Tilburg University On the relationship between arousal and activation, and sensation seeking de ...

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Tilburg University
On the relationship between arousal and activation, and sensation seeking
de Brabander, B.; Boone, C.A.J.J.; Gerits, P.; van Witteloostuijn, A.
Published in:
Personality and Individual Differences
Publication date:
1995
Link to publication in Tilburg University Research Portal
Citation for published version (APA):
de Brabander, B., Boone, C. A. J. J., Gerits, P., & van Witteloostuijn, A. (1995). On the relationship between
arousal and activation, and sensation seeking. Personality and Individual Differences, 18(3), 373-384.
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Person. indiwd. Di%f: Vol. .81 No. .3 pp. .483-373 5991
Coovtieht ni Q 1995 Elsevier ecneicS Ltd
detnirP ta& .niatirB All tights devreser
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RELATIONSHIP BETWEEN AROUSAL AND ACTIVATION,
AND SENSATION SEEKING
treB eD ’,rednabarB ehpotsirhC enooB *, loP ’stireG and nejrA Van ’njiutsoolettiW
ytlucaF‘ fo deilppA ,scimonocE ytisrevinU fo ,ACUR-prewtnA naalmiehleddiM ,I ,prewtnA-0202
muigleB dna etatS‘ ytisrevinU fo ,grubmiL taartsesregnoT 53, 6200 DM ,thcirtsaaM ehT sdnalrehteN
(Received 24 Murch 1994)
Summary-In suoiverp stnemirepxe ew detset eht mgidarap eD( ,rednabarB Perceptual and Motor Skills,
69, 75-82, 1988) taht cisahp lasuora snoitcaer era detcelfer yb dedrater snoitcaer retfa na detcepxenu
yrotaraperp langis ni a laitapsousiv thgir( )erehpsimeh noitanimircsid noitcaer emit .ksat cisahP noitavitca
sesnopser era nwohs yb detarelecca snoitcaer retfa eht yrotaraperp slangis ni a citnames tfel( )erehpsimeh
.ksat nI eht tneserp tnemirepxe eht stluser od niaga etadilav eht .mgidarap ehT niam esoprup fo eht yduts
saw ot esu siht mgidarap ni redro ot etagitsevni rehtehw noitasnes gnikees si detaler ot erom esnetni cisahp
lasuora snoitcaer )ytilibasuora( ro ot erom decnuonorp cisahp noitavitca sesnopser )ytilibatavitca( ro ot .htob
tA tsrif thgis eht stluser etacidni taht ylno ytilibasuora si detaler ot noitasnes ,gnikees yllaicepse moderob
.ytilibitpecsus rehtO elbissop snoitanalpxe fo eht stluser era osla .dessucsid
INTRODUCTION
nI owt tnatropmi weiver selcitra nivraM namrekcuZ ,4891( )0991 has dezirammus and dessucsid eht
sgnidnif no eht lacigoloibohcysp setalerroc fo eht tiart fo noitasnes .gnikees snoitalerroC fo this tiart
htiw lacigoloisyhpohcysp ytivitcaer serusaem and htiw lacimehcoruen serusaem fo ,srettimsnartoruen
cilobatem semyzne and setilobatem devlovni ni eht -niarbdim and metsniarb smetsys hcihw etavitca
lacitroc sessecorp era gnitseretni ni sevlesmeht tub erom os fi yeht nac etadicule eht tnereffid sedom
ni hcihw mammals evah denrael ot tcaer ni an evitpada yaw ot eht lacigoloib ecnatropmi fo yrosnes
.ilumits fI ew daer mih ,llew namrekcuZ sezingocer eerht latnemadnuf sedom )sexelfer( hcihw ew
evah detirehni morf ruo :srotsecna gnitneiro dewollof yb ,noitautibah eht evisnefed xelfer and eht
eltrats mrala( ro )tpurretni .xelfer esehT sexelfer rof snosaer llits ,nwonknu wohs emos ytilibairav
htiw tcepser ot eht ytisnetni and ycneuqerf htiw hcihw yeht era desu ni humans. ehT noitasnes gnikees
tiart is desoppus ot erutpac yllaicepse eht htgnerts fo eht gnitneiro esnopser ot tnaveler levon ilumits
hcihw swolla retteb desucof noitnetta ot eseht .ilumits eW :etouq I“ suggest that laudividni secnereffid
ni ytivitcaer ot esnetni and levon ,noitalumits that edivorp eht basis rof eht noitasnes gnikees ,tiart
yam eb eht dne tluser fo larutan noitairav ni devlove A W smsinahcem ni humans” ,namrekcuZ(
1990, p. .)413 ’A‘ snaem hcaorppa and ’W‘ withdrawal ni s’alrienhcS ygolonimret ,alrienhcS( .)9591
’gnikeeS‘ is eht devlove noisrev fo .hcaorppa And fo ,esruoc evitceffe gnikees fo ,yerp ,doof ,retlehs
etam and gnirpsffo sdeen an trela noitartsiger metsys fo lufgninaem ilumits ro ni rehto sdrow a hgih
yticapac fo desucof .noitnetta ,revewoH ew rednow rehtehw eht eltrats xelfer hcihw is lativ rof lavivrus
is ton neve erom yllacisnirtni detaler ot noitasnes .’gnikees‘ ehT nosaer is that ew esoppus that tsom
fo eht htiw tcepser ot eht lacigoloisyhporuenohcysp sgninniprednu fo noitasnes gnikees suggest
that lasuora hcihw erom and erom smees ot eb eht tceffe fo eht cigrenerdaron noitavitca metsys fo
eht ,niarb is eht elbairav hcihw tseb setaitnereffid hgih and wol noitasnes srekees SSH( and .)SSL
ehT lautpecnoc krowemarf hcihw ot us smees tsom elbatius ot dnatsrednu this tcepsa fo yrosnes
rotom gnilpuoc ni eht ,niarb and eht laudividni snoitairav ,niereht is that fo marbirP and ssenniuGcM
)5791( and rekcuT and nosmailliW .)4891( gnikaT as a starting tniop eht ledom noitnetta desoporp
yb marbirP and ssenniuGcM ,)5791( eht srohtua rekcuT and nosmailliW )4891( desserts eht
ecnatropmi fo eht ecnereffid neewteb cigrenerdaron and cigrenimapod lasuora fo eht .xetroc On eht
basis fo eht neht elbaliava lacigoloruen ,ecnedive hcihw has neeb demrifnoc and dednapxe ecnis
,kralC( ,neffeG & ,neffeG 1987a, b, ,)9891 yeht dedulcnoc that eht owt smetsys trexe tnereffid
lanoitnetta .snoitcnuf ehT main noitcnuf fo eht cigrenerdaron metsys smees ot tsisnoc ni gnitatilicaf
gnitneiro ot levon ilumits and noitautibah ot detaeper noitalumits evitceles( yrosnes .)noitnetta ehT
313
t r e B e D r e d n a b a r B r e a l .
