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BACKGROUND AND PURPOSE: When optimising radiotherapy treatments today, the pharyngeal constrictor muscles and the larynx are usually regarded as the swallowing organs at risk (SWOARs). The purpose of this study was to identify and describe additional, previously undefined groups of muscles (functional units) involved in crucial components of swallowing (hyolaryngeal elevation (HLE), tongue base retraction (TBR) and tongue motion), and to emphasise their relevance in radiation-induced dysphagia.MATERIAL AND METHODS: Based on available literature on human anatomy and swallowing physiology, the functional units of muscles involved in HLE, TBR and tongue motion have been identified and described.RESULTS AND CONCLUSION: Functional swallowing units (FSUs) were defined as groups of swallowing muscles sharing their function, that are in close proximity to each other. Seven FSUs involved in HLE, TBR and tongue motion were identified: floor of mouth, thyrohyoid muscles, posterior digastric/stylohyoid muscles complex, longitudinal pharyngeal muscles, hyoglossus/styloglossus muscles complex, genioglossus muscles, intrinsic tongue muscles. The swallowing physiology and anatomy of the FSUs described in this paper will lead to a greater understanding of radiation-induced dysphagia mechanisms and, consequently, to an improvement in the development of swallowing sparing strategies. This article (PART 1) serves as the theoretical foundation for a subsequent article (PART 2), which provides detailed delineation guidelines for FSUs.

Objectives We aimed to discuss the definition of lingual lymph nodes based on the deep cervical fascia anatomy. Methods A total of 11 cadavers were histopathologically evaluated. Specimens were dissected into serial stepwise cross-sections. The deep cervical fascia and lymph nodes were evaluated by staining the cross-sections with Elastica van Gieson and hematoxylin and eosin stains, respectively. The gross anatomy of the deep cervical fascia was evaluated in 1 cadaver after bilateral dissection. Results A single severely degenerated medial lingual lymph node (MLLN) was identified in the lingual septum of 1 cadaver. A single lateral lingual lymph node (LLLN) was identified in 1 cadaver, between the genioglossus and the hyoglossus. The superficial layer of deep cervical fascia (SLDF) was found to split into two layers encapsulating the submandibular gland. The deeper portion of this fascia was in close contiguity with the hyoglossus and formed the boundary between the mouth and the neck. The SLDF was found to be discontinuous with the area between the sublingual space and the submandibular gland. Conclusions The SLDF could serve as an anatomical landmark for lingual lymph nodes, since it is considered to be the demarcating boundary during neck dissection. When lymph nodes that are located deeper than the SLDF were defined as lingual lymph nodes, 1 MLLN and 1 LLLN were identified.

During primate swallowing, tongue base retraction (TBR) drives the food bolus across the oropharynx towards the esophagus and flips the epiglottis over the laryngeal inlet, protecting against penetration and aspiration of food into the airway. Despite the importance of TBR for swallowing performance, the mechanics of TBR are poorly understood. Using biplanar videoradiography (XROMM) of four macaque monkeys, we tested the extrinsic muscle shortening hypothesis, which posits that shortening of the hyoglossus and styloglossus muscles pulls the tongue base posteriorly, and the muscular hydrostat or intrinsic tongue muscle hypothesis, which suggests that, because the tongue is composed of incompressible fluid, intrinsic muscle shortening increases tongue length and displaces the tongue base posteriorly. Our data falsify these hypotheses. Instead we suggest a novel hydraulic mechanism of TBR: shortening and rotation of suprahyoid muscles compresses the tongue between the hard palate, hyoid and mouth floor, squeezing the midline tongue base and food bolus back into the oropharynx. Our hydraulic mechanism is consistent with available data on human tongue swallowing kinematics. Rehabilitation for poor tongue base retraction might benefit from including suprahyoid muscle exercises during treatment.

The tongue is a dense muscular organ, in which the muscles are arranged in a confusing pattern. The intrinsic muscles were gross anatomically investigated in 25 cadavers to clarify their configuration. The superior longitudinal muscle (SLm) ran beneath the dorsal mucosa and was divided into bundles by the other muscles passing through it to the dorsum. The external bundle of the styloglossus, with the palatoglossus, coursed externally to the hyoglossus. Their fibers spread beneath the SLm or attached to the apex and the inferior longitudinal muscle (ILm) ascended from the root and joined them. The genioglossus and the anterior part of the hyoglossus extended internally and externally to the ILm, respectively, to the dorsum, and the vertical muscle was sandwiched between them. The transverse muscle passed laterally from the lingual septum. The fibers of the posterior part of the hyoglossus converged to the root and spread beneath the SLm. The intersections between these vertical and transverse fibers divided one another into bundles or lamellae. The middle bundle of the styloglossus, passing between the two parts of the hyoglossus, was divided into slips by the intersection with the genioglossus. The internal bundle of the styloglossus, with the glossopharyngeus, descended internally to the posterior part to the root. The findings indicate that the intrinsic and extrinsic muscles of the tongue are not independent groups, and their fibers form a three-dimensional latticework. Each muscle contains numerous bundles or lamellae as functional units that can act separately or cooperate across the muscles.

Introduction The hypoglossal nerve (HN) may be damaged during neck surgeries. A detailed understanding of the anatomy of the hypoglossal nerve in relation to various anatomical landmarks and surrounding structures is important to reduce procedural complications and the risk of nerve damage. Methods The study was carried out using 21 sagittal sections of head and neck of male cadavers (16 right and 5 left). To measure the required parameters the anatomical landmarks taken were the posterior belly of digastric muscle (PBD), descendens hypoglossi of ansa cervicalis (AC), bifurcation of common carotid artery (CCA), tip of the mastoid process (TMP), lingual artery (LA) at the posterior border of hyoglossus muscle, tip of greater cornu of hyoid bone (GH), bifurcation of CCA to the attachment of descendens hypoglossi (CCA-AC). Results The length of PBD was 3.77 ± 1.08 cm on right side and 3.15 ± 0.05 cm on left side. The extended length was measured from the TMP to GH which was 6.7 ± 1.23 cm on right side and 6.75 ± 0.75 on left side. The distance between HN to the GH did not vary much between right and left sides which were 1.21 ± 0.35 and 1.2 ± 0.69 cm respectively. The distance between HN and LA at the posterior border of hyoglossus was 1.12 ± 0.3 cm on right side and 0.5 ± 0.3 cm on left side. Discussion HN dissection is more complicated, as the location of the HN is much deeper. Moreover, there are no bony landmarks around the HN. Therefore PBD, CCA, ICA, ECA, LA were used as landmarks for HN in head and neck surgeries.

Objectives/hypothesis Upper airway stimulation for obstructive sleep apnea (OSA) via implantable hypoglossal nerve stimulation (HGNS) reduces airway obstruction by selectively stimulating nerve fibers that innervate muscles that produce tongue protrusion, while avoiding fibers that produce tongue retraction. This selective stimulation likely depends upon the location, intensity, and type of electrical stimulation delivered. This study investigates the impact of changing stimulation parameters on tongue muscle activation during HGNS using intraoperative nerve integrity monitoring in conjunction with electromyography (EMG). Study design Prospective case series. Methods Ten patients undergoing unilateral HGNS implantation for OSA in a university hospital setting were studied. Data included EMG responses in tongue muscles that produce protrusion (genioglossus), retraction (styloglossus/hyoglossus), and stiffening (transverse/vertical) in response to intraoperative bipolar probe electrical stimulation of lateral and medial branches of the hypoglossal nerve (HGN) and to implantable pulse generator (IPG) unipolar and bipolar settings after placement of the stimulation cuff. Results Stimulation of medial division HGN branches resulted in EMG responses in genioglossus muscles, but not in styloglossus/hyoglossus muscles, whereas stimulation of the lateral division HGN branches drove responses in styloglossus/hyoglossus muscles. Variable responses in transverse/vertical muscles were observed with stimulation of lateral and medial division branches. After electrode cuff placement, unipolar and bipolar HGN stimulation configurations of IPG resulted in unique patterns of muscle activation. Conclusions The relative activation of extrinsic and intrinsic tongue musculature by HGNS is determined by stimulus location, intensity, and type. Intraoperative neurophysiological monitoring of tongue muscle activation enables proper electrode cuff placement and may provide essential data for stimulus optimization. Level of evidence 4 Laryngoscope, 130:1836-1843, 2020.

