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COGNITIVE NEUROPSYCHOLOGY, 2002, 19 (4), 361–380

TRANSIENT BINDING BY TIME:


NEUROPSY CHOLOGICAL EVIDENCE FROM
ANTI-EXTINCTION

Glyn W. Humphreys and M. Jane Riddoch


University of Birmingham, UK

Gudrun Nys
University of Utrecht, The Netherlands

Dietmar Heinke
University of Birmingham, UK

Anti-extinction occurs when there is poor report of a single stimulus presented on the contralesional
side of space, but better report of the same item when it occurs concurrently with a stimulus on the
ipsilesional side (Goodrich & Ward, 1997). We report a series of experiments that examine the factors
that lead to anti-extinction in a patient GK, who has bilateral parietal lesions but more impaired identi-
fication of left-side stimuli. We show a pattern of anti-extinction when stimuli are briefly presented,
which is followed by an extinction effect when stimuli are left for longer in the visual field. In Experi-
ments 1 and 2 we present evidence that the anti-extinction effects are determined by stimuli onsetting
together, and it is not apparent when stimuli are defined by offsets. In Experiments 3 and 4 we report
that performance is not strongly affected by whether the same or different tasks are performed on the
ipsi- and contralesional stimuli, and the anti-extinction effect also survives trials where eye movements
are made to right-side stimuli. Experiment 5 provides evidence that anti-extinction is due to temporal
grouping between stimuli, rather than to increased arousal or cueing attention to the contralesional side.
Experiment 6 demonstrates that anti-extinction dissociates from GK’s conscious perception of when
contra- and ipsilesional stimuli occur together. We interpret the data as indicating that there is uncon-
scious and transient temporal binding in vision.

A basic question confronting vision scientists is chronised and fire at a different time from cells
how a coherent visual world can be coded following responding to other objects. Consistent with this
its initial decomposition by the brain into separable account, there can be time-locked firing of cells
dimensions. One answer to this question, suggested when stimulus elements group but de-synchronised
by neurophysiological data on synchronised neural firing when elements do not group (Singer & Gray,
firing, is that the different features of an object are 1995). There is also some behavioural evidence for
bound by time (e.g., Eckhorn, 1999; Singer & temporal synchronisation in the input being impor-
Gray, 1995). Temporal binding can take place if tant for feature binding in humans (Elliott & Mul-
cells responding to features from one object are syn- ler, 1998; Fahle, 1993). For instance, Elliott and

Requests for reprints should be addressed to Glyn W. Humphreys, Behavioural Brain Sciences, School of Psychology, University
of Birmingham, Birmingham B15 2TT, UK. (Email: g.w.humphreys@bham.ac.uk).
We thank GK for his unending good humour and patience in carrying out this work. The research was supported by grants from
the Medical Research Council (UK) and the European Union (Basic Research Action).

Ó 2002 Psychology Press Ltd 361


http://www.tandf.co.uk/journals/pp/02643294.html DOI:10.1080/02643290143000222
HUMPHREYS ET AL.

Muller reported that detection of a target formed by ipsilesional stimuli (e.g., identify the contralesional
grouped elements was facilitated if the positions of item and detect the presence of the ipsilesional
the elements were pre-cued by features that one), relative to when identical responses were
appeared synchronously, relative to when the pre- made to both (identify both items). Nevertheless,
cued features were presented asynchronously. This we note that trends for anti-extinction were present
facilitation occurred even when observers could not in all cases. We test the effects of having the same
detect when pre-cue features appeared synchro- and different responses in Experiment 3 here, but
nously. In this paper we present converging neuro- fail to find significant effects. Instead we provide
psychological evidence for unconscious, time- evidence that anti-extinction can be due to tran-
based binding of visual elements, based on the sient, temporal grouping between contra- and
phenomenon of “anti-extinction” (Goodrich & ipsilesional items, due to the stimuli having a com-
Ward, 1997). mon onset. The phenomenon provides evidence for
Many patients manifest “extinction” following a time-based binding in human vision.
brain lesion: A contralesional stimulus can be We report six experiments conducted on GK, a
reported when presented in isolation but it is not patient with bilateral parietal lesions following two
reported when an ipsilesional stimulus is exposed at consecutive strokes. Although GK has bilateral
the same time (Karnath, 1988). This is frequently lesions, he shows a strong spatial bias in visual selec-
attributed to a spatial bias in visual selection due to tion, manifesting neglect on many clinical tasks
the brain lesion, which either disrupts input into and, with stimuli presented for relatively long
the selection process or unbalances the selection durations, there is extinction for items on his left
process itself (Duncan, Humphreys, & Ward, (e.g., Boutsen & Humphreys, 2000; Gilchrist,
1997). The bias in selection results in ipsilesional Humphreys, & Riddoch, 1996; Humphreys,
stimuli winning the competition for selection when 1998). Despite extinction occurring with long
ipsi- and contralesional items appear simulta- exposures, we show here that there is anti-
neously (see Heinke & Humphreys, in press; extinction when stimuli are exposed more briefly.
Mozer & Behrmann, 1990, for simulations). The Experiment 1 demonstrates the different time
phenomenon of “anti-extinction” is the opposite of courses of the two effects, with anti-extinction
this. Reported in only one previous paper by Good- being replaced by extinction as the time post-onset
rich and Ward (1997), anti-extinction occurs when increases. Experiment 2 reveals that common
a patient is unable to discriminate a contralesional onsets of contra- and ipsilesional stimuli may be
stimulus presented in isolation but is able to report necessary for anti-extinction, since the effect did
the same stimulus when an ipsilesional item is not occur when the stimuli had common offsets.
exposed simultaneously. Goodrich and Ward Experiment 3 shows that the effect occurred irre-
argued that anti-extinction came about due to a spective of whether the same task was carried out on
form of response linkage between the stimuli. They the contra- and ipsilesional items, and Experiment
suggested that a single contralesional stimulus 4 demonstrates that anti-extinction survives on tri-
alone may not have been sufficient to activate a als on which an eye movement is made to the
response. However, the presence of an ipsilesional ipsilesional item. Experiment 5 contrasts an
stimulus simultaneously primed the same response account of anti-extinction in terms of temporal
to the contralesional item, so that responses could binding with alternatives based on the spread of
then be made to both items. According to Good- spatial attention and on the influence of increased
rich and Ward, this response priming benefit would arousal, and suggests that temporal binding pro-
occur when (1) the contra- and ipsilesional task are vides a better explanation. Finally we assessed
similar to each other, and (2) the same response whether GK was aware when stimuli onset
mechanisms are shared. Consistent with this they together. Experiment 6 evaluated the relation
found that anti-extinction was reduced when their between anti-extinction and “prior entry,” the ten-
patient made different responses to the contra- and dency of some patients with neglect to judge that an

