentration dependence of the CT response to NH4Cl manifests a biphasic nonlinear relationship not observed with KCl or NaCl. Below 0.3 M, the CT response increases as if to approach a saturation value. However, beyond 0.3 M an inflection appears in the CT-concentration curve because of an abrupt increase in CT responses. This kinetic profile is Cl-dependent and is correlated with an increase in transepithelial conductance that displays similar NH4Cl concentration dependence. The biphasic relation to salt concentration is not observed when acetate or hippurate is substituted for Cl-. As with Na+ and K+ salts, less mobile anions than Cl- (Ac- and Hp-) lower the CT responses. However, like Na+ salts, but in contrast to K+ salts, the onset kinetics of CT responses to NH4Ac or NH4Hp remained rapid, even under positive voltage-clamp conditions. Amiloride (100 microM) partially suppresses CT responses within the concentration range of 0.05-0.3 M (48-20% suppression). Amiloride also suppresses the voltage sensitivity of NH4Cl CT responses, but does not eliminate the sensitivity as it does for Na+ salts. In conclusion, the data suggest that taste transduction for NH4 salts is mediated over two NH+ conduction pathways in the taste bud. This is especially evident with NH4Cl, where the CT-concentration curves show two distinct kinetic regimes. Below 0.3 M the saturation with increasing concentration, clamp voltage response dependence, and amiloride sensitivity suggest an apical membrane transduction conductance. Above 0.3 M, the high anion dependence of the response and its amiloride insensitivity indicate participation of the paracellular pathway in transduction.

Amiloride↗

Transganglionic transport of HRP from the circumvallate papilla of the rat.

To learn whether horseradish peroxidase (HRP) injections in gustatory papillae on the tongue can be used to study central topographical projections of taste buds and papillae, injections were made into the circumvallate papilla in rats. Labeled central projections after papilla injections were compared to projections after applying HRP to the cut glossopharyngeal nerve. Papilla injections result in HRP transport by afferent and efferent fibers of the glossopharyngeal nerve, and the pattern of central projections is similar to that after labeling the cut nerve. Projections include a separation in the brainstem of afferent, dorsally located fibers and efferent, ventrally located fibers. Afferent fibers project to the solitary nucleus and the trigeminal system. Efferent projections label muscle motorneurons in the nucleus ambiguus and the cells of origin of parasympathetic preganglionic fibers, which from the inferior salivatory nucleus. The parasympathetic neurons labeled after papilla projections are preganglionic fibers to Remak's ganglia in the tongue; post-ganglionic fibers of these ganglia are the secretomotor supply to the von Ebner's glands. In summary, injections of HRP into gustatory papillae reliably label central projections of the papilla and can be used for studies to discern topography in central projections of the taste system. Injections into the circumvallate papilla also have demonstrated that the parasympathetic neurons innervating von Ebner's glands are located in the inferior salivatory nucleus.

Animals↗

The relative effects of transection of the gustatory branches of the seventh and ninth cranial nerves on NaCl taste detection in rats.

Chorda tympani nerve (CT) transection in rats severely impairs NaCl taste detection. These rats can detect higher concentrations of NaCl, however, suggesting that remaining oral nerves maintain some salt sensibility. Rats were tested in a gustometer with a 2-response operant taste-detection task before and after sham surgery (n = 5), combined transection of the CT and the greater superficial petrosal nerves (GSP; 7x, n = 6), or transection of the glossopharyngeal nerve (GL; 9x, n = 4). Thresholds did not significantly change after sham surgery. Although the GL responds to NaCl and innervates nearly 60% of total taste buds, 9x surgery had no effect. However, 7x surgery increased NaCl detection threshold by approximately 2.5 log(10) units, greater than that reported for CT transection alone. These results suggest that the GSP contributes to NaCl sensitivity in rats and also demonstrate that the GL and perhaps the superior laryngeal and lingual nerve proper can maintain some NaCl detectability at high concentrations. These findings confirm the primacy of the 7th nerve relative to the 9th nerve in sensibility of NaCl in the rat model.

