Allocation of attention across saccades

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1 lloation of attention aross saades Donatas Jonikaitis, Martin Szinte, Martin Rolfs and Patrik Cavanagh J Neurophysiol 19: , 213. First published 5 Deember 212; doi: /jn You might find this additional info useful... This artile ites 75 artiles, 3 of whih you an aess for free at: Updated information and servies inluding high resolution figures, an be found at: dditional material and information about Journal of Neurophysiology an be found at: This information is urrent as of pril 14, 213. Downloaded from at Harvard Univ on pril 14, 213 Journal of Neurophysiology publishes original artiles on the funtion of the nervous system. It is published 24 times a year (twie monthly) by the merian Physiologial Soiety, 96 Rokville Pike, ethesda MD Copyright 213 the merian Physiologial Soiety. ESSN: Visit our website at

2 J Neurophysiol 19: , 213. First published Deember 5, 212; doi:1.1152/jn lloation of attention aross saades Donatas Jonikaitis, 1,2 Martin Szinte, 1,2 Martin Rolfs, 3,4 and Patrik Cavanagh 2 1 llgemeine und Experimentelle Psyhologie, Ludwig-Maximilians Universität Münhen, Munih, Germany; 2 Laboratoire Psyhologie de la Pereption, Université Paris Desartes, Sorbonne Paris Cité, Paris, Frane and Centre National de la Reherhe Sientifique, Unité Mixte de Reherhe 8158, Paris, Frane; 3 Department of Psyhology, New York University, New York, New York; and 4 Laboratoire de Psyhologie Cognitive, Université ix-marseille and Centre National de la Reherhe Sientifique, Marseille, Frane Submitted 3 July 212; aepted in final form 4 Deember 212 Jonikaitis D, Szinte M, Rolfs M, Cavanagh P. lloation of attention aross saades. J Neurophysiol 19: , 213. First published Deember 5, 212; doi:1.1152/jn Whenever the eyes move, spatial attention must keep trak of the loations of targets as they shift on the retina. This study investigated transsaadi updating of visual attention to ued targets. While observers prepared a saade, we flashed an irrelevant, but salient, olor ue in their visual periphery and measured the alloation of spatial attention before and after the saade using a tilt disrimination task. We found that just before the saade, attention was alloated to the ue s future retinal loation, its preditively remapped loation. ttention was sustained at the ue s loation in the world aross the saade, despite the hange of retinal position whereas it deayed quikly at the retinal loation of the ue, after the eye landed. y extinguishing the olor ue aross the saade, we further demonstrate that the visual system relies only on preditive alloation of spatial attention, as the presene of the ue after the saade did not substantially affet attentional alloation. These behavioral results support and extend physiologial evidene showing preditive ativation of visual neurons when an attended stimulus will fall in their reeptive field after a saade. Our results show that traking of spatial loations aross saades is a plausible onsequene of physiologial remapping. remapping; saades; spatial attention; visual stability THE VERGE INTERSCCDIC interval is short, about one-third of a seond, so in everyday situations, we view, study, reognize, and trak objets aross many fixations. Eah objet is therefore enoded from several loations on the retina and then proessed at various times by several different sets of neurons in retinotopi visual proessing areas (Sereno et al. 1995; Gardner et al. 28). Clearly, every time we make an eye movement, the visual system needs to aount for retinal image shifts to maintain objet ontinuity as well as the stability of our visual world (Wurtz 28). Physiologial researh suggests that saade ontrol areas may ompensate for the retinal image shifts aused by eye movements (Duhamel et al. 1992; Sommer and Wurtz 22; Kusunoki and Goldberg 23; Hall and Colby 211). postsaadi target loation is determined by taking into aount the presaadi target position and the metris of the planned saade. This results in the shift of neural ativity from neurons with reeptive fields enoding the presaadi target position to neurons with reeptive fields enoding its postsaadi position (Fig. 1), an effet alled remapping (Duhamel et al. 1992; Nakamura and Colby 22; Sommer and Wurtz 22; Heiser ddress for reprint requests and other orrespondene: D. Jonikaitis, llgemeine und Experimentelle Psyhologie, Leopoldstr 13, 882, Munih, Germany ( donatas.jonikaitis@psy.lmu.de). and Colby 26). Remapping is preditive and, in some areas, even observed before the saade (Duhamel et al. 1992; Kusunoki and Goldberg 23). Remapping-related neural ativity has also been demonstrated in humans (Medendorp et al. 23; Merriam et al. 23; 27; Parks and Corballis 28; 21; Medendorp 211), and it has been proposed that this ativity ould also be seen in behavioral effets (Melher and Colby 28; Cavanagh et al. 21). In partiular, neurophysiologial studies report that only attended or salient stimuli are remapped (Gottlieb et al. 1998) and that saade ontrol areas involved in remapping are also involved in ontrol of spatial attention (Moore and rmstrong 23; Shall 24; wh et al. 26; isley and Goldberg 21). In support of these proposals, a reent study did find presaadi remapping of spatial attention (Rolfs et al. 211). In partiular, these authors reported presaadi pereptual benefits at the loation that an attended stimulus would oupy only after the saade (Rolfs et al. 211). These results implied that remapping spatial attention or attentional pointers enable the visual system to keep trak of relevant objets aross saades (Cavanagh et al. 21; Rolfs et al. 211). However, there has been no test yet of the assumption that the pereptual proessing benefits seen at the remapped loation before the saade (Rolfs et al. 211) are transferred to the spatial loation of the ue after the saade. number of studies of attentional ueing have reported that, after the saade, spatial attention is alloated to either spatial loation of the attended stimulus (suggesting that attention was remapped) or to the retinal loation that the ue oupied before the saade (suggesting that remapping had failed or was inomplete) or to both (Golomb et al. 28, 21b, 211). However, those studies investigated updating of memorized loations, a proess that likely has a different time ourse than the remapping of stimuli present in the immediate visual environment [see Golomb et al. (28) for disussion]. Moreover, those studies did not measure attention alloation both before and after the saade (Golomb et al. 28, 21b, 211), making the link between attentional remapping before the saade (Rolfs et al. 211) and attentional effets after the saade (Golomb et al. 28) open to question. Thus it is unknown whether preditive remapping an be assoiated with spatiotopi alloation of attention aross saades. dditionally, most remapping studies investigated updating of spatial loations, without respet to objet features or identity (Duhamel et al. 1992; Nakamura and Colby 22; Sommer and Wurtz 22; Medendorp et al. 23; Merriam et al. 23; Parks and Corballis 21; Hall and Colby 211). Remapping provides a /13 Copyright 213 the merian Physiologial Soiety 1425

