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Lepauvre & Melloni - The Search For The Neural Correlate of Consciousness. Progress and Challenges (Philosophy and The Mind Sciences, 2021)

Twenty years ago, Thomas Metzinger published the book The Neural Correlates of Consciousness amassing the state of knowledge in the field of consciousness studies at the time from philosophical and empirical perspectives. On the occasion of the 20th anniversary of this impactful publication, we review the progress the field has made since then and the important methodological challenges it faces.

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Lepauvre & Melloni - The Search For The Neural Correlate of Consciousness. Progress and Challenges (Philosophy and The Mind Sciences, 2021)

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The search for the neural correlate of

consciousness: Progress and challenges

Alex Lepauvrea  ([email protected])
Lucia Mellonib  ([email protected])

Abstract
Twenty years ago, Thomas Metzinger published the book The Neural Correlates of Consciousness
amassing the state of knowledge in the field of consciousness studies at the time from philosophical
and empirical perspectives. On the occasion of the 20th anniversary of this impactful publication,
we review the progress the field has made since then and the important methodological challenges
it faces. A tremendous number of empirical studies have been conducted, which has led to the
identification of many candidate neural correlates of consciousness. Yet, this tremendous amount
of work has not unraveled a consensual account of consciousness as of now. Many questions,
some already raised twenty years ago, remain unanswered, and an enormous proliferation of
theories sharply contrasts with the scarcity of compelling data and methodological challenges. The
contrastive method, the foundational method used to study the neural correlate of consciousness
(NCC), has also been called into question. And while awareness in the community of its
shortcomings is widespread, few concrete attempts have been made to go beyond it and/or to
revise existing theories. We propose several methodological shifts that we believe may help to
advance the quest of the NCC program, while remaining uncommitted to any specific theory:
(1) the currently prevalent “contrastive method” should lose its monopoly in favor of methods
that attempt to explain the phenomenology of experience; (2) experimental data should be shared
in centralized, multi-methods databases, transcending the limitations of individual experiments
by granting granularity and power to generalize findings and “distill” the NCC proper; (3) the
explanatory power of theories should be directly pitted against each other to overcome the
non-productive fractioning of the field into insular camps seeking confirmatory evidence for their
theories. We predict these innovations might enable the field to progress towards the goal of
explaining consciousness.
Keywords
Adversarial collaboration ∙ Big data ∙ Consciousness ∙ Contrastive method ∙ Neural correlate of
consciousness ∙ Phenomenology

This article is part of a special issue on “The Neural Correlates of Consciousness,”

edited by Sascha Benjamin Fink and Ying-Tung Lin.

a
Department of Neuroscience, Max Planck Institute for Empirical Aesthetics, Frankfurt am Main
b
Department of Neurology, NYU Grossmann School of Medicine

Lepauvre A., & Melloni L. (2021). The search for the neural correlate of consciousness: Progress
and challenges. Philosophy and the Mind Sciences, 2, 4. https://doi.org/10.33735/phimisci.2021.87
©The author(s). https://philosophymindscience.org ISSN: 2699-0369
Alex Lepauvre and Lucia Melloni 2

1 Introduction
Men ought to know that from the brain, and from the brain only, arise
our pleasures, joys, laughter, and jests, as well as our sorrows, pains,
griefs, and tears. Through it, in particular, we think, see, hear, and
distinguish the ugly from the beautiful, the bad from the good, the
pleasant from the unpleasant.

—Hippocrates of Cos, The Sacred Disease, ~400BC

(translated by W. H. S. Jones, 1959, p. 175)

Since the time of Hippocrates, mankind has attempted to understand what con-
sciousness is, what it does, and how physical systems such as the brain, but not
others such as the heart, can instantiate it. David Chalmers famously referred to
the latter aspect as “The Hard Problem” (Chalmers, 1995): why does matter, such as
the brain, give rise to perceptions and emotions that have subjective, phenomeno-
logical qualities?
As underlined by Metzinger (2000), the difficulty in studying consciousness
empirically lies in the inherent subjectivity of the phenomenon. This is indeed
the main reason why consciousness for a long time remained outside the scope
of modern sciences, whose primary goal is to understand objective phenomena.
Notwithstanding revolutionary discoveries made over the past century by study-
ing cerebral lesions and using electrical stimulation to probe brain function, as in
the pioneering work of Weiskrantz, Penfield, Libet, and Eccles among others (Ec-
cles, 1974; Libet et al., 1967; Penfield, 1958; Weiskrantz, 1980), it was Crick and
Koch’s seminal paper (1990) that provided a scientific framework for the study
of the neural correlates of consciousness that set the field in motion. Crick and
Koch proposed putting aside philosophical and metaphysical considerations and
to focus on the “easy problem of consciousness” (Crick & Koch, 1998), i.e., to study
the neural activity that correlates with consciousness while taking subjectivity as
a dependent variable (Dehaene, 2014). The focus of the science of consciousness
would be narrower, yet more tractable, clearly defined, and theoretically neutral,
i.e., in identifying the neural correlates of consciousness (NCCs): the minimal set of
neural events jointly sufficient for a specific conscious experience (given the appro-
priate enabling conditions, Chalmers, 2000). By quantifying which neural activity
best correlates with conscious experience, one could begin to narrow down the
mechanisms involved in consciousness, and to eventually formulate theories to
answer the hard questions of consciousness (Chalmers, 1995).
This bottom-up strategy has dominated the field for the past thirty years, fur-
ther propelled by the development of ever more refined methods for investigating
the human brain non-invasively, e.g., high-field imaging (7T) with laminar and
columnar resolution, and the rapid development of machine-learning algorithms
that have enabled researchers to track the contents of perception. Thirty years af-
ter Crick and Koch’s findings, we now face an entire cartography of brain areas and

refined spatio-temporal patterns of brain activity, or rather, an urban jungle! Yet,

correlations do not entail explanation. The question is then, are we closer to find-
ing the mechanism(s) responsible for subjective, phenomenal qualities? We will ar-
gue that despite much effort and scientific investigation, one critical component—
the explanation of the phenomenological qualities—remains as elusive as it was
thirty years ago, which contrasts with the wealth of empirical data that have been
amassed meanwhile. We remain optimistic that such an explanation is within
reach, yet it will require us, the scientific community, to embrace a number of
different practices starting from acknowledging serious methodological pitfalls in
the contrastive method—the method that represents the cornerstone of our scien-
tific practice. Expanding the contrastive method will be critical to the success of a
biological research program on consciousness; yet this work will not happen if it
does not include the development of descriptions that address the phenomenologi-
cal level of organization of consciousness. Current theorizing has for the most part
largely disregarded this aspect since Revonsuo (2000) (see also Revonsuo, 2006 for
a more detailed account) made a plea for it; with just a few notable exceptions, e.g.,
the neurophenomenology approach advocated by the late Varela (1996), and the
axiomatic approach of Integrated Information Theory (Tononi, 2004) with its most
recent studies on the phenomenological qualities of space (Haun & Tononi, 2019).
Beyond that, due to the inherent bottom-up approach of the NCC program, as
a community we ought to find ways to aggregate data across multiple studies and
experimental paradigms while also designing studies targeted to test contradictory
predictions of different theories within the same experimental context. Only then,
will we be on a serious track to distill the NCC proper; anything short of this will
render the NCC program futile.
In what follows, we review earlier arguments that we and others have made
regarding the methodologies used to study the NCC, and we advance ideas on
how to bolster the NCC program. With this article, we aim to take a critical look
at Crick and Koch’s framework for studying consciousness, taking the progress
made since then as lessons for a renewed strategy for finding the neural processes
that give rise to consciousness. In this process, we remain neutral regarding the
different hypotheses that have been advanced in the field and their explanatory
power, while at the same we wish to speak to the methodological challenges that
lie ahead of us. We hope addressing the latter will enable us to look at the data
and the proposed theories through a new lens and to gain a unified, and acceptable,
explanation of consciousness in the not-so-distant future.

