Differential effects of visual attention and working memoryon binocular rivalry

Lisa Scocchia $

Frankfurt Institute for Advanced Studies,Johann Wolfgang Goethe University, Frankfurt am Main,

GermanyGiessen University, Department of Psychology, Giessen,

GermanyUniversity of Milano-Bicocca, Department of Psychology,

Milan, Italy

Matteo Valsecchi # $Giessen University, Department of Psychology, Giessen,

Germany

Karl R. Gegenfurtner # $Giessen University, Department of Psychology, Giessen,

Germany

Jochen Triesch # $

Frankfurt Institute for Advanced Studies,Johann Wolfgang Goethe University, Frankfurt am Main,

Germany

The investigation of cognitive influence on binocularrivalry has a long history. However, the effects of visualWM on rivalry have never been studied so far. Weexamined top-down modulation of rivalry perception infour experiments to compare the effects of visual WMand sustained selective attention: In the first threeexperiments we failed to observe any sustained effect ofthe WM content; only the color of the memory probewas found to prime the initially dominant percept. InExperiment 4 we found a clear effect of sustainedattention on rivalry both in terms of the first dominantpercept and of the overall dominance when participantswere involved in a tracking task. Our results provide anexample of dissociation between visual WM andselective attention, two phenomena which otherwisefunctionally overlap to a large extent. Furthermore, ourstudy highlights the importance of the task employed toengage cognitive resources: The observed perceptualepiphenomena of binocular rivalry are indicative ofvisual competition at an early stage, which is notaffected by WM but is still susceptible to attentioninfluence as long as the observer’s attention isconstrained to one of the two rival images via a specificconcomitant task.

Introduction

Binocular rivalry occurs when incompatible monoc-ular images are displayed at the same retinal location inthe two eyes: perception oscillates between the twoalternative interpretations, rather than yielding a stableintermingled percept. As the perceptual epiphenome-non keeps changing over time whereas the physicalstimulus remains invariant, binocular rivalry can beusefully employed as a tool to explore the dynamicalfeatures of visual awareness and how they vary with thestate of the observer. Indeed, binocular rivalry can beinfluenced by a number of cognitive, motivational andaffective factors. Stable characteristics of the observers,such as their past experience and even their sociocul-tural context can affect rivalry dominance: Uprightpictures of human faces prevail over inverted ones(Engel, 1956; Hastorf & Myro, 1959), familiar imagesdominate over novel ones (Goryo, 1960), and emo-tional facial expressions dominate over neutral expres-sions (Alpers & Gerdes, 2007); Jewish and Catholicreligious images dominate for respectively Jewish andCatholic observers (LoSciuto & Hartley, 1963), picturesdepicting violent acts (e.g., rape or murder) are

Citation: Scocchia, L., Valsecchi, M., Gegenfurtner, K. R., & Triesch, J. (2014). Differential effects of visual attention and workingmemory on binocular rivalry. Journal of Vision, 14(5):13, 1-15, http://www.journalofvision.org/content/14/5/13, doi:10.1167/14.5.13.

Journal of Vision (2014) 14(5):13, 1–15 1http://www.journalofvision.org/content/14/5/13

doi: 10 .1167 /14 .5 .13 ISSN 1534-7362 � 2014 ARVOReceived May 17, 2013; published May 30, 2014

perceived more often than are neutral control picturesby policemen and institutionalized offenders than bycontrol observers (Shelley & Toch, 1962; Toch &Schulte, 1961). Binocular rivalry can also be affected bymore volatile cognitive factors: Voluntary intentionand attention are known to affect observers’ ability toincrease alternation rate during rivalry (Lack, 1971;Meng & Tong, 2004; van Ee, van Dam, & Brouwer,2005). However, their effect on rivalry dominance ismore controversial (Chong & Blake, 2006; Chong,Tadin, & Blake, 2005; Lack, 1978; Meng & Tong, 2004;Ooi & He, 1999; van Ee et al., 2005). Furthermore,diverting the observer’s attention to a concurrent taskduring rivalry has been shown to slow down alternationrate (Paffen, Alais, & Verstraten, 2006). Finally,Pearson, Clifford, and Tong (2008) found that endog-enously generated mental images can exert facilitatoryeffects on binocular rivalry stabilization: In case ofintermittent presentation of rivalry displays, perceptualdominance tends to stick to the same stimulus thatappeared dominant on the preceding trial. Whenobservers are required to imagine one of the rivalrystimuli during the blank period between presentations,perception is biased in favor of the imagined stimulus.

The contents of visual working memory (WM) areone of the major, ever-changing aspects that couldaffect the cognitive and perceptual state of theobserver. Indeed, growing experimental evidence sup-ports the notion that holding in WM a visual object notonly influences the processing speed of subsequentlypresented objects (Downing, 2000; Robinson, Manzi, &Triesch, 2008; Pan & Soto, 2010; Soto, Heinke,Humphreys, & Blanco, 2005; Turatto, Vescovi, &Valsecchi, 2008), but can also bias the appearance ofthe new sensory input (Kang, Hong, Blake, &Woodman, 2011; Scocchia, Cicchini, & Triesch, 2013;Scocchia, Valsecchi, Gegenfurtner, & Triesch, 2013).However, in the long history of binocular rivalry thequestion of whether visual WM contents can influenceperceptual dominance has never been posed: Thepresent study investigates precisely this issue. We hadour observers memorize an image and hold it inmemory for a few seconds in the context of a delayedmatch to sample task. In this way, we could testwhether voluntarily maintaining a visual representationin WM has an impact on rivalry dominance.