3 1426 TTENTION CROSS SCCDES Remapped to RF2 2 efore the saade Remapping Flash in RF1 Fixation Flash now in RF2 Planned fter the saade RF1: retinotopi trae Fixation Fig. 1. Preditive remapping. While the observer is preparing a saade from the lighthouse window to the white bird, a flash aptures his or her attention. This flash is registered by a population of neurons with reeptive fields at that loation (RF1: blak dashed irle in ). However, after the saade, the reeptive fields of those neurons are at an irrelevant loation; orresponding to the retinotopi trae of the flash (RF1: blak dashed irle in ). To keep trak of this potentially relevant visual event, the visual system needs to reloalize, or remap, the attention aptured by the flash to the retinotopi loation the flash will have after the saade. This remapping aids visual stability by preativating, before and during the saade, a population of neurons with reeptive fields at the retinal loation (RF2: white dashed irle in ) that will math the loation of the flash after the saade (RF2: white dashed irle in ). predited postsaadi loation for attended objets, but the predition may have some error (Szinte and Cavanagh 211; Szinte et al. 212) and objets may move during a saade. n auxiliary method to establish target loations an all on heking for whether postsaadi objet features math those stored in a transsaadi memory (Deubel et al. 1996, 1998, 21; Crapse and Sommer 212). This proess may take some time after the saade (Zhou and Desimone 211) but would be a viable strategy if spatial updating after the saade takes some time to develop (Golomb et al. 28). However, so far it is not known whether visual remapping is suffiient to eliit spatiotopi attention effets after the saade or whether the visual system atively searhes for partiular ue features to realign attention to the ue following the saade. In the urrent study, we devised a task to investigate these two issues (Fig. 2). While partiipants planned a saade, we indued attentional apture with the onset of an irrelevant olor ue (Müller and Rabbitt 1989; Nakayama and Makeben 1989; Carraso and Yeshurun 29; Shreij et al. 21), as it is known that neural representations of attended objets are remapped aross saades (Gottlieb et al. 1998). We measured the alloation of spatial attention at different loations and at P different times before and after the saade by using a probe disrimination task in whih partiipants had to report a Gabor orientation hange. Indeed, improvements in probe disrimination an be used as a diret measure of attention alloation (Deubel and Shneider 1996; Ling and Carraso 26; Neggers et al. 27; Gersh et al. 29; Carraso 211; Jonikaitis and Deubel 211; Rolfs and Carraso 212). With this novel design, we ould determine whether spatial attention is preditively remapped before the saade and whether, after the saade, it is alloated to the spatial loation of the ue, to the retinotopi loation of the ue, or both. y varying the onset of the tilted Gabor test with respet to the saade, we measured when those attentional benefits appeared or disappeared. In addition to examining the alloation of attention aross saades, we manipulated the status of the attention-apturing olor ue, either keeping it onsreen after the saade or removing it during the saade. This manipulation allowed us to investigate whether the alloation of attention after the saade depends on the ontinuing presene of the olor ue. In partiular, the presene of the olor ue after the saade ould failitate spatiotopi attentional benefits, as the visual system ould use the ue to position attention after the saade. If the presene of the olor ue after the saade failitates attentional alloation, we should observe an inrease of the attentional benefit following the saade as information about the ue s loation builds up. MTERILS ND METHODS Partiipants Fifteen partiipants (age between 21 and 29 yr, 5 females, normal or orreted-to-normal vision) took part in the experiment (11 partiipants ompleted the Transient-ue task, 14 ompleted the Sustainedue task, and ompleted both). ll exept for two of the authors (D. Jonikaitis and M. Szinte) were naive as to the purpose of the study. The experiments were undertaken with the understanding and written onsent of eah subjet and were onduted in aordane with the Delaration of Helsinki. Experiments were arried out in llgemeine und Experimentelle Psyhologie, Ludwig-Maximilians Universität Münhen, Munih, Germany and Laboratoire Psyhologie de la Pereption, Université Paris Desartes, Sorbonne Paris Cité, Paris, Frane. Experiments were designed aording to the ethial requirements speified by orresponding institutions even though no institutional review board (or orresponding institution) ethis approval was needed for experiments that do not involve invasive methods. Setup Partiipants sat in a quiet and dimly illuminated room. We reorded right-eye gaze position with an SR Researh EyeLink 1 desktop mounted eye-traker, alibrated before eah new blok and whenever neessary. Partiipants head movements were minimized using adjustable hin and forehead rests, allowing for an auray of reorded gaze position that is finer than.25 at a sampling rate of 1, Hz. Stimulus presentation and response olletion was ontrolled by an pple omputer and implemented in Matlab (MathWorks, Natik, M) using Psyhophysis and Eyelink toolboxes (Watson and Pelli 1983; rainard 1997; Cornelissen et al. 22). Stimuli were presented at a viewing distane of 6 m, on gamma-linearized sreens, a 21-in. Sony GDM-FR (1, pixels, vertial refresh rate of 85 Hz) in Munih, or a 22-in. Compaq P122 (1, pixels, vertial refresh rate of 12 Hz) in Paris.

