c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7

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Single Case Report

Impaired spatial processing in visual perception,

imagery and art-making following parieto-occipital
infarcts

Jennifer A. Foley a,b,*, Isik Bayraktar e, Harpreet Hyare c,d and

Diana Caine a
a
Neuropsychology, National Hospital for Neurology and Neurosurgery, London, UK
b
UCL Institute of Neurology, London, UK
c
Department of Brain Repair and Rehabilitation, UCL Institute of Neurology, London, UK
d
Imaging Department, UCLH NHS Trust, London, UK
e
c/o National Hospital for Neurology & Neurosurgery, UK

article info abstract

Article history: Visual imagery, like vision as such, is widely thought to be supported by two distinct and
Received 14 June 2019 dissociable processing streams, dedicated to object representation and spatial analysis
Reviewed 30 September 2019 respectively. However, this simple dichotomy has been contested, with recent studies
Revised 27 November 2019 suggesting that impairments in perception-for-action and visuo-spatial imagery may
Accepted 22 January 2020 reflect a more general deficit in space-based attention. Although previous studies have
Published online 20 February 2020 revealed the impact of brain damage on artistic expression, few have examined the impact
on artistic expression in terms of the perceptual and spatial components of either visual
Keywords: processing or visual imagery. Here we present the case of an artist whose artistic
Visual perception expression was dramatically affected following devastating posterior brain damage. Of
Spatial processing particular interest, we demonstrate how these changes relate to impairments in inte-
Imagery grating and aligning different spatial features in both visual processing and visual imagery,
Simultanagnosia suggestive of a general simultanagnosia not previously described.
Art © 2020 Elsevier Ltd. All rights reserved.

contentious. Changes in artistic expression following brain

1. Introduction
Damage to posterior brain regions has been shown to disrupt
both visual processing and visual imagery, with evidence of
dissociable deficits in perceptual and spatial abilities,
although the precise nature of these deficits remains
injury can be exceptionally illuminating of deficits in cognitive
functioning and their neuroanatomical underpinnings
(Annoni, Devuyst, Carota, Bruggimann, & Bogousslavsky,
€ zner & Hennerici, 2007; Blanke & Pasqualini, 2012;
2005; Ba
zina, Botez, & Kaufman, 1985; Chatterjee,
Botez, Olivier, Ve

