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Vertebrates have achieved great evolutionary success due in large part to the anatomical diversification of their jaw complex, which allows them to inhabit almost every ecological niche. While many studies have focused on mechanisms that pattern the jaw skeleton, much remains to be understood about the origins of novelty and diversity in the closely associated musculature. To address this issue, we focused on parrots, which have acquired two anatomically unique jaw muscles: the ethmomandibular and the pseudomasseter. In parrot embryos, we observe distinct and highly derived expression patterns for Scx, Bmp4, Tgfβ2 and Six2 in neural crest-derived mesenchyme destined to form jaw muscle connective tissues. Furthermore, immunohistochemical analysis reveals that cell proliferation is more active in the cells within the jaw muscle than in surrounding connective tissue cells. This biased and differentially regulated mode of cell proliferation in cranial musculoskeletal tissues may allow these unusual jaw muscles to extend towards their new attachment sites. We conclude that the alteration of neural crest-derived connective tissue distribution during development may underlie the spatial changes in jaw musculoskeletal architecture found only in parrots. Thus, parrots provide valuable insights into molecular and cellular mechanisms that may generate evolutionary novelties with functionally adaptive significance.

First arch syndromes correspond to a wide spectrum of human latero-facial congenital anomalies affecting cranial neural crest cells (CNCCs) derivatives of the first pharyngeal arch (PA1). The abnormal traits display variable quantitative expression and are often unilateral. Mandibular skeletal defects are invariably accompanied by hypoplasia or agenesis of masticatory muscles, but no explanation has been proposed for this association. Indeed, during embryonic development, CNCCs give only rise to skeletal components of the head while muscles derive from cephalic myogenic mesodermal cells (CMMCs). Recent studies on animal models have shown that communication between CNCCs and CMMCs is essential for the development of masticatory muscles: genetic lesions affecting only CNCCs can prevent muscularization of the jaws. To evaluate the involvement of CNCC/CMMC interactions in human craniofacial development, we performed a quantitative analysis of masticatory muscle and mandibular bone volumes on craniofacial CT-scans from 8 children, ages 3 months to 16 years, affected by hemifacial microsomia. We found that: (1) in seven patients the masseter muscle is absent in the affected side; (2) the absence of masseter is correlated neither with the age of the patients nor with the volume and shape of the affected ramus; and (3) in all cases the pterygoid and the temporal muscles are either reduced or absent. Our findings suggest that an early developmental event is the origin of the muscular defects in these patients. We propose that the hypoplasia or agenesis of masticatory muscles derives from a defect in the CNCCs/CMMCs communication during early embryonic development., (Copyright © 2011 Wiley-Liss, Inc.)

The morphological events forming the body's musculature are sensitive to genetic and environmental perturbations with high incidence of congenital myopathies, muscular dystrophies and degenerations. Pattern formation generates branching series of states in the genetic regulatory network. Different states of the network specify pre-myogenic progenitor cells in the head and trunk. These progenitors reveal their myogenic nature by the subsequent onset of expression of the master switch gene MyoD and/or Myf5. Once initiated, the myogenic progression that ultimately forms mature muscle appears to be quite similar in head and trunk skeletal muscle. Several genes that are essential in specifying pre-myogenic progenitors in the trunk are known. Pax3, Lbx1, and a number of other homeobox transcription factors are essential in specifying pre-myogenic progenitors in the dermomyotome, from which the epaxial and hypaxial myoblasts, which express myogenic regulatory factors (MRFs), emerge. The proteins involved in specifying pre-myogenic progenitors in the head are just beginning to be discovered and appear to be distinct from those in the trunk. The homeobox gene Pitx2, the T-box gene Tbx1, and the bHLH genes Tcf21 and Msc encode transcription factors that play roles in specifying progenitor cells that will give rise to branchiomeric muscles of the head. Pitx2 is expressed well before the onset of myogenic progression in the first branchial arch (BA) mesodermal core and is essential for the formation of first BA derived muscle groups. Anterior-posterior patterning events that occur during gastrulation appear to initiate the Pitx2 expression domain in the cephalic and BA mesoderm. Pitx2 therefore contributes to the establishment of network states, or kernels, that specify pre-myogenic progenitors for extraocular and mastication muscles. A detailed understanding of the molecular mechanisms that regulate head muscle specification and formation provides the foundation for understanding congenital myopathies. Current technology and mouse model systems help to elucidate the molecular basis on etiology and repair of muscular degenerative diseases.

The development of the mandible was traced on serial sections of 20 human embryos aged 5-8 weeks (developmental stages 13-23). Special consideration was given to the differentiation of skeletal units proposed by Sperber. The first skeletal units, namely the mandibular body, the alveolar unit and the condylar unit, may be distinguished in the 7(th) week. The primordia of all units are identified by the end of the embryonic period (8 weeks).

