Hypothetical and real choice differentially activate common valuation areas.

Hypothetical reports of intended behavior are commonly used to draw conclusions about real choices. A fundamental question in decision neuroscience is whether the same type of valuation and choice computations are performed in hypothetical and real decisions. We investigated this question using functional magnetic resonance imaging while human subjects made real and hypothetical choices about purchases of consumer goods. We found that activity in common areas of the orbitofrontal cortex and the ventral striatum correlated with behavioral measures of the stimulus value of the goods in both types of decision. Furthermore, we found that activity in these regions was stronger in response to the stimulus value signals in the real choice condition. The findings suggest that the difference between real and hypothetical choice is primarily attributable to variations in the value computations of the medial orbitofrontal cortex and the ventral striatum, and not attributable to the use of different valuation systems, or to the computation of stronger stimulus value signals in the hypothetical condition.

Repetitive transcranial magnetic stimulation over the right dorsolateral prefrontal cortex decreases valuations during food choices.

Several studies have found decision-making-related value signals in the dorsolateral prefrontal cortex (DLPFC). However, it is unknown whether the DLPFC plays a causal role in decision-making, or whether it implements computations that are correlated with valuations, but that do not participate in the valuation process itself. We addressed this question by using repetitive transcranial magnetic stimulation (rTMS) while subjects were involved in an economic valuation task involving the consumption of real foods. We found that, as compared with a control condition, application of rTMS to the right DLPFC caused a decrease in the values assigned to the stimuli. The results are consistent with the possibility that the DLPFC plays a causal role in the computation of values at the time of choice.

Mechanisms controlling motor output to a transfer hand after learning a sequential pinch force skill with the opposite hand.

OBJECTIVE: Training to perform a serial reaction-time task (procedural motor learning) with one hand results in performance improvements in the untrained as well as in the trained hand, a phenomenon referred to as intermanual transfer. The aim of this study was to investigate the neurophysiological changes associated with intermanual transfer associated with learning to perform an eminently different task involving fine force control within the primary motor cortex (M1). We hypothesized that intermanual transfer of learning such a task would reveal intracortical changes within M1. METHODS: Speed (time to complete each sequence) and accuracy (% of accuracy errors) of motor performance were measured in both hands before and after right (dominant) hand practice. Transcranial magnetic stimulation (TMS) was used to characterize recruitment curves (RC), short intracortical inhibition (SICI), intracortical facilitation (ICF) and interhemispheric inhibition (IHI) from the left to the right M1. RESULTS: Practice resulted in significant improvements in both speed and accuracy in the right trained hand and in the left untrained hand. RC increased in the left M1, SICI decreased in both M1s, and IHI from the left to the right M1 decreased. No changes were identified in ICF nor in RC in the right M1. CONCLUSIONS: Our results suggest that some neurophysiological mechanisms operating in the M1 controlling performance of an untrained hand may contribute to optimize the procedure for selecting and implementing correct pinch force levels. SIGNIFICANCE: These results raise the hypothesis of a contribution of modulation of SICI and IHI, or an interaction between both to intermanual transfer after learning a sequential pinch force task.

Modulation of effects of intermittent theta burst stimulation applied over primary motor cortex (M1) by conditioning stimulation of the opposite M1.

The excitability of the human primary motor cortex (M1) as tested with transcranial magnetic stimulation (TMS) depends on its previous history of neural activity. Homeostatic plasticity might be one important physiological mechanism for the regulation of corticospinal excitability and synaptic plasticity. Although homeostatic plasticity has been demonstrated locally within M1, it is not known whether priming M1 could result in similar homeostatic effects in the homologous M1 of the opposite hemisphere. Here, we sought to determine whether down-regulating excitability (priming) in the right (R) M1 with 1-Hz repetitive transcranial magnetic stimulation (rTMS) changes the excitability-enhancing effect of intermittent theta burst stimulation (iTBS) applied over the homologous left (L) M1. Subjects were randomly allocated to one of four experimental groups in a sham-controlled parallel design with real or sham R M1 1-Hz TMS stimulation always preceding L M1 iTBS or sham by about 10 min. The primary outcome measure was corticospinal excitability in the L M1, as measured by recruitment curves (RCs). Secondary outcome measures included pinch force, simple reaction time, and tapping speed assessed in the right hand. The main finding of this study was that preconditioning R M1 with 1-Hz rTMS significantly decreased the excitability-enhancing effects of subsequent L M1 iTBS on RCs. Application of 1-Hz rTMS over R M1 alone and iTBS over L M1 alone resulted in increased RC in L M1 relative to sham interventions. The present findings are consistent with the hypothesis that homeostatic mechanisms operating across hemispheric boundaries contribute to regulate motor cortical function in the primary motor cortex.