1.1 How the contrastive methods can help in finding the

NCCs
The main approach taken in finding the NCC, and recently advocated as one of the
hard criteria to evaluate empirical theories of consciousness (Doerig et al., 2021),
has been the contrastive method (Baars, 2005). This method consists in contrasting

conscious and unconscious states and/or contents in consciousness, while main-

taining any other factors constant. The reported perceptual state of the participant
is then taken as a dependent variable and the neural activity associated with the
different perceptual states is contrasted to reveal the NCCs (Dehaene, 2014). As
external factors are supposed to be maintained constant and only perceptual state
should vary, the associated neural activity was assumed to exclusively reflect the
processes involved in consciousness. This method has been employed to compare
the difference between wakefulness/vigilance and non-conscious states in order
to obtain the full NCC, or to compare the neural activity between different percep-
tual contents to yield the content-specific NCCs (Koch et al., 2016). The current
perspective focuses mostly on the latter; but the principles advocated should apply
to any attempt to find a neural correlate of either a state or a content of conscious-
ness.
Several experimental paradigms for erasing stimuli from awareness have been
used to study neural processes associated with both conscious and unconscious
conditions (Axelrod et al., 2015; Kim & Blake, 2005). One example of such an
experimental paradigm is masking, which involves the brief presentation of a crit-
ical stimulus closely preceded and/or followed by another stimulus—a so-called
“mask”—that renders the critical stimulus visible or invisible depending on the in-
terval between the mask and the critical stimulus (Dehaene et al., 2001). Partici-
pants can then report (e.g., verbally or via button press) whether they have seen
a stimulus or not. The neural activity is then compared on the basis of this re-
port. Using this experiment, combined with fMRI and EEG recordings, Dehaene
and colleagues were able to show that seen trials differed from unseen trials by a
widespread fronto-parietal activation (Dehaene et al., 2001). These findings were
further confirmed using other experimental paradigms, such as the attentional
blink, in which participants’ attention is focused on a specific task, rendering a
critical stimulus invisible for the participants (Sergent et al., 2005). They also found
that a late ERP component—the so called P3b—correlated best with the subjective
report of visibility on a continuous perceptual awareness scale (Del Cul et al., 2007).
Another technique that has been used to search for NCCs is binocular rivalry, a
phenomenon in which two incongruent stimuli are presented to each eye, leading
to alternating perception of exclusively one or the other. As the stimuli presented
to both eyes remain constant while perception alternates, this enables researchers
to compare stimuli whose status in consciousness varies while external stimula-
tion remains constant, and thus to dissociate sensation from conscious perception.
Using this method to investigate the NCC in non-human primates with single-cell
recordings, Logothetis and colleagues were able to show that neural activation
in the infero-temporal cortex strongly correlates with conscious perception, i.e.,
what the animal reports “seeing,” whereas neural activation in V1 correlates more
strongly with retinal input, i.e., the two stimuli presented to each eye (Logothetis,
1998). Other studies, however, concluded that the best correlates were found in
early sensory cortices, as opposed to higher-order sensory areas. Using threshold

presentation, Pins and ffytche located the best correlates of consciousness in an

ERP peaking at 100ms in the occipital cortex (Pins & ffytche, 2003). Similarly, in-
vestigating patients with Charles Bonnet syndrome, they were able to show that
the hallucination of percepts correlated with changes of activity in extrastriate
visual areas (ffytche et al., 1998).
This initial surge in empirical data studying the neural correlate of conscious-
ness contrasted with the drought of the decades prior to 1990. The increased avail-
ability of neural recording methods such as fMRI, PET, and MEG, together with
the clear methodology proposed by the contrastive method, seemed to make the
answering of the question of consciousness a goal finally achievable. Soon enough,
as data accumulated, several theories emerged aiming to explain by which mech-
anisms subjective experience emerges from neural activity. Many of these the-
ories are compiled in the seminal anthology of Metzinger (2000). For instance,
the early dichotomy of findings between early and late correlates was logically
reflected in those theories: the recurrent feedback theory proposed by Lamme
(2006) stated that consciousness emerges from local recurrent feedback loop in
early sensory cortex, while one of the variants of the higher-order theory (Lau &
Rosenthal, 2011) stated that consciousness is the result of higher-order processes,
taking place predominantly in prefrontal areas. Similarly, the global workspace
theory first proposed as a cognitive architecture by Baars (1997) was then further
extended by Stanislas Dehaene and colleagues (2001). They provided a biologi-
cally realistic implementation in which consciousness is thought to be the result
of widespread broadcasting of information throughout an interconnected network
of prefrontal-parietal areas and high-level sensory areas, making information avail-
able to a wide range of areas for further processing. Other theories attempted to
explain the “unity of consciousness”—the fact that all senses come together as a uni-
fied whole into one experience—proposing neural synchronization across cortical
areas, predominantly in the gamma frequency, as the critical mechanism explain-
ing the binding of features that gives rise to a unified experience (Crick & Koch,
1990; Edelman & Tononi, 2000; Melloni, Schwiedrzik, Müller, et al., 2011; Varela
et al., 2001). These initial theories were soon followed by more, to the point that
the field today is practically overwhelmed by a mass of available theories trying
to explain consciousness at different levels (Doerig et al., 2021).
Thirty years after the groundbreaking insights of Crick and Koch and twenty
years after Metzinger’s book, the enthusiasm of those early studies’ findings has
been tempered by the realization that the apparent simplicity of the empirical re-
search on NCCs in fact suffers from major methodological challenges. The con-
trastive method by itself was not selective to reveal the NCC proper. In what
follows, we provide a brief description of these methodological challenges, refer-
encing a selected set of studies to demonstrate the pervasiveness of the problem.
Our review of the literature will be from that vantage point and thus be inherently
selective. For recent comprehensive reviews of the empirical findings on the NCC,
we refer the reader to Boly et al. (2017) and Forster et al. (2020).

