Consciousness and Cognition 83 (2020) 102982

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Consciousness and Cognition

journal homepage: www.elsevier.com/locate/concog

A cellular and attentional network explanation of consciousness

T
Gonzalo Munévar
Lawrence Technological University, United States

ARTICLE INFO

Keywords:
Consciousness
Theories of consciousness
Attention
Attention networks
Salience
Brain activation
Neuromodulators

1. Introduction and hypothesis

This paper presents a new theory of how the brain produces individual states of consciousness (in contrast to general alertness),
based on plausible brain cellular and network mechanisms. Its purpose is to explain what Koch, Tononi (2016, 307) and others
specify as the contents of consciousness: “Being conscious means that one is having an experience – the subjective, phenomenal ‘what
it is like’ to see an image, hear a sound, think a thought or feel an emotion … we continue to be conscious when we daydream and
during those periods of sleep when we dream.” As Pitts, Lutsyshyna, and Hillyard (2018, 2) point out, this definition “remains distinct
from enabling states of arousal (e.g. coma versus wakefulness) which are necessary but not sufficient for experiencing conscious
content.”
As is well known, this issue of how the brain brings about such individual conscious states is far from resolved. In this section I
will introduce a new explanation and sketch it by means of an example. I will then discuss some important reasons for dissatisfaction
with the main theories of consciousness and compare them with my new theory. In subsequent sections, I will consider possible
objections, take up the misleading issue of the relationship between self and consciousness (using a Darwinian approach), and suggest
directions for future research.
The hypothesis of the present paper is that consciousness occurs when the salience of a brain event activates an attention network
such that the network successfully inhibits other, potentially competing, networks. What the hypothesis offers is, then, a causal
mechanism that requires the proper causal relationships between its three components for an individual conscious experience to take
place. It also requires that we extend the notion of “salience” beyond exogenous perception to compelling events in experiences well
beyond exogenous perception.
This hypothesis entails, as well, that there are many different neural pathways/forms of attention relevant to consciousness, not
merely those concerned with perception, which are typically the ones investigated experimentally. A proper theory of consciousness
will, thus, require a wider investigation of different forms of attention networks, as will be discussed in this paper.
To be clear, this hypothesis does not equate consciousness with attention, but rather attempts to explain the causal relationship
between them. To be sure, the importance of salience and attention to consciousness have long been discussed, and I will refer to

E-mail address: gmunevar@ltu.edu.

https://doi.org/10.1016/j.concog.2020.102982
Received 7 January 2020; Received in revised form 25 May 2020; Accepted 23 June 2020
1053-8100/ © 2020 Elsevier Inc. All rights reserved.
G. Munévar Consciousness and Cognition 83 (2020) 102982

some of that work below, but, as far as I know, no one has suggested their causal interaction in the sort of mechanism I suggest brings
about individual conscious experiences.
For instance, as will be argued below, claims about consciousness without attention usually involve misinterpretations of ex-
periments using top-down attention. Mainly, the possibility of other types of attention is ignored. And so is, indeed, even the pos-
sibility of different forms of top-down attention (Marchetti, 2012).

