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Colors in the mind’s eye
Abstract
The famous “Piazza del Duomo” paper, published in Cortex in 1978, inspired a considerable amount of research on visual mental imagery in brain-damaged patients. As a consequence, single-case reports featuring dissociations between perceptual and imagery abilities challenged the prevailing model of visual mental imagery. Here we focus on mental imagery for colors. A case study published in Cortex showed perfectly preserved color imagery in a patient with acquired achromatopsia after bilateral lesions at the borders between the occipital and temporal cortex. Subsequent neuroimaging findings in healthy participants extended and specified this result; color imagery elicited activation in both a domain-general region located in the left fusiform gyrus and the anterior color-biased patch within the ventral temporal cortex, but not in more posterior color-biased patches. Detailed studies of individual neurological patients, as those often published in Cortex, are still critical to inspire and constrain neurocognitive research and its theoretical models.
The coordination of different brain regions is required for the visual imagery of complex objects (e.g., faces and places). Short-range connectivity within sensory areas is necessary to construct the mental image. Long-range connectivity between control and sensory areas is necessary to re-instantiate and maintain the mental image. While dynamic changes in functional connectivity are expected during visual imagery, it is unclear whether a category-specific network exists in which the strength and the spatial destination of the connections vary depending on the imagery target. In this magnetoencephalography study, we used a minimally constrained experimental paradigm wherein imagery categories were prompted using visual word cues only, and we decoded face versus place imagery based on their underlying functional connectivity patterns as estimated from the spatial covariance across brain regions. A subnetwork analysis further disentangled the contribution of different connections. The results show that face and place imagery can be decoded from both short-range and long-range connections. Overall, the results show that imagined object categories can be distinguished based on functional connectivity patterns observed in a category-specific network. Notably, functional connectivity estimates rely on purely endogenous brain signals suggesting that an external reference is not necessary to elicit such category-specific network dynamics.
How do attention and consciousness interact in the human brain? Rival theories of consciousness disagree on the role of fronto-parietal attentional networks in conscious perception. We recorded neural activity from 727 intracerebral contacts in 13 epileptic patients, while they detected near-threshold targets preceded by attentional cues. Clustering revealed three neural patterns: first, attention-enhanced conscious report accompanied sustained right-hemisphere fronto-temporal activity in networks connected by the superior longitudinal fasciculus (SLF) II-III, and late accumulation of activity (>300 ms post-target) in bilateral dorso-prefrontal and right-hemisphere orbitofrontal cortex (SLF I-III). Second, attentional reorienting affected conscious report through early, sustained activity in a right-hemisphere network (SLF III). Third, conscious report accompanied left-hemisphere dorsolateral-prefrontal activity. Task modeling with recurrent neural networks revealed multiple clusters matching the identified brain clusters, elucidating the causal relationship between clusters in conscious perception of near-threshold targets. Thus, distinct, hemisphere-asymmetric fronto-parietal networks support attentional gain and reorienting in shaping human conscious experience.
We used ultra-high field 7T fMRI to establish the neural circuits involved in visual mental imagery and perception, and to elucidate the neural mechanisms associated with imagery absence in congenital aphantasia. Ten typical imagers and 10 aphantasic individuals performed imagery and perceptual tasks in five domains: object shape, object color, written words, faces, and spatial relationships. In typical imagers, imagery tasks activated left-hemisphere fronto-parietal areas, a domain-general area in the left fusiform gyrus (the Fusiform Imagery Node), together with the relevant domain-preferring areas in the ventral temporal cortex. In aphantasic individuals, imagery activated similar areas, but the Fusiform Imagery Node was functionally disconnected from fronto-parietal areas. Our results unveil the domain-general and domain-specific circuits of visual mental imagery, their functional disorganization in aphantasia, and support the general hypothesis that subjective visual experience - whether perceived or imagined - depends on the integrated activity of high-level visual cortex and fronto-parietal networks.
Most of us can use our “mind’s eye” to mentally visualize things that are not in our direct line of sight, an ability known as visual mental imagery. Extensive left temporal damage can impair patients’ visual mental imagery experience, but the critical locus of lesion is unknown. Our recent meta-analysis of 27 fMRI studies of visual mental imagery highlighted a well-delimited region in the left lateral midfusiform gyrus, which was consistently activated during visual mental imagery, and which we called the Fusiform Imagery Node (FIN). Here, we describe the connectional anatomy of FIN in neurotypical participants and in RDS, a right-handed patient with an extensive occipito-temporal stroke in the left hemisphere. The stroke provoked right homonymous hemianopia, alexia without agraphia, and color anomia. Despite these deficits, RDS had normal subjective experience of visual mental imagery and reasonably preserved behavioral performance on tests of visual mental imagery of object shape, object color, letters, faces, and spatial relationships. We found that the FIN was spared by the lesion. We then assessed the connectional anatomy of the FIN in the MNI space and in the patient’s native space, by visualizing the fibers of the inferior longitudinal fasciculus (ILF) and of the arcuate fasciculus (AF) passing through the FIN. In both spaces, the ILF connected the FIN with the anterior temporal lobe, and the AF linked it with frontal regions. Our evidence is consistent with the hypothesis that the FIN is a node of a brain network dedicated to voluntary visual mental imagery. The FIN could act as a bridge between visual information and semantic knowledge processed in the anterior temporal lobe and in the language circuits.
