Functional topography of the secondary somatosensory cortex for nonpainful and painful stimulation of median and tibial nerve: an fMRI study

Functional magnetic resonance imaging (fMRI) was used to study the cortical activity of the bilateral secondary somatosensory cortex (SII) during nonpainful (motor threshold) and painful electrical stimulation of median and tibial nerves. fMRI recordings were performed in eight normal young adults. The aim was at evaluating the working hypothesis of a spatial segregation of nonpainful and painful populations not only in the "hand" representation of SII [Ferretti, A., Babiloni, C., Del Gratta, C., Caulo, M., Tartaro, A., Bonomo, L., Rossini, P.M., Romani, G.L., 2003. Functional topography of the secondary somatosensory cortex for nonpainful and painful stimuli: an fMRI study. NeuroImage 20, 1625-1638.] but also in its "foot" representation. Results showed that, in both "hand" and "foot" representations of bilateral SII, the activity elicited by the painful stimulation was localized more posteriorly with respect to that elicited by the nonpainful stimulation. A fine spatial analysis of the SII responses revealed a clear somatotopic organization in the bilateral SII subregion especially reactive to the nonpainful stimuli (i.e., segregation of the hand and foot representations). In contrast, it was not possible to disentangle the "hand" and "foot" representations of SII for painful stimuli. These results extended to the SII "foot" representation previous evidence of a spatial segregation in the SII "hand" representation of subregions for the painful and nonpainful stimuli. Furthermore, they suggest that noxious information is not somatotopically represented in human bilateral SII, at least as inferred from fMRI data at 1.5 T.     

A new device for tactile stimulation during fMRI

Standardized somatosensory stimulation of the face during functional MRI is technically demanding due to the high magnetic field of the MRI scanner and the confined geometry of the head coil. We developed a new computer-controlled MR-compatible stimulation device for mapping somatosensory-evoked brain activations during fMRI. The device employs von Frey-filaments which are commonly used for quantitative sensory testing (QST) to deliver punctate tactile stimuli to the face and other body surfaces with a high spatiotemporal accuracy. Such stimuli were applied to the ipsilateral face and hand of eight volunteers during two different experimental designs to explore the feasibility of the new stimulator for somatosensory mapping. Tactile stimulation activated a distributed neural network including primary (S1) and secondary (S2) somatosensory areas as well as the premotor cortex and the thalamus. An event-related experimental design yielded S1 activation in all subjects despite a smaller total number of stimuli compared to a blocked design where S1 activation was not consistently found in three subjects. In individuals where S1 was significantly activated during both experimental conditions, the punctate tactile stimuli allowed discriminating the face and the hand representation in S1. We conclude that the novel stimulation device appears to be a valuable tool for mapping somatosensory representations. The data suggest that an event-related study design could be beneficial as it better controls for confounding factors such as anticipation, habituation and attention.     

Gender differences in patterns of cerebral activation during equal experience of painful laser stimulation.

A previous functional imaging study demonstrated greater female response in the anterior insula and thalamus and left prefrontal activation in men and right prefrontal activation in women during equal heat intensity but unequal pain experience. For the current study, subjective intensities of noxious heat delivered to the back of the right hand were equalized across subjects, and regional cerebral blood flow was recorded by using positron emission tomography. The female subjects required less laser energy before reporting pain, but the difference was not significant. Correlation of regional cerebral blood flow with subjective pain experience in the whole group showed significant bilateral responses in the parietal, lateral premotor, prefrontal, secondary somatosensory, anterior cingulate and insula cortices, as well as the thalamus. There was significantly greater activation in the left, contralateral, prefrontal, primary and secondary somatosensory, parietal, and insula cortices in the male subjects compared with the female subjects and greater response in the perigenual cingulate cortex in the female subjects. Our study is the first to associate consistent pain experience with gender differences in central response. These differences may relate to differential processing of acute pain with implications for clinical disorders that show a female dominance. The subtle behavioral differences and inconsistent findings across studies, however, suggest the need for caution and further experimentation before speculating further.     

