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 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.

If sensations were behaviorally conceived, as they should be, as complex functional patterns of interaction between overt behavior and the environment, there would be no point in searching for them as instantaneous psychic elements (qualia) within the brain or as internal products of the brain.

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.

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.

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 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.

In Gray's conjecture, mismatches in the subicular comparator (needing problem resolution) and matches (during appetitive approach) have equal prominence in consciousness. In rival cognitive views novelty and difficulty (i.e., information-processing mismatches) especially elicit more conscious modes of cognition and higher levels of self-regulation. The mismatch between Gray's conjecture and these views is discussed.

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

Gray extrapolates from circuit models of psychopathology to propose neural substrates for the contents of consciousness. I raise three concerns: (1) knowledge of synaptic arrangements may be inadequate to fully support his model; (2) latent inhibition deficits in schizophrenia, a focus of this and related models, are complex and deserve replication; and (3) this conjecture omits discussion of the neuropsychological basis for the contents of the unconscious.

An active role for conscious processes in the production of behaviour is proposed, involving top level controls in a hierarchy of behavioural control. It is suggested that by inhibiting or sensitizing lower levels in the hierarchy conscious processes can play a role in the organization of ongoing behaviour. Conscious control can be more or less evident, according to prevailing circumstances.

On the basis of neuropsychological evidence, it is clear that attention should be given a role in any model (or conjecture) 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 thirdperson 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 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.

A hypothesis about sensorimotor integration (the λ model) is described and applied to movement control and kinesthesia. The central idea is that the nervous system organizes positional frames of reference for the sensorimotor apparatus and produces active movements by shifting the frames in terms of spatial coordinates. Kinematic and electromyographic patterns are not programmed, but emerge from the dynamic interaction among the system s components, including external forces within the designated frame of reference. Motoneuronal threshold properties and proprioceptive inputs to motoneurons may be cardinal components of the physiological mechanism that produces positional frames of reference. The hypothesis that intentional movements are produced by shifting the frame of reference is extended to multi-muscle and multi-degrees-of-freedom systems with a solution of the redundancy problem that allows the control of a joint alone or in combination with other joints to produce any desired limb configuration and movement trajectory. The model also implies that for each motor behavior, the nervous system uses a strategy that minimizes the number of changeable control variables and keeps the parameters of these changes invariant. Examples are provided of simulated kinematic and electromyographic signals from single- and multi-joint arm movements produced by suggested patterns of control variables. Empirical support is provided and additional tests of the model are suggested. The model is contrasted with others based on the ideas of programming of motoneuronal activity, muscle forces, stiffness, or movement kinematics.

The spring-like behaviour of a joint following a sudden change of torque is partly a result of the elastic properties of tendons. A large fall in a muscle with a long tendon may be accompanied by tendon recoil causing joint movements as large as 20°.

Two aspects of the target article, (1) the extension of the equilibrium point theory to multi-joint movements, and (2) the consequence that the EMG pattern is not directly controlled by the central nervous system (CNS), are discussed in light of the experiments on upper trunk bending in humans. The principle component kinematic analysis and the analysis of the EMG data, obtained under microgravity and additional loading conditions, support the application of Feldman and Levin's for multi-joint pointing movement to equilibrium control during upper trunk movement.

We emphasize the relevance to cognitive psychology of Feldman and Levin's theoretical position. Traditional views of motor control have failed to clearly separate “production control” at the level of motor command, based on task-independent CV (control variables), from intentional “product control” based on task-dependent parameters. Because F&L's approach concentrates on the first process (trajectory formation), it can distinguish the product control stage.

The conservative strategy proposed by the authors suggests a solution of the degrees-of-freedom problem of the controller. However, several simple motor control tasks cannot be explained by this strategy. A nonconservative strategy, in which more parameters of the control signal vary, can account for these simple motor tasks. However, the simplicity that distinguishes the proposed model from many others is lost.

The lambda model provides a physiologically grounded terminology for describing muscle function and emphasizes the important influence of environmental and reflex-mediated effects on final states. However, lambda itself is only a convenient point on the length-tension curve; its importance should not be overemphasized. Ascribing movement to changes in a lambda-based frame of reference is generally valid, but it leaves unanswered a number of questions concerning mechanisms.