How do attentional networks influence conscious perception? To answer this question, we used magnetoencephalography in human participants and assessed the effects of spatially nonpredictive or predictive supra-threshold peripheral cues on the conscious perception of near-threshold Gabors. Three main results emerged. (i) As compared with invalid cues, both nonpredictive and predictive valid cues increased conscious detection. Yet, only predictive cues shifted the response criterion toward a more liberal decision (i.e. willingness to report the presence of a target under conditions of greater perceptual uncertainty) and affected target contrast leading to 50% detections. (ii) Conscious perception following valid predictive cues was associated to enhanced activity in frontoparietal networks. These responses were lateralized to the left hemisphere during attentional orienting and to the right hemisphere during target processing. The involvement of frontoparietal networks occurred earlier in valid than in invalid trials, a possible neural marker of the cost of re-orienting attention. (iii) When detected targets were preceded by invalid predictive cues, and thus reorienting to the target was required, neural responses occurred in left hemisphere temporo-occipital regions during attentional orienting, and in right hemisphere anterior insular and temporo-occipital regions during target processing. These results confirm and specify the role of frontoparietal networks in modulating conscious processing and detail how invalid orienting of spatial attention disrupts conscious processing.
Purpose of Review
Color provides important information about the identity of the objects we encounter. After early processing stages in the retinal cones, thalamus, and occipital cortex, retinal signals reach the ventral temporal cortex for high-level color and object processing, which links color perception with top-down expectations and knowledge. In the language-dominant hemisphere, some of these regions communicate with the language systems; by assigning verbal labels to percepts, these circuits speed up stimulus categorization, and permit fast and accurate inter-individual communication. This paper provides a review of color processing deficits, from dysfunction of wavelength discrimination in the retinal photoreceptors to deficits of high-level processing in the ventral temporal cortex.
Recent Findings
Neuroimaging evidence defined the existence and localization of color-preferring domains in the ventral occipito-temporal cortex. Evidence from the performance of a brain-damaged patient with color anomia but preserved color categorization demonstrated the independence of color categorization from color naming in the adult brain.
Summary
Evidence from patients with brain damage suggests that high-level color processing may be divided into at least three functional domains: perceptual color experience, color naming, and color knowledge.
Visual mental imagery is the faculty whereby we can “visualize” objects that are not in our line of sight. Longstanding evidence dating back over thirty years has shown that unilateral brain lesions, especially in the left temporal lobe, can impair aspects of this ability. Yet, there is currently no attempt to identify analogies between these neuropsychological findings of hemispheric asymmetry and those from other neuroscientific approaches. Here, we present a critical review of the available literature on the hemispheric laterality of visual mental imagery, by looking at cross-method patterns of evidence in the domains of lesion neuropsychology, neuroimaging, and direct cortical stimulation. Results can be summarized under three main axes. First, frontoparietal networks in both hemispheres appear to be associated with visual mental imagery. Second, lateralization patterns emerge in the temporal lobes, with the left inferior temporal lobe being the most common finding in the literature for endogenously generated images, especially, but not exclusively, when orthographic material is used to ignite imagery. Third, an opposite pattern of hemispheric laterality emerges when visual mental images are induced by exogenous stimulation; direct cortical electrical stimulation tends to produce visual imagery experiences predominantly when applied to the right temporal lobe. These patterns of hemispheric asymmetry are difficult to reconcile with the dominant model of visual mental imagery, which emphasizes the implication of early sensory cortices. They suggest instead that visual mental imagery relies on large-scale brain networks, with a crucial participation of high-level visual regions in the temporal lobes.
The year 2021 marks the 130th anniversary of the untimely death of Heinrich Lissauer (1861-1891). In his thirty years of life, Lissauer managed to put together an impressive number of contributions to neurology and neuroanatomy. Most influential is his famous distinction between apperceptive and associative forms of visual agnosia. It is perhaps less well-known that in the same article, Lissauer outlined a model of possible dissociations between visual perception and visual mental imagery. Drawing on Hermann Munk’s animal experiments, Lissauer proposed that complete destruction of occipital visual cortex would provoke both perception and imagery deficits, whereas its deafferentation resulting from subcortical white matter damage would only affect visual perception, and leave visual memories unimpaired. This proposal resonates with the present debate on the neural bases of visual mental imagery.