2. Elaboration of hypothesis

The following example of what might be called “survival salience” will serve to illustrate the hypothesis. Imagine that you are an
entomologist in a forest observing certain insects on the bark of trees. Your attention to this task is an example of “top-down
attention,” presumably directed in part by working memory. Suddenly, out of the corner of your eye, you detect a motion in the grass
that suggests a big snake, and this forest is known for poisonous snakes. Quite likely, your concern about a possible threat to your life
will override the attention you had been paying to the insects on the trees. Your attention is now riveted on potential signs of the
presence of the snake. This is a case of “bottom-up attention,” which generally reflects the biasing of sensory processing due to
stimulus salience (brightness, movement, size, for example), which causes features to ‘‘pop-out’’ from their surroundings to capture
attention. In this example, the survival-salience of the snake motion activates the attention network based on the inferior parietal lobe
(Corbetta, Patel, & Shulman, 2008), which then inhibits the superior parietal lobe, which was the base for your working-memory
directed observation of insects (Kelley, Serences, Giesbrecht, & Yantis, 2008). Readers will find familiar the described interaction
between these particular regions, since it is found in textbooks (e.g., Banich & Compton, 2018, 303-306).
A review of many studies on the activation and mutual inhibition of neurons and networks relevant to attention and their
connection to consciousness (e.g. Thiele & Bellgrove, 2018, Reynolds, Chelazzi, & Desimone, 1999) indicates that at the network level,
the results are best understood in terms of competing neuronal populations that often end in a “winner take all” (Carandini & Heeger,
2011). This seems an apt description of the example, for we might say that the neuronal population in the inferior parietal lobe wins
its tug-of-war over its counterpart in the superior parietal lobe. But it must be pointed out that, under certain circumstances,
otherwise competing networks can join into a larger network. Let’s modify the example to show how working memory may still retain
a share of the control of attention. Suppose that, instead of looking at insects on trees, you are looking for your small child who is lost
in the forest. It is to be expected that you would be paying extreme attention both to signs of the snake and signs of your small child.
In this case, the inferior parietal lobe and the superior parietal lobe are made to come together into a single network at the in-
traparietal sulcus (Ptak, 2012). See Fig. 1.
Attention, thus, gives causal priority to relevant over irrelevant information by activating certain circuits while inhibiting others.
The causal mechanism that produces individual conscious states is illustrated in Fig. 2. Some clarifications may round out this
account of the hypothesis while helping to prevent misunderstandings.
First, the foregoing does not suggest that only one attention network may produce conscious experience at any given time. The
idea is that within one modality (for that is where the competing networks are more likely to be found) there will be one winner, or
one winner produced by the junction of two or more networks. Attention networks in different modalities, however, are less likely to
so compete (e.g. watching a scene while listening to music). They could even reinforce each other (e.g. hearing a movie’s dialogue better
when subtitles are used). Of course, competition is not entirely ruled out, as when survival salience makes us temporarily forget about
our horrible toothache.
Second, even though for understandable pragmatic experimental reasons most research has concentrated on one type of per-
ceptual attention, there could be many attention networks. We not only perceive objects, but daydream, obsess about love affairs, or
children, are overcome with grief, fantasize about the future, and so on – all different conscious experiences that, if salient enough,
will activate attention networks not grounded on the same regions as the attention networks involved in sensory perception. For

Fig. 1. Brain regions supporting attention networks involved in the consciousness mechanism described in forest survival salience example. SPL:
Superior Parietal Lobe. IPL: Inferior Parietal Lobe. IPS: Intraparietal Sulcus. TPJ: Temporal-Parietal Junction. ANG: Angular Gyrus. Smg:
Supramarginal Gyrus. Art courtesy of Phillip McMurray. Cortex image generated with sLORETA software (Pascual-Marqui, 2002).

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Fig. 2. Cellular-Attentional Model of Consciousness. A salient event activates an attentional network, which inhibits competing networks. This
mechanism results in a conscious experience. Art courtesy of Phillip McMurray.

instance, when engaging in the concentration tasks required in mindful meditation, it would be reasonable to expect that the insula
would play a large role in the attention network, since mindfulness demands attention to the way many parts of the body feel. A
recent fMRI study shows very strong activation of the insula in mindful meditation (Murakami & Hoaki, 2019).
Third, salience is not an exclusive property of sensory stimuli. My (potential) sudden realization that the evidence indicates that
my cancer is terminal would be by far the most salient event and would command my attention, thus becoming conscious. As
Haladjian and Montemayor (2016) point out, this is an introspective form of attention related to self-knowledge and detection of
inner sensations that is often overlooked by researchers, in spite of its importance to consciousness. This issue is also related to
“source monitoring” (Mitchell & Johnson, 2009). At all levels, salience depends on a context that may include goals, episodic
memories, reward history (Watson et al., 2019), intuitive cognition and many other brain functions and regions. Within that context,
the most salient brain phenomena will command attention, and be consciously experienced. The resulting attention-consciousness
network will include those brain functions and regions that contribute to the creation of the context.
In this view, we have different attentional networks acting side by side, some on unconscious stimuli, or phenomena, but inhibited
by the highly activated one. One of those other networks may suddenly become highly activated and inhibit the one that had gained
center stage previously. Top-down attention is no exception in this regard. In the previous example of combined top-down and
bottom-up attention, any sound that might be from your child may immediately concentrate all your attention on any follow-up
sounds, confining any further concerns about the snake to the unconscious.