The idea that the contents of WM can modify theappearance of what we see, especially when sensoryinformation is ambiguous enough to generate bistableperception, stems from a host of experimental evidenceshowing that the same cortical areas that underlieperceptual processing play an important role in main-taining visual information in WM. This overlap betweentheneural networks that encode the visual input and thoseresponsible for itsmaintenance inWMfor several secondsafter its offset has been demonstrated both at late and

early stages of visual processing:Neurons in face-selectiveareas of inferotemporal cortex showed sustained changesin their activity profile when a monkey observer wasretaining a face in WM (Miller, Li, & Desimone, 1993);firing activity in V1, MT, and V4 neurons was alsoobserved during the WM retention interval when theanimal memorized location, motion, and color (orluminance) information respectively (Bisley, Zaksas,Droll, & Pasternak, 2004; Motter, 1994; Super, Spek-reijse, & Lamme, 2001). Along the same lines, humanstudies indicated that activation patterns in V1 duringWM retention reflected the specific attributes thatobservers were required to memorize (Serences, Ester,Vogel,&Awh, 2009), that activity in areasV1 toV4 couldpredict which of two oriented gratings was held in WM(Harrison & Tong, 2009), and that motion informationheld inWMcouldmodulate phosphene reportwhenTMSwas applied to human V5 (Silvanto & Cattaneo, 2010).

If we hypothesize thatWMcontents can impact rivalryperception via the recruitment of sensory areas thatmediate visual processing, we could expect two possibleoutcomes. On one hand, prolonged engagement ofsensory networks could lead to satiation processes (neuralfatigue) in those networks and therefore produce adominance shift to the rival image least overlapping withthememory sample. Suchanoutcomewouldbeconsistentwith recent evidence of repulsive effects observed in non-ambiguous conditions. Kang et al. (2011) employedmoving random dots configurations as stimuli and askedobservers to memorize their direction for delayeddiscrimination:They found that thedirectionofmotionofthe stimulus displayed during the retention interval wasbiased away from thememorized one. Scocchia, Cicchini,and Triesch (2013) required participants to memorizeoriented gratings and involved them in an orientationdiscrimination task during the retention interval: Per-ceived orientationwas repelled away from thememorizedone across two subsequent discrimination judgments. Onthe other hand, sustained engagement of sensory net-works could lower their activation threshold duringrivalry, thus favoring dominance of the rival image mostoverlappingwith thememory sample.Consistentwith thisview, a recent study of ours (Scocchia, Valsecchi,Gegenfurtner, & Triesch, 2013) showed that the contentsof WM can bias bistable perception through facilitation:An ambiguously rotating sphere was perceived tomove inthe samedirectionas apreviously presentedunambiguoussphere significantly more often when participants mem-orized its speed than when they merely attended to it.

Experiment 1

The goal of Experiment 1 was to test whetherholding in WM a specific image could bias perception

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 2

of subsequently presented binocular rivalry stimuli. Tothis purpose, we employed three categories of stimuli(houses, faces, and airplanes) to test WM performancein a delayed match to sample task and a classical house-face rivalry display to be presented during the retentioninterval (see Figure 1A). Importantly, one of the rivalrystimuli could match the memorized stimulus (memorysample) from trial to trial. If WM top-down modula-tion can exert its influence over binocular rivalry via adirect recruitment of neural networks that underpinperception itself, we could expect two possible scenar-ios:

(a) The visual WM load may lead to saturation of thosenetworks. As a result, we should observe prolongedsuppression of the memory sample during rivalry.If, for instance, the observer memorized a face,sustained activity in the fusiform face area due to

WM maintenance could lead to a greater domi-nance of the opposite stimulus, the house, and evenmore so when the rivalry face exactly matches thememorized one.

(b) The WM load may prime the perceptual networksduring WM maintenance, without overloadingthem. If the observer memorized a face, perceptionof the face rival stimulus would prevail in case ofcategory match and even more so in case of exactstimulus match.

Methods

Observers

Observers consisted of 14 undergraduate studentsfrom Giessen University who received payment forparticipation. All observers had normal or corrected-

Figure 1. Experiment 1 stimuli and procedure. (A) Stimuli employed in the WM task. A black frame (not displayed during the

experiment) identifies the WM stimuli that exactly matched the rivalry stimuli. For clarity, stimuli are numbered within each category.

(B) Illustration of a trial sequence of Experiment 1. Observers were required to memorize the initial image, which could belong to one

of three different categories (face, house, or airplane, interleaved across trials). Subsequently, they continuously reported the

perceived image during rivalry (rival stimuli are indicated by a brace). Finally, they judged whether or not the memory test matched

the memory sample. The house-face rival stimuli were the same across trials, whereas the memory stimuli were drawn from the

image pool in a random order. The memory test belonged to the same category as the memory sample and they matched in 50% of

the trials. The display duration is indicated below each stimulus window. Stimuli are not drawn to scale and instructions were

presented as actual sentences.

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 3

to-normal visual acuity, and were naıve to the purposeof the experiment. The experiment was conducted inaccordance with the Declaration of Helsinki.

Apparatus and stimuli

Stimuli were created using Matlab in conjunctionwith the Psychophysics Toolbox (Brainard, 1997) andwere displayed on two 19-in. LCD color monitors(Delle UltraSharp 1907FP) at a viewing distance of55.5 cm. Stimuli for the left eye were presented on theleft monitor screen and stimuli for the right eye on theright monitor screen. Observers viewed the monitorsthrough a Wheatstone mirror stereoscope, consisting oftwo First Surface Mirrors (169 · 194 mm) to bring thetwo views into alignment. Physical luminance equiva-lence between the left and the right display was checkedthrough the mirrors with a Photo Research PR-650spectroradiometer. The observer’s head was stabilizedby a chin and forehead rest.

Rivalry stimuli consisted of a red image of a houseand a green image of a face subtending 3.988 · 3.938 ofvisual angle and were displayed on a gray background(30.0 cd/m2). During rivalry, a white fixation dot (0.098· 0.098) was placed at the center of each image and ablack frame (0.188 thick) surrounded each monocularimage to aid binocular fusion. Rivalry stimuli, memorysamples, and memory tests were drawn from a set offive stimuli for each of the three categories. Whereas thememory sample and test images varied across trials,rivalry stimuli were fixed. Stimuli were displayed asindividual objects, in the absence of backgroundinformation. House and airplane stimuli were createdon the basis of public images downloaded from thewww.flickr.com website, whereas face images weredrawn from the Georgia Tech face database (Nefian,1999), after removal of background information.Illustrative representations of the stimuli are displayedin Figure 1A.

