Relationship between craniofacial skeletal patterns and anatomic characteristics of masticatory muscles: a systematic review and meta-analysis.

BACKGROUND: The anatomic characteristics of the masticatory muscles differ across craniofacial skeletal patterns. OBJECTIVE: To identify differences in the anatomic characteristics of masticatory muscles across different sagittal and vertical craniofacial skeletal patterns. ELIGIBILITY CRITERIA: Studies measuring the thickness, width, cross-sectional area (CSA), volume and orientation of masticatory muscles in healthy patients of different sagittal (Class I, Class II, and Class III) and/or vertical (normodivergent, hypodivergent, and hyperdivergent) patterns. INFORMATION SOURCES: Unrestricted literature searches in 8 electronic databases/registers until December 2023. RISK OF BIAS AND SYNTHESIS OF RESULTS: Study selection, data extraction, and risk of bias assessment with a customised tool were performed independently in duplicate. Random-effects meta-analysis and assessment of the certainty of clinical recommendations with the GRADE approach were conducted. RESULTS: 34 studies (37 publications) were selected with a total of 2047 participants and data from 16 studies were pulled in the meta-analysis. Masseter muscle thickness in relaxation was significantly greater by 1.14 mm (95% CI 0.74-1.53 mm) in hypodivergent compared to normodivergent patients while it was significantly decreased in hyperdivergent patients by - 1.14 mm (95% CI - 1.56 to - 0.73 mm) and - 2.28 mm (95% CI - 2.71 to - 1.85 mm) compared to normodivergent and hypodivergent patients respectively. Similar significant differences were seen between these groups in masseter muscle thickness during contraction as well as masseter muscle CSA and volume. Meta-analyses could not be performed for sagittal categorizations due to insufficient number of studies. CONCLUSIONS: Considerable differences in masseter muscle thickness, CSA and volume were found across vertical skeletal configurations being significantly reduced in hyperdivergent patients; however, results should be interpreted with caution due to the high risk of bias of the included studies. These variations in the anatomic characteristics of masticatory muscles among different craniofacial patterns could be part of the orthodontic diagnosis and treatment planning process. REGISTRATION: PROSPERO CRD42022371187 .

Chewing shrews: Examining the morphology and function of the masticatory musculature in Soricidae via diffusible iodine-based contrast-enhanced computed tomography.

Essential for sustaining a high metabolic rate is the efficient fragmentation of food, which is determined by molar morphology and the movement of the jaw. The latter is related to the jaw morphology and the arrangement of the masticatory muscles. Soricid jaw apparatuses are unique among mammals, as the articulation facet on the condylar process is separated into a dorsal and a ventral part, which has often been linked to more differentiated jaw motions. Soricidae also possess a remarkably elongated angular process. However, the precise function of the unique morphology of soricid jaw apparatuses has not been fully understood yet. By digitally reconstructing the masticatory musculature via the diffusible iodine-based contrast-enhanced computed tomography technique, we show how the unique jaw morphology is reflected in the spatial organization as well as the inner architecture and respective fascicle orientations of the muscles. From the lines of action of the m. masseter and the m. pterygoideus internus, both muscles inserting on the lateral and medial side of the angular process, respectively, we infer that the angular process is substantial for roll and yaw rotations of the mandible. The m. masseter is subdivided into four and the m. pterygoideus internus into five subunits, each exhibiting a slightly different line of action and torque. This enables Soricidae to adjust and adapt these rotational movements according to the properties of the ingested food, allowing for more efficient fragmentation. Additionally, those guided rotational motions allow for precise occlusion despite tooth wear. The temporalis is the largest of the adductor muscles and is mainly responsible for exerting the bite force. Overall, the unique jaw bone morphology in conjunction with the complex muscle arrangement may contribute towards a more efficient energy gain and the maintenance of a high metabolic rate, which is crucial for small-bodied mammals such as shrews.

Contribution of the complex comprising the masticatory fascia, disc, and capsule to temporomandibular joint stabilization: An anatomical study.

