ADHD and the facilitation of neural synchrony by attentional mechanisms


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(Updated 11/26/04)

Lazzaro I, Gordon E, Li W, Lim CL, Plahn M, Whitmont S, Clarke S, Barry RJ, Dosen A, Meares R.
Simultaneous EEG and EDA measures in adolescent attention deficit hyperactivity disorder.
Int J Psychophysiol. 1999 Nov;34(2):123-34.
Adolescent unmedicated ADHD males and age- and sex-matched normal control subjects were examined simultaneously using EEG and EDA measures in a resting eyes-open condition. ADHD adolescents showed increased absolute and relative Theta and Alpha1 activity, reduced relative Beta activity, reduced skin conductance level (SCL) and a reduced number of non-specific skin conductance responses (NS.SCRs) compared with the control subjects. Our findings indicate the continuation of increased slow wave activity in ADHD adolescents and the presence of a state of autonomic hypoarousal in this clinical group. [Abstract]

Lazzaro I, Gordon E, Whitmont S, Plahn M, Li W, Clarke S, Dosen A, Meares R.
Quantified EEG activity in adolescent attention deficit hyperactivity disorder.
Clin Electroencephalogr. 1998 Jan;29(1):37-42.
The aim of this study was to explore elements of the maturational and cortical hypoarousal models in adolescent ADHD, by examining EEG activity in a rest eyes open condition, in 26 adolescent unmedicated ADHD males and 26 age and sex matched normal controls. ADHD adolescents were found to have increased anterior EEG absolute theta activity and reduced posterior relative beta activity compared with controls. These results lend some support to the continuation of a maturational lag and reduced cortical arousal in adolescent ADHD. These measures need to be further explored using concomitant EEG with electrodermal measures of arousal. [Abstract]

Gamma synchrony in ADHD: Project Details by Ms. Hannah Keage

Rubia K, Noorloos J, Smith A, Gunning B, Sergeant J.
Motor timing deficits in community and clinical boys with hyperactive behavior: the effect of methylphenidate on motor timing.
J Abnorm Child Psychol. 2003 Jun;31(3):301-13.
In a previous paper we showed that community children with hyperactive behavior were more inconsistent than controls in the temporal organization of their motor output. In this study we investigated: (1) various aspects of motor timing processes in 13 clinically diagnosed boys with attention deficit hyperactivity disorder (ADHD) who were compared to 11 community boys with hyperactive behavior and to a control group and (2) the effect of methylphenidate on the motor timing processes in the clinical group with ADHD in a double blind, cross-over, medication-placebo design, including 4 weeks of medication. The clinical group with ADHD, like the community group with hyperactivity, showed greater variability in sensorimotor synchronization and in sensorimotor anticipation relative to controls. The clinical group was also impaired in time perception, which was spared in the community group with hyperactivity. The persistent, but not the acute dose, of methylphenidate reduced the variability of sensorimotor synchronization and anticipation, but had no effect on time perception. This study shows that motor timing functions are impaired in both clinical and community children with hyperactivity. It is the first study to show the effectiveness of persistent administration of methylphenidate on deficits in motor timing in ADHD children and extends the use of methylphenidate from the domain of attentional and inhibitory functions to the domain of executive motor timing. [Abstract] [Full Text]

Rubia K, Taylor A, Taylor E, Sergeant JA.
Synchronization, anticipation, and consistency in motor timing of children with dimensionally defined attention deficit hyperactivity behaviour.
Percept Mot Skills. 1999 Dec;89(3 Pt 2):1237-58.
We tested the hypothesis that children with hyperactive behaviour are impaired in the temporal organization of their motor output. The performance of 11 boys, scoring above a cut-off on standard scales of overactivity and inattention, was compared to that of controls in progressively more complex Motor-timing tasks. The tasks administered required self-paced and externally paced Sensorimotor Synchronization and Sensorimotor Anticipation. Deficits at a perceptual level were investigated with a Time-discrimination task. As hypothesized, we found that hyperactive children had no deficits in their perception of time but were impaired in timing their motor output. Hyperactive children were more inconsistent than controls in maintaining a freely chosen tapping rhythm, in synchronizing and in anticipating their motor response to external visual stimulation. [Abstract]

