Differential magnitude coding of gains and omitted rewards in the ventral striatum.

Physiologic studies revealed that neurons in the dopaminergic midbrain of non-human primates encode reward prediction errors. It was furthermore shown that reward prediction errors are adaptively scaled with respect to the range of possible outcomes, enabling sensitive encoding for a large range of reward values. Congruently, neuroimaging studies in humans demonstrated that BOLD-responses in the ventral striatum encode reward prediction errors in similar fashion as dopaminergic midbrain neurons, suggesting that these BOLD-responses may be driven by dopaminergic midbrain activity. However, neuroimaging results are ambiguous with respect to the adaptive scaling of reward prediction errors, leading to the conjecture that under certain circumstances other than dopaminergic midbrain input may drive ventral striatal BOLD-responses. The goal of this study was to substantiate whether BOLD-responses in the ventral striatum rather respond to adaptively scaled reward prediction errors or absolute reward magnitude. In addition, we aimed to identify neuronal structures modulating activity in the ventral striatum. Sixteen healthy participants played a wheel of fortune game, where they could win three differently valued rewards while being scanned. BOLD-responses increased after gaining rewards; this gain was however independent of the absolute reward magnitude. In contrast BOLD-responses upon reward omission decreased with reward magnitude. A psychophysiological interaction analysis identified a cluster in the brainstem in proximity of the dorsal raphe nucleus, a cluster in the lateral orbitofrontal cortex, and a cluster in the rostral cingulate zone. These clusters changed their connectivity with the ventral striatum in relation to the absolute reward magnitude in reward omission trials.

Virtual reality for enhancement of robot-assisted gait training in children with central gait disorders.

OBJECTIVE: To examine the effect of various forms of training interventions, with and without virtual reality, on the initiation and maintenance of active participation during robot-assisted gait training. DESIGN: Intervention study at the Rehabilitation Centre Affoltern a. A., University Children's Hospital, Zurich. SUBJECTS: Ten patients (5 males, mean age 12.47 years, standard deviation 1.84 years) with different neurological gait disorders and 14 healthy children (7 males, mean age 11.76 years, standard deviation 2.75 years). METHODS: All participants walked in the driven gait orthosis Lokomat® in 4 different randomly-assigned conditions. Biofeedback values calculated during swing phases were the primary outcome measure and secondary outcomes were derived from a questionnaire assessing the participant's motivation. RESULTS: Findings revealed a significant main effect for training condition in all participants (p < 0.001), for patients (p < 0.05) and for healthy controls (p < 0.01). Overall, both virtual reality-assisted therapy approaches were equally the most effective in initiating the desired active participation in all children, compared with conventional training conditions. Motivation was very high and differed between the groups only in the virtual navigation condition. CONCLUSION: Novel virtual reality-based training conditions represent a valuable approach to enhance active participation during robot-assisted gait training in patients and healthy controls.

Influence of virtual reality soccer game on walking performance in robotic assisted gait training for children.

BACKGROUND: Virtual reality (VR) offers powerful therapy options within a functional, purposeful and motivating context. Several studies have shown that patients' motivation plays a crucial role in determining therapy outcome. However, few studies have demonstrated the potential of VR in pediatric rehabilitation. Therefore, we developed a VR-based soccer scenario, which provided interactive elements to engage patients during robotic assisted treadmill training (RAGT). The aim of this study was to compare the immediate effect of different supportive conditions (VR versus non-VR conditions) on motor output in patients and healthy control children during training with the driven gait orthosis Lokomat*. METHODS: A total of 18 children (ten patients with different neurological gait disorders, eight healthy controls) took part in this study. They were instructed to walk on the Lokomat in four different, randomly-presented conditions: (1) walk normally without supporting assistance, (2) with therapists' instructions to promote active participation, (3) with VR as a motivating tool to walk actively and (4) with the VR tool combined with therapists' instructions. The Lokomat gait orthosis is equipped with sensors at hip and knee joint to measure man-machine interaction forces. Additionally, subjects' acceptance of the RAGT with VR was assessed using a questionnaire. RESULTS: The mixed ANOVA revealed significant main effects for the factor CONDITIONS (p < 0.001) and a significant interaction CONDITIONS x GROUP (p = 0.01). Tests of between-subjects effects showed no significant main effect for the GROUP (p = 0.592). Active participation in patients and control children increased significantly when supported and motivated either by therapists' instructions or by a VR scenario compared with the baseline measurement "normal walking" (p < 0.001). CONCLUSIONS: The VR scenario used here induces an immediate effect on motor output to a similar degree as the effect resulting from verbal instructions by the therapists. Further research needs to focus on the implementation of interactive design elements, which keep motivation high across and beyond RAGT sessions, especially in pediatric rehabilitation.