BACKGROUND: Anterior displacement of the temporomandibular joint (TMJ) disc is the most typical pathological condition of TMJ disorders. Structures attached to the articular disc may support the disc in various directions and contribute to stabilizing the TMJ. However, the relationship between the articular disc, capsule, and masticatory muscles remains unclear. Therefore, this study aimed to clarify the relationship between the masticatory muscles, related masticatory fascia, articular disc, and capsule. METHODS: We examined 10 halves from adult Japanese cadavers, with five halves macroscopically analyzed and the remaining five histologically analyzed. The TMJ was dissected from the lateral aspect for gross anatomical analysis. For histological analysis, the relationship between the temporal and masseteric fasciae and the articular capsule was observed in the coronal section. Additionally, we evaluated relationships among the disc, capsule, temporal fascia, and masseteric fascia in 10 living and healthy volunteers using magnetic resonance imaging. RESULTS: The articular disc was attached to the capsule without a clear border. The capsule continued into the masseteric and temporal fasciae. Consequently, the articular disc, capsule, masseteric, and temporal fasciae were considered a single complex. CONCLUSIONS: The single complex of the temporalis, masseter, capsule, masticatory fascia, and disc may antagonize the force in the posterolateral direction through the fascia.

Evaluating the muscle attachment hypothesis for sagittal cresting in Gorilla and Pongo.

Primate mandibular morphology is often associated with jaw functionality of the masticatory complex in the context of variation in diets. Recent research into the disparities between the diet and jaw functionality in male and female hominoids is inconclusive and suggests that sexual dimorphism in the mandible may be influenced by external factors such as temporalis and masseter muscle morphology, which in turn may be influenced by sexual selection. As the muscles associated with mastication (i.e., the type of chewing exhibited by primates and other mammals) encompass the mandible as well as the neurocranium, including the sagittal crest among some individuals, this study investigates sex-specific associations between regions of the mandibular ramus and neurocranium associated with mastication in a dentally mature sample of Gorilla and Pongo. A total of four cranial and mandibular variables were measured in two Gorilla taxa (Gorilla gorilla gorilla and Gorilla beringei graueri) and one Pongo taxon (Pongo pygmaeus pygmaeus) (n = 220). For all three taxa, we investigate (a) whether the degree of sexual dimorphism in cranial regions associated with sagittal cresting (sagittal crest size (SCS) and temporalis muscle attachment area (TMAA)) is proportional to the degree of mandibular ramus area (MRA) and coronoid process height (CPH) sexual dimorphism, (b) whether there are sex differences in scaling relationships between TMAA and MRA, and (c) whether there are sex differences in the strength of association between TMAA and CPH. We show that for G. g. gorilla, variables associated with sagittal cresting show higher sexual dimorphism values than our two mandibular ramus variables, which is not the case for G. b. graueri or for P. p. pygmaeus. All three taxa show similar sex-specific scaling relationships between TMAA and MRA, where for males this relationship does not diverge from isometry, and for females there is a negative allometric relationship. Our findings also show intraspecific sex differences in allometric slopes between MRA and TMAA for all three taxa. Only G. g. gorilla shows a significant association between TMAA and CPH, which is observed in both sexes. Although there are some statistical associations between the cranial and mandibular regions associated with mastication, our results show that among male gorillas and orangutans, patterns of variation in the sagittal crest, TMAA, mandibular ramus and the coronoid process cannot be explained by the muscle attachment hypothesis alone. These findings have implications surrounding the associations between social behaviour and the morphology of the craniofacial complex.

Springhares, flying and flightless scaly-tailed squirrels (Anomaluromorpha, Rodentia) are the squirrely mouse: comparative anatomy of the masticatory musculature and its implications on the evolution of hystricomorphy in rodents.

