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7/25/2019 Klein 1999 Inhibition
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346 Copyright © 1999 American Psychological Society VOL. 10, NO. 4, JULY 1999
PSYCHOLOGICAL SCIENCE
Research Article
INHIBITION OF RETURN IS A FORAGING FACILITATOR
IN VISUAL SEARCH
Raymond M. Klein and W. Joseph MacInnes Department of Psychology and Faculty of Computer Science, Dalhousie University, Halifax, Canada
Abstract—Using overt orienting, participants searched a complex
visual scene for a camouflaged target (Waldo from the “Where’s
Waldo?™ ” books). After several saccades, we presented an uncamou-
flaged probe (black disk) while removing or maintaining the scene,
and participants were required to locate this probe by foveating it.
Inhibition of return was observed as a relative increase in the time
required to locate these probes when they were in the general region of
a previous fixation, but only when the search array remained present.
Perhaps also reflecting inhibition of return, preprobe saccades showed
a strong directional bias away from a previously fixated region.
Together with recent studies that replicate the finding of inhibition at
distractor locations following serial but not parallel visual search—so
long as the search array remains visible—these data strongly support the proposal that inhibition of return functions to facilitate visual
search by inhibiting orienting to previously examined locations.
Immediately following a peripheral stimulus (cue), there is often a
short-lived increase in processing efficiency for nearby stimuli. This
phasic improvement has been attributed to automatic (exogenous) ori-
enting of attention toward the cue (see Klein, Kingstone, & Pontefract,
1992; Posner, 1980). In the absence of motivation to maintain atten-
tion at the cued location, this early facilitation is followed (or accom-
panied) by a longer lasting inhibition (Posner & Cohen, 1984) that has
been called inhibition of return (IOR; see Taylor & Klein, 1998b, for
a review).1 Extending Posner and Cohen’s (1984) interpretation of
IOR as an inhibitory mechanism that would encourage the sampling of new information in the visual field, Klein (1988) proposed that IOR
might facilitate visual search when each display item requires an atten-
tion-demanding inspection to determine if it is the target (cf. Treisman
& Gelade, 1980). Inhibitory tagging of display items that have already
been examined attentively would, by repelling attention, help the
observer avoid reinspecting them.
Klein (1988) tested this functional explanation of IOR by present-
ing luminance-detection probes immediately after the subject had per-
formed an easy (preattentive; target “pops out”) or difficult (requiring
serial allocation of attention to array items) visual search (see Fig. 1).
The probes occurred on half of the trials and were presented at loca-
tions where there had been an item in the search display (on probes)
or at locations where no item had been presented (off probes). The
rationale was straightforward: “In serial search if the presumed allo-cation of attention to each item is followed by inhibition of return, then
detection of on-probes should be delayed compared with off-probes”
(Klein, 1988, p. 430). This is precisely what Klein (1988) found using
parallel search to control for other factors, such as masking and
expectancies (see Table 1). This previous study thus provides support
for an intuitively appealing view of IOR as a foraging facilitator.
IOR AS A FORAGING FACILITATOR:
CHALLENGES AND REBUTTALS
Although several challenges to the functional interpretation of IOR
as a foraging facilitator materialized in the years following Klein’s
(1988) article, more recent findings rebut these challenges.
First, evidence began to accumulate that it might not be attention
that was inhibited but rather processes more closely related to
responding. Initial attempts to obtain IOR using nonspatial discrimi-
nation tasks were unsuccessful (Kwak & Egeth, 1992; Pontefract &
Klein, 1988, described in Klein & Taylor, 1994; Tanaka & Shimojo,
1996; Terry, Valdes, & Neill, 1994). Because performance on such
tasks is sensitive to the locus of attention, the fact that IOR did not
influence such tasks challenged the assumption that attention is inhib-
ited in IOR. Indeed, on the basis of this finding and the fact that IOR
does not affect the perceived temporal order of stimuli (Maylor, 1985;
see Klein, Schmidt, & Müller, 1998, for a review), Klein and Taylor
(1994) proposed (see also Schmidt, 1996a, 1996b, and Tanaka & Shi-
mojo, 1996, for similar proposals) that it was not attention that was
inhibited in IOR but rather responding toward stimuli appearing at
locations for which there had previously been an oculomotor program
(cf. Rafal, Calabresi, Brennan, & Sciolto, 1989). If correct, this pro-
posal would undermine the functional interpretation of IOR as a for-
aging facilitator, which is predicated on the inhibition of attention.
