Vision Research 41 (2001) 801– 814

www.elsevier.com/locate/visres

Modulation of spatial attention with unidirectional field motion:

an implication for the shift of the OKN beating field
Katsumi Watanabe *
Computation and Neural Systems, California Institute of Technology, Mail Code 139 -74, Pasadena, CA 91125, USA

Received 15 November 1999; received in revised form 10 August 2000

Abstract

During optokinetic nystagmus (OKN) the mean eye position of gaze (the beating field) shifts in the direction of the fast phases.
The function of this shift may be to re-orient the eyes in the direction of self-motion which optic flow implies (in-coming field).
This idea leads to the hypothesis that visual attention may be directed toward the In-coming field. In Experiment 1, subjects
detected a visual flash presented against unidirectional field motion. The OKN beating field was shifted toward the In-coming
field, and manual reaction times were shorter when the target appeared in the In-coming field. Experiment 2 revealed that this
In-coming field advantage occurred even when OKN (and thus the mean eye-position shift) was suppressed. Subsequent
experiments showed that the In-coming field advantage is not due to a local motion interaction (Experiment 3), survives subject’s
voluntary allocation of attention (Experiment 4), and develops over less than 320 ms after the onset of the motion field
(Experiment 5). These results suggest that unidirectional field motion tends to automatically shift visual attention toward the
In-coming field. © 2001 Elsevier Science Ltd. All rights reserved.

Keywords: Optokinetic nystagmus; Beating field; Attention; Reaction time; Motion field

1. Introduction 1981), in cats (Schweigart & Hoffmann, 1988; Schwei-

A large moving visual field evokes a series of rhyth-
mic reflexive eye movements called optokinetic nystag-
mus (OKN) (Cohen, Matsuo, & Raphan, 1977; van Die
& Collewijn, 1982; Carpenter, 1988; Collewijn, 1991;
Leigh & Zee, 1991; Howard, 1993). Slow phases of
OKN smoothly track the movement of the visual field,
and fast phases (saccades) in the opposite direction
interrupt the slow phases. The purpose of OKN is to
stabilize the retinal image to maintain the high visual
acuity.
Curiously, the average eye position of gaze during
OKN (the beating field) shifts in the direction of the
fast phases, namely, in the direction opposite to the
visual field motion. This is true in man (Jung & Mitter-
maier, 1939; Miyoshi, Shirato, & Hiwatashi, 1978;
Dubois & Collewijn, 1979; Abadi, Howard, & Ohmi,
1999), in monkeys (Kubo, Igarashi, Jensen, & Hormick,
* Tel.: +1-626-395-2362; fax: + 1-626-844-4514.
E-mail address: kw@caltech.edu (K. Watanabe).
gart, 1995), and in rats (Meier & Dieringer, 1993;
Bähring, Meier, & Dieringer, 1994). A similar shift of
the ocular beating field is also observed with vestibular
nystagmus. The mean eye position of vestibular nystag-
mus shifts in the direction of the head rotation (Melvill-
Jones, 1964; Chun & Robinson, 1978; Roucoux,
Crommelinck, Guerit, & Meulders, 1981; Crommelinck,
Roucoux, & Veraart, 1982; Vidal, Berthoz, & Milan-
voye, 1982; Siegler, Israel, & Berthoz, 1998).
Based on these observations of optokinetic and
vestibular nystagmus, some researchers interpreted the
shift of the beating field of nystagmus as a goal-directed
involuntary response, or a reflexive orienting response
toward a ‘center of interest’ (Melvill-Jones, 1964; Chun
& Robinson, 1978; Crommelinck et al., 1982; Vidal et
al., 1982; Meier & Dieringer, 1993; Bähring et al., 1994;
Schweigart, 1995; Siegler et al., 1998). The basic idea is
that the fast phases of involuntary nystagmus may
strategically re-orient the eyes in the direction of self
motion so that the visual system can detect a target
more efficiently in the visual field toward which the
head and/or body move (in-coming field).