Improvement of spatial tactile acuity by transcranial direct current stimulation.

OBJECTIVE: Non-invasive brain stimulation such as transcranial direct current stimulation (tDCS) has been successfully used to induce polarity-specific excitability changes in the brain. However, it is still unknown if anodal tDCS (tDCS(anodal)) applied to the primary somatosensory cortex (S1) can lead to behavioral changes in performance of tactile discriminative tasks. METHODS: Using an accurate tactile discrimination task (grating orientation task: GOT) we tested the hypothesis that application of 1mA of tDCS(anodal) (current density at the electrodes of 0.04mA/cm2) over the left S1 can lead to an improved tactile spatial acuity in the contralateral index-finger (IF). RESULTS: Performance in the GOT task with the contralateral IF but not with the ipsilateral IF was enhanced for about 40min after a 20min application of tDCS(anodal) in the absence of changes with sham stimulation. CONCLUSIONS: These results provide the first evidence that tDCS(anodal) over S1 improves performance in a complex somatosensory task beyond the period of stimulation. SIGNIFICANCE: The ability to induce performance improvement in the somatosensory domain with tDCS applied over S1 could be used to promote functional recovery in patients with diminished tactile perception.

Contribution of transcranial magnetic stimulation to the understanding of cortical mechanisms involved in motor control.

Transcranial magnetic stimulation (TMS) was initially used to evaluate the integrity of the corticospinal tract in humans non-invasively. Since these early studies, the development of paired-pulse and repetitive TMS protocols allowed investigators to explore inhibitory and excitatory interactions of various motor and non-motor cortical regions within and across cerebral hemispheres. These applications have provided insight into the intracortical physiological processes underlying the functional role of different brain regions in various cognitive processes, motor control in health and disease and neuroplastic changes during recovery of function after brain lesions. Used in combination with neuroimaging tools, TMS provides valuable information on functional connectivity between different brain regions, and on the relationship between physiological processes and the anatomical configuration of specific brain areas and connected pathways. More recently, there has been increasing interest in the extent to which these physiological processes are modulated depending on the behavioural setting. The purpose of this paper is (a) to present an up-to-date review of the available electrophysiological data and the impact on our understanding of human motor behaviour and (b) to discuss some of the gaps in our present knowledge as well as future directions of research in a format accessible to new students and/or investigators. Finally, areas of uncertainty and limitations in the interpretation of TMS studies are discussed in some detail.

On-line flexibility of the cognitive tuning of corticospinal excitability: a TMS study in human gait.

Human subjects have been found to be able to cognitively prepare themselves to resist to a TMS-induced central perturbation by selectively modulating the corticospinal excitability (CS). The aim of this study was to investigate the on-line adaptability of this cognitive tuning of CS excitability during human gait. Transcranial magnetic stimulation (TMS) was used both as a central perturbation evoking a movement and as a tool for quantifying the CS excitability before the movement was evoked. TMS was applied at mid-stance (evoking additional hip extension) or at the beginning of the swing (evoking hip flexion) with a random phase, thus evoking unpredictable flexion or extension movement. This was compared to a condition of fixed phase, in which the subjects knew in advance the direction of the evoked movement. In both conditions, we compared the amplitude of the TMS-evoked movement and the motor-evoked potentials (MEPs) of the muscles acting at the hip joint (RF/BF) according to two opposite instructions, either to cognitively prepare to "let go", or to cognitively prepare to "compensate" for the evoked movements. The results showed that the subjects were able to compensate for random TMS-evoked movements, but with a lower performance level in comparison to the fixed TMS-evoked movements. When they succeeded in the random-phase condition, the subjects used the same preparation strategy as in the fixed-phase condition; preparing to compensate resulted in a selective increase in the CS excitability to those muscles which would be involved in counteracting the possible central perturbation. This requires continuous change in the tuning of CS excitability within the stride and thus reveals the high flexibility of the cognitive tuning of CS excitability during gait.