However, in numerous laboratories, IOR has recently been observed to
have an impact on nonspatial discrimination tasks. For example, using
a short (100-ms) cue-target stimulus onset asynchrony (SOA) with a
variety of longer ones, Lupiáñez, Milán, Tornay, Madrid, and Tudela
(1997) found IOR with both simple detection and color discrimination
tasks (though with the discrimination task, IOR appeared at longer
SOAs than with the detection task). In addition, Pratt, Kingstone, and
Khoe (1997) have found evidence for IOR in a form discrimination
task (see also Cheal, Chastain, & Lyon, 1998), and Handy, Jha, and
Mangun (1999) have demonstrated a d ' effect using a go/no-go size
discrimination task. Whatever the cause of the discrepancy,2 the fact
Address correspondence to Raymond Klein, Department of Psychology,
Dalhousie University, Halifax, Nova Scotia B3H 4J1, Canada; e-mail:
1. We use the term IOR (which we think has been overextended recently in
the literature on cognitive inhibition) to refer to inhibition generated by and
measured in spatial behavior.
2. More than one cause is likely responsible. Although a time-course differ-
ence works well for resolving the discrepancy between Lupiáñez et al. (1997) and
Pontefract and Klein (1988), both of whom used a cue-target paradigm, a differ-
ent explanation is needed for the dissociation (IOR with detection and localiza-
tion, but not with color or form discrimination) reported by Tanaka and Shimojo
(1996), who used a target-target paradigm with relatively long intertrial intervals.
In this case, IOR from the preceding target may be masked by a response-repeti-
tion heuristic (cf. Keele, 1973, pp. 91–93). Compared with a simple detection
task, in a complex discrimination task, there will be greater utility for the subject
to bypass the response selection stage by just repeating the previous response
when the subject notices that a similar target has appeared on successive trials.
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PSYCHOLOGICAL SCIENCE
Raymond M. Klein and W. Joseph MacInnes
that IOR can be obtained with nonspatial discriminations reopens the
possibility that attention is inhibited from returning to previouslyinspected locations (see Taylor & Klein, 1998a).
A second challenge came from studies suggesting that IOR could
be maintained at only one location—the one most recently stimulated.
Pratt and Abrams (1995) presented a cue at either one or, in succes-
sion, both possible target locations before presenting a target. In the
two-cue condition, they found inhibition only at the most recently
cued location. Gibson and Egeth (1994) explained this result and
applied it to their own findings by proposing that IOR generated by a
cue was canceled by presentation of a subsequent stimulus at a task-
relevant location, a mechanism they called disinhibition of return.
Although Pratt and Abrams (1995; Abrams & Pratt, 1996) observed
IOR only at the most recently cued location or region, Tipper, Weaver,
and Watson (1996) disputed this finding, and more recently, Kingstone
and his colleagues (Danziger, Kingstone, & Snyder, 1998; Snyder &Kingstone, in press) provided compelling evidence for IOR at more
than one location. Similarly, Klein et al. (1998) showed that disinhibi-
tion of return is an unlikely mechanism that is not needed to explain
Gibson and Egeth’s (1994) pattern of results.
Finally, failures to replicate the original finding of IOR following
serial visual search (Wolfe & Pokorny, 1990; see also Klein & Taylor,
1994) provided a direct challenge to the functional interpretation of
IOR. Recently, the pattern reported by Klein (1988) was replicated in
two different laboratories, but only when the search array remained
visible when the probe was delivered (Müller & von Mühlenen, 1999;
Takeda & Yagi, in press).3 Takeda and Yagi conducted a close replica-
tion of Klein (1988). Their data are shown in Table 1, along with
Müller and von Mühlenen’s remarkably similar pattern. The demon-
stration (Abrams & Dobkin, 1994; Tipper, Driver, & Weaver, 1991;
Tipper, Weaver, Jerreat, & Burak, 1994) that IOR may be tagged to
objects (rather than, or in addition to, environmental locations) pro-
vides a theoretical rationale for the importance of maintaining the dis-play elements in order to observe IOR after a visual search task: If,
during visual search, IOR tags objects in the scene once they have
been inspected, then when the scene is removed (and the objects,
therefore, disappear), the inhibition should likewise be removed.