Purpose Tongue base retraction during swallowing is critical to bolus propulsion in normal physiological swallowing. A better understanding of the hyoglossus and styloglossus, muscles thought to be key to tongue base retraction, will improve the quality of physical rehabilitation in dysphagic patients in addition to preventing iatrogenic damage to structures critical to deglutition. This study utilized muscle functional MRI in healthy adult human subjects in order to determine if the hyoglossus and styloglossus are active during swallowing. Methods and materials Data were collected for 11 subjects with mfMRI before and after swallowing, and after performing the Mendelsohn maneuver. Whole muscle relaxation time profiles (T2 signal in milliseconds) were calculated from weighted averages of multiple dual echo MRI slices, allowing for comparison of physiological response for the muscles in each test condition. Changes in effect size (Cohen’s d) of whole muscle T2 profiles were used to establish whether or not the hyoglossus and styloglossus are utilized during swallowing and during the Mendelsohn maneuver. Results Post-swallowing effect size changes (where a d value of > 0.20 indicates significant activity) for the T2 signal profiles of the hyoglossus and styloglossus were found to be d = 1.19 and 0.22, respectively. The hyoglossus showed an effect size change of d = 0.26 for the Mendelsohn maneuver. Conclusions Muscle functional MRI indicates a physiological response of the hyoglossus and styloglossus during swallowing, and the hyoglossus during the Mendelsohn maneuver.

Three hypothetical models of tongue movement of the walrus during suction feeding are examined. These models encompass the entire range of simple tongue retraction movements possible by examining 1) movement of the tongue directly to the rear following the curvature of the palate, 2) to the rear and ventrally in a straight line, and 3) ventrally in a straight line. The percent of muscular force available from the hyoglossus, genioglossus, and styloglossus that could be applied toward retraction as predicted by each model is calculated. The resistance that the tongue would provide during retraction is calculated using projected tongue areas and is combined with the above data from the muscles to provide an estimate of the percent of the total available force that is needed to retract the tongue for each model. A separate examination of the direction of tongue-induced wear striations on the palatal and lingual aspects of the teeth is used to help support or reject the three models. The model where the tongue is moved directly to the rear is supported by studies of both muscle force and tooth wear. In the mammalian groups that were compared to the walrus, there is a great deal of interspecific variation in movements of the tongue during suction feeding; no two groups can be considered to have identical stereotyped tongue movements.

Tongue musculature in 24 genera of snakes was examined histologically. In all snakes, the tongue is composed of a few main groups of muscles. The M. hyoglossus is a paired bundle in the center of the tongue. The posterior regions of the tongue possess musculature that surrounds these bundles and is responsible for protrusion. Anterior tongue regions contain hyoglossal bundles, dorsal longitudinal muscle bundles and vertical and transverse bundles, which are perpendicular to the long axis of the tongue. The interaction of the longitudinal with the vertical and horizontal muscles is responsible for bending during tongue flicking. Despite general similarities, distinct patterns of intrinsic tongue musculature characterize each infraorder of snakes. The Henophidia are primitive; the Scolecophidia and Caenophidia are each distinguished by derived characters. These derived characters support hypotheses that these latter taxa are each monophyletic. Cylindrophis (Anilioidea) is in some characters intermediate between Booidea and Colubroidea. The condition in the Booidea resembles the lizard condition; however, no synapomorphies of tongue musculature confirm a relationship with any specific lizard family. Although the pattern of colubroids appears to be the most biomechanically specialized, as yet no behavioral or performance feature has been identified to distinguish them from other snakes.

Prey capture in Agama stellio was recorded by high-speed video in combination with the electrical activity of both jaw and hyolingual muscles. Quantification of kinematics and muscle activity patterns facilitated their correlation during kinematic phases. Changes in angular velocity of the gape let the strike be subdivided into four kinematic phases: slow open (SO1 and SOII), fast open (FO), fast close (FC), and slow close-power stroke (SC/PS). The SO1 phase is marked by initial activity in the tongue protractor, the hyoid protractor, and the ring muscle. These muscles project the tongue beyond the anterior margin of the jaw. During the SO11 phase, a low level of activity in the jaw closers correlates with a decline of the jaw-opening velocity. Next, bilateral activity in the jaw openers defines the start of the FO phase. This activity ends at maximal gape. Simultaneously, the hyoid retractor and the hyoglossus become active, causing tongue retraction during the FO phase. At maximal gape, the jaw closers contract simultaneously, initiating the FC phase. After a short pause, they contract again and the prey is crushed during the SC/PS phase. Our results support the hypothesis of tongue projection in agamids by Smith (( 19881 J. Morphol. 196:157-171), and show some striking similarities with muscle activity patterns during the strike in chameleons (Wainwright and Bennett ( 1992al J. Exp. Biol. 168:l-21). Differences are in the activation pattern of the hyoglossus. The agamid tongue projection mechanism appears to be an ideal mechanical precursor for the ballistic tongue projection mechanism of chameleonids; the key derived feature in the chameleon tongue projection mechanism most likely lies in the changed motor pattern controlling the hyoglossus muscle. B 1995 Wiley-Liss, Inc. Lizards capture prey by means of lingual or jaw prehension (Smith, '84, '88). Some authors argue that jaw-mediated prey cap- ture might be the primitive condition in squa- mates (Bels and Goosse, '90; referring to the plesiomorphic condition of jaw prehension in Amphibia, Thexton et al., '77; Lauder and ShafTer, '88; Findeis and Bemis, '90; Nishi- kawa and Cannatella, '91; Anderson, '931, but lingual prey capture is considered by most authors to be more primitive than jaw capture (Schwenk, '88; Schwenk and Bell, '88; Schwenk and Throckmorton, '89). Most functional morphological studies con- cerning prey capture in lizards have empha- sized a few groups belonging to the Iguania (Agamidae: Schwenk and Throckmorton, '89; Kraklau, '91; Iguanidae: Schwenk and Throckmorton, '89; Bels, '90; Bels and Go- osse, '90; Delheusy and Bels, '92; and Cha- maeleonidae: Gnanamuthu, '30; Zoond, '33; Bels and Baltus, '87; Bell, '89, '90; Wain

The purpose of the present study was to classify the courses of the human sublingual artery. For this purpose, the arteries supplying the floor of the mouth and the tongue were gross anatomically investigated using 101 sides of 53 cadavers. The courses were divided into three categories: those passing medial or lateral to the hyoglossus (categories M and L) and that piercing the mylohyoid (category P); they were subdivided into five types. Category M had one type, regarded as the usual one, in which the lingual artery took the usual pattern of distribution. Categories L and P, in which the sublingual artery arose from the facial or submental artery, had the respective two types and were collectively regarded as the unusual type. Sixty-one and 36 of the 101 sides were of the usual and unusual types, respectively, the latter of which included 17 of category L and 19 of category P. The remaining four were variations of the lingual artery itself. On examining the types by gender, the usual type was more often found in females (75.6 %), whereas the unusual type was more often found in males (48.1 %). Bilateral occurrence of the same type was often found in both the usual type (77.4 %) and the unusual type (65.0 %). Existence of the sublingual artery branch significantly increased the thicknesses of the submental arteries. The classification proposed here will conceivably contribute to safer dental implant surgery and more accurate interpretation of angiographic images of arteries in the floor of the mouth.

