A novel computational method to determine subject-specific bite force and occlusal loading during mastication

The evaluation of three-dimensional occlusal loading during biting and chewing may assist in development of new dental materials, in designing effective and long-lasting restorations such as crowns and bridges, and for evaluating functional performance of prosthodontic components such as dental and/or maxillofacial implants. At present, little is known about the dynamic force and pressure distributions at the occlusal surface during mastication, as these quantities cannot be measured directly. The aim of this study was to evaluate subject-specific occlusal loading forces during mastication using accurate jaw motion measurements. Motion data was obtained from experiments in which an individual performed maximal effort dynamic chewing cycles on a rubber sample with known mechanical properties. A finite element model simulation of one recorded chewing cycle was then performed to evaluate the deformation of the rubber. This was achieved by imposing the measured jaw motions on a three-dimensional geometric surface model of the subject’s dental impressions. Based on the rubber’s deformation and its material behaviour, the simulation was used to compute the resulting stresses within the rubber as well as the contact pressures and forces on the occlusal surfaces. An advantage of this novel modelling approach is that dynamic occlusal pressure maps and biting forces may be predicted with high accuracy and resolution at each time step throughout the chewing cycle. Depending on the motion capture technique and the speed of simulation, the methodology may be automated in such a way that it can be performed chair-side. The present study demonstrates a novel modelling methodology for evaluating dynamic occlusal loading during biting or chewing.     

Influence of ground reaction force perturbations on anterior cruciate ligament loading during sidestep cutting

Anterior cruciate ligament (ACL) injury risk is likely increased under unexpected loading conditions. Such situations may arise from mid-air contact with another athlete, or misjudgments in landing height, stride length or surface compliance resulting in an unbalanced landing and unexpected changes in the ground reaction forces (GRFs). The purpose this study was to identify how GRF perturbations influence ACL loading during sidestep cutting. Muscle-actuated simulations of sidestep cutting were generated and analyzed for 20 subjects. Perturbations of 20, 40 and 60% of the nominal value were applied to the posterior, vertical, and medial GRF. Open-loop, forward dynamics simulations were run with no feedback or correction mechanism which allowed deviations from the experimentally measured kinematics as a result of the GRF perturbations. Posterior and vertical GRF perturbations significantly increased ACL loading, although the change was more pronounced with posterior perturbations. These changes were primarily due to the sagittal plane component of ACL loading regardless of perturbation direction. Peak ACL loading occurred almost immediately after initial ground contact, and was thus predicated on initial joint configuration. The results of this study give merit to including knee flexion angle at initial ground contact in the evolving neuromuscular training modalities aimed at preventing non-contact ACL injury.     

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.     

Feeding behaviour and bite force of sabretoothed predators

The feeding behaviour of extinct sabretoothed predators (machaeroidines, nimravids, barbourofelids, machairodonts and thylacosmilines) is investigated using beam theory. Because bite force applied along the mandible should be proportional to the external dimension of the mandibular corpus, patterns of variation in these dimensions at interdental gaps will reflect the adaptation of the jaw to specific loads, related to killing methods. Comparison of the mandibular force profiles of sabretooths to those of extant conical‐toothed carnivorans of known feeding behaviour reveals that sabretooths had a powerful bite, as strong or stronger than extant felids of similar mandibular length. Loads exerted at the lower canine were better constrained in the sagittal plane than in extant conical‐toothed carnivorans, indicating that prey was efficiently restrained when the sabre bite was delivered. The mandibular symphysis is generally better buttressed dorsoventrally in dirk‐toothed sabretooths than in scimitar‐toothed sabretooths, implying different killing strategies for the two ecomorphs: dirktooths delivered powerful sabre bites on prey they restrained with their forelimbs, while scimitartooths delivered slashing sabre bites and may have used their incisor battery to subdue their prey. The mandibular symphysis of Smilodon fatalis is less buttressed dorsoventrally than that of other dirk‐toothed sabretooths, possibly as a consequence of the greater torsional stresses induced while feeding rapidly on carcasses in response to intense competition. The mandibular symphysis of Thylacosmilus atrox is better buttressed dorsoventrally in juveniles than in adults, suggesting that young marsupial sabretooths underwent an extended period of parental care as typically observed in modern felids and inferred for eutherian sabretooths. Finally, machaeroidines and the nimravid Nimravus brachyops are exceptional in exhibiting a degree of dorsoventral buttressing of the mandibular symphysis that is intermediate between advanced sabretooths and conical‐toothed felids but similar to the extant Neofelis nebulosa, suggesting that the latter taxon may be close to the ancestral condition of a new sabretooth radiation. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 145 , 393–426.     

