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Laterality:
Asymmetries of Body,
Brain and Cognition
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Eye Preferences in
Common Marmosets
(Callithrix jacchus):
Influence of Age,
Stimulus, and Hand
Preference
Michelle A. Hook-Costigan & Lesley J.
Rogers
Version of record first published: 18
Oct 2010.

To cite this article: Michelle A. Hook-Costigan & Lesley J. Rogers

(1998): Eye Preferences in Common Marmosets (Callithrix jacchus):
Influence of Age, Stimulus, and Hand Preference, Laterality:
Asymmetries of Body, Brain and Cognition, 3:2, 109-130

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 L A T E R A L IT Y , 1 9 9 8 , 3 (2 ) , 1 0 9 ±1 3 0

E y e P r e fe r e n c e s in C o m m o n M a r m o s e ts (C a llith r ix
ja c c h u s): I n flu e n c e o f A g e , S tim u lu s , a n d H a n d
P r e fe r e n c e

M ic h e lle A . H o o k -C o stig a n a n d L e s le y J . R o g e rs
U n iv e rsity of N ew E n g la nd , A u stra lia
Downloaded by [University of Arizona] at 00:51 03 December 2012

E ye p re fe r en c es o f th e c o m m o n m ar m o se t (C a llith r ix ja c c hu s) w er e e xa m ine d ,
ta k in g in to a cc o u nt a g e, a r o u sa l, a n d h an d p re f er en c e . M o n o c ular e y e u se f or
lo o king thr o u g h a sm a ll h o le a t a s tim u lu s w a s r ec o rd e d. O f 2 1 m a rm o s ets, 20
disp la y e d r ig h t-ey e p r ef e re nc e s f or vie w in g a p ie c e o f fa m ilia r fo od . I n s ub jec ts
te st e d a t 3 ±8 , 1 2 , 1 5 ±1 8 , a n d 2 2 m o n th s, ey e p re f er en c e s w er e c o n s iste nt a c ro ss
ag e . A g ro u p bia s, in d ic ativ e o f r ig h t-ey e d n e ss , w a s a lso f ou n d f o r v ie w in g o th e r
st im u li. T h e st im uli in clu d e d a w atc h, m irr or , m o d el o f a b e e tle, an d th e
ex p e rim en te r ’ s h a n d . H ow e v e r, w h e n th e m a r m o se ts vie w e d a th r e a te n in g
st im u lu s , a m o d el r es e m b lin g tw o r e ar in g sn a k e s , the y d is p lay e d in c re a se d a ro us a l
(ind ica te d b y lo n g er d u ra tio n b etw e en m o n o c ular v ie w ing e v e nts an d inc r ea s ed
in c id e nc e o f a ro u se d v o c a lisa tio ns ) a n d th e e y e p re fe r en c es s h ifte d a w a y fr om a
pr e fe re n ce fo r th e rig ht e y e to eith e r n o p r ef er e nc e o r a lef t-ey e p re f er e n c e . N o
re la tio ns h ip b etw ee n e y e pr e fe re n ce a n d h an d p re fe re n c e f o r h o ld ing f oo d w a s
fo un d . T h e r ef or e , w e s ug g es t th a t e y e p re fe r en c es m a y r ef le c t he m isp h er ic
sp ec ialis a tion s fo r p e rc e p tu al p ro c es si ng , a c c or d in g to th e e m o tio na l va le n c e of th e
st im u lu s .

INTRODUCTION
L a tera lise d fu n ctio nin g o f th e b rain is w id esp re ad a m o ng v erteb ra te s, in clu din g
frog s (B au e r, 1 99 3 ), liz a rds (D e ck e l, 19 9 5 ), b ird s (D h arm are tn a m & A n d rew ,
1 99 4 ; R o g e rs, 19 9 5 ), rod e n ts (D e ne n b erg , 1 9 81 ), a n d p rim a te s (H am ilto n &
V erm eire , 1 9 8 8). A ltho u g h m a n y stud ie s h a ve e x am in ed la te ralisa tio n o f m oto r
fun c tio n , p a rtic u la rly in n o n hu m an p rim a tes, lite ra ture o n a sym m e trie s o f
p erc ep tu al p ro ce ssin g is c o m p a rativ e ly spa rse. M o no c u la r te sts m a y pro v ide a
n on in v asiv e m e th o d for d eterm in in g v isua l la te ralisa tio n. In sp ec ie s w ith

R e q u e s ts fo r re p rin ts sh o u l d b e s e n t to M .A . H o o k -C o s tig a n , D e p a rt m en t o f P h y s io lo g y ,
U n i v e rs it y o f N e w E n g la n d , A rm id a le , N S W 2 3 5 1 , A u s tra li a. E m a il: m h o o k h o @ m e tz .u n e .e d u .a u
L .J. R o g e rs g r at e fu l ly a c k n o w le d g e s s u p p o rt fr o m th e A u s tra l ia n R e s e a rc h C o u n c il (G ra n t N o . A
0 9 3 3 0 9 8 4 ). T h i s re s e a rc h fo rm s p a rt o f t h e re s ea rc h to w a rd s M ic h el le H o o k -C o s tig a n ’ s P h D d e g re e
a n d s h e a c k n o w led g e s th e s u p p o rt o f a U n i v e rs it y o f N e w E n g la n d R e s e ar ch S c h o l a rsh ip .

Ó 1 9 9 8 P s y ch o lo g y P re s s L td
 110 HOOK-COSTIGAN AND ROGERS

la tera lly p lac ed e y es, it is re la tive ly e a sy to d e te rm in e ey e pre fere n ce s fo r

vie w in g d iffe ren t stim uli. F o r ex a m p le, D e c ke l (19 9 5 ) ha s rec o rd ed p refe re n tia l
use o f th e left ey e in th e liz ard , A no lis, to m o n itor an o pp o n en t du rin g a n
ag g ressiv e en c o un te r. In c h ic k s, D ha rm a retn am a n d A nd re w (1 99 4 ) h a v e fo u n d
a rig h t-e y e pre fere n ce fo r v iew ing a liv e he n a nd a le ft-e y e p re fere n ce fo r
vie w in g a n ov e l fla sh in g w h ite lig h t. In prim ates, h o w e v er, the e ye s a re pla ce d
fro n tally an d it is n o t po ssib le to d ete rm in e ey e p refe ren c es fro m the p o stu ra l
orie n ta tio n of th e h e ad . T h e re fo re , e y e p re fere n ce s in p rim a te s h av e b ee n
de te rm ine d in ta sk s re q uirin g the su bje cts to loo k th ro u gh a p e ep h o le o r d o w n a
tu b e (R o g ers, W ard , & S ta fford , 1 9 94 ). T h e te rm ey e p re fere n ce , o r ey e
do m in a n ce , re fe rs to th e co n siste nt ch o ic e o f on e e ye o v er th e o th e r in th e
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m o n o cu lar v iew ing situa tio n .

In p rim a tes e a ch e ye p ro vid es se nso ry inp u t to bo th o f th e c ere bra l
he m isp h ere s. H ow e v e r, th e cro sse d fib re s o f th e op tic n erv e h av e larg e r
dia m e ters, are be tte r m y e lin ate d , a nd tra nsm it se n sory info rm a tio n fa ste r tha n
th e u nc ro ssed fibre s (B ish o p , Je re m y , & L an c e, 1 95 3 ; M a d de ss, 19 7 5 ). In
diu rn al p rim a te s, th ere a re a lso hig he r d e nsities o f p h oto re ce pto rs in th e n asa l
he m ire tin a c om p are d to th e tem po ra l h em ire tin a (C u rc io et a l., 19 8 7 ; P ac k er,
H e n d ric k son , & C u rcio , 1 98 9 ). P h oto rec e pto r d e n sity , w hic h d ec line s w ith
distan c e fro m th e c e ntre of th e fo v ea , d e clin es m ore ste e p ly to th e p e rip h er y in
th e te m p o ra l h e m iretin a th an in th e na sa l h e m iretina (W ik ler, W illiam s, &
R a k ic, 1 9 9 0). G an g lio n ce ll d e nsity also de c re a ses w ith d ista n c e fro m the fo ve a
an d in d iu rn al p rim ates a ste ep e r g ra die nt o f d e cline is fo u nd in th e tem p ora l
he m ire tin a c o m p a red to the n asa l he m ire tin a (W aÈ ssle , G ruÈ n e rt, R oÈ h ren b ec k , &
B o y c ott, 1 9 9 0 ; W ild er, G ruÈ n e rt, L e e , & M artin , 19 9 6 ). A s th ere is ev id en c e fo r a
s tr o n g p o s it iv e c o rr e l a t io n b e t w e e n g a n g l io n c e ll d e n s ity a n d c o r t ic a l
m a g nifica tio n in p rim a tes (W aÈ ssle e t a l., 1 9 9 0 ), in a m o n o cu lar v iew in g
situ atio n th e c o ntra la tera l h e m isp he re re ce ive s q u alita tiv e ly d iffe ren t sen so ry
in fo rm a tio n th an th e ip sila te ral h em isp h ere . It h as b ee n su g ge sted th a t the se
m o rp ho lo gic a l a sy m m e tries m a y resu lt in a stro n ge r c ortica l rep rese n ta tion in
th e h em isp h e re c on tralate ral to a v ie w in g e ye (R o w e , 1 9 9 1; W e isz , B a la z s, &
A d a m , 19 9 4 ), b ec au se th is he m isp h ere re c eiv e s se n so ry info rm a tio n v ia th e
cro sse d o p tic ne rv es o rig in atin g in th e n a sal h em ire tin a . A lso , it ha s b ee n
de m on strate d in h u m a n s th at the re g io n of h ig h est im a ge q ua lity is lo c a te d
ap p ro xim a te ly 5 ° n asa l to the fo v ea (Jen n ing s & C ha rm a n , 1 9 81 ), a n d th a t th e
de c lin e in im ag e qu a lity w ith e cc en tric ity is slo w er in th e n a sal h em ire tin a
co m pa red to th e te m p o ra l he m ire tin a in m a c aq u es (M erig an & K a tz , 1 9 90 ).
T h u s fib re s orig in a tin g in th e n a sal h em ire tin a w ou ld tra n sm it a h igh e r-q u a lity
im ag e to the c on tralate ral h em isph e re . E y e p refe ren c e m ay , th ere fo re , b e
in d ic a tiv e o f late ralisation o f th e h e m isp h ere s fo r p erc ep tu al p ro c essin g (R og e rs
et al., 1 99 4 ).
S tu d ies o f sim p le re a ction tim e in hu m a ns h av e sh ow n th a t in fo rm a tio n
fro m the d o m in an t o r pre fe rre d ey e is p roc esse d m ore rap id ly th an in fo rm a tio n
 EYE PREFERENCES IN MARMOSETS 111

