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Transcranial direct current stimulation (tDCS) is a non-invasive brain stimulation technique which was found to have a positive modulatory effect on online sequence acquisition or offline motor consolidation, depending on the relative role of the associated brain region. Primary motor regions (M1) and dorsolateral prefrontal cortices (DLPFC) have both been related to sequential learning. However, research so far did not systematically disentangle their differential roles in online and offline learning especially in more complex sequential paradigms. In this study, the influence of anodal M1 leg area-tDCS and anodal DLPFC-tDCS applied during complex sequential learning (online and offline) was investigated using a complex whole body serial reaction time task (CWB-SRTT) in 42 healthy volunteers. TDCS groups did not differ from sham tDCS group regarding their response and reaction time (online) and also not in terms of overnight consolidation (offline). Sequence specific learning and the number of recalled items also did not differ between groups. Results may be related to unspecific parameters such as timing of the stimulation or current intensity but can also be attributed to the relative role of M1 and DLPFC during early complex learning. Taken together, the current study provides preliminary evidence that M1 leg area or DLPFC modulation by means of tDCS does not improve complex sequential skill learning. SIGNIFICANCE STATEMENT: Understanding motor learning is helpful to deepen our knowledge about the human ability to acquire new skills. Complex sequential learning tasks have only been studied, sparsely, but are particularly mimicking challenges of daily living. The present study studied early motor learning in a complex serial reaction time task while transcranial direct current stimulation (tDCS) was either applied to leg primary motor cortex or bilateral dorsolateral prefrontal cortex. TDCS did not affect sequential learning, neither directly during performance nor in terms of sequence consolidation. Results provide preliminary information that M1 or bilateral DLPFC modulation does not improve early complex motor learning., (© 2024 The Authors. European Journal of Neuroscience published by Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

Human behavior requires inter-regional crosstalk to employ the sensorimotor processes in the brain. Although external neuromodulation techniques have been used to manipulate interhemispheric sensorimotor activity, a central controversy concerns whether this activity can be volitionally controlled. Experimental tools lack the power to up- or down-regulate the state of the targeted hemisphere over a large dynamic range and, therefore, cannot evaluate the possible volitional control of the activity. We addressed this difficulty by using the recently developed method of spatially bivariate electroencephalography (EEG)-neurofeedback to systematically enable the participants to modulate their bilateral sensorimotor activities. Here, we report that participants learn to up- and down-regulate the ipsilateral excitability to the imagined hand while maintaining constant contralateral excitability; this modulates the magnitude of interhemispheric inhibition (IHI) assessed by the paired-pulse transcranial magnetic stimulation (TMS) paradigm. Further physiological analyses revealed that the manipulation capability of IHI magnitude reflected interhemispheric connectivity in EEG and TMS, which was accompanied by intrinsic bilateral cortical oscillatory activities. Our results show an interesting approach for neuromodulation, which might identify new treatment opportunities, e.g., in patients suffering from a stroke., Competing Interests: MH is employed by LIFESCAPES Inc, KO, NM, RH, TO, MK No competing interests declared, JU is a founder and the Representative Director of the University Startup Company, LIFESCAPES Inc. involved in the research, development, and sales of rehabilitation devices including brain-computer interfaces. He receives a salary from Connect Inc., and holds shares in Connect Inc. This company does not have any relationships with the device or setup used in the current study, (© 2022, Hayashi et al.)

Background . Drastic functional reorganization was observed in the ipsilateral primary motor cortex (M1) of a Paralympic long jumper with a unilateral below-knee amputation in our previous study. However, it remains unclear whether long-term para-sports are associated with ipsilateral M1 reorganization since only 1 athlete with amputation was investigated. Objective . This study aimed to investigate the relationship between the long-term para-sports and ipsilateral M1 reorganization after lower limb amputation. Methods . Lower limb rhythmic muscle contraction tasks with functional magnetic resonance imaging and T1-weighted structural imaging were performed in 30 lower limb amputees with different para-sports experiences in the chronic phase. Results . Brain activity in the ipsilateral primary motor and somatosensory areas (SM1) as well as the contralateral dorsolateral prefrontal cortex, SM1, and inferior temporal gyrus showed a positive correlation with the years of routine para-sports participation (sports years) during contraction of the amputated knee. Indeed, twelve of the 30 participants who exhibited significant ipsilateral M1 activation during amputated knee contraction had a relatively longer history of para-sports participation. No significant correlation was found in the structural analysis. Conclusions . Long-term para-sports could lead to extensive reorganization at the brain network level, not only bilateral M1 reorganization but also reorganization of the frontal lobe and visual pathways. These results suggest that the interaction of injury-induced and use-dependent cortical plasticity might bring about drastic reorganization in lower limb amputees.