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TRANSIENT BINDING BY TIME

ipsilesional stimuli precedes a contralesional one of one or two letters). The experimenter wrote
when they are presented simultaneously (Rorden, down the responses and initiated the next trial by
Mattingley, Karnath, & Driver, 1997). Report of pressing the space bar.
briefly presented stimuli was better with simulta- The procedure for Experiment 2a was the same
neous than with staggered presentations, even except that, prior to the target display, two symmet-
though the contralesional stimulus had to be pre- rical figure-of-eight masks appeared along with the
sented first in order for the stimulus onsets to be fixation asterisk, each centred 7mm from fixation.
equated phenomenally. Hence GK’s conscious The target letters were L, H, T, or F, which could
judgements of temporal order dissociate from the be formed by removing contours from the pre-
effects of temporal simultaneity on stimulus report. masks. The durations were the same as for Experi-
There is unconscious binding of visual events by ment 1, and there were two blocks of 72 trials for
time, to facilitate perceptual report. each duration except 75 ms, for which there were
four trial blocks. Blocks were presented in a random
order. For Experiment 2b, there was no pre-mask
GENERAL METHOD but stimulus displays were followed by a figure-of-
eight post-mask. With this post-mask, the con-
For Experiments 1–3, 5, and 6, the stimuli were tours of letters (when present) remained in the field
presented on an Apple MacIntosh computer using and were added to by the contours of the mask. The
either V-scope (Experiments 1, 3, 5, and 6) or letters were the same as in Experiment 2a, but the
Psyscope (Experiment 2). For Experiment 4, the task was changed from letter identification to
stimuli appeared on an IBM pentium computer detection; GK had to decide whether two, one, or
using MEL software. Letters were presented 7 mm no letters were presented. There was just one stim-
from their centre to fixation, and they were 6 mm x ulus duration: 150 ms. There were three blocks of
6 mm drawn in Times Roman font. The viewing 72 trials.
distance was approximately 80 cm. Experiment 3 contained two subexperiments. In
For Experiment 1 there were four types of dis- Experiment 3a the letters A, B, C, and D were
play: blank trials (no letters present), one-left, one- again used. There were two task conditions: Same
right, and two-letters (bilateral, one left and one task (identify the letters) and Different task (iden-
right). On two-letter displays, letters were never tify any letters on the left and detect any letters on
repeated. Letters were drawn from the set A, B, C, the right). There were two blocks of 72 trials for
and D. Each letter appeared randomly to the left or each task condition (24 blank, 24 single-item—one
right of fixation an equal number of times. The dif- left or one right—and 24 two-item trials), with the
ferent stimulus durations were presented in sepa- blocks presented in ABBA order. The stimulus
rate trial blocks, with there being 72 trials in one duration was 150 ms and no masks were employed.
block: 24 two-letter, 12 one-left, 12 one-right and In Experiment 3b the procedure was the same
24 blank trials. The trials were randomised within except that only the letters X and O were used, rep-
each block. For six stimulus durations there were licating Goodrich and Ward (1997). On two-item
four blocks of trials (75, 300, 450, 600, 750, and 900 trials the letters were identical half the time (either
ms respectively); there were six blocks of trials with OO or XX) and differed on the remaining trials
a stimulus duration of 150 ms. The order of the (OX or XO), for both the Same and the Different
stimulus durations was randomised. GK sat at a task conditions.
viewing distance of about 80 cm. Each trial began In Experiment 4, the procedure was the same as
with a fixation asterisk for 1500 ms in the centre of in Experiment 1 except that there were just two
the screen. GK reported when he was fixated. durations: 180 ms (five blocks of 72 trials) and 450
There followed a blank interval of 150 ms and then ms (four blocks of 72 trials). The blocks were pre-
the stimulus display. GK made a verbal letter iden- sented in random order. Unlike in the other studies,
tification response (either “nothing” or the identity GK’s eye movements were recorded using an

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HUMPHREYS ET AL.

ISCAN remote pupil tracker system (resolution ms or (2) the right led the left by 720 ms. In the
0.3° of visual angle). Prior to each trial block, GK’s temporal judgement task, only displays with two
eye movements were calibrated with dynamic stim- letters were presented and there were 24 trials for
uli (a circle containing a letter that moved from one each of the sequential exposures and 48 trials with
location to the next; GK had to identify the letter simultaneous presentations. The left and right let-
present when the circle was stationary). ters always differed in colour and they were each red
In Experiment 5, the fixation display was on half the trials and green on the remaining. In the
changed to contain three black figure-of-eight colour identification task, the two-letter trials were
pre-masks, one at the centre and the other two accompanied by 24 one-left and 24 one-right letter
each separated by one letter width from the central trials, with the single target letter being red on half
mask, in the left and right visual fields respectively. the trials and green on the other. The blocks with a
GK was asked to fixate the central mask. After 450 ms stimulus onset asynchrony (SOA) were run
1500 ms, contours in the central mask offset to in one session and the blocks with a 720 ms SOA
create a black target letter. Simultaneous with the were run in another. Within each session there were
offsets in the central pre-mask, 0, 1, or 2 “onset” two blocks for each task (temporal order judgement
target letters were presented in the gaps between and colour identification), presented in an ABBA
the three pre-masks. The target letters were drawn order. Letters were presented following the expo-
at random from the set: L, H, T, and F. In Experi- sure of a central fixation cross for 1500 ms, and the
ment 5a there were four blocks of 72 trials in which presentation time for the letters was always 345 ms.
there were 24 displays with no new onsets, 12 with There was no mask.
one onset letter in the left field, 12 with one onset
letter in the right field, and 24 with two onset let-
ters (one right and one left). In blocks 1 and 4, the
onset letters appeared in red, to enhance their CASE REPORT
grouping. In blocks 2 and 3, the onset letters were
black. There was a central offset letter on all trials. GK was 60 years old at the time of testing. Con-
The letter display appeared for 300 ms and GK secutive strokes in 1986 produced occipito-
was asked to identify all the letters present irre- parietal damage in the right hemisphere and bilat-
spective of their colour or how they were created eral parietal-temporal damage. An MRI scan is
(by an offset or onset). In Experiment 5b, the provided in Boutsen and Humphreys (2000). GK
onset letters were always red and the central letter, has a number of neuropsychological deficits
created by the offset of contours from a mask, was including attentional dyslexia (Hall, Humphreys,
again black. In this experiment, GK was asked just & Cooper, 2001) and Balint’s syndrome
to identify the black letter. This provided a test of (simultanagnosia and optic ataxia; Cooper &
whether poor report of the central target, in Humphreys, 2000; Humphreys, Romam, Olson,
Experiment 5a, was due to lateral masking from Riddoch, & Duncan, 1994). Although there is
the onset letters. generally poor selection of multiple visual stimuli,
Experiment 6 examined two tasks: report of the performance is worse on his left-side. There is left
temporal order of red and green letters and report of visual neglect and left-side extinction with relative
the colours of the letters present. For each task a long stimulus exposures (Boutsen & Humphreys,
single block of trials contained three different time 2000; Gilchrist et al., 1996; Humphreys, 1998).
relations between the stimuli when two letters were The left-side extinction effect is modulated by
present. In one block, the letters either (1) appeared grouping based on the form and common surface
simultaneously or (2) the left preceded the right by properties of items in the left and right fields (see
450 ms or (3) the right preceded the left by 450 ms. Humphreys, 1998, for a review). Despite these
In the other block the stimuli again appeared simul- difficulties, GK is well-oriented in time and place
taneously or either (1) the left led the right by 720 and he has good comprehension.