4 TTENTION CROSS SCCDES 1427 Remapped ue loation Remapped ue ontrol loation FT Saade ST Cue loation Cue ontrol loation Future retinotopi trae loation Proedure Probe stream Distrator stream Mask stim. Mask stim. Probe Distr. Mask stim. Cue Distr. Distrator stream + sustained-ue Mask stim. Cue Distr. Distrator stream + transient-ue Mask stim. Cue Probe Probe stream + sustained-ue Mask stim. Cue Probe Probe stream + transient-ue 25 ms Time Main task. Figure 2 depits the display onfiguration. During eah trial, partiipants performed two tasks, a saade task and a probe disrimination task. trial started with partiipants fixating a entral fixation target forming a bull s eye (radius.75 ) on a gray bakground (mean luminane: 39 d/m 2 ). We presented two potential saade targets, filled blak irles (radius:.75 ), 8 to the left and to the right of the fixation. fter a fixation period (mean 1s,SD 3 ms, utoff at 3.3 SD), the fixation target disappeared, and the bull s-eye replaed one of the two potential saade targets. If partiipants did not make a orret saade within 7 ms following saade target appearane, we repeated the trial later during the same experimental session. In addition, six disrimination-task-related objets (radius 2 ) formed two rows omposed of three objets eah, 6 above and below the fixation and the two saade target loations. The objets onsisted of a stream of flikering stimuli, omposed of vertial Gabor pathes (frequeny: 2.5 pd; 1% ontrast; random phase; standard deviation of Gaussian window: 1.1 ; mean luminane: 4 d/m 2 ) alternating with noise masks (eah pixel s gray value from Gaussian distribution; M:.5; SD:.5; ut-off at, blak, and 1, white; mean luminane: 4 d/m 2 ), every ms (3 frames at 12-Hz refresh rate or 2 frames at a 85-Hz refresh rate depending on the setup used). etween 1 ms before and 6 ms after saade target onset (time seleted randomly C FT ST Sustained-ue Transient-ue Pre-sa. probe Post-sa. probe Saade -2 Future retinotopi trae ontrol lo Time relative to saade target onset (ms) Fig. 2. Experimental proedure. : Display setup. Partiipants were instruted to shift their gaze to the saade target (ST), whih ould appear to the right or to the left of the fixation target (FT). We presented 6 visual streams omposed of alternating Gabor and noise pathes above and below the FT and ST. Shortly after the onset of the ST, an attentional ue (green) appeared diretly above or below the FT. t different times relative to the saadi eye movement, a probe (tilted Gabor) appeared within 1 of the 6 visual streams and partiipants reported its orientation. Relative to the position of the ue and to the saade diretion, the probe ould appear at the remapped ue loation (blue frame), the ue loation (red frame), the future retinotopi trae loation (blak frame), or at their respetive ontrol loations mirrored aross the horizontal meridian (respetive light olors). Shemati is not to sale and the olored frames were not visible during the experiment. : visual streams. s a funtion of the loations of the probe and of the ue, the visual streams ould be 1 of 4 different streams. The Distrator stream was omposed of vertial Gabors alternating with noise masks. The Probe stream was idential, exept that at a random time one tilted Gabor appeared. The Distrator stream Sustained-ue, Distrator stream Transient-ue, Probe stream Sustained-ue, and Probe stream Transient-ue streams were similar to the ones desribed above exept that 7 ms after the ST onset all Gabors within these streams were green. Note that no Gabors were shown within any streams after the probe presentation, while the green olor stayed on only in the Sustained-ue ondition. C: stimulus timing. The FT disappeared first with the onset of the ST, followed 7 ms later by the ue. The saade started with an average lateny of 21 ms. The probe appeared either before the saade (presaadi probe) or after the saade (post-saadi probe). Horizontal arrows denote the time interval during whih the probe ould appear. from a uniform distribution), a probe appeared randomly at one of the six loations, with equal probability. t that loation and time, the Gabor hanged orientation for one period of ms followed again by a mask. We seleted the probe orientation based on a threshold proedure explained below. One the probe had appeared, no more Gabor pathes followed at any of the loations and the noise masks now alternated with blanks. Seven-hundred milliseonds after saade target onset, all objets disappeared and the partiipant reported the probe orientation (lokwise or ounterlokwise from vertial) followed by a response feedbak (a beep if inorret). We stressed that the main task was to make aurate and fast saades and told partiipants not to worry if they did not see the probe. The probe disrimination task served as our measure of attention alloation. We summoned attention by presenting an attention-apturing ue (Müller and Rabbitt 1989; Nakayama and Makeben 1989; Carraso and Yeshurun 29; Shreij et al. 21), an abrupt olor onset stimulus presented above or below entral fixation. Speifially, 7 ms after saade target appearane, the Gaussian envelope overing the ued Gabor path hanged olor to green (mean luminane of Gabor green Gaussian envelope 34 d/m 2 ). Partiipants were asked to ignore this olor onset, as the onset loation did not predit the loation of the probe. In the Transient-ue task, we removed the olor ue during the saade (i.e., the Gabor path returned to gray); in the Sustained-ue task, the olor ue remained onsreen after the saade until the end of the trial.

