Modulation of the periodontally evoked masseter reflexes by mechanical stimulation of the face

The current study was designed to determine if facial skin stimulation has a modulatory effect on the jaw reflexes that are elicited by tooth stimulation. This was investigated in eight human volunteers. The testing involved six sessions (three control and three test runs) of 50 identical tooth stimuli of 2 N, at 400 N/s, with 0.5 N preload, to the upper left central incisor. The stimulus typically induced an inhibitory reflex that was immediately followed by an excitatory reflex. All reflexes were recorded by surface electromyogram (SEMG) from the ipsilateral masseter. During the control runs, the tooth stimulus was delivered alone while the test runs involved mechanical stimulation of the skin as the tooth stimulus was repeated. The mechanical skin stimulation was achieved by rubbing a toothbrush with approximately 5 N of force, over the area of the face innervated by the maxillary division of the trigeminal nerve. The effect of skin stimulation on the periodontally induced reflex was measured by comparing the control and test reflex activity. When the SEMG of the test reflex was subtracted from the control reflex, the difference was a net increase in the SEMG in all eight subjects. There was also a significant reduction in the decline of the bite force. It is concluded that the skin stimulation can modulate the reflexes that are induced by tooth stimulus. It is postulated that this modulation may be partly responsible for matching the size and consistency of the food bolus with the appropriate bite force during mastication. Electronic Publication     

Endogenous adrenergic control of reflexes evoked by mechanical stimulation of the heel in the decerebrated rabbit

Our previous findings have indicated that the ventrolateral orbital cortex (VLO) may be involved in modulation of nociception and plays an important role as a higher center of an endogenous analgesic system (a feedback loop) consisting of spinal cord-nucleus submedius (Sm)-VLO-periaqueductal gray (PAG)-spinal cord. To further investigate the neurotransmitter mechanism involved in this nociceptive modulatory pathway, we tested the effects of microinjection of morphine (5 microg, 0.5 microl) into VLO on the tail flick (TF) reflex. The results show that a unilateral microinjection of morphine into VLO dose-dependently suppresses the TF reflex. Furthermore, 6 min after termination of morphine injection, microinjection of opioid receptor antagonist naloxone (1.5 microg, 0.5 microl) into the same VLO site reverses this morphine-evoked inhibition of TF reflex. These results suggest that morphine application to the VLO may directly or indirectly activate VLO neurons projecting to the PAG through the opioid receptor mediation leading to activation of the brainstem descending inhibitory system and depression of the nociceptive inputs at the spinal cord level.     

Origin of orientation-dependent asymmetries in vestibulo-ocular reflexes evoked by caloric stimulation

A caloric stimulus evokes primarily a horizontal vestibulo-ocular reflex (VOR) when subjects are in a supine or prone orientation with the horizontal semicircular canal plane oriented vertically. In both monkeys and humans, the magnitude of VOR eye movements is greater in the supine than in the prone orientation, indicating that some factor or factors, other than the conventionally accepted convective stimulation of the horizontal canals, contributes to the generation of the VOR. We used long-duration caloric irrigations and mathematical models of canal-otolith interactions to investigate factors contributing to the prone/supine asymmetry. Binaural caloric irrigations were applied for 7.5 or 9.5 min with subjects in a null orientation with horizontal canals in the earth-horizontal plane (control trial), or with the subject's pitch orientation periodically changing between null, supine, and prone positions with each orientation held for 30 s (caloric step trial). The control trial responses identified a small response attributable to a direct thermal effect on vestibular afferent activity that accounted for only 15% of the observed prone/supine asymmetry. We show that the gravito-inertial force resolution hypothesis for sensory integration of canal and otolith information predicts that the central processing of canal and otolith information produces an internal estimate of motion that includes both a rotational motion component and a linear acceleration component. These components evoke a horizontal angular VOR and linear VOR, which combine additively in the supine orientation, but subtract in the prone orientation, thus accounting for the majority of the observed prone/supine asymmetry.     