Analysis of Variance↗

Evidence of solitary chemosensory cells in a large mammal: the diffuse chemosensory system in Bos taurus airways.

The diffuse chemosensory system (DCS) of the respiratory apparatus is composed of solitary chemosensory cells (SCCs) that resemble taste cells but are not organized in end organs. The discovery of the DCS may open up new approaches to respiratory diseases. However, available data on mammalian SCCs have so far been collected from rodents, the airways of which display some differences from those of large mammals. Here we investigated the presence of the DCS and of SCCs in cows and bulls (Bos taurus), in which the airway cytology is similar to that in humans, focusing our attention on detection in the airways of molecules involved in the transduction cascade of taste [i.e. alpha-gustducin and phospholipase C of the beta2 subtype (PLCbeta2)]. The aim of the research was to extend our understanding of airway chemoreceptors and to compare the organization of the DCS in a large mammal with that in rodents. Using immunocytochemistry for alpha-gustducin, the taste buds of the tongue and arytenoid were visualized. In the trachea and bronchi, alpha-gustducin-immunoreactive SCCs were frequently found. Using immunocytochemistry for PLCbeta2, the staining pattern was generally similar to those seen for alpha-gustducin. Immunoblotting confirmed the expression of alpha-gustducin in the tongue and in all the airway regions tested. The study demonstrated the presence of SCCs in cows and bulls, suggesting that DCSs are present in many mammalian species. The description of areas with a high density of SCCs in bovine bronchi seems to indicate that the view of the DCS as made up of isolated cells totally devoid of ancillary elements is probably an oversimplification.

Animals↗

Chemoreception in crocodilians: anatomy, natural history, and empirical results.

Anatomical and behavioral aspects of chemoreception in crocodilians are reviewed. Extant crocodilians possess valvular nostrils regulated by smooth muscles; paired nasal cavities, each with three conchae; and flaps closing the rear of the buccopharyngeal passage to channel inspired air through the nasopharyngeal duct. Olfaction is enhanced by gular pumping, whereby the floor of the pharynx is alternately lowered and raised to pulse air through the nasal cavity. Taste buds occur on the tongue, palate, and pharyngeal walls. Crocodilians in nature or in semi-natural enclosures locate distant carrion or concealed meat, implying chemoattraction to food. The American alligator exhibits underwater head-waving and mouth-opening in response to aqueous and chloroform extracts of meat. Alligators increase gular pumping in response to the scents of various meats, indicating that they detect food chemicals by olfaction. Paired gular and paracloacal glands on the integument of crocodilians are thought to produce pheromones. Juvenile crocodilians exhibit heightened gular pumping in response to airborne skin gland secretions. Observations of adults suggest that gular and paracloacal glands produce chemical signals used in mating and/or nesting activities, but behavioral responses to skin gland exudates are too poorly documented to ascribe pheromonal properties.

Alligators and Crocodiles↗

Physiological phenotyping of cholecystokinin-responsive rat taste receptor cells.

The recent discovery that subsets of rat taste receptor cells (TRCs) express the peptide cholecystokinin (CCK) and that subsets of TRCs respond to CCK with altered potassium currents or elevated intracellular calcium via CCK-A receptor has lead to the hypothesis that CCK may play a novel signaling role within the taste bud, perhaps modifying tastant responses by co-transmission with a classic transmitter. To better understand this phenomenon, CCK-responsive TRCs were characterized for sensitivity to two bitter stimuli, quinine or caffeine, or to the neurotransmitter ACh using a ratiometric procedure with the calcium sensitive dye fura-2. In characterizing TRC responses to quinine, it was observed that quinine-induced elevations of intracellular calcium were not due to endogenous fluorescence of the quinine molecule. Most (60-70%) CCK-responsive cells were also sensitive to either bitter stimuli or to cholinergic stimulation. These data suggest that TRCs expressing CCK-receptors also express receptors to bitter stimuli and/or muscarinic receptors. They further support the notion of a putative modulatory role of CCK with convergence of multiple inputs occurring at the level of intracellular calcium.

