Preprint.

Final version is forthcoming in:

Perspectives on Psychological Science.

Perceptual representations
and the vividness of stimulus-triggered
and stimulus-independent experiences

Peter Fazekas1,2,✧, Georgina Nemeth3 & Morten Overgaard2

1 Centre for Philosophical Psychology, University of Antwerp, Belgium
2 Cognitive Neuroscience Research Unit, CFIN, Aarhus University, Denmark

3 Department of Affective Psychology, Eötvös University, Hungary

✧corresponding author, email: fazekas.peter@gmail.com

Summary
In recent years, independent subfields have started to engage with the idea that the
same cortical regions that contribute to online perception are recruited during and
underlie offline activities like information maintenance in working memory, mental
imagery, hallucinations, dreaming, and mind wandering. Accumulating evidence
suggests that in all these cases the activity of posterior brain regions provide the con-
tents of experiences. This paper is interested in moving one step further by exploring
specific links between the vividness of experiences, which is a characteristic feature of
consciousness regardless of its actual content, and certain properties of the content-
specific neural activity patterns. Investigating the mechanisms that underlie mental
imagery and its relation to working memory, and the processes responsible for mind
wandering and its similarities to dreaming form two clusters of research that are in
the forefront of recent scientific study of mental phenomena, yet cross-talk between
these two clusters have been surprisingly sparse. Here our aim is to foster such a
cross-talk by articulating a hypothesis about the fine-grained phenomenological
structure determining subjective vividness and its possible neural basis that allows us
to shed new light on these mental phenomena by bringing them under a common
framework.

Keywords: consciousness; perception; mental imagery; dreaming; mind wandering;

vividness; intensity; specificity.

1. Introduction
Recent efforts comparing conscious experiences that arise in different condi-
tions, e.g. during wakeful perception, mental imagery, mind wandering or
dreaming, have typically concentrated on finding what is common in the under-

1
lying brain processes to identify the possible neural substrate of having con-
scious experiences in general, i.e. to home in on the so-called neural correlate of
consciousness (NCC). In this paper our aim is to move one step beyond this
NCC-centric approach and investigate what can be said about the individual
phenomenological characteristics of such conscious experiences and about the
possible features of brain activity that might underlie these qualities.
The phenomenological characteristic that we will focus on is the vividness
of experiences. Regardless of their kind and content, experiences come with a
certain level of vividness. Although vividness is a quality in terms of which dif-
ferent experiences are routinely compared, it is a vague and ambiguous notion,
without clear definition. Intensity, liveliness, salience, prominence, clarity and
detailedness are but a few examples of those terms that vividness is often
claimed to be synonymous with. It is thus also a specific goal of this paper to
clarify what standard psychological measures of vividness ultimately inform us
about.

2. Background: shared content-specific activity in stimulus-

triggered and stimulus-independent experiences
Whether the ‘seat of consciousness’ is in the back or in the front of the brain is
hotly debated. On the one hand, theoretical approaches that emphasise the im-
portance of cognition in giving rise to conscious experience attribute the critical
role to frontal regions (Brown, Lau, & LeDoux, 2019; Dehaene & Changeux,
2004, 2011; Dehaene, Changeux, Naccache, Sackur, & Sergent, 2006; Dehaene,
Charles, King, & Marti, 2014; Dehaene, Kerszberg, & Changeux, 1998; De-
haene, Lau, & Kouider, 2017; Dehaene & Naccache, 2001; Lau & Rosenthal,
2011; Sergent, 2018). On the other hand, accounts claiming that the major con-
tribution is made by local perceptual processing locate the neural correlates of
consciousness in posterior areas (Block, 2005, 2007; Lamme, 2006, 2010, 2018;
Lamme & Roelfsema, 2000). Lesion, electric stimulation and neuroimaging
studies have been argued to support this latter view (Boly et al., 2017; Koch,
Massimini, Boly, & Tononi, 2016). However, the correct interpretation of these
findings is a matter of dispute (Odegaard, Knight, & Lau, 2017; Phillips, 2018;
Pitts, Lutsyshyna, & Hillyard, 2018).
Be that as it may, there is a relatively broad consensus about the essential
role of the posterior areas (low- and high-level sensory areas, precuneus, pos-
terior cingulate, retrosplenial cortex) in determining the contents of conscious

