[Show abstract][Hide abstract] ABSTRACT: The mechanisms whereby orofacial pain affects motor function are poorly understood. The aims were to determine whether 1) lingual algesic chemical stimulation affected face primary motor cerebral cortex (face MI) excitability defined by intracortical microstimulation (ICMS); and 2) any such effects were limited to the motor efferent MI zones driving muscles in the vicinity of the noxious stimulus. Ketamine-anesthetized Sprague-Dawley male rats were implanted with electromyographic (EMG) electrodes into anterior digastric, masseter, and genioglossus muscles. In 38 rats, three microelectrodes were located in left face MI at ICMS-defined sites for evoking digastric and/or genioglossus responses. ICMS thresholds for evoking EMG activity from each site were determined every 15 min for 1 h, then the right anterior tongue was infused (20 microl, 120 microl/h) with glutamate (1.0 M, n = 18) or isotonic saline (n = 7). Subsequently, ICMS thresholds were determined every 15 min for 4 h. In intact control rats (n = 13), ICMS thresholds were recorded over 5 h. Only left and right genioglossus ICMS thresholds were significantly increased (< or =350%) in the glutamate infusion group compared with intact and isotonic saline groups (P < 0.05). These dramatic effects of glutamate on ICMS-evoked genioglossus activity contrast with its weak effects only on right genioglossus activity evoked from the internal capsule or hypoglossal nucleus. This is the first documentation that intraoral noxious stimulation results in prolonged neuroplastic changes manifested as a decrease in face MI excitability. These changes appear to occur predominantly in those parts of face MI that provide motor output to the orofacial region receiving the noxious stimulation.
Journal of Neurophysiology 07/2008; 100(3):1234-44. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Since there is limited information of the effects of orofacial pain on oromotor behaviour, the aim of this study was to test the effects of injection of the algesic chemical glutamate into orofacial tissues on licking behaviour in rats.
Male Sprague-Dawley rats were trained to carry out stereotyped licking from a water spout connected to a force transducer, and once they met preset licking motor requirements (peak force of licking of 2-20 g, interlick interval of 0.11-0.19s), their masseter muscle or tongue was injected with either isotonic saline (0.9%, 5 microL) or glutamate (1.0M, 5 microL) and several parameters of their licking performance were re-assessed.
There were significant effects (P<0.05; 1-way repeated measures ANOVA) of glutamate injection into the tongue on the number of clusters of licks, the number of licks per cluster, the intercluster period, the peak force of licking and the interlick interval, but there were no significant (P>0.05) effects on licking of isotonic saline injection into the tongue or isotonic saline or glutamate injection into the masseter muscle.
These findings provide the first documentation that noxious stimulation of the tongue, but not of the masseter muscle, has a modulatory effect on licking behaviour in the rat and suggest that the neural substrate for licking may be sensitive to selective nociceptive inputs from the orofacial region.
Archives of Oral Biology 04/2008; 53(4):361-8. · 1.55 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. The lateral pericentral region of the cerebral cortex has been well documented in primates to be important in sensorimotor integration and control and in the learning of new motor skills. 2. The present article provides, first, an overview of limb sensorimotor cortical mechanisms and, second, outlines recent evidence pointing to an important role for the face sensorimotor cortex in semi-automatic, as well as trained, orofacial motor behaviour and to its propensity for neuroplastic changes in association with orofacial motor skill acquisition or an altered oral environment.
Clinical and Experimental Pharmacology and Physiology 02/2005; 32(1-2):109-14. · 2.41 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The present study was undertaken to determine the firing patterns and the mechanoreceptive field (RF) properties of neurons within the face primary motor cortex (face-MI) in relation to chewing and other orofacial movements in the awake monkey. Of a total of 107 face-MI neurons recorded, 73 of 74 tested had activity related to chewing and 47 of 66 neurons tested showed activity related to a trained tongue task. Of the 73 chewing-related neurons, 52 (71.2%) showed clear rhythmic activity during rhythmic chewing. A total of 32 (43.8%) also showed significant alterations in activity in relation to the swallowing of a solid food (apple) bolus. Many of the chewing-related neurons (81.8% of 55 tested) had an orofacial RF, which for most was on the tongue dorsum. Tongue protrusion was evoked by intracortical microstimulation (ICMS) at most (63.6%) of the recording sites where neurons fired during the rhythmic jaw-opening phase, whereas tongue retraction was evoked by ICMS at most (66.7%) sites at which the neurons firing during the rhythmic jaw-closing phase were recorded. Of the 47 task-related neurons, 21 of 22 (95.5%) examined also showed chewing-related activity and 29 (61.7%) demonstrated significant alteration in activity in relation to the swallowing of a juice reward. There were no significant differences in the peak firing frequency among neuronal activities related to chewing, swallowing, or the task. These findings provide further evidence that face-MI may play an important role not only in trained orofacial movements but also in chewing as well as swallowing, including the control of tongue and jaw movements that occur during the masticatory sequence.