The effects of practice distribution upon the regional oscillatory activity in visuomotor learning.

BACKGROUND: The aim of this study was to investigate the effects of a massed compared to a distributed practice upon visuomotor learning as well as upon the regional oscillatory activity in the sensorimotor cortex. METHODS: A continuous visuomotor tracking task was used to assess visuomotor learning; the underlying neuronal correlates were measured by means of EEG. The massed practice group completed a continuous training of 60 minutes, while the distributed practice group completed four 15 minutes practice blocks separated by rest intervals. RESULTS: While the massed and the distributed practice group did not differ in performance, effects of practice distribution were evident in the regional oscillatory activity. In the course of practice, the massed training group showed a higher task-related theta power and a strong task-related power decrease in the upper alpha frequency over the sensorimotor cortex compared to the distributed practice group. CONCLUSIONS: These differences in the regional oscillatory activity indicate a higher cognitive effort and higher attention demands in the massed practice group. The results of this study support the hypothesis, that a distributed practice is superior to a massed practice in visuomotor learning.

Structural neuroplasticity in the sensorimotor network of professional female ballet dancers.

Evidence suggests that motor, sensory, and cognitive training modulates brain structures involved in a specific practice. Functional neuroimaging revealed key brain structures involved in dancing such as the putamen and the premotor cortex. Intensive ballet dance training was expected to modulate the structures of the sensorimotor network, for example, the putamen, premotor cortex, supplementary motor area (SMA), and the corticospinal tracts. We investigated gray (GM) and white matter (WM) volumes, fractional anisotropy (FA), and mean diffusivity (MD) using magnetic resonance-based morphometry and diffusion tensor imaging in 10 professional female ballet dancers compared with 10 nondancers. In dancers compared with nondancers, decreased GM volumes were observed in the left premotor cortex, SMA, putamen, and superior frontal gyrus, and decreased WM volumes in both corticospinal tracts, both internal capsules, corpus callosum, and left anterior cingulum. FA was lower in the WM underlying the dancers' left and right premotor cortex. There were no significant differences in MD between the groups. Age of dance commencement was negatively correlated with GM and WM volume in the right premotor cortex and internal capsule, respectively, and positively correlated with WM volume in the left precentral gyrus and corpus callosum. Results were not influenced by the significantly lower body mass index of the dancers. The present findings complement the results of functional imaging studies in experts that revealed reduced neural activity in skilled compared with nonskilled subjects. Reductions in brain activity are accompanied by local decreases in GM and WM volumes and decreased FA.

Transfer effects of practice for simple alternating movements.

In studies on transfer of practice effects, researchers use simple or complex movements that involve a significant cognitive element. In the present study, the authors studied intermanual and intramanual transfer of practice with a task that can be considered intermediate in difficulty. Using finger tapping as a motor task, 30 participants practiced tapping 6 days per week for 2 weeks with the left or right middle finger in a between-subject design. Compared with controls, the unpracticed middle finger of both hands showed significant improvement, along with all of the other unpracticed digits. There was no significant difference in the strength of transfer from the practiced finger to other fingers of the same (intramanual transfer) or the other (intermanual transfer) hand. The authors did not observe an asymmetry of transfer effects (the degree to which transfer depends on the particular hand trained). Last, in terms of speed and regularity of movement, the digits broke down into 2 different clusters; the thumb, index finger, and middle finger formed 1 cluster superior to that formed by the ring and small fingers.

The architecture of the golfer's brain.

