Selective stimulation of human tooth-pulp with a new stable method: responses and validation.

INTRODUCTION: The aims of this study were to establish a safe technique for selective stimulation of nerves in human tooth-pulp during long experiments and to validate its use even with stimuli of high intensities. METHODS: A custom-made veneer containing 2 silver wire-conductive cream electrodes was attached with cement to the labial surface of an upper central incisor tooth. A variety of stimulus intensities were applied, and sensory and reflex responses from jaw-closing muscles were recorded. RESULTS: In 15 participants, the stimuli evoked predominantly sharp or painful sensations and reflex inhibitions of activity in the jaw muscles. Stimulation of 3 non-vital teeth evoked no sensations or reflexes, even at intensities that evoked maximal reflexes in vital teeth. The electrodes had reasonably stable resistances throughout experiments lasting up to 90 min. CONCLUSION: The method described enables responses to low- or high-intensity stimulation of human pulpal nerves to be investigated in long experiments.

Forces, movements and reflexes produced by pushing human teeth.

Pushing a tooth results in movement of the tooth and reflex inhibition of activity in jaw-closing muscles. The aims of this study were to determine how much tooth movement is required to elicit such reflexes and whether this is dependent on the point of force application to the tooth. Eight experiments were performed on six volunteer subjects. Electromyograms (EMGs) were recorded from a masseter muscle while the subjects produced approximately 12.5 % of the EMG associated with maximal clenching. Reflexes were evoked by pushing at two positions (incisal and cervical) on an upper central incisor. The forces applied and the resulting movements of the tooth were recorded. There was a linear relationship between force and movement regardless of whether the force was incisal or cervical (Pearson's r = 0.91 and r = 0.93 respectively). There were no differences between the slopes or intercepts for these relationships (ANCOVA p = 0.42, p = 0.46 respectively). There were linear relationships between the logarithms of force or movement and the resulting inhibitory reflexes (r = 0.81, 0.79, 0.81 and 0.74 for incisal and cervical forces and incisal and cervical movements, respectively). Again, there were no significant differences between the slopes for these relationships (ANCOVA p = 0.75, p = 0.46 for force and movement, respectively). There were no significant differences between the reflex thresholds for incisal and cervical stimuli in terms of force (0.23 and 0.25 N, ANCOVA p = 0.1) or movement (9.7 and 8.5 µm, ANCOVA p = 0.22). Thus, it appears that neither tooth movements nor jaw reflexes are dependent on the point of force application to a tooth.

Quantification of jaw reflexes evoked by natural tooth contact in human subjects.

Inhibitory jaw reflexes are believed to be important for protecting the teeth and temporo-mandibular structures from damage during sudden or forceful biting or mastication. Accordingly, alterations in these reflexes are sometimes implicated in aetiologies proposed for oro-facial pain syndromes, although the association is not well-established. We now aim to develop a method for quantifying objectively inhibitory jaw reflexes evoked by natural tooth contact. In the longer term, this may provide a new approach to examining the association of altered reflexes and clinical conditions. Eighteen subjects gave their written, informed consent, and were recruited to participate in this study. They were instructed to clench their teeth together in response to visual cues. They performed two such tasks twenty times: from the jaw postural position and from a more open position with the jaws set 10mm apart. Both tasks produced a rapid rise then stabilisation in electromyographic activity in the masseter muscle. This was always interrupted by a large inhibitory reflex starting 11.1±1.5 ms (mean±SD) after tooth contact. The inhibitions produced during the second task were similar but of significantly longer duration (24.3±6.4 vs 18.4±6.5 ms, P=0.0003, paired t-test) and greater magnitude (measured as an integral of the waveform: 1577±478 vs 1279±425%.ms, P=0.007, paired t-test). Interestingly, in a minority (13%) of the tasks, a second inhibition with a longer latency (50.9±0.9 ms) was also observed. Thus reflex responses in the masseter muscle to natural tooth contact usually consist of single inhibitory periods. In this respect they are like those induced by externally applied tooth pushing although occasionally there is a second inhibition, reminiscent of that seen with externally applied tooth taps.

The inhibitory effect of a chewing task on a human jaw reflex.

