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 (V₂), represents the apomorphic gnathostome trait that has evolved in association with the acquisition of an upper jaw. J. Morphol. 275:17–38, 2014. © 2013 Wiley Periodicals, Inc.     

Developmental origins and evolution of jaws: new interpretation of "maxillary" and "mandibular"

Cartilage of the vertebrate jaw is derived from cranial neural crest cells that migrate to the first pharyngeal arch and form a dorsal "maxillary" and a ventral "mandibular" condensation. It has been assumed that the former gives rise to palatoquadrate and the latter to Meckel's (mandibular) cartilage. In anamniotes, these condensations were thought to form the framework for the bones of the adult jaw and, in amniotes, appear to prefigure the maxillary and mandibular facial prominences. Here, we directly test the contributions of these neural crest condensations in axolotl and chick embryos, as representatives of anamniote and amniote vertebrate groups, using molecular and morphological markers in combination with vital dye labeling of late-migrating cranial neural crest cells. Surprisingly, we find that both palatoquadrate and Meckel's cartilage derive solely from the ventral "mandibular" condensation. In contrast, the dorsal "maxillary" condensation contributes to trabecular cartilage of the neurocranium and forms part of the frontonasal process but does not contribute to jaw joints as previously assumed. These studies reveal the morphogenetic processes by which cranial neural crest cells within the first arch build the primordia for jaw cartilages and anterior cranium.     

Notes on the typology of the skull of the Myrmecophagidae (Mammalia, Edentata)

The skulls of Myrmecophaga, Tamandua, and Cyclopes are klinorhynch; the upper jaw is situated rostral to the neurocranium and to a varying degree ventral to the plane of the median basis cranii. The median part of the base of the neurocranium is the structure to which the anatomical modifications in the median plane are referred. The kyphosis, which determines the situation of the upper jaw, is prebasically located either within the upper jaw (Myrmecophaga, Tamandua) or at its basis (Cyclopes).     

Arch form,tooth size,and occlusomandibular kinesis in the Ceboidea

Correlations between dental morphology, arch configuration, and jaw movement patterns were quantitatively investigated in 23 ceboid species to elucidate integrative aspects of occlusal functional anatomy in an adaptive and evolutionary context. Differential maxillary-mandibular arch widths are primary in guiding lateral jaw movements. These movements are characterized according to their associated condylar shifts as either predominantly translatory or rotational. Predominantly translatory movements result from peripheral contact relationships between maxillary arches which are considerably wider posteriorly than their opposing mandibular arches. The greatest degree of mandibular movement is in the molar region in functional association with wide “primitive” maxillary molars, narrow mandibular molars, constricted maxillary intercanine widths, and narrow maxillary incisors. In contrast, predominantly rotational masticatory jaw movements result from differential arch widths which are greatest in the maxillary canine and incisor regions. Here most jaw movement is in the anterior segment and this is reflected in small maxillary-mandibular molar width differences, a high degree of premolarization, wide-set maxillary canine teeth, and wide maxillary incisors. Possible selectional factors in the putative evolution of rotational predominance in mastication from the more primitive translatory pattern are discussed.     

A morphological study of Luciocephalus pulcher,with notes on gular elements in other recent teleosts

The monotypic perciform suborder Luciocephaloidei possesses the following, previously unknown, salient morphological characters: a third joint, the nasopalatopterygoid, between neurocranium and suspensorium; a toothless and dorsally exposed prevomer; no pharyngeal processes on either parasphenoid or basioccipital; a tympanum-covered foramen exoccipitale in the saccular bulla as a hearing organ; a gular ossification; a craniovertebral joint with small exoccipital condyles widely separated from the basioccipital condyle; and a large physoclystic swimbladder with a notable caudal extension. The seemingly functionless gular-like mental ossification is considered a paleomorphic structure with a neogenetic development. Attenuation in longitudinal growth is evident in the derivatives and dermal additions of the mandibular arch and nasal capsule while other regions of the head have remained independent. The primary adaptive significance of the attenuation in the growth of the entire preorbital region is the accomodation of oral incubation. Secondarily the elongate jaws increase both the speed and grasping range of the bite in prey catching. The degree of jaw protrusion depends mainly on the length of the maxillary. The exaggerated length of the ascending processes of the premaxillaries may be the result of a positive differential growth rate within one growth field. The meaning of the preponderance of parallel-fibered cranial muscles is discussed in respect to holding functions, greatest possible excursion of the insertion with minimum loss of force, and muscle fiber length. Based on the overall morphology, the monotypic suborder Luciocephaloidei is retained.     