Journal of Neurophysiology 06/2002; 87(5):2531-41. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Our previous studies have revealed that face primary somatosensory cortex (SI) as well as face primary motor cortex (MI) play important roles in the control of orofacial movements in awake monkeys, and that both face MI and face SI neurons may have an orofacial mechanoreceptive field and show activity related to orofacial movements. Since it is possible that the movement-related activity of face MI neurons could reflect movement-generated orofacial afferent inputs projecting to face MI via face SI, the present study used reversible cold block-induced inactivation of the monkey's face SI to determine if face MI neuronal activity related to a trained tongue-protrusion task, chewing or swallowing was dependent on the functional integrity of the ipsilateral face SI and if inactivation of face SI affects orofacial movements. The effects of face SI cold block were tested on chewing, swallowing and/or task-related activity of 73 face MI neurons. Both task and chewing and/or swallowing-related activity of most face MI neurons was independent of the functional integrity of the ipsilateral face SI since SI cold block affected the movement-related activity in approximately 25% of the neurons. Similarly, unilateral cold block of SI had very limited effects on the performance of the task and chewing, and no effect on the performance of swallowing. These findings suggest that movement-induced reafferentation via face SI may not be a significant factor in accounting for the activity of the majority of ipsilateral face MI neurons related to trained movements, chewing and swallowing.
Somatosensory and Motor Research 02/2002; 19(4):261-71. · 0.93 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Although the cerebral cortex has been implicated in the control of swallowing, the output organization of the cortical swallowing representation, and features of cortically evoked swallowing, remain unclear. The present study defined the output features of the primate "cortical swallowing representation" with intracortical microstimulation (ICMS) applied within the lateral sensorimotor cortex. In four hemispheres of two awake monkeys, microelectrode penetrations were made at </=1-mm intervals, initially within the face primary motor cortex (face-MI), and subsequently within the cortical regions immediately rostral, lateral, and caudal to MI. Two ICMS pulse trains [35-ms train, 0.2-ms pulses at 333 Hz, </=30 microA (short train stimulus, T/S); 3- to 4-s train, 0.2-ms pulses at 50 Hz, </=60 microA (continuous stimulus, C/S)] were applied at </=500-micron intervals along each microelectrode penetration to a depth of 8-10 mm, and electromyographic (EMG) activity was recorded simultaneously from various orofacial and laryngeal muscles. Evoked orofacial movements, including swallowing, were verified by EMG analysis, and T/S and C/S movement thresholds were determined. Effects of varying ICMS intensity on swallow-related EMG properties were examined by applying suprathreshold C/S at selected intracortical sites. EMG patterns of swallows evoked from various cortical regions were compared with those of natural swallows recorded as the monkeys swallowed liquid and solid material. Results indicated that swallowing was evoked by C/S at approximately 20% of 1,569 intracortical sites where ICMS elicited an orofacial motor response in both hemispheres of the two monkeys, typically at C/S intensities </=30 microA. In contrast, swallowing was not evoked by T/S in either monkey. Swallowing was evoked from four cortical regions: the ICMS-defined face-MI, the face primary somatosensory cortex (face-SI), the region lateral and anterior to face-MI corresponding to the cortical masticatory area (CMA), and an area >5 mm deep to the cortical surface corresponding to both the white matter underlying the CMA and the frontal operculum; EMG patterns of swallows elicited from these four cortical regions showed some statistically significant differences. Whereas swallowing ONLY was evoked at some sites, particularly within the deep cortical area, swallowing was more frequently evoked together with other orofacial responses including rhythmic jaw movements. Increasing ICMS intensity increased the magnitude, and decreased the latency, of the swallow-related EMG burst in the genioglossus muscle at some sites. These findings suggest that a number of distinct cortical foci may participate in the initiation and modulation of the swallowing synergy as well as in integrating the swallow within the masticatory sequence.