Indeed, Tipper, who has been a consistent adherent of the view that
IOR functions to facilitate visual search, has made precisely this sug-
gestion (Tipper et al., 1994, pp. 495–496).
DOES INHIBITION OF RETURN GUIDE NATURAL
VISUAL SEARCH?
Our purpose in conducting the present study was twofold: (a) to
determine whether IOR would be observed in the behavior of the ocu-lomotor system when an observer was searching a complex scene for
a camouflaged target and (b) to replicate and extend the finding that
the maintenance of IOR depends on the persistence of the scene being
searched. To accomplish these goals, we presented observers with pic-
tures taken from the “Where’s Waldo?TM” series of picture books,
wherein Waldo (a curly-haired man with a walking stick, red-and-
white striped shirt, and blue slacks) is hidden in a variety of dense and
colorful environments. A long-robed, white-haired wizard is an alter-
nate or additional target in some of the pictures. We presented each
picture superimposed on a central fixation disk whose removal sig-
naled the participants to look for Waldo until they detected a probe
stimulus (reappearance of the disk in a new location), at which time
they were to reacquire it. We placed this probe carefully on the picture
after several saccades had been made; when the probe appeared, thepicture either was simultaneously removed or continued to be dis-
played. The probe was placed on an equi-eccentric circle around the
current fixation such that one of the possible probe locations was the
immediately preceding fixation location (one-back; Experiment 1) or
the fixation immediately preceding that one (two-back; Experiment 2),
and the remaining locations varied in angular separation from this
location (see Figs. 2 and 3). We expected that if IOR were present, sac-
cadic reaction times (SRTs) would increase the closer the probe was to
a previous fixation. If IOR were coded in relation to the scene (rather
than space per se), then maintenance of IOR would depend on main-
tenance of the scene (IOR would not be observed when the scene was
removed).
VOL. 10, NO. 4, JULY 1999 347
Fig. 1. Hypothesized distribution of inhibition of return in visual
search. Filled circles in the top rectangle indicate the locations of dis-
play items in a search task on a target-absent trial. The middle rectan-
gles illustrate that following a difficult search task (serial), inhibitory
tags are left behind at each distractor location “visited” by attention; in
contrast, inhibition is not left behind at distractor locations following
an easy (parallel) search task. Probes presented in new locations
(“Off”) and locations previously occupied by distractors (“On”) are
illustrated in the bottom rectangles. (Redrawn from Klein, 1988.)
3. Klein (1988) found evidence of inhibitory tagging in visual search even
though (as in Wolfe & Pokorny, 1990, and his own later studies) the search
array in this study was removed prior to presentation of the luminance probe.
That maintenance of the search array is necessary to observe IOR following
search suggests (Müller & von Mühlenen, 1999) a possible explanation for this
discrepancy: In Klein’s initial experiments (1988), conditions may have been
optimal for visible persistence of the search scene (high-luminance setting on
a Tektronix 604 oscilloscope with p31 phosphor in a dark room) after the
search array was “turned off.”
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PSYCHOLOGICAL SCIENCE
Inhibition of Return Facilitates Foraging
A general goal of the project was to bring a degree of ecologicalvalidity into the IOR literature. Of course, the “Where’s Waldo?” pic-
tures and task are no more natural than driving a car or conducting a
search on the World Wide Web. However, because they are much more
complex and interesting than the displays and tasks typically used in
studies of attention, they provide an excellent environment for explor-
ing natural search behavior. Finally, besides generating reaction times
to probe stimuli delivered by the experimenter, our method also pro-
vides a rich source of information on oculomotor behavior during the
search task itself.