0042-6989/01/$ - see front matter © 2001 Elsevier Science Ltd. All rights reserved.
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802 K. Watanabe / Vision Research 41 (2001) 801–814
It has been suggested that OKN is mediated in
part by cortical mechanisms (Miles, Kawano, & Opti-
can, 1986; Howard & Simpson, 1989; Boussaoud,
Ungerleider, & Desimone, 1992; Busettini, Masson, &
Miles, 1996; Mestre & Masson, 1997; Watanabe,
1998). Critically for the present study, the shift of the
beating field is under the control of the frontal eye
fields (FEF) in primates (Kubo et al., 1981; Ouchi,
Igarashi, & Kubo, 1981; Fuchs & Mustari, 1993) and
by the FEF homologous structure in rats (dorsome-
dial shoulder of the prefrontal cortex; Meier and
Dieringer (1993); Bähring et al. (1994)). Since the
functional role of the FEF involves selective attention
(Robinson & Fuchs, 1968; Ouchi et al. 1981; Welch
& Stuteville, 1985; Liechnetz & Goldberg, 1988), the
shift of the beating field may be understood as a shift
of spatial attention (Meier & Dieringer; Bähring et
al.).
The purpose of the present study is to directly ex-
amine the attention hypothesis about the shift of the
OKN beating field in humans and to explore the na-
ture of such attentional modulation. In Experiment 1,
it was observed that with the shift of the OKN beat-
ing field, manual reaction times for detecting a small
visual target were shorter when the target occurred in
the In-coming field. Then, Experiment 2 revealed that
this In-coming field advantage happened even when a
stationary fixation stimulus was provided to suppress
OKN, and hence there was no shift of eye position.
Further experiments showed that the In-coming field
advantage is not due to a local motion interaction
(Experiment 3), survives subject’s voluntary allocation
of attention (Experiment 4), and develops over a
Fig. 1. Schematic of stimulus configuration: While the random-dot
background moved leftward or rightward, a bar (Experiments 2 – 5)
or a small square (Experiment 1, not shown in the figure above) was
presented for about 13 ms at either the left or the right of the fixation
stimulus (or, of the display center in Experiment 1). In Experiments
2–5, color and shape of the target were varied randomly. Note that
the fixation stimulus was not presented in Experiment 1.
rather short period after the onset of the background
motion (Experiment 5). These results are interpreted
in terms of the automatic effect of unidirectional field
motion on the distribution of spatial attention and its
relation to the shift of the OKN beating field.
2. Experiment 1: manual reaction time with the OKN
beating-field shift
Experiment 1 was conducted to examine the effect
of the shift of the OKN beating field on manual
reaction time. Spatial selective attention at the loca-
tion of a target reduces manual reaction time for the
target (Posner, 1978, 1980; Posner & Cohen, 1984).
Therefore, if the shift of the OKN beating field is
accompanied by a shift of spatial attention, the reac-
tion time would be shorter when a visual target was
in the In-coming field (e.g. a right target with left-
ward field motion).
2.1. Method
2.1.1. Subjects
Three male subjects (S1 the author, S2, and S3; age
range from 24 to 28 yr) voluntarily participated in
the experiment. All subjects had normal or corrected-
to-normal vision, reported no oculomotor and
vestibular dysfunction, and had previously partici-
pated in other OKN experiments. Except for the au-
thor, the subjects were naı̈ve with regard to the
purpose of the present study. Informed consent was
obtained from each subject before the experiment.
2.1.2. Stimuli
Stimuli were displayed on a CRT color monitor
(frame rate 75 Hz), controlled by a Silicon Graphics
Iris workstation, in an otherwise dark environment
(B0.01 cd/m2). The stimuli were composed of a
background and a target (Fig. 1). The background
was a random-dot pattern moving either leftward or
rightward. The speed of the background moving pat-
terns was 12.9 deg/s. The size of the patterns was
30× 30 deg2 in visual angle (consisting of 800 dots;
dot density 0.89 dot/deg2; individual dot size 0.03
deg). Luminance of the dots was 30 cd/m2 and back-
ground luminance was 0.02 cd/m2. As imaginary
targets cannot suppress OKN (Howard, Giaschi, &
Murasugi, 1989), the display employed in Experiment
1 reliably elicited OKN in all subjects. The target was
a small white square (0.5 deg; 30 cd/m2). It was pre-
sented for one frame (about 13.3 ms) at either 4 deg
to the left or 4 deg to the right of the center of the
display. The delay of the target presentation from the
onset of the background motion was randomized
from 3 to 5 s.
 K. Watanabe / Vision Research 41 (2001) 801–814
Fig. 2. (a) Representative trace of eye position (S3) in Experiment 1.
(b) Histograms of eye position for the leftward background motion
(top panel) and for the rightward background motion (bottom panel).
The horizontal axis indicates eye position in visual angle. Negative
values mean that the eyes were deviated toward left from the primary
position. The vertical axis shows the relative frequency of eye posi-
tion. The eye position (the OKN beating field) was clearly shifted in
the direction opposite to motion, namely, in the In-coming field.
Fig. 3. The effect of the background motion on manual reaction time
for detecting the target in Experiment 1 (in-coming condition versus
Out-going condition). The vertical axis shows the mean reaction time.
Each symbol represents data from an individual subject. T bars
indicate 1 SE.
803
2.1.3. Procedure
The subject observed the stimulus display binocularly
from a distance of 80 cm, with the head stabilized with
a chin and chest support. While trying to keep the eyes
stationary relative to the head and to the stimulus
display, and not to track any particular detail of the
background pattern (so-called ‘stare’ OKN; Honrubia
et al. (1968)), the subject detected the target as soon as
possible by pressing a mouse button. After the subject’s
