23

Vision , Attention , and Action : Inhibition and Facilitation in Sensory -Motor Links Revealed by the Reaction Time and the Line Motion ShinsukeShimojo, YasutoTanaka, Okihide Hikosaka, and Satoru Miyauchi ABSTRACT To investigate critical factors for local inhibition and facilitation in visual-motor tasks, we randomized stimulus dimensions(location, color, and orientation) and response-stimulus interval acrosstrials. The subject performed four different tasks'. Reaction times at the samelocation were longer, that is, "inhibition of return" (lOR ) occurred, in the detection and location discrimination tasks. Reaction times at the samelocation were shorter in the color and orientation discrimination tasks. lOR was observed also in arm-reaching and saccadiceye movement tasks. Moreover , the task-dependent difference of RTs was observed also with popout displays. The results indicate a dissociation of two visual functions: detection/ orienting and fine feature analysis. To investigate whether motor readinesscould draw attention to the target location and have an influenceon visual information processing, we employed the "line motion" illusion; we showed that mere preparation for a motor response, such as arm reaching or saccade , would be sufficient to yield local facilitation at the prepared target location. In this chapter , we discuss issues related to functional and the motor one employing tion

of two

paradigm

links between

the visual

systems . For this purpose , we will present two sets of findings : the reaction time paradigm visual

and indicating

functions , and the other

and indicating

effects of motor

employing

functional

segrega -

the " line motion "

readiness on the visual information

processIng .

0N 23.1 REACTI0 N TIMEAND ATTENTI Spatial attention is an indispensable aspect of visual information processing: indispensable because the brain has a limited capacity, whereas the constant flows of sensory inputs could be infinite in theory . Without selection and filtering by attention , it would be impossible to perceive what is important and to respond to it appropriately Games 1890; Helmholtz 1910; Broadbent 1958). On the other hand, some authors have recently argued that there is no limit to the brain 's capacity for visual information processing (Van der Heijden 1991). Limited capacity may , instead, concern " selection for action ." This view of spatial attention may be a part of the reason why reaction time (RT) has been often employed as a sensitive measure to access the selection

and

filtering

context

processes

For

instance

,

sequentially

,

presented

the

cue

tion

of

-

the

various

( lOR

Egeth

1994

is

ms

where

same

so

target

1500 "

and

.2

revealed

location

. This

return

Kwak

23

has

situation

the

same

and

( 300

ale

it

a at

the

location

large

et

consider

either

at

different of

, and

effects

in

a

spatiotemporal

.

Posner

)

and

1992

or

different

as

the

;

cue

,

R T

when onset

and

the

1984

et

ale

target .

is

When

This

;

the than

or

has

)

and

is

relatively " inhibi

Nissen

Egeth

the onset

called ;

is

at the

SOA

1985

Gibson

target that

been

( Maylor

1994

presented

between ,

) .

are

larger

interval

duplicated

Tassinari

a

asynchrony

Cohen

repeatedly

and

locations cue

particularly ( stimulus

;

a

-

1985

1994

;

;

Tipper

) . 1

INHIBITION

OF

RETURN

AND

ITS

UNDERLYING

MECHANISM

The

underlying

tion

task

effect of

mechanism involves

occurs the

the

.2

target

closely

Some is

to

spatial

( Nissen

occurs good

in

the

visual

instead Valdes

To the employed

of ,

orienting

inhibition

and

,

Neill

understand inhibitory

the effect

by

may

1994

Kwak

and

and

occur

).

,

in

the

is

Egeth

Egeth

Meanwhile

however

it

.

unclear

inhibition

than

process 1980

, ,

that

, rather

Kwak

( Posner

unknown

processing

( Kwak

location ;

is

of

suggest

changed

1985

metaphor

lOR

levels

researchers

alone

related target

of

various

any

1992 for

) .

which ,

),

They

different

location is

that

" spotlight suggest

or

, of effect

would

that

conditions

therefore

this

"

-

the

attributes

argue

a

others

lOR visual

reac

level

when

that

other

the

which

occurs

1992 to

Because at

be

facilitation tasks

a ,

( Terry

,

).

mechanisms

more

in

detection

and

a

simple

Egeth

( 1992

inclusively

, we task

,

which

first

tried was

to similar

duplicate to

that

) .