BACKGROUND: Several recent studies have shown practice-dependent structural alterations in humans. Cross-sectional studies of intensive practice of specific tasks suggest associated long-term structural adaptations. Playing golf at a high level of performance is one of the most demanding sporting activities. In this study, we report the relationship between a particular level of proficiency in playing golf (indicated by golf handicap level) and specific neuroanatomical features. PRINCIPAL FINDINGS: Using voxel-based morphometry (VBM) of grey (GM) and white matter (WM) volumes and fractional anisotropy (FA) measures of the fibre tracts, we identified differences between skilled (professional golfers and golfers with an handicap from 1-14) and less-skilled golfers (golfers with an handicap from 15-36 and non-golfer). Larger GM volumes were found in skilled golfers in a fronto-parietal network including premotor and parietal areas. Skilled golfers revealed smaller WM volume and FA values in the vicinity of the corticospinal tract at the level of the internal and external capsule and in the parietal operculum. However, there was no structural difference within the skilled and less-skilled golfer group. CONCLUSION: There is no linear relationship between the anatomical findings and handicap level, amount of practice, and practice hours per year. There was however a strong difference between highly-practiced golfers (at least 800-3,000 hours) and those who have practised less or non-golfers without any golfing practise, thus indicating a step-wise structural and not a linear change.

Individual preferences modulate incentive values: Evidence from functional MRI.

BACKGROUND: In most studies on human reward processing, reward intensity has been manipulated on an objective scale (e.g., varying monetary value). Everyday experience, however, teaches us that objectively equivalent rewards may differ substantially in their subjective incentive values. One factor influencing incentive value in humans is branding. The current study explores the hypothesis that individual brand preferences modulate activity in reward areas similarly to objectively measurable differences in reward intensity. METHODS: A wheel-of-fortune game comprising an anticipation phase and a subsequent outcome evaluation phase was implemented. Inside a 3 Tesla MRI scanner, 19 participants played for chocolate bars of three different brands that differed in subjective attractiveness. RESULTS: Parametrical analysis of the obtained fMRI data demonstrated that the level of activity in anatomically distinct neural networks was linearly associated with the subjective preference hierarchy of the brands played for. During the anticipation phases, preference-dependent neural activity has been registered in premotor areas, insular cortex, orbitofrontal cortex, and in the midbrain. During the outcome phases, neural activity in the caudate nucleus, precuneus, lingual gyrus, cerebellum, and in the pallidum was influenced by individual preference. CONCLUSION: Our results suggest a graded effect of differently preferred brands onto the incentive value of objectively equivalent rewards. Regarding the anticipation phase, the results reflect an intensified state of wanting that facilitates action preparation when the participants play for their favorite brand. This mechanism may underlie approach behavior in real-life choice situations.

Brain stimulation modulates driving behavior.

BACKGROUND: Driving a car is a complex task requiring coordinated functioning of distributed brain regions. Controlled and safe driving depends on the integrity of the dorsolateral prefrontal cortex (DLPFC), a brain region, which has been shown to mature in late adolescence. METHODS: In this study, driving performance of twenty-four male participants was tested in a high-end driving simulator before and after the application of transcranial direct current stimulation (tDCS) for 15 minutes over the left or right DLPFC. RESULTS: We show that external modulation of both, the left and the right, DLPFC directly influences driving behavior. Excitation of the DLPFC (by applying anodal tDCS) leads to a more careful driving style in virtual scenarios without the participants noticing changes in their behavior. CONCLUSION: This study is one of the first to prove that external stimulation of a specific brain area can influence a multi-part behavior in a very complex and everyday-life situation, therefore breaking new ground for therapy at a neural level.

Chronometric features of processing unpleasant stimuli: a functional MRI-based transcranial magnetic stimulation study.

The quick identification of potentially threatening events is a crucial cognitive capacity to survive in a changing environment. Previous functional MRI data revealed the right dorsolateral prefrontal cortex and the region of the left intraparietal sulcus (IPS) to be involved in the perception of emotionally negative stimuli. For assessing chronometric aspects of emotion processing, we applied transcranial magnetic stimulation above these areas at different times after negative and neutral picture presentation. An interference with emotion processing was found with transcranial magnetic stimulation above the dorsolateral prefrontal cortex 200-300 ms and above the left intraparietal sulcus 240/260 ms after negative stimuli. The data suggest a parallel and conjoint involvement of prefrontal and parietal areas for the identification of emotionally negative stimuli.