Journal of Neurophysiology 09/1999; 82(3):1529-41. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Rhythmical jaw movements can be evoked by intracortical microstimulation within four physiologically defined regions, one of which is the primary face somatosensory cortex (face SI). It has been proposed that these regions may be involved in the selection and/or control of masticatory patterns generated at the brainstem level. The aim here was to determine if mastication is affected by reversible, cooling-induced inactivation of the face SI. Two cranial chambers were chronically implanted in two monkeys (Macaca fascicularis) to allow access bilaterally to the face SI. A thermode was placed on the dura or pia overlying each SI that had been shown with micro-electrode recordings to receive intraoral inputs. A hot or cold alcohol-water solution was pumped through the thermodes while the monkey chewed a small piece of apple or a sultana during precool (thermode temperature, 37 degree C), cool (2-4 degrees C), and postcool (37 degrees C) conditions. Electromyographic (EMG) activity was recorded intramuscularly from the masseter, genioglossus, and anterior digastric. Cooling of SI impaired rhythmical jaw and tongue movements and EMG activity associated with mastication in one monkey (H5), and modified the pattern of EMG activity in the other (H6). The total masticatory time (i.e., time taken for chewing and manipulation of the bolus before swallowing) was increased. This was due principally to an increase in the oral transport time (i.e., time taken for manipulation of bolus after chewing and before swallowing: monkey H6, control, 2.7 sec; cool, 5.2 sec, p < 0.05); the bolus was manipulated by the tongue during this period before swallowing. Within the chewing time (i.e., time during which chewing occurred), cooling resulted in a significant increase in anterior digastric muscle duration, a significant delay in the onset of masseter EMG activity, and a significant increase in the variance of genioglossus EMG duration. The data support the view that the face SI plays a part in modulating the central pattern generator for mastication.
Archives of Oral Biology 03/1998; 43(2):133-41. · 1.55 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Recent studies conducted in our laboratory have suggested that the tongue primary motor cortex (i.e., tongue-MI) plays a critical role in the control of voluntary tongue movements in the primate. However, the possible involvement of tongue-MI in semiautomatic tongue movements, such as those in swallowing, remains unknown. Therefore the present study was undertaken in attempts to address whether tongue-MI plays a role in the semiautomatic tongue movements produced during swallowing. Extracellular single neuron recordings were obtained from tongue-MI, defined by intracortical microstimulation (ICMS), in two awake monkeys as they performed three types of swallowing (swallowing of a juice reward after successful tongue task performance, nontask-related swallowing of a liquid bolus, and nontask-related swallowing of a solid bolus) as well as a trained tongue-protrusion task. Electromyographic activity was recorded simultaneously from various orofacial and laryngeal muscles. In addition, the afferent input to each tongue-MI neuron and ICMS-evoked motor output characteristics at each neuronal recording site were determined. Neurons were considered to show swallow and/or tongue-protrusion task-related activity if a statistically significant difference in firing rate was seen in association with these behaviors compared with that observed during a control pretrial period. Of a total of 80 neurons recorded along 40 microelectrode penetrations in the ICMS-defined tongue-MI, 69% showed significant alterations of activity in relation to the swallowing of a juice reward, whereas 66% exhibited significant modulations of firing in association with performance of the trained tongue-protrusion task. Moreover, 48% showed significant alterations of firing in relation to both swallowing and the tongue-protrusion task. These findings suggest that the region of cortex involved in swallowing includes MI and that tongue-MI may play a role in the regulation of semiautomatic tongue movement, in addition to trained motor behavior. Swallow-related tongue-MI neurons exhibited a variety of swallow-related activity patterns and were distributed throughout the ICMS-defined tongue-MI at sites where ICMS evoked a variety of types of tongue movements. These findings are consistent with the view that multiple efferent zones for the production of tongue movements are activated in swallowing. Many swallow-related tongue-MI neurons had an orofacial mechanoreceptive field, particularly on the tongue dorsum, supporting the view that afferent inputs may be involved in the regulation of the swallowing synergy.