5 1428 TTENTION CROSS SCCDES Saade lateny (ms) Saade lateny Vertial landing site (deg) Saade auray Partiipants ran a minimum of five 1-h sessions for eah of the tasks (if observers performed both tasks, the Sustained-ue ondition preeded the Transient-ue ondition), eah session onsisting of at least 48 trials. efore starting the experiment, eah partiipant ompleted a training session (usually taking 15 min). Threshold proedure. SELINE. efore eah session, we evaluated the probe tilt angles that gave a riterion 82% orret performane for eah probe loations when ued with a 1% valid olor ue and these baseline probe angles were determined for the various eentriities (pre- and postsaade) and presentation times needed to over the possible probe onditions during the main experiment. The purpose of this baseline was to establish the performane level that is ahieved with attention alloated to the probe loation as muh as possible (1% valid ue) so that in the main experiment performane that reahed this level for the baseline probe orientation indiates a strong engagement of attention. We used interleaved QUEST stairases (Watson and Pelli 1983), varying the probe orientation at different loations until partiipants reahed a desired 82% orret disrimination performane. Just like in the main experiment, we asked partiipants to make a saade, and 7 ms after the saade target onset, a ue (abrupt olor hange) appeared. The ue loation ould be any of the six objet loations and probes appeared always at the ued loation. In the threshold proedure, therefore, the ue was 1% valid, instruting partiipants where to shift attention. Three stairases were evaluated for probes presented 1 ms after the ue onset, orresponding to the presaadi period ( ms before the saade started). first stairase was for probes above and below the fixation target (eentriity 6, average tilt angle aross partiipants: 17 ); a seond for the probes above and below the saade target (eentriity 1 and tilt 2 ); and a third for the probes presented opposite of the saade target (eentriity 1 and tilt 2 ). We also measured three other stairases for probes presented 4 ms after ue onset, orresponding to the postsaadi period. Postsaadi probes had different eentriities and thus different orientation thresholds (eentriities 6, 1, and 17.1 ; tilt 14, 2, and 24 ). Only trials in whih a orret saade ourred were used for the threshold proedure. MIN TSK. During the main task, we used the probe orientations based on the 82% thresholds measured in this threshold mapping task. To do so, we traked online eye position, and depending on both probe eentriity and whether the saade started or not, we presented the orresponding baseline probe orientation. This threshold proedure equated baseline probe disrimination performane even if eentriity of probes hanged aross saades, allowing us to ompare probe disrimination aross eentriities as well as before and after the saade. This proedure ensures that 82% orret disrimination in the main task means strong attentional modulation, values below that orrespond to weaker attentional effets and % orresponds to hane performane. Data nalysis We deteted saades offline using an algorithm based on twodimensional eye veloity (Engbert and Mergenthaler 26), omputed from subsequent samples in the eye position series. The thresholds for peak veloity and minimum duration used for saade detetion were 3. SD and 2 ms, respetively. To reate the saade landing error map (Fig. 3), we used a kernel density estimation based on linear diffusion proesses (otev et al. 21). We disarded trials where the saade lateny was 1 ms or ms. We only analyzed trials in whih the saade landed within a 2 radius around its goal. In total we aepted 92% trials:.5% of all trials were rejeted due to blinks, 7% due to inaurate saades, and.1% due to partiipants looking at the olor onset loation. The performane in the probe disrimination task is expressed as the perentage of orret orientation disriminations. s the probe appeared at a random time, we binned probe presentation times into 1-ms time bins before and after the saade for further analysis. In presaadi analyses, eah bin ontained all probes whose presentation ended in a given 1-ms interval; in postsaadi analyses, eah bin ontained all probes whose presentation started in a given 1-ms interval. This analysis thus exluded all probes overlapping with the saade in time. On average for eah partiipant, a bin ontained 6 trials (for distributions of trials for different onditions, see Figs. 5 and 6). Sine there were two response alternatives, the hane level of probe disrimination was at %. For the analysis of probe disrimination performane, we pooled data aross saade diretions. Statistial analyses inluded repeated-measure NOV, and diret omparisons between different time bins were done with paired t-tests. Transient-ue and Sustained-ue tasks were ompared with eah other using a mixed effets NOV, whih allows for omparisons of onditions ontaining partly overlapping partiipant pools. RESULTS The average saade lateny was ms (means SE) in the Sustained-ue task and ms in the Transient-ue task. These latenies math those observed in other studies inves C Saade amplitude (deg) Saade amplitude Probe onset relative saade target onset (ms) Horizontal landing site (deg) Probe onset relative saade target onset (ms) Fig. 3. : Saade lateny as a funtion of probe presentation time and probe loations. Colors indiate the probe loations with respet to saade target and onset ue, as desribed in the small legend ion. Error bars indiate SE. : saade auray for all saade diretions and probe loations. Redder olors represent higher inidene of saades to that loation. C: saade amplitude as a funtion of probe presentation time and probe loation. Same onventions as in. C show data from the Sustained-ue ondition. Data from the Transient-ue (not displayed) ondition were not statistially different.

