On the Physiological Modulation and Potential Mechanisms Underlying Parieto-Occipital Alpha Oscillations

The parieto-occipital alpha (8–13 Hz) rhythm is by far the strongest spectral fingerprint in the human brain. Almost 90 years later, its physiological origin is still far from clear. In this Research Topic I review human pharmacological studies using electroencephalography (EEG) and magnetoencephalography (MEG) that investigated the physiological mechanisms behind posterior alpha. Based on results from classical and recent experimental studies, I find a wide spectrum of drugs that modulate parieto-occipital alpha power. Alpha frequency is rarely affected, but this might be due to the range of drug dosages employed. Animal and human pharmacological findings suggest that both GABA enhancers and NMDA blockers systematically decrease posterior alpha power. Surprisingly, most of the theoretical frameworks do not seem to embrace these empirical findings and the debate on the functional role of alpha oscillations has been polarized between the inhibition vs. active poles hypotheses. Here, I speculate that the functional role of alpha might depend on physiological excitation as much as on physiological inhibition. This is supported by animal and human pharmacological work showing that GABAergic, glutamatergic, cholinergic, and serotonergic receptors in the thalamus and the cortex play a key role in the regulation of alpha power and frequency. This myriad of physiological modulations fit with the view that the alpha rhythm is a complex rhythm with multiple sources supported by both thalamo-cortical and cortico-cortical loops. Finally, I briefly discuss how future research combining experimental measurements derived from theoretical predictions based of biophysically realistic computational models will be crucial to the reconciliation of these disparate findings.

[1]  G. Buzsáki,et al.  Analysis of gamma rhythms in the rat hippocampus in vitro and in vivo. , 1996, The Journal of physiology.

[2]  W. Sannita,et al.  Effects of scopolamine (0.25-0.75 mg i.m.) on the quantitative EEG and the neuropsychological status of healthy volunteers. , 1987, Neuropsychobiology.

[3]  Mark Turmaine,et al.  Thalamic Gap Junctions Control Local Neuronal Synchrony and Influence Macroscopic Oscillation Amplitude during EEG Alpha Rhythms , 2011, Front. Psychology.

[4]  S. Taulu,et al.  Applications of the signal space separation method , 2005, IEEE Transactions on Signal Processing.

[5]  Nancy Kopell Does it have to be this complicated? Focus on "Single-column thalamocortical network model exhibiting gamma oscillations, spindles, and epileptogenic bursts". , 2005, Journal of neurophysiology.

[6]  C. Schroeder,et al.  Neuronal Mechanisms of Cortical Alpha Oscillations in Awake-Behaving Macaques , 2008, The Journal of Neuroscience.

[7]  P. Robinson,et al.  Prediction of electroencephalographic spectra from neurophysiology. , 2001, Physical review. E, Statistical, nonlinear, and soft matter physics.

[8]  Vincenzo Crunelli,et al.  Cellular Dynamics of Cholinergically Induced α (8–13 Hz) Rhythms in Sensory Thalamic Nuclei In Vitro , 2008, The Journal of Neuroscience.

[9]  M. Carandini,et al.  Orientation tuning of input conductance, excitation, and inhibition in cat primary visual cortex. , 2000, Journal of neurophysiology.

[10]  D. Javitt,et al.  Global dynamics of selective attention and its lapses in primary auditory cortex , 2016, Nature Neuroscience.

[11]  Robert Oostenveld,et al.  Localizing human visual gamma-band activity in frequency, time and space , 2006, NeuroImage.

[12]  F. L. D. Silva,et al.  Basic mechanisms of cerebral rhythmic activities , 1990 .

[13]  Keisuke Fukuda,et al.  α Power Modulation and Event-Related Slow Wave Provide Dissociable Correlates of Visual Working Memory , 2015, The Journal of Neuroscience.

[14]  Mingzhou Ding,et al.  Attentional Modulation of Alpha Oscillations in Macaque Inferotemporal Cortex , 2011, The Journal of Neuroscience.

[15]  Y. Kawaguchi,et al.  Serotonin Modulates Fast-Spiking Interneuron and Synchronous Activity in the Rat Prefrontal Cortex through 5-HT1A and 5-HT2A Receptors , 2010, The Journal of Neuroscience.

[16]  B. Saletu,et al.  On central effects of serotonin re-uptake inhibitors: Quantitative EEG and psychometric studies with sertraline and zimelidine , 2005, Journal of Neural Transmission.

[17]  H. Semlitsch,et al.  PHARMACODYNAMICS OF VENLAFAXINE EVALUATED BY EEG BRAIN MAPPING, PSYCHOMETRY AND PSYCHOPHYSIOLOGY. , 1992, British journal of clinical pharmacology.

[18]  M. Danhof,et al.  A comparison of the concentration-effect relationships of midazolam for EEG-derived parameters and saccadic peak velocity. , 1993, British journal of clinical pharmacology.

[19]  W. Klimesch Alpha-band oscillations, attention, and controlled access to stored information , 2012, Trends in Cognitive Sciences.