364 COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4)


TRANSIENT BINDING BY TIME

EXPERIMENT 1: FROM ANTI- contralesional letters on two-item trials; c2 (1) =


EXTINCTION TO EXTINCTION 7.14, 5.01, and 3.94, all p < .05, for exposure dura-
tions of 75, 150, and 300 ms respectively. For dura-
Experiment 1 examined the time course of extinc- tions of 600 ms and above, the opposite pattern
tion and anti-extinction effects by varying the expo- occurred: There was better report of single
sure duration of contra- and ipsilesional stimuli in contralesional letters than of contralesional letters
an identification task. On each trial either no, one, on two-item trials; c2 (1) = 7.25, 14.8, and 55.55, all
or two letters were presented, with single letters p < .01, for exposure durations of 600, 750, and 900
appearing equally often on the contra- and ms respectively). On two-item trials, performance
ipsilesional sides. across durations 75–300 ms was better than that
across durations 450–900 ms, c2 (1) = 5.52, p < .02.
On trials with no letters there were only 2% false
Results and discussion alarms (708/720 correct). On trials with single
The results are given in Figure 1. For stimulus ipsilesional letters there were 21% errors across
exposures under 300 ms there were significant exposures. On two-item trials 62.5% of the errors
anti-extinction effects: There was poor report of were due to report of the right (ipsilesional) letter
single contralesional letters and better report of only. On all trials where two letters were identified,

Figure 1. (a) Example displays from Experiment 1. (b) The percentage correct letter identification responses made by GK on trials with
one-left, one-right, or two (bilateral) letters being presented, as a function of the exposure durations of the letters.

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HUMPHREYS ET AL.

the right was reported first. This held across all the EXPERIMENT 2: THE EFFECTS OF
experiments. COMMON OFFSET
Experiment 1 demonstrates two clear patterns of
performance, which changed as a function of the Experiment 2a: Stimuli defined by offsets
exposure duration of target letters. With brief expo-
Method
sures (of up to 300 ms or so), there was an anti-
The method was the same as for Experiment 1,
extinction effect. There was poor identification of
except that the stimuli were defined by offsets
single left-side letters, but report of the same letters
rather than by onsets. The procedure is outlined in
improved when a second, ipsilesional (right) stimu-
Figure 2a.
lus was presented simultaneously. This mirrors the
results originally reported by Goodrich and Ward
(1997). However, as the stimulus duration Results and discussion
increased, then the pattern of anti-extinction The results are presented in Figure 2b. There was
altered to one of extinction. There was then rela- no evidence for anti-extinction for durations below
tively good identification of single stimuli pre- 300 ms; c2 < 1.0, for all comparisons between single
sented in the left field, but poor identification of the left and two-item trials. In contrast, there was a
same items when an ipsilesional item appeared at clear extinction effect at longer exposures (450 ms
the same time. The results at longer durations repli- and above); c2(1) = 10.33, 7.01, 4.84, and 19.97, all
cate the extinction effect previously documented p < .05, for durations 450, 600, 750, and 900 ms
with this patient (Boutsen & Humphreys, 2000; respectively. On two-item trials GK identified only
Gilchrist et al., 1996; Humphreys, 1998). Indeed, the right-side letter on 73% of the trials. On no-
GK’s performance on two-item trials decreased as item trials there were just 1% false alarms (380/384
the stimulus exposure duration increased—a highly correct).
unusual pattern of performance. Unlike in Experiment 1, there was no clear evi-
Prior studies with normal observers demonstrate dence for anti-extinction at the shorter stimulus
that stimulus onsets can serve as strong cues for exposures here. Performance on two-item trials was
visual attention (Yantis, 1998; Yantis & Jonides, no longer better than on trials with single left tar-
1984, 1990). In Experiment 1, the stimuli were gets. This suggests that common onsets may be
defined by simultaneous onsets, and it may be that necessary to generate anti-extinction (in Experi-
these transient signals led to the anti-extinction ment 1 but not Experiment 2a). In contrast to this,
effect by cueing GK’s attention to the region occu- common offsets seem not to be effective in isolation
pied by both stimuli, on a proportion of two-item (common offsets are not sufficient). Experiment 2b
trials. There remains to explain why extinction sought to test the effects of stimulus offset in more
occurred subsequently, as the stimuli remained in detail, by presenting new stimuli again that were
the field, and we will return to this point in Experi- defined by common onset (as in Experiment 1), but
ment 4. Experiment 2, however, examined whether now replacing these stimuli with post-display
common onsets were important by (1) presenting masks. The post-display masks contained the fea-
the stimuli by means of offsets rather than onsets, so tures present in the letters, plus additional features
removing any effects due to onsets (Experiment that fell around the same spatial area. When such
2a), and (2) defining the stimuli by onsets but fol- masks are used, there is no offset of contours that
lowing them with masks designed so that the con- uniquely make up each letter. However, it is the
tours of the letter targets did not offset (Experiment case that the post-masks used here make perform-
2b). We ask, does the absence of common onsets ance more difficult, relative to when no masks are
disrupt the effect (i.e., are common onsets neces- employed (Experiments 1 and 2a). Consequently,
sary?), and does the effect still arise when the stim- the task changed from letter identification to detec-
uli do not offset together (i.e., are common offsets tion (2 vs. 1 vs. 0 items on a trial?). If the anti-
necessary?)? extinction effect is due to stimulus offsets occurring