6 TTENTION CROSS SCCDES 1429 tigating attentional alloation and saade planning (Deubel and Shneider 1996; Golomb et al. 28; Jonikaitis and Deubel 211; Rolfs et al. 211; Rolfs and Carraso 212), suggesting that the appearane of the olor ue did not have a major impat on saade latenies. dditionally, as our display onsisted of several flikering stimulus streams starting well before the appearane of the saade target, the probe onset itself did not stand out from these bakground events and so did not disrupt saade planning. Figure 3 shows that average saade latenies for probes presented at different loations and at different times after the saade target onset are largely similar. Repeated-measures NOVs did not show an effet of either probe position with respet to the saade target or probe presentation time and this for both tasks (all P.5). Figure 3 shows saade auray. verage saadi errors (as measured in distane from the saade target enter at 8 eentriity) aross partiipants was.23.3 in the Sustained-ue task and.25.5 in the Transient-ue task. Finally, as observed in Fig. 3C, saade landing position did not vary as a funtion of the probe presentation time, or probe position for either of the tasks (all P.5). Next, we analyzed performane in the probe disrimination task. To do so, we omputed probe disrimination performane as the perentage of orret disrimination responses for probes appearing within speified 1-ms time bins loked either to the olor ue onset (Fig. 4) or to the saade onset (Figs. 5 and 6). For eah trial, we defined three positions of interest (ue loation, remapping, and future retinotopi trae) as well their three respetive ontrols, mirrored relative to the saade vetor. We then evaluated the temporal dynamis of attention alloation at these loations, by omparing the atual position with their ontrol for the different time bins. Figure 4 shows probe disrimination performane at the ued loation and at its ontrol loation for two experimental onditions (for this omparison, we looked at the disrimination performane Performane (% orret) Sustained-ue Transient-ue hane level relative to saade ue onset (ms) Fig. 4. Probe disrimination performane at the ue loation before and after the olor ue onset for Sustained-ue and Transient-ue onditions. Colors indiate the probe loations as desribed in the small legend ions for eah panel. We omputed performane in temporal bins separated by 1 ms. For a given temporal bin, filled symbols indiate signifiant differenes between a loation of interest and its ontrol, based on repeated-measures t-tests taken separately for the sustained and the transient-ue ondition. observed for the 1 partiipants who did both onditions). For both Sustained-ue and Transient-ue onditions, we observed the typial effet of transient spatial attention, that is disrimination performane improved at the ue loation, reahing a maximum 1 ms after the ue onset, and then dereased (Müller and Rabbitt 1989; Nakayama and Makeben 1989). Next, we analyzed disrimination performane over time before and after the saade, first for the Sustained-ue (see Fig. 5) and then for the Transient-ue onditions (see Fig. 6). Figure 5, top, shows probe disrimination performane, whereas Fig. 5, bottom, shows the total number of trials used in eah time bin by dividing them with respet to whether the olor ue was already shown (postue trials, plotted upwards) or not (preue trials, plotted downwards). The earliest presaadi time bin (3 2 ms before the saade) ontains mostly preue trials; the latest time bin (1 ms before the saade) ontains mostly postue trials; the intermediate time bin (2 1 ms before the saade) ontains a mix of both. Figure 5 shows that in the Sustained-ue ondition, before the saade began, probe disrimination improved markedly at the ue loation with respet to its ontrol loation. repeatedmeasures NOV (with probe time and its loation as main fators), showed that probe disrimination performane before saade onset was affeted by time [F(2,26) 15.4, P.1], probe loation [F(1,13) 34.36, P.1], and the interation between the two fators [F(2,26) 12.75, P.1]. Probe disrimination performane inreased strongly at the ue loation ompared with ontrol loation for probes presented up to 2 ms before the saade [2 1 ms before saade, t(13) 2.62, P.5; 1 ms before saade, t(13) 9.78, P.1, filled squares in Fig. 5 mark statistially signifiant omparisons]. Figure 5 also shows the disrimination performane for probes presented after the saade has landed. Probe disrimination performane remained higher at the ued loation than at the ue ontrol loation after the saade. Repeated-measures NOV showed the main effet of the probe loation [F(1,13) 11.97, P.1], and an effet of time was approahing signifiane [F(3,39) 2.68, P.6]; the interation was not signifiant [F(3,39) 1.33, P.27]. Paired t-tests showed that probe disrimination at the ue loation was signifiantly better than that at the ontrol loation 2 ms after the saade [ 1 ms after saade, t(13) 2.72, P.5; 1 2 ms after saade, t(13) 2.77, P.1]. Combined, these results show that the ue improved disrimination performane at its loation and that this benefit is sustained, as a spatiotopi attentional benefit, aross the saade. Next, we analyzed probe disrimination performane at other loations in the visual field. The first set of loations we analyzed was the loations above and below the saade target (Fig. 2). efore the saade starts, these two loations are not diretly relevant for the saade task nor are they related to the ue loation, even though it has been suggested that one of those loations is the loation to whih preditive remapping is direted (Mathôt and Theeuwes 21), a view that is no longer maintained (remapping is direted in the opposite diretion see Fig. 1; Krauzlis and Nummela 211; Mathôt and Theeuwes 211; Rolfs et al. 211). fter the saade ends, this loation on the display now orresponds to the retinotopi trae loation for attentional benefits, i.e., the retinal loation the ue had previously oupied (Golomb et al. 28, 21a, 21b, 211). retinotopi trae