Animals↗

G beta association and effector interaction selectivities of the divergent G gamma subunit G gamma(13).

G gamma(13) is a divergent member of the G gamma subunit family considered to be a component of the gustducin G-protein heterotrimer involved in bitter and sweet taste reception in taste bud cells. G gamma(13) contains a C-terminal asparagine-proline-tryptophan (NPW) tripeptide, a hallmark of RGS protein G gamma-like (GGL) domains which dimerize exclusively with G beta(5) subunits. In this study, we investigated the functional range of G gamma(13) assembly with G beta subunits using multiple assays of G beta association and G beta gamma effector modulation. G gamma(13) was observed to associate with all five G beta subunits (G beta(1-5)) upon co-translation in vitro, as well as function with all five G beta subunits in the modulation of Kir3.1/3.4 (GIRK1/4) potassium and N-type (alpha(1B)) calcium channels. Multiple G beta/G gamma(13) pairings were also functional in cellular assays of phospholipase C (PLC) beta 2 activation and inhibition of G alpha(q)-stimulated PLC beta 1 activity. However, upon cellular co-expression of G gamma(13) with different G beta subunits, only G beta(1)/G gamma(13), G beta(3)/G gamma(13), and G beta(4)/G gamma(13) pairings were found to form stable dimers detectable by co-immunoprecipitation under high-detergent cell lysis conditions. Collectively, these data indicate that G gamma(13) forms functional G beta gamma dimers with a range of G beta subunits. Coupled with our detection of G gamma(13) mRNA in mouse and human brain and retina, these results imply that this divergent G gamma subunit can act in signal transduction pathways other than that dedicated to taste reception in sensory lingual tissue.

Amino Acid Sequence↗

Fat perception is related to PROP taster status.

Individuals who are sensitive to the bitter compounds phenylthiocarbamide (PTC) and 6-n-propylthiouracil (PROP) are also more sensitive to selected bitter and sweet substances, to sharp-tasting foods, and to the trigeminal irritant capsaicin. PTC/PROP tasters have a greater density of fungiform taste papillae and it is speculated that PTC/PROP tasters also have more trigeminal innervation. Because oral texture perception is also mediated, in part, by trigeminal fibers, it has been proposed that individual differences in fat perception might also be linked to PTC/PROP taster status and taste bud density. This work tests the hypothesis that individuals who are PROP tasters: 1. have a higher density of fungiform papillae; 2. are more sensitive to capsaicin; and 3. have increased ability to discriminate differences in fat content in salad dressing. Individual subjects were classified as PROP nontasters, medium tasters, or supertasters (n = 25 per group) by comparing their psychophysical function for PROP to that of NaCl. Papillae densities (papillae/cm2) were significantly different among the 3 taster groups (p < or = 0.0001), and were highest among the supertasters. Both medium tasters and supertasters perceived more oral burn from capsaicin than did nontasters at concentrations of 50, 70, and 100 ppm (p < or = 0.0001). Medium tasters and supertasters could also discriminate differences in fat content between 40% fat and 10% fat salad dressings (p < or = 0.005), but the nontasters could not. These data provide the first published evidence that fat perception can be linked to genetic and anatomical differences between individuals.

Adolescent↗

Receptor cell regeneration and connectivity in olfaction and taste.