Color is continuous, yet we group colors into discrete categories associated with color names (e.g., yellow, blue). Color categorization is a case in point in the debate on how language shapes human cognition. Evidence suggests that color categorization depends on top-down input from the language system to the visual cortex. We directly tested this hypothesis by assessing color categorization in a stroke patient, RDS, with a rare, selective deficit in naming visually presented chromatic colors, and relatively preserved achromatic color naming. Multimodal MRI revealed a left occipito-temporal lesion that directly damaged left color-biased regions, and functionally disconnected their right-hemisphere homologs from the language system. The lesion had a greater effect on RDS's chromatic color naming than on color categorization, which was relatively preserved on a nonverbal task. Color categorization and naming can thus be independent in the human brain, challenging the mandatory involvement of language in adult human cognition.
Unlabelled:
The existence of color-processing regions in the human ventral visual pathway (VVP) has long been known from patient and imaging studies, but their location in the cortex relative to other regions, their selectivity for color compared with other properties (shape and object category), and their relationship to color-processing regions found in nonhuman primates remain unclear. We addressed these questions by scanning 13 subjects with fMRI while they viewed two versions of movie clips (colored, achromatic) of five different object classes (faces, scenes, bodies, objects, scrambled objects). We identified regions in each subject that were selective for color, faces, places, and object shape, and measured responses within these regions to the 10 conditions in independently acquired data. We report two key findings. First, the three previously reported color-biased regions (located within a band running posterior-anterior along the VVP, present in most of our subjects) were sandwiched between face-selective cortex and place-selective cortex, forming parallel bands of face, color, and place selectivity that tracked the fusiform gyrus/collateral sulcus. Second, the posterior color-biased regions showed little or no selectivity for object shape or for particular stimulus categories and showed no interaction of color preference with stimulus category, suggesting that they code color independently of shape or stimulus category; moreover, the shape-biased lateral occipital region showed no significant color bias. These observations mirror results in macaque inferior temporal cortex (Lafer-Sousa and Conway, 2013), and taken together, these results suggest a homology in which the entire tripartite face/color/place system of primates migrated onto the ventral surface in humans over the course of evolution.
Significance statement:
Here we report that color-biased cortex is sandwiched between face-selective and place-selective cortex on the bottom surface of the brain in humans. This face/color/place organization mirrors that seen on the lateral surface of the temporal lobe in macaques, suggesting that the entire tripartite system is homologous between species. This result validates the use of macaques as a model for human vision, making possible more powerful investigations into the connectivity, precise neural codes, and development of this part of the brain. In addition, we find substantial segregation of color from shape selectivity in posterior regions, as observed in macaques, indicating a considerable dissociation of the processing of shape and color in both species.
Brain damage can produce acquired deficits of color perception, or cerebral achromatopsia. In these patients, lesions tend to overlap on a restricted region in the ventral occipitotemporal cortex, close to the reported locations of the putative V4 complex and to foci of increased blood-oxygen-level-dependent (BOLD) activity related to color perception in normal participants. Unilateral lesions give rise to achromatopsia in the contralateral visual field (hemiachromatopsia). Here we present a partial English translation of the first case report of a hemiachromatopsic patient with detailed anatomical evidence (Madame R., Verrey, 1888), and discuss these results in relation to a more recent case report (Madame D., Bartolomeo et al., 1997) of a patient with two consecutive hemorrhagic lesions in the occipitotemporal regions of the two hemispheres. Strikingly, Madame D. developed full-field achromatopsia after the second lesion in the right hemisphere, without having shown any signs of hemiachromatopsia after the first lesion in the left hemisphere. Thanks to the comparison of the reconstructed lesion patterns between the two patients and with the putative location of color-related areas in the human brain, we offer a possible, if speculative, account of this puzzling pattern of anatomo-clinical correlations, based on intra- and inter-hemispheric connectivity.
Kosslyn (psychology, Harvard U.) presents a 20-year research program on the nature of high-level vision and mental imagery--offering his research as a definitive resolution of the long-standing "imagery debate," which centers on the nature of the internal representation of visual mental imagery. He combines insights and empirical results from computer vision, neurobiology, and cognitive science to develop a general theory of visual mental imagery, its relation to visual perception, and its implementation in the human brain.
Visual imagery is the creation of mental representations that share many features with veridical visual percepts. Studies of normal and brain-damaged people reinforce the view that visual imagery and visual perception are mediated by a common neural substrate and activate the same representations. Thus, brain-damaged patients with intact vision who have an impairment in perception should have impaired visual imagery. Here we present evidence to the contrary from a patient with severely impaired object recognition (visual object agnosia) but with normal mental imagery. He draws objects in considerable detail from memory and uses information derived from mental images in a variety of tasks. In contrast, he cannot identify visually presented objects, even those he has drawn himself. He has normal visual acuity and intact perception of equally complex material in other domains. We conclude that rich internal representations can be activated to support visual imagery even when they cannot support visually mediated perception of objects.
