PUNISHMENT-RELATED MEMORY-GUIDED ATTENTION:
NEURAL DYNAMICS OF PERCEPTUAL MODULATION
Au ho s: Samuel Suá ez-Suá ez, Soco o Rod íguez Holguín, Fe nando Cada ei a,
Anna Ch is ina Nob e, Sonia Doallo
This is he pee e iewed e sion o he ollowing a icle: Suá ez-Suá ez S; Rod íguez
Holguín S; Cada ei a F; Nob e AC; Doallo S (2019). Punishmen - ela ed memo y-
guided a en ion: Neu al dynamics o pe cep ual modula ion. Co ex, 115, 231-245. doi:
10.1016/j.co ex.2019.01.029.
This a icle may be used o non-comme cial pu poses in acco dance wi h Else ie
Te ms and Condi ions o Use o Sel -A chi ed Ve sions.
2
Punishmen - ela ed memo y-guided a en ion:
Neu al dynamics o pe cep ual modula ion
Samuel Suá ez-Suá eza, Soco o Rod íguez Holguína, Fe nando Cada ei aa,
Anna Ch is ina Nob eb, Sonia Doalloa
a Depa men o Clinical Psychology and Psychobiology, Uni e si y o San iago de
Compos ela, San iago de Compos ela, Spain
b Depa men o Expe imen al Psychology and Ox o d Cen e o Human B ain
Ac i i y, Depa men o Psychia y, Wellcome Cen e o In eg a i e Neu oimaging,
Uni e si y o Ox o d, Ox o d, Uni ed Kingdom
Co esponding au ho :
Sonia Doallo
Depa men o Clinical Psychology and Psychobiology
Uni e si y o San iago de Compos ela
Campus Vida, s/n, 15782 San iago de Compos ela
Galicia-Spain
Telephone: +34 881813916
E-mail: [email p o ec ed]
Decla a ions o in e es : none.
Pos -p in ( inal d a pos - e e eeing)
3
ABSTRACT
Remembe ing he ou comes o pas expe iences allows us o gene a e u u e
expec a ions and shape selec ion in he long- e m. A g owing numbe o s udies has
shown ha lea ned posi i e ewa d alues impac spa ial memo y-based a en ional
biases on pe cep ion. Howe e , whe he memo y-d i en a en ional biases ex end o
punishmen - ela ed alues has ecei ed compa a i ely less a en ion. He e, we
manipula ed whe he ecen spa ial con ex ual memo ies became associa ed wi h
success ul a oidance o punishmen (po en ial mone a y loss). Beha iou al and
elec ophysiological measu es we e collec ed om 27 pa icipan s du ing a subsequen
memo y-based a en ion ask, in which we es ed o he e ec o punishmen a oidance
associa ions. Punishmen a oidance signi ican ly ampli ied e ec s o spa ial con ex ual
memo ies on isual sea ch p ocesses wi hin na u al scenes. Compa ed o non-associa ed
scenes, con ex ual memo ies pai ed wi h punishmen a oidance lead o as e esponses
o a ge s p esen ed a emembe ed loca ions. E en - ela ed po en ials elici ed by a ge
s imuli e ealed ha acqui ed mo i a ional alue o speci ic spa ial loca ions, by i ue
o hei associa ion wi h pas a oidance o punishmen , dynamically a ec ed neu al
signa u es o ea ly isual p ocessing (indexed by la ge P1 and ea lie N1 po en ials)
and a ge selec ion (as indica ed by educed N2pc po en ials). The p esen esul s
ex end ou unde s anding o how memo y, a en ion, and punishmen - ela ed
mechanisms in e ac o op imize pe cep ual decision in eal wo ld en i onmen s.
Keywo ds: Long- e m memo y, isuospa ial a en ion, punishmen , e en - ela ed
po en ials
Pos -p in ( inal d a pos - e e eeing)
4
1. In oduc ion
P e ious esea ch has demons a ed he abili y o spa ial con ex ual long- e m
memo y (LTM) o guide a en ion wi hin scenes, and o modula e neu al signa u es o
ea ly pe cep ual analysis (Summe ield, Rao, Ga side, & Nob e, 2011) and selec ion
(Pa ai, Doallo, & Nob e, 2012) o ele an objec s p esen ed a memo ized loca ions.
This memo y-d i en modula ion o isual p ocessing engages ac i i y in he on al-
pa ie al ne wo k o isual-spa ial o ien ing in conce wi h ac i a ion in o he b ain
egions implica ed in e ie al o objec loca ions wi hin speci ic con ex s (e.g.,
hippocampus) (S okes, A he on, Pa ai, & Nob e, 2012; Summe ield, Lepsien,
Gi elman, Mesulam, & Nob e, 2006). Mo e ecen wo k by Rosen and colleagues shows
ha LTM-guided isuospa ial a en ion ec ui s a ne wo k spanning pa ie al co ical
a eas (la e al in apa ie al sulcus, pos e io p ecuneus and pos e io callosal sulcus) and
subco ical egions ( he cauda e head, mediodo sal halamus and lobule VI/C us I o
ce ebellum) (Rosen, S e n, De aney, & Some s, 2017; Rosen, S e n, Michalka,
De aney, & Some s, 2016). A g owing numbe o s udies ha e also s a ed add essing
he impo an ques ion o whe he and how lea ned ewa d alues impac spa ial
memo y-based a en ional biases on pe cep ion. Recen wo k has shown ha ewa d-
ou come associa ions boos memo ies and a en ional p io i y o speci ic spa ial
loca ions (Chelazzi e al., 2014; Hickey, Chelazzi, & Theeuwes, 2014; see also
Ande son, 2015; Pollmann, Es ocino a, Somme , Chelazzi, & Zinke, 2016), and
dynamically impac di e en le els o isual co ical p ocessing o a ge s p esen ed a
p e iously ewa ded loca ions (Doallo, Pa ai, & Nob e, 2013). These e ec s o ewa d
his o y on a en ional p io i y o space add o accumula ing e idence indica ing ha
ewa d-associa ions modula e bo om-up and op-down a en ional p io i y o s imuli
ea u es and objec ca ego ies (e.g. Ande son, Lau en , & Yan is, 2011; Della Libe a &
Pos -p in ( inal d a pos - e e eeing)
5
Chelazzi, 2006, 2009; Donohue e al., 2016; Ha is e al., 2016; Hickey, Chelazzi, &
Theeuwes, 2010; Hickey, Kaise , & Peelen, 2015; Hickey & Peelen, 2015; Kiss, D i e ,
& Eime , 2009; o e iews see Ande son, 2013; Chelazzi, Pe la o, San and ea, & Della
Libe a, 2013; Failing & Theeuwes, 2018; Pessoa, 2015; Vuilleumie , 2015).
An impo an emaining ques ion is whe he punishmen - ela ed associa ions
also in luence memo y-d i en spa ial-a en ional biases. In con as o he inc easing
numbe o s udies examining posi i e ewa d- ela ed e ec s on hippocampus-dependen
memo y and isuospa ial a en ion, less is known abou whe he simila e ec s a e
obse ed when spa ial loca ions acqui e mo i a ional alue h ough he associa ion wi h
an a e si e ou come (o i s a oidance). The complexi y o his issue inc eases i we
conside ha he ques ion o whe he ewa d and punishmen p ocessing ely on a
common o on dis inc neu al sys ems is s ill deba ed (Bissone e, Gen y, Padmala,
Pessoa, & Roesch, 2014; B ooks & Be ns, 2013; Liu, Hai s on, Sch ie , & Fan, 2011).
