Professor Miles Whittington


Chair in Neuroscience

Role at Hull York Medical School
  • Professor of Neuroscience
  • Preclinical Pharmacology Lead
  • Intercalation Lead

1987-1990 PhD University of Bristol (Pharmacology)

1990-1991 Postdoctoral research assistant (MRC), Bristol

1991- 1995 Postdoc. research assistant (Wellcome), St. Mary’s hospital Medical School, London

1995 - 1998 Governor’s Lecturer, Imperial College School of Medicine.

1998 - 1999 Lecturer, Imperial College School of Medicine

1999 - Senior Lecturer, Univ. Leeds

2001 - Reader in Neuroscience, Univ. Leeds

2003 – Professor of Neuronal Dynamics, Leeds

2005 – Professor of Neuroscience, Newcastle

2012 – Professor of Neuroscience, Hull York Medical School, York


Miles is a member of the Experimental Medicine and Biomedicine research group.

Whittington's research focusses on establishing a mechanistic understanding of the origins and properties of brain rhythms. The driving aim is to be able to understand enough at the molecular, synaptic, cellular and network level to be able to predict primary pathologies from deficits in non-invasive (EEG, MEG) recordings of brain activity in patients with psychiatric illness. His research is multidisciplinary, involving in vitro brain slice models and biologically realistic computational models to provide a link between EEG/ECoG-visible activity patterns and immunocytochemical/electron microscopic changes in neuronal structure along with molecular genetic changes in neuronal function.


Basic Pharmacology Teaching to yr 1 & 2 Medical Students

Introduction to Pharmacology module yr 2 Biomedical Sciences Students

Brain rhythms yr 2 Medical Students


I Books

Traub, R. D. & Whittington, M. A. Oscillations in Health & Disease. pp 431, New York NY, OUP. 2010

Traub, R. D., Jefferys, J. G. R. & Whittington, M. A. Oscillations in Cortical Circuits. pp 324, Cambridge MA, MIT Press. 1999


II Parts of Books

Traub RD, Whittington MA, Cunningham MO. Fast Oscillations and Synchronization Examined with In Vitro Models of Epileptogenesis. In: Noebels JL, Avoli M, Rogawski MA, Olsen RW, Delgado-Escueta AV, editors. Jasper's Basic Mechanisms of the Epilepsies [Internet]. 4th edition. Bethesda (MD): National Center for Biotechnology Information (US); 2012.

Whittington , M.A. & Buhl, E.H. Cortical microcircuits. Book chapter in ‘The hippocampus’. Eds P. Andersen, J. O’Keefe, R. Morris. 2006.

Traub, R. D., Contreras, D. & Whittington, M. A. (2007). Cellular and network mechanisms of oscillations preceding and perhaps initiating epileptic discharges. Eds. I Soltesz, K Staley.  In ‘ Computational Neuroscience in Epilepsy’. Elsevier.

Traub, R. D., Jefferys, J. G. R. & Whittington, M. A. Functionally relevant and functionally disruptive (epileptic) synchronised oscillations in brain slices, eds A. V. Delgado-Escueta, W. A. Wilson, R. W. Olsen & R. J. Porter. In ‘Jasper’s Basic Mechanisms of the Epilepsies third Edition, 709-724, Lippincott Williams & Wilkins, Philadelphia, 1999

Whittington, M. A., Traub, R. D., & Jefferys, J. G. R. . Experimental gamma (40 Hz) oscillations: Mechanisms of generation, function and effects of general anaesthetics, ed. G. Litsher. In ‘Monitoring of Stimulus-Induced 40 Hz Brain Oscillations’, Pabst Science Publishers GmBH, 1998

Traub, R. D., Wong, R. K. S., Jefferys, J. G. R., Whittington, M. A. & Michaelson, H. Generation of temporal correlations in firing of pyramidal cells by networks of inhibitory neurons. eds. BAUDRY, M. & DAVIS, J. In ‘Long term potentiation’, pp. 183-198. Cambridge, MA: MIT Press. 1996


III Full papers in refereed journals

Enhanced interlaminar excitation or reduced superficial layer inhibition in neocortex generates different spike and wave-like electrographic events in vitro. Hall SP, Traub RD, Adams NE, Cunningham MO, Schofield I, Jenkins AJ, Whittington MA. J Neurophysiol. 2017 Sep 27:jn.00516.2017. doi: 10.1152/jn.00516.2017.

Aberrant Network Activity in Schizophrenia. Hunt MJ, Kopell NJ, Traub RD, Whittington MA. Trends Neurosci. 2017 Jun;40(6):371-382. doi: 10.1016/j.tins.2017.04.003. Epub 2017 May 14.

Hetereogeneity in Neuronal Intrinsic Properties: A Possible Mechanism for Hub-Like Properties of the Rat Anterior Cingulate Cortex during Network Activity. Adams NE, Sherfey JS, Kopell NJ, Whittington MA, LeBeau FE. eNeuro. 2017 Mar 7;4(1). pii: ENEURO.0313-16.2017. doi: 10.1523/ENEURO.0313-16.2017. eCollection 2017 Jan-Feb.

The Cognitive Role of the Globus Pallidus interna; Insights from Disease States. Gillies MJ, Hyam JA, Weiss AR, Antoniades CA, Bogacz R, Fitzgerald JJ, Aziz TZ, Whittington MA, Green AL. Exp Brain Res. 2017 May;235(5):1455-1465. doi: 10.1007/s00221-017-4905-8. Epub 2017 Feb 28.

Dorsal vs. ventral differences in fast Up-state-associated oscillations in the medial prefrontal cortex of the urethane-anesthetized rat. Gretenkord S, Rees A, Whittington MA, Gartside SE, LeBeau FE. J Neurophysiol. 2017 Mar 1;117(3):1126-1142. doi: 10.1152/jn.00762.2016. Epub 2016 Dec 21.

GABAB receptor-mediated, layer-specific synaptic plasticity reorganizes gamma-frequency neocortical response to stimulation. Ainsworth M, Lee S, Kaiser M, Simonotto J, Kopell NJ, Whittington MA. Proc Natl Acad Sci U S A. 2016 May 10;113(19):E2721-9. doi: 10.1073/pnas.1605243113. Epub 2016 Apr 26.

