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.

It is widely accepted among philosophers that neuroscientists are conducting a search for the neural correlates of consciousness, or NCC. Chalmers conceptualized this research program as the attempt to correlate the contents of conscious experience with the contents of representations in specific neural populations. A notable claim on behalf of this interpretation is that the neutral language of “correlates” frees us from philosophical disputes over the mind/body relation, allowing the science to move independently. But the experimental paradigms and explanatory canons (...) of neuroscience are not neutral about the mechanical relation between consciousness and the brain. I argue that NCC research is best characterized as an attempt to locate a causally relevant neural mechanism and not as an effort to identify a discrete neural representation, the content of which correlates with some actual experience. It might be said that the first C in “NCC” should stand for “causes” rather than “correlates.”. (shrink)

A spate of recent anti-localizationist publications have re-ignited the old debate about the localization of function. Many of the recent attacks on localization, however, are directed at what I will argue to be a narrow and outmoded view of localization, and thus have little conceptual or empirical impact. What I hope to present here is an analysis of functional localization that more adequately reflects the sophistication and complexity of its use in neuroscientific research, both historically and recently. Proceeding first by (...) way of contrast, I examine theanti-localizationist positions of holism andequipotentiationism. Then, I present a four-fold analysis of localization according to physical scope, physical kind, functional scope, and functional kind. Next, I turn to a discussion of the heuristic value oflocalization in deciphering structure-functionrelationships. Finally, I hope to show that the overall view of functional localization that emerges from these considerations constitutes a much more elusive target than its critics assume. It serves to mitigate, and insome instances even defeat, some forms ofanti-localizationist criticisms. (shrink)

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?

If successive, brief visual images are exposed for recognition or for psychophysical ratings, various effects and phenomena of fast dynamics of conscious perception such as mutual masking, metacontrast, proactive enhancement of contrast, proactive speed-up of the latency of subjective visual experience, the Fröhlich Effect, the Tandem Effect, attentional facilitation by visuospatial precuing, and some others have been found. The theory proposed to deal with these phenomena proceeds from the assumption that two types of brain processes are necessary in order to (...) consciously recognize visual stimuli: fast, specific processes of encoding that allocate and reactivate the stimulus representation which is based on the activity of selected cortical neurons and relatively slower processes of facilitation of the activity of this specific representation that are mediated by the excitatory modulation of the EPSPs of those selected cortical neurons by the ascending input from nonspecific thalamus. Theperceptual retouchconstruct is proposed in order to characterize and analyze the interaction of and . The neurophysiological characteristics of this bifunctional system of afference help to put forward several predictions that are found to be consistent with the empirical regularities of the above-described perceptual-attentional phenomena. These data form a body of converging evidence that is consistent with the predictions of the perceptual retouch approach. (shrink)

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.

Before one can even begin to model consciousness and what exactly it means that it is a subjective phenomenon one needs a theory about what a first-person perspective really is. This theory has to be conceptually convincing, empirically plausible and, most of all, open to new developments. The chosen conceptual framework must be able to accommodate scientific progress. Its ba- sic assumptions have to be plastic as it were, so that new details and empirical data can continuously be fed into (...) the theoretical model as it grows and becomes more refined. This paper makes an attempt at sketching the outlines of such a theory, offering a representationalist analysis of the phenomenal first-person perspective. Three phenomenal target properties are centrally relevant. (shrink)

This is a bibliography of books and articles on consciousness in philosophy, cognitive science, and neuroscience over the last 30 years. There are three main sections, devoted to monographs, edited collections of papers, and articles. The first two of these sections are each divided into three subsections containing books in each of the main areas of research. The third section is divided into 12 subsections, with 10 subject headings for philosophical articles along with two additional subsections for articles in cognitive (...) science and neuroscience. Of course the division is somewhat arbitrary, but I hope that it makes the bibliography easier to use. This bibliography has first been compiled by Thomas Metzinger and David Chalmers to appear in print in two philosophical anthologies on conscious experience . From 1995 onwards it has been continuously updated by Thomas Metzinger, and now is freely available as a PDF-, RTF-, or HTML-file. This bibliography mainly attempts to cover the Anglo-Saxon and German debates, in a non-annotated, fully formatted way that makes it easy to "cut and paste" from the original file. To a certain degree this bibliography also contains items in other languages than English and German - all submissions in other languages are welcome. Last update of current version: July 13th, 2001. (shrink)

In attempts to formulate a computational understanding of brain function, one of the fundamental concerns is the data structure by which the brain represents information. For many decades, a conceptual framework has dominated the thinking of both brain modelers and neurobiologists. That framework is referred to here as "classical neural networks." It is well supported by experimental data, although it may be incomplete. A characterization of this framework will be offered in the next section. Difficulties in modeling important functional aspects (...) of the brain on the basis of classical neural networks alone have led to the recognition that another, general mechanism must be invoked to explain brain function. That mechanism I call " binding." Binding by neural signal synchrony had been mentioned several times in the liter ature before it was fully formulated as a general phenomenon. Although experimental evidence for neural syn chrony was soon found, the idea was largely ignored for many years. Only recently has it become a topic of animated discussion. In what follows, I will summarize the nature and the roots of the idea of binding, especially of temporal binding, and will discuss some of the objec tions raised against it. (shrink)

In the past decade, the notion of a neural correlate of consciousness (or NCC) has become a focal point for scientific research on consciousness (Metzinger, 2000a). A growing number of investigators believe that the first step toward a science of consciousness is to discover the neural correlates of consciousness. Indeed, Francis Crick has gone so far as to proclaim that ‘we … need to discover the neural correlates of consciousness.… For this task the primate visual system seems especially attractive.… No (...) longer need one spend time attempting … to endure the tedium of philosophers perpetually disagreeing with each other. Con- sciousness is now largely a scientific problem’ (Crick, 1996, p. 486).2 Yet the question of what it means to be a neural correlate of consciousness is actually far from straightforward, for it involves fundamental empirical, methodological, and _philosophical _issues about the nature of consciousness and its relationship to the brain. Even if one assumes, as we do, that states of consciousness causally depend on states of the brain, one can nevertheless wonder in what sense there is, or could be, such a thing as a neural correlate of consciousness. (shrink)