* Corresponding author. Department of Neuropsychology, Box 37, National Hospital for Neurology and Neurosurgery, Queen Square,
London, WC1N 3BG, UK.
E-mail address: Jennifer.Foley@ucl.ac.uk (J.A. Foley).
https://doi.org/10.1016/j.cortex.2020.01.023
0010-9452/© 2020 Elsevier Ltd. All rights reserved.
356 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7
2004; Cummings & Zarit, 1987; Finney & Heilman, 2007;
Kaczmarek, 1991; Mell, Howard, & Miller, 2003; Petcu et al.,
2016; Schnider, Regard, Benson, & Landis, 1993; Schott, 2012;
Sherwood, 2012; Smith, Mindelzun, & Miller, 2003; Wapner,
Judd, & Gardner, 1978). Here we describe recent de-
velopments in this debate, as prelude to reporting the case of a
talented young visual artist who suffered a devastating pos-
terior stroke. Through the dual lenses of neuropsychological
investigation and artistic expression, we explore the impact of
bilateral parieto-occipital infarcts upon perceptual and spatial
components of visual processing and visual imagery, and the
theoretical implication of these findings.
1.1. Perceptual and spatial visual cognition:
dissociations and interactions
While about half of all cerebrovascular accidents result in
some form of visual disorder (Rowe et al., 2009), damage to
posterior brain regions in particular can have devastating ef-
fects upon visual processing (Milner et al., 1991). Damage to
the primary visual cortex can result in complete blindness in
the corresponding region of the visual field (cf. Tong, 2003),
and insult to either of the two main pathways linking primary
visual cortex to adjacent areas can result in distinct and
dissociable higher-order visual impairments (cf. Milner &
Goodale, 2008). The precise nature of the processes under-
pinning these impairments remains a focus of significant
debate.
Early neuropsychological studies revealed two parallel
fibre bundles emerging from the occipital cortex. One was a
ventral pathway to inferior temporal areas associated with
visual perception, with damage causing deficits in shape and
colour recognition despite adequate visual acuity (Goodale
et al., 1994; Humphrey, Goodale, Jakobson, & Servos, 1994;
Milner et al., 1991). This finding was later supported by neu-
roimaging (Grill-Spector, 2003; Haxby, Hoffman, & Gobbini,
2000; Ishai, Ungerleider, Martin, Schouten, & Haxby, 1999;
James, Culham, Humphey, Milner, & Goodale, 2003; Malach
et al., 1995) and electrophysiological studies (Cohen, Cross,
Tunik, Grafton, & Culham, 2009; Lee et al., 2002).
The second, a dorsal pathway to the posterior parietal
cortex, was hypothesised as being specialised for spatial
processing, with damage resulting in visuo-spatial disorien-
tation and misreaching (Ungerleider & Mishkin, 1982). Later,
Goodale and Milner recast the dorsal pathway as underpin-
ning transformation of visual information for the control of
spatial action (Goodale, 2017; Goodale & Humphrey, 1998;
Goodale & Milner, 1992; James et al., 2003). Following bilat-
eral lesions in occipital and temporal regions, their patient DF
proved unable to describe or demonstrate the orientation of a
line or slot, but could reach out and post a card through the
same slot without any difficulty. In direct contrast, they
showed that patients with optic ataxia, following lesions to
the parietal regions, had intact visual perception, but made
spatially inaccurate reaches and/or hand rotations when
grasping for objects (Gre a et al., 2002; Jakobson, Archibald,
Carey, & Goodale, 1991; Milner, Dijkerman, McIntosh,
Rossetti, & Pisella, 2003; Pisella et al., 2000).
This apparent double dissociation led Goodale and Milner
to re-characterise the distinct outputs from ventral and dorsal
pathways, as vision-for-perception and vision-for-action
respectively (Goodale & Milner, 1992; Milner & Goodale,
1995). Support for a neural basis for this apparent
perception-action dissociation was amassed from psycho-
physics (e.g., Bridgeman, Lewis, Heit, & Nagle, 1979), neuro-
 goire,
imaging (Culham et al., 2003; Faillenot, Toni, Decety, Gre
& Jeannerod, 1997; Valyear, Culham, Sharif, Westwood, &
Goodale, 2006) and electrophysiology (Lee et al., 2002). This
version of the dual pathway account became the dominant
paradigm of visual processing and has wielded considerable
influence over the past 25 years.
Recently, however, this theoretical framework has been
challenged (Jackson et al., 2005b, 2009; Pisella et al., 2009;
Rosetti et al., 2003, 2017). Several studies have elicited evi-
dence of cross-talk between the two pathways (Pisella et al.,
2009; Rossetti et al., 2017). Moreover, Jackson and colleagues
have demonstrated that patients with optic ataxia are not
only impaired on tasks involving visually-guided action, but
also on all tasks that require comparison of two visual and/or
spatial representations (Jackson, Newport, Mort, & Husain,
2005a, b; 2009; 2018). This implies that the dorsal pathway
has a more general role in the simultaneous coordination of
multiple spatial representations and not just the execution of
visually-guided action. It is known that patients with optic
ataxia often demonstrate simultanagnosia, the inability
visually to perceive more than one object at the same time, but
recent research has shown that these patients are also
impaired on tasks that require matching of two objects, irre-
spective of whether the objects are presented visually or
haptically (Jackson et al., 2018). Whereas simultanagnosia is
thought to reflect a narrowed window of visual attention
(Dalrymple, Barton, & Kingstone, 2013; Rizzo & Robin, 1990),
the current findings hint rather at a supramodal deficit in the
ability to hold more than one representation in the mind's eye.
1.2. Visual imagery: are representational and spatial
components dissociable?
This raises important questions about the relationship be-
tween visual processing and visual imagery. There is certainly
evidence to support the Aristotelian perspective that visual
images are dependent upon visual perception, as if they were
‘residues of the actual [sense] impression’ (De Insomniis 461b;
cf. Rhetorica 137a 28), analogous to impressions in wax.
Perception and imagery paradigms are known to invoke
similar brain regions (Klein et al., 2004; Pearson, Naselaris,
Holmes, & Kosslyn, 2015; Slotnick, Thompson, & Kosslyn,
2005; Thirion et al., 2006), particularly the primary visual
cortex, and the vividness of visual imagery is thought to be
contingent upon this neural overlap (Diijkstra, Bosch & van
Gerven, 2017).
But there is also evidence to suggest that perception and
imagery are conceptually and phenomenologically distinct.
Several studies have reported intact imagery in the face of
significant perceptual disorder (Bartolomeo et al., 1998;
Behrmann, Winocur, & Moscovitch, 1992; Jankowiak,
Kinsbourne, Shalev, & Bachman, 1992) and congenital blind-
ness (Kerr, 1983), as well as the reverse: impaired imagery with
spared perception (Farah, Levine, & Calvanio, 1988; Moro,
Berlucchi, Lerch, Tomaiuolo, & Aglioti, 2008; Riddoch, 1990),
 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7
with evidence that the left temporal and parietal regions are
critically involved in the generation of images (Levine,
Warach, & Farah, 1985; Luzzatti, Vecchi, Agazzi, Cesa-
Bianchi, & Vergani, 1998; Papagno et al., 2002; Sack, Campro-
don, Pascual-Leone, & Goebel, 2005; Winlove et al., 2018).
Furthermore, there are reports of two distinct processing
streams for mental imagery, mirroring those for visual pro-
cessing (Cattaneo et al., 2008). Levine et al. (1985) reported
cases of impaired object imagery with intact spatial imagery
(Patient 1), and impaired spatial imagery with intact object
imagery (Patient 2), with similar dissociations reported by
other groups (Luzzatti et al., 1998; Papagno et al., 2002).
Accordingly, neuroimaging studies have found object imagery
to be associated with ventral occipito-temporal regions (Ishai,
Haxby, & Ungerleider, 2002, 2000; O'Craven & Kanwisher,
2000), and spatial imagery with the dorsal posterior parietal
cortex (Sack et al., 2005, 2008, 2002; Formisano et al., 2002;
Ganis, Thompson, & Kosslyn, 2004; Kawashima, Roland, &
O’Sullivan, 1995; Knauff, Kassubek, Mulack, & Greenlee,
2000; Kosslyn, Digirolamo, Thompson, & Alpert, 1998, 1997;
Thompson, Slotnick, Burrage, & Kosslyn, 2009). This would
suggest that although the generation of mental images relies
upon the left temporal areas, spatial manipulation of them
invokes parietal processes (Sack & Schuhmann, 2012).
1.3. Impact of brain injury on artistic expression
There have been several investigations of the visual cognitive
changes that impact upon artistic ability following brain
damage, mostly limited to unilateral brain insults (Annoni
€ zner & Hennerici, 2007; Blanke & Pasqualini,
et al., 2005; Ba
2012; Cantagallo & Della Sala, 1997; Chatterjee, 2004; Miller &
Hou, 2004; Schott, 2012; Sherwood, 2012; Smith et al., 2003).
Such reports have shown that left hemispheric damage can
disrupt semantic knowledge necessary for drawing objects
from memory (Gainotti, Silveri, Villa, & Caltagirone, 1983), as
well as the symbolic processing involved in the production of
conceptual visual art (Kaczmarek, 1991). Left hemisphere
damage may also release the functions of the right hemi-
sphere, facilitating more emotional and/or creative displays
€ zner & Hennerici, 2007; Mell et al., 2003;
(Annoni et al., 2005; Ba
Miller, Ponton, Benson, Cummings, & Mena, 1996; Miller &
Hou, 2004; Schnider et al., 1993). Right hemispheric damage,
on the other hand, can result in attentional deficits, mani-
festing as drawing omissions, distortions and perseverations
in patients with left-sided neglect (Blanke & Pasqualini, 2012;
Cantagallo & Della Sala, 1997).
Accounts of changed artistic production following poste-
rior brain damage are quite rare. Moreover, the few reports
available have failed to detail the neuropsychological profile,
with often limited or no interrogation of the perceptual and
spatial processes involved in visual processing and visual
imagery. For example, Wapner et al. (1978) discussed the case
of patient and artist JR who suffered bilateral occipital stroke.
Impaired at drawing from memory and poor at copying ob-
jects, JR would frequently lose his place, miss or misplace
features, or redraw parts. This led to erroneous copies, such as
a five-legged rhinoceros. The authors concluded that these
errors bore witness to his profound impairment in object and
spatial processing, but referred only to his performance on the
357
WAIS for corroborative neuropsychology. They also proposed
that JR demonstrated a general deficit in spatial imagery, cit-
ing difficulties matching faces from different perspectives and
describing familiar scenes. Without full neuropsychological
investigation, it is unclear if his deficits reflected impairment
in the generation of images, or rather their subsequent spatial
analysis.
Similarly, Smith et al. (2003) described an artist who
developed difficulties drawing following posterior stroke. Her
paintings became piecemeal, with difficulty centring. She had
difficulty representing a visual scene as a whole, tending to
focus on one part only which, they argued, was suggestive of
simultanagnosia or a deficit in the ability to create the image
in her mind's eye. Neuropsychological assessment was again
limited, here to a brief comment reporting a deficit in sus-
tained attention and constructional apraxia, with a notable
absence of tests of object perception, spatial processing or
mental imagery. Thus, again it remains unclear if her deficits
did indeed reflect an impairment in mental imagery, or spatial
analysis of the image in front of her.
Botez et al. (1985) did perform extensive neuropsychologi-
cal assessment of an amateur artist who complained of life-
long difficulties generating mental images. Born with hypo-
plasia of the right hemisphere, he was found to demonstrate a
striking deficit in the ability to image people, places and ob-
jects, with a corresponding difficulty drawing objects from
memory. However, visual perception was entirely intact and
his copies of the same objects were excellent. Although this
suggests that his drawing performance reflects limited ca-
pacity for imagery, they did not investigate object perception
and imagery comprehensively, omitting entirely any exami-
nation of spatial processing or spatial imagery. This again
precludes understanding of the precise nature of the imagery
deficits demonstrated.
In sum, although extant reports have revealed striking
changes in creative production following brain damage,
hitherto they have failed to address the impact of specific
perceptual and spatial failures upon visual artistic expression.
This study investigated patient IB who demonstrated signifi-
cant changes in both visual processing and imagery following
bilateral parieto-occipital infarcts. We report extensive,
detailed neuropsychology and the correlated and dramatic
changes in her visual art practice.
2. Methods
2.1. Case study
Patient IB is a 32-year-old, right-handed Graphic Designer.
Having attended an international school in her home country
in the Middle East, she came to the UK to attend art school and
following graduation had a flourishing career as a freelancer,
producing intricate patterns and designs for leading brands
(see Fig. 1 for example of her work).
At the age of 29 years, a prolonged episode of viral
myocarditis led to acute heart failure eventually requiring
urgent heart transplant. During this time, she was profoundly
acidotic and hypotensive for several hours, and ventilated
with central extra-corporeal membrane oxygenation. She
358 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7

Fig. 1 e Examples of art work before stroke.

suffered hypoxic-ischaemic encephalopathy, resulting in
watershed infarcts of the parieto-occipital lobes (see Fig. 2).
When she was woken up, she complained of bilateral
blurred vision. Ophthalmologic assessment revealed visual
acuity of 6/36 in each eye. She demonstrated bilateral altitu-
dinal and arcuate field loss, with linear demarcation on the
vertical midline. Eye movements and pupil reflexes were
normal, suggesting intact optic nerve function. She demon-
strated limb weakness, an ataxic gait and intention tremor
worse on the left compared to the right. She also developed
seizures. She was an inpatient for three months and
continued rehabilitation after discharge.
2.2. Materials and procedure
Following discharge from hospital, she underwent extensive
cognitive assessment. General intellectual functioning was
assessed using tasks from the Wechsler Adult Intelligence