366 COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4)


TRANSIENT BINDING BY TIME

Figure 2. (a) Example displays from Experiment 2a (targets defined by offsets). (b) The percentage correct letter identifications made by GK
in Experiment 2a.

close in time to onsets (under the short exposure Experiment 2b confirmed the anti-extinction
conditions of Experiment 1), then the effect should effect with short durations in a detection task, when
not be found here. Just one short exposure duration target letters onset but did not offset together.
was used, 150 ms. From this we conclude that common offsets are not
necessary for anti-extinction (though they still play
a contributory role when common onsets are pres-
Experiment 2b: Eliminating common offsets
ent). In contrast to this, Experiment 2a demon-
Method strated that the effect was eliminated when stimuli
The procedure is outlined in Figure 3a. were defined by offsets. The results suggest that
common onsets are critical.
Results and discussion Goodrich and Ward (1997), when first defining
The data are given in Figure 3b. There was a reli- the anti-extinction phenomenon, did not consider
able anti-extinction effect: single left-side items stimulus factors such as common onsets in great
were detected worse than both letters on two-item detail. Instead they suggested that the effect was
trials, c2(1) = 5.67, p < .025. On single-right item caused by response priming; report of the
trials there were 14% errors, all due to no-item contralesional stimulus was supported by the same
responses. There were 6% false alarms on blank response being made to the ipsilesional item. We
trials. tested this in Experiment 3. GK either had to iden-

COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4) 367


HUMPHREYS ET AL.

Figure 3. (a) Example displays from Experiment 2b (targets followed by masks to prevent contour offsets). (b) Percentage correct detection
responses in Experiment 2b.

tify the letters on a trial (same task for both letters, ent (XX or OO vs. OX or XO). Goodrich and
on two-item trials) or he had to identify the left- Ward reported different effects of item similarity in
side letter and detect the right-side one (present or the Same and Different task conditions. In the
absent?; different task for both letters, on two-item Same task condition (identify both letters), perfor-
trials). According to the response priming account, mance was better when the letters were different
anti-extinction should occur in the same-task but (OX or XO) than when they were the same (OO or
not the different-task condition. XX; see also Baylis, Driver, & Rafal, 1993). In the
Different task condition (detect right, identify left
letter), item similarity did not matter. The contrast-
EXPERIMENT 3: THE EFFECTS OF ing effects of item similarity provides converging
RESPONSE LINKAGE evidence for contrasting response strategies in the
Same and Different task conditions.
Experiment 3 contained two substudies. In Experi-
ment 3a we used the same letters as in the earlier
Experiment 3a: Different letters
studies, with two different letters always being
employed on each trial. In Experiment 3b we Method
employed a procedure more directly modelled on The method was the same as in Experiment 1
Goodrich and Ward’s (1997) study where, on two- except for the change to a detection response for
item trials, the letters were either the same or differ- right-side item in the different response condition.

368 COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4)


TRANSIENT BINDING BY TIME

Results and detected accurately (different task condition).


The results are illustrated in Figure 4a. The data There were 2% false alarms on blank trials (94/96).
were subjected to log-linear analysis with the fac-
tors being response (same vs. different), number of
items (one-left vs. two-items), and accuracy (num-
Experiment 3b: Same and different letters
ber correct vs. error). There were proportionately
more correct to error responses on two-item com- Method
pared with one-left trials, c2 (1) = 7.05, p < .01. This was the same as for Experiment 3a, except
There was also a trend for proportionately more that, on two-item trials, the letters were the same
correct responses when different tasks were per- on half the trials (OO or XX) and different on the
formed on left- and right-field stimuli, relative to remainder (OX or XO).
when the same task was performed, c2(1) = 1.24, p >
.05. There was no evidence of an interaction
between number of items and the task factor, or for Results
a three-way interaction (both c2 < 1.0). Single-right The results are presented in Figure 4b. The data
letters were both identified (same task condition) were again subject to a log linear with the factors
being response (same vs. different), number of
items (one- left vs. two-items), and accuracy (cor-
rect or error). There was a reliable interaction
between number of items and the proportion of
correct to error responses, c2 (1) = 9.49, p < .01, and
a trend for an interaction between response and
accuracy, c2 (1) = 2.35, p > 0.10. There were rela-
tively more correct to error responses on two-item
trials compared with single-left trials. There also
tended to be proportionately more correct
responses on different- than on same-response tri-
als. There was no evidence for an interaction
between response and number of items, or for a
three-way interaction between response, number of
items, and accuracy (both c2 < 1.0). The advantage
for two-item over single-left trials held irrespective
of whether the same or a different response was car-
ried out on letters in the left and right fields. There
was no difference in the accuracy of responses to
single-right letters in the two response conditions
(c2 < 1.0). There was just one false alarm on blank
trials (95/96 correct).
Performance on two-item trials was broken
down according to whether the letters were the
Figure 4. (a) Percentage correct responses in Experiment 3a. In the same (OO or XX) or different (OX or XO). On
same task condition, GK made identification responses to left and same-response trials there were 3/24 correct
right-side letters. In the different task condition, GK identified any responses to two identical letters and 15/24 to two
left-side letters and made a detection response to any right-side different letters. On different-response trials there
letters. The letters on each trial were always different. (b)
Percentage correct responses in Experiment 3b. The procedure was
were 11/24 correct responses on trials with identical
the same as in Experiment 3a, except that the letters were identical letters and 13/24 correct responses on trials with
on half the trials and different on the remaining. different letters. The effect of item similarity

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HUMPHREYS ET AL.