Potential Mechanisms Underlying Intercortical Signal Regulation via Cholinergic Neuromodulators. Lee JH, Whittington MA, Kopell NJ. J Neurosci. 2015 Nov 11;35(45):15000-14. doi: 10.1523/JNEUROSCI.0629-15.2015.

Effects of NMDA receptor antagonists and antipsychotics on high frequency oscillations recorded in the nucleus accumbens of freely moving mice. Hunt MJ, Olszewski M, Piasecka J, Whittington MA, Kasicki S. Psychopharmacology (Berl). 2015 Dec;232(24):4525-35. doi: 10.1007/s00213-015-4073-0. Epub 2015 Oct 8.

Unbalanced Peptidergic Inhibition in Superficial Neocortex Underlies Spike and Wave Seizure Activity. Hall S, Hunt M, Simon A, Cunnington LG, Carracedo LM, Schofield IS, Forsyth R, Traub RD, Whittington MA. J Neurosci. 2015 Jun 24;35(25):9302-14. doi: 10.1523/JNEUROSCI.4245-14.2015.

Subregional differences in the generation of fast network oscillations in the rat medial prefrontal cortex (mPFC) in vitro. Glykos V, Whittington MA, LeBeau FE. J Physiol. 2015 Aug 15;593(16):3597-615. doi: 10.1113/JP270811. Epub 2015 Jun 30.

Near-field electromagnetic holography for high-resolution analysis of network interactions in neuronal tissue. Kjeldsen HD, Kaiser M, Whittington MA. J Neurosci Methods. 2015 Sep 30;253:1-9. doi: 10.1016/j.jneumeth.2015.05.016. Epub 2015 May 28.

Aberrant high frequency oscillations recorded in the rat nucleus accumbens in the methylazoxymethanol acetate neurodevelopmental model of schizophrenia. Goda SA, Olszewski M, Piasecka J, Rejniak K, Whittington MA, Kasicki S, Hunt MJ. Prog Neuropsychopharmacol Biol Psychiatry. 2015 Aug 3;61:44-51. doi: 10.1016/j.pnpbp.2015.03.016. Epub 2015 Apr 7.

Brain rhythms connect impaired inhibition to altered cognition in schizophrenia. Pittman-Polletta BR, Kocsis B, Vijayan S, Whittington MA, Kopell NJ. Biol Psychiatry. 2015 Jun 15;77(12):1020-30. doi: 10.1016/j.biopsych.2015.02.005. Epub 2015 Feb 14.

Electrographic waveform structure predicts laminar focus location in a model of temporal lobe seizures in vitro. Adams C, Adams NE, Traub RD, Whittington MA. PLoS One. 2015 Mar 23;10(3):e0121676. doi: 10.1371/journal.pone.0121676. eCollection 2015.

Beyond the connectome: the dynome. Kopell NJ, Gritton HJ, Whittington MA, Kramer MA. Neuron. 2014 Sep 17;83(6):1319-28. doi: 10.1016/j.neuron.2014.08.016.

What is a seizure network? Very fast oscillations at the interface between normal and epileptic brain. Traub RD, Cunningham MO, Whittington MA. Adv Exp Med Biol. 2014;813:71-80. doi: 10.1007/978-94-017-8914-1_6.

Virtual Electrode Recording Tool for Extracellular potentials (VERTEX): comparing multi-electrode recordings from simulated and biological mammalian cortical tissue. Tomsett RJ, Ainsworth M, Thiele A, Sanayei M, Chen X, Gieselmann MA, Whittington MA, Cunningham MO, Kaiser M. Brain Struct Funct. 2015 Jul;220(4):2333-53. doi: 10.1007/s00429-014-0793-x. Epub 2014 May 27.

A unified approach to linking experimental, statistical and computational analysis of spike train data. Meng L, Kramer MA, Middleton SJ, Whittington MA, Eden UT. PLoS One. 2014 Jan 17;9(1):e85269. doi: 10.1371/journal.pone.0085269. eCollection 2014.

Neurosystems: brain rhythms and cognitive processing. Cannon J, McCarthy MM, Lee S, Lee J, Börgers C, Whittington MA, Kopell N. Eur J Neurosci. 2014 Mar;39(5):705-19. doi: 10.1111/ejn.12453. Epub 2013 Dec 13.

Gap junction networks can generate both ripple-like and fast ripple-like oscillations. Simon A, Traub RD, Vladimirov N, Jenkins A, Nicholson C, Whittaker RG, Schofield I, Clowry GJ, Cunningham MO, Whittington MA. Eur J Neurosci. 2014 Jan;39(1):46-60. doi: 10.1111/ejn.12386. Epub 2013 Oct 14.

Top-down beta rhythms support selective attention via interlaminar interaction: a model. Lee JH, Whittington MA, Kopell NJ. PLoS Comput Biol. 2013;9(8):e1003164. doi: 10.1371/journal.pcbi.1003164. Epub 2013 Aug 8.

A neocortical delta rhythm facilitates reciprocal interlaminar interactions via nested theta rhythms. Carracedo LM, Kjeldsen H, Cunnington L, Jenkins A, Schofield I, Cunningham MO, Davies CH, Traub RD, Whittington MA. J Neurosci. 2013 Jun 26;33(26):10750-61. doi: 10.1523/JNEUROSCI.0735-13.2013.

Rates and rhythms: a synergistic view of frequency and temporal coding in neuronal networks.

Ainsworth M, Lee S, Cunningham MO, Traub RD, Kopell NJ, Whittington MA. Neuron. 2012 Aug 23;75(4):572-83. doi: 10.1016/j.neuron.2012.08.004.

Fast Oscillations and Synchronization Examined with In Vitro Models of Epileptogenesis. Traub RD, Whittington MA, Cunningham MO. In: Noebels JL, Avoli M, Rogawski MA, Olsen RW, Delgado-Escueta AV, editors. Jasper's Basic Mechanisms of the Epilepsies [Internet]. 4th edition. Bethesda (MD): National Center for Biotechnology Information (US); 2012.