S imuli ha unde go a e si e condi ioning ha e been shown o exhibi enhanced
p ocessing unde challenging condi ions (Padmala & Pessoa, 2008), o cap u e a en ion
au oma ically du ing isual sea ch (Schmid , Belopolsky, & Theeuwes, 2015), and o
coun e ac he a en ional blink (Lim, Padmala, & Pessoa, 2009). G owing e idence
sugges s ha s imulus ea u es and objec s associa ed wi h a mone a y loss h ough
associa i e lea ning also a ec a en ional and pe cep ual p ocesses. Face s imuli
associa ed o he eceip o a mone a y loss a e mo e likely o be ecognized han o he
aces in a subsequen a en ional blink ask (Raymond & O’B ien, 2009) and ecei e
enhanced isual p ocessing in a masked ecogni ion ask (O’B ien & Raymond, 2012).
S onge eo ien ing p ocesses o ab up -onse (exogenous) colo ed cues linked o a
mone a y punishmen ha e also been epo ed (Bucke & Theeuwes, 2016; bu see
Ru he o d, O'B ien, & Raymond, 2010), as well as inc eased a en ional cap u e by
Pos -p in ( inal d a pos - e e eeing)

6
colo ed single on dis ac o s associa ed o e asion o a mone a y loss in a isual sea ch
ask (Wen u a, Mülle , & Ro he mund, 2014). Recen da a e eal ha pe cep ual
ea u es associa ed o eceip o punishmen boos p ima y isual co ex esponses (as
e lec ed by he highe ampli ude o he C1 po en ial), ela i e o ewa d-associa ed and
neu al ones (Rossi e al., 2017), al hough e idence o an enhanced ep esen a ion in
isual en al a eas o gain- e sus loss-associa ed s imuli has been ound o symbol
cues du ing an economic decision-making ask (San Ma ín, Appelbaum, Hue el, &
Woldo , 2014) and o objec ca ego ies p esen ed in images o na u al scenes
(Ba ba o, Peelen, & Hickey, 2017).
Though e idence o modula o y e ec s o punishmen associa ions in isual
pe cep ual and a en ional asks, whe he i can be obse ed o speci ic loca ions in
clu e ed na u alis ic con ex s emains an open ques ion; u he mo e, i is unclea
whe he i has a las ing e ec ha can modula e LTM-based a en ion.
He e, we aimed o shed ligh on whe he and how punishmen - ela ed
mechanisms in luence he abili y o LTM o guide pe cep ion in na u alis ic con ex s.
We used a simila expe imen al app oach as in ou p e ious s udy (Doallo e al., 2013)
o disen angle whe he spa ial expec a ions om LTM inco po a e alue signals
speci ically ela ed o pas a oidance o an a e si e ou come o magni y expe ience-
based biases upon pe cep ual decisions on ele an objec s in clu e ed scenes. E en -
ela ed po en ials (ERPs) we e eco ded o e eal he ime cou se o e which
punishmen -associa ed memo y-guided a en ion exe s i s e ec s on he ongoing neu al
ac i i y in isual co ical a eas. We analysed he P1, N1 and N2pc componen s. These
po en ials e lec isual p ocessing (P1 and N1) and a ge selec ion (N2pc). The P1
po en ial, a posi i e de lec ion peaking abou 100 ms a e s imulus p esen a ion, is
modula ed by isuospa ial a en ion and hough o e lec a senso y gain con ol
Pos -p in ( inal d a pos - e e eeing)
7
mechanism wi hin ex as ia e isual co ex (Heinze e al., 1994; Hillya d, Vogel, &
Luck, 1998; Ma ínez e al., 1999). The N1 po en ial is a nega i e de lec ion,
subsequen o he P1, which is also modula ed by a en ion. The N1 a en ion e ec is
belie ed o index a op-down modula ion o disc imina i e p ocessing in a eas o he
en al isual s eam (Hop , Vogel, Woodman, Heinze, & Luck, 2002; Luck, 1995;
Vogel & Luck, 2000). The N2pc is an enhanced nega i i y a pos e io elec ode si es
con ala e al o he loca ion o an a ended a ge , ypically eme ging 200-300 ms a e
a ge onse . I is hough o e lec a en ional selec ion o a a ge among dis ac e s
(Eime , 1996; Luck & Hillya d, 1994) and is gene a ed in pa ie al and in e io occipi al-
empo al co ical a eas (Hop e al., 2000). These ERP componen s ha e been
p e iously shown o be modula ed by memo y-guided o ien ing (Doallo e al., 2013;
Pa ai e al., 2012; Summe ield e al., 2011) and ha e also been used o in es iga e
alue-based modula ion o pe cep ual p ocessing and a en ional selec ion in o he ypes
o asks (Donohue e al., 2016; Ha is e al., 2016; Hickey e al., 2010; I hipu ipa , Cha,
Rangsipa , & Se ences, 2015; McLean & Giesb ech , 2015; Qi, Zeng, Ding, & Li,
2013).
In he p esen expe imen , pa icipan s i s pe o med a lea ning ask du ing
which hey lea n he spa ial loca ion o a p ede ined a ge (a small key) embedded
wi hin isual scenes. Punishmen - ela ed associa ions we e manipula ed by punishing
bad pe o mance du ing he las block o he lea ning ask on a p opo ion o ials.
Twen y- ou hou s la e , hey comple ed an LTM-cued isual sea ch ask in which hey
had o disc imina e he p esence o absence o a ge key s imuli wi hin he p e iously
s udied scenes while ERPs we e eco ded. The ini ial p esen a ion o he scene (wi hou
he a ge ) se ed as a con ex ual memo y-based cue o o ien spa ial a en ion in each
ial. Con ex ual scene cues could ei he ha e been associa ed wi h a oidance o an
Pos -p in ( inal d a pos - e e eeing)
8
a e si e ou come (i.e. po en ial mone a y loss) o ha e had no ou come associa ion.
Based on ou p io wo k (Doallo e al., 2013), we hypo hesized ha i expe ience-
dependen a en ional biases on pe cep ion a e in luenced by punishmen a oidance
associa ions, his would lead, speci ically, o (i) enhanced beha iou al pe o mance, as
e ealed by imp o ed eac ion imes and accu acy, and (ii) modula ion o neu al
signa u es o ea ly isual p ocessing (exp essed as enhanced ampli udes o he P1
po en ial and ea lie la encies o N1) and a ge selec ion (indexed by educed
ampli udes o N2pc).
2. Ma e ials and Me hods
2.1. Pa icipan s
Thi y- wo heal hy s uden s om he Uni e si y o San iago de Compos ela (Galicia,
Spain) pa icipa ed in his s udy o mone a y compensa ion. Pa icipan s comple ed a
ques ionnai e ega ding pe sonal his o y o neu ological and/o psychia ic disease,
exis ing ch onic disease and/o cu en pha macological ea men . All pa icipan s ga e
w i en consen . Da a om i e pa icipan s we e disca ded du ing EEG p ep ocessing
because o excessi e a i ac s in hei EEG eco ding. The emaining 27 pa icipan s (18
women, age ange 19-27, mean age 20.89 ± 2.02) we e all igh -handed and had no mal
o co ec ed- o-no mal ision. The s udy was app o ed by he Bioe hics Commi ee o
he Uni e si y o San iago de Compos ela.
2.2. Expe imen al p ocedu e
The ask used in his s udy was a modi ied e sion o he expe imen al design employed
in Doallo e al. (2013). The e we e wo phases o he expe imen . Pa icipan s i s
pe o med a lea ning ask (see Figu e 1), ollowed on he second day by a memo y-cued
Pos -p in ( inal d a pos - e e eeing)
9
o ien ing ask (in which EEG ac i i y was eco ded) (Figu e 2A) and a spa ial memo y
ecall ask (Figu e 2B).