Feige B, Aertsen A, Kristeva-Feige R.
Dynamic synchronization between multiple cortical motor areas and muscle activity in phasic voluntary movements.
J Neurophysiol. 2000 Nov;84(5):2622-9.
To study the functional role of synchronized neuronal activity in the human motor system, we simultaneously recorded cortical activity by high-resolution electroencephalography (EEG) and electromyographic (EMG) activity of the activated muscle during a phasic voluntary movement in seven healthy subjects. Here, we present evidence for dynamic beta-range (16-28 Hz) synchronization between cortical activity and muscle activity, starting after termination of the movement. In the same time range, increased tonic activity in the activated muscle was found. During the movement execution a low-frequency (2-14 Hz) synchronization was found. Using a novel analysis, phase-reference analysis, we were able to extract the EMG-coherent EEG maps for both, low- and high-frequency beta range synchronization. The electrical source reconstruction of the EMG-coherent EEG maps was performed with respect to the individual brain morphology from magnetic resonance imaging (MRI) using a distributed source model (cortical current density analysis) and a realistic head model. The generators of the beta-range synchronization were not only located in the primary motor area, but also in premotor areas. The generators of the low-frequency synchronization were also located in the primary motor and in premotor areas, but with additional participation of the medial premotor area. These findings suggest that the dynamic beta-range synchronization between multiple cortical areas and activated muscles reflects the transition of the collective motor network into a new equilibrium state, possibly related to higher demands on attention, while the low-frequency synchronization is related to the movement execution. [Full Text]

Kristeva-Feige R, Fritsch C, Timmer J, Lucking CH.
Effects of attention and precision of exerted force on beta range EEG-EMG synchronization during a maintained motor contraction task.
Clin Neurophysiol. 2002 Jan;113(1):124-31.
OBJECTIVE: The present study was aimed at investigating the effect of attention and precision level of exerted force on beta range EEG-EMG synchronization. METHODS: We simultaneously recorded cortical electrical activity (EEG) in a bipolar manner from the contralateral sensorimotor areas and surface electromyographic (EMG) activity from the flexor digitorum superficialis muscle in 10 healthy subjects during a maintained motor contraction task at 8% of the maximal voluntary contraction (MVC) force level. The coherence between oscillatory processes in the EEG and EMG was calculated. Three different conditions were investigated: (i) performing the task with high precision (HP); (ii) performing the task with high precision and simultaneously performing a mental arithmetic task (HPAT), i.e. attention was divided between the motor task and the mental arithmetic task; and (iii) performing the task with low precision (LP). RESULTS: We have found that the amount of beta range EEG-EMG synchronization decreases below the 95% confidence level when attention is divided between the motor task and the mental arithmetic task. The results also show that the frequency of beta range synchronization is higher with a higher level of precision but still lies within the beta frequency range (15-30 Hz). CONCLUSIONS: The data indicate that beta range synchronization represents a state of the cortico-muscular network when attention is directed towards the motor task. The frequency of synchronization of this network is associated with, and possibly encodes, precision in force production. [Abstract]

Lee KM, Ahn TB, Jeon BS, Kim DG.
Change in phase synchronization of local field potentials in anesthetized rats after chronic dopamine depletion.
Neurosci Res. 2004 Jun;49(2):179-84.
In order to investigate temporal characteristics of oscillatory neural activity and the effect of chronic dopamine depletion on it, local field potentials were measured in anesthetized rats without or with a 6-OHDA lesion at either the ventral tegmental area or the substantia nigra compacta, using a pair of electrodes that were separated by 120 microm. Coupling of neural activity in this mesoscopic scale was measured by a synchronization index that quantified the distribution of differences in instantaneous phase between the two potentials recorded. Phase synchrony was significantly stronger at deep basal ganglia sites, more so than at cortical sites, over a gamma range (30-120 Hz) in normal rats. After chronic dopamine depletion, this synchrony was no longer observable, especially after a substantia nigra lesion, while there was an increase in phase synchrony in the cortex at a lower frequency band (10 Hz). These findings are consistent with previously reported findings that the effect of dopaminergic innervation on oscillatory neural activity varies from synchronizing to desynchronizing, depending on the structure innervated and the frequency of the oscillation. [Abstract]

