REVIEWS

The human imagination: the cognitive

neuroscience of visual mental imagery
Joel Pearson   
Abstract | Mental imagery can be advantageous, unnecessary and even clinically disruptive.
With methodological constraints now overcome, research has shown that visual imagery
involves a network of brain areas from the frontal cortex to sensory areas, overlapping with
the default mode network , and can function much like a weak version of afferent perception.
Imagery vividness and strength range from completely absent (aphantasia) to photo-​like
(hyperphantasia). Both the anatomy and function of the primary visual cortex are related
to visual imagery. The use of imagery as a tool has been linked to many compound cognitive
processes and imagery plays both symptomatic and mechanistic roles in neurological and
mental disorders and treatments.

School of Psychology,
The University of New South
Wales, Sydney, Australia.
e-​mail: jpearson@
unsw.edu.au
https://doi.org/10.1038/
s41583-019-0202-9
What is the human imagination? What is this amazing
ability, which most of us have, that allows us to travel
through space and time, testing out different virtual
worlds, objects, foods, fears and pleasures? When we
think about the sensory characteristics of something
like an apple or a sunset, most of us have a conscious
visual experience of those things. We literally become
conscious of some version of the apple or sunset, albeit
a degraded, fuzzy or weak experience.
Mental imagery has played a central role in discus­
sions of mental function for thousands of years, first by
philosophers, then by psychologists, and now by neuro­
scientists. Almost any behaviour or cognitive process
that might gain from sensory simulation tends to utilize
mental images.
However, to a minority of individuals, the idea that
people can have a voluntary conscious sensory experi­
ence in their mind’s eye comes as a complete surprise;
these individuals lack the ability to voluntarily form
mental images: aphantasia1. On the other side of the
spectrum, strong imagery plays a core role in many
anxiety disorders, depression, schizophrenia and
Parkinson disease, and is increasingly being harnessed
as a uniquely powerful tool for psychological treatment.
Mapping out imagery’s seemingly contradictory contri­
butions to human cognition, whereby imagery can be
both advantageous and clinically disruptive, or even
possibly unnecessary (aphantasia), offers exciting novel
insights into the human mind.
When people talk about the mind’s eye, they typi­
cally refer to the voluntary experience of creating a
conscious sensory experience at will. However, there are
many examples of involuntary sensory experiences that
are equally decoupled from direct sensory input. For
example, in synaesthesia and in many visual illusions,
individuals can experience vivid colour without colour
information stimulating the retina2. In post-​traumatic
stress disorder (PTSD), individuals experience flash­
backs or vivid, intrusive memories of trauma expe­
rienced as involuntary imagery 3. Such conscious,
involuntary sensory experiences, without a direct cor­
responding sensory input, have elsewhere been dubbed
‘phantom perceptions’2. One proposed overarching
framework is that internal experiences like imagery
can be divided into two types of imagery-​like experi­
ences, where one is voluntary and the other involuntary.
These two categories are analogous to the subtypes of
attention — endogenous and exogenous4 — and, as in
the early days of research into attention, current work
is just beginning to shed light on the commonalities
and differences between these two forms of phantom
perception2.
Importantly, mental imagery covers all five senses;
however, visual mental imagery research has tended to
dominate, as is also the case with perception research.
Focusing research on one faculty can make the endeav­
our more tractable and studying vision has several
advantages over studying the other senses. Humans
are visual creatures; this is clear when we look at the
proportion of cortical tissue assigned to processing
visual information as compared to the other senses.
Additionally, visual perception is vivid and full of
detailed information, often making visual stimuli
informative tools, as is the case with many illusions that
induce forms of involuntary phantom vision. However,
it is worth noting the limitations this focus on vision
has on our overall understanding of mental imagery
and it is perhaps too often assumed that findings
from visual imagery will naturally generalize to the
other senses.

Nature Reviews | Neuroscience

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Reverse directionality
Neural mechanisms of visual imagery
The reverse direction of neural Mental imagery involves activity across a large neural
information flow, for example, network spanning frontal areas right back to primary
from the top-​down, as opposed sensory areas. The neuroscience of imagery can be
to the bottom-​up.
divided into work looking at the mechanisms involved
in triggering the imagery process, mechanisms of gener­
ation or manipulation, as well as mechanisms underlying
the strength and vividness of visual imagery content and
its overlap with sensory perception. However, imagining
an apple, doing a mental rotation task or imagining per­
forming a sporting activity are three very different tasks
that too often get referred to simply as imagery. Much
work has been done investigating visual imagery in
particular, and the role of the primary visual cortex
(V1), for reasons relating to the imagery debate (Box 1).
Further, more recently, the large individual differences
in imagery strength and vividness have been harnessed
to obtain information about the neural mechanisms of
imagery. Here, a brief overview is given of the evidence
for the involvement of different neural areas and how
imagery is processed across the network of these areas.
Imagery creation: vision in reverse
It is hypothesized that voluntary mental imagery is based
on combinations of information retrieved from stored
memory. Accordingly, a very simple yet intuitive model
of functional voluntary imagery has been proposed as
depicted in Fig. 1: a reverse visual hierarchy5,6. In addi­
tion to the empirical evidence, this model also makes
intuitive sense as it is difficult to imagine content you
have never been exposed to and therefore have no mem­
ories of. Typically, in imagery generation, we are com­
bining different content that our senses have previously
been exposed to and is stored in our memory.