Early cortical activities evoked by noxious stimulation in humans

Lasers can selectively activate the nociceptors of A-delta fibers. Since nociceptors in the skin are activated via temperature conduction by the laser beam, a latency jittering of cortical responses among trials would affect results obtained with a conventional averaging (C-AVE) technique. We therefore used a new method, latency-adjusted averaging (L-AVE), to investigate cortical responses to noxious laser stimulation in normal subjects. L-AVE was done by averaging trials after adjusting the latency so that the peak latency of an activity in the temporal region of all trials matched on the time axis. Both in C-AVE and in L-AVE, clear activations were found in the contralateral primary somatosensory cortex (SI) and bilateral parasylvian regions, whose activities peaked 163–181 ms after the stimulation. In addition to these three main activities, weak activities peaking at around 109–119 ms could be identified in only L-AVE in similar cortical regions. Since the direction of the source differed between early and main activities, we considered that the early weak activities were cancelled out by the later main activities with an opposite orientation. The results suggested that early cortical processing of noxious information occurs earlier than previous neurophysiological studies have estimated and that the temporal sequence of activations should be reconsidered.     

Wiping reflexes and nerve impulse patterns evoked by electrical stimulation of the skin in frogs.

Electric shocks applied to the skin evoke well aimed wiping reflexes in the frogs, Discoglossus pictus and Rana esculenta. In skin-grafted animals, moreover, wipes were misdirected from those areas which gave misdirected responses to tactile stimulation. Recordings from cutaneous sensory neurons revealed that only one action potential, with varying latency, was evoked per shock in any given unit. It is concluded that the capacity for sensory localization in anurans cannot depend upon specific patterns of afferent nerve impulses.     

Jaw stretch reflexes in children

The substantial morphological transformations that occur during human development present the nervous system with a considerable challenge in terms of motor control. Variability of skilled motor performance is a hallmark of a developing system. In adults, the jaw stretch reflex contributes to the functional stability of the jaw. We have investigated the response properties of the jaw stretch reflex in two groups of young children and a group of young adults. Response latencies increased with development, and all age groups produced stimulus-magnitude-dependent increases in reflex gain and resulting biting force. Reflex gain was largest for the older children (9–10 years), yet net increases in resulting biting force were comparable across age groups. These data and earlier experiments suggest that oral sensorimotor pathways mature throughout childhood in concert with the continued acquisition of complex motor skills.     

Modulation of heat evoked nociceptive withdrawal reflexes by painful intramuscular conditioning stimulation

Convergence between cutaneous heat nociceptors and muscles afferents was investigated by applying a phasic, conditioning electrical stimulus to the tibialis anterior muscle (a train of five 1 ms pulses over 21 ms) at varying time intervals relative to a thermal test stimulus used for evoking the withdrawal reflex in humans. The 200 ms thermal stimulus was applied on the dorsum of the foot at an intensity of two times the pain threshold. The conditioning electrical stimulus was applied at an intensity of two times the pain threshold via a set of intramuscular needle electrodes. The conditioning-test interval was varied between –400 ms and 8,000 ms at 17 different intervals. The mean reflex onset latency of reflexes evoked by thermal stimuli alone was 354 ± 9 ms. A facilitation of the reflex was seen when the conditioning stimulus was applied 275 ms (174 ± 30% compared to control) and 300 ms (162 ± 32% compared to control) after the test stimulus onset indicating sensory convergence between muscle afferents (group I–III) and cutaneous Aδ heat nociceptors arriving simultaneously at the spinal cord.     

Excitatory and inhibitory electrodermal reflexes evoked by cutaneous stimulation in acute spinal cats

The electrodermal reflex (EDR) was recorded from the paws of acute spinal cats by a DC potential recording method. The EDR was produced either by pinching the skin in various areas of the trunk, or by electrical stimulation of cutaneous afferents of various spinal segments. It was found that not only excitatory but also inhibitory EDRs were produced and that occurrence of the excitatory and inhibitory EDRs depended on both the segmental position and the laterality of the stimulated area. Cutaneous stimulation generally produced excitatory EDRs more frequently than inhibitory ones, except when afferent inputs entered the spinal cord at or close to the segments of sudomotor outflow; in the latter case inhibitory EDRs were comparatively frequent and strong. Maximal inhibitory EDRs were elicited by stimulation of the group II afferent fibers of the cutaneous nerves only, whereas maximal excitatory EDRs were elicited after stimulation of the group II, III and IV afferent fibers of cutaneous origin.     

