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Anina N. Rich et al- Neural correlates of imagined and synaesthetic colours

Anina N. Rich et al- Neural correlates of imagined and synaesthetic colours

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Neuropsychologia 44 (2006) 2918–2925
Neural correlates of imagined and synaesthetic colours
Anina N. Rich
, Mark A. Williams
, Aina Puce
, Ari Syngeniotis
,Matthew A. Howard
, Francis McGlone
, Jason B. Mattingley
Cognitive Neuroscience Laboratory, School of Behavioural Science, University of Melbourne, Victoria 3010, Australia
Visual Attention Laboratory, Brigham & Women’s Hospital, Harvard University, 64 Sidney Street, Cambridge, MA 02139, USA
 McGovern Institute, MIT, Cambridge, MA 02139, USA
Centre for Advanced Imaging, Department of Radiology, West Virginia University, USA
 Brain Sciences Institute, Swinburne University of Technology, Australia
 Brain Research Institute, Austin & Repatriation Medical Centre, Australia
Unilever Research Laboratories, Port Sunlight, UK 
Received 23 January 2006; received in revised form 13 June 2006; accepted 15 June 2006Available online 9 August 2006
The experience of colour is a core element of human vision. Colours provide important symbolic and contextual information not conveyed byform alone. Moreover, the experience of colour can arise without external stimulation. For many people, visual memories are rich with colourimagery.Intheunusualphenomenonofgrapheme-coloursynaesthesia,achromaticformssuchasletters,wordsandnumberselicitvividexperiencesof colour. Few studies, however, have examined the neural correlates of such internally generated colour experiences. We used functional magneticresonance imaging (fMRI) to compare patterns of cortical activity for the perception of external coloured stimuli and internally generated coloursin a group of grapheme-colour synaesthetes and matched non-synaesthetic controls. In a voluntary colour imagery task, both synaesthetes andnon-synaesthetes made colour judgements on objects presented as grey scale photographs. In a synaesthetic colour task, we presented letters thatelicited synaesthetic colours, and asked participants to perform a localisation task. We assessed the neural activity underpinning these two differentforms of colour experience that occur in the absence of chromatic sensory input. In both synaesthetes and non-synaesthetes, voluntary colourimagery activated the colour-selective area, V4, in the right hemisphere. In contrast, the synaesthetic colour task resulted in unique activity forsynaesthetes in the left medial lingual gyrus, an area previously implicated in tasks involving colour knowledge. Our data suggest that internallygenerated colour experiences recruit brain regions specialised for colour perception, with striking differences between voluntary colour imageryand synaesthetically induced colours.© 2006 Elsevier Ltd. All rights reserved.
Colour imagery; Synaesthesia; fMRI; Visual imagery; V4; Occipitotemporal cortex
1. Introduction
Colour perception is a fundamental aspect of human vision(Gegenfurtner & Sharpe, 1999).Although several neuroimag- ing studies have explored the involvement of early visualareas in visual imagery (e.g.,D’Esposito et al., 1997;Kosslyn, Thompson,Kim,&Alpert,1995;Kosslyn,Thompson,&Alpert,1997;Mellet et al., 2000;Slotnick, Thompson, & Kosslyn, 2005),few investigations have specifically examined the neu-ral correlates of colour imagery. Several areas in the ventral
Corresponding author. Tel.: +1 617 768 8815; fax: +1 617 768 8816.
 E-mail address:
occipitalcortexareactivewhenparticipantsviewcolouredstim-uli, or perform colour-related cognitive tasks. These includearea(s) in the posterior fusiform gyrus, most often called areaV4 (Zeki, 1973; Zeki et al., 1991),and more medial ventral occipital regions that are active particularly during tasks thatrequire retrieval of object-colour knowledge (Chao & Martin,1999;Martin, Haxby, Lalonde, Wiggs, & Ungerleider, 1995; Price, Moore, Humphreys, Frackowiak, & Friston, 1996).Here,we used fMRI to examine neural activity in these regions intwo distinct types of internally generated colour experience.First,weexamined
byhavingparticipantsimag-ine the colours of familiar objects. Second, we investigated theinvoluntarycolourexperiencesthatcharacterisetheunusualphe-nomenon of 
grapheme-colour synaesthesia.