tended to be larger on same-response trials (identify Baylis et al. (1993) and Goodrich and Ward (1997).
both the left and right letters) than on different- Baylis et al. interpreted their results as indicating
response trials (identify the left letter and detect the impaired formation of “object tokens,” when stim-
right), c2(1) = 2.75, p > .05. On same-response tri- uli have the same response-relevant feature. How-
als, performance was better with different than with ever, the error patterns shown by GK suggest that
identical letters, c2(1) = 10.76, p < .01. On differ- his results could reflect a response bias away from
ent-response trials letter identity had no significant giving letters the same identity. Nevertheless, this
effect (c2 < 1.0). On different-response trials 19 bias was not apparent in the different-task condi-
errors were due to GK only detecting the right-side tion. There was a shift in the response strategy on
letter; on the remaining 5 trials he detected the different- compared with same-response trials.
right-side letter and assigned the wrong identity to So far we have presented evidence that two stim-
the left letter (3 on same-identity trials and 2 on dif- ulus factors are important for the anti-extinction
ferent-identity trials). On same-response trials GK effect with GK: stimulus exposure and common
made 20 errors by identifying only the right letter; onset of the items. There was no effect of response
on the remaining 10 trials he identified the right- linkage. The effects of common onsets may arise
side letter and misidentified the left letter. There because of at least three factors: (1) the onsets may
were nine such misidentifications on same-identity cue attention to a spatial area subtended by the let-
trials and one on different-identity trials. This sug- ters, enabling left- as well as right-side items to be
gests that GK was biased to guess that the identity reported (Yantis & Jonides, 1990); (2) the onsets
of the left letter was different to that of the right let- may increase arousal, and this enables both letters
ter, when he had to assign identities to both items to be reported more accurately than when only a
(on same-response trials). single-left stimulus appears (cf., Robertson &
Manley, 1999); or (3) letters than onset together are
Discussion bound by their common temporal signal (Singer &
In both Experiments 3a and 3b we observed an Gray, 1995), and this form of grouping enables the
anti-extinction effect; left items were identified left as well as the right letter to be identified. In
more accurately on two-item trials than on single- Experiment 5 we return to provide an explicit test
left trials. Also, in both experiments this effect was between these different accounts. In Experiment 4,
almost identical when the same (identification) task though we explore one factor that might underlie
was performed on left- and right-field items as the effects of stimulus exposure on GK’s perfor-
when different tasks were used (identify left and mance: Whether identification changes at longer
detect right). There was no evidence for a response exposure durations because GK then makes an eye
priming effect. In Experiment 3a it might be argued movement to the right item (on two-item trials). It
that GK adopted the same response strategy irre- is possible that eye movements are responsible for
spective of whether different tasks were formally this shift from anti-extinction to extinction. For
required to left- and right-side stimuli. However, in example, GK may show extinction at long stimulus
Experiment 3b we found contrasting effects of item durations because of a bias to make a right eye
similarity on same- and different-response trials, movement, which then reduces his identification of
with item similarity only affecting performance on left-side items when two letters are present. Short
same-response trials. This provides converging evi- exposures may reduce the likelihood that eye move-
dence that GK did adopt different response proto- ments are made, enabling an anti-extinction effect
cols under the two task conditions. The finding that to emerge. To test this in Experiment 4, GK
identification performance was better when the let- received blocks of trials with stimulus durations of
ters differed, relative to when they were the same, 180 and 450 ms, during which we recorded his eye
matches the pattern of data reported previously by movements.

370 COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4)


TRANSIENT BINDING BY TIME

EXPERIMENT 4: THE EFFECTS OF


EYE MOVEMENTS

Method
The method was the same as that employed in
Experiment 1, except that GK’s eye movements
were recorded whilst he undertook the task. Letter
identification performance was scored according to
whether a rightwards eye movement was made
whilst the stimulus was displayed.

Results and discussion


Data were summed across the two exposure dura-
tions, since they were similar in both cases. In Fig-
ure 5 we present the number of correct letter
identifications as a function of trials where either a
rightwards eye movement occurred (Figure 5a) or
there was no eye movement (Figure 5b). There was
a rightwards eye movement during the stimulus
presentation on 106/540 (20%) of the trials and no
eye movement on 409 trials (76%). There were just
25 trials where a left eye movement occurred (less Figure 5. (a) Percentage correct letter identifications made on
than 5% of the time). Trials with left eye move- trials when a rightwards eye movement occurred when letters were
ments were not analysed further. present. (b) Percentage correct letter identifications made on trials
There was a reliable anti-extinction effect both when no eye movements occurred whilst the letters were exposed.
on trials where no eye movement occurred and on
trials where a rightwards eye movement was initi- from anti-extinction to extinction, is not simply due
ated. For trials without an eye movement, c2 (1) = to a change in the probability of an eye movement
4.52 for one-left trials (6/85) vs. two-item correct occurring. Whatever the factor is that determines
trials (28/156). For trials with a rightwards eye the better report of the left letter on two-item trials,
movement, Fisher exact probability = .03 for one- it survives an eye movement being made to the
left (0/17) vs. two-item correct trials (11/49). On right.
blank trials there were 9% false alarm errors on trials
without an eye movement (73/80 correct) and 15%
errors (17/20 correct) when a rightwards eye move- EXPERIMENT 5: TEMPORAL
ment occurred. On one-right trials performance BINDING, ATTENTIONAL CUEING,
was slightly better when there was a rightwards eye OR AROUSAL
movement relative to when no eye movement took
place (12/20—60% vs. 41/80—51%). Experiments 1–4 reveal a reliable anti-extinction
The results were quite clear. The anti-extinction effect when left- and right-field stimuli onset
effect survives even when an eye movement is initi- together and are presented for a short duration. We
ated to the right during stimulus presentation. have discussed three accounts of the onset effect:
Hence the time course of performance, the change that it is due to (1) cueing attention to a common

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HUMPHREYS ET AL.