4 1 3
e n i m a p o d s ’ m e t s y s main , n o i t c n u f , r e v e w o h s m e e s r e h t a r d e r a e g a t e h t n o i t a r a p e r p a n d n o i t a t i l i c a f
f o e h t g n i l p u o c f o e h t t h g i r e s n o p s e r o t i l u m i t s d e s u c o f ( ) n o i t n e t t a e e s [ o s l a r e n i e W ) 0 9 9 1 ( r o f a n
e v i s n e t x e n o i s s u c s i d f o e s e h t s m e t s y s a n d r i e h t e l o r n i t n e t a l . ] n o i t i b i h n i e h T c i g r e n e r d a r o n m e t s y s
i s e h t l a r u e n e t a r t s b u s f o w h a t m a r b i r P a n d s s e n n i u G c M ) 5 7 9 1 ( e v a h d e l l a c e h t l a s u o r a ‘ ’ m e t s y s
s a e r e h w e h t c i g r e n i m a p o d m e t s y s s e d i c n i o c h t i w r i e h t d e l l a c o s n o i t a v i t c a ‘ . ’ m e t s y s r e k c u T a n d
n o s m a i l l i W ) 4 8 9 1 ( o s l a e t i c e m o s e c n e d i v e that e h t c i g r e n e r d a r o n a n d c i g r e n i m a p o d s m e t s y s e r a
y l l a i t n e r e f f i d d e k n i l o t e h t s n o i t c n u f f o e h t t h g i r a n d t f e l . e r e h p s i m e h e h T l a s u o r a m e t s y s i s d e s o p p u s
o t s u s t a i n e h t l a i c e p s y r o s n e s s e i t i c a p a c f o e h t t h g i r l a t e i r a p e r e h p s i m e h e l i h w e h t n o i t a v i t c a m e t s y s
d l u o w r e h t a r t r o p p u s e h t r o t o m - y r o s n e s s e i t i c a p a c f o e h t t f e l l a t n o r f . e r e h p s i m e h
s a e r e h W l a c i m o t a n a - o r u e n e c n e d i v e r o f n e v e n u s n o i t u b i r t s i d f o c i g r e n e r d a r o n a n d c i g r e n i m a p o d
e v r e n s g n i d n e n i e h t x e t r o c f o a n i m a l s i s a b u n d a n t , h u m a n e c n e d i v e i s e c r a c s a n d s e o d t o n y l t c e r i d
t r o p p u s e h t n o i t a z i l a r e t a l s i s e h t o p y h , r a p s a G ( , r e g r e B , t e r v b e F y n g i V & , y r n e H . ) 9 8 9 1 r e v e w o H t h i s
s e o d t o n y l p m i that a l a n o i t c n u f n o i t c e n n o c n e e w t e b e h t o w t l a c i t r o c l a s u o r a s m e t s y s a n d e h t
s e r e h p s i m e h i s . d e d u l c x e r o F , e c n a t s n i , r e l t t i K z t i w e k r u T a n d g r e b d l o G ) 9 8 9 1 ( d e v r e s b o a n l a i t i n i t f e l
l a u s i v d l e i f e g a t n a v d a n i y l t c e r r o c g n i z i n g o c e r y l l a r e t a l a n d y l l a c i p o c s o t s i h c a t d e t c e j o r p x e l p m o c
l a u s i v s n r e t t a p ) i j n a k ( a n d a t f i h s s d r a w o t t h g i r l a u s i v d l e i f e g a t n a v d a n e h w e h t s e r u g i f e m a c e b e r o m
. r a i l i m a f s i h T g n i d n i f i s e l b i t a p m o c h t i w e h t a e d i that e h t t h g i r e r e h p s i m e h h c i h w i s d e z i l a i c e p s n i
k c i u q c i t s i l o h n o i t i n g o c e r f o l a u s i v s t c e j b o w a h s d a r B ( & , n o t e l t t e N 198 ; 1 i h c m i K & , v a h r e M , ) 1 9 9 1
s t i u r c e r e h t l a i c e p s s e i t i c a p a c f o e h t l a s u o r a m e t s y s o t e c n a h n e e h t e s i o n / s u l u m i t s o i t a r f o e h t
l a u t p e c r e p m e t s y s n e h w e h t i l u m i t s e r a . l e v o n i t t o i L a n d r e k c u T ) 2 9 9 1 ( d n u o f a g n i w o l s f o n o i t c a e r
s e m i t n i S s h t i w d e c u d n i e v i s s e r p e d d o o m y l n o o t t f e l l a u s i v d l e i f c i p o c s o t s i h c a t . i l u m i t s t I h a s n e e b
d e t a r t s n o m e d that e v i s s e r p e d d o o m i s d e t a i c o s s a h t i w d e c n a h n e e v i t a l e r t h g i r e r e h p s i m e h l a s u o r a
, r e f f a h c S ( n o s d i v a D & , n o r a S ; 3 8 9 1 s e u q i r n e H & , n o s d i v a D ; 0 9 9 1 , t o n i t r a M , y d r a H , e n i l e F , t e r u H
, r e y o z a M , y v e L - r a t t A P a p p a t a & , a t o r y S ; 0 9 9 1 , n e k r a m o T n o s d i v a D & , s e u q i r n e H ; 0 9 9 1 s e u q i r n e H
& , n o s d i v a D n i . ) s s e r p , n e e r G , s i r r o M , n i e t s p E t s e W a n d , r e l g n E ) 2 9 9 1 ( d n u o f a e v i t i s o p n o i t a l e r r o c
n e e w t e b t h g i r s v t f e l l a c i t r o c l a s u o r a n i e h t G E E a n d a t h g i r s v t f e l l a u s i v d l e i f e g a t n a v d a s i d a s
d e r u s a e m y b s e c n e r e f f i d n i n o i t c a e r s e m i t n i a l a c i x e l n o i s i c e d t a s k . U s i n g e h t , m g i d a r a p - p o o r t S
l h a d h g u H a n d n o z n a r F ) 7 8 9 1 ( d e v r e s b o l a i t i n i e t a r - t r a e h n o i t a r e l e c e d g n i r u d e h t t s r i f l a i r t k c o l b y l n o
o t i l u m i t s y l l a c i p o c s o t s i h c a t d e t n e s e r p o t e h t t h g i r . e r e h p s i m e h e t a r - t r a e H n o i t a r e l e c e d y l l a c i p y t s r u c c o
n i s n o i t a u t i s g n i v l o v n i g n i t n e i r o . r o i v a h e b e h T s r o h t u a o s l a n o i t n e m e h t g n i d n i f f o a n n o i t a i c o s s a
n e e w t e b d e s a e r c n i t h g i r e r e h p s i m e h d e k o v e s l a i t n e t o p a n d e t a r - t r a e h , n o i t a r e l e c e d y b r e k l a W a n d
S a n d m a n . ) 9 7 9 1 ( l l A t h i s e c n e d i v e s u g g e s t s that e r e h t t h g i m e b a l a i c e p s k n i l n e e w t e b e h t s n o i t c n u f
f o e h t t h g i r e r e h p s i m e h a n d e h t l a s u o r a m e t s y s h c i h w s e t a t i l i c a f g n i t n e i r o o t l e v o n . i l u m i t s A n
n o i t a c i d n i f o a l a i t n e r e f e r p n o i t a i c o s s a n e e w t e b e h t r o t o m - y r o s n e s g n i l p u o c n o i t c n u f f o e h t t f e l