from th e no n p re fe rre d ey e (M in u cc i & C o n n o rs, 1 96 4 ). C o re n an d P o ra c

(19 7 6 ) rep o rted , fo r h u m a n sub jec ts, th at th ere is size e nh a nc e m e n t o f a
stim ulu s p re se nte d to th e d o m ina n t e y e, b u t n o t o f a stim u lus p re se n ted to th e
n on p re fe rred ey e . T h e im a g e from th e d om in an t e y e h as also be en de sc ribe d a s
cle are r, m o re sa tia ted in co lou r (P ora c & C or en , 1 98 1 ) an d a s pro d u cin g a
m o re stab le retina l im a g e tha t tak e s lo ng e r to fa de (P o ra c & C o ren , 1 98 2 ;
W a d e, 1 9 75 ). T h ese o b serv a tio n s in dic a te th at th e re are a sy m m etries in th e
efficie nc y o f p roc e ssing a n d p erc ep tion b e tw e e n th e p refe rre d a n d n o n pre fe rre d
ey e s.
V ery fe w stud ies of fun c tio n al la te ralisa tio n in n on h u m a n p rim ate s h av e
ex a m in e d e y e p re fe re nc e. T h ere ha s b ee n o nly o ne stu dy o f e ye p refe re n c e in
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p ro sim ia n s. R o g e rs e t a l. (1 99 4 ) e x am in ed th e ey e p re fe re n ce s of the sm a ll-e are d

b ush b a by , O tole m u r g arn e ttii. T h e fiv e sub jec ts teste d d isp lay ed sig n ifica n t left-
ey e p re fere n ce s w h e n v iew in g fo od . H o w ev e r, w h en th ree of th e b ush b a bie s
w ere p rese n ted w ith a m ore a rou sin g stim u lu s (th eir b ab ie s h eld in th e
ex p e rim en ter’ s h an d ) th e ir ey e pre fere n ce s ch a ng e d; tw o su b je cts d isp lay e d
eq u a l u se o f bo th e y es w h ile th e th ird su b je ct d isp la ye d a w ea k er left-ey e
p re fere n c e. T h is su g g es ts th at e ye p refe ren c e m ay b e affe cte d b y aro u sa l a nd /o r
the stim ulu s b ein g vie w e d .
T h e re h a v e b ee n so m e stu d ie s ex a m in in g e y e p re fere n ce in O ld W o rld
p rim ate s bu t the re is n o cle ar info rm atio n a s to w h eth e r th ese sp ec ie s h a v e le ft o r
rig h t e y ed n e ss. C o le (1 9 5 7) fo u nd a te nd e nc y to w a rds rig ht p re fe re n ce s in a
g ro u p o f se v e n a d u lt m a c aq u es , M a c a ca n e m e strin a , te ste d b y s c o rin g
m o n o cu lar lo o k in g th rou g h a tub e . B y c o ntrast, K rup e r, B o y le , an d P a tto n
(19 6 6 ) fou n d left ey e dn e ss in a g ro up o f 1 9 n aiv e, im m a ture m a c aq u e s (M a ca c a
m ula tta ) a nd n o b ias fo r eith er e y e in a gro u p of sev en a d u lt m a c aq u e s. S m ith
(19 7 0 ) a nd K o u n in (1 9 3 8 ) also re p orte d le ft-e y e p re fere n c es, ea c h testing on e
im m a tur e rh esu s m a c aq u e .
T h e re h a ve b e en o n ly tw o stu d ies of e y e p re fe re nc e in th e pla ty rrh in e sp ec ies.
K ou n in (1 93 8 ) m ea su red ey e p refe ren c es in th re e im m a ture N e w W o rld
p rim ate s (C e bu s sp p). W he n th e su b jec ts h e ard a ``s lig h t’ ’ no ise th ey w ere
req u ire d to loo k th rou g h a tub e a t a pie ce o f fo od . T h e y d isp lay e d rig h t-ey e
p re fere n c es. M c F erra n (1 99 2 ) in v e stig ate d e ye p re fe ren ce in a g ro up o f 3 8
co tton -to p ta m a rin s, S a gu in u s oe d ipu s, a lso lo ok in g m o no cu larly a t fo o d
thro u gh a v iew in g ho le. O n ly se v en o f th e 3 8 sub je cts te sted d isp lay e d
sign ific a nt e ye p re fe ren c es, six p re fe rrin g th e le ft e ye a nd o n e the rig ht.
A lth o ug h th e se re sults m ay ind ica te in co n siste n cy in the p re se n ce an d
d ire ction o f latera l b ias for ey e p refe ren c e am o n g N e w W o rld prim a tes,
m e tho d o log ica l d iffere n ce s be tw ee n th e tw o e xp e rim en ts m a y h a v e a ffe c te d th e
resu lts. F or e xa m p le , th e intro du c tio n of a n a u d ito ry c u e m ay h av e in flu e n ce d
the re sults o f K o u n in (1 9 3 8) b y ind ire ctly c o nd itio n in g th e sub jec ts’ resp o n ses.
In M cF e rra n ’ s stu dy , testin g in a g ro up situ atio n m a y h av e in flu en c ed th e
resu lts, as a ro us al m a y ha v e b e en in du c ed b y co m pe tition b e tw e en su b jec ts a t
 112 HOOK-COSTIGAN AND ROGERS

th e pe e ph o le. In fac t, M cF e rran (1 99 2 , p .3 8 ) states th at in tere st in the p ee p ho le

w a s h ig h w ith se ve ral a nim als jostlin g for a n op p o rtu n ity to v iew .
W e d ec id e d to in v e stig ate e ye p refe ren c e s in an o th e r sp ec ies of N e w W o rld
prim ate, C a llithrix ja cc h u s, ta k in g in to a cc o un t th e e ffe c ts o f a g e, a rou sa l, th e
ha n d u sed by th e e xp e rim en te r to pre sen t th e re w a rd, a nd th e h an d p re fe ren c es
of th e m arm ose ts.
A re lation sh ip b etw ee n e ye a n d h a n d pre fere n ce s m ay in dic ate th a t th e sa m e
ev o lutio na ry an d e n viro nm e nta l fa cto rs in flu e nc e b oth form s o f la tera lity , b u t
sev e ral p rev io us stu d ies h a ve fo un d n o co rre la tio n b e tw e en e y e a n d ha n d
pre fe re n ce s in no n h u m a n p rim ate s (C o le , 1 9 57 ; K ru p e r e t al., 19 6 6 ; R og e rs e t
al., 1 99 4 ). E arly stu d ies a lso re po rted n o re latio nsh ip be tw e e n ha n d an d ey e
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p r e fe r e n c e i n h u m a n s ( G r o n w a ll & S a m p s o n , 1 9 7 1 ) b u t m o re r e c e n t
in v e stig atio n s ha v e fou n d a re lation sh ip b etw ee n th e tw o typ e s o f la tera l
pre fe re n ce (B ou ra ssa, M cM a n u s, & B ry d en , 1 9 96 ; B row n & T a y lor, 19 8 8 ;
H a rris & L o ve g rov e , 1 98 4 ; M eta lis & N ie m ic , 19 8 4 ). O ur aim w as to fu rth e r
in v e stig ate the re latio nsh ip b etw e en e y e p refe re n ce a n d h a nd p re fere n ce in a
N e w W o rld p rim a te , b e c a u s e n o s u c h p o t e n t ia l re la ti o n s h ip h a d b e e n
in v e stig ate d in an y N ew W o rld prim a te sp ec ies .