The capacity of adult mammalian gustatory and olfactory receptor cells to regenerate and make synaptic reconnections provides examples that may be useful in initiating replacement of other kinds of sensory receptor cells. The sensory code for taste quality may not be degraded by taste receptor cell turnover because axons probably recouple to the appropriate type of new receptor cell by axon-receptor cell affinity. Experiments on the development and regeneration of taste receptor cells suggest that they regenerate and turn over by recapitulating the late but not the early steps in taste bud development. To evaluate the replacement of vertebrate olfactory receptors, we began by characterizing the spatial pattern of primary olfactory projections in rainbow trout. Contiguous clusters of HRP-labeled olfactory receptor neurons (ORN) make highly divergent projections to the olfactory bulb. Retrograde transport of fluorescent latex beads revealed that a given restricted site in the glomerular layer received axons from ORNs widely scattered in the epithelium. Hence, ORN axons do not form point-to-point or regional topographic maps. Rather, the olfactory epithelial sheet makes a plane-to-point or holographic-like projection, since any given point in the glomerular layer receives information from the entire olfactory epithelial plane. Receptor cells that reacted with the lectin pokeweed agglutinin were highly dispersed in the olfactory epithelium with axons widely scattered in the olfactory nerve. Yet, as a consequence of the extensive reaggregation of axons at the nerve-bulb interface, the lectin-positive axons fasciculated and converged into a subregion of the glomerular layer.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Detection of NaCl and KCl in TRPV1 knockout mice.

Both amiloride-sensitive and -insensitive mechanisms contribute to NaCl taste transduction. The amiloride-sensitive mechanism relies on the epithelial Na(+) channel ENaC, which is widely expressed on the apical membrane of fungiform taste cells. The amiloride-insensitive mechanism, which predominates in circumvallate and foliate taste buds, was recently reported to involve a variant of the nonselective cation channel TRPV1. We performed 2-bottle preference and threshold experiments with TRPV1 knockout mice and wild-type (C57BL/6J) controls to test for NaCl preference and detection thresholds in the presence and absence of amiloride. Surprisingly, TRPV1 knockout mice not only detected NaCl in the presence of amiloride but they preferred NaCl over water at concentrations avoided by the wild-type mice. NaCl detection thresholds were between 2 and 3 mM for both genotypes. Amiloride increased the detection thresholds of wild-type mice but not knockout mice. The knockout mice also preferred 100 mM KCl compared with wild-type controls, suggesting that TRPV1 receptors may mediate a general aversive response to salts. Analyses of consumption data also revealed that TRPV1 knockout mice ingested more of the NaCl, with and without amiloride, and KCl solutions than the wild-type mice. However, comparisons of preference ratios and consumption volumes indicated that both wild-type and TRPV1 knockout mice avoided citric acid in quite a similar manner, suggesting that TRPV1 receptors do not mediate the detection of citric acid. These data, taken together, suggest that additional mechanisms must contribute to the amiloride-insensitive NaCl response.

Amiloride↗

Kainate-activated cobalt uptake in the primary gustatory nucleus in goldfish: visualization of the morphology and distribution of cells expressing AMPA/kainate receptors in the vagal lobe.

Gustatory afferent fibers of the vagus nerve that innervate taste buds of the oropharynx of the goldfish, Carassius auratus, project to the vagal lobe, which is a laminated gustatory nucleus in the dorsal medulla. As in the mammalian gustatory system, responses by second-order cells in the goldfish medulla are mediated by N-methyl-D-aspartate (NMDA) and non-NMDA ionotropic glutamate receptors. We utilized a cobalt uptake technique to label vagal lobe neurons that possess cobalt-permeable ionotropic glutamate receptors. Vagal lobe slices were bathed in kainate (40 microM) or glutamate (0.5 or 1 mM) in the presence of CoCl(2), which can pass into cells through the ligand-gated cation channels of non-NMDA receptors made up of certain subunit combinations. Cobalt-filled cells and dendrites were observed in slices that were activated by kainate or glutamate, but not in control slices that were bathed in CoCl(2) alone, nor in slices that were bathed with the non-NMDA receptor antagonist 6,7-dinitroquinoxaline-2,3-dione (10 microM) in addition to an agonist. Likewise, simple depolarization of the cells with KCl failed to induce cobalt loading. Cobalt-filled round unipolar cells, elongate or globular bipolar cells, and multipolar cells with elongate or polygonal perikarya were distributed throughout the cell layers in the sensory zone of the vagal lobe. Numerous labeled neurons had dendrites spanning layers IV and VI, the two principal layers of primary afferent input. Apical and basal dendrites often extended radially through neighboring laminae, but many cells also extended dendrites tangential to the lamination of the sensory zone. In the motor layer, cell bodies and proximal dendrites of small, multipolar neurons, and large motoneurons were regularly loaded with cobalt.

