theta entrainment of gamma modules: effects of heterogeneity and non-stationarity

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POSTER PRESENTATION Open Access Theta entrainment of gamma modules: effects of heterogeneity and non-stationarity Ruben A Tikidji-Hamburyan 1,2* , Carmen C Canavier 1 From Twenty First Annual Computational Neuroscience Meeting: CNS*2012 Decatur, GA, USA. 21-26 July 2012 Transient synchronization of neurons in the gamma range (30-100 Hz) is believed to play an important role in attention, memory tasks[1] and other cognitive func- tions[2]. In many brain regions, gamma oscillations are modulated by the lower frequency theta rhythm[3]. There are at least two open questions with respect to gamma frequency synchronization: how this synchroni- zation might occur between distal regions without direct connections and whether inputs from slow theta oscilla- tions can synchronize populations of neurons firing at gamma frequency. It is well known that a low frequency input is capable of phase-locking a higher frequency oscillator, thus a population of gamma neurons with similar frequencies may synchronize due to common theta input. Here we exploit maps based upon the phase resetting curves [4] (PRCs) to examine minimal model of a feed- forward network of oscillating neurons and to derive existence and stability criteria for phase-locking of high frequency oscillators by a common theta input. Then we extend these results for heterogeneous network with Gaussian-distributed periods and show good agreement among direct simulation of 200,000 Morris-Lecar model neurons, results obtained by PRC mapping of the same population, and theoretical predictions. These results may be extended to a special case when biological noise is approximated by treating the period of each gamma oscillator as non-stationary, specifically as an Ornstein- Uhlenbeck process (OUP) with a fixed time constant and variance. Good agreement between theory and the PRC mapping results were obtained for small variance and a sufficiently large time constant of the OUP such that the time scale is slower than the rate of convergence of gamma neurons to their preferred phase. However, the limitation of the method is that the predicted and actual distributions diverge with increasing OUP variance as well as with decreasing time constant. For example, if the PRC has two stable branches and the period varies rapidly compared to the speed of attraction of the stable locking point, phases corresponding to the unstable branches will have an inflated probability density com- pared to our theoretical predictions due to transient crossing of these phases as the fixed point switches between branches. Finally we adopt this theory for the case in which there are multiple theta inputs to the gamma population and show that regions of existence and stability can thereby be significantly extended. Moreover, this result suggests a possible control scheme for switching the activity of a gamma population between synchrony, a cluster solution or a completely desynchronized pattern by varying the phase between theta inputs, which could theoretically lead to theta gamma nesting, which is widely observed experimentally. Author details 1 Department of Cell Biology and Anatomy, Louisiana State University, New Orleans, Louisiana 70005, USA. 2 A.B. Research Institute for Nurocybernatics, Southern Federal University, Rostov-on-Don, 344090, Russia. Published: 16 July 2012 References 1. Jensen O, Kaiser J, Lachaux JP: Human gamma-frequency oscillations associated with attention and memory. Trends Neurosci 2007, 30:317-324. 2. Gray CM: Synchronous oscillations in neuronal systems: Mechanisms and functions. J.Comput. Neurosci 1994, 1:11-38. 3. Buzsaki G: Rhythms of the brain. New York: Oxford University Press Inc; 2006. 4. Canavier CC, Achuthan S: Pulse coupled oscillators and the phase resetting curve. Math. Biosci 2010, 226:77-96. * Correspondence: [email protected] 1 Department of Cell Biology and Anatomy, Louisiana State University, New Orleans, Louisiana 70005, USA Full list of author information is available at the end of the article Tikidji-Hamburyan and Canavier BMC Neuroscience 2012, 13(Suppl 1):P170 http://www.biomedcentral.com/1471-2202/13/S1/P170 © 2012 Tikidji-Hamburyan and Canavier; licensee BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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Page 1: Theta entrainment of gamma modules: effects of heterogeneity and non-stationarity

POSTER PRESENTATION Open Access

Theta entrainment of gamma modules: effects ofheterogeneity and non-stationarityRuben A Tikidji-Hamburyan1,2*, Carmen C Canavier1

From Twenty First Annual Computational Neuroscience Meeting: CNS*2012Decatur, GA, USA. 21-26 July 2012