Opioidergic inhibition of reflexes evoked by selective stimulation of sural nerve C fibres in the rabbit.

Reflexes were evoked in the gastrocnemius medialis (GM) muscle nerve by selective electrical stimulation of the non-myelinated C fibres of the ipsilateral sural nerve of decerebrated, spinalized rabbits. The opioid antagonist (-)-quadazocine (555 micrograms/kg i.v.) enhanced responses to sural C fibre stimulation to an average of 236% of pre-drug levels. In addition, C fibre-evoked reflexes were depressed for 7-9 min after repetitive activation of the high threshold axons of the common peroneal nerve, and this effect was reversed after quadazocine. Thus, GM responses to stimulation of non-myelinated sural afferent fibres are suppressed by endogenous opioid peptides, but the degree of inhibition does not appear to be as profound as that previously reported for reflexes evoked by myelinated fibres.     

Recording of spinal reflexes evoked by dorsal root stimulation in unanesthetized free moving cats

Summary A simple technique is presented which allows electrical stimulation of dorsal roots in awake cats via electrodes fixed extradurally in vertebral arches. Spinal reflexes are recorded with bipolar electrodes from several muscles.Similar to results obtained in experiments in anesthetized cats, conditioning stimulation of the contralateral red nucleus enlarged reflexes mainly to flexor muscles. In accordance with the literature, arousal reactions were generally accompained by an increase in both flexor and extensor reflex amplitudes.Supported by the Deutsche Forschungsgemeinschaft (Br 242/3).     

Tuning the gain of somato-sympathetic reflexes by stimulation of the thoracic spine in humans

In animals, somatic stimulation of the limbs can evoke sympathetic reflexes of supraspinal origin. In addition, spinal reflexes can be elicited by stimulation of somatic tissues of the trunk. However, limited evidence is available concerning the specific modulation of sympathetic reflexes by afferents from the thoracic spine. This has also been largely overlooked in healthy humans. The aim of the present study was to determine whether tonic noxious heat (NH) applied to the skin over T3–T5 could segmentally increase supraspinal sympathetic reflexes (skin conductance responses – SCRs) induced by phasic electrical stimulation of the sural nerve. In addition, the effect of spinal manipulation (SM) on SCR amplitude and SCR amplification by NH was investigated. During the control session, palmar and plantar SCR amplitude was stable, showing no significant modulation. During NH and SM, however, palmar SCR amplitude was respectively increased and decreased in comparison to baseline, leading to a robust difference in SCR amplitude between the 2 conditions (p < 0.001). Moreover, these changes were also significantly and marginally different compared to the control session (p = 0.041 and p = 0.053, respectively). Interestingly, when applied immediately before NH, SM had a preventive effect on palmar SCR amplification induced by NH. In sharp contrast, changes in plantar SCRs were not significantly different between sessions (p = 0.42). Altogether, these results indicate that somatic stimulation of the thoracic spine may modulate somato-sympathetic reflexes segmentally in conscious, healthy volunteers.     

Tuning the gain of somato-sympathetic reflexes by stimulation of the thoracic spine in humans

An outbred rat model of novelty-seeking phenotype can differentiate between rats that show high rates (high responders; HRs) versus low rates (low responders; LRs) of locomotor reactivity to a novel environment. In the present study, LR and HR rats were exposed to a regimen of environmental and social stimuli (ESS) consisting of 14 random exposures of isolation, crowding or novel environment, once per day during the peripubertal-juvenile period (postnatal days 28-41) or handled as controls. Twenty-four hours after the last ESS exposure or control handling, all animals were tested on the forced swim and social interaction tests for depressive-like and social anxiety-like behaviors respectively. The ESS exposure during the peripubertal-juvenile period led to antidepressive-like effects on the forced swim test associated with increase in acetylation of histones 3 and 4 at the promoter regions P2 and P4 of the brain-derived neurotrophic factor (BDNF) gene in the dorsal hippocampus of HRs. Moreover, epigenetic activation of the hippocampal BDNF in the HRs following ESS exposure was accompanied by increase in the supra-pyramidal mossy fibre (SP-MF) and total mossy fibre terminal field volumes compared to handled controls. These findings suggest that the ESS exposure in the peripubertal-juvenile period may constitute an example of environmental induction of the hippocampal BDNF, and may mimic behavioral effects of exogenous antidepressants in the HR phenotype.     

