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In this study we asked whether asymmetries related to handedness in right-handers (n=10) and left-handers (n=10) could influence hemispheric motor facilitation when the preferred or less-preferred hand is engaged in haptic sensing. Transcranial magnetic stimulation (TMS) was used to assess corticomotor excitability when young participants performed a precision grip under two conditions, i.e., with or without a haptic sensing component. In the grip condition participants were required to grasp a 5-mm thick plate between thumb and index fingers, whereas in the haptic condition they performed the same action but were also required to make judgements about the plate's thickness, i.e., either thin (5 mm) or thick (10 mm). Analysis of task-related variations in motor evoked potentials (MEP) amplitude recorded in hand muscles revealed a significant task×hand/hemisphere interaction only in the group of right-handers when the left hand/right hemisphere was engaged in haptic sensing as opposed to simply gripping. Collectively, these results are congruent with other TMS reports describing a greater degree of hemispheric asymmetry in right-handers, who are typically more strongly lateralised than left-handers. Our results with regard to a rightward hemispheric asymmetry for MEP facilitation with haptic sensing are also congruent with the predominant role ascribed to the right sensorimotor cortex in the processing of proprioceptive information.

Background: Haptic sensing with the fingers represents a unique class of manipulative actions, engaging motor, somatosensory and associative areas of the cortex while requiring only minimal forces and relatively simple movement patterns. Using transcranial magnetic stimulation (TMS), we investigated task-related changes in motor evoked potential (MEP) amplitude associated with unimanual haptic sensing in two related experiments. In Experiment I, we contrasted changes in the excitability of the hemisphere controlling the task hand in young and old adults under two trial conditions, i.e. when participants either touched a fine grating (smooth trials) or touched a coarse grating to detect its groove orientation (grating trials). In Experiment II, the same contrast between tasks was performed but with TMS applied over the hemisphere controlling the resting hand, while also addressing hemispheric (right vs. left) and age differences., Results: In Experiment I, a main effect of trial type on MEP amplitude was detected (p = 0.001), MEPs in the task hand being ~50% larger during grating than smooth trials. No interaction with age was detected. Similar results were found for Experiment II, trial type having a large effect on MEP amplitude in the resting hand (p < 0.001) owing to selective increase in MEP size (~2.6 times greater) for grating trials. No interactions with age or side (right vs. left) were detected., Conclusions: Collectively, these results indicate that adding a haptic component to a simple unilateral finger action can elicit robust corticomotor facilitation not only in the working hemisphere but also in the opposite hemisphere. The fact that this facilitation seems well preserved with age, when task difficulty is adjusted, has some potential clinical implications.

Background: Active contraction leads to facilitation of motor responses evoked by transcranial magnetic stimulation (TMS). In small hand muscles, motor facilitation is known to be also influenced by the nature of the task. Recently, we showed that corticomotor facilitation was selectively enhanced when young participants actively discriminated tactile symbols with the tip of their index or little finger. This tactile-dependent motor facilitation reflected, for the large part, attentional influences associated with performing tactile discrimination, since execution of a concomitant distraction task abolished facilitation. In the present report, we extend these observations to examine the influence of age on the ability to produce extra motor facilitation when the hand is used for sensory exploration., Methods: Corticomotor excitability was tested in 16 healthy seniors (58-83 years) while they actively moved their right index finger over a surface under two task conditions. In the tactile discrimination (TD) condition, participants attended to the spatial location of two tactile symbols on the explored surface, while in the non discrimination (ND) condition, participants simply moved their finger over a blank surface. Changes in amplitude, in latency and in the silent period (SP) duration were measured from recordings of motor evoked potentials (MEP) in the right first dorsal interosseous muscle in response to TMS of the left motor cortex., Results: Healthy seniors exhibited widely varying levels of performance with the TD task, older age being associated with lower accuracy and vice-versa. Large inter-individual variations were also observed in terms of tactile-specific corticomotor facilitation. Regrouping seniors into higher (n = 6) and lower performance groups (n = 10) revealed a significant task by performance interaction. This latter interaction reflected differences between higher and lower performance groups; tactile-related facilitation being observed mainly in the former group. Latency measurements and SP durations were not affected by task conditions., Conclusions: The present findings provide further insights into the factors influencing task-dependent changes in corticomotor excitability in the context of aging. Our results, in particular, highlight the importance of adjusting task demands and controlling for attention when attempting to elicit task-specific motor facilitation in older persons engaged in fine manual actions. Such information could be critical in the future for planning interventions to re-educate or maintain hand function in the presence of neurological impairments.