responses, the background motion field was turned off.
For each combination of the background motion direc-
tion and the target location, 36 trials were repeated
randomly [background motion direction (2)× target lo-
cation (2)× repeat (36)= 144 trials]. The stimulus con-
ditions were classified into two categories: In-coming or
Out-going conditions. If the target was presented in the
visual field from which the background motion came, it
was called the In-coming condition. Otherwise, it was
defined as the Out-going condition. Each condition
contained 72 trials. A full experiment was conducted in
one session. For each subject, reaction times faster or
slower than the mean reaction time by two standard
deviations were excluded.
2.1.4. Eye mo6ement recording
Horizontal positions of the subject’s right eye were
recorded with an infrared eye-tracker (Permobil,
Ober2 ). Eye position data were sampled at a 100 Hz
rate. Data from the onset of the background motion to
the onset of the target for each trial were stored for
off-line analysis. At the beginning of a session the
eye-tracker was calibrated with 10-deg horizontal
saccades.
2.2. Results
Fig. 2 shows a representative trace of eye position
(S3) and histograms of eye positions for all subjects.
The trace of eye position indicates the clear occurrence
of OKN. The histograms confirm that the mean posi-
tions of gaze tended to be shifted in the direction
opposite to the background motion (i.e. toward the
In-coming field). The reaction time results of Experi-
ment 1 are shown in Fig. 3. All three subjects re-
sponded more quickly when the visual target appeared
in the In-coming field than when it appeared in the
Out-going field (two-tailed t-test within each subject,
PB 0.05).
2.3. Discussion
The OKN beating field was shifted toward the In-
coming field (Fig. 2), which is consistent with other
OKN studies (Jung & Mittermaier, 1939; Miyoshi et
al., 1978; Dubois & Collewijn, 1979; Abadi et al., 1999).
The new finding in Experiment 1 was that the manual
804 K. Watanabe / Vision Research 41 (2001) 801–814
reaction times were shorter when the visual target ap-
peared in the In-coming field (In-coming field advan-
tage). This result partially supports the attention
hypothesis for the shift of the OKN beating field be-
cause it implies a link between spatial visual attention
and the mean eye-position during OKN. However, the
results of Experiment 1 are equivocal with respect to
whether the shift of spatial attention is the cause or
result of the beating-field shift. Experiment 2 was
aimed to tease apart these two alternatives.
3. Experiment 2: In-coming field advantage without the
OKN beating-field shift
Experiment 1 demonstrated a positive correlation
between the shortening of reaction time and the shift
of the OKN beating field. The strong version of the
attention hypothesis proposes that the optokinetic
stimulation alters the spatial distribution of attention
so that attention is directed toward the In-coming
field. This shift of attention may then result in the
shift of the mean eye position of OKN in the direction
opposite to the optokinetic stimulus motion. If this is
the case, even when OKN is suppressed with a fixation
Fig. 4. (a) Representative trace of eye position (S3) in Experiment 2.
The large spike-like amplitudes were eye-blink artifacts, which were
excluded for the eye-movement analysis. (b) Histograms of eye posi-
tion. There was no significant shift of eye position, showing that the
fixation stimulus and instruction in Experiment 2 were effective to
suppress OKN.
stimulus, the results would be similar to those without
fixation. Additionally, Experiment 2 involved a manip-
ulation of task requirement. Four different tasks (de-
tection, localization, color discrimination, and shape
discrimination) were employed.
3.1. Methods
3.1.1. Subjects
The same three subjects as in Experiment 1 took
part in the experiment.
3.1.2. Stimuli
The stimuli were identical to those used in Experi-
ment 1 except for the following. A yellow fixation
cross (1.5 deg; 18.7 cd/m2) and a black disc (3.0 deg;
0.02 cd/m2) were presented at the center of the display
throughout a session, both of which occluded the
background motion (Fig. 1). A target was either a
horizontal or vertical bar, the color of which was
either red or green, and the location of which was at
either the left or right of the fixation stimulus. The
luminance of the target was 19 cd/m2 for both colors.
The length and the width of the target were 1.2 and
0.5 deg, respectively.
3.1.3. Procedure
Each subject was instructed to firmly stare at the
fixation cross throughout each trial. The subject’s task
was to report either (1) the appearance of a target by
pressing a mouse button as quickly as possible (detec-
tion), (2) the target location (left or right) by pressing
one of the two mouse buttons as quickly and accu-
rately as possible (localization), (3) the target color
(red or green) by button pressing (color discrimina-
tion), or (4) the target shape (horizontal or vertical) by
button pressing (shape discrimination). For each com-
bination of the background motion direction and the
target type, nine trials were repeated randomly [back-
ground motion direction (2)× target location (2)×
target color (2)× target shape (2)× repeat (9)=144
trials]. The subject performed different tasks in sepa-
rate sessions. Trials were classified as in Experiment 1
(In-coming versus Out-going), resulting in 72 trials for
each condition. Reaction times faster or slower than
the mean reaction time by two standard deviations,
and trials with erroneous responses, were excluded
from analysis. The other procedures were the same as
those of Experiment 1.
3.2. Results
The direction of the background motion did not
bias the eye position of gaze (Fig. 4), confirming that
the fixation stimulus and instruction successfully sup-
pressed OKN in Experiment 2 (OKN suppression:
Murphy, Kowler, and Steinman (1975), Barnes and
 K. Watanabe / Vision Research 41 (2001) 801–814 805