Segregation of axonal and somatic activity during fast network oscillations. Dugladze T, Schmitz D, Whittington MA, Vida I, Gloveli T. Science. 2012 Jun 15;336(6087):1458-61. doi: 10.1126/science.1222017.

Axonal properties determine somatic firing in a model of in vitro CA1 hippocampal sharp wave/ripples and persistent gamma oscillations. Traub RD, Schmitz D, Maier N, Whittington MA, Draguhn A. Eur J Neurosci. 2012 Sep;36(5):2650-60. doi: 10.1111/j.1460-9568.2012.08184.x. Epub 2012 Jun 15.

Glissandi: transient fast electrocorticographic oscillations of steadily increasing frequency, explained by temporally increasing gap junction conductance. Cunningham MO, Roopun A, Schofield IS, Whittaker RG, Duncan R, Russell A, Jenkins A, Nicholson C, Whittington MA, Traub RD. Epilepsia. 2012 Jul;53(7):1205-14. doi: 10.1111/j.1528-1167.2012.03530.x. Epub 2012 Jun 12.

Dynamical changes in neurological diseases and anesthesia. McCarthy MM, Ching S, Whittington MA, Kopell N. Curr Opin Neurobiol. 2012 Aug;22(4):693-703. doi: 10.1016/j.conb.2012.02.009. Epub 2012 Mar 23.

Long-range-projecting GABAergic neurons modulate inhibition in hippocampus and entorhinal cortex. Melzer S, Michael M, Caputi A, Eliava M, Fuchs EC, Whittington MA, Monyer H. Science. 2012 Mar 23;335(6075):1506-10. doi: 10.1126/science.1217139.

Study protocol: a randomized controlled trial of a computer-based depression and substance abuse intervention for people attending residential substance abuse treatment. Kelly PJ, Kay-Lambkin FJ, Baker AL, Deane FP, Brooks AC, Mitchell A, Marshall S, Whittington M, Dingle GA. BMC Public Health. 2012 Feb 10;12:113. doi: 10.1186/1471-2458-12-113.

Cognitive dysfunction in psychiatric disorders: characteristics, causes and the quest for improved therapy. Millan MJ, Agid Y, Brüne M, Bullmore ET, Carter CS, Clayton NS, Connor R, Davis S, Deakin B, DeRubeis RJ, Dubois B, Geyer MA, Goodwin GM, Gorwood P, Jay TM, Joëls M, Mansuy IM, Meyer-Lindenberg A, Murphy D, Rolls E, Saletu B, Spedding M, Sweeney J, Whittington M, Young LJ. Nat Rev Drug Discov. 2012 Feb 1;11(2):141-68. doi: 10.1038/nrd3628.

Dual γ rhythm generators control interlaminar synchrony in auditory cortex. Ainsworth M, Lee S, Cunningham MO, Roopun AK, Traub RD, Kopell NJ, Whittington MA. J Neurosci. 2011 Nov 23;31(47):17040-51. doi: 10.1523/JNEUROSCI.2209-11.2011. Erratum in: J Neurosci. 2012 Feb 22;32(8):2911.

Natural splice variant of MHC class I cytoplasmic tail enhances dendritic cell-induced CD8+ T-cell responses and boosts anti-tumor immunity. Rodríguez-Cruz TG, Liu S, Khalili JS, Whittington M, Zhang M, Overwijk W, Lizée G. PLoS One. 2011;6(8):e22939. doi: 10.1371/journal.pone.0022939. Epub 2011 Aug 10.

Circuits and brain rhythms in schizophrenia: a wealth of convergent targets. Whittington MA, Roopun AK, Traub RD, Davies CH. Curr Opin Pharmacol. 2011 Oct;11(5):508-14. doi: 10.1016/j.coph.2011.04.010.

Impaired mitochondrial function abolishes gamma oscillations in the hippocampus through an effect on fast-spiking interneurons. Whittaker RG, Turnbull DM, Whittington MA, Cunningham MO. Brain. 2011 Jul;134(Pt 7):e180; author reply e181. doi: 10.1093/brain/awr018. Epub 2011 Mar 4.

Neuronal assembly dynamics in the beta1 frequency range permits short-term memory. Kopell N, Whittington MA, Kramer MA. Proc Natl Acad Sci U S A. 2011 Mar 1;108(9):3779-84. doi: 10.1073/pnas.1019676108. Epub 2011 Feb 14.

Fast oscillatory activity induced by kainate receptor activation in the rat basolateral amygdala in vitro. Randall FE, Whittington MA, Cunningham MO. Eur J Neurosci. 2011 Mar;33(5):914-22. doi: 10.1111/j.1460-9568.2010.07582.x. Epub 2011 Jan 24.

Chemical synaptic and gap junctional interactions between principal neurons: partners in epileptogenesis. Traub RD, Cunningham MO, Whittington MA. Neural Netw. 2011 Aug;24(6):515-25. doi: 10.1016/j.neunet.2010.11.007. Epub 2010 Dec 1.

Multiple origins of the cortical γ rhythm. Whittington MA, Cunningham MO, LeBeau FE, Racca C, Traub RD. Dev Neurobiol. 2011 Jan 1;71(1):92-106. doi: 10.1002/dneu.20814.

Are different rhythms good for different functions? Kopell N, Kramer MA, Malerba P, Whittington MA. Front Hum Neurosci. 2010 Nov 2;4:187. doi: 10.3389/fnhum.2010.00187. eCollection 2010.

Cholinergic neuromodulation controls directed temporal communication in neocortex in vitro. Roopun AK, Lebeau FE, Rammell J, Cunningham MO, Traub RD, Whittington MA. Front Neural Circuits. 2010 Mar 22;4:8. doi: 10.3389/fncir.2010.00008. eCollection 2010.