e r e h p s i m e h a n d e h t n o i t a v i t c a m e t s y s i s r o f e c n a t s n i e h t g n i d n i f y b d d i K a n d l l e w o P ) 3 9 9 1 ( that s n o s r e p
h t i w a d e c n u o n o r p l a c i p y t o z i h c s y t i l a n o s r e p d e d n e t s d r a w o t e v i t a l e r t f e l e r e h p s i m e h , n o i t a v i t c a r e v o
a s d e r u s a e m y b d e h s i n i m i d t f e l e r e h p s i m e h d e t a l e r - k s a t r e w o p - a n o i t c u d e r n i e h t . s e v a w - G E E t I h a s
n e e b y l t n e i c i f f u s d e h s i l b a t s e that a i n e r h p o z i h c s i s d e t a i c o s s a h t i w c i g r e n i m a p o d n o i t a v i t c a r e v o
, n e s a e r d n A ( ; 8 8 9 1 , r e n i e W . ) 0 9 9 1 y l t n e u q e s n o C e n o y a m a s s u m e that a i n e r h p o z i h c s e n o r p s n o s r e p
e v a h t h i s t f e l e r e h p s i m e h n o i t a v i t c a r e v o e s u a c e b f o a n d e t a v i t c a r e v o c i g r e n i m a p o d n o i t a v i t c a . m e t s y s
e r o M t c e r i d e c n e d i v e f o a k n i l n e e w t e b e h t c i g r e n i m a p o d m e t s y s a n d e h t t f e l e r e h p s i m e h i s d e t s e g g u s
y b e h t s t l u s e r f o e h t m e t r o m - t s o p s i s y l a n a f o n i a r b t i s s u e s f o c i n e r h p o z i h c s s t n e i t a p y b s d l o n y e R ) 3 8 9 1 (
o h w d e v r e s b o a c i f i c e p s e s a e r c n i f o e n i m a p o d n i e h t a l a d g y m a f o e h t t f e l . e r e h p s i m e h t I h a s o s l a n e e b
d e t a r t s n o m e d that t n e t a l n o i t i b i h n i , ) I L ( a n o n e m o n e h p h c i h w i s d e l l o r t n o c y b c i g r e n i m a p o d n o i t a v i t c a
a i v e h t s u e l c u n s n e b m u c c a , r e n i e W ( , ) 0 9 9 1 i s d e t a u n e t t a n i c i t o h c y s p e n o r p s l a u d i v i d n i n i h c i h w a
e r o m d e t a v e l e n o i t a v i t c a f o e h t t f e l e r e h p s i m e h n a c e b a s s u m e d , h c u r a B ( y e l s m e H & , y a r G ; 8 8 9 1
, w o b u L , s h c a S - g r e b g n I a d r O - n i e t s l a Z & , z t i w r e G . ) 2 9 9 1 e h T s g n i d n i f f o , t f a r C , h c t i v o r u o G , n o t w o D
n o t n a w S a n d e t r o f n o B ) 2 9 9 1 ( e r a o s l a t n a v e l e r r o f t h i s issue. y e h T d n u o f that e l a m S s h t i w
l a t n e m p o l e v e d e n i m a p o d n o i t e l p e d ) a i r u n o t e k l y n e h p ( d e w o h s a t h g i r l a u s i v d l e i f t n e m r i a p m i n i
g n i g a g n e s i d n o i t n e t t a m o r f a n d i l a v n i e u c n i a n o i t c a e r e m i t . t s e t y e h T d e t e r p r e t n i t h i s n o n e m o n e h p
a s a n o i t p u r s i d f o t f e l e r e h p s i m e h . n o i t c n u f c i g r e n i m a p o D , n o i t a v i t c a n i , l a r e n e g s m e e s o t t r o p p u s a n d
e t a t i l i c a f e h t n o i t c e l e s a n d n o i t a z i n a g r o f o r o t o m - y r o s n e s s n o i t a i c o s s a y b s n a e m f o g n i c n a h n e e h t
e c i o n - s u l u m i t s o i t a r f o d e n r a e l l a r o i v a h e b t u p t u o s m a r g o r p n i e h t y r a t n e m e l p p u s r o t o m s a e r a f o e h t
x e t r o c n e h o C ( & , r e b i e r h c S - n a v r e S . ) 3 9 9 1 e n i m a p o D s n o r u e n e r i f r o i r p o t a n d g n i r u d e h t n o i t u c e x e
,lasuorA noitavitca dna noitasnes gnikees 375
fo denrael stnemevom tub yldrah wohs yna ytivitca ni noitaicossa htiw suoenatnops stnemevom
rebierhcS-navreS( & ,nehoC .)2991
eW evah enod lareves stnemirepxe ot tset eht ytidilav fo eht mgidarap tuoba eht noitaicossa neewteb
eht lasuora metsys and eht thgir erehpsimeh no eht eno hand, and neewteb eht noitavitca metsys and
eht tfel erehpsimeh no eht rehto hand. nI eht stnemirepxe owt noitanimircsid noitcaer emit tasks era
,desu eno hcihw is rehtar elbitapmoc htiw eht seiticapac fo eht thgir erehpsimeh gnisserp(
sehctiwsorcim nehw owt ylevisseccus gniraeppa ilumits no a rotinom neercs era ni a denimretederp
)noitisop and thus ylekil ot eb dellortnoc yb that ,erehpsimeh and eno tfel erehpsimeh task gnitcaer(
nehw owt srettel ylevisseccus gniraeppa no eht neercs era ton htob slewov fo .)stnanosnoc nI htob
tasks, ni ylmodnar nesohc trials, an detcepxenu yrotaraperp signal sraeppa a tnemom erofeb eht
noitcaer stimulus neewteb( 300 and 100 .).cesm gnidroccA ot eht denibmoc sesehtopyh fo marbirP
and ssenniuGcM )5791( and rekcuT and nosmailliW )4891( eht nosrep dluohs eb deltrats nehw eht
detcepxenu signal sesuora eht nosrep ni eht thgir erehpsimeh task and yltneuqesnoc eht noitcaer emit
dluohs eb .regnol On eht yrartnoc nehw eht nosrep is detavitca ni eht tfel erehpsimeh task an
detarelecca noitcaer (as a sign fo detavele desucof noitnetta and laroivaheb )ssenderaperp dluohs eb
.devresbo nI eerht evisseccus stnemirepxe this smees ot eb eht esac eD( ,rednabarB ;8891 eD
rednabarB & ,enooB ;9891 eD ,rednabarB stireG & ,enooB .)0991 oslA ni eht elpmas fo this yduts
eht mgidarap smees ot eb dilav ees( eht Results .)noitces
nI his 1984 weiver fo hcraeser tuoba noitasnes gnikees ni ehT‘ laroivaheB and niarB ’secneicS
namrekcuZ sezirammus ni elbaT 1 eht lacigoloib setalerroc fo noitasnes gnikees and rieht laroivaheb
setalerroc ni humans and animals. noitcepsnI fo eht list fo senimaoib and srotalugeroruen shows that
ylno esoht detaler ot eht senimalohcetac enimapod )AD( and enilanerdaron )AN( era yltnacifingis
detaler ot noitasnes .gnikees SSH mees ot evah rewol slevel fo teletalp esadixO-enimA-onoM (MAO,
eht emyzne hcihw sezilobatac ,)AN desaerced ytivitca fo esalyxordyH-ateB-enimapoD ,HBD( eht
emyzne hcihw sezilobatem enimapod ot )AN and rewol diulF-lanipS-orbereC )FSC( slevel fo NA.