Corticomotor facilitation was investigated using transcranial magnetic stimulation in healthy young adults when they actively stroked either their own left palm (intraactive) or the experimenter's palm (interactive touch) with their right index finger. We predicted, based on the sensory cancellation hypothesis, that corticomotor facilitation would be lower with intraactive touch. Motor evoked potential amplitude in the right first dorsal interosseous was affected by mode of touch, but not by sex. In contrast to our prediction, motor evoked potential facilitation was larger (mean 14%) during intraactive touch. The present results are in agreement with recent evidence suggesting that self-touch represents a unique class of sensorimotor experiences that are critical for the elaboration of internal body structure representation.

This study examined tactile pattern recognition performance in human observers (N = 44) in the context of a letter recognition task at the fingertip. Participants were recruited from three different age groups (youth, n = 17; young adults, n = 14; seniors, n = 13) to examine age-related differences in performance. The influence of gender (males vs females) and hand (right vs left) was also examined. Performance was characterized in terms of both response accuracy and associated response times (RTs). Patterns of confusion between letters were also examined. Results showed that age was the most important factor in determining the capacity of our participants to perform fast and accurate pattern recognition. In this respect, younger participants (i.e., youth and young adults) clearly outperformed seniors by showing not only better accuracy and less confusion but also 2-3 times faster RT. By comparison, the combined influence of "hand" and "gender" on recognition performance was only marginal. These results indicate that the ability to perform complex tactile pattern recognition is already well established in youth 10-14 years of age with only minor refinements occurring later in early adulthood. With advancing age, such ability becomes far less efficient, as judged by the drastic increase in RT observed in seniors, in spite of a relatively good accuracy. This suggests that alterations not only at the peripheral receptor level but also at the central processing level might play an important role in limiting the ability of seniors to perform fast and efficient pattern recognition at the fingertip.

In the present report, we extend our previous observations on the effect of age on postural stabilization from fingertip contact (Exp Brain Res 157 (2004) 275) to examine the possible influence of sensory thresholds measured at the fingertip on the magnitude of contact forces. Participants (young, n=25, 19-32 years; old, n=35, 60-86 years) underwent psychophysical testing of the right index finger to determine thresholds for spatial acuity, pressure sensitivity and kinesthetic acuity. Spatial acuity was determined from the ability to detect gaps of different widths, while Semmes-Weinstein monofilaments were used for pressure sensitivity. Kinesthetic acuity was determined by asking participants to discriminate plates of different thicknesses using a thumb-index precision grip. These tests were selected on the basis that each reflected different sensory coding mechanisms (resolution of spatial stimuli, detection of mechanical forces and integration of multi-sensory inputs for hand conformation) and thus provided specific information about the integrity and function of mechanoreceptive afferents innervating the hand. After log transformation, thresholds were first examined to determine the influence of age (young and old) and gender (male, female) on tactile acuity. Sensory thresholds were then entered into multiple linear regression models to examine their ability to predict fingertip contact forces (normal and tangential) applied to a smooth surface when subjects stood with eyes closed on either a firm or a compliant support surface. As expected, age exerted a significant effect (p<0.01) on all three thresholds, but its impact was greater on spatial acuity than on pressure sensitivity or kinesthetic acuity. Gender had a marginal impact on pressure sensitivity thresholds only. The regression analyses revealed that tactile thresholds determined at the index fingertip accounted for a substantial proportion of the variance (up to 30%) seen in the contact forces deployed on the touch-plate, especially those exerted in the normal direction. The same analyses further revealed that much of the variance explained by the models arose from inter-individual differences in tactile spatial acuity and not from differences in pressure sensitivity or in kinesthetic acuity. Thus, of all three tests, the spatial acuity task was the most sensitive at detecting differences in hand sensibility both within and between age groups and, accordingly, was also better at predicting the magnitude of fingertip forces deployed for postural stabilization. Since spatial acuity is critically dependent upon innervation density, we conclude that the degree of functional innervation at the fingertip was likely an important factor in determining the capacity of older participants to use contact cues for stability purposes, forcing the most affected individuals to exert unusually high pressures in order to achieve stabilization in the presence of reduced tactile inputs arising from contact with the touched surface.

