Morphological evolution of the mammalian jaw adductor complex

The evolution of the mammalian jaw during the transition from non‐mammalian synapsids to crown mammals is a key event in vertebrate history and characterised by the gradual reduction of its individual bones into a single element and the concomitant transformation of the jaw joint and its incorporation into the middle ear complex. This osteological transformation is accompanied by a rearrangement and modification of the jaw adductor musculature, which is thought to have allowed the evolution of a more‐efficient masticatory system in comparison to the plesiomorphic synapsid condition. While osteological characters relating to this transition are well documented in the fossil record, the exact arrangement and modifications of the individual adductor muscles during the cynodont–mammaliaform transition have been debated for nearly a century. We review the existing knowledge about the musculoskeletal evolution of the mammalian jaw adductor complex and evaluate previous hypotheses in the light of recently documented fossils that represent new specimens of existing species, which are of central importance to the mammalian origins debate. By employing computed tomography (CT) and digital reconstruction techniques to create three‐dimensional models of the jaw adductor musculature in a number of representative non‐mammalian cynodonts and mammaliaforms, we provide an updated perspective on mammalian jaw muscle evolution. As an emerging consensus, current evidence suggests that the mammal‐like division of the jaw adductor musculature (into deep and superficial components of the m. masseter, the m. temporalis and the m. pterygoideus) was completed in Eucynodontia. The arrangement of the jaw adductor musculature in a mammalian fashion, with the m. pterygoideus group inserting on the dentary was completed in basal Mammaliaformes as suggested by the muscle reconstruction of Morganucodon oehleri. Consequently, transformation of the jaw adductor musculature from the ancestral (‘reptilian’) to the mammalian condition must have preceded the emergence of Mammalia and the full formation of the mammalian jaw joint. This suggests that the modification of the jaw adductor system played a pivotal role in the functional morphology and biomechanical stability of the jaw joint.     

Morphology of the Jaw-Closing Musculature in the Common Wombat (Vombatus ursinus) Using Digital Dissection and Magnetic Resonance Imaging

Wombats are unique among marsupials in having one pair of upper incisors, and hypsodont molars for processing tough, abrasive vegetation. Of the three extant species, the most abundant, the common wombat (Vombatus ursinus), has had the least attention in terms of masticatory muscle morphology, and has never been thoroughly described. Using MRI and digital dissection to compliment traditional gross dissections, the major jaw adductor muscles, the masseter, temporalis and pterygoids, were described. The masseter and medial pterygoid muscles are greatly enlarged compared to other marsupials. This, in combination with the distinctive form and function of the dentition, most likely facilitates processing a tough, abrasive diet. The broad, flat skull and large masticatory muscles are well suited to generate a very high bite force. MRI scans allow more detail of the muscle morphology to be observed and the technique of digital dissections greatly enhances the knowledge obtained from gross dissections.     

Morphological analysis of tendinous structure in the American alligator jaw muscles

In the American alligator, the jaw muscles show seven bundles of tendinous structure: cranial adductor tendon, mandibular adductor tendon, lamina anterior inferior, trap-shaped lamina lateralis, lamina intramandibularis, lamina posterior, and depressor mandibular tendon (originating from the musculus depressor mandibulae, m. pseudotemporalis, m. adductor mandibulae posterior, m. adductor mandibulae externus, m. intramandibularis, m. pterygoideus anterior, and m. pterygoideus posterior). These tendinous structures are composed of many collagen fibrils and elastic fibers; however, the distributions and sizes of the fibers in these tendinous components differ in comparison with those of other masticatory muscles. The differences of these properties reflect the kinetic forces or the stretch applied to each tendon by the muscle during jaw movements in spite of the simple tendon-muscle junctions. © 1993 Wiley-Liss, Inc.     

The shrew tamed by Wolff's law: Do functional constraints shape the skull through muscle and bone covariation?

Bone is a highly plastic tissue that reflects the many potential sources of variation in shape. Here, we focus on the functional aspects of bone remodeling. We choose the skull for our analyses because it is a highly integrated system that plays a fundamental role in feeding and is thus, likely under strong natural selection. Its principal mechanical components are the bones and muscles that jointly produce bite force and jaw motion. Understanding the covariations among these three components is of interest to understand the processes driving the evolution of the feeding apparatus. In this study, we quantitatively and qualitatively compare interactions between these three components in shrews from populations known to differ in shape and bite force. Bite force was measured in the field using a force transducer and skull shape was quantified using surface geometric morphometric approaches based on µCT‐scans of the skulls of same individuals. The masseter, temporalis, pterygoideus, and digastricus muscles of these individuals were dissected and their cross sectional areas determined. Our results show strong correlations between bite force and muscle cross sectional areas as well as between bite force and skull shape. Moreover, bite force explains an important amount of skull shape variation. We conclude that interactions between bone shape and muscle characteristics can produce different morpho‐functional patterns that may differ between populations and may provide a suitable target for selection to act upon. J. Morphol. 276:301–309, 2015. © 2014 Wiley Periodicals, Inc.     