The development of suckling behavior is a fundamental characteristic of mammalian development. The occurrence of this behavior across mammals allows us to extrapolate information from animal models to better understand normal and abnormal masticatory development in infants. This review focuses on prenatal cell, molecular, and morphological changes in rat and/or mouse masticatory muscles, trigeminal motoneurons (Mo5) and mesencephalic trigeminal neurons (Me5) that accompany the development of suckling behavior. A special emphasis is placed on N-methyl-d-aspartate (NMDA) receptor subunit expression because of the important role that NMDA receptors play in the production of rhythmical jaw movements and neuronal development. Prenatally the timing of NMDA subunit changes follows neuromuscular junction formation in masticatory muscles, and is coincident with the emergence of rhythmical jaw movements and in vitro rhythmical trigeminal activity. Our data suggest that NMDA receptor subunit changes in Mo5 and Me5 are synchronized with the emergence of rhythmical jaw movements and trigeminal motor activity.

This review summarizes findings concerning the unique developmental characteristics of mouse head muscles (mainly the masticatory and tongue muscles) and compares their characteristics with those of other muscles. The developmental origin of the masticatory muscles is the somitomeres, whereas the tongue and other muscles, such as the trunk (deep muscles of the back, body wall muscles) and limb muscles, originate from the somites. The program controlling the early stages of masticatory myogenesis, such as the specification and migration of muscle progenitor cells, is distinctly different from those in trunk and limb myogenesis. Tongue myogenesis follows a similar regulatory program to that for limb myogenesis. Myogenesis and synaptogenesis in the masticatory muscles are delayed in comparison with other muscles and are not complete even at birth, whereas the development of tongue muscles proceeds faster than those of other muscles and ends at around birth. The regulatory programs for masticatory and tongue myogenesis seem to depend on the developmental origins of the muscles, i.e., the origin being either a somite or somitomere, whereas myogenesis and synaptogenesis seem to progress to serve the functional requirements of the masticatory and tongue muscles.

The current genetic data stress the importance of musculo-skeletal connections in the development of a coherent system connecting the tendons and aponeurosis muscles with the osseous parts. The observations in tomodensitometry of musculo-skeletal connections in otomandibular dysostosis make it possible qualitatively to observe the development of the muscles and their functions.

Four pairs of branchial arch appear apparently in the neck region of human embryo about 32 days after fertilization. Maxillary prominence of the first branchial arch gives rise to the maxilla and zygomatic bone etc., and mandibular prominence forms the mandible and so on. Muscles for mastication are also derived from 1st branchial arch, into which fifth cranial nerves grow from the brain. Thus, human embryo improves the 1st branchial arches into the upper and lower jaws, and forms digestive organs needed for intake, mastication, and swallowing of foods. Finally they develop the brain for integral treatment of sensory information from eyes, tongue, nose, and ears.

Parrots have developed novel head structures in their evolutionary history. The appearance of two new muscles for strong jaw adduction is especially fascinating in developmental and evolutionary contexts. However, jaw muscle development of parrots has not been described, despite its uniqueness. This report first presents the normal developmental stages of the cockatiel (Nymphicus hollandicus), comparable to that of the chick. Next, the peculiar skeletal myogenesis in the first visceral arch of parrots is described, mainly focusing on the development of two new jaw muscles. One of the parrot-specific muscles, M. ethmomandibularis, was initially detected at Nymphicus Stage 28 (N28) as the rostral budding of M. pterygoideus. After N32, the muscle significantly elongates rostrodorsally toward the interorbital septum, following a course lateral to the palatine bone. Another parrot-specific muscle, M. pseudomasseter, was first recognized at N36. The muscle branches off from the posteromedial M. adductor mandibulae externus and grows in a dorsolateral direction, almost covering the lateral surface of the jugal bar. The upper tip of the muscle is accompanied by condensed mesenchyme, which seems to be derived from cephalic neural crest cells., (Copyright 2003 Wiley-Liss, Inc.)

Members of the MyoD family of basic helix-loop-helix (bHLH) transcription factors control the formation of all skeletal muscles in vertebrates, but little is known of the molecules or mechanisms that confer unique identities to different types of skeletal muscles. MyoR and capsulin are related bHLH transcription factors expressed in specific facial muscle precursors. We show that specific facial muscles are missing in mice lacking both MyoR and capsulin, reflecting the absence of MyoD family gene expression and ablation of the corresponding myogenic lineages. These findings identify MyoR and capsulin as unique transcription factors for the development of specific head muscles.