2
experiences regardless of the presence or absence of matching stimuli (Boly et
al., 2017; Siclari et al., 2017).
In the case of stimulus-triggered online perception these occipital, tempor-
al and parietal regions process the incoming information and form content-spe-
cific neural representations that are selectively activated by colours, faces, places,
etc. (Dehaene & Changeux, 2011; Dehaene et al., 2006; Lamme, 2006). Accord-
ing to recent sensory recruitment theories of working memory, the very same
regions are also responsible for maintaining information after stimulus offset,
i.e. the neural representations that are active during the processing of incoming
information are also active during the maintenance of this information in work-
ing memory (Christophel, Klink, Spitzer, Roelfsema, & Haynes, 2017; D'Es-
posito & Postle, 2015; Emrich, Riggall, LaRocque, & Postle, 2013; Postle, 2015;
Riggall & Postle, 2012).
This information-maintenance component of working memory (Fazekas
& Nemeth, 2018) has recently been tightly linked to mental imagery by a study
that found evidence of common representations in early visual areas shared by
mental imagery and visual working memory (Albers, Kok, Toni, Dijkerman, &
de Lange, 2013), and by the subsequent interpretation that the phenomenon of
mental imagery is, in fact, the dynamic component of visual working memory
(Tong, 2013). Moreover, several recent studies have found a large neural overlap
between perception and mental imagery, and showed in particular that the con-
tent of mental imagery is subserved by the activation of the same posterior areas
that are active during online perception (Dijkstra, Bosch, & van Gerven, 2017a,
2019; Ishai, Ungerleider, & Haxby, 2000; Johnson & Johnson, 2014; Naselaris,
Olman, Stansbury, Ugurbil, & Gallant, 2015; O'Craven & Kanwisher, 2000;
Reddy, Tsuchiya, & Serre, 2010; Stokes, Thompson, Cusack, & Duncan, 2009).
The perceptual contents of conscious experiences occurring during dream-
ing have recently also been associated with the activity of the posterior regions
in question. Horikawa and colleagues (2013) could accurately identify the broad
categories of content-elements of sleep onset mentations using decoding models
that had been trained on fMRI-measured stimulus-induced brain activity in
visual cortical areas. This demonstrated that the neural representations underly-
ing the contents of experiences were shared by stimulus-triggered perception
and sleep onset mentations. Siclari and colleagues (Siclari et al., 2017) extended
this link to experiences occurring in REM sleep, by showing that dream experi-
ences with faces, definite spatial setting, a sense of movement, and speech cor-
related with increased high-frequency EEG activity in regions that are associ-
ated with these content-types during wakeful perception.

3
 That is, the claim that the contents of conscious experiences are subserved
by neural activity in posterior areas is well supported by findings from many
independent strands of research. This claim serves as the starting point for the
rest of our paper. Given that it is known where one should look for the neural
activity underlying specific contents of consciousness, our question is whether it
is possible to move one step further and try to link features of this neural activ-
ity with phenomenological features of conscious experiences.

3. The vividness of experiences

The phenomenological feature of consciousness that we will focus on is the sub-
jective vividness of experiences. Vividness is a central phenomenal feature of a
broad range of (if not all) experiences: for instance, pains, felt emotions, visual,
auditory, olfactory, tactile experiences all come with a certain level of vividness.
The vividness of these experiences plays an important role in structuring our
phenomenal life (Morales, 2019) and has significant cognitive effects. For ex-
ample, vivid dream contents are easier to remember (Cohen & MacNeilage,
1974), vivid mental imagery elicits incidental recall (D'Angiulli et al., 2013) and
can even lead to false remembering (Gonsalves et al., 2004), meta-consciousness
in mind wandering is more sensitive to vivid experiences (Schooler, 2002), and
vivid hallucinations feel more realistic (Stephan-Otto et al., 2017).

3.1 What is vividness?

To see how subjective vividness is operationalised in psychological investiga-
tions, a good starting point is provided by the questionnaires that are designed
and used to measure the level of vividness in different contexts.
Versions of the so-called Vividness of Visual Imagery Questionnaire
(VVIQ) have been widely used since 1973 to measure the vividness of conscious
experiences occurring during mental imagery tasks (Marks, 1973). In its original
form, VVIQ asks subjects to rate the vividness of their imagery on a 5-point
scale where 1 means ‘perfectly clear, and as vivid as normal vision’, 2 means
‘clear and reasonably vivid’, 3 means ‘moderately clear and vivid’, 4 means
‘vague and dim’, and 5 means ‘no image at all’. Although routinely used in ex-
perimental practice, the notion of vividness that VVIQ tries to measure (VVIQ-
vividness, for short) is notoriously problematic as it is only intuitively defined
with the use of other concepts like clarity, detail, brightness, intensity, etc.
which themselves are then left unexplained (see e.g. Cornoldi et al., 1991; Denis,
1995; Kind, 2017; McKelvie, 1995a, 1995b). Even whether VVIQ-vividness is

4
supposed to be a single feature of experience or a construct with more than one
components is debated (McKelvie, 1995a).
In his enormously rich review and analysis, Stuart McKelvie (1995a,
1995b) argues that VVIQ-vividness is a combination of two factors: clarity and
liveliness, where the former characterises the sharpness and detailedness, while
the latter characterises the illumination, brightness and intensity of the content
of experience. McKelvie also emphasises that, as it is suggested by the fact that
the maximum value of VVIQ-vividness is related to the clarity and liveliness of
normal vision, VVIQ asks subjects to measure their imagery against their full-
blown visual experiences, i.e. VVIQ is indicative of how similar one’s imagery
is to one’s normal (stimulus-triggered) visual experience.

