Anatomy of the lingual nerve: Application to oral surgery

The purpose of this research is to obtain morphological information about the traveling route, branching pattern, and distribution within the tongue of the lingual nerve, all of which are important for oral surgical procedures. Using 20 sides from 10 Japanese cadaveric heads, we followed the lingual nerve from its merging point with the chorda tympani to its peripheral terminal in the tongue. We focused on the collateral branches in the area before reaching the tongue and the communication between the lingual and hypoglossal nerves reaching the tongue. The collateral branches of the lingual nerve were distributed in the oral mucosa between the palatoglossal arch and the mandibular molar region. Two to eight collateral branches arose from the main trunk of the nerve, and the configuration of branching was classified into three types. More distally, the lingual nerve started to communicate with the hypoglossal nerve before passing the anterior border of the hyoglossus muscle. Nerve communications were also found in the main body and near the apex of the tongue. A thorough understanding of the collateral branches near the tongue, and the communication with the hypoglossal nerve inside the tongue, will help to prevent functional disorders from local anesthesia and oral surgical procedures associated with the lingual nerve. Clin. Anat. 32:635–641, 2019. © 2019 Wiley Periodicals, Inc.     

Main trajectories of nerves that traverse and surround the tympanic cavity in the rat

To guide surgery of nerves that traverse and surround the tympanic cavity in the rat, anatomical illustrations are required that are topographically correct. In this study, maps of this area are presented, extending from the superior cervical ganglion to the otic ganglion. They were derived from observations that were made during dissections using a ventral approach. Major blood vessels, bones, transected muscles of the tongue and neck and supra and infrahyoid muscles serve as landmarks in the illustrations. The course of the mandibular, facial, glossopharyngeal, vagus, accessory and hypoglossal nerves with their branches, and components of the sympathetic system, are shown and discussed with reference to data available in the literature. Discrepancies in this literature can be clarified and new data are presented on the trajectories of several nerves. The course of the tympanic nerve was established. This nerve originates from the glossopharyngeal nerve, enters the tympanic cavity, crosses the promontory, passes the tensor tympani muscle dorsally, and continues its route intracranially to the otic ganglion as the lesser petrosal nerve after intersecting with the greater petrosal nerve. Auricular branches of the glossopharyngeal and of the vagus nerve were noted. We also observed a pterygopalatine branch of the internal carotid nerve, that penetrates the tympanic cavity and courses across the promontory.     

Discussion of clinical anatomy of the lingual nerves

The tongue has various functions, such as gustation, pronunciation, mastication, and deglutition. The nerve fibers are complexly intermingled, and communications between the lingual nerve and the hypoglossal nerve have been reported. Fifteen Japanese heads (30 sides) donated to the 2nd-year students dissection course at the Nippon Dental University School of Life Dentistry, Niigata, were studied with regard to the following aspects: 1. relation of the bifurcation between the lingual and the inferior alveolar nerves close to the oval foramen; 2. distance between the oral foramen and the bifurcation of the lingual and inferior alveolar nerves; and 3. communication between the lingual and hypoglossal nerves. Three types of bifurcation were observed. The standard bifurcation was observed in 21 cases (70.1%). A high bifurcation was observed in 5 cases (16.6%), and a communicating bifurcation was observed in 4 cases (13.3%). The average distance between the oval foramen and the bifurcation of the lingual and inferior alveolar nerves was 8.7 +/- 4.2 mm (minimum: 0 mm/maximum: 14 mm). An anterior-type communication between the lingual and hypoglossal nerves was observed in 8 cases (26.6%), a posterior-type was observed in 17 cases (56.7%), and no communication was observed in 5 cases (17.7%).     

Distribution pattern of the human lingual nerve

The tongue is an intricate organ with many functions. Despite the knowledge of the presence of muscular and neural connections in the tongue, a detailed neuroanatomical depiction of the nerves' topography in the tongue has not been demonstrated. The topography, branching patterns and neuronal interconnections of the lingual nerve were studied in five postmortem human tongues. They were stained with Sihler's stain, a technique that renders most of the tongue tissue translucent while counterstaining nerves. The lingual nerve reaches the tongue posterolaterally. There are two main branches off of the main trunk: the medial branch sends 2-4 small branches to the medial part of the ventrolateral tongue and the lateral branch runs along the lateral tongue border and sends 3-4 large branches to the anterior tip of tongue. Each subdivision gives off 2-5 distal branches. Both medial and lateral branches have interconnections with the proximal part of the hypoglossal nerve. One of the unexpected discoveries in this study was the high density of nervous fibers in the lateral aspect of the tongue as compared to the midline region. The average diameter of the main trunk of the lingual nerve is 3.5 mm. The medial and lateral branches average 1 mm in diameter, the more distal subdivisions measure 0.5-0.75 mm, and the lingual-hypoglossal interconnections measure 0.125-0.250 mm. In summary, this study provides the first detailed depiction of the topography of the human lingual nerve and its branches in situ, confirmation of lingual-hypoglossal nerve connection, and the first depiction of the high density of lingual nerve innervation in the lateral tongue.     