Ontogenetic scaling of bite force in lizards and turtles

Because selection on juvenile life-history stages is likely strong, disproportionately high levels of performance (e.g., sprint speed, endurance, etc.) might be expected. Whereas this phenomenon has been demonstrated with respect to locomotor performance, data for feeding are scarce. Here, we investigate the relationships among body dimensions, head dimensions, and bite force during growth in lizards and turtles. We also investigate whether ontogenetic changes in bite performance are related to changes in diet. Our analyses show that, for turtles, head dimensions generally increase with negative allometry. For lizards, heads scale as expected for geometrically growing systems. Bite force generally increased isometrically with carapace length in turtles but showed significant positive allometry relative to body dimensions in lizards. However, both lizards and turtles display positive allometric scaling of bite force relative to some measures of head size throughout ontogeny, suggesting (1) strong selection for increased relative bite performance with increasing head size and (2) intrinsic changes in the geometry and/or mass of the jaw adductors during growth. Whereas our data generally do not provide strong evidence of compensation for lower absolute levels of performance, they do show strong links among morphology, bite force, and diet during growth.     

A method of bite force measurement in primates

A bite force transducer consisting of two differential strain beams with four strain gages in a full bridge configuration was modified for measuring occlusal forces in rhesus monkeys. A procedure of muscle stimulation (20-50 V, 60 Hz, and 0.8 ms duration) produced maximal unilateral masticatory muscle contraction when stimulating electrodes were placed in the masseter muscle. Tests of this procedure revealed reproducible results and a potential for use in studies of the force of isometric contraction of the masticatory muscles in normal and experimentally altered macaques and other primates.     

Design and construction of a transducer for bite force registration

This study describes the development of a system for quantification of human biting forces by (1) determining the mechanical properties of an epoxy resin reinforced with carbon fiber, (2) establishing the transducer's optimal dimensions to accommodate teeth of various widths while minimizing transducer thickness, and (3) determining the optimal location of strain gages using a series of mechanical resistance and finite element (FE) analyses. The optimal strain gage location was defined as the position that produced the least difference in strain pattern when the load was applied by teeth with two different surface areas. The result is a 7.3-mm-thick transducer with a maximum load capacity beyond any expected maximum bite force (1500 N). This system includes a graphic interface that easily allows acquisition and registration of bite force by any health-sciences or engineering professional.     

Feeding biomechanics and theoretical calculations of bite force in bull sharks (Carcharhinus leucas) during ontogeny

Evaluations of bite force, either measured directly or calculated theoretically, have been used to investigate the maximum feeding performance of a wide variety of vertebrates. However, bite force studies of fishes have focused primarily on small species due to the intractable nature of large apex predators. More massive muscles can generate higher forces and many of these fishes attain immense sizes; it is unclear how much of their biting performance is driven purely by dramatic ontogenetic increases in body size versus size-specific selection for enhanced feeding performance. In this study, we investigated biting performance and feeding biomechanics of immature and mature individuals from an ontogenetic series of an apex predator, the bull shark, Carcharhinus leucas (73–285 cm total length). Theoretical bite force ranged from 36 to 2128 N at the most anterior bite point, and 170 to 5914 N at the most posterior bite point over the ontogenetic series. Scaling patterns differed among the two age groups investigated; immature bull shark bite force scaled with positive allometry, whereas adult bite force scaled isometrically. When the bite force of C. leucas was compared to those of 12 other cartilaginous fishes, bull sharks presented the highest mass-specific bite force, greater than that of the white shark or the great hammerhead shark. A phylogenetic independent contrast analysis of anatomical and dietary variables as determinants of bite force in these 13 species indicated that the evolution of large adult bite forces in cartilaginous fishes is linked predominantly to the evolution of large body size. Multiple regressions based on mass-specific standardized contrasts suggest that the evolution of high bite forces in Chondrichthyes is further correlated with hypertrophication of the jaw adductors, increased leverage for anterior biting, and widening of the head. Lastly, we discuss the ecological significance of positive allometry in bite force as a possible “performance gain” early in the life history of C. leucas.     