Anomaluromorpha is a particularly puzzling suborder of Rodentia. Endemic to Africa, this clade includes the extant genera Idiurus, Anomalurus, Zenkerella, and Pedetes. These rodents present an hystricomorphous condition of the skull, characterized by a large infraorbital foramen, which evolved independently within the mouse-related clade over a span of approximately 57 million years. They exhibit a high disparity in craniomandibular and dental morphology that has kept their phylogenetic affinities disputed for a long time. Given the past significance of masticatory morphotypes in establishing the classification of Rodentia, we propose to explore variations in the masticatory apparatus of Anomaluromorpha in order to evaluate whether its related features can offer additional data for systematics and contribute to our understanding of the complexity of hystricomorphy. In order to do so, we used traditional dissection and diffusible iodine-based contrast-enhanced computed tomography (diceCT) to accurately describe and compare the anatomy of the specimens. We found that the muscle morphology displays clear differentiation among each anomaluromorph taxonomic unit. Specifically, the masseteric complex of Anomaluromorpha exhibits distinctive synapomorphies such as the infraorbital part of the zygomaticomandibularis muscle being separated into a rostral and orbital part and an absence of a posterior part of the zygomaticomandibularis. Additionally, the orbital portion of the infraorbital part originates from a well-marked ridge and fossa at the level of its area of origin on the anteromedial wall of the orbital cavity, a feature that is absent in other members of the mouse-related clade. This evident bony feature, among others, is strongly associated with muscular anatomy and can contribute to ascertaining the taxonomic status of extinct representatives of the clade. Finally, we showed that the hystricomorphy of Anomaluromorpha largely differs from those of Ctenohystrica and Dipodoidea and that the definition of this morphotype is complex and cannot be reduced simply to the size of the opening of the infraorbital foramen.

Three-dimensional Analysis of the Muscles Related to the So-Called "Pterygomandibular Raphe": An Anatomical and Histological Study.

The pterygomandibular raphe (PMR) is a tendinous bundle between the bucinator (BM) and the superior constrictor of pharynx (SC) and has been considered essential for swallowing. Despite its functional significance, previous studies reported that the PMR is not always present. Another study reported presence of the connecting fascia between the BM and deep temporalis tendon (dTT). Therefore, the present study analyzed the three-dimensional relationship between the BM, SC, and dTT. We examined 13 halves of 11 heads from adult Japanese and Caucasian cadavers: eight halves macroscopically and five halves histologically. There was no clear border between the BM and SC in any specimens macroscopically. The BM attachment varied depending on its levels. At the level of the superior part of the internal oblique line, the BM fused with the SC with no clear border. At the level of the midpart of the internal oblique line of the mandible, the BM attached to the dTT directly, and the SC attached to the dTT via collagen fibers and the BM. Based on these results, these muscles should be described as the BM/dTT/SC (BTS) complex. The three-dimensional relationship of the BTS complex might result in the so-called "pterygomandibular raphe." The BTS complex could be important as a muscle coordination center in chewing and swallowing.

Masticatory myology of the llama (Lama glama, Camelidae) and comparisons with other camelids and euungulates.

Camelids are the only living representatives of the Suborder Tylopoda, and present a unique set of osteo-myological masticatory features, differing from all other extant euungulates. They combine selenodont dentition and rumination with a fused symphysis, and roughly plesiomorphic muscle proportions. Despite its potential relevance as an euungulate model in comparative anatomy studies, the available data is strikingly scarce. The present study represents the first description of the masticatory muscles of a Lamini, analyzing the functional morphology of Lama glama and other camelids in a comparative framework. Both sides of the head of three adult specimens from Argentinean Puna were dissected. Descriptions, illustrations, muscular maps, and weighing of all masticatory muscles were performed. Some facial muscles are also described. The myology of llamas confirms that camelids possess relatively large temporalis muscles, with Lama being less extreme than Camelus. This plesiomorphic feature is also recorded in suines and some basal euungulates. Conversely, the direction of the fibers of the M. temporalis is mainly horizontal, resembling grinding euungulates such as equids, pecorans, and some derived suines. Although the M. masseter of camelids and equids do not reach the particularly modified configuration of pecorans, in which it is rostrally extended and arranged horizontally, the posterior sectors of Mm. masseter superficialis and pterygoideus medialis have acquired relatively horizontal disposition in the former lineages, suitable for protraction. The pterygoidei complex presents several bundles, and its relative size is intermediate between suines and derived grinding euungulates. The whole masticatory muscles are relatively light when compared to jaw weight. The evolution of the masticatory muscles and chewing of camelids implied that grinding abilities were reached with less extreme modifications of the topography and/or proportions than pecoran ruminants and equids. A relatively large M. temporalis recruited as a powerful retractor during the power stroke is a key feature of camelids. The relaxed pressure on chewing derived from the acquisition of rumination explains the slenderer build masticatory musculature of camelids compared to other euungulates except ruminants.