Mitochondrial DNA mutations affect calcium handling in differentiated neurons. Trevelyan AJ, Kirby DM, Smulders-Srinivasan TK, Nooteboom M, Acin-Perez R, Enriquez JA, Whittington MA, Lightowlers RN, Turnbull DM. Brain. 2010 Mar;133(Pt 3):787-96. doi: 10.1093/brain/awq023. Epub 2010 Mar 5.

Spatiotemporal patterns of electrocorticographic very fast oscillations (> 80 Hz) consistent with a network model based on electrical coupling between principal neurons. Traub RD, Duncan R, Russell AJ, Baldeweg T, Tu Y, Cunningham MO, Whittington MA. Epilepsia. 2010 Aug;51(8):1587-97. doi: 10.1111/j.1528-1167.2009.02420.x. Epub 2009 Dec 7.

A nonsynaptic mechanism underlying interictal discharges in human epileptic neocortex. Roopun AK, Simonotto JD, Pierce ML, Jenkins A, Nicholson C, Schofield IS, Whittaker RG, Kaiser M, Whittington MA, Traub RD, Cunningham MO. Proc Natl Acad Sci U S A. 2010 Jan 5;107(1):338-43. doi: 10.1073/pnas.0912652107. Epub 2009 Dec 4.

Transmitochondrial embryonic stem cells containing pathogenic mtDNA mutations are compromised in neuronal differentiation. Kirby DM, Rennie KJ, Smulders-Srinivasan TK, Acin-Perez R, Whittington M, Enriquez JA, Trevelyan AJ, Turnbull DM, Lightowlers RN. Cell Prolif. 2009 Aug;42(4):413-24. doi: 10.1111/j.1365-2184.2009.00612.x. Epub 2009 Jun 23.

Temporal Interactions between Cortical Rhythms. Roopun AK, Kramer MA, Carracedo LM, Kaiser M, Davies CH, Traub RD, Kopell NJ, Whittington MA. Front Neurosci. 2008 Dec 15;2(2):145-54. doi: 10.3389/neuro.01.034.2008. eCollection 2008 Dec.

NMDA receptor-dependent switching between different gamma rhythm-generating microcircuits in entorhinal cortex. Middleton S, Jalics J, Kispersky T, Lebeau FE, Roopun AK, Kopell NJ, Whittington MA, Cunningham MO. Proc Natl Acad Sci U S A. 2008 Nov 25;105(47):18572-7. doi: 10.1073/pnas.0809302105. Epub 2008 Nov 7.

Model of very fast (> 75 Hz) network oscillations generated by electrical coupling between the proximal axons of cerebellar Purkinje cells. Traub RD, Middleton SJ, Knöpfel T, Whittington MA. Eur J Neurosci. 2008 Oct;28(8):1603-16. doi: 10.1111/j.1460-9568.2008.06477.x.

Period concatenation underlies interactions between gamma and beta rhythms in neocortex. Roopun AK, Kramer MA, Carracedo LM, Kaiser M, Davies CH, Traub RD, Kopell NJ, Whittington MA. Front Cell Neurosci. 2008 Apr 8;2:1. doi: 10.3389/neuro.03.001.2008. eCollection 2008.

Detecting seizure origin using basic, multiscale population dynamic measures: preliminary findings. Roopun AK, Traub RD, Baldeweg T, Cunningham MO, Whittaker RG, Trevelyan A, Duncan R, Russell AJ, Whittington MA. Epilepsy Behav. 2009 Jan;14 Suppl 1:39-46. doi: 10.1016/j.yebeh.2008.09.008. Epub 2008 Oct 31.

Rhythm generation through period concatenation in rat somatosensory cortex. Kramer MA, Roopun AK, Carracedo LM, Traub RD, Whittington MA, Kopell NJ. PLoS Comput Biol. 2008 Sep 5;4(9):e1000169. doi: 10.1371/journal.pcbi.1000169.

High-frequency network oscillations in cerebellar cortex. Middleton SJ, Racca C, Cunningham MO, Traub RD, Monyer H, Knöpfel T, Schofield IS, Jenkins A, Whittington MA. Neuron. 2008 Jun 12;58(5):763-74. doi: 10.1016/j.neuron.2008.03.030.

Region-specific changes in gamma and beta2 rhythms in NMDA receptor dysfunction models of schizophrenia. Roopun AK, Cunningham MO, Racca C, Alter K, Traub RD, Whittington MA. Schizophr Bull. 2008 Sep;34(5):962-73. doi: 10.1093/schbul/sbn059. Epub 2008 Jun 9.

Characterization of a CNS penetrant, selective M1 muscarinic receptor agonist, 77-LH-28-1. Langmead CJ, Austin NE, Branch CL, Brown JT, Buchanan KA, Davies CH, Forbes IT, Fry VA, Hagan JJ, Herdon HJ, Jones GA, Jeggo R, Kew JN, Mazzali A, Melarange R, Patel N, Pardoe J, Randall AD, Roberts C, Roopun A, Starr KR, Teriakidis A, Wood MD, Whittington M, Wu Z, Watson J. Br J Pharmacol. 2008 Jul;154(5):1104-15. doi: 10.1038/bjp.2008.152. Epub 2008 May 5.

Can brain rhythms inform on underlying pathology in schizophrenia? Whittington MA. Biol Psychiatry. 2008 Apr 15;63(8):728-9. doi: 10.1016/j.biopsych.2008.02.007.

The source of afterdischarge activity in neocortical tonic-clonic epilepsy. Trevelyan AJ, Baldeweg T, van Drongelen W, Yuste R, Whittington M. J Neurosci. 2007 Dec 5;27(49):13513-9.

Impairment of hippocampal gamma-frequency oscillations in vitro in mice overexpressing human amyloid precursor protein (APP). Driver JE, Racca C, Cunningham MO, Towers SK, Davies CH, Whittington MA, LeBeau FE. Eur J Neurosci. 2007 Sep;26(5):1280-8.

Gap junctions on hippocampal mossy fiber axons demonstrated by thin-section electron microscopy and freeze fracture replica immunogold labeling. Hamzei-Sichani F, Kamasawa N, Janssen WG, Yasumura T, Davidson KG, Hof PR, Wearne SL, Stewart MG, Young SR, Whittington MA, Rash JE, Traub RD. Proc Natl Acad Sci U S A. 2007 Jul 24;104(30):12548-53. Epub 2007 Jul 18.

