Online research in philosophy

Recent experiments have shown that the amplitudes of cortical gamma band oscillatory activities that occur during anesthesia are often greater than amplitudes of similar activities that occur without anesthesia. This result is apparently at odds with the hypothesis that synchronized oscillatory activities constitute the neural correlate of consciousness. We argue that while synchronization and oscillatory patterning are necessary conditions for consciousness, they are not sufficient. Based on the results of a binocular rivalry study of Fries et al. (1997), we propose that the degrees of oscillatory strength and synchronization of neuronal activities determine the degree of awareness those activities produce. On the other hand, the overal firing rates of neurons in cortical sensory areas are not correlated with the degree of awareness the activities of those neurons produce. The results of the experiment of Fries et al. (1997) appear to conflict with the results of another binocular rivalry experiment, in which monkeys were trained to pull a lever in order to report which stimulus object was being perceived (Leopold & Logothetis, 1996). In the latter experiment, it was demonstrated that the firing rates of neurons in striate cortex did not change during perceptual alterations, while 90% of neurons in inferior and superior temporal cortices changed their firing rate when the perceived image changed. This result led to the conclusion that activities in temporal cortex are correlated with visual awareness, but those in striate cortex are not. We argue that activities in temporal cortex contribute little, if anything, to perceptual awareness, and that their primary function is computational. Thus the correlation between the firing rates of neurons in these areas and the responses of the monkeys is due to the recognition of a particular stimulus object, which in turn is due to the computations made there.

Neuronal aggregates involved in conscious awareness are not evenly distributed throughout the CNS but comprise key components referred to as the neural network correlates of consciousness (NNCC). A critical node in this network is the posterior cingulate, precuneal, and retrosplenial cortices. The cytological and neurochemical composition of this region is reviewed in relation to the Brodmann map. This region has the highest level of cortical glucose metabolism and cytochrome c oxidase activity. Monkey studies suggest that the anterior thalamic projection likely drives retrosplenial and posterior cingulate cortex metabolism and that the midbrain projection to the anteroventral thalamic nucleus is a key coupling site between the brainstem system for arousal and cortical systems for cognitive processing and awareness. The pivotal role of the posterior cingulate, precuneal, and retrosplenial cortices in consciousness is demonstrated with posterior cingulate epilepsy cases, midcingulate lesions that de-afferent this region and are associated with unilateral sensory neglect, observations from stroke and vegetative state patients, alterations in blood flow during sleep, and the actions of general anesthetics. Since this region is critically involved in self reflection, it is not surprising that it is similarly a site for the NNCC. Interestingly, information processing during complex cognitive tasks and during aversive sensations such as pain induces efforts to terminate self reflection and result in decreased processing in posterior cingulate and precuneal cortices.

What are the neural correlates of conscious visual awareness? Tackling this question requires contrasting neural correlates of stimulus processing culminating in visual awareness with neural correlates of stimulus processing unaccompanied by awareness. To contrast these two neural states, one must be able to erase an otherwise visible stimulus from awareness. This paper describes and critiques visual phenomena involving dissociation of physical stimulation and conscious awareness: degraded stimulation, visual masking, visual crowding, bistable figures, binocular rivalry, motion-induced blindness, inattentional blindness, change blindness and attentional blink. While no single strategy stands above the others, those producing changing visual awareness despite invariant physical stimulation are clearly preferable.

It has been shown that visual awareness in the blind hemifield of hemianopic cats that have undergone unilateral ablations of visual cortex can be restored by sectioning the commissure of the superior colliculus or by destroying a portion of the substantia nigra contralateral to the cortical lesion (the Sprague effect). We propose that the visual awareness that is recovered is due to synchronized oscillatory activities in the superior colliculus ipsilateral to the cortical lesion. These oscillatory activities are normally partially suppressed by the inhibitory, GABAergic contralateral nigrotectal projection, and the destruction of the substantia nigra, or the sectioning of the collicular commissure, disinhibits the collicular neurons, causing an increase in the extent of oscillatory activity and/or synchronization between activities at different sites. This increase in the oscillatory and synchronized character is sufficient for the activities to give rise to visual awareness. We argue that in rodents and lower vertebrates, normal visual awareness is partly due to synchronized oscillatory activities in the optic tectum and partly due to similar activities in visual cortex. It is only in carnivores and primates that visual awareness is wholly due to cortical activities. Based on von Baerian recapitulation theory, we propose that, even in humans, there is a period in early infancy when visual awareness is partially due to activities in the superior colliculus, but that this awareness gradually disappears as the nigrotectal projection matures.

Based on theoretical considerations of Aurell (1979) and Block (1995), we argue that object recognition awareness is distinct from purely sensory awareness and that the former is mediated by neuronal activities in areas that are separate and distinct from cortical sensory areas. We propose that two of the principal functions of neuronal activities in sensory cortex, which are to provide sensory awareness and to effect the computations that are necessary for object recognition, are dissociated. We provide examples of how this dissociation might be achieved and argue that the components of the neuronal activities which carry the computations do not directly enter the awareness of the subject. The results of these computations are sparse representations (i.e., vector or distributed codes) which are activated by the presentation of particular sensory objects and are essentially engrams for the recognition of objects. These final representations occur in the highest order areas of sensory cortex; in the visual analyzer, the areas include the anterior part of the inferior temporal cortex and the perirhinal cortex. We propose, based on lesion and connectional data, that the two areas in which activities provide recognition awareness are the temporopolar cortex and the medial orbitofrontal cortex. Activities in the temporopolar cortex provide the recognition awareness of objects learned in the remote past (consolidated object recognition), and those in the medial orbitofrontal cortex provide the recognition awareness of objects learned in the recent past. The activation of the sparse representation for a particular sensory object in turn activates neurons in one or both of these regions of cortex, and it is the activities of these neurons that provide the awareness of recognition of the object in question. The neural circuitry involved in the activation of these representations is discussed.