[20]  Nikolaus Weiskopf,et al.  Flexible head-casts for high spatial precision MEG , 2017, Journal of Neuroscience Methods.

[21]  Emery N. Brown,et al.  Electroencephalogram signatures of loss and recovery of consciousness from propofol , 2013, Proceedings of the National Academy of Sciences.

[22]  R. VanRullen,et al.  Ongoing EEG Phase as a Trial-by-Trial Predictor of Perceptual and Attentional Variability , 2011, Front. Psychology.

[23]  P. Celada,et al.  The Hallucinogen DOI Reduces Low-Frequency Oscillations in Rat Prefrontal Cortex: Reversal by Antipsychotic Drugs , 2008, Biological Psychiatry.

[24]  D. Leopold,et al.  Ongoing Alpha Activity in V1 Regulates Visually Driven Spiking Responses , 2017, Cerebral cortex.

[25]  Robert Oostenveld,et al.  Neuronal Oscillations with Non-sinusoidal Morphology Produce Spurious Phase-to-Amplitude Coupling and Directionality , 2016, Front. Comput. Neurosci..

[26]  A. Craig,et al.  TIME COURSE AND REGIONAL DISTRIBUTION OF CORTICAL CHANGES DURING ACUTE ALCOHOL INGESTION , 2004, The International journal of neuroscience.

[27]  A. Engel,et al.  Cortical Hypersynchrony Predicts Breakdown of Sensory Processing during Loss of Consciousness , 2011, Current Biology.

[28]  Wolf Singer,et al.  Ketamine Dysregulates the Amplitude and Connectivity of High-Frequency Oscillations in Cortical-Subcortical Networks in Humans: Evidence From Resting-State Magnetoencephalography-Recordings. , 2015, Schizophrenia bulletin.

[29]  Saskia Haegens,et al.  Laminar Profile and Physiology of the α Rhythm in Primary Visual, Auditory, and Somatosensory Regions of Neocortex , 2015, The Journal of Neuroscience.

[30]  Ole Jensen,et al.  A biologically plausible mechanism for neuronal coding organized by the phase of alpha oscillations , 2016, The European journal of neuroscience.

[31]  C. Rosenberg,et al.  Electroencephalography: Basic Principles, Clinical Applications, and Related Fields, 3rd Ed. , 1994 .

[32]  G Pfurtscheller,et al.  Event-related desynchronization (ERD) during visual processing. , 1994, International journal of psychophysiology : official journal of the International Organization of Psychophysiology.

[33]  R. Shapley,et al.  Stochastic Generation of Gamma-Band Activity in Primary Visual Cortex of Awake and Anesthetized Monkeys , 2012, The Journal of Neuroscience.

[34]  R. VanRullen,et al.  An oscillatory mechanism for prioritizing salient unattended stimuli , 2012, Trends in Cognitive Sciences.

[35]  J. Lisman,et al.  Oscillations in the alpha band (9-12 Hz) increase with memory load during retention in a short-term memory task. , 2002, Cerebral cortex.

[36]  B. Connors,et al.  Intrinsic oscillations of neocortex generated by layer 5 pyramidal neurons. , 1991, Science.

[37]  F. D. da Silva,et al.  Organization of thalamic and cortical alpha rhythms: spectra and coherences. , 1973, Electroencephalography and clinical neurophysiology.

[38]  J. Palva,et al.  Functional Roles of Alpha-Band Phase Synchronization in Local and Large-Scale Cortical Networks , 2011, Front. Psychology.

[39]  P. Anderer,et al.  Effects of the South American Psychoactive Beverage Ayahuasca on Regional Brain Electrical Activity in Humans: A Functional Neuroimaging Study Using Low-Resolution Electromagnetic Tomography , 2004, Neuropsychobiology.

[40]  Anticholinergic drug effects on quantitative electroencephalogram, visual evoked potential, and verbal memory , 1992, Biological Psychiatry.

[41]  R. Lorente de Nó,et al.  Action potential of the motoneurons of the hypoglossus nucleus. , 1947, Journal of cellular and comparative physiology.

[42]  Ernst Fernando Lopes Da Silva Niedermeyer,et al.  Electroencephalography, basic principles, clinical applications, and related fields , 1982 .

[43]  Emiliano Ricciardi,et al.  Cholinergic enhancement differentially modulates neural response to encoding during face identity and face location working memory tasks , 2013, Experimental biology and medicine.

[44]  P. Roelfsema,et al.  Alpha and gamma oscillations characterize feedback and feedforward processing in monkey visual cortex , 2014, Proceedings of the National Academy of Sciences.

[45]  Alexander Thiele,et al.  Muscarinic signaling in the brain. , 2013, Annual review of neuroscience.

[46]  Verner J. Knott,et al.  The Cholinergic Basis of the Smoking-Induced EEG Activation Profile , 1998, Neuropsychobiology.