Modulation of H reflexes in the forearm during voluntary teeth clenching in humans

We investigated whether there is any modulation of the H reflex in the forearm during teeth clenching and how any correlation that may be found is modulated. The H reflexes of the flexor carpi radialis (FCR) and the extensor carpi radialis (ECR) muscles were evoked on the right side in five healthy adult volunteers. The H reflexes of the FCR and ECR muscles were facilitated in association with voluntary teeth clenching in a force-dependent manner (r=0.46–0.663, P<0.05). The increase in amplitude of the H reflex of the FCR muscle associated with teeth clenching started before the onset of the EMG activity of the masseter muscle. The results of the present study demonstrate that oral motor activity exerts strong influences on the motor activity of the forearm.A part of this study was presented at the World Congress on Sports Dentistry and Dental Traumatology, 22 June 2001, Boston, Massachusetts, USA     

Eye movements evoked by proprioceptive stimulation along the body axis in humans

Proprioceptive input arising from torsional body movements elicits small reflexive eye movements. The functional relevance of these eye movements is still unknown so far. We evaluated their slow components as a function of stimulus frequency and velocity. The horizontal eye movements of seven adult subjects were recorded using an infrared device, while horizontal rotations were applied at three segmental levels of the body [i.e., between head and shoulders (neck stimulus), shoulders and pelvis (trunk stimulus), and pelvis and feet (leg stimulus)]. The following results were obtained: (1) Sinusoidal leg stimulation evoked an eye response with the slow component in the direction of the movement of the feet, while the response to trunk and neck stimulation was oriented in the opposite direction (i.e., in that of the head). (2) In contrast, the gain behavior of all three responses was similar, with very low gain at mid- to high frequencies (tested up to 0.4 Hz) but increasing gain at low frequencies (down to 0.0125 Hz). We show that this gain behavior is mainly due to a gain nonlinearity for low angular velocities. (3) The responses were compatible with linear summation when an interaction series was tested in which the leg stimulus was combined with a vestibular stimulus. (4) There was good correspondence of the median gain curves when eye responses were compared with psychophysical responses (perceived body rotation in space; additionally recorded in the interaction series). However, correlation of gain values on a single-trial basis was poor. (5) During transient neck stimulation (smoothed position ramp), the neck response noticeably consisted of two components – an initial head-directed eye shift (phasic component) followed by a shift in the opposite direction (compensatory tonic component). Both leg and neck responses can be described by one simple, dynamic model. In the model the proprioceptive input is fed into the gaze network via two pathways which differ in their dynamics and directional sign. The model simulates either leg or neck responses by selecting an appropriate weight for the gain of one of the pathways (phasic component). The interaction results can also be simulated when a vestibular path is added. This model has similarities to one we recently proposed for human self-motion perception and postural control. A major difference, though, is that the proprioceptive input to the gaze-stabilizing network is weak (restricted to low velocities), unlike that used for perception and postural control. We hold that the former undergoes involution during ontogenesis, as subjects depend on the functionally more appropriate vestibulo-ocular reflex. Yet, the weak proprioceptive eye responses that remain may have some functional relevance. Their tonic component tends to stabilize the eyes by slowly shifting them toward the primary head position relative to the body support. This applies solely to the earth-horizontal plane in which the vestibular signal has no static sensitivity. Received: 10 October 1997 / Accepted: 22 January 1998     

Facilitation and inhibition of jaw reflexes evoked by electrical stimulation of the cat's cerebral cortex