3. Neuronal processing

At the cellular level, neuromodulators lead to significant changes in cognitive function, including consciousness, for they affect
firing rates of excitatory and inhibitory neurons (Mitchell, Sundberg, & Reynolds, 2007). Lowering variability of firing rates reduces
noise and improves information, which is crucial to consciousness. The relevant attention network areas of the brain depend on
neuromodulators released by brain-stem neurons (Bichot, Heard, DeGennaro, & Desimone, 2015) (Marder, 2012). In higher cognitive
functions some crucial neurotransmitters are: Acetylcholine (Ach), dopamine (DA), noradrenaline (NA), and serotonin (5H-T) (Thiele
& Bellgrove, 2018). The newly activated attention network sends feedback to the appropriate brain-stem area, which in turn increases
the network’s activation.
Judith Ford’s view on hallucinations in schizophrenia (Whitford, Ford, Mathalon, Kubicki, & Shenton, 2012) gives an interesting
illustration of the role of feedback between attention networks and brain-stem areas where neuromodulators are produced. According
to Ford, myelin abnormalities in the white tracks from the frontal lobe to other regions of the cortex lead to a loss of neural syn-
chronization. This means that signals that are generated by the brain itself (say, in an imaginary conversation) do not arrive at the
appropriate areas of the brain at the appropriate time. This lack of synchronization in turn, then, leads to prediction errors in self-
generated actions and thoughts, which in turn lead to confusion about the origin of thoughts, and thus are experienced as external
(hallucinations). That is, when a brain-generated fragment of conversation arrives late so its self-generated origin is no longer
identified, it seems to be a novel perception and is thereby salient. The response to this salience is increased dopamine modulation by
brain-stem neurons, which highlights the hallucination’s salience even more. Additional myelin abnormalities in the corpus callosum
and other brain regions are correlated with negative symptoms in schizophrenia (Esaki et al., 2020).
Neuronal processing is affected by both bottom-up and top-down influences, as well as other kinds of attention in many areas of
the brain. Incidentally, top-down processing is often taken to reflect the voluntary (and presumably conscious) guidance of attention
to locations, features, or objects in the environment. This is somewhat misleading, however, for the processing may be largely un-
conscious. It may also be generated by unconscious memories or drives from cortical areas and then influence perception and the

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generation of new memories.