METHOD
Participants
Eight adult volunteers, including one of the authors (W.J.M.), par-
ticipated in Experiment 1; 6 new adult volunteers participated in
Experiment 2. All were either students or employees of Dalhousie
University.
Stimuli and Apparatus
Stimuli were eight scenes scanned from three “Where’s Waldo?TM”
books4 and modified for our requirements. We chose pictures in which
there was a relatively high density of distracting information, in which
there was no black and very little dark gray, and in which Waldo, when
present (in half the pictures), and the potential distractors were rela-
tively small. The fixation and probe stimuli were black circles thatsubtended 0.5° of visual angle and had transparent centers 0.1° in
diameter. Stimuli were presented on a 17-in., SVGA,ViewSonic mon-
itor (640 × 480 pixels, 256 colors) in an area measuring 25.0° (width)× 19.4° (height) at a viewing distance of 71 cm. Scenes were drawn or
modified in a single screen refresh.
An EyeLink TM video-based eye-tracking system was used to mon-
itor each participant’s direction of gaze every 4 ms with a resolution of
0.1° or better. With this system, information about changes in direction
of gaze was available to the experimental program within approxi-
mately 20 ms.
Design and Procedure
Each experiment consisted of a block of 288 trials in which each of
the eight Waldo pictures was used equally often. The sequence of
events in a typical trial is shown in Figure 2. Each trial began when the
subject pressed the space bar to indicate that he or she was looking atthe fixation circle at the center of the screen. If fixation was stable, a
picture was displayed. The fixation stimulus remained present for an
additional 700 ms, and participants were instructed to maintain fixa-
tion until it disappeared and then to begin searching. Participants were
told to search for the wizard if they found Waldo in a particular pic-
ture. Examples of Waldo and the wizard were shown to subjects who
were not already familiar with them. Waldo was present in half the pic-
tures, and the wizard in none, so that participants would continue
searching pictures that had become familiar. Participants were
informed that on most trials, after a random period, the fixation disk
would be presented somewhere on the screen, and that when it reap-
peared they should stop searching and shift their gaze to it as quickly
as possible. On half the trials, at the time of this probe presentation,
the search picture remained present until the probe was fixated; on theremaining half of the trials, the picture was removed, leaving only the
probe present on a gray screen.
Any saccades or drifts (of more than 1°) that occurred before the
removal of the fixation dot resulted in a warning beep and recycling of
the trial. Once the fixation dot was removed, subjects were free to
search the picture as they saw fit. After a variable number of saccades
(minimum of 4), the probe was presented (approximately 20 ms after
saccade termination) at a location determined by the current and pre-
vious fixation positions according to the following procedures (illus-
trated in Fig. 3a). In Experiment 1 (one-back condition), the current
348 VOL. 10, NO. 4, JULY 1999
4. The displayed scenes were excerpted from three books by Martin Hand-
ford (Grolier Ltd., London).From Where’sWaldo?” (1987), we used theBeach,
theMuseum, theDepartment Store (withWaldo), andthe Fair; from FindWaldo
Now (1988), we used Once Upon a Saturday Morning (with Waldo) and The
Last Days of the Aztecs (with Waldo); and from The Giant Waldo Search
(1989), we used The Great Ball Game Players (with Waldo) and the Deep Sea
Divers.Additional information about the scenes is available from the authors.
Table 1. Difference in probe reaction time (in milliseconds) between targets presented at locationsoccupied by distractors and targets presented at new locations following serial and parallel search inrecent studies
Search Inferred
Study Serial Parallel inhibition of return
Klein (1988, Experiment 1) 39 7 32Klein (1988, Experiment 2) 62 16 46
Display maintainedTakeda and Yagi (in press, Experiment 2) 63 36 27Takeda and Yagi (in press, Experiment 3) 68 35 33Müller and von Mühlenen (1999, Experiment 2) 108 71 37
Display removedTakeda and Yagi (in press, Experiment 1) 54 49 5Müller and von Mühlenen (1999, Experiment 1) 42 39 3
Note. The data are from target-absent trials and are collapsed across set size (see Klein, 1988, for a justification).