2.2.1. S imuli
Two hund ed and ou digi al images o scenes we e ob ained om he image se used
in ou p e ious s udy (Doallo e al., 2013). A se o 192 scenes was used in
expe imen al ials, and addi ional 12 scenes we e used o amilia iza ion and p ac ice
ials. Each scene was p epa ed, using Ma lab (Ma hwo ks, Na ick, MA), in wo
di e en o ma s, one o he lea ning ask and one o he o ien ing ask. Fo
coun e balancing pu poses, ou lea ning ask e sions we e p epa ed o each scene
wi h he key (15 x 29 pixels, equi alen o 0.3º x 0.7º) placed in one o each o he ou
isual quad an s, p e e ably in a hidden loca ion. The assignmen o scenes o di e en
expe imen al condi ions, key p esence o absence (in he o ien ing ask) and key
loca ion wi hin scenes we e coun e balanced ac oss pa icipan s. Fo he o ien ing ask,
he scenes wi h keys we e emade o include a la ge and b igh e key (25 x 49 pixels;
0.6º x 1.1º) in he loca ion o he o iginal key o make i isible wi hin he b ie ly
displayed a ge scene. Scene s imuli we e p esen ed using P esen a ion
(Neu obeha io al Sys ems, Albany, CA) and sub ended 22º x 17º o isual angle a a
iewing dis ance o 100 cm.
2.2.2. Lea ning ask
Du ing he lea ning ask, pa icipan s iewed 192 na u alis ic scenes epea ed in andom
o de o e i e blocks (Figu e 1A). Pa icipan s explo ed he scenes o e ly o sea ch
o a small gold key a ge (p esen in all he scenes). Once loca ed, hey ac i a ed he
mouse cu so wi h a le -side mouse click and indica ed he loca ion o he key by
posi ioning he cu so on he loca ion o he key and making a second le -sided mouse
click. A e a esponse o a e he a ailable sea ch ime expi ed, isual w i en eedback
Pos -p in ( inal d a pos - e e eeing)
16
we e hus co ec ed o mul iple compa isons. Addi ional analyses should be conside ed
as explo a o y and hypo hesis-gene a ing. Fo all p alues epo ed h oughou he
pape , we p o ide nominal p alues and, whe e app op ia e, co ec ed p alues using
FDR, as well as e ec sizes calcula ed as pa ial e a-squa ed (ŋ2p) alues.
2.5. Media ion analysis
Media ion analyses we e conduc ed o examine whe he modula ion o wa e o m
po en ials media ed he e ec o condi ion on pe o mance (i.e. whe he he bene i s in
a oidance s. sa e ials we e media ed by modula ions o isual co ical p ocessing).
We pe o med sepa a e media ion analyses o epea ed-measu es designs using he
MEMORE mac o o SPSS (Mon oya & Hayes, 2017), wi h condi ion (a oidance s.
sa e) as he independen a iable, ERPs (ampli ude and/o la ency pa ame e s) as he
media o s, and beha iou (RT, accu acy and d’) as he dependen a iables. The
p ocedu e desc ibed by Mon oya and Hayes (2017) concep ualizes media ion analysis
as a pa h-analy ic amewo k in which media ion is assessed by a single es o he
indi ec e ec (a*b; i.e. he conjunc ion o he e ec o condi ion on he po en ial
media o [pa h a] and he e ec o he po en ial media o on beha io al pe o mance
[pa h b]). The di ec e ec o condi ion on beha io al pe o mance ha does no ope a e
h ough he media o is also calcula ed (pa h c’). Media ion analyses we e conduc ed
using he pe cen ile boo s ap me hod wi h 10,000 i e a ions. The indi ec e ec was
conside ed s a is ically signi ican i he con idence in e al (CI 95%) excluded ze o.
3. Resul s
3.1. Fo ma ion o obus LTMs o a ge loca ions wi hin na u al scenes
Pos -p in ( inal d a pos - e e eeing)

17
Beha iou al analysis o he lea ning ask con i med ha pa icipan s we e able o
es ablish obus memo ies o he spa ial loca ions a which a ge s imuli we e
p esen ed. O e he cou se o he lea ning blocks, pa icipan s loca ed an inc easing
numbe o a ge s, wi h inc easing speed (Block 1: mean accu acy ± SEM = 92.4 ±
1.1%, mean sea ch imes ± SEM = 4.2 ± 0.2 sec; Block 5: 97.8 ± 0.4%, 1.2 ± 0.1 sec)
(Figu e 1B). ANOVAs es ing o linea dec eases in sea ch imes e ealed a signi ican
linea con as o e he lea ning blocks, F(1, 26) = 386.64, p < .001, ŋ2p = .94. A
signi ican linea inc ease in accu acy o e he lea ning blocks was simila ly e ealed,
F(1, 26) = 48.53, p < .001, ŋ2p = .65. No di e ence was ound be ween a oidance and
sa e scenes in he inal (six h) block ei he in sea ch ime (a oidance: 1.1 ± 0.05; sa e:
1.1 ± 0.05; F(1, 26) = .07, p = .79, ŋ2p = .003) o in accu acy (a oidance: 97.7 ± 0.5;
sa e: 96.9 ± 0.5; F(1, 26) = 3.34, p = .08, ŋ2p = .11).
The ecall ask pe o med immedia ely a e he o ien ing ask con i med ha
pa icipan s e ained s ong memo ies o he key loca ions wi hin he lea ned scenes on
he day a e he lea ning ask. On a e age, pa icipan s could explici ly ecall he
co ec loca ions o a ge s on 75% o scenes (± 2.7 SEM). In addi ion, subjec s'
con idence a ings a ied sys ema ically wi h he esponse dis ance om ac ual a ge
loca ion, F(1.6, 41.6) = 64.96, p < .001, ɛ = .8, ŋ2p = .71, e ealing ha highe
con idence a ings we e associa ed wi h mo e accu a e memo ies [mean dis ance in
pixels ± SEM; Ra e 1: 81.4 ± 3.9; Ra e 2: 60.7 ± 3.5; Ra e 3: 36.9 ± 1.6] (see Figu e
2B). Recall did no di e be ween a oidance-associa ed and sa e key loca ions
(accu acy: 75.9 ± 2.6% s. 73.8 ± 3%, F(1, 26) = 1.93, p = .18, ŋ2p = .07; o e all
dis ance in pixels: 58.4 ± 3 s. 60.9 ± 2.9, F(1, 26) = .39, p = .54, ŋ2p = .02; con idence
a ings: 2.67 ± .04 s. 2.68 ± .04, F(1, 26) = .01, p = .92, ŋ2p < .001).
Pos -p in ( inal d a pos - e e eeing)
18
3.2. One single punishmen a oidance- ela ed associa ion enhances he abili y o
spa ial memo ies o d i e isual sea ch in na u al scenes
RT and accu acy le els o a ge -p esen and a ge -absen ials in a oidance and sa e
condi ions in he o ien ing ask a e summa ized in Table 1.