Ontogenetic and in silico models of spatial-packing in the hypermuscular mouse skull.

Networks linking single genes to multiple phenotypic outcomes can be founded on local anatomical interactions as well as on systemic factors like biochemical products. Here we explore the effects of such interactions by investigating the competing spatial demands of brain and masticatory muscle growth within the hypermuscular myostatin-deficient mouse model and in computational simulations. Mice that lacked both copies of the myostatin gene (-/-) and display gross hypermuscularity, and control mice that had both copies of the myostatin gene (+/+) were sampled at 1, 7, 14 and 28 postnatal days. A total of 48 mice were imaged with standard as well as contrast-enhanced microCT. Size metrics and landmark configurations were collected from the image data and were analysed alongside in silico models of tissue expansion. Findings revealed that: masseter muscle volume was smaller in -/- mice at day 1 but became, and remained thereafter, larger by 7 days; -/- endocranial volumes begin and remained smaller; -/- enlargement of the masticatory muscles was associated with caudolateral displacement of the calvarium, lateral displacement of the zygomatic arches, and slight dorsal deflection of the face and basicranium. Simulations revealed basicranial retroflexion (flattening) and dorsal deflection of the face associated with muscle expansion and abrogative covariations of basicranial flexion and ventral facial deflection associated with endocranial expansion. Our findings support the spatial-packing theory and highlight the importance of understanding the harmony of competing spatial demands that can shape and maintain mammalian skull architecture during ontogeny.

Fatigue resistant jaw muscles facilitate long-lasting courtship behaviour in the southern alligator lizard (Elgaria multicarinata).

The southern alligator lizard (Elgaria multicarinata) exhibits a courtship behaviour during which the male firmly grips the female's head in his jaws for many hours at a time. This extreme behaviour counters the conventional wisdom that reptilian muscle is incapable of powering high-endurance behaviours. We conducted in situ experiments in which the jaw-adductor muscles of lizards were stimulated directly while bite force was measured simultaneously. Fatigue tests were performed by stimulating the muscles with a series of tetanic trains. Our results show that a substantial sustained force gradually develops during the fatigue test. This sustained force persists after peak tetanic forces have declined to a fraction of their initial magnitude. The observed sustained force during in situ fatigue tests is consistent with the courtship behaviour of these lizards and probably reflects physiological specialization. The results of molecular analysis reveal that the jaw muscles contain masticatory and tonic myosin fibres. We propose that the presence of tonic fibres may explain the unusual sustained force properties during mate-holding behaviour. The characterization of muscle properties that facilitate extreme performance during specialized behaviours may reveal general mechanisms of muscle function, especially when done in light of convergently evolved systems exhibiting similar performance characteristics.

Deep learning vs. atlas-based models for fast auto-segmentation of the masticatory muscles on head and neck CT images.

BACKGROUND: Impaired function of masticatory muscles will lead to trismus. Routine delineation of these muscles during planning may improve dose tracking and facilitate dose reduction resulting in decreased radiation-related trismus. This study aimed to compare a deep learning model with a commercial atlas-based model for fast auto-segmentation of the masticatory muscles on head and neck computed tomography (CT) images. MATERIAL AND METHODS: Paired masseter (M), temporalis (T), medial and lateral pterygoid (MP, LP) muscles were manually segmented on 56 CT images. CT images were randomly divided into training (n = 27) and validation (n = 29) cohorts. Two methods were used for automatic delineation of masticatory muscles (MMs): Deep learning auto-segmentation (DLAS) and atlas-based auto-segmentation (ABAS). The automatic algorithms were evaluated using Dice similarity coefficient (DSC), recall, precision, Hausdorff distance (HD), HD95, and mean surface distance (MSD). A consolidated score was calculated by normalizing the metrics against interobserver variability and averaging over all patients. Differences in dose (∆Dose) to MMs for DLAS and ABAS segmentations were assessed. A paired t-test was used to compare the geometric and dosimetric difference between DLAS and ABAS methods. RESULTS: DLAS outperformed ABAS in delineating all MMs (p < 0.05). The DLAS mean DSC for M, T, MP, and LP ranged from 0.83 ± 0.03 to 0.89 ± 0.02, the ABAS mean DSC ranged from 0.79 ± 0.05 to 0.85 ± 0.04. The mean value for recall, HD, HD95, MSD also improved with DLAS for auto-segmentation. Interobserver variation revealed the highest variability in DSC and MSD for both T and MP, and the highest scores were achieved for T by both automatic algorithms. With few exceptions, the mean ∆D98%, ∆D95%, ∆D50%, and ∆D2% for all structures were below 10% for DLAS and ABAS and had no detectable statistical difference (P > 0.05). DLAS based contours had dose endpoints more closely matched with that of the manually segmented when compared with ABAS. CONCLUSIONS: DLAS auto-segmentation of masticatory muscles for the head and neck radiotherapy had improved segmentation accuracy compared with ABAS with no qualitative difference in dosimetric endpoints compared to manually segmented contours.