Box 1 | imagery debate
starting in the 1970s and running until the 2000s, a debate over the nature of mental
representation dominated mental imagery research21,134. the debate hinged on the
question of what formats the brain could represent information in. One side of the
debate argued that information about visual objects was stored in a symbolic, and even
language-​like, format135, whereas the other side argued that such information can be
stored in a number of different ways, including depictive formats. accordingly,
the debate largely focused on the two representational options: propositional and
depictive. a propositional format would be something similar to a descriptive format
used in a language. Depictive representations are often referred to as pictorial for
shorthand, as depictive representations require a functional space like a two-​dimensional
XY coordinate plane or picture (see ref.21 for examples).
throughout the 1970s and 1980s, various behavioural paradigms produced evidence
that imagery could be represented in a depictive format, only for the other side of the
debate to point out alternative interpretations of the data136,137. when neuroimaging
became available in the 1990s, the debate focused on showing a blood oxygenation
level-​dependent or positron emission tomography response to imagery in the primary
visual cortex, following the logic that the primary visual cortex represents visual
information depictively.
in 2015, Pearson and Kosslyn made the strong claim that the so-​called imagery debate
was over21 and the abundance of evidence that imagery can be depictive, while still
involving semantic information, is now overwhelming. evidence spans from behavioural
evidence that imagery is represented locally in retinotopic visual and orientation space
to functional magnetic resonance imaging voxel-​wise Gabor wavelet models trained on
depictive visual features (perceptual retinotopic location, spatial frequency and
orientation); see ref.21 for a full discussion. Few public debates have lasted so many
decades; this unique dialogue adds to the rich and often controversial history of
imagery research.
Due largely to the imagery debate (Box 1), the major­
ity of neural imaging work on mental imagery has
focused on early visual areas and the overlap or similar­
ities between visual imagery and visual perception. As
such, not much brain imaging work has investigated this
systemic model of voluntary imagery in operation as a
whole, though evidence from various different studies
does appear to support it.
Frontal areas. Forming or manipulating a mental image
does appear to involve activity in the frontal areas7–10.
However, the activity in frontal areas seems largely inde­
pendent of the precise content of the imagery9–11. This
suggests that frontal areas play more of a general organ­
izational or executive role in coordinating spatial and
sensory areas, rather than holding imagery representa­
tions or content per se. The frontal cortex shows up as
an active area in many functional magnetic resonance
imaging (fMRI) studies7,9–12. However, despite some
evidence of reverse directionality through the processing
hierarchy5,6, it remains unclear how the different areas
throughout the brain cooperate dynamically to achieve
the goal of an imagery experience. Part of the difficulty
in parsing out the isolated role of frontal areas is due
to the diversity of types of imagery tasks performed in
fMRI experiments. For example, imagining an apple,
doing a mental rotation task or imagining a physical
activity are three very different tasks that logically should
involve different brain areas, yet all get referred to under
the umbrella term ‘imagery’13,14.
Hippocampus. Some evidence suggests that the hippo­
campus might be required to form complex or spatially
distributed images. When individuals form such images,
a blood oxygen level-​dependent (BOLD) response has
been documented in the hippocampus15,16. Likewise,
individuals with hippocampal damage show spatially
related impairments in constructing imagined experi­
ences and their narrative descriptions lack richness and
spatial coherence17. When single neurons are measured
directly in humans, a small percentage of hippocampal
cells do respond to imagery18; however, other work has
failed to show a role for the hippocampus in imagery cre­
ation19,20. In sum, although probably involved in spatial
and memory elements of imagery, the exact role of the
hippocampus in imagery remains unclear.
Visual cortex. Questions about the role of the early visual
cortex, in particular V1, in mental imagery have domi­
nated brain imaging work on mental imagery for the last
25 years. From the early 1990s to late 2000s, over 20 studies
investigated the role of area V1 in imagery3. Most of
these studies were motivated by what is referred to as the
imagery debate, now generally considered to be over21
(Box 1). However, those studies were not without con­
troversy; although many did not find significant BOLD
responses above baseline in the early visual cortex10,22–27,
many others did13,28–38. In hindsight, these discrepancies
can be explained by inter-​study differences in imagery
tasks, imagery content (for example, a single visual fea­
ture versus a complex image), as well as by individual
differences in imagery strength or vividness3,35,39.
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Voxel-​wise models of
perception
Magnetic resonance imaging
and functional magnetic
resonance imaging decoding
methods that are constrained
by or based on individual voxel
responses to perception, which
are then used to decode
imagery.
Spatial transformations
Transformations in a spatial
domain.
Qualia
The conscious sense or feeling
of something, different from
detection.
Nature Reviews | Neuroscience
With the advent of multivariate decoding or pre­
dictive data analysis techniques in fMRI, there is now
clear evidence that the content of mental imagery can
be decoded from early visual areas, including V1 and V2
(refs38,40–42). It is interesting that this appears to be the
case despite relatively low BOLD amplitude responses.
The decoding algorithms used by these approaches can
be trained on visual perception, visual working mem­
ory content and, of course, imagery, and in all cases can
accurately decode the content of mental imagery40,42.
Further, when these algorithms are trained on perceptual
images and constrained by voxel-​wise models of perception
(the decoding model is based on perceptual information
such as contrast and spatial frequency), imagery can still
be accurately decoded, suggesting imagery in the early
visual cortex is composed of the same visual features as
afferent sensory perception42.
