Electromyographic analysis of the superior belly of the omohyoid muscle and anterior belly of the digastric muscle in tongue and head movements.

The participation of the superior belly of the omohyoid muscle and anterior belly of the digastric muscle in tongue and head movements was studied eletromyographically in 20 normal young volunteers. A pair of monopolar electrodes was used in each muscle for simultaneous recording of their actions. The muscles act in the following tongue movements: protrusion, right and left lateral movements, placement of the tip of the tongue on soft and hard palates and on the floor of the mouth. The strongest levels of activity of the superior belly of the omohyoid muscle were observed in the placement of the tip of the tongue on the soft palate, coincidentally with a greater dislocation of hyoid bone. Both of the muscles studied did not participate in the head's kinesiology.     

The time course of taste bud regeneration after glossopharyngeal or greater superficial petrosal nerve transection in rats

We previously have published data detailing the time course of taste bud regeneration in the anterior tongue following transection of the chorda tympani (CT) nerve in the rat. This study extends the prior work by determining the time course of taste bud regeneration in the vallate papilla, soft palate and nasoincisor ducts (NID) following transection of either the glossopharyngeal (GL) or greater superficial petrosal (GSP) nerve. Following GL transection in rats (n = 6 per time point), taste buds reappeared in the vallate papilla between 15 and 28 days after surgery, and returned to 80.3% of control levels (n = 12) of taste buds by 70 days postsurgery. The first appearance and the final percentage of the normal complement of regenerated vallate taste buds after GL transection resembled that seen previously in the anterior tongue after CT transection. However, in the latter case, regenerated taste buds reached asymptotic levels by 42 days after surgery, whereas within the time frame of the present study, a clear asymptotic return of vallate taste buds was not observed. In contrast to the posterior (and anterior) tongue, only 25% of the normal complement of palatal taste buds regenerated by 112 days and 224 days after GSP transection (n = 9). The difference in regenerative capacity might relate to the surgical approach used to transect the GSP. These experiments provide useful parametric data for investigators studying the functional consequences of gustatory nerve transection and regeneration.     

REGIONAL VARIATION IN SWEET SUPPRESSION

Differences in the sweet‐blocking efficacy of 2‐(4‐methoxyphenoxy) propanoic acid (PMP) for different sweeteners (sucrose and aspartame) and for various exposure areas of the mouth were found. Twenty participants rated sweetener solutions with and without PMP for sweetness, sourness, saltiness, bitterness and umami for stimulation of anterior tongue, posterior tongue and whole‐mouth areas. For sweetness ratings, suppression was significant for all stimulation areas. In the presence of PMP, stimulation of the posterior tongue yielded significantly higher sweetness ratings than stimulation of the anterior tongue for aspartame but not for sucrose. Sourness and bitterness ratings were significantly higher for anterior tongue than posterior tongue stimulations for aspartame but not for sucrose. The increases in sourness ratings in the presence of PMP were likely because of the sour taste PMP has at the concentration used. Results imply a difference between the front and the back of the tongue in the mechanisms involved in the perception of sweetness.     

Tongue evolution in lungless salamanders, family plethodontidae. III. Patterns of peripheral innervation

Innervation of the tongue and associated musculature in plethodontid salamanders was studied using Palmgren stained sectioned materials, fresh dissection, and whole mounts of experimental specimens treated with horseradish peroxidase (HRP). Species studied were chosen to represent modes of tongue projection recognized by Lombard and Wake ('77). Special attention was given to species of the genera Plethodon, Batrachoseps, Pseudoeurycea, and Hydromantes, but representatives of other genera were investigated. As expected we found that cranial nerves IX and X and spinal nerve 1 supplied the muscles involved in tongue movement. The peripheral courses of the nerves were traced, and both functionally related and phylogenetically determined routes were found. As relative projection length increases, the nerves supplying the tongue tip also increase in length. When the tongue is at rest the long nerves are stored in coils. The coil of ramus lingualis lies between the ceratobranchials, but that of ramus hypoglossus is more variable, although constant within a species. Ramus hypoglossus bifurcates into separate branches to tongue and anterior musculature of the floor of the mouth. In generalized, presumably primitive, modes the bifurcation and coiling are far anterior. In most of the tongue projection modes bifurcation is relatively posterior, but in one, bifurcation is anterior, but coiling is relatively posterior in position. The most unusual condition is in Hydromantes, in which bifurcation is relatively posterior and a coiled ramus hypoglossus joins a coiled ramus lingualis to form a unique, coiled common ramus to the tongue tip. Hydromantes has the greatest projection distance of any salamander.     