ehT lacigoloisyhp pihsnoitaler fo eht MAO and HBD serusaem htiw lartnec enimalohcetac ytivitca
era ton .raelc teY ereht smees ot eb emos noitalerroc neewteb eht ecneserp ni plasma fo eht
senimalohcetac and rieht setilobatem and FSC slevel ,namrekcuZ( ;4891 llergeD and ,ygaN .)0991
oslA ti sraeppa that larehpirep noitalumits fo citehtapmys sevren hcihw ylbissop stceffa larehpirep
,ytivitca-HBD at eht emas emit stceffa ytivitca-HBD ni eht lartnec suovren metsys eD( ,rettoP Chanh,
eD temS & eD ,revyrdpeahcS .)6791 fI ew yam etaluceps that teletalp MAO is detalerroc htiw eht
etar fo lartnec enimalohcetac cilobatem ytivitca and that plasma ytivitca-HBD is yltrap detaler ot
lartnec DA ot NA ,noisrevnoc eht stluser dezirammus yb namrekcuZ thgim eb an noitacidni fo eht
tcaf that ni ,SSH ylbissop esuaceb fo decuder noitazilobatem as llew as a rehgih etar fo ,erutpacer NA
and DA serots ni eht lartnec cigrenimalohcetac sevren yam eb ylevitaler rehgih than ni SSL hcihw
seilpmi a retaerg noissimsnart yticapac and yltneuqesnoc osla decnahne ssenevitceffe ni gnisuora and
gnitavitca eht .laudividni SSH osla mees ot eb sretnemgua fo lacitroc dekove slaitnetop saerehw SSL
dnet ot eb srecuder ,namrekcuZ( ;0991 ,namrekcuZ ;4891 ,sakuL .)7891 ,yllaniF noitasnes gnikees
osla smees ot eb detaler ot erom decnuonorp lacigoloisyhpohcysp gnitneiro lamredortcele( ,esnopser
etar-traeh )noitareleced ot tnaveler ilumits llaB( & ,namrekcuZ ;2991 namrekcuZ )4891 nekaT
rehtegot eseht stluser suggest that ni smret fo ruo mgidarap ew nac tcepxe that SSH lliw wohs erom
lasuora noitadrater( fo noitcaer emit retfa eht detcepxenu signal ni eht thgir erehpsimeh task) and erom
noitavitca noitarelecca( fo noitcaer emit retfa eht detcepxenu signal ni eht tfel erehpsimeh task), than
.SSL
ruO mgidarap tuoba eht tnereffid tceffe fo detcepxenu yrotaraperp signals no noitcaer semit ni a
thgir sv tfel erehpsimeh task, swolla us ot distinguish eltrats morf desucof noitnetta sesnopser fi eno
nac redisnoc that eht noitadrater fo eht noitcaer ni eht thgir erehpsimeh task is a eltrats esnopser and
that eht noitarelecca ni eht tfel erehpsimeh task is eht ecneuqesnoc fo a cisahp esaercni ni desucof
noitnetta ro gnitneiro esnopser ot eht tnaveler stimulus. fI noitasnes gnikees stcelfer a lacigoloib
noitisopsiderp sdrawot retaerg ytivitcaer fo htob eht lasuora and noitavitca ,smetsys ew nac tcepxe
eht serocs no eht noitasnes gnikees elacs ot etalerroc ylevitisop htiw noitcaer syaled retfa an
detcepxenu yrotaraperp signal ni eht thgir erehpsimeh task and ylevitagen htiw hcus syaled ni eht tfel
erehpsimeh task.
t r e B e D r e d n a b a r B
6 7 3
e t al.
D O H T E M
Subjects and procedure
e h T t n e m i r e p x e w a s d e t a r o p r o c n i n i a e e r h t k e e w ‘ s k i l l s g n i n i a r t ’ e s r u o c n i e m a g y r o e h t r o f e t a u d a r g
s t n e d u t s f o s c i m o n o c e a t e h t y t i s r e v i n U f o g r u b m i L e h T ( . ) s d n a l r e h t e N e h T e s r u o c d e t r a t s h t i w a
y r a n e l p n o i s s e s n i h c i h w 52 , s t n e d u t s o h w d e b i r c s b u s o t e h t s k i l l s g n i n i a r t , e s r u o c e r e w d e k s a o t l l i f
n i a e r i a n n o i t s e u q o t a s s e s s n o i t a s n e s . g n i k e e s
g n i n r e c n o C e h t e s o p r u p f o e h t , t n e m i r e p x e e w y l n o d e c n u o n n a that t i w a s d e n g i s e d o t n e p e e d r u o
a n d r i e h t g n i d n a t s r e d n u f o r o i v a h e b n i a e m a g . g n i t t e s e h T s t n e d u t s e r e w d e s i m o r p k c a b d e e f n o e h t
r o j a m s g n i d n i f f o e h t h c r a e s e r t c e j o r p r e t f a n o i t e l p m o c f o e h t . e s r u o c e W o s l a d e e t n a r a u g t c i r t s
y t i l a i t n e d i f n o c f o e h t n o i t a m r o f n i d e d i v o r p y b e h t . s e r i a n n o i t s e u q
e h T n o i t c a e r e m i t t n e m i r e p x e k o o t e c a l p n i e h t d r i h t k e e w f o e h t . e s r u o c At that , e m i t 12 s t n e d u t s
y d a e r l a d e t t i u q e h t , e s r u o c g n i v a e l a l a t o t f o 40 s n o i t a v r e s b o 6 2 ( , s e l a m 14 . ) s e l a m e f t I d l u o h s e b
d e n o i t n e m that s t n e d u t s a t e h t y t i s r e v i n U f o g r u b m i L e r a d e w o l l a o t t i u q a e s r u o c a t y n a . t n e m o m t I
i s y l e k i l n u that e s o h t s t n e d u t s e v a g up e s u a c e b f o e h t , t n e m i r e p x e a s e h t t i u q o i t a r s e o d t o n r e f f i d m o r f
e s o h t f o r e h t o . s e s r u o c e h T g n i n i a m e r 5 s e r e w y l m o d n a r a s s i g n e d o t s p u o r g f o 7 - 5 . s n o s r e p e h T s p u o r g
e r e w y l e v i s s e c c u s d e t i v n i o t a m o o r g n i n i a t n o c t h g i e l a n o s r e p s r e t u p m o c ) s C P ( r o f e h t l a t n e m i r e p x e
. n o i s s e s
h c a E S h a d o t m r o f r e p o w t n o i t c a e r e m i t t a s k s n o a : C P a l a i t a p s o u s i v a n d a c i t n a m e s t a s k . h t o B
t a s k s t s i s n o c f o 100 trials. O n h c a e l a i r t a e t i h w r e t t e l i s d e y a l p s i d t s r i f n o e h t t n i o p d i m f o e h t P C
r o t i n o m . n e e r c s e h T r e t t e l s y a t s e r e h t l i t n u e h t d n e f o e h t . l a i r t r e t f A a m o d n a r l a v r e t n i n e e w t e b 5 a n d
1.5 sets a d n o c e s e t i h w r e t t e l i s d e t c e j o r p r e d n u r o e v o b a e h t t s r i f . r e t t e l n I e h t l a i t a p s o u s i v t a s k e h t
S i s d e t c u r t s n i o t s s e r p e h t r a b e c a p s f o e h t d r a o b y e k a s y l k c i u q a s e l b i s s o p y l n o f i e h t d n o c e s r e t t e l