3.2 Vividness as the quality of experience

In the context of whether consciousness is an all-or-nothing phenomenon or it
comes in degrees, the so-called Perceptual Awareness Scale (PAS) has been used
to measure how degraded people’s conscious experiences of a masked stimulus
can be (Ramsøy & Overgaard, 2004; Sandberg & Overgaard, 2015; Sandberg,
Timmermans, Overgaard, & Cleeremans, 2010). By using PAS subjects can rate
on a 4-point scale whether they had no impression of the stimulus (‘not seen’),
whether they have a feeling that something had been shown but cannot specify
any features of the stimulus (‘weak glimpse’), whether they had an ambiguous
experience of the stimulus with more vivid impressions of some stimulus as-
pects and less vivid impression of others (‘almost clear experience’), or whether
they had a clear, specific, unambiguous experience of the stimulus (‘clearly
seen’).
As comparing the descriptions of PAS and VVIQ reveals, what PAS meas-
ures in the context of stimulus-triggered visual experiences is very similar to
what VVIQ measures in the context of mental imagery: whether experiences of
masked stimuli in the former case and of one’s imagery in the latter case are as
clear and as vivid (lively) as full-blown visual experiences are. That is, in line
with McKelvie’s analysis, VVIQ and PAS measure the similarity between specif-
ic experiences that are degraded for some reason and normal, full-blown experi-
ences — i.e. how reduced these specific experiences in question are in quality.
According to a recently proposed theoretical model, consciousness can be
reduced in quality along many different dimensions (Fazekas & Overgaard,
2016, 2018a, 2018b). The way the content-elements entering consciousness ap-
pear in a conscious experience — what is called the quality of the experience in
this context — is determined by a number of factors, many of which are inde-

5
pendent. In fact, these different factors form factor-families determining high-
level characteristics of experiences like their subjective intensity, specificity and
stability. Subjective intensity is determined by how much the content-element in
question stands out from the perceived background. More intense content-ele-
ments have more strength (Morales, 2019) and more liveliness (McKelvie,
1995a). Subjective specificity is determined by how distinguishable a content-ele-
ment is from other content-elements. A less specific experience of a content-
element is more generic, more vague, and more ambiguous. Subjective stability is
determined by how long a content-element is present in the experience. Less
stable content-elements occur in an experience only for a shorter period of time
(Fazekas & Overgaard, 2016, 2018a, 2018b).
As common characteristics of factor-families, these features can change in
many different ways. For example, a content-element can visually appear in an
experience with a higher subjective intensity if its contrast is increased, or if its
saturation is higher, or if its brightness is amplified. Similarly, a content-ele-
ment’s subjective specificity can change along many different sub-dimensions: it
can be less ambiguous by being highly precise (specific shade of red vs. generic
red), or by being sharper and less blurry, or by being rich in details (photograph
vs. line drawing). Note that these are independent factors that can be modulated
either individually or in any combination.
Standard measures of vividness (VVIQ, PAS) conflate these different
factors. What they measure is the quality of the conscious experience in ques-
tion, but their resolution is not fine-grained enough to uncover the different
levels of degradation along these different dimensions. This conclusion is sup-
ported by studies aiming to find the neural underpinnings of subjective vivid-
ness ratings.

4. The neural bases of the factors of vividness

The neural mechanisms underlying at least some of the different factors that af-
fect the vividness (quality) of experiences are well known. In what follows, the
focus will be on subjective intensity and subjective specificity. For more discus-
sion of the temporal factor see (Fazekas & Overgaard, 2018a).

4.1 Subjective intensity

Subjective intensity in the contrast sense, i.e. apparent contrast, is associated
with the strength of the response of edge detectors: allocating attention to a
Gabor-patch increases the apparent contrast of the stimulus by inducing an in-
crease in the amplitude of the response function of the corresponding popula-

6
tion of orientation sensitive neurons (Carrasco, Ling, & Read, 2004). Attention
has a similar effect on subjective intensity in the saturation sense, i.e. apparent
saturation, as well (Carrasco, 2011; Carrasco & Barbot, 2019; Fazekas & Nanay,
2018): allocating attention to a colour patch increases the apparent saturation of
the colour by inducing an increase in the amplitude of the response function of
the corresponding population of hue sensitive neurons, which associates appar-
ent saturation with the strength of the response of hue detectors (Fuller & Car-
rasco, 2006). In a very similar vein, subjective intensity in the brightness sense
has been associated with the strength of the response of neurons representing
object surfaces: change in apparent brightness can be evoked by modulating the
luminance of the surround of the object surface in question that induces a
phase-shifted change in the strength of the neural response of the population
representing the object surface (Rossi & Paradiso, 1999).
These findings suggest that factors of subjective intensity — contrast, sat-
uration and brightness, all prothetic dimensions with meaningful zero values
and inherent directionality (Fuller & Carrasco, 2006; Stevens & Galanter, 1957)
— are neurally encoded in a similar fashion, by the strength of the response
functions of populations of neurons that represent specific metathetic features
(with qualitative differences and without inherent less-to-more directionality)
like orientation, hue, and object surfaces. As increased strength of neural re-
sponse functions means increased firing rates, these findings predict that vari-
ance in subjective intensity along these sub-dimensions (individually or com-
bined) could in theory be detected as modulations in the BOLD signal and in
high frequency EEG activity (Le Van Quyen et al., 2010; Masuda & Doiron,
2007; Panzeri, Macke, Gross, & Kayser, 2015).