7 143 TTENTION CROSS SCCDES Remapping Cue C Future retinotopi trae Performane (% orret) hane level # trials Post-ue Pre-ue exists only after the saade; thus in the presaadi period we refer to that loation as the future retinotopi trae loation. Probe disrimination performane inreased at both the future retinotopi trae loation and at its ontrol loation [Fig. 5C; effet of time was signifiant F(2,26) 5.54, P.1], but there was no signifiant differene between the two loations [F(1,13).93, P.35] nor an interation between the two fators [F(2,26).44, P.65]. Paired t-tests showed no signifiant differenes between the two loations at any time point before the saade (all P.5). Thus probe disrimination inreased at both loations with a similar time ourse and magnitude, probably due to their proximity to the saade target (Gersh et al. 29). This disrimination performane before the saade therefore fails to show the advantage reported by Mathôt and Theeuwes (Mathôt and Theeuwes 21). possible explanation for this disrepany is that Mathôt and Theeuwes s stimulus reated a strong pereption of apparent motion between the presaadi ue and the attentional probe. reent repliation of their experiment revealed general reation time benefits along the path onneting the ue and target (as ompared with eentriity-mathed ontrol loations) strengthening this onjeture (Harrison et al. 212). fter the saade, performane at the retinotopi trae loation and its ontrol (Golomb et al. 28) was affeted by probe presentation time [F(3,39) 3.53, P.5] but not by probe position [F(1,13).9, P.35] and probe position did not interat with probe timing [F(3,39) 1.28, P.29]. Nevertheless, planned t-tests showed that probe disrimination at the Post-ue Pre-ue Post-ue Pre-ue Fig. 5. Probe disrimination performane before and after saades for the Sustained-ue ondition is shown at top: Performane for probes presented at the remapped loation and its ontrol (), at the ue loation and its ontrol (), and at the future retinotopi trae loation and its ontrol (C). Colors indiate the probe loations as desribed in the small legend ions for C. We omputed performane in temporal bins separated by 1 ms both for probes appearing before the saade started and for probes appearing after the saade finished. For a given temporal bin, filled symbols indiate signifiant differenes between a loation of interest and its ontrol, based on repeated-measures t-tests. Numbers of trials when probe was shown either before the olor ue (preue trials, plotted downwards) or after the olor ue (postue trials, plotted upwards) are shown at bottom. Same olor onventions as in the upper panels. The number of preue and postue trials for eah bin sums up to the total number of trials available for that time bin. Error bars are SE. retinotopi trae loation was better than at its ontrol loation [t(13) 2.25, P.5] over the interval 1 ms after the saade but not beyond. In other words, there was a short-lived performane advantage at the retinotopi trae loation after the saade in support of earlier reports by Golomb and olleagues (Golomb et al. 28, 21a, 21b, 211). Finally, we analyzed the two loations on the other side of the saade goal. These two loations were related neither to the saade target nor to the ue and were in the opposite visual hemifield from the saade target. However, before the saade starts, the loation at the same vertial position as the ue is the loation on the retina that the ue will oupy after the saade (see Fig. 2) and is therefore the remapped loation of the ue (Duhamel et al. 1992; Kusunoki and Goldberg 23; Hall and Colby 211; Krauzlis and Nummela 211; Rolfs et al. 211). Thus our presaadi analysis was entered on finding whether the spatial attention aptured by the olor ue is preditively remapped to this loation before the saade (Fig. 5). This would put in plae the attention that would subsequently align with the ued loation after the saade supporting the postsaadi pereptual benefits that we found there (postsaadi ue loation). repeated-measures NOV showed a signifiant effet of probe position [F(1,13) 2.17, P.1]; the probe presentation time effet was marginally signifiant [F(3,39) 3.18, P.6]; and the interation between the two fators was not signifiant [F(3,39).68, P.51]. Paired t-tests revealed that probe disrimination was

8 TTENTION CROSS SCCDES 1431 Remapping Cue C Future retinotopi trae Performane (% orret) hane level # trials Post-ue Pre-ue better at the remapped loation than at the ontrol loation in time intervals 2 1 ms before saade onset [t(13) 3.12, P.1] and 1 to ms before saade onset [t(13) 5.45, P.1]. fter the saade was finished, these two loations on the sreen, furthest from the saade target have no relevane for the effets of either the abrupt onset or the saade planning. The data show that the postsaadi probe disrimination was still affeted by probe position [F(1,13) 9.78, P.1] but not by probe presentation time [F(3,39) 1.17, P.33] and that there was no interation [F(3,39).82, P.48]. The main effet of probe position was unexpeted and turned out not to hold up in the Transient-ue ondition (see below). Disrimination performane in the Transient-ue ondition (Fig. 6), where the ue was presented only before the saade, showed largely similar effets. Disrimination at the ue and ue ontrol loations before the saade depended on probe presentation time [F(2,2) 12.37, P.1] and probe loation [F(1,1) 15.51, P.1], but there was no interation between these two fators [F(2,2) 2.36, P.12]. We obtained the same results after the saade [probe presentation time, F(3,3) 12.1, P.1; probe position, F(1,1) 5.18, P.5; and interation, F(2,2).6, P.97]. Thus probe disrimination was better at the spatial ue position both, before and after the saade. In partiular, probe disrimination at the ue loation was better than at the ontrol loation for the first 1 ms after the saade [t(1) 2.6, P.5]. gain, we did not observe any benefits at the future retinotopi trae loation before the saade [probe position, F(1,1).28, P.6; probe time, F(2,2) 3.22, P.6; and interation, F(2,2).14, P.86]. s in the Sustained-ue experiment, during the first 1 ms following the Post-ue Pre-ue Post-ue Pre-ue Fig. 6. Probe disrimination performane before and after saades for the Transient-ue ondition. ll onventions are the same as in Fig. 5. saade, disrimination performane was better at the retinotopi trae loation than at the ontrol loation [t(1) 3.36, P.1; probe position, F(1,1).3, P.59; probe time, F(3,3) 8.68, P.1; and interation, F(3,3) 4.37, P.5]. Thus, in the Transient-ue ondition, we repliated the attentional retinotopi trae benefits observed in the Sustained-ue ondition. Finally, we observed a benefit at the remapped loation before the saade [probe position, F(1,1) 21.32, P.1; time, F(2,2) 1.5, P.36; and interation, F(2,2) 1.81, P.18] but not after (all P values.5). To evaluate whether the presene of the olor ue after the saade affeted the alloation of spatial attention, we ompared Sustained-ue and Transient-ue tasks diretly, using mixed effets NOV with three fators: probe presentation time, position, and task. If ue presene after the saade had an effet on the disrimination benefit at the ue loation ompared with the ontrol loation, then one would observe an interation between the three fators. We found no interation neither between the probe presentation time and position [F(3,69).73, P.43] nor an interation among task, position, and time [F(3,69).78, P.36]. Thus the presene or absene of an attention apturing ue after the saade did not affet the attentional benefits at the ue loation. This means that feature-based information did not affet disrimination benefits observed at the ue loation after the saade. DISCUSSION We investigated the remapping of spatial attention aross saadi eye movements and report the following findings. First, performane at the ue loation inreased substantially, relative to its ontrol, demonstrating the lassi attentional benefit of a task-

