Transcranial magnetic stimulation over sensorimotor cortex disrupts anticipatory reflex gain modulation for skilled action

Skilled interactions with new environments require flexible changes to the transformation from somatosensory signals to motor outputs. Transcortical reflex gains are known to be modulated according to task and environmental dynamics, but the mechanism of this modulation remains unclear. We examined reflex organization in the sensorimotor cortex. Subjects performed point- to- point arm movements into predictable force fields. When a small perturbation was applied just before the arm encountered the force field, reflex responses in the shoulder muscles changed according to the upcoming force field direction, indicating anticipatory reflex gain modulation. However, when a transcranial magnetic stimulation (TMS) was applied before the reflex response to such perturbations so that the silent period caused by TMS overlapped the reflex processing period, this modulation was abolished, while the reflex itself remained. Loss of reflex gain modulation could not be explained by reduced reflex amplitudes nor by peripheral effects of TMS on the muscles themselves. Instead, we suggest that TMS disrupted interneuronal networks in the sensorimotor cortex, which contribute to reflex gain modulation rather than reflex generation. We suggest that these networks normally provide the adaptability of rapid sensorimotor reflex responses by regulating reflex gains according to the current dynamical environment

Mapping the invisible hand: a body model of a phantom limb

After amputation, individuals often have vivid experiences of their absent limb (i.e., a phantom limb). Therefore, one’s conscious image of one’s body cannot depend on peripheral input only (Ramachandran & Hirstein, 1998). However, the origin of phantom sensations is hotly debated. Reports of vivid phantoms in the case of congenital absence of the limb show that memory of former body state is not necessary (Brugger et al., 2000). According to one view, phantoms may reflect innate organization of sensorimotor cortices (Melzack, 1990). Alternatively, phantoms could reflect generalization from viewing other people’s bodies (Brugger et al., 2000), a sensorimotor example of the classic theory that understanding oneself follows from understanding the “generalized other” (Mead, 1934, p. 154). Because phantom limbs cannot be stimulated, sensory testing cannot directly compare visual and somatosensory influences on representations of phantom limbs. Consequently, empirical investigation of phantoms is limited

Experimental investigations of control principles of involuntary movement: a comprehensive review of the Kohnstamm phenomenon

The Kohnstamm phenomenon refers to the observation that if one pushes the arm hard outwards against a fixed surface for about 30 s, and then moves away from the surface and relaxes, an involuntary movement of the arm occurs, accompanied by a feeling of lightness. Central, peripheral and hybrid theories of the Kohnstamm phenomenon have been advanced. Afferent signals may be irrelevant if purely central theories hold. Alternatively, according to peripheral accounts, altered afferent signalling actually drives the involuntary movement. Hybrid theories suggest afferent signals control a centrally-programmed aftercontraction via negative position feedback control or positive force feedback control. The Kohnstamm phenomenon has provided an important scientific method for comparing voluntary with involuntary movement, both with respect to subjective experience, and for investigating whether involuntary movements can be brought under voluntary control. A full review of the literature reveals that a hybrid model best explains the Kohnstamm phenomenon. On this model, a central adaptation interacts with afferent signals at multiple levels of the motor hierarchy. The model assumes that a Kohnstamm generator sends output via the same pathways as voluntary movement, yet the resulting movement feels involuntary due to a lack of an efference copy to cancel against sensory inflow. This organisation suggests the Kohnstamm phenomenon could represent an amplification of neuromotor processes normally involved in automatic postural maintenance. Future work should determine which afferent signals contribute to the Kohnstamm phenomenon, the location of the Kohnstamm generator, and the principle of feedback control operating during the aftercontraction

Discreteness of the volume of space from Bohr-Sommerfeld quantization

A major challenge for any theory of quantum gravity is to quantize general relativity while retaining some part of its geometrical character. We present new evidence for the idea that this can be achieved by directly quantizing space itself. We compute the Bohr-Sommerfeld volume spectrum of a tetrahedron and show that it reproduces the quantization of a grain of space found in loop gravity.Comment: 4 pages, 4 figures; v2, to appear in PR

Bodily Illusions Modulate Tactile Perception

Touch differs from other exteroceptive senses in that the body itself forms part of the tactile percept. Interactions between proprioception and touch provide a powerful way to investigate the implicit body representation underlying touch. Here, we demonstrate that an intrinsic primary quality of a tactile object, for example its size, is directly affected by the perceived size of the body part touching it. We elicited proprioceptive illusions that the left index finger was either elongating or shrinking by vibrating the biceps or triceps tendon of the right arm while subjects grasped the tip of their left index finger. Subjects estimated the distance between two simultaneous tactile contacts on the left finger during tendon vibration. We found that tactile distances feel bigger when the touched body part feels elongated. Control tests showed that the modulation of touch was linked to the perceived index-finger size induced by tendon vibration. Vibrations that did not produce proprioceptive illusion had no effect on touch. Our results show that the perception of tactile objects is referenced to an implicit body representation and that proprioception contributes to this body representation. We also provide, for the first time, a quantitative, implicit measure of distortions of body size

Sensorimotor signals underlying space perception: An investigation based on self-touch