Different strategies do not moderate primary motor cortex involvement in mental rotation: a TMS study.

BACKGROUND: Regions of the dorsal visual stream are known to play an essential role during the process of mental rotation. The functional role of the primary motor cortex (M1) in mental rotation is however less clear. It has been suggested that the strategy used to mentally rotate objects determines M1 involvement. Based on the strategy hypothesis that distinguishes between an internal and an external strategy, our study was designed to specifically test the relation between strategy and M1 activity. METHODS: Twenty-two subjects were asked to participate in a standard mental rotation task. We used specific picture stimuli that were supposed to trigger either the internal (e.g. pictures of hands or tools) or the external strategy (e.g. pictures of houses or abstract figures). The strategy hypothesis predicts an involvement of M1 only in case of stimuli triggering the internal strategy (imagine grasping and rotating the object by oneself). Single-pulse Transcranial Magnetic Stimulation (TMS) was employed to quantify M1 activity during task performance by measuring Motor Evoked Potentials (MEPs) at the right hand muscle. RESULTS: Contrary to the strategy hypothesis, we found no interaction between stimulus category and corticospinal excitability. Instead, corticospinal excitability was generally increased compared with a resting baseline although subjects indicated more frequent use of the external strategy for all object categories. CONCLUSION: This finding suggests that M1 involvement is not exclusively linked with the use of the internal strategy but rather directly with the process of mental rotation. Alternatively, our results might support the hypothesis that M1 is active due to a 'spill-over' effect from adjacent brain regions.

A network for audio-motor coordination in skilled pianists and non-musicians.

Playing a musical instrument requires efficient auditory and motor processing. Fast feed forward and feedback connections that link the acoustic target to the corresponding motor programs need to be established during years of practice. The aim of our study is to provide a detailed description of cortical structures that participate in this audio-motor coordination network in professional pianists and non-musicians. In order to map these interacting areas using functional magnetic resonance imaging (fMRI), we considered cortical areas that are concurrently activated during silent piano performance and motionless listening to piano sound. Furthermore we investigated to what extent interactions between the auditory and the motor modality happen involuntarily. We observed a network of predominantly secondary and higher order areas belonging to the auditory and motor modality. The extent of activity was clearly increased by imagination of the absent modality. However, this network did neither comprise primary auditory nor primary motor areas in any condition. Activity in the lateral dorsal premotor cortex (PMd) and the pre-supplementary motor cortex (preSMA) was significantly increased for pianists. Our data imply an intermodal transformation network of auditory and motor areas which is subject to a certain degree of plasticity by means of intensive training.

Converging evidence of ERD/ERS and BOLD responses in motor control research.

In this chapter we summarize findings of our group in which we studied the neural underpinnings of finger tapping control using different methods (functional magnetic resonance imaging: fMRI, electroencephalography: EEG, transcranial magnetic stimulation: TMS, and behavioural experiments). First, we found that maximum finger tapping speed is a matter of training as shown for professional musicians. Secondly, we demonstrated that different finger tapping speeds are accompanied by different hemodynamic responses in the primary hand motor area (M1), the cerebellum and partly in pre-motor areas. With increasing tapping speed there is an increase of hemodynamic response in these areas (rate effect). Thirdly, the effect measured with fMRI is substantiated by rate effects measured by means of task-related power decreases in the upper alpha-band (10-12 Hz) over the primary motor cortex. In case of sequential finger movement learning, we observed decreases in task-related alpha-power in lateral PMC (event-related desynchronization: ERD) and simultaneous alpha-power increases in SMA (event-related synchronization: ERS) that came along with training-induced increases in movement rate. This pattern is discussed in relation to the "focal ERD/surround ERS" phenomenon suggested by Pfurtscheller and Lopes da Silva. Finally, we demonstrated that finger tapping speed was slowed by selectively inhibiting the primary hand motor area using TMS. Taken together, these studies demonstrate on the basis of converging evidence that the primary hand motor area is the basic control centre for controlling the movement parameter tapping speed. However, the neural efficiency to control finger tapping speed (as measured with hemodynamic responses or ERD/ERS patterns) is a matter of training.