Brown P.
Oscillatory nature of human basal ganglia activity: relationship to the pathophysiology of Parkinson's disease.
Mov Disord. 2003 Apr;18(4):357-63.
Alterations of basal ganglia physiology in parkinsonism may consist of two elements, an increase in the firing rate of neurones and a change in the pattern of synchronisation of discharges between neurones. Recent findings suggest the presence of two principal modes of synchronised activity within the human subthalamo-pallidal-thalamo-cortical circuit, at <30 Hz and >60 Hz. These oscillations are dynamically and systematically modulated by task, thereby suggesting a functional role in movement. More importantly, the two frequency modes are inversely affected by movement, consistent with opposing actions, and differentially expressed according to the prevailing level of dopaminergic activity. It is argued that the balance between these modes determines the effects of basal ganglia-thalamocortical projections on the motor areas of the cortex. The lower frequency oscillations facilitate slow idling rhythms in the motor areas of the cortex, whereas synchronisation at high frequency restores dynamic task-related cortical ensemble activity in the gamma band. [Abstract]

Ahveninen J, Kahkonen S, Tiitinen H, Pekkonen E, Huttunen J, Kaakkola S, Ilmoniemi RJ, Jaaskelainen IP.
Suppression of transient 40-Hz auditory response by haloperidol suggests modulation of human selective attention by dopamine D2 receptors.
Neurosci Lett. 2000 Sep 29;292(1):29-32.
Cognitive processes including selective attention may depend on synchronous activity of neurons at the gamma-band (around 40Hz). To determine the effect of neuroleptic challenge on transient auditory evoked 40-Hz response, simultaneous measurement of 122-channel magnetoencephalogram (MEG) and 64-channel electroencephalogram (EEG) was used. Either 2mg of dopamine D(2)-receptor antagonist haloperidol or a placebo was administered orally to 11healthy subjects in a double-blind randomized crossover design in two separate sessions. The subjects attended to tones presented to one ear and ignored those presented to the other ear. Haloperidol significantly suppressed the transient 40-Hz electric response to the attended stimuli, while no significant effect was observed in the electric responses to the unattended tones or in the magnetic responses. The present result suggests that dopamine D(2) receptors modulate selective attention. [Abstract]

Yasumoto F, Negishi T, Ishii Y, Kyuwa S, Kuroda Y, Yoshikawa Y.
Endogenous dopamine maintains synchronous oscillation of intracellular calcium in primary cultured-mouse midbrain neurons.
Cell Mol Neurobiol. 2004 Feb;24(1):51-61.
We demonstrated synchronous oscillation of intracellular Ca2+ in cultured-mouse mid-brain neurons. This synchronous oscillation was thought to result from spontaneous and synchronous neural bursts in a synaptic neural network. We also examined the role of endogenous dopamine in neural networks showing synchronous oscillation. Immunocytochemical study revealed a few tyrosine hydroxylase (TH)-positive dopaminergic neurons, and that cultured neurons expressed synaptophysin and synapsin I. Western blot analyses comfirmed synaptophysin, TH, and 2 types of dopamine receptor (DR), D1R and D2R expression. The synchronous oscillation in midbrain neurons was abolished by the application of R(-)-2-amino-5-phosphonopentanoic acid (AP-5) as an N-methyl-D-aspartate receptor (NMDAR) antagonist. This result suggests that the synchronous oscillation in midbrain neurons requires glutamatergic transmissions, as was the case in previously reported cortical neurons. SCH-12679, a D1R antagonist, inhibited synchronous oscillation in midbrain neurons, while raclopride, a D2R antagonist, induced a transient increase of intracellular Ca2+ and inhibited synchronous oscillation. We consider that endogenous dopamine maintains synchronous oscillation of intracellular Ca2+ through D1R and D2R, and that these DRs regulate intracellular Ca2+in distinctly different ways. Synchronous oscillation of midbrain neurons would be a useful tool for in vitro researches into various neural disorders directly or indirectly caused by dopaminergic neurons. [Abstract]

