Relationships between the size and spatial morphology of human masseter and medial pterygoid muscles, the craniofacial skeleton, and jaw biomechanics

The relationship between human craniofacial morphology and the biomechanical efficiency of bite force generation in widely varying muscular and skeletal types is unknown. To address this problem, we selected 22 subjects with different facial morphologies and used magnetic resonance imaging, cephalometric radiography, and data from dental casts to reconstruct their craniofacial tissues in three dimensions. Conventional cephalometric analyses were carried out, and the cross-sectional sizes of the masseter and medial pterygoid muscles were measured from reconstituted sections. The potential abilities of the muscles to generate bite forces at the molar teeth and mandibular condyles were calculated according to static equilibrium theory using muscle, first molar, and condylar moment arms. On average, the masseter muscle was about 66% larger in cross section than the medial pterygoid and was inclined more anteriorly relative to the functional occlusal plane. There was a significant positive correlation (P less than 0.01) between the cross-sectional areas of the masseter and medial pterygoid muscles (r = 0.75) and between the bizygomatic arch width and masseter cross-sectional area (r = 0.56) and medial pterygoid cross-sectional area (r = 0.69). The masseter muscle was always a more efficient producer of vertically oriented bite force than the medial pterygoid. Putative bite force from the medial pterygoid muscle alone correlated positively with mandibular length and inversely with upper face height. When muscle and tooth moment arms were considered together, a system efficient at producing force on the first molar was statistically associated with a face having a large intergonial width, small intercondylar width, narrow dental arch, forward maxilla, and forward mandible. There was no significant correlation between muscle cross-sectional areas and their respective putative bite forces. This suggests that there is no simple relationship between the tension-generating capacity of the muscles and their mechanical efficiency as described by their spatial arrangement. The study shows that in a modern human population so many combinations of biomechanically relevant variables are possible that subjects cannot easily be placed into ideal or nonideal categories for producing molar force. Our findings also confirm the impression that similar bite-force efficiencies can be found in subjects with disparate facial features.     

Chewing pattern and muscular activation in open bite patients

Different studies have indicated, in open bite patients, that masticatory muscles tend to generate a small maximum bite force and to show a reduced cross-sectional area with a lower EMG activity. The aim of this study was to evaluate the kinematics parameters of the chewing cycles and the activation of masseters and anterior temporalis muscles of patients with anterior dental open bite malocclusion. There have been no previous reports evaluating both kinematic values and EMG activity of patients with anterior open bite during chewing. Fifty-two young patients (23 boys and 29 girls; mean age±SD 11.5±1.2 and 10.2±1.6years, respectively) with anterior open bite malocclusion and 21 subjects with normal occlusion were selected for the study. Kinematics parameters and surface electromyography (EMG) were simultaneously recorded during chewing a hard bolus with a kinesiograph K7-I Myotronics-Usa. The results showed a statistically significant difference between the open bite patients and the control group for a narrower chewing pattern, a shorter total and closing duration of the chewing pattern, a lower peak of both the anterior temporalis and the masseter of the bolus side. In this study, it has been observed that open bite patients, lacking the inputs from the anterior guidance, that are considered important information for establishing the motor scheme of the chewing pattern, show narrower chewing pattern, shorter lasting chewing cycles and lower muscular activation with respect to the control group.     

Pressure distribution measurement in biting surimi gels with molars using a multiple-point sheet sensor

The bite force of three surimi gels with molars was measured in the mouth using a multiple-point sheet sensor. A peak force appeared at the breaking point of each sample, and then the force increased again, accompanied by a decrease in the opening between the upper and lower teeth. Low values in the peak force, pressure, and time at the first peak, the time at which the maximum contact area was engaged, impulse, and slope of bite curve were observed in samples with low breaking force and low breaking deformation found by the mechanical measurement of gel strength, and with less toughness in the sensory assessment. The duration of the bite force, the second peak time, and active bite pressure at the second peak did not change with a change in the surimi texture. The active pressure at the breaking point of each gel was affected by gel strength, while that at the second peak was independent of the gel strength.     