Little is known about the anatomy and musculature of the gray whale (Eschrichtius robustus), especially related to the anatomy of the tongue and hyoid region. The recovery of an extremely fresh head of a neonatal female gray whale provided an opportunity to conduct the first in-depth investigation of the musculoskeletal features of the tongue and hyoid apparatus. Unlike other mysticetes, the gray whale tongue is strong, muscular, and freely mobile inside the buccal cavity. In particular, the genioglossus and hyoglossus muscles are extremely large and robust making up the majority of the body of the tongue. In addition, the genioglossus had a unique position and fiber orientation in the tongue compared to other mammals. The structure of the hyoid apparatus differs between E. robustus and other mysticete species, although there are similarities among individual elements. We provide the first documentation of fungiform papillae that may be associated with taste buds in Mysticeti. The highly mobile, robust tongue and the presence of well-defined tongue and hyoid musculature are in keeping with observations of gray whale feeding that suggest this group of whales utilize oral suction to draw benthic prey into the buccal cavity. Anat Rec, 298:660–674, 2015. © 2014 Wiley Periodicals, Inc.

The purpose of this study was to show the benefit of visualizing a tree-dimensional (3D) image of the tongue's muscle structure, which until now has been regarded as fully understood. Until now, no suitable 3D observation methods have been developed for soft tissue, such as the tongue, using histological magnification. For this purpose, this study used a micro-computed tomographic method (micro-CT) and image processing after the fixation, decalcification, and dehydration of a mouse tongue. Results: 3D rendered images of tongue muscles obtained by micro-CT showed every mus- cle and their relationships to each other. The superior longitudinal and the hyoglossus muscles of the tongue made up one group, while the inferior longitudinal and the styloglossus muscles of the tongue made up another. The boundary of the two muscles in each group was difficult to distin- guish. On the other hand, what appear to be newly described muscles were identified. These re- sults indicate that our micro-CT method is beneficial and that classical knowledge of tongue mus- cles derived from two-dimensional (2D) images does not fully describe the actual complexity of the tongue muscles. In our opinion, 3D rendered images mixed with raw structure can provide a more in-depth picture of the tongue from an integrated as well as an analytical perspective.

The hypoglossal or twelfth cranial nerve is the motor nerve to the extrinsic and intrinsic muscles of the tongue, and the superior root of the ansa cervicalis and the thyrohyoid and geniohyoid branches are delivered through the nerve. This study investigated the muscular branches of the hypoglossal nerve to clarify their spatial relationships with the muscles of the tongue and the neighboring structures. The muscles and the nerve were gross anatomically examined in 42 cadavers. The superior root and the thyrohyoid branch left the nerve near the occipital and lingual arteries, respectively. The extrinsic muscles consisted of some components, and the geniohyoid branch and the lingual branches arose on the hyoglossus. The ascending lingual branches formed a plexus on the anterior part of the hyoglossus and were divided into the proximal and distal groups. They supplied the two parts of the hyoglossus, the three bundles of the styloglossus and the superior and inferior longitudinal muscles and communicated with the lingual nerve. The descending lingual branches supplied the inferior part of the genioglossus, and the terminal branches gave intramuscular twigs to its main part and the transverse and vertical muscles. The findings indicated that the branching pattern of the hypoglossal nerve is characterized by the positional relationships to the components of the extrinsic muscles. The hyoid bone can be an effective marker to identify the branches and affected position if it was used in combination with the morphology of the extrinsic muscles, and the knowledge of their variations is also beneficial. Anat Rec, 302:558-567, 2019. © 2018 Wiley Periodicals, Inc.

Although respiratory muscle motor units have been studied during natural breathing, simultaneous measures of muscle force have never been obtained. Tongue retractor muscles, such as the hyoglossus (HG), play an important role in swallowing, licking, chewing, breathing, and, in humans, speech. The HG is phasically recruited during the inspiratory phase of the respiratory cycle. Moreover, in urethane anesthetized rats the drive to the HG waxes and wanes spontaneously, providing a unique opportunity to study motor unit firing patterns as the muscle is driven naturally by the central pattern generator for breathing. We recorded tongue retraction force, the whole HG muscle EMG and the activity of 38 HG motor units in spontaneously breathing anesthetized rats under low-force and high-force conditions. Activity in all cases was confined to the inspiratory phase of the respiratory cycle. Changes in the EMG were correlated significantly with corresponding changes in force, with the change in EMG able to predict 53–68% of the force variation. Mean and peak motor unit firing rates were greater under high-force conditions, although the magnitude of discharge rate modulation varied widely across the population. Changes in mean and peak firing rates were significantly correlated with the corresponding changes in force, but the correlations were weak ( r2 = 0.27 and 0.25, respectively). These data indicate that, during spontaneous breathing, recruitment of HG motor units plays a critical role in the control of muscle force, with firing rate modulation playing an important but lesser role.

Presence of accessory submandibular salivary gland (ASSG) is an extremely rare variation. Knowledge of its relations could be very useful to oral and maxillofacial surgeons, head and neck surgeons, and radiologists. During dissection classes, an ASSG was noted between the mylohyoid and hyoglossus muscles. The main submandibular salivary gland had superficial and deep parts. The deep part was narrow and measured about 5 cm. The lingual nerve passed between the superficial and deep parts. The accessory submandibular gland was situated below and parallel to the deep part of SSG. It also measured 5 cm. The ASSG had its own duct, which joined the duct of main gland. The ASSG and the deep part of the SSG were united at the lateral border of geniohyoid muscle to give a characteristic ‘‘horseshoe’’ appearance. The ASSG overlapped both lingual and hypoglossal nerves.

The pharyngeal muscles overlap each other and some of their parts have different areas of origin. Such arrangements make the interrelationships among pharyngeal muscles complicated. This study investigated the attachments of the middle constrictor to clarify its configuration and re-examine its functions. The gross anatomies of the pharyngeal and neighboring muscles were examined in 41 cadavers. The middle constrictor arose from the stylohyoid ligament and the hyoid bone, and its fibers were divided into three overlapping groups. The anterosuperior group ascended posterosuperiorly from the ligament and the lesser horn and fanned out. The middle group ascended posterosuperiorly from the greater horn and fanned out. The posteroinferior group fanned out from the posterior part of the greater horn, while the middle constrictor arose internally to the hyoglossus; some fibers often passed externally, and their fibers sometimes intersected around the lingual artery, which ran between them. Some fibers attached to the hyoglossus, occasionally to the stylohyoid and the posterior belly of the digastric, but seldom to the lingual artery and the triticeal cartilage in the thyrohyoid ligament. The three groups were inserted into the pharyngeal raphe, and the descending fibers joined the longitudinal pharyngeal muscles. The ascending and descending fibers rarely reached the top of the pharynx and the thyroid cartilage, respectively. The fiber arrangement suggested that, besides constriction of the pharynx, the ascending and descending fibers of the middle constrictor can act as an elevator muscle, and the irregular attachments could affect the functions of the muscles and vessels. Clin. Anat. 603–609, 2014. © 2013 Wiley Periodicals, Inc.

The TNM classification for oral malignancies has been criticized for its upstaging to T4a when tumour involves styloglossus, hyoglossus, palatoglossus and genioglossus. The aims of this study were to (1) create an anatomical computer atlas of extrinsic tongue musculature, and (2) reassess the original staging of pre-treatment archived magnetic resonance images (MRI) of tongue carcinomas using the strict extrinsic muscle criteria. The anatomy of the extrinsic tongue muscles was mapped using images from the Visible Human Project (VHP) to create a computer model of the extrinsic tongue muscles. This was co-registered with 87 archived pre-staging MRI scans of tongue carcinomas to assess tumour ingress of the extrinsic tongue muscles. Of the 87 image sets reviewed, 16 were of superficial tumours not visible on MRI. In the remaining 71 cases that showed positive extrinsic muscle tumour ingress, 52% were upstaged from T1/2/3 tumours to cT4a based upon this finding. Extrinsic lateral and genioglossus muscle invasion did not predict occult cervical lymph node invasion or disease-related survival. In conclusion, tumour invasion of styloglossus or hyoglossus would result in the majority of lateral tongue tumours being staged T4a. Such stratification is of little clinical relevance, and an alternative more reliable method is required.