4. Best regarded theories of consciousness

I will briefly discuss the two most prominent theories of how the brain produces conscious states (this discussion will be further
summarized in Table 1) and contrast them with the theory proposed in this paper. The first is the Global Workspace Theory of
Consciousness (Baars, 2005), or Global Neuronal Workspace Theory (Dehaene, 2015). In this theory, consciousness would be a way of
coordinating and integrating functions of the brain such as sensory perception, long-term memory, attention, setting goals, planning
actions, determining value, and many others. This integration into a whole is achieved by having, say, a visual stimulus enter a global
workspace in which its contents become available to those other brain functions. By doing so, the stimulus moves from unconscious
visual processing to conscious experience. Essentially, according to Dehaene, consciousness is a process of brain-wide information
sharing. Very importantly, the global workspace is to be found mainly in the higher association areas, the frontal-parietal areas, with
key roles played by the frontal lobe.
As an example, Dehaene discusses the conscious recognition of a face. The right fusiform gyrus receives inputs from several brain
areas and integrates them, but all this activity is unconscious. Consciousness occurs because “long distance axons allow the visual
information to be dispatched to virtually any corner of the brain” and those other regions themselves participate in “a broader
network of workspace areas” underpinned by strong anatomical connections. Ironically, a recent finding of his research program
(Demertzi et al., 2019) is that (1) whereas in the loss of consciousness, coordinated brain activity becomes largely restricted to
interareal coordination that follows the anatomical connections, (2) the return to consciousness provides for a complex configuration
of interareal coordination that goes beyond the interareal anatomical connections and permits interaction at long distances. This
result seems to undermine the emphasis on strong anatomical connections that underpins the network of workspace areas.
Leaving that aside, we would still want to know from Dehane why at a particular time Ann’s face is perceived consciously but no
other faces also integrated by the fusiform face area are. Dehaene claims, correctly, that the brain has evolved to determine relevance.
Nonetheless, how is this face more relevant than others? This problem is solved by my model by its reliance on its notion of salience,
which gives us an advantage when explaining why we consciously perceive certain faces but not others (threatening, attractive,
familiar, etc.). If Ann, for example, was a friend in college, then memories tinged by positive emotions might make it much more
salient than other faces, thus activating an attention network and so on.
Moreover, the global neuronal workspace theory is contradicted by fMRI experiments whose results presumably show that the
“frontal areas are associated with active report and introspection” rather than with the actual perception (Frässle, Sommer, Jansen,
Naber, & Einhäuser, 2014). Tononi, Boly, Massimini, and Koch (2016) as well as Tononi and Koch (2016) also make an extensive case
to similar effect. Pitts et al. (2018) find certainly suggestive fMRI studies such as Farooqui and Manly (2017) who “reported deac-
tivations of frontal cortical areas for non-target stimuli that were clearly seen.” Nevertheless, Pitts et al. and Safavi, Kapoor,
Logothetis, and Panagiotaropoulos (2014) make an appeal for additional and more refined experiments. See also the comments by
Naber and Brascamp (2015). Of greater concern, however, is the finding that people and animals without a frontal lobe, and indeed
much of the rest of the cortex, are able to have some conscious experiences (Aleman & Merker, 2014).
Since it is clear that we can have specific instances of conscious experience in which the frontal lobe plays no role, we then need to
look elsewhere for a theory of how the brain creates individual conscious experiences. My own theory is designed to fulfill that need.
The activation of an attention network by salience would work even in cases where the frontal lobe is absent. The thalamus and other
subcortical areas should provide enough attention networks (even if somewhat diminished by comparison with many of those in a
normal human brain) as well as the ability to determine many instances of salience.
Nevertheless, some credit may be given to the neuronal global workspace theory. It does not provide the mechanism of conscious
states, but it may account for the richness of the contents of many conscious states, since those contents may be influenced by
learning, memories, and so on to be found in many areas of the brain, and particularly in the frontal cortex. After all, a full, rich top-
down attention is very important to the mental life of human beings and other animals.
The other best regarded theory of consciousness is Giulio Tononi’s ingenious Information Integration Theory (Tononi et al., 2016),
also championed by Christof Koch (2012). The basic idea is that consciousness occurs when processes in the brain achieve a good
ratio of differentiation to integration, for such a ratio gives us the most information. At first this seems to fit well with perceptual
processes, at least visual perception. The brain receives information from the retina and then parcels out information about the color
patterns of an object, its motion, its inclination, its shape, etc. If all these differentiated properties came to consciousness separately at
the same time, we would have a very confusing experience. Instead the brain integrates these different visual properties into a
perception of the object as a whole. The brain indeed has modules that very successfully integrate differentiated properties, e.g. the
perception of faces (from eyes, noses, lips, etc.) in the fusiform gyrus – the fusiform face area, which Tononi says comes close to
qualifying as a content-specific neural correlate of consciousness area (NCC). As Koch asserts in agreement, achieving the right ratio
of differentiation and integration (Φ) gives the most information. Too much differentiation without integration, as happens in an-
esthesia, eliminates consciousness. Too much integration without differentiation, as it happens in grand mal epileptic seizures also
eliminates consciousness. On the other hand, when Φ is high, the brain is given holistic information that brings together and goes
beyond the total amount presented by the sum of the individual constituents of the perception. The greater Φ is, the higher the degree
of consciousness.
Koch also argues that a merely descriptive theory of consciousness, such as the Global Neuronal Workspace Theory, is deficient. A
truly satisfactory theory should be prescriptive. And by “prescriptive” Koch means that “it must give necessary and sufficient con-
ditions for consciousness to occur” (p. 121). Now, is the Information Integration Theory itself prescriptive in this sense?

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Let us return to the perception of faces. Most of the many “face integrations” in the fusiform face area of the brain are never
perceived consciously. For example, in a busy city street you see lots of faces, but you perceive consciously only a few (e.g. those of
your friend walking towards you and of the man shouting that the world will end tomorrow). Why those and not others? Apparently,
the face with the highest Φ, to put it crudely, is the one chosen to be broadcast widely in the brain. Unfortunately, it would seem that
the ratio of differentiation to integration in many of those faces would be the same. Thus, at best, achieving the ideal ratio of
information and integration would be a necessary condition for conscious perception, but not a sufficient condition.
It is not a necessary condition either. Presumably, according to Koch, the more highly an experience possesses the right com-
bination of information and integration, the more likely it is to be conscious. Such an experience is thus likely to be highly structured
(again, as in faces). But there are many types of intense conscious experiences, acute experiences, in which it is difficult to detect what
possible structure is revealed by the conscious experience. A clear example is pain, which is no less conscious than the perception of a
face. If anything, particularly with chronic pain, we wish to be far less conscious of it. The “right” combination of differentiation and
integration seems to be irrelevant to whether we experience pain or not. Likewise, no such holistic structure imposed on differ-
entiated aspects seems to be revealed when we are conscious of an intense emotion, let alone of vague feelings. Like attention,
consciousness comes in different forms (see Haladjian & Montemayor, 2015, 2016; Montemayor & Haladjian, 2015). It is not in the
least obvious that we consciously perceive the snake motion over the search for the bugs on the trees because perceiving the snake
motion offers a higher Φ. Not at all. Its salience is why we perceive it. And the same answer applies to the previous discussion of what
counts as a sufficient condition: We consciously perceive those faces that are most salient to us.
Koch counters that even the perception of the darkness in a pitch-black room is highly informative because it implies that you are
not looking at well-lit rooms, mountains and a trillion other things. His reply is hard to square with Tononi’s claim that consciousness
requires a single integrated process with a great many highly differentiated states. In addition to this problem, Koch also makes the
computation of Φ absurdly complicated, for according to him the numbers involved may be of the order of ten followed by hundreds
of zeroes, as he acknowledges in the case of C. elegans, and thus likely thousands, if not millions of zeroes in the case of human brains.
It seems unreasonable to suppose that the brain can choose which event is to be perceived consciously when the difference between,
say, two faces can be insignificant, given the extremely large numbers involved. As for his example of the dark room, what is not
experienced becomes truly informative only when the number of relevant states is quite limited. To illustrate his point, Koch tells us a
story in which Sherlock Holmes deduces that the dog in a house knew the intruder, for it did not bark. That is, the absence of certain
information was on the whole quite informative. But this sort of example only works when the number of choices is quite limited,
unlike the case of the perception of the dark room.