The development of the muscles in the floor of the mouth is described in 10 human embryos and fetuses ranging from 6.9 to 76 mm CRL by means of computer-aided graphical 3D-reconstructions. All primordia of the muscles in the floor of the mouth could be identified from the 15.6 mm CRL stage on. The proportions and insertion lines of the early muscles were found to be different from adult anatomy. Each muscle first inserted in the medial surface of Meckels cartilage, but during the developmental period between 19 and 68 mm CRL the insertion lines were gradually transposed to the bony ridges of the mandible which progrediently embraced Meckels cartilage. The fibers of the mylohyoid muscles left the anterior region near the symphysis mentalis free during all stages of this study. The digastric muscle revealed only one belly with a constriction of its continuous fibers where it passed the hyoid bone primordium. There was no attachment of digastric muscle fibers to the hyoid; only geniohyoid and mylohyoid fibers. Geniohyoid and genioglossus muscles basically correspond to their definite arrangement, but they underwent proportional changes. Individual specimens embodied irregularities such as accessory geniohyoid and hyoid portions and muscle fibers separate from the mylohyoide muscle.

The Krox-20 null mutation results in a loss of rhombomeres 3 and 5, which give rise to neurons that are essential to oral motor behaviors. Thus, the Krox-20 null mutant is an excellent model to investigate the development of oral motor circuitry. Our morphological examination of embryonic and neonatal Krox-20 null mutants revealed that a significant reduction of anterior digastric and mylohyoid muscles, the primary jaw openers, occurs between embryonic days 15 and 19. There are no gross morphological alterations in other masticatory muscles. These findings demonstrate that Krox-20 expression is critical for the normal development of the primary jaw opener musculature and they help explain previous studies documenting a reduction in jaw opening in Krox-20 null mutants. Since jaw opening is the power stroke of suckling behavior, our data help explain the reduction of colostrum/milk ingestion in Krox-20 null mutant neonates., (Copyright 2001 S. Karger AG, Basel)

Purpose: In this study, the development of human embryonic temporomandibular joint (TMJ) and masticatory muscles were investigated by using computed 3-dimensional reconstructions., Materials and Methods: Sixteen human embryos and fetuses, ranging from 6.5 to 107 mm crown-rump length, were examined., Results: At 10 weeks, a band of mesenchyme extending from the attachment of the lateral pterygoid muscle to the condylar process was observed to pass through the medial side of the condylar process to attach to the malleus. The temporal, masseter, and pterygoid muscles develop from the so called "temporal muscle" primordium, and the temporal muscle was in continuity with the masseter muscle until 14 weeks of fetal life., Conclusions: The study shows that the muscles of mastication arise from a single primordium. It also confirms the presence of a ligamentous attachment between the lateral pterygoid muscle and the malleus.

Inactivation of neurotrophin-3 (NT3) completely blocks the development of limb proprioceptive neurons and their end organs, the muscle spindles. We examined whether cranial proprioceptive neurons of the trigeminal mesencephalic nucleus (TMN) require NT3, brain-derived neurotrophic factor (BDNF) or neurotrophin-4 (NT4) for their development. Complements of TMN neurons and masticatory muscle spindles were decreased by 62% in NT3 null mutants, 33% in BDNF null mutants, and 10% in NT4 null mutant mice at birth. The extent of proprioceptive deficiencies differed among different masticatory muscles, particularly in NT3 null mice. Masticatory muscles of embryonic mice heterozygous for the NT3(lacZneo) or BDNF(lacZ) reporter genes expressed both NT3 and BDNF, consistent with target-derived neurotrophin support of TMN neurons. Although more than 90% of TMN neurons expressed TrkB as well as TrkC receptor proteins by immunocytochemistry in wild-type newborns, TrkC or TrkB null mice exhibited only partial proprioceptive deficiencies similar to those present in NT3 or BDNF;NT4 null mice. Thus, in terms of the survival outcome, two main subpopulations of TMN neurons may exist during embryogenesis, one dependent on TrkC/NT3 functioning and the other utilizing TrkB/BDNF signaling. The differential dependence of TMN neurons on neurotrophins may reflect differential accessibility of the neurons to limiting amounts of NT3, BDNF, or NT4 in target tissues, especially if the tissue distribution or levels of BDNF, NT3, and NT4 were dynamically regulated both spatially and temporally., (Copyright 2000 Wiley-Liss, Inc.)

The digastric muscles of 99 cadavers were examined bilaterally. In 5 of them, the digastric muscles showed different types of complex abnormalities. These complex anomalies should be kept in mind in functional studies involving the floor of mouth and evaluating the same structures with CT and MR imaging. Secondly, the embryological basis of the isolated anomalies of the anterior digastric muscle with respect to its occurrence together with the mylohyoid muscle stressed in this article needs further studies of their development from the first branchial arch.

