An investigation of the effectiveness of neurofeedback training on motor performance in healthy adults: A systematic review and meta-analysis.

Neurofeedback training (NFT) refers to a training where the participants voluntarily aim to manipulate their own brain activity using the sensory feedback abstracted from their brain activity. NFT has attracted attention in the field of motor learning due to its potential as an alternative or additional training method for general physical training. In this study, a systematic review of NFT studies for motor performance improvements in healthy adults and a meta-analysis on the effectiveness of NFT were conducted. A computerized search was performed using the databases Web of Science, Scopus, PubMed, JDreamIII, and Ichushi-Web to identify relevant studies published between January 1st, 1990, and August 3rd, 2021. Thirty-three studies were identified for the qualitative synthesis and 16 randomized controlled trials (374 subjects) for the meta-analysis. The meta-analysis, including all trials found in the search, revealed significant effects of NFT for motor performance improvement examined at the timing after the last NFT session (standardized mean difference = 0.85, 95% CI [0.18-1.51]), but with the existence of publication biases and substantial heterogeneity among the trials. Subsequent meta-regression analysis demonstrated the dose-response gradient between NFTs and motor performance improvements; more than 125 min of cumulative training time may benefit for the subsequent motor performance. For each motor performance measure (e.g., speed, accuracy, and hand dexterity), the effectiveness of NFT remains inconclusive, mainly due to its small sample sizes. More empirical NFT studies for motor performance improvement may be needed to show beneficial effects on motor performance and to safely incorporate NFT into real-world scenarios.

Evidence that endpoint feedback facilitates intermanual transfer of visuomotor force learning by a cognitive strategy.

Humans continuously adapt their movement to a novel environment by recalibrating their sensorimotor system. Recent evidence, however, shows that explicit planning to compensate for external changes, i.e., a cognitive strategy, can also aid performance. If such a strategy is planned in external space, it should improve performance in an effector-independent manner. We tested this hypothesis by examining whether promoting a cognitive strategy during a visual-force adaptation task performed in one hand can facilitate learning for the opposite hand. Participants rapidly adjusted the height of visual bar on screen to a target level by isometrically exerting force on a handle using their right hand. Visuomotor gain increased during the task and participants learned the increased gain. Visual feedback was continuously provided for one group, whereas for another group only the endpoint of the force trajectory was presented. The latter has been reported to promote cognitive strategy use. We found that endpoint feedback produced stronger intermanual transfer of learning and slower response times than continuous feedback. In a separate experiment, we found evidence that aftereffects are reduced when only endpoint feedback is provided, a finding that has been consistently observed when cognitive strategies are used. The results suggest that intermanual transfer can be facilitated by a cognitive strategy. This indicates that the behavioral observation of intermanual transfer can be achieved either by forming an effector-independent motor representation or by sharing an effector-independent cognitive strategy between the hands.NEW & NOTEWORTHY The causes and consequences of cognitive strategy use are poorly understood. We tested whether a visuomotor task learned in a manner that may promote cognitive strategy use causes greater generalization across effectors. Visual feedback was manipulated to promote cognitive strategy use. Learning consistent with cognitive strategy use for one hand transferred to the unlearned hand. Our result suggests that intermanual transfer can result from a common cognitive strategy used to control both hands.

Touch inhibits touch: sanshool-induced paradoxical tingling reveals perceptual interaction between somatosensory submodalities.

Human perception of touch is mediated by inputs from multiple channels. Classical theories postulate independent contributions of each channel to each tactile feature, with little or no interaction between channels. In contrast to this view, we show that inputs from two sub-modalities of mechanical input channels interact to determine tactile perception. The flutter-range vibration channel was activated anomalously using hydroxy-α-sanshool, a bioactive compound of Szechuan pepper, which chemically induces vibration-like tingling sensations. We tested whether this tingling sensation on the lips was modulated by sustained mechanical pressure. Across four experiments, we show that sustained touch inhibits sanshool tingling sensations in a location-specific, pressure-level and time-dependent manner. Additional experiments ruled out the mediation of this interaction by nociceptive or affective (C-tactile) channels. These results reveal novel inhibitory influence from steady pressure onto flutter-range tactile perceptual channels, consistent with early-stage interactions between mechanoreceptor inputs within the somatosensory pathway.

Specificity of action selection modulates the perceived temporal order of action and sensory events.