Fig. 5. The effect of the background motion on manual reaction time for detecting (left top), localizing (right top), discriminating color (left
bottom), and discriminating shape of (right bottom) the target in Experiment 2. The vertical axis shows the mean reaction time. Each symbol
represents data from an individual subject. T bars indicate 1 SE.

Crombie (1985), Murasugi, Howard, and Ohmi (1986),
Pola and Wyatt (1993)). Nonetheless, the In-coming
field advantage was still observed with the detection
and the localization tasks (Fig. 5; two-tailed t-test
within each subject, P B 0.05). However, the reaction
times for the color discrimination and the shape dis-
crimination did not show a significant In-coming field
advantage (Fig. 5; two-tailed t-test within each subject,
P \0.05).
3.3. Discussion
When the subject’s task was the detection or the
localization of the target, the unidirectional motion
field affected the spatial distribution of attention (as
measured with manual reaction time), suggesting that
attention was directed toward the visual field from
which the motion came (In-coming field advantage).
The effect size in the detection task of Experiment 2
was almost the same as that in Experiment 1, which is
also similar to the effect size observed in many studies
of the attentional effect on manual reaction times
(Jonides, 1981). Hence, the shift of the OKN beating
field is not necessary for the In-coming field advantage
to occur. This result strengthens the attention hypothe-
sis that visual attention is modulated by unidirectional
field motion.
Interestingly, in Experiment 2, the In-coming field
advantage was found only with the orienting tasks
(detection and localization), but not with the identifica-
tion tasks (color- and shape-discrimination). If subjects
deliberately shifted visual attention, all the tasks should
have exhibited the In-coming field advantage. Since this
was not the case, it is inferred that the attentional shift
toward the In-coming field may be caused mainly by an
automatic mechanism for orienting responses.
Researchers have suggested that the characteristics of
attention may differ for different task requirements,
especially between orienting tasks (detection and local-
ization) and identification tasks (e.g., color discrimina-
tion and shape discrimination) (Nakayama &
Mackeben, 1989; Tanaka & Shimojo, 1996). Crudely
806 K. Watanabe / Vision Research 41 (2001) 801–814
speaking, identification may depend heavily on en-
dogenous (voluntary, top-down) components of atten-
tion, whereas orientation may depend more on
exogenous (automatic, bottom-up) components of at-
tention. Much evidence now suggests that cortical
mechanisms for localization (or orientation) and iden-
tification of ob-jects are distinct, and so are atten-
tional mechanisms. Starting from occipital cortex, the
ventral pathway to temporal cortex mediates identifi-
cation (‘what an object is’: Haxby et al. (1991),
Grady et al. (1992)), whereas the dorsal pathway to
the parietal region processes orientation and spatial
attention (‘where an object is’ and ‘where to look’:
Haxby et al. (1991), Grady et al. (1992), Corbetta,
Shulman, Miezin, and Peterses (1995)). The results of
Experiment 2 may be related to the exogenous-dorsal-
orientation versus endogenous-ventral-identification
distinction.
Overall, the results of Experiment 2 are consistent
with the attention hypothesis about the shift of the
OKN beating field. But, there is a potential artifact
due to local motion interaction. With unidirectional
background motion, there is the inherent asymmetry
of the direction of the background motion. For ex-
ample, with rightward background motion, the pat-
tern in the left visual field moves from peripheral to
Fig. 6. Schematic of stimulus configuration of Experiment 3. The
random-dot background moved inward or outward. The other stimu-
lus configurations were identical to those of Experiment 2.
foveal region, whereas the pattern in the right visual
field moves from foveal to peripheral region. In other
words, the results of Experiment 2 could be inter-
preted as indicating that the reaction time to detect
or localize the visual target might be faster in the
peripheral-fovea background motion than in the
fovea-peripheral background motion.
4. Experiment 3: rejecting an artifact due to local
motion interaction
It is known that visual latencies for detecting a
small moving target are shorter for motion towards
the fovea than for motion away (Mateeff & Hohns-
bein, 1988; Mateeff et al., 1991). If the briefly pre-
sented target was perceived to move in the same
direction as the background motion (motion capture;
Ramachandran and Inada (1985), Ramachandran and
Cavanagh (1987), Yo and Wilson (1992), Murakami
and Shimojo (1993)), the results of Experiment 2
might be simply due to some interactions of local
motion processes. Experiment 3 was designed to ex-
amine whether motion interactions could be responsi-
ble for the difference in manual reaction time. For
