There is considerable evidence that the hippocampal system contributes both to (1) the temporary maintenance of memories and to (2) the processing of a particular type of memory representation. The findings on amnesia suggest that these two distinguishing features of hippocampal memory processing are orthogonal. Together with anatomical and physiological data, the neuropsychological findings support a model of cortico-hippocampal interactions in which the temporal and representational properties of hippocampal memory processing are mediated separately. We propose that neocortical association areas maintain (...) shortterm memories for specific items and events prior to hippocampal processing as well as providing the final repositories of long-term memory. The parahippocampal region supports intermediate-term storage of individual items, and the hippocampal formation itself mediates an organization of memories according to relevant relationships among items. Hippocampal-cortical interactions produce (i) strong and persistent memories for events, including their constituent elements and the relationships among them, and (ii) a capacity to express memories flexibly across a wide range of circumstances. (shrink)

Gray's comparator model fails to provide an adequate explanation of consciousness for two reasons. First, it is based on a narrow definition of consciousness that excludes basic phenomenology and active functions of consciousness. Second, match/mismatch decisions can be made without producing an experience of consciousness. The model thus violates the sufficiency criterion.

Gray extrapolates from circuit models of psychopathology to propose neural substrates for the contents of consciousness. I raise three concerns: knowledge of synaptic arrangements may be inadequate to fully support his model; latent inhibition deficits in schizophrenia, a focus of this and related models, are complex and deserve replication; and this conjecture omits discussion of the neuropsychological basis for the contents of the unconscious.

Gray's neuropsychological model of consciousness uses a hierarchical feedback loop framework that has been extensively discussed by many others in psychology. This commentary therefore urges Gray to integrate with, or at least acknowledge previous models. It also points out flaws in his feedback model and suggests directions for further theoretical work.

Gray has expanded his account of schizophrenia to explain consciousness as well. His theory explains neither phenomenon adequately because he treats individual minds in isolation. The primary function of consciousness is to permit high level interactions with other conscious beings. The key symptoms of schizophrenia reflect a failure of this mechanism.

It is accepted that “primary awareness” may emerge from the integration of two classes of information. It is unclear, however, why this cannot take place within the comparator rather than in conjunction with feedback to the perceptual systems. The model has plausibility in relation to the continuity of conscious experience in the normal waking state and may be extended to encompass certain aspects of the “sense of self” which are frequently disrupted in psychotic patients.

Perspective and reflection have each been considered in some way basic to phenomenal consciousness. Each has possible evolutionary value, though neither seems sufficient for consciousness. Consider an account of consciousness in terms of the combination of perspective and reflection, its relationship to the problem of other minds, and its capacity to inherit evolutionary explanation from its components.

Gray's account is remarkable in its depth and scope but too little attention is paid to poor correspondences with the literature on hippocampal/subicular damage, the theta rhythm, and novelty detection. An alternative account, focusing on hippocampal involvement in organizing memories in a way that makes them accessible to conscious recollection but not in access to consciousness per se, avoids each of these limitations.

Gray hypothesizes that the contents of consciousness correspond to the outputs of a subicular (hippocampal/temporal lobe) comparator that compares the current state of the organism's perceptual world with a predicted state. I argue that Gray has identified a key contributing system to conscious awareness, but that his model is inadequate for explaining how conscious contents are generated in the brain. An alternative model is offered.

Drawing on previous models of anxiety, intermediate memory, the positive symptoms of schizophrenia, and goal-directed behaviour, a neuropsychological hypothesis is proposed for the generation of the contents of consciousness. It is suggested that these correspond to the outputs of a comparator that, on a moment-by-moment basis, compares the current state of the organism's perceptual world with a predicted state. An outline is given of the information-processing functions of the comparator system and of the neural systems which mediate them. The hypothesis (...) appears to be able to account for a number of key features of the contents of consciousness. However, it is argued that neitherthis nor any existing comparable hypothesis is yet able to explain why the brain should generate conscious experience of any kind at all. (shrink)

In this commentary, I point out some weaknesses in Gray's target article and, in the light of that discussion, I attempt to delineate the kinds of problem a cognitive neuroscience of consciousness faces on its way to a scientific understanding of subjective experience.

To explore the mechanism of sensation correlations between EP component amplitude and signal detection indices were studied. The time of sensation coincided with the peak latency of those EP components that showed a correlation with both indices. The components presumably reflected information synthesis in projection cortical neurons. A mechanism providing the synthesis process is proposed.