[47]  Alexander L. Green Cortical Oscillations in Health and Disease. Oxford Univ. Press, New York (2010), June, Color plates, Hard cover, 448 pp, $74.95., ISBN: 978-0-19-534279-6 , 2010 .

[48]  Dominique L. Pritchett,et al.  Quantitative analysis and biophysically realistic neural modeling of the MEG mu rhythm: rhythmogenesis and modulation of sensory-evoked responses. , 2009, Journal of neurophysiology.

[49]  A. Thiele,et al.  Attention – oscillations and neuropharmacology , 2009, The European journal of neuroscience.

[50]  F. H. Lopes da Silva,et al.  The cortical source of the alpha rhythm. , 1977, Neuroscience letters.

[51]  S. Hughes,et al.  Thalamic Mechanisms of EEG Alpha Rhythms and Their Pathological Implications , 2005, The Neuroscientist : a review journal bringing neurobiology, neurology and psychiatry.

[52]  R. Traub,et al.  Synchronized oscillations in interneuron networks driven by metabotropic glutamate receptor activation , 1995, Nature.

[53]  O. Jensen,et al.  Shaping Functional Architecture by Oscillatory Alpha Activity: Gating by Inhibition , 2010, Front. Hum. Neurosci..

[54]  Chun-I Yeh,et al.  Laminar analysis of visually evoked activity in the primary visual cortex , 2012, Proceedings of the National Academy of Sciences.

[55]  Judith E. Hall,et al.  Enhanced Stimulus-Induced Gamma Activity in Humans during Propofol-Induced Sedation , 2013, PLoS ONE.

[56]  J. Palva,et al.  New vistas for α-frequency band oscillations , 2007, Trends in Neurosciences.

[57]  João Ricardo Sato,et al.  Acute Biphasic Effects of Ayahuasca , 2015, PloS one.

[58]  Krish D. Singh,et al.  Significant reductions in human visual gamma frequency by the gaba reuptake inhibitor tiagabine revealed by robust peak frequency estimation , 2016, Human brain mapping.

[59]  Terrence J. Sejnowski,et al.  Neuronal Synchronization and Thalamocortical Rhythms in Sleep, Wake and Epilepsy , 2012 .

[60]  P. Denise,et al.  Localization of Scopolamine Induced Electrocortical Brain Activity Changes, in Healthy Humans at Rest , 2013, Journal of clinical pharmacology.

[61]  P. Dayan Fast oscillations in cortical circuits , 2000 .

[62]  A. Bond,et al.  A comparison of the psychotropic profiles of tofisopam and diazepam , 2004, European Journal of Clinical Pharmacology.

[63]  Th R Knösche,et al.  Transformation of Whole-Head MEG Recordings Between Different Sensor Positions / Transformation von Ganzkopf-MEG-Messungen zwischen verschiedenen Sensorpositionen , 2002, Biomedizinische Technik. Biomedical engineering.

[64]  B. Orser,et al.  The General Anesthetic Propofol Slows Deactivation and Desensitization of GABAA Receptors , 1999, The Journal of Neuroscience.

[65]  J. M. Hupé,et al.  Cortical feedback improves discrimination between figure and background by V1, V2 and V3 neurons , 1998, Nature.

[66]  G. Barnes,et al.  Neuronal network pharmacodynamics of GABAergic modulation in the human cortex determined using pharmaco‐magnetoencephalography , 2009, Human brain mapping.

[67]  Eric Maris,et al.  Rhythmic Components in Extracranial Brain Signals Reveal Multifaceted Task Modulation of Overlapping Neuronal Activity , 2016, PloS one.

[68]  Judith E. Hall,et al.  Evidence that Subanesthetic Doses of Ketamine Cause Sustained Disruptions of NMDA and AMPA-Mediated Frontoparietal Connectivity in Humans , 2015, The Journal of Neuroscience.

[69]  R. Romo,et al.  α-Oscillations in the monkey sensorimotor network influence discrimination performance by rhythmical inhibition of neuronal spiking , 2011, Proceedings of the National Academy of Sciences.

[70]  Petroc Sumner,et al.  Acute Effects of Alcohol on Stimulus-Induced Gamma Oscillations in Human Primary Visual and Motor Cortices , 2014, Neuropsychopharmacology.

[71]  L. Soufflet,et al.  Pharmacodynamic Effects of Acamprosate on Markers of Cerebral Function in Alcohol-Dependent Subjects Administered as Pretreatment and during Alcohol Abstinence , 2004, Neuropsychobiology.

[72]  Verner Knott,et al.  Effects of Ketamine on Resting-State EEG Activity and Their Relationship to Perceptual/Dissociative Symptoms in Healthy Humans , 2016, Front. Pharmacol..

[73]  Y. Okada,et al.  Contributions of principal neocortical neurons to magnetoencephalography and electroencephalography signals , 2006, The Journal of physiology.

[74]  J. Gates,et al.  Benzodiazepines in epilepsy: pharmacology and pharmacokinetics , 2008, Acta neurologica Scandinavica.