Taken together, the abovementioned brain imaging
work involving the visual cortex suggests there are pat­
terns of activity common to perception and imagery that
emerge as early as V1, but become increasingly similar
with ascension up the visual processing hierarchy3. This
visual hierarchy trend fits well into the overall model of
imagery dynamics (Fig. 1). If voluntary imagery is a top-​
down, triggered instance of sensory–memory recall that
uses the visual cortex (and depending on imagery content,
hippocampal and/or parietal areas), then it naturally fol­
lows that higher-​level visual areas (for example, visual area
3 and fusiform face area) that are physically and/or synap­
tically closer to the trigger source (frontal and medial tem­
poral lobe), would have stronger or more perception-​like
representations than more distant areas like V1 (Fig. 2).
Default mode network and imagery
The default mode network (DMN) is a networked group
of brain areas that regularly show up during periods of
non-​task rest, it is largely defined by the functional con­
nectivity or temporal relationships between activity in
spatially remote areas such as the posterior cingulate
cortex, dorsal medial prefrontal cortex and hippocam­
pus43. However, the DMN also shows similarities with
networks identified as being active during cognitively
demanding tasks44. Interestingly, the networks known
to be involved in reexperiencing the past or creating
possible future experiences overlap with the DMN45–48
and has become known for its association with such
processes. One of the few studies to look at conscious
manipulation, construction and destruction of mental
images showed that multiple areas, including the dorso­
lateral prefrontal cortex, posterior parietal cortex, pos­
terior precuneus and occipital cortex act together as a
network to construct and deconstruct abstract mental
images, that is, images that are not representations from
specific episodic memories7. In sum, evidence suggests
that the DMN is involved in imagery generation or con­
struction; however, because most of these studies have
investigated complex scenes from episodic memory or
future scenarios, it is hard to separate the relationships
between network elements and temporal and memory
components from the isolated act of forming the pure
sensory elements of a mental image. Although consider­
ing that many individuals are likely engaging in imagery
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1
3
2
Fig. 1 | A top-​down general model of voluntary mental
imagery: a reverse hierarchy. The initial mental action to
create a mental image ‘begins’ high in the cortical processing
hierarchy , in the frontal cortex (step 1). This triggers a
cascade of neural events, running ‘backwards’, that next
retrieves stored information or memories from more
posterior regions such as medial temporal areas (step 2)20,
and sensory and spatial representations of the imagery
content are then formed (steps 3). If the ‘requested’
representation involves movement and spatial locations,
then other areas like the middle temporal area and parietal
lobes would also be involved.
during a resting state scan, when they are typically asked
to relax and think about whatever they like, it should not
be surprising that the DMN relates to imagery. Future
work should compare the DMN with measures of pure
visual imagery of abstract features or objects and probe
mind-​wandering imagery content during these resting
scans to divulge the role of the DMN in imagery.
Ventral and dorsal streams
A common way to functionally divide the visual sys­
tem is division into the two processing streams that run
ventrally and dorsally. The ventral pathway is associated
with properties of objects and, when areas in this stream
are damaged, object perception and the ability to vis­
ualize shape are both disrupted. Damage to the dorsal
pathway, by contrast, disrupts the ability to visualize
locations or spatial transformations14,49. In sum, it seems
that the data support parallels between perceptual and
imagery processing regarding these two streams of visual
processing. Indeed, recent work on aphantasia (see later
section), suggests that spatial manipulations are normal
in individuals who have no object-​based imagery50.
Involuntary imagery
As mentioned above, there are other types of imagery
that are considered involuntary, for example, those trig­
gered by an association. These images are often referred
to as phantom perception2 as they involve sensory qualia
without concurrent or direct sensory stimulation.
Animal work has demonstrated that, as learning
occurs between two stimuli, sensory neurons in areas like
the middle temporal or inferior temporal cortex begin
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In sum, much like voluntary imagery, involuntary

Top-down images seem to involve the appropriate, correspon­
imagery signal ding sensory cortices. For example, involuntary motion
Imagery
imagery will involve the middle temporal area, whereas
involuntary colour imagery will involve V1 and V4.

Strong representations
(High-level brain areas)
More overlap What currently remains unknown, is the involvement,
between if any, of higher-​level, non-​sensory areas like the frontal
imagery and and parietal cortices (Fig. 3). Although it seems logical
perception
areas that these areas would not be involved in generating
involuntary imagery, we lack good datasets to show this.

Neural mechanisms of imagery strength

At least since the 1800s, people have wondered why

Fig. 2 | Graphical depiction showing the two streams — bottom-​up perception and
top-​down voluntary imagery. Voluntary imagery and perception have a greater overlap technique) (Box 2) has shown that the surface size of
in high-​level areas (dark blue) than in lower-​level areas (light blue). This graphical depiction
may well not hold for involuntary imagery.
Schizotypal personality
disorder
A mental disorder
characterized by social anxiety,
thought disorder, paranoid
ideation, derealization and
transient psychosis.
Strong representations
imagery differs so much from person to person. Now
Less overlap that the imagery debate is resolved21, the mechanisms
between of imagery, how an image is formed and why some
imagery and
perception are stronger than others (both inter-​individually and
areas intra-​individually) remains one of the primary research
Perception questions.