Animals↗

Afferent signals regulating food intake.

Food intake is a regulated system. Afferent signals provide information to the central nervous system, which is the centre for the control of satiety or food seeking. Such signals can begin even before food is ingested through visual, auditory and olfactory stimuli. One of the recent interesting findings is the demonstration that there are selective fatty acid taste receptors on the tongue of rodents. The suppression of food intake by essential fatty acids infused into the stomach and the suppression of electrical signals in taste buds reflect activation of a K rectifier channel (K 1.5). In animals that become fat eating a high-fat diet the suppression of this current by linoleic acid is less than that in animals that are resistant to obesity induced by dietary fat. Inhibition of fatty acid oxidation with either mercaptoacetate (which blocks acetyl-CoA dehydrogenase) or methylpalmoxirate will increase food intake. When animals have a choice of food, mercaptoacetate stimulates the intake of protein and carbohydrate, but not fat. Afferent gut signals also signal satiety. The first of these gut signals to be identified was cholecystokinin (CCK). When CCK acts on CCK-A receptors in the gastrointestinal tract, food intake is suppressed. These signals are transmitted by the vagus nerve to the nucleus tractus solitarius and thence to higher centres including the lateral parabrachial nucleus, amygdala, and other sites. Rats that lack the CCK-A receptor become obese, but transgenic mice lacking CCK-A receptors do not become obese. CCK inhibits food intake in human subjects. Enterostatin, the pentapeptide produced when pancreatic colipase is cleaved in the gut, has been shown to reduce food intake. This peptide differs in its action from CCK by selectively reducing fat intake. Enterostatin reduces hunger ratings in human subjects. Bombesin and its human analogue, gastrin inhibitory peptide (also gastrin-insulin peptide), reduce food intake in obese and lean subjects. Animals lacking bombesin-3 receptor become obese, suggesting that this peptide may also be important. Circulating glucose concentrations show a dip before the onset of most meals in human subjects and rodents. When the glucose dip is prevented, the next meal is delayed. The dip in glucose is preceded by a rise in insulin, and stimulating insulin release will decrease circulating glucose and lead to food intake. Pyruvate and lactate inhibit food intake differently in animals that become obese compared with lean animals. Leptin released from fat cells is an important peripheral signal from fat stores which modulates food intake. Leptin deficiency or leptin receptor defects produce massive obesity. This peptide signals a variety of central mechanisms by acting on receptors in the arcuate nucleus and hypothalamus. Pancreatic hormones including glucagon, amylin and pancreatic polypeptide reduce food intake. Four pituitary peptides also modify food intake. Vasopressin decreases feeding. In contrast, injections of desacetyl melanocyte-stimulating hormone, growth hormone and prolactin are associated with increased food intake. Finally, there are a group of miscellaneous peptides that modulate feeding. beta-Casomorphin, a heptapeptide produced during the hydrolysis of casein, stimulates food intake in experimental animals. In contrast, the other peptides in this group, including calcitonin, apolipoprotein A-IV, the cyclized form of histidyl-proline, several cytokines and thyrotropin-releasing hormone, all decrease food intake. Many of these peptides act on gastrointestinal or hepatic receptors that relay messages to the brain via the afferent vagus nerve. As a group they provide a number of leads for potential drug development.

Adipose Tissue↗

Secondary connections of the dorsal and ventral facial lobes in a teleost fish, the rockling (Ciliata mustela).

