Michela Bassolino

Extended Multisensory Space in Blind Cane Users

In the present work, we investigated whether an auditory peripersonal space exists around the hand and whether such a space might be extended by a brief tool-use experience or by long-term experience using a tool in everyday life. To this end, we studied audio-tactile integration in the space around the hand and in far space, in blind subjects who regularly used a cane to navigate and in sighted subjects, before and after brief training with the cane. In sighted subjects, auditory peripersonal space was limited to around the hand before tool use, then expanded after tool use, and contracted backward after a resting period. In contrast, in blind subjects, peri-hand space was immediately expanded when they held the cane but was limited to around the hand when they held a short handle. These results suggest that long-term experience with the cane induces a durable extension of the peripersonal space.

Everyday use of the computer mouse extends peripersonal space representation.

Auditory and tactile stimuli are integrated within a limited space around the body to form an auditory peripersonal space (APPS). Here we investigate whether the APPS representation around the hand can be extended through the use of a common technological tool such as the computer mouse. When using a mouse, an action occurring in the space around the hand has a distal effect in the space defined by the computer screen; thus, the mouse virtually links near and far space. Does prolonged experience with the mouse durably extend APPS representation to the far space? We examined 16 habitual mouse users to determine whether a sound presented near the right hand or near the computer screen affected reaction times to a tactile target at the hand. When subjects sat in front of the computer, without holding the mouse, they responded faster to tactile stimuli when sounds were presented near the hand rather than near the screen, consistent with a normal segregation of APPS around the hand. In contrast, when subjects either actively used or even passively held the mouse, the difference between the effects of near and far sounds disappeared, thus showing an extension of the APPS toward the far space. This effect was selective for the effector used to operate the mouse: if tactile stimuli were presented on the left hand, rarely used to act upon the mouse, a sound presented near the hand speeded up reactions times when subjects both held and did not hold the mouse in their left hand.

Tool-use reshapes the boundaries of body and peripersonal space representations

Interaction with objects in the environment typically requires integrating information concerning the object location with the position and size of body parts. The former information is coded in a multisensory representation of the space around the body, a representation of peripersonal space (PPS), whereas the latter is enabled by an online, constantly updated, action-orientated multisensory representation of the body (BR). Using a tool to act upon relatively distant objects extends PPS representation. This effect has been interpreted as indicating that tools can be incorporated into BR. However, empirical data showing that tool-use simultaneously affects PPS representation and BR are lacking. To study this issue, we assessed the extent of PPS representation by means of an audio-tactile interaction task and BR by means of a tactile distance perception task and a body-landmarks localisation task, before and after using a 1-m-long tool to reach far objects. Tool-use extended the representation of PPS along the tool axis and concurrently shaped BR; after tool-use, subjects perceived their forearm narrower and longer compared to before tool-use, a shape more similar to the one of the tool. Tool-use was necessary to induce these effects, since a pointing task did not affect PPS and BR. These results show that a brief training with a tool induces plastic changes both to the perceived dimensions of the body part acting upon the tool and to the space around it, suggesting a strong overlap between peripersonal space and body representation.

Use-dependent hemispheric balance.

In the human brain, homologous regions of the primary motor cortices (M1s) are connected through transcallosal fibers. Interhemispheric communication between the two M1s plays a major role in the control of unimanual hand movements, and the strength of this connection seems to be dependent on arm activity. For instance, a lesion in the M1 can induce an increase in the excitability of the intact M1 and an abnormal high inhibitory influence onto the damaged M1. This can be attributable to either the disuse of the affected limb or the overuse of the unaffected one. Here, to directly investigate cortical modifications induced by an abnormal asymmetric use of the two limbs, we studied both the excitability of the two M1s and transcallosal interaction between them in healthy subjects whose right hand was immobilized for 10 h. The left “not-immobilized” arm was completely free to move in one group of participants (G1) and limited in the other one (G2). We found that the non-use reduced the excitability of the left M1 and decreased the inhibitory influence onto the right hemisphere in the two groups. However, an increase in the excitability of right M1 and a deeper inhibitory interaction onto the left hemisphere were evident only in G1. Thus, modifications in the right M1 were not directly produced by the non-use but would depend on the overuse of the “not-immobilized” arm. Our findings suggest that the balance between the two M1s is strongly use dependent.

Training the Motor Cortex by Observing the Actions of Others During Immobilization

Limb immobilization and nonuse are well-known causes of corticomotor depression. While physical training can drive the recovery from nonuse-dependent corticomotor effects, it remains unclear if it is possible to gain access to motor cortex in alternative ways, such as through motor imagery (MI) or action observation (AO). Transcranial magnetic stimulation was used to study the excitability of the hand left motor cortex in normal subjects immediately before and after 10 h of right arm immobilization. During immobilization, subjects were requested either to imagine to act with their constrained limb or to observe hand actions performed by other individuals. A third group of control subjects watched a nature documentary presented on a computer screen. Hand corticomotor maps and recruitment curves reliably showed that AO, but not MI, prevented the corticomotor depression induced by immobilization. Our results demonstrate the existence of a visuomotor mechanism in humans that links AO and execution which is able to effect cortical plasticity in a beneficial way. This facilitation was not related to the action simulation, because it was not induced by explicit MI.