(Low-level brain areas) Recent work has begun to investigate the relation­
Bottom-up ship between V1 surface size and imagery character­
perception istics59. V1 surface size, like imagery, can vary between
individuals substantially60. Research using objective
perceptual measures of imagery (the binocular rivalry
both V1 and V2, but not V3, negatively predicts the
sensory strength of mental imagery59. In other words,
those with stronger imagery had smaller primary visual
to respond to an absent stimulus as if it were actually cortices. Further, the binocular rivalry technique allows
pre­sent51–53. Nevertheless, despite these data suggesting imagery precision to be tested across a number of visual
that the animal might have a conscious imagery-​like features. In this study, the precision of visual imagery
experience of the absent stimulus, we simply do not know. was measured by the spread, or generalization, of
Related work with humans has demonstrated that imagery across both spatial orientation and retinotopic
colour memory of colour-​specific fruit (for example, an visual space. Both these measures of imagery precision
orange is typically orange; a banana is typically yellow) showed a positive predictive relationship with V1 sur­
can be decoded from the BOLD response in V1 when face size59, much like perception does61. Interestingly,
humans view grey-​scale images54. These brain-​imaging those with the strongest imagery and smallest V1s did
studies are supported by behavioural research suggest­ not have the most precise imagery, suggesting that, as
ing we have a colour experience when viewing such imagery becomes stronger, it also becomes less precise.
grey-​scale colour-​specific images55 (Fig. 3). Other work This relationship between more vivid and stronger
has demonstrated that the perceptual ‘filling in’ of low-​ imagery and a smaller visual cortex is also an emerg­
contrast moving texture patterns leads to activity in V1 ing trend in the clinical literature. Human brain his­
and secondary V2 (ref.56) and, likewise, colour filling or tology shows that the size of V1 and its total number
‘neon colour spreading’ is associated with greater activity of neurons are reduced in schizophrenia compared
in V1 for the corresponding regions of the visual field to normal healthy control brains by around 25%62.
where the illusion is experienced57. This is compared to the overall brain size reduction
Prior expectations about an upcoming visual stim­ in patients with schizophrenia of only around 2%.
ulus can have a strong effect on subsequent sensory Individuals with schizophrenia show enhanced self-​
perception; indeed, fMRI work has demonstrated that reported vividness of mental imagery63 and individuals
a sensory template of the upcoming stimulus is for­ with schizotypal personality disorder show stronger sen­
med in V1 even when the perceptual stimulus is never sory imagery in the objective binocular rivalry imagery
presented58. Interestingly, it remains unknown how paradigm compared to controls64. Likewise, in PTSD,
voluntary or involuntary such a representation is; are patients who report more vivid voluntary self-​initiated
individuals taking up a strategy of using voluntary imagery65 also show signs of reduced visual cortex
imagery to aid the upcoming perceptual task? Or are size66,67. Furthermore, stimulant-​dependent individuals
these representations purely reflexive and involuntary? who demonstrate strong involuntary imagery that plays
Are these representations conscious? New work suggests an inductive role in substance dependencies68 also show
that images can form prior to conscious voluntary effort, a reduction in grey matter around the visual cortex69.
suggesting the existance of non-​conscious involuntary It is worth noting, however, that the link between
images 41
. Fig. 3 shows a proposed overview of some of imagery strength and V1 size is correlational, and so,
the types of imagery and their possible overlap in the despite this relationship showing up across both clinical
visual cortex. and non-​clinical populations, it does not clearly specify a

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functional mechanism. Interestingly, the data suggest a
reciprocal relationship between V1 and frontal cortex
size, with a smaller V1 predicting larger frontal areas70.
If frontal areas are responsible for the top-​down imagery
signal, then a scenario with larger frontal areas and
smaller V1 might enable more control over the visual
cortex (Fig. 4). Moreover, V1 surface area is influenced,
in part, by genetic differences71–73, suggesting a possible
indirect genetic contribution to imagery strength.
In addition to the anatomy of early visual cortex, evi­
dence suggests that cortical activity metrics also predict
imagery vividness35,38. Further, recent behavioural work
has shown that imagery vividness is not a static trait-​
like characteristic of imagery, but dynamically changes
from moment to moment within an individual59,74–76, and
these changes are predicted by a network of activity over
the whole brain, including the degree of overlap with
perception in the visual system38.
Some recent data suggest that the excitability of the
visual cortex might play a role in imagery strength77.
This work shows correlational evidence that uses trans­
cranial magnetic stimulation-​induced phosphenes and
fMRI resting activity as proxies of cortical excitability.
This study shows that causative transcranial direct cur­
rent stimulation, which alters resting excitability, had a
weak effect on imagery strength by either up or down­
regulating it77. These multiple data sources suggest that
stronger visual imagery is associated with lower resting
activity in the visual cortex. These data, although still
preliminary, suggest that imagery, along with other expe­
riences of phantom vision, like synaesthesia78 (which
itself is linked to stronger imagery79), might depend on
visual cortex excitability.
Association-induced image
Voluntary Involuntary CS Imagery of US
(associative)
=
Top-down Top-down
Lateral
Involuntary
Overlap? (perceptual)
Sensory cortex
representations
Fig. 3 | Mapping out the different types of visual imagery — voluntary , involuntary
(associative) and involuntary (local perceptual). The involuntary associative imagery
would be the product of associative learning. For example, after thousands of co-​occurrences
seeing tomatoes as red, a black and white image of a tomato is seen as slightly red due to
red involuntary imagery; your brain literally fills in the colour. Likewise, the classic case of
conditioning with Pavlov’s dogs, does the sound of the ringing bell induce an image of the
food in the dog’s mind? Here, the speaker symbol, the conditioned stimulus (CS) illustrates
this sound-​induced involuntary image, the unconditioned stimulus (US). Perceptual
involuntary imagery , or phantom vision, is when the structure of low-​level perceptual brain
areas produces involuntary vision like the colour spreading in neon-​colour spreading illusion.
Adapted with permission from ref.2, Elsevier.