i s d e y a l p s i d w o l e b d n a ( t o n ) e v o b a e h t t s r i f . r e t t e l O n e h t c i t n a m e s t a s k e h t S m u s t t c a e r y l n o f i e h t
d n o c e s r e t t e l s e o d t o n g n o l e b o t e h t e m a s y r o g e t a c a s e h t . t s r i f e h T s e i r o g e t a c e r a s l e w o v a n d
. s t n a n o s n o c e h T d n o c e s r e t t e l s r a e p p a r e d n u r o e v o b a e h t t s r i f g n i d r o c c a o t a m o d n a r e m e h c s a n d i s
f o e h t e m a s c i t n a m e s y r o g e t a c a s e h t t s r i f n i 50 f o e h t 100 trials. e h T m u m i x a m e m i t d e w o l l a o t t c a e r
i s 0.4 t e s n i e h t l a i t a p s o u s i v t a s k a n d 0.5 t e s n i e h t c i t n a m e s t a s k , a s e h t r e t t a l t a s k i s t a h w e m o s e r o m
t l u c i f f i d t h a n e h t . r e m r o f
O n a r e b m u n f o trials y l m o d n a r n e s o h c y b e h t r e t u p m o c m a r g o r p g n i d r o c c a o t a % 0 5 , e t a r a n
n o i t a m a l c x e , k r a m t u o b a 5 m c h g i h a n d 2 m c , e d i w s r a e p p a a t 1 0 ” f o l a u s i v e l g n a t f e l a n d t h g i r f o
e h t r e t t e l n o e h t t n i o p d i m f o e h t . n e e r c s e s e h T y r o t a r a p e r p s i g n a l s e r a d e y a l p s i d a t m o d n a r s t n e m o m
n e e w t e b e h t e m i t f o 0.3 t e s r e t f a e h t y a l p s i d f o e h t t s r i f r e t t e l a n d 0.1 t e s e r o f e b e h t y a l p s i d f o e h t
d n o c e s . r e t t e l y a l p s i D s t n e m o m e r a d e t a r e n e g g n i d r o c c a o t a r a l u g n a t c e r . n o i t u b i r t s i d e h T y a l p s i d f o
e s e h t s i g n a l s l a s t s 0.18 . c e s
r e t f A h c a e l a i r t n i h t o b s e p y t f o t a s k s e h t g n i w o l l o f k c a b d e e f w a s d e d i v o r p o t e h t S s . f I e h t S d i d
t o n t c a e r n e h w a n o i t c a e r w a s , d e r i u q e r o o t ‘ ’ e t a l i s d e y a l p s i d n o e h t . n e e r c s ’ t c e r r o C ‘ s r a e p p a n o e h t
n e e r c s r e t f a , y l e m i t t c e r r o c s n o i t c a e r a n d ’ r o r r e ‘ r e t f a , y l e m i t t c e r r o c n i . s n o i t c a e r
At e h t t e s n o f o e h t t s r i f p u o r g , n o i s s e s e n o f o e h t s r o h t u a y l m o d n a r d e n i m r e t e d h c i h w t a s k e h t S s
h a d o t y a l p . t s r i f m o r F n e h t , s d r a w n o e h t t a s k r e d r o w a s d e t a n r e t l a n i h c a e f o e h t r e h t o x i s . s p u o r g
n e h W a p u o r g d e v i r r a a t e h t l a t n e m i r e p x e , m o o r a P C w a s a s s i g n e d o t h c a e f o e h t S s . , t x e N e h t
r e t n e m i r e p x e e v a g s n o i t c u r t s n i g n i n r e c n o c n e h w a n d w o h o t t c a e r n i e h t t s r i f e p y t f o t a s k u s i n g a
r e t u p m o c o m e d f o e h t n o i t c a e r . e m a g e h T r e t n e m i r e p x e d e t a v i t c a e h t r e t u p m o c m a r g o r p a n d e h t 5 s
d e m r o f r e p e h t t s r i f t a s k . y l s u o e n a t l u m i s r e t f A n o i t e l p m o c f o e h t t s r i f t a s k , e h t e m a s e r u d e c o r p w a s
d e s u r o f e h t d n o c e s t a s k .
e W d e t a v i t o m e h t S s y b s n a e m f o a e s i m o r p that e h t p o t e e r h t s r e m r o f r e p n i h t o b f o e h t t a s k s d l u o w
e v i e c e r a n e k o t r o f c i s u m . s d r o c e r e c n a m r o f r e P k c a b d e e f w a s o s l a d e s u n i r e d r o o t e t a l u m i t s k c i u q
a n d t c e r r o c . s e s n o p s e r y l l a u t c A e h t t n u o m a f o s u o e n o r r e s e s n o p s e r w a s y r e v , w o l % 6 n o . e g a r e v a
e c n a m r o f r e P w a s d e t a u l a v e y b g n i t a l u c l a c e h t o i t a r f o e h t r e b m u n f o t c e r r o c s n o i t c a e r d e d i v i d y b e h t
e g a r e v a y c n e t a l f o e s e h t . s n o i t c a e r s i h T o i t a r s e t a l u m i t s h t o b t c e r r o c a n d y d e e p s . s n o i t c a e r e W o s l a
d e l a e p p a o t e h t l a i c o s n o i t a v i t o m r o f e g i t s e r p y b g n i l l e t e h t S s that e h t e c n a m r o f r e p g n i k n a r d l u o w
e b d e c n u o n n a n i c i l b u p n i a l a n i f y r a n e l p n o i s s e s a t e h t d n e f o e h t . e s r u o c
,lasuorA noitavitca dna noitasnes gnikees 377
Measures
ehT dellac-os tsjiletfeohebsgninnapS‘ ’)LBS( depoleved yb jieF and nav neliuZ )4891( was desu
ot assess noitasnes .gnikees ehT LBS is a hctuD noitatpada fo eht naciremA noitasneS gnikeeS ,elacS
erom yllacificeps eht mrof-SSS V ,namrekcuZ( .)4891 ehT LBS stsisnoc fo 67 ,smeti 51 fo hcihw era
dengised ot erusaem noitasnes gnikees )SS( eht( rehto smeti era rellif .)smeti ehT stnednopser era deksa
ot etacidni no a evif tniop elacs ot what tnetxe yeht eerga htiw hcae fo 5 1 stnemetats 1( = ylgnorts
,eergasid 2 = yletaredom ,eergasid 3 = t’nod ,wonk 4 = yletaredom ,eerga 5 = ylgnorts .)eerga ehT
serocs fo 15 smeti era ,desrever os that a hgih erocs sdnopserroc htiw hgih noitasnes .gnikees
gniognO hcraeser suggests that noitasnes gnikees is yltrap a lanoisnemid-itlum ytilanosrep
tcurtsnoc dezilanoitarepo yb selacs-bus ,htiw ,ylbissop tnereffid cipytoneg snoitadnuof ,namrekcuZ(
1979a, b, ;c jieF and nav ,neliuZ ;4891 ,jieF ,ekebelrO madneznaG & nav ,neliuZ .)5891 ehT gniwollof
snoisnemid nac eb dehsiugnitsid ,namrekcuZ( 1979a, b, ;c jieF & nav ,neliuZ :)4891
)1( llirhT and erutnevdA gnikeeS :)SAT( a erised ot egagne ni lacisyhp seitivitca
gnivlovni emos lacisyhp regnad ro risk.