Differences in the Mode of Growth Inhibition of Baker’s Yeast Brought about by Arsenate and Methanearsonate

The bite force of three surimi gels with molars was measured in the mouth using a multiple-point sheet sensor. A peak force appeared at the breaking point of each sample, and then the force increased again, accompanied by a decrease in the opening between the upeer and lower teeth. Low values in the peak force, pressure, and time at the first peak, the time at which the maximum contact area was engaged, impulse, and slope of bite curve were observed in samples with low breaking force and low breaking deformation found by the mechanical measurement of gel strength, and with less toughness in the sensory assessment. The duration of the bite force, the second peak time, and active bite pressure at the second peak did not change with a change in the surimi texture. The active pressure at the breaking point of each gel was affected by gel strength, while that at the second peak was independent of the gel strength.     

Scaling of bite force in the blacktip shark Carcharhinus limbatus

Although bite force is a frequently studied performance measure of feeding ecology, changes in bite force over ontogeny have rarely been investigated. Biting by the blacktip shark Carcharhinus limbatus was theoretically modeled over ontogeny to investigate the scaling of bite force, the morphological basis of the observed scaling relationship, the ecological consequences of ontogenetic changes in performance, and whether cranial morphometrics can be used as an accurate proxy for bite force. Theoretical bite force, which was positively allometric with respect to total length (TL), ranged from 32 N (61 cm TL) to 423 N (152 cm TL) at the anterior tips of the jaws and from 107 (61 cm TL) to 1083 N (152 cm TL) at the posterior teeth. This observation is attributed to positive allometry in the mechanical advantage of the jaw-adducting mechanism and the cross-sectional area of all four jaw-adducting muscles. Theoretical bite force was accurately predicted by cranial morphometrics including prebranchial length and head width as well. Although positive allometry of bite force in C. limbatus would seem to indicate an ecological necessity for this phenomenon, dietary analyses do not necessarily indicate any ontogenetic shift in prey types requiring larger bite forces. The positively allometric increase in theoretical bite force may be associated with numerous other selective pressures including maintenance of an apical position within the ecosystem.     

Jaw-muscle activity in ferrets, Mustela putorius furo.

Electromyographical (EMG) activity was recorded bilaterally from the masseter and temporalis muscles of alert ferrets (Mustela putorius furo) during mastication and crushing. Electromyographic activity was also recorded during biting while a bite-force transducer placed between the carnassial teeth registered forces ranging from 1.5 to 48.8 N. Linear regression analysis demonstrates that temporalis and masseter EMG activity are linearly related to bite force. Electromyographic activity from the balancing-side muscles is nearly equal to EMG activity of the working-side muscles during bone crushing with the carnassial teeth. It is hypothesized that a high percentage of balancing-side muscle activity in ferrets can be recruited during carnassial biting because the postglenoid process prevents ventral displacement of the working-side mandibular condyle.     

A new model for calculating muscle forces from electromyograms

A muscle model is described that uses electromyogram (EMG), muscle length and speed of contraction to predict muscle force. Physiological parameters are the Hill constants and the shape of the twitch response to a single stimulus. The model was incorporated in a jaw model of the rabbit and tested by predicting the bite force produced by the jaw muscles during mastication. The time course of the calculated force appeared to match the bite force, measured in vivo by a strain gauge, applied to the bone below the teeth. The variation in peak strain amplitude from cycle to cycle correlated with the variation predicted by the model. The peak amplitude of the integrated EMGs of individual jaw muscles showed an average correlation with peak strain of 0.41. Use of the sum of the available peak amplitudes, weighted according to their effect upon the bite force increased the correlation to 0.46; the model predicted bite forces showed a correlation of 0.57 with the strain. The increase in correlation was statistically significant. The muscle forces were calculated using a minimum number of easily obtainable constants.     