[75]  V. Knott,et al.  COMT polymorphism modulates the resting‐state EEG alpha oscillatory response to acute nicotine in male non‐smokers , 2015, Genes, brain, and behavior.

[76]  C G Link,et al.  Effects of granisetron and lorazepam, alone and in combination, on the EEG of human volunteers. , 1991, British journal of clinical pharmacology.

[77]  V. Feshchenko,et al.  Propofol-Induced Alpha Rhythm , 2004, Neuropsychobiology.

[78]  John J. Foxe,et al.  The Role of Alpha-Band Brain Oscillations as a Sensory Suppression Mechanism during Selective Attention , 2011, Front. Psychology.

[79]  E. Seifritz,et al.  Psilocybin-induced spiritual experiences and insightfulness are associated with synchronization of neuronal oscillations , 2015, Psychopharmacology.

[80]  Julian Keil,et al.  The role of alpha oscillations for illusory perception , 2014, Behavioural Brain Research.

[81]  Verner Knott,et al.  Acute Nicotine Administration in Alzheimer’s Disease: An Exploratory EEG Study , 2000, Neuropsychobiology.

[82]  W. Klimesch,et al.  EEG alpha oscillations: The inhibition–timing hypothesis , 2007, Brain Research Reviews.

[83]  Lutz Jäncke,et al.  Activation of Serotonin 2A Receptors Underlies the Psilocybin-Induced Effects on α Oscillations, N170 Visual-Evoked Potentials, and Visual Hallucinations , 2013, The Journal of Neuroscience.

[84]  Jonas Obleser,et al.  Alpha Rhythms in Audition: Cognitive and Clinical Perspectives , 2011, Front. Psychology.

[85]  Robert Oostenveld,et al.  Online and offline tools for head movement compensation in MEG , 2013, NeuroImage.

[86]  E. Brown,et al.  Thalamocortical Mechanisms for the Anteriorization of Alpha Rhythms during Propofol-Induced Unconsciousness , 2013, The Journal of Neuroscience.

[87]  J. F. Alonso,et al.  Inhibition of alpha oscillations through serotonin-2A receptor activation underlies the visual effects of ayahuasca in humans , 2016, European Neuropsychopharmacology.

[88]  B. Connors,et al.  Two types of network oscillations in neocortex mediated by distinct glutamate receptor subtypes and neuronal populations. , 1996, Journal of neurophysiology.

[89]  O. Jensen,et al.  Asymmetric Amplitude Modulations of Brain Oscillations Generate Slow Evoked Responses , 2008, The Journal of Neuroscience.

[90]  E. Domino,et al.  Tobacco smoking produces widespread dominant brain wave alpha frequency increases. , 2009, International journal of psychophysiology : official journal of the International Organization of Psychophysiology.

[91]  B. Antkowiak,et al.  Different actions of general anesthetics on the firing patterns of neocortical neurons mediated by the GABA(A) receptor. , 1999, Anesthesiology.

[92]  R. Ilmoniemi,et al.  The interplay of lorazepam-induced brain oscillations: microstructural electromagnetic study , 2004, Clinical Neurophysiology.

[93]  Nico Bunzeck,et al.  Dopaminergic stimulation facilitates working memory and differentially affects prefrontal low theta oscillations , 2014, NeuroImage.

[94]  J. L. Slangen,et al.  Effects of oxazepam on event-related brain potentials, EEG frequency bands, and vigilance performance , 1995, Psychopharmacology.

[95]  A. T. Gulledge,et al.  Serotonin and Prefrontal Cortex Function: Neurons, Networks, and Circuits , 2011, Molecular Neurobiology.

[96]  H. Kennedy,et al.  Visual Areas Exert Feedforward and Feedback Influences through Distinct Frequency Channels , 2014, Neuron.

[97]  Mikko Pohja,et al.  On the human sensorimotor-cortex beta rhythm: Sources and modeling , 2005, NeuroImage.

[98]  D. Albe-Fessard,et al.  Electrophysiological studies of some deep cerebral structures in man. , 1966, Journal of the neurological sciences.

[99]  Bethany Routley,et al.  The effects of AMPA blockade on the spectral profile of human early visual cortex recordings studied with non-invasive MEG , 2016, Cortex.

[100]  E. Brown,et al.  Thalamocortical model for a propofol-induced α-rhythm associated with loss of consciousness , 2010, Proceedings of the National Academy of Sciences.

[101]  Bart Gips,et al.  Temporal coding organized by coupled alpha and gamma oscillations prioritize visual processing , 2014, Trends in Neurosciences.

[102]  Tali Leibovich,et al.  Quantities, Amounts, and the Numerical Core System , 2012, Front. Hum. Neurosci.

[103]  Emery N. Brown,et al.  Tracking brain states under general anesthesia by using global coherence analysis , 2011, Proceedings of the National Academy of Sciences.

