Biomechanics of the masticatory apparatus of Pteropus giganteus (Megachiroptera)

Jaw mechanics in Pteropus were studied by means of a three-dimensional model. The model included several parameters of muscle architecture, combined with quantified movement and electromyographical data. Estimates of the nature of the applied forces that act upon the mandible during a chewing cycle, and subsequent estimates of reaction forces at the bite point and joints during the powerstroke, were thus obtained for different food consistencies. The resultant muscle force (relative to the palate) shifts from upward and slightly backward at large gapes to upward and markedly backward at the end of closing. The resultant simultaneously moves anteriorly. During the powerstroke it retains a constant position and orientation along the thickened anterior edge of the coronoid process. The early stages of opening are guided by the slope of the teeth and mandibular fossa; during the remaining part of opening the working line of the resultant crosses the skull behind the joint and thus acquires an opening moment. The bite force has downward and forward components, and a slight transverse component. For a given applied muscular force its magnitude is larger in more posteriorly positioned bite points. Both joints are loaded, the contralateral one more than the ipsilateral. Food consistency affects magnitude and orientation of the applied force, and hence, magnitude and orientation of the bite force and magnitude of the joint reaction forces. The magnitude of masseter activity relative to temporalis activity appears to be the key factor for the orientation of the bite force, and hence for the mechanical optimal position of the food. The adaptive value of the general topography of the masticatory muscles in Pteropus is discussed.     

Three-dimensional finite element modelling of muscle forces during mastication

This paper presents a three-dimensional finite element model of human mastication. Specifically, an anatomically realistic model of the masseter muscles and associated bones is used to investigate the dynamics of chewing. A motion capture system is used to track the jaw motion of a subject chewing standard foods. The three-dimensional nonlinear deformation of the masseter muscles are calculated via the finite element method, using the jaw motion data as boundary conditions. Motion-driven muscle activation patterns and a transversely isotropic material law, defined in a muscle-fibre coordinate system, are used in the calculations. Time-force relationships are presented and analysed with respect to different tasks during mastication, e.g. opening, closing, and biting, and are also compared to a more traditional one-dimensional model. The results strongly suggest that, due to the complex arrangement of muscle force directions, modelling skeletal muscles as conventional one-dimensional lines of action might introduce a significant source of error.     

Mutual influences of upper and lower extrimities during cyclic movements

The possibility of muscle activation of passive arm during its cyclic movements, imposed by active movements of contralateral arm or by experimenter was studied, as well as the influence of lower extremities cyclic movements onto arm muscles activity. In addition to that the activity of legs muscles was estimated in dependence on motor task condition for arms. Ten healthy supine subjects carried out opposite movements of arms with and without stepping-like movements of both legs. The experiment included three conditions for arm movements: 1) the active movements of both arms; 2) the active movements of one arm, when other entirely passive arm participated in the movement by force; 3) passive arm movement caused by experimenter. In the condition 2) additional load on active arm was applied (30 N and 60 N). In all three conditions the experiment was carried out with arms movements only or together with legs movements. The capability of passive moving arm muscles activation depended on increasing afferent inflow from muscles of contralateral arm was demonstrated. Emerging electrical activity was modulated in the arms movements cycle and depended on the degree of active arm loading. During combined active movements of arms and legs the reduction of activity in the flexor muscles of shoulder and forearm was observed. Concomitant arms movements increased the magnitude ofelectromiographic bursts during passive stepping-like movements in the most of recorded muscles, and the same increasing was only observed in biceps femoris and tibialis anterior muscles during active legs movement. The increasing of loading of one arm caused essential augmentation of EMG-activity in the majority of recording legs muscles. The data obtained are the additional proof of existence of functionally significant neuronal interaction both between arms and between upper and lower extremities, which is evidently depend on the intraspinal neuronal connections.     