Two distinct projection areas from tongue nerves in the frontal operculum of macaque monkeys as revealed with evoked potential mapping

Projection areas in the cerebral cortex from tongue nerves were investigated in anesthetized macaque monkeys (Macaca fuscata). Three tongue nerves (lingual nerve, chorda tympani, and a lingual branch of the glossopharyngeal nerve) were electrically stimulated and thereby evoked mass field potentials were recorded with microelectrodes roving through the exposed and buried frontal operculum. Two distinct tongue nerve projection areas were thus located; one in the lower part of the precentral gyrus ventral to the inferior precentral sulcus, and the other in the medial part of the buried frontal operculum. The former corresponds to the "cortical masticatory area", previously defined in macaque monkeys as involved in somatic sensation as well as mastication. However, the prominent projection from the chorda tympani and glossopharyngeal nerve branch, containing taste fibers, to this area suggests its involvement in taste sensation. The latter corresponds to the "pure gustatory area" defined in squirrel monkeys in both location and cytoarchitecture. However, projection to this area from the lingual nerve, which contains only somatic nerve fibers, suggests its involvement in somatic sensation, as well.     

The origins of the afferent fibers to the lingual muscles of the dog,a retrograde labeling study with horseradish peroxidase

The origin of the afferent fibers to the lingual muscles of the dog was investigated by means of retrograde transport of horseradish peroxidase (HRP) from injection sites in the tongue and the extrinsic lingual muscles. Intralingual injections were not satisfactory because the enzyme diffused beyond the intrinsic lingual muscles to include virtually all tissues within the tongue. Thus, the resultant retrograde labeling of cell bodies of the trigeminal, geniculate, glossopharyngeal, vagal, and first cervical (C₁) spinal ganglia represented a composite of lingual sensory innervation. In order to confine HRP uptake to intramuscular nerve endings, injections were limited to surgically isolated extrinsic lingual muscles, i.e., the genioglossus, hyoglossus, and styloglossus. After these intramuscular injections, labeled neurons appeared ipsilaterally in the C₁ spinal ganglion, the proximal vagal (jugular) ganglion, and trigeminal ganglion. Earlier suggestions that the lingual proprioceptive neurons of the dog reside in the distal vagal (nodose) ganglion and hypoglossal ganglia were not confirmed. The mesencephalic nucleus of the trigeminal nerve failed to label after enzyme injections into the tongue or the extrinsic lingual muscles. The retrograde labeling of cell bodies in the C₁ spinal ganglion was abolished when HRP injections into the extrinsic lingual muscles were preceded by surgical interruption of the ansa cervicalis or distal section of the hypoglossal nerve. Retrograde labeling of neurons in the proximal vagal ganglion persisted after hypoglossal nerve transections.     

The origins of the afferent fibers to the lingual muscles of the dog, a retrograde labelling study with horseradish peroxidase

The origin of the afferent fibers to the lingual muscles of the dog was investigated by means of retrograde transport of horseradish peroxidase (HRP) from injection sites in the tongue and the extrinsic lingual muscles. Intralingual injections were not satisfactory because the enzyme diffused beyond the intrinsic lingual muscles to include virtually all tissues within the tongue. Thus, the resultant retrograde labeling of cell bodies of the trigeminal, geniculate, glossopharyngeal, vagal, and first cervical (C1) spinal ganglia represented a composite of lingual sensory innervation. In order to confine HRP uptake to intramuscular nerve endings, injections were limited to surgically isolated extrinsic lingual muscles, i.e., the genioglossus, hyoglossus, and styloglossus. After these intramuscular injections, labeled neurons appeared ipsilaterally in the C1 spinal ganglion, the proximal vagal (jugular) ganglion, and trigeminal ganglion. Earlier suggestions that the lingual proprioceptive neurons of the dog reside in the distal vagal (nodose) ganglion and hypoglossal ganglia were not confirmed. The mesencephalic nucleus of the trigeminal nerve failed to label after enzyme injections into the tongue or the extrinsic lingual muscles. The retrograde labeling of cell bodies in the C1 spinal ganglion was abolished when HRP injections into the extrinsic lingual muscles were preceded by surgical interruption of the ansa cervicalis or distal section of the hypoglossal nerve. Retrograde labeling of neurons in the proximal vagal ganglion persisted after hypoglossal nerve transections.     

