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The tactile sensitivity of the teeth, and associated periodontium, serves important sensory and motor functions. Microneurographic recordings from human periodontal ligament mechanoreceptor (PDLM) nerves, in response to tooth loading, reveal discharge patterns with sole slowly adapting (SA) II-type characteristics, highlighting the unique role of PDLMs in oral sensory processes. Here we investigate these receptors' properties, psychophysically and with neuroimaging (fMRI), in response to varying frequencies of dynamic (vibrotactile) stimulation. The finding of increased activity in primary (SI) and secondary (SII) somatosensory cortices (SI and SII) at low frequencies of stimulation (20 Hz) as compared with higher frequencies (50 and 100 Hz), shows an increased entrainment of the PDLMs at this lower frequency in line with expected SA II-type response properties. At the highest frequency (100 Hz), no significant activity was found in SI or SII, suggesting this frequency is outside the range of activity of PDLMs. An activation matrix is mapped that includes SI, SII, insular, inferior frontal gyrus, inferior parietal lobe and supplementary motor area as well as middle frontal gyrus and cerebellum. We compared the responses to tooth stimulation with those produced by identical vibrotactile stimulation of the finger. The results strongly suggest that the PDLMs play a significant role in the specification of the forces used to hold and manipulate food between teeth, and in these respects, the masticatory system appears analogous to fine finger-control mechanisms used during precision manipulation of small objects. Because fMRI reveals activations in posterior insular cortex, we also speculate that PDLMs, and SA II-type receptors in general, may be involved in one aspect of the feeling of body ownership.

Microneurographic recordings were obtained from 20 periodontal mechanoreceptive afferents in the inferior alveolar nerve while force profiles of different amplitudes and rates were applied to a premolar or the first molar in the most sensitive direction. The majority of afferents (17/20) showed a hyperbolic relationship between the steady-state discharge rate and the amplitude of the stimulating force, featuring a pronounced saturation tendency. These afferents were also characterized by a similar decline in dynamic sensitivity with increasing amplitude of background force. However, a few afferents (3/20) showed nearly linear stimulus-response relationships and a small decline in dynamic sensitivity with increasing tooth load. Quantitative models developed for all afferents successfully predicted the afferent discharge rates for novel force stimulations. Application of the transfer function to chewing forces predicted that the discharge rates of periodontal afferents rapidly increased at initial tooth contact and continued to discharge as long as the tooth was loaded. However, due to the marked saturation tendencies at higher forces, most periodontal afferents poorly encoded the magnitude of the strong chewing forces. In addition, the discharge rates of a minority of afferents continued to reflect the force profile during high chewing forces. The results revealed that periodontal afferents of posterior teeth were less sensitive at low tooth loads compared with afferents of anterior teeth. During each chewing cycle, periodontal afferents may provide information about the mechanical properties of food shortly after tooth contact that can be used to scale the muscle commands of the upcoming power phase.