Relationships between the paradoxical painful and nonpainful sensations induced by a thermal grill

The simultaneous application of innocuous cutaneous warm and cold stimuli with a thermal grill can induce both paradoxical pain and paradoxical warmth (heat). The goal of this study was to investigate further the relationships between these paradoxical sensations. Stimuli were applied to the palms of the right hands of 21 volunteers with a thermode consisting of 6 bars, the temperature of which was controlled by Peltier elements. We assessed the quality and intensity of the sensations evoked by series of stimuli consisting of progressively colder temperatures combined with a series of given warm temperatures. We applied a total of 116 series of stimuli, corresponding to 785 combinations of warm and cold temperatures. The 2 paradoxical phenomena were reported for most of the series of stimuli (n = 66). In each of these series, the 2 phenomena occurred in the same order: paradoxical warmth followed by paradoxical pain. The difference between the cold–warm temperatures eliciting paradoxical warmth was significantly smaller than that producing paradoxical pain. The intensities of the warmth and unpleasantness evoked by the stimuli were directly related to the magnitude of the warm–cold differential. Our results suggest that there is a continuum between the painful and nonpainful paradoxical sensations evoked by the thermal grill that may share pathophysiological mechanisms. These data also confirm the existence of strong relationships between the thermoreceptive and nociceptive systems and the utility of the thermal grill for investigating these relationships.     

Quantitative evaluation of hypnotically suggested hyperaesthesia and analgesia by painful laser stimulation

The ability to reduce both clinically and experimentally induced pain by hypnotic suggestion of analgesia is well known. However, the nature of hypnotic analgesia still remains uncertain. Attempts to demonstrate and identify specific psychophysiological mechanisms have, so far, been unsatisfactory. Methodological problems in inducing pain and monitoring physiological responses may be the reason for this lack of success. In the present study, we have attempted to eliminate some of these methodological problems. The sensory and pain thresholds to laser stimulation were determined, and the laser-evoked brain potentials were measured for 8 highly hypnotically susceptible subjects in 3 conditions: (1) waking state, (2) suggestion of hyperaesthesia, (3) suggestion of analgesia. The thresholds were reduced during induced hyperaesthesia and increased during analgesia. During hyperaesthesia sensations could be evoked by laser intensities which were below intensities that could be perceived in the awake state. The amplitude of the evoked brain potentials increased during hyperaesthesia and decreased during analgesia. The latency of the potential remained constant. The perception of pain during hypnosis can change very fast, indicating that slow endogenous mechanisms may play only a minor role in suggested hyperaesthesia/analgesia.     

Functional topography of the secondary somatosensory cortex for nonpainful and painful stimuli: an fMRI study

The regional activity of the contralateral primary (SI) and the bilateral secondary (SII) somatosensory areas during median nerve stimulations at five intensity levels (ranging from nonpainful motor threshold to moderate pain) was studied by means of functional magnetic resonance imaging (fMRI). The aim was to characterize the functional topography of SII compared to SI as a function of the stimulus intensity. Results showed that the galvanic stimulation of the median nerve activated the contralateral SI at all stimulus intensities. When considered as a single region, SII was more strongly activated in the contralateral than in the ipsilateral hemisphere. When a finer spatial analysis of the SII responses was performed, the activity for the painful stimulation was localized more posteriorly compared to that for the nonpainful stimulation. This is the first report on such a SII segregation for transient galvanic stimulations. The activity (relative signal intensity) of this posterior area increased with the increase of the stimulus intensity. These results suggest a spatial segregation of the neural populations that process signals conveyed by dorsal column-medial lemniscus (nonpainful signals) and neospinothalamic (painful signals) pathways. Further fMRI experiments should evaluate the functional properties of these two SII subregions during tasks involving sensorimotor integration, learning, and memory demands.     

Oscillatory interaction between human motor cortex and trunk muscles during isometric contraction.

We investigated oscillatory interaction between magnetoencephalographic signals of the human motor cortex and surface electromyogram from the paraspinal (PS) and abdominal (ABD) muscles. The results were compared with data obtained during contraction of the first dorsal interosseus (FDI) and tibialis anterior (TA) muscles. Significant coherence at 15-35 Hz was observed for both PS and ABD muscles in all subjects but the coherence was weak compared with that for FDI and TA. The cortical sources for both the PS and the ABD coherences were located in the motor cortex between the source areas for the FDI and TA coherences, thereby agreeing with the classical trunk area of the motor homunculus previously determined by invasive studies. The sources were strictly contralateral for PS but bilateral for ABD contractions. Our results indicate that during isometric contractions descending motor commands are modulated by cortical oscillations for both limb and trunk muscles, although the modulation is weaker and may be bilateral for trunk muscles.     

Suppression of motor evoked potentials in a hand muscle following prolonged painful stimulation.