Nature Reviews | Neuroscience
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One possible way to conceptualize the imagery net­
work and characteristics of V1, is that, at least in terms
of visual imagery, the visual cortex is something akin to
a ‘representational blackboard’ that can form representa­
tions from either the bottom-​up or top-​down inputs77.
The brain is not in a silent state waiting for sensory
stimulation, but always active, showing intrinsic activ­
ity dynamics, which emerge as spontaneous or ongoing
activity80. The degree of this ongoing activity noise in this
‘representational blackboard’ will impact imagery, and so
too will the strength of the top-​down signal (analogous
to the strength or contrast of ‘chalk’ in this blackboard
metaphor). A given individual might have low levels of
noise activity in V1, but a weak top-​down signal, and
therefore the overall degree of imagery strength, might
be weak. Another individual, by contrast, might have a
very strong top-​down signal, but their visual cortex is
much noisier, hence the imagery representation becomes
corrupted (Fig. 4). In this scenario, an individual with
both a strong top-​down signal and low noise, or quieter
sensory cortex, should have the strongest imagery (bot­
tom left Fig. 4). Presently, this is only a theory, and there
are many other clues suggesting imagery might have a
complex multi-​component mechanism, as it is clear that
imagery can be affected by many other factors such as
memory, hormones81, multiple neurological and mental
disorders, and even psychedelics82.
Imagery, like a weak form of perception
The conscious experience of imagining and perceiving
something are clearly very different for almost everyone.
However, evidence from the past 100 years of behavioural
research suggests that visual imagery can have a func­
tional effect on sensory processing akin to a weak form of
visual perception3. For example, both imagining and per­
ceiving oriented lines can have a similar effect on subse­
quent perception83,84. Imagery content can also undergo
different types of learning. Imagining visual content, just
like perceptual content, can induce visual per­ceptual
learning, which improves subsequent visual sensitivity to
those specific types of stimuli85. Likewise, the content of
visual imagery can undergo associative learning with an
emotional stimulus that transfers to perceptual stimuli in
a manner specific to early visual processing86. In addition,
the brightness of imagined stimuli corresponds to pre­
dictable changes in pupil diameter, without concurrent
sensory stimulation87.
There is now substantive evidence that visual imagery
can have a facilitative effect on visual perception of a
relevant visual stimulus74,76,83,84. Indeed, even when
imagery and perception are separated in time by another
demanding cognitive task, these facilitatory effects
remain83. Weak perceptual stimuli (low contrast or lumi­
nance) also show this same facilitatory priming effect
on subsequent perception83,88,89. The energy (strength or
duration) of a prior stimulus seems to dictate whether
that prior stimulus has a facilitatory (priming for weak
images) or suppressive (adaptation for strong images)
effect on subsequent perception. Likewise, longer
durations, or epochs, of imagery have a stronger prim­
ing effect on subsequent perception41,74,83. Hence, the
degree to which imagery affects subsequent perception
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has been taken as a measure of the sensory strength or
visual energy of imagery76 (see Box 2 on measurement
techniques). Just like visual perception, visual imagery
seems to be largely local in retinotopic and orientation
space42,59,79,83,90,91, lending further support to the similarities
between weak visual perception and visual imagery.
A criticism of some imagery research was the lack of
methodology to differentiate visual imagery and visual
attention. For example, if an individual imagines a par­
ticular colour while they are performing a perceptual
task involving colour, they will be differentially attending
to the colour they imagine. Hence, any effects of imagery
and attention will be difficult to separate. Importantly,
in some of this research, the effect of prior imagery and
prior visual attention can be dissociated in a number of
ways such as through the timing of these effects and their
susceptibility to sensory disruption83; this provides fur­
ther evidence that these sensory-​like effects are driven by
imagery and not attention. In addition, these perceptual
priming effects are linked to the reported imagery vivid­
ness on a trial-​by-trial basis74, suggesting the effects are
coupled to the imagery generation process.
Brain imaging work also provides evidence that
imagery can be thought of as a form of top-​down weak
perception. Multiple studies have now used BOLD acti­
vation patterns during afferent visual perception to later
decode the contents of mental images42,91,92, suggesting
an overlap in neural representation. Further, deep learn­
ing algorithms trained on a limited set of perceptual fea­
tures can be used to decode untrained features in mental
Box 2 | New measurement techniques
Like dreaming or hallucinations, research into mental imagery has been constrained by
a lack of reliable objective measurement methods. From the early use of questionnaires
by Galton in the 1800s110, self-​report questionnaires have been the gold standard used
to measure imagery. However, recent years have seen a flurry of new and more objective
measurement techniques.
Behavioural techniques
in 2008, we published the first work showing how you could use a visual illusion, called
binocular rivalry, to measure imagery. this technique involves randomly cuing an
individual to imagine a coloured object before a brief rivalry presentation. the content
of the imagery primes dominance in the rivalry illusion in a manner that reflects the
imagery vividness both on a trial-​by-trial and individual differences basis74,83. importantly,
this technique works even when a cognitively demanding task is added between the
imagery period and the rivalry presentation. this technique can be made even more
objective, in terms of removing direct subjective reports, by measuring rivalry
dominance with an embedded probe task138. In this case, rivalry dominance is assessed
using an embedded probe task, where the probe is easier to detect when it is
embedded in the dominant rivalry pattern, and then the influence of imagery on rivalry
can be determined by the level of probe detection. using such a performance-​based
dependent variable instead of rivalry dominance reports means that imagery can be
measured without any subjective reports. in addition, such methods can be used to
measure imagery of different visual features such as pure colour or motion imagery, or
even complex object imagery, using the one dependent measure90,133,138, or even
compare imagery and synaesthesia using the one common measure79.
imaging techniques
recent functional magnetic resonance imaging research has clearly shown that
imagery content can be objectively investigated using brain imaging40,42,139. what
someone imagines can be decoded or predicted based only on the brain’s response to
the afferent perceptual versions of that same content. in fact, imagery content can now
be decoded with perceptual training data from different object classes93, and other
methods like electroencephalography and pupil dynamics can also be used.
imagery93, and takes researchers one step closer to
being able to recreate the sensory contents of the mind.