A simple morphological predictor of bite force in rodents

Bite force was quantified for 13 species of North American rodents using a piezo-resistive sensor. Most of the species measured (11) formed a tight relationship between body mass and bite force (log 10(bite force)=0.43(log 10(body mass))+0.416; R ²>0.98). This high correlation exists despite the ecological (omnivores, grazers and more carnivorous) and taxonomic (Cricetidae, Heteromyidae, Sciuridae and Zapodidae) diversity of species. Two additional species, Geomys bursarius (Geomyidae) and a Sciurus niger (Sciuridae), bit much harder for their size. We found a simple index of strength based on two measurements of the incisor at the level of the alveolus ( Z_i =((anterior-posterior length)²× (medial-lateral width))/6) that is highly predictive of bite force in these rodents (R²>0.96). Z_i may be useful for prediction of bite force (log10 (Bite Force)=0.566log10 ( Z_i )+1.432) when direct measurements are not available.     

Maximum bite force after the replacement of complete dentures

Objective: The aim of the present study was to investigate whether the maximum bite force ( MBF ) can be improved by the replacement of complete dentures for elderly people. Design: Nine edentulous volunteers, mean age 74.2 (± 5.5) years and average denture experience 19.4 ± 19.5 years (1 to 50 years) had replacement dentures made. After a rehearsal session, MBF was recorded with the old dentures, and with the new dentures immediately at insertion, at 3, 8 days, 2–3 weeks, 1, 2, 3 and 6–10 months post insertion ( p‐i. ). MBF was recorded with the central bearing point method using a full‐bridge strain gauge load cell. Data were analysed off‐line using the mean of two peak readings per patient per session. Results: The results indicate that MBF tended to be impaired when replacement dentures were first fitted (n.s.). However, this trend reversed during the first month p‐i. for patients with a “moderate” lower ridge resorption of Atwood grade 3 or 4 (n = 5). Patients with more severe lower ridge resorption Atwood grade 5 or 6 (n = 4) showed a significantly lower MBF over the entire observation period (p0.05) and took longer to regain bite strength. Only patients with moderate bone resorption exceeded their pre‐insertion level of MBF within the observation period of 6–10 months p‐i. Conclusion: The present pilot study suggests that, at least for elderly patients with severe bone resorption, delayed improvement of MBF should be expected with replacement complete dentures.     

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.     

The maximum average bite force for a given jaw length

Major osteological landmarks were used to prepare idealized drawings of mammalian and reptilian lower jaws. Measurements from these drawings allowed the average output or bite force, along the entire jaw, to be calculated for many different anteroposterior positions of the input or muscle force. In the mammalian drawing, the maximum average bite force is exerted when the resultant force is located at about 30% of the way along the jaw from the joints. Because of geometric differences in the reptilian drawing, a resultant positioned at 20% of the way along the jaw exerts the maximum average bite force; a maximum force that is smaller than that in the mammalian case. The estimated location of the muscle resultant in actual cases corresponds to these calculated positions. Therefore, in real animals, the muscles are located in the position that produces the largest average force for any jaw length. The geometric changes necessary to transform the idealized reptilian drawing, with a smaller maximum average bite force, into that of the mammalian drawing, with a larger maximum force, are the same as those changes seen in the fossil record of the reptile/mammal transition. This finding suggests that the morphological changes that occurred in the jaws increased the average bite force in the primitive mammals.     

Diet,bite force and skull morphology in the generalist rodent morphotype

For many vertebrate species, bite force plays an important functional role. Ecological characteristics of a species' niche, such as diet, are often associated with bite force. Previous evidence suggests a biomechanical trade‐off between rodents specialized for gnawing, which feed mainly on seeds, and those specialized for chewing, which feed mainly on green vegetation. We tested the hypothesis that gnawers are stronger biters than chewers. We estimated bite force and measured skull and mandible shape and size in 63 genera of a major rodent radiation (the myomorph sigmodontines). Analysis of the influence of diet on bite force and morphology was made in a comparative framework. We then used phylogenetic path analysis to uncover the most probable causal relationships linking diet and bite force. Both granivores (gnawers) and herbivores (chewers) have a similar high bite force, leading us to reject the initial hypothesis. Path analysis reveals that bite force is more likely influenced by diet than the reverse causality. The absence of a trade‐off between herbivores and granivores may be associated with the generalist nature of the myomorph condition seen in sigmodontine rodents. Both gnawing and chewing sigmodontines exhibit similar, intermediate phenotypes, at least compared to extreme gnawers (squirrels) and chewers (chinchillas). Only insectivorous rodents appear to be moving towards a different direction in the shape space, through some notable changes in morphology. In terms of diet, natural selection alters bite force through changes in size and shape, indicating that organisms adjust their bite force in tandem with changes in food items.     