Cognitive tuning of corticospinal excitability during human gait: adaptation to the phase.

The aim of this study was to investigate how the cognitive tuning of corticospinal (CS) excitability adapts to the type of evoked-movement (Flexion vs. Extension) during human gait. Transcranial magnetic stimulation (TMS) was used both as a central perturbation evoking a movement and as a tool for quantifying the CS excitability of the muscles under study (RF/BF). In the first condition (Dst), the TMS occurred at mid-stance, inducing hip extension, whereas in the second condition (Dsw), the TMS occurred at the beginning of the swing phase, inducing hip flexion. In both conditions, the subjects were asked to cognitively prepare to either not intervene (NINT) or to compensate (COMP) for the evoked-movements. The results showed that, regardless of the type of evoked-movement, preparing to compensate resulted in a selective increase in the CS excitability to those muscles that would be involved in counteracting the possible central perturbation, i.e. the hip extensor muscle (BF) to compensate for an evoked flexion during the swing phase or the hip flexor muscle (RF) to compensate for an evoked extension during the stance phase. This latter result offers the first evidence of a modulation in CS excitability to the proximal muscles during the stance phase. In conclusion, the cognitive tuning of CS excitability was found to adapt to the gait phases. Moreover, the same selective preparation strategy was observed whether the central perturbation occurred during the stance or the swing phase of the step cycle.

Direct evidence for a binding between cognitive and motor functions in humans: a TMS study.

During voluntary motor actions, the cortico-spinal (CS) excitability is known to be modulated, on the one hand by cognitive (intention-related) processes and, on the other hand, by motor (performance-related) processes. Here, we studied the way these processes interact in the tuning of CS excitability during voluntary wrist movement. We used transcranial magnetic stimulation (TMS) both as a reliable tool for quantifying the CS excitability, through the motor-evoked potentials (MEPs), and as a central perturbation evoking a movement (because the stimulation intensity was above threshold) with subjects instructed to prepare (without changing their muscle activation) either to "let go" or to "resist" to this evoked movement. We studied the simultaneous evolution of both the motor performance and the MEPs in the wrist flexor and extensor, separately for the successful trials (on average, 66% of the trials whatever the condition) and the unsuccessful trials; this allowed us to dissociate the intention- and performance-related processes. To their great surprise, subjects were found able to cognitively prepare themselves to resist a TMS-induced central perturbation; they all reported an important cognitive effort on the evoked movement. Moreover, because TMS only evoked short-latency MEPs (and no long-latency components), the amplitude of these short-latency MEPs was found to be related in a continuous way to the actual movement whatever the prior intention. These results demonstrate that prior intention allows an anticipatory modulation of the CS excitability, which is not only selective (as already known) but also efficient, giving the intended motor behavior a real chance to be realized. This constitutes a direct evidence of the role of the CS excitability in the binding between cognitive and motor processes in humans.

Task-induced modulation of motor evoked potentials in upper-leg muscles during human gait: a TMS study.

The aim of this study was to determine the relative involvement of the corticospinal (CS) pathway in voluntarily controlled walking compared to unconstrained walking. In the voluntarily controlled walking condition, subjects had to walk at the same speed as in unconstrained walking with a mechanical constraint, which is known to affect specifically the upper-leg muscles. The motor cortex was activated transcranially using a focal magnetic stimulation coil in order to elicit motor evoked potentials (MEPs) in the rectus femoris (RF) and the biceps femoris (BF). The magnetic stimulation was delivered at the end of the swing (at 90% of the cycle duration), when the EMG backgrounds were similar in the two experimental conditions. For each subject in each condition, MEPs were measured for several stimulus intensities in order to establish the input/output (I/O) curve (MEPs amplitude plotted against stimulus strength). The results showed a significant increase in the MEPs amplitude of both the RF and BF in voluntarily controlled walking compared to unconstrained walking, which is the first evidence of cofacilitation of MEPs in antagonist upper-leg muscles during human gait. In conclusion, although a lot of studies have emphasized a privileged input of the corticospinal pathway to the distal lower-leg muscles, this study shows that, if a locomotory task requires fine control of the proximal upper-leg muscles, a selective facilitation of MEPs is observed in these muscles.