Cortical Networks for Correcting Errors in Sensorimotor Synchronization Depend on the Direction of Asynchrony.

Recent work provides clues that different cortical mechanisms may be employed when correcting for errors in sensorimotor synchronization that increase tap-tone asynchrony compared with those that decrease it. The authors tested this hypothesis by recording 64-channel electroencephalography while participants synchronized with an auditory metronome. We systematically introduced positive and negative phase-shift perturbations that were either liminal (10%) and subliminal (3%). We used a distributed source modeling approach to evaluate oscillatory activity and connectivity of discrete cortical sources. Three key findings support our hypothesis. First was a theta band response indicative of error detection and top-down control observed in frontomedial presupplementary motor area (pre-SMA) and anterior cingulate for liminal positive perturbations. Second was an increase in theta band coupling between the SMA and contralateral motor cortex exclusively for positive perturbations suggesting a top-down modulation of motor parameters. Third, when compared with other conditions, liminal positive perturbations result in an increase in postmovement beta rebound within contralateral primary motor cortex. The authors propose that frontomedial motor areas exert a top-down inhibitory influence over the primary motor cortex to effectively lengthen tap intervals in response to lengthening tap-tone asynchronies.

Early and late event-related potentials are modulated by infant and adult faces of high and low attractiveness.

The processing of infant faces may be somewhat distinct from that of adult faces. Indeed, recent neuroimaging studies have provided evidence of an early, "baby-specific" neural response whereby infant faces are perceived more rapidly than adult faces. Using event-related potentials, the present study aimed to determine whether the preferential response to infant faces is present at both early and late stages of face processing, and to investigate the effects of esthetic appearance on the processing of adult and infant faces by directly manipulating the perceived attractiveness or cuteness within a given face identity. Here, we find evidence for enhanced processing of infant faces, relative to adult faces, at both early (N170, P2) and late (LPC) stages of face processing. We also find that the esthetic appearance of both infant and adult faces modulates early neural responses, with enhanced responses to less attractive/cute faces as compared to more attractive/cute faces. Overall, our results provide additional evidence for a preferential response to infant faces at early stages of processing, and provide new evidence that this preferential response occurs at later stages of face processing as well, independent of the esthetic quality of the face or observer sex.

Dorsal stream activity and connectivity associated with action priming of ambiguous apparent motion.

Theories proposing a bidirectional influence between action and perception are well supported by behavioral findings. In contrast to the growing literature investigating the brain mechanisms by which perception influences action, there is a relative dearth of neural evidence documenting how action may influence perception. Here we show that action priming of apparent motion perception is associated with increased functional connectivity between dorsal cortical regions connecting vision with action. Participants manually rotated a joystick in a clockwise or counter-clockwise direction while viewing ambiguous apparent rotational motion. Actions influenced perception when the perceived direction of the ambiguous display was the same as manual rotation. For comparison, participants also rotated the joystick while viewing non-ambiguous apparent motion and in the absence of apparent motion. In a final control condition, participants viewed ambiguous apparent motion without manual rotation. Actions influence on perception was accompanied by a significant increase in alpha and beta band event related desynchronization (ERD) in contralateral primary motor cortex, superior parietal lobe and middle occipital gyrus. Increased ERD across these areas was accompanied by an increase in gamma band phase locking between primary motor, parietal, striate and extrastriate regions. Similar patterns were not observed when action was compatible with perception, but did not influence it. These data demonstrate that action influences perception by strengthening the interaction across a broad sensorimotor network for the putative purpose of integrating compatible action outcomes and sensory information into a single coherent percept.

A parametric fMRI investigation of context effects in sensorimotor timing and coordination.

Mounting evidence suggests that information derived from environmental and behavioral sources is represented and maintained in the brain in a context-dependent manner. Here we investigate whether activity patterns underlying movements paced according to an internal temporal representation depend on how that representation is acquired during a previous pacing phase. We further investigate the degree to which context dependence is modulated by different time delays between pacing and continuation. BOLD activity was recorded while subjects moved at a rate established during a pacing interval involving either synchronized or syncopated coordination. Either no-delay or a 3, 6 or 9s delay was introduced prior to continuation. Context-dependent regions were identified when differences in neural activity generated during pacing continued to be observed during continuation despite the intervening delay. This pattern was observed in pre-SMA, bilateral lateral premotor cortex, bilateral declive and left inferior semi lunar lobule. These regions were more active when continuation followed from syncopation than from synchronization regardless of the delay length putatively revealing a context-dependent neural representation of the temporal interval. Alternatively, task related regions in which coordination-dependent differences did not persist following the delay, included bilateral putamen and supplementary-motor-area. This network may support the differential timing demands of coordination. A classic prefrontal-parietal-temporal working memory network was active only during continuation possibly providing mnemonic support for actively maintaining temporal information during the variable delay. This work provides support for the hypothesis that some timing information is represented in a task-dependent manner across broad cortical and subcortical networks.