Faces, houses, and airplanes were presented on agreen, red, and yellow background, respectively.Stimuli had the same mean luminance, and the rootmean square contrast within each figure was fixed at20%.

Calibration procedure

Each participant underwent a preliminary calibra-tion procedure of the rivalry stimuli in order to identifythe RMS contrast that yielded equivalent subjectivedominance rates for the two stimuli during a baselinerivalry task. This procedure served to determine thecontrast at which the face stimulus should be displayedto match the house stimulus in baseline exclusivedominance during the main experiment. The calibra-tion procedure was conducted before every experiment

employing house-face rivalry presented from then on.On the basis of pilot data, we tested seven levels ofRMS contrast for the face stimulus (18.3%, 16.7%,15%, 13.3%, 11.7%, 10%, and 8.3%) against a fixed 20%RMS contrast for the house stimulus. Each facecontrast was presented five times to each eye in acounterbalanced fashion, over 70 trials. Each trialstarted with the presentation of a white fixation dot(0.098 · 0.098) for 0.5 s, followed by the display of therivalry stimuli for 30 s. Participants were required tokeep fixation on the white dot and to press the left orthe right arrow key only when they exclusivelyperceived the house or the face, respectively, as long asexclusive perception lasted. They were asked to releasethe button in case of mixed dominance. Participants’responses were sampled at 100 Hz. At the end of therivalry stimulus presentation, a blank gray window wasdisplayed until the observer pressed a key to proceed tothe next trial. Individual proportions of ‘‘face domi-nant’’ responses were computed at each level of facestimulus contrast. For each observer, face dominanceproportions were linearly regressed over stimuluscontrast to find the contrast level at which baselinedominance for house and face were equal. Participantswho at this stage reported problems with fusing the twoimages, whose pattern of responses clearly deviatedfrom linearity or direct proportionality, or for whomthe interpolation procedure yielded values below 5% ofRMS contrast were excluded from the main experi-ment. On this basis, two participants were excluded andthe experiment was conducted on the remaining 12participants.

Experimental procedure

Participants were instructed to memorize an imagethat could belong to one of three different categories(face, house, or airplane) for a 2AFC delayed match tosample. Figure 1B illustrates a trial sequence ofExperiment 1. At the beginning of each trial, theinstruction to memorize the stimulus was presented for0.7 s, followed by a 1 s blank interval and then by thememory sample, which was displayed for 0.5 s.Afterwards, the fixation dot was shown for 3 s,followed by the 30 s rivalry stimulus. During itspresentation, participants were required to monitorsubjective dominance by pressing the left or the rightarrow key (as they did during the calibration session).After a 1 s blank interval, the memory test wasdisplayed until participants provided their response:They pressed the up arrow key to indicate that thememory test matched the memory sample and thedown arrow key to indicate that they were different.The WM test matched the sample in 50% of cases; inthe remaining 50% it was randomly selected from thesame category set. A 1.5 s blank interval followed

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participants’ response before the next trial began:Observers were allowed to take a break every five trials.

Each WM sample was presented 16 times, for a totalamount of 240 trials. Since the two rivalry stimuli weredriven from the same image pool as the memory andthe test stimuli, the memory sample matched one of therival images in 32 trials. Individual data were collectedover five sessions (including calibration session) lastingabout one hour each.

Data analysis

In the four experiments reported in this study, wecalculated dominance proportions for each participantand each experimental factor as the ratio of mutuallyexclusive responses (e.g., ‘‘house’’ or ‘‘face,’’ sampledat 100 Hz) over periods of exclusive dominance,collapsed across trials. In other words, all theinstantaneous keypresses relative to a particularobserver and experimental factor yielded a singleproportion dominance value that was computed afterexcluding periods of mixed dominance. The use ofdominance proportion has the advantage of combin-ing the effects of the number of episodes of exclusiveperceptual dominance across trials and of theiraverage duration. These measures are known to showa large intersubjective variability (Levelt, 1965).Instead, the employment of a comprehensive measureas dominance proportion was meant to control forindividual differences in alternation rate and toimprove statistical sensitivity to possible differencesbetween conditions. We analyzed the observers’reports in terms of dominance proportion rather thanof dominance durations: Dominance durations are nota robust measure in our paradigm since single episodeswere often truncated (between 3% and 23% of thetimes, depending on participant and experimentalcondition), because the display time of the rivalrystimulus was considerably shorter in our experimentsthan what is typically reported in the literature, whererivalry is displayed for 1 min or longer (e.g.,Brascamp, van Ee, Noest, Jacobs, & van den Berg,2006; Chong et al., 2005; Levelt, 1965; Meng & Tong,2004). However, results do not change significantlywhen the analyses are conducted on normalizeddominance durations.1 For sake of clarity, we willreport dominance proportions in terms of prevalenceof house responses from now on. In order todistinguish between sustained and immediate effects ofthe WM sample presentation on perceptual domi-nance during rivalry, we also analyzed the firstpercepts of each trial: We reported the number oftrials where ‘‘house’’ was perceived first over the totalnumber of trials for each participant, WM identityand category.

Results and discussion

Participants understood and adequately performedthe task, scoring on average 93.4% correct responses atthe memory test. On average, observers reported seeingthe house for 53.1% of time during exclusive domi-nance and they perceived exclusive dominance for40.85% of the time during rivalry. The left panel ofFigure 2 shows the proportion of ‘‘house’’ responsesaveraged across observers, separately for each WMstimulus and category. A one-way, repeated-measuresANOVA on the proportion of relative dominanceacross categories failed to highlight any significanteffect, F(2, 22) ¼ 1.76, p ¼ 0.2. We also compared theproportion of ‘‘house’’ responses given when the WMstimulus matched the rivalry image and the average ofthe other stimuli within the same category in order totest the effects of stimulus identity. Dominanceproportion did not differ according to identity matchboth within the house, t(11) ¼ 1.01, p ¼ 0.33, and theface category, t(11)¼ 0.73, p¼ 0.48. The right panel ofFigure 2 illustrates the proportion of trials whereparticipants reported the house as the first dominantpercept. Data are averaged across the 12 observers andpresented separately for WM stimulus category andidentity. A one-way, repeated-measures ANOVA onthe proportion of ‘‘house first’’ responses highlighted asignificant difference across categories, F(2, 22)¼ 4.92,p¼ 0.017. The Duncan post-hoc tests further showedthat observers were more likely to report ‘‘house’’ as afirst percept after memorizing a house as opposed toboth a face (p , 0.01) and an airplane (p , 0.05).Direct comparisons of the proportion of ‘‘house first’’responses between the WM stimulus that matched therivalry image and the average of the other stimuliwithin category also failed to highlight any effect ofstimulus identity, t(11) ¼ 1.3, p ¼ 0.22 for the housecategory; t(11) ¼�0.22, p¼ 0.83 for the face category.