Wrobel A.
Beta activity: a carrier for visual attention.
Acta Neurobiol Exp (Wars). 2000;60(2):247-60.
The alpha (8-13 Hz), beta (15-25 Hz) and gamma (30-60 Hz) bands of the EEG have been long studied clinically because of their putative functional importance. Old experimental results indicate that repetitive stimulation of the visual pathway evokes synchronous responses at the cortical level with a gain that depends on frequency; oscillations within relevant bands are less damped at subsequent processing levels than others. Our current results show in the cat that cortico-geniculate feedback has a build-in potentiation mechanism that operates at around the beta frequency and activates thalamic cells thus lowering the threshold for visual information transmission. We have also shown that enhanced beta activity is propagated along this feedback pathway solely during attentive visual behavior. This activity consists of 300-1,000 ms bursts that correlate in time with gamma oscillatory events. Beta-bursting activity spreads to all investigated visual centers, including the lateral posterior and pulvinar complex and higher cortical areas. Other supporting data are discussed that are concerned with the enhanced beta activity during attentive-like behavior of various species, including humans. Finally, we put forward a general hypothesis which attributes the appearance of oscillations within the alpha, beta and gamma bands to different activation states of the visual system. According to this hypothesis, alpha activity characterizes idle arousal of the system, while beta bursts shift the system to an attention state that consequently allows for gamma synchronization and perception. [Abstract] [PDF]

Aston-Jones G, Rajkowski J, Cohen J.
Role of locus coeruleus in attention and behavioral flexibility.
Biol Psychiatry 1999 Nov 1;46(9):1309-20
"Previous findings have implicated the noradrenergic locus coeruleus (LC) system in functions along the dimension of arousal or attention. It has remained uncertain what role this system has in attention, or what mechanisms may be involved. We review our recent work examining activity of LC neurons in monkeys performing a visual discrimination task that requires focused attention. Results indicate that LC cells exhibit phasic or tonic modes of activity, that closely correspond to good or poor performance on this task, respectively. A computational model was used to simulate these results. This model predicts that alterations in electrotonic coupling among LC cells may produce the different modes of activity and corresponding differences in performance. This model also indicates that the phasic mode of LC activity may promote focused or selective attention, whereas the tonic mode may produce a state of high behavioral flexibility or scanning attentiveness. The implications of these results for clinical disorders such as attention-deficit hyperactivity disorder, stress disorders, and emotional and affective disorders are discussed." [Abstract]

Editor's note: if both norepinephrine and beta range synchronization are involved in attention, then what is the role of norepinephrine in beta range synchronization? How does dopamine influence beta range synchronization? Several genetic polymorphisms that may increase susceptibility to ADHD influence noradrenergic and/or dopaminergic systems; see my compilation of ADHD genetic research elsewhere on for a literature review.

Singer W.
Consciousness and the binding problem.
Ann N Y Acad Sci. 2001 Apr;929:123-46.
It is proposed that phenomenal awareness, the ability to be aware of one's sensations and feelings, emerges from the capacity of evolved brains to analyze their own cognitive processes by iterating and reapplying on them-selves the very same cortical operations that they use for the interpretation of signals from the outer world. Search for the neuronal substrate of awareness therefore converges with the search for the cognitive mechanisms through which brains analyze their environment. The hypothesis is put forward that the mammalian brain generates continuously highly dynamic states that, when modulated by input signals, rapidly converge towards points of transient stability that correspond to the respective input constellation. It is proposed that these states are characterized by the dynamic binding of feature-specific cells into functionally coherent cell assemblies which as a whole represent the constellation of features defining a particular perceptual object. Arguments are presented that favor the notion that the cognitive operations supporting awareness consist of an iteration of such dynamic binding processes which then lead to the formation of higher-order assemblies that correspond to the contents of conscious awareness. Experimental data are reviewed relating to the questions of how assemblies are formed and which signatures define the relations among the responses of distributed neurons. It is argued that assemblies self-organize through reciprocal interactions of neurons coupled by reentrant loops and that the signature of relatedness consists of the transient synchronization of the discharges of the respective neurons. Evidence is presented that these synchronization phenomena depend on the same state variables as awareness: Both require for their manifestation activated brain states characterized by desynchronized EEGs. It is concluded that phenomenal awareness is amenable to neurobiological reductionism; but it is also proposed that self-consciousness requires a different explanatory approach because it emerges from the dialogue between different brains and hence has the quality of a cultural construct.

