Neural Correlates of Consciousness - Forward Versus Feedback Projections

Forward Versus Feedback Projections

Many actions in response to sensory inputs are rapid, transient, stereotyped, and unconscious. They could be thought of as cortical reflexes and are characterized by rapid and somewhat stereotyped responses that can take the form of rather complex automated behavior as seen, e.g., in complex partial epileptic seizures. These automated responses, sometimes called zombie behaviors, could be contrasted by a slower, all-purpose conscious mode that deals more slowly with broader, less stereotyped aspects of the sensory inputs (or a reflection of these, as in imagery) and takes time to decide on appropriate thoughts and responses. Without such a consciousness mode, a vast number of different zombie modes would be required to react to unusual events.

A feature that distinguishes humans from most animals is that we are not born with an extensive repertoire of behavioral programs that would enable us to survive on our own ("physiological prematurity"). To compensate for this, we have an unmatched ability to learn, i.e., to consciously acquire such programs by imitation or exploration. Once consciously acquired and sufficiently exercised, these programs can become automated to the extent that their execution happens beyond the realms of our awareness. Take, as an example, the incredible fine motor skills exerted in playing a Beethoven piano sonata or the sensorimotor coordination required to ride a motorcycle along a curvy mountain road. Such complex behaviors are possible only because a sufficient number of the subprograms involved can be executed with minimal or even suspended conscious control. In fact, the conscious system may actually interfere somewhat with these automated programs.

From an evolutionary standpoint it clearly makes sense to have both automated behavioral programs that can be executed rapidly in a stereotyped and automated manner, and a slightly slower system that allows time for thinking and planning more complex behavior. This latter aspect may be one of the principal functions of consciousness.

It seems possible that visual zombie modes in the cortex mainly use the dorsal stream in the parietal region. However, parietal activity can affect consciousness by producing attentional effects on the ventral stream, at least under some circumstances. The conscious mode for vision depends largely on the early visual areas (beyond V1) and especially on the ventral stream.

Seemingly complex visual processing (such as detecting animals in natural, cluttered scenes) can be accomplished by the human cortex within 130–150 ms, far too brief for eye movements and conscious perception to occur. Furthermore, reflexes such as the oculovestibular reflex take place at even more rapid time-scales. It is quite plausible that such behaviors are mediated by a purely feed-forward moving wave of spiking activity that passes from the retina through V1, into V4, IT and prefrontal cortex, until it affects motorneurons in the spinal cord that control the finger press (as in a typical laboratory experiment). The hypothesis that the basic processing of information is feedforward is supported most directly by the short times (approx. 100 ms) required for a selective response to appear in IT cells.

Conversely, conscious perception is believed to require more sustained, reverberatory neural activity, most likely via global feedback from frontal regions of neocortex back to sensory cortical areas that builds up over time until it exceeds a critical threshold. At this point, the sustained neural activity rapidly propagates to parietal, prefrontal and anterior cingulate cortical regions, thalamus, claustrum and related structures that support short-term memory, multi-modality integration, planning, speech, and other processes intimately related to consciousness. Competition prevents more than one or a very small number of percepts to be simultaneously and actively represented. This is the core hypothesis of the global workspace theory of consciousness.

In brief, while rapid but transient neural activity in the thalamo-cortical system can mediate complex behavior without conscious sensation, it is surmised that consciousness requires sustained but well-organized neural activity dependent on long-range cortico-cortical feedback.