1.2 Methodological challenge: More than one NCC

The core assumption of the contrastive method is that when comparing between
conditions in which the external input is controlled for and only perception varies,
the resulting difference directly reflects neural activity involved in consciousness.
We and others have, however, argued that this strategy is too simplistic (Aru, Bach-
mann, et al., 2012; Graaf et al., 2012). We will briefly rehearse the main argument
(for a longer treatment of the issue we refer the reader to the original work Aru,
Bachmann, et al., 2012; see also Miller, 2015, where this problem is referred to as
the correlation/constitution distinction problem).
When comparing neural responses associated with two perceptual states, as
the contrastive method does, two other families of processes co-occur with the
ones directly reflecting consciousness: the NCC-precursors (NCC-pr) and the
NCC-consequences (NCC-co, Aru, Bachmann, et al., 2012, see Fig 1). NCC-pr
refers to processes that precede the NCC proper. They might enable a given
stimulus to reach consciousness but are not conscious themselves. The NCC-co
refer to processes that might follow the NCC proper. They result from conscious-
ness but are not conscious. A typical example of NCC-pr is attention. While the
role of attention in consciousness is still being debated (Koch & Tsuchiya, 2007;
Koivisto & Revonsuo, 2007; Pitts et al., 2018), it is now widely agreed that they
are indeed two different mechanisms (though that was not always the case, see
Posner, 1994). In classical contrastive paradigms, covert shifts of attention may
explain why some stimuli are seen while others are not. For example, in masking
paradigms, differential engagement of attention across trials might well explain
why an otherwise identical physical stimulus is perceived in some trials but
missed in others. The NCC-co are processes triggered by conscious experience
but not responsible for it. Some theories of consciousness assign consciousness a
function. Therefore, the fact that a critical stimulus was perceived must entail that
additional processes will follow it. Such processes include encoding in working
memory (Soto & Silvanto, 2014) and/or episodic memory, reflecting about the
perceived stimulus, and in the case of most experimental paradigms employing
the contrastive method, reporting about it (Tsuchiya et al., 2015).
This methodological caveat seriously called into question the wealth of early
findings, given that by themselves they do not enable the NCC-pr and NCC-co to
be distinguished from the NCC proper. As such, there is no guarantee that NCC
findings that attribute, for instance, the prefrontal cortex a role in consciousness
may not instead be reflecting NCC-co associated with report, task performance,
or encoding in working memory and/or episodic memory. The same holds true
for findings that state that early sensory activity constitutes an NCC, which could
instead merely reflect NCC-pr, e.g., attentional shift, expectations, etc. As findings
of NCC in early contrastive studies fostered the development of several theories of
consciousness, the impact of these methodological challenges and how much they
affect the proposed theories is to date unclear.

Figure 1: The contrastive method is implemented by experimentally comparing

conscious versus unconscious conditions e.g., via masking, binocular rivalry, etc.
The neural activations induced in the conscious vs. non-conscious condition are
contrasted to reveal the NCC. While apparently straightforward, this method was
argued to be insufficient to fully isolate the so-called NCC-proper (2). The results
of this simple subtraction are confounded by the occurrence of temporally flanked
mechanisms, termed NCC-pr (1) and NCC-co (3) (see also Aru, Bachmann, et al.,
2012). The former refers to the mechanisms enabling a stimulus to reach conscious-
ness without directly accounting for consciousness itself (yellow in the figure),
while the latter refers to mechanisms that are consequences of the conscious per-
ception of a stimulus but need not be conscious themselves (red in plot). Examples
of NCC-pr are attention and prior expectations (Aru, Axmacher, et al., 2012; Aru,
Bachmann, et al., 2012; Melloni, Schwiedrzik, Muller, et al., 2011). As most theories
of consciousness assign a function to consciousness, typical NCC-co accompany
it: encoding in memory, decision making to name a few. Additionally, as many
contrastive studies rely on report to differentiate conscious versus non-conscious
conditions, motor responses and planning, as well as self-monitoring, oftentimes
confound the NCC-proper (Tsuchiya et al., 2015).

The previously accepted NCCs therefore had to be reviewed by experimentally

controlling more thoroughly, ideally in the same experimental setting, for covary-
ing processes in order to isolate the neural activity directly related to conscious-
ness. We called this process the “distillation of the NCC,” as in effect it implies
removing all confounding variables until the remaining core neural activity that
correlates with consciousness is found. While many processes precede and suc-
ceed a conscious experience (and will have their own neural correlates), there is
one such confound of previous studies that recently has received particular atten-
tion, namely report. This relates to the fact that participants have had to overtly
report the perceptual state. Report can be performed verbally or via button press
(ffytche et al., 1998; Kleinschmidt et al., 2002; Logothetis, 1998; Pitts & Britz, 2011)
using either forced choice tasks (Dehaene et al., 2001; Gaillard et al., 2009; Hen-
son et al., 2008) or the perceptual awareness scale (Melloni, Schwiedrzik, Muller,
et al., 2011; Overgaard et al., 2006). When subjects report what they perceive on
a trial-by-trial basis, they engage additional processes not directly reflecting con-
scious perception: they have to monitor their own state, make decisions about
what to report depending on the afferent input, keep a template in their mind to
use for reporting, and both engage in motor planning and emit a motor response
(Tsuchiya et al., 2015). Introspective report has also been challenged on theoretical
grounds: Ned Block has put forward the view that reports only yield information
about higher-order thought processes, not about consciousness itself (Block, 2005).
To circumvent or minimize the engagement of post-perceptual processes, sev-
eral paradigms have been used in which report is not directly needed in order to
assess perception; these are referred to as no-report paradigms. One such example
is reliance on optokinetic nystagmus (i.e., eye movements) to estimate perceptual
content in binocular rivalry paradigms, which has been successfully implemented
both in humans (Frässle et al., 2014) and other animals, e.g., cats (Fries et al., 1997)
and monkeys (Kapoor et al., 2020). Using this methodology in combination with
fMRI, Frässle et al. (2014) showed that when drifting sinusoidal gratings moving
in different directions are exposed to each eye of the participant, the optokinetic
nystagmus closely tracks the current percept, as validated by comparison with
participant reports. When comparing the neural activation associated with per-
ceptual changes in the absence of report (i.e., using optokinetic nystagmus mea-
sure with that observed under overt report) Frässle and colleagues observed that
while parietal activation still tracked perceptual changes, frontal activation, found
under reporting conditions, no longer correlated with the perceptual state (see
Figure 3a and 3b in Frässle et al., 2014). Similarly, Farooqui & Manly (2018) using
fMRI showed that when consciously perceived stimuli are not relevant for task
performance, frontal regions get deactivated. These studies seem to indicate that
prefrontal activity indexes NCC-co as opposed to the NCC proper.
In a set of experiments using EEG to investigate the electrophysiological cor-
relates of consciousness with and without report using inattentional blindness of
shapes (Pitts et al., 2012), words (Schelonka et al., 2017), and faces (Shafto & Pitts,
2015), Pitts and colleagues found P3b responses putatively indexing consciousness

only when subjects were reporting. Comparable results were obtained in forward
and backward masking paradigms, whereby a robust P3b was found in the report
condition, but was absent when subjects did not need to report (Cohen et al., 2020).
Instead, they found that an earlier (200–300ms) negative component over posterior
areas correlated with consciousness independently from report. This component,
named the visual awareness negativity, has been found to correlate with conscious-
ness across a range of studies (Koivisto & Revonsuo, 2010), making it a prime NCC
candidate (but see Melloni, Schwiedrzik, Müller, et al., 2011 for a more nuanced
discussion of the temporal dynamics related to the NCC). Moreover, P3b has also
been shown to differ between frequent and rare stimuli in oddball paradigms, even
when all the stimuli were unconsciously perceived, which calls into question P3b
activation as an exclusive marker of conscious perception (Silverstein et al., 2015).
Such observations are not limited to the visual domain, as Sergent et al. (2021)
recently showed that the P3b correlates with awareness of auditory stimuli during
the active report condition, but not during the no-report task.1
While these studies seem to relegate late, frontal activation to post-perceptual
processes only, other studies employing invasive and more sensitive methods, such
as electro-corticography (ECoG) and single-unit recording, have revealed a more
nuanced picture. For instance, Noy et al. (2015), using ECoG in epilepsy patients,
showed that while the late-onset frontal activation was present under reporting
conditions only, an initial glow spreading through all cortical areas, including the
pre-frontal cortex, was observed regardless of report. Additionally, Kapoor et al.
(2020), using single-unit recordings in macaque monkeys, showed that in the ab-
sence of overt report the content of consciousness can be accurately decoded from
the prefrontal cortex in binocular rivalry. Thus, while some of the previously ob-
served prefrontal activation might index confounding factors such as report, lead-
ing to an exacerbated prefrontal cortex activation, a selective portion might still
correspond to the NCC proper. As such, while no-report paradigms might deal
well with disregarding post-perceptual processes at the level of the experimental
design, by themselves they do not resolve the inherent sensitivity issue faced by
most non-invasive methodologies, e.g., fMRI, PET, M-EEG. Multimodal methodolo-
gies, and in particular a combination of invasive and non-invasive techniques, will
be necessary to arbitrate on the role of prefrontal cortex in consciousness, which
has been challenged by no-report paradigms.
Nevertheless, not all confounds relate to report (Melloni, Schwiedrzik, Muller,
et al., 2011). Prior expectation also influences neural activations in contrastive
paradigms (Aru, Axmacher, et al., 2012; Mayer et al., 2016; Melloni, Schwiedrzik,
Muller, et al., 2011). Therefore, the mechanisms of expectation and prior knowl-
edge are also confounding the search for NCC. In paradigms with passive view-
ing, such as in the experiment by Cohen et al. (2020), encoding in working mem-
ory occurs in the seen but not in the unseen condition, also confounding the em-
1
In that study, the best correlate of conscious access was found in a component overlapping in time
with the P3-like component but with a bilateral positivity over temporal regions.