How finger tapping practice enhances efficiency of motor control.

Maximum-speed movements have been suggested to put maximum neural control demands on the primary motor cortex; hence, we are asking how primary motor cortex function changes to enable enhanced maximum movement rates induced by long-lasting practice. Cortical function was assessed by recording task-related spectral electroencephalogram alpha-power. Low-resolution brain electromagnetic tomography was used to localize intracortical neuronal sources. The main result is a decrease in neural activity in the left hemisphere (ipsilateral to trained hand) from pretraining to posttraining, whereas right hemispheric activity remained constant across training. This likely reflects the initially limited capacity of the right hemisphere to control demanding left-hand movements, but also highlights its ability to become more efficient with training, indicated by reduced involvement of the left primary motor cortex after training.

Neural control of playing a reversed piano: empirical evidence for an unusual cortical organization of musical functions.

Using functional magnetic imaging techniques and neuropsychological tests, we studied a young male musician (C.S.) who performs at a professional level both on a regular piano keyboard and on a reverse keyboard (reversed right to left). The participant was left-handed, had left dominance for language but, remarkably, right dominance for the control of piano playing on both keyboards. With respect to music perception, C.S. showed left-sided activation dominance within the left superior temporal sulcus, which is normally associated with higher order auditory processing and right-sided activations in the secondary sensory cortex extending into the supramarginal gyrus. We suggest that C.S.'s pattern of functional asymmetry, characterized by audio-motor control using a right-sided network, could be a factor in his exceptional piano-playing ability on both the standard and reversed keyboard.

A network for sensory-motor integration: what happens in the auditory cortex during piano playing without acoustic feedback?

Playing a musical instrument requires efficient auditory as well as motor processing. We provide evidence for the existence of a neuronal network of secondary and higher-order areas belonging to the auditory and motor modality that is important in the integration of auditory and motor domains.

Bimanual versus unimanual coordination: what makes the difference?

Using fMRI, we investigated the neuronal structures controlling bimanual coordination applying a visuomotor coordination task. Recent studies suggest the existence of a widespread network for the neuronal control of bimanual coordination including primary sensorimotor cortices (M1/S1), lateral and medial premotor cortices (PMC, SMA), cingulate motor area (CMA), and cerebellum (CB). In the present study, subjects performed bimanual and unimanual tasks requiring the coordination of two fingers at a time to navigate a cursor on a computer screen. Thus, in contrast to previous studies, we are using appropriate unimanual control (UNI) tasks. By using this new motor task, we identified a similar activation network for uni- and bimanual movements. Subjects exhibited bilateral activations in PMC, SMA, posterior-parietal cortex (PPC), occipital, and inferiotemporal cortex, as well as in the contralateral M1/S1 and ipsilateral CB. We did not find any additional activation when comparing bimanual with unimanual conditions. The lack of significant activation in the comparison "bimanual > unimanual" gives reason to suggest that this network is not limited to the control of bimanual motor actions, but responsible for unimanually coordinated movements as well. Interestingly, we found stronger activations for unimanual as compared to bimanual coordination. We hypothesize that task difficulty (degrees of freedom to control, e.g., number of limbs) is more important in determining which network components are activated and to what extent, compared to the factor of bimanuality. It even seemed to be less demanding for the motor system to control the cursor bimanually compared to the unimanual performance with two adjacent fingers.

Long-term training affects cerebellar processing in skilled keyboard players.

We studied cerebellar hemodynamic responses in highly skilled keyboard players and control subjects during complex tasks requiring unimanual and bimanual finger movements. Both groups showed strong hemodynamic responses in the cerebellum during the task conditions. However, non-musicians showed generally stronger hemodynamic responses in the cerebellum than keyboard players. We conclude that, due to long-term motor practice a different cortical activation pattern can be visualized in keyboard players. For the same movements fewer neurons need to be recruited. The different volume of the activated cortical areas might therefore reflect the different effort necessary for motor performance in both groups.