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PSYCHOLOGICAL SCIENCE
Raymond M. Klein and W. Joseph MacInnes
gaze position became the center of an imaginary circle with a radius
equal to the distance between the current and penultimate (immedi-
ately preceding) fixation. Possible probe locations were centered on
the circumference of this circle, appearing equally often at the follow-
ing six angles relative to the previous fixation location: 0°, 60°, 120°,
180°, 240°, and 300°. Probe presentation was delayed if the calculat-
ed radius was less than 1° or would place any one of the possible
probes off the screen. If no suitable gaze position could be found with-
in 12 saccades, the trial was terminated without a warning beep. In
Experiment 2 (two-back condition), probe locations were determined
in the same manner except that the radius was based on the ante-
penultimate fixation (i.e., preceding the one used in Experiment 1; see
Fig. 3a).
A saccade was said to acquire the probe if the landing coordinates
of the saccade were within 1° of the probe’s center. All other fixationlocations were classified as misses. The primary dependent variable
was the SRT to acquire the probe when it was acquired within one sac-
cade.
RESULTS AND DISCUSSION
Probe Acquisition
Trials were excluded from this analysis if there was a blink (4.1%,
Experiment 1; 11.9%, Experiment 2) or a probe could not be delivered
(5.6%, Experiment 1; 20.4%, Experiment 2). Across the remaining tri-
als, probes were acquired in one saccade more frequently when the
scene was removed (63.1%) than when it remained present (44.6%),
F (1, 12) = 69.6, p < .001, a difference likely due to increased salience
of probes that were not superimposed on a complex scene. The reac-
tion times of these saccades provided the primary data for testing the
hypotheses.
Mean SRT for trials on which the probe was acquired in the first
saccade (see Fig. 4) were subjected to a 2 (one- vs. two-back) × 2
(scene removed vs. maintained) × 4 (angular distance5 from the previ-
ous fixation) mixed analysis of variance. There was a significant main
VOL. 10, NO. 4, JULY 1999 349
A
B1-back
2-back
3-back
Potential Probe Locationsin Experiment 1 (one back)
Gaze Position attime of probe
Saccade Path0°
-120°
180°
-60°
120°
2
3
60°
1
Fig. 3. Placement of probes (a) and method of analyzing the direction
of freely executed saccades as a function of the angular difference
between a saccade and preceding fixation locations (b). In (a), the last
three saccades prior to probe presentation are shown. The empty cir-
cle shows the location of gaze at the time of probe presentation, and
the solid circles show the six possible locations for the probe target in
Experiment 1. Only one probe location was selected for each trial.
Each of these six locations was used equally often, but for purposes of analysis, the locations were collapsed on the basis of the absolute dif-
ference in direction between the probe target and the penultimate fix-
ation location (marked 0°). The checkerboard-filled circle marks the
location of the antepenultimate fixation that was used to generate the
probe locations in Experiment 2. In (b), four saccades preceding the
delivery of the probe target are shown, along with the angular differ-
ence between the last saccade in this sequence and each of the three
preceding fixations.
5. For present purposes, the six different angles were collapsed into four
angular distances: 0°, ±60°, ±120°, and 180° (as shown in Fig. 3a).
Fig. 2. Sequence of displays in the experiments. The participants indi-
cated when they were ready while fixating the disk at the center of a
blank display (a). The search array (shown here as a brick-wall pat-
tern; see the text for a description) was then added (b). Removal of the
fixation disk was the signal to begin the search (c). After four or more
saccades (d), a probe stimulus (reappearance of the fixation disk) was
presented on an imaginary circle that was centered about the present
fixation and had a radius equal to the distance between the current and
penultimate (Experiment 1) or antepenultimate (Experiment 2) fixa-
tions. When the probe appeared, the search array either remained or
was removed (e). See the text and Figure 3 for further details.