Shaping Motor Cortex Plasticity Through Proprioception

Short-term upper limb disuse induces a hemispheric unbalance between the primary motor cortices (M1s). However, it is still unclear whether these changes are mainly attributable to the absence of voluntary movements or to the reduction of proprioceptive information. The goal of this work was to investigate the role of proprioception in modulating hemispheric balance during a short-term right arm immobilization. We evaluated the 2 M1s excitability and the interhemispheric inhibition (IHI) between M1s in 3 groups of healthy subjects. Two groups received during the immobilization a proprioceptive (P-VIB, 80 Hz) and tactile (T-VIB, 30 Hz) vibration to the right hand, respectively. Another group did not receive any conditioning sensory inputs (No-VIB). We found that in the No-VIB and in the T-VIB groups immobilization induced a decrease of left M1 excitability and IHI from left to right hemisphere and an increase of right M1 excitability and IHI from right to left hemisphere. Differently, only a partial decrease in left M1 excitability, no change in right M1 excitability and in IHI was observed in the P-VIB group. Our findings demonstrate that the maintenance of dynamic proprioceptive inputs in an immobilized arm through muscle vibration can prevent the hemispheric unbalance induced by short-term limb disuse.

Dissociating effect of upper limb non-use and overuse on space and body representations

Accurate and updated representations of the space where the body acts, i.e. the peripersonal space (PPS), and the location and dimension of body parts (body representation, BR) are essential to perform actions. Because both PPS and BR are involved in motor execution and display the same plastic proprieties after the use of a tool to reach far objects, it has been suggested that they overlap in a unique representation of the body in a space devoted to action. Here we determined whether manipulating actions in space, without modifying body metrics, i.e. through immobilization, induces a dissociation of the plastic properties of PPS and BR. In 39 healthy subjects we evaluated PPS and BR for the non-used right limb and the overused left limb before and after 10 h of right arm immobilization. We observed that non-use reduces PPS representation around the immobilized arm, without affecting the metric representation (i.e. perceived length) of that limb. In contrast, overuse modulates the metric representation of the free arm, leaving PPS unchanged around that limb. These results suggest that the plasticity in PPS and BR depends on different mechanisms; while PPS representation is shaped as a function of the dimension of the acting space, metric characteristics of BR are forged on a complex interplay between visual and sensorimotor information related to the body. This behavioral dissociation between PPS and BR defines a new scenario for the role of action in shaping space and body representations.

Functional effect of short-term immobilization: Kinematic changes and recovery on reaching-to-grasp

Although previous investigations agree in showing significant cortical modifications related to short-term limb immobilization, little is known about the functional changes induced by non-use. To address this issue, we studied the kinematic effect of 10h of hand immobilization. In order to prevent any movement, right handed healthy participants wore on their dominant hand a soft bandage. They were requested to perform the same reaching-to-grasping task immediately after immobilization, 1 day before (baseline 1) and in other two following days without non-use (baseline 2 and baseline 3). While no differences were found among baseline conditions, an increase of the total duration of reaching movement together with an anticipation of the time to peak velocity were observed in the first trial after immobilization. Interestingly, these initial effects decreased quickly trial-by-trial, following an exponential function till reaching values equal to those observed in the control conditions. The present findings show firstly that the transport phase of the reaching-to-grasp task was affected by a temporary reduction of sensory and motor information. Secondly, a trial-by-trial recovery of the immobilization-related changes, likely driven by the sensory inputs and motor outputs associated to the repetition of the movement has been observed. All together these results confirm a fundamental role of a continuous stream of sensorimotor signals in maintaining motor efficiency and in driving recovery process.

Quantifying the role of motor imagery in brain-machine interfaces.

Despite technical advances in brain machine interfaces (BMI), for as-yet unknown reasons the ability to control a BMI remains limited to a subset of users. We investigate whether individual differences in BMI control based on motor imagery (MI) are related to differences in MI ability. We assessed whether differences in kinesthetic and visual MI, in the behavioral accuracy of MI, and in electroencephalographic variables, were able to differentiate between high- versus low-aptitude BMI users. High-aptitude BMI users showed higher MI accuracy as captured by subjective and behavioral measurements, pointing to a prominent role of kinesthetic rather than visual imagery. Additionally, for the first time, we applied mental chronometry, a measure quantifying the degree to which imagined and executed movements share a similar temporal profile. We also identified enhanced lateralized μ-band oscillations over sensorimotor cortices during MI in high- versus low-aptitude BMI users. These findings reveal that subjective, behavioral, and EEG measurements of MI are intimately linked to BMI control. We propose that poor BMI control cannot be ascribed only to intrinsic limitations of EEG recordings and that specific questionnaires and mental chronometry can be used as predictors of BMI performance (without the need to record EEG activity).

Activating the motor system through action observation: is this an efficient approach in adults and children?

Observing an action performed by another person to learn a new movement is a frequent experience in adult daily life, such as in sports. However, it is an especially common circumstance during the development of motor skills in childhood. Studies on healthy humans indicate that action observation induces a facilitation in the observers motor system. This effect is supported by an action‐perception matching mechanism available both in adults and in children. Because of the simplicity of action observation, it has been proposed to apply this method in clinical contexts. After a brief, non‐exhaustive introduction of the essential features underlying action observation in healthy people, we review recent studies reporting beneficial effects of rehabilitative training based on a combination of action perception and execution. We focus on therapeutic interventions for patients with upper‐limb motor disabilities such as adults after stroke or children with hemiplegia due to cerebral palsy. Further, we consider data from basic science demonstrating that the facilitation induced by visual perception of the action can be modulated by the combination of multimodal stimuli related to the movement (e.g. visual and acoustic action‐related inputs). In line with this, we discuss possible new directions to improve basic knowledge and therapeutic applications of action observation.