This study was undertaken to investigate whether an inhibitory jaw reflex could be modulated by experimentally controlled conditions that mimicked symptoms of temporomandibular disorders. Reflecting on previous work, we anticipated that these conditions might suppress the reflex. Electromyographic recordings were made from a masseter muscle in 18 subjects, while electrical stimuli were applied to the upper lip. An inhibitory reflex wave (mean latency 47 ms) was identified and quantified. Immediately following an accelerated chewing task, which in most cases produced muscle fatigue and/or pain, the size of the reflex wave decreased significantly by about 30%. The suppression of inhibitory jaw reflexes by fatigue and pain may result in positive feedback, which may contribute to the symptoms of temporomandibular disorders. Future studies of temporomandibular disorder sufferers will help to determine whether such reflex changes reflect the underlying etiology and/or are a result of the temporomandibular disorder itself.

Mastication and swallowing: 2. control.

UNLABELLED: In recent years, it has become clear that the neural mechanisms controlling chewing and swallowing emanate from centres in the brainstem. However, these activities may be modulated by conscious processes and by feedback from peripheral nerves.This review relates this knowledge to clinical dentistry and, in particular, to the possible relationship between craniomandibular dysfunctions and impaired control of the masticatory system. CLINICAL RELEVANCE: Dentists should understand the neuromuscular control of the masticatory apparatus as this can be relevant to the general health of patients as well as to craniomandibular disorders involving the jaw muscles and joints.

Mastication and swallowing: 1. Functions, performance and mechanisms.

UNLABELLED: The process of mastication involves movements of the tongue, lips and cheeks as well as the more obvious actions of the teeth and jaws. In recent years there have been significant advances in our knowledge of the relationships between these movements in human beings and of how the processes of mastication are related to the associated events of swallowing. In this, the first of two papers, we review the role of mastication in food processing and nutrition and the effect of tooth loss on masticatory performance.The paper also reviews new information on masticatory and swallowing functions in human beings eating naturally. The review relates this knowledge to clinical dentistry, notably to the relevance of a good dentition to the digestive process and practical considerations in the replacement of missing teeth. CLINICAL RELEVANCE: Dentists should understand the process of mastication because an adequate dentition can facilitate the general health and well-being of their patients.This understanding can also inform the clinical management of patients with a depleted dentition or otherwise impaired masticatory system.

Differences between inhibitory jaw reflexes evoked by stimulation of tooth pulp and across the adjacent alveolar process in man.

OBJECTIVE: In humans, stimulation of nerves in or around teeth can evoke inhibitory jaw reflexes. Previous studies had suggested that there may be subtle differences in the timings of the responses. The aim of the present study was to investigate this by comparing reflexes evoked by electrical stimulation of a tooth and of the adjacent tissues in individual subjects. DESIGN: Experiments were performed on 9 volunteers (3 male, 6 female). EMG recordings were made from the masseter muscle ipsilateral to the stimuli, whilst the subjects maintained a steady level of activity in the muscle. Reflexes were evoked by applying stimuli to an incisor tooth (pulpal stimuli) or across the adjacent alveolar process (transalveolar stimuli), using bipolar electrodes. RESULTS: Two inhibitory responses were evoked in most (8/9) subjects. The first occurred at a shorter latency after transalveolar than after pulpal stimulation (12.3+/-0.5 ms vs 19.4+/-1.5 ms; P=0.0014, paired t-test). For technical reasons, it was not possible to make such comparisons for the second inhibitory responses in all the subjects. In 5 subjects where such a comparison was possible, the mean latency of the transalveolar-evoked response was again shorter than that of the pulpal-evoked response (56.4+/-2.8 ms and 58.8+/-5.3 ms, respectively), but this difference was not significant (P=0.5). CONCLUSIONS: It appears that inhibitory jaw reflexes evoked from around the teeth are faster than those from the dental pulp. This observation could be due to differences between the peripheral afferent and/or the central pathways mediating the reflexes.

Modulation of an inhibitory jaw reflex by remote noxious stimulation: effects of spatial conditioning factors.