Earlier investigations have shown that stimulation of peripheral afferent nerves induces prolonged changes in the excitability of the human motor cortex. The present study compared the effect of experimental pain and non-painful conditioning stimulation on motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS) in the relaxed first dorsal interosseous (FDI) and flexor carpi ulnaris (FCU) muscles. The MEPs were measured in 10 healthy subjects, and stimulus-response curves were generated before and after each of four stimulation paradigms conducted in random order on separate occasions: (a) control; (b) "dual stimulation" consisting of electrical stimulation of the FDI motor point paired with TMS; (c) painful infusion of hypertonic saline in the FDI muscle; and (d) pain combined with dual stimulation. There were no significant changes in FDI MEPs following the control paradigm, and dual stimulation induced an increase in the FDI MEPs only inconsistently. In contrast, the painful stimulation and the combined pain and dual stimulation paradigms were followed by significant suppression of the FDI MEPs at higher stimulus intensities. No changes were observed in the FCU MEPs following the four paradigms. In two additional subjects, the responses evoked in FDI by direct stimulation of the descending corticospinal tracts were significantly depressed following painful stimulation of the FDI, although the ulnar-evoked M-waves remained constant. It is concluded that muscle pain is followed by a period with profound depression of MEPs amplitudes in the resting muscle, but that these changes are at least in part due to a lasting depression of the excitability of the motoneurones in the spinal cord. Hence, painful stimulation differs from non-painful, repetitive stimulation, which facilitates the corticomotor pathway.     

Cardiocirculatory coupling during sinusoidal baroreceptor stimulation and fixed-frequency breathing

The question of whether respiratory sinus arrhythmia (RSA) originates mainly from a central coupling between respiration and heart rate, or from baroreflex mechanisms, is a subject of controversy. If there is a major contribution of baroreflexes to RSA, cardiocirculatory coupling during breathing and during cyclic baroreflex stimulation should show similarities. We applied a sinusoidal stimulus to the carotid baroreceptors and generated heart rate fluctuations of the same magnitude as RSA with a frequency similar to, but different from, the breathing frequency (0.2 Hz, compared with 0.25 Hz), and at 0.1 Hz, in 17 supine healthy subjects (age 28-39 years). The data were analysed using discrete Fourier-transform and transfer function analysis. Respiratory fluctuations in systolic blood pressure preceded RSA with a time lag equal to that between baroreceptor stimulation and oscillations in RR interval (0.62+/-0.18 s compared with 0.57+/-0.28 s at 0.2 Hz neck suction). The response of systolic blood pressure to neck suction at 0.2 Hz was 5 times less than the respiratory blood pressure fluctuations. Neck suction at 0.1 Hz largely increased fluctuations in blood pressure and RR interval, whereas the spontaneous phase relationship between blood pressure and RR interval remained unchanged. Our results are not consistent with the hypothesis that the origin of RSA is predominantly a central phenomenon which secondarily generates fluctuations in blood pressure, but suggest that, under the condition of fixed-frequency breathing at 0.25 Hz, baroreflex mechanisms contribute to respiratory fluctuations in RR interval.     

The cerebellum contributes to somatosensory cortical activity during self-produced tactile stimulation.

We used fMRI to examine neural responses when subjects experienced a tactile stimulus that was either self-produced or externally produced. The somatosensory cortex showed increased levels of activity when the stimulus was externally produced. In the cerebellum there was less activity associated with a movement that generated a tactile stimulus than with a movement that did not. This difference suggests that the cerebellum is involved in predicting the specific sensory consequences of movements and providing the signal that is used to attenuate the sensory response to self-generated stimulation. In this paper, we use regression analyses to test this hypothesis explicitly. Specifically, we predicted that activity in the cerebellum contributes to the decrease in somatosensory cortex activity during self-produced tactile stimulation. Evidence in favor of this hypothesis was obtained by demonstrating that activity in the thalamus and primary and secondary somatosensory cortices significantly regressed on activity in the cerebellum when tactile stimuli were self-produced but not when they were externally produced. This supports the proposal that the cerebellum is involved in predicting the sensory consequences of movements. In the present study, this prediction is accurate when tactile stimuli are self-produced relative to when they are externally produced, and is therefore used to attenuate the somatosensory response to the former type of tactile stimulation but not the latter.     