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PSYCHOLOGICAL SCIENCE
Inhibition of Return Facilitates Foraging
effect of scene removal, F (1, 12) = 28.5, p < .001, reflecting longer
SRTs when the scene was removed (257.7 ms) than when it remained
present (209.9 ms) during probe presentation. Sensory or motor mech-
anisms could be responsible for this difference. As an example of the
former, the global transient associated with removal of the scene might
have masked the local transient associated with the onset of the probe
(as occurs in change blindness; see Simons & Levin, 1997, p. 263),
thus delaying its perception. Alternatively, removing the visual data
used to compute oculomotor parameters for the Waldo search might
have caused a refractory period for the programming of subsequent
saccades, until clearing and resetting of the visual-motor representa-
tions was completed. The results strongly favor this possibility: Sac-
cades that were made immediately after probe presentation but were
not directed to the probe showed the same cost (46.4 ms) of scene
removal as did saccades that immediately acquired the probe.
The main effect of angular distance was significant, F (3, 36) =8.03, p < .001, as was the predicted interaction between angular dis-
tance and scene removal, F (3, 36) = 8.15, p < .001. This pattern was
further qualified by a three-way interaction involving experiment, F (3,
36) = 3.299, p < .05. To determine whether the results supported the
predictions outlined earlier, we analyzed the two conditions (scene
removed and scene maintained) separately for the presence of IOR.
There were no significant effects or interactions (all F s < 1) when
the scene was removed: The time to find the target was unaffected by
its angular distance from the immediately preceding fixation (one-
back) or the fixation before that one (two-back). In contrast, when the
scene was maintained, the effect of angular distance was significant,
F (3, 36) = 16.24, p < .001, as was the interaction with experiment,F (3,
36) = 3.12, p < .05.
We expected that when the scene was maintained, the slowestSRTs would be to probes presented in the same location as a previous
fixation (0°), and that there would be a monotonic decrease in SRT
with increasing angular distance from this location. Precisely this pat-
tern was observed in the two-back condition; in the one-back condi-
tion, the pattern was similar except that SRTs to targets presented at
the immediately prior fixation (0°) were faster than expected. The two-
back data (which were collected to help address this unexpected find-
ing) suggest that one or more processes were operating in the one-back
condition to mask the inhibitory effect in the 0° location. The most
recently encoded region of the scene is likely to be the most accurate-
ly represented, so changes (e.g., probe presentation) in this region may
be more conspicuous than other changes. Alternatively, facilitation,
akin to that elicited in a peripheral cuing paradigm, might mask the
inhibition in this region. Although there are reasons to favor the con-spicuity account,6 whatever the explanation, the important point is that
in the two-back condition, when both processes would be less likely to
be operating, the data pattern obtained was precisely the pattern pre-
dicted by the proposal that IOR was operating in this task.
Saccades Prior to Probe Presentation
If IOR is elicited following each fixation, then the “freely” gener-
ated saccades made by our participants prior to the appearance of the
probe should show evidence of this inhibition. We explored this possi-
350 VOL. 10, NO. 4, JULY 1999
Fig. 4. Saccadic reaction time when the first posttarget saccade
acquired the probe target. Results are shown separately for Experiment
1 (a), with probes at the one-back location, and Experiment 2 (b), with
probes at the two-back location. Within-subjects 95% confidence
intervals, appropriate for comparing different angular distances from
the same curve, were generated according to Loftus and Masson
(1994) using the Distance × Subject MSE s from analyses of variance
done separately for the scene-maintained and scene-removed condi-
tions of each experiment.
6. First, the viability of the facilitation account depends on the spread of
inhibition exceeding the spread of facilitation, a relation that has not yet beendemonstrated. Second, if facilitation were operating, we ought to have seen evi-
dence of it in the scene-removed condition. Finally, we split SRTs to probe tar-
gets (in the scene-maintained condition) in the one-back study on the basis of
the preceding fixation duration. The analysis supports the conspicuity account
in that following longer-than-average fixations (which permit more encoding),
there was a 20-ms advantage for targets at the location of preceding fixation
compared with targets ±60° from this location; this advantage was not present
following shorter-than-average fixations. This is not what would be expected
from a facilitation account, because if facilitation is initiated when a location is
fixated, and begins to decay from this point, then measured IOR should be larg-
er following longer-than-average fixation durations.