[104]  Wolfgang Klimesch,et al.  Prestimulus amplitudes modulate P1 latencies and evoked traveling alpha waves , 2015, Front. Hum. Neurosci..

[105]  Judith E. Hall,et al.  Ketamine amplifies induced gamma frequency oscillations in the human cerebral cortex , 2015, European Neuropsychopharmacology.

[106]  S. Cole,et al.  Brain Oscillations and the Importance of Waveform Shape , 2017, Trends in Cognitive Sciences.

[107]  D. Leopold,et al.  Layer-Specific Entrainment of Gamma-Band Neural Activity by the Alpha Rhythm in Monkey Visual Cortex , 2012, Current Biology.

[108]  Roshan Cools,et al.  GABAergic Modulation of Visual Gamma and Alpha Oscillations and Its Consequences for Working Memory Performance , 2014, Current Biology.

[109]  R. Ilmoniemi,et al.  Magnetoencephalography-theory, instrumentation, and applications to noninvasive studies of the working human brain , 1993 .

[110]  N. Jaworska,et al.  Effects of acute nicotine administration on behavioral and neural (EEG) correlates of working memory in non-smokers , 2012, Brain Research.

[111]  Rufin VanRullen,et al.  Visual Trails: Do the Doors of Perception Open Periodically? , 2011, PLoS biology.

[112]  Mathias Schreckenberger,et al.  The thalamus as the generator and modulator of EEG alpha rhythm: a combined PET/EEG study with lorazepam challenge in humans , 2004, NeuroImage.

[113]  F. H. Lopes da Silva,et al.  Relative contributions of intracortical and thalamo-cortical processes in the generation of alpha rhythms, revealed by partial coherence analysis. , 1980, Electroencephalography and clinical neurophysiology.

[114]  Robert Oostenveld,et al.  Oscillatory dynamics of response competition in human sensorimotor cortex , 2013, NeuroImage.

[115]  Robert Oostenveld,et al.  Competitive interactions in sensorimotor cortex: oscillations express separation between alternative movement targets. , 2014, Journal of neurophysiology.

[116]  H. Berger Über das Elektrenkephalogramm des Menschen , 1938, Archiv für Psychiatrie und Nervenkrankheiten.

[117]  A Babloyantz,et al.  A model of the inward current Ih and its possible role in thalamocortical oscillations. , 1993, Neuroreport.

[118]  Mathias Schreckenberger,et al.  Anterior limbic alpha-like activity: a low resolution electromagnetic tomography study with lorazepam challenge , 2005, Clinical Neurophysiology.

[119]  Gustavo Deco,et al.  The Neuronal Basis of Attention: Rate versus Synchronization Modulation , 2008, The Journal of Neuroscience.

[120]  Ole Jensen,et al.  Alpha Oscillations Correlate with the Successful Inhibition of Unattended Stimuli , 2011, Journal of Cognitive Neuroscience.

[121]  N. Jaworska,et al.  Effects of acute nicotine administration on resting EEG in nonsmokers. , 2012, Experimental and clinical psychopharmacology.

[122]  D. Millett Hans Berger: From Psychic Energy to the EEG , 2001, Perspectives in biology and medicine.

[123]  M. Scanziani,et al.  Instantaneous Modulation of Gamma Oscillation Frequency by Balancing Excitation with Inhibition , 2009, Neuron.

[124]  D. Liley,et al.  Drug-induced modification of the system properties associated with spontaneous human electroencephalographic activity. , 2003, Physical review. E, Statistical, nonlinear, and soft matter physics.

[125]  菊知 充 EEG changes following scopolamine administration in healthy subjects : quantitative analysis during rest and photic stimulation , 1999 .

[126]  W. Kirch,et al.  Pharmacokinetic‐Pharmacodynamic Modeling of the Electroencephalogram Effects of Scopolamine in Healthy Volunteers , 2001, Journal of clinical pharmacology.

[127]  R. Dolan,et al.  Cholinergic Enhancement of Visual Attention and Neural Oscillations in the Human Brain , 2012, Current Biology.

[128]  George A. Mashour,et al.  Neurophysiologic Correlates of Ketamine Sedation and Anesthesia: A High-density Electroencephalography Study in Healthy Volunteers , 2017, Anesthesiology.

[129]  P. Fries Rhythms for Cognition: Communication through Coherence , 2015, Neuron.

[130]  Oral intake of γ-aminobutyric acid affects mood and activities of central nervous system during stressed condition induced by mental tasks , 2012, Amino Acids.

[131]  M. D’Esposito,et al.  Inverted-U–Shaped Dopamine Actions on Human Working Memory and Cognitive Control , 2011, Biological Psychiatry.

[132]  Á. Pascual-Leone,et al.  α-Band Electroencephalographic Activity over Occipital Cortex Indexes Visuospatial Attention Bias and Predicts Visual Target Detection , 2006, The Journal of Neuroscience.

[133]  Alan Gevins,et al.  Tracking the Cognitive Pharmacodynamics of Psychoactive Substances with Combinations of Behavioral and Neurophysiological Measures , 2002, Neuropsychopharmacology.