pirical findings. Attention level constitutes yet another such confounding factor.
As observed by Pitts and colleagues (2014), in the case of inattentional blindness
paradigms, seen and unseen trials might also differ in the level of attention. Neural
processes related to decision making can also obscure the NCC proper as demon-
strated in a recent study by Mazzi et al. (2020). Here, while both the visual aware-
ness negativity (VAN) and P3b (termed LP, i.e., late positivity in the text) correlated
with awareness, P3b was tightly correlated with decision-making, i.e., with deci-
sion criteria. Siclari and colleagues have also recently addressed confounds present
in state paradigms, i.e., those investigating neural correlates of wakefulness and
sleep in REM vs. NREM states, which are confounded by differences in neuromod-
ulation across states (Siclari et al., 2017). They have extended the methodology to a
no-task, within-state paradigm, in which subjects are serially awakened and asked
to report dreams both during REM and NREM, enabling investigators to contrast
‘consciousness’ to lack thereof (i.e., reported presence vs. absence of dream in both
REM and non-REM sleep) while controlling for both the state and the neuromod-
ulation milieu. Thus, there are a number of confounds, some already known, for
instance: report, prior expectations, working memory, episodic memory, task de-
mands, attention, decision making, and neuromodulatory state. This list is not ex-
haustive. As more studies are carried out and our understanding of the processes
uniquely related to consciousness is improved, this list is likely to be expanded.
Aru, Bachmann, et al. (2012) have previously proposed that, in order to isolate the
NCC proper, it will first be necessary to control for all of these potential (known)
confounds in order to look for the neural activity that systematically relates to
consciousness. In a sense, this pursuit can be framed as a problem of generaliza-
tion: the search for brain responses that systematically covary with consciousness
while manipulating all other nuisance variables, e.g., stimuli, task, experimental
paradigms, and other cognitive processes, etc. (Yarkoni, 2020). Certainly, no sin-
gle study is expected to manipulate all these variables at once. Instead, in addition
to controlling for as many confounding factors as possible within a single study,
it will be necessary to perform multiple studies to cross-validate the results: a re-
search program that will take years and significant collective effort to complete.
This problem has been noted and acknowledged by the scientific community, and
its impact on the theories is just now starting to be grasped (Yaron et al., 2021). For
instance, there is growing empirical evidence that the P3b, a marker once thought
to reflect the broadcasting of information into the global workspace (Sergent et al.,
2005), might instead reflect post-perceptual processes (Forster et al., 2020). These
significant developments have led to theoretical revisions of the theories—e.g.,
Global Neuronal Workspace theory (Sergent et al., 2021)—while at the same time
driving a slow convergence on the tighter link of other earlier, ERP components,
such as the VAN, with consciousness. Despite that encouraging progress, concrete
actions for addressing the challenges faced by the contrastive methods have lagged
behind, and most studies continue using that methodology as originally defined.
Moreover, to this day, there exists no database accumulating electrophysiological

and neuroimaging data from consciousness studies available to the community,

thus limiting the integration across different studies that is so dearly needed in
order to disentangle the NCC proper from all potential confounds. In its absence,
the performance of isolated, local studies will be highly inefficient and the search
for the NCC proper could prove to be a century-long enterprise.
Far from being a gloomy depiction of the state of the field, we believe the cur-
rent situation is a natural sign of its maturity. As the field has evolved, research
questions have crystallized even more, and both experimental challenges and their
potential solutions have manifested. In the next section we cover some proposals
for addressing these challenges and more. The list is not exhaustive, yet we hope
it may inspire readers to seek creative alternatives that can propel the field even
further over the next twenty years.

2 Homework for the field

2.1 Development of new methods for investigating the NCC
The contrastive method, recently extended by no-report paradigms, has dominated
the field of consciousness research. Even though it has been argued that the com-
parison of conscious vs. unconscious conditions is crucial for finding the NCC
(Doerig et al., 2021), it is not the only way to better understand consciousness
(Noy et al., 2015) and possibly not even the most desirable one as it suffers from
a severe problem of indeterminacy. In a typical experiment, we ask subjects to
report what they see, offering them binary choices, e.g., seen/unseen, or alterna-
tives such as the perceptual awareness scale (Melloni, Schwiedrzik, Muller, et al.,
2011; Overgaard et al., 2006). Yet in either case those reports tell almost nothing
about what subjects have actually experienced (unless they report on their phe-
nomenology; more on this point in the next section). When they say they have
seen a stimulus: What do they mean? What have they actually experienced? On
what basis are they judging the stimulus visibility? Is it the vividness of the stimu-
lus, its contrast, motion, color, shape, etc? As experimenters, we attempt to define
a “template” for the subjects to report on what they see, but we can never know
what they are actually experiencing. We sometimes use tricks, for instance, in
some trials we present nothing, expecting that subjects will report that they see
“nothing.” If they report having seen something in those trials, we have a dilemma:
either we are excited, and use those “false alarm” trials to look for the correlates of
consciousness without the confounds of sensory stimulation (Gosselin & Schyns,
2003; Kim & Blake, 2005), or we discard those trials/subjects, interpreting them as
featuring a “bias” (shift in criterion in signal detection theory) and not real per-
ception (Aru, Axmacher, et al., 2012; Dehaene et al., 2001; Mazzi et al., 2020). In a
nutshell, the reports that are typically obtained in most studies investigating the
NCC are complicated to interpret; they lead to a number of confounds and might
easily explain differences across studies, as “instructing” subjects similarly is not