Medial pterygoid muscle activity during the closing and compressive phases of human mastication

The lack of specific data correlating activity in the human medial pterygoid muscle with displacement of the jaw during mastication, and the hint of possible differences in function between certain mammalian species, prompted a study of unilateral mastication in six adult subjects. Muscle activity in the medial pterygoid, masseter, and anterior temporal muscles was recorded simultaneously with three-dimensional movement of an incisor point on the mandible. Signals from muscles and displacement transducer were sampled by a disc-based computer system programmed to analyze data averaged over 30 chewing cycles on each side and in some instances over 30 open-close and clench cycles. Patterns of medial pterygoid activity were consistent for the group as a whole, demonstrating activation of both muscles early in the closing cycle with strong ipsilateral muscle activity before and throughout the intercuspal phase of mastication. By contrast contralateral activity ceased during the crushing phase of the cycle, reappearing in some subjects just before the end of intercuspation. Medial pterygoid activity mirrored masseter and anterior temporal activity only during certain phases of the closing cycle, suggesting that these muscles should be considered as being selectively coactivated with, rather than synergists of, the major elevators of the jaw. The muscles were active during horizontal components of movement of the incisor teeth in chewing, but were inactive during the open-close and clench task despite vigorous contraction of the masseter muscles. Overall, the observations complement previous reports of medial pterygoid muscle activity in humans. They also confirm, for these muscles at least, a general similarity between man and the little brown bat, a relationship hitherto suspected but unsubstantiated.     

Mutual influences of upper and lower extremities during cyclic movements

The possibility for the activation of muscles in a passive arm during its cyclic movements imposed by active movements of the contralateral arm or by an experimenter and the effect that the movements of lower extremities have on the activity of the arm muscles have been studied. In addition, the activity of the leg muscles was studied as dependent on the motor task performed by the arms. Ten healthy subjects performed antiphase arm movements with and without stepping-like movements of both legs in the supine position. The experiment was performed under three conditions for the arm movements: (1) both arms performed active movements; (2) one arm performed active movements, and the contralateral arm, being entirely passive, was forced to participate in movements; (3) the movement of the passive arm was caused by an experimenter. Under condition (2), additional loadings of 30 and 60 N were applied to the active arm. Under all conditions, the arm movements were performed with and without leg movements. The possibility for the activation of muscles in the arm performing passive movements has been demonstrated. To a large extent, this is possible due to an increase in the afferent inflow from the muscles of the contralateral arm. The electrical activity was modulated during cyclic arm movements and depended on the level of loading of the active arm. During the combined active movements of the arms and legs, the reduction in the activity of the flexor muscles of the shoulder and forearm was observed. In the case of passive stepping-like movements, the concomitant arm movements increased the magnitude of electromyographic bursts in most of the examined leg muscles. During active leg movements, a similar increase in electromyographic bursts was observed only in the m. biceps femoris (BF) and the anterior tibial muscle. An increase in the loading of one arm caused a significant increase in the EMG activity in most examined muscles of the legs. The data obtained provide additional proof for the existence of a functionally significant neuronal interaction between the arms, as well as between the upper and lower extremities, which is probably due to intraspinal neuronal connections.     

Structure and direction of jaw adductor muscles as herbivorous adaptations in <Emphasis Type="Italic">Neotoma mexicana</Emphasis> (Muridae,Rodentia)

The herbivorous adaptations of the jaw adductor muscles in Neotoma mexicana were clarified by a comparative study with an unspecialized relative, Peromyscus maniculatus. In P. maniculatus, the anterior part of the deep masseter arises entirely from the lateral side of an aponeurosis, i.e., superior zygomatic plate aponeurosis, whereas N. mexicana has an additional aponeurosis for this part of the muscle, and the fibers attach on both sides of the superior zygomatic plate aponeurosis. Although the structure of the temporalis muscle is nearly identical in the two genera, a clear aponeurosis of origin occurs only in N. mexicana. These characteristics allow fibrous tissues to be processed with a large occlusal force. The deep masseter, internal pterygoid, and external pterygoid muscles of N. mexicana incline more anterodorsally than those of P. maniculatus. The transverse force component of these muscles relative to whole muscle force is smaller in N. mexicana than in P. maniculatus, with the exception of the internal pterygoid. The anterior part of the temporalis muscle of N. mexicana is specialized to produce occlusal pressure. These findings suggest that in N. mexicana a large anterior force is required to move the heavy mandible, due to the hypsodont molars, against frictional force from food, and that the posterior pull of the temporalis, which adjusts the forward force by the other jaw adductor muscles to a suitable level, need not be large for the mandibular movement.     

The structure and function of the jaw muscles in the rat (Rattus norvegicus L.)

