Experimental elicitation of jaw closure reflex & lock-jaw in cats.
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Morphology and evolution of the jaw suspension in lamniform sharks.
The morphology of the jaw suspension and jaw protrusion mechanism in lamniform sharks is described and mapped onto a cladogram to investigate how changes in jaw suspension and protrusion have evolved. This has revealed that several evolutionary modifications in the musculoskeletal apparatus of the jaws have taken place among lamniform sharks. Galeomorph sharks (Carcharhiniformes, Lamniformes, Orectolobiformes, and Heterodontiformes) have paired ethmopalatine ligaments connecting the ethmoid process of the upper jaw to the ethmoid region of the cranium. Basal lamniform sharks also acquired a novel single palatonasal ligament connecting the symphysis of the upper jaw to the cranium mid-ventral to the nasal capsule. Sharks in the family Lamnidae subsequently lost the original paired ethmopalatine ligament while retaining the novel palatonasal ligament. Thus, basal lamniform taxa (Mitsukurina owstoni, Carcharius taurus, Alopias vulpinnis) have increased ligamentous support of the lateral region of the upper jaw while derived species (Lamnidae) have lost this lateral support but gained anterior support. In previous studies the morphology of the jaw suspension has been shown to play a major role in the mechanism of upper jaw protrusion in elasmobranchs. The preorbitalis is the primary muscle effecting upper jaw protrusion in squalean (sister group to galeomorphs) and carcharhiniform (sister group to lamniforms) sharks. The preorbitalis originates from the quadratomandibularis muscle and inserts onto the nasal capsule in squalean and carcharhiniform sharks. Carcharhiniform sharks have evolved a subdivided preorbitalis muscle with the new division inserting near the ethmoid process of the palatoquadrate (upper jaw). Alopid sharks have also independently evolved a partially subdivided preorbitalis with the new division inserting at the base of the ethmoid process and surrounding connective tissue. Lamnid sharks have retained the two preorbitalis divisions but have modified both of the insertion points. The original ventral preorbitalis division now inserts onto the connective tissue surrounding the mid-region of the upper jaw, while the new dorsal preorbitalis division inserts onto the surrounding connective tissue and skin at a more posterior position on the upper jaw. The retractor muscle of the jaws, the levator hyomandibularis, has also been modified during the evolution of lamniform sharks. In most sharks, including basal lamniforms, the levator hyomandibularis inserts onto the hyomandibula and functions to retract the jaws after protrusion. In alopid and lamnid sharks the levator hyomandibularis inserts primarily onto the upper and lower jaws around the jaw joint and is a more direct route for retracting the jaws. Thus, there has been at least one instance of character loss (ethmopalatine ligament), acquisition (palatonasal ligament), subdivision (preorbitalis), and modification (ventral preorbitalis, dorsal preorbitalis, and levator hyomandibularis) in the ligaments and muscles associated with the jaw suspension and jaw protrusion mechanism in lamniform sharks. While derived lamniform sharks (Lamna nasus, Carcharodon carcharius, and Isurus oxyrinchus) lost the ancestral passive lateral support of the ethmoid articulation of the upper jaw, they simultaneously acquired muscular support by way of the levator hyomandibularis, which provides a dynamic mechanism for lateral support. The evolution of multiple divisions of preorbitalis insertions onto the palatoquadrate and modification of the levator hyomandibularis insertion directly onto the jaws provides an active mechanism for multiple protractions and retractions of the upper jaw, which is advantageous in those sharks that gouge or saw pieces from large oversized prey items.
Animals↗
Dopaminergic and cholinergic stimulation of the ventrolateral striatum elicit rat jaw movements that are funnelled via distinct efferents.