Altered control of submaximal bite force during bruxism in humans

The control of bite force during varying submaximal loads was examined in patients suffering from bruxism compared to healthy humans not showing these symptoms. The subjects raised a bar (preload) with their incisor teeth and held it between their upper and lower incisors using the minimal bite force required to keep the bar in a horizontal position. Further loading was added during the preload phase. A sham load was also used. Depending on the session, the teeth were loaded by the experimenter or the subject and in one session the subject did not see the load (no visual feedback). The bite force was measured continuously using a calibrated force transducer. In all the subjects, the bite force increased with increasing load. Following the addition of the load, the level of the tonic bite force was reached rapidly with no marked overshoot. The patients with bruxism used significantly higher bite forces to hold the submaximal loads compared to the control subjects. In the control subjects, the holding forces for each submaximal load were identical in the men and the women and were independent of subject maximal bite force. Sham loading evoked no marked responses in biting force. Whether the subject or the experimenter added the load or whether the subject had visual feedback or not were not significant factors in determining the level of bite force. The results indicated that the patients with bruxism used excessively large biting forces for each given submaximal load. This study showed no evidence that the inappropriate control of bite force by patients with bruxism was due to an abnormality in the higher cortical circuits that regulates the function of trigeminal motoneurons in the brainstem. This was shown by a lack of abnormality in coordination of voluntary hand movement with biting force, a lack of abnormal anticipation response to a sham load and a lack of any effect of visual feedback. The results were in line with the hypothesis that afferent input from oral (periodontal or masticatory muscle) tissues does not provide an appropriate control of motor command in bruxism.     

Comparison of force and EMG measures in normal and reinnervated tibialis anterior muscles of the rat.

The relationship between motor unit force and the recorded voltage produced by activated muscle unit fibres (electromyogram, EMG) was examined in normal and reinnervated rat tibialis anterior muscles. The number, cross-sectional area, and radial distance from the recording electrode of muscle fibres in a given unit, obtained directly from a sample of glycogen-depleted motor units, were analysed in relation to the magnitude of the EMG signal produced by that unit. EMG peak to peak amplitude and area varied as approximately the square root of twitch force in both normal and reinnervated units. Furthermore, the EMG amplitude increased approximately as the total cross-sectional area of the motor unit (number of muscle fibres x the average cross-sectional area of the fibres) and inversely with approximately the square root of the distance of fibres from the recording electrodes on the surface of the muscle.     

Motor pattern control for increasing crushing force in the striped burrfish (Chilomycterus schoepfi)

The relationship between muscular force modulation and the underlying nervous system control signals has been difficult to quantify for in vivo animal systems. Our goal was to understand how animals alter muscle activation patterns to increase bite forces and to evaluate how accurate these patterns are in predicting crushing forces. We examined the relationship between commonly used measures of cranial muscle activity and force production during feeding events of the striped burrfish (Chilomycterus schoepfi), a mollusc crushing specialist. We quantified the force required to crush a common gastropod prey item (Littorina irrorata) of burrfish using a materials testing device. Burrfish were fed these calibrated prey items while we recorded electromyograms (EMGs) from the main jaw closing muscles (adductor mandibulae A1beta, A2alpha, and A2beta). We quantified EMG activity by measuring the burst duration, rectified integrated area, and then calculated the intensity of activity from these two variables. Least squares regressions relating force to crush (Fcrush) and all EMG variables were calculated for each fish. Multiple regression analyses were used to determine how much of the variation in Fcrush could be explained by muscle activation patterns. We found that 20 cm burrfish are capable of generating extremely high crushing forces (380 N peak force) primarily by increasing the duration of muscle activity. EMG variables explained 71% of the total variation in force production. After accounting for the inherent variation in Fcrush of snails, EMGs do a very good job of predicting bite forces for these fish.     