A Pilot Study A single target was presentedin either the top left or top right position of the display, while the subject fixated at a point (FP) in the bottom center. The location of the target was randomized between these two locations across trials. The interval between the button-pushing responseand the next target appearance(reaction stimulusinterval, or RSI) was also randomizedacrossthe trials (200/ 400/ 1,000/ 2,000 ms). The distancebetweenthe target and FP was 12 degrees. Note that there was no cue in this experiment, and we were mostly interestedin the positional effect of the previous target on RT to the present target. Four subjects(two naive and two nonnaive (authors)) were askedto detect a target, and to pressa mousebutton as quickly as possible. In results, strong inhibition of return was obtained for all subjectsat all RSIs, except the longest (2,000 ms). Thus the cue/ target distinction is not a necessarycondition for lOR, as originally suggestedby Tassinariand his colleagues(Tassinariet al. 1987).

598

Shimoj0 et at.

23.3

TASK DEPENDENCY OF INHIBITION AND FACILITATION

What are the critical conditions for the lOR effect, then? Could the distinction between detection and discriminationbe critical? Or rather, could the type of visual information, say, spatial versus nonspatial attributes be critical (Kwak and Egeth 1992; Tassinari et al. 1994; Terry, Valdes, and Neill 1994)? To addresstheseissues, we conductedexperiment 1.1. Experiment 1.1: Task Dependency of Inhibition and Facilitation of -

Return in Button Pressing Subjects Six subjects participated in the experiment : four naive and two nonnaive (authors). Stimuli and Apparatus Stimulus configuration was similar to that employed in the pilot study ; as in the pilot study , there was no cue. Location (left / right ), color (red/ green), and orientation of the target (vertically / horizontally elongated ) were all randomized across trials (see fig . 23.1). The stimuli were presented on the CRT display , controlled by a personal computer (Commodore Amiga 500). Target size was 0.5 degree x 0.4 degree, and its luminance was 6.4 cd/ m2 (hue: red (.555, .344); green (.320, .555)). The fixation point ' s size was 0.4 x 0.4 . Luminance of the background was 0.01 cd/ m2. The distance between the fixation point and the target was 6.0 degrees. Viewing distance was 114 cm. Subject' s head was

0

to

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previous target

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+ \

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.....

..... "'"

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... ... ... ...

t2

d ,

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Respon e Stimulus Interval (RSI )

360 "

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Figure 23.1 Stimulusconfigurationand sequencefor RT experimentwith single target (experiment1.1).

599

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fixed

by

using

the

of

the

chin

Ober

1 .3 dark

Procedure to

mouse

on

location (4 ) was

the

ms

in

response

in

four

only

subject

was

Subjects

;

analyzed

and

total

present

were

Results

,

found

same

).

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Cue LeadTime (ms) Line Motion Reaching Figure 23.18 Resultsof the arm-reachingline motion experiment , with the cuelead time randomized(test session4). Rateof line motionfrom cued(target) sideis plotted againstcue leadtime for onesubjectasexample(solidcurve). Meanperformance time of arm-reachingat eachcue lead time was also calculatedfrom arm-reachingtrials and plotted in samepanel (dottedcurve).

ep~s peno WOJ ,:j UO!~ V'J eu!l e~et:J

performance time of reaching, which has been calculated from the reaching trials in the same test session (the dotted curve in the figure ). As obvious in the figure , the rising time course of line motion was inversely correlated with the performance time of reaching, as the waiting time increased. The results were similar in this regard, though somewhat noisier , for the other subjects. This suggests that the attentional mechanism reflected in the line motion might in fact be related to the motor readiness for reaching . 23 .11

SACCADE READINESS WITH THE LINE MOTION

To see if the finding could be generalized to another kind of motor response, we employed a saccadic eye movement task.

622

Shirnojo et at.

Saccade Training Session

Fixation Point (FP) On

.

67

FP Color

..

~R 1 m

Cue Lead Ti ~~ 1330

G m

'"

i

ms

Change

Saccade to center of the target

I

:

TargetsOn

- - -

-

~

Time

Figure 23.19 Stimulus configuration and sequencefor saccadetraining (experiment 2.2). Stimulus parameterswere very similar to those in arm-reaching experiments. Subject' s task was to move eyes as quickly as possible to relevant target, which was indicated by color change of fixation.

Experiment2.2: Line Motion from the Goal Locationof SaccadicEye Movement Subjects

Three

subjects

Stimuli

, apparatus

were

virtually

stimulus a

training which

was

done ( up

percent

and

in

the

experiment

line

details

of

experimental

the

color

course quite figure

/ line

was

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as

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of line

motion

) . This

was

the

the

in the

saccade

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so for

two

other

Vision, Attention , and Action

of goal

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session

( the

subjects

as well

the thick

session .

the make

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session the

same

as in 4 ).

results

. When

period ) , then

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a in

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same

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with

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. The

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fig . 23 .9B

task , which

. Subjects

were

experiment

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and

as

the

( see

fig . 23 . 19 ) , and

time

presented

unfold

comparable

trials

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of

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; see

lead

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nature

target

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probe

the

experiment

the

the

trials

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probe

appeared

623

of

the 200

in

for

to

by to

All

those

movement

arm - reaching

line

the

participated

procedure

) , except

session 50

the

time

authors

to

( see fig . 23 .20 ) . The

Results

was

and

identical

eye

recording

trials

the

configuration

saccadic

the

of

.