)2( ecneirepxE gnikeeS :)SE( eht erised ot kees levon secneirepxe hguorht ,levart
,cisum ,tra and gnimrofnocnon .elyts-efil
)3( noitibihnisiD :)SID( eht deen ot kees esaeler ni detibihninu laicos seitivitca htiw
ro tuohtiw eht aid fo .lohocla
)4( moderoB ytilibitpecsuS :)SB( an noisreva rof evititeper secneirepxe fo yna ,dnik
enituor ,krow ro elbatciderp .elpoep
ehT LBS is dengised ot assess htob eseht ruof snoisnemid-bus fo noitasnes gnikees as llew as a latot
noitasnes gnikees ehT.erocs rebmun fo smeti ot assess ,SAT ,SE SB and SID era 12, 14, 13 and 12,
.ylevitcepser serocS rof eseht snoisnemid-bus era deniatbo yb summing eht sesnopser ot eht laudividni
.smeti suhT eht mumixam serocs rof ,SAT ,SE SB and SID era 60,70, 65 and 60, .ylevitcepser ehT
latot noitasnes gnikees erocs is eht sum fo eht segareva fo eht ruof snoisnemid-bus muminim( 4,
mumixam )02 ees( jieF and nav ,neliuZ .)4891 ehT gniwollof stnemetats era selpmaxe fo emos
:smeti-LBS I“ dluow ekil ot nrael ot ”ylf ,)meti-SAT( I“ ekil ot rednaw dnuora ni a egnarts ytic no
ym ,nwo neve fi ti snaem gnitteg ”tsol ,)meti-SE( I“ yllausu t’nod yojne a eivom ro yalp nehw I nac
tciderp what lliw neppah ni ”ecnavda )meti-SB( and I“ ekil ,dliw detibihninu ”seitrap .)meti-SID(
evisnetxE hcraeser gnoma gnuoy adults suggests that eht LBS is a elbailer and dilav tnemerusaem
tnemurtsni jieF( & nav ,neliuZ .)4891 ehT hcabnorC stneiciffeoc ni eht tneserp yduts era 0.64 rof
a
SAT n( = 40, rebmun fo smeti ,)21 0.71 rof SE n( = 39, rebmun fo smeti ,)41 0.67 rof SB n( = 40,
rebmun fo smeti )31 0.69 rof SID n( = 40, rebmun fo smeti )21 and 0.78 rof latot noitasnes gnikees
SS n( = 39, rebmun fo smeti .)15 esehT seulav era tahwemos rewol than esoht detroper yb jieF and
nav neliuZ ,)4891( tub llew evoba eht rewol stimil fo ,ytilibatpecca yllareneg deredisnoc ot eb dnuora
06.0-05.0 ,yllannuN( .)8791
ehT tnedneped selbairav era eht :gniwollof
owT serusaem fo
l arousal by the preparatory signals in the visuospatial task (right
hemisphere task):
fRA = eht egareva noitcaer emit ni tcerroc trials nierehw a yrotaraperp signal is deyalpsid minus
eht egareva noitcaer emit ni trials tuohtiw yrotaraperp signal, dedivid yb eht egareva fo eseht egareva
noitcaer ,semit deilpitlum yb 100. ehT noitcaer semit dedulcni ni eht snoitaluclac era all tcerroc trials
etal( snoitcaer )dedulcni nierehw eht S is deriuqer ot .tcaer fRA is detaluclac no eht basis fo eht tsrif
50 trials.
AR1 = is ralimis ot fRA tub desab no eht last 50 trials.
ehT noitcnitsid neewteb fRA and AR1 is edam ni redro ot kcehc rehtehw eht 5s wohs noitautibah
ot eht yrotaraperp signals. fI eht deyaled snoitcaer retfa a yrotaraperp signal era yllaer eht tceffe fo
a cisahp esnopser yb eht cigrenerdaron lasuora metsys gnisuac a eltrats ,noitcaer neht ew dluohs eb
elba ot evresbo a rellams tceffe ni eht last 50 trials. sihT noitatcepxe is desab no lacigoloruen ecnedive
detic[ ni marbirP & ssenniuGcM ])5791( gniwohs that cigrenerdaron sevren gnidnecsa morf eht sucol
sueluroec esaerced gnirif setar ni esnopser ot detaeper .noitalumits ,eromrehtruF ereht is erom than
t r e B e D r e d n a b a r B t e
8 7 3
al.
a b u n d a n t e c n e d i v e o o t ( h c u m o t ) e t i c e c n i s s ’ v o l o k o S y l r a e s t n e m i r e p x e that l a c i g o l o i s y h p o h c y s p
g n i t n e i r o s e s n o p s e r d e l l o r t n o c y b e h t c i t e h t a p m y s s u o v r e n m e t s y s e s a e r c e d h t i w s t i m u l u s . n o i t i t e p e r
a o w T s e r u s a e m f o
activation by the preparatory s i g n a l s in the s e m a n t i c task ( l e f t
h e m i s p h e r e task):
f C A i s s u o g o l a n a o t f R A h t i w t h i s e c n e r e f f i d that e h t e g a r e v a n o i t c a e r s e m i t n i e h t a l u m r o f n r e c n o c
e h t t s r i f 50 trials n i e h t c i t n a m e s t a s k . e h T r e w o l f C A e h t r e t a e r g e h t e e r g e d f o e v i t a t u p c i g r e n i m a p o d
. n o i t a v i t c a
A C 1 i s r a l i m i s o t f C A t u b d e s a b n o e h t l a s t 50 trials.
e c n i S c i g r e n i m a p o d n o i t a v i t c a s e o d t o n y l p m i n o i t a u t i b a h o t , i l u m i t s n o e h t y r a r t n o c c i g r e n i m a p o d
s e v r e n m e e s o t e s a e r c n i g n i r i f s e t a r r e t f a d e t a e p e r n o i t a l u m i t s e c n e d i v e [ d e t i c n i m a r b i r P &
s s e n n i u G c M J ) 5 7 9 1 ( e w o s l a t c e p x e A C 1 o t e b r e w o l t h a n f C A r o n i r e h t o s d r o w e w t c e p x e e h t c i s a h p
n o i t a v i t c a y b e h t y r o t a r a p e r p i l u m i t s o t e s a e r c n i h t i w d e t a e p e r . e r u s o p x e
e r u g i F 1 s t c i p e d e h t l a u t p e c n o c l e d o m a n d w o h e h t s e s e h t o p y h t i f . n i
Analysis of data
v o m i m S - v o r o g o m l o K s t s e t r o f s s e n d o o g f o t i f d e t a c i d n i that all e h t s e l b a i r a v d e s u n i e h t s e s y l a n a
n a c e b d e r e d i s n o c a s d e v i r e d m o r f a l a m r o n . n o i t u b i r t s i d y l t n e u q e s n o C c i r t e m a r a p l a c i t s i t a t s s t s e t e r a
. d e s u e h T s i s y l a n a s t s i s n o c f o s t s e t f o e h t g n i w o l l o f g n i k r o w s e s e h t o p y h h c i h w w o l l o f m o r f r u o
l a c i t e r o e h t s n o i s s u c s i d n i e h t y r o t c u d o r t n i . n o i t c e s
f R A a n d A R 1 e r a y l e v i t i s o p d e t a l e r r o c h t i w . S S
Hypothesis 1.
f C A a n d A C 1 e r a y l e v i t a g e n d e t a l e r r o c h t i w . S S n a m r e k c u Z , a 9 7 9 1 ( b, , c ) 4 8 9 1
Hypothesis 2.
d e t s e g g u s that s e c i d n i f o c i g r e n e r d a r o n a n d c i g r e n i m a p o d y t i v i t c a t h g i m e b d e t a l e r y l t n e r e f f i d o t
t n e r e f f i d s e l a c s b u s f o e h t n o i t a s n e s g n i k e e s . e l a c s f I t h i s n o i t s e g g u s i s e u r t a n d g n i r r e f n i m o r f t n e r r u c
e g d e l w o n k f o e h t l a c i g o l o h c y s p s t c e f f e f o e h t n o i t a v i t c a r o n o i t i b i h n i f o e h t N A a n d D A s m e t s y s y b
s u o i r a v d r u g s e n o i s d e n i l c n i o t t c e p x e that c i s a h p l a s u o r a y l b a b o r p l l i w e b d e t a l e r e r o m y l g n o r t s o t
e h t m o d e r o b y t i l i b i t p e c s u s a n d e h t n o i t i b i h n i s i d s e l a c s b u s e l i h w c i s a h p n o i t a v i t c a d l u o c e b d e t a l e r
e r o m y l g n o r t s o t e h t l l i r h t a n d e r u t n e v d a a n d e c n e i r e p x e g n i k e e s . s e l a c s b u s , r e v e w o H e w r e f e r p o t
r e d i s n o c e s e h t s n o i t a t c e p x e a s . y r o t a r o l p x e
f R A i s r e h g i h t h a n f C A a n d A R 1 i s r e h g i h t h a n A C l .