In this study, we compared the ability of young (n=10, 19-32 years) and older subjects (n=35, 60-86 years) to use fingertip contact as a balance aid during quiet stance under various conditions to determine whether aging would influence contact strategies. Experimental trials (duration, 60 s) included two visual conditions (vision; no vision), three fingertip contact conditions (no touch; smooth touch; rough touch) and two support surface conditions (firm; foam). In trials with contact, participants were required to maintain a light contact with their right index fingertip on an instrumented touch-plate. Subjects were not constrained to exert minimal contact force, although they were aware that the touch-plate was not designed for physical support. From displacements of the centre of foot pressure (COP), mean sway amplitude (MSA) was computed in the anterior-posterior (COP(AP)) and medio-lateral (COP(ML)) directions. Subjective estimates of stability were also obtained by asking participants to rate perceived stability on a visual analog scale in each condition. Mean normal force (FN) and mean resultant tangential force (F(TAN)) were computed from contact force data applied on the touch plate. In both age groups, touch conditions had a substantial effect on MSA in the AP direction under both support surface conditions, with reductions averaging between 40-55% when touch was allowed. Reductions in the ML direction, though less important (8-12% on average), were nevertheless highly significant, especially in the older subjects when standing on the foam. In the two groups, vision and texture had only marginal impact on MSA computed on both support surfaces. Contrasting with sway measurements, stability ratings were highly influenced by visual conditions in both age groups. Only in conditions of deficient support (foam surface) and absent vision did the perceived effect of touch exceed that of vision. Age had a major impact, however, on contact forces deployed during trials with touch. While individuals in the young group typically produced forces of <1 N (mean FN, 0.32+/-0.15 N) to achieve postural stabilization, older subjects tended to use higher, though not too excessive, contact forces (mean FN, 1.21+/-0.75 N) under the same conditions. From these findings, we conclude that the ability to use contact cues from the fingertip as a source of sensory information to improve postural stability is largely preserved in healthy older adults. The increase in contact force deployed by older individuals to achieve postural stabilization is interpreted as a compensatory strategy to help overcome age-related loss in tactile sensation, an issue that will be further addressed in a companion paper.

In this study, we sought to better define the limit of spatial resolution at the fingertips of elderly participants (n = 30, age 60-95 years) using an extended set of JVP grating domes, incorporating four new grating dimensions (2.5-, 3.5-, 4.0- and 4.5-mm width). A secondary aim was to examine whether deficits in tactile acuity could be related to hand dysfunction in older adults. Spatial resolution thresholds were determined by the finest grating whose orientation (dominant index finger) could be reported reliably. Manual dexterity was assessed with the Grooved Pegboard Test (GPT). The extended set of domes improved threshold measurements in a majority of participants (21/30). Still, accurate threshold estimates could not be obtained in one third of the participants, mostly in the older age group (8/9, 74-95 years). Grating resolution thresholds at the index finger were strongly correlated (r = 0.66, p<0.01) with dexterity scores derived from the GPT. From these results, we conclude that the 2.5- and 3.5-mm grating domes are suitable additions when assessing spatial acuity at the fingertips of older subjects between 60 and 70 years of age (mean threshold, 2.7+/-0.6 mm). For the older ones, the 4.0- and 4.5-mm domes can improve threshold measurements but interpretation of values can be complicated by the presence of undiagnosed pathologies (e.g., diffuse polyneuropathy) as people advance in age. The strong relationship between grating resolution thresholds and dexterity scores indicates that an impaired spatial acuity at the fingertips may translate into great difficulties in tasks requiring fine manipulations. These findings have important implications for the assessment of hand function in older adults.