Our results point out that the effect of the memorysample on subsequent rivalry dominance is short-livedand can be detected only in terms of the influence of theWM content on the stimulus perceived first duringrivalry. Instead, when considering the overall amountof exclusive dominance perceived during rivalry, nodifferences emerged across conditions. Furthermore,the effect of the WM sample on initial dominance waslimited to the WM stimulus category and did notencompass its identity. Although our goal in thisexperiment was to test the effect of voluntarily holdingan object in visual WM on subsequent rivalryperception, we cannot exclude that we actually tackledimplicit memory processes, such as priming. Thisaccount of Experiment 1 results seems plausible if weconsider that the WM task turned out to be relativelyeasy for participants: As such, it may have posed onlylimited cognitive demands, which would not require

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 5

holding an active visual representation of the stimulusin WM. However, the experimental outcome seemsincompatible with stimulus priming, as stimulus identitydid not play a role in the observed effect. Rather, wecould explain our results in terms of color priming, asthe WM samples and the rivalry stimuli shared thesame color background (see Figure 1) in the case ofhouse and face images. Experiment 2 was designed tobetter address whether visual WM contents may affectrivalry perception and whether color priming couldexplain the observed results.

Experiment 2

In the first experiment we could not detect a cleareffect of the WM stimulus identity; nevertheless, weobserved an influence of its category on subsequentinitial rivalry dominance. The conclusions we canderive from these results are limited for two reasons:First, participants were on average 93.4% accurate atthe memory task. Such a ceiling effect indicates thatthe task was not challenging enough, and it may nothave been necessary for participants to hold an activevisual representation in WM to solve it. Second, as thehouse and the face images had the same colorbackground both when used as rivalry and as WMstimuli, the observed effect of category on rivalryinitial dominance may be due to color priming. InExperiment 2, we employed a different and larger setof stimuli and a more challenging WM testingprocedure, which aimed to favor holistic visualencoding of the WM samples and yielded an average74.2% memory accuracy. Furthermore, we employedgrayscale stimuli for the WM task to exclude thepossibility of color priming.

Methods

Observers

A new pool of seven paid volunteers participated inthis experiment. All of them had normal or corrected-to-normal visual acuity, and were naıve to the purposeof the experiment. The experimental setup was the sameas described in Experiment 1.

Stimuli and procedure

Two hundred and forty different grayscale images ofhouses, faces, and cars (80 for each category) wereemployed as WM samples and tests. Face images werea selection from the face database provided by theMax-Planck Institute for Biological Cybernetics inTuebingen, Germany (Troje & Bulthoff, 1996): Theyrepresented male heads with no hair, glasses, beard,moustaches, or paraphernalia. The head models of thedatabase were obtained by morphing real scans toavoid close resemblances to actual individuals. The carsand houses databases were created ad hoc from thewww.flickr.com website. All the stimuli were displayedas single objects, in the absence of any informativebackground. WM stimuli subtended 6.128 · 68 ofvisual angle, whereas rivalry stimuli subtended 3.988 ·3.938. Rivalry stimuli were surrounded by a blackframe (0.188 thick) and a white fixation dot (0.098 ·0.098) was placed at the center of each monocularimage. The rival house and face stimuli were displayedon a red and on a green background, respectively, andthe baseline contrast for the face stimulus wasindividually determined via the calibration procedure,as described for Experiment 1.

The experimental procedure is illustrated in Figure 3.The rival stimuli were constant across trials and theywere drawn from the same image databases as the

Figure 2. Results of Experiment 1. Left panel: proportion of ‘‘house’’ responses as a function of memory sample. Right panel: the

proportion of trials participants reported the house as the first percept during rivalry is plotted as a function of the WM sample,

separately for each stimulus category. Red and green arrows indicate respectively the house and the face WM sample that

corresponded to the rival stimulus. WM stimuli within each category are numbered as is in Figure 1. Error bars represent the SEM.

and black, dashed lines indicate chance level.

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 6

house and the face WM stimuli: At variance withExperiment 1, no WM sample matched any of the rivalstimuli. On each trial, a new WM sample waspresented: The WM test stimulus was equal to the WMsample in 50% of trials; in the remaining 50% itrandomly matched one of the other memory sampleswithin the same category.

The WM test display was partially occluded andpresented as the upper left, upper right, lower left, orlower right corner of the image, in a counterbalancedand randomly ordered fashion for each category. Thismeasure was taken to render the WM task morechallenging and to favor holistic visual encoding of theWM sample. The displayed corner of the WM testcovered 49% of the original stimulus (including neutralbackground): This value was chosen on the basis of apilot experiment where eight naıve participants wereinvolved in a two-back memory task on the same set ofstimuli. The test image was displayed at eight differentlevels of occlusion (70%, 63%, 56%, 49%, 41%, 34%,27%, or 20% visible) and the 49% level yieldedcomparable performance across categories and ob-servers, averaging 73.5% correct. The experimentalprocedure was the same as in Experiment 1, except thatthe display time of the rivalry stimulus was limited to15 s. Altogether, individual data were collected in foursessions (including calibration session) lasting about45–60 min each.