It has been reported that two distinct types of jaw movements can be elicited by bilateral injections of drugs into the ventrolateral striatum: (1) dopamine receptor-mediated jaw movements that are elicited by a mixture of (+/-)-6-chloro-7,8-dihydroxy-3-allyl-1-phenyl-2,3,4,5-tetrahydro-1H-3-benzazepine-7,8-diol (SKF 82958; 5 microg) and quinpirole (10 microg), and (2) acetylcholine receptor-mediated jaw movements that are elicited by carbachol (2.5 microg). In the present study, electromyographic analysis was used to characterise these movements: the dopamine receptor-mediated jaw movements were marked by a dominant digastric activity during jaw opening and a dominant masseter activity during jaw closing (digastric/masseter type), whereas the acetylcholine receptor-mediated jaw movements were marked by a dominant digastric activity during jaw opening without any significant change in masseter activity during jaw closing (digastric type). The main goal was to (in)validate the hypothesis that these two types of jaw movements are funnelled via distinct gamma-aminobutyric acid (GABA)ergic output channels. Bilateral injections of both muscimol (25 and 50 ng/0.2 microl per side) and bicuculline (50 and 150 ng/0.2 microl per side) into the ventral pallidum, entopeduncular nucleus or dorsolateral part of the substantia nigra pars reticulata essentially inhibited dopamine receptor-mediated jaw movements to various degrees. In contrast, acetylcholine receptor-mediated jaw movements were inhibited by muscimol given into the entopeduncular nucleus and dorsolateral part of the substantia nigra pars reticulata, whereas these movements were enhanced by bicuculline. The acetylcholine receptor-mediated jaw movements were not affected by muscimol injections into the ventral pallidum, but were inhibited by bicuculline injections. Studies on such injections into the ventral pallidum, entopeduncular nucleus or dorsolateral part of the substantia nigra pars reticulata of naive rats revealed that jaw movements of the digastric/masseter type were elicited either by muscimol injections into the dorsolateral part of the substantia nigra pars reticulata or by combined injections of muscimol and bicuculline into the entopeduncular nucleus, and that jaw movements of the digastric type were elicited only by combined injections of muscimol and bicuculline into the entopeduncular nucleus. Together, the data allow the conclusion that dopamine receptor-mediated and acetylcholine receptor-mediated jaw movements are two distinct types of jaw movements that are funnelled via separate GABAergic output channels. It is suggested that the three different profiles of responses to GABAergic drugs in animals showing either dopamine receptor-mediated or acetylcholine receptor-mediated jaw movements reflect the involvement of three distinct types of output neurons of the striatum, namely: type I neurons with collateralised axons to the ventral pallidum, entopeduncular nucleus and dorsolateral part of the substantia nigra pars reticulata, mediating the dopamine receptor-mediated jaw movements; type II neurons with collateralised axons to the globus pallidus that, in turn, project to the entopeduncular nucleus and the dorsolateral part of the substantia nigra pars reticulata, mediating directly the acetylcholine receptor-mediated jaw movements; and type III neurons with a single axon to the ventral pallidum, mediating indirectly the acetylcholine receptor-mediated movements. It is evident that future studies are required to provide direct evidence in favour of the latter hypothesis.
Animals↗
Comparison of external load compensation during rhythmic arm movements and rhythmic jaw movements in humans.
Experiments were performed on human elbow flexor and extensor muscles and jaw-opening and -closing muscles to observe the effect on rhythmic movements of sudden loading. The load was provided by an electromagnetic device, which simulated the appearance of a smoothly increasing spring-like load. The responses to this loading were compared in jaw and elbow movements and between expected and unexpected disturbances. All muscles showed electromyographic responses to unexpected perturbations, with latencies of approximately 65 ms in the arm muscles and 25 ms in the jaw. When loading was predictable, anticipatory responses started in arm muscles approximately 200 ms before and in jaw muscles 100 ms before the onset of loading. The reflex responses relative to the anticipatory responses were smaller for the arm muscles than for the jaw muscles. The reflex responses in the arm muscles were the same with unexpected and expected perturbations, whereas anticipation increased the reflex responses in the jaw muscles. Biceps brachii and triceps brachii showed similar sensory-induced responses and similar anticipatory responses. Jaw muscles differed, however, in that the reflex response was stronger in masseter than in digastric. It was concluded that reflex responses in the arm muscles cannot overcome the loading of the arm adequately, which is compensated by a large centrally programmed response when loading is predictable. The jaw muscles, particularly the jaw-closing muscles, tend to respond mainly through reflex loops, even when loading of the jaw is anticipated. The differences between the responses of the arm and the jaw muscles may be related to physical differences. For example, the jaw was decelerated more strongly by the load than the heavier arm. The jaw was decelerated strongly but briefly, <30 ms during jaw closing, indicating that muscle force increased before the onset of reflex activity. Apparently, the force-velocity properties of the jaw muscles have a stabilizing effect on the jaw and have this effect before sensory induced responses occur. The symmetrical responses in biceps and triceps indicate similar motor control of both arm muscles. The differences in reflex activity between masseter and digastric muscle indicate fundamental differences in sensory feedback to the jaw-closing muscle and jaw-opening muscle.