The base of tongue plays an important role in the pathophysiology of obstructive sleep apnea (OSA), and addressing the tongue base can be an alternative for continuous positive airway pressure and some surgeries done such as uvulopalatopharyngoplasty to treat OSA. One of the options to address the tongue base to treat the OSA is stimulation of human hypoglossal nerve with implantable devices. Thus, one needs to know the physiology of the human hypoglossal nerve to understand the mechanism of such devices. The human hypoglossal nerve is the primary motor nerve that innervates 7 of the 8 extrinsic and intrinsic muscles of the tongue on each side, except palatoglossus muscle, which is innervated by the ipsilateral vagus nerve. At the level of the hyoid bone, the human hypoglossal nerve (the proximal and midthird) is normally unfasciculated, becoming fascicular only in its distal third where it branches. The lateral branch has many small, independent branches that supply the styloglossus, hyoglossus, and inferior longitudinalis muscles. The medial branch innervates the genioglossus, superior longitudinalis, transversalis, and verticalis muscles.

This work was carried out on the heads of sixteen apparently healthy adult domestic cats of various sex- es. The available literatures dealing with the gross anatomy of the hyoid apparatus of various species of domestic animals were reviewed. The Manual dissection, dry bony specimens as well as histological exami- nation were used according to the purpose under investigation. The study described the six segments forming the cat hyoid apparatus; Tympanohyoideum, Stylohyoide- um, Epihyoideum, Ceratohyoideum, Basihyoideum and Thyrohyoideum.Elevenpairs of muscles where attached to the hyoid apparatus. The most enlarged of these muscles were those that retract the hyoid bone caudally, the sternohyoid muscle and rostrally, the geniohyoid muscle as well as those controlling the tongue movement, styloglossus and hyoglossus. The histological studies showed that the Tympano-hyoideum and Ceratohyoideum were mostly hyaline cartilage, the Epihyoideum was partly hyaline and partly fibrous, the Stylohyoideum and Thyrohyoideum were partly hyaline and partly osseous while the Basihyoideum was completely ossified. The results obtained were photographed, described and discussed with those of other domestic animals.

The salivary glands comprise the parotid, the submandibular and the sublingual glands as well as small subsidiary glands scattered beneath the mucosa of the buccal cavity. The largest, the parotid, drains by its duct into the mouth at the level of the second upper molar tooth. It is traversed by the facial nerve, which may be invaded by a parotid carcinoma or damaged during parotid surgery. The submandibular gland has a superficial part, at the angle of the jaw, wedged between the mandible and the mylohyoid muscle. Its deep part projects deep to mylohyoid along the hyoglossus. Its duct opens alongside the fraenum of the tongue, where it is easily visible, and is crossed by the lingual nerve. The sublingual gland lies in the floor of the mouth into which it drains directly by a series of short ducts.

The tongue is a highly muscular organ, and the extrinsic muscles of the tongue overlap one another, which makes their configuration intricate. The aim was to clarify their spatial interrelationships. The extrinsic muscles of the tongue were gross anatomically investigated in 42 cadavers. The hyoglossus consisted of two parts arising from the hyoid body and the greater horn, respectively. They ascended to the tongue external to the genioglossus. The quadrilateral anterior part contained several parallel bundles, and the triangular posterior part comprised of three bundles converging to the insertion. The genioglossus consisted of many compartments. Although its fibers radiated from the mental spine to the tongue internal to the inferior longitudinal muscle, the lateral fibers of the inferior parts passed externally to it, and the most inferior part attached to the hyoid bone and the epiglottis. The chondroglossus arising from the lesser horn ascended internally to the lateral fibers of the inferior parts. The styloglossus arising from the styloid process coursed externally and internally to the hyoglossus and between its two parts. The external and internal bundles fused with the fibers of the palatoglossus and the superior pharyngeal constrictor, respectively. Three bundles were inserted into the apex with the inferior longitudinal muscle or joined the inferior parts of the genioglossus. The extrinsic muscles are subdivided into small groups of fibers depending on their spatial interrelationships, suggesting that they contain many functional units. Some units act separately, and others cooperate across the traditional boundaries of the muscles.

This study reports an investigation of the well-known context-dependent variation in English /r/ using a biomechanical tongue-jaw-hyoid model. The simulation results show that preferred /r/ variants require less volume displacement, relative strain, and relative muscle stress than variants that are not preferred. This study also uncovers a previously unknown mechanism in tongue biomechanics for /r/ production: Torque in the sagittal plane about the mental spine. This torque enables raising of the tongue anterior for retroflexed [Symbol: see text] by activation of hyoglossus and relaxation of anterior genioglossus. The results provide a deeper understanding of the articulatory factors that govern contextual phonetic variation.

There are many reports on the variation of origin site of the lingual artery branching from the external carotid artery. However, there are few reports systematically investigating the course of the lingual artery in detail from branching site to the body of tongue. The purpose of this study is to classify systematically the courses of the lingual artery including variations. Using 111 body sides of 63 Japanese cadavers for dissection practices, the lingual artery and the surrounding structures were investigated gross anatomically. The courses of the lingual artery were classified into five types based on the positional relationships with the hyoglossus and the mylohyoid as follows; type M: coursing medial to the hyoglossus (normal course, 104 sides), type L: coursing lateral to the hyoglossus (2 sides), type T: transferring its course from lateral to medial to the hyoglossus (2 sides), type P: penetrating the mylohyoid (2 sides), and type C: the coinciding of types M and P (1 side). Types L, T, P, and C were considered to be variant lingual arteries. Types M and T, type L, and type P arose from the external carotid, facial, and submental arteries, respectively. In types L and P, adding to the variant lingual artery, the remnant of the normal lingual artery was also observed. The present study provides detailed information on the courses of lingual artery which will be of clinical importance especially in the super-selective arterial angiography.

A wide range of pathologic processes may involve the floor of the mouth, the part of the oral cavity that is located beneath the tongue. They include lesions that arise uniquely in this location (eg, ranula, submandibular duct obstruction) as well as various malignancies, inflammatory processes, and vascular abnormalities that may also occur elsewhere in the head and neck. Some lesions that arise in superficial tissues such as the mucosa may be easily diagnosed at physical examination. However, computed tomography, magnetic resonance imaging, or ultrasonography may be necessary for a reliable assessment of lesion extension to deeper structures. In such cases, knowledge of the complex muscular, vascular, glandular, ductal, and neural anatomy of the region is important for accurate diagnosis and treatment planning. Familiarity with the radiologic imaging appearances of the floor of the mouth and recognition of anatomic landmarks such as the mylohyoid and hyoglossus muscles are especially useful for localizing disease within this region.

UNLABELLED With 8 figures SUMMARY This study aimed at revealing the origin, course and distribution of the hypoglossal nerve in 20 adult male New Zealand rabbits. In all the animals dissected, the hypoglossal nerve arose from the ventrolateral side of the medulla oblongata with two main roots and gave off a descending branch to the ansa cervicalis before reaching the division of the common carotid artery. This branch was not seen on the right side of only one case. At the lateral aspect of the hyoglossus muscle, the nerve then divided into the lateral and medial main branches, sent branches to the styloglossus, hyoglossus, genioglossus and geniohyoideus muscles and terminated in the intrinsic tongue muscles. A communicating branch was observed between the hypoglossal and accessory nerves in the right side of one animal and between the hypoglossal nerve and the ganglion nodosum in the right retropharyngeal area of another animal. An additional branch was observed innervating the stylohyoideus muscle in one animal only. A lateral lingual-hypoglossal communication was also seen between the lateral branch of the hypoglossal nerve and terminal branches of the lingual nerve.