[134]  D. Gilbert,et al.  Effects of nicotine and caffeine, separately and in combination, on EEG topography, mood, heart rate, cortisol, and vigilance. , 2000, Psychophysiology.

[135]  S. Andersson,et al.  Physiological basis of the alpha rhythm , 1968 .

[136]  G. V. Simpson,et al.  Anticipatory Biasing of Visuospatial Attention Indexed by Retinotopically Specific α-Bank Electroencephalography Increases over Occipital Cortex , 2000, The Journal of Neuroscience.

[137]  Ole Jensen,et al.  On the relationship between cortical excitability and visual oscillatory responses — A concurrent tDCS–MEG study , 2016, NeuroImage.

[138]  Monica Rojas-Martínez,et al.  Evaluation of multiple comparison correction procedures in drug assessment studies using LORETA maps , 2015, Medical & Biological Engineering & Computing.

[139]  Kevin Murphy,et al.  Neural correlates of the LSD experience revealed by multimodal neuroimaging , 2016, Proceedings of the National Academy of Sciences.

[140]  Ole Jensen,et al.  Spatiotemporal Dynamics of Cortical Representations during and after Stimulus Presentation , 2016, Front. Syst. Neurosci..

[141]  V. Knott,et al.  Effects of Haloperidol Pretreatment on the Smoking-Induced EEG/Mood Activation Response Profile , 2001, Neuropsychobiology.

[142]  D. Liley,et al.  Modeling the effects of anesthesia on the electroencephalogram. , 2005, Physical review. E, Statistical, nonlinear, and soft matter physics.

[143]  Emery N. Brown,et al.  The BRAIN Initiative: developing technology to catalyse neuroscience discovery , 2015, Philosophical Transactions of the Royal Society B: Biological Sciences.

[144]  F. L. D. Silva,et al.  Dynamics of the human alpha rhythm: evidence for non-linearity? , 1999, Clinical Neurophysiology.

[145]  Con Stough,et al.  The effects of nicotine on the 13 Hz steady-state visually evoked potential , 2000, Clinical Neurophysiology.

[146]  G. V. Simpson,et al.  Parieto‐occipital ∼1 0Hz activity reflects anticipatory state of visual attention mechanisms , 1998 .

[147]  David J. Nutt,et al.  The effects of elevated endogenous GABA levels on movement-related network oscillations , 2013, NeuroImage.

[148]  H. Kennedy,et al.  Alpha-Beta and Gamma Rhythms Subserve Feedback and Feedforward Influences among Human Visual Cortical Areas , 2016, Neuron.

[149]  M. Corsi-Cabrera,et al.  Diazepam-Induced Changes in EEG Oscillations During Performance of a Sustained Attention Task , 2011, Journal of clinical neurophysiology : official publication of the American Electroencephalographic Society.

[150]  M. Fink,et al.  Blood levels and electroencephalographic effects of diazepam and bromazepam , 1976, Clinical pharmacology and therapeutics.

[151]  M. Iyo,et al.  Effects of a small dose of triazolam on P300 and resting EEG , 1996, Psychopharmacology.

[152]  Karl J. Friston,et al.  Canonical Microcircuits for Predictive Coding , 2012, Neuron.

[153]  S. Golombok,et al.  The psychopharmacological effects of premazepam, diazepam and placebo in healthy human subjects. , 1984, British journal of clinical pharmacology.

[154]  J. Jefferys,et al.  High‐frequency gamma oscillations coexist with low‐frequency gamma oscillations in the rat visual cortex in vitro , 2010, The European journal of neuroscience.

[155]  M. Jarvis,et al.  Effect of subcutaneous nicotine injections on EEG alpha frequency in non-smokers: a placebo-controlled pilot study , 1994, Psychopharmacology.

[156]  I. Fried,et al.  Coupling between Neuronal Firing Rate, Gamma LFP, and BOLD fMRI Is Related to Interneuronal Correlations , 2007, Current Biology.

[157]  W. Kirch,et al.  Effects of physostigmine on scopolamine–induced changes in quantitative electroencephalogram and cognitive performance , 1998 .

[158]  Michael Okun,et al.  Instantaneous correlation of excitation and inhibition during ongoing and sensory-evoked activities , 2008, Nature Neuroscience.

[159]  V. Knott,et al.  The separate and combined effects of monoamine oxidase A inhibition and nicotine on resting state EEG , 2016, Journal of psychopharmacology.

[160]  Steven Lemm,et al.  A novel mechanism for evoked responses in the human brain , 2007, The European journal of neuroscience.

[161]  Ned T. Sahin,et al.  Dynamic circuit motifs underlying rhythmic gain control, gating and integration , 2014, Nature Neuroscience.

[162]  Seppo P. Ahlfors,et al.  Parieto‐occipital ∼1 0Hz activity reflects anticipatory state of visual attention mechanisms , 1998 .