Impulses in 45 single mechanoreceptive afferents were recorded from the human inferior alveolar nerve with permucosally inserted tungsten microelectrodes. All afferents responded to mechanical stimulation of one or more premolar or molar teeth and most likely innervated their periodontal ligaments. For each afferent, isolated "ramp-and-hold" shaped force profiles of similar magnitudes (252 +/- 24 mN; mean +/- SD) were applied to the lower first premolar, the second premolar, and the first molar on the recording side. The tooth loads were applied in six directions: lingual, facial, mesial, and distal in the horizontal plane and up and down in the vertical direction of the tooth. The afferents response during the static phase of the stimulus was analyzed. All afferents were slowly adapting, discharging continuously in response to static forces in at least one stimulation direction. Twenty-nine afferents (64%) were spontaneously active, exhibiting an ongoing discharge in the absence of external stimulation. Stimulation of a single tooth was found to excite each afferent most strongly. The most sensitive tooth (MST) was the first premolar for 23, the second premolar for 13, and the first molar for 9 afferents. About half of the afferent population also responded to loading of one or two more teeth. The response profiles of these afferents indicated that the multiple-teeth receptive fields were due to mechanical coupling between the teeth rather than branching of single afferents to innervate several teeth. The afferent responses to loading the mesial and distal halves of the first molars were very similar. Thus both intensive and directional aspects of the afferent response when loading one side of the tooth was preserved to a great extent when loading the other side. When loading the MST, the afferents typically showed excitatory responses in two to four of the six stimulation directions, i.e., the afferents were broadly tuned to direction of tooth loading. In the horizontal plane, the afferent populations at the premolar teeth expressed no clear directional preferences. The afferents at the molar, however, showed a strong directional bias in the distal-lingual direction. In the vertical plane, there was a preference for downward-directed forces with a gradually decreasing sensitivity distally along the dental arch. The present results demonstrate that human periodontal afferents supplying anterior and posterior teeth differ in their capacity to signal horizontal and vertical forces, respectively.

Intrafascicular multiunit activity and impulses in single mechanoreceptive afferents were recorded from the human lingual nerve with permucosally inserted tungsten microelectrodes. Nylon filaments and blunt glass probes were used for mechanical stimulation of the mucosa of the dorsal surface of the tongue. The innervation territories of nine nerve fascicles were mapped during multiunit recordings. All fascicle fields included the tip of the tongue, suggesting a particularly high innervation density for this area. Thirty-three single mechanoreceptive afferents were isolated and studied. Of these afferents, 22 were characterized by very small mucosal receptive fields (range: 1-19.6 mm2; geometric mean: 2.4 mm2) and responded to extremely low mechanical forces (force threshold range: 0.03-2 mN; geometric mean: 0.15 mN). As such, it was concluded that these "superficial" units terminated near the surface of the tongue. The remaining 11 units responded to probing of large areas of the tongue (> 200 mm2) and exhibited high force thresholds (> or = 4 mN). It was concluded that these "deep" units terminated in the muscle mass of the tongue. Fourteen of the superficial units were classified as rapidly adapting and resembled the fast-adapting type I afferents described for the glabrous skin of the human hand. The rapidly adapting units responded both during the application and removal of, but not during maintenance of, the mechanical stimuli on the receptive field. Two types of slowly adapting responses were observed. One type (characteristic of only 2 units) was characterized by a pronounced sensitivity to force change during the application and removal of the mechanical stimuli and an irregular static discharge during maintenance of the stimulus on the receptive field. In contrast, the other six units exhibited a weak sensitivity to force change, a highly regular static discharge, and spontaneous activity. As such, these two types of slowly adapting units resembled the slowly adapting I and II afferents, respectively, described for the hand. All 11 deep units were slowly adapting, and 7 were, in addition, spontaneously active. The units were not equally sensitive to the application and removal of the mechanical stimuli, suggesting at least two different modes of termination in tongue muscle. The deep units reliably encoded information about tongue movements in the absence of direct contact with the receptive field. In contrast, the superficial units responded vigorously when the tongue was moved to bring the receptive field into physical contact with other intraoral structures.

Dense multichannel recordings of scalp electroencephalogram were obtained in the vicinity of primary somatosensory cortex, time-locked to repetitive train microstimulation of single, physiologically characterized skin mechanoreceptive afferents in the median nerve of a single human subject. Frequency-domain analysis of cross-trial averages for fast-adapting type one and slowly adapting type one afferents revealed prominent, topographically organized "driving" responses in the electroencephalogram at the frequency of stimulation, which vanished under various statistical and experimental control conditions. The responses also exhibited systematic declines in amplitude both across and within trials, and orderly changes in scalp topography as a function of the location of afferents' receptive fields on the hand. The observed response properties are tentatively explained in terms of characteristics of the pattern of afferent drive impressed on the cortex by microstimulation.