this purpose, the stimulus display was configured to
produce the same level of motion signal and interac-
tion as the display of Experiment 2, but not to pro-
duce any consistent OKN signal.
4.1. Method
4.1.1. Subjects
The same three subjects as in the previous experi-
ments took part in Experiment 3.
4.1.2. Stimuli
There were two conditions of the background mo-
tion in Experiment 3 (Fig. 6). In half of the trials, the
random-dot pattern in the left visual field moved
rightward and that in the right visual field moved
leftward (Inward condition). In the other half of the
trials, the random-dot pattern in the left visual field
moved leftward and that in the right visual field
moved rightward (Outward condition). Note that the
local stimulus relationship between the target and the
background was identical between the Inward condi-
tion and the In-coming condition in Experiment 2,
and between the outward condition and the Out-go-
ing condition in Experiment 2. The other stimulus
parameters were the same as those of Experiment 2.
4.1.3. Procedure
The procedures were identical to those of Experi-
ment 2.
 K. Watanabe / Vision Research 41 (2001) 801–814
Fig. 7. (a) Representative trace of eye position (S3) in Experiment 3.
The large spike-like amplitudes were eye-blink artifacts, which were
excluded for the eye-movement analysis. (b) Histograms of eye posi-
tion. With both the inward and the outward background motions, no
significant deviation of eye position from the primary position was
observed.
4.2. Results
The eye movement recording did not show any
systematic deviation of gaze while the subjects
performed the task (Fig. 7). The results of Experiment
3 are shown in Fig. 8. For all the tasks, the reaction
times did not differ significantly between the Inward
and the Outward conditions (two-tailed t-test within
each subject, P\ 0.1).
4.3. Discussion
The local motion fields in which the target was
presented were identical between the In-coming condi-
tion in Experiment 2 and the Inward condition in
Experiment 3, and that between the Out-going condi-
tion in Experiment 2 and the Outward condition in
Experiment 3. Yet, Experiment 3 did not produce con-
sistent differences in the manual reaction time. These
results clearly eliminate the possibility that the In-com-
ing field advantage observed in Experiment 2 was due
to a local interaction between the flashed target and the
background motion. Together, the results of Experi-
ment 2 and 3 suggest that the translation motion field
807
modulates spatial attention, as manifested in the differ-
ence in manual reaction time in the detection and
localization tasks (i.e. in orienting tasks).
5. Experiment 4: automaticity of the In-coming field
advantage
So far, the automatic nature of the In-coming field
advantage has been speculative from the fact that only
the orienting tasks led to the significant effect. Experi-
ment 4 was conducted to obtain a direct evidence for
the involuntary shift of visual attention in the In-com-
ing field advantage. With the combination of the In-
coming versus Out-going field paradigm (Experiment
2), a directional cue was provided in Experiment 4 so
that subjects would have knowledge regarding the
probable location of the target and could allocate atten-
tion endogenously (Posner, 1980; Jonides, 1981; Posner
& Cohen, 1984). If the In-coming field advantage is
caused by an automatic mechanism, it should be ob-
served even with an endogenous shift of attention.
5.1. Method
5.1.1. Subjects
The same three subjects took part in the experiment.
5.1.2. Stimuli
The stimuli were almost the same as those used in
Experiment 2. However, a directional cue was added
for manipulating the subject’s endogenous attention.
One side of the horizontal bar of the fixation cross was
elongated by 0.75 deg for 1 s. The onset of the cue was
randomized from 1.5 to 2.5 s before the target presenta-
tion. In 80% of trials, the elongated side of the fixation
cross and the target location were in the same side
(valid cue). In the remaining 20% of trials, the cue was
in the opposite side of the target (invalid cue).
5.1.3. Procedure
The target location, color, and shape were deter-
mined randomly. There were four types of conditions:
(1) In-coming condition with a valid cue, (2) In-coming
condition with an invalid cue, (3) Out-going condition
with a valid cue, and (4) Out-going condition with an
invalid cue. Each condition with a valid cue consisted
of 192 trials, while each condition with an invalid cue
consisted of 48 trials. Therefore, the total number of
trials was 480 trials (divided into two sessions). Since
the In-coming field advantage was expected only for the
detection and the localization tasks, the color discrimi-
nation and the shape discrimination tasks were not
involved in Experiment 4. The detection task and the
localization task were performed in separate sessions.
The subject was informed that 80% of the directional
808 K. Watanabe / Vision Research 41 (2001) 801–814