Non-specific esterases and esterproteases in masticatory muscles from the muscular dystrophic mouse

With the aid of histochemical and electrophoretic techniques activities for esterase and esterprotease were investigated in the digastric and masseter muscles from normal and dystrophic mice. The substrates used were -naphthyl acetate and N-acetyl-l-alanine -naphthyl ester. According to the microscopic observations of the dystrophic muscles the histopathological changes in the masseter muscle were much more pronounced than in the digastric muscle. The connective tissue surrounding the myofibers of the dystrophic masseter contained a large number of cells with pronounced enzyme activity. Among them were mast cells that were strongly stained for esterprotease. The connective tissue of the dystrophic digastricus was much less infiltrated with cellular elements reacting for esterprotease. In zymograms the normal digastricus, the dystrophic masseter and the dystrophic digastricus showed a strong activity for certain isoenzymes that were absent or weakly expressed in the normal masseter.This study was supported by grand No. 12-6516 from the Danish Medical Research Council     

Aging affects different human muscles in various ways. An image analysis of the histomorphometric characteristics of fiber types in human masseter and vastus lateralis muscles from young adults and the very old

This study is an attempt to objectively evaluate age-related changes in human muscles by use of histomorphometric methods. Aging in humans induces dramatic transformations in the skeletal muscles but little is known as to whether or not the aging processes per se may affect all muscles equally. In this study aging of two human muscles with different functions, origin and nerve supply is compared. Sections were cut from masseter and vastus lateralis muscles obtained from young adults aged 18-24 years and from the very old aged 90-102 years. Muscle fiber types were classified with the traditional myofibrillar ATPase staining. Various histomorphometric parameters of the different fiber types in human masseter and vastus lateralis muscle sections were obtained by image analyses to evaluate the age-related changes in the muscle fibers. The following variables were calculated: the number of each fiber type per photographed area; the area of each fiber and two indicators for the shape of the muscle fibers. In the aging muscles there was no relative preferential loss of a fiber type. High numbers of intermediate ATPase-stained fibers (IM fibers) were found in some old vastus muscles but were only sporadic in young vastus muscles. However, there was no change in the percentage distribution of intermediate ATPase-stained fibers when young and very old human masseter muscles were compared. Incubation of the sections with antimyosin antibodies showed that the IM fibers in old masseter and old vastus contained different myosin heavy chains. Thus ATPase activity and anti-myosin staining displayed a somewhat different pattern of fiber type distribution. The main changes in the shape and area indicated that type I fibers in the masseter became more circular while in the vastus they decreased significantly in size. The type II fibers in the vastus became very small and deviated significantly from circularity whereas the type II fibers in the masseter only exhibited a decrease in the size of the fibers. Histomorphometric measurements show that aging affects different human muscles in various ways.     

The relationships among jaw‐muscle fiber architecture,jaw morphology,and feeding behavior in extant apes and modern humans

The jaw‐closing muscles are responsible for generating many of the forces and movements associated with feeding. Muscle physiologic cross‐sectional area (PCSA) and fiber length are two architectural parameters that heavily influence muscle function. While there have been numerous comparative studies of hominoid and hominin craniodental and mandibular morphology, little is known about hominoid jaw‐muscle fiber architecture. We present novel data on masseter and temporalis internal muscle architecture for small‐ and large‐bodied hominoids. Hominoid scaling patterns are evaluated and compared with representative New‐ (Cebus) and Old‐World (Macaca) monkeys. Variation in hominoid jaw‐muscle fiber architecture is related to both absolute size and allometry. PCSAs scale close to isometry relative to jaw length in anthropoids, but likely with positive allometry in hominoids. Thus, large‐bodied apes may be capable of generating both absolutely and relatively greater muscle forces compared with smaller‐bodied apes and monkeys. Compared with extant apes, modern humans exhibit a reduction in masseter PCSA relative to condyle‐M₁ length but retain relatively long fibers, suggesting humans may have sacrificed relative masseter muscle force during chewing without appreciably altering muscle excursion/contraction velocity. Lastly, craniometric estimates of PCSAs underestimate hominoid masseter and temporalis PCSAs by more than 50% in gorillas, and overestimate masseter PCSA by as much as 30% in humans. These findings underscore the difficulty of accurately estimating jaw‐muscle fiber architecture from craniometric measures and suggest models of fossil hominin and hominoid bite forces will be improved by incorporating architectural data in estimating jaw‐muscle forces. Am J Phys Anthropol 151:120–134, 2013. © 2013 Wiley Periodicals, Inc.     