Adult↗
Micro- and macroevolutionary decoupling of cichlid jaws: a test of Liem's key innovation hypothesis.
The extent to which elements of functional systems can change independently (modularity) likely influences the diversification of lineages. Major innovations in organismal design, like the pharyngeal jaw in cichlid fishes, may be key to a group's success when they relax constraints on diversification by increasing phenotypic modularity. In cichlid fishes, pharyngeal jaw modifications that enhanced the ability to breakdown prey may have freed their oral jaws from serving their ancestral dual role as a site of both prey capture and prey processing. This functional decoupling that allowed the oral jaws to become devoted solely to prey capture has been hypothesized to have permitted the two sets of cichlid jaws to evolve independently. We tested the hypothesis that oral and pharyngeal jaw mechanics are evolutionarily decoupled both within and among Neotropical Heroine cichlids. In the trophically polymorphic species Herichthys minckleyi, molariforms that exhibit enlarged molarlike pharyngeal jaw teeth were found to have approximately 400% greater lower jaw mass compared to H. minckleyi with the alternative papilliform pharyngeal morphology. However, oral jaw gape, lower jaw velocity ratios, anterior jaw linkage mechanics, and jaw protrusion did not differ between the morphotypes. In 40 other Heroine species, there was a weak correlation between oral jaw mechanics and pharyngeal jaw mass when phylogenetic history was ignored. Yet, after expansion of the cytochrome b phylogeny for Heroines, change in oral jaw mechanics was found to be independent of evolutionary change in pharyngeal jaw mass based on independent contrasts. Evolutionary decoupling of oral and pharyngeal jaw mechanics has likely played a critical role in the unparalleled trophic diversification of cichlid fishes.
Animals↗
A dynamic model of mouth closing movements in clariid catfishes: the role of enlarged jaw adductors.
Some species of Clariidae (air breathing catfishes) have extremely large (hypertrophied) jaw closure muscles. Besides producing higher bite forces, the enlarged muscles may also cause higher accelerations of the lower jaw during rapid mouth closure. Thus, jaw adductor hypertrophy could potentially also enable faster mouth closure. In this study, a forward dynamic model of jaw closing is developed to evaluate the importance of jaw adductor hypertrophy on the speed of mouth closure. The model includes inertia, pressure, tissue resistance and hydrodynamic drag forces on the lower jaw, which is modelled as a rotating half-ellipse. Simulations are run for four clariid species showing a gradual increase in jaw adductor hypertrophy (Clarias gariepinus, Clariallabes longicauda, Gymnallabes typus and Channallabes apus). The model was validated using data from high-speed videos of prey captures in these species. In general, the kinematic profiles of the fastest mouth closure from each species are reasonably well predicted by the model. The model was also used to compare the four species during standardized mouth closures (same initial gape angle, travel distance and cranial size). These simulations suggest that the species with enlarged jaw adductors have an increased speed of jaw closure (in comparison with the non-hypertrophied C. gariepinus) for short lower jaw rotations and when feeding at high gape angles. Consequently, the jaw system in these species seems well equipped to capture relatively large, evasive prey. For prey captures during which the lower jaw rotates freely over a larger distance before impacting the prey, the higher kinematic efficiency of the C. gariepinus jaw system results in the fastest jaw closures. In all cases, the model predicts that an increase in the physiological cross-sectional area of the jaw muscles does indeed contribute to the speed of jaw closure in clariid fish.
Animals↗
Control of jaw orientation and position in mastication and speech.
