On the maxillary nerve.

The trigeminal, the fifth cranial nerve of vertebrates, represents the rostralmost component of the nerves assigned to pharyngeal arches. It consists of the ophthalmic and maxillomandibular nerves, and in jawed vertebrates, the latter is further divided into two major branches dorsoventrally. Of these, the dorsal one is called the maxillary nerve because it predominantly innervates the upper jaw, as seen in the human anatomy. However, developmentally, the upper jaw is derived not only from the dorsal part of the mandibular arch, but also from the premandibular primordium: the medial nasal prominence rostral to the mandibular arch domain. The latter component forms the premaxillary region of the upper jaw in mammals. Thus, there is an apparent discrepancy between the morphological trigeminal innervation pattern and the developmental derivation of the gnathostome upper jaw. To reconcile this, we compared the embryonic developmental patterns of the trigeminal nerve in a variety of gnathostome species. With the exception of the diapsid species studied, we found that the maxillary nerve issues a branch (nasopalatine nerve in human) that innervates the medial nasal prominence derivatives. Because the trigeminal nerve in cyclostomes also possesses a similar branch, we conclude that the vertebrate maxillomandibular nerve primarily has had a premandibular branch as its dorsal element. The presence of this branch would thus represent the plesiomorphic condition for the gnathostomes, implying its secondary loss within some lineages. The branch for the maxillary process, more appropriately called the palatoquadrate component of the maxillary nerve (V(2)), represents the apomorphic gnathostome trait that has evolved in association with the acquisition of an upper jaw.

Accessory osseous passages in the vicinity of foramen rotundum in a dry adult human skull: morphological evaluation, embryological correlation and clinical reappraisal.

An unusual bony depression with accessory osseous passages in the right side of middle cranial fossa was observed in a dry adult human skull. It was located below the superior orbital fissure and optic canal. The bony depression was divided into two parts ,medial and lateral fossa, by an incomplete oblique bony septum. The foramen rotundum and an accessory bony canaliculus were present in the medial fossa whereas the lateral fossa was blind. The foramen rotundum led as usual into the pterygopalatine fossa . The presence of such an abnormal bony depression in the right middle cranial fossa with accessory osseous passages in the vicinity of the foramen rotundum is highlighted in the present case, with embryological correlation and clinical relevance. Anatomical awareness of such variations in the middle cranial fossa are important for the present day radiologist who interprets imaging for this area, neurosurgeons , who operate in this area in the vicinity of cavernous sinus and anaesthetists, during regional block anaesthesia.

Prenatal morphogenesis of the human incisive canal.

The literature describing the formation of the incisive canal is very bizarre. The fusion of the primary and secondary palatal processes leads to formation of a triangular seam, which erroneously has been taken for the future incisive canal. If so, the nasopalatine (incisive) nerve and its accompanying vessels were to run through the primary oronasal cavity, which is not compatible with our biological experience. This study was undertaken to shed light on this region of fusion. We focus on the formation of the incisive canal; the neighboring nasopalatine ducts, which are a transient formation, are mentioned where present. A series of seven horizontal cross-sections of human embryos and fetuses from the 7th to the 24th week of pregnancy (between 25 and 225 mm CRL, crown-rump-length) were examined histologically and partly reconstructed in 3D applying the software analySIS (Soft Imaging Systems, Münster, Germany). The incisive canal did not develop at the junction of the primary and the secondary palate, but within the primary palate rostral to that location. The nasopalatine nerve and the nasopalatine artery are structures that exist before ossification starts in the area of the future incisive canal. The neighboring nasopalatine ducts were found in regions laterally and anterolaterally of the nasopalatine nerve, and it was mostly separated from it by bone. In advanced stages of development, the nasopalatine duct only existed as single epithelial remnants and was prone to obliteration.

Development of a morphologically-based scoring system for postimplantation New Zealand White rabbit embryos.

Rodent whole-embryo culture (WEC) systems are well-established, as are several corresponding morphological scoring systems. Recently, WEC techniques for rabbits have been developed, creating the need for a morphological evaluation system in this species. Consequently, we developed a gestational-age-based quantitative morphology evaluation system for rabbit embryos. Detailed descriptions of 21 embryonic structures, as collected from gestational day (gd) 9-13 rabbit embryos, formed the basis for this evaluation system. These descriptions were then developed into specific criteria for assigning numerical scores to quantify the degree of development of each embryonic structure. The overall morphologic score was calculated as the average of the individual structure scores. To make the system as informative as possible, the numerical scale of the scoring system was gestationally age-based (i.e., range of potential scores was 9.0-13.0). The scoring system was then applied in the evaluation of New Zealand White (NZW) rabbit embryos explanted on gd 9 and cultured for 48 hr. Embryos grown in vitro developed normally, but at a slightly slower rate in vitro than in vivo, as evidenced by the lower morphology score (10.4 in vitro, 11.0 in vivo) and measures of growth (somite number, total protein, and head length). This work firmly establishes the normal archetype of embryonic development in the gd 9-13 NZW rabbit and provides an important tool for the advancement of mechanistic studies of rabbit embryos developing both in vivo and in vitro.