Fig. 2 e MRI at the time of the ischaemic insult demonstrates absence of parenchymal signal abnormality or focal atrophy
(AeC). Five months later, there is mild widening of the intra-parietal sulcus bilaterally (D), widening of the calcarine sulcus
(E) and cortical signal abnormality affecting the frontal and parietal lobes bilaterally (F), right greater than left, consistent
with ischaemic injury. Follow up imaging at two years (GeI) demonstrates progression of the parietal and occipital lobe
atrophy (E,F), which remains stable at three years (JeL).
 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7
Scale e Third Edition (WAIS-III; (Wechsler, 1997), pro-rated to
generate scores for both verbal (VIQ) and non-verbal intel-
lectual abilities (PIQ). Further neuropsychological examina-
tion was as follows: verbal recognition and recall memory
(Words Recognition Memory Test short form [RMT;
Warrington, 1984]; Adult Memory and Information Processing
Battery Story subtest [AIMPB, Coughlan & Hollows, 1985]);
language (Oldfield Naming Test [(Oldfield and Wingfield,
1965)]; semantic fluency); visual processing (Cortical Visual
Screening Test [CORVIST; James, Plant, & Warrington, 1991;
Visual Object and Space Perception Battery [VOSP; Warrington
& James, 1991]); other parietal functions (Oral Graded Diffi-
culty Spelling [Baxter & Warrington, 1994]; arithmetic tests
[Jackson & Warrington, 1986]); executive functioning (phone-
mic fluency; Cognitive Estimates [MacPherson et al., 2014];
Hayling Sentence Completion Test [Burgess & Shallice, 1997]);
and speed of information processing (Counting Backwards
from 30).
IB also underwent tests of visual imagery, as described
below. On one of these, 11 healthy participants were also
asked to complete the test. The sample size for this control
group was determined through convenience and available
resources. In comparison to patient IB, these healthy partici-
pants were of a similar age (mean ¼ 31.9 years) and educa-
tional background (mean ¼ 18.5 years).
No part of the study procedures or analyses was pre-
registered prior to the research being conducted.
We report how we determined our sample size for the
control group, all data exclusions, all inclusion/exclusion
criteria, whether inclusion/exclusion criteria were established
prior to data analysis, all manipulations, and all measures in
the study.
3. Results
3.1. General cognitive assessment
Initially, five months following her stroke, cognitive assess-
ment revealed she was fully orientated to time, place and
person, and e remarkably - verbal memory recognition and
recall were intact. Language functioning was also well pre-
served. She did demonstrate significant executive dysfunction
(Cognitive Estimates), slow speed of processing (Counting 30
to 1) and poor arithmetic (Oral Graded Difficulty Arithmetic
Test). However, by far the most striking feature of her profile
was significant abnormalities in visual processing (see Table 1,
with impaired scores highlighted in bold).
3.2. Visual cognition immediately following stroke
While she had little difficulty on tests of basic visual acuity
(CORVIST Symbol Acuity) and visual sensory capacity (VOSP
Shape Detection), performance on tests of higher-order visual
processing was impaired.
She failed all trials assessing the ability to discriminate
subtle colour differences (CORVIST Hue Discrimination), and
she was poor at identifying shapes, fragmented letters and
numbers (CORVIST Shape Discrimination, CORVIST Shape
Detection, VOSP Incomplete Letters, CORVIST Fragmented
359
Numbers). Reading was poor, with frequent visual errors
(CORVIST Word Reading, e.g., ‘mouse’ for ‘museum’) and ev-
idence of visual crowding (CORVIST Crowding).
Similarly, on tests of visuo-spatial processing, she was
completely unable to discriminate between different sizes
(CORVIST Size Discrimination), make same-different judge-
ments of position (VOSP Position Discrimination), or count
dots on a page (CORVIST Scattered Dots, VOSP Dot Counting).
She was also unable to copy intersecting pentagons (Fig. 3).
She described her visual experience as, ‘when you lie
beneath a tree and look up to the sky, it's how you look though
the leaves'.
3.3. Visual cognition three years after stroke
Three years after her stroke, her vision and visual fields have
remained broadly stable although her subjective experience is
that there has been significant improvement. She continues to
demonstrate bilateral hemianopia, more dense in the right
hemifield. She is unable to count fingers in right upper, right
lower and left lower quadrants, but can in the left upper
quadrant. Automated perimetry confirms a stable pattern of
visual field loss, but with some minor improvement in that it
has become less dense and more patchy.
Repeat neuropsychological assessment revealed minor
improvement in visual perception (see Table 1). Performance
had improved on two tests of object perception (CORVIST
Shape Discrimination, Shape Detection), but remained
significantly impaired when discriminating between different
colours, and identifying fragmented letters and numbers.
There was also ongoing evidence of acquired dyslexia and
visual crowding. Performance also remained grossly impaired
on all tests of visuo-spatial processing.
When asked to describe a visual scene (Boston Diagnostic
Aphasia Examination Cookie Theft Picture, see Box 1), she
made several visual errors (such as describing the woman at
the sink as ‘washing a cat’). She also failed to describe the
overall scene (who was in the picture, where it was set), and
some of the salient features (e.g., the perilous stool). This,
along with her poor reading and visual crowding, suggests a
degree of simultanagnosia.
Box 1
IB's description of the Cookie Theft scene.
“I do not know if this is her husband or the little girl's
brother… I think the brother is taking a cookie from the
cupboard from a jar and handing it to his little sister…
while the mum is washing the dishes and the water is
spilling from the sink everywhere, but she doesn't seem
to be aware or maybe she is aware. She is wearing high
heels and curtains are open and I can see some trees
outside. Is she washing dishes or is she washing a cat?
She would not be washing the cat e I think she is
washing dishes, but there is something in the sink. What
is it? There is something in the sink, what is it? There is
something in the sink. I think it might be because of the
spillage it looks like an animal.”
360 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7

Table 1 e Performance on neuropsychological assessment five months and three years post-stroke.

Domain Test Five months post-stroke Three years post-stroke

General Intellectual Functioning WAIS VIQ 88 85
WAIS PIQ 60 63
Memory Short RMT Words 25/25 (>50th percentile) 25/25 (>50th percentile)
AMIPB Story Immediate 26/56 (10e25th percentile) 40/56 (50e75th percentile)
AMIPB Story Delayed 31/56 (25e50th percentile) 38/56 (50e75th percentile)
Naming Oldfield Naming Test 21/30 23/30
Visual CORVIST Symbol Acuity 36/36 36/36
CORVIST Shape Discrimination 3/4 4/4
CORVIST Size Discrimination 0/2 1/2
CORVIST Shape Detection 4/8 8/8
CORVIST Hue Discrimination 0/4 1/4
CORVIST Scattered Dot 1/4 1/4
CORVIST Fragmented Numbers 5/8 6/8
CORVIST Word Reading 12/16 14/16
CORVIST Face Perception 8/8 7/8
CORVIST Crowding 2/4 3/4
VOSP Shape Detection 18/20 (>5th percentile) 19/20 (>5th percentile)
VOSP Incomplete Letters 6/20 (<5th percentile) 15/20 (<5th percentile)
VOSP Dot Counting 2/10 (<5th percentile) 2/10 (<5th percentile)
VOSP Position Discrimination 7/20 (<5th percentile) 12/20 (<5th percentile)
Parietal Oral Graded Spelling 7/30 (7th percentile) 11/30 (12th percentile)
Oral Graded Arithmetic 2/24 (impaired) 5/24 (low average)
Executive Phonemic fluency ‘S’ 15 (31e34th percentile) 15 (31e34th percentile)
Category fluency - Animals 18 (25th percentile) 18 (25th percentile)
Cognitive Estimates 10 11
Hayling 13 (low average) 17 (average)
Speed Counting 30 > 1 26 s, 1 error 31 s

WAIS-III FSIQ: Wechsler Adult Intelligence Scale e Third Edition Full Scale IQ (Wechsler, 1997); RMT: Recognition Memory Test (Warrington,
1984); AMIPB: Adult Memory and Information Processing Battery (Coughlan & Hollows, 1985); Oldfield Naming Test (Oldfield & Wingfield,
1965); CORVIST: Cortical Vision Screening Test (James et al., 1991); VOSP: Visual Object and Space Perception Battery (Warrington & James,
1991); Oral Graded Difficulty Spelling Test (Baxter & Warrington, 1994); Oral Graded Difficulty Arithmetic Test (Jackson & Warrington, 1986);
Cognitive Estimates (MacPherson et al., 2014); Hayling: Hayling Sentence Completion Test (Burgess & Shallice, 1997).