Recruitment of parvalbumin-positive interneurons determines hippocampal function and associated behavior. Fuchs EC, Zivkovic AR, Cunningham MO, Middleton S, Lebeau FE, Bannerman DM, Rozov A, Whittington MA, Traub RD, Rawlins JN, Monyer H. Neuron. 2007 Feb 15;53(4):591-604.

Beta rhythms (15-20 Hz) generated by nonreciprocal communication in hippocampus. Bibbig A, Middleton S, Racca C, Gillies MJ, Garner H, Lebeau FE, Davies CH, Whittington MA. J Neurophysiol. 2007 Apr;97(4):2812-23. Epub 2007 Feb 7.

A beta2-frequency (20-30 Hz) oscillation in nonsynaptic networks of somatosensory cortex. Roopun AK, Middleton SJ, Cunningham MO, LeBeau FE, Bibbig A, Whittington MA, Traub RD. Proc Natl Acad Sci U S A. 2006 Oct 17;103(42):15646-50. Epub 2006 Oct 9.

Low-dimensional maps encoding dynamics in entorhinal cortex and hippocampus. Pervouchine DD, Netoff TI, Rotstein HG, White JA, Cunningham MO, Whittington MA, Kopell NJ. Neural Comput. 2006 Nov;18(11):2617-50. Erratum in: Neural Comput. 2007 Jan;19(1):302.

Neuronal metabolism governs cortical network response state. Cunningham MO, Pervouchine DD, Racca C, Kopell NJ, Davies CH, Jones RS, Traub RD, Whittington MA. Proc Natl Acad Sci U S A. 2006 Apr 4;103(14):5597-601. Epub 2006 Mar 24.

Region-specific reduction in entorhinal gamma oscillations and parvalbumin-immunoreactive neurons in animal models of psychiatric illness. Cunningham MO, Hunt J, Middleton S, LeBeau FE, Gillies MJ, Davies CH, Maycox PR, Whittington MA, Racca C. J Neurosci. 2006 Mar 8;26(10):2767-76. Erratum in: J Neurosci. 2006 May 17;26(20)

Combined experimental/simulation studies of cellular and network mechanisms of epileptogenesis in vitro and in vivo. Traub RD, Contreras D, Whittington MA. J Clin Neurophysiol. 2005 Oct;22(5):330-42.

Orthogonal arrangement of rhythm-generating microcircuits in the hippocampus. Gloveli T, Dugladze T, Rotstein HG, Traub RD, Monyer H, Heinemann U, Whittington MA, Kopell NJ. Proc Natl Acad Sci U S A. 2005 Sep 13;102(37):13295-300. Epub 2005 Sep 2.

Slow and fast inhibition and an H-current interact to create a theta rhythm in a model of CA1 interneuron network. Rotstein HG, Pervouchine DD, Acker CD, Gillies MJ, White JA, Buhl EH, Whittington MA, Kopell N. J Neurophysiol. 2005 Aug;94(2):1509-18. Epub 2005 Apr 27.

Transient depression of excitatory synapses on interneurons contributes to epileptiform bursts during gamma oscillations in the mouse hippocampal slice. Traub RD, Pais I, Bibbig A, Lebeau FE, Buhl EH, Garner H, Monyer H, Whittington MA. J Neurophysiol. 2005 Aug;94(2):1225-35. Epub 2005 Feb 23.

Induction by kainate of theta frequency rhythmic activity in the rat medial septum-diagonal band complex in vitro. Garner HL, Whittington MA, Henderson Z. J Physiol. 2005 Apr 1;564(Pt 1):83-102. Epub 2005 Jan 27.

Structure/function correlates of neuronal and network activity--an overview. LeBeau FE, Whittington MA. J Physiol. 2005 Jan 1;562(Pt 1):1-2. Epub 2004 Nov 11.

Oscillatory activity within rat substantia gelatinosa in vitro: a role for chemical and electrical neurotransmission. Asghar AU, Cilia La Corte PF, LeBeau FE, Al Dawoud M, Reilly SC, Buhl EH, Whittington MA, King AE. J Physiol. 2005 Jan 1;562(Pt 1):183-98. Epub 2004 Nov 4.

Single-column thalamocortical network model exhibiting gamma oscillations, sleep spindles, and epileptogenic bursts. Traub RD, Contreras D, Cunningham MO, Murray H, LeBeau FE, Roopun A, Bibbig A, Wilent WB, Higley MJ, Whittington MA. J Neurophysiol. 2005 Apr;93(4):2194-232. Epub 2004 Nov 3.

Persistent gamma oscillations in superficial layers of rat auditory neocortex: experiment and model. Traub RD, Bibbig A, LeBeau FE, Cunningham MO, Whittington MA. J Physiol. 2005 Jan 1;562(Pt 1):3-8. Epub 2004 Oct 15.

Differential involvement of oriens/pyramidale interneurones in hippocampal network oscillations in vitro. Gloveli T, Dugladze T, Saha S, Monyer H, Heinemann U, Traub RD, Whittington MA, Buhl EH. J Physiol. 2005 Jan 1;562(Pt 1):131-47. Epub 2004 Oct 14.

Alpha 5 subunit-containing GABAA receptors affect the dynamic range of mouse hippocampal kainate-induced gamma frequency oscillations in vitro. Towers SK, Gloveli T, Traub RD, Driver JE, Engel D, Fradley R, Rosahl TW, Maubach K, Buhl EH, Whittington MA. J Physiol. 2004 Sep 15;559(Pt 3):721-8. Epub 2004 Jul 29.

Coexistence of gamma and high-frequency oscillations in rat medial entorhinal cortex in vitro. Cunningham MO, Halliday DM, Davies CH, Traub RD, Buhl EH, Whittington MA. J Physiol. 2004 Sep 1;559(Pt 2):347-53. Epub 2004 Jul 14.

Gap junctions, fast oscillations and the initiation of seizures. Traub RD, Michelson-Law H, Bibbig AE, Buhl EH, Whittington MA. Adv Exp Med Biol. 2004;548:110-22.