However, despite the evidence suggesting commonali­
ties between weak perception and imagery, the two are
clearly very different conscious experiences and future
work should aim to map out why non-​clinical imagery
is not experienced with the full conscious experience of
afferent perception.
Imagery in normal cognitive function
Recent years have produced many publications linking
mental imagery to a range of cognitive processes from
episodic and visual working memory, spatial navigation,
reading comprehension to creativity and moral decision-​
making (Fig. 5). If mental imagery is a sensory simula­
tion of events, objects or scenarios in isolation from the
afferent sense organ-​induced versions of these things,
then how and when do we use imagery in everyday life?
Memory and imagery
Despite the fact that the fields of mental imagery, visual
working and autobiographical episodic memory have
developed independently, evidence suggests that many
people utilize imagery to perform most forms of visual
memory tasks94–97. Indeed, in the episodic memory liter­
ature, thinking about the future is referred to as imagin­
ing the future. Depending on who you are and probably
how strong your imagery is, the role of imagery across
different types of memory might seem obvious and self-​
evident. However, the fields of imagery and working
memory have diverged and until recently have remained
largely separate98.
In regard to visual working memory, when people are
asked how they complete such tasks, they typically report
using two primary strategies. One strategy is to pick
out salient features in a test array that the person is
required to remember and encode them in a pro­positio­
nal or phonological form, which is then com­pared to
the test99. The other strategy commonly des­cribed is the
creation of detailed mental images during the retention
interval, which are then compared to the sub­sequent
test arrays94,95,99,100. These descriptions are synonymous
with descriptions of mental imagery and it is likely that
these individuals use mental imagery as a mnemonic
tool to retain visual information. Both behavioural and
brain imaging work now suggests that sensory imagery
representations are used by people who have imagery to
complete some forms of visual working memory
tasks40,94,95,101. Additionally, both imagery and visual
working memory can change perception83,102, and they
show similar capacity functions103,104. However, research
also suggests that not all people use visual imagery
to solve visual working memory tasks, with irrele­
vant visual information impairing only some partic­
ipant’s performance on visual working memory tasks.
Specifically, only participants with good imagery appear
to be affected by irrelevant visual information94,95,105, sug­
gesting that only they use visual imagery and low-​level
visual regions to perform these tasks. The idea that only
good imagers use visual imagery, and hence the visual
cortex, to perform visual working memory tasks may
help explain much of the current controversy in both the
www.nature.com/nrn

Low
Noise level in visual cortex
High
Low
Weakest imagery
Top-down signal strength
Strongest imagery
High
Fig. 4 | Theoretical representation of visual imagery of a square, showing possible
interaction between the strength of the top-​down imagery signal and noise in the
visual cortex. A strong top-​down signal and low noise (bottom left) gives the strongest
mental image (square), whereas a high level of neural noise and a weak top-​down
imagery signal would produce the weakest imagery experience (top right).
visual working memory modelling and neuroimaging
literature105.
Likewise, to anyone who has good imagery it is
immediately obvious that imagery would be an impor­
tant experiential tool when referencing autobiographi­
cal memories or future projections. When I think of my
birthday last year, visual flashes of people sitting around
a table sharing food and chatting involuntarily flash
through my mind. Imagery has long been discussed as a
crucial element to autobiographical thinking. However,
the inherent difficulties in measuring imagery are per­
haps dwarfed by the very personal, bespoke and complex
phenomenology involved in autobiographical memory
measurement. Needless to say, investigating the role of
imagery in episodic memory brings many methodological
challenges.
Despite a primary reliance on self-​report techniques,
research suggests that individuals with vivid imagery do
re-​experience episodes from their life via memory dif­
ferently to those with weak imagery106–109. Data suggest
that individuals with vivid imagery report more sensory
details both in episodic and future event construction108,
and also rate their experience of reliving the event as
more real than those with poor imagery106, while more
recent work has shown that object and spatial imagery
predict different dimensions of episodic memory96,109.
Interestingly, people without any imagery (aphantasics),
can still perform visual working memory tasks and seem
to have episodic memories of their lives (see section
below on aphantasia). Hence, it would seem that using
imagery for such compound cognitive tasks is just one of
many available strategies, underpinning the importance
of probing the particular strategy an individual might
use for memory tasks.
Nature Reviews | Neuroscience
Reviews
Why does it matter that imagery is utilized across
such a large range of cognitive processes? We have
known since Francis Galton’s seminal paper on mental
imagery in 1880 that the spectrum of imagery vivid­
ness spans from aphantasia to hyperphantasia (highly
vivid, almost realistic imagery)110. Accordingly, if many
cognitive processes use imagery, but imagery can vary
so much across the population, then it follows that the
strength of imagery will have a strong knock-​on effect for
how someone performs any dependent tasks. Anything
that might affect imagery strength might also have an
effect on visual working and episodic memory, reading
comprehension, the veracity of eye witness memory,
moral decisions and maybe even creativity, not to men­
tion dimensions of mental health (Fig. 5). Therefore, one
cannot fully understand any of these dependent com­
pound cognitive processes without taking into account
or understanding the variations in mental imagery
strength. Accordingly, understanding imagery and meas­
uring it accurately holds the potential key to unlocking
the mysteries of multiple cognitive processes105.