1. The kinematics of sagittal-plane jaw motion were assessed in mastication and speech. The movement paths were described in joint coordinates, in terms of the component rotations and translations. The analysis focused on the relationship between rotation and horizontal translation. Evidence was presented that these can be separately controlled. 2. In speech, jaw movements were studied during consonant-vowel utterances produced at different rates and volumes. In mastication, bolus placement, compliance, and size as well as chewing rate were manipulated. Jaw movements were recorded using the University of Wisconsin X-ray microbeam system. Jaw rotation and translation were calculated on the basis of the motion of X-ray tracking pellets on the jaw. 3. The average magnitudes of jaw rotation and translation were greater in mastication than in speech. In addition, in speech, it was shown that the average rotation magnitude may vary independent of the horizontal translation magnitude. In mastication, the average magnitude of vertical jaw translation was not dependent on the magnitudes of jaw rotation or horizontal jaw translation. 4. The magnitude of rotation and horizontal jaw translation tended to be correlated when examined on a trial by trial basis. Some subjects also showed a correlation between jaw rotation and vertical jaw translation. However, the proportion of variance accounted for was greater for all subjects in the case of rotation and horizontal translation. 5. Joint space paths in both mastication and speech were found to be straight. The pattern was observed at normal and fast rates of speech and mastication and for loud speech as well. Straight line paths were also observed when subjects produced utterances that had both the syllabic structure and the intonation pattern of speech. The findings suggest that control may be organized in terms of an equilibrium jaw orientation and an equilibrium jaw position. 6. Departures from linearity were also observed. These were typically associated with differences during jaw closing in the end time of rotation and translation. Start time differences were not observed in jaw closing and the movement paths were typically linear within this region.
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Evolution and mechanics of long jaws in butterflyfishes (family Chaetodontidae).
We analyzed the functional morphology and evolution of the long jaws found in several butterflyfishes. We used a conservative reanalysis of an existing morphological dataset to generate a phylogeny that guided our selection of seven short- and long-jawed taxa in which to investigate the functional anatomy of the head and jaws: Chaetodon xanthurus, Prognathodes falcifer (formerly Chaetodon falcifer), Chelmon rostratus, Heniochus acuminatus, Johnrandallia nigrirostris, Forcipiger flavissimus, and F. longirostris. We used manipulations of fresh, preserved, and cleared and stained specimens to develop mechanical diagrams of how the jaws might be protruded or depressed. Species differed based on the number of joints within the suspensorium. We used high-speed video analysis of five of the seven species (C. xanthurus, Chel. rostratus, H. acuminatus, F. flavissimus, and F. longirostris) to test our predictions based on the mechanical diagrams: two suspensorial joints should facilitate purely anteriorly directed protrusion of the lower jaw, one joint should allow less anterior protrusion and result in more depression of the lower jaw, and no joints in the suspensorium should constrain the lower jaw to simple ventral rotation around the jaw joint, as seen in generalized perciform fishes. We found that the longest-jawed species, F. longirostris, was able to protrude its jaws in a predominantly anterior direction and further than any other species. This was achieved with little input from cranial elevation, the principal input for other known lower jaw protruders, and is hypothesized to be facilitated by separate modifications to the sternohyoideus mechanism and to the adductor arcus palatini muscle. In F. longirostris the adductor arcus palatini muscle has fibers oriented anteroposteriorly rather than medial-laterally, as seen in most other perciforms and in the other butterflyfish studied. These fibers are oriented such that they could rotate the ventral portion of the quadrate anteriorly, thus projecting the lower jaw anteriorly. The intermediate species lack modification of the adductor arcus palatini and do not protrude their jaws as far (in the case of F. flavissimus) or in a purely anterior fashion (in the case of Chel. rostratus). The short-jawed species both exhibit only ventral rotation of the lower jaw, despite the fact that H. acuminatus is closely related to Forcipiger.
Animals↗
Tremulous characteristics of the vacuous jaw movements induced by pilocarpine and ventrolateral striatal dopamine depletions.