METHOD
Subjects and Housing
A to tal o f 2 1 m a rm o se ts (8 m a les a nd 1 3 fem ales) fro m th re e fam ily g ro u p s w e re
te sted . E a ch fam ily w as h o use d in a sep a ra te ro o m . In itia lly , th e fa m ilie s w e re
ho u sed a s he te ro se x ua l pa irs w ith th eir o ffsp ring . W h en b ree d in g p airs ha d sire d
tw o to th re e se ts of tw in s o r trip lets, th e fa m ilie s w e re se p ara ted in to sin g le-se x
gro u ps . G ro u ps, c o m p risin g tw o to six m arm o sets, w ere h o use d in ca g es
1.8 ´ 1 .5 ´ 1.6 m h ig h . O n ly o ne m a le w as h o use d in d iv id ua lly fro m 3 3 m on ths
of a g e in a c ag e 0 .9 ´ 0 .8 ´ 2 .2 m hig h .
L igh t a n d d a rk cy c le s w e re re gu la te d ; lig h ts o n a t 7.0 0 h o u rs an d o ff at 1 9 .0 0
ho u rs. T h e m a rm o sets w e re ex p ose d to u ltrav io le t lig h t (35 0 ±3 90 n m ) fo r 3 0
m inu te s p er d ay to su p p le m en t V itam in D inta k e. T h e re w as a lso a sk ylig h t in
ea ch roo m to pro v id e a n a tu ra l sou rc e o f su nligh t. T h e m arm ose ts w e re fe d a
va rie ty o f foo d s tw ice da ily a t a pp ro xim ate ly 10 .3 0 a nd 1 5.3 0 h o urs. W ate r w a s
av a ila b le a d lib itu m . R o o m te m p e ratu res w e re m a in ta in e d b etw ee n 2 1 °C a n d
25 °C .

General method

T e sts w e re co n d uc ted in the h o m e ca g es. E a ch sub jec t w a s te sted in d iv id ua lly ,

th e ca g em ate s be in g p artitio n ed into o n e h alf of th e c ag e w h ile th e su b je ct w a s
be in g tes te d .
 EYE PREFERENCES IN MARMOSETS 113

T h e m e th o d u se d ha s b ee n d e scrib ed p rev io us ly b y R o ge rs et a l. (1 9 9 4),

alth ou g h the testin g a p p ara tus w a s m od ifie d for th e sm a lle r siz e of th e m a rm o se ts.
T h e te stin g a p pa ra tu s co n siste d o f a she e t o f g a lv a nise d ste el (5 6 ´ 5 6 cm ) w ith a
stee l b o x (1 5 cm de ep ´ 1 7 cm ´ 1 7 cm ) se c ure d on th e ce n tre ba ck , fa c in g th e
ex p e rim en ter (Fig . 1 ). T h ere w a s a ho rizo n ta l slit (1 cm ´ 0 .5 c m ) in bo th th e fron t
an d ba c k o f th e bo x , allow in g th e sub je cts to lo o k into th e b ox th ro u gh a slit to
v ie w a stim u lu s. T he ex p er im en ter lo o ke d thro u gh th e o th er slit to m o n ito r th e ey e
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FIG. 1. A p p a ra tu s u s e d t o te s t e y e p re fe re n c e . T h e s y m b o ls re fe r to (a ) sh e e t o f g a lv an is e d s te e l, (b )
s te e l b o x s ec u re d o n c e n tre b a c k o f s h e e t, (c ) h in g e d d o o r a llo w in g e x p er i m e n te r to p l a c e h a n d in
b o x , (d ) p a n e l w i th c e n tra ll y lo c a t ed h o le w h ic h w a s s l id in f ro n t o f th e h o riz o n ta l s lit t o a llo w
d e t e rm in a t io n o f m o n o c u l ar v ie w in g p re fe re n ce s in te s ti n g . A s e c o n d s lit w a s l o c a te d o p p o s i te , in th e
fr o n t s id e o f t h e b o x , al lo w in g th e e x p e rim e n te r to m o n i to r th e ey e u se d b y t h e s u b j e ct s , (e ) h a n d le
u s e d to sl id e t h e p a n e l (d ) in fr o n t o f th e h o ri z o n ta l s lit .
 114 HOOK-COSTIGAN AND ROGERS

use d b y th e su b jec t. T h e su bje cts w e re tra ine d to lo o k th ro ug h th e h orizo n ta l slit to

vie w a p ie ce o f b a na n a w ith b o th e y es. A fte r the b in oc u lar g a ze w a s h eld fo r
ap p ro xim a te ly tw o se co n d s, th e su b je ct w a s re w ard e d w ith a p iec e o f b an a na . A
hin g ed d oo r, in th e le ft sid e o f th e b ox , a llo w e d th e ex p er im en ter to h old th e
re w ar d direc tly in fro nt o f th e ho rizo n ta l slit u sin g he r left h an d . T h e
ex p erim en ter’ s h a nd w a s use d to p res en t stim uli be c au se it w as fa m iliar to th e
m a rm o se ts an d b e ca u se it fac ilita ted tra nsfe r o f th e rew a rd th ro u g h th e slit. It
sho u ld be n o te d tha t th e se m a rm o se ts a re v e ry re lax e d in th e p re sen c e o f h u m a n s
an d are v e ry fam ilia r w ith the ex p erim e n te r. T he sub je cts w e re train ed in a
m inim u m o f 20 0 tria ls o v er 5 ±1 0 d ay s.
In the te stin g trials th e slit w as re pla ce d b y a ce n tra lly lo c a te d h o le (0.5 c m
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dia m e ter) by slid ing ac ro ss a pa n el. T h is allo w ed on ly m on o c u la r v ie w in g o f th e

stim ulu s a n d th e ey e u se d to v ie w w a s sc o re d . T h e in ne r c an th i o f th e le ft o r
rig h t e y e w as v isib le w he n a su b jec t lo ok e d th rou g h th e m o n o cu lar v ie w ing h ole
an d th is a llo w ed th e e x pe rim e n te r to d e te rm in e w h ic h e y e w as u sed to vie w a
stim ulu s. M o n oc u lar e y e u se w as no t rew a rd ed . R e w ard e d trials o f b in o c ula r ey e
use w e re ra n d om ly d istrib u te d b etw e en m o no c u la r e ye -u se trials to m a inta in th e
m a rm o se t’ s in tere st in th e ta sk. S u b jec ts w ere re qu ired to disp la y a m in im u m o f
tw o in c id e n ce s o f m o n o cu lar ey e u se b e fore a b in o cu la r e ye -u se tria l w a s
pre se nte d. O n av e ra g e fo u r in c id e n ce s o f no n rew a rd ed m o n o cu la r v iew in g
e v e n ts w e re r e c o rd e d b e fo re a r e w a rd e d b in o cu la r tria l w a s p re se n te d .
R e w a rd in g sub je cts fo r bin o c ula r ey e use o n ly en su red th at e ye p re fe ren c es
w e re n ot c on d itio n ed . T e stin g w as c o nd u c te d o ve r a m inim um o f eig h t d a ys a n d
5±1 5 sco re s w ere co lle c ted in e ac h testing se ssio n . A m in im u m of 1 0 0
m o n o cu la r e y e -u se sc o re s p e r s u b je c t w e re c o lle cte d fo r e a c h stim u lu s
pre se nte d.

Eye Preference and Age

T h e ey e p re fe ren c es o f the m a rm o se ts w e re m on itore d at 3±8 m o n th s (1 5
sub je cts), 1 2 m o n ths (1 3 su b je cts), 15 ±1 8 m o nth s (1 1 sub je cts), a n d 22 m on ths
(1 7 su bje cts). A t ea ch a ge su bje cts w ere re q u ire d to v iew a pie ce o f fa m ilia r fo o d
(b a n an a ). E le ve n su bje c ts w ere tested a t a ll fo ur a g es.

Presentation of Different Stimuli

O n co m ple tio n o f th e e ye -p refe ren c e te sts a t th e v ariou s ag e s, a d d itio na l
m o n o cu lar v ie w in g te sts w e re c o nd u c te d to d ete rm in e w h eth e r th e stim u lu s
vie w e d in flu e nc e d th e e ye p refe re nc es of th e m a rm o se ts. A tota l of 1 4 a du lt
sub je cts (o lde r th an 22 m o nth s), 4 m a les an d 1 0 fe m a le s, w ere te ste d in tw o te st
series . In th e first te st se rie s fo u r stim u li w e re p re sen ted , a go ld w atc h, a sm all
m irror (6c m ´ 3 cm ), a sm a ll w oo d e n b ee tle w ith m o v in g a pp e nd a g es, a n d a
pie c e o f fa m ilia r fo o d (b an a n a), th e latte r a s a c on trol. F o r th e se co n d te st se rie s,
th re e stim u li w ere p re sen ted , th e e xp e rim e nte r’ s left h an d , a m od e l re sem b lin g
 EYE PREFERENCES IN MARMOSETS 115

tw o re a ring sna k es (sna k e m od e l), a n d, a g ain , a p ie c e o f fa m ilia r fo o d a s a

co n tro l. F o r b o th tes t se rie s the o rd e r o f p re sen tatio n of th e stim u li w a s
ran d o m ise d w ith resp e ct to th e testin g pe rio d a n d for ea ch in div id ua l. A ll stim uli
w ere h eld in the ex p erim e n ter’ s left h a n d, p o sition e d c en trally in th e b o x,
d ire ctly in fro n t o f th e p e ep h o le a nd 5 ±1 5 sco re s w e re c o lle cte d p er d aily te stin g
sessio n . T h e tim e tak e n to c o n du c t th e e ye p re fe re n ce te sts v aried b etw ee n
su bje cts, stim u li, an d a cro ss da y s. T h e aim w as to co lle ct th e re qu ired n u m b e r o f
sco re s pe r d ay irre sp ec tiv e of th e tim e ta k en .
F or e a ch stim u lu s the m ea n d ura tio n b e tw e en in c id e n ce s o f m on o cu la r ey e
u se, ca lc ula te d b y d iv id ing th e to ta l tim e ta ke n to c on d u c t th e test b y the n u m be r
o f m o n o cu lar e y e-u se sco re s, w a s u se d as an in dic ato r o f aro u sa l. V oc a lisa tio n s
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ind ica tive o f a rou sa l (``tsik’ ’ , ``eg g ’ ’ , a nd ``o c k’ ’ : E p p le, 1 9 68 ) w e re also