Anatomy of the feeding apparatus of the lemon shark,Negaprion brevirostris

The anatomy of the feeding apparatus of the lemon shark, Negaprion brevirostris, is investigated by gross dissection, computer axial tomography, and histological staining. The muscles and ligaments of the head associated with feeding are described. The upper and lower jaws are suspended by the hyoid arch, which in turn is braced against the chondrocranium by a complex series of ligaments. In addition, various muscles and the integument contribute to the suspension and stability of the jaws. The dual jaw joint is comprised of lateral and medial quadratomandibular joints that resist lateral movement of the upper and lower jaws on one another. This is important during feeding involving vigorous head shaking. An elastic ethmoplatine ligament that unites the anterior portion of the upper jaw to the neurocranium is involved with upper jaw retraction. The quadratomandibularis muscle is divided into four divisions with a bipinnate fiber arrangement of the two large superficial divisions. This arrangement would permit a relatively greater force per unit volume and reduce muscle bulging of the jaw adductor muscle in the spatially confined cheek region. Regions of relatively diffuse integumental ligaments overlying the adductor mandibulae complex and the levator palatoquadrati muscle, interspersed with localized regions of longer tendonlike attachments between the skin and the underlying muscle, permit greater musculoskeletal movement relative to the skin. The nomenclature of the hypobranchial muscles is discussed. In this shark they are comprised of the unsegmented coracomandibularis and coracohyoideus, and the segmented coracoarcualis. © 1995 Wiley-Liss, Inc.     

The oral health‐related quality of life in edentulous patients treated with Conventional complete dentures

Objective: To compare the oral health‐related quality of life (OHRQoL) between patients with both maxillary and mandibular complete denture and those with either the maxillary or the mandibular complete denture. Background: Satisfaction of denture wearers can be estimated using the OHRQoL questionnaires like the OHIP‐EDENT and the Geriatric Oral Health Assessment Index (GOHAI). Methods: Two questionnaires were used to compare the OHRQoL between edentulous patients who had conventional removable complete denture on both jaws and those who had on either one of the jaws. Result: The age of the participants ranged from 42 to 75 years, with the mean age of 58 ± 8.12 years. The mean OHIP‐EDENT scores were significantly high among those who wore conventional dentures in both jaws (54.12 ± 5.21), compared with the participants who only had denture either on upper or lower jaw (46.52 ± 7.35). It was noticed that the mean GOHAI score was significantly lower (p < 0.05) among participants who had conventional denture on both upper and lower jaw (28.25 ± 3.67), as compared to those who had conventional denture only on one arch (35.12 ± 2.11). Conclusion: Patients with complete dentures in both jaw (Group I) were less satisfied than patients with single complete denture (Group II). The result obtained in this study shows dissatisfaction with conventional dentures among edentulous patients.     

Evolution of the vertebrate jaw from developmental perspectives

Attainment of the biting jaw is regarded as one of the major novelties in the early history of vertebrates. Based on a comparison between lamprey and gnathostome embryos, evolutionary developmental studies have tried to explain this novelty as changes in the developmental patterning of the mandibular arch, the rostralmost pharyngeal arch, at the molecular and cellular levels. On the other hand, classical theories in the field of comparative morphology assumed the involvement of hypothetical premandibular arch(es) that ancestral animals would have possessed rostral to the mandibular arch, in the transition from agnathan to gnathostome states. These theories are highly biased toward the segmental scheme of the vertebrate head, and the concept of premandibular “arches” is no longer accepted by the current understanding. Instead, the premandibular domain has now become of interest in the understanding of cranial development, especially in its rostral part. As newer theories that consider involvement of the premandibular domain, the neoclassical and heterotopy theories are here compared from evolutionary developmental perspectives, in conjunction with the development of nasal and hypophyseal placodes, in the context of the evolutionary acquisition of the jaw. Given recent advances in understanding of the lamprey development, evolution of the Dlx code is also discussed together with the evolutionary scenario of jaw acquisition.     

Jaw transformation with gain of symmetry after Dlx5/Dlx6 inactivation: Mirror of the past?

In modern vertebrates upper and lower jaws are morphologically different. Both develop from the mandibular arch, which is colonized mostly by Hox-free neural crest cells. Here we show that simultaneous inactivation of the murine homeobox genes Dlx5 and Dlx6 results in the transformation of the lower jaw into an upper jaw and in symmetry of the snout. This is the first homeotic-like transformation found in this Hox-free region after gene inactivation. A suggestive parallel comes from the paleontological record, which shows that in primitive vertebrates both jaws are essentially mirror images of each other. Our finding supports the notion that Dlx genes are homeotic genes associated with morphological novelty in the vertebrate lineage.     