The perceived temporal order of actions and changes in the environment is crucial for our inferences of causality. Sensory events presented shortly after an action are more likely considered as self-generated compared to the same events occurring before action execution. However, the estimation of when an action or a sensory change occurred is a challenge for the human brain. This estimation is formed from available sensory information combined with internal representations. Researchers suggested that internal signals associated with action preparation drive our awareness of initiating an action. This study aimed to directly investigate this hypothesis. Participants performed a speeded action (left or right key-press) in response to a go-signal (left or right arrow). A flash was presented at different time points around the time of the action, and participants judged whether it was simultaneous with the action or not. To investigate the role of action preparation in time perception, we compared trials where a cue indicated which action to perform in response to a later go signal presentation, and trials with a neutral cue where participants did not know until the time of the go signal which action to perform. We observed that a flash presented before the action was reported as simultaneous with the action more frequently when actions were cued than when they were uncued. This difference was not observed when the action was replaced by a tactile stimulation. These results indicate that precued actions are experienced earlier in time compared to unprepared actions. Further, this difference is not due to mere non-motor expectation of an event. The experience of initiating an action is driven by action preparation process: when we know what to do, actions are perceived ahead of time.

Functional Organization of Flash-Induced V1 Offline Reactivation.

The primary visual cortex exhibits a late, long response with a latency of >300 ms and an immediate early response that occurs ∼100 ms after a visual stimulus. The late response is thought to contribute to visual functions such as sensory perception, iconic memory, working memory, and forming connections between temporally separated stimuli. However, how the visual late response is generated and organized is not completely understood. In the mouse primary visual cortex in vivo, we isolated long-delayed responses by using a brief light-flash stimulus for which the stimulus late response occurred long after the stimulus offset and was not contaminated by the instantaneous response evoked by the stimulus. Using whole-cell patch-clamp recordings, we demonstrated that the late rebound response was shaped by a net-balanced increase in excitatory and inhibitory synaptic conductances, whereas transient imbalances were caused by intermittent inhibitory barrage. In contrast to the common assumption that the neocortical late response reflects a feedback signal from the downstream higher-order cortical areas, our pharmacological and optogenetic analyses demonstrated that the late responses likely have a thalamic origin. Therefore, the late component of a sensory-evoked cortical response should be interpreted with caution. SIGNIFICANCE STATEMENT: The long-delayed responses of neocortical neurons are thought to arise from cortical feedback activity that is related to sensory perception and cognition. The mechanism of neocortical late responses was investigated using multiple electrophysiological techniques and the findings indicate that it actually arises from the thalamus. In addition, during the late response, excitation and inhibition are balanced, but inhibition is dominant in patterning action potentials.

Constructing Visual Perception of Body Movement with the Motor Cortex.

The human brain readily perceives fluent movement from static input. Using functional magnetic resonance imaging, we investigated brain mechanisms that mediate fluent apparent biological motion (ABM) perception from sequences of body postures. We presented body and nonbody stimuli varying in objective sequence duration and fluency of apparent movement. Three body postures were ordered to produce a fluent (ABC) or a nonfluent (ACB) apparent movement. This enabled us to identify brain areas involved in the perceptual reconstruction of body movement from identical lower-level static input. Participants judged the duration of a rectangle containing body/nonbody sequences, as an implicit measure of movement fluency. For body stimuli, fluent apparent motion sequences produced subjectively longer durations than nonfluent sequences of the same objective duration. This difference was reduced for nonbody stimuli. This body-specific bias in duration perception was associated with increased blood oxygen level-dependent responses in the primary (M1) and supplementary motor areas. Moreover, fluent ABM was associated with increased functional connectivity between M1/SMA and right fusiform body area. We show that perceptual reconstruction of fluent movement from static body postures does not merely enlist areas traditionally associated with visual body processing, but involves cooperative recruitment of motor areas, consistent with a "motor way of seeing".

Decoding sequential finger movements from preparatory activity in higher-order motor regions: a functional magnetic resonance imaging multi-voxel pattern analysis.

Performing a complex sequential finger movement requires the temporally well-ordered organization of individual finger movements. Previous behavioural studies have suggested that the brain prepares a whole sequence of movements as a single set, rather than the movements of individual fingers. However, direct neuroimaging support for this hypothesis is lacking and, assuming it to be true, it remains unclear which brain regions represent the information of a prepared sequence. Here, we measured brain activity with functional magnetic resonance imaging while 14 right-handed healthy participants performed two types of well-learned sequential finger movements with their right hands. Using multi-voxel pattern analysis, we examined whether the types of the forthcoming sequence could be predicted from the preparatory activities of nine regions of interest, which included the motor, somatosensory and posterior parietal regions in each hemisphere, bilateral visual cortices, cerebellum and basal ganglia. We found that, during preparation, the activity of the contralateral motor regions could predict which of the two sequences would be executed. Further detailed analysis revealed that the contralateral dorsal premotor cortex and supplementary motor area were the key areas that contributed to the prediction consistently across participants. These contrasted with results from execution-related brain activity where a performed sequence was successfully predicted from the activities in the broad cortical sensory-motor network, including the bilateral motor, parietal and ipsilateral somatosensory cortices. Our study supports the hypothesis that temporary well-organized sequences of movements are represented as a set in the brain, and that preparatory activity in higher-order motor regions represents information about upcoming motor actions.