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PSYCHOLOGICAL SCIENCE
Raymond M. Klein and W. Joseph MacInnes
bility by examining the angle between a preprobe saccade’s vector and
the vector connecting the launching point for that saccade with the
previous fixation for all preprobe saccades that would have allowed
the delivery of a probe (see Method section). The relative angles were
first sorted into six equal pie-shaped regions, with one centered on the
vector from the preceding fixation (0°) and the remaining five going
around an imaginary circle (as in Fig. 3a), and the six regions werethen collapsed as in the SRT analysis (see footnote 5). This process
was repeated (see Fig. 3b) using the fixation before the immediately
preceding fixation (two-back) and the fixation before that one (three-
back). As can be seen in Figure 5, there was a very strong tendency for
saccades made during search to be launched in directions away from
previous landing positions. The proportion of saccades whose direc-
tion would shift gaze in the direction of previous fixations was rela-
tively low, and there was, in each case, a significant effect of angular
distance in the form of a monotonic gradient—one-back: F (3, 36) =
41.1, p < .001; two-back: F (3, 36) = 25.2, p < .001; three-back: F (3,
36) = 13.18, p < .001.
Although it is theoretically appealing to assert that IOR causes this
directional bias in oculomotor search, it is also possible that the ocu-
lomotor bias causes IOR. Because oculomotor responses are likelyinitiated by a winner-take-all algorithm mediated by lateral inhibition
(and implemented in the superior colliculus), any asymmetric prepa-
ration would result in inhibition of the least-prepared saccades.
Although we believe that both behavioral and neurophysiological
studies may be able to distinguish between these alternatives,7 for pre-
sent purposes we need not be disturbed by the cause-effect ambiguity,
because either way the visual system is keeping track of regions of the
scene that need not be reexamined.
SUMMARY AND CONCLUSION
There are three main findings from this study of oculomotor
behavior during search of complex scenes. First and foremost, when
probe targets were presented in a scene during search, participants
were slower to find (saccade to) them when they were in the gener-
al region of preceding fixations than when they were in a new
region. This finding is precisely what is predicted by the proposal
that the function of IOR is to facilitate foraging (Klein, 1988; Pos-
ner & Cohen, 1984; Tipper et al., 1994). Moreover, the monotonic
function relating SRT to angular distance (particularly in Experiment
2) is consistent with the idea that there is a gradient of inhibition
around a previously attended (Maylor & Hockey, 1985) or fixated
region.
Second, this inhibition was not observed if the search array was
removed when the probe was delivered. This finding is consistent with
evidence showing that IOR is attached to objects in a scene (e.g.,
Abrams & Dobkin, 1994; Tipper et al., 1991, 1994) and also replicates
and extends recent demonstrations (Müller & von Mühlenen, 1999,
Experiments 1 and 2; Takeda & Yagi, in press) that IOR measured in
manual reaction time to probes following serial search depends on
maintenance of the search array (as predicted by Tipper et al., 1994).
Third, the freely executed saccades of participants prior to the pre-
sentation of the probe showed the same bias that was evident in the
time to acquire the probe; that is, each saccade was more likely to
repeat the previous direction than to reverse it. This is precisely what
one would expect if the IOR that was seen in the time to find the probe
was operating equivalently on the freely made saccades during search.
And it is precisely such freely made saccades during visual search of
a scene that ought to be influenced by inhibition serving as a foragingfacilitator.
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Acknowledgments—The research described here was supported by a Nat-ural Sciences and Engineering Research Council of Canada CollaborativeProjects Grant (“Selection of Objects by the Primate Oculomotor System”)to R. Klein (principal investigator), D. Munoz, P. McMullen, and T.Trappenberg.
7. A behavioral approach we are pursuing using the same “Where’s
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(RECEIVED 8/24/98; REVISION ACCEPTED 1/12/99)
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T h i s d o c u m e n t i s a s c a n n e d c o p y o f a p r i n t e d d o c u m e n t . N o w a r r a n t y i s g i v e n a b o u t t h e
a c c u r a c y o f t h e c o p y . U s e r s s h o u l d r e f e r t o t h e o r i g i n a l p u b l i s h e d v e r s i o n o f t h e m a t e r i a l .