Reflex control of cat extrinsic and intrinsic tongue muscles exerted by intraoral receptors

The effects of mechanical and noxious stimulation of the palatal and lingual surfaces on the activity of the extrinsic and intrinsic tongue muscles have been studied in cats. Stimulation of the hard palate produced mainly activation of extrinsic tongue muscles while inhibition was elicited by stimulating the soft palate. Longlasting pressure on the hard palate caused rhythmic tongue flapping by intermittent genioglossal activity. The intrinsic tongue muscles, m. transversus and m. verticalis, were activated by noxious stimuli applied to the hard palate, the effects apparently being mediated by high-threshold afferents. Mechanical and noxious stimulation applied to the dorsal and ventral lingual surfaces of the tongue either activated or inhibited the extrinsic tongue muscles depending on the reflex area stimulated. The intrinsic tongue muscles were activated by noxious stimuli applied to the tongue surfaces. The anastomoses running between the hypoglossal and lingual nerves were found to mediate mainly nociceptive afferent impulses travelling from the hypoglossal to the lingual nerve. The intrinsic muscles were found to be controlled by anastomosal nociceptive afferents.     

Neural circuits underlying tongue movements for the prey-catching behavior in frog: distribution of primary afferent terminals on motoneurons supplying the tongue

The hypoglossal motor nucleus is one of the efferent components of the neural network underlying the tongue prehension behavior of Ranid frogs. Although the appropriate pattern of the motor activity is determined by motor pattern generators, sensory inputs can modify the ongoing motor execution. Combination of fluorescent tracers were applied to investigate whether there are direct contacts between the afferent fibers of the trigeminal, facial, vestibular, glossopharyngeal-vagal, hypoglossal, second cervical spinal nerves and the hypoglossal motoneurons. Using confocal laser scanning microscope, we detected different number of close contacts from various sensory fibers, which were distributed unequally between the motoneurons innervating the protractor, retractor and inner muscles of the tongue. Based on the highest number of contacts and their closest location to the perikaryon, the glossopharyngeal–vagal nerves can exert the strongest effect on hypoglossal motoneurons and in agreement with earlier physiological results, they influence the protraction of the tongue. The second largest number of close appositions was provided by the hypoglossal and second cervical spinal afferents and they were located mostly on the proximal and middle parts of the dendrites of retractor motoneurons. Due to their small number and distal location, the trigeminal and vestibular terminals seem to have minor effects on direct activation of the hypoglossal motoneurons. We concluded that direct contacts between primary afferent terminals and hypoglossal motoneurons provide one of the possible morphological substrates of very quick feedback and feedforward modulation of the motor program during various stages of prey-catching behavior.     

Morphological Features of the Branching Pattern of the Hypoglossal Nerve

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.     

Morphologic studies of the venous drainage of the tongue

Summary The aim of this work was to study the role of the venous drainage of the tongue in tongue inspection in traditional Chinese medicine and tongue-flap surgery. Thirty-two adult cadavers were observed, including 7 venous corrosivecast specimens. The decreasing order of venous drainage of the tongue, based on the diameter of the vein and size of its drainage area, was the accompanying v. of the hypoglossal n., the epiglottic valleculate v., the accompanying v. of lingual n., the lingual root v. and the accompanying v. of the lingual a. The veins constituting the picture of the tongue seen in sublingual collateral inspection were the companion vv. of the hypoglossal and lingual nn. The pedicle of a tongue flap must maintain efficient venous drainage canal.This work was supported by the National Natural Science Fund     

舌咽神经、迷走神经和副神经断层解剖与MRI

目的：探讨舌咽神经、迷走神经和副神经的走行和毗邻关系,为影像学诊断及临床开展该区手术提供形态学依据。方法：利用36例成尸头部连续横断层标本和15例成尸头部连续冠状断层标本,并与10例志愿者的3D—CISS序列MR图像进行对照,观察舌咽神经、迷走神经和副神经在颅内的走行规律及其与周围结构的位置关系。结果：舌咽、迷走和副神经由上而下从延髓的橄榄后沟发出,跨过延池,穿颈静脉孔出颅。根据走行,可分其为延髓内段、脑池段和颈静脉孔段。在脑池段,舌咽神经走行在上方,迷走神经和副神经在下方且结合紧密;在颈静脉孔段,这3对脑神经及其与颈内静脉和颈内动脉的关系是：颈内动脉位于最前方,颈内静脉位于最外侧,舌咽神经走行在前内上方,有单独的硬脊膜包绕,迷走神经和副神经位于其后外下方,形成迷走、副神经复合体。结论：在标本的连续断面和对应的MR图像上能够清楚地显示舌咽、迷走和副神经的走行和毗邻关系。     