Decision uncertainty as a context for motor memory.

The current view of perceptual decision-making suggests that once a decision is made, only a single motor programme associated with the decision is carried out, irrespective of the uncertainty involved in decision making. In contrast, we show that multiple motor programmes can be acquired on the basis of the preceding uncertainty of the decision, indicating that decision uncertainty functions as a contextual cue for motor memory. The actions learned after making certain (uncertain) decisions are only partially transferred to uncertain (certain) decisions. Participants were able to form distinct motor memories for the same movement on the basis of the preceding decision uncertainty. Crucially, this contextual effect generalizes to novel stimuli with matched uncertainty levels, demonstrating that decision uncertainty is itself a contextual cue. These findings broaden the understanding of contextual inference in motor memory, emphasizing that it extends beyond direct motor control cues to encompass the decision-making process.

Food vibrations: Asian spice sets lips trembling.

Szechuan pepper, a widely used ingredient in the cuisine of many Asian countries, is known for the tingling sensation it induces on the tongue and lips. While the molecular mechanism by which Szechuan pepper activates tactile afferent fibres has been clarified, the tingling sensation itself has been less studied, and it remains unclear which fibres are responsible. We investigated the somatosensory perception of tingling in humans to identify the characteristic temporal frequency and compare this to the established selectivity of tactile afferents. Szechuan pepper was applied to the lower lip of participants. Participants judged the frequency of the tingling sensation on the lips by comparing this with the frequencies of mechanical vibrations applied to their right index finger. The perceived frequency of the tingling was consistently at around 50 Hz, corresponding to the range of tactile RA1 afferent fibres. Furthermore, adaptation of the RA1 channel by prolonged mechanical vibration reliably reduced the tingling frequency induced by Szechuan pepper, confirming that the frequency-specific tactile channel is shared between Szechuan pepper and mechanical vibration. Combining information about molecular reactions at peripheral receptors with quantitative psychophysical measurement may provide a unique method for characterizing unusual experiences by decomposing them into identifiable minimal units of sensation.

Learning to like it: aesthetic perception of bodies, movements and choreographic structure.

Appreciating human movement can be a powerful aesthetic experience. We have used apparent biological motion to investigate the aesthetic effects of three levels of movement representation: body postures, movement transitions and choreographic structure. Symmetrical (ABCDCBA) and asymmetrical (ABCDBCA) sequences of apparent movement were created from static postures, and were presented in an artificial grammar learning paradigm. Additionally, "good" continuation of apparent movements was manipulated by changing the number of movement path reversals within a sequence. In an initial exposure phase, one group of participants saw only symmetrical sequences, while another group saw only asymmetrical sequences. In a subsequent test phase, both groups rated all sequences on an aesthetic evaluation scale. We found that posture, movement, and choreographic structure all influenced aesthetic ratings. Separate ratings for the static body postures presented individually showed that both groups preferred a posture that maximized spatial symmetry. Ratings for the experimental sequences showed that both groups gave higher ratings to symmetrical sequences with "good" continuation and lower ratings to sequences with many path reversals. Further, participants who had been initially familiarized with asymmetrical sequences showed increased liking for asymmetrical sequences, suggesting a structural mere exposure effect. Aesthetic preferences thus depend on body postures, apparent movement continuation and choreographic structure. We propose a hierarchical model of aesthetic perception of human movement with distinct processing levels for body postures, movements and choreographic structure.

Does decision confidence reflect effort?