To es ou main hypo heses ega ding beha iou al pe o mance (imp o ed RT
and accu acy), we examined he main e ec o condi ion on hese wo dependen
a iables. The e was a signi ican e ec on RT, F(1, 26) = 6.67, p = .016, p(FDR
co ec ed) = .048, ŋ2p = .204, e ealing ha a ge disc imina ion was as e in
a oidance (700.03 ± 31.13) e sus sa e (709.07 ± 31.22) ials. The main e ec o
condi ion on accu acy was no signi ican (a oidance: 0.91 ± 0.01, sa e: 0.92 ± 0.01;
F(1, 26) = .09, p = .77, p(FDR co ec ed) = .81, ŋ2p = .003). In addi ion, RT and
accu acy ANOVAs showed he ollowing esul s. Ta ge disc imina ion was as e in
a ge -p esen ials (p esen : 651.58 ± 32.3, absen : 757.53 ± 32.37; F(1, 26) = 36.59, p
< .001, ŋ2p = .59), bu no signi ican in e ac ion was ound be ween condi ion and a ge
p esence on RT, F(1, 26) = .58, p = .45, ŋ2p = .02. Analysis o accu acy e ealed
ma ginally signi ican e ec s o a ge p esence (p esen : 0.91 ± 0.01, absen : 0.93 ±
0.01; F(1, 26) = 3.93, p = .058, ŋ2p = .13) and o he in e ac ion be ween condi ion and
a ge p esence, F(1, 26) = 3.89, p = .059, ŋ2p = .13, which showed a end o highe
accu acy o sa e s. a oidance condi ions in a ge absen ials. To es he possibili y
ha RT e ec s e lec ed a speed-accu acy ade-o , Pea son’s co ela ion be ween RT
and accu acy e ec s (calcula ed by sub ac ing sa e ials om a oidance ials) was
pe o med. A signi ican nega i e co ela ion ( = -.48, p = .006) indica ed ha as e
esponses in a oidance s. sa e scenes we e associa ed wi h highe le els o accu acy.
Pos -p in ( inal d a pos - e e eeing)
19
Finally, he analysis o pe cep ual sensi i i y showed ha he d’ measu e was
equi alen be ween a oidance and sa e ials (a oidance: 3.27 ± 0.63, sa e: 3.2 ± 0.61;
(26) = .58, p = .57).
3.3. Punishmen a oidance-associa ed memo ies modula e a ge - ela ed neu al
ac i i y
3.3.1. Ea ly isual p ocessing (P1 and N1 po en ials)
Ta ge -p esen scenes elici ed he expec ed isual po en ials P1 and N1 o e pa ie o-
occipi al scalp egions in all condi ions (Figu e 3).
As explained in he In oduc ion sec ion, d awing on ou p e ious wo k (Doallo
e al., 2013; see also Summe ield e al., 2011), we expec ed ha punishmen a oidance-
associa ed memo y-based o ien ing would esul in enhanced ampli udes o he P1
po en ial and ea lie la encies o N1. As p edic ed, he e was a signi ican main e ec o
condi ion on P1 ampli ude, F(1, 26) = 4.31, p = .048, ŋ2p = .14, e ealing ha P1 was
la ge o a ge s appea ing a a oidance-associa ed emembe ed loca ions ela i e o
sa e loca ions. A e ha ing iden i ied ha one ol age alue was ou side 3SD om he
mean, we e i ied ha he main e ec o condi ion on P1 ampli ude emained
signi ican when he ou lie was eplaced by he se ies’ mean, F(1, 26) = 4.68, p = .040,
ŋ2p = .15. This esul , howe e , did no su i e he FDR co ec ion, p(FDR co ec ed) =
.067. Rega ding he N1 la ency, he e was no a signi ican main e ec o condi ion, F(1,
26) = 2.45, p = .13, p(FDR co ec ed) = .81, ŋ2p = .09. The ANOVAs ca ied ou on he
P1 ampli ude and N1 la ency also e ealed he ollowing esul s. A signi ican main
e ec o elec ode on P1 ampli ude, F(2.03, 52.66) = 3.73, p = .03, ɛ = .338, ŋ2p = .13,
indica ed ha i was maximal o e elec odes P5/6 and PO7/8, bu no o he main e ec s
o in e ac ions eached signi icance. The analysis o N1 la ency showed ha i peaked
Pos -p in ( inal d a pos - e e eeing)
20
ea lie o e he mo e pos e io si es (O1/2, PO3/4; main e ec o elec ode, F(3.09,
80.27) = 10.77, p < .001, ɛ = .515, ŋ2p = .29), and o e con ala e al (149 ± 2 msec)
e sus ipsila e al (151 ± 2 msec) si es (main e ec o hemisphe e: F(1, 26) = 4.72, p =
.039, ŋ2p = .15). The e also was a signi ican hemisphe e x condi ion in e ac ion on N1
la ency, F(1, 26) = 6.31, p = .019, ŋ2p = .195, which showed he la encies o be ea lies
o a ge s p eceded by a oidance-associa ed memo y cues (149 ± 2 msec) han o
hose p eceded by sa e memo y cues (153 ± 3 msec) o e he ipsila e al hemisphe e (p =
.030). The pos -hoc compa isons also e ealed he e ec o hemisphe e being
signi ican o sa e ials (con ala e al s. ipsila e al: 150 ± 3 s. 153 ± 3 msec; p =
0.009) bu no o a oidance ials (149 ± 2 s. 149 ± 2 msec; p = 0.708).
The analysis o P1 la ency showed ea lie la encies a pos e io elec odes (O1/2,
PO3/4; main e ec o elec ode: F(3.13, 81.29) = 10.28, p < .001, ɛ = .521, ŋ2p = .28),
bu no signi ican e ec s o hemisphe e o condi ion; only subsidia y ANOVAs on a
signi ican 3-way in e ac ion be ween elec ode, hemisphe e and condi ion, F(4, 104) =
2.48, p = .049, ɛ = .667, ŋ2p = .09, e ealed a signi ican hemisphe e x condi ion
in e ac ion a PO9/10 elec odes, F(1, 26) = 5.43, p = .028, ŋ2p = .17, showing a end
o ea lie P1 la encies in a oidance (95 ± 2 msec) e sus sa e (100 ± 3 msec) ials o e
ipsila e al si es (p = 0.053).
The analysis o N1 ampli ude showed ha i was la ge a pos e io elec odes
(O1/2, PO3/4; main e ec o elec ode, F(3.33, 86.59) = 12.83, p < .001, ɛ = .555, ŋ2p =
.33), bu nei he a main e ec no an in e ac ion in ol ing he ac o condi ion we e
ound, indica ing ha he ampli ude o he N1 componen was una ec ed by ype o
memo y cue.
Pos -p in ( inal d a pos - e e eeing)
21
___________________________________________
Inse Figu e 3 a ound he e
___________________________________________
3.3.2. Ta ge selec ion (N2pc)
The N2pc po en ial was elici ed by a ge -p esen scenes a pa ie o-occipi al elec odes
con ala e al o he side o he a ge (Figu e 4). The eliabili y o he N2pc was
con i med by a main e ec o hemisphe e, F(1,26) = 32.28, p < .001, ŋ2p = .55, on mean
ampli udes 190-270 msec a e a ge onse . I as p edic ed, based on ou p e ious
esea ch, N2pc ampli ude is modula ed by he mo i a ional alue o he p eceding cue,
his should esul in an in e ac ion be ween hemisphe e and condi ion. The p edic ed
wo-way in e ac ion was indeed signi ican , e en a e FDR co ec ion, F(1,26) = 6.23,
p = .019, p(FDR co ec ed) = .048, ŋ2p = .19, indica ing ha , as expec ed, he ampli ude
o N2pc became a enua ed by punishmen a oidance associa ed-memo y cues. Pos -
hoc analysis indica ed ha a signi ican N2pc was elici ed in sa e ials (p < .001) and,
al hough smalle , was also p esen in a oidance ials (p = .001).
To con i m ha he N2pc was ela ed o he selec ion o he a ge in i s scene
con ex and o ule ou ha i may ha e been d i en simply by he o ien ing o spa ial
a en ion, we also es ed o he p esence o he N2pc in a ge -absen ials. In hese
cases, he e was no signi ican main e ec o hemisphe e, F(1,26) = .73, p = .40, ŋ2p =
.03, on mean ampli udes 190-270 a e scene onse , con i ming ha N2pc was no
obse ed when no a ge was p esen .