Perception of space has puzzled scientists since antiquity, and is among the foundational questions of scientific psychology. Classical “local sign” theories assert that perception of spatial extent ultimately derives from efferent signals specifying the intensity of motor commands. Everyday cases of self-touch, such as stroking the left forearm with the right index fingertip, provide an important platform for studying spatial perception, because of the tight correlation between motor and tactile extents. Nevertheless, if the motor and sensory information in self-touch were artificially decoupled, these classical theories would clearly predict that motor signals – especially if self-generated rather than passive – should influence spatial perceptual judgements, but not vice versa. We tested this hypothesis by quantifying the contribution of tactile, kinaesthetic, and motor information to judgements of spatial extent. In a self-touch paradigm involving two coupled robots in master-slave configuration, voluntary movements of the right-hand produced simultaneous tactile stroking on the left forearm. Crucially, the coupling between robots was manipulated so that tactile stimulation could be shorter, equal, or longer in extent than the movement that caused it. Participants judged either the extent of the movement, or the extent of the tactile stroke. By controlling sensorimotor gains in this way, we quantified how motor signals influence tactile spatial perception, and vice versa. Perception of tactile extent was strongly biased by the amplitude of the movement performed. Importantly, touch also affected the perceived extent of movement. Finally, the effect of movement on touch was significantly stronger when movements were actively-generated compared to when the participant's right hand was passively moved by the experimenter. Overall, these results suggest that motor signals indeed dominate the construction of spatial percepts, at least when the normal tight correlation between motor and sensory signals is broken. Importantly, however, this dominance is not total, as classical theory might suggest

Somatosensory evoked potentials that index lateral inhibition are modulated according to the mode of perceptual processing: comparing or combining multi-digit tactile motion

Many perceptual studies focus on the brain’s capacity to discriminate between stimuli. However, our normal experience of the world also involves integrating multiple stimuli into a single perceptual event. Neural mechanisms such as lateral inhibition are believed to enhance local differences between sensory inputs from nearby regions of the receptor surface. However, this mechanism would seem dysfunctional when sensory inputs need to be combined rather than contrasted. Here, we investigated whether the brain can strategically regulate the strength of suppressive interactions that underlie lateral inhibition between finger representations in human somatosensory processing. To do this, we compared sensory processing between conditions that required either comparing or combining information. We delivered two simultaneous tactile motion trajectories to index and middle fingertips of the right hand. Participants had to either compare the directions of the two stimuli, or to combine them to form their average direction. To reveal preparatory tuning of somatosensory cortex, we used an established event-related potential design to measure the interaction between cortical representations evoked by digital nerve shocks immediately before each tactile stimulus. Consistent with previous studies, we found a clear suppression between cortical activations when participants were instructed to compare the tactile motion directions. Importantly, this suppression was significantly reduced when participants had to combine the same stimuli. These findings suggest that the brain can strategically switch between a comparative and a combinative mode of somatosensory processing, according to the perceptual goal, by preparatorily adjusting the strength of a process akin to lateral inhibition

Information about action outcomes differentially affects learning from self-determined versus imposed choices

The valence of new information influences learning rates in humans: good news tends to receive more weight than bad news. We investigated this learning bias in four experiments, by systematically manipulating the source of required action (free versus forced choices), outcome contingencies (low versus high reward) and motor requirements (go versus no-go choices). Analysis of model-estimated learning rates showed that the confirmation bias in learning rates was specific to free choices, but was independent of outcome contingencies. The bias was also unaffected by the motor requirements, thus suggesting that it operates in the representational space of decisions, rather than motoric actions. Finally, model simulations revealed that learning rates estimated from the choice-confirmation model had the effect of maximizing performance across low- and high-reward environments. We therefore suggest that choice-confirmation bias may be adaptive for efficient learning of action–outcome contingencies, above and beyond fostering person-level dispositions such as self-esteem

Latent awareness: Early conscious access to motor preparation processes is linked to the readiness potential

An experience of intention to move accompanies execution of some voluntary actions. The Readiness Potential (RP) is an increasing negativity over motor brain areas prior to voluntary movement. Classical studies suggested that the RP starts before intention is consciously accessed as measured by offline recall-based reports, yet the interpretation of the RP and its temporal relation to awareness of intention remain controversial. We designed a task in which self-paced actions could be interrupted at random times by a visual cue that probed online awareness of intention. Participants were instructed to respond by pressing a key if they felt they were actively preparing a self-paced movement at the time of the cue (awareness report), but to ignore the cue otherwise. We show that an RP-like activity was more strongly present before the cue for probes eliciting awareness reports than otherwise. We further show that recall-based reports of the time of conscious intention are linked to visual attention processes, whereas online reports elicited by a probe are not. Our results suggest that awareness of intention is accessible at relatively early stages of motor preparation and that the RP is specifically associated with this conscious experience

Interhemispheric communication during haptic self-perception

During the haptic exploration of a planar surface, slight resistances against the hand's movement are illusorily perceived as asperities (bumps) in the surface. If the surface being touched is one's own skin, an actual bump would also produce increased tactile pressure from the moving finger onto the skin. We investigated how kinaesthetic and tactile signals combine to produce haptic perceptions during self-touch. Participants performed two successive movements with the right hand. A haptic force-control robot applied resistances to both movements, and participants judged which movement was felt to contain the larger bump. An additional robot delivered simultaneous but task-irrelevant tactile stroking to the left forearm. These strokes contained either increased or decreased tactile pressure synchronized with the resistance-induced illusory bump encountered by the right hand. We found that the size of bumps perceived by the right hand was enhanced by an increase in left tactile pressure, but also by a decrease. Tactile event detection was thus transferred interhemispherically, but the sign of the tactile information was not respected. Randomizing (rather than blocking) the presentation order of left tactile stimuli abolished these interhemispheric enhancement effects. Thus, interhemispheric transfer during bimanual self-touch requires a stable model of temporally synchronized events, but does not require geometric consistency between hemispheric information, nor between tactile and kinaesthetic representations of a single common object