The anatomy of the intralingual neural interconnections

Purpose

The intrinsic lingual neural interconnections are overlooked. It was hypothesized that intralingual anatomically well defined anastomoses interconnect the somatic and autonomic neural systems of the tongue. It was thus aimed to evaluate the intralingual neural scaffold in human tongues.

Methods

Human tongue samples (ten adult and one pediatric) were microdissected (4.5 magnification).

Results

In the interstitium between the genioglossus and hyoglossus muscles, the branches of the lingual nerve (LN) and the medial trunk of the hypoglossal nerve (HN) had a layered disposition of the outer and inner side, respectively, of the lingual artery with its periarterial plexus. Anastomoses of these three distinctive neural suppliers of tongue were recorded, as also were those of the LN with the lateral trunk of the HN and the anastomoses between successive terminal branches of the LN. Successive ansae linguales were joining the LN branches and the medial trunk of the HN.

Conclusion

The intrinsic neural system of the tongue supports integrative functions and allows a better retrospective understanding of various experimental studies. The topographical pattern is useful for an accurate diagnosis of intralingual nerves on microscopic slides.     

Development of glossopharyngeal nerve branches in the early chick embryo with special reference to morphology of the Jacobson''s anastomosis

Summary The development of glossopharyngeal nerve branches was studied by an immunohistochemical technique which stains the whole nervous system in situ. Prior to the formation of the ramus (r.) lingualis IX, pre- and post-trematic branches developed just beneath the pharyngeal ectoderm. This mode of development resembled that of the chorda tympani. The post-trematic nerve seemed to be a precursor of the r. lingual. IX. In addition to the r. pharyngeus dorsalis IX, another branch, r. pharyng. posterior IX, appeared. Both these branches formed an anastomosis with the facial and vagus beneath the dorsal aorta. The term Jacobson's anastomosis seemed to be most suitable to refer to an anastomosis made up of these dorsal pharyngeal branches of cranial nerves VII, IX and X. The primary anastomosis between the facial and the glossopharyngeal nerves in the chick is only temporarily present and is comparable to the similar anastomosis in a shark in which the sympathetic system is not present in the cranial region.     

The three-dimensional architecture of the human styloglossus especially its posterior muscle bundles.

The arrangement of the lingual muscles in the interior of the human tongue, particularly the course of the posterior muscle bundles of the styloglossus, was studied by gross anatomical examination and SEM, and its relationship with tongue functions was considered. The styloglossus divided into anterior and posterior fiber bundles. The bilateral anterior fiber bundles ran anteriorly, and fused at the median region of the lower surface of the tongue, forming a large arched structure. The posterior bundles divided into 10 or more smaller bundles and entered the interior of the tongue. These muscle bundles passed through the spaces between the inferior longitudinal and genioglossus and inserted in the lingual septum, forming a small arched structure. These posterior muscle bundles passed near the midpoint between the central third and dorsal third of the line between the mental spine and the dorsal surface of the tongue in the upper half of the root of the tongue, showing a multilayer structure. In many of the areas in which posterior muscle bundles were distributed, muscle bundles in the tongue were arranged along the posterior muscle bundles of the styloglossus, glossopharyngeal bundles of the superior pharyngeal constrictor muscle, and transverse muscle of the tongue from the deep layer to the dorsal surface of the tongue.     

Parasympathetic postganglionic cells in the glossopharyngeal nerve trunk and their relationship to unmyelinated nerve fibers in the fungiform papillae of the frog

The glossopharyngeal nerve of the frog is made up of afferent nerve fibers and efferent, parasympathetic and sympathetic nerve fibers. The precise origin and course of the parasympathetic efferent nerve fibers in the fungiform papillae of the frog's tongue were investigated. We found the ganglionic cells in the lingual branch of the frog glossopharyngeal nerve. The surface of the ganglionic cell bodies was partly covered by synaptic endings that impinged upon it. Synaptic endings contained clear synaptic vesicles and large dense-cored vesicles. After cutting of the glossopharyngeal nerve proximal to the jugular ganglion, synaptic endings were found to show definite signs of degeneration. These findings led us to the conclusion that the ganglionic cells in the lingual branch of the glossopharyngeal nerve are the parasympathetic postganglionic cells. After cutting of the glossopharyngeal nerve distal to the jugular ganglion, some unmyelinated nerve fibers in the fungiform papillae and postganglionic cells in the lingual branch remained intact. These results strongly suggest that the origin of some of the unmyelinated nerve fibers is the parasympathetic postganglionic cell in the lingual branch.     