Vacuous jaw movements induced by the muscarinic agonist pilocarpine and striatal dopamine depletions were examined using a slow motion videotape system. With this procedure, rats were videotaped in a Plexiglas tube so that the profile of the head region could be seen. Vacuous jaw movements were analyzed by examining the tape at 1/6 normal speed. An observer recorded each jaw movement using a computer, and the computer program re-calculated the temporal characteristics of jaw movement responses back to normal speed. The interresponse time was recorded for each jaw movement, and each jaw movement interresponse time was assigned to a 50 ms wide time bin. Thus, the distribution of interresponse times could be used to analyze the temporal characteristics of jaw movement responses. In the first experiment, rats were administered saline vehicle, 1.0 mg/kg and 2.0 mg/kg pilocarpine. The rats were videotaped 10-15 min after injection, and the data were analyzed as described above. Pilocarpine induced very high levels of vacuous jaw movements, and the vast majority of all movements occurred in "bursts" with interresponse times of 1.0 s or less. Analysis of the interresponse time distributions showed that most of the jaw movements were within the 150-350 ms range. The modal jaw movement interresponse time was in the 150-200 ms range, which corresponds to a local frequency of 5-6.66 Hz. In the second experiment, the neurotoxic agent 6-hydroxydopamine was injected directly into the ventrolateral striatum in order to produce a local dopamine depletion. The dopamine-depleted rats were observed for jaw movements 7 days after surgery. The overall level of jaw movement activity resulting from dopamine-depletion was much lower than that produced by pilocarpine. There was a significant inverse correlation between ventrolateral striatal dopamine levels and total number of vacuous jaw movements. Videotape analysis indicated that the temporal characteristics of jaw movements induced by dopamine depletions were similar to those shown with pilocarpine. These experiments indicate that vacuous jaw movements induced by pilocarpine and striatal dopamine depletion occur in a frequency range similar to that shown in parkinsonian tremor.
Animals↗
Behavioral and electromyographic characterization of the local frequency of tacrine-induced tremulous jaw movements.
Rats were implanted with fine-wire electromyograph (EMG) electrodes and were videotaped to identify the local frequency characteristics and muscle activity associated with tacrine-induced tremulous jaw movements. All rats received intraperitoneal injections of 2.5 mg/kg tacrine. The videotape sessions were played back in slow motion (i.e., one-sixth normal speed), and an observer entered each jaw movement into a computer program that recalculated the interresponse time and the local frequency (in hertz) for each movement within a burst. Analyses of the distribution of frequencies showed that the peak frequency of jaw movements was in the 3- to 5-Hz frequency range, with an average frequency of 4.0 Hz. EMG electrodes were implanted into three jaw muscles: temporalis, anterior belly of digastricus, and masseter. Tremulous jaw movements were not accompanied by consistent changes in masseter activity. The anterior belly of digastricus showed bursts of EMG activity during some jaw movements, although the temporal relation between jaw movements and EMG activity was somewhat inconsistent. The muscle that showed activity most closely related to tremulous jaw movements was the temporalis. During bursts of jaw movements, temporalis muscles across several different rats showed bursts of EMG activity. Sections of videotape corresponding to bursts of EMG activity were reanalyzed by freeze-frame examination of the tape; typically, the temporalis showed a burst for each jaw movement, with the burst of activity occurring during the jaw-closing phase and the transition between jaw closing and opening. These results indicate that the local frequency of tremulous jaw movements is within the 3- to 7-Hz frequency that is typically associated with parkinsonian tremor. Moreover, the EMG data suggest that temporalis is a major contributor to the muscle activity that underlies tremulous jaw movements.
Animals↗
Coordination of cortically induced rhythmic jaw and tongue movements in the rabbit.
1. Rhythmic movements of the jaw, tongue, and hyoid that were induced by stimulation of the cortical masticatory area (CMA) were recorded cineradiographically in the anesthetized rabbit. Jaw movements were also recorded by a laser position detector. 2. The evoked jaw movements were classified into four types: small circular (type A), large circular (type B), large vertical (type C), and crescent-shaped (type D). Among these, types B and D resembled the jaw movements of the food transport cycle and those of the chewing cycle in a masticatory sequence. 3. Each type of jaw movement was associated with a particular pattern of tongue and hyoid movements. In general, the tongue protruded during jaw opening and retracted during jaw closure. The hyoid generally moved upward and forward during jaw opening but downward and backward during jaw closure. 4. Electromyograms (EMGs) were recorded from jaw muscles [masseter (Ma) and digastric (Di) muscles], extrinsic tongue muscles [styloglossus (Sg) and genioglossus (Gg) muscles], and hyoid muscles [sternohyoid (Sh) and geniohyoid (Gh) muscles] during cortically induced rhythmic jaw and tongue movements (CRJTMs). These muscles were classified into two groups: group 1 was activated mainly in the jaw opening phase, and group 2 was activated mainly in the jaw closing and power phases. The Di, Gg, and Gh were included in the former, and the Ma, Sg, and Sh were included in the latter. 5. The timings of EMG activation to a jaw movement cycle were relatively constant for the muscles of group 1, irrespective of the types of CRJTMs, whereas those for the muscles of group 2 altered considerably with the different types of CRJTMs. 6. Relationships of the integrated muscle activity between the Di and Gg and between the Di and Gh were significant, whereas those between the Ma and Sg and between the Ma and Sh were not. 7. When a small strip of polyurethane form of various degrees of hardness was inserted between the opposing molars during CRJTMs, EMG activity of the muscles of group 2 increased with the hardness of the strip. On the other hand, EMG activities of the muscles of group 1 were less affected by the same intraoral stimuli. 8. Two conclusions were reached: first, physiological properties of the CRJTMs and cortically induced rhythmic movements of the hyoid were essentially similar to those observed in natural mastication. This fictive mastication might thus be regarded as a suitable model for simulating natural mastication.(ABSTRACT TRUNCATED AT 400 WORDS)
Animals↗
[Biomechanics of the jaw apparatus in the horn-shark (Heterodontus portusjacksoni = Heterodontus philippi)].