Mandibular arch musculature of anuran tadpoles, with comments on homologies of amphibian jaw muscles

This study analyzes the structure of the mandibular arch musculature in larval, metamorphic, and postmetamorphic anurans of 26 species and makes comparisons with larvae of three caudate and one gymnophione species. Major transformations in early evolution of anuran larvae comprise, for example, the powering of the larval upper jaw cartilages by relocating insertion sites of mandibular arch levators; splitting of some larval muscles into two muscles or muscle heads (m. intermandibularis, m. lev. mand. externus, m. lev. mand. longus); evolution of a muscle invading the lower lip of the oral disk (m. mandibulolabialis), and shift of origin of the internus and longus muscles from dorsal on the cranium to sites on the ventral otic capsule and palatoquadrate, respectively. In all these characters, Ascaphus truei shares the plesiomorphic conditions with caudates. The larva of Xenopus laevis is remarkable because the insertion pattern of three larval mandibular muscles anticipates the postmetamorphic condition of frogs in general and also resembles the caudate condition. Discoglossids, bombinatorids, pelobatids, and neobatrachians are largely similar in their muscle arrangements. The filter-feeding microhylids, however, have most clearly modified the general neobatrachian pattern. Past conflicts in the interpretation and naming of muscles can be attributed to the implicit or explicit homology assumptions used. In particular, the muscles' relations to the branches of the trigeminal nerve have been the dominant criteria for inferring homology and has led to inconsistencies. This concept is questioned herein. It is observed that the relative position of the ramus mandibularis (V(3)) is more variable interspecifically in anuran larvae than previously thought. The relations of the nerve branches and muscles in larvae are maintained during metamorphosis. Considering the muscle pattern to be more conserved in interspecific comparisons than the position of the nerve branches results in a new interpretation of muscle homologies and a hypothesis of jaw muscle evolution in amphibians that is more parsimonious than earlier views. A new, simplified terminology for the jaw musculature is proposed that is applicable for larvae and adults. It maximizes information content and reflects the hypothesized homologies of amphibian jaw muscles.     

Endothelin-A receptor-dependent and -independent signaling pathways in establishing mandibular identity

The lower jaw skeleton is derived from cephalic neural crest (CNC) cells that reside in the mandibular region of the first pharyngeal arch. Endothelin-A receptor (Ednra) signaling in crest cells is crucial for their development, as Ednra(-/-) mice are born with severe craniofacial defects resulting in neonatal lethality. In this study, we undertook a more detailed analysis of mandibular arch development in Ednra(-/-) embryos to better understand the cellular and molecular basis for these defects. We show that most lower jaw structures in Ednra(-/-) embryos undergo a homeotic transformation into maxillary-like structures similar to those observed in Dlx5/Dlx6(-/-) embryos, though lower incisors are still present in both mutant embryos. These structural changes are preceded by aberrant expansion of proximal first arch gene expression into the distal arch, in addition to the previously described loss of a Dlx6/Hand2 expression network. However, a small distal Hand2 expression domain remains. Although this distal expression is not dependent on either Ednra or Dlx5/Dlx6 function, it may require one or more GATA factors. Using fate analysis, we show that these distal Hand2-positive cells probably contribute to lower incisor formation. Together, our results suggest that the establishment of a 'mandibular identity' during lower jaw development requires both Ednra-dependent and -independent signaling pathways.     

Fishing for jaws in early vertebrate evolution: a new hypothesis of mandibular confinement

The evolutionary origin of the vertebrate jaw persists as a deeply puzzling mystery. More than 99% of living vertebrates have jaws, but the evolutionary sequence that ultimately gave rise to this highly successful innovation remains controversial. A synthesis of recent fossil and embryological findings offers a novel solution to this enduring puzzle. The Mandibular Confinement Hypothesis proposes that the jaw evolved via spatial confinement of the mandibular arch (the most anterior pharyngeal arch within which the jaw arose). Fossil and anatomical evidence reveals: (i) the mandibular region was initially extensive and distinct among the pharyngeal arches; and (ii) with spatial confinement, the mandibular arch acquired a common pharyngeal pattern only at the origin of the jaw. The confinement occurred via a shift of a domain boundary that restricted the space the mesenchymal cells of the mandibular arch could occupy. As the surrounding domains replaced mandibular structures at the periphery, this shift allowed neural crest cells and mesodermal mesenchyme of the mandibular arch to acquire patterning programs that operate in the more posterior arches. The mesenchymal population within the mandibular arch was therefore no longer required to differentiate into specialized feeding and ventilation structures, and was remodelled into a jaw. Embryological evidence corroborates that the mandibular arch must be spatially confined for a jaw to develop. This new interpretation suggests neural crest as a key facilitator in correlating elements of the classically recognized vertebrate head ‘segmentation’.     