Summary The effects of electrical stimulation of the cerebral cortex on the monosynaptic jaw closing and the disynaptic jaw opening reflexes were studied in cats anaesthetized with chloralose. The time course of the reflex effects was recorded. Similar rhythmic sequences of facilitation and inhibition were observed in both reflexes (Fig. 3). The sequence could start with facilitation or inhibition. The latency of the initial effects was short (2.5 ms) indicating a minimum of two synapses in the descending path. The period of the rhythmic sequence was approximately 10 ms.Optimal parameters for the conditioning cortical stimuli were found to be: trains of 3–5 surface anodal pulses, 0.5 ms, 400 Hz. The threshold of the cortical effects on the reflexes was 0.3 mA. A single pulse evoked effects.The cortical origin of the effects was located and related to the somatosensory projections, and to the cytoarchitecture. The effects of largest amplitude and most complex time course were evoked from the oral and perioral projections to areas 3a and 3b. Effects evoked from areas 4, 5a, and 6a were less complex and of lower amplitude.It is suggested that a trigemino-cortico-trigeminal loop via 3a may function in reflex modulation of the jaw movements. In addition area 3a may contribute to cortico-cortical motor elaborations via U-fiber connections to area 4.This work was supported by the Swedish Medical Research Council (Proj B80-14X-00045-16A)     

Human interlimb reflexes evoked by electrical stimulation of cutaneous nerves innervating the hand and foot

There is some discrepancy over the extent to which reflex pathways from different cutaneous nerves in the hand and foot link the cervical and lumbar spinal cord in neurologically intact humans. The present experiments were designed to determine whether stimulation of a cutaneous nerve in the foot or in the hand evoked reflexes in the non-stimulated limbs (interlimb reflexes). Reflexes were elicited by stimulating (5x1-ms pulses at 300 Hz) the superficial peroneal (SP; innervates the foot dorsum) or superficial radial (SR; innervates the dorsolateral portion of the hand) nerve while subjects (n=10) performed focused contractions of different upper and lower limb muscles. Reflex responses were divided into early (<75 ms), middle (75-120 ms), and late (>120 ms) epochs as determined from averages of 50 sweeps of stimulus-locked electromyographic activity. Significant interlimb reflexes were found at the early latency in 44/106 and 44/103 muscles sampled after SP and SR nerve stimulation, respectively. At the middle latency, significant interlimb reflexes were seen in 89/106 and 87/103 muscles sampled after SP and SR nerve stimulation, respectively. Interlimb reflexes were seen when stimulating at the wrist (i.e. SR nerve) and when stimulating at the ankle (i.e. SP nerve) with an equal probability. The results show that interlimb cutaneous reflexes are widely distributed in humans. The mean latency of the earliest response was quite short and may be mediated by a propriospinal pathway. Functionally, these pathways may provide a substrate for transferring information to coordinate movements between the limb segments.     

Capsaicin differentially influences somatosensory cortical responses evoked by peripheral electrical or mechanical stimulation.

The effects of capsaicin injected intraperitoneally (200 micrograms/kg) or applied locally to the cortical surface (10(-5) M) were studied on cortical potentials evoked by peripheral electrical or mechanical stimulation. Capsaicin treatment (i.p.) differentially influenced the cortical evoked potentials depending on the type of stimulation. Just after both types of capsaicin application, the responses to both kinds of stimuli decreased in amplitude. Additionally, during this time a short fall in blood pressure was observed. Half an hour later, however, only in the case of interperitoneal application the potentials evoked by electrical stimulation were facilitated, while the potentials evoked by vibrissa deflection had recovered and stayed around the control levels thereafter. In addition, the responsive cortex area activated by electrical stimulation became enlarged after the i.p. injection of capsaicin, while that of the cortex region activated by mechanical stimulation did not change significantly. Capsaicin applied locally to the cortex resulted neither in the facilitation of evoked potentials nor in the enlargement of the responsive cortical area. The present findings are the first to demonstrate that the i.p. (but not local) administration of capsaicin, in low dosage, differentially influences the cortical responses evoked by electrical and mechanical stimulation of somatosensory afferents.