section_path: Scientific study/Neural correlates
section_name: Neural correlates
paragraph_text: Neural correlates
  
  A major part of the scientific literature on consciousness consists of studies that examine the relationship between the experiences reported by subjects and the activity that simultaneously takes place in their brains—that is, studies of the neural correlates of consciousness. The hope is to find that activity in a particular part of the brain, or a particular pattern of global brain activity, which will be strongly predictive of conscious awareness. Several brain imaging techniques, such as EEG and fMRI, have been used for physical measures of brain activity in these studies.
  
  Another idea that has drawn attention for several decades is that consciousness is associated with high-frequency (gamma band) oscillations in brain activity. This idea arose from proposals in the 1980s, by Christof von der Malsburg and Wolf Singer, that gamma oscillations could solve the so-called binding problem, by linking information represented in different parts of the brain into a unified experience. Rodolfo Llinás, for example, proposed that consciousness results from recurrent thalamo-cortical resonance where the specific thalamocortical systems (content) and the non-specific (centromedial thalamus) thalamocortical systems (context) interact in the gamma band frequency via synchronous oscillations.
  
  A number of studies have shown that activity in primary sensory areas of the brain is not sufficient to produce consciousness: it is possible for subjects to report a lack of awareness even when areas such as the primary visual cortex (V1) show clear electrical responses to a stimulus. Higher brain areas are seen as more promising, especially the prefrontal cortex, which is involved in a range of higher cognitive functions collectively known as executive functions. There is substantial evidence that a "top-down" flow of neural activity (i.e., activity propagating from the frontal cortex to sensory areas) is more predictive of conscious awareness than a "bottom-up" flow of activity. The prefrontal cortex is not the only candidate area, however: studies by Nikos Logothetis and his colleagues have shown, for example, that visually responsive neurons in parts of the temporal lobe reflect the visual perception in the situation when conflicting visual images are presented to different eyes (i.e., bistable percepts during binocular rivalry). Furthermore, top-down feedback from higher to lower visual brain areas may be weaker or absent in the peripheral visual field, as suggested by some experimental data and theoretical arguments; nevertheless humans can perceive visual inputs in the peripheral visual field arising from bottom-up V1 neural activities. Meanwhile, bottom-up V1 activities for the central visual fields can be vetoed, and thus made invisible to perception, by the top-down feedback, when these bottom-up signals are inconsistent with the brain's internal model of the visual world.
  
  Modulation of neural responses may correlate with phenomenal experiences. In contrast to the raw electrical responses that do not correlate with consciousness, the modulation of these responses by other stimuli correlates surprisingly well with an important aspect of consciousness: namely with the phenomenal experience of stimulus intensity (brightness, contrast). In the research group of Danko Nikolić it has been shown that some of the changes in the subjectively perceived brightness correlated with the modulation of firing rates while others correlated with the modulation of neural synchrony. An fMRI investigation suggested that these findings were strictly limited to the primary visual areas. This indicates that, in the primary visual areas, changes in firing rates and synchrony can be considered as neural correlates of qualia—at least for some type of qualia.
  
  In 2013, the perturbational complexity index (PCI) was proposed, a measure of the algorithmic complexity of the electrophysiological response of the cortex to transcranial magnetic stimulation. This measure was shown to be higher in individuals that are awake, in REM sleep or in a locked-in state than in those who are in deep sleep or in a vegetative state, making it potentially useful as a quantitative assessment of consciousness states.
  
  Assuming that not only humans but even some non-mammalian species are conscious, a number of evolutionary approaches to the problem of neural correlates of consciousness open up. For example, assuming that birds are conscious—a common assumption among neuroscientists and ethologists due to the extensive cognitive repertoire of birds—there are comparative neuroanatomical ways to validate some of the principal, currently competing, mammalian consciousness–brain theories. The rationale for such a comparative study is that the avian brain deviates structurally from the mammalian brain. So how similar are they? What homologs can be identified? The general conclusion from the study by Butler, et al., is that some of the major theories for the mammalian brain also appear to be valid for the avian brain. The structures assumed to be critical for consciousness in mammalian brains have homologous counterparts in avian brains. Thus the main portions of the theories of Crick and Koch, Edelman and Tononi, and Cotterill seem to be compatible with the assumption that birds are conscious. Edelman also differentiates between what he calls primary consciousness (which is a trait shared by humans and non-human animals) and higher-order consciousness as it appears in humans alone along with human language capacity. Certain aspects of the three theories, however, seem less easy to apply to the hypothesis of avian consciousness. For instance, the suggestion by Crick and Koch that layer 5 neurons of the mammalian brain have a special role, seems difficult to apply to the avian brain, since the avian homologs have a different morphology. Likewise, the theory of Eccles seems incompatible, since a structural homolog/analogue to the dendron has not been found in avian brains. The assumption of an avian consciousness also brings the reptilian brain into focus. The reason is the structural continuity between avian and reptilian brains, meaning that the phylogenetic origin of consciousness may be earlier than suggested by many leading neuroscientists.
  
  Joaquin Fuster of UCLA has advocated the position of the importance of the prefrontal cortex in humans, along with the areas of Wernicke and Broca, as being of particular importance to the development of human language capacities neuro-anatomically necessary for the emergence of higher-order consciousness in humans.
  
  A study in 2016 looked at lesions in specific areas of the brainstem that were associated with coma and vegetative states. A small region of the rostral dorsolateral pontine tegmentum in the brainstem was suggested to drive consciousness through functional connectivity with two cortical regions, the left ventral anterior insular cortex, and the pregenual anterior cingulate cortex. These three regions may work together as a triad to maintain consciousness.