Journal of Neurophysiology 10/1997; 78(3):1516-30. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: This study sought to characterise the electromyographic (EMG) activity patterns of orofacial-muscles during trained tongue-protrusion and biting tasks in two awake monkeys (Macaca fascicularis). Chronic or acute EMG electrodes were placed in the anterior digastric (DIG), genioglossus (GG), masseter (MASS), platysma (PLAT), zygomaticus major (ZYGO), orbicularis oris superior (OOS), and orbicularis oris inferior (OOI) muscles and their EMG activity as well as the force signals of the tongue-protrusion and biting tasks were recorded. A total of 327 tongue-protrusion task trials and a total of 210 biting-task trials were successfully completed in several recording sessions and the EMG patterns were generally consistent between the different sessions. For the tongue task, the mean onset time of increase in GG activity significantly (p < 0.0001) led the mean onset time of increase in the force. The DIG, GG, and OOI (and also the OOS in one of the monkeys) showed a significant (p < 0.0001) increase in mean EMG amplitude during the holding phase, but the GG in both monkeys had the highest mean EMG amplitude ratio (MAR), i.e. the mean EMG amplitude during the holding or dynamic phase divided by the mean EMG amplitude during the pre-trial period. A similar EMG pattern was documented for different directions of the tongue-protrusion task (right, symmetrical, and left) and changes in the levels of EMG activities occurred in GG and OOI as the direction of the tongue-protrusion task changed from left to right. The task at different forces was associated with no apparent change in MAR for the holding phase for each muscle recorded. However, during the dynamic phase, only the GG showed a significant increase in EMG activity as the forces were increased. For the biting task, the mean onset times of the MASS activity and force were not significantly different. The MASS and ZYGO muscles (and the PLAT in one of the monkeys) showed a significant increase in mean EMG amplitude during the holding phase compared with the pre-trial period, and the MASS showed the highest MAR. It was also the only muscle showing a significant increase in the EMG activity when the bite-force level was increased. These findings reveal that certain orofacial muscles are selectively recruited during the two different orofacial tasks.(ABSTRACT TRUNCATED AT 400 WORDS)
Archives of Oral Biology 11/1994; 39(11):955-65. · 1.55 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. In previous papers we have presented evidence suggesting an important role for the face primary somatosensory cortex (SI) in the fine control of tongue movements. These findings, plus our earlier evidence that many neurons in face motor cortex (MI) may exhibit firing rates related to the direction of tongue protrusion, led us to test the hypothesis that variations in the direction of a tongue-protrusion movement would be associated with variations in the activity of different face SI neurons. 2. Two monkeys were trained to perform a tongue-protrusion task in each of three directions: the task transducer was positioned at 0 degrees, 30 degrees to the left, or 30 degrees to the right from the midsagittal plane. The latter two positions were termed asymmetrical tongue-protrusion task positions. Single-neuron activity was recorded from face SI during trials of the tongue-protrusion task at each of two or three of the above positions. In addition, the mechanoreceptive field (RF) was delineated for each neuron. 3. Directional relations were found in 25 (58%) of the 43 neurons studied; this included 20 neurons showing a significant direction-by-time interaction in firing rate, i.e., the change of firing rate from the pretrial period to the task period was significantly different between different directions, and 5 showing no direction-by-time interaction but a significant difference in firing rate between different directions of the tongue-protrusion task. 4. Of the 43 neurons investigated, 21 and 20 had a RF on the tongue and lips, respectively ("tongue RF" and "lip RF" neurons), and the remaining 2 received mechanosensitive afferent inputs from other orofacial regions. There was no significant difference in the incidence of directional sensitivity between the neurons with a tongue RF and those with a lip RF (12/21 and 11/20, respectively). 5. Eight of the 25 "directional" neurons were located in area 3b and 17 in area 1. There was no significant difference in the proportion of directional neurons between areas 3b and 1. 6. The increase in discharge frequency at the preferred direction was, on the average for the 25 directional neurons, 39% over the mean discharge frequency observed during the task period for all directions of the tongue-protrusion task. Eight directional neurons showed a significant increase in firing rate during the tongue-protrusion task up to 130 ms before the onset of genioglossus electromyographic activity.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 07/1994; 71(6):2391-400. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. We have demonstrated recently that reversible, cooling-induced inactivation of the face primary somatosensory cortex (SI) severely impairs the successful performance of a tongue-protrusion task but has relatively minor effects on the performance of a biting task. In an attempt to establish a neuronal correlate for these different behavioral relations, the present study was initiated to document the mechanoreceptive field properties of a population of face SI neurons and their activity during the tongue-protrusion and biting tasks. 2. Within SI, the representation of the face was found immediately lateral to that of the hand, and there was a clear somatotopic pattern of organization within face SI: the periorbital or nose region was located most medially in the face SI, then followed laterally in sequence the representation of the upper lip, lower lip, and intraoral area. A mechanoreceptive field (RF) was identified for 253 neurons, which included 162 "lip RF" neurons receiving mechanosensitive afferent inputs from the upper lip, lower lip, or both; 72 "tongue RF" neurons that received mechanosensitive afferent inputs from the tongue; 11 "periodontium RF" neurons receiving periodontal inputs; and 8 neurons that received inputs from other orofacial regions. 3. Nearly all (249/253) of the face SI neurons responded to light tactile stimuli, and most of them received contralateral inputs (78%) and showed a rapidly adapting (RA) response to tactile stimulation (82%). There was no significant difference in the ratio of slowly adapting (SA) to RA neurons in areas 3b and 1. 4. For 193 neurons studied in one or both of the orofacial tasks, 113 were found, on the basis of histological reconstruction, to be distributed in area 1, 61 in area 3b, and 19 in area 2. 5. The firing rate of most tongue RF (79% of 56) neurons and lip RF (60% of 93) neurons tested was significantly altered during the tongue-protrusion task. Only some (14% of 36 tongue RF neurons and 34% of the 92 lip RF neurons tested) showed a significant change in firing rate during the biting task. Three of 7 periodontium RF neurons studied in the tongue-protrusion task altered their firing rate and 5 of 10 altered their firing rate during the biting task. 6. Most of the 116 face SI neurons studied during both tasks exhibited a preferential relation to the tongue-protrusion task as distinct from the biting task, and none showed task-related activity during the biting task only.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 07/1994; 71(6):2377-90. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. Studies using ablation, intracortical microstimulation (ICMS) and surface stimulation, and single-neuron recordings have suggested that the primate primary somatosensory cortex (SI) may play an important role in movement control. Our aim was to determine whether bilateral inactivation of face SI would indeed interfere with the control of tongue or jaw-closing movements. 2. Effects of reversible inactivation by cooling of face SI was investigated in two monkeys trained to perform both a tongue-protrusion task and a biting task. The cooling experiments were carried out after the orofacial representation within SI was identified by systematically defining the mechanoreceptive field of single neurons recorded in face SI. The deficits in the tongue or jaw-closing movement were evaluated by the success rates for the monkeys' performance of both tasks and by the force and electromyographic (EMG) activity recorded from the masseter, genioglossus, and digastric muscles associated with the tasks. 3. During bilateral cooling of face SI, there was a statistically significant reduction in the success rates for the performance of the tongue-protrusion task in comparison with control series of trials while the thermodes used to cool face SI were kept at 37 degrees C. Detailed analyses of force and EMG activity showed that the principal deficit was the inability of the monkeys to maintain a steady tongue-protrusive force in the force holding period during each trial and to exert a consistent tongue-protrusion force between different trials. The task performance returned to control protocol levels at 4 min after commencement of rewarming. 4. Identical cooling conditions did not significantly affect the success rates for the performance of the biting task. Although the extent of the deficit was not severe enough to cause a significant reduction in successful rates for the biting task, cooling did significantly affect the ability of the monkeys to maintain a steady force in the holding period during each trial and to exert a consistent force between different trials. In one monkey the success rate of the biting task was also not affected by bilaterally cooling of face SI with a pair of larger thermodes placed on the dura over most of the face SI, face primary motor cortex (face MI), and adjacent cortical regions.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 10/1993; 70(3):985-96. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. We have recently demonstrated that reversible, cooling-induced inactivation of the face motor cortex results in a severe impairment in the ability of monkeys (Macaca fascicularis) to perform a tongue-protrusion task but produces only relatively minor effects on the performance of a biting task by the same monkeys. To establish a neuronal correlate for these different behavioral relations, the present study has detailed the afferent input and intracortical microstimulation (ICMS)-defined output features of a population of face motor cortical neurons, and in a subsequent study we have documented the activities of the same population of neurons during the performance of the tongue-protrusion and biting tasks. 2. Of the 231 single neurons recorded within the face motor cortex, 163 were located at sites from which ICMS (less than or equal to 20 microA) could evoke tongue movements (i.e., "tongue-MI" sites) at the lowest threshold for eliciting orofacial movements. The remainder were located at sites from which ICMS evoked jaw movements ("jaw-MI" sites), face movements ("face-MI" sites), or at a few sites, tongue movements and, at the same threshold intensity, either a jaw movement or a facial movement. 3. We confirmed the general organizational features of the face motor cortex that have been defined in previous studies, but we documented in detail the organizational features for tongue-MI. Thus we found that tongue movements were well represented, whereas jaw-closing movements were poorly represented; the representations for face, jaw, and tongue movements were overlapped; the same ICMS-evoked tongue movement could be multiply represented within tongue-MI; tongue-MI was characterized by a prominent input from superficial mechanosensory afferents, whereas there was little evidence for deep input; a close spatial match was found between ICMS-defined motor output and somatosensory afferent input for tongue-MI. 4. A variety of tongue movements could be evoked by ICMS at tongue-MI sites and were categorized into protrusion, retrusion, laterally directed, and other types of tongue movement. Low-threshold (i.e., less than or equal to 5 microA) ICMS-defined tongue-MI sites, which were considered to represent "efferent zones" projecting relatively directly to motoneurons, were reconstructed three dimensionally to provide insights into the spatial organization of tongue-MI. Examples of each of the four low-threshold efferent-zone categories were usually found throughout the ICMS-defined tongue-MI without any apparent preferential distribution. Furthermore, different low-threshold efferent-zone categories had close spatial relationships to each other in cortex.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 04/1992; 67(3):747-58. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. The previous paper has described in detail the input and output features of single neurons located at sites within primate face motor cortex from which intracortical microstimulation (ICMS, less than or equal to 20 microA) evoked tongue movements at the lowest threshold ("tongue-MI" sites); for comparative purposes, we also reported on the input and output features of a smaller number of neurons recorded at sites from which ICMS could evoke jaw movements ("jaw-MI" sites), facial movements ("face-MI" sites), or, at a few sites, tongue movements and, at the same threshold intensity, either a jaw movement or a facial movement. 2. Our findings of an extensive and diverse representation of sites within face motor cortex of monkeys for the generation of elemental components of tongue movement, and the relatively few sites from which jaw-closing movements could be evoked, were consistent with our recent observations that reversible, cooling-induced inactivation of the face motor cortex severely impaired the performance by monkeys of a tongue-protrusion task but had only relatively minor effects on the performance of a biting task. In an attempt to establish a neuronal correlate for these different behavioral relations, the present study has documented the task-related activities of those single neurons that were characterized in the previous paper in terms of afferent input and ICMS-defined output features. 3. Each task required the development and maintenance by each monkey of a fixed force level for a minimum period of time to obtain a fruit-juice reward. During one or both of these tasks, we characterized the activities of 231 single face motor cortical neurons that were located at the above-mentioned ICMS-defined sites. Neurons were said to be related to a particular task if they showed statistically significant differences in firing rates during the task in comparison with a control pretrial period (PTP). 4. In tongue-MI, there was a significantly higher proportion of neurons (63% of 156 neurons tested) that were related to the tongue-protrusion task than to the biting task (15% of 65). However, in jaw-MI the proportion of neurons that were biting task-related (63% of 19) was significantly higher than the proportion related to the tongue-protrusion task (11% of 9); the proportion of biting task-related neurons at ICMS-defined jaw-closing sites was also higher than that at jaw-opening sites.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 04/1992; 67(3):759-74. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. In previous papers we presented evidence pointing to an important role for face motor cortex in the control of tongue movements. Intracortical microstimulation (ICMS) at many sites within face motor cortex evoked different types of tongue movement, and many neurons at these "tongue-MI" sites received intraoral mechanosensitive afferent input, and their activity was related to a tongue-protrusion task performed by a monkey. In view of the synergistic action of the various tongue muscles during tongue movement, we hypothesized that these different tongue-MI sites are recruited to effect the appropriate change in tongue shape and position during a tongue-protrusion movement. A prediction from this hypothesis is that variations in the direction of a tongue-protrusion movement should be associated with variations in the activities within the different tongue-MI efferent zones. Differences in efferent-zone activity should be reflected in differences in the firing rates of neurons that are located at these tongue-MI sites. 2. We trained two monkeys to perform a tongue-protrusion task at each of three directions. The tongue-protrusion task transducer was positioned at 0 degrees (Ts), 30 degrees to the left (Tlt), or 30 degrees to the right (Trt) from the midsagittal plane; the latter two positions were termed asymmetric tongue-protrusion task positions. Single neurons were recorded from tongue-MI during trials of tongue-protrusion task at each of two or three of the above positions. Some of the neurons were also studied during a biting task. In addition, neurons were tested for possible mechanosensitive afferent input. 3. Of the 66 neurons studied, 31 (45%) exhibited directional relations; that is, the change in firing rate between the pretrial period (PTP) and the task period for the tongue-protrusion task was significantly different for each neuron depending on the direction in which the activity of the neuron was studied. 4. The "directional" neurons exhibited a single preferred direction of firing in that the mean firing rate during one direction of tongue-protrusion task was significantly greater than for any other direction. Of the 20 neurons studied at all three directions of tongue-protrusion task, the mean firing rate of each of 18 was highest at one of the asymmetric positions, and 12 of these 18 neurons exhibited a monotonic decrease in absolute firing frequency from one asymmetric task direction to the other. 5. Thirteen of the neurons were also studied while the monkey performed the biting task. Most tongue-MI directional neurons were not related to the biting task.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 04/1992; 67(3):775-85. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. Intracortical microstimulation (ICMS) and surface stimulation studies of primate face motor cortex have shown an extensive representation within face motor cortex devoted to movements of the tongue and face; only a very small representation for jaw-closing movements has ever been demonstrated. These data suggest that face motor cortex plays a critical role in the generation of tongue and facial movements but is less important in the generation of jaw-closing movements. Our aim was to determine whether disruption of primate face motor cortical function would indeed interfere with the generation of tongue movements but would not interfere with the generation of jaw-closing movements. 2. The face motor cortex was reversibly inactivated with the use of cooling in two monkeys that were trained to perform both a tongue-protrusion task and a biting task. Recording of single neuronal activity in the cortex beneath the thermode confirmed the reversible inactivation of the cortex. Each task involved a series of trials in which the monkey was required to produce a preset force level for a 0.5-s force holding period; the monkey received a fruit-juice reward if it successfully completed a task trial. Cooling of the ICMS-defined face motor cortex was achieved bilaterally or, in one experiment, unilaterally by circulating coolant through thermodes placed either on intact dura overlying face motor cortex in both monkeys or directly on the exposed pia in one of the monkeys;thermode temperature was lowered to 3-5 degrees C during cooling. Electromyographic (EMG) recordings were also made from masseter, genioglossus, and digastric muscles. 3. During bilateral cooling of the thermodes on the dura overlying the face motor cortex, there was a significant reduction in the success rates for the performance of the tongue-protrusion task in comparison with control series of trials (i.e., precool and postcool) in which the thermodes were kept at 37 degrees C. Quantitative analyses of force and EMG activity showed that the principal deficit was an inability of each monkey to exert sufficient force with its tongue for a sufficient length of time onto the tongue-protrusion task transducer; this deficit was paralleled by a reduction in the level of genioglossus and digastric EMG activity. At 4 min after commencement of rewarming, task performance had returned to control, precool levels.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 04/1991; 65(3):511-30. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. The lateral part of the pericentral cortex of both hemispheres in three awake monkeys was explored with intracortical microstimulation (ICMS) using short trains (T/S; 200-microseconds pulses at 333 Hz for 35 ms, less than or equal to microA) and long trains (C/S; 200-microseconds pulses at 50 Hz for 3 s, less than or equal to 60 microA). In both hemispheres of one of these monkeys, the responsiveness of single cortical neurons to stimulation of the orofacial region was tested at the same intracortical sites where ICMS was applied. 2. Movements were evoked from four physiologically defined cortical regions: the primary face motor cortex (MI), the primary face somatosensory cortex (SI), the principal part of the cortical masticatory area (CMAp) which was located in the precentral gyrus lateral to MI, and a deep part of the cortical masticatory area (CMAd) which was located in the inferior face of the frontal operculum. 3. Two types of cortically induced movements were observed: a single twitch movement and EMG activity of the orofacial muscles that was evoked by T/S at a short latency (10-45 ms) and rhythmical jaw movements (RJMs) which were only evoked by C/S. 4. RJMs were evoked at C/S frequencies ranging from 20 to 300 Hz. At movement threshold, the frequency of the cortically induced RJMs varied from 0.7 to 1.5 Hz and usually increased with the increase of C/S intensity up to 2 times movement threshold. The vertical amplitude of RJMs was also stimulus dependent, and at movement threshold it ranged from 3 to 9 mm. 5. The movement patterns of the cortically induced RJMs remained constant during the course of C/S but could be differentiated in the frontal plane into ipsilateral- (RJMi), vertical-(RJMv), and contralateral- (RJMc) directed movements. These three different patterns of RJMs were associated with different patterns of masticatory muscle activity. 6. Each cortical region contained many sites from which RJMs could be induced (so-called RJM sites). The RJMi sites were more numerous than RJMc sites in all regions except SI and were located anterolateral or lateral to the RJMc sites in each region; the RJMv sites were scattered throughout each cortical region. 7. In MI, C/S elicited RJMs from 94 intracortical sites from which short-latency twitch movements could also be evoked by T/S.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 04/1989; 61(3):635-50. · 3.30 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 1. The technique of intracortical microstimulation (ICMS), supplemented by single-neuron recording, was used to carry out an extensive mapping of the face primary motor cortex. The ICMS study involved a total of 969 microelectrode penetrations carried out in 10 unanesthetized monkeys (Macaca fascicularis). 2. Monitoring of ICMS-evoked movements and associated electromyographic (EMG) activity revealed a general pattern of motor cortical organization. This was characterized by a representation of the facial musculature, which partially enclosed and overlapped the rostral, medial, and caudal borders of the more laterally located cortical regions representing the jaw and tongue musculatures. Responses were evoked at ICMS thresholds as low as 1 microA, and the latency of the suprathreshold EMG responses ranged from 10 to 45 ms. 3. Although contralateral movements predominated, a representation of ipsilateral movements was found, which was much more extensive than previously reported and which was intermingled with the contralateral representations in the anterior face motor cortex. 4. In examining the fine organizational pattern of the representations, we found clear evidence for multiple representation of a particular muscle, thus supporting other investigations of the motor cortex, which indicate that multiple, yet discrete, efferent microzones represent an essential organizational principle of the motor cortex. 5. The close interrelationship of the representations of all three muscle groups, as well as the presence of a considerable ipsilateral representation, may allow for the necessary integration of unilateral or bilateral activities of the numerous face, jaw, and tongue muscles, which is a feature of many of the movement patterns in which these various muscles participate. 6. In six of these same animals, plus an additional two animals, single-neuron recordings were made in the motor and adjacent sensory cortices in the anesthetized state. These neurons were electrophysiologically identified as corticobulbar projection neurons or as nonprojection neurons responsive to superficial or deep orofacial afferent inputs. The rostral, medial, lateral, and caudal borders of the face motor cortex were delineated with greater definition by ICMS and these electrophysiological procedures than by cytoarchitectonic features alone. We noted that there was an approximate fit in area 4 between the extent of projection neurons and field potentials anti-dromically evoked from the brain stem and the extent of positive ICMS sites.(ABSTRACT TRUNCATED AT 400 WORDS)
Journal of Neurophysiology 04/1988; 59(3):796-818. · 3.30 Impact Factor