On other tests, arithmetic has improved, but there remain
impairments on tests of executive function (Cognitive Esti-
mates) and speed of processing (Counting 30 to 1).
In sum, IB suffers gross deficits in visual processing, which
are confirmed upon testing. Extensive assessment revealed
impairment in visual perception and a particularly marked
deficit in visuo-spatial processing. Although she has also
sustained other cognitive deficits the impairments in visual
processing are by far the most marked and the most disabling.

3.4. Visual imagery three years after stroke

In contrast with the profound impairment in visual processing,

Fig. 3 e Copy of intersecting pentagons.
she denies any change in visual imagery. Three years after the
stroke, she reports that she is able to imagine and dream ob-
jects with correct form, size and colour. She was able to draw
clearly recognisable objects from memory, despite problems
with visuo-spatial aspects of the construction (see Fig. 4).
She also performed well on tests assessing object form and
size imagery (see Table 2). She was able to state correctly if a
given animal was larger or smaller than a goat, and if the ears
protruded upwards or downwards (Kosslyn, Farah, Holtzman,
& Gazzaniga, 1985). Performance was flawless on a test
assessing letter imagery: when asked if lower-case letters
contained an ascender (e.g., ‘b’, ‘h’), descender (e.g., ‘p’, ‘g’), or
neither (e.g., ‘m’, ‘e’; Weber & Castleman, 1970). Furthermore,
despite dyschromatopsia, she demonstrated intact colour
imagery. Following Riddoch and Humphreys (1987), perfor-
mance was also nearly flawless on a series of questions con-
cerning the colour of objects, both for those with strong verbal
 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7 361

Fig. 4 e Drawing from memory.

colour associations (e.g., ‘What is the colour of snow?’) and
those without (e.g., ‘What is the colour of a frog?’).
However, comparison of imagined faces was less robust.
As shown in Table 3, when asked to make judgements about
the similarity between three imagined faces (e.g., ‘Who looks
more like Bob Dylan: Johnny Depp or Arnold Schwarzeneg-
ger?’), she made correct judgements in 6/8 trials (Young,
Humphreys, Riddoch, Hellawell, & de Haan, 1994). Although
Table 2 e Performance on tests of visual imagery e form,
size and colour.
this was comparable to the scores of 11 healthy controls (see
supplementary file for raw scores), there were important
qualitative differences. Similar to patient HJA (Riddoch,
Humphreys, Gannon, Blott, & Jones, 1999), IB seemingly
drew upon piecemeal or semantic information in order to
make judgements. For example, she likened David Cameron
to Boris Johnson rather than Leonardo Di Caprio because ‘he
had become fat’.
With regard to navigation (see Table 4), we assessed her
ability to generate familiar and novel routes (as described by

Domain Test Score Description Table 3 e Performance of IB and 11 healthy controls on a

test involving comparison between three imagined faces.
Object and Animal size 18/18 Normal
letter imagery Animal ears 17/19 Normal Domain IB Control
Letters 12/12 Normal
Mean SD Range
Colour e associations 9/10 Normal
Colour e no associations 10/10 Normal Comparison of imagined faces 6/8 6.27 1.10 5e8
362 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7

Table 4 e Performance on tests of visual imagery e Art came to my help as a means of therapy. As my vision
navigation and mental rotation (with impaired scores was what I have been using the most and was mourning
highlighted in bold).
the lack of the most, I started with that. Drawing eyes
Domain Test Score Description was one of the first things I did. As couldn't see most of
Navigation Routes e familiar 2/2 Normal what I was drawing, not much was aligned. To begin
Routes e unfamiliar 0/2 Impaired with I could only see someone's eye and a little bit of
Mental Rotation Manikin Test e Total 15/32 Impaired their shoulder. So it could have been a world of single
Manikin Test e Upright 7/16 Impaired eyes floating in space if I hadn't known better. Some-
Manikin Test e Inverted 8/16 Impaired times I draw brains and symbols of electricity as an
attribute to the electrical malfunctioning in my brain. I
Burgess, Trinkler, King, Kennedy, & Cipolotti, 2006). She was should probably use more symbology for blood as it was
able to describe two familiar, remembered routes in great the lack of it in my brain that demolished my coordina-
detail (the bus journey she takes to the hospital in Queen tion and memory.
Square, and her walk to Queen Square from where she alights I have been told that I could recover substantially but
the bus), but was completely unable to generate new routes in they didn't know how long it would take or and they
familiar areas (such as the journey from Euston to Queen couldn't tell me the limitation of my recovery. All I could
Square, or from her house to King's Cross station). She also do was to wait the time to pass. Tried to solve puzzles, fill
performed at chance levels on a test of mental rotation in Sudoku and do crosswords unsuccessfully. I couldn't
(Manikin Test; Ratcliff, 1979). solve anything neither could use the tools that could
Thus, in contrast to her difficulties with both visuo- help me get around things and improve my neuro-
perceptual and visuo-spatial processing IB denies any connections. It was a Catch 22. So, I drew lines to sym-
change in imagery and indeed performs well on tests thought bolize the painfully slow passing of time.
to assess imagery of object form, size and colour. She did, I am starting to be happy. Depression is harmful to the
however, perform poorly on imagery tasks that require addi- brain. One cannot think clearly. Everything is distorted
tional spatial processing, generation of novel routes and and muggy. Being grateful saves oneself from falling into
spatial rotation of mental percepts and on tests involving self-pity. Neuroplasticity is something that I firmly
comparison of imagined faces. There is, therefore, a dissoci- believe in. After all, how much do we really know and
ation within the field of visual imagery, with imagery for understand the human brain? What do we truly know of
perceptual features other than faces relatively preserved, its capabilities?
while spatial imagery is impaired.