rec o rd e d fo r ea c h te stin g se ssion . T h e tsik , e g g , a n d o c k v oc a lisatio n s o f th e
co m m o n m arm o se t e vo k e in c rea se d alertn e ss in th e g ro u p a n d are em itte d w h e n
m o b b in g a p o ten tial p re da to r an d du ring h ostile en co u n ters b etw ee n m arm ose ts
(E p p le , 1 9 68 ). T h e refo re, w e re co rd ed a ll o f the v o c alisation s h ea rd in th e
testin g ro om d u rin g a se ssio n . T h e n u m b e r o f v oc a lisa tio n s co u nte d for e ac h
testin g se ssio n w as d ivid ed b y th e to ta l tim e tak e n to c o n du c t th e te st to p ro d uc e
a stan d ard ised v o ca lisa tio n ra te (vo c alisatio ns/se co n d ). B o th d u ra tion s b etw ee n
the e v en ts o f m on o c ula r e y e u se a n d the v o c alisation s m a d e in e a ch te stin g
sessio n w ere d ete rm in ed from p o st h o c v id e o a n aly ses . E ac h te sting se ssion w a s
rec o rd e d u sin g a S o ny H i8 vid eo c am era .

Hand Use of Experimenter

In th e tes ts o u tline d ea rlie r, th e e xp e rim en te r’ s left h an d w a s p la ce d in to th e bo x
to h o ld th e stim ulu s. It w a s rec o gn ise d th at th is m igh t b ias th e ey e pre fere n ce s o f
the m a rm o se ts. T he refo re , w e co n du c te d fu rth er e y e-p ref ere nc e te sts on 1 0
su bje cts (4 m a les , 6 fem a le s) u sin g a re plic a o f the in itia l a p pa ratu s, b u t w ith th e
h in g ed do o r o n the rig ht sid e, a llo w in g th e ex p erim e n te r to u se th e rig h t h a nd to
p re se nt th e p iec e of fa m iliar foo d . M o n o cu lar e y e u se w he n v ie w ing a p ie ce o f
fam ilia r fo o d w a s sco re d.
A pe rio d o f 6±1 4 m o n th s sep ara ted th e tests in w hic h th e stim ulu s w as h e ld in
the e xp e rim e nte r’ s le ft h a nd a n d th o se in w h ic h it w a s h eld in th e rig h t h a n d.
H ow e v e r, it re m a ine d p o ssib le th a t the m arm o sets’ e ye p refe ren c es w ere
ind u ce d b y ra p id le a rn in g in the first tests, in w h ich th e e x pe rim e nte r use d th e
left h a nd . T o in ve stig a te the po ssib ility o f su ch rap id ly ac q u ire d late ralisatio n,
w e co m p are d th e first a n d se co n d 5 0 in c id e n ce s o f m o n o cu lar ey e u se w ithin
ea c h te st. T h is pro c ed u re w a s ap p lie d fo r v ie w in g e ac h o f the stim u li.

Simple Hand-preference Tests

H an d p re fere nc e s w e re a ssesse d a t 3 ±8 , 1 2 , 18 , a n d 2 2 m o n ths o f a g e,
co rresp o n d in g to th e a ge s a t w h ic h ey e pre fere n ce s w ere m e asu re d, u sin g th e
 116 HOOK-COSTIGAN AND ROGERS

m e th o d o f H oo k -C ostig an a n d R o g ers (19 9 5 ). H a nd u se w as re c o rd e d w he n a

sub je ct h e ld a p ie ce o f fo od in on e h a nd an d to o k it to th e m o uth . A m in im u m o f
10 0 un im a nu al h o ld in g sc or es pe r sub jec t w e re co lle c ted o v e r a m in im u m o f
eig h t d a ys. S am p le siz e s w e re th e sa m e a s in th e ey e -p re fere n ce tests.

Statistical Analysis
O n e -sa m p le ru n tests w ere a pp lie d to e sta b lish w h e th e r su cc essiv e o b serv atio n s
of ey e a n d h a nd use w e re ind e p en d en t o r oc c urre d in b o u ts. T h e seq u en c es o f
sco re s on e ac h d a y o f testing a n d fo r ea ch su b jec t w e re e xa m in e d.
T h e sig n ifica n ce o f th e e y e-p refe ren c e sc o re de te rm ine d for ea c h in div id ua l
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w a s asse ssed u sin g a z-sco re te st (E d w a rds, 1 9 63 ). T h e rela tiv e bia s w a s

de te rm ine d b y c om p arin g th e z-sc o re c a lcu lated w ith re spe c t to th e e xp e c te d
bin o m ia l d istrib ution o f tota l re spo n ses . T h e z-sco re e q ua tion is, z = (X -M )/
Ö N pq , in w hic h X = th e n u m b e r o f le ft o r rig h t re sp on se s, N = th e tota l n u m b e r
of resp o nse s, M = N /2 , a n d p a nd q = 0.5 . A n in d iv id ua l w a s c on sid ere d to h av e a
sig n ific an t ey e o r h a nd p refe ren c e if th e z sco re w a s ø 1 .9 6 (P ø .0 5 ).
C h i-s qu a red (if nu m b er of su bje cts ù 1 5) a nd b in o m ia l (if n u m b e r o f
sub je cts < 1 5) sign ifica nc e te sts w e re u sed to d ete rm in e rela tiv e b ias o f
distrib u tion o f rig h t- a nd left-ey e pre fere n ce s w ith in g ro up s.
R e la tion sh ips b etw ee n p re fere nc e s d isp la y ed ac ross a ge , for ey e a nd ha n d
pre fe re n ce s, a n d a c ro ss stim u li w e re a n aly sed u sing th e F rie d m an statistic
(S ie g el & C aste lla n , 1 9 9 4). S tatistic ally sig nific a n t differe n ce s in g ro up m e d ia n s
w e re ac c ep ted if P ø .0 5 . S ig nifica n t re su lts o f F ried m an tests w e re fu rth e r
an a lyse d usin g p o st ho c W ilco x on sig n ed ran k an a ly se s.
T h e Sp e arm an ra n k c o rrela tio n co e fficie n t w a s c alcu la te d to d e te rm in e th e
de g ree o f rela tio ns h ip b etw e en e y e p re fe ren ce s d isp lay e d w he n the e xp e rim e nte r
use d th e le ft a nd rig h t h an d to p rese nt th e stim u lu s. T h e S p e arm an ra n k
co rre la tio n ca lcu lation w a s a lso use d to d ete rm ine the d eg re e o f re latio nsh ip
be tw e e n ey e pre fere n ce an d h an d pre fere n ce .

RESULTS
Eye Preference and Age
T h e m a rm o sets d isp lay e d a gro u p b ias to w ard s rig h t e y ed n e ss fo r vie w in g th e
fa m iliar fo o d a t the fo u r ag e s e x am in e d (see F ig . 2). A t 3 ±8 m o nth s o f a g e, 1 2 o f
th e 15 m a rm o se ts te ste d disp la y e d sig n ifica n t righ t-e ye p re fere n ce s, tw o
pre fe rre d th e le ft e ye , a n d o n e d id no t d isp lay a p re fe ren c e (z-sc o re test). T hu s,
th e re w as a g ro up b ia s to w ard s rig h t e y ed n ess ( c = 1 4.8 , P ø .0 05 ). M o st o f
2

th e se su bje cts m ain tain ed th e se ey e p refe ren c es at 1 2 m on th s o f a ge a n d the re

w a s sign ifica n t righ t-e ye d n ess in th e g ro u p (b in om ia l, P = .0 3 8 , F ig . 2).
A sign ifica n t righ t-ey e bia s w a s a lso fo u nd a t the g ro u p le v el at 15 ±1 8
m o n th s (Fig . 2 ; bin o m ia l: P = .0 2 2 ) a n d at 2 2 m o nth s ( c = 1 3 .2 4 , P ø .0 01 ).
2
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FIG. 2. P e rce n t l e ft-e y e u s e w h e n v ie w in g b a n a n a . T h e p er ce n ta g e o f le ft-e y e u s e fo r e a c h in d iv id u a l w h e n v i e w i n g b a n a n a is

p l o t te d fo r th e fo u r a g e s te s te d . A g e s a re d e p i ct e d o n th e y -a x i s. T h e o p e n s y m b o ls ( s ) in d i c ate s ig n i fic a n c e at P ø .0 5 , fi lle d
s y m b o l s ( ) re p re s e n t n o n s ig n if ica n t s c o re s . 2 x a n d 3 x in d ic a t e th a t 2 a n d 3 s u b j e ct s, re sp e ct iv e ly , h a d t h e p e rc e n t e y e -
·

117
p r efe re n ce s c o r e rep re s en te d b y th e s n e x t to th e se s y m b o ls.
 118 HOOK-COSTIGAN AND ROGERS