spatial region; (2) increased arousal; or (3) temporal last). In contrast, on trials with a single left onset
binding of the letters. These alternative accounts letter, the right bias in report should lead to the cen-
were tested in Experiment 5. tral letter being reported prior to, and more accu-
Each trial began with a display of three pre- rately than, the left letter.
masks, one at fixation and two flankers (Figure 6a). Experiment 5b used a procedure identical to that
The critical left and right target letters appeared as in Experiment 5a, except that the central letter
onsets in the gaps between the central and the outer always differed in colour from the more peripheral
two masks. At the same time as these onset letters letters (it was black and they were red), and GK was
were presented, a central letter was created by off- asked only to identify the central (black) letter. This
setting contours in the pre-mask. In Experiment study was run as a control to test if poor report of the
5a, GK was asked to report any letters he could, central letter on two-onset trials in Experiment 5a
irrespective of their colour, spatial location, or was due to lateral masking of the central letter by
whether they were created by an onset or an offset. the more peripheral onsets. Such masking should
In all the prior experiments, GK adopted a right- arise irrespective of whether the onset letters have
to-left report procedure reflecting his spatial bias in to be reported.
selection, and we expected the same here. With
these conditions, and this order of report, the
Experiment 5a: Report all the letters
grouping account makes a different prediction to
the attentional cueing and arousal proposals. The Method
cueing account holds that there is improved report The main details of the procedure are given in the
on two-letter trials because GK’s attention is cued General Method. There were four blocks of trials.
to the spatial region covering the two onsets. It fol- In two blocks, the onset letters were in red and the
lows that GK should show good report of the cen- pre-masks and offset letters were in black, to
tral letter on trials when he identifies a left-side emphasise grouping differences between the onset
letter created by an onset, since the central letter and offset items. In the other two blocks the onset
should be selected prior to the left one (as it falls fur- and offset letters were all black. In all blocks, GK
ther to the right, within the attended area of space). was simply asked to identify all the letters present.
This same prediction is made by the arousal
account. The two onsets, on two-item trials, should Results and discussion
increase GK’s arousal, and this should improve his Figure 6b presents the overall percentage correct
ability to report all the letters present. On this reports of the flanking onset letters. There was a
account, report of the central letter on trials with reliable anti-extinction effect: Identification was
two onsets should be better than on trials with one higher on two-item than on one-left trials, c2 (1) =
or no onsets. In contrast, according to the binding 4.10, p < .05. There were no false alarms on blank
proposal, the left and right letters should be trials. Performance did not differ whether the onset
grouped and selected together without the central letters were black (17/48 vs. 4/24, on two-item and
letter necessarily being selected too (since the cen- one-left trials) or red (16/48 and 4/24 on two-item
tral letter is created by an offset rather than an and one-left trials). Figure 6c gives the percentage
onset). Given GK’s right-side bias, then the group of reports of the central offset letter, as a function of
falling further into the right field should be selected the number of onset letters present. Performance
first—the onset group. If both group members are was better on one-left than on two-item trials, c2 (1)
identified before any other stimuli present, then it is = 23.41, p < .01. There was also a trend for worse
even possible that the left-side letter (grouped with report of the central letter on two-item trials than
the right letter) will be identified before the central on trials with only right letters (Figure 6c). Poor
letter. This should lead to trials on which the left report of the central letter on two-item trials could
(onset) letter is reported and the central (offset) let- be due to the right onset letter being identified
ter is not (e.g., because the central letter is reported first–although in this case we would not expect the

372 COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4)


TRANSIENT BINDING BY TIME

central letter should be good on trials on which the


left, onset letter is identified. This should hold for
one- and two-item trials alike. Contrary to this
account, identification of the central letter
remained better on one-item than on two-item
trials even when the left letter was identified (8/8 vs.
2/33 trials; Fisher exact probability, p = .000). On
the one-item trials when the left letter was identi-
fied, GK always reported the central letter first (8/
8). On the two-items trials when the left letter was
identified, he never identified the central letter
before the left letter (0/33). On two-item trials,
report of the central letter did not differ whether the
onset letters were red (1 correct) or black (1 correct),
with the left letter identified.
The data indicate that, on one-item trials, GK
identified the central letter on all occasions where
he able to name the left-side target. Also, on all of
these trials, GK identified the central letter before
the left letter was reported. In contrast, on two-item
trials, GK was poor at reporting the central offset
letter even when he identified the left onset letter
correctly. He then also always reported the left let-
ter before the central one. These data are consistent
with the binding account rather than the
attentional cueing and arousal accounts. According
to the binding account, the anti-extinction effect
occurs because the left and right letters are grouped
by common onset. GK is biased to select the
rightmost items for report and, in this case, there is
selection of the onset group prior to the central let-
ter since members of the onset group fall in his right
field. This leads to the left-side letter being selected
prior to the central letter, and to it being identified
Figure 6. (a) Example displays from Experiment 5, with “offset”
even when the central letter is not reported. In con-
as well as “onset” target letters. (b) Percentage correct identification trast, there was no evidence for attention spreading
of the onset letters. (c) Percentage correct identification of the across the spatial region covering the two onsets
central offset letter, as a function of the whether no, one-left, one- (due either to cueing visual attention or to increased
right, or two onset letters were presented. arousal). If attention simply spread across the
region, GK’s tendency to report from right-to-left
central item to suffer more on two-item than on would lead him to reporting the central letter before
one-right trials. More critical is to assess report of the left. On two-item trials where he identifies the
the central letter on two-item and one-left trials left letter, he should also typically identify the cen-
when the left onset letter was identified. The tral letter. He did not—even though this was the
“spreading attention” and arousal accounts main- pattern on single-left trials.
tain that the central letter should typically be identi- Other accounts of the data from Experiment 5a
fied prior to the left-side letter; hence report of the are not easy to sustain. One possibility is that GK

COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4) 373


HUMPHREYS ET AL.