Goal directed behaviour requires transformation of sensory input to decision, and then to output action. How the sensory input is accumulated to form the decision has been extensively studied, however, the influence of output action on decision making has been largely dismissed. Although the recent emerging view postulates the reciprocal interaction between action and decision, still little is known about how the parameters of an action modulates the decision. In this study, we focused on the physical effort which necessarily entails with action. We tested if the physical effort during the deliberation period of the perceptual decision, not the effort required after deciding a particular option, can impact on the process of forming the decision. Here, we set up an experimental situation where investing effort is necessary for the initiation of the task, but importantly, is orthogonal to success in task performance. The study was pre-registered to test the hypothesis that the increased effort will decrease the metacognitive accuracy of decision, without affecting the decision accuracy. Participants judged the direction of a random-dot motion stimuli, while holding and maintaining the position of a robotic manipulandum with their right hand. In the key experimental condition, the manipulandum produced force to move away from its position, requiring the participants to resist the force while accumulating the sensory evidence for the decision. The decision was reported by a key-press using the left-hand. We found no evidence that such incidental (i.e., non-instrumental) effort may influence the subsequent decision process and most importantly decision confidence. The possible reason for this result and the future direction of the research are discussed.

Ready steady slow: action preparation slows the subjective passage of time.

Professional ball game players report the feeling of the ball 'slowing-down' before hitting it. Because effective motor preparation is critical in achieving such expert motor performance, these anecdotal comments imply that the subjective passage of time may be influenced by preparation for action. Previous reports of temporal illusions associated with action generally emphasize compensation for suppressed sensory signals that accompany motor commands. Here, we show that the time is perceived slowed-down during preparation of a ballistic reaching movement before action, involving enhancement of sensory processing. Preparing for a reaching movement increased perceived duration of a visual stimulus. This effect was tightly linked to action preparation, because the amount of temporal dilation increased with the information about the upcoming movement. Furthermore, we showed a reduction of perceived frequency for flickering stimuli and an enhanced detection of rapidly presented letters during action preparation, suggesting increased temporal resolution of visual perception during action preparation. We propose that the temporal dilation during action preparation reflects the function of the brain to maximize the capacity of sensory information-acquisition prior to execution of a ballistic movement. This strategy might facilitate changing or inhibiting the planned action in response to last-minute changes in the external environment.

Am I seeing my hand? Visual appearance and knowledge of controllability both contribute to the visual capture of a person's own body.

When confronted with complex visual scenes in daily life, how do we know which visual information represents our own hand? We investigated the cues used to assign visual information to one's own hand. Wrist tendon vibration elicits an illusory sensation of wrist movement. The intensity of this illusion attenuates when the actual motionless hand is visually presented. Testing what kind of visual stimuli attenuate this illusion will elucidate factors contributing to visual detection of one's own hand. The illusion was reduced when a stationary object was shown, but only when participants knew it was controllable with their hands. In contrast, the visual image of their own hand attenuated the illusion even when participants knew that it was not controllable. We suggest that long-term knowledge about the appearance of the body and short-term knowledge about controllability of a visual object are combined to robustly extract our own body from a visual scene.

Consensus paper: roles of the cerebellum in motor control--the diversity of ideas on cerebellar involvement in movement.

Considerable progress has been made in developing models of cerebellar function in sensorimotor control, as well as in identifying key problems that are the focus of current investigation. In this consensus paper, we discuss the literature on the role of the cerebellar circuitry in motor control, bringing together a range of different viewpoints. The following topics are covered: oculomotor control, classical conditioning (evidence in animals and in humans), cerebellar control of motor speech, control of grip forces, control of voluntary limb movements, timing, sensorimotor synchronization, control of corticomotor excitability, control of movement-related sensory data acquisition, cerebro-cerebellar interaction in visuokinesthetic perception of hand movement, functional neuroimaging studies, and magnetoencephalographic mapping of cortico-cerebellar dynamics. While the field has yet to reach a consensus on the precise role played by the cerebellum in movement control, the literature has witnessed the emergence of broad proposals that address cerebellar function at multiple levels of analysis. This paper highlights the diversity of current opinion, providing a framework for debate and discussion on the role of this quintessential vertebrate structure.

Importance of precentral motor regions in human kinesthesia: a single case study.

Prompted by our neuroimaging findings in 60 normal people, we examined whether focal damage to the hand section of precentral motor regions impairs hand kinesthesia in a patient, and investigated brain regions related to recovery of kinesthetic function. The damage impaired contralateral kinesthesia. The peri-lesional cerebral motor region, together with the ipsilateral intermediate cerebellum, participated in the recovered kinesthetic processing. The study confirmed the importance of precentral motor regions in human kinesthesia, and indicated a contribution of the peri-lesional cerebral region in recovered kinesthesia after precentral damage, which conceptually fits with cases of recovery of motor function.

Visuokinesthetic perception of hand movement is mediated by cerebro-cerebellar interaction between the left cerebellum and right parietal cortex.