Does visual imagery engage some of the same representations used in visual perception? The evidence collected by cognitive psychologists in support of this claim has been challenged by three types of alternative explanation: Tacit knowledge, according to which subjects use nonvisual representations to simulate the use of visual representations during imagery tasks, guided by their tacit knowledge of their visual systems; experimenter expectancy, according to which the data implicating shared representations for imagery and perception is an artifact of experimenter expectancies; and nonvisual spatial representation, according to which imagery representations are partially similar to visual representations in the way they code spatial relations but are not visual representations. This article reviews previously overlooked neuropsychological evidence on the relation between imagery and perception, and discusses its relative immunity to the foregoing alternative explanations. This evidence includes electrophysiological and cerebral blood flow studies localizing brain activity during imagery to cortical visual areas, and parallels between the selective effects of brain damage on visual perception and imagery. Because these findings cannot be accounted for in the same way as traditional cognitive data using the alternative explanations listed earlier, they can play a decisive role in answering the title question.
We report here the use of positron emission tomography (PET) to reveal that the primary visual cortex is activated when subjects close their eyes and visualize objects. The size of the image is systematically related to the location of maximal activity, which is as expected because the earliest visual areas are spatially organized. These results were only evident, however, when imagery conditions were compared to a non-imagery baseline in which the same auditory cues were presented (and hence the stimuli were controlled); when a resting baseline was used (and hence brain activation was uncontrolled), imagery activation was obscured because of activation in visual cortex during the baseline condition. These findings resolve a debate in the literature about whether imagery activates early visual cortex and indicate that visual mental imagery involves 'depictive' representations, not solely language-like descriptions. Moreover, the fact that stored visual information can affect processing in even the earliest visual areas suggests that knowledge can fundamentally bias what one sees.
A patient with complete cortical blindness after bilateral posterior cerebral artery infarctions denied her blindness. Her pretended visual experiences could frequently be traced back to synaesthetic translations of acoustic or tactile perceptions into mental visual images. Possibly, the belief to see resulted from a confusion of mental visual images with real percepts. The patient manifested preserved visual imagery also by correct responses to questions concerning the shapes of letters and the shapes and colours of objects. MRI showed an almost complete destruction of primary visual cortex with sparing of only small remainders of cortex at the occipital tip of the left upper calcarine lip. In the literature there are a few cases of denial of blindness with similarly severe damage to primary visual cortex but none with unequivocal evidence of complete destruction of primary visual cortex. We conclude that severe damage to primary visual cortex is compatible with visual imagery but that there is a possibility that islands of visual cortex must be spared to permit the generation of mental visual images.
We report on two brain-damaged subjects who exhibit the uncommon pattern of loss of object color knowledge, but spared color perception and naming. The subject P.C.O., as in previously reported patients, is also impaired in processing other perceptual and functional properties of objects. I.O.C., in contrast, is the first subject on record to have impaired object color knowledge, but spared knowledge of object form, size and function. This pattern of performance is consistent with the view that semantic information about color and other perceptual properties of objects is grounded in modality-specific systems. Lesion analysis suggests that such grounding requires the integrity of the mesial temporal regions of the left hemisphere.
Different individuals experience varying degrees of vividness in their visual mental images. The distribution of these variations across different imagery domains, such as object shape, color, written words, faces, and spatial relationships, remains unknown. To address this issue, we conducted a study with 117 healthy participants who reported different levels of imagery vividness. Of these participants, 44 reported experiencing absent or nearly absent visual imagery, a condition known as "aphantasia". These individuals were compared to those with typical (N = 42) or unusually vivid (N = 31) imagery ability. We used an online version of the French-language Battérie Imagination-Perception (eBIP), which consists of tasks tapping each of the above-mentioned domains, both in visual imagery and in visual perception. We recorded the accuracy and response times (RTs) of participants' responses. Aphantasic participants reached similar levels of accuracy on all tasks compared to the other groups (Bayesian repeated measures ANOVA, BF = .02). However, their RTs were slower in both imagery and perceptual tasks (BF = 266), and they had lower confidence in their responses on perceptual tasks (BF = 7.78e5). A Bayesian regression analysis revealed that there was an inverse correlation between subjective vividness and RTs for the entire participant group: higher levels of vividness were associated with faster RTs. The pattern was similar in all the explored domains. The findings suggest that individuals with congenital aphantasia experience a slowing in processing visual information in both imagery and perception, but the precision of their processing remains unaffected. The observed performance pattern lends support to the hypotheses that congenital aphantasia is primarily a deficit of phenomenal consciousness, or that it employs alternative strategies other than visualization to access preserved visual information.
The ventral temporal cortex hosts key regions for the high-level visual processing of object shape and color. These areas represent nodes of large-scale neural circuits dedicated to object recognition. In the language-dominant hemisphere, some of these regions communicate with the language systems; by assigning verbal labels to percepts, these circuits speedup stimulus categorization, and permit fast and accurate interindividual communication. By impairing the functioning of these circuits, neurological damage may provoke disabling disorders of the processing of visual objects and of their colors. Brain damage of vascular, degenerative, toxic, or traumatic origin can induce deficits at different levels of visual processing, from the building of shape- or wavelength-invariant percepts, to their connections with semantic knowledge and with the appropriate lexical entry. After an overview of the neuroimaging of domain-preferring regions for object shape and color in the ventral temporal cortex, this chapter reviews evidence from historical and recent cases of acquired visual agnosia and color processing deficits. A recurrent motif emerging from patients’ patterns of performance and lesion locations is the existence of caudo-rostral gradients in the ventral occipito-temporal cortex, spanning from more perceptual to more cognitive stages of processing.