opted to report the peripheral onsets before the


central offset letter, perhaps because of an
attentional bias to stimuli created by onsets rather
than offsets (e.g., Yantis, 1998). However, we
would expect this bias to hold across the one- and
two-item trials here, which occurred randomly.
This was not the case. GK did not identify a single
left onset letter before the central offset letter. A
more viable possibility is that the central letter was
difficult to report on two-onset trials because of
masking from the onsets1. A single left onset may Figure 7. The percentage of correct identifications of the central
not generate such a strong masking effect, enabling offset letter as a function of whether no, one-left, one-right, or two
the central letter then to be reported before the left onset letters occurred in Experiment 5b.
flanker. To test this, we ran Experiment 5b. This
study was the same as Experiment 5a, except that and there were reports of the right rather than the
GK was asked just to report the central letter. To central letter on 10/36 of the errors on trials with
help him select the central item, it always differed in two flankers. On all such occasions GK maintained
colour from the flanking onset letters. The masking that he reported the black letter. Prior studies have
effect should be caused by the presentation of the demonstrated that GK is susceptible to illusory
onsets and so it should occur even without report of conjunctions of letters and colour (Humphreys,
the flankers. On the other hand, if the relatively Cinel, Wolfe, Olson, & Klempen, 2000), and these
poor report of the central letter was due to GK incorrect letter reports may reflect illusory binding
selecting the two flankers for report, then this effect of the central colour with the identity of a flanker
should be lessened here. letter. More importantly for our present purposes,
the decreased report of the central item was less
apparent here than when flanker letters had to be
Experiment 5b: Reporting only the central
selected for report (Experiment 5a). The decreased
black letter
report on the central letter in the two-item condi-
Method tion of Experiment 5a does not seem to be due to
This was the same as for Experiment 5a, except that lateral masking.
the central letter was always black and the flankers
red.
EXPERIMENT 6: TEMPORAL ORDER
Results JUDGEMENTS AND SEQUENTIAL
Figure 7 shows the number of correct reports of the LETTER PRESENTATIONS
central black letter as a function of whether no
(blank), one-left, one-right, or two onsets occurred. Experiments 1–5 have demonstrated an anti-
Unlike Experiment 5a, there was only a small, non- extinction effect, where, under brief exposure con-
significant trend for identification of the central let- ditions, report of a left-side letter was facilitated by
ter to be worse on two onset trials relative to trials in simultaneous presentation of a right-side letter. We
which only a single left flanker was presented (c2 < have proposed that anti-extinction is caused by
1.0). On one-left trials GK made 1/14 error by temporal binding based on common onset of the
reporting the left letter rather than the right letter. letters. According to this account, the stimuli may
On one-right trials there were 7/19 errors due to need to occur simultaneously in order for the anti-
reporting the right letter rather than the left letter, extinction effect to occur. We examined this possi-

1
Our thanks to Rob Ward for this suggestion.

374 COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4)


TRANSIENT BINDING BY TIME

bility in Experiment 6. In so doing, we also evalu- third of the trials had simultaneous presentations,
ated the relations between the anti-extinction effect on another third the left letter preceded the right
and GK’s conscious judgements of when onsets one, and on the remaining third the right preceded
were simultaneous. To do this, we compared GK’s the left.
identification and temporal order judgements.
Rorden et al. (1997) demonstrated that patients
Results and discussion
with extinction frequently show biased temporal
order judgements so that, when ipsi- and The results for the temporal order judgements are
contralesional stimuli are presented simulta- presented in Figure 8a, and Figure 8b gives the per-
neously, they judge that the ipsilesional item pre- centage of correct colour identifications. For colour
cedes the contralesional one. In order for ipsi- and identification, there was an anti-extinction effect
contralesional events to be judged as simultaneous, when the stimuli appeared simultaneously (report
the contralesional event needs to lead the on two-item trials was better than on one-left trials;
ipsilesional one by some time period: the phenome- c2(1) = 6.71, p < .01. There was no reliable effect
non of “prior entry”. If GK manifested prior entry, when the onsets of the letters were staggered (c2 <
then we expected him to be poor at judging that 1.0 for left-first and for right-first). In the temporal
simultaneously occurring letters did onset at the order judgement task, though, GK showed a strong
same time, and that we would need to present the effect of “prior entry”, judging that the right letter
left letter first for judgements of simultaneity to be appeared before the left on the vast majority of tri-
made. In contrast, the report of briefly presented als. This tendency varied across the different tem-
left letters may be best under simultaneous presen- poral presentation conditions, but even when the
tations. There may be a dissociation between GK’s left letter led the right one by 720 ms, GK reported
conscious perception of simultaneity and the effects that the right came first on over half the trials.
of simultaneous onsets on letter identification. This Experiment 5 shows that the anti-extinction
would suggest that GK is not conscious of temporal effect occurred when right and left letters appeared
binding, even though it affects his conscious letter simultaneously, but not when they were staggered
report. in time. In contrast, under these same conditions
GK consistently judged that the right letter was
presented first and the left letter had to precede by
Method
at least 720 ms for the onsets to be judged as simul-
The procedure is outlined in the General Method. taneous. The effects of time on anti-extinction dis-
For the temporal order judgements, GK was pre- sociated from the effects on conscious temporal
sented with one red and one green letter on each order judgements.
trial, and he had to decide which colour appeared Across Experiments 1–5, when the onset letters
first. We required report of the temporal order of always had the same colour, the advantage varied
the colours, rather than of the left and right letters, from 12–22% for the identification of left letters on
in order to eliminate possible response biases two-item relative to one-left trials. In Experiment
favouring the right-side letter. The letters were pre- 6, when the onset letters differed in colour, the
sented simultaneously or they were staggered in advantage was 14.5%. The colour difference
time—the left preceding or following the right by between the letters had little influence on the effect.
450 ms or by 720 ms (presented in separate trial
blocks). To match the attributes being reported, the
identification task required GK to report the GENERAL DISCUSSION
colours present. The two-item trials were the same
as for the temporal order judgement task but, for We have demonstrated a reliable anti-extinction
the identification task, there were also single colour effect when left and right stimuli are presented
trials (one-left or one-right). Within a block one simultaneously for brief exposure durations in

COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4) 375


HUMPHREYS ET AL.

Figure 8. (a) Percentage of trials on which GK reported that the letter on the right appeared first, as a function of the stimulus onset
asynchrony (SOA) between the left and right letters. (b) Percentage of correct colour identifications across the different SOAs and on one-left
and one-right letter trials. + indicates that the left letter preceded the right letter; – indicates the opposite.