Contour analysis of masticatory jaw movements and muscle activity in Macaca mulatta

Frontal plane mandibular movements during mastication and the associated electromyographic (EMG) activity for left and right superficial masseter, posterior temporalis, anterior temporalis, and anterior belly of the digastric (ABD) were studied for two adult male Macaca mulatta by the new technique of “contour” analysis. Contour analysis allowed graphic and quantitative portrayal of multiple chew cycle patterns of mandibular movement and EMG activity during active mastication. A series of computer programs (ATS, ATSED, ATSXYZ) facilitated the collection, editing and definition, and finally processing of these masticatory data into contour plots. These preliminary data indicated the essential symmetry of mandibular movement patterns, high chew cycle variability inferior to occlusion, multiple centers of intense EMG activity for balancing-side superficial masseter, and no difference between working-side anterior and posterior temporalis EMG patterns. Maximum EMG amplitude was found in the area of buccal phase power stroke (BPS). Maximum EMG amplitude for ABD was located medial and inferior to occlusion; all other muscle maximum amplitudes were buccal and inferior to occlusion. The location of maximum EMG amplitudes for superficial masseter and ABD were closer to occlusion (more superior) during mastication of carrot than were maximum amplitudes during biscuit mastication. The absence of any detectable shift of EMG maximum amplitude location between biscuit and carrot for posterior and anterior temporalis suggested, along with the continuous EMG activity of working-side posterior temporalis, a secondary role for the temporalis (compensation for superficial masseter activity) during active mastication.     

Masticatory Muscle Anatomy and Feeding Efficiency of the American Beaver, <Emphasis Type="Italic">Castor canadensis</Emphasis> (Rodentia,Castoridae)

Beavers are well-known for their ability to fell large trees through gnawing. Yet, despite this impressive behavior, little information exists on their masticatory musculature or the biomechanics of their jaw movements. It was hypothesized that beavers would have a highly efficient arrangement of the masticatory apparatus, and that gnawing efficiency would be maintained at large gape. The head of an American beaver, Castor canadensis, was dissected to reveal the masticatory musculature. Muscle origins and insertions were noted, the muscles were weighed and fiber lengths measured. Physiological cross-sectional areas were determined, and along with the muscle vectors, were used to calculate the length of the muscle moment arms, the maximum incisor bite force, and the proportion of the bite force projected along the long axis of the lower incisor, at occlusion and 30° gape. Compared to other sciuromorph rodents, the American beaver was found to have large superficial masseter and temporalis muscles, but a relatively smaller anterior deep masseter. The incisor bite force calculated for the beaver (550–740 N) was much higher than would be predicted from body mass or incisor dimensions. This is not a result of the mechanical advantage of the muscles, which is lower than most other sciuromorphs, but is likely related to the very high percentage (>96 %) of bite force directed along the lower incisor long axis. The morphology of the skull, mandible and jaw-closing muscles enable the beaver to produce a very effective and efficient bite, which has permitted beavers to become highly successful ecosystem engineers.     

Mastication and the postorbital ligament: dynamic strain in soft tissues

Although the FEED database focuses on muscle activity patterns, it is equally suitable for other physiological recording and especially for synthesizing different types of information. The present contribution addresses the interaction between muscle activity and ligamentary stretch during mastication. The postorbital ligament is the thickened edge of a septum dividing the orbital contents from the temporal fossa and is continuous with the temporal fascia. As a tensile element, this fascial complex could support the zygomatic arch against the pull of the masseter muscle. An ossified postorbital bar has evolved repeatedly in mammals, enabling resistance to compression and shear in addition to tension. Although such ossification clearly reinforces the skull against muscle pull, the most accepted explanation is that it helps isolate the orbital contents from contractions of the temporalis muscle. However, it has never been demonstrated that the contraction of jaw muscles deforms the unossified ligament. We examined linear deformation of the postorbital ligament in minipigs, Sus scrofa, along with electromyography of the jaw muscles and an assessment of changes in pressure and shape in the temporalis. During chewing, the ligament elongated (average 0.9%, maximum 2.8%) in synchrony with the contraction of the elevator muscles of the jaw. Although the temporalis bulged outward and created substantial pressure against the braincase, the superficial fibers usually retracted caudally, away from the postorbital ligament. In anesthetized animals, stimulating either the temporalis or the masseter muscle in isolation usually elongated the ligament (average 0.4-0.7%). These results confirm that contraction of the masticatory muscles can potentially distort the orbital contents and further suggest that the postorbital ligament does function as a tension member resisting the pull of the masseter on the zygomatic arch.     