Fig. 8. The effect of the background motion on manual reaction time for detecting (left top), localizing (right top), discriminating color (left
bottom), and discriminating shape of (right bottom) the target in Experiment 3. The vertical axis shows the mean reaction time. Each symbol
represents data from an individual subject. T bars indicate 1 SE.

cues were valid. The other experimental procedures
were identical to those of Experiment 2.
5.2. Results
Fig. 9 shows the results of Experiment 4. There was
a clear effect of cueing: the valid cue shortened manual
reaction times in all subjects under most of the condi-
tions (one-tailed t-test, P B0.05; except for S3’s local-
ization performance under the Out-going condition,
where P = 0.21). The In-coming field advantage sur-
vived even with the cueing effect (one-tailed t-test,
P B 0.05, except for S2’s detection and localization
performance with invalid cues, where P = 0.42 and
P = 0.21, respectively).
(Posner, 1980; Jonides, 1981; Posner & Cohen, 1984;
Müller & Rabbit, 1989; Nakayama & Mackeben, 1989;
Remington, Johnston, & Yantis, 1992; Shimojo,
Tanaka, & Watanabe, 1996; Yantis, 1996). This is in
accordance with the claim that exogenous and endoge-
nous attention show several different characteristics
and may be mediated by separate neural mechanisms
(Butter, 1987; Nakayama & Mackeben; Klein, King-
stone, & Pontefract, 1992; Klein, 1994; Robinson &
Kertzman, 1995; Riggio & Kirsner, 1997; Briand, 1998;
Coull, Frith, Büchel, & Nobre, 2000).
6. Experiment 5: time-course of the In-coming field
advantage

5.3. Discussion In the previous experiments, subjects were exposed to

The results of Experiment 4 suggest that the In-com-
ing field advantage involves processes separate from
endogenous (top-down) attention, and may be due to a
process classified as exogenous (bottom-up) attention
the background motion for 3–5 s prior to the presenta-
tion of the target. The results have suggested that the
translation motion field automatically shifts visual at-
tention toward the In-coming field for the detection and
the localization tasks. Obviously, when the target is
 K. Watanabe / Vision Research 41 (2001) 801–814
presented at the moment the background motion
appears, there will be no difference in manual reaction
time between the In-coming and the Out-going
conditions. This raised an interesting question: How
does the In-coming field advantage develop over time?
Experiment 5 addressed this question in order to
further characterize the In-coming field advantage and
to have an idea about possible underlying mechanisms.
6.1. Method
6.1.1. Subjects
The author (S1) and one naı̈ve subject (S4) partici-
pated. Both had normal vision, and reported no oculo-
motor and vestibular dysfunction.
6.1.2. Stimuli
The stimuli were almost identical to those used in
Experiment 2, except that the target appeared at various
Fig. 9. The effect of the background motion on manual reaction time
for the detection task (left column) and for the localization task (right
column) in Experiment 4. The vertical axis shows the mean reaction
time. Each symbol represents data from an individual subject. T bars
indicate 1 SE. Even with the presence of the cueing effect (endoge-
nous shift of attention), the In-coming field advantage [out-going
(white square) − In-coming (black square)] tended to be preserved.
809
timings after the onset of the background motion (0, 40,
80, 160, 320 and 1280 ms after the onset).
6.1.3. Procedure
The target location, color and shape were determined
at random. The subject’s task was to localize the target
as quickly and accurately as possible. For each combi-
nation of the background motion condition (In-coming
or Out-going) and the target presentation timing, 40
trials were presented in random order [background
motion condition (2)× target timing (6)× repeat (40)=
480 trials]. A full experiment was divided into four
sessions. As a control experiment, the identical proce-
dure was repeated by using the inward –outward back-
ground motion display used in Experiment 3.
6.2. Results
The results of Experiment 5 are presented in Fig. 10.
When the unidirectional motion field was used, the
In-coming field advantage was observed in both sub-
jects. However, the reaction time difference between the
In-coming condition and the Out-going condition did
not show significance until some time after the onset of
the background motion (one-tailed t-test; PB0.01, af-
ter 160 ms for S1; after 320 ms for S4). There was no
significant difference between the Inward versus the
Outward conditions, irrespective of the target timing
(one-tailed t-test; P\ 0.1). In general, the reaction time
became shorter as the duration of the background
increased (one-way ANOVA, F\ 3, PB 0.001 for both
subjects and for all conditions), probably reflecting a
standard foreperiod effect (Mowrer, 1940; Bertelson,
1967).
6.3. Discussion
The In-coming field advantage did develop over time.
It took about 160 –320 ms for the effect to reach the
significant level. These rather short latencies suggest
that visually-induced self-motion perception (vection)
may not be responsible for the In-coming field
advantage.
Although the magnitude of vection was not mea-
sured, it was quite unlikely that vection occurred under
the stimulus conditions used in all the present experi-
ments. The stimulus visual field was restricted to 30-deg
of the central visual field in all experiments. The dura-
tion of the background motion was 5 s at most (Exper-
iment 1–4). The significant In-coming field advantage
was achieved with the background motion duration less
than 320 ms (Experiment 5). These values are known to
be inefficient to induce a strong sensation of self-motion
(Brandt, Dichgans, & Koening, 1973; Berthoz, Pavard,
& Young, 1975; Johansson, 1977; Dichgans & Brandt,
1978; Wong & Frost, 1987; Telford & Frost, 1993;
Kennedy, Hettinger, Harm, Ordy, & Dunlap, 1996).
810 K. Watanabe / Vision Research 41 (2001) 801–814