In humans, inhibitory jaw reflexes can be depressed by painful stimulation of remote parts of the body. The underlying mechanisms may involve diffuse noxious inhibitory controls (DNIC). Animal experiments have shown that the neurons which may mediate DNIC show spatial encoding (i.e. their responses vary in relation to the size of the body area being stimulated). The aim of this study was to investigate whether the modulation of an inhibitory jaw reflex shows similar spatial dependency. Electromyographic recordings were made in 9 subjects, from a masseter muscle that was activated to a level equivalent to 10% of that obtained during a maximum voluntary contraction. Reflex inhibitions were evoked by electrical stimuli to the upper lip, either alone (controls) or during the application of conditioning stimuli (47 degrees C water) to the fingers, the hand, the half forearm or the whole forearm. Conditioning stimuli applied to the larger but not to the smaller areas resulted in significant modulations of the reflex. There was a significant correlation between stimulus area and reflex magnitude. These results demonstrate a spatial dependency for the modulation of an inhibitory jaw reflex by painful stimuli -- a further parallel with DNIC as studied on single neurons in animals.

Dissociation of nociceptive modulation of a human jaw reflex from the influence of stress.

In human beings, inhibitory jaw reflexes can be depressed by painful stimulation of remote parts of the body. Since similar effects can be produced by the stress of anticipating pain, we wished to investigate whether the effects of remote painful stimuli are dependent on stress. EMG recordings were made from a masseter muscle while subjects maintained activity in the muscle at approximately 12.5% of maximum using visual feedback. The protocols involved three sequences: (1) "standard controls" in which reflexes were evoked by electrical test stimuli applied to the upper lip; (2) "standard conditioning" in which painful electrical conditioning stimuli were applied over the sural nerve 100 ms before the test stimuli; (3) "random sequences" in which test-only and conditioning-test combinations were employed in a double-blind, random, order. Data are presented as means +/- SEMs. In the standard controls, the stimuli evoked clear inhibitory reflexes (latency 37 +/- 1.3 ms, duration 62 +/- 5.6 ms; n = 10) in all the subjects. During standard conditioning, the reflex magnitude was reduced significantly (by 50.0 +/- 8.5%, P = 0.0002, one-sample t-test). When the test-only and conditioning-test responses were extracted from the random sequences, there was also a significant reduction in the reflex magnitude following conditioning (by 34.6 +/- 5.5%, P = 0.0002, one-sample t-test) albeit less so than between the standard sequences (P = 0.03, paired t-test). A second series of experiments suggested that these lesser effects during the random sequences were not substantially due to any loss of temporal summation of the conditioning mechanisms. The evidence for this was that application of pairs of conditioning stimuli did not produce a significantly greater effect than single conditioning stimuli within a random sequence (39.9 +/- 9.6% as opposed to 32.7 +/- 9.1% reductions in the reflex, P = 0.117, paired t-test). Therefore since any stress in the random sequences would not have been "tied" to the conditioned responses alone, the effects of remote painful stimuli on this inhibitory jaw reflex cannot be entirely secondary to stress.

Modulation of human jaw reflexes: heterotopic stimuli and stress.

The inhibitory reflexes in jaw elevator muscles, which are the predominant muscle responses to stimuli in or around the human mouth, are subject to modulation by nociceptive stimulation of remote parts of the body. The evidence for, and nature of, these modulatory effects are reviewed with particular emphasis on the reflex inhibition of masseteric activity evoked by electrical stimulation of the upper lip. This reflex is markedly reduced in magnitude by noxious stimulation of remote parts of the body surface or deeper tissues. Qualitatively similar effects on this reflex have been evoked by experimental stress and other psychological manipulations. However, recent research has eliminated the possibility that the modulatory effects of remote noxious stimuli act by inducing stress; it is more likely that they are a manifestation of the phenomena known as diffuse noxious inhibitory controls which act via inhibitory pathways originating in the medulla and ultimately producing post-synaptic inhibition of interneurones in the trigeminal nuclei.

Functional roles of oral reflexes in chewing and biting: phase-, task- and site-dependent reflex sensitivity.

Because loading during chewing is not totally predictable and jaw-closing muscles are strong and act over short distances, feedback from oral receptors is important in the control of mastication. Information on such feedback can be obtained by studying reflexes in jaw muscle EMGs. This review will deal with the contribution of reflex mechanisms to modifying motor neuron activity during chewing, and the dependency of reflex sensitivity on motor task, phase of movement, and site of stimulation.