easy. Complicating the matter, most studies do not even thoroughly report how
subjects were instructed and what measures were taken to ensure consistent be-
havior across subjects. Anyone who has tried to reproduce a study in a different
laboratory immediately notices how much the “context” of the research matters,
including how the investigator instructed the subjects. But little attention is paid
to making this information accessible in publications. What other approaches can
then be taken in order to move beyond the contrastive method?
Haynes (2009) has argued for the use of multivariate analysis to decode the
content of consciousness, thus providing a much finer identification of NCC than
the simple study of the dichotomy between presence and absence of critical stimuli.
Recent studies using this approach have shown that decodability of stimulus orien-
tation was highest from occipital sources in early time windows (130–320ms). Fur-
thermore, decoding performances increased with clarity of the percept as reported
on a perceptual awareness scale (Andersen et al., 2016). Salti et al. (2015), on the
other hand, showed decoding from frontal regions at later time points—270ms post
stimulus onset—of the spatial location of perceived targets. While decoding might
be an exciting approach for unravelling the neural correlate of consciousness, the
results are not easily interpretable; for example, they might spuriously suggest
widespread patterns that are contradicted when tested further with causal manip-
ulations (Bouton et al., 2018). As such, multivariate decoding analysis should be
considered with caution.
Other approaches relate to focusing not just on what happens following the
entry of a given stimulus into consciousness, but on their maintenance and, more
generally, on a defining feature of consciousness, i.e., the experience of flow and
temporality. With just a few notable exceptions (Gerber et al., 2017; Golan et al.,
2016), most direct empirical work has aimed at understanding the entry of stimuli
into consciousness, thus obviating the questions of how and why a conscious
percept is maintained in consciousness. In this regard, perceptual continuity is
an intriguing concept: we perceive the world as stable despite constant sensory
disruptions caused by eye movements and blinks (Golan et al., 2016). The latter, for
instance, causes a massive disruption in the sensory flow as the eyelid occludes the
pupil for approximately 100–150 ms, yet we do not perceive the world as blanking
out. Such a natural experiment occurs approximately 1000 times an hour (Cruz et
al., 2011), amounting to roughly 3–4% of our waking life spent with our eyes closed,
but without having noticed that the world has disappeared. A promising and more
naturalistic approach to investigating the mechanisms related to consciousness
is to focus on understanding what maintains a stimulus in awareness, i.e., why
we fail to update our phenomenology in light of massive changes in sensory
stimulation. As we have argued elsewhere (Melloni, 2015), understanding the flow
of consciousness—i.e., moving from episodes (the snapshot view that has domi-
nated studies on consciousness) to events (integrations across moments in time)
central to the experience of such “flow”—is another crucial, but hitherto neglected,

aspect of conscious perception. This resonates with early attempts by Varela (1999)
to address the experience of temporality, called “specious present” by James (1890,
p. 377), and with the proposition that internal brain dynamics, in particular the
synchronization of widely distributed neural assemblies in the gamma band and
phase transitions across states, lies at the core of time consciousness. Support for
this idea comes from studies investigating binocular rivalry (Cosmelli & Thompson,
2007; Doesburg et al., 2009, among others).
Another promising approach to investigating consciousness in populations
with restricted responsiveness, such as infants and patients with cerebral lesions,
is the use of neural frequency tagging, which we initially developed to evaluate
language capacities and access to meaning in neurotypical individuals (Ding et al.,
2016). Recently, we and others have used the frequency tagging approach for both
retrospective (Gui et al., 2020) and prospective (Sokoliuk et al., 2021) classification
on unresponsive patients, and the results have been encouraging.
Despite the clear advantages of the contrastive method, its mostly exclusive use
in the research of consciousness may have constrained the field too much, becom-
ing an obstacle to finding NCCs. It is expected that broadening the experimental
methods will help us to better understand the phenomenon.

2.2 Addressing the central problem of consciousness:

Phenomenology
While the previously discussed approaches go beyond the contrastive method, they
all fall short in one important aspect: the final goal of a science of consciousness
should be to understand its phenomenology and relate it to physical systems. The
ultimate frontier is to explain why we feel the way we do (akin to Levine’s (1983)
explanatory gap): why does vision feel different from audition, why does space
feel extended, why do we feel time as flowing?
Revonsuo (2000), in his visionary chapter, eloquently called for the develop-
ment of a science of consciousness that systematically describes the phenomenal
level of organization on its own terms—from the first-person point of view. Such
descriptions should go beyond cognitive and/or computational descriptions, and
they will need to be developed from within the science of consciousness. In the
spirit of previous traditions in phenomenology, the goal should be to describe the
structures of consciousness as experienced from the first-person point of view.
Progress in this area in the past decades has been meager. Instead, the field has
been taken up by a cognitivist/functionalist view that leaves little room for expla-
nations of the phenomenal qualities.
One important conceptual disagreement in the field refers to the distinction
between phenomenal (P) and access (A) consciousness (Block, 1995). Phenom-
enal consciousness refers to “what it is like to be in that state,” whereas access
consciousness refers to the availability of information to perform cognitive opera-
tions, such as reasoning about an experience, reporting it to an external observer,

etc. Phenomenal consciousness exists in and of itself, for our own inner spectacle,
while access consciousness is defined by the functions it serves. A shorthand for
the distinction between “being” and “doing.” This fundamental distinction led to
a clear dichotomy in the different theories of consciousness, with some assuming
independence of the two and some assuming them to be one and the same.
For some theories, such as Global Neuronal Workspace, phenomenal qualities
of experience are fully captured by access (see Naccache, 2018). Under this as-
sumption, studying access is equivalent to studying its phenomenology. Other
theories, in contrast, posit that P-consciousness is distinct and independent from
the collection of cognitive processes related to access (Tononi, 2004; Tsuchiya et
al., 2015). The debate on whether phenomenology overflows access is ongoing (Fu
et al., 2021). Notwithstanding the empirical challenge in arbitrating between these
different conceptions, it is important to refocus the debate to the main aim of the
science of consciousness, i.e., what theories of consciousness ultimately aim to ex-
plain is the phenomenological qualities of experiences, the “what it is like to be in
that state” (Revonsuo, 2006).
As a matter of fact, one reason why the debate between access and phenomenal
consciousness will not easily be settled empirically is that we lack clear descrip-
tions of the phenomenological qualities of experience assumed by the different
theories of consciousness. For instance, take the debate on the richness vs. sparse-
ness of consciousness (often discussed in concert with the above-mentioned debate
of A- vs. P-consciousness, see Block, 2011). One side posits that the richness of ex-
perience is an illusion and that what can actually be perceived and remembered is
limited. But what does it mean for something not to be rich? For instance, if a sin-
gle number, say the number 15 in red ink, is foveally presented for enough time to
be perceived, say 100 ms, and at a size that makes it easy to be perceived, what do
the different theories assume is consciously perceived? Just the number, just the
color, just the location, or the font, or the orientation of the letters, or all of those
features? This is not a naïve question, given recent work on attribute amnesia
(Chen & Wyble, 2016; Wang et al., 2021), which demonstrates that subjects fail to
report specific attributes of stimuli, e.g., color, that are otherwise reported as per-
ceived. Can we interpret these results as evidence that phenomenology overflows
access? Not easily, as (some) theories of consciousness have not made explicit
what they assume to be experienced (this is a separate question from whether P-
consciousness is fully subsumed under in A-consciousness). Some could claim that
subjects experience the letter but not its color (Fu et al., 2021). Thus, in the absence
of explicit assumptions from the theories about the phenomenological qualities of
consciousness, the debate between access and phenomenal consciousness will not
easily be settled empirically.
Nevertheless, settling the P- vs. A-consciousness debate, while representing a
significant advance in the field, will not provide understanding of the phenomeno-
logical qualities of experience and their generation by the brain. To that end, clear
and standardized methods for studying phenomenology in cognitive neuroscience