SUMMARYA combination of dissection and computed tomography scanning has provided significant novel insights into the structure and function of the Dermochelys coriacea larynx and its associated muscles. Several previously unknown features of the laryngeal aditus (glottis) are described and their functional significance in its opening and closure are considered. The tongue plays an essential part in producing and maintaining closure during dives and feeding bouts. Closure is brought about by compression of the glottis under the action of the two hyoglossus muscles. The tongue thus plays the role of the epiglottis of mammals, sealing the entrance to the larynx. As is already clear, opening is brought about by abduction of the arytenoid cartilages. In addition, there is a powerful mechanism for maintaining the larynx in close apposition to the hyoid plate during feeding and neck flexion, thereby enhancing the efficiency of feeding.

The human tongue muscle hyoglossus (HG) muscle is active in oro-motor behaviors encompassing a wide range of tongue movement speeds. Here we test the hypothesis that the human HG is composed of “uncommon” myosin heavy chain (MHC) isoforms MHCembryonic, MHCneonatal and MHCslow tonic as has been reported for other head and neck muscles active during kinematically diverse behaviors. Following reaction of human HG with antibodies specific for MHCI, MHCIIA, MHCII, MHCembryonic, MHCextraocular, MHCneonatal and MHCslow tonic only antibodies to MHCI, MHCIIA and MHCIIA-X label more than occasional muscle fibers. These antibodies describe five phenotypes with prevalence MHCIIA>MHCI>MHCI-IIX>MHCI-IIA>MHCIIX. In MHC composition, the human HG is thus similar to human appendicular muscles, the human tongue muscle styloglossus and many human head and neck muscles but different from human masseter and extraocular muscles which contain five or more MHC isoforms.

The pharyngeal swallow can be elicited as an isolated event but, in normal animals, it occurs within the context of rhythmic tongue and jaw movement (RTJM). The response includes activation of the multifunctional geniohyoid muscle, which can either protract the hyoid or assist jaw opening; in conscious nonprimate mammals, two bursts of geniohyoid EMG activity (GHemg) occur in swallow cycles at times consistent with these two actions. However, during experimentally elicited pharyngeal swallows, GHemg classically occurs at the same time as hyoglossus and mylohyoid activity (short latency response) but, when the swallow is elicited in the decerebrate in the absence of RTJM, GHemg occurs later in the swallow (long latency response). We tested the hypothesis that it was not influences from higher centers but a brain stem mechanism, associated with RTJM, which caused GHemg to occur earlier in the swallow. In 38 decerebrate piglets, RTJM occurred sporadically in seven animals. Before RTJM, GHemg had a long latency, but, during RTJM, swallow related GHemg occurred synchronously with activity in hyoglossus and mylohyoid, early in the swallow. Both early and late responses were present during the changeover period. During this changeover period, duplicate electrodes in the geniohyoid could individually detect either the early or the late burst in the same swallow. This suggested that two sets of geniohyoid task units existed that were potentially active in the swallow and that they were differentially facilitated or inhibited depending on the presence or absence of rhythmic activity originating in the brain stem.

The present study evaluated the effect of coactivation of tongue protrusors and retractors on pharyngeal patency in patients with obstructive sleep apnea. The effect of genioglossus (GG), hyoglossus (HG), and coactivation of both on nasal pressure (Pn):flow relationships was evaluated in a sleep study (SlS, n = 7) and during a propofol anesthesia study (AnS, n = 7). GG was stimulated with sublingual surface electrodes in SlS and with intramuscular electrodes in AnS, while HG was stimulated with surface electrodes in both groups. In the AnS, the cross-sectional area (CSA):Pn relationships was measured with a pharyngoscope to estimate velopharyngeal compliance . In the SlS, surface stimulation of GG had no effect on the critical pressure (Pcrit), HG increased Pcrit from 2.8 ± 1.7 to 3.7 ± 1.6 cmH2O, but coactivation lowered Pcrit to 0.2 ± 1.9 cmH2O ( P < 0.01 for both). In the AnS, intramuscular stimulation of GG lowered Pcrit from 2.6 ± 1.3 to 1.0 ± 2.8 cmH2O, HG increased Pcrit to 6.2 ± 2.5 cmH2O ( P < 0.01), and coactivation had a similar effect to that of GG (Pcrit = 1.2 ± 2.4 cmH2O, P < 0.05). None of the interventions affected significantly velopharyngeal compliance. We conclude that the beneficial effect of coactivation depends on the pattern of GG fiber recruitment: although surface stimulation of GG failed to protrude the tongue, it prevented the occlusive effect of the retractor, thereby improving pharyngeal patency during coactivation. Stimulation of deeper GG fibers with intramuscular electrodes enlarged the pharynx, and coactivation had no additive effect.

Dermoid or epidermoid cysts usually appear as a midline swelling in the sublingual region or the floor of the mouth. There have been several case reports of lateral neck epidermoid cysts, which are considered to arise as median cysts and extend to the space between the mylohyoid and hyoglossus muscles. We report a rare case of epidermoid cyst arising between the submandibular gland and the platysma. A 40-year-old woman was referred to our hospital for evaluation of an asymptomatic swelling in the submandibular region. The MR imaging showed a well-defined cystic mass between the submandibular gland and the platysma. The lesion was extirpated under general anesthesia. No adhesion to the surrounding tissues was observed. White caseous material was contained in the lesion. Histopathologically, the cyst lumen was filled with keratin and lined by stratified squamous epithelium. There was no skin appendage, and a final diagnosis of epidermoid cyst was made. The postoperative course has been good, without any complications.

The histology and morphology characteristics of the tongue in Scincella tsinlingensis were studied by light and electronic microscopy. Under light microscopy, the tongue consists of tip, lingual body and radix in sequence. Numerous lingual papillae widely distribute on the surface of the dorsal and ventral flanks in the tongue, in addition to some regions of the tip. The papillae's surface is covered with the epithelial layer. The lamina propria and dense connective tissue are distinct existing under the epithelial layer. There are many lingual glands spread over the lamina propria. Tongue muscle is developed and composed of distinct intrinsic muscle, hyoglossus and genioglossus. By scanning electron microscopy, at higher magnification, the epithelial cells of the dorsal surface in the divaricate tongue tips show numerous microvilli, micro-ridges and micro-pores. The surface of dorsal side of the papillae in lingual body is covered with abundant of micro-ridges and taste bud lacuna. On the surface of the papillae in radix, micro-facets and micro-ridges are compactly distributed, as well as scattered mucilage-pores. The lingual epithelium is divided into four layers observed by the transmission electron microscope. Cells of basal layer are irregularly elliptical in shape, with sparse organelles in the cytoplasm. The deep intermediate layer is not always distinct. Small numbers of organelles are scattered into the cytoplasm. The cells of the superficial intermediate layer gradually flatten, as do their nuclei. The cytoplasm contains many keratohyalin granules. Cell membranes are formed processes around cells and joined by abundant desmosomes to the cell membranes of adjacent cells. The cells located on the extreme free-surface side of the keratinized layer have fallen off. The basal lamina is intercalated between the basal layer and the lamina propria. The lamina propria of lingual body contains lingual gland. A large part of the cytoplasm is occupied by mucus granules which located in the distal part of the cell. The connective tissue contains myelinated nerve fibers, vessel and muscle cells.