'Tonic' and 'phasic' muscle fibre types can be distinguished histologically, using either histochemical techniques or by staining for lipid with Sudan black B. As muscles of mastication not only move the lower jaw of the rat, a 'phasic' action, but also suspend it from the cranium, a 'tonic' activity, some indication of the contribution of the major muscles to these functions has been gained from an examination of the fibre content of transverse frozen sections stained with Sudan black B. The numbers of 'pale' ('phasic') and 'dark' ('tonic') fibres were counted using a montage at a magnification of 60. Results suggest that the anterior temporal, deep masseter and external pterygoid have an important tonic action in stabilizing the position of the lower jaw as well as contributing to the production of movement; and that the superficial masseter and posterior temporal, in particular, have an almost completely phasic action. These conclusions are entirely consistent with the probable functions of the muscles inferred from their anatomy.     

Temporalis function in anthropoids and strepsirrhines: an EMG study

The major purpose of this study is to analyze anterior and posterior temporalis muscle force recruitment and firing patterns in various anthropoid and strepsirrhine primates. There are two specific goals for this project. First, we test the hypothesis that in addition to transversely directed muscle force, the evolution of symphyseal fusion in primates may also be linked to vertically directed balancing-side muscle force during chewing (Hylander et al. [2000] Am. J. Phys. Anthropol. 112:469-492). Second, we test the hypothesis of whether strepsirrhines retain the hypothesized primitive mammalian condition for the firing of the anterior temporalis, whereas anthropoids have the derived condition (Weijs [1994] Biomechanics of Feeding in Vertebrates; Berlin: Springer-Verlag, p. 282-320). Electromyographic (EMG) activities of the left and right anterior and posterior temporalis muscles were recorded and analyzed in baboons, macaques, owl monkeys, thick-tailed galagos, and ring-tailed lemurs. In addition, as we used the working-side superficial masseter as a reference muscle, we also recorded and analyzed EMG activity of the left and right superficial masseter in these primates. The data for the anterior temporalis provided no support for the hypothesis that symphyseal fusion in primates is linked to vertically directed jaw muscle forces during mastication. Thus, symphyseal fusion in primates is most likely mainly linked to the timing and recruitment of transversely directed forces from the balancing-side deep masseter (Hylander et al. [2000] Am. J. Phys. Anthropol. 112:469-492). In addition, our data demonstrate that the firing patterns for the working- and balancing-side anterior temporalis muscles are near identical in both strepsirrhines and anthropoids. Their working- and balancing-side anterior temporalis muscles fire asynchronously and reach peak activity during the power stroke. Similarly, their working- and balancing-side posterior temporalis muscles also fire asynchronously and reach peak activity during the power stroke. Compared to these strepsirrhines, however, the balancing-side posterior temporalis of anthropoids appears to have a relatively delayed firing pattern. Moreover, based on their smaller W/B ratios, anthropoids demonstrate a relative increase in muscle-force recruitment of the balancing-side posterior temporalis. This in turn suggests that anthropoids may emphasize the duration and magnitude of the power stroke during mastication. This hypothesis, however, requires additional testing. Furthermore, during the latter portion of the power stroke, the late activity of the balancing-side posterior temporalis of anthropoids apparently assists the balancing-side deep masseter in driving the working-side molars through the terminal portion of occlusion.     

Analysis of the tendinous structure in human masticatory muscles.

In the masticatory muscles, the development of bundles of the tendon was examined: they were composed of many collagen fibers and a few elastic fibers. In the masseter muscle, the property of the tendon differs in the distribution and size of collagen fibers and elastic fibers in comparison with those of other masticatory muscles. This difference is concerned with the kinetic force for the stress or the stretch of each tendon and muscle during jaw movement.     

Three-dimensional model to predict muscle forces and their relation to motor variances in reaching arm movements

A three-dimensional (3-D) arm movement model is presented to simulate kinematic properties and muscle forces in reaching arm movements. Healthy subjects performed reaching movements repetitively either with or without a load in the hand. Joint coordinates were measured. Muscle moment arms, 3-D angular acceleration, and moment of inertias of arm segments were calculated to determine 3-D joint torques. Variances of hand position, arm configuration, and muscle activities were calculated. Ratios of movement variances observed in the two conditions (load versus without load) showed no differences for hand position and arm configuration variances. Virtual muscle force variances for all muscles except deltoid posterior and EMG variances for four muscles increased significantly by moving with the load. The greatly increased variances in muscle activity did not imply equally high increments in kinematic variances. We conclude that enhanced muscle cooperation through synergies helps to stabilize movement at the kinematic level when a load is added.     