Fig. 10. The effect of the duration of the background motion on the In-coming field advantage (Experiment 5). The top panels show the results
with the incoming– outgoing unidirectional motion. The bottom panels show the results with the inward– outward motion. The vertical axis shows
the mean reaction time. T bars indicate 1 SE. The In-coming field advantage is conspicuous with the unidirectional motion field and develops over
time.

Moreover, when asked after the experiments, no subject 7. General discussion

reported a compelling sensation of self-motion. There-
fore, the attentional shift underlying the In-coming field
advantage may occur automatically before conscious
experience of self-motion. This parallels the suggestions
that the threshold for the perception of self-motion may
be higher than the threshold for automatic orienting
response such as postural reflex (body sway) induced by
a visual field motion (Stoffregen, 1985), and that the
latency for self-motion experience may be also larger
than the latency for postural reflex (Previc & Mullin,
1991). The reflexive postural adjustment and the atten-
tional shift observed in the present study may share the
same mechanism for motion analysis1.
1
Thilo, Guerraz, Bronstein, and Gresty (2000) have recently shown
that self-motion perception due to a prolonged observation of an
optokinetic motion field enhances the shift of the OKN beating field.
Their effect may be explained by another component besides the
automatic shift of visual attention, presumably an additional compo-
nent of endogenous attention.
The initial motivation of the present study was to
examine the relationship between the shift of the OKN
beating field and spatial visual attention. The results
were consistent with the idea that the subject’s attention
is shifted along with the shift of the OKN beating field
(Experiment 1). Moreover, the present study directly
demonstrated the effect of unidirectional field motion
on the distribution of spatial attention. Spatial atten-
tion, measured as manual reaction times, tends to be
shifted toward the In-coming field (i.e. toward the
direction from which the motion originates) without the
large shift of the mean position of gaze (Experiment 2
and Experiment 3).
Optic flow contains rich information about the move-
ment of the observer and the three-dimensional envi-
ronmental structure (Gibson, 1950, 1954; Gibson,
Olum, & Rosenblatt, 1955). Unidirectional field mo-
tion, as one type of optic flow, implies that the observ-
er’s head and/or body translates in the direction
 K. Watanabe / Vision Research 41 (2001) 801–814
opposite to the field motion. The system of spatial
attention may utilize such information to enhance ori-
enting responses for stimuli that appear suddenly in
the visual field toward which the head and/or body
move.
The attention hypothesis might be related to other
previous findings which concerned other types of optic
flow. During simulated forward motion with a radial
optic flow stimulus, the distribution of gaze tends to be
clustered near the focus of expansion (Lappe, Pekel, &
Hoffmann, 1998; Niemann, Lappe, Büscher, & Hoff-
mann, 1999). Since the heading judgment becomes
poor with increasing angle between gaze and the focus
of radial flow (Warren & Kurtz, 1992), this gaze strat-
egy may also reflect an involuntary attentional shift,
which would reduce the error in the heading judgment.
Similarly, during driving, driver’s gaze tends to be
directed at informative places in scene (e.g. ‘tangent
point’ on the inside of a road curve; Land (1992),
Land & Lee (1994)). These strategic shifts of eye
movements seem to occur automatically and might
reflect the utilization of global motion signal by the
attention system. Thus the modulation of attention by
a motion field may not be restricted to translation
motion, but could be generalized to various other
types of optic flow patterns. However, further investi-
gation is required for the issue of generality.
At this point, it is unavoidable that any explanation
about the underlying mechanism of the In-coming field
advantage is speculative. However, there are two phe-
nomena, which seem to be related to the In-coming
field advantage observed in the present experiments.
One is the directional asymmetry of visual attention
during smooth pursuit eye movements (Tanaka,
Yoshida, & Fukushima, 1998; van Donkelaar, 1999).
During smooth pursuit eye movements, saccades in the
same direction as the preceding pursuit have shorter
latencies than those in the opposite direction (Tanaka
et al., 1998). Also, manual reaction times are shorter
for stimuli flashed ahead of the pursuit target than for
those presented behind (van Donkelaar, 1999). Thus,
the distribution of visual attention is asymmetric along
the line of the eye movement. If one considers the
retinal event during smooth pursuit eye movements
against a structured field, it is identical to the retinal
event during active fixation against an optokinetic mo-
tion field. Plus, the attentional shift during smooth
pursuit and that during active fixation (in the present