Morphology and function of the tongue and hyoid apparatus in Varanus (Varanidae, Lacertilia)

The morphology and function of the tongue and hyoid apparatus in Varanus were examined by anatomical and experimental techniques. Morphological features unique to Varanus include a highly protrusible tongue that has lost a roughened dorsal surface, an exceptionally strong and mobile hyobranchial apparatus, a well-defined joint between the ceratohyal and anterior process, and a series of distinct muscles inserting at the anterior hyobranchial region. Varanus is also unusual among lizards in a number of feeding behaviors; it ingests prey entirely by inertial feeding, as the tongue does not participate in food transport. Further specializations include an increased reliance on hyobranchial movements in drinking and pharyngeal packing and compression. The long, narrow tongue is most likely related to the mechanics of tongue protrusion; the increased amount, strength, and complexity of hyobranchial movement is related to the fact that the hyobranchium in Varanus replaces the tongue in many functions. Previous hypotheses for the origin of these adaptations are discussed, and the difficulties of attributing these specializations to any specific scenario of adaptation or constraint are emphasized.     

Differential sensitivity of tongue areas and palate to electrical stimulation: a suprathreshold cross-modal matching study

A cross-modal matching procedure was used, in twelve subjects,to evaluate regional differences in suprathreshold sensitivityof the oral cavity to electrogustometric stimulation. Stimulationof five loci on each side of the oral cavity was performed:tongue tip (one cm from the midline), anterior tongue side (2.5cm from tip on lateral margin), posterior tongue side (regionof the foliate papillae), posterior medial tongue (one cm frommidline on circumvallate papillae), and soft palate (one cmfrom midline, one cm above superior pole of anterior palatinearch). The tip of the tongue was significantly more sensitivethan the other areas to electric stimulation, as evidenced bythe slope and absolute position of the psychophysical powerfunctions. Strong correlations were observed in the sensitivitymeasures across tongue loci and between tongue and palate sides.No effects of subject gender or mouth side were found.     

Cyclopamine and jervine in embryonic rat tongue cultures demonstrate a role for Shh signaling in taste papilla development and patterning: fungiform papillae double in number and form in novel locations in dorsal lingual epithelium

From time of embryonic emergence, the gustatory papilla types on the mammalian tongue have stereotypic anterior and posterior tongue locations. Furthermore, on anterior tongue, the fungiform papillae are patterned in rows. Among the many molecules that have potential roles in regulating papilla location and pattern, Sonic hedgehog (Shh) has been localized within early tongue and developing papillae. We used an embryonic, tongue organ culture system that retains temporal, spatial, and molecular characteristics of in vivo taste papilla morphogenesis and patterning to study the role of Shh in taste papilla development. Tongues from gestational day 14 rat embryos, when papillae are just beginning to emerge on dorsal tongue, were maintained in organ culture for 2 days. The steroidal alkaloids, cyclopamine and jervine, that specifically disrupt the Shh signaling pathway, or a Shh-blocking antibody were added to the standard culture medium. Controls included tongues cultured in the standard medium alone, and with addition of solanidine, an alkaloid that resembles cyclopamine structurally but that does not disrupt Shh signaling. In cultures with cyclopamine, jervine, or blocking antibody, fungiform papilla numbers doubled on the dorsal tongue with a distribution that essentially eliminated inter-papilla regions, compared with tongues in standard medium or solanidine. In addition, fungiform papillae developed on posterior oral tongue, just in front of and beside the single circumvallate papilla, regions where fungiform papillae do not typically develop. The Shh protein was in all fungiform papillae in embryonic tongues, and tongue cultures with standard medium or cyclopamine, and was conspicuously localized in the basement membrane region of the papillae. Ptc protein had a similar distribution to Shh, although the immunoproduct was more diffuse. Fungiform papillae did not develop on pharyngeal or ventral tongue in cyclopamine and jervine cultures, or in the tongue midline furrow, nor was development of the single circumvallate papilla altered. The results demonstrate a prominent role for Shh in fungiform papilla induction and patterning and indicate differences in morphogenetic control of fungiform and circumvallate papilla development and numbers. Furthermore, a previously unknown, broad competence of dorsal lingual epithelium to form fungiform papillae on both anterior and posterior oral tongue is revealed.     