need to be developed. Phenomenology has been extensively discussed in philoso-

phy and efforts have been made to naturalize its investigation in the language of
the cognitive neurosciences (see Gallagher, 2012, for a more detailed account).
Without a doubt, obtaining phenomenological insights is a challenging task
(LeVasseur, 2003; Schwitzgebel, 2008), but it is a task that the science of conscious-
ness cannot shy away from inasmuch as it constitutes our very own object of
study. While we wait for the perfect methodology to naturalize phenomenology,
assuming that this is even achievable, there are, meanwhile, a number of areas in
which progress can be made.
For a start, by making explicit the assumptions that theories of conscious-
ness themselves have about phenomenological qualities of experience, these as-
sumptions can be used as a source for empirical investigation. In practice, such
assumptions can be tested through the front-loaded phenomenology, which con-
sists in using phenomenological considerations to design experimental paradigms
(Gallagher, 2003). By building experimental designs around phenomenological as-
sumptions, these can be tested directly and their neural correlates investigated.
Another advantage of this approach is that it makes direct comparison between
theories easier since it brings the comparison with regard to the “being” into focus,
suspending the question of whether “being” and “doing” are the same or different
depending on the theoretical account. There are a number of interesting areas in
which this approach can be tested, such as the temporality of experience, given
that divergent ideas have already been formulated (Dennett & Kinsbourne, 2011;
Varela, 1999). Other areas concern assumptions of primary versus secondary at-
tributes of objects. To return to our previous example regarding attribute amnesia,
it is possible that theories differ in what they assume to be mandatory conscious
perception of objects, e.g., an object’s color might not be considered an intrinsic at-
tribute while others, such as its location, might be. Similarly, when it comes to the
conscious perception of a face, which features are expected to be automatically
available to consciousness? Could a face be perceived without experiencing its
orientation or skin color, etc? Being explicit about the “grounding,” i.e., what the
theories assume experiences to be, will help us not only to refocus on what we aim
to understand, i.e., the phenomenal aspect of consciousness, but also to test such
assumptions and thereby improve our understanding. As such, the front-loaded
method is probably the one most readily compatible with cognitive neurosciences,
given that it requires neither an extensive conceptual or methodological shift, nor
extensive training of subjects (Gallagher, 2003).
An alternative, yet more loaded, approach could be the more extensive use of
neurophenomenology introduced by the late Varela (1996). This approach consists
in standardizing the phenomenological qualities of experience in mathematical
terms, under the working hypothesis that phenomenology and cognitive science
are engaged in a relation of reciprocal constraints, by which phenomenological
accounts should drive the cognitive neuroscience investigation of consciousness
and in return be driven by it. More specifically, this approach consists in isolating

invariants of conscious experience, formalized through mathematics to investigate

the neural mechanism accounting for them (Gallagher, 2003). The search for in-
variants has, for example, been used to investigate the commonalities of the
experience of time (Varela, 1999). This approach has been further operationalized
experimentally in a number of studies by training subjects to reliably report their
experiences (Bockelman et al., 2013; Lutz et al., 2002). While similar to the report
in contrastive paradigms, this method differs in that it uses introspection (stan-
dardized through training) to guide analysis, rather than fitting the reports into
pre-defined categories of perceptual states in order to analyze the data (Bockelman
et al., 2013). Using this method, Lutz and colleagues were able to show that the
inter-trial neural measurements variability correlates with reported cognitive con-
text from trained subjects, factors otherwise very difficult to control for (Lutz et al.,
2002). However, this approach presents a number of challenges already exposed by
Bayne (2004); for example, it is not immediately clear how neurophenomenology
naturalizes phenomenology, and this requires further conceptualization, as does
the effect of training as well as the extent to which it affects the experiences them-
selves. These are crucial questions; and while some of them have been addressed by
Antoine Lutz with his concept of generative passages (Lutz, 2002), and newer ideas
have been proposed in line with a computational approach to neurophenomenol-
ogy (Ramstead et al., 2021), there remains much more to be done. An interesting
proposition by Fink and colleagues in this issue, akin to the description of phe-
nomenal qualities through their invariants, puts forward that the mathematical
structure of phenomenal domains ought to be preserved in the NCC proper, thus
constraining the NCC proper beyond simple correlation, offering promising em-
pirical perspectives (Fink et al., 2021).
Another approach is to take insights from phenomenology at face value
and to use them to guide our understanding of whether and how a physical
system might be able to express phenomenal qualities. This is the approach
taken by the Integrated Information Theory (IIT), in which phenomenology is
used as a basis for the theory’s axioms—identified as invariants of any possible
experience—which are then translated into postulates in an attempt to address the
hard question of consciousness (Tononi, 2004; Tononi et al., 2016). IIT has been
used principally to evaluate consciousness in unresponsive individuals, as well as
to make predictions about which brain areas might participate in consciousness
and which ones might not (e.g., cerebellum Tononi et al., 2016). In a recent
interesting development, the machinery of IIT was used in an attempt to explain
the experience of space, i.e., why space feels extended (Haun & Tononi, 2019).
Extending the axiom-driven approach used in the development of the IIT, Haun
and Tononi derived basic principles that must be true in order for the experience
of space to feel extended. These principles were then derived into physical
properties that the cause-effect power of a system must satisfy in order to account
for the properties of spatial experience. Using the assumptions underlying the IIT,

they constructed a system of 8 binary units interconnected in a grid-like fashion,

and they were able to demonstrate that under certain configurations the unfolded
cause-effect structure of the system can account for the properties of spatial experi-
ence. Their demonstration constitutes a very interesting approach to bridging the
explanatory gap (Levine, 1983) by proposing an “explanatory identity” between
the phenomenology and its physical substrate.
In sum, a remaining challenge in the field is to diversify our experimental
approaches, explicitly incorporating the phenomenological dimension, which in-
cludes developing a systematic framework to describe the different ways in which
experience can be structured in normal and abnormal/pathological states. Only
then will we be closer to finding contact points between the mind and the brain.

2.3 The need for big data

Advancing research on the neural correlates of consciousness might require not
only improving in our methodology, i.e., the contrastive method, putting the fo-
cus on phenomenology, but also means for aggregating the abundance of rich
multimodal data that have been (and will be) gathered. This data encompass not
only correlative, invasive, and non-invasive recordings in both healthy subjects
and patients, but also causal measures, such as electrical stimulation and lesion
studies, in both humans and animals, e.g., macaque monkeys, mice, and, more re-
cently, corvids (Boly et al., 2017; Nieder et al., 2020; Palmiter, 2011; Redinbaugh
et al., 2020). As such, when it comes to empirical research on the neural corre-
late of consciousness, it is fair to say that the field has progressed tremendously
(for reviews of the literature, see Forster et al., 2020; Koch et al., 2016; Mashour et
al., 2020). We are far from those early days when studying consciousness within
the cognitive sciences was met with hesitation (Michel, 2019). Yet most of these
studies have been performed and reported in isolation, with little aggregation of
data to common databases (for recent efforts to create an aggregated database on
confidence, see Rahnev et al., 2020), which would enable discoveries and proper
evaluation of the reliability of the findings across experimental paradigms. Most
data aggregation has occurred at the level of reviews written primarily from the
perspective of the proponents of the theories, without careful treatment of data
contradicting a theory’s core postulates (Dehaene & Changeux, 2011; Koch et al.,
2016; Lau & Rosenthal, 2011). One example of the need for data aggregation and
metaanalysis relates to P3b as a marker of perceptual awareness. As previously dis-
cussed, recent evidence suggests that P3b indexes post-perceptual processes (Co-
hen et al., 2020; Forster et al., 2020), rather than perceptual awareness, as initially
thought; yet no systematic meta-analysis of the data has so far been carried out,
leaving open the question as to whether the results generalize across different set
of stimuli, tasks, experimental paradigms, etc., or might instead indicate that P3b
indexes post-perceptual processes under certain conditions, but not under others.
A similar example concerns the role of prefrontal cortex (PFC) in consciousness.