experiment) are in the same direction, namely, toward
the In-coming field. These similarities imply a common
mechanism for active fixation and smooth pursuit. In
fact, recent studies have shown that the smooth-pur-
suit system and the fixation system may share common
mechanisms to some extent (Munoz & Wultz, 1993a,b;
Tam & Ono, 1994; Krauzlis & Miles, 1996), although
these systems are not identical (Luebke & Robinson,
811
1988; Goldreich, Krauzlis, & Lisberger, 1992; Schwartz
& Lisberger, 1994). For example, some cells in the
rostral superior colliculus in monkeys become active
both during smooth pursuit and fixation (Munoz &
Wultz, 1993a,b; Krauzlis, Basso, & Wultz, 1997).
The other related phenomenon is the residual
smooth eye movement under OKN suppression (Tam-
minga & Collewijn, 1981; Wyatt & Pola, 1984;
Collewijn & Tamminga, 1986; Waespe & Schwarz,
1986, 1987; van den Berg & Collewijn, 1987; Pola,
Wyatt, & Lustgarten, 1995; Wyatt, Pola, Lustgarten, &
Aksionoff, 1995). When the optokinetic motion field
changes in a sinewave fashion, small residual smooth
eye movements, which are roughly counter-phase to
the field motion (i.e., toward the in-coming field), can
be observed (Wyatt & PolaPola, Wyatt, & Lustgarten,
1992). These residual eye movements are enhanced
with a foveally stabilized target and background mo-
tion (open-loop condition) and depends critically on
stimulus predictability (Wyatt et al.). Although the size
of eye movements are quite small, the residual eye
movements could be detectable even with unidirec-
tional field motion (Waespe & Schwarz, 1987)2. Based
on extensive explorations of this phenomenon, Pola et
al. suggested that the OKN suppression with a station-
ary fixation stimulus may be mediated by automatic
mechanisms that process relative motion between the
fixation target and the motion field and depend on the
subject’s attention. This tendency for the eyes to devi-
ate into the In-coming field under the OKN suppres-
sion may be closely related to the shift of the OKN
beating field.
The suggested automaticity of the residual smooth
eye movements (Pola et al., 1995) parallels the results
of Experiment 4 which showed that the In-coming field
advantage survives the subject’s endogenous allocation
of attention. Also, the dependency of the residual eye
movements on stimulus predictability (Wyatt et al.,
1995) parallels that of smooth pursuit (Yasui &
Young, 1984) and, presumably, the results of Experi-
ment 5. It took, at least, 150 ms for the In-coming
field advantage to be significant, suggesting that this
duration of predictable motion stimulation is neces-
sary. The shared characteristics among smooth pursuit,
active fixation, the residual eye movement during
OKN, the shift of the OKN beating field, and the
In-coming field advantage are suggestive of the exis-
tence of a mechanism which processes relative motion
signals in the retinal coordinate and quickly regulates
various orienting responses.
2
In fact, a close examination of Fig. 4 would suggest that there
might be a slight tendency of deviation toward the in-coming field.
This could be due to the residual slow eye movements. However, it is
quite unlikely that such small deviations of gaze were the main cause
of the in-coming field advantage in Experiment 2.
812 K. Watanabe / Vision Research 41 (2001) 801–814
Finally, the In-coming field advantage was found
only for the detection task and the localization task (i.e.
orienting tasks), not for the color discrimination task
and the shape discrimination task (i.e. identification
tasks). Although a clear conclusion warrants more in-
vestigation, it seems to be consistent with the idea that
identification depends mainly on endogenous compo-
nents of attention whereas orientation depends mainly
on exogenous components of attention (Nakayama &
Mackeben, 1989). However, if the In-coming field ad-
vantage is a form of exogenous attention, the sustained
nature of it (Experiment 5) is puzzling. This is because,
it has been suggested that endogenous attention is slow
and sustained, and exogenous attention is fast and
transient. (Nakayama & Mackeben, 1989). Therefore,
further investigation should be conducted to examine
the relationship between other forms of exogenous at-
tention and the In-coming field advantage.
8. Conclusion
The present study demonstrated that unidirectional
field motion tends to shift visual attention in the visual
field from which the motion field originates (In-coming
field advantage). The In-coming field advantage seems
to be based on a relatively quick automatic mechanism,
which analyzes motion field and modulates spatial at-
tention accordingly. Such a mechanism might allow a
faster orienting response for the target appeared in the
direction of self-motion and thus provide behavioral
benefits.
Acknowledgements
Thanks are due to Romi Nijhawan and Rory Sayres
for comments on an earlier version of the manuscript
and two anonymous reviewers for valuable comments.
This work was supported by the Sloan Foundation
Fellowship for Theoretical Neurobiology.
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