Association between the lack of teeth and the expression of myosins in masticatory muscles of microphthalmic mouse

The purposes of the present study were to elucidate the influences of the deficiency of teeth on masticatory muscles, such as the masseter, temporalis and digastric muscles and compare the influence among masticatory muscles. We analysed the expressions of myosin heavy chain (MyHC) isoform messenger RNA (mRNA) and protein in these muscles in the microphthalmic (mi/mi) mouse, whose teeth cannot erupt because of a mutation in the mitf gene locus. The expression levels of MyHC mRNA and protein in the masseter, temporalis, digastric, tibialis anterior and gastrocnemius muscles of +/+ and mi/mi mice were analysed with real‐time polymerase chain reaction and sodium dodecyl sulfate‐polyacrylamide gel electrophoresis, respectively. The mi/mi masseter muscle at 8 weeks of age expressed 4·1‐fold (p < 0·05) and 3.3‐fold (p < 0·01) more MyHC neonatal mRNA and protein than that in the +/+, respectively; the expression level of MyHC neonatal protein was 19% of the total MyHC protein in the masseter muscle of mi/mi mice. In the digastric muscle, the expression levels of MyHC I mRNA and protein in the mi/mi mice were 4·7‐fold (p < 0·05) and 5‐fold (p < 0·01) higher than those in the +/+ mice. In the temporalis, tibialis anterior and gastrocnemius muscles, there was no significant difference in the expression levels of any MyHC isoform mRNA and protein between +/+ and mi/mi mice. These results indicate associations between the lack of teeth and the expression of MyHC in the masseter and digastric muscles but not such associations in the temporalis muscle, suggesting that the influence of tooth deficiency varies among the masticatory muscles. Copyright © 2011 John Wiley & Sons, Ltd.     

Functional assessment of subfamily variation in maxillomandibular morphology among Old World monkeys

Among Old World monkeys, subfamily variation in maxillomandibular form is commonly attributed to divergent dietary and social behaviors. However, our knowledge of any musculoskeletal adaptations for gape in cercopithecines, and folivory in colobines, is incomplete. Such data are requisite to a more informed perspective on the evolutionary morphology of these taxa. Structural analyses of gape and biomechanical efficiency were applied to a representative sample of adult cercopithecids. Factors pertaining to the biomechanical scaling of cranial structures were evaluated with least-squares bivariate regression techniques. To assess subfamily differences in masticatory efficiency, analyses of covariance were made between relevant factors. Cercopithecines achieve increased gape and relative canine size mainly with strong positive allometry of the facial skull, combined with a larger gonial angle. Colobines possess a relatively long masseter lever arm and short facial skull, as well as an enlargened masseter-medial pterygoid complex. Subfamily differences in temporalis lever arm scaling are negligible. Biomechanical comparisons within and between subfamilies suggest that the mechanical advantage of the temporalis is relatively greater than that of the masseter, while the mechanical advantage of both muscles increases with face length. Evidence is presented to stress the need for adequate consideration of the dependent variable in allometric investigations of skull form.     

Expression of Masticatory-specific Isoforms of Myosin Heavy-chain,Myosin-binding Protein-C and Tropomyosin in Muscle Fibers and Satellite Cell Cultures of Cat Masticatory Muscle