Hypothesis 3.
A R 1 i s r e w o l t h a n . f R A
Hypothesis 4.
A C 1 i s r e w o l t h a n . f C A
Hypothesis 5.
s i s e h t o p y H 3 i s , y l t c e r i d a n d 4 a n d 5 e r a y l t c e r i d n i d e r r e f n i m o r f r u o c i s a b . m g i d a r a p r e b m e m e R
that e h t r e h g i h f R A a n d A R 1 e h t r e h g i h e h t c i s a h p l a s u o r a y b e h t y r o t a r a p e r p s i g n a l n i e h t l a i t a p s o u s i v
t h g i r ( ) e r e h p s i m e h t a s k i s d e s o p p u s o t , e b a n d that e h t r e w o l f C A a n d A C 1 e h t r e h g i h e h t c i s a h p
n o i t a v i t c a y b e h t y r o t a r a p e r p s i g n a l n i e h t c i t n a m e s t f e l ( ) e r e h p s i m e h t a s k i s a s s u m e d o t . e b r u O t s r i f
n o i t a l u m r o f f o e h t m g i d a r a p w a s e h t : g n i w o l l o f
“ . . . , f I . j u s t e r o f e b a s t i m u l u s , s r a e p p a a t r o h s y r o t a r a p e r p s t i m u l u s i s d e y a l p s i d e h t n i a r b
y l n e d d u s s e v i e c e r a y r a t n e m e l p p u s g n i s s e c o r p - n o i t a m r o f n i . d a o l f I e w a s s u m e that e h t
e m a s e r e h p s i m e h h c i h w y l i r a m i r p s d n e t t a o t e h t t a s k y b ( h c i h w e w n a e m that t i i s
e l b i s n o p s e r r o f g n i d o c n e e h t e s n o p s e r s t i m u l u s ) o s l a s e d o c n e e h t y r o t a r a p e r p s i g n a l ,
n e h t e w n a c o s l a a s s u m e that e h t t c e f f e f o t h i s y r o t a r a p e r p s i g n a l n o p u t a s k e c n a m r o f r e p
s t c e l f e r e h t t c e f f e f o a n e d d u s e s a e r c n i n i y t i v i t c a f o e h t e r e h p s i m e h g n i m r o f r e p e h t t a s k .
s i h T i s a s s u m e d o t t l u s e r n i r e h t i e d e s a e r c n i n o i t a v i t c a n e h w e h t t f e l e r e h p s i m e h s l o r t n o c
t a s k e c n a m r o f r e p r o n i d e s a e r c n i l a s u o r a n e h w e h t t h g i r e r e h p s i m e h i s n i . l o r t n o c t u B
w o h n a c s e s a e r c n i n i l a s u o r a s u s r e v n o i t a v i t c a e b ? d e t c e t e d m o r F r i e h t s i s y l a n a f o
l a r o i v a h e b o r u e n d a t a m a r b i r P a n d s s e n n i u G c M ) 5 7 9 1 ( e d u l c n o c that e h t l a s u o r a m e t s y s
s t c a n o s m s i n a h c e m - p o t s f o r o i v a h e b e l i h w e h t n o i t a v i t c a m e t s y s s t c a n o
, s m s i n a h c e m - o g . e . i t i y l u r t . s e t a v i t c a f I t h i s e r e w e h t e s a c n i h u m a n , s t c e j b u s n o i t a v i t c a
d l u o h s e t a t i l i c a f s e s n o p s e r a n d l a s u o r a d l u o h s ” . t i b i h n i e D ( , r e d n a b a r B 1988, pp.
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0 5 ; s l a i r t , f R A l a s u o r a s e r o c s n i t s r i f 0 5 ; s l a i r t , I R A l a s u o r a s e r o c s
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e h T e r o m e v i t a g e n e h t n o i t a v i t c a s e r o c s e h t r e g r a l e h t e e r g e d f o
. n o i t a v i t c a
n o i t a t i l i c a F f o a e s n o p s e r shows n i a r e t r o h s n o i t c a e r e m i t a n d n o i t i b i h n i s t l u s e r n i a d e y a l e d n o i t c a e r
. e m i t
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e l b a T 1 s t c i p e d e h t s n o i t a l e r r o c n e e w t e b e h t t n e d n e p e d s e l b a i r a v a n d e h t s e r o c s n o e h t s e l a c s b u s
a n d e h t l a t o t e r o c s n o e h t n o i t a s n e s g n i k e e s . e r i a n n o i t s e u q e h T r e b m u n f o s e s a c n o h c i h w e h t
s n o i t a l e r r o c e r a d e s a b i s less t h a n 40 r o r e f f i d e s u a c e b f o e s i w r i a p m i s s i n g d a t a .
m o r F e h t m o t t o b w o r t i i s t n e r a p p a that y l n o s i s e h t o p y H 1 i s e l b i s u a l p a n d that s i s e h t o p y H 2 m u s t
e b . d e t c e j e r s i h T s m e e s o t y l p m i that n o i t a s n e s g n i k e e s i s y l n o d e t a l e r o t y t i l i b a s u o r a f o e h t
c i g r e n e r d a r o n l a s u o r a . m e t s y s f I e w k o o l a t e h t s t l u s e r f o e h t s n o i t a l e r r o c f o e h t t n e d n e p e d s e l b a i r a v
h t i w e h t s e r o c s n o e h t , s e l a c s b u s S A T a n d , S E t p e c x e r o f e n o y l e c r a c s y l l a c i t s i t a t s t n a c i f i n g i s
, t n e i c i f f e o c o d t o n m e e s o t w o h s y n a p i h s n o i t a l e r h t i w r e h t i e l a s u o r a r o , n o i t a v i t c a r o e r o m y l l a c i f i c e p s
h t i w e h t c i s a h p s n o i t c a e r o t d e t c e p x e n u i l u m i t s f o e s e h t . s m e t s y s , S I D t u b t s o m y l t n a c i f i n g i s S B
m o d e r o b ( ) y t i l i b i t p e c s u s i s y l r a e l c d e t a l e r o t e h t c i s a h p l a s u o r a s e r u s a e m n i e h t d e t c e p x e . e s n e s e h T
p i h s n o i t a l e r i s r e k a e w h t i w A R 1 t h a n h t i w f R A h c i h w i s e l b i t a p m o c h t i w e h t a e d i that e h t e r o m
d e c n u o n o r p l a s u o r a s n o i t c a e r f o m o d e r o b e l b i t p e c s u s S s y l l a u d a r g r a e w f f o e s u a c e b f o , n o i t a u t i b a h
e n O n o i t a l e r r o c h t i w e h t c i s a h p n o i t a v i t c a s e r u s a e m s e h c a e r a s u o i r e s e e r g e d f o e c n a c i f i n g i s , y l e m a n
e h t e v i t i s o p n o i t a l e r r o c n e e w t e b S B a n d A C l . s i h T t l u s e r i s n i n o i t c i d a r t n o c h t i w s i s e h t o p y H 2 h c i h w
y d a e r l a h a d o t e b . d e t c e j e r t u B y h w t h i s d e t c e p x e n u ? g n i d n i f n I , t c e f f e t i s m e e s o t e t a r o b o r r o c e h t
n o i s u l c n o c that y l n o e h t g n i n o i t c n u f f o e h t l a s u o r a m e t s y s i s d e t a l e r o t n o i t a s n e s . g n i k e e s e h T t s o m