The constructional morphology of the jaw apparatus in the horn-shark Heterodontus has been investigated. The origines and insertions of the jaw muscles have been delimited and the lines of action were determined. The individual muscles have been weighed, in order to get on the basis of their masses an estimate of the forces which are exerted by the components of the jaw musculature. The mandibular joints, the occlusion of the jaws, morphological details of the upper and lower jaws as well as the "subodontium" have been subject to macroscopic and microscopic study. The joint between palatoquadrate and mandibular consists of a medial and a lateral compartment. Only hinge movements are possible. If the jaws are closed, the contacts between upper and lower jaws are confined to the large crushing teeth of only one or two tooth families. The teeth are fixed to the jaw cartilage by ligamentous structures. Three layers can be discerned histologically: The uppermost layer, beneath the bases of the teeth, is composed of the fibrae interdentales and of the fibrae subbasales. By these, the individual teeth are firmly connected to form a continuous pavement. - The middle layer is characterised by the great number of cell nuclei. - The fibre system which constitutes the lowermost layer is arranged according to its function. The subdental layer of fibrous tissue grows faster than the one adjacent to the jaws. Thus a particular growth structure is formed. - Two constructive principles are realised: 1. The biting forces or "loads", applied to one or two teeth, are split and distributed on all teeth of the same family which are lingual of the loaded one. 2. As in the thecodont mammals, the compressive biting (= occlusal) force is transformed into a tensile force by the tooth-fixing apparatus and by the shapes of the jaws and this is sustained by fibrous structures. Magnitude and directions of the stresses which appear in the upper and lower jaw during biting are compared with stress patterns evoked in consoles and in beams of technical constructions. The moments of resistence have been calculated for 9 or 8 cross sections, respectively, through the upper and the lower jaw. The forms of both jaws are comparable to a beam of equal strength on two supports, namely, bitten object and joints; while the load is represented by the muscular force. Where the bending moments are at a maximum, opposite to the muscle insertions, a strong ligamentous reinforcement of the perichondrium extends parallel to the tooth rows and fades out towards the joint and towards the symphysis. The strength properties of cartilage and of collagenous fibres in the species under consideration have been measured. The lines of action of the jaw muscles intersect in the area where the moments of resistence reach their maximum. The largest crushing teeth are located in this same section, so that the muscles exert most force on the teeth, while the joint is exposed to only moderate compression.
Animals↗
Regenerative capability of upper and lower jaws in the newt.