Cellular mechanisms of neuronal population oscillations in the hippocampus in vitro. Traub RD, Bibbig A, LeBeau FE, Buhl EH, Whittington MA. Annu Rev Neurosci. 2004;27:247-78.

A role for fast rhythmic bursting neurons in cortical gamma oscillations in vitro. Cunningham MO, Whittington MA, Bibbig A, Roopun A, LeBeau FE, Vogt A, Monyer H, Buhl EH, Traub RD. Proc Natl Acad Sci U S A. 2004 May 4;101(18):7152-7. Epub 2004 Apr 21.

Interneuron diversity series: inhibitory interneurons and network oscillations in vitro. Whittington MA, Traub RD. Trends Neurosci. 2003 Dec;26(12):676-82.

The role of electrical signaling via gap junctions in the generation of fast network oscillations. LeBeau FE, Traub RD, Monyer H, Whittington MA, Buhl EH. Brain Res Bull. 2003 Nov 15;62(1):3-13.

Gamma oscillations induced by kainate receptor activation in the entorhinal cortex in vitro. Cunningham MO, Davies CH, Buhl EH, Kopell N, Whittington MA. J Neurosci. 2003 Oct 29;23(30):9761-9.

GABA-enhanced collective behavior in neuronal axons underlies persistent gamma-frequency oscillations. Traub RD, Cunningham MO, Gloveli T, LeBeau FE, Bibbig A, Buhl EH, Whittington MA. Proc Natl Acad Sci U S A. 2003 Sep 16;100(19):11047-52. Epub 2003 Sep 5.

A novel network of multipolar bursting interneurons generates theta frequency oscillations in neocortex. Blatow M, Rozov A, Katona I, Hormuzdi SG, Meyer AH, Whittington MA, Caputi A, Monyer H. Neuron. 2003 Jun 5;38(5):805-17.

The effects of general anaesthetics on carbachol-evoked gamma oscillations in the rat hippocampus in vitro. Dickinson R, Awaiz S, Whittington MA, Lieb WR, Franks NP. Neuropharmacology. 2003 Jun;44(7):864-72.

Sharp wave-like activity in the hippocampus in vitro in mice lacking the gap junction protein connexin 36. Pais I, Hormuzdi SG, Monyer H, Traub RD, Wood IC, Buhl EH, Whittington MA, LeBeau FE. J Neurophysiol. 2003 Apr;89(4):2046-54. Erratum in: J Neurophysiol. 2003 Aug;90(2):1362.

Fast rhythmic bursting can be induced in layer 2/3 cortical neurons by enhancing persistent Na+ conductance or by blocking BK channels. Traub RD, Buhl EH, Gloveli T, Whittington MA. J Neurophysiol. 2003 Feb;89(2):909-21.

Contrasting roles of axonal (pyramidal cell) and dendritic (interneuron) electrical coupling in the generation of neuronal network oscillations. Traub RD, Pais I, Bibbig A, LeBeau FE, Buhl EH, Hormuzdi SG, Monyer H, Whittington MA. Proc Natl Acad Sci U S A. 2003 Feb 4;100(3):1370-4. Epub 2003 Jan 13.

New roles for the gamma rhythm: population tuning and preprocessing for the Beta rhythm. Olufsen MS, Whittington MA, Camperi M, Kopell N. J Comput Neurosci. 2003 Jan-Feb;14(1):33-54.

Long-range synchronization of gamma and beta oscillations and the plasticity of excitatory and inhibitory synapses: a network model. Bibbig A, Traub RD, Whittington MA. J Neurophysiol. 2002 Oct;88(4):1634-54.

A model of atropine-resistant theta oscillations in rat hippocampal area CA1. Gillies MJ, Traub RD, LeBeau FE, Davies CH, Gloveli T, Buhl EH, Whittington MA. J Physiol. 2002 Sep 15;543(Pt 3):779-93.

Fast network oscillations induced by potassium transients in the rat hippocampus in vitro. LeBeau FE, Towers SK, Traub RD, Whittington MA, Buhl EH. J Physiol. 2002 Jul 1;542(Pt 1):167-79.

Axonal gap junctions between principal neurons: a novel source of network oscillations, and perhaps epileptogenesis. Traub RD, Draguhn A, Whittington MA, Baldeweg T, Bibbig A, Buhl EH, Schmitz D. Rev Neurosci. 2002;13(1):1-30.

A comparison of the efficacy of carbamazepine and the novel anti-epileptic drug levetiracetam in the tetanus toxin model of focal complex partial epilepsy. Doheny HC, Whittington MA, Jefferys JG, Patsalos PN. Br J Pharmacol. 2002 Mar;135(6):1425-34.

Fast network oscillations in the rat dentate gyrus in vitro. Towers SK, LeBeau FE, Gloveli T, Traub RD, Whittington MA, Buhl EH. J Neurophysiol. 2002 Feb;87(2):1165-8.

Altered dentate filtering during the transition to seizure in the rat tetanus toxin model of epilepsy. Finnerty GT, Whittington MA, Jefferys JG. J Neurophysiol. 2001 Dec;86(6):2748-53.

Gap junctions between interneuron dendrites can enhance synchrony of gamma oscillations in distributed networks. Traub RD, Kopell N, Bibbig A, Buhl EH, LeBeau FE, Whittington MA. J Neurosci. 2001 Dec 1;21(23):9478-86.

Self-organized synaptic plasticity contributes to the shaping of gamma and beta oscillations in vitro. Bibbig A, Faulkner HJ, Whittington MA, Traub RD. J Neurosci. 2001 Nov 15;21(22):9053-67.

Impaired electrical signaling disrupts gamma frequency oscillations in connexin 36-deficient mice. Hormuzdi SG, Pais I, LeBeau FE, Towers SK, Rozov A, Buhl EH, Whittington MA, Monyer H. Neuron. 2001 Aug 16;31(3):487-95.