0028-3932/$ – see front matter © 2006 Elsevier Ltd. All rights reserved.doi:10.1016/j.neuropsychologia.2006.06.024
 A.N. Rich et al. / Neuropsychologia 44 (2006) 2918–2925
Individuals with grapheme-colour synaesthesia experiencevivid sensations of colour when they see or hear particularletters or digits, and these experiences are highly consistentover time (e.g.,Baron-Cohen, Wyke, & Binnie, 1987;Rich, Bradshaw, & Mattingley, 2005;Rich & Mattingley, 2002). Crucially, synaesthetic colours arise as an internally generatedcolour experience without voluntary effort, and are difficult tofully suppress even when there is a desire to do so (Dixon,Smilek, Cudahy, & Merikle, 2000;Mattingley, Rich, Yelland,& Bradshaw, 2001;Mills, Boteler, & Oliver, 1999;Odgaard, Flowers, & Bradman, 1999;Wollen & Ruggiero, 1983).Thus, grapheme-colour synaesthesia provides a unique opportunityto examine the neural correlates of an internally generated butinvoluntary colour experience.The neuroanatomical loci of colour imagery remain contro-versial. A number of neuropsychological studies have docu-mented an association between impairments of colour percep-tion and deficits in colour imagery in patients with damage tothe fusiform gyrus (for review, seeFarah, 1988).Other authors, however, argue for a double dissociation between the two func-tions based on patients who seem to have intact imagery in thecontextofcolourperceptiondeficitsor
(forreview,seeBartolomeo, 2002).Very few studies have explicitly examinedthe neural correlates of colour imagery in healthy individuals,andthedatatheyhaveyieldedaresomewhatambiguous.Howardet al. (1998)used fMRI to investigate brain activity during acolourimagerytask.Theypresentednon-synaestheteswithtasksthat required them to imagine colour (e.g., to judge whether acanary is a darker yellow than a banana) or to imagine a set of spatial relations (e.g., to judge the relative angle between thehour and minute hands on a clock face showing “20 to 7”).Howard et al. found that although coloured stimuli elicited sig-nificant activity in the posterior fusiform gyrus, correspondingto area V4, colour imagery yielded no significant activity in thesame region. It should be noted, however, that the authors useda whole-brain analysis, which may have reduced the likelihoodof detecting a subtle change in V4 activity.In a more recent study,Nunn et al. (2002)ailed to find sig- nificant V4 activation in a group of non-synaesthetes trainedon associations between coloured squares and spoken words. Intheir study, participants were asked to imagine the colour theyhad been trained to associate with each word, and then to esti-mate the percentage of colour associations they had correctlyimagined. Unfortunately, Nunn et al. did not include an objec-tive measure of their participants’ performance. Moreover, itis not possible to determine whether they were able to imag-ine the colours, or simply recalled the colour name associatedwith the spoken word. The same criticism holds for an ear-lier study using similar methodology (Goldenberg et al., 1989).The hypothesis that colour imagery recruits regions involved incolourperceptionthereforeneedstobetestedwithaparadigminwhich performance can be measured objectively, and in whichparticipants must imagine colour, rather than merely recallingcolour names.There have also been a small number of studies investigatingthe neural correlates of involuntary colour experiences. Patientswith Charles Bonnet syndrome, who experience visual hallu-cinations following peripheral visual pathology, show activityin the posterior fusiform gyrus for hallucinations that involvecolour (Ffytche et al., 1998).For individuals with synaesthe- sia, who experience colours without chromatic stimulation inthe absence of nervous system pathology, the findings of imag-ing studies have been mixed. Recent studies indicate that, atleast in some individuals, area V4 is active during grapheme-colour synaesthesia.Sperling, Prvulovic, Linden, Singer, andStirn (2006)ound increased activity in retinotopically mappedV4 when synaesthetes viewed achromatic letters that inducedinvoluntarycoloursrelativetoaconditionwithlettersthatdidnotinduce coloured synaesthesia. This pattern was significant fortwo of the four synaesthetes when analysed individually. Sim-ilarly,Hubbard, Annan, Ramachandran, and Boynton (2005)reported that synaesthetes had greater V4 activity than con-trols for synaesthetic inducers relative to characters that did notinduce synaesthesia. Activity in left V4 has also been docu-mented in a late-blind synaesthete, with more anterior fusiformactivation in the right hemisphere when the same participantwas asked to imagine colours (Steven, Hansen, & Blakemore,2006).Finally,Nunnetal.