Bite force and occlusal stress production in hominin evolution

Maximum bite force affects craniofacial morphology and an organism's ability to break down foods with different material properties. Humans are generally believed to produce low bite forces and spend less time chewing compared with other apes because advances in mechanical and thermal food processing techniques alter food material properties in such a way as to reduce overall masticatory effort. However, when hominins began regularly consuming mechanically processed or cooked diets is not known. Here, we apply a model for estimating maximum bite forces and stresses at the second molar in modern human, nonhuman primate, and hominin skulls that incorporates skeletal data along with species‐specific estimates of jaw muscle architecture. The model, which reliably estimates bite forces, shows a significant relationship between second molar bite force and second molar area across species but does not confirm our hypothesis of isometry. Specimens in the genus Homo fall below the regression line describing the relationship between bite force and molar area for nonhuman anthropoids and australopiths. These results suggest that Homo species generate maximum bite forces below those predicted based on scaling among australopiths and nonhuman primates. Because this decline occurred before evidence for cooking, we hypothesize that selection for lower bite force production was likely made possible by an increased reliance on nonthermal food processing. However, given substantial variability among in vivo bite force magnitudes measured in humans, environmental effects, especially variations in food mechanical properties, may also be a factor. The results also suggest that australopiths had ape‐like bite force capabilities. Am J Phys Anthropol 151:544–557, 2013. © 2013 Wiley Periodicals, Inc.     

Comparative finite element analysis of the cranial performance of four herbivorous marsupials

Marsupial herbivores exhibit a wide variety of skull shapes and sizes to exploit different ecological niches. Several studies on teeth, dentaries, and jaw adductor muscles indicate that marsupial herbivores exhibit different specializations for grazing and browsing. No studies, however, have examined the skulls of marsupial herbivores to determine the relationship between stress and strain, and the evolution of skull shape. The relationship between skull morphology, biomechanical performance, and diet was tested by applying the finite element method to the skulls of four marsupial herbivores: the common wombat (Vombatus ursinus), koala (Phascolarctos cinereus), swamp wallaby (Wallabia bicolor), and red kangaroo (Macropus rufus). It was hypothesized that grazers, requiring stronger skulls to process tougher food, would have higher biomechanical performance than browsers. This was true when comparing the koala and wallaby (browsers) to the wombat (a grazer). The cranial model of the wombat resulted in low stress and high mechanical efficiency in relation to a robust skull capable of generating high bite forces. However, the kangaroo, also a grazer, has evolved a very different strategy to process tough food. The cranium is much more gracile and has higher stress and lower mechanical efficiency, but they adopt a different method of processing food by having a curved tooth row to concentrate force in a smaller area and molar progression to remove worn teeth from the tooth row. Therefore, the position of the bite is crucial for the structural performance of the kangaroo skull, while it is not for the wombat which process food along the entire tooth row. In accordance with previous studies, the results from this study show the mammalian skull is optimized to resist forces generated during feeding. However, other factors, including the lifestyle of the animal and its environment, also affect selection for skull morphology to meet multiple functional demands. J. Morphol. 276:1230–1243, 2015. © 2015 Wiley Periodicals, Inc.     

Masticatory ability in 80-year-old subjects and its relation to intake of energy,nutrients and food items

Objective: The aim of the present study was to analyse the relationship between masticatory ability (self‐assessed masticatory ability and bite force) and intake of energy, nutrients and food items in a population sample of elderly subjects. Design and Subjects: From a population sample of 80‐year‐old people, 160 individuals (74 men and 86 women) took part in an odontological study. Main Outcome Measures: A dental examination including bite force recording, a questionnaire focusing on self‐assessed masticatory ability, and a dietary interview. Setting: Department of Geriatric Medicine, Göteborg University, Sweden. Results: The dental status among the participants varied much (from edentulous in both jaws ‐ 22% ‐ to more than 20 natural teeth ‐ 30%). The mean maximum bite force was higher in men (165 N) than in women (105 N). Bite force was significantly correlated to the Eichner index and to the number of teeth. One third of the subjects reported no masticatory problem, whereas 18% identified 3 such problems. The intake of energy and nutrients varied much but the means were well above recommended values. The correlations between dental status and bite force on one side and dietary intake on the other side were in general weak and most often statistically non‐significant. Impaired general health and reduced dentition were both associated with more masticatory problems. Conclusion: The examined sample of 80‐year‐old subjects had a great variation in dental status, bite force and self‐assessed masticatory ability, but these factors had only a minor influence on dietary selection and intake, which on average were well above recommended values.     