Transient synchronization of neurons in the gammarange (30-100 Hz) is believed to play an important rolein attention, memory tasks[1] and other cognitive func-tions[2]. In many brain regions, gamma oscillations aremodulated by the lower frequency theta rhythm[3].There are at least two open questions with respect togamma frequency synchronization: how this synchroni-zation might occur between distal regions without directconnections and whether inputs from slow theta oscilla-tions can synchronize populations of neurons firing atgamma frequency. It is well known that a low frequencyinput is capable of phase-locking a higher frequencyoscillator, thus a population of gamma neurons withsimilar frequencies may synchronize due to commontheta input.Here we exploit maps based upon the phase resetting

curves [4] (PRCs) to examine minimal model of a feed-forward network of oscillating neurons and to deriveexistence and stability criteria for phase-locking of highfrequency oscillators by a common theta input. Then weextend these results for heterogeneous network withGaussian-distributed periods and show good agreementamong direct simulation of 200,000 Morris-Lecar modelneurons, results obtained by PRC mapping of the samepopulation, and theoretical predictions. These resultsmay be extended to a special case when biological noiseis approximated by treating the period of each gammaoscillator as non-stationary, specifically as an Ornstein-Uhlenbeck process (OUP) with a fixed time constant andvariance. Good agreement between theory and the PRCmapping results were obtained for small variance and asufficiently large time constant of the OUP such that thetime scale is slower than the rate of convergence of

gamma neurons to their preferred phase. However, thelimitation of the method is that the predicted and actualdistributions diverge with increasing OUP variance aswell as with decreasing time constant. For example, if thePRC has two stable branches and the period variesrapidly compared to the speed of attraction of the stablelocking point, phases corresponding to the unstablebranches will have an inflated probability density com-pared to our theoretical predictions due to transientcrossing of these phases as the fixed point switchesbetween branches.Finally we adopt this theory for the case in which

there are multiple theta inputs to the gamma populationand show that regions of existence and stability canthereby be significantly extended. Moreover, this resultsuggests a possible control scheme for switching theactivity of a gamma population between synchrony, acluster solution or a completely desynchronized patternby varying the phase between theta inputs, which couldtheoretically lead to theta – gamma nesting, which iswidely observed experimentally.

Author details1Department of Cell Biology and Anatomy, Louisiana State University, NewOrleans, Louisiana 70005, USA. 2A.B. Research Institute for Nurocybernatics,Southern Federal University, Rostov-on-Don, 344090, Russia.

Published: 16 July 2012

References1. Jensen O, Kaiser J, Lachaux JP: Human gamma-frequency oscillations

associated with attention and memory. Trends Neurosci 2007, 30:317-324.2. Gray CM: Synchronous oscillations in neuronal systems: Mechanisms and

functions. J.Comput. Neurosci 1994, 1:11-38.3. Buzsaki G: Rhythms of the brain. New York: Oxford University Press Inc;

2006.4. Canavier CC, Achuthan S: Pulse coupled oscillators and the phase

resetting curve. Math. Biosci 2010, 226:77-96.* Correspondence: [email protected] of Cell Biology and Anatomy, Louisiana State University, NewOrleans, Louisiana 70005, USAFull list of author information is available at the end of the article

Tikidji-Hamburyan and Canavier BMC Neuroscience 2012, 13(Suppl 1):P170http://www.biomedcentral.com/1471-2202/13/S1/P170

© 2012 Tikidji-Hamburyan and Canavier; licensee BioMed Central Ltd. This is an Open Access article distributed under the terms of theCreative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, andreproduction in any medium, provided the original work is properly cited.

Page 2: Theta entrainment of gamma modules: effects of heterogeneity and non-stationarity

doi:10.1186/1471-2202-13-S1-P170Cite this article as: Tikidji-Hamburyan and Canavier: Theta entrainmentof gamma modules: effects of heterogeneity and non-stationarity. BMCNeuroscience 2012 13(Suppl 1):P170.

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Tikidji-Hamburyan and Canavier BMC Neuroscience 2012, 13(Suppl 1):P170http://www.biomedcentral.com/1471-2202/13/S1/P170

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