We outline fresh findings that show that our macroscopic electrocorticographic (ECoG) simulations can account for synchronous multiunit pulse oscillations at separate, simultaneously activated cortical sites and the associated gamma-band ECoG activity. We clarify our views on the approximations of dynamic class applicable to neural events at macroscopic and microscopic scales, and the analogies drawn to classes of ANN behaviour. We accept the need to introduce memory processes and detailed anatomical and physiological information into any future developments of our simulations. On (...) the issue of intrinsic cortical stability and the role of extrinsic fibre systems in maintaining stability, we argue that this position is not in extreme contradiction to those of our commentators, and that the mechanisms implicit in our simulations' properties imply rich computational possibilities. We discuss some of the reasons for and against the existence of significant global resonances in the brain and explain why such behaviour appears absent in our simulations. Last, we discuss other phenomena, such as rhythmic driving of the cortex, which have not yet been introduced into our models, and indicate lines for future development of the simulations. (shrink)

There is some complementarity of models for the origin of the electroencephalogram (EEG) and neural network models for information storage in brainlike systems. From the EEG models of Freeman, of Nunez, and of the authors' group we argue that the wavelike processes revealed in the EEG exhibit linear and near-equilibrium dynamics at macroscopic scale, despite extremely nonlinear – probably chaotic – dynamics at microscopic scale. Simulations of cortical neuronal interactions at global and microscopic scales are then presented. The simulations depend (...) on anatomical and physiological estimates of synaptic densities, coupling symmetries, synaptic gain, dendritic time constants, and axonal delays. It is shown that the frequency content, wave velocities, frequency/wavenumber spectra and response to cortical activation of the electrocorticogram (ECoG) can be reproduced by a “lumped” simulation treating small cortical areas as single-function units. The corresponding cellular neural network simulation has properties that include those of attractor neural networks proposed by Amit and by Parisi. Within the simulations at both scales, sharp transitions occur between low and high cell firing rates. These transitions may form a basis for neural interactions across scale. To maintain overall cortical dynamics in the normal low firing-rate range, interactions between the cortex and the subcortical systems are required to prevent runaway global excitation. Thus, the interaction of cortex and subcortex via corticostriatal and related pathways may partly regulate global dynamics by a principle analogous to adiabatic control of artificial neural networks. (shrink)

I would like to emphasize the significance of chaotic dynamics at both local and macroscopic levels in the cortex. The basic notions dealt with in this commentary will be noise-induced order, chaotic “itinerancy” and dissipative structure. Wright & Laley's theory would be partially misleading, since emergent nonlinearity rather than the linearity at even a macroscopic level can actually subserve cortical functions.

Some dichotomies related to modeling electrocortical activities are analyzed. Attractor neural networks versus biologically motivated models, near-equilibrium versus nonequilibrium processes, linear and nonlinear dynamics, stochastic and chaotic patterns, local and global scale simulation of cortical activities are discussed.

Wright & Liley contrast their theory that the global dynamics of the EEG are linear with that of Freeman, who hypothesizes an EEG governed by (nonlinear) deterministic-chaotic dynamics. A “call for further discussion” on the part of the authors is made as to how either theory fits with experimental findings indicating that EEG dynamics are non-linear but stochastic.

For some years there has been a controversy about whether brain state variables such as EEG or neuronal spike trains exhibit chaotic behaviour. Wright & Liley claim that the local dynamics measured by spike trains or local field potentials exhibit chaotic behaviour, but global measures like EEG should be governed by linear dynamics. We propose a different scheme. Based on simulation studies and various experiments, we suggest that the pointwise dimension of EEG time series may provide some valuable information about (...) underlying neuronal generators. (shrink)

The unification of microscopic and macroscopic models of brain behaviour is of paramount importance and Wright & Liley's target article provides some important groundwork. In this commentary, I propose that a useful approach for the future is to incorporate a developmental perspective into such models. This may be an important constraint, providing a key to understanding the nature of macroscopic measures of brain function such as functional measures like ERP.

Multiscale dynamics, linear approximations, global boundary conditions, experimental verification, and global influences on local cell assemblies are considered in the context of Wright & Liley's work. W&L provide a nice introduction to these issues and a reasonable simulation of intermediate scale dynamics, but the model does not adequately simulate combined local and global processes.

Event-related potentials (ERPs) – neglected almost entirely by Wright & Liley – allow objective investigation of information processing in the brain. The application of chaos theory to such an analysis broadens this possibility. Through the use of the point correlation dimension (PD2) accurate dimensional analysis of different Event-Related Potential components such as the P3 wave is possible.

Simulation of brain dynamics requires the use of accurate empirical data. This commentary points out major errors in some of the empirical data used in Wright & Laley's simulation. The simulation is quantitatively very different from the real cortex, and may also have important qualitative differences.

We present simulation results of an olfactory cortex model complementing the results presented in Wright & Liley's target article. We show how the cortical dynamics as expressed in EEG can be regulated by neuromodulation and discuss how the system can attain global stability without cortical-subcortical interaction, as presumed necessary by Wright & Liley. Network structure is shown to be crucial.

We focus on one aspect of Wright & Liley's target article: the linearity of the EEG. According to the authors, some nonlinear models of the cortex can be reduced (approximated) to the linear case at the millimetric scale. We argue here that the statement about the linear character of EEG is too strong and that EEG exhibits nonlinear features which cannot be ignored.

To model the organization of levels' of cortical dynamics, at least some general scheme for hierarchy, functional diversity, and proper intrinsic control must be provided. Rhythmic control forces the system to iterate its state by short trajectories, which makes it much more stable and predictable without discarding the desirable ability of chaotic systems to make rapid phase transitions. Rhythmic control provides a fundamentally different systems dynamics, one not provided by models that allow the emergence of continuous trajectories in the systems (...) state space. (shrink)

The work in progress reported by Wright & Liley shows great promise, primarily because of their experimental and simulation paradigms. However, their tentative conclusion that macroscopic neocortex may be considered (approximately) a linear near-equilibrium system is premature and does not correspond to tentative conclusions drawn from other studies of neocortex.

Although this is an impressive piece of modeling work, I worry that the two models that Wright & Liley have created do not yet provide us with useful empirical information regarding brain processing.

Two hypotheses concerning nonlinear elements in complex systems are contrasted: that neurons, intrinsically unstable, are stabilized through embedding in networks and populations; and, conversely, that cortical neurons are intrinsically stable, but are destabilized through embedding in cortical populations and corticostriatal feedback systems. Tests are made by piecewise linearization of nonlinear dynamics at nonequilibriumoperating points, followed by linear stability analysis.

Wright & Liley provide an advance in addressing the interaction of multiple scales of processing in the brain. It should address in more detail the biological evidence that underlies the models it proposes to replace.

Does the Wright & Liley model predict: (1) that subdural and hippocampal EEGs coherence tend to rise and fall in parallel for many frequencies, (2) that it is locally high or low within 10mm and falls steeply on average or, (3) that it is in constant flux, mostly rising and falling within 5–15 sec?

Some concepts relating to learned, structured functioning of local modules in neocortex are clarified in order to ensure that the integration from the small scale to the global attempted by Wright & Liley does not miss the target.