4.2 Subjective specificity

The neural underpinnings of the different factors of subjective specificity are
more diverse. Allocating attention to a particular hue, say, crimson, increases
subjective specificity of the apparent colour in the precision sense, i.e. decreases
the ambiguity between different shades of red by sharpening the population re-
sponse function (decreasing its variance), and thus increasing the precision of
the population code (Martinez-Trujillo & Treue, 2004; Maunsell & Treue,
2006), thereby providing more unique neural representations coding for the fea-
ture in question (Carrasco, 2011; Fazekas & Nanay, 2018). This suggests a link
between the subjective precision of the experience and the precision of the un-
derlying neural representation.

7
 Subjective specificity in the blurriness sense has been associated with in-
formation in high spatial frequency channels, i.e. information about fine-
grained edges. Such information might not be available due to at least three dif-
ferent reasons: stimulus blur, when the stimulus itself is blurry (as in the case of
using of a low-pass filter to produce the stimulus); optical blur, when optical
problems with the eye (scattering and optical aberrations) result in blurry vis-
ion; and ‘neural blur’, i.e. relatively low level of activity of neurons with small
receptive fields, which can result from low neural sensitivity to higher spatial
frequencies in the case of stimulus-triggered perception or from a low level of
recruitment of early visual representations in the case of stimulus-independent
percepts (Webster, Georgeson, & Webster, 2002; Webster & Marcos, 2017). Sim-
ilarly to the case of precision, the allocation of attention can modulate apparent
blur as it increases visual acuity by shrinking the effective receptive field of
neurons thereby enhancing their spatial resolution and increasing their sensitiv-
ity to high spatial frequencies (Abrams, Barbot, & Carrasco, 2010; Fazekas &
Nanay, 2018; Gobell & Carrasco, 2005).
Relatedly, subjective specificity in the detailedness sense, i.e. the amount
of fine-grained detail in the experience is determined by the processing of high
spatial frequency information that is associated with activity at the lower levels
of the visual hierarchy in the occipital cortex (Lu et al., 2018). The rapid pro-
cessing of low spatial frequency (coarse grained) information along the dorsal
visual stream results in a coarse parsing of the visual scene, which then, via
feedback to primary visual cortex, guides the slower analysis of high spatial fre-
quency information along the ventral stream (Kauffmann, Ramanoël, & Peyrin,
2014; Musel et al., 2014).
According to the emerging picture, thus, higher levels of subjective spe-
cificity both in the blurriness and the detailedness sense require broader in-
volvement of early visual areas, i.e. require a higher level of recruitment of early
visual processing.

5. The neural signatures of vividness in stimulus-triggered and

stimulus-independent experiences
The links between features of neural activity patterns and factors of subjective
vividness established above allow us to move beyond the resolution standard
interpretations of empirical data offer with regard to how vividness changes dur-
ing different varieties of stimulus-triggered and stimulus-independent experi-
ences.

8
5.1 Online perception
In the case of stimulus-triggered online perception the relation between brain
activity and vividness as measured by subjective PAS ratings has been extens-
ively explored.
In an early study, Christensen and colleagues investigated how the fMRI-
detected BOLD signal changed with different levels of subjective vividness
(measured with a 3-point PAS-like scale; Christensen, Ramsøy, Lund, Madsen,
& Rowe, 2006). They found that the characteristics of the activity in areas that
subserve the content of consciousness correlate with the phenomenal features of
how the specific content-elements appeared in one’s conscious experience. In
particular, they identified a correlation between the level of subjective vividness
of the experience and the intensity of the activity of these areas: vivid experi-
ences were accompanied by a significantly higher level of brain activity than
non-vivid experiences (both in a direct comparison and in comparisons to no
experiences). This finding, thus, suggests that masked stimuli appear in con-
scious experience with a degraded level of vividness in the sense of having a
lower level of subjective intensity.
More recent studies point towards a similar conclusion, and provide more
accurate localisation of the relevant neural activity both in time and in space
(relative to the content of the conscious percept). Andersen and colleagues
showed that in the case of simple geometrical figures the different levels of
vividness reported using the PAS-scale could be decoded from MEG signals
from the occipital lobe within the VAN (Visual Awareness Negativity) time
window using multivariate classification algorithms (Andersen, Pedersen, Sand-
berg, & Overgaard, 2016). Others also found an association between the intens-
ity of neural signals and the PAS-measured vividness of corresponding experi-
ences, and reinforced that the intensity of VAN activity reflects the level of
vividness of phenomenal awareness (Fu et al., 2017; Tagliabue, Mazzi, Bagattini,
& Savazzi, 2016).
Note that while these findings do establish a correlation between vividness
and the level of neural activity they are not able to provide information about
the more fine-grained structure of the factors of vividness. As Andersen and col-
leagues conclude, their findings show that differences between different PAS-
scores “are best explained by the conglomerate activity of the neurons in the
occipital lobe during the VAN time range” (Andersen et al., 2016). This result,
however, is compatible with two scenarios: (1) higher PAS ratings are underlain
by bigger proportions of activated occipital lobe, i.e. the intensity of the activity

9
does not change, it only spreads through the occipital lobe, and (2) higher PAS
ratings are underlain by more active occipital lobe subregions, i.e. the region of
activity does not change, just its intensity. That is, these results are indecisive
with regard to whether variations of PAS-scores reflect variations in subjective
intensity or in subjective specificity.