Studies using no-report paradigms, e.g., indexing perception through eye move-
ments (optokinetic nystagmus) and fMRI in humans, have shown that activity in
PFC may reflect report, whereas other studies in non-human primates—also using
no-report paradigms in a binocular rivalry paradigm, but measuring direct electro-
physiological activity (single unit)—reported decoding of the content of perception
from PFC, which is not explained by report per se. Aggregating data across stud-
ies in a common database would help to make sense of these disparate results.
The best approximation to date is Neurosynth (https://www.neurosynth.org/),
which provides a synthesis of human functional neuroimaging data only (Yarkoni
et al., 2011), built over automated extraction of activation coordinates. While it
addresses a critical need, and as such falls short of being a revolutionary resource,
Neurosynth suffers from a number of shortcomings. First, it currently includes
only neuroimaging data and does not include data from other modalities, such
as M-EEG, ECoG, single units, or lesion/stimulation data. Second, it does not in-
clude unthresholded data, which are critical for reliable meta-analytical results,
especially considering the enormous variability in analytical choices by the inves-
tigators (Botvinik-Nezer et al., 2020). Third, it currently incorporates descriptors
reflecting broad cognitive states, which will need to be refined to incorporate more
nuanced descriptions of the tasks and experimental paradigms.
Distilling the NCC or finding any meaningful correlate of consciousness
presupposes generalization across variables such as a stimulus’s parameters,
tasks, research site, etc. While the scientific aim is clear, i.e., generalization over
non-critical parameters, it is not done in practice, and is also not taken into
account statistically. Yarkoni (2020) has recently provided the clearest discussion
of this problem, calling attention to the need to align verbal descriptions (hypoth-
esis/theories) with statistical expressions in order to avoid current problems in
psychology/neuroscience, such as the replication crisis. As the number of vari-
ables to include in a statistical model is daunting, the most reasonable strategy is
to aggregate across multiple studies. In principle, the prescriptive strategy seems
rather simple: researchers openly share data, run a complex statistical model over
the data, and voilà—the NCC are discovered. However, it is not that simple, even
when leaving aside the conceptual disputes as to how to define consciousness.
Meaningful data aggregation requires far richer and more detailed descriptions of
the experiment and experimental setting than are commonly provided in publica-
tions (or even shared datasets). Recognizing such limitations, some researchers in
the community have made efforts to provide more comprehensive descriptions
of the study materials, e.g., through platforms such as protocols.io and/or lab
protocols, study protocols recently launched at PloS One, as well as through the
development of community standards, such as Neuroimaging Data Model (NIDM)
experiment. The major limitation for data aggregation is the insufficient metadata
descriptor of the study materials, without which no meaningful statistical data
aggregation enabling generalization is possible. Hence, sharing data, while useful,

is an insufficient measure unless it is accompanied by the corresponding metadata.

Thus, an initial, apparently trivial but quite important step towards data aggrega-
tion would be to share a clear and detailed description of the experimental materi-
als that is aligned with standard classifications, e.g., NIDM-experiments and tasks
ontologies (Poldrack & Yarkoni, 2016) when available, and, ideally, linked with a
persistent identifier (DOI) to make it findable and accessible, e.g., protocols.io.
Expanding such standard classification and ontologies to cover domain-specific
metadata for our research community will enrich the usability of these data, as in-
formation will be properly tagged and thus meaningfully digested and aggregated
by our community. A good start could be to expand the suite of tasks defined
in the cognitive atlas (Poldrack et al., 2011, accessible online at http://www.co
gnitiveatlas.org) to cover those tasks typically used in our community, and to
define a catalogue of the cognitive processes engaged in each of them. Another
challenge will be data aggregation across modalities, i.e., fMRI, M-EEG, ECoG, etc.,
and species, not least because these data need to be brought together into common
atlases (Bjerke et al., 2018); but there are efforts in this direction. Notwithstanding
the challenges that lie ahead, and while requiring development and community
efforts, data aggregation across studies seems to be a tractable problem. An initial
step towards this goal will simply be to aggregate the preexisting data reported
in the literature into common databases, gathering data across different studies,
including all research modalities, in order to create an interactive, community-
augmented meta-analysis tool for the science of consciousness (for similar efforts
in other communities, see http://metalab.stanford.edu). This effort will enable
a completely new cartography of the field as well as a theoretically neutral per-
spective allowing us to identify tendencies, such as whether theories rely on all
experimental paradigms or only a handful of them, etc (Yaron et al., 2021). The
ultimate goal, however, is much more ambitious: the capacity to share raw and/or
unthresholded data across investigators (including metadata). We and others (Aru,
Bachmann, et al., 2012; Graaf et al., 2012) have argued that the isolation of the NCC
proper from the confounds of NCC-pr and NCC-co will require the performance
of several studies to account for as many confounds as possible. The existence of
a centralized database would provide the community with a “mindscope,” which
would greatly facilitate this enterprise, enabling analysis across studies and taking
the specific experimental condition of separate studies as regressors to disentangle
pre- and post-perceptual mechanisms from the NCC proper. This will, however,
require a major sociological change in research culture in order to break the habit
of investigators of not sharing data, to embrace team science, and to collect meta-
data, thus enabling data reuse (FAIR data). The slow but steady progress in the
open science revolution and ongoing changes in incentives for data sharing will
certainly facilitate these initiatives.

2.4 Relying on theories and their inferential power: Com-

paring theories

While the aggregation of data to centralized databases will be crucial to the

progress of consciousness research, the technological and methodological chal-
lenges it implies are considerable, and it could take years before such resources
become easily and meaningfully available. On the other hand, the generalization
of the investigation of phenomenological qualities will require that the theories
clearly state what they assume those qualities to be. Furthermore, the obstacles
to obtaining reliable phenomenological insights attest to the difficulty of this
problem. Therefore, it will take considerable time before the above-proposed
development in the field of consciousness research can enable empirical progress.
One approach that can be implemented in the meantime, however, is to rely on
the inferential power of existing theories of consciousness, e.g., Local Recurrent
Processing (LRP) theory (Lamme & Roelfsema, 2000), Global Neuronal Workspace
Theory (GNWT, Baars, 2005; Dehaene & Naccache, 2001), Higher-Order Theory
(HOT, Lau & Rosenthal, 2011), and Integrated Information Theory (IIT, Tononi,
2004), to name a few. Inasmuch as theories specify a neural mechanism that
generates subjective experience, they all formulate explicit predictions and
boundary conditions for consciousness to emerge. The comparison of the theories
predictions could, therefore, serve as guiding principles for either testing theories
core predictions or comparing their predictive power against other accounts,
but most importantly, for guiding discoveries, given that theories should also
make predictions about novel observations/experiments (comparative testing of
theories predictions regarding the NCCs was extensively discussed by Fink, 2016).
The reliance on theories’ inferential power is, however, more challenging than
it seems. In the current state of the field many theories exist at different levels of
explanation and those are rarely compared directly (Doerig et al., 2021; Signorelli
et al., 2021). As discussed by Michel (2019), theories disagree not only on the mech-
anisms giving rise to consciousness but also on their background assumptions. A
critical divergence refers to the detection rule, i.e., what is considered conscious
versus not. For example, higher-order theories (HOT) assume that the failure to
report a stimulus amounts to the absence of consciousness. In contrast, other theo-
ries, such as IIT, LRP, and more generally first-order theories, state that conscious-
ness can indeed occur in the absence of report. Therefore, while the findings of
no-report paradigms may challenge HOT, proponents of HOT can easily dismiss
them by rejecting the background assumption that stimuli were conscious despite
the absence of report, thereby leaving their theory intact.
The comparison of theories is challenging, but it is not unsolvable. Doerig et
al. (2021), recently advanced four so-called hard criteria for evaluating theories
of consciousness. We commend the proposal for finding agreed-upon criteria, yet
some might be too restrictive, as in the case of the blind reliance on the contrastive
method (e.g., criterium#1, “paradigm cases of consciousness and the unconscious