We test the hypothesis that cat jaw satellite cells belong to a distinct lineage preprogrammed to express masticatory-specific isoforms of myosin heavy-chain (m-MyHC), myosin-binding protein-C (m-MBP-C), and tropomyosin (m-Tm) during myogenesis in vitro. A monoclonal antibody (MAb) against m-MyHC and MAbs raised here against cat m-MBP-C and m-Tm were used to stain cryostat sections of cat masseter muscle and cultured myotubes derived from satellite cells of cat temporalis and limb muscles, using peroxidase immunohistochemistry. MAbs against m-MBP-C bound purified m-MBP-C in Western blots. MAbs against m-Tm failed to react with m-Tm in Western blots, but reacted with native m-Tm in gel electrophoresis–derived ELISA. In cat masseter sections, MAbs against m-MyHC, m-MBP-C, and m-Tm stained all masticatory fibers, but not the jaw-slow fibers. Cat jaw and limb muscle cultures mature significantly more slowly relative to rodent cultures. However, at 3 weeks, all three MAbs extensively stained temporalis myotubes, whereas they apparently stained isolated myotubes weakly in cat limb and rat jaw cultures. We conclude that satellite cells of masticatory fibers are preprogrammed to express these isoforms during myogenesis in vitro. These results consolidate the notion that masticatory and limb muscle allotypes are distinct. (J Histochem Cytochem 58:623–634, 2010)     

The functional significance of the squamosal suture in Australopithecus boisei.

A juvenile Australopithecus boisei specimen from the Omo basin, southern Ethiopia, is found to exhibit and extraordinarily large overlap of the temporal squama on the parietal, a phenomenon shared with at least two adult specimens of A. boisei. An attempt is made to interpret the overlap as a structural (bony/ligamentous) adaptation necessitated by the unique combination of certain components of the masticatory system of A. boisei. These are: (1) the massiveness and strength of the temporalis muscle, (2) its relatively anterior location, and (3) the lateral position of the masseter muscle due to the flaring of the zygomatic arches. The effect of the temporalis muscle is to create excessive pressure on the portion of the squamosal suture along the parietal, while the lateral placement of the masseter and the resultant increase of pressure on the temporal squama via the zygomatic arch tend to "loosen" the contact between the temporal and parietal bones.     

Internal architecture, origin-insertion site, and mass of jaw muscles in Old World hamsters

The jaw muscle (i.e., masticatory, suprahyoid, and extrinsic tongue) anatomy and mass were examined in four genera of Old World hamsters (cricetine murids), Mesocricetus, Cricetulus, Tscherskia, and Phodopus. The masseter was the largest and most complicated of the muscles examined. In the superficial layer, a few ventral fibers form a small medially turned portion with an insertion site more similar to those of sciurids than of other murids. In Mesocricetus, the superficial layer has a discrete anteroventral portion that has not been reported for other murid rodents. Examination of the fiber attachment sites indicated that the deep layer contains four parts and the medial layer contains three parts. The deep layer originates from two aponeuroses that are firmly connected to each other at their anterior ends and lie along the zygomatic arch. The aponeurosis of insertion for the deep layer is situated along the masseteric ridge and the dorsal border of the angular process, but is absent in its middle part, consistent with reports in two relatives, sigmodontine and arvicoline murids. In cricetine murids, unlike in other rodents, fibers insert on the dorsal narrow strip of the posterior mandibular aponeurosis, not on its broad medial aspect. The relative mass of some masticatory and suprahyoid muscles is related to body mass. Small species (Cricetulus and Phodopus) have relatively larger masseter and mylohyoid muscles and smaller temporalis and geniohyoid muscles than large species (Mesocricetus and Tscherskia).     

睡眠剥夺对大鼠行为学及咀嚼肌肌电图的影响

目的：颞下颌关节紊乱病是口腔科的一种常见病和多发病,精神心理因素是颞下颌关节紊乱病的一个主要病因。本文通过观察睡眠剥夺对大鼠行为学及咀嚼肌肌电图的影响,探讨睡眠剥夺在颞下颌关节紊乱病发病中的作用。方法：35只Wistar大鼠,随机分为5组：睡眠剥夺1d组、5d组、9d组、正常对照组和大平台对照组。采用改良多平台睡眠剥夺法（modified multiple plat—formmethod,MMPM）建立大鼠SD模型,观察大鼠行为学及咀嚼肌肌电图的变化。结果：睡眠剥夺1d组和5d组在旷场实验水平得分和垂直得分上均高于对照组,而睡眠剥夺9d组均低于对照组;睡眠剥夺1d、5d和9d组在松弛状态和紧咬状态时颞肌前后束及咬肌的电位均明显高于对照组,且两侧无明显差别,同时,睡眠剥夺组双侧颞肌和咬肌的肌电图静息期较对照组显著延长。结论：睡眠剥夺可使大鼠行为学发生改变并对咀嚼肌肌电图造成影响,这可能是颞下颌关节紊乱病的病因之一,为我们对颞下颌关节紊乱病的预防和治疗提供了一定的理论指导。     