The ontogeny of maximum bite force in humans.

Ontogenetic changes in the human masticatory complex suggest that bite force, a key measure of chewing performance, increases throughout growth and development. Current published bite force values for humans exist for molar and incisal biting, but few studies measure bite forces across all tooth types, or measure bite force potentials in subjects of different ages. In the absence of live data, models of bite force such as the Constrained Lever Model (CLM), are employed to predict bite force at different bite points for adults, but it is unclear whether such a model can accurately predict bite force potentials for juveniles or subadults. This study compares theoretically derived bite forces and live bite force data, and places these within an ontogenetic context in humans. Specifically, we test whether (1) patterns of maximum bite force increase along the tooth row throughout ontogeny, (2) bite force patterns estimated using the CLM match patterns observed from live bite force data, and (3) changes in bite forces along the tooth row and throughout ontogeny are associated with concomitant changes in adductor muscle leverage. Our findings show that maximum bite forces increase throughout ontogeny and change along the tooth row, with the highest forces occurring at the posterior dentition. These findings adhere to the expectations under the CLM and validate the model's utility in predicting bite force values throughout development. Furthermore, adductor muscle leverage values reflect this pattern, with the greatest leverage values occurring at the posterior dentition throughout ontogeny. The CLM informs our study of mammalian chewing mechanics by providing a model of how morphological changes of the masticatory apparatus during ontogeny affect bite force distribution along the tooth row. Furthermore, the decreased bite force magnitudes observed in juveniles and subadults compared with adults suggest that differences in juvenile and subadult diets may partially be due to differences in bite force production potentials.

Functional morphology of the jaw adductor muscles in the Canidae.

Cranial form is closely allied to diet and feeding behavior in the Canidae, with the force and velocity of jaw-closing depending on the bony morphology of the skull and mandible, and the mass, architecture, and siting of the jaw adductor muscles. Previously, little has been reported on the details of the form and function of canid jaw adductor muscles, with earlier studies basing functional hypotheses on data derived from dry skull specimens. Here we use empirically derived muscle data from fresh-frozen specimens to explore the architecture of the muscles, and to inform finite element analyses models that predict bite force and strain energy in 12 species of wild canid. The inclusion of muscle architectural detail is shown to influence masticatory muscle force production capability calculations, indicating that muscles with longer fascicles were disadvantaged compared to muscles with shorter fascicles. No clear patterns of allometry were detected. Dietary groups were differentiated by temporalis fascicle angles, which, when allied with the differentiation of rostral length reported in previous studies, may further contribute to specializations of fast jaw-closing or forceful jaw-closing species. The most biomechanically demanding masticatory function is canine biting, and the highest strain energy values were reported in this loading condition, particularly in the zygomatic arches and caudal rostrum. Specific head shapes may be constrained by size, with scaled strain energy models predicting that some bony morphologies may only be viable in species with small body masses.

DiceCT Analysis of the Extreme Gouging Adaptations Within the Masticatory Apparatus of the Aye-Aye (Daubentonia madagascariensis).