Suboccipital muscles in the cat neck: Morphometry and histochemistry of the rectus capitis muscle complex

The morphometry, histochemistry, and biomechanical relationships of rectus capitis muscles were examined in adult cats. This family of muscles contained six members on the dorsal, ventral, and lateral aspects of the upper cervical vertebral column. Three dorsal muscles (rectus capitis posterior major, medius, and minor) formed a layered complex spanning from C1 and C2 to the skull. Rectus capitis posterior major was composed predominantly of fast fibers, but the other two deeper muscles contained progressively higher proportions of slow fibers. One ventral muscle, rectus capitis anterior major, was architecturally complex. It originated from several cervical vertebrae and appeared to be divided into two different heads. In contrast, rectus capitis anterior minor and rectus capitis lateralis were short, parallel-fibered muscles spanning between the skull and C1. The ventral muscles all had nonuniform distributions of muscle-fiber types in which fast fibers predominated. Dorsal and ventral muscle groupings usually had cross-sectional areas of 0.5 cm² or more, reflecting a potential capacity to generate maximal tetanic force in excess of 9 N. Biomechanical analyses suggested that one muscle, rectus capitis lateralis, had its largest moment in lateral flexion, whereas the other muscles had large, posturally dependent moment arms appropriate for actions in flexion-extension. The observation that most rectus muscles have relatively large cross-sectional areas and high fast-fiber proportions suggests that the muscles may have important phasic as well as postural roles during head movement. © 1993 Wiley-Liss, Inc.     

Three-dimensional temporomandibular joint muscle attachment morphometry and its impacts on musculoskeletal modeling

In musculoskeletal models of the human temporomandibular joint (TMJ), muscles are typically represented by force vectors that connect approximate muscle origin and insertion centroids (centroid-to-centroid force vectors). This simplification assumes equivalent moment arms and muscle lengths for all fibers within a muscle even with complex geometry and may result in inaccurate estimations of muscle force and joint loading. The objectives of this study were to quantify the three-dimensional (3D) human TMJ muscle attachment morphometry and examine its impact on TMJ mechanics. 3D muscle attachment surfaces of temporalis, masseter, lateral pterygoid, and medial pterygoid muscles of human cadaveric heads were generated by co-registering measured attachment boundaries with underlying skull models created from cone-beam computerized tomography (CBCT) images. A bounding box technique was used to quantify 3D muscle attachment size, shape, location, and orientation. Musculoskeletal models of the mandible were then developed and validated to assess the impact of 3D muscle attachment morphometry on joint loading during jaw maximal open-close. The 3D morphometry revealed that muscle lengths and moment arms of temporalis and masseter muscles varied substantially among muscle fibers. The values calculated from the centroid-to-centroid model were significantly different from those calculated using the ‘Distributed model’, which considered crucial 3D muscle attachment morphometry. Consequently, joint loading was underestimated by more than 50% in the centroid-to-centroid model. Therefore, it is necessary to consider 3D muscle attachment morphometry, especially for muscles with broad attachments, in TMJ musculoskeletal models to precisely quantify the joint mechanical environment critical for understanding TMJ function and mechanobiology.     

Adaptation of muscle coordination to altered task mechanics during steady-state cycling

The objective of this work was to increase our understanding of how motor patterns are produced during movement tasks by quantifying adaptations in muscle coordination in response to altered task mechanics. We used pedaling as our movement paradigm because it is a constrained cyclical movement that allows for a controlled investigation of test conditions such as movement speed and effort. Altered task mechanics were introduced using an elliptical chainring. The kinematics of the crank were changed from a relatively constant angular velocity using a circular chainring to a widely varying angular velocity using an elliptical chainring. Kinetic, kinematic and muscle activity data were collected from eight competitive cyclists using three different chainrings--one circular and two different orientations of an elliptical chainring. We tested the hypotheses that muscle coordination patterns (EMG timing and magnitude), specifically the regions of active muscle force production, would shift towards regions in the crank cycle in which the crank angular velocity, and hence muscle contraction speeds, were favorable to produce muscle power as defined by the skeletal muscle power-velocity relationship. The results showed that our hypothesis with regards to timing was not supported. Although there were statistically significant shifts in muscle timing, the shifts were minor in absolute terms and appeared to be the result of the muscles accounting for the activation dynamics associated with muscle force development (i.e. the delay in muscle force rise and decay). But, significant changes in the magnitude of muscle EMG during regions of slow crank angular velocity for the tibialis anterior and rectus femoris were observed. Thus, the nervous system used adaptations to the muscle EMG magnitude, rather than the timing, to adapt to the altered task mechanics. The results also suggested that cyclists might work on the descending limb of the power-velocity relationship when pedaling at 90 rpm and sub-maximal power output. This finding might have important implications for preferred pedaling rate selection.     