9 1432 TTENTION CROSS SCCDES irrelevant onset (see Carraso 211, for a review; Theeuwes 1994; Yantis and Hillstrom 1994). Importantly, before the saade, the disrimination performane also inreased at the remapped loation of the onset ue, whih demonstrates with behavioral measures the remapping of attention aptured by a salient stimulus. This remapping of attention ourred regardless of whether the ue disappeared or stayed onsreen aross the saade. fter the saade, we observed a short-lived improvement in performane at the retinotopi loation that the ue oupied before the saade. This retinotopi trae of attention dissipated within the first 1 ms after the saade. Moreover, an attentional benefit at the ue loation on the sreen persisted after the saade, meaning that transient attention, invoked at the remapped loation by an abrupt onset stimulus before saade, is orretly alloated to its intended spatiotopi loation immediately after the saade. This effet ours beause the retinal image shift during the saade brings the ue s loation into alignment with the remapped loation, thus reating a spatiotopi attentional benefit, and this effet was seen whether or not the olor ue was present after the saade. Our findings demonstrate plausible behavioral onsequenes of the remapping of neural ativity reported in neurophysiologial studies (see Fig. 7). Neural responses to stimuli appearing outside of neurons visual reeptive fields, but at loations that those reeptive fields will oupy after the saade, have been observed in several areas involved in saade planning - the frontal eye fields (Umeno and Goldberg 1997; Sommer and Wurtz 26), the lateral intraparietal ortex (Duhamel et al. 1992; Kusunoki and Goldberg 23; Heiser and Colby 26), and the superior olliulus (Walker et al. 1995; Churan et al. 211). Suh remapping of visual ativity has also been observed in a number of human funtional MRI (Medendorp et al. 23; Merriam et al. 23) and EEG studies (Parks and Corballis 28; 21; Peterburs et al. 211). While funtional MRI studies, due to the sluggishness of signal, demonstrate remapping of memorized stimuli after the saade (Medendorp et al. 23; Merriam et al. 23, 27), EEG and single ell reording studies demonstrate preditive remapping of attended stimuli before the saades (Duhamel et al. 1992; Kusunoki and Goldberg 23; Parks and Corballis 28, 21). The role of attention in remapping has been frequently disounted (Duhamel et al. 1992; Sommer and Wurtz 26; Hall and Colby 211; Melher 211). However, our result is in agreement with the established physiology of attention and eye movements. Frontal and parietal areas as well as the superior olliulus, all of whih show preditive remapping ativity (Duhamel et al. 1992; Walker et al. 1995; Kusunoki and Goldberg 23; Sommer and Wurtz 26), are also involved in attentional shifts (Shall 22; isley and Goldberg 23, 21; Liu et al. 21; Lovejoy and Krauzlis 21). Consequently, we argue that it is the loations of spatial attention, attentional pointers, that are remapped aross saade (Cavanagh et al. 21). Indeed, typially, remapping ours only for attended stimuli (Gottlieb et al. 1998) and both behavioral and neurophysiologial studies demonstrate that saade targets, whih are strongly attended, are remapped regardless of whether partiipants plan a single saade (Collins et al. 29; Rolfs et al. 211) or sequenes of saades (Sommer and Wurtz 22; Ostendorf et al. 21; Rolfs et al. 211). Finally, several studies have shown that the loation of hand movement targets is also remapped aross eye movements (Medendorp and Crawford 22; van Pelt and Medendorp 28), an expeted finding given that planning hand movements to an objet leads to the automati C Shift of neuron s reeptive field Remapping of attention pointer Neural reording Fixation Saade Eletrophysiology Visual sene Disrimination performane ehavior Fig. 7. Remapping in neurophysiology and behavior. While the observer is preparing a saade from the lighthouse window to the white bird, a flash aptures his or her attention (visual sene, ). oth eletrophysiologial () and behavioral (C) studies investigated how the visual system traks an attention-apturing stimulus, here the flash, despite intervening eye movements. : in eletrophysiologial studies of preditive remapping, an attentionapturing ue is presented at a loation (marked by an asterisk) that falls outside the neuron s reeptive field (solid irle). Just before the saade onset, the neuron beomes responsive to the attention-apturing ue, if the reeptive field of that neuron will fall on the ued loation after the saade. This finding has been interpreted as a reeptive field shift in the diretion of the saade, here marked by the arrow, to what has been termed the ell s future reeptive field (dotted irle). C: in the urrent study we presented an attention-apturing ue (a green Gaussian blob) and measured disrimination performane at the loation indiated by the tilted Gabor path. We interpret the inreased pereptual disrimination performane as preditive remapping of visual attention in the diretion opposite the saade, marked by the arrow. oth neurophysiologial and behavioral studies measured loations or reeptive fields that maintain the same relationship: the attention-grabbing stimulus is presented at one loation, and the response is measured at the loation or reeptive field where the ued loation will land after the saade. These are two equivalent desriptions of the same proess that we argue is best desribed as a transfer of ativation from the initial loation of the ue to its future retinal position (Cavanagh et al. 21). alloation of spatial attention to reah targets (aldauf and Deubel 28, 21; Jonikaitis et al. 21; Jonikaitis and Deubel 211). Our study demonstrates both the remapping of spatially ued attention before the saade and alloation of attention at the ue s loation in the world after the saade. This lends support to the hypothesis that the remapping of visual attention ontributes to spatiotopi attention alloation aross saades: even though the objet is present in the reeptive fields of different visual neurons before and after saade, the transfer of attention from the ued loation to the remapped loation before the saade will bring that attention bak to the ue s spatial loation one the saade lands (Cavanagh et al. 21). Importantly, we observed disrimination benefits for the first 1 ms after the saade, regardless of whether the olor ue was still present in the visual field or not. If pereptual benefits during the first 1 ms after the saade had ourred ontingent on the presene of the olor ue aross the two fixations, then spatiotopi ueing effets should have been observed only when the ue was visible after the saade and not when it was