Differentiation of nociceptive- from stress-induced modulatory influences on human reflexes.

This paper describes a new protocol that addresses the question of whether, in human experiments, modulatory effects of remote nociceptive conditioning stimuli on reflex responses are mediated by the stress induced by the conditioning stimuli. The protocol has been illustrated by a study into the effect of a remote nociceptive conditioning stimulus on an inhibitory jaw reflex. Electromyograms were recorded from an active masseter muscle and inhibitory reflexes were evoked by applying electrical stimuli to the upper lip. This protocol utilised the application of discrete electrical conditioning stimuli applied to the sural nerve prior to the test stimulus. A preliminary experiment determined that the optimal interval between the conditioning and test stimuli, which produced modulatory effects was 100 ms. In the definitive study, computer software was used to deliver control and conditioned sweeps in a double-blind randomised sequence. This resulted in a "stress-equal" protocol in which the level of stress would be the same for both control and conditioned sweeps. Therefore any observed modulatory effects on the reflexes could not have been wholly secondary to stress. This protocol could be adapted to the study of the modulation of other reflexes or evoked sensations by nociceptive conditioning stimuli.

Topical review: modulation of trigeminal sensory input in humans: mechanisms and clinical implications.

In this review, the modulatory effects of tooth and implant loading, orofacial pain, and psychological factors on somatosensory and jaw-motor function in humans are assessed. Experimental studies on the control of jaw actions have revealed that patients with prostheses supported by osseointegrated implants show an impairment of fine motor control of the mandible. One possibility is that this may be related to the loss of afferent information from periodontal ligament mechanoreceptors, which results in considerably higher and more variable forces to hold and manipulate food between the teeth. However, psychophysical investigations have shown that patients still perceive mechanical stimuli exerted on osseointegrated implants in the jawbone. The use of somatosensory evoked potentials may revealed what specific receptor groups are responsible for this so-called osseoperception phenomenon. Orofacial pain is another modulator of trigeminal system functioning. Experimental jaw muscle pain has several effects on the somatosensory and motor function of the masticatory system, all of them serving to warn the individual about the ongoing damaging of tissues. Finally, the influence of mental state on the sensory and motor functions of the trigeminal system will be addressed. While some animal studies suggest that psychological stress can reduce acute pain, less speculative are the findings in human subjects that the anticipation of receiving a painful stimulus or undertaking difficult mental tasks can modulate jaw reflexes, including those evoked by mechanical stimuli applied to the teeth. Since such stimuli occur regularly during normal oral activities, the study of the resulting motor effects may yield clinically meaningful results in the context of other variables that modulate mandibular function.

Remote noxious stimuli modulate jaw reflexes evoked by activation of periodontal ligament mechanoreceptors in man.

The purpose of the study was to investigate whether jaw reflexes evoked by selective stimulation of periodontal ligament me canoreceptors are susceptible to modulation by remote noxious stimulation. Experiments were performed on 10 volunteer subjects. Skin surface recordings were made from the jaw-closing masseter muscle. The subjects activated the muscle to approximately 10% of maximum by biting on a rubber impression of their molar teeth while they received visual feedback of the electromyogram (EMG) of the muscle. Reflexes were produced by the application of gentle mechanical stimuli to an upper central incisor tooth. The stimuli were in the form of 'ramp and hold' forces with a 5 ms rise-time and a 1.5 N plateau which lasted 350 ma. The resulting reflexes were recorded both under control conditions and while the subjects received a remote noxious stimulus (immersion of a hand in water at 3 degrees C). In all 10 subjects, the stimuli produced a single period of inhibition of masseteric activity (latency, 12.8 t 04 ms; duration, 18.1+/-1.3 ms; means +/- S.E.M.), which was usually followed by a period of increased masseteric activity. The period of inhibition constituted a downward wave in full-wave rectified, averaged signals. The integrals of such waves were significantly smaller (by 17+/- 6.5 %; P = 0.027; Student's t test) when the reflex was evoked during remote noxious stimulation rather than under control conditions. As such reflexes are beLieved to play a modulatory role during normal oral function this finding maybe relevant to disorders of mastication associated with pain.