Form and function of the masticatory musculature in the tree sloths, Bradypus and Choloepus

The tree sloths, Bradypus and Choloepus, show unusual masticatory specializations, compared to each other and to other mammals. Both have an incomplete zygomatic arch with descending jugal process, a complex superficial masseter, a large temporalis and medial pterygoid musculature, and a lateral pterygoid with two heads. In Choloepus the deep masseter and zygomaticomandibularis are typical when compared to other mammals. However, in Bradypus there is an ascending jugal process from which enlarged and vertically oriented deep masseter and zygomaticomandibularis muscles originate. Although both sloths are folivores, the anterior teeth in Choloepus are caniniform, while those of Bradypus have lost such elongation. In both sloths the glenoid cavity is similarly located; however, in Bradypus the craniomandibular joint is raised above the occlusal plane, and the pterygoid flanges are elongated. Prediction of the evolutionary sequence of cranial changes from Choloepus-like (primitive) to Bradypus-like (derived) morphology is based upon the most parsimonious model of masseter-medial pterygoid complex changes for masticatory efficiency improvement. The model proposes that the condylar neck in Bradypus was elongated and that this single change predicated a series of other structural changes. Mandibular movement patterns in both sloths showed anteromedially directed unilateral power strokes as in other mammals. Puncture-crushing, tooth-sharpening, and chewing cycles are distinct in Choloepus, less so in Bradypus. The masticatory rate is slow in sloths compared to other mammals of similar body size, averaging 590 ms per cycle for Choloepus and 510 ms for Bradypus.     

Masticatory function in patients with xerostomia

The effects of reduced salivary output in patients suffering from xerostomia on masticatory function has not been previously studied. This study compares masticatory performance and kinematic activity of patients suffering from xerostomia with age-, sex-, and number of occluding pairs-matched healthy controls. Masticatory function was evaluated by assessment of chewing motion and muscle activity during chewing an artificial food (CutterSil®), chewing gum and swallowing a bolus of almond. Chewing motion was recorded with the Optotrak® computer system. Bilateral muscle activity of both masseter and anterior temporalis was recorded using surface electrodes. Results of this study revealed significant differences between patients and controls in their ability to process food and masticatory muscle activity. The majority of patients could not break down the artificial food, others had a larger median particle size than the controls. A significant difference was also observed in the number of chewing cycles required to swallow almonds, the patients required more than twice as many chews as the controls, P<0.001. The right masseter muscle displayed significantly less activity for the patient than the controls. These findings suggest that patients with xerostomia exhibit reduced ability to process food. The observed decline in masticatory performance is probably due to reduced activity of the muscles of mastication.     

Mechanism of paradoxical resistance of the digastric muscle to the action of tetanus toxin

Excitability parameters of m. masseter and m. digastricus fibers (resting minute potential--MP, action potential--AP, critical level of depolarization--CLD and rheobase currents) were determined in anesthetized rats (nembutal, 40 mg/kg) on days 3-4 and 10-15 after local intramuscular tetanus toxin injection (20 and 40 mouse Dlm, respectively). The development of local tetanus in m. masseter was characterized by typical excitability changes--a decrease in MP and AP. Local spasm in m. digastricus under analogous conditions of tetanus toxin injection was absent, while excitability tests demonstrated MP, AP and CLD augmentation. As m. digastricus motoneurons are devoid of inhibitory synaptic regions, these nervous cells are not affected by tetanus toxin.     

Selective expression of the type 3 isoform of ryanodine receptor Ca2+ release channel (RyR3) in a subset of slow fibers in diaphragm and cephalic muscles of adult rabbits

The expression pattern of the RyR3 isoform of Ca2+ release channels was analysed by Western blot in neonatal and adult rabbit skeletal muscles. The results obtained show that the expression of the RyR3 isoform is developmentally regulated. In fact, RyR3 expression was detected in all muscles analysed at 2 and 15 days after birth while, in adult animals, it was restricted to a subset of muscles that includes diaphragm, masseter, pterygoideus, digastricus, and tongue. Interestingly, all of these muscles share a common embryonic origin being derived from the somitomeres or from the cephalic region of the embryo. Immunofluorescence analysis of rabbit skeletal muscle cross-sections showed that RyR3 staining was detected in all fibers of neonatal muscles. In contrast, in those adult muscles expressing RyR3 only a fraction of fibers was labelled. Staining of these muscles with antibodies against fast and slow myosins revealed a close correlation between expression of RyR3 and fibers expressing slow myosin isoform.