Functional MRI reveals the existence of modality and coordination-dependent timing networks.

Growing evidence suggests that interval timing in humans is supported by distributed brain networks. Recently, we demonstrated that the specific network recruited for the performance of rhythmic timing is not static but is influenced by the coordination pattern employed during interval acquisition. Here we expand on this previous work to investigate the role of stimulus modality and coordination pattern in determining the brain areas recruited for performance of a self-paced rhythmic timing task. Subjects were paced with either a visual or an auditory metronome in either a synchronized (on the beat) or syncopated (off the beat) coordination pattern. The pacing stimulus was then removed and subjects continued to move based on the required interval. When compared with networks recruited for auditory pacing and continuation, the visual-specific activity was observed in the classic dorsal visual stream that included bilateral MT/V5, bilateral superior parietal lobe, and right ventral premotor cortex. Activity in these regions was present not only during pacing, when visual information is used to guide motor behavior, but also during continuation, when visual information specifying the temporal interval was no longer present. These results suggest a role for modality-specific areas in processing and representing temporal information. The cognitive demands imposed by syncopated coordination resulted in increased activity in a broad network that included supplementary motor area, lateral pre-motor cortex, bilateral insula, and cerebellum. This coordination-dependent activity persisted during the subsequent continuation period, when stimuli were removed and no coordination constraints were imposed. Taken together, the present results provide additional evidence that time and timing are served by a context-dependent distributed network rooted in basic sensorimotor processes.

Neural substrates of real and imagined sensorimotor coordination.

Much debate in the behavioral literature focuses on the relative contribution of motor and perceptual processes in mediating coordinative stability. To a large degree, such debate has proceeded independently of what is going on in the brain. Here, using blood oxygen level-dependent measures of neural activation, we compare physically executed and imagined rhythmic coordination in order to better assess the relative contribution of hypothesized neuromusculoskeletal mechanisms in modulating behavioral stability. The executed tasks were to coordinate index finger to thumb opposition movements of the right hand with an auditory metronome in either a synchronized (on the beat) or syncopated (off the beat) pattern. Imagination involved the same tasks, except without physical movement. Thus, the sensory stimulus and coordination constraints were the same in both physical and imagination tasks, but the motoric requirements were not. Results showed that neural differences between executed synchronization and syncopation found in premotor cortex, supplementary motor area, basal ganglia and lateral cerebellum persist even when the coordinative patterns were only imagined. Neural indices reflecting behavioral stability were not abolished by the absence of overt movement suggesting that coordination phenomena are not exclusively rooted in purely motoric constraints. On the other hand, activity in the superior temporal gyrus was modulated by both the presence of movement and the nature of the coordination, attesting to the intimacy between perceptual and motoric processes in coordination dynamics.

Spatiotemporal analysis of the neuromagnetic response to rhythmic auditory stimulation: rate dependence and transient to steady-state transition.

OBJECTIVE: Whole head magnetoencephalography was used to investigate the spatiotemporal dynamics of neuromagnetic brain activity associated with rhythmic auditory stimulation. METHODS: In order to characterize the evolution of the auditory responses we applied a Karhunen-Loève decomposition and k-means cluster analysis to globally compare spatial patterns of brain activity at different latencies and stimulation rates. Tones were presented binaurally at 27 different stimulation rates within a perceptually and behaviorally relevant range from 0.6 to 8.1 Hz. RESULTS: Over this range, we observed a linear increase of the amplitude of the main auditory response at 100 ms latency (N1m) with increasing inter-stimulus interval, and qualitative changes of the overall spatiotemporal dynamics of the auditory response. In particular, a transition occurred between a transient evoked response at low frequencies, and a continuous steady-state response at high frequencies. CONCLUSIONS: We show the onset of temporal overlap between responses to successive tones that leads to this transition. Response overlap begins to occur near 2 Hz, marking the onset of a continuous perceptual representation.

Practice-dependent modulation of neural activity during human sensorimotor coordination: a functional Magnetic Resonance Imaging study.