( after the

. The

line rising

in fig . 23 .21 ) thin

curve

in

Saccade Test Session

Fixation Point (FP) On

!

67

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.

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Time

Varied (16.7 - 1600 ms ) Targets On In 50 % Trials

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g

.

~

~

------ -.--.-....-....-..... A line probe, and the subject should judge the direction of illusory motion

Task

: saccade

to

the

center

of

target as fast as possible

Figure 23.20 Stimulus configuration and sequencefor saccadetask, with line motion (experiment 2.2). Just as in arm-reaching experiment (2.1; see fig. 23.17), line probe was presented with varying cue lead time beforeblink of fixation point as GO sign. These line probe trials were 50 percent of total trials, and were again randomized with the saccadetrials.

Thus the rising time coursesof focal attention indicatedby the line motion were very similar between two types of motor responses , saccadiceye movement and arm reaching. It is consistentwith the idea that readinessor preprogramming for responseto a particular target location is alone sufficient to yield a local facilitation strong enough to induce an illusory line motion. Some previous studies also have provided similar data (e.g., Rizzolatti et al. 1987; Klein 1980), though without the line motion as a measure.

23.12 A COMMONATTENTIONMECHANISM ? On the other hand, this doesnot necessarilymeanthat the line motion, which is a visual effect, and the motor programming, which is by definition nonvisual, could not be dissociablein terms of the relevant attention mechanism. We have several reasons for this skepticism. First, our subjects had been trained visually in the first training sessions ; that is, their motor performance was guided by visual input and feedback. And even in the later training and

624

Shimojo et al.

Line Motion by Readiness for Reaching & Saccade

88

1 .0

0 .9

0 .8

0 .7

0 .6 -- -- - -- ----- -- - - --- - -- - -- -- -- - -- - - - - -- - - - --- - -- - -- - - - () . ~

- - -- - -

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0.2

c

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0.1

.

Saccade

0 .0 16 . 7

100

200

400

600

800

12001600

Cue Lead Time (ms) Figure 23.21 Resultsof saccade line motion experiment , with cue lead time randomized (experiment2.2). Rateof line motionfrom the cued(target) sideis plottedagainstcueleadtime for one subjectasan example(thick curve). Rateof line motion for samesubjectin previous arm-reachingexperiment(2.1; seefig. 23.18) is replottedfor comparison (thin curve).

ep!s peno WOJ .:J UO!~ V'J eU!l e~e'tj

tests, their performancewas heavily based on visual memory. This could potentially explain why motor readinessinevitably triggered local facilitation at the target location in the visualfield, which in turn led to a line motion. Second, we actually conducted another subexperiment, where stimulus configuration and sequencewere similar to the previous experiments. The only differenceswere that top half or the bottom half of the target was randomly chosen and made slightly brighter than the other half, and that subjectshad to indicate which of the halves was brighter by a button-pressing response(a two-alternative, forced-choice task). Thus subjectsstill had to decode the meaning of color change of the fixation point to decide which target would be task-relevant, but also to constantly fixate on the fixation point and simply to do a visual discrimination task as fast as possiblewhen the GO sign (= the blink of the fixation point) was given. Thus in order for the fastestresponse, subjectshad to develop a visual expectation as to which location the relevant target would be at, and mentally "wait right there." We

625

Vision, Attention , and Action

then

randomized

ination

50

trials

by

experiments

.

percent

of

which

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trials

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one the

underly ,

,

it

the

it

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motor

find

,

this

such

Campsall

possibility and

these

line

is

motion

leads

to

23

a

. 13

We

be

visual

found

the

that

.

tasks

,

required

a

as

detailed

even

when

seemed "

the

feature

task

likely

it

the

second

. The

the

observed

cue

or

nonvisual

Shimojoet al.

answer line

under

some

sensory

have

;

confirmed

controversial

such

also

required it

in

the

color

,

global

was

detection

different

arm

kinds

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return

shape

as

, ,

or

before

.

of

tasks

when

.

the

vernier This

task offset

,

dichotomy

dichotomy

feedback

global

inhibition

kind

,

.

between

of

This

task

the

- dependent

indicates

location rather

underlying

is

than

that

based

on

facilitation

the

parallel .

inhibition

even

Thus

of

it

return

.