Results and discussion

Accuracy at the WM task averaged 74.2% correctresponses, implying that the task engaged participants’cognitive resources. On average, observers reportedseeing the house for 51.88% of the time during exclusivedominance, and they perceived exclusive dominance31.84% of the time during rivalry. The left panel ofFigure 4 illustrates the relative dominance of ‘‘house’’responses as a function of stimulus category held inWM. We tested the influence of WM stimulus categoryon dominance proportion via a one-way, repeated-measures ANOVA and observed no effect, F(2, 12) ¼

0.41, p¼ 0.67. A direct comparison between the houseand the face category also failed to reach statisticalsignificance, t(6)¼ 0.73, p¼ .49. The right panel ofFigure 4 illustrates the proportion of trials whereparticipants reported the house as the first dominantpercept, separately for each WM stimulus category.

A one-way, repeated-measures ANOVA on theproportion of ‘‘house first’’ responses also failed tohighlight any significant difference between categories,F(2, 12) ¼ 1.23, p ¼ 0.33, as did the direct comparisonbetween the house and the face category, t(6)¼ 1, p ¼0.35.

These results indicate that holding in WM a visualrepresentation does not favor nor hinder perception ofthe rival stimulus belonging to the same category.Rather, we conclude that Experiment 1 results arebetter described in terms of color priming, which wouldalso explain why stimulus identity did not play a role inthe observed effect.

Experiment 3

The null result reported in Experiment 2 stands instark contrast to the findings of our previous study(Scocchia, Valsecchi, Gegenfurtner, et al., 2013), wherewe showed that WM contents can affect the perceiveddirection of ambiguous structure-from-motion. In thatstudy, participants had to memorize the speed of anunambiguously rotating sphere and were shown anambiguous transparent sphere during the retentioninterval: They reported the perceived direction ofmotion on a moment-to-moment basis, as in theexperiments described here. Dominance systematicallyshifted towards the direction of motion that matchedthe memorized stimulus. In Experiments 1 and 2 of thecurrent study, rivalry dominance was not influenced bythe memory of object identity (house, face), a muchmore high-level representation compared to the motionspeed which was memorized in our previous study(Scocchia, Valsecchi, Gegenfurtner, et al., 2013).However, it is still possible that if the memorized

Figure 3. Trial procedure of Experiment 2. Participants were required to memorize the stimulus for a delayed match to sample. During

the retention interval, they were presented with the rivalry stimuli and continuously monitored perceptual dominance. The memory

test was displayed as the upper left, upper right, lower left, or lower right corner of the test image, covering 49% of its original area.

The display duration is indicated below each stimulus window. Stimuli are not drawn to scale and instructions were presented as

actual sentences.

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 7

stimulus were encoded in terms of low-level features(e.g., speed, spatial frequency, orientation. . .) ratherthan face or object identity, binocular rivalry domi-nance could be affected. Therefore, in Experiment 3 wetested whether a direct match in low-level visualfeatures between the WM and either of the rival stimuliwould impact rivalry dominance. We asked partici-pants to memorize the spatial frequency of a gratingand displayed two rival orthogonal gratings during theretention interval: The memory sample had theorientation of either rival stimulus. To favor directcomparison, we employed a similar paradigm to theone reported in our previous study (Scocchia, Valsec-chi, Gegenfurtner, et al., 2013) and added a controlcondition where participants attended to, but did notmemorize, the first grating stimulus.

Methods

Observers

A new pool of 12 paid volunteers participated in thisexperiment. All of them had normal or corrected-to-normal visual acuity, and were naıve to the purpose ofthe experiment. The experimental setup was the same asdescribed in Experiment 1.

Stimuli and procedure

The rivalry stimuli consisted of a green and of a red6458 sinusoidal gratings at 50% Michelson contrastthat subtended 18 of visual angle and were displayed ona grey background. The average luminance of each ofthe two gratings and of the background was 13.8 cd/m2.The spatial frequency of the gratings was 3.25 cyclesper degree (cpd). Eye of presentation, grating color,and orientation were independently manipulated. Thenon-rival stimuli (i.e.: the memory sample and tests inthe WM condition and the attention stimulus in thecontrol condition) were grayscale gratings (18, 13.8 cd/

m2) that matched the orientation of either rivalstimulus, in a counterbalanced fashion with a randompresentation order. All the stimuli were displayedwithin a 1.58 checkerboard frame to aid fixation.

In the WM condition, the memory sample and testhad the same orientation within a trial. The spatialfrequency of the sample ranged randomly between 1.94and 4.95 cpd, whereas the spatial frequency of thememory test was determined by a staircase algorithmaiming at 75% of correct responses as participants wereinvolved in a delayed match to sample. The experi-mental procedure of the WM condition is depicted inFigure 5A: Each trial began with the presentation ofthe checkerboard, which was displayed thereafter untilthe rivalry stimulus offset. After 2 s the memory samplewas shown for 0.5 s; then the rivalry stimulus waspresented for 30 s after a 3 s inter-stimulus interval.Participants monitored perceptual dominance withtheir right hand as in the previous experiments: Theypressed the right or the left arrow key according to theperceived orientation of the stimulus. They were askedto refrain from responding in case they perceived mixeddominance. The memory test appeared after a 1 s blankinterval and was displayed until response. Participantsjudged the test stimulus spatial frequency (i.e.: theywere asked whether the distance between the gratingbands was the same as in the memory stimulus); theyused their left hand to press the ‘‘s’’ or the ‘‘d’’ key toprovide ‘‘same’’ or ‘‘different’’ responses, respectively.Afterwards a blank window was presented for 1.5 s;then started the next trial.

The experimental procedure in the attention conditionis depicted in Figure 5B; it was the same as in the WMcondition, with the following exceptions: First, partici-pants were not required to memorize the unambiguousstimulus but to judge a rapid change in its spatialfrequency (i.e.: they were asked whether the distancebetween the grating bands increased or decreased), thusproviding their response after its offset. For this, a 100ms change in the unambiguous stimulus spatial fre-

Figure 4. Results of Experiment 2. Left panel: proportion of ‘‘house’’ responses as a function of stimulus category held in WM. Each

data point represents the average proportion for one subject. Right panel: proportion of times participants reported the house as the

first percept during rivalry as a function of WM category. Error bars represent the SEM and black, dashed lines indicate chance level.