Table 1
Comparison of Standard and New Theory of Consciousness.
Features Addressed Global Neuronal Workspace Information-Integration Activated Attention

Frontal lobe required for consciousness Yes No No

Explains role of relevance in consciousness No No Yes
Salience central No No Yes
Gives necessary and sufficient conditions for consciousness No No Yes
Connection between consciousness and adaptation Yes Yes Yes (more precisely)
Multiplicity of attention networks No No Yes

5. Consciousness and evolution

Many potential sensory perceptions are helped or hindered long before the cortical circuits favored by the two theories of
consciousness criticized above even come into play. The brain constantly abstracts from the deluge of stimuli that floods our senses.
Only a very small percentage of the signals coming from the retina go beyond the lateral geniculate nucleus to the cortex. Much of the
processing (i.e. filtering) of potential perceptual consciousness takes places in subcortical areas. Francis Crick’s suggestion of the
thalamic reticular nucleus (1984) has been recently vindicated by M. Halassa and his team (Wimmer et al., 2015), which has
individuated a complicated neural circuit that also includes the basal ganglia. Very importantly, most of the activity of this circuit
concerning perception is the massive inhibition of incoming signals. Furthermore, the signals that are favored involve movement,
size, and bright coloring, as well as other properties that would be relevant to being able to survive or adapt to the environment (e.g.
for inhibition of the detection of large objects in favor of the perception of the motion of small objects see Tadin et al., 2019). Insofar
as this process of inhibition is part and parcel of sensory perception, it seems that evolution has pretty much hardwired us to favor
salience in conscious perception. The focus of attention does diminish slightly four times per second (Fiebelkorn & Kastner, 2019),
which increases the probability that a new and more salient phenomenon may replace the current perception.
In this manner the salience hypothesis captures more precisely the connection between consciousness and adaptation suggested
by both the Global Neuronal Workspace Theory and the Information-Integration Theory, in that, by making the conscious phe-
nomenon widely available to many parts of the brain, greater sensory and other information may effectively guide action.
Of course, there is much more to consciousness than sensory perception. Moreover, phenomena processed in non-perceptual
systems may be salient enough, as far as the brain is concerned, to override salient sensory signals, as we saw in the variation of the