Further evaluation of an EMG technique for assessment of the deep cervical flexor muscles

A novel surface electromyographic (EMG) technique was recently described for the detection of deep cervical flexor muscle activity. Further investigation of this technique is warranted to ensure EMG activity from neighbouring muscles is not markedly influencing the signals recorded. This study compared deep cervical flexor (DCF) muscle activity with the activity of surrounding neck and jaw muscles during various anatomical movements of the neck and jaw in 10 volunteer subjects. DCF EMG activity was recorded with custom electrodes inserted via the nose and fixed by suction to the posterior mucosa of the oropharynx. Surface electrodes were placed over the sternocleidomastoid, anterior scalene, masseter and suprahyoid muscles. Positioned in supine, subjects performed isometric cranio-cervical flexion, cervical flexion, right and left cervical rotation, jaw clench and resisted jaw opening. Across all movements examined, EMG amplitude of the DCF muscles was greatest during neck movements that would require activity of the DCF muscles, particularly during cranio-cervical flexion, their primary anatomical action. The actions of jaw clench and resisted jaw opening demonstrated significantly less DCF EMG activity than the cranio-cervical flexion action (p < 0.05). Across all other movements, the neighbouring neck and jaw muscles demonstrated greatest EMG amplitude during their respective primary anatomical actions, which occurred in the absence of increased EMG amplitude recorded from the DCF muscles. The finding of substantial EMG activity of the DCF muscles only during neck actions that would require their activity, particularly cranio-cervical flexion, and not during actions involving the jaw, provide further assurance that the majority of myoelectric signals detected from the nasopharyngeal electrode are from the DCF muscles.     

Anterior hip joint force increases with hip extension, decreased gluteal force, or decreased iliopsoas force

Abnormal or excessive force on the anterior hip joint may cause anterior hip pain, subtle hip instability and a tear of the acetabular labrum. We propose that both the pattern of muscle force and hip joint position can affect the magnitude of anterior joint force and thus possibly lead to excessive force and injury. The purpose of this study was to determine the effect of hip joint position and of weakness of the gluteal and iliopsoas muscles on anterior hip joint force. We used a musculoskeletal model to estimate hip joint forces during simulated prone hip extension and supine hip flexion under four different muscle force conditions and across a range of hip extension and flexion positions. Weakness of specified muscles was simulated by decreasing the modeled maximum force value for the gluteal muscles during hip extension and the iliopsoas muscle during hip flexion. We found that decreased force contribution from the gluteal muscles during hip extension and the iliopsoas muscle during hip flexion resulted in an increase in the anterior hip joint force. The anterior hip joint force was greater when the hip was in extension than when the hip was in flexion. Further studies are warranted to determine if increased utilization of the gluteal muscles during hip extension and of the iliopsoas muscle during hip flexion, and avoidance of hip extension beyond neutral would be beneficial for people with anterior hip pain, subtle hip instability, or an anterior acetabular labral tear.     

Functional heterogeneity in a multipinnate muscle.

Many mammalian muscles have a complex internal architecture. This type of structure could allow a single muscle to produce a variety of force vectors through selective regional contractions. This hypothesis was tested electromyographically in the multipinnate pig masseter by recording simultaneously from several intramuscular sites. It was found that the activity in different portions of the masseter varied systematically during the various phases of mastication. anatomical correlates of the differential activity included fasciculus orientation and length, sarcomere length in specific jaw positions, and histochemical fiber type. The usual assumptions made about muscles for biomechanical analysis, such as uniform contraction and constant line of action, are inappropriate for complex muscles such as the pig masseter.     

Coactivation of jaw muscles: recruitment order and level as a function of bite force direction and magnitude

The aim of this study was to obtain insight into the coactivation behaviour of the jaw muscles under various a priori defined static loading conditions of the mandible. As the masticatory system is mechanically redundant, an infinite number of recruitment patterns is theoretically possible to produce a certain bite force. Using a three-component force transducer and a feedback method, subjects could be instructed to produce a bite force of specific direction and magnitude under simultaneous registration of the EMG activity of anterior and posterior temporal, masseter and digastric muscles on each side. Forces were measured at the second premolars. Vertical, anterior, posterior, lateral and medial force directions were examined; in each direction force levels between 50 N and maximal voluntary force were produced. The results show that for all muscles the bite force-EMG relationship obeys a straight-line fit for forces exceeding 50 N. The relationship varies with bite force direction, except in the case of the digastric muscles. Variation is small for the anterior temporal and large for the posterior temporal and masseter muscles. The relative activation of muscles for a particular force in a particular direction in unique, despite the redundancy.     