Relative to all other primates, the aye-aye (Daubentonia madagascariensis) exists at the extremes of both morphology and behavior. Its specialized anatomy-which includes hypselodont incisors and highly derived manual digits-reflects a dietary niche, unique among primates, which combines tap-foraging with gouging to locate and extract wood-boring larvae. Here, we explore the impact of this extreme dietary ecology upon the masticatory musculature of this taxon with reference to a second, similarly sized but highly generalist lemuriform-the mongoose lemur (Eulemur mongoz). Using non-destructive, high-resolution diffusible iodine-based contrast-enhanced computed tomography techniques, we reconstruct the three-dimensional volumes of eight masticatory muscles, and, for the first time in strepsirrhines, isolate and visualize their constituent muscle fascicles in situ and in three dimensions. Using these data, we report muscle volumes, forces, and fascicle lengths from each muscle portion, as well as their orientation relative to two standardized anatomical planes. Our findings demonstrate the overbuilt nature of the aye-aye's masticatory apparatus, in which each muscle possesses an absolutely and relatively larger muscle volume and PCSA than its counterpart in the mongoose lemur. Likewise, for several adductor muscles, aye-ayes also possess relatively greater fascicle lengths. Finally, we note several unusual features within the lateral pterygoid of the aye-aye-the muscle most responsible for jaw protrusion-that relate to force maximization and reorientation. As this jaw motion is critical to gouging, we interpret these differences to reflect highly specific specializations that facilitate the aye-aye's extreme subsistence strategy. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:282-294, 2020. © 2019 American Association for Anatomy.

Masticatory Apparatus Performance and Functional Morphology in the Extremely Large Mice from Gough Island.

Since their arrival approximately 200 years ago, the house mice (Mus musculus) on Gough Island (GI) rapidly increased in size to become the largest wild house mice on record. Along with this extreme increase in body size, GI mice adopted a predatory diet, consuming significant quantities of seabird chicks and eggs. We studied this natural experiment to determine how evolution of extreme size and a novel diet impacted masticatory apparatus performance and functional morphology in these mice. We measured maximum bite force and jaw opening (i.e., gape) along with several musculoskeletal dimensions functionally linked to these performance measurements to test the hypotheses that GI mice evolved larger bite forces and jaw gapes as part of their extreme increase in size and/or novel diet. GI mice can bite more forcefully and open their jaws wider than a representative mainland strain of house mice. Similarly, GI mice have musculoskeletal features of the masticatory apparatus that are absolutely larger than WSB mice. However, when considered relative to body size or jaw length, as a relevant mechanical standard, GI mice show reduced performance, suggesting a size-related decrease in these abilities. Correspondingly, most musculoskeletal features are not relatively larger in GI mice. Incisor biting leverage and condylar dimensions are exceptions, suggesting relative increases in biting efficiency and condylar rotation in GI mice. Based on these results, we hypothesize that evolutionary enhancements in masticatory performance are correlated with the extreme increase in body size and associated musculoskeletal phenotypes in Gough Island mice. Anat Rec, 2019. © 2018 American Association for Anatomy.

The Ontogeny of Masticatory Muscle Architecture in Microcebus murinus.

The masticatory apparatus has been the focus of many studies in comparative anatomy-especially analyses of skulls and teeth, but also of the mandibular adductor muscles which are responsible for the production of bite force and the movements of the mandible during food processing and transport. The fiber architecture of these muscles has been correlated to specific diets (e.g., prey size in felids) and modes of foraging (e.g., tree gouging in marmosets). Despite the well-elucidated functional implications of this architecture, little is known about its ontogeny. To characterize age-related myological changes, we studied the masticatory muscles in a large (n = 33) intraspecific sample of a small, Malagasy primate, Microcebus murinus including neonatal through geriatric individuals. We removed each of the mandibular adductors and recorded its mass as well as other linear measurements. We then chemically dissected each muscle to study its architecture-fascicle length and physiological cross-sectional area (PCSA) which relate to stretch (gape) and force capabilities, respectively. We observed PCSA and muscle mass to increase rapidly and plateau in adulthood through senescence. Fascicle lengths remained relatively constant once maximal length was reached, which occurred early in life, suggesting that subsequent changes in PCSA are driven by changes in muscle mass. Quadratic curvilinear models of each of the architectural variables of all adductors combined as well as individual muscles regressed against age were all significant. Anat Rec, 303:1364-1373, 2020. © 2019 American Association for Anatomy.