___________________________________________
Inse Figu e 4 a ound he e
___________________________________________
Pos -p in ( inal d a pos - e e eeing)

22
Visual inspec ion o he wa e o ms also e ealed a la e la e alized e ec , ollowing he
N2pc wi h opposi e pola i y (see Figu e 4), which has been also epo ed in ou p e ious
s udies examining LTM-guided isual sea ch in na u alis ic scenes (labeled as pos e io
con ala e al posi i i y, PCP; Doallo e al., 2013; Pa ai e al., 2012). This e ec was an
enhanced posi i i y o e pos e io con ala e al ( ela i e o ipsila e al) scalp loca ions o
he a ge side in he la ency window be ween 320 and 380 msec pos s imulus. An
ANOVA analyzing ERP mean ampli udes h ough his la ency window o e PO9/10,
PO7/8, PO3/4 and O1/2 con i med he p esence o his la e alized e ec (main e ec o
hemisphe e: F(1,26) = 14.67, p < .001, ŋ2p = .36), bu i was no di e en ially
modula ed by punishmen a oidance-associa ed con ex ual memo ies (hemisphe e x
condi ion: F(1,26) = .45, p = .508, ŋ2p = .02).
3.4. Rela ionship be ween beha io al and elec ophysiological measu es
Media ion analyses showed ha he P1 ampli ude media ed he e ec o
condi ion on RT, ab = -4.227, boo SE = 2.739, CI95% [-10.499, -.102], which was
he only indi ec e ec signi ican ly di e en om ze o (see also Supplemen a y Figu e
1). This esul sugges s ha as e esponses in a oidance ela i e o sa e condi ions
may occu h ough changes in senso y p ocessing e lec ed by he P1 po en ial (i.e. P1
ampli ude seems o p edic a signi ican p opo ion o he co- a ia ion be ween
condi ion and RT). The comple e pa e n o esul s om he media ion analyses is
shown in Supplemen a y Tables 1, 2 and 3.
4. Discussion
The indings o his expe imen demons a e he ole o pas a oidance o an a e si e
ou come in magni ying memo y-d i en a en ional biases upon pe cep ual decisions on
Pos -p in ( inal d a pos - e e eeing)
23
ele an objec s embedded in clu e ed na u al scenes. Impo an ly, ou indings
e ealed he abili y o one single punishmen -a oidance associa ion in memo y o
enhance a en ion and isual sea ch p ocesses in scenes and o modula e ongoing
p ocessing in isual co ical a eas.
Beha io al esul s showed ha spa ial expec a ions om LTM associa ed o
success ul a oidance o punishmen con e ed beha io al bene i s, as e ealed by as e
esponses o a ge s placed in punishmen a oidance- e sus sa e- ela ed emembe ed
loca ions. These indings eplica e and ex end ou p e ious esul s (Doallo e al., 2013)
by showing ha punishmen - ela ed LTM, simila ly o wha is obse ed when posi i e
ewa d is in ol ed, allows a en ion o each he a ge loca ion mo e apidly compa ed
wi h when memo ies no associa ed o mo i a ional alues guide spa ial o ien ing. The
obse ed e ec s in beha io hus indica e ha punishmen a oidance can bias a en ion
h ough associa ions in memo y bu in he absence o immedia e po en ially nega i e
consequences (i.e. no punishmen was a s ake in he isual sea ch ask). The p esen
da a expand upon he g owing beha io al e idence showing ha s imuli associa ed
h ough lea ning o punishmen a oidance (Wen u a e al., 2014) o o an ac ual
mone a y loss (Bucke & Theeuwes, 2016) can in luence a en ional deploymen and
ecei e acili a ed p ocessing (O'B ien & Raymond, 2012). Unlike ou p io s udy
manipula ing posi i e ewa ds, we did no ind signi ican e ec s on accu acy. Taking
in o accoun ha he e was also a end o highe accu acy in sa e ela i e o a oidance
ials o a ge absen scenes, i is ha d o say, om he p esen esul s, i his di e ence
may be a ibu able o speci ically manipula e a oidance- ela ed alues, leading o
di e en ial beha io al e ec s han when gains a e in ol ed. In his sense, RT e ec s in
he absence o accu acy ones, ela ed o punishmen -associa ed s imuli, ha e been
epo ed p e iously (Bucke & Theeuwes, 2016).
Pos -p in ( inal d a pos - e e eeing)
24
The momen -by-momen eco d o a ge - ela ed ERPs showed ha he
beha io al imp o emen d i en by a oidance- ela ed associa ions was accompanied by
modula ion o mul iple s ages o isual p ocessing. Simila ly o he pa e n o esul s
obse ed in ou p e ious s udy, he ea lies e ec was obse ed on he P1 po en ial, an
index o a en ional gain in ex as ia e isual co ex, which showed la ge ampli udes a
pa ie o-occipi al si es o a ge s appea ing a punishmen a oidance-associa ed
emembe ed loca ions ( ela i e o sa e- ela ed loca ions), in he absence o e ec s on
he P1 la ency. These indings p o ide e idence ha a oidance-associa ed memo y-
d i en a en ion biases ea ly pe cep ual analysis o ele an objec s embedded in
c owded eal-wo ld scenes. Al hough he main e ec o condi ion on P1 ampli ude,
which passed a nominal alpha h eshold o .05, did no su i e FDR co ec ion, he
inding ha he P1 ampli ude modula ion signi ican ly media ed he ela ionship
be ween punishmen - ela ed associa ions in memo y and bene i s in he eac ion ime
u he unde sco es he signi icance o his po en ial in he p io i ized p ocessing o
s imuli a spa ial loca ions linked o e asion o punishmen in he pas .
Ou indings seem o di e om some ecen epo s o ERP modula ions by
acqui ed posi i e and nega i e mo i a ional salience. Baye e al. (2017) in es iga ed
he in e play be ween ewa d p ospec (i.e. pe o mance-based mone a y incen i es)
du ing a cued pa e n disc imina ion ask on he ac i i y o he p ima y and ex as ia e
isual co ex. Mo i a ional ele ance con eyed by he cue ( ela ed o bo h ewa d
app oach and punishmen a oidance) inc eased he ampli ude o he C1 po en ial
( e lec ing ea ly pe cep ual p ocessing in V1) bu had no e ec s a he P1 le el. On he
o he hand, Hamme schmid , Sennhenn-Reulen, and Schach (2017) ound ha neu al
aces p e iously associa ed wi h h ee di e en mone a y ou comes (gain, loss o ze o)
impac ed di e en ly he ampli ude o he P1 po en ial: he acili a ed senso y p ocessing
Pos -p in ( inal d a pos - e e eeing)
25
o s imuli wi h associa ed mo i a ional salience, as e lec ed by la ge P1 ampli udes,
was con ined o ewa d- ela ed aces ( ela i e o he neu al, ze o-ou come, ones). When
add essing he di e ences be ween ea lie s udies and ou s, i is impo an o ake in o
accoun ha indings such as hose om Hamme schmid e al. (2017) indica e a
spa ially unspeci ic e ec ( a he han limi ed o speci ic spa ial posi ions) o acqui ed
mo i a ional salience. The di e ences be ween he esul s epo ed by Baye e al.
(2017) and hose o he cu en s udy could e lec di e en ial e ec s o isual spa ial
cues con eying in o ma ion abou pe o mance-based mone a y incen i es and
con ex ual memo y cues which acqui ed alue in a p io lea ning expe ience. Ano he
con ibu ing ac o o hese di e ences could be he challenging pe cep ual condi ions in
ou ask. Ou a ge s imuli we e embedded wi hin an associa ed clu e ed scene, which
may a o p io i iza ion e ec s (as e lec ed by highe ampli udes o P1) by which
s imuli appea ing a spa ial loca ions associa ed wi h a oidance o an a e si e ou come
in pas encoun e s win ep esen a ion a he expense o o he s imuli.