The Puerto Rican direct-developing frog Eleutherodactylus coqui (Leptodactylidae) displays a novel mode of jaw muscle development for anuran amphibians. Unlike metamorphosing species, several larval-specific features never form in E. coqui; embryonic muscle primordia initially assume an abbreviated, mid-metamorphic configuration that is soon remodelled to form the adult morphology before hatching. Also lacking are both the distinct population of larval myofibres and the conspicuous, larval-to-adult myofibre turnover that are characteristic of muscle development in metamorphosing species. These modifications are part of a comprehensive alteration in embryonic cranial patterning that has accompanied life history evolution in this highly speciose lineage. Embryonic 'repatterning' in Eleutherodactylus may reflect underlying developmental mechanisms that mediate the integrated evolution of complex structures. Such mechanisms may also facilitate, in organisms with a primitively complex life cycle, the evolutionary dissociation of embryonic, larval, and adult features.

Degenerating myofibers have been reported in the embryos and neonates of a number of birds and mammals, but neither the pervasiveness of the phenomenon nor the spatio-temporal patterns of degeneration has been examined in detail. Using transmission electron microscopy, we determined the patterns of muscle cell death in the chick biventer cervicis, a head extensor muscle. Cell death is most pronounced at incubation days 10 through 15, and occurs throughout the muscle. This is the period during which many myofiber clusters segregate into individual fibers, each with a separate basal lamina, and secondary myofibers become demarcated. Cells of largest diameter, presumably the primary myofibers, are preferentially affected. Degenerating cells exhibit a cohort of cytological features consistent with apoptosis, including the presence of dense, darkly-staining, hypercontracted myofibrils, misshapen nuclei with irregular chromatin condensations along the nuclear envelope, and scores of cytoplasmic vesicles and vacuoles. In cross section some large diameter muscle cells are characterized by sparse, flocculent cytoplasm that is devoid of myofibrils and organelles. Some show disintegrating cell membranes. In longitudinal section 200-300 microns long regions of hypercontracted myofibrils alternate with areas devoid of fibrils; this arrangement suggests that the myofibrils break into segments that are in register along one part of a muscle fiber and entirely absent from the adjacent length of fiber. We have observed similar patterns of muscle cell degeneration in the complexus, splenius cervicis, depressor mandibulae, and branchiomandibularis muscles. By day 18 of incubation most signs of degeneration are absent and by hatching (day 21) the muscle fibers all appear healthy. Many of these cytological changes in embryonic head muscle cells are characteristic of programmed cell death. We hypothesize that large-scale death of myocytes is a normal part of avian myogenesis and an important mechanism for affecting the transformation from embryonic to hatching muscle patterning.

The arrangement of the pterygospinosus muscle was analyzed in 5 human fetuses. The pterygospinosus muscle extends from the posterior border of the lateral lamina of the pterygoid process to Meckel's cartilage. Such an arrangement would permit its action on the joint formed by Meckel's cartilage and the incus of the middle ear. The pterygospinosus muscle is a remnant of the masticatory muscle group. The relationships of the pterygospinosus muscle with the mandibular nerve and its branches and the maxillary artery and its branches were analyzed.

The calcification levels of the mandible and the temporal bone of human fetuses, which ranged from 12 to 32 weeks of gestation, were systematically investigated with a soft X-ray analyzer linked to an image analyzer. The profile of the condylar process (head) revealed high levels of calcification, in contrast to that in the mandibular fossa of the temporal bone. The basal portion of the condylar process and the mandibular notch exhibited moderate calcification from 12 weeks of gestation. The weight and the cross-sectional areas of the muscle and the muscle fibers in masticatory muscles (masseter, temporal, medial, and lateral pterygoid muscles) are all increased gradually during development from 12 to 32 weeks of gestation. These changes in calcification and in cross-sectional area of muscle suggest that muscle development may be related to bone calcification during formation of the mandible.

A study of the relationships of the temporomandibular joint disc and the lateral pterygoid, temporalis, and masseter muscles during the human fetal period and in the adult was conducted. The superior head of the lateral pterygoid muscle was seen to insert into the anteromedial two thirds of the temporomandibular joint disc. The fibers of the posterior one third of the temporalis muscle and fibers of the deep bundle of the masseter muscle were attached on the anterolateral one third of the disc. The attachment of these muscles to the disc was through the anterior extension of the disc, also known as the premeniscal or prediscal lamina. The possible functional role of these muscle attachments is discussed.