O n e in d ivid u al d isp lay ing a left-e ye p re fe re n ce a t 3 ±8 m on th s h a d n o p re fe re nc e

at 1 2 an d 1 5±1 8 m o n th s, a nd a righ t-ey e p re fere n ce a t 2 2 m on th s. O n ly o n e
sub je ct disp lay e d a co n sisten t le ft-e y e p refe ren c e a c ro ss the fo ur a g e tests (F ig .
2). T hu s, irre sp ec tiv e o f a ge , 2 0 of th e 2 1 su bje cts te ste d d isp la y ed sig n ifica n t
rig h t-e y e p refe re nc e s an d on e sub je ct w a s left-ey e p re fere n t.
E lev e n su bje c ts w e re teste d ac ro ss the fo u r a g e te sts. T h e F rie d m a n statistic
sho w e d th at the re w a s n o sig n ific a nt ch a n ge in e ye p re fere n ce w ith ag e
( c = 4 .3 2 , P = .2 2 9 ).
2

Presentation of Different Stimuli

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T h e d a ta co lle c te d to m ea su re a ro u sa l w ere an a ly se d se pa rate ly fo r th e tw o te st

series (F ig. 3 ). T h e m e an d ura tion b etw ee n m o no c u lar vie w in g ev e nts an d th e
nu m b er o f a ro u se d vo c alisatio ns pe r se co n d w ere co m p are d ac ro ss stim uli (F ig .
3). In th e first te st serie s, a slig h t b u t sig n ific an t d iffere nc e in th e m e an du ratio n
be tw e e n m o no c u lar vie w in g ev e nts w as fo un d (F rie d m a n statistic , c = 11 .5 7 ,
2

P = .0 09 ; F ig . 3 a ). It w as de m on strate d tha t the m arm ose ts d ispla y ed lo ng e r

in terv a ls be tw ee n vie w ing e ve n ts w he n the m irro r o r be etle w e re p re se nte d , in
+
co m pa rison to p rese n ta tio n o f the g o ld w a tch (m irro r an d w a tc h , T = 11 ,
+
P = .0 3 6 , b e e tle an d w a tc h , T = 1 1 , P = .0 1 9 ). H o w e v e r, th e re w e re n o
diffe re n ce s in th e m ea n du ratio n be tw e e n m o no c u la r vie w in g ev e n ts fo r th e
+
ba n an a co m p a red to th e o the r th ree stim u li (b an a n a an d m irro r, T = 4 , P = .0 96 ,
+ +
ba n an a a n d w a tc h , T = 5 , P = .1 7 7 , b a n an a an d b ee tle , T = 3 , P = .0 6 4 ). T h ere
w a s n o effec t of stim u lu s o n th e vo c alisatio n ra te s fou n d in the first test series
(F ried m a n sta tistic, c = 1.2 1 , P = .7 5 12 ; F ig . 3 b). T h u s, a lth o u gh th e re m ig h t
2

ha v e b e en sligh t d iffe ren c es in m o tiv a tio n fo r v iew in g th e stim u li p re sen te d in

th is first series , th e re w e re n o m ar ke d d iffere n ce s in a ro u sa l e x pre sse d
co lle c tiv ely in v iew in g p a tte rn o r v oc a lisa tio n ra te.
In the sec o nd test se rie s, the sna k e m od e l w a s stro n gly aro u sing (F ig s. 3 a
an d b). C o m pa rison b etw ee n the stim uli pre se nte d in th e se c on d test series
(e x p erim e n ter’ s ha n d , b an an a , a n d sn a ke m o d el) re v ea le d sign ific a n t e ffe cts o f
stim ulu s fo r b oth the m ea n d u ratio n b e tw e e n m o n o cu lar v iew in g ev e nts a n d
vo c alisa tio n rate ( c = 16 .7 1 , P = .0 0 02 , c = 11 .1 1 , P = .0 0 39 re sp ec tiv e ly).
2 2

P re se n ta tio n o f th e sna k e m o d el re su lte d in sig n ific a ntly lo n g er d u ratio n s

be tw e e n m o n o cu lar v ie w in g e v en ts w h e n co m pa red to th e ba n an a an d ha n d
+
(T = 1 3, 1 3 , re spe c tiv e ly , P < .0 0 2 ; F ig. 3a ) an d a hig h er rate o f p ro d u cin g
+
a r o u s e d v o c a l is a ti o n s (T = 1 0 , 1 2 , r e s p e c t iv e l y , P < .0 5 ; F i g . 3 b ). N o
diffe re n ce s w ere fo u n d fo r eithe r m e a sure o f a ro u sal w h en the h an d a n d
ba n an a w ere p re sen ted (m e an d u ra tion b etw ee n m o no c u lar vie w in g ev e nts,
+ +
T = 1 0 , P = .2 20 9 , v oc a lisa tio n ra te , T = 5 , P = .7 2 9 9). It is no ted , in F ig. 3 ,
th a t hig h er v o ca lisation ra tes w e re fou n d in th e sec o n d te st serie s c om p are d
w ith the first. T h e rate of p ro d uc in g a ro us ed v o ca lisa tio n s in the se co n d series
co u ld h a ve b ee n ca u se d b y the ran d o m p rese n tatio n o f th e ba n an a , h a nd , a n d
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FIG. 3. (a ) T h e m e a n d u ra t io n b e tw e e n m o n o c u l ar v ie w in g e v e n t s d e te rm in ed a t t h e g ro u p le v el is
p l o t te d . T h e m e a n d u ra t io n o f t h e in te rv a l b e t w e e n m o n o c u la r v i ew i n g e v e n t s w a s d e te rm i n e d fo r
e a ch in d i v i d u a l a n d ea c h st im u lu s. In d iv i d u a l m e an d u ra ti o n s b e tw e e n m o n o c u la r v ie w in g e v e n ts
w e re u s e d to c a lc u la te th e g ro u p m ea n s ( ± S E M ) d e p ic te d in th e g ra p h . (b ) T h e ra te s o f p ro d u c in g
a ro u s e d v o c a lis a ti o n s w e re d e te rm in e d fo r e a c h s tim u lu s , a n d fo r e ac h in d iv id u a l, a n d a re p re s e n t e d
a s g ro u p m e a n s w ith s ta n d a rd e rr o rs (s ee 3 a ).

119
 120 HOOK-COSTIGAN AND ROGERS

sna k e m o d el. A ro u sa l in d u ce d b y v ie w in g th e sna k e m o d e l m ay h a v e c a rrie d

ov e r so th at th ere w as inc re ase d pro d u ctio n o f aro u se d v o ca lisation s to th e
oth e r stim u li.
A n a lysis o f th e d istribu tio n o f e ye p refe ren c es fo r v ie w ing th e stim u li th a t d id
no t e lic it a rou se d b eh a vio ur (th e w atch , m irror, w oo d e n b ee tle , e x pe rim e nte r’ s
ha n d, an d th e p iec e o f fa m ilia r fo o d ) d em o n stra te d n o diffe re n ce s be tw e e n
m e d ia n s (F ried m a n sta tistic, c = 8 .8 6, P = .1 1 4 9 ). O f th e 1 4 su b je c ts tested , 1 2
2

to 1 3 d isp la y ed sign ifica n t rig h t-e y e p re fere n ce s fo r vie w ing e ac h o f the se

stim uli (se e F ig . 4 ). F o r e ac h stim u lus th e re w a s a sign ific a n t g ro up b ias fo r
rig h t-e y ed n e ss (b in om ial: P < .0 02 , .0 0 4 , .0 06 ).
W h en th e m arm o sets w ere p rese n te d w ith the sn ak e m o d e l, th re e su b jec ts
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disp lay e d sig nific a n t le ft-e y e p re fe re n c es, e ig h t d id n ot disp la y a pre fere nc e fo r

eithe r ey e , a n d th ree ha d a rig ht-ey e p re fere n ce . T h us, no g rou p b ia s tow a rd s
eithe r lef t o r rig ht e y ed n e ss w as fou n d w h en th e sna k e m o d el w a s v iew e d
(b in o m ial: P = .6 8 8 ). C om p ariso n b e tw e en th e e y e p refe re nc es d ispla y ed w h e n
vie w in g the n eu tral (no t aro u sin g ) stim u li an d tho se disp lay e d w h e n v iew in g th e
sna k e m od e l re ve a le d th at aro u sal affe cte d the distrib u tion of ey e p re fere nc e s a t
th e g rou p le v el (F ried m an sta tistic, c = 3 1.9 9 , P < .0 0 0 1). S u bse q u en t p a ire d
2

co m pa rison s, usin g W ilc o x on sig ne d ran k tests, b e tw e e n th e e y e p re fe ren c es

disp lay e d w h e n v iew in g th e sn ak e m o d el an d for v ie w in g e ac h o f th e n e u tra l
+
stim uli rev e ale d sig n ific an t differ en c es in all ca se s (T = 1 4, 1 3, 13 , 12 , 14 ,
re sp e ctive ly, P < .0 0 0 5 ). T h is m e an s th a t w he n th e sn ak e m o d e l w as p re se n te d
th e re w a s a sign ifica n t sh ift a w a y from rig h t-e y e p re fe ren c es .