Rega ding he N1 po en ial, an elec ophysiological ma ke o disc imina i e
p ocesses in a eas o he en al isual s eam (Hop e al., 2002; Luck, 1995; Vogel &
Luck, 2000), ou esul s showed a signi ican in e ac ion be ween condi ion and
hemisphe e on i s la ency, indica ing ha o ien ing o spa ial a en ion based on loss
a oidance end o sho ened he la ency o he N1 componen o e ipsila e al si es.
Al hough ou p ima y hypo hesis es ing did no e eal a main e ec o condi ion on N1
la ency, his in e ac ion sugges s ha con ex ual memo ies associa ed o a oidance o
punishmen a e also able o speed up he disc imina ion o a ge s appea ing a hese
loca ions in subsequen encoun e s, in line wi h ou p io indings (Doallo e al., 2013).
This esul , howe e , should be in e p e ed wi h cau ion because i was no subjec ed o
mul iple es ing co ec ion.
Pos -p in ( inal d a pos - e e eeing)
32
his issue, i is impo an o ake in o accoun ha he es ic ed s a is ical powe , mainly
due o he cha ac e is ics o his ype o expe imen al design, makes necessa y o
balance he likelihood o Type II e o s. We should no e ha he ecological alidi y
achie ed by he use o memo ies o speci ic a ge loca ions in eal scenes comes a a
cos . I limi s he numbe o scenes we can use, and p e en s us om epea ing scenes
du ing he pe cep ual disc imina ion ask (wi hou possibly comp omising he s a e o
he memo ies being in es iga ed). This consequen ly educes he numbe o ials
a ailable and limi s he s a is ical powe ha can be achie ed in o he ypes o designs.
To deal wi h his limi a ion and balance he likelihood o Type I and Type II e o s, we
co ec ed ou p ima y hypo heses o mul iple compa isons, as epo ed h oughou his
s udy.
In conclusion, he p esen s udy p o ides new e idence ha memo y-dependen
spa ial a en ional biases on pe cep ion a e in luenced by punishmen a oidance
associa ions, leading o enhanced beha io al pe o mance and modula ion o neu al
signa u es o a ge p ocessing. I also ex ends ou unde s anding o he ole o acqui ed
mo i a ional alue in selec i ely p io i izing speci ic spa ial loca ions when sea ching
o objec s in na u alis ic con ex s.
Acknowledgmen s
This esea ch was suppo ed by a P ojec G an o S.D. om he Conselle ía de
Educación e O denación Uni e si a ia (Xun a de Galicia, Spain) (EM2012-017). S.S-S
was suppo ed by a g an o p edoc o al con ac s om he Spanish Minis y o
Economy and Compe i i eness (BES-2016-076298). A.C.N. is suppo ed by Wellcome
Pos -p in ( inal d a pos - e e eeing)

33
T us Senio In es iga o Awa d (104571/Z/14/Z). We hank Kenia Co ea o
assis ance in EEG da a collec ion and p ep ocessing.
Da a s a emen
We epo how we de e mined ou sample size, all da a exclusions, all
inclusion/exclusion c i e ia, whe he inclusion/exclusion c i e ia we e es ablished p io
o da a analysis, all manipula ions, and all measu es in he s udy. No pa o he s udy
p ocedu es o analyses was p e egis e ed p io o he esea ch being unde aken. Due o
policy o he esea ch g oup, public a chi ing o da a and digi al s udy ma e ials will be
done a he ime ou ins i u ion makes a ailable an ins i u ional eposi o y o ha
pu pose. Un il hen, eade s seeking access o he da a should con ac he co esponding
au ho .
REFERENCES
Adcock, R. A., Thanga el, A., Whi ield-Gab ieli, S., Knu son, B., & Gab ieli, J. D.
(2006). Rewa d-mo i a ed lea ning: Mesolimbic ac i a ion p ecedes memo y
o ma ion. Neu on, 50(3), 507-517. doi: 10.1016/j.neu on.2006.03.036
Ande son, B. A. (2013). A alue-d i en mechanism o a en ional selec ion. Jou nal o
Vision, 13(3), 7-7. doi:10.1167/13.3.7
Ande son, B. A. (2015). Value-d i en a en ional cap u e is modula ed by spa ial
con ex . Visual Cogni ion, 23(1-2), 67-81. doi: 10.1080/13506285.2014.956851
Ande son, B. A. (2016). The a en ion habi : how ewa d lea ning shapes a en ional
selec ion. Annals o he New Yo k Academy o Sciences, 1369(1), 24-39. doi:
10.1111/nyas.12957
Pos -p in ( inal d a pos - e e eeing)
34
Ande son, B. A., Kuwaba a, H., Wong, D. F., Gean, E. G., Rahmim, A., B ašic, J. R.,
… Yan is, S. (2016). The ole o dopamine in alue-based a en ional o ien ing.
Cu en Biology, 26(4), 550-555. doi: 10.1016/j.cub.2015.12.062
Ande son, B. A., Kuwaba a, H., Wong, D. F., Robe s, J., Rahmim, A., B ašic, J. R., &
Cou ney, S. M. (2017). Linking dopamine gic ewa d signals o he de elopmen
o a en ional bias: A posi on emission omog aphic s udy. Neu oimage, 157, 27-
33. doi: 10.1016/j.neu oimage.2017.05.062
Ande son, B. A., Lau en , P. A., & Yan is, S. (2011). Value-d i en a en ional cap u e.
P oceedings o he Na ional Academy o Sciences, U.S.A., 108, 10367-10371. doi:
10.1073/pnas.1104047108
Ande son, B. A., Lau en , P. A., & Yan is, S. (2014). Value-d i en a en ional p io i y
signals in human basal ganglia and isual co ex. B ain Resea ch, 1587, 88-96.
doi: 10.1016/j.b ain es.2014.08.062
Ba ba o, L., Peelen, M. V., & Hickey, C. (2017). Valence, no u ili y, unde lies ewa d-
d i en p io i iza ion in human ision. Jou nal o Neu oscience, 37(43), 10438-
10450. doi: 10.1523/JNEUROSCI.1128-17.2017
Baye , M., Rossi, V., Vanlessen, N., G ass, A., Schach , A., & Pou ois, G. (2017).
Independen e ec s o mo i a ion and spa ial a en ion in he human isual co ex.
Social Cogni i e and A ec i e Neu oscience, 12(1), 146-156.
doi:10.1093/scan/nsw162
Benjamini, Y., & Hochbe g, Y. (1995). Con olling he alse disco e y a e: a p ac ical
and powe ul app oach o mul iple es ing. Jou nal o he Royal S a is ical Socie y
Se ies B, 57, 289–300. h p://www.js o .o g/s able/2346101.
Bissone e, G. B., Gen y, R. N., Padmala, S., Pessoa, L., & Roesch, M. R. (2014).
Impac o appe i i e and a e si e ou comes on b ain esponses: Linking he
Pos -p in ( inal d a pos - e e eeing)
35
animal and human li e a u es. F on ie s in Sys ems Neu oscience, 8, 24.
doi:10.3389/ nsys.2014.00024
B ignani, D., Lepsien, J., & Nob e, A. C. (2010). Pu ely endogenous cap u e o
a en ion by ask-de ining ea u es p oceeds independen ly om spa ial a en ion.