The dominant neural model of visual mental imagery (VMI) stipulates that memories from the medial temporal lobe acquire sensory features in early visual areas. However, neurological patients with damage restricted to the occipital cortex typically show perfectly vivid VMI, while more anterior damages extending into the temporal lobe, especially in the left hemisphere, often cause VMI impairments. Here we present two major results reconciling neuroimaging findings in neurotypical subjects with the performance of brain-damaged patients: (1) a large-scale meta-analysis of 46 fMRI studies, of which 27 investigated specifically visual mental imagery, revealed that VMI engages fronto-parietal networks and a well-delimited region in the left fusiform gyrus. (2) A Bayesian analysis showing no evidence for imagery-related activity in early visual cortices. We propose a revised neural model of VMI that draws inspiration from recent cytoarchitectonic and lesion studies, whereby fronto-parietal networks initiate, modulate, and maintain activity in a core temporal network centered on the fusiform imagery node, a high-level visual region in the left fusiform gyrus.
Color provides valuable information about the environment, yet the exact mechanisms explaining how colors appear to us remain poorly understood. Retinal signals are processed in the visual cortex through high-level mechanisms that link color perception with top-down expectations and knowledge. Here, we review the neuroimaging evidence about color processing in the brain, and how it is affected by acquired brain lesions in humans. Evidence from patients with brain-damage suggests that high-level color processing may be divided into at least three modules: perceptual color experience, color naming, and color knowledge. These modules appear to be functionally independent but richly interconnected, and serve as cortical relays linking sensory and semantic information, with the final goal of directing object-related behavior. We argue that the relations between colors and their objects are key mechanisms to understand high-level color processing.
We investigated object-colour knowledge in RDS, a patient with impaired colour naming after a left occipito-temporal stroke. RDS’s colour perception, object naming and verbal colour-knowledge (the ability to verbally say the typical colour of an object) were relatively spared. RDS was also able to state if an object was appropriately coloured or not. However, he could neither match colour names to coloured objects, nor match colour patches to grey-scale objects. Thus, RDS’s colour-naming deficit was associated with an impaired ability to conceptually relate visually presented object shapes and colours. These results suggest that objects in their typical colour are processed holistically in the visual modality, and that abilities important for colour naming may also be involved in abstracting colours from visual objects. We discuss these findings in the context of developmental psychology and linguistic anthropology, and propose a model of neuro-functional organization of object-colour knowledge.
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. Mental imagery plays a role in a variety of cognitive processes such as memory recall. In this review, Joel Pearson discusses recent insights into the neural mechanisms that underlie visual imagery, how imagery can be objectively and reliably measured, and how it affects general cognition.
One of the founding principles of human cognitive neuroscience is the so-called universality assumption, the postulate that neurocognitive mechanisms do not show major differences among individuals. Without negating the importance of the universality assumption for the development of cognitive neuroscience, or the importance of single-case studies, here we aim at stressing the potential dangers of interpreting the pattern of performance of single patients as conclusive evidence concerning the architecture of the intact neurocognitive system. We take example from the case of Leonardo Botallo, an Italian surgeon of the Renaissance period, who claimed to have discovered a new anatomical structure of the adult human heart. Unfortunately, Botallo’s discovery was erroneous, because what he saw in the few samples he examined was in fact the anomalous persistence of a fetal structure. Botallo’s error is a reminder of the necessity to always strive for replication, despite the major hindrance of a publication system heavily biased towards novelty. In the present paper, we briefly discuss variations and anomalies in human brain anatomy and introduce the issue of variability in cognitive neuroscience. We then review some examples of the impact on cognition of individual variations in (1) brain structure, (2) brain functional organization and (3) brain damage. We finally discuss the importance and limits of single case studies in the neuroimaging era, outline potential ways to deal with individual variability, and draw some general conclusions.
When we try to remember whether we left a window open or closed, do we actually see the window in our mind? If we do, does this mental image play a role in how we think? For almost a century, scientists have debated whether mental images play a functional role in cognition. The Case for Mental Imagery presents a complete and unified argument that mental images do depict information, and that these depictions do play a functional role in human cognition. It outlines a specific theory of how depictive representations are used in information processing, and shows how these representations arise from neural processes. To support this theory, it weaves together conceptual analyses and the many varied empirical findings from cognitive psychology and neuroscience. In doing so, the book presents the conceptual grounds for positing this type of internal representation, summarizing and refuting arguments to the contrary. Its argument also serves as a historical review of the imagery debate from its earliest inception to its most recent phases, and provides evidence that significant progress has been made in our understanding of mental imagery. In illustrating how scientists think about one of the most difficult problems in psychology and neuroscience, this book goes beyond the debate, to explore the nature of cognition and to draw out implications for the study of consciousness. © 2006 by Stephen M. Kosslyn, William L. Thompson, and Giorgio Ganis. All rights reserved.