Microbial Analysis of Bite Marks by Sequence Comparison of Streptococcal DNA

Bite mark injuries often feature in violent crimes. Conventional morphometric methods for the forensic analysis of bite marks involve elements of subjective interpretation that threaten the credibility of this field. Human DNA recovered from bite marks has the highest evidentiary value, however recovery can be compromised by salivary components. This study assessed the feasibility of matching bacterial DNA sequences amplified from experimental bite marks to those obtained from the teeth responsible, with the aim of evaluating the capability of three genomic regions of streptococcal DNA to discriminate between participant samples. Bite mark and teeth swabs were collected from 16 participants. Bacterial DNA was extracted to provide the template for PCR primers specific for streptococcal 16S ribosomal RNA (16S rRNA) gene, 16S–23S intergenic spacer (ITS) and RNA polymerase beta subunit (rpoB). High throughput sequencing (GS FLX 454), followed by stringent quality filtering, generated reads from bite marks for comparison to those generated from teeth samples. For all three regions, the greatest overlaps of identical reads were between bite mark samples and the corresponding teeth samples. The average proportions of reads identical between bite mark and corresponding teeth samples were 0.31, 0.41 and 0.31, and for non-corresponding samples were 0.11, 0.20 and 0.016, for 16S rRNA, ITS and rpoB, respectively. The probabilities of correctly distinguishing matching and non-matching teeth samples were 0.92 for ITS, 0.99 for 16S rRNA and 1.0 for rpoB. These findings strongly support the tenet that bacterial DNA amplified from bite marks and teeth can provide corroborating information in the identification of assailants.     

Comparison between in vivo and theoretical bite performance: Using multi-body modelling to predict muscle and bite forces in a reptile skull

In biomechanical investigations, geometrically accurate computer models of anatomical structures can be created readily using computed-tomography scan images. However, representation of soft tissue structures is more challenging, relying on approximations to predict the muscle loading conditions that are essential in detailed functional analyses. Here, using a sophisticated multi-body computer model of a reptile skull (the rhynchocephalian Sphenodon), we assess the accuracy of muscle force predictions by comparing predicted bite forces against in vivo data. The model predicts a bite force almost three times lower than that measured experimentally. Peak muscle force estimates are highly sensitive to fibre length, muscle stress, and pennation where the angle is large, and variation in these parameters can generate substantial differences in predicted bite forces. A review of theoretical bite predictions amongst lizards reveals that bite forces are consistently underestimated, possibly because of high levels of muscle pennation in these animals. To generate realistic bites during theoretical analyses in Sphenodon, lizards, and related groups we suggest that standard muscle force calculations should be multiplied by a factor of up to three. We show that bite forces increase and joint forces decrease as the bite point shifts posteriorly within the jaw, with the most posterior bite location generating a bite force almost double that of the most anterior bite. Unilateral and bilateral bites produced similar total bite forces; however, the pressure exerted by the teeth is double during unilateral biting as the tooth contact area is reduced by half.     

Comparative analysis of methods for determining bite force in the spiny dogfish Squalus acanthias

Many studies have identified relationships between the forces generated by the cranial musculature during feeding and cranial design. Particularly important to understanding the diversity of cranial form amongst vertebrates is knowledge of the generated magnitudes of bite force because of its use as a measure of ecological performance. In order to determine an accurate morphological proxy for bite force in elasmobranchs, theoretical force generation by the quadratomandibularis muscle of the spiny dogfish Squalus acanthias was modeled using a variety of morphological techniques, and lever-ratio analyses were used to determine resultant bite forces. These measures were compared to in vivo bite force measurements obtained with a pressure transducer during tetanic stimulation experiments of the quadratomandibularis. Although no differences were found between the theoretical and in vivo bite forces measured, modeling analyses indicate that the quadratomandibularis muscle should be divided into its constituent divisions and digital images of the cross-sections of these divisions should be used to estimate cross-sectional area when calculating theoretical force production. From all analyses the maximum bite force measured was 19.57 N. This relatively low magnitude of bite force is discussed with respect to the ecomorphology of the feeding mechanism of S. acanthias to demonstrate the interdependence of morphology, ecology, and behavior in organismal design.     