The late fetal development of rat extra-ocular and masticatory muscles was examined by myosin immunohistochemistry. The pattern of slow and neonatal myosin isoform expression in primary and secondary myotubes in these muscles was generally similar to that seen by others in limb muscles. We observed a consistent difference between the Sprague-Dawley and Wistar rats in the degree of maturity reached by all muscles studied at a particular age. In both strains, extra-ocular muscles were also about one day in advance of the masticatory muscles. Thus, secondary myotubes were first seen at E17 in Wistar extraocular muscles, at E18 in Sprague-Dawley extra-ocular muscles and Wistar masticatory muscles, and at E19 in Sprague-Dawley masticatory muscles. There was a strikingly early and complete type differentiation of primary myotubes in extraocular muscles, and tonic myosin first appeared before birth in presumptive extrafusal tonic fibres in the orbital layer of the oculorotatory muscles. Throughout the late fetal period, retractor bulbi was composed of fast myotubes only, but these myotubes were not arranged in classical clusters. In the masticatory muscles at E17/E18 some slow primary myotubes started to express tonic myosin, and these presumptive spindle bag2 fibres were located only in regions of the muscles known to contain spindles in the adult. Presumptive bag1 fibres appeared about a day later (initially without tonic myosin), and in the region of the spindle cluster in anterior deep masseter extrafusal secondary myotube production appeared to be suppressed.

Age changes of morphometrical parameters of the masticatory muscles have been analyzed in domestic sheep and pigs of white large breed in the following age groups: 2-, 3-, 4-month-old fetuses, newborns, 4-month-old lambs, 10-month-old pigs, 18-month-old lambs, mature she-sheep and brood-sows. Uneven weight growth of the masticatory muscles in the sheep and pigs during the prenatal ontogenesis should be considered as a consequence of recapitulation of their phylogenesis, and in the postnatal ontogenesis it depends on changes in life conditions, type of nutrition, character of food and type of life. In newborn sheep the digastric, lateral, pterygoid and temporal muscles grow intensively, and in pigs--medial pterygoid and temporal ones. When they pass to roughage, in the former the mass of the musculus masseter major and medial pterygoid muscle increases, and in the latter--that of the musculus masseter major and temporal one. The masticatory muscles of the species studied increase in their mass especially intensively during the middle of the prenatal ontogenesis and during suckling period of their development. This should be taken into consideration in stock-breeding practice. In domestic pigs there is only one muscular belly in the digastric muscle. In sheep there are two bellies, separated one from another by means of a tendinous intersection, owing to crossing of the latter by the stylohyoid muscle.

Previous results have shown that the adult human masseter muscle contains myosin isoforms that are specific to early stages of development in trunk and limb muscles, i.e. embryonic and fetal (neonatal) myosin heavy chains (MHC) and embryonic myosin light chain (MLC1emb). We wanted to know if this specific pattern is the result of a late maturation or of a distinct evolution during development. We show here that the embryonic and the fetal MHC and the MLC1emb are expressed throughout perinatal and postnatal masseter development. Our results also demonstrate that MLC1emb accumulation increases considerably during the postnatal period. In addition, both the slow MLCs and the slow isoform of tropomyosin are expressed later in the masseter than quadriceps and the fast skeletal muscle isoform MLC3 is not detected during fetal and early postnatal development in the masseter whereas it is expressed throughout fetal development in the quadriceps. Our results thus confirm previous histochemical data and demonstrate that the masseter muscle displays a pattern of myosin and tropomyosin isoform transitions different to that previously described in trunk and limb muscles. This suggests that control of masseter muscle development involves mechanisms distinct from other body muscles, possibly as a result of either its craniofacial innervation or of a possibly different embryonic origin.

This study was prompted by the renewed clinical interests in understanding the natural history or early morphogenesis of the human temporomandibular joint. Using histologic preparations of 52 representative human embryos and fetuses, each of the major components of the joint was systematically assessed for its changing structure and related to an approximate time scale. The emergence and continued morphogenesis of the joint articular fossa, mandibular condyle, disk, capsule, lateral pterygoid muscle fibers, and both joint cavities occur in regular temporal and spatial pattern. A key observation from this study of embryos and fetuses ranging in age from 32 days to 22 weeks is that each of the component parts of the TMJ progressively emerge with some kind of continuity from a common mass of embryonic mesenchyme interposed between the future temporal bone and mandibular regions. The observations of this study lead to the suggestion that significant developmental disturbances to this common tissue mass or "developmental field" can lead to anomalous morphogenesis of those structures expected to emerge over time from the "developmental field" of the temporomandibular joint. Timing and the identification of a critical time period for the joint are important variables. This study identifies the critical period in the early morphogenesis of TMJ structures as generally falling between the early 7th and 11th prenatal weeks.