3.5. Artistic expression after stroke

In the first place, where there was relatively little repre-
sentation of the human form or features there has been since
There has been a dramatic change in her art practice and
her illness a preoccupation with face/eyes/brain, revealing
production. Prior to her stroke, she worked as a graphic artist
repetition of a different order. Where previously there was a
making digital art in a style that was linear and complex, with
focus on pattern now there is a preoccupation with the site
strong geometric organisation and often bold coloration. The
and impact of trauma, which began as an event but the effects
work was highly patterned, the pattern often repeating (see
of which recur moment by moment in her life; eyes which
Fig. 1). Since her stroke, she has continued with an undimin-
could see and through which she could imagine with a bril-
ished drive to create art, but now as a painter. The work she
liance now reduced in their capacity so that every plan or
has produced since her illness is strikingly different in several
intention, in life and in her work, is tethered by that limita-
ways (see Fig. 5 for examples and Box 2 for her description of
tion. Nevertheless, she makes art. Simpler in form, but at the
this).
same time, more communicative: images that represent her
Box 2 loss, and mourning, images of eyes that both can and cannot
IB's description of her life and artwork after stroke see what it is they are looking at. No longer made for a client,
the work is now suffused with emotion and meaning.
In the immediate aftermath of her illness there were
Upon exiting the hospital grounds, I was left with a vi-
repeated images of a face with eyes, with other features less
sual impairment; an ill-functioning brain and a poorly
distinct. At first the face appeared alone on the page, eyes
functioning right arm and pointing finger that I could not
sometimes weeping, sometimes in sombre colouring. Several
feel. That meant my career as a graphic designer was
paintings have scored out eyes, reflecting her loss of visual
over. This in itself was devastating enough. As I re-
abilities. The mood portrayed by the paintings made soon
learned how to walk, tie my shoelaces and wash
after her illness is bleak, reflecting suffering and despair. Over
myself, grew strong enough to lift a pint glass, I tried to
time colour and other elements began to appear. An occa-
live a normal life. I soon realized that I lost more than I
sional tree, a hint of sunlight as her mood changed. But also
thought. I had very little spatial awareness or sense of
tally marks, as if counting up the days ‘without vision’, like a
direction, was suffering from acute fatigue therefore
prison sentence but one that may or may not end. Sometimes
sleeping all the time and as a result, also started
eyes fill the page. The images are much simpler: the small and
suffering from major depression.
repeated patterns replaced now by fewer objects set upon
 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7
Fig. 5 e Examples of art work after stroke.
plainer backgrounds. A narrowed thematic range echoes the
kind of repeated preoccupation often seen in artists' work.
Here such changes may reflect the fronto-parietal brain
damage, observed to be important for artistic creativity (Shi,
Cao, Chen, Zhuang, & Qiu, 2017). More recently an element
of playfulness has entered the work, with brighter colours and
humour. The use of colour, is, though, reduced possibly
reflecting her deficits in colour discrimination, as corrobo-
rated by her neuropsychology.
As a result of her deficits, the materials have had to change.
Whereas her graphic design relied on digital tools to construct
intricate detailed patterns, she is now much less able to use a
computer both because of the difficulty successfully shifting
vision from keyboard to screen and back, and because of the
difficulties following and understanding computer software.
Now she draws and paints using media that she encountered
first during her childhood and early schooling and that were
offered to her once more during her inpatient stay: pencils,
watercolours and A4 paper.
Most strikingly, the images created after her illness betray
the visual processing deficits: items, including facial features,
misaligned in space. She reports particular difficulty aligning
eyes in space, in relation to one another and to the remainder
of the face Other facial elements are often absent if not mis-
placed just as in her drawings of objects from memory (Fig. 4):
363
the head above the tie, for instance, has too high a mouth.
There is also separation of individual elements; the owl's legs
are separate to its body, and the saw's serrated edge is sepa-
rated from its handle. This impairment in drawing from
memory is mirrored by her difficulty in copying, as demon-
strated in Fig. 3.
4. Discussion
4.1. Dissociation between representational and spatial
components of visual imagery
IB is a young woman for whom making art has always been
central to her life and who was on the cusp of a stellar career
as a graphic artist when she sustained bilateral watershed
infarcts, which have had a dramatic impact on her life, her
visual cognitive abilities and her art making. Formal neuro-
psychological investigation demonstrated impaired visual
perception and reduced capacity for visuospatial processing.
These impairments reflect damage to both the ventral and
dorsal pathways supporting higher-order visual function. In
contrast, we demonstrated a striking dissociation between
intact visual perceptual imagery for colours, shapes and let-
ters and impaired performance on all tests that required
364 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7
imagined spatial processing or manipulation, specifically, on
tasks involving novel route generation and mental rotation.
This discrepancy between object and spatial imagery supports
the claim that, like visual processing, mental visual imagery is
also supported by two distinct processing streams (Cattaneo
et al., 2008).
This dissociation between impaired object perception and
intact object imagery confirms earlier reports of distinguish-
able processes involved in the detection and generation of
visual images (Bartolomeo et al., 1998; Behrmann, Moscovitch,
& Winocur, 1994; Farah et al., 1988; Jankowiak et al., 1992;
Moro et al., 2008; Riddoch, 1990). Conversely, despite broadly
intact face perception, performance was impaired on the task
that required comparison of different imagined faces.
Although face analysis draws upon perception of basic and
largely invariant configurations (Haxby et al., 2000), compari-
son of different facial features across individuals demands a
greater degree of spatial attention and holistic representation
(Taubert & Alais, 2009). Recourse to reliance upon gross
perceptual features as opposed to configural information to
perform this task is indicative of her deficits in the spatial
processing of imagined facial features.
4.2. A deficit in space-based attention?
Our patient's visuo-spatial deficit affecting both the process-
ing of the visuospatial aspects of external stimuli and visuo-
spatial imagery is suggestive of a more general deficit in
space-based attention. It has been proposed that the
occipito-parietal regions may be crucially involved in the
simultaneous coordination of multiple spatial representations
(Jackson et al., 2005a, b; 2009; 2018). In keeping with findings
from optic ataxia patients, the data from our patient is also
suggestive of a marked deficit in holding more than one
spatial representation in the mind's eye.
Kosslyn's (1980) influential model of imagery proposed
three componential processes: generation, inspection and
transformation of an image within short-term memory. Our
patient seems to have intact generation, with satisfactory
recollection of object form, size and colour from long-term
memory. However, inspection appears impaired, with poor
processing of spatial features and incoherence of the overall
percept, precluding subsequent image transformation or
rotation (Sack & Schuhmann, 2012). Previous research has
suggested that whereas the left temporal and parietal cortices