A variety of neural substrates are implicated in the initiation, coordination, and stabilization of voluntary movements underpinned by adaptive contraction and relaxation of agonist and antagonist muscles. To achieve such flexible and purposeful control of the human body, brain systems exhibit extensive modulation during the transition from resting state to motor execution and to maintain proper joint impedance. However, the neural structures contributing to such sensorimotor control under unconstrained and naturalistic conditions are not fully characterized. To elucidate which brain regions are implicated in generating and coordinating voluntary movements, we employed a physiologically inspired, two-stage method to decode relaxation and three patterns of contraction in unilateral finger muscles (i.e., extension, flexion, and co-contraction) from high-density scalp electroencephalograms (EEG). The decoder consisted of two parts employed in series. The first discriminated between relaxation and contraction. If the EEG data were discriminated as contraction, the second stage then discriminated among the three contraction patterns. Despite the difficulty in dissociating detailed contraction patterns of muscles within a limb from scalp EEG signals, the decoder performance was higher than chance-level by 2-fold in the four-class classification. Moreover, weighted features in the trained decoders revealed EEG features differentially contributing to decoding performance. During the first stage, consistent with previous reports, weighted features were localized around sensorimotor cortex (SM1) contralateral to the activated fingers, while those during the second stage were localized around ipsilateral SM1. The loci of these weighted features suggested that the coordination of unilateral finger muscles induced different signaling patterns in ipsilateral SM1 contributing to motor control. Weighted EEG features enabled a deeper understanding of human sensorimotor processing as well as of a more naturalistic control of brain-computer interfaces., Competing Interests: Declaration of Competing Interest J.U. is one of the founders and the Representative Director of the University Startup Company, Connect Inc. for the research, development, and sales of rehabilitation devices including brain-computer interface. J.U. has received a salary from Connect Inc., and has held the shares of Connect Inc. This company does not have any relationship with the present study., (Copyright © 2020 The Author(s). Published by Elsevier Inc. All rights reserved.)

The present study examined corticospinal excitability of the contralateral and ipsilateral hemispheres during actual (ACT) and imaginary (IMG) unilateral hand force-matching tasks of different difficulty. Seventeen young male adults (21.2 ± 2.2 yrs) actually and imaginarily matched their left index finger abduction force to a displayed target force. Task difficulty was manipulated by varying the acceptable force range about each mean target force (5 and 15% MVC for ACT, 15% MVC for IMG). Specifically, easy (EASY) and difficult (DIFF) tasks were assigned an acceptable force range of ±7% and ±0% of target force, respectively. Single pulse transcranial magnetic stimulation was applied to the both hemispheres in ACT and over the left hemisphere in IMG. Motor evoked potentials (MEPs) were collected from the first dorsal interosseous muscle during tasks. In ACT, MEPs in both the contracting and resting hands were significantly larger (P < 0.05) during DIFF than EASY when collapsed across target force levels. In IMG, MEPs in the resting right hand were significantly larger (P < 0.05) during DIFF than during EASY. The relative change in MEP amplitude in the right hand from EASY to DIFF in ACT was positively correlated (r = 0.63) with that in IMG. These results indicate that greater task difficulty increases corticospinal excitability of the contralateral hemisphere in ACT, and increases corticospinal excitability of the ipsilateral hemisphere in both ACT and IMG. The relative changes in corticospinal excitability of the ipsilateral hemisphere with increasing task difficulty are correlated between ACT and IMG., (Copyright © 2018 IBRO. Published by Elsevier Ltd. All rights reserved.)