The demonstration of an implication of attentional/eye gaze systems in visual mental imagery might help to understand why some patients with visual neglect, who suffer from severe attentional deficits, also show neglect for mental images. When normal participants generate mental images of previously explored visual scenes, their oculomotor behavior resembles that used during visual exploration. However, this could be a case of encoding specificity, whereby the probability of retrieving an event increases if some information encoded with the event (in this case its spatial location) is present at retrieval. In the present study, normal participants were invited to conjure up a mental image of the map of France and to say whether auditorily presented towns or regions were situated left or right of Paris. A perceptual version of the task was administered after the imaginal condition. Thus, in the imaginal condition participants had to retrieve information from long-term memory. Vocal response times and, unbeknownst to participants, also eye movements were recorded. Participants tended to produce similar eye movements on the imaginal and on the perceptual conditions of the task. We concluded that some mechanisms involved in spontaneous oculomotor behavior may be shared in exploration of visuospatial mental images. Deficits of these common processes participating in the oculomotor exploration might contribute to imaginal neglect.
The relationships between spatial neglect for perceptual objects and representational for imagined items are difficult to explore because of several methodological problems, including the dearth of comparable tests for real and imagined scenes. We asked 19 patients with right brain damage and 12 healthy controls to say whether an auditorily presented French geographical location was left or right of Paris, and recorded their vocal response times. Afterwards, participants performed a similar test with visually presented items. Although several patients had asymmetries of performance on the perceptual version of the test, only one patient was more accurate for right-sided than for left-sided imagined stimuli, thus showing evidence for imaginal neglect. However, this patient performed normally on place description and on mental number line bisection, perhaps as a consequence of different strategies he employed for these tasks. Overall, our results confirm previous evidence showing that imaginal neglect is less frequent than, and often occurs in association with, perceptual neglect. Imaginal neglect may result from the contribution of deficits partly distinct from those implicated in perceptual neglect, such as impaired endogenous orienting of attention or deficits of spatial working memory.
We describe the case of a patient with right hemisphere damage and left unilateral neglect. The patient was asked to draw from memory common objects, either with or without visual feedback. In the conditions without visual feedback the patient was either blindfolded or he made "invisible" drawings using a pen with the cap on, the drawings being recorded with carbon paper underneath. Results showed more neglect without than with visual feedback, contrary to previously published cases. This patient's pattern of performance may result from the contribution of a deficit of spatial working memory. Alternatively or in addition, the patient, who was undergoing cognitive rehabilitation for neglect, may have found easier to compensate for his neglect with visual feedback, which allowed him to visually explore the left part of his drawings.
A case of loss of mental imagery following a vascular lesion of the left occipital lobe is described and discussed. The findings support a twofold (analogue and propositional) theory of neural representations of the external world. It is argued that sense-specific representations may be preserved in spite of the reported loss of imagery in the corresponding modality. the possibility that this disorder may reflect a functional disconnection between brain centres is discussed.RésuméOn décrit et on discute un cas de perte de l'imagerie mentale à la suite d'une lésion vasculaire du lobe occipital gauche. Ces constatations sont en faveur d'une théorie à 2 plans (analogue et propositionnel) des représentations nerveuses du monde extérieur. On admet que les représentations spécifiques des modalités sensorielles peuvent être préservées malgré la perte de l'imagerie dans la modalité correspondante. On discute la possibilité que ce désordre traduise une disconnexion fonctionnelle entre centres cérébraux.ZusammenfassungNach einer vaskulären Läsion im linken Occipitallappen verlor ein Patient die Fähigkeit zur bildlichen Vorstellung. Die Beobachtung unterstützt die Theorie einer zweifachen, analogen und propositionellen neuronalen Repräsentation der Auβenwelt. Es wird erörtert, daβ sinnesspezifische Repräsentationen erhalten bleiben können, auch wenn die Vorstellungskraft in der entsprechenden Modalität verloren geht. Die Möglichkeit wird diskutiert, daβ diese Funktionsstörung eine funktionelle Leitungsstörung zwischen Hirnzentren anzeigt.
Using a quantitative measure, we analyzed the relationship between visuospatial and representational neglect in right- and left-brain-damaged patients and found signs of representational neglect only in right-brain-damaged patients. Although representational neglect was always associated with visuospatial neglect, suggesting that the two forms share a common underlying mechanism, the most frequent finding in right-brain--damaged patients was that of visuospatial neglect in isolation. A strong influence of the phenomenon of attentional attraction toward space ipsilateral to the lesion in visuospatial, as opposed to imaginal, tasks can account for this finding.