Influence of bite force and tongue pressure on oro-pharyngeal residue in the elderly

Objective: To investigate the influence of maximal bite force, maximal tongue pressure, number of mastications and swallowing on the oro‐pharyngeal residue in the elderly. Background: Oro‐pharyngeal residue in the elderly is an indication of dysphagia. Pharyngeal residue is especially critical as it may cause aspiration pneumonia, which is one of the major causes of death in elderly. Materials and methods: Videofluorographic recordings were performed on 14 elderly volunteers (six males, eight females, age range 65–93 years) without any history or symptoms of dysphagia. The subjects were instructed to consume 9 g of barium containing bread in two manners; free mastication and swallow (FMS: masticate and swallow freely), and limited mastication and swallow (LMS: swallow once after 30 chewing actions). The amount of oral and pharyngeal residue was evaluated using a 4‐point rating scale. Maximal occlusal force was measured by a pressure sensitive sheet, and maximal tongue pressure using a handy probe. Multiple regression analysis was performed to examine the influence of these items on the amount of oral and pharyngeal residue in FMS and LMS. Results: In FMS, age was found to be a factor which increased oral residue (p = 0.053), and the number of swallowing (p = 0.017) and the state of the prosthesis (p = 0.030) reduced the pharyngeal residue. In LMS, tongue pressure was a factor which reduced oral residue (p = 0.015) and increased pharyngeal residue (p = 0.008). Conclusion: It is suggested that in the elderly tongue pressure contributed to propulsion of the food bolus from oral cavity into the pharynx, and multiple swallowing contributed to the reduction in the amount of pharyngeal residue.     

Insularity affects head morphology,bite force and diet in a Mediterranean lizard

Island environments differ with regard to numerous features from the mainland and may induce large‐scale changes in most aspects of the biology of an organism. In this study, we explore the effect of insularity on the morphology and performance of the feeding apparatus, a system crucial for the survival of organisms. To this end, we examined the head morphology and feeding ecology of island and mainland populations of the Balkan green lizard, Lacerta trilineata. We predicted that head morphology, performance and diet composition would differ between sexes and habitats as a result of varying sexual and natural selection pressures. We employed geometric morphometrics to test for differences in head morphology, measured bite forces and analysed the diet of 154 adult lizards. Morphological analyses revealed significant differences between sexes and also between mainland and island populations. Relative to females, males had larger heads, a stronger bite and consumed harder prey than females. Moreover, island lizards differed in head shape, but not in head size, and, in the case of males, demonstrated a higher bite force. Islanders had a wider food niche breadth and included more plant material in their diet. Our findings suggest that insularity influences feeding ecology and, through selection on bite force, head morphology. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 469–484.     

Intramuscular pressure, force and blood flow in rabbit tibialis anterior muscles during single and repetitive contractions

The elevated intramuscular pressure (IMP) associated with sustained muscle contraction can affect blood flow, and could influence the long-term viability of functional skeletal muscle grafts. We therefore examined the relationship between force, peak IMP and blood flow in the tibialis anterior muscle of the anaesthetized rabbit. During isometric contractions, IMP was related linearly to force, and only the slope of the relationship varied between animals. During isotonic contractions, however, the highest values of IMP were found at the lowest force levels, and IMP appeared to be related to the amount and speed of shortening. During repeated isometric contractions, the ratio of IMP to force varied with time, stimulation pattern and subject. Mean blood flow did not differ appreciably between␣repetitive isometric contractions at duty cycles of 10–40%, and was unrelated to integrated pressure, integrated force, or depth from the surface. We conclude: (1) that IMP is unlikely to affect mean blood flow during cyclic activity that has a duty cycle less than 40%; and (2) that the clinical use of IMP as a predictor of muscle force appears to be justified only for single isometric contractions, and needs to be interpreted cautiously when contractions involve shortening or fatigue. Accepted: 17 November 1997     

Incisal bite force direction in humans and the functional significance of mammalian mandibular translation

Incisal bite force direction was recorded and analyzed in ten human subjects using a specially designed force transducer. In all ten subjects the maxillary incisal bite force was vertically and anteriorly directed both during static biting and during biting associated with simultaneous mandibular translation and rotation. Since the resultant muscle force could not have been equal and opposite to the mandibular bite force, the mandibular condyles must have been loaded. These data demonstrate that the mandible acts as a lever during incisal biting and that there is no consistent relationship between incisal bite force direction and object size. In some individuals the bite force direction was more vertical during biting on a large transducer (30 mm high), while in other subjects it was more vertical during biting on a small transducer (10 mm high).