Genetically altered AMPA-type glutamate receptor kinetics in interneurons disrupt long-range synchrony of gamma oscillation. Fuchs EC, Doheny H, Faulkner H, Caputi A, Traub RD, Bibbig A, Kopell N, Whittington MA, Monyer H. Proc Natl Acad Sci U S A. 2001 Mar 13;98(6):3571-6.

A possible role for gap junctions in generation of very fast EEG oscillations preceding the onset of, and perhaps initiating, seizures. Traub RD, Whittington MA, Buhl EH, LeBeau FE, Bibbig A, Boyd S, Cross H, Baldeweg T. Epilepsia. 2001 Feb;42(2):153-70.

Differential expression of synaptic and nonsynaptic mechanisms underlying stimulus-induced gamma oscillations in vitro. Whittington MA, Doheny HC, Traub RD, LeBeau FE, Buhl EH. J Neurosci. 2001 Mar 1;21(5):1727-38.

Inhibition-based rhythms: experimental and mathematical observations on network dynamics. Whittington MA, Traub RD, Kopell N, Ermentrout B, Buhl EH. Int J Psychophysiol. 2000 Dec 1;38(3):315-36.

A model of gamma-frequency network oscillations induced in the rat CA3 region by carbachol in vitro. Traub RD, Bibbig A, Fisahn A, LeBeau FE, Whittington MA, Buhl EH. Eur J Neurosci. 2000 Nov;12(11):4093-106.

Pathway-specific habituation of induced gamma oscillations in the hippocampal slice. Doheny HC, Faulkner HJ, Gruzelier JH, Baldeweg T, Whittington MA. Neuroreport. 2000 Aug 21;11(12):2629-33.

Gamma and beta frequency oscillations in response to novel auditory stimuli: A comparison of human electroencephalogram (EEG) data with in vitro models. Haenschel C, Baldeweg T, Croft RJ, Whittington M, Gruzelier J. Proc Natl Acad Sci U S A. 2000 Jun 20;97(13):7645-50.

Neuronal fast oscillations as a target site for psychoactive drugs. Whittington MA, Faulkner HJ, Doheny HC, Traub RD. Pharmacol Ther. 2000 May;86(2):171-90.

Gamma rhythms and beta rhythms have different synchronization properties. Kopell N, Ermentrout GB, Whittington MA, Traub RD. Proc Natl Acad Sci U S A. 2000 Feb 15;97(4):1867-72.

Anaesthetic/amnesic agents disrupt beta frequency oscillations associated with potentiation of excitatory synaptic potentials in the rat hippocampal slice. Faulkner HJ, Traub RD, Whittington MA. Br J Pharmacol. 1999 Dec;128(8):1813-25.

Functionally relevant and functionally disruptive (epileptic) synchronized oscillations in brain slices. Traub RD, Jefferys JG, Whittington MA. Adv Neurol. 1999;79:709-24.

Blood and cerebrospinal fluid pharmacokinetics of the novel anticonvulsant levetiracetam (ucb L059) in the rat. Doheny HC, Ratnaraj N, Whittington MA, Jefferys JG, Patsalos PN. Epilepsy Res. 1999 Apr;34(2-3):161-8.

On the mechanism of the gamma --> beta frequency shift in neuronal oscillations induced in rat hippocampal slices by tetanic stimulation. Traub RD, Whittington MA, Buhl EH, Jefferys JG, Faulkner HJ. J Neurosci. 1999 Feb 1;19(3):1088-105.

Gamma frequency oscillations gate temporally coded afferent inputs in the rat hippocampal slice. Burchell TR, Faulkner HJ, Whittington MA. Neurosci Lett. 1998 Oct 23;255(3):151-4.

Disruption of synchronous gamma oscillations in the rat hippocampal slice: a common mechanism of anaesthetic drug action. Faulkner HJ, Traub RD, Whittington MA. Br J Pharmacol. 1998 Oct;125(3):483-92.

Gamma-frequency oscillations: a neuronal population phenomenon, regulated by synaptic and intrinsic cellular processes, and inducing synaptic plasticity. Traub RD, Spruston N, Soltesz I, Konnerth A, Whittington MA, Jefferys GR. Prog Neurobiol. 1998 Aug;55(6):563-75.

Morphine disrupts long-range synchrony of gamma oscillations in hippocampal slices. Whittington MA, Traub RD, Faulkner HJ, Jefferys JG, Chettiar K. Proc Natl Acad Sci U S A. 1998 May 12;95(10):5807-11.

Recurrent excitatory postsynaptic potentials induced by synchronized fast cortical oscillations. Whittington MA, Traub RD, Faulkner HJ, Stanford IM, Jefferys JG. Proc Natl Acad Sci U S A. 1997 Oct 28;94(22):12198-203.

Spatiotemporal patterns of gamma frequency oscillations tetanically induced in the rat hippocampal slice. Whittington MA, Stanford IM, Colling SB, Jefferys JG, Traub RD. J Physiol. 1997 Aug 1;502 (Pt 3):591-607.

Simulation of gamma rhythms in networks of interneurons and pyramidal cells. Traub RD, Jefferys JG, Whittington MA. J Comput Neurosci. 1997 Apr;4(2):141-50.

Review of the role of inhibitory neurons in chronic epileptic foci induced by intracerebral tetanus toxin. Jefferys JG, Whittington MA. Epilepsy Res. 1996 Dec;26(1):59-66.

A mechanism for generation of long-range synchronous fast oscillations in the cortex. Traub RD, Whittington MA, Stanford IM, Jefferys JG. Nature. 1996 Oct 17;383(6601):621-4.

Effects of intravenous anaesthetic agents on fast inhibitory oscillations in the rat hippocampus in vitro. Whittington MA, Jefferys JG, Traub RD. Br J Pharmacol. 1996 Aug;118(8):1977-86.

Ethanol withdrawal hyperexcitability in vivo and in isolated mouse hippocampal slices. Ripley TL, Whittington MA, Butterworth AR, Little HJ. Alcohol Alcohol. 1996 Jul;31(4):347-57.