Attentional mechanisms could impose a coherent subthreshold modulation on neurons in cortical areas that need to participate in the execution of the anticipated task and thereby permit rapid synchronization of selected responses using the synchronizing mechanisms described above. According to this scenario, the attentional mechanisms would induce what one might call a state of expectancy in the respective cortical areas by imposing on them a specific, task related dynamic activation pattern which then, once stimulus-driven input becomes available, acts like a dynamic filter that causes rapid synchronization of selected responses, thereby accomplishing the required grouping and binding of responses and in addition assuring rapid transmission. [PDF]

Gross J, Schmitz F, Schnitzler I, Kessler K, Shapiro K, Hommel B, Schnitzler A.
Modulation of long-range neural synchrony reflects temporal limitations of visual attention in humans.
Proc Natl Acad Sci U S A. 2004 Aug 31;101(35):13050-5. Epub 2004 Aug 24.
Because of attentional limitations, the human visual system can process for awareness and response only a fraction of the input received. Lesion and functional imaging studies have identified frontal, temporal, and parietal areas as playing a major role in the attentional control of visual processing, but very little is known about how these areas interact to form a dynamic attentional network. We hypothesized that the network communicates by means of neural phase synchronization, and we used magnetoencephalography to study transient long-range interarea phase coupling in a well studied attentionally taxing dual-target task (attentional blink). Our results reveal that communication within the fronto-parieto-temporal attentional network proceeds via transient long-range phase synchronization in the beta band. Changes in synchronization reflect changes in the attentional demands of the task and are directly related to behavioral performance. Thus, we show how attentional limitations arise from the way in which the subsystems of the attentional network interact. [Abstract] [PDF]

Gomez CM, Vazquez M, Vaquero E, Lopez-Mendoza D, Cardoso MJ.
Frequency analysis of the EEG during spatial selective attention.
Int J Neurosci. 1998 Jul;95(1-2):17-32.
In this study, we recorded the event-related potentials (ERPs) elicited by stimuli appearing at attended and unattended locations. The voltage amplitudes and latencies of the P1, N1, P2, N2 and P3 visual components showed statistically significant differences in the attended condition with respect to the unattended one. The power spectral density of the EEG following stimulus onset was calculated. The difference between the spectral densities of the attended and unattended conditions was computed. Statistically significant differences were found in the decrease of alpha (9-11 Hz) and the increase of beta (15-17 Hz) frequencies during the attention condition with respect to the unattended condition. These results suggest that the arrival of a visual stimulus during the attended condition generates a complex reorganization of neuronal activity in both time and frequency domains.

Bekisz M, Wrobel A.
Attention-dependent coupling between beta activities recorded in the cat's thalamic and cortical representations of the central visual field.
Eur J Neurosci. 2003 Jan;17(2):421-6.
We have previously proposed that enhanced 16-24 Hz (beta) local field potential activity in the primary visual cortex and lateral geniculate nucleus may be an electrophysiological correlate of the attentional mechanism that increases the gain of afferent visual information flow to the cortex. In this study, we measured coupling between beta signals recorded in the thalamic (i.e. lateral geniculate or perigeniculate) and cortical representations of the central visual field (within 5 degrees from area centralis), during visual and auditory attentive situations. Signal coupling was calculated in two ways: (i) by means of crosscorrelation between raw beta activities, which depends primarily on phase coherence, and (ii) by phase-independent crosscorrelation between amplitude envelopes of beta activities. Mean amplitudes of raw signal cross correlations obtained for thalamo-cortical recording pairs were not significantly different when calculated during behavioural demands for either visual or auditory attention. In contrast, amplitudes of envelope cross correlations obtained during behaviour requiring visual attention were, on average, two times higher than those calculated during the auditory task. This attention-related coupling emerged from synchronized amplitude modulation of beta oscillatory activity that occurs within the cortico-thalamic circuit involved in central vision. [Abstract]

Olufsen MS, Whittington MA, Camperi M, Kopell N.
New roles for the gamma rhythm: population tuning and preprocessing for the Beta rhythm.
J Comput Neurosci. 2003 Jan-Feb;14(1):33-54.
Gamma (30-80 Hz) and beta (12-30 Hz) oscillations such as those displayed by in vitro hippocampal (CA1) slice preparations and by in vivo neocortical EEGs often occur successively, with a spontaneous transition between them. In the gamma rhythm, pyramidal cells fire together with the interneurons, while in the beta rhythm, pyramidal cells fire on a subset of cycles of the interneurons. It is shown that gamma and beta rhythms have different properties with respect to creation of cell assemblies. In the presence of heterogeneous inputs to the pyramidal cells, the gamma rhythm creates an assembly of firing pyramidal cells from cells whose drive exceeds a threshold. During the gamma to beta transition, a slow outward potassium current is activated, and as a result the cell assembly vanishes. The slow currents make each of the pyramidal cells fire with a beta rhythm, but the field potential of the network still displays a gamma rhythm. Hebbian changes of connections among the pyramidal cells give rise to a beta rhythm, and the cell assemblies are recovered with a temporal separation between cells firing in different cycles. We present experimental evidence showing that such a separation can occur in hippocampal slices. [Abstract] [PDF]