Fgf and Bmp signals repress the expression of Bapx1 in the mandibular mesenchyme and control the position of the developing jaw joint

The development of the jaw joint between the palatoquadrate and proximal part Meckel's cartilage (articular) has recently been shown to involve the gene Bapx1. Bapx1 is expressed in the developing mandibular arch in two distinct caudal, proximal patches, one on either side of the head. These domains coincide later with the position of the developing jaw joint. The mechanisms that result in the restricted expression of Bapx1 in the mandibular arch were investigated, and two signaling factors that act as repressors were identified. Fibroblast growth factors (Fgfs) expressed in the oral epithelium restrict expression of Bapx1 to the caudal half of the mandibular arch, while bone morphogenetic proteins (Bmps) expressed in the distal mandibular arch restrict expression of Bapx1 to the proximal part of the mandible. Application of Fgf8 and Bmp4 beads to the proximal mesenchyme led to loss of Bapx1 expression and later fusion of the quadrate and articular as the jaw joint failed to form. In addition to fusion of the jaw joint, loss of Bapx1 lead to loss of the retroarticular process (RAP), phenocopying the defects seen after Bapx1 function was reduced in the zebrafish. By manipulating these signals, we were able to alter the expression domain of Bapx1, resulting in a new position of the jaw joint.     

Modeling the jaw mechanism of Pleuronichthys verticalis: The morphological basis of asymmetrical jaw movements in a flatfish

Several flatfish species exhibit the unusual feature of bilateral asymmetry in prey capture kinematics. One species, Pleuronichthys verticalis, produces lateral flexion of the jaws during prey capture. This raises two questions: 1) How are asymmetrical movements generated, and 2) How could this unusual jaw mechanism have evolved? In this study, specimens were dissected to determine which cephalic structures might produce asymmetrical jaw movements, hypotheses were formulated about the specific function of these structures, physical models were built to test these hypotheses, and models were compared with prey capture kinematics to assess their accuracy. The results suggest that when the neurocranium rotates dorsally the premaxillae slide off the smooth, rounded surface of the vomer (which is angled toward the blind, or eyeless, side) and are “launched” anteriorly and laterally. The bilaterally asymmetrical trajectory of the upper jaw is determined by the orientation of the “launch pad,” the vomer. During lower jaw depression, the mandibles rotate about their articulations with the quadrate bones of the suspensoria. The quadrato‐mandibular joint is positioned farther anteriorly on the eye side than on the blind side, and this asymmetry deflects the lower jaw toward the blind side. Asymmetry in the articular surfaces of the lower jaw augments this effect. Thus, it appears that fish with intermediate forms of this asymmetrical movement could have evolved from symmetrical ancestors via a few key morphological changes. In addition, similar morphological modifications have been observed in other fish taxa that also produce jaw flexion during feeding, which suggests that there may be convergence in the basic mechanism of asymmetry. J. Morphol. 256:1–12, 2003. © 2003 Wiley‐Liss, Inc.     

Description of a new diphyllidean parasite of triakid sharks from the deep Red Sea

Specimens of Echinobothrium diamanti n. sp. (Cestoda: Diphyllidea) were recovered from the spiral intestine of Iago omanensis and Mustelus mosis (Carcharhiniformes: Triakidae), in the Gulf of Aqaba, Red Sea. The new species can be distinguished from all other species in Echinobothrium by the presence of a conspicuous vaginal sphincter. Echinobothrium diamanti possesses a corona of spines between the apical armature and the bothria, as in Echinobothrium notoguidoi, Echinobothrium musteli, and Echinobothrium scoliodoni, also parasites of sharks. However, E. diamanti possesses more testes per proglottid than E. notoguidoi and E. scoliodoni, and it is larger and has more spines per column on the cephalic peduncle than E. musteli and E. notoguidoi, and it also has circum-medullary vitelline follicles rather than distributed in lateral columns. Echinobothrium diamanti is the first species of diphyllidean reported from the triakid genus Iago.     

Hind limb movements of the American alligator (Alligator mississippiensis) and postural grades

Middle ear ontogeny in Monodelphis domestica is investigated to understand better both the immediate consequences for suckling in a neonate marsupial and the epigenetic factors that constrain morphogenesis. Neonates of Monodelphis possess neither mammalian (dentarysquamosal) nor reptilian (quadrate-articular) jaw articulations, nor does the contact between the incus and crista parotica offer a joint surface ( contra Maier, 1987). Elasticity in Meckel's cartilage allows minimal deflection of the lower jaw. Observation of the developmental rate of the individual elements reveals that mandibular arch derivatives (malleus, incus and tensor tympani) are on a slightly faster ontogenetic schedule than hyoid arch derivatives (stapes and m. stapedius).