Analysis of gamma rhythms in the rat hippocampus in vitro and in vivo. Traub RD, Whittington MA, Colling SB, Buzsáki G, Jefferys JG. J Physiol. 1996 Jun 1;493 ( Pt 2):471-84.

Neuronal networks for induced '40 Hz' rhythms. Jefferys JG, Traub RD, Whittington MA. Trends Neurosci. 1996 May;19(5):202-8.

Erosion of inhibition contributes to the progression of low magnesium bursts in rat hippocampal slices. Whittington MA, Traub RD, Jefferys JG. J Physiol. 1995 Aug 1;486 ( Pt 3):723-34.

Synchronized oscillations in interneuron networks driven by metabotropic glutamate receptor activation. Whittington MA, Traub RD, Jefferys JG. Nature. 1995 Feb 16;373(6515):612-5.

Rescue of neurophysiological phenotype seen in PrP null mice by transgene encoding human prion protein. Whittington MA, Sidle KC, Gowland I, Meads J, Hill AF, Palmer MS, Jefferys JG, Collinge J.  Nat Genet. 1995 Feb;9(2):197-201.

Increased NMDA receptor and calcium channel activity underlying ethanol withdrawal hyperexcitability. Whittington MA, Lambert JD, Little HJ. Alcohol Alcohol. 1995 Jan;30(1):105-14.

Epileptic activity outlasts disinhibition after intrahippocampal tetanus toxin in the rat. Whittington MA, Jefferys JG. J Physiol. 1994 Dec 15;481 ( Pt 3):593-604.

A branching dendritic model of a rodent CA3 pyramidal neurone. Traub RD, Jefferys JG, Miles R, Whittington MA, Tóth K. J Physiol. 1994 Nov 15;481 ( Pt 1):79-95.

Scrapie infection of transgenic mice leads to network and intrinsic dysfunction of cortical and hippocampal neurones. Jefferys JG, Empson RM, Whittington MA, Prusiner SB. Neurobiol Dis. 1994 Nov;1(1-2):25-30.

Enhanced NMDA conductance can account for epileptiform activity induced by low Mg2+ in the rat hippocampal slice. Traub RD, Jefferys JG, Whittington MA. J Physiol. 1994 Aug 1;478 Pt 3:379-93.

Prion protein is necessary for normal synaptic function. Collinge J, Whittington MA, Sidle KC, Smith CJ, Palmer MS, Clarke AR, Jefferys JG. Nature. 1994 Jul 28;370(6487):295-7.

Epileptic focus induced by intrahippocampal cholera toxin in rat: time course and properties in vivo and in vitro. Williams SF, Colling SB, Whittington MA, Jefferys JG. Epilepsy Res. 1993 Oct;16(2):137-46.

Epileptic focus induced in rat by intrahippocampal cholera toxin: neuronal properties in vitro. Watts AE, Whittington MA, Jefferys JG. Neuroscience. 1993 Jul;55(1):45-56.

Changes in voltage-operated calcium channels modify ethanol withdrawal hyperexcitability in mouse hippocampal slices. Whittington MA, Little HJ. Exp Physiol. 1993 May;78(3):347-70.

Increases in synaptic activation of calcium current as a mechanism for generation of ethanol withdrawal seizures. Whittington MA, Lambert JD, Little HJ. Alcohol Alcohol Suppl. 1993;2:391-4.

Calcium channel antagonists prevent adaptive responses to ethanol. Little HJ, Dolin SJ, Whittington MA. Alcohol Alcohol Suppl. 1993;2:263-7.

Changes in intrinsic inhibition in isolated hippocampal slices during ethanol withdrawal; lack of correlation with withdrawal hyperexcitability. Whittington MA, Little HJ, Lambert JD. Br J Pharmacol. 1992 Oct;107(2):521-7.

The effects of chronic treatment with the dihydropyridine, Bay K 8644, on hyperexcitability due to ethanol withdrawal, in vivo and in vitro. Whittington MA, Butterworth AR, Dolin SJ, Patch TL, Little HJ. Br J Pharmacol. 1992 Feb;105(2):285-92.

Evidence that changes in hippocampal excitability in vitro are caused by withdrawal from chronic in vivo ethanol administration. Morton B, Ripley TL, Whittington MA, Butterworth AR, Little HJ. Alcohol Alcohol. 1992 Jan;27(1):71-9.

Nitrendipine, given during drinking, decreases the electrophysiological changes in the isolated hippocampal slice, seen during ethanol withdrawal. Whittington MA, Little HJ. Br J Pharmacol. 1991 Jul;103(3):1677-84.

Chronic dihydropyridine treatment can reverse the behavioural consequences of and prevent adaptations to, chronic ethanol treatment. Whittington MA, Dolin SJ, Patch TL, Siarey RJ, Butterworth AR, Little HJ. Br J Pharmacol. 1991 Jul;103(3):1669-76.

A calcium channel antagonist stereoselectively decreases ethanol withdrawal hyperexcitability but not that due to bicuculline, in hippocampal slices. Whittington MA, Little HJ. Br J Pharmacol. 1991 Jun;103(2):1313-20.

Patterns of changes in field potentials in the isolated hippocampal slice on withdrawal from chronic ethanol treatment of mice in vivo. Whittington MA, Little HJ. Brain Res. 1990 Jul 23;523(2):237-44.

Action of chlormethiazole in a model of ethanol withdrawal. Green AR, Davies EM, Little HJ, Whittington MA, Cross AJ. Psychopharmacology (Berl). 1990;102(2):239-42.

Effects of dihydropyridine calcium channel antagonists in ethanol withdrawal; doses required, stereospecificity and actions of Bay K 8644. Littleton JM, Little HJ, Whittington MA. Psychopharmacology (Berl). 1990;100(3):387-92.


Prof R. D. Traub, Dept. of Physiology, SUNY, New York

Prof N. Kopell, Dept. Mathematics, Univ. Boston

Prof H. Monyer, Max Planck Institute, Heidelberg

Dr Fiona Randall, Eisai Inc. Andover

Prof Antal Rot, QMU, London

Dr Martin Gillies, John Radcliffe Hospital, Oxford

Prof tengis Gloveli, Charite, Berlin