are involved in the generation of images (Levine et al., 1985;
Luzzatti et al., 1998; Papagno et al., 2002; Sack et al., 2005),
the right parietal cortex is thought to be responsible for the
spatial comparison of such imagined content (Sack et al.,
2005). As our patient suffered damage to both parietal lobes,
we propose that this bilateral insult has resulted in her
marked disorder in spatial imagery.
It may be that our findings simply reflect differing levels of
task difficulty. Some studies have shown that vividness of
mental imagery is associated with the function of the V1
(Bergmann, Genç, Kohler, Singer, & Pearson, 2016; Dijkstra,
Bosch, & van Gerven, 2017): thus it is possible that although
our patient passes our tests of object imagery, her difficulty
generating visual images may be magnified in tasks of higher
cognitive load, such as mental rotation, leading to what ap-
pears to be a selective deficit in space-based attention. How-
ever, such an argument can be countered by the previous case
studies reporting the reverse pattern of deficits: impaired im-
agery with intact novel route generation (Levine & Calvanio,
1989; Levine et al., 1985). This apparent double dissociation
between mental imagery and mental spatial computations
demonstrates that these are separable processes, not simply
explained by levels of difficulty. Moreover, the notion that our
patient's observed deficits are simply a consequence of task
difficulty is a circular one. Tasks deemed more ‘difficult’ may be
considered so because of a greater attentional load. As we
propose our patient has significant impairment in spatial
attention, tasks drawing upon such reduced attentional re-
sources will therefore necessarily be more difficult.
Similarly, mental imagery is not reducible to just attention.
Thompson, Hsiao, and Kosslyn (2011) found that manipulating
the size of the stimuli differentially affects performance on
imagery and attentional tasks, confirming that these are also
dissociable processes. This is in line with our findings of pre-
served performance on simple tasks of mental imagery, but
impaired performance on tasks that place greater demands
upon attention, namely those that involve spatial processing.
4.3. Effect upon artistic expression
Unlike all previous case reports to date, we can also observe
how our patient's deficits are made manifest through her
artistic expression. Her deficit in space-based attention is
notably reflected in her visual art, with a change in style from
highly constrained and geometrically organised to much
simpler, more representational, and with misaligned ele-
ments. The combination of impaired spatial processing with
impaired object perception likely explains her severely
impaired copy of intersecting pentagons, whereas the execu-
tion of drawing objects from memory is perhaps relatively
more preserved because of her intact object imagery. How-
ever, the impaired spatial processing is still apparent, partic-
ularly in the misalignment of multiple elements and fewer
identical repeating objects.
Although previous reports have sought to illustrate the role
of impaired spatial processing in the production of visual art,
these have been limited by insufficient corroborative evidence
from neuropsychology (Smith et al., 2003; Wapner et al., 1978).
In our study, we have shown that our patient's impairments in
visuo-spatial processing and spatial imagery are clearly illu-
minated by her drawings, both when attempting to draw from
models and from memory.
4.4. Conclusion
We may conclude that our patient's deficits in integrating and
aligning different spatial features, as witnessed by both neu-
ropsychological assessment and artistic expression, illustrate
the presence of impaired simultaneous coordination of multi-
ple spatial representations. As this deficit is shared across
visuo-processing and visual imagery, it supports the recent
claims by Jackson et al. (2018) that occipito-parietal regions are
crucially involved in the ability to hold more than one spatial
 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7
representation in the mind's eye. Although simultanagnosia is
a concept currently limited to the deficient perception of two
simultaneous representations, the deficits discussed here
appear to reflect simultanagnosia for stimuli, either perceived
or imaginal, as explained by a severely limited spatial window
of attentional processing (Dalrymple et al., 2013).
Data availability statement
All data concerning patient IB's behavioural performance is
contained in the manuscript. The conditions of our medical
ethics do not permit public archiving of patient IB's anatom-
ical MRI scan, which cannot be shared with any individual
outside the author team under any circumstances.
Due to legal copyright restrictions, the test materials used
in this study are not publicly archived and can be obtained
from the copyright holders in the cited references.
Credit author statement
Jennifer A. Foley: Conceptualization; Data curation; Formal
analysis; Investigation; Methodology; Project administration;
Resources; Roles/Writing - original draft; Writing - review &
editing.
Isik Bayraktar: Data curation; Resources; Writing - review
& editing.
Harpreet Hyare: Data curation; Formal analysis; Writing -
review & editing.
Diana Caine: Conceptualization; Data curation; Investiga-
tion; Methodology; Project administration; Supervision; Visu-
alization; Roles/Writing - original draft; Writing - review &
editing.
Supplementary data
Supplementary data to this article can be found online at
https://doi.org/10.1016/j.cortex.2020.01.023.
references
Annoni, J. M., Devuyst, G., Carota, A., Bruggimann, L., &
Bogousslavsky, J. (2005). Changes in artistic style after minor
posterior stroke. Journal of Neurology, Neurosurgery & Psychiatry,
76, 797e803.
Bartolomeo, P., Bachoud-Le vi, A., De Gelder, B., Denes, G., Dalla
Barba, G., Brugieres, P., et al. (1998). Multiple-domain
dissociation between impaired visual perception and
preserved mental imagery in a patient with bilateral
extrastriate lesions. Neuropsychologia, 36, 239e249.
Baxter, D. M., & Warrington, E. K. (1994). Measuring dysgraphia: A
graded-difficulty spelling test. Behavioural Neurology, 7,
107e116.
€ zner, H., & Hennerici, M. G. (2007). Painting after right-
Ba
hemisphere stroke e case studies of professional artists.
Frontiers in Neurology and Neuroscience, 22, 1e13.
Behrmann, M., Moscovitch, M., & Winocur, G. (1994). Intact visual
imagery and impaired visual perception in a patient with
365
visual agnosia. Journal of Experimental Psychology: Human
Perception and Performance, 20, 636e637.
Behrmann, M., Winocur, G., & Moscovitch, M. (1992). Dissociation
between mental imagery and object recognition in a brain-
damaged patient. Nature, 359, 636e637.
Bergmann, J., Genç, E., Kohler, A., Singer, W., & Pearson, J. (2016).
Smaller primary visual cortex is associated with stronger, but
less precise mental imagery. Cerebral Cortex, 26, 3838e3850.
Blanke, O., & Pasqualini, I. (2012). The riddle of style changes in
the visual arts after interference with the right brain. Frontiers
in Human Neuroscience, 5, 154.
Botez, M. I., Olivier, M., Vezina, J., Botez, T., & Kaufman, B. (1985).
Defective revisualization: Dissociation between cognitive and
imagistic thought case report and short review of the
literature. Cortex, 21, 375e389.
Bridgeman, B., Lewis, S., Heit, G., & Nagle, M. (1979). Relation
between cognitive and motor-oriented systems of visual
position perception. Journal of Experimental Psychology: Human
Perception and Performance, 5, 692e700.
Burgess, P. W., & Shallice, T. (1997). The hayling and brixton tests.
Bury St Edmunds, UK: Thames Valley Test Company.
Burgess, N., Trinkler, I., King, J., Kennedy, A., & Cipolotti, L. (2006).
Impaired allocentric spatial memory underlying topographical
disorientation. Reviews in the Neurosciences, 17, 239e251.
Cantagallo, A., & Della Sala, S. (1997). Preserved insight in an
artist with extrapersonal spatial neglect. Cortex, 34,
163e189.
Cattaneo, Z., Vecchi, T., Cornoldi, C., Mammarella, I., Bonino, D.,
Ricciardi, E., et al. (2008). Imagery and spatial processes in
blindness and visual impairment. Neuroscience and