,

purely both

same

employed

of to

effect

the

was

chapter

yes

as

found

pathway

was

task

of

identical

the

readiness .

whether

, facilitation

corlicallevels

to

,

neurophysiological

display

the

or

and motor

DIRECTIONS

occurred

movement

attentional

conditions

the

task

single

that

when

also

,

a

.

we

lead

motion

have

particular

communication

FUTURE

of

were the

neural

of

a

programming

least

namely

pathways

early

can

the owing

that

to

highly

motor

nature

target

leads

the

-

same

,

literature

that

conclude

information still

half

alone

the

popout

involves

readiness motion

,

that

significantly

In

,

- demanded

analysis

seems

eye

,

the

the

nature

prior

still

only

return

saccade

with "

the

conclude

occurred the

tendency

surprisingly when

return of

of

" what

. be

by

in

studies

is

AND

parameters

consistent the

Somewhat dichotomy when

the

stimulus

be

and

at

of

feature

the to

" where

of

regardless

opposite

But

personal

could

CONCLUSIONS

Inhibition

such

found

,

it

however

underlying

target

other

- guided

, we

inhibition ,

the

);

to

memory

effect

,

1980

premature

attentional

discrimination

We

seems

conservative

information

motor

626

it

SUMMARY

spatial or

,

Klein

,

) .

underlying

. To

;

1984

, or

. in

while

motion

mechanism .

visual

training

Driver

),

line

50

- reaching

attentional

be

in

mechanism

motion

studies

to

target

arm

-

previous

directions

single line

the

( Jon 1993

two

discrim

the

of

the

attentional

still

shift

saccade

1980

Pierce

reasons

mechanism

a

course

mechanism

some

Herdman

( Posner

( Remington

For

and

a

the

in

been

attentional

as

,

is ,

might

/ memory have

in of

visual in the

in

attentional

readiness

for

lead

and

of

did

time

obtained

. That

the

rest

we without

rising

that

could

tasks

feedback

as

commonality

that

there

evidence

response

Stelmach

a

result

the

just

presented

to

the

motor

visual ,

was

programming

the

,

obtained

visual

with

time

similar

suggest

finally

trials

lead

we

This

the

during

and

probe ,

motor also

heavily

line

suggest

and

probe

cue

highly .

the

line

the

result

be

could

could

Third

failed

the

a

to

motor

motion

to

of

experiments

could

line

out

hand

tween

token

is , . As

turned

the

percent

varying

we

raise

visual for could .

the not

the attentional

question

of effect

arm

- reaching

and

be

attributed

to

, the any

whether that

motor is ,

the

saccade kind

line tasks

of

visual

;

This does not necessarilyindicate that the same attention mechanismis shared in the two

cases, the motor

readiness and the line motion . It could

simply mean that motor readinessalone is sufficient to yield local facilitation of visual processing, which is responsiblefor the line motion, particularly when the subjectshave been trained to perform the motor task basedupon visual memory .

Also , our data have not yet addressed the original issue, namely , at which level the inhibition

and facilitation

of return occur , whether

sensory , sensory -

motor, or motor. In this regard, it would be interesting to compare the reaction time paradigm with the line motion paradigm, holding stimulus parametersas close as possible. This seemsto be a promising way to resolve the prickly issuein the field: that is, of how many IIattention mechanisms " we have to deal with . A CKN 0 WLEDG

MENT

The researchprojects reported here have been supportedpartly by Grant-inAid for Creative Basic Research from the Ministry of Education , Science, and Culture of Japan, the Human Frontier Science Program , and Nissan Cam-

bridge BasicResearch . NOTES

1. It is believed that lOR occurs only when the peripheral cue is not informative, that is, when it does not predict the location of the target (Posner 1980; Posner and Cohen 1984). However, Tanaka and Shimojo (1993) systematically manipulated the probability that the target was presentedat the samelocation as the previous target, successfullyisolating the lOR from the predictability effect. For example, they compared two independent sessions: one in which the target was presentedat the samelocation with p = .8 (the opposite location with p = .2), and the other in which it was presented at the opposite location with p = .8 (the same location with p = .8). They found that the RTs at the same locations in the former session tended to be

larger than the RTs at the opposite locations in the latter session, even though the predictabilities

were the same ( p = .8). Thus the " location priming " (i .e., lOR ) could be isolated

from the "probability priming." 2. In fact, recent studies suggest that there might be two types of lOR, one related to eye movements and the other related to stimulus detection (Abrams and Dobkin , 1994 ; Tipper et al . 1994 ).

3. Goodale and Milner (1992 ) considered both the " action " and the "recognition " functions

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