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 8

quency occurred after 400 ms from its onset: Partici-pants pressed the ‘‘a’’ or the ‘‘s’’ key to indicate anincrease or a decrease, respectively. The change in spatialfrequency was determined by a staircase algorithmaiming at 75% of correct responses. Second, no memorytest stimulus was presented, and the rivalry stimulus wasimmediately followed by a 1.5 s blank interval.

Each experimental condition comprised 32 trials,and participants were allowed to take a break everyeight trials. After a brief familiarization with theexperimental procedure, participants underwent bothconditions, in a counterbalanced order, in a uniqueexperimental session that lasted about 75 min.

Results and discussion

Participants scored 69.5% correct at the WM and68.2% at the attention control task, meaning that thetask was challenging and of comparable difficulty in thetwo conditions. The overall proportion of exclusivedominance during rivalry was 82.5% in the WM and84.5% in the attention condition.

We sorted data by orientation of the unambiguousstimulus: Perception during rivalry was defined to beeither congruent or incongruent, depending on whetherthe reported percept matched the orientation of theunambiguous stimulus or not. By analogy with theprevious experiments, we calculated dominance pro-portion for each participant in the two experimentalconditions as the amount of ‘‘congruent’’ and ‘‘incon-

gruent’’ responses (sampled at 100 Hz) over periods ofexclusive dominance, collapsed across trials.

The left panel of Figure 6 depicts the relativedominance of ‘‘congruent’’ responses as a function ofexperimental condition. On average, the dominanceproportion of congruent percepts was 49.85% in theWM and 50.34% in the attention condition. Such adifference was not significant at the two-tailed, paired-sample t test conducted on the data, t(11)¼�0.66, p¼0.52. The right panel of Figure 6 represents theproportion of trials where participants reported acongruent percept as the first dominant percept in thetwo conditions. The first rival percept matched theunambiguous stimulus in 49.2% of the trials in the WMand in 50% in the attention condition, a statisticallynegligible difference, t(11) ¼�0.48, p ¼ 0.5.

These results provide no evidence for the hypothesisthat holding in WM a low-level visual feature couldmodulate rivalry perception, at least in this experi-mental paradigm. Rather, they are in line with theoutcome of Experiment 2 which showed no influence ofWM on binocular rivalry.

Experiment 4

Notwithstanding the careful experimental design andthe measures taken to engage participants in achallenging WM task based on holistic visual encodingof the memory samples, Experiments 2 and 3 did nothighlight any significant effect of visual WM contents

Figure 5. Trial procedure in the WM (A) and attention condition (B) of Experiment 3. In (A) participants were asked to memorize the

grating spatial frequency for a delayed match to sample. During the retention interval, they were presented with the rivalry stimuli

and reported perceptual dominance on a moment-to-moment basis. In (B) participants were required to judge a 0.1 s change in the

unambiguous grating spatial frequency. They provided their responses during the 3 s inter stimulus interval and then monitored

rivalry dominance as in (A) The display duration is indicated below each stimulus window.

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 9

on binocular rivalry. As there is evidence that WMcontents can influence perception of structure-from-motion ambiguous stimuli (Scocchia, Valsecchi, Ge-genfurtner, et al., 2013), it is possible that the outcomeof Experiments 2 and 3 was determined by differentmechanisms underlying the two types of bistablepercepts, with rivalry being a more automatic form ofvisual completion and therefore being less susceptibleto top-down influence. Indeed, it has been shown thatthe instruction to attend to one of the two alternativeinterpretations of the ambiguous stimulus can sub-stantially influence perception of the Necker cube butexerts only a modest effect on face-house rivalry stimuli(Meng & Tong, 2004). On the other hand, endogenousattention strongly affects binocular rivalry whenparticipants are engaged in a tracking task, such asmonitoring a continuous change in one of the rivalstimuli (Chong et al., 2005; Chong & Blake, 2006).

The discrepancy between the results of Meng andTong (2004) and the ones of Chong and Blake (2006)and Chong et al. (2006) might be the result of the taskemployed to elicit endogenous attention (explicit verbalinstructions vs. tracking) but also of the differencebetween the kind of stimuli being used: Whereas Mengand Tong (2004) employed high-level house-facestimuli during rivalry, Chong and Blake (2006) andChong et al. (2005) made use of gratings or radialcheckerboard versus bull’s eye stimuli.

Experiment 4 investigated whether house-face rivalryis susceptible to cognitive influence other than the oneexerted by WM. In particular, we tested the effect ofsustained endogenous attention with the same type ofstimuli whose perception proved not to be affected bythe contents of WM.

Methods

Observers

A new pool of seven paid volunteers participated inthis experiment. All of them had normal or corrected-to-normal visual acuity, and were naıve to the purposeof the experiment. The experimental setup was the sameas described in Experiment 1.

Stimuli and procedure

The rivalry stimuli were the same as in Experiment 2,and all participants underwent the calibration proce-dure prior to testing, as described above. Theexperimental procedure is depicted in Figure 7. NoWM stimulus was displayed: Instead, participants wereasked to monitor either the face or the house rivalimage, in a blocked fashion, in order to detect a rapidchange in its appearance. The change consisted of a 1

Figure 6. Results of Experiment 3. Left panel: proportion of ‘‘congruent’’ responses as a function of experimental condition. The rival

percept is defined to be congruent to the unambiguous stimulus if it shared the same orientation. Each data point represents the

results of one subject. Right panel: proportion of times participants reported a congruent percept as the first percept of a trial. Error

bars represent the SEM and black, dashed lines indicate chance level.

Figure 7. Trial procedure of Experiment 4. Participants were

required to report the dominant percept while tracking one of

the rival images in order to detect a 200 ms change. 0.8 s after

rivalry offset, an instruction window prompted participants to

judge whether a change occurred in the target image. The

display duration is indicated below each stimulus window.

Stimuli are not drawn to scale, and instructions were presented

as actual sentences.