Neu oImage, 51(2), 859-866. doi: 10.1016/j.neu oimage.2010.03.029
B ooks, A. M., & Be ns, G. S. (2013). A e si e s imuli and loss in he
mesoco icolimbic dopamine sys em. T ends in Cogni i e Sciences, 17(6), 281-
286. doi:10.1016/j. ics.2013.04.001
Bucke , B., & Theeuwes, J. (2016). Appe i i e and a e si e ou come associa ions
modula e exogenous cueing. A en ion, Pe cep ion & Psychophysics, 78(7), 2253-
2265. doi:10.3758/s13414-016-1107-6
Chelazzi, L., Eš očino á, J., Calle i, R., Lo Ge o, E., Sani, I., Della Libe a, C., &
San and ea, E. (2014). Al e ing spa ial p io i y maps ia ewa d-based lea ning.
Jou nal o Neu oscience, 34(25), 8594-8604. doi:10.1523/JNEUROSCI.0277-
14.2014
Chelazzi, L., Pe la o, A., San and ea, E., & Della Libe a, C. (2013). Rewa ds each
isual selec i e a en ion. Vision Resea ch, 85, 58-72.
doi:10.1016/j. is es.2012.12.005
Della Libe a, C., & Chelazzi, L. (2006). Visual selec i e a en ion and he e ec s o
mone a y ewa ds. Psychological Science, 17(3), 222-227. doi: 10.1111/j.1467-
9280.2006.01689.x
Della Libe a, C., & Chelazzi, L. (2009). Lea ning o a end and o igno e is a ma e o
gains and losses. Psychological Science, 20(6), 778-784. doi: 10.1111/j.1467-
9280.2009.02360.x
Pos -p in ( inal d a pos - e e eeing)
36
Doallo, S., Pa ai, E. Z., & Nob e, A. C. (2013). Rewa d associa ions magni y memo y-
based biases on pe cep ion. Jou nal o Cogni i e Neu oscience, 25(2), 245-257.
doi:10.1162/jocn_a_00314
Donohue, S. E., Hop , J-M., Ba sch, M. V., Schoen eld, M. A., Heinze, H-J., &
Woldo , M. G. (2016). The apid cap u e o a en ion by ewa ded objec s.
Jou nal o Cogni i e Neu oscience, 28(4), 529-541. doi: 10.1162/jocn_a_00917
Eime , M. (1996). The N2pc componen as an indica o o a en ional selec i i y.
Elec oencephalog aphy and Clinical Neu ophysiology, 99(3), 225-234.
doi:S0921884X96957112
Engelmann, J. B., Dama aju, E., Padmala, S., & Pessoa, L. (2009). Combined e ec s o
a en ion and mo i a ion on isual ask pe o mance: ansien and sus ained
mo i a ional e ec s. F on ie s in Human Neu oscience, 3, 4. doi:
10.3389/neu o.09.004.2009
Failing, M., & Theeuwes, J. (2018). Selec ion his o y: How ewa d modula es
selec i i y o isual a en ion. Psychonomic Bulle in & Re iew, 25, 514-538. doi:
10.3758/s13423-017-1380-y
Ghazizadeh A., G iggs, W., Leopold, D. A., & Hikosaka, O. (2018). Tempo al-
p e on al co ical ne wo k o disc imina ion o aluable objec s in long- e m
memo y. P oceedings o he Na ional Academy o Sciences o he Uni ed S a es o
Ame ica, 115(9), 2135-2144. doi: 10.1073/pnas.1707695115
G iggs, W. S., Ami a, H., Gopal, A., & Hikosaka, O. (2018). Visual neu ons in he
supe io colliculus disc imina e many objec s by hei his o ical alues. F on ie s
in Neu oscience, 12, 396. doi: 10.3389/ nins.2018.00396
Pos -p in ( inal d a pos - e e eeing)
37
Hamme schmid , W., Sennhenn-Reulen, H., & Schach , A. (2017). Associa ed
mo i a ional salience impac s ea ly senso y p ocessing o human aces.
Neu oImage, 156, 466-474. doi:S1053-8119(17)30325-7
Ha is, J. A., Donohue, S. E., Schoen eld, M. A., Hop , J-M., Heinze, H-J., & Woldo ,
M. G. (2016). Rewa d-associa ed ea u es cap u e a en ion in he absence o
awa eness: E idence om objec -subs i u ion masking. Neu oImage, 137, 116-
123. doi: 10.1016/j.neu oimage.2016.05.010
Heinze, H. J., Mangun, G. R., Bu che , W., Hin ichs, H., Scholz, M., Mün e, T. F., …
Hillya d, S. A. (1994). Combined spa ial and empo al imaging o b ain ac i i y
du ing isual selec i e a en ion in humans. Na u e, 372(6506), 543-546. doi:
10.1038/372543a0
Hickey, C., Chelazzi, L., & Theeuwes, J. (2010). Rewa d changes salience in human
ision ia he an e io cingula e. Jou nal o Neu oscience, 30(33), 11096-11103.
doi: 10.1523/JNEUROSCI.1026-10.2010
Hickey, C., Chelazzi, L., & Theeuwes, J. (2014). Rewa d-p iming o loca ion in isual
sea ch. PLoS One, 9(7), e103372. doi: 10.1371/jou nal.pone.0103372
Hickey, C., Di Lollo, V., & McDonald, J. J. (2009). Elec ophysiological indices o
a ge and dis ac o p ocessing in isual sea ch. Jou nal o Cogni i e
Neu oscience, 21(4), 760-775. doi:10.1162/jocn.2009.21039
Hickey, C., Kaise , D., & Peelen, M. V. (2015). Rewa d guides a en ion o objec
ca ego ies in eal-wo ld scenes. Jou nal o Expe imen al Psychology: Gene al,
144, 264-273. doi: 10.1037/a0038627
Hickey, C., & Peelen, M. (2015). Neu al mechanisms o incen i e salience in
na u alis ic human ision. Neu on, 85, 512-518. doi:
10.1016/j.neu on.2014.12.049
Pos -p in ( inal d a pos - e e eeing)

38
Hickey, C., & Peelen, M. V. (2017). Rewa d selec i ely modula es he linge ing neu al
ep esen a ion o ecen ly a ended objec s in na u al scenes. Jou nal o
Neu oscience, 37(31), 7297-7304. doi: 10.1523/JNEUROSCI.0684-17.2017
Hilimi e, M. R., Moun s, J. R., Pa ks, N. A., & Co ballis, P. M. (2010). E en - ela ed
po en ials dissocia e e ec s o salience and space in biased compe i ion o isual
ep esen a ion. PloS One, 5(9), e12677. doi: 10.1371/jou nal.pone.0012677
Hillya d, S. A., Vogel, E. K., & Luck, S. J. (1998). Senso y gain con ol (ampli ica ion)
as a mechanism o selec i e a en ion: elec ophysiological and neu oimaging
e idence. Philosophical T ansac ions o he Royal Socie y o London, Se ies B,
Biological Sciences, 353(1373), 1257-1270. doi: 10.1098/ s b.1998.0281
Hop , J. M., Luck, S. J., Gi elli, M., Hagne , T., Mangun, G. R., Scheich, H., & Heinze,
H. J. (2000). Neu al sou ces o ocused a en ion in isual sea ch. Ce eb al Co ex,
10(12), 1233-1241.
Hop , J. M., Vogel, E., Woodman, G., Heinze, H. J., & Luck, S. J. (2002). Localizing
isual disc imina ion p ocesses in ime and space. Jou nal o Neu ophysiology,
88(4), 2088-2095. doi:10.1152/jn.2002.88.4.2088
I hipu ipa , S., Cha, K., Rangsipa , N., & Se ences, J. T. (2015). Value-based
a en ional cap u e in luences con ex -dependen decision-making. Jou nal o
Neu ophysiology, 114(1), 560-569. doi: 10.1152/jn.00343.2015
Je de, T. A., Me iam, E. P., Riggall, A. C., Hedges, J. H., & Cu is, C. E. (2012).