Identification of maxillary factor, a maxillary process-derived chemoattractant for developing trigeminal sensory axons.

Trigeminal sensory axons project to several epithelial targets, including those of the maxillary and mandibular processes. Previous studies identified a chemoattractant activity, termed Maxillary Factor, secreted by these processes, which can attract developing trigeminal axons in vitro. We report that Maxillary Factor activity is composed of two neurotrophins, neurotrophin-3 (NT-3) and Brain-Derived Neurotrophic Factor (BDNF), which are produced by both target epithelium and pathway mesenchyme and which are therefore more likely to have a trophic effect on the neurons or their axons than to provide directional information, at least at initial stages of trigeminal axon growth. Consistent with this, the initial trajectories of trigeminal sensory axons are largely or completely normal in mice deficient in both BDNF and NT-3, indicating that other cues must be sufficient for the initial stages of trigeminal axon guidance.

Anatomic relationships of the orbital muscle of Müller in human fetuses.

A study was carried out in twenty human fetuses on the relationships of the orbital muscle of Müller. This muscle forms a lamina of smooth muscle fibres that cover the inferior orbital fissure. The latter is very wide during the fetal period because ossification of the bones that delimit this region is still incomplete. Some fibres of the orbital muscle extend along the superior orbital fissure under the inferior ophthalmic vein and the lower wall or floor of the cavernous sinus. This association suggests a possible influence on autonomically mediated vascular dynamics.

Developmental pattern of axonal pathways in the house shrew maxillary nerve.

The topographic patterns of peripheral receptors and effectors seem to contribute to the construction of the neuronal circuit in the central nervous system (CNS) in mammals. Many patterns replicating those of the periphery have been found in the CNS, and fasciculation has been regarded as having a central role in the pattern replication. The house shrew, Suncus murinus, is an excellent species in which to study this topic because it has a vibrissae system arranged in a single ordered fashion and extraordinarily well-developed trigeminal spinal tracts. Using immunostaining and retrograde-tracing techniques, we examined the developmental pattern of the maxillary nervous system in the house shrew. The results indicate that the basic pattern of axonal extension reiterates with a parallel arrangement throughout the course of development except at a site in the brainstem where the central processes bifurcate into ascending and descending branches. Dorsoventral inversion of the peripheral pattern in the spinal tract occurs with this dual-leveled bifurcation in association with the mediolaterally ordered entry of the central processes into the brainstem. The basic pattern of the central processes is established prior to the appearance of the vibrissae, indicating that the basic topographic pattern of the maxillary nerve is not related to the vibrissae system. The fasciculation pattern does not correspond to the overall layout of the arrays of vibrissae, and there are frequent exchanges of axons between fascicles both in the periphery and centrally. The parallel organization of the majority of the processes, together with the free exchange of processes between fascicles, suggests that these processes have an important role in the formation of the fasciculation and somatotopic patterns.

Is Far a Hox mutation?

The mouse First arch mutation, Far, causes a severe syndrome of craniofacial defects described previously. All of the known defects are derived from the anterior first arch, and to a very small extent, the dorsal second arch. Recently Far has been shown to be closely linked to Ulnaless on chromosome 2, and therefore in the vicinity of the Hox-4 gene cluster. This paper reports the results of several studies focused on the development origin of the most consistently expressed dominant effect caused by Far, an abnormal major bifurcation of the maxillary nerve. Nerve-stained whole-mount preparations of day 12 embryos showed that in Far mutants the maxillary nerve appears to have a central wedge missing from the normal single-stalked fan shape, and that the nerve defect in Far/Far and +/Far may be equally severe. The effect of retinoic acid on the development of the maxillary nerve was tested. Maternal treatment with 5 mg/kg retinoic acid on day 9 of gestation had no detectable effect on the maxillary nerve of +/Far embryos, and similar treatment with a teratogenic dosage (20 mg/kg) on day 8 or 9 produced no Far-like maxillary nerve defects in genetically normal embryos. The neural crest cells that give rise to nerves and mesenchyme of the first arch originate from specific rhombomeres, discrete segments of the developing head. The rhombomeres of 15 embryos at the 14-23 somite stages, of which 75% are expected to be +/Far or Far/Far, were examined. There was no detectable defect in segmentation or morphology of the rhombomeres compared with controls. The significance of ectopic cartilage in the palate of Far/Far mutants in relation to nerve bifurcation was explored. In histological studies, five out of six Far/Far day-15 fetuses had a rod of ectopic cartilage lateral to the posterior palate, running parallel to, and morphologically similar to, Meckel's cartilage, and lying between the two trunks of the abnormally bifurcated maxillary nerve. None of six +/Far day-15 fetuses examined had detectable ectopic cartilage in this region. We hypothesize that the maxillary nerve defects in Far mutants may be explained by the presence of an ectopic precartilaginous blastema that does not always further develop into detectable cartilage. The ectopic cartilage found in Far/Far resembles the epibranchial cartilage expressed in more posterior branchial arches and in the first arch of lower organisms, and therefore may represent an atavistic posteriorization of the anterior first arch in Far mutants.(ABSTRACT TRUNCATED AT 400 WORDS)

Peripheral development of avian trigeminal nerves.