Labeling afferent nerves in the tongue by peripheral transganglionic transport of HRP and WGA-HRP in the rat

Peripheral transganglionic transport of horseradish pcroxidase(HRP) and wheat germ agglutinin–horseradish peroxidaseconjugate (WGA–HRP) was used to label afferent fibersin the taste buds and lingual epithelium of the rat. Microinjectionsof the tracer were made in the brain stem central projectionarea of the afferent nerves to the tongue. Optimal labelingof nerve endings in the tongue was obtained when 2 µlof 20% HRP was injected into the brain stem and postinjectionsurvival times of 24–36 h were used. The distributionof single nerves was studied by using this tracing procedurein combination with strategic transections of the various afferentnerves supplying the tongue. Labeled nerve fibers from the combinedchorda tympani–lingual nerve were found in the epitheliumand in taste buds in the fungiform and anterior foliate papillaeof the anterior 3/4 of the tongue. Labeled nerve fibers in theepithelium of the anterior 2/3 of the tongue but none in tastebuds were found when the lingual nerve alone was studied, althoughnumerous perigeminal fibers were found. The glossopharyngealnerve was found to innervate die posterior 1/4 of the tongueepithelium including the taste buds of the circumvallate papillae.The glossopharyngeal nerve on one side was found to innervatethe taste buds on both sides of the midline. The results showthat this tracing procedure can be a useful supplement to othermethods for studying afferent nerves in the tongue.     

Filiform papillae of human, rat and swine tongue

In order to study the structure of filiform papillae (FP), tissue specimens were taken from the anterior part of the tongues of 8 humans, 8 rats and 8 swine. The formalin-fixed samples were processed routinely for scanning electron microscopy (SEM) and light microscopy. With SEM, FP of human tongue contained 5-12 hairs which were covered with a massive plaque of micro-organisms. FP of rat tongue, on the other hand, contained one papillary projection with smooth surface structure. Colonization of micro-organisms was seen on the anterior part of the body of FP, but not on the hairs. In the cross-section of FP of the rat tongue, the cells of the papilla were close to each other and no micro-organisms were seen within the papillae. On the contrary, the spaces between the squamous epithelial cells of the hairs of human FP contained numerous micro-organisms. The structure of FP of the swine tongue resembled that of the rat tongue. The hairs were smooth, and some micro-organisms were seen on the cell surface of the interpapillary areas. The structure of FP is discussed from the standpoint of different keratinization and colonization of micro-organisms.     

Intramural mechanics of the human tongue in association with physiological deformations

Contraction of the tongue musculature during speech and swallowing is associated with characteristic patterns of tissue deformation. In order to quantify local deformation (strain) in the human tongue, we used a non-invasive NMR tagging technique that represents tissue as discrete deforming elements. Subjects were studied with a fast gradient echo pulse sequence (TR,TE 2.3/0.8 ms, slice thickness 10 mm, and effective spatial resolution 1.3x1.3 mm). Individual elements were defined by selectively supersaturating bands of magnetic spills in resting tongue tissue along the antero-posterior and superior inferior directions of the mid-sagittal plane, resulting in a rectilinear square grid. Axial and shear strains relative to the rest condition were determined for each clement and represented by two-dimensional surface strain maps. During forward protrusion, the anterior tongue underwent positive antero posterior strain (elongation) (maximum 200%) and symmetrical negative medial lateral and superior inferior strain (contraction). During sagittal curl directed to the hard palate, the tongue exhibited positive asymmetrical antero posterior strain (maximum 160%) that increased radially as a function of distance from the center of curvature (r = 0.9216, p<0.0005), and commensurate negative strain in the medial lateral direction. Similarly, the magnitude of anterior posterior strain during left-directed tongue curl was proportional to the distance from the curved inner surface (r = O.8978, p<0.0005). We conclude that the regulation of tongue position for the motions studied was related to regional activation of the intrinsic lingual musculature.     