5.2 Working memory maintenance

Moving away from stimulus-triggered experiences, consider the maintenance of
perceptual information in working memory after stimulus offset.
In an experiment where subjects were asked to retain the orientation of a
full-contrast square-wave grating presented for 1 sec in memory and report it by
turning a probe grating after a 12 sec delay interval, Ester and colleagues, rely-
ing on fMRI data from areas V1 and V2 and using a forward encoding model of
orientation selectivity (Brouwer & Heeger, 2009, 2011), generated a set of ori-
entation selective response functions (tuning profiles) to evaluate the features of
the neural representations maintained by populations of neurons during the
delay period (Ester, Anderson, Serences, & Awh, 2013). These tuning profiles
showed the amplitudes of the population response as a function of orientation,
and they peaked around the orientation stored in working memory. It was
found that individual differences in the dispersion (precision), but not the amp-
litude (intensity), of the tuning profiles were robust predictors of the parti-
cipants’ mean recall error (the difference between the orientation of the target
and the orientation of the probe they set). That is, broader tuning profiles, i.e.
less precise neural representations of the target stimulus maintained during
working memory storage, were associated with greater error. The delay period
signals from areas V1 and V2 used to estimate the tuning profiles disappeared
when subjects received a cue at stimulus offset instructing them to ‘drop’ the
presented item (i.e. that no recall test would be run with that particular item)
indicating that these signals were indeed results of working memory storage,
and not only lingering effects of stimulus encoding (see also D'Esposito &
Postle, 2015).
This finding supports a link between the precision of the distributed
neural representation maintained during working memory storage and the pre-
cision of the mental representation that is assumed to support the recall per-
formance (Postle, 2015). Although whether the content of this mental repres-
entation was conscious or not was not directly evaluated in the original study,
keeping a target orientation in mind is a conscious effort and working memory
representations are typically considered to be conscious (for work on uncon-

10
scious working memory representations see e.g. Bergström & Eriksson, 2018;
King, Pescetelli, & Dehaene, 2016; Persuh, LaRock, & Berger, 2018). It is, thus,
a probable conclusion that higher precision neural representations maintained
during the delay period were accompanied by experiences with higher subject-
ive specificity in the precision sense.

5.3 Mental imagery

The parallel research domains of mental imagery and visual working memory
have recently been tightly linked by a study that found evidence of common
representations in early visual areas shared by mental imagery and visual work-
ing memory, and the subsequent interpretation that the phenomenon of mental
imagery is, in fact, the dynamic component of visual working memory (Albers
et al., 2013; Dijkstra et al., 2019; Tong, 2013).
Mental imagery studies, using VVIQ have established that at the trait level
— measuring how vivid someone’s mental imagery usually is — reported vivid-
ness correlated with the average of the relative visual cortex signal measured by
fMRI in visualisation tasks, i.e. that people who had more vivid visual imagery
showed higher activity in the visual cortex during imagery (Amedi, Malach, &
Pascual-Leone, 2005; Cui, Jeter, Yang, Montague, & Eagleman, 2007).
This association has further been reinforced at the level of trial-by-trial
variations (measuring how vivid a particular mental imagery experience is) as
well. It was found that subjective mental imagery vividness correlates with ‘im-
agery strength’, i.e. the priming effect of mental imagery on the dominant stim-
ulus in a succeeding binocular rivalry presentation (Bergmann, Genç, Kohler,
Singer, & Pearson, 2016; Pearson, Rademaker, & Tong, 2011; Rademaker &
Pearson, 2012), which can be seen as an indicator of the intensity of the neural
code underlying the imagined content (James, Humphrey, Gati, Menon, &
Goodale, 2000; Pearson & Brascamp, 2008; Pearson, Clifford, & Tong, 2008;
Wiggs & Martin, 1998).
Even more directly, in experimental paradigms where subjects had to rate
the vividness of their experienced mental imagery (on a 4 point scale: 1 - not
vivid at all; 4 - very vivid) after each trial of an imagery task it was found that
variations in the moment-to-moment experienced vividness of visual imagery
correlated with the intensity of the neural activity (strength of simultaneously
recorded fMRI signal) in a series of posterior brain regions. For instance, Dijk-
stra and colleagues (Dijkstra et al., 2017a) found a correlation between experi-
enced imagery vividness and increased activity in the early visual cortex, the
precuneus, the right parietal cortex, and the medial frontal cortex. Similarly,