Development of the trigeminal nerve branches was studied in stage -17 to -27 chick embryos stained with an antibody to neurofilament protein. The following findings were obtained. 1) Ectopic ganglia transiently appeared in the ectoderm of the supraorbital region and were considered as remnant ophthalmic-placode-derived ganglia. 2) Most of the cutaneous sensory branches of the maxillomandibular nerve arose from a loosely arborized mass of neurites, provisionally termed the maxillomandibular reticulum, in which the fibers intermingled in a seemingly random fashion. 3) The growth of the trigeminal branches was mainly correlated with the development of the facial processes; however, irregular communications between different groups of branches were observed, suggesting that topographical organization of the peripheral branches is not rigid in early stages. 4) From the ophthalmic nerve around stage 23, transient dorsal rami developed and were distributed in the mesenchymal space, the cavum epiptericum, and passed near the ectoderm. Their homology with the rr. tentorii in human anatomy is suggested.

Correlated appearance of ossification and nerve tissue in human fetal jaws.

The factors initiating the onset of desmal jaw formation are not known. The purpose of the present report was to examine the correlation between the appearance of ossification and nerve tissue in human fetal jaws. This was done through elaboration of similarities in occurrence of tissue types at four different sites of initial bone formation in the jaws. Radiological and histochemical methods applied to the jaws of 26 human embryos/fetuses revealed that nerve tissue appeared in the jaws before bone tissue. Early bone formation occurred in close relation to the mandibular nerve, the maxillary nerve, the palatine nerve, and the naso-palatine nerve. It is suggested that the foramina (mental foramen, infraorbital foramen, palatal foramen, and incisive foramen) are the areas of incipient bone formation, and that the sequence in bone formation corresponds to the sequence in the development of nerve fibers from the trigeminal ganglion.

The post-injury changes that occur in transected foetal nerves; an experimental study in the rat.

The findings of this experimental foetal rat study have shown that transection of the peripheral branches of the maxillary and facial nerves on the 18th day of gestation is followed by axonal degeneration and regeneration. Although these processes are similar to those which occur in the adult animal, they occur more rapidly in the foetus. In this study the regenerating axons had crossed the wound and entered the neural tubes of the distal nerve segments by 24 hours after injury.

Fasciculation in the early mouse trigeminal nerve is not ordered in relation to the emerging pattern of whisker follicles.

Detailed reconstructions of lengths of the embryonic mouse maxillary nerve were made from serial light and electron microscope sections to determine whether there is any correspondence between the arrangement of fasciculi in the developing nerve and the emerging pattern of whisker follicles on the snout. There was neither correspondence nor obvious pattern in the arrangement of fasciculi at either E11, when trigeminal nerve fibers first contact the presumptive whisker pad, or at E12, when the pattern of developing whisker follicles becomes apparent. Fasciculi merged and branched to form an intricate plexus. Furthermore, the ratio of the number of nerve fibers in one fasciculus to the number in another prior to their merger differed significantly from the ratio of fiber numbers in the two fasciculi after their separation, which indicates that nerve fibers are freely exchanged between fasciculi. Our findings suggest that the somatotopic representation of the whisker follicle pattern in the brainstem does not develop by nerve fiber growth augmenting an initially ordered pattern of fasciculi.

The development of the peripheral trigeminal innervation in Xenopus embryos.

The development of the ophthalmic, maxillary and mandibular nerves has been followed in Xenopus laevis embryos from the first emergence of growth cones from the trigeminal ganglia until the establishment of functional innervation of the skin or cement gland. The course of each main nerve is highly predictable and follows pre-existing openings between blocks of other tissues. The development of the mandibulary nerve was observed most easily. Like that of the other trigeminal nerves it falls into three stages: (1) A pioneer neurite emerges and a nerve forms as other, later neurites fasciculate with this. (2) On reaching the inside surface of the cement gland the neurites separate and penetrate holes in the basal lamina. (3) The neurites grow between the cells they will innervate and form free nerve endings. The scanning EM observations have been confirmed by electrical recordings from trigeminal neurones. The role of pioneer fibres and substrate guidance are discussed.