Biobehavioral Reviews, 32, 1346e1360.
Chatterjee, A. (2004). The neuropsychology of visual artistic
production. Neuropsychologia, 42, 1568e1583.
Cohen, N. R., Cross, E. S., Tunik, E., Grafton, S. T., & Culham, J. C.
(2009). Ventral and dorsal stream contributions to the online
control of immediate and delayed grasping: A TMS approach.
Neuropsychologia, 47, 1553e1562.
Coughlan, A. K., & Hollows, S. E. (1985). The adult memory and
information processing battery. Leeds, UK: St James’ University
Hospital.
Culham, J. C., Danckert, S. L., DeSouza, J. F. X., Gati, J. S.,
Menon, R. S., & Goodale, M. A. (2003). Visually-guided grasping
produces activation in dorsal but not ventral stream brain
areas. Experimental Brain Research, 153, 180e189.
Cummings, J. L., & Zarit, J. M. (1987). Probable Alzheimer's disease
in an artist. Journal of the American Medical Association, 258,
2731e2734.
Dalrymple, K. A., Barton, J. J. S., & Kingstone, A. (2013). A world
unglued: Simultanagnosia as a spatial restriction of attention.
Frontiers in Human Neuroscience, 7, 145.
Dijkstra, N., Bosch, S. E., & van Gerven, M. A. J. (2017). Vividness of
visual imagery depends of the neural overlap with perception
in visual areas. Journal of Neuroscience, 37, 1367e1373.
Faillenot, I., Toni, I., Decety, J., Gregoire, M. C., & Jeannerod, M.
(1997). Visual pathways for object-oriented action and object
recognition: Functional anatomy with PET. Cerebral Cortex, 7,
77e85.
Farah, M. J., Levine, D. N., & Calvanio, R. (1988). A case study of
mental imagery deficit. Brain and Cognition, 8, 147e164.
Finney, G. R., & Heilman, K. M. (2007). Artwork before and after
onset of progressive nonfluent aphasia. Cognitive and
Behavioural Neurology, 20, 7e10.
Formisano, E., Linden, D. J., Di Salle, F., Trojano, L., Esposito, F.,
Sack, A. T., et al. (2002). Tracking the mind's image in the
brain: W. Time-resolved fMRI during visuospatial imagery.
Neuron, 35, 185e194.
Gainotti, G., Silveri, M. C., Villa, G., & Caltagirone, C. (1983).
Drawing objects from memory in aphasia. Brain, 106, 613e622.
366 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7
Ganis, G., Thompson, W. L., & Kosslyn, S. M. (2004). Brain areas
underlying visual mental imagery and visual perception: An
fMRI study. Cognitive Brain Research, 20, 226e241.
Goodale, M. A. (2017). Duplex vision: Separate cortical pathways
for conscious perception and the action of control. In
S. Schneider, & M. Velmans (Eds.), The blackwell companion to
consciousness. Malden, MA: Blackwell.
Goodale, M. A., & Humphrey, G. K. (1998). The objects of action
and perception. Cognition, 67, 181e207.
Goodale, M. A., Meenan, J. P., Bülthoff, H. H., Nicolle, D. A.,
Murphy, K. J., & Racicot, C. I. (1994). Separate neural pathways
for the visual analysis of object shape in perception and
prehension. Current Biology, 4, 604e610.
Goodale, M. A., & Milner, A. D. (1992). Separate visual pathways
for perception and action. Trends in Neuroscience, 15, 20e25.
Grea, H., Pisella, L., Rossetti, Y., Desmurget, M., Tilikete, C.,
Grafton, S., et al. (2002). A lesion of the posterior parietal
cortex disrupts on-line adjustments during aiming
movements. Neuropsychologia, 40, 2471e2480.
Grill-Spector, K. (2003). The neural basis of object perception.
Current Opinions in Neurobiology, 13, 159e166.
Haxby, J. V., Hoffman, E. A., & Gobbini, M. I. (2000). The distributed
human neural system for face perception. Trends in Cognitive
Sciences, 4, 223e233.
Humphrey, G. K., Goodale, M. A., Jakobson, L. S., & Servos, P.
(1994). The role of surface information in object recognition:
Studies of a visual form agnosic and normal subjects.
Perception, 23, 1457e1481.
Ishai, A., Haxby, J. V., & Ungerleider, L. (2002). Visual imagery of
famous faces: Effects of memory and attention revealed by
fMRI. Neuroimage, 17, 1729e1741.
Ishai, A., Ungerleider, L., & Haxby, J. V. (2000). Distributed neural
systems for the generation of visual images. Neuron, 28,
979e990.
Ishai, A., Ungerleider, L. G., Martin, A., Schouten, J. L., &
Haxby, J. V. (1999). Distributed representation of objects in the
human ventral visual pathway. Proceedings of the National
Academy of Sciences, 96, 9379e9384.
Jackson, S. R., Condon, L. A., Newport, R. W., Pears, S., Husain, M.,
Bajaj, N., et al. (2018). Optic ataxia and the dorsal visual stream
re-visited: Impairment in bimanual haptic matching
performed without vision. Cortex, 98, 60e72.
Jackson, S. R., Newport, R., Husain, M., Fowlie, J. E.,
O'Donoghue, M., & Bajaj, N. (2009). There may be more to
reaching than meets the eye: Re-thinking optic ataxia.
Neuropsychologia, 47, 1397e1408.
Jackson, S. R., Newport, R., Mort, D., & Husain, M. (2005a). Where
the eye looks, the hand follows: Limb-dependent magnetic
misreaching in optic ataxia. Current Biology, 15, 42e46.
Jackson, S. R., Newport, R., Mort, D., Husain, M., Jackson, G. M.,
Swainson, R., et al. (2005b). Action binding and parietal lobes:
Some new perspectives on optic ataxia. In G. W. Humphreys,
& M. J. Riddoch (Eds.), Attention in action: Advances from cognitive
neuroscience. Hove, UK: Psychology Press.
Jackson, M., & Warrington, E. K. (1986). Arithmetic skills in
patients with unilateral cerebral lesions. Cortex, 22, 611e620.
Jakobson, L. S., Archibald, Y. M., Carey, D. P., & Goodale, M. A.
(1991). A kinematic analysis of reaching and grasping
movements in a patient recovering from optic ataxia.
Neuropsychologia, 29, 803e809.
James, T. W., Culham, J., Humphey, K., Milner, A. D., &
Goodale, M. A. (2003). Ventral occipital lesions impair object
recognition but not object-directed grasping: An fMRI study.
Brain, 126, 2463e2475.
James, M., Plant, G. T., & Warrington, E. K. (1991). Cortical vision
screening test. Oxford, UK: Thames Valley Test Company.
Jankowiak, J., Kinsbourne, M., Shalev, R. S., & Bachman, D. L.
(1992). Preserved visual imagery and categorization in a case
of associative visual agnosia. Journal of Cognitive Neuroscience, 4,
119e131.
Kaczmarek, B. L. J. (1991). Aphasia in an artist: A disorder of
symbolic processing. Aphasiology, 5, 361e371.
Kawashima, R., Roland, P. E., & O'Sullivan, B. T. (1995). Visuo-
motor learning: A positron emission tomography study.
Cerebral Cortex, 5, 111e122.
Kerr, N. H. (1983). The role of vision in ‘visual imagery’
experiments: Evidence from the congentially blind. Journal of
Experimental Psychology: General, 112, 265e277.
Klein, I., Dubois, J., Mangin, J., Kherif, F., Flandin, G., Poline, J.,
et al. (2004). Retinotopic organization of visual mental images
as revealed by functional magnetic resonance imaging.
Cognitive Brain Research, 22, 26e31.
Knauff, M., Kassubek, J., Mulack, T., & Greenlee, M. W. (2000).
Cortical activation evoked by visual mental imagery as
measured by fMRI. Neuroreport, 11, 3957e3962.
Kosslyn, S. M. (1980). Image and mind. Cambridge, MA: Harvard
University Press.
Kosslyn, S. M., Digirolamo, G. J., Thompson, W. L., & Alpert, N. M.
(1998). Mental rotation of objects versus hands: Neural
mechanisms revealed by positron emission tomography.
Psychophysiology, 35, 151e161.
Kosslyn, S. M., Farah, M. J., Holtzman, J. D., & Gazzaniga, M. S.
(1985). A computational analysis of mental image generation:
Evidence from functional dissociations in split-brain patients.
Journal of Experimental Psychology: General, 114, 311e341.
Kosslyn, S. M., Thompson, W. L., & Alpert, N. M. (1997). Neural
systems shared by visual imagery and visual perception: A
positron emission tomography study. Neuroimage, 6, 320e334.
Lee, J., & van Donkelaar, P. (2002). Dorsal and ventral visual
stream contributions to perception-action interactions during
pointing. Experimental Brain Research, 143, 440e446.
Levine, D. N., & Calvanio, R. (1989). Prosopagnosia: A defect in
visual configural processing. Brain and Cognition, 10, 149e170.
Levine, D. N., Warach, J., & Farah, M. (1985). Two visual systems in
mental imagery: Dissociation of “what” and “where” in
imagery disorders due to bilateral posterior cerebral lesions.
Neurology, 35, 1010e1018.
Luzzatti, C., Vecchi, T., Agazzi, D., Cesa-Bianchi, M., & Vergani, C.
(1998). A neurological dissociation between preserved visual
and impaired spatial processing in mental imagery. Cortex, 34,
