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The neural correlates of memory formation in humans have long been investigated by exposing subjects to diverse material and comparing responses to items later remembered to those forgotten. Tasks requiring memorization of sensory sequences afford unique possibilities for linking neural memorization processes to behavior, because, rather than comparing across different items of varying content, each individual item can be examined across the successive learning states of being initially unknown, newly learned, and eventually, fully known. Sequence learning paradigms have not yet been exploited in this way, however. Here, we analyze the event-related potentials of subjects attempting to memorize sequences of visual locations over several blocks of repeated observation, with respect to pre- and post-block recall tests. Over centro-parietal regions, we observed a rapid P300 component superimposed on a broader positivity, which exhibited distinct modulations across learning states that were replicated in two separate experiments. Consistent with its well-known encoding of surprise, the P300 deflection monotonically decreased over blocks as locations became better learned and hence more expected. In contrast, the broader positivity was especially elevated at the point when a given item was newly learned, i.e., started being successfully recalled. These results implicate the Broad Positivity in endogenously-driven, intentional memory formation, whereas the P300, in processing the current stimulus to the degree that it was previously uncertain, indexes the cumulative knowledge thereby gained. The decreasing surprise/P300 effect significantly predicted learning success both across blocks and across subjects. This presents a new, neural-based means to evaluate learning capabilities independent of verbal reports, which could have considerable value in distinguishing genuine learning disabilities from difficulties to communicate the outcomes of learning, or perceptual impairments, in a range of clinical brain disorders.

Adapting movements to a visual rotation involves the activation of right posterior parietal areas. Further performance improvement requires an increase of slow wave activity in subsequent sleep in the same areas. Here we ascertained whether a post-learning trace is present in wake EEG and whether such a trace is influenced by sleep slow waves.In two separate sessions, we recorded high-density EEG in 17 healthy subjects before and after a visuomotor rotation task, which was performed both before and after sleep. High-density EEG was recorded also during sleep. One session aimed to suppress sleep slow waves, while the other session served as a control.After learning, we found a trace in the eyes-open wake EEG as a local, parietal decrease in alpha power. After the control night, this trace returned to baseline levels, but it failed to do so after slow wave deprivation. The overnight change of the trace correlated with the dissipation of low frequency (8 Hz) NREM sleep activity only in the control session.Visuomotor learning leaves a trace in the wake EEG alpha power that appears to be renormalized by sleep slow waves.These findings link visuomotor learning to regional changes in wake EEG and sleep homeostasis.

We investigated whether dopamine influences the rate of adaptation to a visuomotor distortion and the transfer of this learning from the right to the left limb in human subjects. We thus studied patients with Parkinson disease as a putativein vivomodel of dopaminergic denervation. Despite normal adaptation rates, patients showed a reduced transfer compared with age-matched healthy controls. The magnitude of the transfer, but not of the adaptation rate, was positively predicted by the values of dopamine-transporter binding of the right caudate and putamen. We conclude that striatal dopaminergic activity plays an important role in the transfer of visuomotor skills.

Motor sequence learning is not a unitary phenomenon, but involves optimizing different components that include declarative and procedural aspects. In this work we designed an experimental approach that allows monitoring all the aspects of sequence learning using a finger opposition task and a movement-by-movement analysis. Subjects performed a visuomotor sequence learning paradigm with (Explicit) or without (Implicit) instructions and we measured response time (RT) and touch duration (TD) for each finger opposition movement of the sequence. Our results indicated that sequence learning induced a double-faced effect on motor performance: a decrease of RT and an increase of TD. However, the above changes manifested differently among subjects: all subjects that, by the end of session, had complete recall of the sequence order, reached an equal level of performance by the last sequence block while in those who had on average only a poor recall of the sequence order, learning was evident only as a slight decrease of RT across sequence blocks, while no kinematic changes (i.e., changes in TD) occurred. Our results indicate that, in the absence of specific instructions, learning evolves from an early stage in which only small decreases of RT are observed to a phase in which progressive knowledge of the sequential structure allows for dramatic changes of RT, together with a progressive change of motor performance (i.e., changes in TD).

To determine whether the process involved in movement preparation of patients in the early stages of Parkinson's disease (PD) shares attentional resources with visual learning, we tested 23 patients with PD and 13 normal controls with two different tasks. The first was a motor task where subjects were required to move as soon as possible to randomly presented targets by minimizing reaction time. The second was a visual learning task where targets were presented in a preset order and subjects were asked to learn the sequence order by attending to the display without moving. PD patients showed higher reaction and movement times, while visual learning was reduced compared to controls. For PD patients, reaction times, but not movement times, displayed an inverse significant correlation with the scores of visual learning. We conclude that visual declarative learning and movement preparation might share similar attentional and working memory resources.