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snake motion example. Learning and personal history also play important roles in wiring the brain for all sorts of conscious ex-
periences, thus leading to the preliminary inhibition or suppression of some signals in favor of others, initially facilitating the
recognition of salience and the activation of an attention network that will inhibit the still extant potential competing attention
networks. Finally, once consciousness appeared, it began to serve many functions in brains of increasing complexity. For a further
exploration on the evolution of attention and consciousness, see Haladjian and Montemayor (2015).
A proper account of the evolutionary value of consciousness, however, should require a consideration of decades of research on
implicit learning, which have identified a variety of brain mechanisms for the non-conscious processing information about the
environment, mechanisms that constitute the neurocognitive foundations of intuition. As Reber, Beeman, and Paller (2013) point out,
deliberate deduction and implicit intuition “depend on separate types of memory supported by distinct brain networks” (p. 474). This
intuition is not guided by awareness of the processes or events that bring it about, and indeed may be inhibited by conscious
deliberation, as anyone who plays a team sport can confirm. When scoring a goal with a scissors kick (flying horizontal on my back
and kicking the ball over my head) I would perform the acrobatic play without any conscious decision to do so. This shows, in-
cidentally, that such unconscious intuitive behavior is not just purely automatic, born from lots of practice. I never practiced scissor
kicks nor met anyone who did. It is a moment of, largely, proprioreceptor intuition. Clearly, there is much adaptive value to such
intuitive skills, as they must have come in handy in fights with predators and in hunting, as well as in many other aspects of human
interactions with their environments. Cleeremans, Allakhverdov, and Kuvaldina (2019) provide important additional discussions of
implicit learning; see also Alamia et al., 2016).
Incidentally, there is a connection with the adaptive value of consciousness as well. Reber at al. narrate the case of a lieutenant
firefighter who ordered his men out of a building on fire because he “sensed” something wrong. Indeed, the floor was about to
collapse and have the raging fire below kill him and his men. His was an intuitive reaction, without awareness, let alone conscious
understanding at the moment of the clues that, given his vast experience, made him sense mortal danger. Still, the sensing itself was a
conscious experience: A conscious experience of extreme salience that led to survival.

6. Some potential objections

In some experiments (Matthews, Schroder, Kaunitz, van Boxtel, & Tsuchiya, 2018), the subject pays attention to a section of the
screen, but nevertheless becomes aware of salient images, e.g. faces, in unattended sections. This, Matthews concludes, is a case of
consciousness without attention. But we need not jump to that conclusion. All that strictly follows is that you can have consciousness
without voluntary Top-Down Attention. But we may nevertheless have attention in those cases: Bottom-Up Attention that leads to the
consciousness of salient phenomena, for example. I do not mean to suggest, however, that Bottom-Up Attention is the only me-
chanism that can overrule Top-Down Attention. The key factor is salience, and salience can come from a great many regions of the
brain. As Giorgio Marchetti (2012) argues, the claim that consciousness can exist without some form of attention ignores the many
varieties of both attention and consciousness. Since faces, say, attract exogenous attention, such dual-task experiments (Reddy,
Wilken, & Koch, 2004) could be interpreted as a combination of exogenous attention and some form of diffused top-down con-
sciousness. Or as Kouider, de Gardelle, Sackur, and Dupoux (2010, p. 304) observe, “the possibility of consciousness without at-
tention is usually based on a restrictive definition that does not take into account the possibility of residual attention at lower (i.e.
sensory, non-conceptual) levels of processing.”
Even in top-down attention, stimuli can be processed preconsciously with a minimal level of attention, as can be seen in ex-
periments on implicit learning and Stroop effect (Hartman, Knopman, & Nissen, 1987), dichotic listening and shadowing tasks
(MacKay, 1973; Treisman, 1964), and visual masking (Marcel, 1983). And in some experiments (Logan, 1995; Neuman, 1984),
subjects are known to pay marginal attention to stimuli they have been instructed to ignore. This marginal attention is achieved by
widely distributing the focus of attention, as Marchetti (2012) explains, and experiments by McCormick (1997) exemplify.
At any rate, an integral aspect of the salience hypothesis of consciousness is that, during a particular instance of consciousness,
there might have several potentially competing attention networks. Generally, only one will be activated by the salient brain event,
although as we have seen, the connections created by the activation may lead to the combination of two attention networks (perhaps
more) into one, as explained in the example of the father looking for his son in the forest.
Tallon-Baudry (2012) argues that a decision can be made at the sensory level, before attention mechanisms can shape or enhance
the signal, and thus the mechanisms of consciousness and attention may be distinct, even if complementary at times. This is not incompatible
with the salience hypothesis, which only claims that, at some point, a signal may activate a network of attention that then inhibits
potential competitors. Moreover, it should not be surprising that salience is often determined at the sensory level.
It is important to consider as well how, during sleep, particularly dreaming, or under general anesthesia, consciousness might be
affected, for the brain reduces the neuromodulator support normally given to attention during alertness. During REM sleep, for
example, regions such as the inferior parietal cortex, the precuneus, the orbitofrontal cortex (OFC), and the dorsolateral prefrontal
cortex (dlPFC) are deactivated, relative to the waking state, and such deactivation may lead to “disorientation, illogical thinking,
reduced cognitive control, and impaired working memory” (Perogamvros, Dang-Vu, Desseilles, & Schwartz, 2013). At the same time
the amygdala interacts with the brain stem and activates its connections with the thalamus, the anterior cingulate cortex (ACC), the
medial prefrontal cortex, the septal nuclei and the hippocampus. This combination of deactivation of cognitive areas and activation of
areas involved in strong emotions and the affective value of memories would seem to account for the half-haphazard story telling that
take place in dreaming. Less-than-fully activated attention networks may not be able to inhibit all potential competitors for long,
while the salience of this or that image may suddenly be enhanced by varying emotional influences, such as fear and anxiety, which
would result in frequent shifts in the contents of conscious experience, as is characteristic of dreaming.

