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      Neural masses and fields in dynamic causal modeling

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          Abstract

          Dynamic causal modeling (DCM) provides a framework for the analysis of effective connectivity among neuronal subpopulations that subtend invasive (electrocorticograms and local field potentials) and non-invasive (electroencephalography and magnetoencephalography) electrophysiological responses. This paper reviews the suite of neuronal population models including neural masses, fields and conductance-based models that are used in DCM. These models are expressed in terms of sets of differential equations that allow one to model the synaptic underpinnings of connectivity. We describe early developments using neural mass models, where convolution-based dynamics are used to generate responses in laminar-specific populations of excitatory and inhibitory cells. We show that these models, though resting on only two simple transforms, can recapitulate the characteristics of both evoked and spectral responses observed empirically. Using an identical neuronal architecture, we show that a set of conductance based models—that consider the dynamics of specific ion-channels—present a richer space of responses; owing to non-linear interactions between conductances and membrane potentials. We propose that conductance-based models may be more appropriate when spectra present with multiple resonances. Finally, we outline a third class of models, where each neuronal subpopulation is treated as a field; in other words, as a manifold on the cortical surface. By explicitly accounting for the spatial propagation of cortical activity through partial differential equations (PDEs), we show that the topology of connectivity—through local lateral interactions among cortical layers—may be inferred, even in the absence of spatially resolved data. We also show that these models allow for a detailed analysis of structure–function relationships in the cortex. Our review highlights the relationship among these models and how the hypothesis asked of empirical data suggests an appropriate model class.

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          Most cited references60

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          Dynamic causal modelling.

          In this paper we present an approach to the identification of nonlinear input-state-output systems. By using a bilinear approximation to the dynamics of interactions among states, the parameters of the implicit causal model reduce to three sets. These comprise (1) parameters that mediate the influence of extrinsic inputs on the states, (2) parameters that mediate intrinsic coupling among the states, and (3) [bilinear] parameters that allow the inputs to modulate that coupling. Identification proceeds in a Bayesian framework given known, deterministic inputs and the observed responses of the system. We developed this approach for the analysis of effective connectivity using experimentally designed inputs and fMRI responses. In this context, the coupling parameters correspond to effective connectivity and the bilinear parameters reflect the changes in connectivity induced by inputs. The ensuing framework allows one to characterise fMRI experiments, conceptually, as an experimental manipulation of integration among brain regions (by contextual or trial-free inputs, like time or attentional set) that is revealed using evoked responses (to perturbations or trial-bound inputs, like stimuli). As with previous analyses of effective connectivity, the focus is on experimentally induced changes in coupling (cf., psychophysiologic interactions). However, unlike previous approaches in neuroimaging, the causal model ascribes responses to designed deterministic inputs, as opposed to treating inputs as unknown and stochastic.
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            The free-energy principle: a rough guide to the brain?

            This article reviews a free-energy formulation that advances Helmholtz's agenda to find principles of brain function based on conservation laws and neuronal energy. It rests on advances in statistical physics, theoretical biology and machine learning to explain a remarkable range of facts about brain structure and function. We could have just scratched the surface of what this formulation offers; for example, it is becoming clear that the Bayesian brain is just one facet of the free-energy principle and that perception is an inevitable consequence of active exchange with the environment. Furthermore, one can see easily how constructs like memory, attention, value, reinforcement and salience might disclose their simple relationships within this framework.
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              Voltage oscillations in the barnacle giant muscle fiber.

              Barnacle muscle fibers subjected to constant current stimulation produce a variety of types of oscillatory behavior when the internal medium contains the Ca++ chelator EGTA. Oscillations are abolished if Ca++ is removed from the external medium, or if the K+ conductance is blocked. Available voltage-clamp data indicate that the cell's active conductance systems are exceptionally simple. Given the complexity of barnacle fiber voltage behavior, this seems paradoxical. This paper presents an analysis of the possible modes of behavior available to a system of two noninactivating conductance mechanisms, and indicates a good correspondence to the types of behavior exhibited by barnacle fiber. The differential equations of a simple equivalent circuit for the fiber are dealt with by means of some of the mathematical techniques of nonlinear mechanics. General features of the system are (a) a propensity to produce damped or sustained oscillations over a rather broad parameter range, and (b) considerable latitude in the shape of the oscillatory potentials. It is concluded that for cells subject to changeable parameters (either from cell to cell or with time during cellular activity), a system dominated by two noninactivating conductances can exhibit varied oscillatory and bistable behavior.
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                Author and article information

                Journal
                Front Comput Neurosci
                Front Comput Neurosci
                Front. Comput. Neurosci.
                Frontiers in Computational Neuroscience
                Frontiers Media S.A.
                1662-5188
                21 February 2013
                28 May 2013
                2013
                : 7
                : 57
                Affiliations
                [1] 1Wellcome Trust Centre for Neuroimaging, Institute of Neurology, University College London London, UK
                [2] 2Virginia Tech Carilion Research Institute, Virginia Tech Roanoke, VA, USA
                [3] 3Bradley Department of Electrical and Computer Engineering, Virginia Tech Blacksburg, VA, USA
                Author notes

                Edited by: Peter Robinson, University of Sydney, Australia

                Reviewed by: Peter Robinson, University of Sydney, Australia; James Roberts, Queensland Institute of Medical Research, Australia

                *Correspondence: Rosalyn Moran, Virginia Tech Carilion Research Institute, Virginia Tech, 2 Riverside Drive, Roanoke, 24016 VA, USA e-mail: rosalynj@ 123456vtc.vt.edu

                †These authors have contributed equally to this work.

                Article
                10.3389/fncom.2013.00057
                3664834
                23755005
                a43724ac-36af-4b32-bf73-9d08c410174b
                Copyright © 2013 Moran Pinotsis and Friston.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in other forums, provided the original authors and source are credited and subject to any copyright notices concerning any third-party graphics etc.

                History
                : 16 January 2013
                : 21 April 2013
                Page count
                Figures: 3, Tables: 0, Equations: 1, References: 90, Pages: 12, Words: 10078
                Categories
                Neuroscience
                Review Article

                Neurosciences
                dynamic causal modeling,electroencephalography,magnetoencephalography (meg),local field potential (lfp),neural mass models

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