Hand Use of Experimenter

T h e h a n d u se d b y th e ex p erim e n ter to p rese n t th e rew a rd did no t affe ct th e ey e

pre fe re n ce s o f th e m a rm o se ts. T h e re w a s a stro n g c o rre la tio n be tw e e n th e
pre fe re n ce s d isp la y ed w h e n th e ex p e rim en ter u se d the le ft v e rsu s rig h t ha n d
(r s = 0 .9 1 , P = .0 06 2 ). T h e refo re, th e la te ral bia s o f th e ex p erim e nt er a n d te stin g
ap p ara tus d id no t affe ct th e ey e pre fere n ce s o f th e m arm ose ts.
T h e d a ta w ere in v estig ated fo r p o ssible effe cts o f le arn in g on e y e p refe re n ce s.
F irst, co m p a riso n w a s m ad e b etw ee n th e p e rce n t le ft-e y e u se d isp lay e d w he n
vie w in g th e ba n an a in th e first 5 0 an d th e la st 5 0 sco re s of e ac h te st (F ig . 5 a).
In d iv id u al W ilc ox o n sig n e d ra nk s tes ts re ve a le d no sig nifica n t d iffe ren c e s
be tw e e n p erc en t le ft-ey e use in th e first a nd la st 50 sc or es a t 3 ±8 , 1 2 , a n d 15 ±1 8
+
m o n th s (T = 7 , 6, 4 , resp e ctive ly , P = .5 7 5 6 , .7 7 94 , .8 2 3 1 ), b u t th ere w a s a
te n d en c y fo r a low e r pe rce n t le ft-e y e u se in th e first 5 0 sco re s a t 22 m on ths
+
(T = 5 , P = .0 5 6 8 ). If this re p rese n ts w ith in -test le arn in g, it is a m in o r e ffe c t, a s
th e m ea n p erc en t left-ey e u se ch a n ge s b y o n ly 5 % . T h e first a nd la st 5 0 sco re s
w e re c o m p a re d for a ll of the oth e r stim u li, in clu d in g th e ex tra tests w ith b a na n a
(F ig . 5 b ). T h e re w ere no sign ific a nt d iffe ren c es b etw ee n p erc en t le ft-e ye u se in
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FIG. 4. P e rc e n t le ft -e y e u s e w ith p re s en ta ti o n o f d iffe re n t s ti m u li. T h e p e rc en ta g e o f le ft -e y e u s e (% L ) fo r v ie w in g a v a ri et y

o f s tim u li is p lo t te d o n th e x a x i s fo r e a c h i n d i v i d u a l ( s ) . S tim u li a re p l o t te d o n th e y a x is a n d th e te s t s e ri e s a re d e n o te d a s
o u tli n e d in t e x t . T h e o p e n s y m b o ls ( s ) in d ic a te s ig n ific a n c e a t P ø .0 5 , fill e d s y m b o l s ( ) re p re se n t n o n si g n ific a n t s c o re s .

121
·
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FIG. 5. C o m p a ris o n b e t w e e n t h e p e rc e n t le ft-e y e u s e i n th e firs t a n d la s t 5 0 in c id en c e s o f

m o n o c u la r e y e -u s e a t e a ch o f t h e ag e s t es t ed . ( a) T h e a g e a t w h ic h th e e y e p re fe re n c e te s ts w e re
c o n d u c te d a re p re s e n te d o n th e x -a x i s a n d t h e p e rc e n t l ef t -e y e u s e i s d e n o te d o n th e y -a x is . T h e o p e n
c o lu m n s re p r es e n t t h e m e an p e rc e n t le ft -e y e u s e ( ± S E M ) fo r th e fi rs t 5 0 i n c id e n ce s o f m o n o c u la r
e y e u s e in e a c h te s t, an d th e fi lle d c o l u m n s th e m e a n p e rc e n ta g e o f l ef t-e y e u s e ( ± S E M ) in t h e l a st
5 0 s c o re s o f e a c h te s t. (b ) C o m p a ri so n b e tw e e n p er ce n t le ft -e y e u s e in th e firs t 5 0 i n c i d e n c es o f
m o n o c u la r e y e u s e a n d th e p e rc e n t le ft- ey e u se in th e la s t 5 0 s c o re s w h e n v ie w in g d iffe re n t s ti m u l i
a re p re s e n te d (s e e 5 a ). T h e s tim u lu s v ie w e d i s d e n o t e d o n th e x -a x is .

122
 EYE PREFERENCES IN MARMOSETS 123
+
the first a nd la st 5 0 sc o re s, e xc e pt fo r th e m irro r (m irro r, T = 2 , P = .0 1 7 , in all
o th e r ca ses P > .0 5 ) b ut, a g ain , this is a m in o r ch a ng e o f a p pro x im a te ly 5 % .

Simple Hand Preference Tests

A lth o u gh th e m arm o sets d ispla y ed sign ific a nt in d iv id ua l h an d p re fere n c es fo r
h old in g fo o d, th e re w a s n o e vid en c e fo r h an d ed n e ss at th e g ro u p lev e l (F ig. 6 ).
O f the 15 m a rm o se ts, teste d a t 3 ±8 m o n ths, 6 su b jec ts d isp la y ed sig n ific an t
rig h t-h a n d p refe ren c es, 7 p re ferre d the le ft h an d , an d 2 did n o t d ispla y a
p re fere n c e fo r e ith er h a nd . A ll o f th e su b jec ts tested a g ain a t 12 m on th s
m a inta ine d th eir h an d pre fere n ce s (F ig . 6 ). T he 11 su b je cts teste d a t b o th 1 8 an d
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2 2 m o nth s disp lay e d th e sa m e h an d p re fere n c es a t e ac h ag e (F ig . 6 ).

U sin g th e h a n d-p re fe ren c e sc ore s o f 1 1 su b je cts e x am in ed a t all fo ur ag e s, it
w as fo un d tha t no sign ific a n t ch a n ge s in h a nd p refe re n c es o cc u rred w ith ag e
(F rie dm a n sta tistic, c = 1.0 5 7 , P = .7 87 ). H o w e ve r, a t 3±8 m o n ths, tw o su b je cts
2

(no t inc lud e d in the F ried m a n an a lysis a s th ey h a v e n o t y e t b e en teste d a t a ll fo u r

ag e s) ex h ib ite d n o sign ifica n t h a nd p re fe re nc e s (Fig . 6 ), an d a t 2 2 m o n ths,
an o the r thre e su b jec ts h a d ha n d-p re fere nc e sco res b e tw e e n 4 5 % a n d 6 1 % (tw o
n on sig n ifica n t). T h e se th ree su b jec ts w e re fou n d in g m e m b e rs o f the c olo n y an d
w ere te sted fo r ey e pre fere n ce o nly at 2 2 m o nth s. W e sug g es t th at d iffe ren t
ex p e rie n ce s m a y h av e a ffec ted th e d e ve lo p m e nt o f th eir h an d pre fe re n ce s, a s
the y h a d be e n ra ise d w ith le ss m ate rna l c o nta ct a n d w ith fe w er o pp o rtu n itie s to
clim b co m pa red to th e o th er m e m b e rs o f o u r c olo n y. H o w e ve r, as the fo c us o f
this stu d y w a s o n ey e p refe ren c e, w h ich d id no t ap p ea r to be affec te d b y th ese
ex p e rie n ce s, th e su b jec ts w e re in clu de d in o u r an aly ses .
E y e a nd h a n d p refe re n ce sco res co u ld no t b e c o rre late d at 1 2 a n d 15 ±1 8
m o n th s o f ag e be ca u se the da ta w ere c lu ste red ; the re w a s a b im o d al d istrib u tio n
o f h a n d pre fere n ce s. H o w ev e r, a t 3±8 m on ths an d 2 2 m o n th s o f a ge , w ith th e
ad d itio n al sub je cts ju st disc usse d , the ha n d- pre fere n ce da ta w ere c o n tin u ou sly
d istrib u te d. T h e da ta co lle c ted a t th ese tw o a g es w ere teste d fo r c orre latio n an d
n o relatio n ship b e tw e en e y e p re fere n c e an d h a nd p re fere nc e fo r fo od h o ldin g
w as fo un d at eith er ag e (3 ±8 m ths, r s = 0 .1 4 , P = .6 0 , 2 2 m ths, r s = 0 .0 1 , P = .9 6).

Runs Analysis
O nly 2 .2 % o f th e d aily seq u e nc e s o f le ft/righ t m o n o c ula r vie w in g e ve n ts w ere
n on ra n do m (i.e . 1 1 o f the 5 00 se qu e n ce s ex a m in ed ). A g rea te r n um b er o f ru n s
tha n e xp e cte d b y ch a nc e w as pre sen t in 1 .8 % o f the d a ily se qu e n ce s, in d ic a tin g
altern a tin g u se of th e le ft an d righ t e y e, an d fe w er ru n s th an e xp e c te d b y ch a nc e
o cc u rre d in 0 .4 % of th e se q ue n c es, in dic ativ e of b ou ts o f lo o k in g w ith th e le ft o r
rig h t e y e. A n a ly sis of the tota l se q ue n ce s for e ac h su b jec t sh o w ed tha t th e sin gle
an o m a lo us d a y s did no t affe ct the resu lts of th e c o m p le te te sts. N o n e o f th e to ta l
seq u e n ce an a ly se s re v e aled n o n ra n d o m seq u e nc e s.
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124
FIG. 6. S im p le h a n d p re fer en c e s a c ro s s a g e . T h e p e rce n ta g e o f le ft-h an d u s e fo r f o o d h o l d in g a t 3 ±8 , 1 2 , 1 8 , a n d 2 2
m o n th s o f a g e is p lo tte d fo r e a c h in d i v id u a l ( s ). A g e s a re d e p ic t e d o n th e y -a x is . T h e o p e n s y m b o ls ( s ) in d ic a te
s ig n i fic an c e a t P ø .0 5 , f ill ed s y m b o ls ( ) re p re s e n t n o n s ig n if ica n t s c o re s . 2 x a n d 3 x in d ic a t e th a t 2 a n d 3 su b je ct s ,
· s
re s p e c tiv e ly , h a d th e p e rc e n t e y e -p ref ere n c e sc o re re p re s e n te d b y th e n e x t to th e s e s y m b o ls .
 EYE PREFERENCES IN MARMOSETS 125

F or th e h a nd p re fe re n ce te sts, o n ly 0 .8 % o f d a ily se qu e n ce s w e re n on ra n do m
(i.e. 4 o f the 5 0 0 se qu e nc e s a na lyse d ). A ll o f the n o n ra n d om se q ue n ce s w e re du e
to a g rea ter n um b er o f run s tha n ex p ec ted by ch a nc e , in d ica ting altern atio n o f
left- a nd rig h t-ha n d u se . A g a in , n o n e o f the to ta l te st a n aly ses re ve a le d
n on ra n do m se q ue n c es. W e su g g est tha t th e lo w in c id e n ce o f n on ra n do m
seq u e n ce s fo u nd for bo th th e ey e -p re fe re n ce a n d h a nd -p refe ren c e tests m igh t b e
ex p e cte d b y ch a nc e .