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pixel horizontal stretch displayed for 200 ms, whichcould occur at any time of stimulus presentation apartfrom the first and last second. Participants wererequired to report subjective dominance while moni-toring one of the two images in order to detect thechange. The trial procedure is depicted in Figure 7: Atthe beginning the rivalry stimulus was displayed for 15s, followed by a blank window for 0.8 s and then by aprompt window that signaled participants to reportwhether a change occurred in the face (or in the house)stimulus. Once participants entered their response, ablank window was displayed for 0.7 s and the trialsequence resumed from the beginning. The experi-mental trials were divided into three sessions of 100trials each: There were 40 catch trials per session, wherethe change actually occurred 20 times on the house and20 on the face stimulus. Half of the participants wererequired to monitor the house during the first 50 trialsof each experimental session and the face during thelast 50 trials; the other half did the opposite. Aninstruction window reminded participants which stim-ulus to track at the beginning and at the middle of theexperimental session, after they took a compulsorybreak. Furthermore, participants were allowed to takea break every five trials. Participants were tested in fourseparate sessions (including calibration session) held ondifferent days, and the whole procedure lasted ap-proximately 3.5 hr. As exogenous transients in therivalry display are known to affect subjective domi-nance (Walker & Powell, 1979; Wilson, Blake, & Lee,2001), catch trials were not included in the analyses.

Results and discussion

Overall accuracy in the two tracking tasks was87.71%, indicating that the task was sufficientlydifficult to require sustained attention. On average,observers perceived exclusive dominance for 48.33% ofthe time during rivalry. When they tracked the faceimage, house relative dominance decreased by about

30% compared to when they tracked the house image.This difference proved to be significant at thedependent samples, two-tailed t test conducted ondominance proportion, t(6)¼�4.43, p¼ 0.004. Resultsare illustrated in the left panel of Figure 8. The rightpanel shows the proportion of times participantsreported the house as the first percept as a function ofthe tracked stimulus. Similar to the results ondominance proportion, observers reported ‘‘house’’ asthe first percept of a trial about 30% more often whenthey were tracking the house compared to the faceimage. The difference between tracking conditions wasstatistically significant, t(6)¼�4.61, p ¼ 0.004.1

One might surmise that the increase in relativedominance of the attended image derived from a bias toreport intermingled percepts as exclusive ones whenparts of the attended stimulus were visible. Althoughthe level of mixed dominance in Experiment 4 wasrelatively high, we are quite confident that the observedresult is not contingent on our specific experimentalsetup or on the level of mixed dominance, as it has beenpreviously described in the literature (Chong & Blake,2006; Chong et al., 2005; Ooi & He, 1999). Instead, wedid not find any effect of WM content regardless of thelevel of mixed dominance (which ranged from 68% inExperiment 2 to 17% in Experiment 3).

Our data provide additional evidence for theinfluence of endogenous attention on rivalry perception(Chong et al., 2005) when attention is engaged byrequiring observers to actively perform a task on one ofthe rival stimuli. Furthermore, they extend the validityof those findings to complex, meaningful stimuli whichproved insensitive to attentional influence in previousstudies (Meng & Tong, 2004).

General discussion

In four experiments we assessed the amenability totop-down influence of binocular rivalry using both

Figure 8. Results of Experiment 4. Left panel: proportion of ‘‘house’’ responses as a function of the stimulus tracked during rivalry.

Each data point represents the results of one subject. Right panel: proportion of times participants reported the house as the first

percept as a function of the tracked stimulus. Error bars represent the SEM and black, dashed lines indicate chance level.

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 11

complex (houses and faces) and simple stimuli (sinu-soidal gratings): We showed that rivalry dominance canbe strongly influenced by sustained attention but not byvisual working memory load, at least in this experi-mental paradigm.

In Experiment 1 we tested whether holding in memorythe image of a face, of a house, or of a neutral stimulus(airplane) could affect subsequent rivalry perception: Wefound that when the WM and one of the rival stimulibelonged to the same category, initial dominance of thematching stimulus prevailed. However, category matchdid not affect the global amount of dominance acrosstrials. Instead, an identitymatch between theWMsampleand either of the rival stimuli had no effect beyond the oneof class membership. Experiment 2 suggests that theresults of Experiment 1 could be explained in terms ofcolor priming, as no effect of WM was observed whencolor information was removed from the WM samples.Experiment 3 extended the findings of Experiment 2 to adifferent kind of visual stimulus: The lack of aWM effectwas replicated with grating stimuli, providing additionalevidence against the idea that a direct match in low-levelvisual features between the stimulus held in memory andeither of the monocular ones may influence rivalryperception. Finally, Experiment 4 highlighted the effect ofsustained attention on both rivalry initial dominance andoverall dominance proportion when participants wereinvolved in a tracking task.

Overall, the present results do not support the idea thatWM contents can influence binocular rivalry. Cautionshould be used in drawing conclusions from a null result,which might be limited to the experimental paradigmutilized in this study, however. In a previous study(Scocchia, Valsecchi, Gegenfurtner, et al., 2013), in whichwe employed a very similar experimental paradigm, weobserved antithetical results:WM contents were found toaffect the perception of an ambiguously rotating sphere.In that case, observers were required to memorize anunambiguously moving sphere in one condition and toperform a change discrimination task on the sphere inanother condition. They tended to perceive the ambigu-ous stimulus as moving in the same direction as theunambiguous stimulus both in theWMand in the controlcondition, but consistently more so in the WM one.Furthermore, relative dominance dropped to chancelevels towards the end of the trials in the attentioncondition, whereas the effect in the WM condition wassustained over time. Thus, where a specificWM influenceon bistable perception has been observed, it wascharacterized as a long-lived effect. The fact that inExperiment 1of the present study the WM stimuluscategory had only a short-lived effect on rivalry domi-nance, togetherwith theevidence collected inExperiments2 and 3, favors an interpretation of the result in terms ofpriming rather than of WM effects.