[163]  Rikkert Hindriks,et al.  Thalamo-cortical mechanisms underlying changes in amplitude and frequency of human alpha oscillations , 2013, NeuroImage.

[164]  V. Knott,et al.  Neural expression of nicotine's antidepressant properties during tryptophan depletion: An EEG study in healthy volunteers at risk for depression , 2012, Biological Psychology.

[165]  Hans-Peter Landolt,et al.  The Functional Val158Met Polymorphism of COMT Predicts Interindividual Differences in Brain α Oscillations in Young Men , 2009, The Journal of Neuroscience.

[166]  V. Knott,et al.  Nicotine and smoker status moderate brain electric and mood activation induced by ketamine, an N-methyl-d-aspartate (NMDA) receptor antagonist , 2006, Pharmacology Biochemistry and Behavior.

[167]  Karl J. Friston,et al.  Broadband Cortical Desynchronization Underlies the Human Psychedelic State , 2013, The Journal of Neuroscience.

[168]  Vince D. Calhoun,et al.  EEGIFT: Group Independent Component Analysis for Event-Related EEG Data , 2011, Comput. Intell. Neurosci..

[169]  T. Sejnowski,et al.  Cortical Enlightenment: Are Attentional Gamma Oscillations Driven by ING or PING? , 2009, Neuron.

[170]  S Taulu,et al.  Artifact and head movement compensation in MEG. , 2007, Neurology, neurophysiology, and neuroscience.

[171]  Karl J. Friston,et al.  a K.E. Stephan, a R.B. Reilly, , 2007 .

[172]  S. Muthukumaraswamy The use of magnetoencephalography in the study of psychopharmacology (pharmaco-MEG) , 2014, Journal of psychopharmacology.

[173]  K. Mathewson,et al.  Pulsed Out of Awareness: EEG Alpha Oscillations Represent a Pulsed-Inhibition of Ongoing Cortical Processing , 2011, Front. Psychology.

[174]  A. Compston The Berger rhythm: potential changes from the occipital lobes in man. , 2010, Brain : a journal of neurology.

[175]  V. Feshchenko,et al.  Comparison of the EEG effects of midazolam, thiopental, and propofol: the role of underlying oscillatory systems. , 1997, Neuropsychobiology.

[176]  G. Gücer,et al.  Thalamic EEG recordings in patients with chronic pain , 1978, Journal of Neurology.

[177]  D. Nutt,et al.  Differences between magnetoencephalographic (MEG) spectral profiles of drugs acting on GABA at synaptic and extrasynaptic sites: A study in healthy volunteers , 2015, Neuropharmacology.

[178]  Linda Drijvers,et al.  Hearing and seeing meaning in noise: Alpha, beta, and gamma oscillations predict gestural enhancement of degraded speech comprehension , 2018, Human brain mapping.

[179]  B. Connors,et al.  Cellular Mechanisms of the Augmenting Response: Short-Term Plasticity in a Thalamocortical Pathway , 1996, The Journal of Neuroscience.

[180]  Y. Saalmann,et al.  The Pulvinar Regulates Information Transmission Between Cortical Areas Based on Attention Demands , 2012, Science.

[181]  J. Lieberman,et al.  Serotonergic basis of antipsychotic drug effects in schizophrenia , 1998, Biological Psychiatry.

[182]  C. Moore,et al.  Neural mechanisms of transient neocortical beta rhythms: Converging evidence from humans, computational modeling, monkeys, and mice , 2016, Proceedings of the National Academy of Sciences.

[183]  M. Puig,et al.  Serotonin Modulation of Prefronto-Hippocampal Rhythms in Health and Disease. , 2015, ACS chemical neuroscience.

[184]  Khalid Hamandi,et al.  Elevating Endogenous GABA Levels with GAT-1 Blockade Modulates Evoked but Not Induced Responses in Human Visual Cortex , 2013, Neuropsychopharmacology.

[185]  Vadim V. Nikulin,et al.  Effects of alcohol on spontaneous neuronal oscillations: A combined magnetoencephalography and electroencephalography study , 2005, Progress in Neuro-Psychopharmacology and Biological Psychiatry.

[186]  L. Hazrati,et al.  In vitro recordings of human neocortical oscillations. , 2015, Cerebral cortex.

[187]  Emery N Brown,et al.  Neural oscillations demonstrate that general anesthesia and sedative states are neurophysiologically distinct from sleep , 2017, Current Opinion in Neurobiology.

[188]  Joachim Gross,et al.  Good practice for conducting and reporting MEG research , 2013, NeuroImage.

[189]  R. Desimone,et al.  Modulation of Oscillatory Neuronal Synchronization by Selective Visual Attention , 2001, Science.

[190]  R. Desimone,et al.  Laminar differences in gamma and alpha coherence in the ventral stream , 2011, Proceedings of the National Academy of Sciences.

[191]  F. H. Lopes da Silva Neural mechanisms underlying brain waves: from neural membranes to networks. , 1991, Electroencephalography and clinical neurophysiology.