Bite Force and Masticatory Muscle Architecture Adaptations in the Dietarily Diverse Musteloidea (Carnivora).

Dietary ecology and its relationship with both muscle architecture and bite force potential has been studied in many mammalian (and non-mammalian) taxa. However, despite the diversity of dietary niches that characterizes the superfamily Musteloidea, the masticatory muscle fiber architecture of its members has yet to be investigated anatomically. In this study, we present myological data from the jaw adductors in combination with biomechanical data derived from craniomandibular measurements for 17 species representing all four families (Ailuridae, Mephitidae, Mustelidae, and Procyonidae) of Musteloid. These data are combined to calculate bite force potential at each of three bite points along the dental row. Across our sample as a whole, masticatory muscle mass scaled with isometry or slight positive allometry against both body mass and skull size (measured via a cranial geometric mean). Total jaw adductor physiological cross-sectional area scaled with positive allometry against both body mass and skull size, while weighted fiber length scaled with negative allometry. From a dietary perspective, fiber length is strongly correlated with dietary size such that taxa that exploit larger foods demonstrated myological adaptations toward gape maximization. However, no consistent relationship between bite force potential and dietary mechanical resistance was observed. These trends confirm previous findings observed within the carnivoran family Felidae (as well as within primates), suggesting that the mechanisms by which masticatory anatomy adapts to dietary ecology may be more universally consistent than previously recognized. Anat Rec, 302:2287-2299, 2019. © 2019 American Association for Anatomy.

Dry versus wet and gross: Comparisons between the dry skull method and gross dissection in estimations of jaw muscle cross-sectional area and bite forces in sea otters.

Bite force is a measure of feeding performance used to elucidate links between animal morphology, ecology, and fitness. Obtaining live individuals for in vivo bite-force measurements or freshly deceased specimens for bite force modeling is challenging for many species. Thomason's dry skull method for mammals relies solely on osteological specimens and, therefore, presents an advantageous approach that enables researchers to estimate and compare bite forces across extant and even extinct species. However, how accurately the dry skull method estimates physiological cross-sectional area (PCSA) of the jaw adductor muscles and theoretical bite force has rarely been tested. Here, we use an ontogenetic series of southern sea otters (Enhydra lutris nereis) to test the hypothesis that skeletomuscular traits estimated from the dry skull method accurately predicts test traits derived from dissection-based biomechanical modeling. Although variables from these two methods exhibited strong positive relationships across ontogeny, we found that the dry skull method overestimates PCSA of the masseter and underestimates PCSA of the temporalis. Jaw adductor in-levers for both jaw muscles and overall bite force are overestimated. Surprisingly, we reveal that sexual dimorphism in craniomandibular shape affects temporalis PCSA estimations; the dry skull method predicted female temporalis PCSA well but underestimates male temporalis PCSA across ontogeny. These results highlight the importance of accounting for sexual dimorphism and other intraspecific variation when using the dry skull method. Together, we found the dry skull method provides an underestimation of bite force over ontogeny and that the underlying anatomical components driving bite force may be misrepresented.

Visualization and Quantification of Digitally Dissected Muscle Fascicles in the Masticatory Muscles of Callithrix jacchus Using Nondestructive DiceCT.