Background Neurostimulation of the hypoglossal nerve has shown promising results in the treatment of obstructive sleep apnea. This anatomic study describes the detailed topography of the hypoglossal nerve's motor points as a premise for super-selective neurostimulation in order to optimize results and minimize the risk of complications related to main nerve trunk manipulation. Methods Thirty cadaveric hypoglossal nerves were dissected and characterized by number of branches, arborization pattern, and terminal branch motor point location. For each motor point, the distance to cervical midline (x axis), distance to posterior aspect of the symphysis (y axis), and depth from the plane formed by the inferior border of symphysis and anterior border of hyoid (z axis) were recorded. Results The average number of distal branches for each hypoglossal nerve was found to be 9.95 ± 2.28. The average number of branches per muscle was found to be 3.3 ± 1.5 for the hyoglossus muscle, 1.8 ± 0.9 for the geniohyoid muscle, and 5.0 ± 1.6 for the genioglossus muscle. It was found that branches to the genioglossus and geniohyoid muscles were located closer to midline (relative lengths of 0.19 ± 0.07 and 0.19 ± 0.05, respectively) while hyoglossus branches were located more laterally (0.38 ± 0.10 relative length). On the y-axis, the branches to the genioglossus were the most anterior and therefore closest to the posterior symphysis of the mandible (relative length of 0.48 ± 0.11), followed by the geniohyoid (0.66 ± 0.09), and the hyoglossus (0.76 ± 0.16). The branches to the geniohyoid were the most superficial (relative length of 0.26 ± 0.06), followed by the genioglossus (0.36 ± 0.09), and finally, the hyoglossus branches (0.47 ± 0.11), which were located deeply. Conclusion A topographical map of the hypoglossal nerve terminal motor points was successfully created and could provide a framework for the optimization of the neurostimulation techniques.

Tongue muscles are difficult to study by dissection and imaging methods because of the intermeshing of the muscular fibers. The study of the architecture of the tongue was based on 853 tongue sections of a fetus aged 32 weeks after conception. The analysis of the sections allowed demonstration of the different intrinsic and extrinsic muscles of the tongue, to determine their situation from the palatoglossus arch to the apex of the tongue. The tongue muscles are organized in different layers from cranial to caudal and from lateral to medial, one medial for the genioglossus muscle, one paramedial for the hyoglossus, the styloglossus and the inferior longitudinal muscles. In the anterior third of the tongue, the muscles presented a concentric organization.

The purpose of this study was to examine the influence of reduced tongue activity by artificial rearing on the morphology of motoneurons innervating the extrinsic tongue retrusors. Artificially reared rat pups were fed via gastric cannula from postnatal day 3 to postnatal day 14. Artificially reared animals and dam-reared controls had cholera toxin (subunit B) conjugate of horseradish peroxidase injected into the styloglossus to label motoneurons innervating hyoglossus and styloglossus on postnatal day 13 and postnatal day 59, artificially reared animals and dam-reared controls. Following perfusion on postnatal days 14 and 60, serial transverse sections treated with tetramethyl benzidine and counter-stained neutral red were used to analyze motoneuron morphology. The shorter diameter of hyoglossus motoneurons increased with age for the dam-reared but not the artificially reared group. There was a tendency for a similar pattern for styloglossus motoneurons across the two rearing groups. The changes in form factor reflected the changes in shorter diameter for both motoneuron pools. Therefore, reducing suckling activity during normal postnatal development leads to diminished motoneuron somal growth in rats. This may also be the case in premature infants necessarily fed artificially.

Rationale: Historically, respiratory-related research in sleep apnea has focused exclusively on the extrinsic tongue muscles (i.e., genioglossus, hyoglossus, and styloglossus). Until recently, the respiratory control and function of intrinsic tongue muscles (i.e., inferior and superior longitudinalis, transverses, and verticalis), which comprise the bulk of the tongue, were unknown. Objectives: The current study sought to determine if extrinsic and intrinsic tongue muscles are coactivated in conditions of hypoxemia comparable to that experienced by adults with obstructive sleep apnea. Measurements: Esophageal pressure and EMG activity of an extrinsic (hyoglossus) and an intrinsic (superior longitudinal) tongue muscle were studied in anesthetized, tracheotomized, spontaneously breathing rats. Average EMG activity was compared in a control gas condition (PaO2, 160 ± 12 mm Hg) and in mild isocapnic hypoxia (PaO2, 69 ± 7.2 mm Hg), with and without brief (3-breath) airway occlusions, pre- and postbilateral vagotomy. Main Results: (1) intrinsic and extrinsic tongue muscles are coactivated in mild hypoxia, (2) airway occlusion increased the activities of intrinsic retractor muscles in mild hypoxia, and (3) extrinsic retractor muscles have a steeper rate of rise of activity and an earlier burst onset relative to intrinsic retractor activities in mild hypoxia. Conclusions: These findings support our working hypothesis that airway patency is maintained not simply by activation of extrinsic tongue muscles but by the coactivation of intrinsic and extrinsic protrudor and retractor muscles.

Gross and microscopic examination of the tongue and hyolingual apparatus of the sperm whale (Physeter macrocephalus) revealed numerous dis-tinct differences from those of other toothed whales and dolphins, largely reflecting the tongue’s atypi-cal position, relations, and size, and its primary role in suction ingestion, rather than prey prehen-sion or transport, as in many other odontocetes. Unlike other odontocetes, the sperm whale has a short, wide tongue that is uniquely situated at the rear of the open oral cavity. Since the tongue does not extend to the tooth row, which runs along the elongated median mandibular symphysis, it cannot easily reorient grasped prey items, yet it can posi-tion them to be swallowed or sucked directly into the oropharyngeal opening. The scarcity of intrin-sic lingual musculature (m. lingualis proprius), coupled with the relatively large paired extrinsic muscles inserting in the tongue—notably the m. hyoglossus, whose profuse fibers comprise much of the tongue root, and the m. genioglossus—suggests the tongue mainly undergoes positional, rather than shape, changes as it is retracted by the hyoid to generate negative intraoral pressures to capture and ingest prey items via suction. The tongue possesses numerous longitudinal folds or plicae, but almost no free tip; its slightly convex dorsum bears deep fissures and few sensory receptors in a multilayered and predominantly aglandular horny epithelium.Key Words: sperm whale, Physeter macrocepha-lus, tongue, hyoid, myology, morphology, suction feeding

The muscular-hydrostat model of tongue function proposes a constant interaction of extrinsic (external bony attachment, insertion into base of tongue) and intrinsic (origin and insertion within the tongue) tongue muscles in all tongue movements (Kier WM and Smith KK. Zool J Linn Soc 83: 207-324, 1985). Yet, research that examines the respiratory-related effects of tongue function in mammals continues to focus almost exclusively on the respiratory control and function of the extrinsic tongue protrusor muscle, the genioglossus muscle. The respiratory control and function of the intrinsic tongue muscles are unknown. Our purpose was to determine whether intrinsic tongue muscles have a respiration-related activity pattern and whether intrinsic tongue muscles are coactivated with extrinsic tongue muscles in response to respiratory-related sensory stimuli. Esophageal pressure and electromyographic (EMG) activity of an extrinsic tongue muscle (hyoglossus), an intrinsic tongue muscle (superior longitudinal), and an external intercostal muscle were studied in anesthetized, tracheotomized, spontaneously breathing rats. Mean inspiratory EMG activity was compared at five levels of inspired CO2. Intrinsic tongue muscles were often quiescent during eupnea but active during hypercapnia, whereas extrinsic tongue muscles were active in both eupnea and hypercapnia. During hypercapnia, the activities of the airway muscles were largely coincident, although the onset of extrinsic muscle activity generally preceded the onset of intrinsic muscle activation. Our findings provide evidence, in an in vivo rodent preparation, of respiratory modulation of motoneurons supplying intrinsic tongue muscles. Distinctions noted between intrinsic and extrinsic activities could be due to differences in motoneuron properties or the central, respiration-related control of each motoneuron population.