Mental and neurological disorders
Mental imagery plays a major role in much of psycho­
pathology, perhaps unsurprisingly given experimental
findings that visual thought produces more emotion than
verbal thought111. Intrusive, affect-​laden mental imagery
causes functional distress across a range of mood dis­
orders, for example, by subserving ruminative and worry
states112. In depression, individuals have difficulty form­
ing positive future imagery113 and imagining suicidal acts
may increase the risk of suicide114. Patients with bipolar
disorder show high spontaneous use of imagery and
intrusive imagery115,116, and future-​oriented imagery
‘flash forwards’ to suicidal acts115. Involuntary intru­
sive visual images are a key component of cravings in
addiction117–119 and of flashbacks in PTSD. The realness,
or ‘nowness’, of intrusive images predicts the severity
of PTSD symptoms and outcomes120 and the ease with
which these intrusions can be triggered through viewing
related images predicts PTSD severity120. An individual’s
non-​traumatic visual imagery predicts PTSD sympto­
mology, with more vivid imagery being associated with
more frequent flashbacks65. Intervention research has
also suggested that disrupting the visuospatial sketch­
pad by playing the game Tetris after viewing traumatic
footage reduces subsequent intrusive imagery121. It has
been theorized that playing Tetris occupies the visual
areas of the brain (or the visuospatial sketchpad or rep­
resentational blackboard of the mind), and this impairs
an individual’s ability to replay the event during memory
consolidation.
Both schizophrenia and Parkinson disease have been
associated with stronger sensory imagery, measured
using the objective binocular rivalry technique64,122. In
fact, vivid imagery has been proposed as a trait maker
for schizophrenia63. As mentioned above, it is interesting
that both in the normal population and in schizophre­
nia, a smaller V1 is associated with stronger imagery.
In patients with Parkinson disease, the strength of an
individual’s visual imagery predicts the severity of their
visual hallucinations122.
Reviews
In addition to featuring alongside or as a symptom of
the disorder, visual imagery increasingly plays a role in
evidence-​based psychological treatment. For example,
cognitive behavioural therapy often includes ‘imaginal
exposure’, which involves the patient repeatedly imag­
ining a feared object or context until their anxiety level
subsides123. Imagery rescripting has been somewhat
successful in treating PTSD, social and snake phobias,
and a variety of other psychological disorders124. It typi­
cally involves the practice of rescripting the imagery in a
sequence to a more positive or adaptive outcome.
As the evidence grows that visual imagery can trigger
a larger emotional response than propositional or sym­
bolic thoughts125, the understanding of imagery both as a
symptom and as a strategic therapeutic treatment grows.
Clinical research should aim to leverage data and theory
from fundamental brain research on imagery as soon as
possible, as there are many promising avenues for uti­
lizing or manipulating imagery as a novel therapeutic
intervention.
Extreme differences in visual imagery
Aphantasia. Since Galton’s seminal paper in the 1800s110,
the idea of a category of people whose mind is com­
pletely blind was largely forgotten until recently1,126.
Initially based just on self-​report, it seemed that a
Prospective Eye witness
memory memory
Arithmetic
False memory
Navigation
Episodic Mental Visual working
memory imagery memory Moral decisions
Sports
Creativity
Motor control
Reading
Mind wandering
comprehension
Fig. 5 | Graphical depiction of the cognitive processes related to mental imagery.
The cognitive processes shown in the dark blue zone — visual working and episodic
memory — are there because reasonable evidence exists that, in people with mental
imagery (excluding people with aphantasia), they are largely reliant on mental imagery
for normal functioning and there is a ‘good’ amount of evidence to support this
relationship94,95. Evidence has shown that those with strong imagery will utilize it as a
mnemonic tool to perform visual working memory tasks94,95. This evidence is beyond
correlational and has used sensory perturbation methodologies (see section on
memory)94,95. Likewise, episodic memory research has specifically looked at the rich
sensory nature of remembered past experiences as well as the vividness of projecting
oneself into possible future episodic experiences96,109. Across a range of different
methodologies, effect sizes and degree of evidence, the cognitive processes shown in
the lighter blue zone have been hypothesized to involve mental imagery , although the
degree of evidence is less than that for the dark blue zone. The outer lighter blue zone
shows moral decisions140, eye witness memory141, false memories142, arithmetic,
navigation143, sports144, reading comprehension145, mind wandering146 and creativity147.
small proportion of otherwise healthy people report
having no visual experience at all when they attempt
to imagine something. In other words, their minds are
completely blind — no matter how hard they try, they
do not have a conscious visual experience of mental
imagery; this phenomenon has only recently been given
the name congenital aphantasia1. Initially, it was unclear
if aphantasia was more of an issue of imagery reporting,
in that people with aphantasia might still have func­
tional imagery, but not be aware of it or have a differ­
ent criterion for reporting it. However, recent research
goes beyond subjective self-​reports of aphantasia and
has shown that, along with floor scores on imagery
questionnaires, people with aphantasia score signifi­
cantly lower on the sensory binocular rivalry measure
of visual imagery50 (Box 2); in other words, the sensory
strength of their imagery is also lacking, suggesting that
aphantasia is more than poor metacognition of visual
imagery.