376 COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4)


TRANSIENT BINDING BY TIME

patient GK, who has a bias favouring right-side These results suggest that there can be grouping
stimuli after bilateral parietal damage. The effect is based on common onset between visual stimuli.
dependent on the stimuli having simultaneous This form of binding dissociates from GK’s con-
onsets (Experiments 2 and 6), and is unaffected by scious judgements about the temporal order of
eliminating the common offset of the stimuli occurrence of the stimuli (Experiment 6). Thus the
(Experiment 2b), by varying the tasks performed on temporal code that “binds” the left and right letters
left and right-side items (Experiment 3), and by a was not consciously available to GK. The findings
rightwards eye movement occurring whilst stimuli are consistent with the notion that that temporal
are exposed (Experiment 4). The anti-extinction codes are important for binding in vision (Eckhorn,
effect reduced as the time since stimulus onset 1999; Singer & Gray, 1995), and with one signal
increased, being replaced by left-side extinction at for binding being based on actual (rather than per-
longer stimulus durations (Experiments 1 and 2). It ceived) synchronous onsets (Elliott & Muller,
also dissociated from GK’s conscious judgements of 1998; Fahle, 1993). Once bound, both left and
temporal order; the anti-extinction effect occurred right stimuli could be selected for report even when
when letters appeared simultaneously whereas left an eye movement was first made to the item on the
letters had to precede right letters by 720 ms or right side.
more before GK tended to judge that the letters Our evidence also indicates that binding by com-
appeared simultaneously (Experiment 6). mon onset produces only a transient influence on
Experiment 5 provided evidence that the anti- visual selection. Thus the anti-extinction effect
extinction effect was due to temporal binding lasted for stimuli presented for up to 400–500 ms,
rather than to increased arousal or due to a spread before being replaced by extinction as the exposure
of spatial attention across a region activated by the duration of the stimuli increased. With longer expo-
left and right stimuli. In that study, we presented a sures single left letters could be identified, but there
central letter defined by an offset between the left was then biased selection favouring the right letter
and right letters, which were defined by onsets. on two-item trials. How can we explain this shift to
We found that, on two-item trials where GK an extinction effect at longer stimulus durations?
reported the left as well as the right letter, he was To account for the effects of stimulus duration,
poor at identifying the central letter. On one-left we suggest that there is a decay in the temporal tag
trials, however, he could report the central letter in that binds the right and left stimuli, even though
addition to the left one (indeed he identified the the letters may remain present in the field. This idea
central one first). This result was found both when is illustrated in Figure 9, in which we separate two
the onset and offset letters had the same colours forms of binding, a temporary effect based on co-
and when they had different colours, so that col- occurrence of a transient signal generated by the
our grouping did not appear important. We con- onsets of the stimuli (perhaps also reinforced by off-
clude that GK did not attend to the region sets, when offsets occur relatively close in time to
subtended by the left and right onset letters and the onsets), and a longer-lasting effect based on
report the letters there in right to left order. sustained (form and colour) information derived
Instead, the left and right letters were grouped by from stimuli. In other studies with the patient
common onset and reported together, prior to the tested here, GK, we have shown that the magnitude
central letter being selected. Consistent with this, of extinction (at longer stimulus durations) can be
there was not a reliable drop in performance on modulated by grouping based on the form and sur-
two-item relative to one-left trials when only the face properties of objects (Boutsen & Humphreys,
central letter had to be identified, when selection 2000; Gilchrist et al., 1996; Humphreys, 1998).
of the flankers was not required (Experiment 5b). The anti-extinction effect, though, was little
Identification of the central letter dropped partic- affected by whether onset letters had the same or
ularly when onset letters were selected for a different colours (Experiment 1–5 vs. Experiment
response. 6). This is consistent with different forms of bind-

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HUMPHREYS ET AL.

Figure 9. An illustration of two proposed neural signals for binding in vision: a transient signal that links stimuli having common onsets,
and a more sustained signal determined by the form and surface details of objects. Note that the strength of the transient signal decreases over
time even if the stimulus remains in the field.

ing between visual stimuli, operating across con- sustained signal), GK tends to select the item on the
trasting time courses. There is transient binding right before the one on the left, generating an
between stimuli that have synchronous onsets. This extinction effect. The exception to this is if the
may provide the visual system with a “quick but stimuli group by form or surface property, in which
dirty” representation of the world, useful perhaps case they may be selected together (Boutsen &
for fast actions. However, since stimuli that onset Humphreys, 2000; Gilchrist et al., 1996;
together may have different form and surface prop- Humphreys, 1998). Due to this spatial bias in using
erties, binding by common onset may be of little use sustained information from a stimulus, the anti-
for object recognition. In order to construct a more extinction effect is replaced over time by extinction.
accurate representation, the transient binding may If there is temporary binding based on common
be replaced over time by a more sustained binding onsets of stimuli, we might ask why anti-extinction
process, sensitive to form and surface-based simi- effects have not been found more frequently in
larities between stimuli. patients. There may be several reasons for this. One
To account for the extinction effect at longer is that, in patients showing poor report of single
duration, we propose that GK has a spatial bias in contralesional items, experimenters may not exam-
using the sustained information for selection. Thus ine performance with double simultaneous stimu-
if two stimuli are presented bilaterally for relatively lation due to an assumption that two-item trials
long durations (long enough to establish the would in any case be more difficult than single-item

378 COGNITIVE NEUROPSYCHOLOGY , 2002, 19 (4)


TRANSIENT BINDING BY TIME

trials. As we have demonstrated, this assumption is contralesional item having to precede the
not necessarily valid. A second reason relates to the ipsilesional one in order for them to be judged as
magnitude of the effects. The differences between occurring simultaneously. It may be that any com-
two-item and one-left trials reported here (and also petition for selection between developing sustained
by Goodrich & Ward, 1997) are quite subtle, with a codes for ipsi- and contralesional stimuli is maxi-
benefit of about 15% on two-item trials. It is possi- mised when the codes begin to develop at the same
ble that such a small difference may not be detected time (with simultaneous stimulus exposures), or it
when small numbers of trials are collected. A third may be that this competition is pronounced when
possibility is that other patients may have more there is an initial temporal tag to bind the stimuli.
rapid development of sustained information from Whichever is the case, their result suggests again
the stimulus than is the case for GK, and they too that the effect of time on perceptual report is inde-
may have a spatial bias in using the sustained infor- pendent of conscious awareness of the temporal
mation. Consequently, there may not be a sufficient relations between events.
time window for anti-extinction to be demon-
strated before extinction effects emerge. A fourth Manuscript received 5 June 2001
suggestion is that such effects depend on Revised manuscript received 7 November 2001
subcortical structures, sensitive to dynamic changes Revised manuscript accepted 21 November 2001
in visual stimulation. Effects at this level may only
become apparent in patients like GK, with rela-
tively large lesions, who are impaired at responding
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