We investigated corticospinal excitability during positive (execution) and negative (suppression) imageries for the right and left upper and lower limbs. In the Positive Imagery tasks, sixteen subjects were instructed to repeatedly imagine rotation of the index finger of the right or left hand, or the ankle of the right or left foot. In the Negative Imagery tasks, they were asked to imagine the suppression of movements for the index finger of the right or left hand, or the ankle of the right or left foot. A single-pulse transcranial magnetic stimulation was delivered over the left hand primary motor cortex, and motor evoked potentials (MEPs) were recorded from the right first dorsal interosseous (FDI) muscle under all conditions. The MEP amplitudes of the FDI were significantly larger in the Positive and Negative Imagery tasks than in the resting control task during motor imagery of the right hand, left hand, and left foot, but not during that of right foot. Our results indicate that imageries of suppressing hand and foot movements enhanced corticospinal excitability., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Our previous studies showed that corticospinal excitability during imagery of squeezing a foam ball was enhanced by somatosensory input generated by passively holding the ball. In the present study, using the same experimental model, we investigated whether corticospinal excitability was influenced by holding the object with the hand opposite to the imagined hand. Corticospinal excitability was assessed by monitoring motor evoked potentials (MEPs) in the first dorsal interosseous muscle following transcranial magnetic stimulation over the motor cortex during motor imagery. Subjects were asked to imagine squeezing a foam ball with the right hand (experiment 1) or the left hand (experiment 2), while either holding nothing (Null condition), a ball in the right hand (Right condition) or a ball in the left hand (Left condition). The MEPs amplitude during motor imagery was increased, only when the holding hand and the imagined hand were on the same side. These results suggest that performance improvement and rehabilitation exercises will be more effective when somatosensory stimulation and motor imagery are done on the same side., (Copyright © 2012 Elsevier Ireland Ltd. All rights reserved.)

Excitability of the corticospinal pathway increases during observation of an action. However, how corticospinal excitability changes during observation of sequential actions in the course of acquiring novel skills (observational learning) remains unexplored. To investigate this, we used a previously unpracticed sequence of ten hand postures. Participants were asked to repeat observation and replication of the sequence. This block of observation and replication was repeated 5 times. During observation of a given hand posture (OK sign), motor-evoked potentials (MEPs) elicited by transcranial magnetic stimulation were recorded from hand muscles. In experiment 1, the OK sign appeared in the 9th position of the sequence. Almost all participants could replicate the OK sign only at the 5th block of the experiment. MEP amplitude was greater than that in the control, and decreased with the stages. This suggested that during observational learning of sequential hand postures MEP changed with the progress of the learning. To evaluate this idea, we performed two additional experiments. In experiment 2, the OK sign appeared in the 2nd position. Almost all participants replicated the OK sign even in the 1st block. The MEP amplitude did not change across stages. In experiment 3, the OK sign appeared in the 9th position, but the order of other signs was randomized in every stage. Many participants were not able to replicate the OK sign even during the 5th block of the experiment. The MEP amplitude did not change across stages. These results suggest that: (1) During observational learning modulation of corticospinal excitability is associated with the learning process. (2) Corticospinal excitability decreases as learning progresses.

Excitability of the corticospinal pathway increased during observation of an action. The present study investigated whether corticospinal excitability during an action observation is modulated by physical movement executed by the observers themselves. Participants alternately repeated the observation of a pinching action with the thumb and index finger and execution (30 times) of the same action. Motor-evoked potentials (MEPs) induced by transcranial magnetic stimulation were recorded from the first dorsal interosseous muscle during observation of the action. MEP amplitude gradually increased with the number of executions of the same action. However, MEP amplitude did not change when the participants only repeated observation of the pinching action or when the participants alternately repeated observation of the pinching action and execution of the isometric abduction of the index finger. These findings suggest that enhancement of cotricospinal excitability is dependent on the execution of a particular action that is very similar to the observed action.

We investigated whether corticospinal excitability during the imagery of an action involving an external object was influenced by actually touching the object. Corticospinal excitability was assessed by monitoring motor evoked potentials (MEPs) in the first dorsal interosseous muscle following transcranial magnetic stimulation over the motor cortex during imagery of squeezing a ball-with or without passively holding the ball. The MEPs amplitude during imagery when the ball was held was larger than that when the ball was not held. The MEPs amplitude was not modulated just by holding the ball. In the same experimental condition, the somatosensory evoked potentials (SEPs) in response to the stimulation of median nerve were not modulated by motor imagery or by holding the ball. These results suggest that the corticospinal excitability during imagery of squeezing a ball is enhanced with the real touch of the ball, and the enhancement would be caused by some changes along the corticospinal pathway itself and not by the change in responsiveness along the afferent pathway to the primary somatosensory cortex.