This paper summarizes recent work showing that tactile roughness appreciation increases in a nearly linear fashion as tactile element spacing or spatial period (SP, distance centre-to-centre between raised dots in these experiments) is increased from 1.5 to 8.5 mm. Although a previous study had reported a U-shaped psychophysical function peaking at a nominal SP of 3.2 mm, differences in the surfaces (including changing SP in only one dimension as compared with two and higher dot heights that minimized contact with the smooth floor) likely contributed to the difference in the results. Roughness estimates were also unaffected by a 2-fold change in scanning speed (50 vs. 95 mm/s). Parallel recordings from neurones in primary somatosensory cortex (SI) during a texture discrimination task indicate that the discharge frequency of many SI cells shows a monotonic relation with SP (up to 5 mm tested). For some cells, the texture signals were ambiguous because discharge frequency co-varied with both texture and the scanning speed, as has also been reported for the peripheral mechanoreceptors that are activated by textured surfaces. Yet other SI cells showed a speed-invariant response to surface texture, consistent with perceptual constancy for roughness over a range of scanning speeds. We suggest that such a discharge pattern could be based on a simple intensive, or mean rate, code: an invariant central representation of surface texture could be obtained by subtracting a speed-varying signal from the ambiguous signals that co-vary with roughness and speed.

Recent studies suggest that sensory input generated during highly repetitive tasks can degrade the sensory representation of the hand and eventually lead to sensory and motor problems. In this study, we investigated whether early changes in tactile perception and manual dexterity could be detected in persons exposed to computer tasks. Performance in tests designed to assess tactile perception (grating orientation task for spatial acuity and roughness discrimination) and manual dexterity (grooved pegboard test) was compared between two groups of healthy individuals, matched for age, gender, and experience, who differed in terms of computer habits. One group consisted of frequent users (FU, > 2 h/day, n = 36) and the other of non or occasional users (OU, < 2 h/day, n = 28). Comparison of performance between groups with subjects sorted by gender revealed significant differences (t-test, p < 0.05) in female, but not male, participants. Grating resolution thresholds at the tip on the second and fifth digits were, on average, 40% higher in female FU (n = 13) than in female OU (n = 10) and performance scores on the dexterity test were significantly higher for the left hand. The results of this study indicate that early signs of deterioration in hand function can be present in persons constantly exposed to computer tasks and that these signs are more readily apparent in women than in men. The loss of tactile spatial acuity found in female FU possibly reflect an early consequence of the degraded sensory representation of the hand resulting from constant repetitions of fine motor tasks.

Two rhesus monkeys were trained to discriminate, with the use of passive touch, a standard surface [rectangular arrays of raised dots with a spatial period (SP) of 2 mm across the rows and columns] from three modified surfaces in which the SP between rows was increased to 3, 4, or 5 mm over the second half of the surface. After the surface presentation (to digit tips 3 and 4 of one hand) the monkeys indicated the presence or absence of a change in texture by pulling or pushing a lever, respectively, with the opposite hand. Of 193 neurons recorded from primary somatosensory cortex (SI, 3 hemispheres) and 94 neurons from secondary somatosensory cortex (SII, 1 hemisphere), all contralateral to the stimulated hand, the discharge of 51 SI and 19 SII neurons was classified as texture related. Two types of texture-related responses were obtained. Graded neurons showed a linear relationship between mean discharge frequency and SP; nongraded neurons showed a significant change in discharge over the modified half of the surfaces but the discharge did not distinguish between the three modified surfaces. The distribution of these texture responses was significantly different in SI and SII: whereas most of the texture-related neurons in SI (44 of 51, 86%) were graded, the majority of those in SII (12 of 19, 63%) were nongraded. The results were interpreted as suggesting that the nongraded responses reflect feature extraction in SII, signaling the presence of a change in texture but not its magnitude, and so support the notion that texture signals are processed sequentially, first in SI and then in SII.