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It was recently discovered in Fan Wang’s lab at Duke that general anesthesia activates firing neurons in the supraoptic nucleus of
the hypothalamus (Jiang-Xie et al., 2019). This small nucleus has long projections to the pituitary gland that lead to the release of
vasopressin and other hormones into the bloodstream. This connection between the nervous and endocrine systems creates a slow-
wave sleep, a deep slumber associated with unconsciousness. Exciting the neurons in that nucleus makes mice fall sleep. Deactivating
them makes the mice unable to sleep. Thus, it is not surprising to see some similarities between the effects of many anesthetics and
sleep. This finding fulfills many of the criteria anticipated by cognitive unbinding as an explanation of unconsciousness in anesthesia
(Mashour, 2013). Some neural activity can persist despite anesthesia, but there will be temporal fragmentation of that activity. This
isolation of neural activity will impair the effective communication between brain regions that, as we have seen, attention provides in
bringing about consciousness. In the case of dreaming, the fragmented attention may lead to a seemingly illogical narrative. In the
case of anesthesia, as Mashour points out, primary sensory networks processing may persist, but it cannot be taken in the context of
other cognitive activity that employs cortical-cortical or cortical-subcortical connectivity. Thus, the anesthetized patient is unable to
perceive consciously. Nonetheless, the similarities are strong enough that it should not be surprising that some patients (26% on
average) report dreaming while under anesthesia (Mashour, 2011). A new study by Wang, Eguchi, Takayuki Yamashita, and
Tomoyuki (2020) indicates that volatile anesthetics selectively inhibit high-frequency neurotransmission, which will attenuate in-
tegral neuronal functions, with minimal inhibition of low-frequency life-supporting basal neurotransmission. This result points to
possible avenues for experimentation not only for the question of dreaming under anesthesia but also for the consciousness me-
chanism proposed by my model.

7. Consciousness of self

The Cartesian “ghost in the machine” has led some to the notion that to be conscious is, at least in part, to be conscious of
ourselves, as well as to talk about the centralizing role of the “self” – and of the unity of the self. Even distinguished researchers like
Crick and Koch (2003) proposed that, when we are immersed in certain intense experiences (in which we do not think about
ourselves), nonetheless the frontal lobe still monitors such experiences and performs the centralizing role of informing the brain that
such experiences belong to it. An fMRI study by Goldberg, Harel, and Malach (2006), however, showed that demanding perceptual
tasks actually lower the activation of the frontal regions hypothesized by Crick and Koch as having such self-monitoring role.
Moreover, according to Gillihan and Farah’s (2005) meta-analysis of neuroscientific studies of self-knowledge, the data do not
“implicate common brain areas across different aspects of the self. This is not surprising because there is generally little clustering
even within specific aspects of the self” (p. 94). And Llinas (2001) concludes that, since there is no centralizing brain structure that
corresponds to a Cartesian self, the self is just an internal perception, an illusion, or as some would say, a construction. It is common
to believe that episodic, autobiographical, memories give us our sense of self, since they are created by the very actions that sculpt
our personality. But Stanley Klein (2004) has demonstrated that patients who cannot form episodic memories because they have lost
their hippocampi still have a good sense of their personality, even when the personality changes.
Evolutionary biology, however, offers an alternative non-Cartesian account that explains the apparently perplexing results of the
neuroscientific research on the self (Munevar, 2011, 2014), and that is compatible with the previous remarks in this paper about
salience and attention. To survive in their environments, organisms need, among other things, the ability to tell self from others. In a
great many animals, that need is met by the function of the immune and nervous systems. In higher animals, the brain plays an
important role, for it has evolved to coordinate external information with internal information about the states of the organism. In
doing so it is guided by its genetic inheritance and its previous experience. This coordination is of necessity very complex, since we
have very complex interactions with the environment, and thus the brain should perform a large variety of different tasks in telling
self from other. As explained in the references, the brain has evolved to perform many of the functions ascribed to the self. The self is
as the brain does.
Therefore, instead of a centralized function, of a unified self, we should thus expect a distributive function of the self, which a
highly distributive brain, as the human brain is, would thus be best able to perform. The task of recognizing photos of ourselves is
quite different from the task of distinguishing ourselves from others on the basis of traits that we ascribe to ourselves and to those
others, and therefore it should be expected that different regions of the brain should be activated.
To take up this last task as an illustration, when we ascribe traits to ourselves and close and distant others, the results are much in
line with a Darwinian approach. For example, in an fMRI study my colleagues and I found that the areas activated in distinguishing
between self and a distant other (a celebrity) were the same as those activated when distinguishing between best friend and distant
other – particularly the ACC, which is involved in Theory of Mind – although the level of activation was somewhat smaller for the best
friend case (Munevar & Cole, 2014). Surely, since we are social animals, we should be expected to identify with those who are closest
to us, although not as much as with ourselves. Self and best friend are nonetheless distinguished in several areas, but most im-
portantly in Broadman Area 31, in which the brain also makes a distinction between geocentric orientation (to my right or left, etc.)
and allocentric orientation, i.e. where it “objectifies” its physical position with respect to the world (e.g. by using landmarks).
It should be noted, incidentally, that most of these neural activities of the “self” are unconscious, which should come as no
surprise since most of the brain’s activities are unconscious. The brain generally builds up its connections with priority to what is
important to the organism. Particularly salient items will become conscious, as we have seen, by their activation of an attention
network that involves the most relevant connections while inhibiting potential competing networks.
Awareness of self is, then, not a requirement for consciousness.