[192]  Nancy Kopell,et al.  Alpha-Frequency Rhythms Desynchronize over Long Cortical Distances: A Modeling Study , 2000, Journal of Computational Neuroscience.

[193]  M. Greenlee,et al.  Nicotine facilitates memory consolidation in perceptual learning , 2013, Neuropharmacology.

[194]  N. Kopell,et al.  Thalamic model of awake alpha oscillations and implications for stimulus processing , 2012, Proceedings of the National Academy of Sciences.

[195]  G. Price,et al.  The effects of dexamphetamine on the resting‐state electroencephalogram and functional connectivity , 2016, Human brain mapping.

[196]  Nico Bunzeck,et al.  Acetylcholine modulates human working memory and subsequent familiarity based recognition via alpha oscillations , 2016, NeuroImage.

[197]  R. Pigeau,et al.  Distinctive effects of modafinil and d-amphetamine on the homeostatic and circadian modulation of the human waking EEG , 2003, Psychopharmacology.

[198]  O. Jensen,et al.  Rhythmic Pulsing: Linking Ongoing Brain Activity with Evoked Responses , 2010, Front. Hum. Neurosci..

[199]  V. Knott,et al.  Naltrexone alteration of the nicotine-induced EEG and mood activation response in tobacco-deprived cigarette smokers. , 2007, Experimental and clinical psychopharmacology.

[200]  Erika E. Fanselow,et al.  Behavioral Modulation of Tactile Responses in the Rat Somatosensory System , 1999, The Journal of Neuroscience.

[201]  C. Schroeder,et al.  Neuronal Mechanisms and Attentional Modulation of Corticothalamic Alpha Oscillations , 2011, The Journal of Neuroscience.

[202]  S. Hughes,et al.  Temporal Framing of Thalamic Relay-Mode Firing by Phasic Inhibition during the Alpha Rhythm , 2009, Neuron.

[203]  R. O’Connell,et al.  Donepezil Impairs Memory in Healthy Older Subjects: Behavioural, EEG and Simultaneous EEG/fMRI Biomarkers , 2011, Alzheimer's & Dementia.

[204]  Lucas C. Parra,et al.  Subject position affects EEG magnitudes , 2013, NeuroImage.

[205]  M Danhof,et al.  Pharmacokinetic‐pharmacodynamic modeling of midazolam effects on the human central nervous system , 1988, Clinical pharmacology and therapeutics.

[206]  Robert Oostenveld,et al.  FieldTrip: Open Source Software for Advanced Analysis of MEG, EEG, and Invasive Electrophysiological Data , 2010, Comput. Intell. Neurosci..

[207]  F. L. D. Silva,et al.  Event-related EEG/MEG synchronization and desynchronization: basic principles , 1999, Clinical Neurophysiology.

[208]  P. Anderer,et al.  Comparative pharmacodynamic studies with the novel serotonin uptake-enhancing tianeptine and — inhibiting fluvoxamine utilizing EEG mapping and psychometry , 2005, Journal of Neural Transmission.

[209]  S. Hughes,et al.  Synchronized Oscillations at α and θ Frequencies in the Lateral Geniculate Nucleus , 2004, Neuron.

[210]  Matti S. Hämäläinen,et al.  MRI-constrained spectral imaging of benzodiazepine modulation of spontaneous neuromagnetic activity in human cortex , 2007, NeuroImage.

[211]  M. Fink EEG and human psychopharmacology. , 1969, Annual review of pharmacology.

[212]  N. Jaworska,et al.  Effects of nicotine on electroencephalographic (EEG) and behavioural measures of visual working memory in non-smokers during a dual-task paradigm , 2013, Pharmacology Biochemistry and Behavior.

[213]  Jyrki Ahveninen,et al.  Effects of scopolamine on MEG spectral power and coherence in elderly subjects , 2003, Clinical Neurophysiology.

[214]  P. Matthews,et al.  The Effects of Nicotine Replacement on Cognitive Brain Activity During Smoking Withdrawal Studied with Simultaneous fMRI/EEG , 2011, Neuropsychopharmacology.

[215]  P. Anderer,et al.  Topographic pharmaco-EEG mapping of the effects of the South American psychoactive beverage ayahuasca in healthy volunteers. , 2002, British journal of clinical pharmacology.

[216]  R. Traub,et al.  Neuronal Networks of the Hippocampus , 1991 .

[217]  Emery N. Brown,et al.  Electroencephalogram signatures of ketamine anesthesia-induced unconsciousness , 2016, Clinical Neurophysiology.

[218]  F H da Silva,et al.  Essential differences between alpha rhythms and barbiturate spindles: spectra and thalamo-cortical coherences. , 1973, Electroencephalography and clinical neurophysiology.

[219]  R. VanRullen Perceptual Cycles , 2016, Trends in Cognitive Sciences.

[220]  D. Albe-Fessard Electrophysiological methods for the identification of thalamic nuclei , 1973, Zeitschrift für Neurologie.