461e469.
MacPherson, S. E., Wagner, G. P., Murphy, P., Bozzali, M.,
Cipolotti, L., & Shallice, T. (2014). Bringing the cognitive
estimation task into the 21st century: Normative data on two
new parallel forms. Plos One, 9, e92554.
Malach, R., Reppas, J. B., Benson, R. R., Kwong, K. K., Jiang, H.,
Knnedy, W. A., et al. (1995). Object-related activity revealed by
functional magnetic resonance imaging in human occipital
cortex. Proceedings of the National Academy of Sciences, 92,
8135e8139.
Mell, J. C., Howard, S. M., & Miller, B. L. (2003). Art and the brain:
The influence of frontotemporal dementia on an
accomplished artist. Neurology, 60, 1707e1710.
Miller, B. L., & Hou, C. E. (2004). Portraits of artists: Emergence of
visual creativity in dementia. Archives of Neurology, 61,
842e844.
Miller, B. L., Ponton, M., Benson, D. F., Cummings, J., & Mena, I.
(1996). Enhanced artistic creativity with temporal lobe
degeneration. Lancet, 348, 1744e1745.
Milner, A. D., Dijkerman, H. C., McIntosh, R. D., Rossetti, Y., &
Pisella, L. (2003). Delayed reaching and grasping in patients
with optic ataxia. Progress in Brain Research, 142, 225e242.
Milner, A. D., & Goodale, M. A. (1995). The visual brain in action.
Oxford, UK: Oxford University Press.
Milner, A. D., & Goodale, M. A. (2008). Two visual systems re-
viewed. Neuropsychologia, 46, 774e785.
 c o r t e x 1 2 6 ( 2 0 2 0 ) 3 5 5 e3 6 7
Milner, A. D., Perrett, D. I., Johnston, R. S., Benson, P. J.,
Jordan, T. R., Heeley, D. W., et al. (1991). Perception and action
in ‘visual form agnosia’. Brain, 114, 405e428.
Moro, V., Berlucchi, G., Lerch, J., Tomaiuolo, F., & Aglioti, S. M. (2008).
Selective deficit of mental visual imagery with intact primary
visual cortex and visual perception. Cortex, 44, 109e118.
Oldfield, R. C., & Wingfield, A. (1965). A series of pictures for use in
object-naming. MRC Psycholinguistic Research Unit Special
Report No. PLU/65/19.
O'Craven, K. M., & Kanwisher, N. (2000). Mental imagery of faces
and places activates corresponding stimulus-specific brain
regions. Journal of Cognitive Neuroscience, 12, 1013e1023.
Papagno, C. (2002). Progressive impairment of constructional
abilities: A visuospatial sketchpad deficit? Neuropsychologia, 40,
1858e1867.
Pearson, J., Naselaris, T., Holmes, E. A., & Kosslyn, S. M. (2015).
Mental imagery: Functional mechanisms and clinical
applications. Trends in Cognitive Sciences, 19, 590e602.
Petcu, E. B., Sherwood, K., Popa-Wagner, A., Buga, A. M., Aceti, L.,
& Miroiu, R. I. (2016). Artistic skills recovery and compensation
in visual artists after stroke. Frontiers in Neurology, 7, 76.
a, H., Tilikete, C., Vighetto, A., Desmurget, M.,
Pisella, L., Gre
Rode, G., et al. (2000). An automatic pilot for the hand in the
human posterior parietal cortex toward a reinterpretation of
optic ataxia. Nature Neuroscience, 3, 729e736.
Pisella, L., Sergio, L., Blangero, A., Torchin, H., Vighetto, A., &
Rossetti, Y. (2009). Optic ataxia and the function of the dorsal
stream: Contributions to perception and action.
Neuropsychologia, 47, 3033e3044.
Ratcliff, G. (1979). Spatial though, mental rotation and the right
cerebral hemisphere. Neuropsychologia, 17, 49e54.
Riddoch, M. J. (1990). Loss of visual imagery: A generation deficit.
Cognitive Neuropsychology, 7, 249e273.
Riddoch, M. J., & Humphreys, G. W. (1987). A case of integrative
visual agnosia. Brain, 110, 1431e1462.
Riddoch, M. J., Humphreys, G. W., Gannon, T., Blott, W., &
Jones, V. (1999). Memories are made of this: The effect of time
on stored visual knowledge in a case of visual agnosia. Brain,
122, 537e559.
Rizzo, M., & Robin, D. A. (1990). Simultanagnosia: A defect of
sustained attention yields insights on visual information
processing. Neurology, 40, 447.
Rosetti, Y., Pisella, L., & McIntosh, R. D. (2017). Rise and fall of two
visual systems theory. Annals of Physical and Rehabilitation
Medicine, 60, 130e140.
Rossetti, Y., Vighetto, A., & Pisella, L. (2003). Optic ataxia revisited:
Immediate motor control versus visually guided action.
Experimental Brain Research, 153, 171e179.
Rowe, F. J., Hepworth, L. R., Howard, C., Hanna, K. L.,
Cheyne, C. P., & Currie, J. (2009). High incidence and
prevalence of visual problems after acute stroke: An
epidemiology study with implications for service delivery. Plos
One, 14, e0213035.
Sack, A. T., Camprodon, J. A., Pascual-Leone, A., & Goebel, R.
(2005). The dynamics of interhemispheric compensatory
processes in mental imagery. Science, 308, 702e704.
Sack, A. T., Jacobs, C., De Martino, F., Staeren, N., Goebel, R., &
Formisano, E. (2008). Dynamic premotor-to-parietal
interactions during spatial imagery. Journal of Neuroscience, 28,
8417e8429.
Sack, A. T., & Schuhmann, T. (2012). Hemispheric differences
within the fronto-parietal network dynamics underlying
spatial imagery. Frontiers in Psychology, 3, 214.
367
Sack, A. T., Sperling, J. M., Prvulovic, D., Formisano, E., Goebel, R.,
Di Salle, F., et al. (2002). Tracking the mind's image in the
brain: II. Transcranial magnetic stimulation reveals parietal
asymmetry in visuospatial imagery. Neuron, 35, 195e204.
Schnider, A., Regard, M., Benson, D. F., & Landis, T. (1993). Effects
of a right-hemisphere stroke on an artist's performance.
Neuropsychiatry, Neuropsychology, and Behavioral Neurology, 6,
249e255.
Schott, G. D. (2012). Pictures as a neurological tool: Lessons from
enhanced and emergent artistry in brain disease. Brain, 135,
1947e1963.
Sherwood, K. (2012). How a cerebral haemorrhage altered my art.
Frontiers in Human Neuroscience, 6, 55.
Shi, B., Cao, X., Chen, Q., Zhuang, K., & Qiu, J. (2017). Different
brain structures associated with artistic and scientific
creativity: A voxel-based morphometry study. Scientific reports,
7, 42911.
Slotnick, S. D., Thompson, W. L., & Kosslyn, S. M. (2005). Visual
mental imagery induces retinotopically organized activation
of early visual areas. Cerebral Cortex, 15, 1570e1583.
Smith, W. S., Mindelzun, R. E., & Miller, B. (2003). Simultanagnosia
through the eyes of an artist. Neurology, 60, 1832e1834.
Taubert, J., & Alais, D. (2009). The composite illusion requires
composite face stimuli to be biologically plausible. Vision
Research, 49, 1877e1885.
Thirion, B., Duchesnay, E., Hubbard, E., Dubois, J., Poline, J. B.,
Lebihan, D., et al. (2006). Inverse retinotopy: Inferring the
visual content of images from brain activation patterns.
Neuroimage, 33, 1104e1116.
Thompson, W. L., Hsiao, Y., & Kosslyn, S. M. (2011). Dissociation
between visual attention and visual mental imagery. Journal of
Cognitive Psychology, 23, 256e263.
Thompson, W. L., Slotnick, S. D., Burrage, M. S., & Kosslyn, S. M.
(2009). Two forms of spatial imagery: Neuroimaging evidence.
Psychological Science, 20, 1245e1253.
Tong, F. (2003). Primary visual cortex and visual awareness.
Nature Reviews Neuroscience, 4, 219e229.
Ungerleider, L. G., & Mishkin, M. (1982). Two cortical visual
systems. In D. J. Ingle, M. A. Goodale, & R. J. W. Mansfield
(Eds.), Analysis of visual behaviour. Cambridge, Massachusetts:
The MIT Press.
Valyear, K. F., Culham, J. C., Sharif, N., Westwood, D., &
Goodale, M. A. (2006). A double dissociation between
sensitivity to changes in object identity and object orientation
in the ventral and dorsal visual streams: A human fMRI study.
Neuropsychologia, 44, 218e228.
Wapner, W., Judd, T., & Gardner, H. (1978). Visual agnosia in an
artist. Cortex, 14, 343e364.
Warrington, E. K. (1984). Recognition memory test. Windsor, UK:
NEFR-Nelson.
Warrington, E. K., & James, M. (1991). The visual object and space
perception battery. Bury St Edmunds, UK: Thames Valley Test
Company.
Weber, R. J., & Castleman, J. (1970). The time it takes to imagine.
Perception and Psychophysics, 8, 165e168.
Wechsler, D. (1997). In WAIS-III. San Antonio, TX, USA:
Psychological Corporation.
Winlove, C. I. P., Milton, F., Ranson, J., Fulford, J., MacKisack, M.,
Macpherson, F., et al. (2018). The neural correlates of visual
imagery: A co-ordinate-based meta-analysis. Cortex, 105, 4e25.
Young, A. W., Humphreys, G. W., Riddoch, M. J., Hellawell, D. J., &
de Haan, E. H. F. (1994). Recognition impairments and face
imagery. Neuropsychologia, 32, 693e702.