Laboratory rats were acutely implanted with an electrode array composed of eight independently controllable contacts applied to ventral and dorsal aspects of the left and right hypoglossal nerves (HGNs) and their branches. Bipolar intramuscular electromyographic (EMG) electrodes were implanted into the left and right genioglossus, hyoglossus and styloglossus muscles to identify which muscles were activated during stimulation via the contacts. Elicited movements, including changes in the position of the tongue and in the size and the shape of the airway, were documented video-graphically through a surgery microscope and an endoscope. Constant current electrical stimulation activated various combinations of electrode contacts and the stimulation patterns were correlated with corresponding oral movements, airway sizes, and EMG activities. Results demonstrate that graded responses and differential activation of the various tongue muscles are achievable by stimulation of specific contacts in the electrode array. These effects are interpreted to result from the targeted activation of regions of the nerve lying under and between the electrodes. Further testing established that the muscle responses elicited by unilateral electrical stimulation with the present approach can be smoothly graded, that the muscle responses resulted in opening of the airway and could be reliably maintained for long durations.

Obstructive sleep apnea (OSA) patients have increased upper airway muscle activity, including such lingual muscles as the genioglossus (GG), geniohyoid (GH), and hyoglossus (HG). This adaptation partially protects their upper airway against obstructions. Rodents are used to study the central neural control of sleep and breathing but they do not naturally exhibit OSA. We investigated whether, in chronically instrumented, behaving rats, disconnecting the GH and HG muscles from the hyoid (H) apparatus would result in a compensatory increase of other upper airway muscle activity (electromyogram, EMG) and/or other signs of upper airway instability. We first determined that, in intact rats, lingual (GG and intrinsic) muscles maintained stable activity levels when quantified based on 2 h-long recordings conducted on days 6 through 22 after instrumentation. We then studied five rats in which the tendons connecting the GH and HG muscles to the H apparatus were experimentally severed. When quantified across all recording days, lingual EMG during slow-wave sleep (SWS) was modestly but significantly increased in rats with surgically altered upper airway [8.6 ± 0.7% (SE) vs. 6.1 ± 0.7% of the mean during wakefulness; p = 0.012]. Respiratory modulation of lingual EMG occurred mainly during SWS and was similarly infrequent in both groups, and the incidence of sighs and central apneas also was similar. Thus, a weakened action of selected lingual muscles did not produce sleep-disordered breathing but resulted in a relatively elevated activity in other lingual muscles during SWS. These results encourage more extensive surgical manipulations with the aim to obtain a rodent model with collapsible upper airway.

International audience; Agamid lizards use tongue prehension for capturing all types of prey. The purpose of this study was to investigate the functional relationship between tongue structure, both surface and musculature, and function during prey capture in Pogona vitticeps. The lack of a detailed description of the distribution of fibre-types in the tongue muscles in some iguanian lizards has hindered the understanding of the functional morphology of the lizard tongue. Three methodological approaches were used to fill this gap. First, morphological analyses were performed (i) on the tongue surface through scanning electron microscopy, and (ii) on the lingual muscle by histological coloration and histochemistry to identify fibre-typing. Secondly, kinematics of prey capture was quantified by using high-speed video recordings to determine the movement capabilities of the tongue. Finally, electromyography (EMG) was used to identify the motor pattern tongue muscles during prey capture. Morphological and functional data were combined to discuss the functional morphology of the tongue in agamid lizards, in relation to their diet. During tongue protraction, M. genioglossus contracts 420 ± 96 ms before tongue-prey contact. Subsequently, Mm. verticalis and hyoglossus contract throughout tongue protraction and retraction. Significant differences are found between the timing of activity of the protractor muscles between omnivorous agamids (Pogona sp., this study) and insectivorous species (Agama sp.), despite similar tongue and jaw kinematics. The data confirm that specialisation toward a diet which includes more vegetal materials is associated with significant changes in tongue morphology and function. Histoenzymology demonstrates that protractor and retractor muscles differ in fibre composition. The proportion of fast glycolytic fibres is significantly higher in the M. hyoglossus (retractor muscle) than in the M. genioglossus (protractor muscle), and this difference is proposed to be associated with differences in the velocity of tongue protrusion and retraction (5 ± 5 and 40 ± 13 cm s(-1) , respectively), similar to Chamaeleonidae. This study provides a way to compare fibre-types and composition in all iguanian and scleroglossan lizards that use tongue prehension to catch prey.

This anatomical investigation in the rat was designed to illustrate the detailed organization of the tongue's muscles and their innervation in order to elucidate the actions of the muscles of the higher mammalian tongue and thereby clarify the protrusor subdivision of the hypoglossal-tongue complex. The hypoglossal innervated, extrinsic styloglossus, hyoglossus, and genioglossus and the intrinsic transversus, verticalis and longitudinalis linguae muscles were observed by microdissection and analysis of serial transverse-sections of the tongue. Sihler's staining technique was applied to whole rat tongues to demonstrate the hypoglossal nerve branching patterns. Dissections of the tongue demonstrate the angles at which the extrinsic muscles act on the base of the tongue. The Sihler stained hypoglossal nerves demonstrate branches to the styloglossus and hyoglossus emanating from its lateral division while branches to the genioglossus muscle exit from its medial division. The largest portions of both XIIth nerve divisions can be seen to enter the body of the tongue to innervate the intrinsic muscles. Transverse sections of the tongue demonstrate the organization of the intrinsic muscle fibers of the tongue. Longitudinal muscle fibers run along the entire circumference of the tongue. Alternating sheets of transverse lingual and vertical lingual muscles can be observed to insert into the circumference of the tongue. Most importantly in clarifying tongue protrusion, we demonstrate the transversus muscle fibers enveloping the most superior and inferior portions of the longitudinalis muscles. Longitudinal muscle fascicles are completely encircled and thus are likely to be compressed by transverse muscle fascicles resulting in elongation of the tongue. We discuss our findings in relation to biomechanical studies, that describe the tongue as a muscular hydrostat and thereby define the "elongation-protrusion apparatus" of the mammalian tongue. In so doing, we clarify the functional organization of the hypoglossal-tongue complex.

The rhombencephalic neural crest play several roles in craniofacial development. They give rise to the cranial sensory ganglia and much of the craniofacial skeleton, and are vital for patterning of the craniofacial muscles. The loss of Hoxa1 or Hoxa2 function affects the development of multiple neural crest-derived structures. To understand how these two genes function together in craniofacial development, an allele was generated that disrupts both of these linked genes. Some of the craniofacial defects observed in the double mutants were additive combinations of those that exist in each of the single mutants, indicating that each gene functions independently in the formation of these structures. Other defects were found only in the double mutants demonstrating overlapping or synergistic functions. We also uncovered multiple defects in the attachments and trajectories of the extrinsic tongue and hyoid muscles in Hoxa2 mutants. Interestingly, the abnormal trajectory of two of these muscles, the styloglossus and the stylohyoideus, blocked the attachment of the hyoglossus to the greater horn of the hyoid, which in turn correlated exactly with the presence of cleft palate in Hoxa2 mutants. We suggest that the hyoglossus, whose function is to depress the lateral edges of the tongue, when unable to make its proper attachment to the greater horn of the hyoid, forces the tongue to adopt an abnormal posture which blocks closure of the palatal shelves. Unexpectedly, in Hoxa1/Hoxa2 double mutants, the penetrance of cleft palate is dramatically reduced. We show that two compensatory defects, associated with the loss of Hoxa1 function, restore normal attachment of the hyoglossus to the greater horn thereby allowing the palatal shelves to lift and fuse above the flattened tongue.