11
Fulford and colleagues (Fulford et al., 2017) reported that subjective imagery
vividness judged image by image correlated positively with activations of the
precuneus, the posterior cingulate, and higher order visual association cortex.
As we have seen, these correlations suggest that in these mental imagery tasks
experienced vividness changes along the subjective intensity dimension.
In addition to these findings, it has also been reported that VVIQ mea-
sures of vividness correlate with the overlap between the activity in visual areas
during mental imagery and perception: bigger overlap between the activation of
the occipital cortex during imagery and perception predicts more vivid imagery
experience (Albers et al., 2013; Cui et al., 2007; Dijkstra et al., 2017a). Moreover,
studying directional connectivity shows that vividness modulates top-down
connectivity to early visual areas: the vividness of visual mental imagery posi-
tively correlates with the strength of top-down recruitment of early visual areas
(Dijkstra et al., 2019; Dijkstra, Zeidman, Ondobaka, van Gerven, & Friston,
2017b). These findings suggest that experienced vividness of mental imagery
changes along the subjective specificity (blurriness / detailedness) dimension as
well.
To summarise, recent efforts aiming at detecting the neural correlates of
visual imagery vividness seem to support the claim that VVIQ does indeed con-
flate the different factors — subjective intensity and subjective specificity — that
together determine the vividness of the experiences occurring during mental
imagery. By focusing on the distinct neural signatures of these different factors,
however, it becomes possible to provide a more fine-grained analysis of how
vividness changes in different conditions.

6. Using the neural signatures of vividness to form alternative

interpretations
Over and above providing access to a more fine-grained picture with regard to
the factors determining the vividness of experiences, the foregoing analysis also
allows us to shed new light on conscious experiences occurring during dreaming
and mind wandering.

6.1 The vividness of dreams

The visual qualities of dreams vary along similar dimensions that determine the
overall vividness of consciousness experiences. In a now classic strand of studies
dream experiences were investigated by using variations of a single photograph,
where each variation was reduced along one or more dimensions like bright-
ness, contrast, colour saturation, figure clarity, background clarity, overall hue,

12
etc. (Antrobus, Hartwig, Rosa, Reinsel, & Fein, 1987; Antrobus, Kondo, Rein-
sel, & Fein, 1995; Antrobus & Wamsley, 2009; Fosse, 2000; Kerr, 1993;
Rechtschaffen & Buchignani, 1983, 1992). Subjects had to find that element of a
set of manipulated photographs that best matched the appearance of a preceding
dream experience. It was found that REM dreams, in general, were more intense
(higher brightness and contrast) and more specific (higher clarity) than NREM
mentations (Antrobus, 1991; Kerr, 1993; Rechtschaffen & Buchignani, 1992).
Additionally, better quality dream experiences were reported from phasic than
from tonic periods of REM sleep (Foulkes & Pope, 1973; Kahn, Pace-Schott, &
Hobson, 1997; Molinari & Foulkes, 1969; Pivik, 1991; Rechtschaffen & Buchig-
nani, 1992). Importantly from our present perspective, it has also been implic-
ated that enhanced quality dream experiences were underlain by increased neur-
al activity (Antrobus et al., 1995; Nielsen, 2017; Rechtschaffen & Buchignani,
1992).
However, up until very recently, little has been known about the qualitat-
ive characteristics of individual dreams, or about the fine-grained changes of
such content characteristics as the vividness of dream experiences. An import-
ant example of the prevalence of categorical thinking and coarse-grained charac-
terisation in this regard is the case of so-called white dreams. When subjects re-
port white dreams they are certain that they had a dream prior to awakening
but are unable to recall any detail about these dream experiences (Cohen, 1972;
De Gennaro & Violani, 1990; Monday, Montplaisir, & Malo, 1987; Montplaisir,
Cote, Laverdiere, & St-Hilaire, 1985; Strauch & Meier, 1996). White dreaming is
an intriguing phenomenon so much the more as it is quite frequent: approxim-
ately 30% of post-awakening reports described white dreams (Cohen, 1972;
Siclari, LaRocque, Postle, & Tononi, 2013).
White dreams are traditionally interpreted as forgotten dreams due to
problems with the retrieval of dream experiences: subjects undergo ‘proper’
dream experiences, these experiences are then stored in memory, but then sub-
jects are unable to recall the content of these dreams because they cannot access
the stored memory traces (Cohen, 1974; Nir & Tononi, 2010). Alternatively, it
has recently been proposed that white dreams might be ‘contentless’ or ‘image-
less’ in the sense that subjects experience only a minimal form of conscious
presence with no narrative structure, and no specific percepts, bodily sensations
or thoughts occurring during such experiences (Windt, Nielsen, & Thompson,
2016).
A recent high dimensional EEG study (Siclari et al., 2017) directly ad-
dressing this question for the first time provided both content and neural activ-