Observation of people performing movements facilitates motor planning, execution and memory formation. Tempo, a crucial aspect involved in the execution of rhythmic movements, is normally perceived and learned through auditory channels. In this work, we ascertained whether: first, the frequency of self-paced finger movements (SPMs), which in normal subjects is around 2 Hz, is modified by prior observation of movements performed at either 1 or 3 Hz; second, such changes are lasting; third, there is an effect of time interval between observation and performance. We finally determined the effect of providing explicit information about the upcoming motor task. Seventy-two normal subjects (12 groups) performed a simple finger sequence at different intervals after observation of videos of either landscapes or finger opposition movements. Both with and without information about the upcoming task, observation influenced the tempo of SPMs and led to memory formation. With knowledge of the upcoming task, such changes occurred at all observation-execution intervals, while without instructions, changes took place only when SPMs were performed immediately after observation. Compared to explicit instructions, the absence of instructions produced tempo's changes that more closely resembled the observed rhythms. We conclude that learning requires a prompt comparison between visual and sensorimotor representations of movements; moreover, learning with explicit instructions is more efficient, as activity in both the dorsal and ventral streams might be potentiated by the chatecholaminergic attentional systems that promote long-term potentiation. These results provide the bases for novel neurorehabilitation strategies in terms of temporal re-organization of movement.

The ability to perform accurate sequential movements is essential to normal motor function. Learning a sequential motor behavior is comprised of two basic components: explicit identification of the order in which the sequence elements should be performed and implicit acquisition of spatial accuracy for each element. Here we investigated the time course of learning of these components for a first sequence (SEQA) and their susceptibility to interference from learning a second sequence (SEQB). We assessed explicit learning with a discrete index, the number of correct anticipatory movements, and implicit learning with a continuous variable, spatial error, which decreased during learning without subject awareness. Spatial accuracy to individual sequence elements reached asymptotic levels only when the whole sequence order was known. Interference with recall of the order of SEQA persisted even when SEQB was learned 24 h after SEQA. However, there was resistance to interference by SEQB with increased initial training with SEQA. For implicit learning of spatial accuracy, SEQB interfered at 5 min but not 24 h after SEQA. As in the case of sequence order, prolonged initial training with SEQA induced resistance to interference by SEQB. We conclude that explicit sequence learning is more susceptible to anterograde interference and implicit sequence learning is more susceptible to retrograde interference. However, both become resistant to interference with saturation training. We propose that an essential feature of motor skill learning is the process by which discrete explicit task elements are combined with continuous implicit features of movement to form flawless sequential actions.

We used a sequence-learning task to assess whether: 1. The time interval between awakening and training equally affects the rate of acquisition of sequence order, a declarative component, and the kinematic optimization process, an implicit component; 2. Sleep enhances the retention of both these aspects of sequence learning.For aim 1, we compare the acquisition rate of a new motor sequence in a group trained in the morning and another in the evening. For aim 2., we tested retention of the same motor sequence twelve hours later, either without sleep (normal day activity or a night of sleep deprivation) or with interposed sleep (afternoon napping or regular full night sleep).Training and Testing were performed in a controlled laboratory setting.Thirty-six right-handed normal subjects (age range 18-24 years; 16 women).During the training, acquisition rate of the sequence order was significantly higher in the AM-trained than in the PM-trained group, without differences in the kinematic optimization processes. Both declarative and implicit learning indices were significantly higher in the subjects tested after sleep compared to those tested without interposed sleep.The best time for fast and efficient acquisition of new declarative material is the morning, while the kinematic aspects of skill acquisition are not sensitive to the time of day. However, better retention of both declarative material and motor skills requires two conditions: a period of post-training sleep and the achievement of performance saturation during training.

In this study, we characterized the patterns and timing of cortical activation of visually guided movements in a task with critical temporal demands. In particular, we investigated the neural correlates of motor planning and on-line adjustments of reaching movements in a choice-reaction time task. High-density electroencephalograohy (EEG, 256 electrodes) was recorded in 13 subjects performing reaching movements. The topography of the movement-related spectral perturbation was established across five 250-ms temporal windows (from prestimulus to postmovement) and five frequency bands (from theta to beta). Nine regions of interest were then identified on the scalp, and their activity was correlated with specific behavioral outcomes reflecting motor planning and on-line adjustments. Phase coherence analysis was performed between selected sites. We found that motor planning and on-line adjustments share similar topography in a fronto-parietal network, involving mostly low frequency bands. In addition, activities in the high and low frequency ranges have differential function in the modulation of attention with the former reflecting the prestimulus, top-down processes needed to promote timely responses, and the latter the planning and control of sensory-motor processes.

C. Ghez, J. Gordon, and M. F. Ghilardi (1995; J. Gordon, M. F. Ghilardi, & C. Ghez, 1995; R. L. Sainburg, M. F. Ghilardi, H. Poizner, & C. Ghez, 1995) have found that proprioceptive deafferentation impairs feedforward and feedback mechanisms that control reaching movements. In the present study, the authors found immobilization-induced changes in limb kinematics, including joint motion, in 32 healthy participants who performed out-and-back movements before and after 0, 6, or 12 hr of immobilization of the left arm. Control participants did not undergo the arm immobilization procedure. Immobilization for 12 hr, but not 6 hr, caused trajectories with increased hand-path areas and altered interjoint coordination. The abnormalities were smaller in amplitude but similar in quality to those reported in deafferented patients (R. L. Sainburg et al.). In addition, movement onset point significantly drifted after immobilization. Thus, short-term limb disuse can affect interjoint coordination by acting on feedforward mechanisms. These behavioral alterations are potentially related to cortical plastic changes.