Laser evoked responses to painful stimulation persist during sleep and predict subsequent arousals

We studied behavioural responses and 32-channel brain potentials to nociceptive stimuli during all-night sleep in 12 healthy subjects, using sequences of thermal laser pulses delivered over the dorsum of the hand. Laser stimuli less than 20 dB over perception threshold had clear awakening properties, in accordance with the intrinsic threatening value of nociceptive signals. Even in cases where nociceptive stimulation did not interrupt sleep, it triggered motor responses in 11% of trials. Only four subjects reported dreams, and on morning questionnaires there was no evidence of incorporation to dreams of nociceptive stimuli. Contrary to previous reports suggesting the absence of cortical nociceptive responses during sleep, we were able to record brain-evoked potentials to laser (LEPs) during all sleep stages. Sleep LEPs were in general attenuated, but their morphology was sleep-stage-dependent: in stage 2, the weakened initial response was often followed by a high-amplitude negative wave with typical features of a K-complex. During paradoxical sleep (PS) LEP morphology was similar to that of waking, but frontal components showed strong attenuation, consistent with the reported frontal metabolic deactivation. A late positive component (450–650 ms) was recorded in both stage 2 and PS, the amplitude of which was significantly enhanced in trials that were followed by an arousal. This response appeared functionally related to the P3 wave, which in waking subjects has been associated to conscious perception and memory encoding.     

Modulation of an inhibitory reflex in single motor units in human masseter by tonic painful stimulation

Perioral electrical stimuli cause inhibitory reflex responses in single motor-units (SMU) and surface electromyographic (EMG) recordings from voluntary contracted human jaw-closing muscles. Tonic experimental masseter pain has recently been shown to reduce the inhibitory reflex response in surface EMG recordings but the effect on SMU activity has not been described. In this study, motor-unit action potentials were recorded with wire electrodes inserted into the left masseter in eleven subjects. The subjects kept the SMU firing rate around 10 Hz by feedback. Ninety-nine electrical stimuli were applied sequentially to the left mental nerve with increasing stimulus delays in steps of 1 ms after the preceding motor unit action potential. The inhibitory reflex in SMU was recorded before, during and after infusion of hypertonic saline (5%) into the ipsilateral masseter muscle. Spike train data were used to calculate (1) the mean pre- and post-stimulus inter-spike-intervals (ISI) in all of the 99 trials, (2) cumulative changes in firing probability, and (3) estimation of the compound inhibitory post-synaptic potential (IPSP) in the masseter motoneuron. Tonic masseter pain did not change pre-stimulus SMU firing characteristics but the mean ISI for the first post-stimulus discharge (158.2+/-9.2 ms) was significantly decreased compared to the pre-pain (175.8+/-11.3 ms, P<0.05) and post-pain conditions (172. 6+/-11.6 ms, P<0.05). The post-stimulus firing probability was significantly increased and the relative amplitude of the estimated IPSP significantly decreased during tonic masseter pain compared to pre-pain and post-pain conditions. In conclusion, this study indicates that tonic masseter pain has a net excitatory effect on the inhibitory jaw-reflexes, which could be mediated by presynaptic mechanisms on the involved motoneurons.     

The influence of simultaneous ratings on cortical BOLD effects during painful and non-painful stimulation

This fMRI study investigates the influence of a rating procedure on BOLD signals in common pain-activated cortical brain regions. Painful and non-painful mechanical impact stimuli were applied to the left hand of healthy volunteers. Subjects performed ratings of the perceived intensity during every second stimulation period by operating a visual analogue scale with the right hand. During every other stimulus period the subjects rested passively. Pain and touch stimuli were found to activate the same cortical areas previously defined as the "cortical pain matrix". General Linear Models were used to calculate contrasts between cortical activations during the "rating" and "non-rating" paradigm. In most brain regions activation following pain and touch was stronger during "rating" compared to "non-rating" conditions. Only the responses in the S1 projection field of the stimulated hand following pain were not influenced by the rating procedure. Furthermore, activations in the right and left posterior insular cortex and in the left superior frontal gyrus showed an opposite pattern, namely a stronger BOLD signal during "non-rating". We concluded: (1) Cortical areas regularly activated by painful stimuli may also be activated by touch stimulation. (2) Enhancement of the BOLD contrast by a rating procedure is probably an effect of closer stimulus evaluation and attention focussing. (3) In contrast to most other cortical regions, the posterior insular cortex, which is crucial for the integration of interoceptive afferent input, shows stronger responses in the absence of ratings, which points to a unique role of this region in the processing of somato-visceral information.     