e l p m i s n o i t a n a l p x e i s that n e v e n e h w a t c e j b u s i s g n i m r o f r e p a t f e l e r e h p s i m e h t a s k , l a s u o r a y a m
y l l a u d a r g m l e h w r e v o , n o i t a v i t c a a n d that m o d e r o b e l b i t p e c s u s s n o s r e p e r a e r o m e l b a n e m a o t t h i s
. n o n e m o n e h p n I n o m m o c e s n e s e g a u g n a l e n o d l u o c e b i r c s e d t h i s n o n e m o n e h p a s . s w o l l o f y l l a i t i n I
m o d e r o b s e l b i t p e c s u s a n d s n o s r e p o h w e r a less , o s e r a y l l a u q e d e t a v i t c a y b e h t t n a v e l e r r i y r o t a r a p e r p
i l u m i t s n i e h t t f e l e r e h p s i m e h t a s k . e t o N ( that e h t n o i t a l e r r o c n e e w t e b S B a n d f C A s l a u q e ) . 5 0 . 0
, r e v e w o H r e t f a a e l i h w e h t l a r u t a n y c n e d n e t f o e h t m o d e r o b s e l b i t p e c s u s o t k e e s n o i t a l u m i t s s e m o c r e v o
e h t y c n e d n e t o t t r e v i d n o i t n e t t a m o r f e h t t n a v e l e r r i y r o t a r a p e r p i l u m i t s a n d y e h t e m o c e b y l l a u d a r g
e r o m d e s u o r a y b . m e h t
e l b a T 2 s y a l p s i d e h t s t l u s e r f o d e r i a p s t s e t - f s n o s i r a p m o c ( e r a a r t n i ) t c e j b u s d e d e e n o t t s e t
s e s e h t o p y H 3,4 a n d 5. e h T r e b m u n f o s e s a c s t n u o m a y l n o o t 39 e s u a c e b f o e h t t c a f that r o f e n o S
e r e h t e r e w t o n h g u o n e s n o i t a v r e s b o n i e n o l a t n e m i r e p x e n o i t i d n o c o t e t a l u c l a c a e u l a v r o f e h t
t n e d n e p e d . e l b a i r a v
, l a s u o r A n o i t a v i t c a d n a n o i t a s n e s g n i k e e s 381
e l b a T . 2 d e r i a P s e l p m a s d e d i s - e n o s t s e t - F ‘ n e e w t e b e g a r e v a s e r o c s n o e h t
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: s e t o N , f C A n o i t a v i t c a s e r o c s n i t s r i f 0 5 ; s l a i r t . I C A n o i t a v i t c a s e r o c s n i
t s a l O S ; s l a i r t , f R A l a s u o r a s e r o c s n i t s r i f O S ; s l a i r t , I R A l a s u o r a s e r o c s
n i t s a l 0 5 . s l a i r t
e h T e r o m e v i t a g e n e h t n o i t a v i t c a s e r o c s e h t r e g r a l e h t e e r g e d f o . n o i t a v i t c a
e n O n a c e e s that f R A a n d f C A e r a y l l a c i t s i t a t s y l t n a c i f i n g i s t n e r e f f i d n i e h t d e t c e p x e . n o i t c e r i d o s l A
e h t s i g n s f o e h t s e g a r e v a e r a n i t n e m e e r g a h t i w r u o . m g i d a r a p n I e h t t h g i r e r e h p s i m e h t a s k a n
d e t c e p x e n u s t i m u l u s s m e e s o t t i b i h n i e h t t n e u q e s b u s n o i t c a e r e l i h w e h t y r a r t n o c s m e e s o t e b e u r t n i
e h t t f e l e r e h p s i m e h t a s k . e h T e c n e r e f f i d n e e w t e b A R 1 a n d A C 1 d e s a b ( n o e h t l a s t 50 ) s l a i r t i s t o n
t n a c i f i n g i s . e r o m y n a s i h T s m e e s o t e b e u d o t e h t d e n i b m o c t c e f f e f o n o i t a u t i b a h f o e h t l a s u o r a m e t s y s
a n d f o e h t d e s a e r c n i n o i t a v i t c a n i e h t l a s t 50 trials. e h T e g a r e v a l a s u o r a s n o i t c a e r e s a e r c e d m o r f 2.78
o t - 1.68, a y l h g i h t n a c i f i n g i s . e c n e r e f f i d e h T e g a r e v a l a s u o r a s n o i t c a e r n i e h t l a s t 50 trials, , r e v e w o h
e r a t o n y l l a c i t s i t a t s y l t n a c i f i n g i s t n e r e f f i d m o r f o r e z r ( = . ) 4 9 . 0 n o i t a v i t c A n o e h t y r a r t n o c s e s a e r c n i
m o r f - 0.74 ) f C A ( o t - 2.57 ) l C A ( t ( = 1.94, P < . ) 6 6 0 . 0 r e b m e m e R that e h t r e w o l f C A a n d A C 1
e h t r e t a e r g e h t n o i t a v i t c a i s d e s o p p u s o t . e b A C l , r e v e w o h i s y l t n a c i f i n g i s t n e r e f f i d m o r f o r e z t ( = 1.94,
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y r o t a r a p e r p s i g n a l s y l l a u d a r g e m o c e b less d e s u o r a y b , m e h t a n d e r o m . d e t a v i t c a e h T d e s a e r c e d
, l a s u o r a r e v e w o h s m e e s o t e b r e g n o r t s t h a n e h t d e s a e r c n i n o i t a v i t c a o s that e h t e c n e r e f f i d n e e w t e b
l a s u o r a ) I R A ( a n d n o i t a v i t c a ) l C A ( n i e h t l a s t 50 trials i s t o n t n a c i f i n g i s . e r o m y n a , o S e w n a c e d u l c n o c
that s i s e h t o p y H 3 i s y l l a i t r a p . d e t a d i l a v s i s e h t o p y H 4 i s y l r a e l c d e t a d i l a v a n d s i s e h t o p y H 5 s m e e s o t
e b d i l a v t u b e h t e c n e d i v e i s t o n y r e v . g n o r t s e h T t s a e l e n o n a c , y a s , r e v e w o h i s that all e h t d a t a e r a
e l b i t a p m o c h t i w e h t c i s a b m g i d a r a p h c i h w s t a t e s that d e t c e p x e n u y r o t a r a p e r p s i g n a l s n i a t h g i r
e r e h p s i m e h t a s k e r a e l b i s n o p s e r r o f c i s a h p l a s u o r a s n o i t c a e r h c i h w y l l a u d a r g r a e w f f o a n d r o f
y l l a u d a r g g n i s a e r c n i c i s a h p n o i t a v i t c a s n o i t c a e r n i a t f e l e r e h p s i m e h t a s k . e h T g n i s a e r c e d l a s u o r a a n d
g n i s a e r c n i n o i t a v i t c a e r a e h t e c n e u q e s n o c f o e h t l a c i p y t l a n o i t c n u f s c i t s i r e t c a r a h c f o e h t c i g r e n e r d a r o n
a n d c i g r e n i m a p o d l a r t n e c n o i t a v i t c a s m e t s y s h c i h w y l e v i t c e p s e r t c e f f e e h t l a s u o r a a n d n o i t a v i t c a f o
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