DISCUSSION
O f th e 2 1 m a rm o sets tes te d , 2 0 d isp lay e d sig n ific an t righ t-ey e p re fere n c es fo r
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v ie w in g a pie ce of fa m ilia r fo o d he ld in the ex p er im en ter’ s h a nd , th e re m a inin g

m a rm o se t d isp la y in g a sig nifica n t left-ey e p re fere n ce . T h e fin d in g o f a rig h t-ey e
b ia s w ithin th e m a rm o se t gro u p c on c u rs w ith th e find in g s o f K ou n in (1 93 8 ) fo r
ca p u ch in s. H ow e ve r, M cF e rra n (19 9 2 ) re po rte d m o re le ft- th a n rig h t-ey e
p re fere n c es fo r co tto n -top tam ar in s, a lth o ug h m os t su b jec ts d isp la y e d n o
p re fere n c e fo r e ith er ey e . A s M c F erra n (1 99 2 ) teste d th e tam arin s in a g ro u p
situ a tio n , co m pe tition b e tw e en su b je cts a s th e y trie d to lo o k th rou g h th e
m o n o cu lar v iew in g h o le m ay h av e in cre ase d a ro u sa l. W e h a ve e v id e nc e
ind ica ting th a t in cre ase d aro u sal le v e ls in M cF e rra n ’ s stu d y m ay e xp lain th e
d iffe re n t resu lt, a s w e fo u n d th a t v iew in g an a rou sin g stim ulu s c au se d th e
m a rm o se ts to sh ift aw a y from righ t-ey e d ne ss. T h is o cc u rre d w h e n th e y w ere
p re se nte d w ith the m o d el re se m b lin g tw o re ar in g sn a ke s. T he sh ift a w ay from
rig h t-e y e p refe re nc e s w h e n v ie w in g a fe ar -in d uc ing stim u lu s m a y re fle c t a
co n flic t b e tw e e n u se o f th e p refe rred ey e a n d u se o f th e rig h t h em isph e re w h ic h
is d o m in an t for th e p ro du c tio n o f e m o tion a l re sp on se s. T h ere is e vid en c e of rig h t
h em isp h e re d om ina n c e fo r th e m ed iation o f em otion a l re sp o nse s in m a n y
sp ec ie s. F o r e x am p le , D e ck e l (19 9 5 ) ha s fo un d a left-ey e (rig h t h e m isp h ere ) bia s
for th e m e d ia tio n o f ag g re ssiv e b eh a vio u rs in th e liz ard (A no lis) a n d rig h t
h em isp h e re co n tro l fo r n eg a tiv e e m o tio n a l resp o nse s h as be en sho w n in c h ic k s
(P hillip s & Y o un g re n, 1 9 8 6), rats (G a rba n a ti e t al., 1 9 8 3), rh esu s m o nk e y s
(H au se r, 19 9 3 ), a n d h u m a ns (S ac k e im , G ur, & S a uc y , 1 97 7 ). A ctiva tio n o f th e
rig h t h e m isp h ere w ith aro us al m a y m a sk or e ve n re v e rse latera l asy m m e tries .
O ve rall, the resu lts o f th is stu d y d e m o n stra te th at, a s fo r O to lem u r g a rne ttii
(R o g er s et a l., 1 9 94 ), th e ey e dn e ss o f th e m a rm o se ts m ay d e pe n d o n th e stim u lu s
b ein g v ie w ed a nd /o r the sta te o f a rou sa l. T h e d em on stra te d e ffec t o f a ro usa l o n
this form o f la te ralisatio n e m p h asise s th a t c a ution is req u ired w he n d esig nin g
ex p e rim en tal stu d ie s o f la tera lisa tio n , a nd a lso th e im p orta nc e of m ain tain ing a
stab le e n viro n m e n t in the d aily ca re a n d m a in ten a n ce o f ex p e rim e n tal su b jec ts.
T h e e y e p refe ren c es re po rted fo r th e m a rm o se ts a re o pp o site to th ose fou n d
for O tole m u r g a rne ttii (R o g ers e t a l., 1 99 4 ). W h e re a s m a rm o sets d isp lay e d
rig h t-e y ed n e ss for v iew in g n o n aro u sing stim u li a nd a sy m m etric a l d istrib u tio n
o f e ye pre fere n ce s w ith inc rea sin g a rou sa l, th e b ush b a b ie s w e re left-ey e
 126 HOOK-COSTIGAN AND ROGERS

pre fe re n t fo r v iew in g n e utra l stim u li a nd d isp lay e d a sym m e tric a l d istribu tio n o f
ey e p refe re nc es w he n a rou s ed . T h e o pp o site la tera lisa tio n o f th e m a rm o sets,
co m pa red to th e p ro sim ia n s, m a y rela te to ev o lu tio n a ry a d ap tation s fo r the ir
re sp e ctive d iurn a l an d n oc tu rn a l en v iro n m e n ts. A s R o g ers e t al. (1 9 9 4) sug g es t,
ey e pre fere nc e s m ay h av e sh ifted thro u g ho u t th e c o urse o f p rim ate e v o lution .
T h e re is so m e ev id en c e th at th e a sy m m etrica l distrib u tio n o f ga n glio n ce lls, w ith
de n sitie s h igh e r in th e na sa l tha n th e tem po ra l h em ire tin a , rep o rte d fo r
m a rm o se ts a nd oth er d iu rn al prim ate s, m a y n o t b e p re se nt in p ro sim ia n s
(D eB ruy n , W ise, & C a seg ran d e, 1 9 80 ). In G a la g o sen g a len sis, a p ro sim ia n
spe c ie s clo se ly rela ted to O tole m u r g a rn ettii, th e g an g lion c ell d istrib u tio n
ap p ea rs to b e m a rgin a lly b iase d tow a rd a slow e r d e cline in d en sity in th e
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te m p o ra l, an d no t th e na sa l, he m ire tin a (D e B ru y n e t al., 19 8 0 ). T h u s, in th e

no c turn a l p ro sim ia n sp ec ie s, th ere m a y b e a slig h tly b e tte r c o rtica l re pre se nta -
tio n in th e ipsilate ra l h em isp he re , w h e re a s in m a rm o sets th ere w o u ld b e a
stro n g er c ortica l rep rese n tation in th e he m isp h ere co n tra late ral to th e v iew in g
ey e (R o w e , 1 9 9 1; W eisz e t al., 1 99 4 ). T h e h ig h est im a g e q ua lity w o u ld b e
tra nsm itte d to the le ft h em isp h e re by v ie w in g w ith the le ft ey e b y O tole m u r
ga rn et tii a n d w ith th e rig h t ey e b y C a llith rix ja c c hu s. T his retina l o rg an isa tio n
m a y ex p la in ey e d om in a n ce d iffe ren c e s be tw e e n diu rna l an d n o ctu rna l sp ec ie s.
T h is h y p oth e sis c an b e te ste d b y d e te rm in ing ey e pre fere nc e s in m o re n o ctu rna l
an d d iu rn al sp ec ie s.
C a n ta lu po , B isa z z a, a n d V allo rtig a ra (19 9 5 ) su g g este d th at th e te le o st fish
(G ira rd inu s falc atu s) m a y d isp la y co m p le m e n ta ry h e m isp he ric sp ec ialisa tio ns
w h e n ca teg o rising thre a ten in g a nd n o n th re ate n in g stim u li. T h e y fo un d a
po p u lation b ia s fo r tu rnin g rig h t to esc ap e w h e n a sim u late d p re da tor w a s first
pre se nte d. T h e fish v iew e d th e p re da to r w ith th e le ft ey e w h ile tu rnin g rig h t to
esc a pe . H o w e ve r, on su b se qu e n t p re sen tatio ns th e re w a s a sh ift to le ftw a rd
tu rn ing , v iew in g th e pre d ato r w ith th e rig h t e ye . T h e y su g g este d th at the se
asy m m e trie s m a y ref le ct a n in bo rn righ t h em isp he re bia s fo r resp o n din g to
frig hte n in g stim uli a n d a le ft he m isp h ere bia s fo r c a te g orisa tio n o f n e u tra l
stim uli. L e ft h em isph e re sp ec ialisation for ca teg o risatio n ha s also b e en fo u n d in
ch ic ks (A n d rew , 1 99 1 ) a nd in h u m a n s (B ra d sh aw , G ates & N ettle to n , 19 7 7 ;
G r oss, 19 7 2 ; Jon e s, 1 97 9 ). T h e d iffe ren tial e ye p refe re n c es d isp la y ed b y th e
m a rm o se ts a p pe a r to re flec t th e sa m e d ire c tio na l spe cia lisa tio n s a s te le o st fish
an d c h ic k s. C ateg o risation o f stim uli m ay b e pe rform e d b y the le ft h em isp h e re,
pro d uc in g righ t-ey e d o m ina n c e so th a t the m or e d e ta ile d c o rtica l re pre sen ta tio n
is tra n sm itted to th e le ft h em isph e re. E q ua l u se o f b o th e y es w h en vie w in g a n
aro u sing stim u lus co u ld allow bo th he m isp h e re s to rec e ive d ire ct an d rap id inp u t
of in fo rm a tio n (i.e . n o t v ia th e co rp us c allosu m ), a n d th u s th e sep a rate
spe c ia lisatio n s w o uld co n trib u te to a sse ssin g a th rea ten in g situa tio n .
In ad d itio n to th e effe cts o f a rou sa l o n ey e pre fere nc e , w e e x am ine d th e
in flu en c e (on la tera lise d e ye use ) o f ag e, th e m a n u al b ias o f th e ex p erim en ter,
an d th e h a nd pre fere n ce s of th e su b je c ts. A g e d id n o t a ffe ct la tera lise d e ye u se.
 EYE PREFERENCES IN MARMOSETS 127