Heat pain thresholds and cerebral event-related potentials following painful CO2 laser stimulation in chronic tension-type headache

Current opinion concerning the pathophysiology of tension-type headache (TTH) and its related pericranial muscle tenderness proposes a primary role of central sensitization at the level of dorsalhorn/trigeminal nucleus as well as the supraspinal level. Investigation of these phenomena can be conducted using laser-evoked potentials (LEPs), which are objective and quantitative neurophysiological tools for the assessment of pain perception. In the present study we examined features of LEPs, as well as cutaneous heat-pain thresholds to laser stimulation, in relation to the tenderness of pericranial muscles in chronic TTH resulting from pericranial muscle disorder, during a pain-free phase. Twelve patients with TTH and 11 healthy controls were examined using the Total Tenderness Scoring (TTS) system. The stimulus was a laser pulse generated by a CO(2) laser. The dorsum of the hand and the cutaneous zones corresponding to pericranial muscles were stimulated. Subjective perception of stimulus intensity was assessed by a visual analogue scale. Two responses, the earlier named N2a and the last named P2, were considered; the absolute latency was measured at the highest peak of each response. The N2a-P2 components' peak-to-peak amplitude was detected. The heat pain threshold was similar in TTH patients and controls at the level of both the hand and pericranial skin. The TTS scores at almost all pericranial sites were higher in TTH patients than in normal controls. The amplitude of the N2a-P2 complex elicited by stimulation of the pericranial zone was greater in TTH patients than in controls; the amplitude increase was significantly associated with the TTS score. Our findings suggest that pericranial tenderness may be a primary phenomenon that precedes headache, and is mediated by a greater pain-specific hypervigilance at the cortical level.     

BOLD effects in different areas of the cerebral cortex during painful mechanical stimulation

The contribution of four cortical areas (S1, S2, insular cortex and gyrus cinguli) to pain processing was assessed by functional magnetic resonance imaging (fMRI). Phasic (mechanical impact) and tonic stimuli (squeezing) were applied to the back of a finger, both at two different strengths. Stimuli were adjusted to inflict weak and strong pain sensations. It had been shown before that stronger noxious mechanical stimuli induce a weaker input from myelinated mechanoreceptors, but a more vigorous input from nociceptive primary afferents, and vice versa. Sizes of activation clusters and percent increase of the blood oxygenation level dependent (BOLD) signal during activation were compared in the areas of interest. Phasic stimulus patterns were more closely reflected in the time course of the MR signal in S1, S2 and the cingulate than tonic patterns, since the tonic stimuli tended to induce slow MR signal increase also during the resting periods which is in parallel to the persisting character of the tonic pain sensations. In S1 only the contralateral side was activated in most cases, and the more painful stimuli did not induce greater BOLD responses compared to the less painful stimuli in this area. Paradoxically, more painful stimuli produced smaller activation clusters in S1, both in tonic and phasic stimulus trials. In contralateral S2 more painful phasic stimuli induced significantly stronger BOLD responses than the weaker stimuli. The responses to tonic stimuli did not differentiate painfulness and were significantly smaller than the phasic. Activation clusters in this area were also smaller for tonic stimuli. In the gyrus cinguli more painful phasic stimuli induced stronger BOLD responses, but no difference was seen between tonic stimulation of different strength. Though the insular cortex was often bilaterally activated, no significant differences between stimulus quality or intensity were found. Our results provide evidence for a contribution of the S2 projection area and of the cingulate cortex to the processing of the intensity dimension of phasic mechanical pain. Such evidence was not found for the S1 area, which probably receives dominant input from non-nociceptive mechanoreceptors.     

Prospective diary study of nonpainful and painful phantom sensations in a preselected sample of child and adolescent amputees reporting phantom limbs

OBJECTIVE: To prospectively study factors associated with the occurrence of phantom sensations and pains in a pre-selected sample of child and adolescent amputees reporting phantom limbs. DESIGN: Prospective diary study over 1 month. PARTICIPANTS: Fourteen child and adolescent amputees from 10-18 years of age who were missing a limb due to trauma (n = 12) or congenital limb deficiency (n = 2), and who had previously reported having phantom sensations and pain. MAIN OUTCOME MEASURE: Diary used to assess the occurrence of non-painful and painful phantom sensations. Items included age, sex, location and cause of amputation, past experience with stump pain and pre-amputation pain, and intensity, quality, duration, and triggers of the sensations and pains. RESULTS: Thirteen amputees reported having 104 incidents of non-painful phantom sensations with an average intensity of 4.17 (SD = 2.14) on a 0-10 rating scale. Fifty-three incidents of phantom pain with an average intensity of 6.43 (SD = 1.76) were recorded by 8 amputees. Both amputees with a congenital limb deficiency reported phantom phenomena. Girls reported more psychosocial triggers than did boys whereas boys were more likely than girls to report that they could not identify a trigger (P = 0.0001). Boys also reported a higher proportion of physical triggers than psychosocial triggers while there were no differences for girls (P = 0.0001). DISCUSSION: Child and adolescent amputees experience phantom sensations and pains on a regular basis over a 1-month period. Differences in triggers of phantom phenomena between boys and girls may be due to differences in activities, awareness, attribution, and willingness to report psychosocial triggers.     