(2002)reportedincreasedV4activity when synaesthetes heard words that elicited colours comparedwithnon-inducingtones.Incontrast,otherstudiesofsynaesthe-sia have failed to find significant activation in colour-selectiveregions of cortex, and have instead documented activations invisual association cortex, including inferior temporal and infe-rior parietal areas (Paulesu et al., 1995;Weiss, Zilles, & Fink, 2005).These findings suggest that colour-selective areas of cor-texarerecruitedduringinternallygeneratedcolourexperiences,butthattheremaybedifferencesbetweencolourimageryontheone hand, and synaesthetic experiences on the other.In the present study we used fMRI to characterise brainactivation patterns associated with internally generated colourexperiences in both synaesthetes and non-synaesthetic con-trols. We first identified regions in the ventral occipital cortexthat responded selectively to colour by presenting large mul-ticoloured rectangles (‘Mondrians’) in alternating blocks withgrey-scale luminance equivalent stimuli (McKeefry & Zeki,1997).Using the peak activations as centres for regions of inter-est (ROIs), we then analysed data within these regions fromseparate colour imagery and synaesthetic colour experiments toexamine the neural correlates of these colour experiences in theabsence of chromatic stimulation.In one experiment, we had participants perform a
colour imagery
task, to examine the neural correlates of voluntar-ily generated colour experiences. Participants viewed greyscalephotographsofpairsofobjectsthathaveatypicalor’canonical’colour (e.g., a banana and a cob of corn;Fig. 1A). Participants wereaskedtojudgewhichobject,ifseeninitsnaturalchromaticform, would be darker in colour (colour imagery task); in a con-trol condition they were asked to indicate which object wouldbe larger in size (size baseline task). The colour imagery task wasdesignedtomaximisethelikelihoodthatparticipantswouldvoluntarily evoke colour images rather than simply drawing onstored verbal colour labels concerning familiar objects. Thus,for example, simply knowing that bananas are yellow shouldnot help in determining whether they are a darker yellow than a
A.N. Rich et al. / Neuropsychologia 44 (2006) 2918–2925
Fig. 1. Example stimuli and fMRI activations for the voluntary colour imagery experiment. (A) Example stimuli for the colour imagery and size (baseline) tasks.Participants judged which object would be darker in colour or larger in size. (B) Activity within the functionally defined ROIs for the colour imagery vs. baselinecomparison. Brain slices show regions of significantly greater activity for control participants (yellow) and synaesthetes (red). For controls significant activationoccurred in the right inferior occipital lobe (
: [26
=0.016); for synaesthetes significant activation occurred in the right inferior occipital lobe (
<0.001) and right fusiform gyrus (
: [30
=0.001). Coordinates in this and the subsequent figure have been calculated using the MNIthree-dimensional space.
cob of corn. The baseline task, on the other hand, was designedto require a similar judgement of a stimulus attribute that wasnot related to colour. We used identical pairs of pictures in thetwo tasks to ensure that any changes in brain activity could notbe due to low-level luminance, intensity, contrast, or featuraldifferences between stimuli.In a second experiment, synaesthesia-inducing letters werepresented either in the matching or ‘congruent’ colour foreach synaesthete, or in shades of grey (‘achromatic’) equatedfor luminance. These conditions were compared with a base-line condition of greyscale placeholders (Fig. 2A). A group of  sex-,age-andhandedness-matchednon-synaesthetesviewedtheidentical stimuli. Participants were required to detect the brief disappearance of one stimulus in the set of four presented ineach display in a localisation task.We hypothesized that if regions of the brain specialised forcolour perception are also involved in colour imagery, bothsynaesthetes and controls should show significant activationwithin the ROIs defined by the colour localiser. Similarly, if synaesthetic colours recruit areas involved in colour perception,significantactivationshouldbepresentwithinthesameROIsforsynaesthetes, but not for controls, during presentation of achro-matic letters.
2. Methods
2.1. Participants
Seven grapheme-colour synaesthetes (6 female; 5 right-handed; meanage=28years,S.D.
7.1,range:18–37years),andsevennon-synaestheticcon-trols matched for age, sex and handedness (mean age=27 years; S.D.=7.4;

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