The organization and length of a muscle's fascicles imparts its contractile properties. Longer fascicles permit increased muscle excursion, whereas changes in fascicle orientation relate to the overall vector of contractile force. Collecting data on fascicle architecture has traditionally involved destructive and irreversible gross dissection. In recent years, however, new imaging modalities have permitted muscles and their fascicles to be visualized nondestructively. Here, we present data from a primate (Callithrix jacchus), in which, for the first time, individual muscle fascicles are digitally "dissected" (segmented and reconstructed) using nondestructive, high-resolution diffusible iodine-based contrast-enhanced computed tomography (DiceCT) techniques. We also present quantitative data on the length and orientation of these fascicles within 10 muscle divisions of the jaw adductor and abductor musculature (superficial, deep, and zygomatic portions of temporalis and masseter; medial and lateral pterygoid; anterior and posterior digastric) and compare these digitally measured lengths to fascicular lengths measured using traditional gross and chemical dissection. Digitally derived fascicle lengths correspond well to their dissection-derived counterparts. Moreover, our analyses of changes in fascicle orientation across the adductor complex enable us to visualize previously uncharacterized levels of detail and highlight significant variation between adjacent muscle layers within muscle groups (e.g., between superficial, deep, and zygomatic portions of masseter and temporalis). We conclude that this technique offers great potential to future research, particularly for questions centered around the visualization and quantification of obscured and often-overlooked muscles such as the pterygoid and digastric muscles, and for deriving more accurate models of the masticatory system as a whole. Anat Rec, 302:1891-1900, 2019. © 2019 American Association for Anatomy.

Cephalometric studies of the mandible, its masticatory muscles and vasculature of growing Göttingen Minipigs-A comparative anatomical study to refine experimental mandibular surgery.

Over many decades, the Göttingen Minipig has been used as a large animal model in experimental surgical research of the mandible. Recently several authors have raised concerns over the use of the Göttingen Minipig in this research area, observing problems with post-operative wound healing and loosening implants. To reduce these complications during and after surgery and to improve animal welfare in mandibular surgery research, the present study elucidated how comparable the mandible of minipigs is to that of humans and whether these complications could be caused by specific anatomical characteristics of the minipigs' mandible, its masticatory muscles and associated vasculature. Twenty-two mandibular cephalometric parameters were measured on CT scans of Göttingen Minipigs aged between 12 and 21 months. Ultimately, we compared this data with human data reported in the scientific literature. In addition, image segmentation was used to determine the masticatory muscle morphology and the configuration of the mandibular blood vessels. Compared to data of humans, significant differences in the mandibular anatomy of minipigs were found. Of the 22 parameters measured only four were found to be highly comparable, whilst the others were not. The 3D examinations of the minipigs vasculature showed a very prominent deep facial vein directly medial to the mandibular ramus and potentially interfering with the sectional plane of mandibular distraction osteogenesis. Damage to this vessel could result in inaccessible bleeding. The findings of this study suggest that Göttingen Minipigs are not ideal animal models for experimental mandibular surgery research. Nevertheless if these minipigs are used the authors recommend that radiographic techniques, such as computed tomography, be used in the specific planning procedures for the mandibular surgical experiments. In addition, it is advisable to choose suitable age groups and customize implants based on the mandibular dimensions reported in this study.

The effect of the masticatory muscle physiological cross-sectional area on the structure of the temporomandibular joint in Carnivora.

We compared the temporomandibular joint structure between species of the order Carnivora and investigated its variation among family lineages. We also investigated the effect of the masticatory muscle physiological cross-sectional area (PCSA) on temporomandibular joint structure. The masticatory muscle is composed of multiple muscles, which contract in different directions and exert pressure on the temporomandibular joint. We investigated the effect of the ratio of each muscle's PCSA-an indicator of muscle force-and muscle size relative to body size on temporomandibular joint structure. The temporalis PCSA relative to body size showed the highest correlation with temporomandibular joint structure. When the temporalis PCSA is large relative to body size, the preglenoid projects caudally, the postglenoid projects rostrally and the pre-postglenoid angle interval is small, indicating that the condyle is locked in the fossa to reinforce the temporomandibular joint. Most Carnivora use blade-like carnassial teeth when slicing food. However, dislocation occurs when the carnassial teeth are used by the temporalis muscle. Our results suggest that the temporomandibular joint is reinforced to prevent dislocation caused by the temporalis muscle. In Mustelidae, the temporomandibular joint with a rostrally projecting postglenoid is suitable for carnassial biting using the temporalis muscle. In Felidae, the force of the masseter onto the carnassial teeth is diverted to the canine by tightening the temporomandibular joint. In Canidae, the masticatory muscle arrangement is well-balanced, enabling combined action. Hence, reinforcement of the temporomandibular joint by bone structure is unnecessary.