TONGUE MORPHOLOGY AND AFFINITIES OF THE HAWAIIAN HONEYCREEPER MELAMPROSOPS PHAEOSOMA

Melamprosops phaeosoma has a thick straight tongue with an anterior spoon-like trough, but without any indication of a tube. Its hyoid skeleton has an elongated, flattened basihyale, stout parallel paraglossalia and a concavity along the dorsal surface of the ceratobranchiale. All the tongue muscles could be dissected but the superficial M. mylohyoideus and M. serpihyoideus were too damaged to figure and describe. The M. ceratoglossus and the M. hypoglossus obliquus are large; the latter muscle inserts mainly on the basihyale and ceratobranchiale, but a small number of fibres pass beneath the basihylale. The M. hypoglossus anterior is absent. The morphology of the tongue apparatus of Melamprosops is very similar to that of Ciridops and of Loxops and supports placement of Melamprosops in the Drepanididae. Further it suggests that Melamprosops evolved from an ancestor with a tubular tongue, supporting the suggestion that it evolved from Loxops.     

Light and scanning electron microscopic study of the tongue in the cormorant Phalacrocorax carbo (Phalacrocoracidae, Aves)

The tongue of the cormorant Phalacrocorax carbo is a small, immobile structure with a length of 1.4 cm, situated in the middle part of the elongated lower bill. The uniquely shaped tongue resembles a mushroom, with a short base and an elongated dorsal part with sharpened anterior and posterior tips. A median crest can be observed on the surface of the tongue. Examination by light and scanning electron microscopy revealed that the whole tongue is formed by a dense connective tissue with many bundles of elastic fibers. The lingual mucosa is covered by a multilayered keratinized epithelium. The thickest, horny layer of the lingual epithelium was observed on the surface of the median crest and on the posterior tip of the tongue. Lingual glands are absent in cormorants. The framework of the tongue is composed of a hyoid cartilage incorporated into the base. The localization and structure of the tongue in the cormorant show that it is a rudimentary organ and that the lingual body, usually well-developed in birds, is conserved.     

Form and function of the tongue in agamid lizards with comments on its phylogenetic significance

The morphology of the tongue of agamid lizards is reviewed and discussed in the context of its functional and phylogenetic significance. It is shown that in several features, including the development of the central musculature of the tongue into a ring muscle and the presence of a genioglossus internus muscle in adults, the tongue in most agamids is derived relative to that in other squamates. In some features, such as the vertical connective tissue septa, agamids share primitive features with Sphenodon. Some conditions found in agamids are also found in anoline iguanids. Two genera, Uromastyx and Leiolepis, differ significantly from other agamids in intrinsic tongue musculature. The functional significance of the unique tongue morphology is that agamids utilize a different mechanism of tongue protrusion from that of other lizards. This mechanism involves the production of force against the lingual process, leading to an anterior slide of the tongue, and is detailed in this paper. Finally, I discuss the mechanical basis for the transformation series of tongue protrusion mechanisms from agamids to chamaeleonids. It is suggested that the mechanism of tongue protrusion in chamaeleonids is not unique, but is a highly derived state of the condition found in agamids.     

Effects of chorda tympani nerve anesthesia on taste responses in the NST

Human clinical and psychophysical observations suggest that the taste system is able to compensate for losses in peripheral nerve input, since patients do not commonly report decrements in whole mouth taste following chorda tympani nerve damage or anesthesia. Indeed, neurophysiological data from the rat nucleus of the solitary tract (NST) suggests that a release of inhibition (disinhibition) may occur centrally following chorda tympani nerve anesthesia. Our purpose was to study this possibility further. We recorded from 59 multi- and single- unit taste-responsive sites in the rat NST before, during and after recovery from chorda tympani nerve anesthesia. During anesthesia, average anterior tongue responses were eliminated but no compensatory increases in palatal or posterior tongue responses were observed. However, six individual sites displayed increased taste responsiveness during anesthesia. The average increase was 32.9%. Therefore, disinhibition of taste responses was observed, but infrequently and to a small degree in the NST At a subset of sites, chorda tympani-mediated responses decreased while greater superficial petrosal-mediated responses remained the same during anesthesia. Since this effect was accompanied by a decrease in spontaneous activity, we propose that taste compensation may result in part by a change in signal-to-noise ratio at a subset of sites.      