Bibbig A, Traub RD, Whittington MA.
Long-range synchronization of gamma and beta oscillations and the plasticity of excitatory and inhibitory synapses: a network model.
J Neurophysiol. 2002 Oct;88(4):1634-54.
The ability of oscillating networks to synchronize despite significant separation in space, and thus time, is of biological significance, given that human gamma activity can synchronize over distances of several millimeters to centimeters during perceptual and learning tasks. We use computer simulations of networks consisting of excitatory pyramidal cells (e-cells) and inhibitory interneurons (i-cells), modeling two tonically driven assemblies separated by large (>or=8 ms) conduction delays. The results are as follows. 1) Two assemblies separated by large conduction delays can fire synchronously at beta frequency (with i-cells firing at gamma frequency) under two timing conditions: e-cells of (say) assembly 2 are still inhibited "delay + spike generation milliseconds" after the e-cell beat of assembly 1; this means that the e-cell inhibitory postsynaptic potential (IPSP) cannot be significantly shorter than the delay (2-site effect). This implies for a given decay time constant that the interneuron --> pyramidal cell conductances must be large enough. The e-cell IPSP must last longer than the i-cell IPSP, i.e., the interneuron --> pyramidal cell conductance must be sufficiently large and the interneuron --> interneuron conductance sufficiently small (local effect). 2) We define a "long-interval doublet" as a pair of interneuron action potentials-separated by approximately "delay milliseconds"-in which a) the first spike is induced by tonic inputs and/or excitation from nearby e-cells, while b) the second spike is induced by (delayed) excitation from distant e-cells. "Long-interval population doublets" (long-interval doublets of the i-cell population) are necessary for synchronized firing in our networks. Failure to produce them leads to almost anti-phase activity at gamma frequency. 3) An (almost) anti-phase oscillation is the most stable oscillation pattern of two assemblies that are separated by axonal conduction delays of approximately one-half a gamma period (delays from 8 to 17 ms in our simulations) and that are firing at gamma frequency. 4) Two assemblies separated by large conduction delays can synchronize their activity with the help of interneuron plasticity. They can also synchronize without pyramidal cell --> pyramidal cell connections being present. The presence of pyramidal cell --> pyramidal cell connections allows, however, for synchronization if other parameters are at inappropriate values for synchronization to occur. 5) Synchronization of two assemblies separated by large conduction delays with the help of interneuron plasticity is not simply due to slowing down of the oscillation frequency. It is reached with the help of a "synchronizing-weak-beat," which induces sudden changes in the oscillation period length of the two assemblies. [Full Text]

Brovelli A, Ding M, Ledberg A, Chen Y, Nakamura R, Bressler SL.
Beta oscillations in a large-scale sensorimotor cortical network: directional influences revealed by Granger causality.
Proc Natl Acad Sci U S A. 2004 Jun 29;101(26):9849-54. Epub 2004 Jun 21.
Previous studies have shown that synchronized beta frequency (14-30 Hz) oscillations in the primary motor cortex are involved in maintaining steady contractions of contralateral arm and hand muscles. However, little is known about the role of postcentral cortical areas in motor maintenance and their patterns of interaction with motor cortex. We investigated the functional relations of beta-synchronized neuronal assemblies in pre- and postcentral areas of two monkeys as they pressed a hand lever during the wait period of a visual discrimination task. By using power and coherence spectral analysis, we identified a beta-synchronized large-scale network linking pre- and postcentral areas. We then used Granger causality spectra to measure directional influences among recording sites. In both monkeys, strong Granger causal influences were observed from primary somatosensory cortex to both motor cortex and inferior posterior parietal cortex, with the latter area also exerting Granger causal influences on motor cortex. Granger causal influences from motor cortex to postcentral sites, however, were weak in one monkey and not observed in the other. These results are the first, to our knowledge, to demonstrate in awake monkeys that synchronized beta oscillations bind multiple sensorimotor areas into a large-scale network during motor maintenance behavior and carry Granger causal influences from primary somatosensory and inferior posterior parietal cortices to motor cortex. [Full Text]

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