In the rockling, Ciliata mustela (Teleostei), a portion of the dorsal fin is a specialized chemosensory organ possessing solitary chemoreceptor cells innervated by a recurrent branch of the facial nerve. Previous studies have demonstrated that the specialized solitary chemoreceptor cell system is represented in the dorsal segment of the medullary facial lobe (DFL), whereas the taste buds in the remainder of the facial-nerve-innervated skin are represented in the ventral division of the lobe (VFL). The carbocyanine dye DiI was used to investigate the secondary and higher order brain connections of these two distinct subdivisions of the facial lobe. Both segments of the facial lobe sent fibers into the contralateral DFL via a dorsocaudal facial commissure and to the contralateral vagal lobes and VFL via fibers arching ventrally through the reticular formation. Ascending fibers from both facial lobe segments were traced into the secondary gustatory nucleus and into the lateral superficial facial nucleus, a small area in the dorsolateral brainstem laterally adjacent to the nucleus medialis of the octavolateral complex. Additionally, the VFL had reciprocal connections with a newly described nucleus adjacent to the incoming facial nerve root. Both DFL and VFL had descending fibers reaching two portions of the funicular nuclear complex, although the VFL contribution to this area is far more extensive than the DFL input. Thus, substantial overlap exists in the connections of the two facial subsystems; i.e., the solitary chemoreceptor information is not processed in nuclei distinct from those making up the usual gustatory lemniscus.

Animals↗

Ultrastructure of Weber's salivary glands of the root of the tongue in the rat.

BACKGROUND: The mammalian tongue encompasses several sizeable agglomerations of minor salivary glands. The ultrastructure of the various glands in the body of that organ has already been determined. In contrast, almost nothing is known of the structure of Weber's glands, a collection of salivary glands in the root of the tongue. METHODS: The entire tongue was extirpated from anesthetized adult male rats that had been perfused with fixative and specimens of tissue that included Weber's glands were dissected from the lingual root. These were prepared for electron microscopic examination by conventional means. RESULTS: Weber's glands in the rat are mixed glands, consisting of long mucous tubules that often are capped by serous demilunes. There appear to be no ducts per se, the mucous tubules increasing in caliber and approaching the crypts of the dorsal lingual surface while still retaining their mucous secretory character. As these ducts near the surface, their lining changes to stratified squamous epithelium. The mucous cells are cytologically similar to those in the sublingual glands of the same animal. Mucous droplets undergo vertical fusion to form centrally situated intracellular channels through which mucus is exocytosed, much in the same manner as goblet cells. The serous cells, which have all of the hallmarks of protein-secreting cells, probably empty their secretory granules via cellular extensions that directly reach the tubule lumen since there are no intercellular canaliculi between adjacent mucous cells. CONCLUSIONS: The mucus elaborated by Weber's glands undoubtedly aids in swallowing dry food. We postulate that the serous cells in these glands, as in the more anterior von Ebner's glands, might play a role in the mechanism of taste, especially where posteriorly situated, nonlingual taste buds are concerned.

Animals↗

Tongue flicking in agamid lizards: morphology, kinematics, and muscle activity patterns.

We wanted to examine whether a relation between foraging strategy, morphology, the mechanics of tongue protrusion, and prey chemical detection and discrimination exists in agamid lizards. Tongue-flick behavior was observed in two species of this family: Uromastix acanthinurus and Plocederma stellio. Potential prey chemical discrimination by means of tongue flicking was examined by using applicator tests. Tongue flicks were subsequently recorded by high-speed video in combination with the electrical activity of a number of jaw and hyolingual muscles. The kinematics of jaws and tongue and the muscle activity patterns were quantified. To investigate if the observed differences in tongue-flick behavior (mainly in the frequency of use) are translated into corresponding differences in tongue morphology, the tongues of both species were examined by light and scanning electron microscopy. The species differed mainly in the surface morphology of the foretongue and in the abundance and distribution of taste buds on the tongue and oral cavity. These differences can be related to behavioural observations; whereas U. acanthinurus readily uses tongue flicks to detect and discriminate between food items, P. stellio does not. However, differences in tongue-flick mechanics (kinematics, electromyograms) between both species were minor. Based on the data gathered in this study and from previously published data, an evolutionary transformation series leading to the complex tongue-flick cycles as observed in snakes is proposed. The required morphological and mechanical changes that accompany such an evolutionary sequence are discussed.

Animals↗