P io i ized maps o space in human on opa ie al co ex. Jou nal o Neu oscience,
32(48), 17382-17390. doi: 10.1523/JNEUROSCI.3810-12.2012
Kahn , T., Pa k, S. Q., Haynes, J. D., & Toble , P. N. (2014). Disen angling neu al
ep esen a ions o alue and salience in he human b ain. P oceedings o he
Pos -p in ( inal d a pos - e e eeing)
39
Na ional Academy o Sciences o he Uni ed S a es o Ame ica, 111(13), 5000-
5005. doi: 10.1073/pnas.1320189111
Kim, H. F., Ghazizadeh, A., & Hikosaka, O. (2015). Dopamine neu ons encoding long-
e m memo y o objec alue o habi ual beha io . Cell, 163(5), 1165-1175. doi:
10.1016/j.cell.2015.10.063
Kiss, M., D i e , J., & Eime , M. (2009). Rewa d p io i y o isual a ge single ons
modula es ERP signa u es o a en ional selec ion. Psychological Science, 20(2),
245-251. doi: 10.1111/j.1467-9280.2009.02281.x
Kuo, B. C., Rao, A., Lepsien, J., & Nob e, A. C. (2009). Sea ching o a ge s wi hin he
spa ial layou o isual sho - e m memo y. Jou nal o Neu oscience, 29(25),
8032-8038. doi:10.1523/JNEUROSCI.0952-09.2009
LaBa , K. S., & Cabeza, R. (2006). Cogni i e neu oscience o emo ional memo y.
Na u e Re iews Neu oscience, 7(1), 54-64. doi: 10.1038/n n1825
Lim, S. L., Padmala, S., & Pessoa, L. (2009). Seg ega ing he signi ican om he
mundane on a momen - o-momen basis ia di ec and indi ec amygdala
con ibu ions. P oceedings o he Na ional Academy o Sciences o he Uni ed
S a es o Ame ica, 106(39), 16841-16846. doi:10.1073/pnas.0904551106
Liu, X., Hai s on, J., Sch ie , M., & Fan, J. (2011). Common and dis inc ne wo ks
unde lying ewa d alence and p ocessing s ages: A me a-analysis o unc ional
neu oimaging s udies. Neu oscience & Biobeha io al Re iews, 35(5), 1219-1236.
doi:10.1016/j.neubio e .2010.12.012
Luck, S. J. (1995). Mul iple mechanisms o isual-spa ial a en ion: Recen e idence
om human elec ophysiology. Beha iou al B ain Resea ch, 71(1-2), 113-123.
Pos -p in ( inal d a pos - e e eeing)
40
Luck, S. J., & Hillya d, S. A. (1994). Spa ial il e ing du ing isual sea ch: E idence
om human elec ophysiology. Jou nal o Expe imen al Psychology. Human
Pe cep ion and Pe o mance, 20(5), 1000-1014.
MacLean, M. H., & Giesb ech , B. (2015). Neu al e idence e eals he apid e ec s o
ewa d his o y on selec i e a en ion. B ain Resea ch, 1606, 86-94. doi:
10.1016/j.b ain es.2015.02.016
Ma ínez, A., Anllo-Ven o, L., Se eno, M. I., F ank, L. R., Bux on, R. B., Dubowi z, D.
J., … Hillya d, S. A. (1999). In ol emen o s ia e and ex as ia e isual co ical
a eas in spa ial a en ion. Na u e Neu oscience, 2(4), 364-369. doi: 10.1038/7274
Mohan y, A., Gi elman, D. R., Small, D. M., & Mesulam, M. M. (2008). The spa ial
a en ion ne wo k in e ac s wi h limbic and monoamine gic sys ems o modula e
mo i a ion-induced a en ion shi s. Ce eb al Co ex, 18(11), 2604-2613. doi:
10.1093/ce co /bhn021
Mon oya, A. K., & Hayes, A. F. (2017). Two-condi ion wi hin-pa icipan s a is ical
media ion analysis: A pa h-analy ic amewo k. Psychological Me hods, 22(1), 6-
27. doi: 10.1037/me 0000086
Mu y, V. P., LaBa , K. S., & Adcock, R. A. (2012). Th ea o punishmen mo i a es
memo y encoding ia amygdala, no midb ain, in e ac ions wi h he medial
empo al lobe. Jou nal o Neu oscience, 32(26), 8969-8976. doi:
10.1523/JNEUROSCI.0094-12.2012
Mu y, V. P., Ri chey, M., Adcock, R. A., & LaBa , K. S. (2010). MRI s udies o
success ul emo ional memo y encoding: A quan i a i e me a-analysis.
Neu opsychologia, 48(12), 3459-3469. doi: 10.1016/
j.neu opsychologia.2010.07.030
Pos -p in ( inal d a pos - e e eeing)
41
O'B ien, J. L., & Raymond, J. E. (2012). Lea ned p edic i eness speeds isual
p ocessing. Psychological Science, 23(4), 359-363.
doi:10.1177/0956797611429800
Padmala, S., & Pessoa, L. (2008). A ec i e lea ning enhances isual de ec ion and
esponses in p ima y isual co ex. Jou nal o Neu oscience, 28(24), 6202-6210.
doi:10.1523/JNEUROSCI.1233-08.2008
Pa ai, E. Z., Doallo, S., & Nob e, A. C. (2012). Long- e m memo ies bias sensi i i y
and a ge selec ion in complex scenes. Jou nal o Cogni i e Neu oscience, 24(12),
2281-2291. doi:10.1162/jocn_a_00294
Pessoa, L. (2015). Mul iple in luences o ewa d on pe cep ion and a en ion. Visual
Cogni ion, 23(1-2), 272-290. doi:10.1080/13506285.2014.974729
Pollmann, S., Es ocino a, J., Somme , S., Chelazzi, L., & Zinke, W. (2016). Neu al
s uc u es in ol ed in isual sea ch guidance by ewa d-enhanced con ex ual
cueing o he a ge loca ion. Neu oImage, 124(P A), 887-897.
doi:10.1016/j.neu oimage.2015.09.040
P ak, R. (2012). The on opa ie al a en ional ne wo k o he human b ain: ac ion,
saliency, and a p io i y map o he en i onmen . The Neu oscien is , 18(5), 502-
515. doi: 10.1177/1073858411409051
Qi, S., Zeng, Q., Ding, C., & Li, H. (2013). Neu al co ela es o ewa d-d i en
a en ional cap u e in isual sea ch. B ain Resea ch, 1532, 32-43. doi:
10.1016/j.b ain es.2013.07.044
Raymond, J. E., & O'B ien, J. L. (2009). Selec i e isual a en ion and mo i a ion: The
consequences o alue lea ning in an a en ional blink ask. Psychological Science,
20(8), 981-988. doi:10.1111/j.1467-9280.2009.02391.x
Pos -p in ( inal d a pos - e e eeing)
Figu e1
Click he e o download high esolu ion image
Pos -p in ( inal d a pos - e e eeing)

Figu e 2
Click he e o download high esolu ion image
Pos -p in ( inal d a pos - e e eeing)
Figu e 3
Click he e o download high esolu ion image
Pos -p in ( inal d a pos - e e eeing)
Figu e 4
Click he e o download high esolu ion image
Pos -p in ( inal d a pos - e e eeing)
Suppl. ma e ial o online publica ion only
Click he e o download Suppl. ma e ial o online publica ion only: Supplemen a y da a.docx
Pos -p in ( inal d a pos - e e eeing)