13
ity specific data about dreaming with high spatial resolution. The study found a
difference between white dreams and dreams with recallable content in high-
frequency EEG activity over the medial and lateral frontal areas associated with
memory encoding. Consequently, the authors proposed that white dreams are
‘normal’ dream experiences (just like the reportable ones), but due to problems
with memory encoding their content is unavailable for the awakened subjects
(Siclari et al., 2017).
However, a closer reflection on the data revealed that the content-specific
neural activity occurring during dreaming was also different in the white dream
and normal ‘contentful’ dream conditions (Fazekas, Nemeth, & Overgaard,
2019). The content of dreams are underlain by neural activity in content-specific
posterior regions (Horikawa et al., 2013; Siclari et al., 2017; see also §2). In the
case of white dreams (compared to no dream experiences) the high-frequency
component of the EEG signal showed a local increase over posterior content
specific regions (similar to the ones indicated by the mental imagery studies of
Dijkstra et al., 2017a; and Fulford et al., 2017, see above), which was smaller
than the local increase that was found in the case of contentful dreams (Fazekas
et al., 2019). That is, white dreams turn out to be associated with an existing
(non-zero) posterior brain activity that is diminished in intensity compared to
the activity characteristic of contentful dreams. It follows that white dreams are
not really contentless (contra Windt et al., 2016), neither are they full-fledged
dream experiences that later become forgotten (contra Cohen, 1974; and Siclari
et al., 2017). Rather, on the basis of the link between subjective intensity and
the level of activity of associated brain areas (see §4.1), white dreams are experi-
ences with degraded vividness (quality) along the subjective intensity dimen-
sion. Dream experiences during white dreams, thus, are less vivid than during
contentful dreams. This reduced vividness results in lower meta-awareness,
which might be responsible for the inability to report these experiences
(Schooler, 2002).
The fact that both white and contentful dreams are reported from the
same sleep stages (Siclari et al., 2017; Siclari et al., 2013) indicates that the vivid-
ness of dreams fluctuates, i.e. changes on (at least) a dream-by-dream basis —
similarly to how mental imagery vividness changes on a trial-by-trial basis (see
§5.3).

6.2 Vividness during mind wandering

When the mind freely wanders — similarly to dreaming, and unlike in the case
of the intentional construction of mental imagery — the self-generated images

14
occurring in conscious experience can spontaneously change as the process of
image construction is less constrained (Christoff, Irving, Fox, Spreng, & An-
drews-Hanna, 2016). Instead of a top-down control characteristic of voluntary
mental imagery, and the typically bottom-up, salience driven nature of online
perception, the internal generation of the contents of mind wandering is under
the influence of a part of the so-called default mode network that increases its
activity in the absence of tasks or stimuli requiring externally oriented cognitive
efforts (Raichle, 2015; Raichle et al., 2001). According to a recent neural model
(Christoff et al., 2016), the source of the variability in the content of mind wan-
dering is a subsystem of the default mode network that includes the hippocam-
pal formation, parahippocampal cortex, retrosplenial cortex, ventral medial pre-
frontal cortex, and the posterior inferior parietal lobule.
Thought-like content is prevalent in mind wandering (Perogamvros et al.,
2017), yet a significant proportion of spontaneous mental activity unfolds in the
form of mental images (Andrews-Hanna et al., 2013; Chou et al., 2017; Delamil-
lieure et al., 2010). Although during mind wandering the primary sensory areas
are decoupled from the default mode network, which may underlie the relative
independence of this self-generated activity from perception (Schooler et al.,
2011), higher order sensory association areas are nevertheless very much active
supporting the image-like nature of the contents of the wandering mind (An-
drews-Hanna, Irving, Fox, Spreng, & Christoff, 2018; Fox, Nijeboer, So-
lomonova, Domhoff, & Christoff, 2013; Fox, Spreng, Ellamil, Andrews-Hanna,
& Christoff, 2015).
In a now classic study, Mason and colleagues could identify key brain
areas the activity of which correlated with the self-reported frequency of mind
wandering (Mason et al., 2007). Periods of high-incidence mind wandering were
established by training subjects on certain working memory tasks. fMRI-meas-
ured brain activity during performing practiced task sequences (high-incidence
condition) was contrasted with brain activity during performing similar but
novel tasks (low-incidence condition). This data-set was then compared to a
questionnaire-based score (Singer & Antrobus, 1972) of the subjects’ general
mind wandering proclivity. Mason and colleagues (Mason et al., 2007) found
that the frequency of mind wandering positively correlated with greater activity
in regions including the medial prefrontal cortex, the anterior cingulate, the
posterior cingulate, the precuneus, the left angular gyrus, the insula, and areas
in the superior and middle temporal gyri (for a recent confirmation of the in-
creased activity of these regions by a meta-analytic review, see Fox et al., 2015).