The first claim in the target article was that there is as yet no transparent, causal account of the relations between consciousness and brain-and-behaviour. That claim remains firm. The second claim was that the contents of consciousness consist, psychologically, of the outputs of a comparator system; the third consisted of a description of the brain mechanisms proposed to instantiate the comparator. In order to defend these claims against criticism, it has been necessary to clarify the distinction between consciousness-as-such and the (...) contents of consciousness, to widen the description of the neural machinery instantiating the comparator system, and to clarify the relationship between the contents of consciousness in the here-and-now and episodic memory. (shrink)

Robust theories concerning the connection between consciousness and brain function should derive not only from empirical evidence but also from a well grounded inind-body ontology. In the case of the comparator hypothesis, Gray develops his ideas relying extensively on empirical evidence, but he bounces irresolutely among logically incompatible metaphysical theses which, in turn, leads him to excessively skeptical conclusions concerning the naturalization of consciousness.

As Gray insists, his comparator model proposes a brute correlation only – of consciousness with septohippocampal output. I suggest that the comparator straddles a feedback loop that boosts the activation ofnovelrepresentations, thus helping them feature in present or recollected experience. Such a role in organizing conscious contents would transcend correlation and help explain how consciousness emerges from brain function.

Gray, like other recent authors, seeks a scientific approach to consciousness, but fails to provide a biologically convincing description, partly because he implicitly bases his model on a computationalist foundation that embeds the contents of thought in irreducible symbolic representations. When patterns of neural activity instantiating conscious thought are shorn of homuncular observers, it appears most likely that these patterns and the circuitry that compares them with memories and plans should be found distributed over large regions of neocortex.

Gray takes an information-processing paradigm as his departure point, invoking a comparator as part of the system. He concludes that consciousness is to be found “in” the comparator but is unable to point to how the comparison takes place. Thus, the comparator turns out not to be an entity arising out of brain research per se, but out of the logic of the paradigm. In this way, Gray both reinvents dualism and remains trapped in the language game of his own (...) model – ending up dealing with the unknowable. (shrink)

Segmentalized consciousness in schizophrenia reflects a loss of the normal Gestalt organization and contextualization of perception. Grays model explains such segmentalization in terms of septohippocampal dysfunction, which is consistent with known neuropsychological impairment in schizophrenia. However, other considerations suggest that everyday perception and its failure in schizophrenia also involve prefrontal executive mechanisms, which are only minimally elaborated by Gray.

The communicative aspects of the contents of consciousness are analyzed in the framework of a neural network model of animal communication. We discuss some issues raised by Gray, such as the control of the contents of consciousness, the adaptive value of consciousness, conscious and unconscious behaviors, and the nature of a model's consciousness.

Gray's integration of the different levels of description and explanation in his theory is problematic: The introduction of consciousness into his theorising consists of the mind-brain identity assumption, which tells us nothing new. There need not be correlations between levels of description. Gray's account does not extend beyond “brute” correlation. Integration must be achieved in a principled, mutually constraining way.

Gray unwisely melds together two distinguishable contributions of consciousness: one to epistemology, the other to evolution. He also renders consciousness needlessly invisible behaviorally.

Colonius suggests that, in using standard set theory as the language in which to express our computational-level theory of human memory, we would need to violate the axiom of foundation in order to express meaningful memory bindings in which a context is identical to an item in the list. We circumvent Colonius's objection by allowing that a list item may serve as a label for a context without being identical to that context. This debate serves to highlight the value of (...) specifying memory operations in set theoretic notation, as it would have been difficult if not impossible to formulate such an objection at the algorithmic level. (shrink)

On the basis of neuropsychological evidence, it is clear that attention should be given a role in any model of consciousness. What is known about the many instances of dissociation between explicit and implicit knowledge after brain damage suggests that conscious experience might not be linked to a restricted area of the brain. Even if it were true that there is a single brain area devoted to consciousness, the subicular area would seem to be an unlikely possibility.

This commentary elaborates on Gray's conclusion that his neurophysiological model of consciousness might explain how consciousness arises from the brain, but does not address how consciousness evolved, affects behaviour or confers survival value. The commentary argues that such limitations apply to all neurophysiological or other third-person perspective models. To approach such questions the first-person nature of consciousness needs to be taken seriously in combination with third-person models of the brain.

The commentary argues that we cannot be sure that human consciousness has survival value and that in order to understand the origins and, perhaps, the function of consciousness, we should examine the behavioural and neural precursors to consciousness in nonhumans. An example is given of research on the role of context in decisions regarding fleeing from probable predators in the Mongolian gerbil.