This anatomical investigation was prompted by the incomplete knowledge of the myotopic organization of the dorsal subdivison of the hypoglossal nucleus. Intrinsic muscle motoneurons were not segregated and labeled previously with regard to the lateral division of the hypoglossal nerve. Also, motoneuron number and cell size, in relation to the individual retrusor tongue musculature, were rarely addressed previously. Retrograde labeling of retrusor muscle motoneurons in the dorsal subdivision of the rat hypoglossal nucleus was done. Cholera toxin conjugate horseradish peroxidase (CTHRP) was injected into the retrusor tongue muscles with only the lateral division of the hypoglossal nerve intact. The dorsal subdivision of the hypoglossal nucleus contained approximately 800 motoneurons ranging in cell body size from 19 to 41 μm. When either the styloglossus, hyoglossus, superior longitudinal, or inferior longitudinal muscle was isolated and injected with CTHRP, a separate motoneuron pool for each muscle was seen. The extrinsic muscle motoneurons, styloglossus and hyoglossus, were found rostrolateral and caudolateral respectively. In contrast, the intrinsic superior and inferior longitudinal muscle motoneurons were found more central and medial in the nucleus. Extrinsic muscle motoneurons were larger (≈30 μm) than intrinsic muscle motoneurons (≈26 μm; P < .0001). Intrinsic muscle motoneurons account for a great majority of the motoneurons in the dorsal aspect of the hypoglossal nucleus and their axons have been shown to be contained in the lateral (retrusor) division of the hypoglossal nerve. This study revealed the myotopic organization of the retrusor subdivision of the rat hypoglossal nucleus. Anat Rec 254:222–230, 1999. © 1999 Wiley-Liss, Inc.

"#$%!&$ The movements of the tongue, hyoid, and jaw were recorded cineradiographically in preweaning pigs as they suckled bariumized milk from a veterinary teat or drank it from a bowl. The movements were quantified by measuring the X, Y coordinates of radioopaque markers embedded in the tongue and attached to both jaws and to the hyoid. EMG activity in masseter, anterior digastric, geniohyoid, genioglossus, hyoglossus, sternohyoid, stylohyoid, and omohyoid muscles was recorded synchronously with cineradiography at 100 frames/sec. In both suckling and drinking, the movements were characterized by minimal movements of the jaw and hyoid but extensive movements of the tongue. In suckling, the movements were largely confined to the mid- posterior part of the tongue. A seal was formed between the posterior tongue and soft palate while a depression formed in the mid-tongue; this was associated with fluid moving into the depression probably because of a reduced intraoral pressure. The depression was associated with increased EMG activity in the genioglossus muscle and overlapping activity in digastric, geniohyoid, hyoglos- sus, and sternohyoid muscles. In drinking cycles, significant movement occurred in all parts of the tongue; milk ingestion was associated with tongue movements that combined elements character- istic both of suckling (mid-tongue depression with a posterior seal) and of lapping (extensive antero- posterior movements within the tongue itself). In drinking, compared to suckling, there was a major reduction in EMG activity in masseter, digastric, geniohyoid, and sternohyoid muscles. Af- ter milk had accumulated in the valleculae, swallows usually occurred in every other cycle during suckling and in every third or fourth cycle during drinking. The emptying of the valleculae was an event that was embedded in the early jaw-opening phase of an otherwise normal suckling or drinking cycle. Emptying of the valleculae was associated with posteriorly directed movement of the back of the tongue and increased EMG activity in hyoglossus, styloglossus, and omohyoid muscles. No differences were noted in the kinematics associated with swallowing in the two activi- ties, but, in the normalized and averaged EMG data, there were significant differences in the timing of genioglossus activity and in the relative balance of hyoglossal and stylohyoid activity. '( )*+(,-../(,01234 © 1998 Wiley-Liss, Inc.

Physiological articulatory models have evolved from simpler forms to complex ones, while recent models preserve traits of oversimplification and anatomical unreality. This work combines MRI observations at ATR-BAIC and Johns Hopkins University to point to the issue for advancing extrinsic tongue muscle modeling. The genioglossus, previously thought to arise from the genial tubercle of the mandible, has direct fiber attachments on the short tendon of the tongue. The posterior genioglossus, being regarded as a functional division of the muscle, can be given anatomical definition to the horizontal bundle arising from the inferior aspect of the short tendon. The styloglossus has been modeled as linear strings traveling 'free in air' before inserting into the tongue, but the extralingual part is actually restrained by the surrounding soft tissues to lack mobility. The intralingual styloglossus forms anterior and posterior slings in the tongue tissue, possibly with the distal fibers of the hyoglossus. Combined styloglossus and hyoglossus shortening via the slings may be a factor shaping the tongue into various forms.

Summary The distribution in the chicken of motoneurons innervating the hyolingual muscles, i.e. the Mm. hyoglossus rostralis (HR), hyoglossus obliquus (HO), ceratoglossus (CG), interceratobranchialis (CB). stylohyoideus (YH), serpihyoideus (PH) and cricohyoideus (CR), and the laryngotracheal muscles, comprising the Mm. tracheolateralis (TL), cleido-hyoideus (CL) and sternotrachealis (ST), was examined by retrograde transport of horseradish peroxidase conjugated with wheat-germ agglutinin. Labelled motoneurons are only found in the hypoglossal nucleus. The rostrocaudal distributions of motoneurons projecting to hyolingual muscles are restricted in the hypoglossal nucleus cranial to the obex, and those projecting to laryngotracheal muscles are distributed in the more caudal part of hypoglossal nucleus. Detailed analysis of the data showed that the most rostrally positioned motoneurons in the hypoglossal nucleus supplied to the PH, followed by the CG, CB. HR, YH, HO, CR, TL, CL and ST in that order, overlapping each other. In the hypoglossal nucleus motoneurons innervating the PH and YH have the smallest perikarya. Of the motoneurons in the hypoglossal nucleus, those supplying the laryngotracheal muscles (CL and TL) have the largest perikarya. Motoneurons innervating the other muscles are intermediate in size.

The influence of lingual muscle activity on airflow dynamics in the upper airway was examined in nine patients with obstructive sleep apnea. Muscles that retract the tongue (hyoglossus and styloglossus) and protrude the tongue (genioglossus) were selectively stimulated electrically during sleep with fine wire electrodes placed intramuscularly transorally. We confirmed that stimulation with 50 Hz and 40-microseconds pulse duration did not elicit changes in electroencephalographic patterns or heart rate or alter airflow after the stimulation burst had ceased. The highest stimulus intensity that did not arouse patients from sleep was then utilized to examine the effect of lingual muscle recruitment on airflow dynamics during steady-state periods of inspiratory airflow limitation. When applying a stimulus burst during single inspirations, maximal inspiratory airflow decreased by 239 +/- 177 ml/s (P < 0.05) during retractor stimulation, whereas maximal inspiratory airflow increased by 217 +/- 93 ml/s during protrusor stimulation (P < 0.001) compared with breaths immediately before and after the stimulated breath. When consecutive inspirations were stimulated repeatedly, protrusor stimulation decreased the frequency of obstructive breathing episodes in four patients breathing at 3.9 +/- 3.4 (SD) cmH2O nasal pressure. The findings suggest that stimulation of the lingual muscles can increase or decrease airflow depending on the specific muscles stimulated without arousing patients from sleep.

Surgical techniques were used to isolate the extrinsic bellies of the styloglossus and hyoglossus muscles from the body of the tongue for cholera toxin HRP injection. An average of 53 styloglossus and 121 hyoglossus motoneurons in the dorsal subdivision of the hypoglossal nucleus were demonstrated using tetramethyl benzidine histochemistry. Styloglossus motoneurons were restricted to the rostral 25% of the nucleus while hyoglossus motoneurons occupied other regions of the dorsal nucleus.