Application and validation of a three-dimensional mathematical model of the human masticatory system in vivo.

A previously described three-dimensional mathematical model of the human masticatory system, predicting maximum possible bite forces in all directions and the recruitment patterns of the masticatory muscles necessary to generate these forces, was validated in in vivo experiments. The morphological input parameters to the model for individual subjects were collected using MRI scanning of the jaw system. Experimental measurements included recording of maximum voluntary bite force (magnitude and direction) and surface EMG from the temporalis and masseter muscles. For bite forces with an angle of 0, 10 and 20 degrees relative to the normal to the occlusal plane the predicted maximum possible bite forces were between 0.9 and 1.2 times the measured ones and the average ratio of measured to predicted maximum bite force was close to unity. The average measured and predicted muscle recruitment patterns showed no striking differences. Nevertheless, some systematic differences, dependent on the bite force direction, were found between the predicted and the measured maximum possible bite forces. In a second series of simulations the influence of the direction of the joint reaction forces on these errors was studied. The results suggest that they were caused primarily by an improper determination of the joint force directions.     

Amplification of Oral Streptococcal DNA from Human Incisors and Bite Marks

Challenges to the evidentiary value of morphometric determinations have led to a requirement for scientifically substantiated approaches to the forensic analysis of bite marks. Human teeth support genotypically distinctive populations of bacteria that could be exploited for forensic purposes. This study explored the feasibility of directly amplifying bacterial DNA from bite marks for comparison with that from teeth. Samples from self-inflicted experimental bite marks (n = 24) and human incisors were amplified by PCR using primers specific for streptococcal 16S ribosomal DNA. Amplicon profiles (resolved by denaturing gradient gel electrophoresis) from bite mark samples aligned significantly more closely with profiles generated from the teeth responsible than with those from other teeth. Streptococcal amplicons were generated from dental samples applied to excised porcine skin for up to 48 h. These findings indicate that streptococcal DNA can be amplified directly from bite marks, and have potential application in bite mark analysis.     

ORAL ASSESSMENT OF HARDNESS BETWEEN ELASTIC AND PLASTIC PRODUCTS

This study examined how people compare the hardness of different materials. Simple models with either elastic or plastic behavior were prepared and controlled mechanically. Six elastomers were compared to 29 plastic dental waxes of different hardness. For each elastic sample, a psychophysical staircase method was used to determine its perceived hardness in comparison to the plastic samples. Single bites were performed and the forces were recorded by a small load-cell placed between the sample and the teeth. Nine subjects. free of dental pathology, participated in this study. Subjects were able to match samples of both materials with a specific stress ratio which depended on the hardness of the elastic samples. Results indicated that people do not use the same sensory cues and, in this case, no clear role of the bite force was established.     

A functional analysis of carnassial biting

The jaw mechanism of carnivores is studied using an idealized model (Greaves, 1978). The model assumes: (i) muscle activity on both sides of the head, and (ii) that the jaw joints and the carnassial teeth are single points of contact between the skull and the lower jaw during carnassial biting. The model makes the following predictions: (i) in carnivores with carnassial teeth the resultant force of the jaw muscles will be positioned approximately 60% of the way from the jaw joint to the tooth—this arrangement delivers the maximum bite force possible together with a reasonably wide gape (remembering that bite force and gape cannot both be maximized); (ii) in an evolutionary sense, if greater bite force is required at the carnassial tooth, either the animal will get larger so as to deliver an absolutely larger bite force or the architecture of the muscles may change, becoming more pinnate, for example, but jaw geometry (i.e. the relative positions of the jaw joints, the carnassial tooth, and the muscle resultant force) will not change; (iii) if greater gape is required, the animal will get larger so as to have longer jaws and therefore an absolutely wider gape or change its muscle architecture allowing for greater stretch while the geometry remains unchanged; and (iv) in animals with a longer shearing region (e.g. the extinct hyaenodonts) the shearing region will be approximately 20% of jaw length and the muscle resultant force will be positioned approximately 60% of the way from the jaw joint to the most anterior shearing tooth.