The regenerating amphibian jaw represents an important model for studying pattern formation and the mechanisms underlying regeneration of facial structures. We have studied regeneration of upper and lower jaws in the urodele amphibian, Notophthalmus viridescens, using whole mount preparations stained for bone and cartilage, scanning electron microscopy and immunocytochemistry to further characterize these regenerating systems. In addition, we have investigated whether lower jaws of adults and larvae display similar regenerative ability. Although in adult animals the original shape of both the lower and upper jaws is rather faithfully reproduced following amputation, and the teeth and oral mucosa with its specialized sensory organs fully regenerate, significant differences in the regenerative ability of the various skeletal elements are observed. In fact, only tooth-bearing skeletal elements ossify, while the other elements of the regenerated skeleton remain cartilaginous for as long as 5 months after amputation. In contrast, a regenerated lower jaw in the larva is indistinguishable from an unamputated one at the same stage of development. Interestingly, regenerating adult jaws form directly bicuspid teeth, which are the type of teeth normally found in the adult, rather than the monocuspid teeth characteristic of larval jaws, indicating that jaw regeneration is not a recapitulation of development, in that an adult jaw blastema directly regenerates an adult jaw. Finally, we have studied the expression of tissue specific markers in normal and regenerating upper and lower jaws to establish whether the blastemal cells, which will form the missing part of the jaw, express any of these markers of the differentiated state, or are undifferentiated as suggested by their morphological appearance. Under our experimental conditions, no expression of markers of the differentiated state, such as those for muscle, cartilage and glands is detectable in early regenerates. On the contrary, the mesenchymal marker 22/31, whose expression in normal jaws is restricted to dermal fibroblasts and the dental pulp, is expressed in at least a half of the blastemal cells. The significance of these observations in relation to the origin of blastemal cells in the jaw will be discussed.
Animals↗
Phasic and tonic stretch reflexes in muscles with few muscle spindles: human jaw-opener muscles.
We investigated phasic and tonic stretch reflexes in human jaw-opener muscles, which have few, if any, muscle spindles. Jaw-unloading reflexes were recorded for both opener and closer muscles. Surface electromyographic (EMG) activity was obtained from left and right digastric and superficial masseter muscles, and jaw orientation and torques were recorded. Unloading of jaw-opener muscles elicited a short-latency decrease in EMG activity (averaging 20 ms) followed by a short-duration silent period in these muscles and sometimes a short burst of activity in their antagonists. Similar behavior in response to unloading was observed for spindle-rich jaw-closer muscles, although the latency of the silent period was statistically shorter than that observed for jaw-opener muscles (averaging 13 ms). Control studies suggest that the jaw-opener reflex was not due to inputs from either cutaneous or periodontal mechanoreceptors. In the unloading response of the jaw openers, the tonic level of EMG activity observed after transition to the new jaw orientation was monotonically related to the residual torque and orientation. This is consistent with the idea that the tonic stretch reflex might mediate the change in muscle activation. In addition, the values of the static net joint torque and jaw orientation after the dynamic phase of unloading were related by a monotonic function resembling the invariant characteristic recorded in human limb joints. The torque-angle characteristics associated with different initial jaw orientations were similar in shape but spatially shifted, consistent with the idea that voluntary changes in jaw orientation might be associated with a change in a single parameter, which might be identified as the threshold of the tonic stretch reflex. It is suggested that functionally significant phasic and tonic stretch reflexes might not be mediated exclusively by muscle spindle afferents. Thus, the hypothesis that central modifications in the threshold of the tonic stretch reflex underlie the control of movement may be applied to the jaw system.
Electromyography↗
Vacuous jaw movements induced by acute reserpine and low-dose apomorphine: possible model of parkinsonian tremor.
Coadministration of the monoamine-depleting agent reserpine with a low dose of apomorphine has been shown to result in high levels of vacuous jaw movements in rats. Two experiments were conducted to study the pharmacologic and motoric characteristics of the vacuous jaw movements induced by 5.0 mg/kg reserpine plus 0.1 mg/kg apomorphine. The first experiment was undertaken to determine whether the vacuous jaw movements induced by reserpine plus apomorphine could be reduced by coadministration of the muscarinic antagonist scopolamine. Injections of scopolamine produced a dose-related decrease in vacuous jaw movements induced by reserpine plus apomorphine, with the two highest doses (0.5 and 1.0 mg/kg scopolamine) producing significant differences relative to the control group that received reserpine plus apomorphine. In the second experiment, a slow-motion videotape system was used to study the temporal characteristics of the vacuous jaw movements induced by reserpine and apomorphine, and to study the effects of 1.0 mg/kg scopolamine on these movements. Most of the vacuous jaw movements shown by rats treated with reserpine and apomorphine occurred in rapid bursts of jaw movement. Analysis of the interresponse times (i.e., time between each jaw movement) showed that most of the jaw movements had a local frequency in the range of 2.86-6.67 Hz. Cotreatment with scopolamine significantly affected several measures of jaw movements. Thus, the vacuous jaw movements induced by reserpine plus apomorphine can be reversed by anticholinergic treatment, and these movements tend to occur as periodic oscillations of the lower jaw with a frequency of 3-7 Hz. Vacuous jaw movements in rats share some characteristics with parkinsonian symptoms and may represent an animal model of parkinsonian tremor.