alternative”) for the reasons previously outlined, or the second criterion of the
unfolding argument (e.g., criterium#2, “the unfolding argument”). In the view of
Doerig et al. (2021), because any input-output function of a recurrent network can
be approximated by an equivalent feedforward network, any theory of conscious-
ness that assumes the necessity of a specific causal structure for consciousness to
emerge is either false or not within the reach of science. This purely functionalist
perspective, which equates consciousness with its input-output function, leaves
little room for rich, phenomenological states, such as dreams. Others have ar-
gued that the treatment of the causal structure as a black box is largely unjustified
(Tsuchiya et al., 2019). Overall, the reductionist strictness of the hard criteria devel-
oped by Doerig et al. (2021) runs the risk of locking down the field to a similar state
as what it was in in its early stages and at a time when it could benefit the most
from a broadening of experimental methodologies (see Tsuchiya et al., 2019, for a
more detailed argumentation against Doerig’s hard criteria for consciousness).
Until agreed-upon criteria emerge, we believe that mechanisms such as
“adversarial collaboration” might offer a viable alternative for arbitrating among
competing theories, as proven by the current existence of several such projects
in the field (such as Melloni et al., 2021). This practice consists in bringing
advocates of two or more opposing theories together to work on a common
experiment or set of experiments (Cowan et al., 2020). By doing so, the different
sides agree beforehand on rules that cannot be broken without direct implications
for their theories. In the framework of consciousness research, for instance,
higher-order and first-order theories could agree on detection rules for a given
experiment (or set of experiments), such that the condition in which a stimulus
is consciously perceived is agreed upon by both sides, side-stepping the problem
of under-determination (Michel, 2019). As each theory has a specific set of
predictions about which patterns of neural activity will lead to consciousness, the
experiment(s) should be designed in such a way that direct comparison of oppos-
ing predictions between the theories is possible. Findings will thus result in the
confirmation or disproof of specific aspects of a theory. While a winner-takes-all
situation, in which one of the theories is fully discredited, is unlikely to occur, such
projects lead to efficient identifications of core vs. auxiliary predictions (Lakatos,
1973). But perhaps more importantly, by explicitly stating these core and auxiliary
predictions and pre-registering them prior to data collection (as recommended
by Cowan et al., 2020), the need to patch-up theories by transforming core into
auxiliary predictions of the data is made evident and is therefore tractable. Such
an effort could not only aid in settling standing quarrels in the field (Michel,
2019), it could also identify the most promising theories, while sidestepping
some of the challenges of the contrastive method and its indeterminacy. This is
without a doubt a challenging process, yet it is a fruitful one and is already imple-
mentable. Currently, there are two consortia, one focused on testing contradictory
predictions of the Global Neuronal Workspace Theory and the Integrated Infor-
mation Theory, and the other pitting first-order, recurrent processing theory (RPT),

against higher-order theories of consciousness; but more are expected to come as

part of the Accelerating Research on Consciousness program spearheaded by the
Templeton World Charity Foundation. We are excited about these developments
and the impact they will have on the field and the theories in the future.

3 Conclusion
We have come a long way in the research of consciousness over the past thirty
years. The initial enthusiasm and the abundance of data obtained through the
contrastive method were followed by the development of a number of theories at-
tempting to explain how consciousness fits into the physical world. As the field
advanced, that initial enthusiasm faded and methodological as well as theoretical
and philosophical disputes clouded the horizon. At the same time, more and more
innovative approaches have emerged, expanding the range of possibilities beyond
the traditional use of the contrastive method. We would like to advocate for these
innovative methods, since the best way through what appears to be an impasse
in the field will be via methodological and theoretical shifts in current practices.
We believe it will be key for future discovery to transcend the basic dichotomy of
seen vs. unseen by bringing phenomenology into the center of the empirical effort.
This will be accomplished, first, by expanding efforts to describe the structure of ex-
perience and using those descriptions to guide empirical research, and second, by
inviting theories to make explicit their assumptions about phenomenological prop-
erties of experience so that they are then accessible to the front-loaded approach.
Furthermore, we believe great advances will be facilitated by the community’s ac-
tive effort to gather empirical data in common databases, to enable cross-studies
analyses, and to increase the spatio-temporal grain. Beyond that, the existence of
common databases will help narrow the search for the NCC by disentangling the
NCC-pr and NCC-co from the NCC proper. While no agreed-upon theory has so
far emerged, adversarial collaboration could offer a powerful tool for testing the
explanatory power of the theories, while also leveraging the theories’ inferential
power to fuel novel discoveries about what the function of consciousness might
be and how experience arises in the soggy matter of the brain.

Acknowledgments
We would like to thank Caspar M. Schwiedrzik and the Cogitate Consortium for discus-
sions, to Diana Koch for the figure and proofreading of earlier versions and to William
Martin for the final proofreading. We would also like to thank the Max Planck Society and
the Templeton World Charity Foundation, Inc. (TWCF0389) for their generous support.
The opinions expressed in this publication are those of the authors and do not necessarily
reflect the views of the Templeton World Charity Foundation, Inc.

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Lepauvre & Melloni - The Search For The Neural Correlate of Consciousness. Progress and Challenges (Philosophy and The Mind Sciences, 2021)

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Lepauvre & Melloni - The Search For The Neural Correlate of Consciousness. Progress and Challenges (Philosophy and The Mind Sciences, 2021)

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The search for the neural correlate of

consciousness: Progress and challenges

This article is part of a special issue on “The Neural Correlates of Consciousness,”

—Hippocrates of Cos, The Sacred Disease, ~400BC

refined spatio-temporal patterns of brain activity, or rather, an urban jungle! Yet,

1.1 How the contrastive methods can help in finding the

conscious and unconscious states and/or contents in consciousness, while main-

presentation, Pins and ffytche located the best correlates of consciousness in an

1.2 Methodological challenge: More than one NCC

Figure 1: The contrastive method is implemented by experimentally comparing

The previously accepted NCCs therefore had to be reviewed by experimentally

and neuroimaging data from consciousness studies available to the community,

2 Homework for the field

2.2 Addressing the central problem of consciousness:

need to be developed. Phenomenology has been extensively discussed in philoso-

invariants of conscious experience, formalized through mathematics to investigate

they constructed a system of 8 binary units interconnected in a grid-like fashion,

2.3 The need for big data

is an insufficient measure unless it is accompanied by the corresponding metadata.

2.4 Relying on theories and their inferential power: Com-

While the aggregation of data to centralized databases will be crucial to the

against higher-order theories of consciousness; but more are expected to come as

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