Growth and Activities of Sulfate-Reducing and Methanogenic Bacteria in Human Oral Cavity

Viable counts and activities of sulfate-reducing bacteria (SRB) and methanogenic bacteria were determined in the oral cavities of eight volunteers. Of these, seven harbored viable SRB populations, and six harbored viable methanogenic bacterial populations. Two volunteers classified as type III periodontal patients had both SRB and methanogenic bacteria. Six separate sites were sampled: posterior tongue, anterior tongue, mid-buccal mucosa, vestibular mucosa, supragingival plaque, and subgingival plaque. The SRB was found in all areas in one volunteer, and it was mostly present in posterior tongue, anterior tongue, supragingival, and subgingival plaques in many volunteers. The methanogenic bacteria were mostly found in supragingival and subgingival plaques. The activities of sulfate reduction and methane production were determined in randomly selected isolates. Received: 27 July 2002 / Accepted: 27 August 2002     

SEM study on the dorsal lingual surface of the common tree shrew, Tupaia glis.

The dorsal lingual surface of the common tree shrew was examined by SEM after treating it with HCl to remove the mucous substance. Filiform (FI), fungiform (FU) and circumvallate papillae (CI) were observed. The FI exhibited a small circular bulge surrounded by anterior and posterior filamentous processes. FU were scattered among the FI. There were 3 CI separating the anterior 4/5 from the posterior 1/5 of the tongue. In addition, a group of conical projections with caudal orientation was found anterior to the palatoglossal fold on each side of the tongue. Microridges were widely observed on the entire dorsal lingual surface, except on the free surface of FI processes.     

Observations on the structure of the subepithelial nerve plexus in the tongue

The intrinsic innervation of the anterior two thirds of the tongue in adult dogs of both sexes was studied in paraffin sections stained with Bodian, Holmes, cholinesterase and other stains. In all the sections, a subepithelial plexus of nerve fibres and cells was always seen on the dorsum of the tongue. Nerve endings were seen extending in between the epithelial cells on the dorsum of the tongue. The nerve cells were usually spindle-shaped and collected to form numerous ganglia in the submucosa. There were other ganglia in the tongue whose structure was very much similar to terminal autonomic ganglia. The significance of the ganglia consisting of biopolar nerve cells is being discussed.     

Separate and distinctive roles for Wnt5a in tongue, lingual tissue and taste papilla development

Although canonical Wnt signaling is known to regulate taste papilla induction and numbers, roles for noncanonical Wnt pathways in tongue and taste papilla development have not been explored. With mutant mice and whole tongue organ cultures we demonstrate that Wnt5a protein and message are within anterior tongue mesenchyme across embryo stages from the initiation of tongue formation, through papilla placode appearance and taste papilla development. The Wnt5a mutant tongue is severely shortened, with an ankyloglossia, and lingual mesenchyme is disorganized. However, fungiform papilla morphology, number and innervation are preserved, as is expression of the papilla marker, Shh. These data demonstrate that the genetic regulation for tongue size and shape can be separated from that directing lingual papilla development. Preserved number of papillae in a shortened tongue results in an increased density of fungiform papillae in the mutant tongues. In tongue organ cultures, exogenous Wnt5a profoundly suppresses papilla formation and simultaneously decreases canonical Wnt signaling as measured by the TOPGAL reporter. These findings suggest that Wnt5a antagonizes canonical Wnt signaling to dictate papilla number and spacing. In all, distinctive roles for Wnt5a in tongue size, fungiform papilla patterning and development are shown and a necessary balance between non-canonical and canonical Wnt paths in regulating tongue growth and fungiform papillae is proposed in a model, through the Ror2 receptor.     

Dynamic interaction between the tongue and soft palate during obstructive apnea in anesthetized patients with sleep-disordered breathing.

Little is known about the mechanisms of persistence of obstructive apnea. Structurally, the dorsum of the tongue locates anterior to the soft palate. On the basis of the observation of posterior displacement of the tongue during obstructive apnea, we hypothesized that the dorsum of the tongue pushes the anterior wall of the soft palate posteriorly during inspiratory efforts, maintaining closure at the retropalatal airway. To test this hypothesis, we measured the pressure between dorsum of the tongue and anterior wall of the soft palate (PT&P) during experimentally induced obstructive apneas in anesthetized patients with sleep-disordered breathing. P(T&P) changes during the obstruction significantly depended on collapsibility of the retroglossal airway. Progressive increase in the P(T&P) during obstructive apnea was observed only in patients with highly collapsible retroglossal airways. Significant increase in the P(T&P) during inspiratory effort in accordance with positive deflection pattern of P(T&P) tracing was evident in the patients with highly collapsible retroglossal airways. The results indicate significant dynamic interaction between the tongue and soft palate during both obstructive apnea and each inspiratory effort, possibly maintaining closure at the retropalatal airway.