We investigated the degree to which differences in the pattern of blood oxygen level dependent activity (BOLD) between syncopated and synchronized coordination patterns are altered by practice. Baseline levels of BOLD activity were obtained from eight subjects while they syncopated or synchronized with an auditory metronome at 1.25 Hz. Subjects then practiced syncopation at the same rate for four consecutive sessions. Post practice scans of the two coordination patterns were then performed. Before practice, baseline syncopation activated a much broader network of both cortical and subcortical regions than synchronization that included Supplementary Motor Area (SMA), bilateral putamen, left thalamus, bilateral superior temporal gyrus as well as the vermis. This pattern of activity is hypothesized to reflect the extra timing and attention requirements of syncopation. After practice, activity in superior temporal gyrus and vermis were no longer observed during syncopation reflecting a reduction in the need for attention and the use of sensory feedback for guiding behavior. Surprisingly, post practice synchronization resulted in additional significant activations in SMA, inferior frontal gyrus and superior temporal gyrus as well as small activations in bilateral putamen. Practice with the more difficult syncopation task thus had a dual effect of decreasing the number of active regions during syncopation and increasing the number of active regions during synchronization. Since overt syncopation performance did not change significantly as a result of practice, these observed neural changes appear to be due to context- and history-dependent factors, rather than behavioral learning per se.

Neuromagnetic activity in alpha and beta bands reflect learning-induced increases in coordinative stability.

OBJECTIVE: To investigate how learning induced increases in stability on a syncopation task are manifest in the dynamics of cortical activity. METHOD: Magnetoencephalography was recorded from 143 sensors (CTF Systems, Inc). A pre-training procedure determined the critical frequency (F(c)) for each subject (n=4). Subjects either syncopated or synchronized to a metronome that increased in frequency from 1.2 to 3.0 Hz in 0.2 Hz steps. The F(c) was the point at which subjects spontaneously switched from syncopation to synchronization. Subjects then underwent 100 training trials (with feedback) at F(c). Following the learning phase the pre-training procedure was repeated. RESULTS: An increase in the F(c) occurred indicating that practice improved the stability of syncopation. The transition delay was also observed in the phase of the time-averaged signal in sensors over the contralateral sensorimotor area and in power analysis in the 8-12 Hz and 18-24 Hz frequency bands. Initially, reduced power was observed bilaterally during syncopation compared to synchronization. Following training, these differences were reduced or eliminated. CONCLUSION: Pre-training power differences can be explained by the greater difficulty of the syncopation task. The reduction in power differences following training suggests that at the cortical level, syncopation became more similar to synchronization possibly reflecting a decrease in task and/or attention demands.

Neural network classifications and correlation analysis of EEG and MEG activity accompanying spontaneous reversals of the Necker cube.

It has recently been suggested that reentrant connections are essential in systems that process complex information [A. Damasio, H. Damasio, Cortical systems for the retrieval of concrete knowledge: the convergence zone framework, in: C. Koch, J.L. Davis (Eds.), Large Scale Neuronal Theories of the Brain, The MIT Press, Cambridge, 1995, pp. 61-74; G. Edelman, The Remembered Present, Basic Books, New York, 1989; M.I. Posner, M. Rothbart, Constructing neuronal theories of mind, in: C. Koch, J.L. Davis (Eds.), Large Scale Neuronal Theories of the Brain, The MIT Press, Cambridge, 1995, pp. 183-199; C. von der Malsburg, W. Schneider, A neuronal cocktail party processor, Biol. Cybem., 54 (1986) 29-40]. Reentry is not feedback, but parallel signalling in the time domain between spatially distributed maps, similar to a process of correlation between distributed systems. Accordingly, it was expected that during spontaneous reversals of the Necker cube, complex patterns of correlations between distributed systems would be present in the cortex. The present study included EEG (n=4) and MEG recordings (n=5). Two experimental questions were posed: (1) Can distributed cortical patterns present during perceptual reversals be classified differently using a generalised regression neural network (GRNN) compared to processing of a two-dimensional figure? (2) Does correlated cortical activity increase significantly during perception of a Necker cube reversal? One-second duration single trials of EEG and MEG data were analysed using the GRNN. Electrode/sensor pairings based on cortico-cortical connections were selected to assess correlated activity in each condition. The GRNN significantly classified single trials recorded during Necker cube reversals as different from single trials recorded during perception of a two-dimensional figure for both EEG and MEG. In addition, correlated cortical activity increased significantly in the Necker cube reversal condition for EEG and MEG compared to the perception of a non-reversing stimulus. Coherent MEG activity observed over occipital, parietal and temporal regions is believed to represent neural systems related to the perception of Necker cube reversals.

Motor cortex activity and predicting side of movement: neural network and dipole analysis of pre-movement magnetic fields.

Neuromagnetic fields were recorded from human subjects during the performance of left and right voluntary finger movements. Modeling of current dipole sources indicated symmetric activation of both motor cortices beginning 600 ms prior to movement onset. This activity became lateralized to the contralateral hemisphere 200-300 ms prior to movement onset, the period during which an artificial neural network showed increased ability to predict side of movement within single trials. The results describe the mechanism of lateralization of cortical brain activity preceding voluntary movement and provide further evidence of the involvement of ipsilateral motor cortex in unilateral movements.