Two cases (G.G. and A.V.) are described of cognitive impairment resulting from herpes simplex infection. Both cases demonstrated anomic disorders and impairments in drawing but only in G.G.'s drawings was there a reliable selective impairment for items from natural categories. Both cases, however, showed an impairment for the retrieval of knowledge concerning the colours of objects. The impairment has, in the past, been ascribed to interference from colour anomia; this was not so for the present cases. For G.G. and A.V., impairments in object-colour retrieval were related to errors in picture naming. More errors were associated with items that induced circumlocutions than to those that were correctly named. The impairment was also present for some items that were named correctly. The patients' impairments are discussed within a model in which object-colour knowledge is functionally situated between an object's shape description and its output phonology but on a separate route from other associated object knowledge.
Mental visual images can be employed for widely different tasks and they can depict widely different kinds of visual entities. There is evidence that the context and content of mental visual images matter for the determination of their neural basis. Single case studies demonstrate that there are at least five kinds of visual entities whose imagery can be affected independently from each other: shapes of objects, colours of objects, faces, letters and spatial relationships.
Disorders in perceiving and exploring the visual space contralateral to a brain lesion have been frequently described. Many patients with hemi-neglect for extrapersonal space also show neglect in a representational domain when the task requires imagining a well-known piazza from a given vantage point or comparing two visual images. Cognitive and psychophysiological studies show a functional parallelism between the perceptual and imaginative domain, indicating that spatial perception and imagery share the same neural substrata. Here we describe a patient with a persistent disorder in visual imagery for familiar piazzas in the absence of any neglect for stimuli located in a far or near space or on his own body. Contrary to previous cases involving imagery disorders, computerized tomography scans showed a lesion confined to the right frontal lobe, suggesting the role of the frontal lobe in some specific types of mental imagery.
We investigated the ability of a patient (D.F.) with profound visual form agnosia to perform a variety of tasks requiring visual imagery. Despite her inability to discriminate between objects and patterns of different shapes, sizes, and orientations, D.F. showed quite normal visual imagery involving these same 'visual' properties when the images were drawn from long-term memory. Thus, she was able both to scan mental images in search of particular features and to form new images by combining several known images. While there is growing evidence that perception and imagery share common neural substrates, the fact that D.F. shows intact visual imagery in the face of a massive perceptual deficit in form vision challenges recent suggestions that these two psychological processes share common input pathways in early vision. It is suggested that regions in the occipitotemporal pathway may be important for the generation of visual images while regions in the posterior parietal system might be involved in the manipulation of these images.
Bisiach and Luzzatti (1978) provided evidence that unilateral spatial neglect is not only a disorder of visual perception, but also can affect mental representations such that patients fail to report the left side of scenes or objects in mental imagery. However in case reports of representational neglect generally it is accompanied by perceptual neglect. We report a rare occurrence of a patient (NL) who presents a persistent unilateral neglect which appears to be limited to visual imagery. The deficit appeared in tasks which require the formation and manipulation of new visuo-spatial representations as well as those which require access to information about familiar scenes. The patient, who had a lesion in the right parietal lobe, showed no evidence of perceptual or personal neglect, although there was some evidence of visual extinction. We argue that the concept of visuo-spatial working memory can provide a framework within which to interpret aspects of the representational form of neglect, whether or not it is accompanied by perceptual neglect.
We report the case of a patient who, after sequential bilateral strokes in the occipital regions sparing the primary visual cortex, developed a severe deficit of colour perception. At variance with other reports of acquired achromatopsic patients, she showed a perfectly vivid visual imagery for colours. These findings, together with similar data in domains other than colour processing, challenge the theories which posit that the same cognitive processes are involved in both the perception and the retrieval from memory of a given stimulus.
A brain-damaged patient is described whose pattern of performance provides insight into both the functional mechanisms and the neural structures involved in visual mental imagery. The patient became severely agnosic, alexic, achromatopsic and prosopagnosic following bilateral brain lesions in the temporo-occipital cortex. However, her mental imagery for the same visual entities that she could not perceive was perfectly preserved. This clear-cut dissociation held across all the major domains of high-level vision: object recognition, reading, colour and face processing. Our findings, together with other reports on domain-specific dissociations and functional brain imaging studies, provide evidence to support the view that visual perception and visual mental imagery are subserved by independent functional mechanisms, which do not share the same cortical implementation. In particular, our results suggest that mental imagery abilities need not be mediated by early visual cortices.
Signor Piazza, a patient with a left parieto-occipital haemorrhage and a right thalamic stroke, showed severe right personal neglect (e.g. touching own body parts) and right perceptual neglect in tasks with (e.g. cancelling tasks) or without (e.g. description of a complex picture) motor response. He had also right-sided neglect dyslexia (including single words), without language impairments. However, the patient also presented with a clear left-sided deficit in the representational domain (e.g. imagery tasks). Signor Piazza's pattern of performance suggests dissociation between imagery and perception within the neglect syndrome.