Interestingly, although people with aphantasia scored
lower than controls on all imagery vividness or content
measures, they actually scored slightly higher than con­
trols on spatial imagery questionnaires50. This suggests
that, although the ‘what’ or content of imagery typically
associated with the ventral stream of visual processing
is lost, the ‘where’, or spatial properties, associated with
dorsal areas might still be intact. It also suggests that
aphantasia could dissociate along the classic ‘what and
where’ processing pathways in the brain127. Accordingly,
areas of the brain that process spatial properties, includ­
ing the hippocampus, may not be the underlying cause
of aphantasia. Other work provides evidence that peo­
ple with aphantasia can perform easy and medium, but
not hard visual working memory tasks128. Likewise,
a case report of two people with aphantasia showed
that they scored low on ratings of ‘reliving’ their epi­
sodic memories, although they still reported believing
these memories106. Many individuals with aphantasia
describe visual imagery during dreams1, suggesting that
involuntary forms of imagery might still be intact.
Although the popular media has shown a fascination
with aphantasia, it is still early days in terms of scientific
research. Perhaps the most poignant question surrounds
the idea that aphantasia could either be just one end of a
spectrum of imagery abilities or it could be in a category
all of its own, different to people with very weak imagery.
Further research is needed to shed light on these two
possibilities.
The fact that individuals with aphantasia can still per­
form many daily tasks that non-​aphantasics use men­
tal imagery for, for example, visual working memory,
suggests they use a different strategy and brain mech­
anisms to do so. Presently, it is unknown what mem­
ory aids they use, perhaps semantic encoding or some
non-​representational geometry or symbols. It is worth
thinking about the possible implications of this, if one
was to simply go by performance on a compound task
like visual working memory, it would be very difficult to
differentiate a person with aphantasia and a person with
normal mental imagery. However, it is safe to say that
the underlying neural mechanism used to perform the
task by these two individuals would be quite different.
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 Reviews

Such a scenario underlines the importance of testing
imagery ability when investigating cognitive processes
like visual working or episodic memory. Further, this
suggests that using imagery as a cognitive tool is not
funda­mental to performing other cognitive functions
and has interesting implications for the way we should
theorize about imagery as a cognitive tool and its possible
role in evolution.
Hyperphantasia or eidetic imagery. Hyperphantasia,
or what has previously been called eidetic imagery, is
at the other end of the imagery spectrum — consisting
of strong and vivid imagery. Early in the 20th century,
the topic of eidetic imagery attracted much attention,
commonly described as a highly detailed, almost photo-​
realistic image experienced directly after seeing the
object, but also days or weeks later129–132. Eidetic imagery,
like aphantasia, is only observed in a small percentage
of the population, mainly in children, with frequency
estimates from 0–11% in children129. Unlike afterim­
ages, eidetic images remain still while the eye moves,
are experienced in positive colours, projected out into
space (much like a form of synaesthesia) and individuals
use present, not past tense language, when referring to
the image.
Eidetic imagery has been assessed in a number of
ways, from verbal descriptions to more objective meth­
ods such as showing individuals random-​dot stereo­
grams, pictures of hundreds of randomly placed dots,
with those in the centre shifted to give the perception
of depth when the two images are shown one to each
eye simultaneously; however, for eidetic imagers the
depth patch could be seen clearly, even when the two
images were shown up to 24 hours apart, by combin­
ing the first or eidetic image with the second perceptual
image130. Because modern day psychophysics, fMRI
and transcranial magnetic stimulation methods have
not been used to investigate eidetic imagery, we do not
have a lot of data to suggest a possible mechanism or
why it seems more prevalent in children. Hopefully,
as imagery research gains popularity, researchers will
reignite eidetic and hyperphantasic imagery research.
The future of imagery research
Now that the imagery debate is over21, and we have mul­
tiple new objective methods to investigate imagery, what
should imagery researchers focus on?
Visual perception is a constructive affair; it involves
both feedforward and feedback signals working closely
and interactively together. Hence, isolating the effects of
bottom-​up feedforward or feedback signals during per­
ception is very difficult. Voluntary imagery is perhaps
the only pure form of sensory representation that can
be solely due to feedback signals and can occur in com­
plete sensory isolation (although not always). Hence, it
represents a unique window to study the dynamics of
feedback signals in the brain and understand the con­
structive nature of visual perception — an important
step of reductionism to understand the mechanisms
behind sensory perception. As perception is interactive,
and hence cannot be understood without parsing out the
mechanisms of feedback, it can be argued that imagery
is the key to understanding normal visual perception. It
will be interesting to see how imagery research can be
harnessed to understand visual perception.
Several pertinent questions regarding imagery
remain, including the issue of why imagery rarely feels
as strong and vivid as afferent sensory perception; the
causes underlying aphantasia and the large range of
individual differences in the experience of imagery;
whether there exists a training protocol that might
improve imagery strength; whether imagery indeed
induces more activity in visual cortical areas, as opposed
to being just modulatory; and whether voluntary or
involuntary imagery could ever exist unconsciously133.
As many of the new methods to investigate imagery are
only newly available, it is exciting to think over all the
future imagery research waiting to be performed.
Published online xx xx xxxx

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Acknowledgements
The author thanks R. Keogh, R. Koenig-​Robert and A. Dawes
for helpful feedback and discussion on this paper. This paper,
and some of the work discussed in it, was supported by
Australian National Health and Medical Research Council
grants APP1024800, APP1046198 and APP1085404,
a Career Development Fellowship APP1049596 and an
Australian Research Council discovery project grant
DP140101560.
Competing interests
The author declares no competing interests.
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