The effect of hypnotically induced emotional states on brain potentials evoked by painful argon laser stimulation.

The relationship between pain perception and emotional states is well known. However, the nature of this relationship and how different emotional states affect sensory and cognitive dimensions of pain remains uncertain. Results from experimental investigations are often contradictory, which may be due to methodological difficulties in inducing pain and monitoring physiological responses. In addition, most studies have focused on a single emotion, and data on the relative effects of different emotional states are lacking. In the present study we attempted to eliminate some of these methodological problems. Laser evoked potentials were used as a quantitative correlate to pain perception and were measured in 12 highly hypnotically susceptible subjects during seven conditions: (a) a prehypnotic baseline condition; (b) a neutral hypnotic control condition; (c-e) hypnotically recalled anger, fear, and depression in randomized order; (f) a hypnotically recalled happy condition, and (g) a posthypnotic awake control condition. The pain evoked potentials were significantly decreased in the angry condition and significantly increased in the depressed condition compared with baseline. No differences could be detected for either the happy or the fear-related condition compared with the baseline or neutral hypnotic condition. A significant positive correlation between the subjective intensity of depression and the increase in evoked potentials was found, but none for the other three emotions. The results support earlier findings that clinical depression is related to increased pain perception, and findings that the expression of anger can inhibit the experience of pain.(ABSTRACT TRUNCATED AT 250 WORDS)     

Transcranial direct current stimulation over the primary motor cortex during fMRI

Measurements of motor evoked potentials (MEPs) have shown that anodal and cathodal transcranial direct current stimulations (tDCS) have facilitatory or inhibitory effects on corticospinal excitability in the stimulated area of the primary motor cortex (M1). Here, we investigated the online effects of short periods of anodal and cathodal tDCS on human brain activity of healthy subjects and associated hemodynamics by concurrent blood-oxygenation-level-dependent (BOLD) functional magnetic resonance imaging (fMRI) at 3T. Using a block design, 20s periods of tDCS at 1 mA intensity over the left M1 altered with 20s periods without tDCS. In different fMRI runs, the effect of anodal or cathodal tDCS was assessed at rest or during finger tapping. A control experiment was also performed, in which the electrodes were placed over the left and right occipito-temporo-parietal junction. Neither anodal nor cathodal tDCS over the M1 for 20s stimulation duration induced a detectable BOLD signal change. However, in comparison to a voluntary finger tapping task without stimulation, anodal tDCS during finger tapping resulted in a decrease in the BOLD response in the supplementary motor area (SMA). Cathodal stimulation did not result in significant change in BOLD response in the SMA, however, a tendency toward decreased activity could be seen. In the control experiment neither cathodal nor anodal stimulation resulted in a significant change of BOLD signal during finger tapping in any brain area including SMA, PM, and M1. These findings demonstrate that the well-known polarity-dependent shifts in corticospinal excitability that have previously been demonstrated using measurements of MEPs after M1 stimulation are not paralleled by analogous changes in regional BOLD signal. This difference implies that the BOLD signal and measurements of MEPs probe diverse physiological mechanisms. The MEP amplitude reflects changes in transsynaptic excitability of large pyramidal neurons while the BOLD signal is a measure of net synaptic activity of all cortical neurons.     

Effects of intensity and locus of painful stimulation on postural stability

Stimulation of small diameter afferents can influence motor behavior. Little is known about how a prolonged painful stimulation of these small afferents may affect essential motor behavior such as the maintenance of an erect stance. The present study documents the effects of 10-s weak, moderate and extreme painful stimulations applied to the dorsum of the feet on the postural stability. Also, the moderate painful stimulation was applied to the metacarpal heads to determine if a painful stimulation to a limb not involved in the maintenance of the erect stance affects the postural control mechanisms. Increasing the intensity of the painful stimulation applied to the feet yielded larger postural oscillations whereas stimulation to the hands did not affect the control of posture. This suggests that the painful stimulation mainly affected the postural control mechanisms via sensorimotor processes rather than via cognitive resources related to the perception of pain.