15
 According to the original interpretation, the increased neural activity in
these areas tracks with the subjects’ mind wandering frequency, i.e. with how
often subjects find themselves being engaged in mind wandering (Mason et al.,
2007). Note, however, that the areas implicated in the Mason et al. study include
regions that in mental imagery studies show increased activity with increased
imagery vividness (Dijkstra et al., 2017a; Fulford et al., 2017). Therefore, it
might be the case that the increased activity in the same regions occurring dur-
ing mind wandering, instead of being directly linked with the general frequency
of self-generated mental activity, is, in fact, indicative of the vividness — in the
sense of subjective intensity (see §4.1) — of the conscious content of mind wan-
dering.
Mason and colleagues already acknowledge that the questionnaire-based
measure of the frequency of mind wandering might assess subjects’ meta-aware-
ness regarding their mind wandering episodes rather than their propensity to
engage in such episodes. Tying the increased brain activity uncovered to the
vividness (subjective intensity) of the content of mind wandering is compatible
with this meta-awareness-linked interpretation, given that more vivid content is
more salient, captures attention to a greater extent, and thus elicits a higher level
of meta-awareness and is easier to remember (Schooler, 2002). This interpreta-
tion also fits better with the idea that mind-wandering is a psychological
baseline emerging when the brain is otherwise unoccupied (Mason et al., 2007),
as according to this understanding (as opposed to the frequency-based one), the
mind does return into this baseline state automatically — the difference is that
in the case of certain individuals the vividness (in the sense of subjective intens-
ity) of the accompanying experiences are usually higher, which results in more
robust memories about these states that are then reflected in their answers to
retrospective questionnaires.

7. Future directions
Shifting the focus from the neural correlates of consciousness per se to the cor-
relates of more fine-grained characteristics of experiences sets the course of fu-
ture research by suggesting novel research questions. In this paper we have pro-
posed hypothetical correspondence relations between phenomenological charac-
teristics and neural features. In many contexts these hypothetical correspond-
ence relations still need to be experimentally confirmed. Our discussion concen-
trated on the visual modality — the questions whether and how our hypotheses
can be extended to other modalities need to be addressed by future investiga-

16
tions. Important additional questions also arise and require further empirical
investigations and more extensive analysis.
For instance, if visual subjective specificity in the detailedness sense is as-
sociated with primary visual cortex involvement, then does the fact that mind
wandering is associated with sensory decoupling (attenuated sensory cortex
activity; Schooler et al., 2011; Smallwood, Beach, Schooler, & Handy, 2008)
mean that mind wandering experiences in general are less vivid in the specificity
(detailedness) sense than similar mental imagery experiences?
Relatedly, it has been reported that mind wandering frequency in indi-
viduals with Parkinson’s Disease who have hallucinatory experiences is strongly
associated with the coupling between primary visual cortex and dorsal default
mode network (Walpola et al., 2019). Is this an indication of more vivid (de-
tailed) phenomenology?
The vividness of hallucinations might be elevated in the sense of subjective
intensity as well. It has been claimed that both visual and auditory hallucina-
tions were underlain by the hyperactivation of the sensory cortices providing
the content of the hallucinatory experiences (Zmigrod & Hommel, 2011), and
that the sensory strength of mental imagery predicts the frequency of visual hal-
lucinations (Shine et al., 2015). The mental images formed during hallucinations
might thus be more vivid (both in the intensity and the specificity sense) than
during non-hallucinatory mental imagery. This might be a reason why indi-
viduals with hallucinations don’t question the reality of the contents of these
mental images (Stephan-Otto et al., 2017). The vividness of conscious experi-
ences, thus, might provide an important clue for metacognitive judgments about
whether the contents presented in the experience are real or not.

8. Conclusion
Once it is known where to look for the neural underpinnings of the contents of
conscious experiences, it becomes possible to start exploring potential corres-
pondence relations between features of these content-specific neural activity pat-
terns and certain phenomenological characteristics of subjective experiences. As
recent evidence converges on the claim that it is activity in posterior brain areas
that correlates with the contents of consciousness, here our aim was to try to
link features of this neural activity with the level of vividness with which differ-
ent contents of consciousness appear in experiences. Vividness is a common
characteristic of all experiences, regardless of their content. We have argued that
vividness, neither from a phenomenological perspective nor in the sense of
what existing psychological tools measure, is a monolithic phenomenon. We

17
have distinguished between two major components of vividness, subjective in-
tensity and subjective specificity, and identified further factors that determine
these major components: contrast, saturation and brightness in the case of in-
tensity, and precision, sharpness and detail in the case of specificity. At least
some of the fine-grained structure of vividness can be mapped onto distinct
neural activity patterns, i.e. some of the different factors that together determ-
ine the vividness of an experience have unique neural underpinnings. The
factors determining subjective intensity correlate with the strength of the under-
lying neural activity, whereas the factors determining subjective specificity cor-
relate with the precision of the corresponding neural population codes and with
the recruitment of early visual areas. We used these specific neural signatures to
shed new light on the more specific phenomenological counterparts of many
recent empirical findings about online perception, working memory mainten-
ance and mental imagery. These correspondence relations also helped us formu-
late new interpretations and draw novel conclusions with regard to those kinds
of experiences that are hard to access, like the ones occurring during dreaming
and mind wandering.

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