The previous study in this series demonstrated that the ingrowth of the central axons of the trigeminal (V) ganglion is prerequisite to V motor axon outgrowth and somatic translocation. In the present experiment we determined whether further interactions with V ganglion cell bodies were required by V motoneurons after the V ganglion innervates the brainstem. Soon after the ganglion axons had penetrated the brainstem they were severed, and a barrier, either permeable or impermeable, was placed between the ganglion cell bodies and the metencephalon. V motor axons grew along aberrant pathways to circumvent the impermeable barriers, many rerouting to reach the V ganglion. Only those V motor nerves which contacted the V ganglion distal to the barrier reached their target musculature in the mandible. The pattern of migration of V motoneurons was normal regardless of the V motor nerve trajectory, but the cell bodies of those axons which did not reach a muscle were not fully differentiated. When permeable barriers (Millipore filters) were implanted, the nerves followed two types of trajectories. If the pore size of the filter was small (0.45 and 0.025 microns), the V motor nerves grew identically to those observed in embryos in which impermeable barriers had been implanted. If the pore size of the filter was large (8.0 and 0.08 microns), the V motor nerve grew along its normal path directly to the barrier. Small axonal bundles from these nerves frequently grew into the filter toward the distal V ganglion. These results indicate that V motor axons preferentially grow to the V ganglion perikarya after exiting from the brainstem. Contact with the V ganglion always results in V motor nerve growth to the mandible while growth of the V motor axons to aberrant target sites only occurs when the axons fail to contact the V ganglion cells distal to the barrier.

The mesenchymal origin muscular tissues entailing some difficulties in the knowledge of certain muscles embryogeny, two methods are therefore applied: the comparison of the ontogenesis data with those of comparative anatomy and the use as a guide of the functional motor unit with which the muscle combines. In resorting to these methods necessary precautions are precisely defined. The origins of the concerned muscles are, at first, situated among those of the whole musculature subjected to will, which are of two types "somitic" and "branchial". The realities lying under these two misleading terms are analysed. Then the usual data on each of the muscular groups embryogeny reviewed and compared, if necessary, with recent works. For the facial muscles a confusion results from the use of the term platysma both in comparative anatomy and in embryology, in pursuance of transposition, exact on that particular point, of the philogenic development of these muscles in ontogenesis. The development of these muscles comes in the scope of the extensive general character of the superficial hyoïd arch derivatives, and their topographic and functional particularities should be brought together with the disposition of the facial nerve nucleus and its arising fibres. Among the muscles in action in the mastication, a link appears between their precise role in this function and their embryogeny. For the neck muscles, the spinal nerve systematisation and the variations of the sterno-mastoïd muscle in mammals agree in assigning to this muscle an entire somitic origin lying exactly at the junction with the so-called branchial muscles. The somitic origin of the tongue muscles and the sharing between the somitic and branchial origins of those of the soft palate bring to light the place in the organism of these two anatomic structures. Then, the conjunction of muscles proceeding thus from two origins in these anatomic structures carries a particular signification in man by reason of the language.

Horseradish peroxidase-wheat germ agglutinin conjugate (HRP) was injected into masticatory and eye muscles of 6- to 15-day-old chick embryos and posthatched chicks to establish the timetable of axonal outgrowth and distribution of central and peripheral terminations of mesencephalic trigeminal neurones (MTN). HRP-labelled MTN first appeared on the 10th day of incubation and by the 14th day most of MTN became labelled, indicating that axonic outgrowth to peripheral targets occurred between the 10th and 14th days of incubation. Peripheral targets included the pterygoideus lateralis and medialis, protractor quadratus, pseudotemporalis superficialis and profundus, and adductor mandibulae of jaw-closing muscles. HRP-filled central axonic processes of MTN were identified first on the 13th day of incubation and they terminated exclusively in the motor nucleus of the trigeminal nerve. Between the 10th and 13th days of incubation, at the peak of naturally occurring cell death in the MTN pool, a consistently lower percentage of neurones could be labelled with HRP than in older embryos and in posthatched chicks. This finding suggests that many MTN die before establishing contact with peripheral targets.