A comparison between the results of this and of ourprevious study (Scocchia, Valsecchi, Gegenfurtner, etal., 2013) highlights two crucial aspects: First, visualWM effects on ambiguous perception depend on thekind of stimulus being used, with binocular rivalrybeing resistant and ambiguous structure-from-motionperception being amenable to WM modulation. Sec-ond, the influence of attention over bistable perceptionis multi-faceted: Even when focusing on selective visualattention, a clear distinction emerges between theeffects of transient and sustained attention. Whenattention is directed to a specific visual feature andreleased before the ambiguous stimulus onset (as in thecase of the control condition in Experiment 3 and inScocchia, Valsecchi, Gegenfurtner, et al., 2013), itseffect is limited even for those stimuli that prove to bemore suitable to cognitive modulation. Instead, atten-tive tracking can influence the perception of stimuli thattypically involve more automatic forms of visualcompetition, as binocular rivalry, and that proveresistant to transient focusing of attention. (See alsoDieter & Tadin, 2011 and Paffen & Alais, 2011 forinsightful reviews on the topic of attentional modula-tion of binocular rivalry).

Previous studies reported diverging evidence on theeffects of endogenous selective attention on binocularrivalry (Chong et al., 2005; Meng & Tong, 2004). On onehand, a possible explanation of this discrepancy couldrely on the different attention tasks participantsperformed in the two experimental paradigms: In Mengand Tong (2004) observers were instructed to attend toone of the two stimuli, whereas in Chong et al. (2005)they were actively engaged in a tracking task. On theother hand, the difference between paradigms that led todissimilar results could be due to the rivalry stimuluscomplexity: Meng and Tong (2004) employed mean-ingful, high-level house-face rivalry, whereas Chong andcolleagues (2005) used gratings or patterned stimuli. Ourexperimental results allow unequivocally discarding theaccount based on stimulus complexity, as we observed aconsistent effect of tracking on house-face rivalry: Thetask employed to engage attentional resources clearlyplays a major role in modulating top-down influenceover binocular rivalry. Interestingly, Helmholtz (1866/1925) himself observed that he could favor dominance ofone of two rival gratings by mentally counting the linesor comparing the spaces within it. As Chong andcolleagues (2005) proposed, performing some kind ofmental operation on one stimulus during rivalry couldbe the critical factor underlying cognitive control over itsprevalence. When participants’ attentional focus is notbound to one of the stimuli via a specific concomitanttask, rivalry dominance seems to be constrained bybottom-up (such as RMS contrast, in our experiments)rather than top-down factors.

Journal of Vision (2014) 14(5):13, 1–15 Scocchia, Valsecchi, Gegenfurtner, & Triesch 12

Importantly, our results show that the effects of WMand of sustained attention on binocular rivalry are notthe same. There is no definitive agreement on thedistinction between WM and sustained selective atten-tion in the literature, and it has been proposed thatvisual WM is one and the same thing as visual attentionsustained internally over time (Chun, 2011). Indeed, therequirement of holding a visual stimulus in WM forsubsequent recall has been commonly shown to act asan endogenous attentional cue: Visuospatial attentionis attracted towards the location occupied by a stimulusthat corresponds to the WM template (Downing, 2000;Soto et al., 2005, Turatto et al., 2008). Our experimentsunderscore that sustained visual attention and visualWM have dissimilar effects on the same bistablestimuli, thus suggesting that they rely on distinctoperating mechanisms. Although this conclusionshould be limited to the experimental paradigmemployed in this study, it is in agreement with researchindicating that a match between a visual search itemand the content of WM do not automatically elicitattention allocation when paying attention to the WMmatching item would impair performance (Downing &Dodds, 2004; Woodman & Luck, 2007). Furthermore,other studies showed that feature-based WM andfeature-based attention have additive effects on theperception of motion direction (Mendoza, Schneider-man, Kaul, & Martinez-Trujillo, 2011), and thatcomparable multiple object tracking and visual WMloads pose different dual-task costs to a concurrentvisual WM task (Fougnie & Marois, 2006).

To conclude, the fact that visual WM contents biasrelative dominance during ambiguous structure-from-motion perception but not during rivalry points todifferent neural mechanisms implied in the two types ofbistable perception, with binocular rivalry involvingcompetition at an earlier stage of visual processing—atleast for the stimuli used in our experiments. Our dataare in keeping with previous evidence (Meng & Tong,2004) showing a substantial difference between theeffects of selective attention on perceptual dominanceduring rivalry and other, higher-level kinds of visualcompetition (i.e., Necker cube). The results areconsistent with the idea that binocular rivalry entails amore automatic form of visual competition thanambiguous structure-from-motion, which would be atthe origin of the dissimilar effects of visual WM on thetwo types of bistable stimuli. Indeed, our studysupports the notion that the perceptual epiphenomenaof binocular rivalry reflect competition at an earlystage, where the signals from the two eyes are not yetcombined (Blake, 1989; Haynes, Deichmann, & Rees;2005; Lee, Blake, & Hegeer, 2005; Lunghi, Burr, &Morrone, 2011; Tong & Engel, 2001; Wunderlich et al.,2005). Such a low-level form of visual competitionseems resistant to an influence of visual WM contents,

but is still amenable to a top-down modulation as longas the observer’s attention is constrained to one of thetwo rival images via a specific concomitant task.

Keywords: binocular rivalry, visual working memory,selective attention, top-down control

Acknowledgments

LS and JT have been supported by the JohannaQuandt Foundation, MV has been supported by theAlexander von Humboldt Foundation.

Commercial relationships: none.Corresponding author: Lisa Scocchia.Email: lisagiorgia@gmail.com.Address: Department of Psychology, University ofMilano-Bicocca, Milan, Italy.

Footnote

1 For all the reported experiments, additionalanalyses on normalized dominance durations wereperformed: A first analysis examined the durations ofall single percepts for each observer; a second analysisfocused on the duration of the first percept of each trial.Durations were normalized based on each observer’saverage duration of all percepts and of all first percepts,respectively. The results of both types of analysesoverlapped the ones obtained by analyzing dominanceproportions. In particular, no effect whatsoever of WMor of its interactions was found in Experiments 1, 2,and 3. The only significant result observed was aninteraction effect in Experiment 4: a 2-way repeatedmeasures ANOVA with Condition (attend house orattend face) and Reported Percept (house or face) asfactors revealed that the interaction between Conditionand Reported Percept was significant for both theanalysis of mean normalized durations (F[1, 6]¼ 29,98,p , .01) and for the analysis of first percepts durations(F[1, 16] ¼ 12,74, p , .05), thus confirming that theattention condition influenced binocular rivalry.

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