Mandible position and activation of submental and masseter muscles during sleep.

Movement of the mandible could influence pharyngeal airway caliber because the mandible is attached to the tongue and to muscles that insert on the hyoid bone. In normal subjects and patients with obstructive sleep apnea (OSA) we measured jaw position during sleep with strain gauges, as well as masseter and submental electromyograms, airflow, esophageal pressure, oximetry, electroencephalograms, and electrooculograms. Jaws of patients with OSA were open more than those of normal subjects at end expiration and opened further at end inspiration, particularly at the termination of apneas when the masseter and submental muscles contracted. Masseter activation occurred only in patients with OSA and in a pattern similar to that of submental muscles. Jaw opening at end expiration could narrow the upper airway, whereas opening at end inspiration could reflect efforts to expand the airway with tracheal tug and with submental muscle activation and efforts to open the mouth to allow mouth breathing. Masseter contraction does not close the jaw but may serve to stabilize it.     

Analyzing surface EMG signals to determine relationship between jaw imbalance and arm strength loss

ABSTRACT: BACKGROUND: This study investigated the relationship between dental occlusion and arm strength; in particular, the imbalance in the jaw can cause loss in arm strength phenomenon. One of the goals of this study was to record the maximum forces that the subjects can resist against the pull-down force on their hands while biting a spacer of adjustable height on the right or left side of the jaw. Then EMG measurement was used to determine the EMG-Force relationship of the jaw, neck and arms muscles. This gave us useful insights on the arms strength loss due to the biomechanical effects of the imbalance in the jaw mechanism. METHODS: In this study to determine the effects of the imbalance in the jaw to the strength of the arms, we conducted experiments with a pool of 20 healthy subjects of both genders. The subjects were asked to resist a pull down force applied on the contralateral arm while biting on a firm spacer using one side of the jaw. Four different muscles -- masseter muscles, deltoid muscles, bicep muscles and trapezoid muscles -- were involved. Integrated EMG (iEMG) and Higuchi fractal dimension (HFD) were used to analyze the EMG signals. RESULTS: The results showed that (1) Imbalance in the jaw causes loss of arm strength contra-laterally; (2) The loss is approximately a linear function of the height of the spacers. Moreover, the iEMG showed the intensity of muscle activities decreased when the degrees of jaw imbalance increased (spacer thickness increased). In addition, the tendency of Higuchi fractal dimension decreased for all muscles. CONCLUSIONS: This finding indicates that muscle fatigue and the decrease in muscle contraction level leads to the loss of arm strength.     

Neural control exploits changing mechanical advantage and context dependence to generate different feeding responses in <Emphasis Type="Italic">Aplysia</Emphasis>

How does neural control reflect changes in mechanical advantage and muscle function? In the Aplysia feeding system a protractor muscle's mechanical advantage decreases as it moves the structure that grasps food (the radula/odontophore) in an anterior direction. In contrast, as the radula/odontophore is moved forward, the jaw musculature's mechanical advantage shifts so that it may act to assist forward movement of the radula/odontophore instead of pushing it posteriorly. To test whether the jaw musculature's context-dependent function can compensate for the falling mechanical advantage of the protractor muscle, we created a kinetic model of Aplysia's feeding apparatus. During biting, the model predicts that the reduction of the force in the protractor muscle I2 will prevent it from overcoming passive forces that resist the large anterior radula/odontophore displacements observed during biting. To produce protractions of the magnitude observed during biting behaviors, the nervous system could increase I2's contractile strength by neuromodulating I2, or it could recruit the I1/I3 jaw muscle complex. Driving the kinetic model with in vivo EMG and ENG predicts that, during biting, early activation of the context-dependent jaw muscle I1/I3 may assist in moving the radula/odontophore anteriorly during the final phase of protraction. In contrast, during swallowing, later activation of I1/I3 causes it to act purely as a retractor. Shifting the timing of onset of I1/I3 activation allows the nervous system to use a mechanical equilibrium point that allows I1/I3 to act as a protractor rather than an equilibrium point that allows I1/I3 to act as a retractor. This use of equilibrium points may be similar to that proposed for vertebrate control of movement.