10 TTENTION CROSS SCCDES 1433 erased. Instead, we observed that removing the olor ue during the saade had little or no effet on attentional benefits at the ue loation after the saade, suggesting that spatiotopi benefits aross saades mainly depend on spatial attention that was appropriately remapped before the saade landing. If the ued loation had to be redisovered following the saade, we would expet some delay in the appearane of the attentional benefits. For example, reent single ell reording studies show that after a saade visual seletivity to attended features suh as olor or shape whih were present before the saade takes 1 ms to build up in attention-modulated visual areas V4 and frontal eye fields (ihot et al. 25; Zhou and Desimone 211). Moreover, reent evidene suggests that frontal eye field neurons detet a stimulus that hanges aross a saade (inluding hanges in loation, olor, or size) but this seletivity also takes some time to develop (Crapse and Sommer 212). Thus, if the visual system had to detet feature information following the saade to loalize spatial attention, spatiotopi benefits would take more time to emerge than is observed in single ell reording studies of remapping (Duhamel et al. 1992; Kusunoki and Goldberg 23). Even though we do not disount that feature-based visual proessing ould potentially aid in loalizing objets in some situations after the saade, our data demonstrate that preditive remapping of attended targets and spatiotopi attentional alloation after the saades is the default mode of funtion in the visual system, even when there are no task demands to update information aross saades and even when the unique feature of an objet is extinguished during the saade. Our findings address urrent ontroversies onerning the distribution of spatial attention after saades. Golomb and olleagues have reported that after a saade there is a strong attentional benefit at the retinotopi loation that had been oupied by a memorized stimulus before the saade (retinotopi trae loation) and that in some ases it took up to a 1 ms for spatiotopi attention effets to our (Golomb et al. 28, 21a; 21b). Our results onfirm the presene of the retinotopi trae after the saade; however, we also observed lear spatiotopi effets immediately after the saade. Contrary to our task, the studies of Golomb and olleagues used loation memory to investigate spatiotopi and retinotopi proessing benefits after the saade. The time ourse of spatial updating aross saades for memorized loations might be different, as there is no urgeny for the visual system to update information about the stimulus that has already disappeared from the visual field. dditionally, those studies did not measure attention alloation before the saade, whih leaves open the possibility that there is no presaadi remapping of attention in a loation memory task. This is in ontrast to the presaadi remapping of spatial attention in response to an attention-grabbing, urrently visible stimulus. Future researh would need to ompare both situations diretly. Our design is similar to neurophysiologial studies that demonstrate that abrupt onset stimuli are remapped (Duhamel et al. 1992; Sommer and Wurtz 22; Kusunoki and Goldberg 23; Merriam et al. 23). The findings from neurophysiologial studies and our urrent results indiate that salient, attention-apturing stimuli are automatially remapped aross saades, even in ases when there is no task-related benefit to do so. However, less is known about whether remapping also ours for voluntary attentional shifts. Studies that investigated spatial attention alloation before saades found that voluntary attention shift to loations that are not saade targets is impaired (Kowler et al. 1995; Deubel and Shneider 1996; Deubel 28). The ompetition between voluntary and saade-triggered attention shifts ould influene the nature and timing of the remapped of voluntary attention aross saades. To our knowledge, this question has not been investigated. Our results, ombined with those of previous studies, portray a dynami piture of attention alloation before and after saadi eye movements. ttention drawn to salient objets before a saade is remapped around the time the eyes move in the opposite diretion of the saade. s a onsequene, attention is ontinuously alloated to the spatial loations of attended objets in the world aross saades, orreting for the large position shifts that eye movements ause for these objets on the retina and throughout retinotopi orties. Moreover, the retinal positions of attended presaadi stimuli shows brief attentional benefits after a saade (Golomb et al. 28), suggesting that these benefits annot be immediately extinguished. Combined, suh spatial updating of attention may help quikly follow attended targets, despite perpetual eye movements. GRNTS This researh was supported by Chaire d Exellene Grant (to P. Cavanagh), Frenh Ministère de l Enseignement Supérieur et de la Reherhe Grant (to M. Szinte), and Deutshe Forshungsgemeinshaft Grant JO 98/1-1 (to D. Jonikaitis). M. Rolfs is supported by the 7th Framework Program of the European Union (Marie Curie International Outgoing Fellowship, projet number ). DISCLOSURES No onflits of interest, finanial or otherwise, are delared by the author(s). UTHOR CONTRIUTIONS uthor ontributions: D.J., M.S., M.R., and P.C. oneption and design of researh; D.J. and M.S. performed experiments; D.J. analyzed data; D.J., M.S., M.R., and P.C. interpreted results of experiments; D.J. and M.S. prepared figures; D.J. and P.C. drafted manusript; D.J., M.S., M.R., and P.C. edited and revised manusript; D.J., M.S., M.R., and P.C. approved final version of manusript. REFERENCES wh E, rmstrong KM, Moore T. Visual and oulomotor seletion: links, auses and impliations for spatial attention. Trends Cogn Si 1: , 26. aldauf D, Deubel H. Visual attention during the preparation of bimanual movements. Vision Res 48: , 28. aldauf D, Deubel H. ttentional landsapes in reahing and grasping. Vision Res : , 21. ihot NP, Rossi F, Desimone R. Parallel and serial neural mehanisms for visual searh in maaque area V4. Siene 38: , 25. isley JW, Goldberg ME. Neuronal ativity in the lateral intraparietal area and spatial attention. Siene 299: 81 86, 23. isley JW, Goldberg ME. ttention, intention, and priority in the parietal lobe. nnu Rev Neurosi 33: 1 21, 21. otev ZI, Grotowski JF, Kroese DP. Kernel density estimation via diffusion. nnu Stat 38: , 21. rainard DH. The Psyhophysis Toolbox. Spat Vis 1: , Carraso M, Yeshurun Y. Covert attention effets on spatial resolution. Prog rain Res 176: 65 86, 29. Carraso M. Visual attention: the past 25 years. Vision Res 51: , 211. Cavanagh P, Hunt R, fraz, Rolfs M. Visual stability based on remapping of attention pointers. Trends Cogn Si 14: , 21. Churan J, Guitton D, Pak CC. Context dependene of reeptive field remapping in superior olliulus. J Neurophysiol 16: , 211. Collins T, Rolfs M, Deubel H, Cavanagh P. Post-saadi loation judgments reveal remapping of saade targets to non-foveal loations. J Vis 9: , 29.

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