Animals↗
Predicting masticatory jaw movements from chin movements using multivariate linear methods.
Previously, we have used bivariate correlations of maximum and minimum displacement, velocity and acceleration variables to compare masticatory chin and jaw movements (J. Prosthet. Dent. 81 (1999) 179). This previous study represented a first step in exploring the hypothesis that the chin contained useful information regarding jaw kinematics. The current study extends our understanding of the relationship between masticatory chin and jaw movements by: (1) reconstructing and evaluating a more continuous trajectory of chin and jaw movements, and (2) performing multivariate correlations comparing chin and jaw movements at discrete points along the trajectory in order to gain insight into the coupling of chin and jaw movements during a chewing cycle. Results indicated that chin and jaw movement trajectories were visually similar in the lateral, vertical, and anteroposterior axes. The adjusted R(2) results in the lateral, vertical, and anteroposterior dimensions averaged 0.74, 0.78, and 0.89, respectively. Within chewing cycles, the lowest correlations between chin and jaw movements in the lateral and vertical dimensions occurred when the jaw was relatively closed, whereas the lowest correlations between chin and jaw movements in the anteroposterior dimension occurred while the jaw was opening from a closed position. The results indicated that jaw and chin movements were qualitatively similar and that at least 74% of the variation in jaw movements could be accounted for by multivariate linear models of chin movement.
Adult↗
Dynamics of the human masticatory muscles during a jaw open-close movement.
The movements of the human jaw are controlled by the forces produced by the masticatory muscles. As the jaw moves, these muscles change in length and their force producing units, the sarcomeres, change in length simultaneously. The lengths and length changes of the sarcomeres are determinants for the forces they are able to produce. Hence, masticatory muscle force and jaw movement influence each other which makes it difficult to study their mutual relationship. In this paper, lengths and contraction velocities of the sarcomeres of the human jaw-opening and jaw-closing muscles are presented as well as the consequences for force production during jaw open-close movements simulated with a biomechanical model. Jaw-opening muscles acted almost synchronic in terms of sarcomere length, contraction velocity and force production. They were able to produce the largest isometric forces at relatively small jaw openings at the cost of reduced force production capabilities in wide open positions. In contrast, the jaw-closing muscles acted more differently. They were able to sustain active muscle force throughout a large range of the closing movement. Within this group the masseter and medial pterygoid contracted excentrically during a short time. The lateral pterygoid muscle portions behaved differently with respect to both groups. The jaw-opening muscles produced negligible passive forces during jaw closing. The passive forces of the jaw-closing muscles, however, contributed significantly to a limitation of the jaw-opening movement.
Humans↗
The jaw open-close movements predicted by biomechanical modelling.
The aim of this study was to analyse unloaded jaw-opening and jaw-closing movements in humans. For this purpose a dynamical 6-degree-of-freedom mathematical model of the human masticatory system was developed. It incorporated morphology, muscle architecture and dynamical muscle properties. Various symmetrical jaw-opening and jaw-closing movements were simulated based upon different muscle activation schemes. It was found that the balance between swing and slide of the mandibular condyle at the onset of a jaw-opening movement was predominantly dependent on the level of activation of the digastric and inferior lateral pterygoid muscles. The level of activation of the temporalis muscle parts was of critical importance for the jaw-closing movements. The amount of jaw opening was limited by the passive forces of the jaw-closing muscles. In contrast, the influence of the passive forces of the jaw-opening muscles on the jaw-closing movement was neglectable. Throughout the movements the temporomandibular joints remained loaded. The average torques generated by the jaw-opening or jaw-closing muscles with respect to the centre of gravity of the lower jaw had similar orientations and can be considered to be responsible for joint stabilization. The average direction of their lines of action, however, was about opposite, and this can be considered as the major discriminant between a movement in opening or closing direction.
Biomechanical Phenomena↗