T h e e y e p re fe ren c es o f th e su b je c ts d id n o t h av e an y re lation sh ip to w h ich h an d

the ex p erim e n ter use d : th e m arm o se ts d isp la y ed the sam e e ye p re fere n ce s w h e n
the ex p erim en ter u sed the le ft o r rig h t h a nd . It is u n lik ely th a t th e rig h t-ey e
p re fere n c es of th e m arm o se ts w ere e x p erim e n ta lly in d u ce d e arly in the te stin g
sessio n s in w h ich th e e xp e rim en te r p rese n te d th e stim u lu s usin g h e r left h a nd , a s
the re w as n o c on v in c in g in d ic a tio n o f a sh ift in ey e p re fe ren c e fo r v ie w ing th e
b an a na w ith a ge (i.e . su cc e ssiv e te stin g ) an d n o stro ng e v id e n ce o f w ithin -
sessio n lea rnin g e ffe c ts o n e ye p refe ren c e. O u t o f e lev e n ca se s an aly se d , in on ly
tw o ca se s w a s th e re a sig nifica n t, or n ea r sig n ific an t, bu t sm a ll (5 % ) in c rea se in
p erc en t le ft-e y e u se fro m th e first to last 5 0 sc o res in a tes t. A n y m in or sh ifts in
ey e p re fere n ce , w h e n th ey d o o c cu r, m ig h t b e a re su lt o f c ha n g ed a ro u sa l ra th e r
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tha n le arn in g .
T h e ey e p ref ere nc e s o f the m a rm o sets w e re also n o t re late d to the ir h an d
p re fere n c es. T h is re su lt c o n cu rs w ith the fin din g s of o the r n o nh u m an p rim ate
sp ec ie s (C o le, 19 5 7 ; K ru pe r e t a l., 1 9 66 ; R o g ers e t al., 19 9 4 ) b u t n o t w ith th e
ev id en c e su gg e stin g th at th ere is a re latio n sh ip b etw ee n e ye pre fere n ce a n d h an d
p re fere n c e in h u m a n s (B ro w n & T a y lo r, 1 9 88 ; H arris & L ov e gr ov e , 1 98 4 ;
M etalis & N iem ic, 1 9 8 4). It m a y b e arg u ed tha t va riab les su ch a s th e ty p e o f ta sk
u sed to m e a sur e h a nd pre fere n ce s o r th e p o sture a d op ted d u rin g th e p erfo rm a nc e
o f a task m a y h av e h a d an influ en c e o n the re lation sh ip s b etw e en ha n d a nd ey e
p re fere n c es. A s th e h an d u se o f n o nh u m a n prim a tes is ta sk -de p e nd e n t (H oo k -
C ostiga n & R og e rs, 1 9 9 5; Fa g o t & V a u cla ir, 1 9 91 ) it is p ossib le th a t h an d
p re fere n c es m ea su red in o th e r ta sk s m ay c o rre late w ith e ye p refe ren c e. H ow e v e r,
it is su g g este d th a t th e se h an d p refe re n c es w o u ld re flec t u nd e rly in g h em isp h e ric
sp ec ia lisation s, as is a rg u e d fo r e ye p refe re nc es, an d thu s w o u ld n o t in d ic a te a
d ire ct c a usa l re la tio n sh ip b e tw e e n m a n ua l sp ec ia lisa tio n a nd ey e do m in a n ce p e r
se , as h a s be e n sho w n in h u m a n s. In a d dition , altho u g h p o stu re h as b e en sh o w n to
effe ct the d ire ction o f ha n d p refe ren c es in so m e n o nh u m an p rim a te sp e cie s
(F ag o t, D re a, & W a lle n , 19 9 1 ; H op k in s, 1 9 9 3; P a rr, H o p k in s, & D e W a al, 1 99 7 ;
S an fo rd , G u in , & W a rd, 19 8 4 ), w e ha v e fo un d th a t th is v a ria ble d o e s n o t in flu e n ce
the d ire ction o f ha n d p re fe ren c es in m a rm o se ts (H o ok -C os tig an & R o g ers, 1 9 9 7).
T h e refo re, it is u nlik ely th at the po stu re a do p ted d urin g fee d in g a ffe cts the la c k o f
co rrela tio n fou n d in this stud y b etw ee n e ye an d ha n d pre fere n ce s.
T h e la ck o f rela tio ns h ip be tw e e n e y e a nd ha n d p re fe re nc es in th e m arm ose t
su gg e sts tha t the se tw o form s o f late ra lisation de p en d o n d iffe ren t m ec h an ism s,
an d th at e ye p re fere n ce s c a nn o t b e sim p ly a p ro du c t o f la te ralise d m o to r c on tro l.
T h is re su lt su p p o rts th e h y p oth esis th a t e ye p re fere nc e s re fle c t h em isp h e ric
sp ec ia lisation s fo r p e rce p tua l a n d no t m o tor fu nc tion .
G iv e n th e w id e spre ad o c cu rre nc e of la te ralisatio n of p erc ep tu a l p ro ce ssin g in
b ird s a n d lo w e r m a m m a ls, w e sug g e st th a t th e sp ec ialisa tio n of th e left
h em isp h e re fo r m a nu a l fu n ctio n in h u m a ns m ay h a ve e vo lv ed from a pre -
ex istin g sp ec ialisatio n fo r p erc ep tu al p roc e ssin g . T h is h y p oth e sis c o n tra dic ts th e
p ro p o se d ``p o stura l orig ins th e ory ’ ’ o f M ac N eila ge , S tu d de rt-K en n ed y , an d
 128 HOOK-COSTIGAN AND ROGERS

L in db lo m (19 8 7 ), w ho sug g e sted th at h em isph e ric sp e cia lisa tio n e v o lve d in

no n h u m a n p rim a tes to o v erc o m e p ro b lem s of p ostu ra l c on tro l w h e n fe ed ing ,
re su ltin g in m an u a l sp e cialisatio n s th a t p rec e d ed the e vo lu tio n o f m a nu a l a n d
la n g ua g e sp ec ia lisa tio n in hu m an s. U n lik e la te ra l h a nd u se, w h ic h a pp e a rs to
va ry in d irec tio n b e tw e e n sp ec ies, h e m isp h eric asy m m e trie s o f pe rce p tua l
fu n c tio n a p p ea r to b e d ire c tio n ally c o nsisten t ac ro ss spe cie s, su g ge sting tha t
th e y m ay h av e e vo lv ed b efo re h a n de d ne ss.
F u rth er rese arc h in to p e rce p tua l fu n ctio n s in n o n hu m a n p rim a te s sp ec ie s is
ne e de d to d e te rm in e h o w a n d w h y h em isp he ric spe c ia lisatio n ev o lve d . E y e
pre fe re n ce te sts m ay be on e w a y o f e x am in in g sp ec ia lisa tio ns o f c o gn itiv e a n d
pe rc ep tua l fu n ctio ns. T h e p rese n ce of a d o m ina n t e y e in h um a ns a nd no n h u m a n
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spe c ie s th ro u gh o u t the ev o lution a ry c o ntin uu m w o u ld sug g e st th at th e re m u st b e

a se lec tiv e a d va n tag e for this fo rm o f fu n ctio na l la tera lisa tio n .

M an u sc ri p t r ec e i v e d 2 8 O c t o b e r 1 9 9 6
R e v i se d m an u sc ri p t re c e iv e d 5 M a rc h 1 9 9 7

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 EYE PREFERENCES IN MARMOSETS 129

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J o u r n a l o f P s y c h o lo g y , 6 2 , 1 7 5 ±1 8 5 .
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R e s e a rc h , 2 1 , 4 4 5 ±4 5 5 .
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 130 HOOK-COSTIGAN AND ROGERS

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R o g e rs , L .J. , W a rd , J .P ., & S t a ff o rd , D . (1 9 9 4 ). E y e d o m in an c e i n t h e s m a ll-e a re d b u s h b a b y ,
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