Functional anatomy of the hypoglossal innervated muscles of the rat tongue: a model for elongation and protrusion of the mammalian tongue

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.     

Peripheral facial nerve communications and their clinical implications

The facial nerve (CN VII) nerve follows a torturous and complex path from its emergence at the pontomedullary junction to its various destinations. It exhibits a highly variable and complicated branching pattern and forms communications with several other cranial nerves. The facial nerve forms most of these neural intercommunications with branches of all three divisions of the trigeminal nerve (CN V), including branches of the auriculotemporal, buccal, mental, lingual, infraorbital, zygomatic, and ophthalmic nerves. Furthermore, CN VII also communicates with branches of the vestibulocochlear nerve (CN VIII), glossopharyngeal nerve (CN IX), and vagus nerve (CN X) as well as with branches of the cervical plexus such as the great auricular, greater, and lesser occipital, and transverse cervical nerves. This review intends to explore the many communications between the facial nerve and other nerves along its course from the brainstem to its peripheral branches on the human face. Such connections may have importance during clinical examination and surgical procedures of the facial nerve. Knowledge of the anatomy of these neural connections may be particularly important in facial reconstructive surgery, neck dissection, and various nerve transfer procedures as well as for understanding the pathophysiology of various cranial, skull base, and neck disorders.     

Lingual mechanoreceptors activated by muscle twitch

1. Two groups of mechanoreceptors in the tongue have been identified by recording afferent discharges in single nerve fibres dissected from the lingual nerves of anaesthetized cats. The group of superficially situated, rapidly adapting mechanoreceptors had faster conduction velocities than the presumed deeply situated, slowly adapting endings.2. Stretch of the tongue musculature did not prove to be as efficient in activating the endings as local deformation, although it was possible to excite some of the presumed deeply situated endings in this way.3. No mechanoreceptor fibres could be identified in filaments dissected from the hypoglossal nerves in the same experiments.4. Twitches of the tongue musculature produced by stimulation of the hypoglossal nerve were able to cause discharge of superficial and presumed deeply situated mechanoreceptors during the tension change.5. The timing of the discharges in response to twitch was such that it could account for the delay in synaptic potentials produced in hypoglossal motoneurones when supramaximal stimuli were applied to the hypoglossal nerve.6. The possible significance of mechanoreceptor discharges in reflex activation of tongue motor units is discussed.     

Target pioneering and early morphology of the murine chorda tympani

Many studies demonstrate that differentiation of certain sensory receptors during development is induced by their nerve supply. Thus the navigational accuracy of pioneering fibres to their targets is crucial to this process. The special gustatory elements of the facial and glossopharyngeal nerves are used extensively as model systems in this field. We examined the chorda tympani, the gustatory component of the facial nerve, to determine the precise time course of its development in mice. The transganglionic fluorescent tracer DiI was injected into the anterior aspect of the mandibular arch of fixed embryos aged between 30 and 50 somites (E10–E12). It was allowed to diffuse retrogradely via the geniculate ganglion to the brainstem for 4 wk, before the distribution of DiI was determined using confocal laser scanning microscopy. Geniculate ganglion cells were first labelled at the 34 somite stage (E10). Pioneering chorda tympani fibres that arise from these cells passed peripherally and followed an oblique course as they grew towards the mandibular arch. At the 36 somite stage (E10.5), the peripheral component followed an intricate postspiracular course and passed anteriorly to arch over the primitive tympanic cavity, en route to the lingual epithelium. From the 36 to 50 somite stages (E10.5–E12), it consistently traced in the fashion of a ‘U’ bend. The central fascicle also traced at the 36 somite stage (E10.5) and just made contact with the brainstem. At the 40 somite stage (E11), the central fibres clearly chose a route of descent into the spinal trigeminal tract and branched into the solitary tract. Pioneering chorda tympani fibres contact the lingual epithelium when the target is primordial. The lingual epithelium may be a source of a neurotropic factor that attracts peripheral chorda tympani fibres to the sites of putative papillae. However, the chorda tympani is probably not a vital influence on the subsequent differentiation of gustatory papillae, since the papillae are elaborated 5 d later at E15 in murine embryos. The early morphology of the nerve is true to the amniote vertebrate phenotype.