We describe a patient, VSB, whose reading was impaired as a consequence of a left temporal-parietal lesion, whereas writing was relatively preserved. At variance with other pure alexic patients described in the literature, VSB claimed to have become unable to mentally visualise letters and words. Indeed, his performance on a series of tests tapping visual mental imagery for orthographic material was severely impaired. However, performance on the same tests was dramatically ameliorated by allowing VSB to trace each item with his finger. Visual mental imagery for non-orthographic items was comparatively spared. The pattern of dissociation shown by VSB between impaired visual mental imagery and relatively preserved motor-based knowledge for orthographic material lends support to the view that separate codes, respectively based on visual appearance and on motor engrams, may be used to access knowledge of the visual form of letters and words.
Visual perception and visual mental imagery, the faculty whereby we can revisualise a visual item from memory, have often been regarded as cognitive functions subserved by common mechanisms. Thus, the leading cognitive model of visual mental imagery holds that visual perception and visual imagery share a number of mental operations, and rely upon common neural structures, including early visual cortices. In particular, a single visual buffer would be used "bottom-up" to display visual percepts and "top-down" to display internally generated images. The proposed neural substrate for this buffer consists of some cortical visual areas organised retinotopically, that is, the striate and extrastriate occipital areas. Empirical support for this model came from the report of brain-damaged patients showing an imagery deficit which parallels a perceptual impairment in the same cognitive domain. However, recent reports of patients showing double dissociations between perception and imagery abilities challenged the perception-imagery equivalence hypothesis from the functional point of view. From the anatomical point of view, the available evidence suggests that occipital damage is neither necessary nor sufficient to produce imagery deficits. On the other hand, extensive left temporal damage often accompanies imagery deficits for object form or colour. Thus, visual mental imagery abilities might require the integrity of brain areas related to vision, but at an higher level of integration than previously proposed.
A patient with unilateral neglect had to evoke mentally the map of France in two different conditions. In the first condition, he was asked to build an iconic representation of the map of France and to list all the towns that he could 'see' on this mental image within two minutes. In the second condition, he had to remember and name as many French towns as possible within two minutes, without being instructed to form a mental image. Left representational neglect was observed in the first condition only, i.e., when an iconic representation was required. These findings, which were replicated four months later, suggest a dual mode of coding, retrieval, or both, of geographic information and show that, although topographic, geographic data has to be spatialized to be neglected.
A complex link exists between vision and unilateral spatial neglect (USN). Firstly, USN is not a perceptual deficit, secondly, USN is not necessarily accompanied by a visual deficit and finally, USN can be observed in non-visual modalities as well as in mental spatial imagery. This apparent supramodality of USN stands in sharp contrast to the fact that neglect signs are often more severe and more durable in the visual than in other sensory modalities (Chokron et al., 2002). The influence of vision on spatial representation has rarely been studied. In the present study we assessed six right brain-damaged patients suffering from left USN on two tasks involving spatial representations: a clock-drawing task and a drawing from memory task in two experimental conditions, with and without visual control. We confirm that even in mental imagery, the absence of visual feedback may decrease and even suppress left neglect signs (Bartolomeo and Chokron, 2001b; 2002). Since vision is largely involved in the orientation of attention in space, suppressing visual control could reduce the magnetic attraction towards the right ipsilesional hemispace and in this way could allow a re-orientation of attention towards the left neglected hemispace. We discuss the theoretical and therapeutic implications of these findings.
When describing known places from memory, patients with left spatial neglect may mention more right- than left-sided items, thus showing representational, or imaginal, neglect. This suggests that these patients cannot either build or explore left locations in visual mental imagery. However, in place description there is no guarantee that patients are really employing visual mental imagery abilities, rather than verbal-propositional knowledge. Thus, patients providing symmetrical descriptions might be using other strategies than visual mental imagery. To address this issue, we devised a new test which strongly encourages the use of visual mental imagery. Twelve participants without brain damage and 12 right brain-damaged patients, of whom 7 had visual neglect, were invited to conjure up a visual mental image of the map of France. They subsequently had to state by pressing a left- or a right-sided key whether auditorily presented towns or regions were situated to the left or right of Paris on the imagined map. This provided measures of response time and accuracy for imagined locations. A further task, devised to assess response bias, used the words "left" or "right" as stimuli and the same keypress responses. Controls and non-neglect patients performed symmetrically. Neglect patients were slower for left than for right imagined locations. On single-case analysis, two patients with visual neglect had a greater response time asymmetry on the geographical task than predicted by the response bias task, but with symmetrical accuracy. The dissociation between response times and accuracy suggests that, in these patients, the left side of the mental map of space was not lost, but only "explored" less efficiently.
Two patients showing left unilateral neglect were asked to describe imagined perspectives of familiar surroundings. Left-sided details were largely omitted in the descriptions. Some theoretical implications of the occurrence of unilateral neglect in representational space are briefly considered.