Since experimental and clinical evidence supports some role of musculature in determining the form and size of facial bones during the active periods of growth after birth, this study addresses the same basic relationships between muscle and bone during the periods of active growth before birth. The relationship between the masseter muscle and the mandible, including its ramal and body components, was chosen as the model for study in nineteen human fetuses (ages 16 to 36 weeks). Cross-sectional cephalometric data indicated that, although increases in the size of the muscle and mandible were linearly related to increasing age, the ramal portion of the mandible was more closely related to changes in the masseter muscle than to changes in the mandibular body. Moreover, it appears that reorientation of the muscle anteriorly and downward precedes a similar reorientation of the ramus, with the combination of both fetal events leading to the typical relationships of the two structures expected after birth. Although this study does not get to cause-and-effect relationships, and although the fetal specimens cannot be monitored longitudinally over time, the parallelisms between our prenatal findings and those reported for postnatal periods certainly lend further support to the observation that many aspects of morphogenesis and growth are continual processes spanning the periods on either side of birth.

The skeletal muscle tissue of the lip, tongue, soft palate and masticatory muscles have been morphometrically studied in 70 human fetuses (4-10-month-old). In the muscles mentioned decrease in the nuclear volume and in the nuclear-cytoplasmic ratio is observed. Increase of the muscle fibres in the area of transversal sections is especially intensive during last three months. The labial and glossal fibres reach the greatest thickness. On the 6th month and further, there is an essential difference in number of myons with nuclei in the transversal section of the tongue and lips, as compared with the masticatory muscle and soft palate. In 9-10-month-old fetuses the index on saturation of the glossal and labial muscle fasciculi with capillaries is much greater than that of the soft palate and the masticatory muscle. There exists a statistically important and rather high positive correlation between the area of transversal sections of the muscle fibres, the number of nuclei in them and the density of capillaries in the muscle fasciculi. The relation between histochemical profile of the muscles under investigation and the number of capillaries in them is discussed.

The objective of these experiments was to determine the embryonic origins of craniofacial and cervical voluntary muscles and associated connective tissues in the chick. To accomplish this, suspected primordia, including somitomeres 3-7, somites 1-7, and cephalic neural crest primordia have been transplanted from quail into chick embryos. Quail cells can be detected by the presence of a species-specific nuclear marker. The results are summarized as follows: (table; see text) These results indicate that muscles associated with branchial arch skeletal structures are derived from paraxial mesoderm, as are all other voluntary muscles in the vertebrate embryo. Thus, theories of vertebrate ontogeny and phylogeny based in part on proposed unique features of branchiomeric muscles must be critically reappraised. In addition, many of these cephalic muscles are composites of two separate primordia: the myogenic stem cells of mesodermal origin and the supporting and connective tissues derived from the neural crest or lateral plate mesoderm. Defining these embryonic origins is a necessary prerequisite to understanding how the mesenchymal primordia of cephalic muscles and connective tissues interact to form patterned, species-unique musculoskeletal systems.

The muscular dysgenesis (mdg) mutation in mice causes developmental arrest of myogenesis at the myotube stage. Stained and cleared skeletal preparations and histological sections of the heads of mdg/mdg and normal mouse fetuses were compared in order to determine the effects of the mutation on craniofacial morphology. Muscles of mastication were more severely affected than tongue muscles. Mutant crania had more domed vaults and slender zygomatic arches that were displaced dorsally. The posterior part of the dysgenic mandible was bent dorsally and compressed dorsoventrally. These shape changes may be the distorting effects of persistent cervical kyphosis. Measurements of camera lucida tracings indicated that the lengths of the total mandible, the angular cartilage and process, and the condylar cartilage and process were similar between mutant and control mice. However, the widths of the condylar and angular cartilages were greatly decreased in the mutants. Moreover, the cartilages of dysgenic mice showed no size increase between 18 and 20 days of gestation. The diminished cartilages presumably result from the absence of mechanical loading from prenatal muscle function. For both condylar and angular cartilages, normal muscle activity should produce compression along the long axis of the cartilage and bending/shearing perpendicular to the long axis. It is argued that these forces are more important in promoting growth in cartilage width than length, accounting for the selective reduction of widths in the dysgenic mutant.

The pterygomandibular raphe as described in current anatomy textbooks is not supported by actual observations in cadavers. A study was made on 60 adult Caucasian and Negro cadavers (52 right and 58 left sides, giving a total of 110 sides) providing comparison with an earlier study on Japanese specimens. In addition, 50 fetuses (25 mm crown-rump length to term) were examined to determine the arrangement of the raphe prenatally. Variations in the morphology of the raphe region were classified into three types: Type A--only the upper portion of the raphe could be identified and had a broad, triangular shape. Type B--the buccinator and superior pharyngeal constrictor muscles were widely separated by a broad, fascial region. Type C--the raphe was absent with complete continuity of the buccinator and superior pharyngeal constrictor muscles. A prominent, narrow, tendinous band with attachments as described in current textbooks was never found in adults. There was a complete absence of the raphe in 36% of the specimens resulting in continuity of the buccinator and superior pharyngeal constrictor muscles (type C). However, the remaining adult specimens (64%) exhibited some form of a broad, fascial region that either completely (type B, 36%) or partially (type A, 28%) separated the two muscles. All of the fetuses exhibited the type B arrangement exclusively, indicating that changes in the shape of the raphe occur postnatally. The frequency of appearance of the raphe types in adults differs significantly according to race.

