Directing attention to some acoustic features of a sound has been shown repeatedly to modulate the stimulus-induced neural responses. On the contrary, little is known about the neurophysiological impact of auditory attention when the auditory scene remains empty. We performed an experiment in which subjects had to detect a sound emerging from silence (the sound was detectable after different durations of silence). Two frontal activations (right dorsolateral prefrontal and inferior frontal) were found, regardless of the side where sound was searched for, consistent with the well established role of these regions in attentional control. The main result was that the superior temporal cortex showed activations contralateral to the side where sound was expected to be present. The area extended from the vicinity of Heschl's gyrus to the surrounding areas (planum temporale/anterior lateral areas). The effect consisted of both an increase in the response to a sound delivered after attention was directed to detect its emergence and a baseline shift during the silent period. Thus, in absence of any acoustic stimulus, the search for an auditory input was found to activate the auditory cortex.
Using two-dimensional (2D) angles composed of two straight, 8-cm-long arms that formed an angle, we investigated the importance of cutaneous feedback from the exploring index finger, and proprioceptive feedback from the shoulder (scanning movements made with the outstretched arm), to the human ability to discriminate small differences in the angles. Using a two-alternative forced-choice paradigm, subjects identified the larger angle in each pair explored (standard angle, 90 degrees; comparison angles, 91 degrees to 103 degrees). Subjects were tested under four experimental conditions: (1) active touch (reference condition); (2) active touch with digital anaesthesia; (3) passive touch (a computer-controlled device displaced the angle under the subject's immobile digit); and (4) passive touch with digital anaesthesia. When only proprioceptive feedback from the shoulder was available (condition 2), there was a significant increase in discrimination threshold, from 4.0 degrees in the reference condition (condition 1) to 7.2 degrees, indicating that cutaneous feedback from the exploring digit contributed to task performance. When only cutaneous feedback from the finger was available (condition 3), there was also a significant increase in threshold from 4.2 degrees in the active condition to 8.7 degrees. This suggested that proprioceptive feedback from the shoulder, potentially from a variety of deep (muscle and joint) but also cutaneous receptors, contributed to the ability to discriminate small changes in 2D angles. When both sources of feedback were eliminated (condition 4), subjects were unable to discriminate even the largest difference presented (13 degrees). The results suggest that this sensory task is truly an integrative task drawing on sensory information from two different submodalities and so, following the definition of Gibson, is haptic in nature. The results are discussed in relation to the potential neural mechanisms that might underlie a task that requires integration across two anatomically separate body parts and two distinct modalities.
The human ability to recognize objects on the basis of their shape, as defined by active exploratory movements, is dependent on sensory feedback from mechanoreceptors located both in the skin and in deep structures ( haptic feedback). Surprisingly, we have little information about the mechanisms for integrating these different signals into a single sensory percept. With the eventual aim of studying the underlying central neural mechanisms, we developed a shape discrimination test that required active exploration of objects, but was restricted to one component of shape, two-dimensional (2D) angles. The angles were machined from 1-cm-thick Plexiglas, and consisted of two 8-cm-long arms that met to form an angle of 90 degrees (standard) or 91 degrees to 103 degrees (comparison angles). Subjects scanned pairs of angles with the index finger of the outstretched arm and identified the larger angle of each pair explored. Discrimination threshold (75% correct) was 4.7 degrees (range 0.7 degrees to 12.1 degrees), giving a precision of 5.2% (0.8-13.4%: difference/standard). Repeated blocks of trials, either in the same session or on different days, had no effect on discrimination threshold. In contrast, the motor strategy was partly modified: scanning speed increased but dwell-time at the intersection did not change. Finally, 2D angle discrimination was not significantly modified by rotating the orientation of one of the angles in the pair (0 degrees, 4 degrees or 8 degrees rotation towards the midline, in the vertical plane), providing evidence that subjects evaluated each angle independently in each trial. Subject reports indicated that they relied on cutaneous feedback from the exploring digit (amount of compression of the finger at the angle) and mental images of the angles, most likely arising from proprioceptive information (from the shoulder) generated during the to-and-fro scans over the angle. In terms of shoulder angles, the mean discrimination threshold here was 0.54 degrees (range 0.08 degrees to 1.36 degrees). These values are lower than previous estimates of position sense at the shoulder. In light of the subjects' strategies, it therefore seems likely that both cutaneous and proprioceptive (including both dynamic and static position-related signals) feedback contributed to the haptic discrimination of 2D angles.
Beneficial effects of mental practice likely arise because motor imagery involves largely similar neural networks as physical execution of the same movement. While it is known that the involvement of the motor system is favoured by focusing on the kinaesthetic modality and by the first person perspective, little is known about the impact of these factors on the somatosensory system. The present paper examines the effects on the somatosensory excitability of both perspective (the point of view of the person imagining a motor act) and modality (visual versus kinaesthetic) during mental practice. Seventeen healthy subjects participated. Quality of mental practice was controlled using chronometric tests and a subjective questionnaire. Excitability of the somatosensory system was assessed through the steady-state electroencephalographical response to a continuous train of electrical stimuli applied to the radial nerve, at the same time subjects were instructed to perform one of five tasks designed to separate the effects of perspective, modality and motor versus non-motor imagery. Kinaesthetic motor imagery exerts the largest effect on somatosensory excitability whereas visual motor imageries (1st and 3rd person perspectives) produce the same lower effect that static visual imagery does. Strikingly, specific effect of kinaesthetic motor imagery correlates with the selfselected speed to imagine and execute the same movement. These findings suggest a key role of the kinaesthetic content of motor imagery in recruiting the sensorimotor system.
Observation of hand movements has been repeatedly demonstrated to increase the excitability of the motor cortical representation of the hand. Little attention, however, has been devoted to its effect on somatosensory processing. Movement execution is well known to decrease somatosensory cortical excitability, a phenomenon termed 'gating'. As executed and observed actions share common cortical representations, we hypothesized that action observation (hand movements) should also modulate the cortical response to sensory stimulation of the hand. Seventeen healthy subjects participated in these experiments in which electroencephalographic (EEG) recordings of the somatosensory steady-state response (SSSR) were obtained. The SSSR provides a continuous measure of somatosensory processing. Recordings were made during a baseline condition and five observation conditions in which videos showed either a: (1) hand action; (2) passive stimulation of a hand; (3) static hand; (4) foot action; or (5) static object. The method employed consisted of applying a continuous 25 Hz vibratory stimulation to the index finger during the six conditions and measuring potential gating effects in the SSSR within the 25 Hz band (corresponding to the stimulation frequency). A significant effect of condition was found over the contralateral parietal cortex. Observation of hand actions resulted in a significant gating effect when compared to baseline (average gating of 22%). Observation of passive touch of the hand also gated the response (17% decrease). In conclusion, the results show that viewing a hand performing an action or being touched interferes with the processing of somatosensory information arising from the hand.
This electroencephalography study demonstrates with a quick, easily implementable, and noninvasive paradigm that the change in cortical somatosensory excitability during pain observation is limb-specific, and confirms from a neuroscience perspective that being exposed to others' pain implies more than the sharing of an affective experience.
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