7
G. Munévar Consciousness and Cognition 83 (2020) 102982

8. Directions for future research

The hypothesis of this paper was inspired by a wealth of neuroscientific studies about consciousness and attention networks, many
of which have found their way to a variety of excellent reviews, such as Thiele and Bellgrove (2018), as well as in related fields, e.g.,
neuropsychology: Berlucchi and Marzi (2019). As it is often the case in neuroscience, future research should focus on clarifying in
detail the key relationships involved in the model under examination. In order to achieve this aim, there is a great need for a broader
study of attention networks and of conscious experience that extends beyond the study of perception. That restricted standard
approach has born much fruit, but there is much more to the diversity and richness of our consciousness and the possible attention
networks connected with it. It may be particularly fruitful, in the next few years, to combine qEEG and neuroimaging to study
instances of consciousness beyond the standard perceptual cases. The reason for this combination is that we need not only good
spatial resolution, as provided by fMRI, but also excellent temporal resolution so as to be better able to determine what may cause
what in a particular instance of consciousness.
With that aim in mind, I hope to begin the experimental phase of this research by examining two instances of consciousness that
go beyond those already studied. One of them is synesthesia. Many synesthetes experience some letters as being colored, say the “F”
and all words that begin with it as being red. But this phenomenon is not a mere question of perception, for a synesthete will perceive
an F as red only after identifying it as an F to begin with. The second would be the study of volitional consciousness. I intend to redo,
in the combined qEEG-neuroimaging environment, Libet’s famous (1985) experiment in which he measured the timing of a subject’s
conscious decision to flex a hand, the timing of the readiness potential in the brain to initiate the action, and the flexing of the hand
itself. Much to Libet’s shock, the readiness potential happened first, followed on the average 350 ms later by the conscious decision,
which was followed 200 ms later by the flexing of the hand. Although I have been highly critical of the “free-will controversy” that
ensued from Libet’s experiment (see Munevar, 2011), his time determinations can prove useful. My proposal is to capture the neural
correlate of volitional consciousness by looking at what happens in the brain around 200 ms before the flexing of the hand. In this and
the synesthesia study, the aim will be to determine the activating salient event and the relevant attention network, as well as to
measure the inhibition of specific neuronal populations.
I do hope that these proposed experiments will be supplemented by many other researchers’ attempts to expand our investigation
of consciousness and attention. At the very least, they would add to the continuing work on perception, and contribute to a better
understanding of the rich mental life the brain makes possible.

Acknowledgments

I am grateful to Eugene Brooks, Henry Bucthel, Adnan Jaber, Douglas MacDonald, George Mashour, and David Paulsen for their
very helpful comments on earlier drafts of this paper. I also wish to thank Phillip McMurray for his artwork in Fig. 1 and Fig. 2, and
Mathew Cole for his help with Table 1.

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