The existence of meniscocapsular insertions of the temporal, masseter and external pterygoid muscles complicates the scheme of capsulo-meniscal dynamics. Our findings do indeed agree with those of DUBECQ (Bordeaux); but we think that the insertions of the masseter and the temporal are not only fine tracts. In the embryon, the meniscus is the preglossal mekelian conjunctival blastema, which receives the 3 masticatory muscles on its anterior border. In the adult, menisco-capsulo-muscular relationships are not modified; inspite of considerable functional adaptation of the articulation to varied stimuli, the menisco-capsular apparatus seems to be triply controlled by 3 musculo-masticatory bands, owing to the anterior premeniscal tendinous lamina, in histological continuity with the meniscus and rich in corpuscles of deep sensitivity. The resultant of the tridirectional muscular traction of the masseter, external pterygoid and temporal is a force in the postero-anterior oblique direction, downwards and forwards, which allows the meniscus to stretch, as was shown by Pr Delaire, and thus to have a sub-temporal sliding pathway of 8 to 12 mm. The three muscle bundles external pterygoid, temporal and masseter constitute the dynamic complex of the meniscus.

The purpose of this investigation was: (a) to study the developmental relationships of the temporal muscle and the coronoid process during the critical initial stages of morphogenesis and (b) to correlate the developmental stages of the muscle and the bone with data describing the functioning of the muscles of mastication in utero. The heads of 41 human embryos and fetuses, 6 to 11 weeks, estimated fertilization age, were sectioned and examined under light microscopy. The findings are described in terms of six successive stages, each characterized by a major developmental change occurring during that stage. The data indicate that the temporal and masseter muscle anlagen begin to develop prior to the skeleton to which they ultimately become attached. The coronoid process differentiates subsequently as a discrete entity within the mass of the temporal muscle anlage at an estimated fertilization age of 7 to 7.5 weeks (23-24 mm CRL). At approximately eight weeks of age, the coronoid process unites with the main portion of the mandibular ramus. The findings here presented do not support the conclusion that the coronoid process is self-differentiating as Washburn ('47) contended. Instead, the development of this feature of the human mandible represents a response that follows the differentiation of the temporal muscle. This conclusion is consistent with the observations drawn from a number of investigations concerning structural and functional development of the face.

The relationship of muscle to periosteum, tendon and ultimately to bone was studied in histological sections of rat mandibles aged 19 days in utero to 56 days postnatal. The following stains were employed: Van Gieson, Masson trichrome, Couciero and Friere, Gomori, orcinol-new fuchsin and Fullmer. Three modes of muscle attachment to the growing rat mandible were described according to the arrangement of collagen and reticular fibres at the interface between the bone and muscles. The morphological characteristics of the three modes persisted even in 56-day-old rats. The structure of the muscle-bone interface suggested a single mechanism of adaptation of muscles to bone during growth. With maturation, the size, number and variety of cells decreased and the size and thickness of collagen fibres increased in the muscle attachments. Each muscle was attached to the mandible by one or more types of attachment defined in this study. A schematic map of the distribution of the different attachment modes associated with muscles of the rat mandible was constructed. Distinction between these modes of attachment may be important in relating the form of a bone to its function.

The canine masticatory muscles contain a unique adult fiber type composition and different contractile protein isoforms than do adult limb muscles. To determine when these characteristic proteins are expressed during development, samples from canine temporalis (masticatory) and pectineus (limb) muscles were compared between 55 days gestation and 60 days postpartum by histochemical, biochemical, and immunocytochemical analysis. At 55 days gestation and 3 days postpartum, both muscles contained identical histochemical type 2C fibers, native myosin isozymes, and myosin light and heavy chains. By 14 days postpartum, fiber-type expression in these muscles diverged, with resultant formation of type 1 and type 2M fibers in the temporalis muscle and type 1 and 2A fibers in the pectineus muscle. The distinctive myosin isoforms, light chains, and heavy chain of the temporalis muscle were also expressed 2 weeks postpartum. Based on the methods used in this study, we conclude that (1) the temporalis muscle develops from embryonic fibers that initially contain a myosin indistinguishable from embryonic limb muscle fibers, suggesting they have a common precursor, and (2) the myosin light chains and heavy chain unique to the temporalis muscle are initially expressed 2 weeks postpartum.