Combination of visual and kinesthetic information is essential to perceive bodily movements. We conducted behavioral and functional magnetic resonance imaging experiments to investigate the neuronal correlates of visuokinesthetic combination in perception of hand movement. Participants experienced illusory flexion movement of their hand elicited by tendon vibration while they viewed video-recorded flexion (congruent: CONG) or extension (incongruent: INCONG) motions of their hand. The amount of illusory experience was graded by the visual velocities only when visual information regarding hand motion was concordant with kinesthetic information (CONG). The left posterolateral cerebellum was specifically recruited under the CONG, and this left cerebellar activation was consistent for both left and right hands. The left cerebellar activity reflected the participants' intensity of illusory hand movement under the CONG, and we further showed that coupling of activity between the left cerebellum and the "right" parietal cortex emerges during this visuokinesthetic combination/perception. The "left" cerebellum, working with the anatomically connected high-order bodily region of the "right" parietal cortex, participates in online combination of exteroceptive (vision) and interoceptive (kinesthesia) information to perceive hand movement. The cerebro-cerebellar interaction may underlie updating of one's "body image," when perceiving bodily movement from visual and kinesthetic information.

Activity in the posterior parietal cortex mediates visual dominance over kinesthesia.

When both visual and kinesthetic information of a limb are available, vision is usually the dominant source of information used to perceive the spatial location. In this study, we conducted behavioral and functional magnetic resonance imaging (fMRI) experiments to examine the brain mechanisms underlying the visual dominance over kinesthesia in perceiving the position of a hand. We used tendon vibration to induce an illusory percept of flexion movement of an immobile hand, while the participants viewed a live image of either the vibrated or nonvibrated static hand through an on-line video camera. The intensity of illusory movement was significantly attenuated (for both the left and right hands) only when the participants viewed the static image of the vibrated hand. The fMRI study showed that the posterior parietal cortex (PPC) is specifically involved in the attenuation of illusory movement and that the activity of the PPC was associated with the degree of attenuation. This indicates that PPC is involved in the multisensory processing that occurs when vision overrules simultaneously available kinesthetic information for estimating the spatial location of a limb. It is thus suggested that the human parietal cortex may play a critical role in the maintenance of a coherent body image when the brain receives potentially conflicting multisensory information from the body.

Perceptual decisions are biased by the cost to act.

Perceptual decisions are classically thought to depend mainly on the stimulus characteristics, probability and associated reward. However, in many cases, the motor response is considered to be a neutral output channel that only reflects the upstream decision. Contrary to this view, we show that perceptual decisions can be recursively influenced by the physical resistance applied to the response. When participants reported the direction of the visual motion by left or right manual reaching movement with different resistances, their reports were biased towards the direction associated with less effortful option. Repeated exposure to such resistance on hand during perceptual judgements also biased subsequent judgements using voice, indicating that effector-dependent motor costs not only biases the report at the stage of motor response, but also changed how the sensory inputs are transformed into decisions. This demonstrates that the cost to act can influence our decisions beyond the context of the specific action.

Sanshool on The Fingertip Interferes with Vibration Detection in a Rapidly-Adapting (RA) Tactile Channel.

An Asian spice, Szechuan pepper (sanshool), is well known for the tingling sensation it induces on the mouth and on the lips. Electrophysiological studies have revealed that its active ingredient can induce firing of mechanoreceptor fibres that typically respond to mechanical vibration. Moreover, a human behavioral study has reported that the perceived frequency of sanshool-induced tingling matches with the preferred frequency range of the tactile rapidly adapting (RA) channel, suggesting the contribution of sanshool-induced RA channel firing to its unique perceptual experience. However, since the RA channel may not be the only channel activated by sanshool, there could be a possibility that the sanshool tingling percept may be caused in whole or in part by other sensory channels. Here, by using a perceptual interference paradigm, we show that the sanshool-induced RA input indeed contributes to the human tactile processing. The absolute detection thresholds for vibrotactile input were measured with and without sanshool application on the fingertip. Sanshool significantly impaired detection of vibrations at 30 Hz (RA channel dominant frequency), but did not impair detection of higher frequency vibrations at 240 Hz (Pacinian-corpuscle (PC) channel dominant frequency) or lower frequency vibrations at 1 Hz (slowly adapting 1 (SA1) channel dominant frequency). These results show that the sanshool induces a peripheral RA channel activation that is relevant for tactile perception. This anomalous activation of RA channels may contribute to the unique tingling experience of sanshool.