Neuron, Vol.

14, 217-228, February, 1995, Copyright 1995 by Cell Press

Principles of Human Brain
Organization Derived from
Split-Brain Studies
Review
Michael S. Gazzaniga
Cent er f or Neur osci ence
Uni ver si t y of Cal i f orni a, Davi s
Davi s, Cal i f or ni a 95616
St udi es of damaged human br ai ns have al ways i nt r i gued
neur osci ent i st s. The ri ch sympt omat ol ogy t hat can resul t
is r i vet i ng to al l who exper i ence exami nat i on of such pa-
t i ent s. Yet , one of t he concer ns of basi c sci ent i st s is t hat
st r uct ur e- f unct i on correl at es are di f f i cul t t o make i n t he
damaged brai n, si nce t he l esi ons are nat ur al l y occur r i ng
and usual l y qui t e di f f use. In addi t i on to t he probl em of
quant i t at i ng t he l esi on, t her e has al ways been concer n
whet her t he i nf or mat i on gai ned about t he brai n i n t he pres-
ence of l esi ons is al l t hat usef ul i n under st andi ng normal
br ai n mechani sms. Ar e obser ved ef f ect s due to t he dam-
age of speci f i c areas or to di st ant ef f ect s? These and ot her
i ssues have l i mi t ed t he gener al accept ance of t he f i ndi ngs
from cl i ni cal st udi es.
One of t he i mmedi at e appeal s of t he st udy of pat i ent s
wi t h sur gi cal di vi si on of t he f or ebr ai n is t hat t he separat e
f unct i ons of t he t wo cerebral hemi spher es can be st udi ed
r eadi l y i n t he absence of f ocal damage. Al so, t he cal l osal
sur ger y, whi l e pr oduci ng damage to t he brai n, was di s-
cret e damage t o a f i ber syst em and not to nucl ear areas.
In 1961, R. W. Sperry and hi s col l eagues commenced a
seri es of st udi es on pat i ent s who had under gone sur gi cal
sect i on of t he cerebral commi ssur es i n an effort to cont rol
t hei r ot her wi se i nt r act abl e epi l epsy (Bogen et al ., 1965).
St udi es cont i nue on t he or i gi nal pat i ent s as wel l as ot hers
and sti l l shed l i ght on t he nat ur e of both cor t i cal and sub-
cor t i cal neur al net works. In what f ol l ows, I r evi ew some
aspect s of t hi s wor k to i l l ust r at e how t hi s i ni t i al wor k has
progressed from t he ear l y si xt i es unt i l t he t i me of Sperry' s
deat h i n t he spr i ng of 1994.
Pat i ent s who under go so-cal l ed spl i t - br ai n sur ger y al l
suf f er from i nt r act abl e epi l epsy. Pri or t o t hei r surgery, ex-
t ensi ve at t empt s are made to cont r ol t hei r sei zur es medi -
cal l y. Fai l ure to do so, al ong wi t h ot her cl i ni cal cri teri a,
f i nds t hem candi dat es f or sur gi cal di vi si on of t he corpus
cal l osum, and i n some cases t he ant er i or commi ssur e as
wel l . Al t hough t hese pat i ent s are not nor mal , t he onset of
t hei r epi l epsy var i es and has di f f er ent et i ol ogi es. It has
not been possi bl e to cor r el at e any of t hese var i at i ons i n
t hei r neur ol ogi c hi st or y wi t h t he pat t ern of resul t s obt ai ned
from t he cogni t i ve st udi es. It is known from ot her dat a t hat ,
i n order f or t her e t o be si gni f i cant changes i n t he normal
pat t er ns of cer ebr al l at er al i zat i on, a l arge l esi on must oc-
cur t o one si de of t he brai n i n ear l y chi l dhood (Rasmussen
and Mi l ner , 1977).
In t he or i gi nal st udi es on t he Cal i f or ni a seri es of pat i ent s,
i t was necessar y to rel y on sur gi cal not es f or det er mi nat i on
of t he compl et eness of t he sur gi cal sect i ons. In more re-
cent years, both magnet i c r esonance i magi ng (MRI) and
ot her el ect r i cal brai n mappi ng t echni ques have pr ovi ded a
more accur at e r epr esent at i on of t he ext ent of t he sur gi cal
sect i ons. Accu rate document at i on of t he ext ent of cal l osal
sect i on becomes cr uci al f or under st andi ng t he or gani za-
t i on of t he cer ebr al commi ssur e.
The mai n met hods of t est i ng t he per cept ual and cogni -
t i ve f unct i ons of each hemi spher e have not changed i n
pr i nci pl e over t he years. However, t here have been si gni f i -
cant advances i n t he t echnol ogi es used to pr esent st i mul i
to t he sur gi cal l y separat ed hemi spher es. In t he ear l y
years, i nf or mat i on was vi sual l y l at er al i zed t o one or t he
ot her hemi spher es by qui ckl y f l ashi ng st i mul i to one or
t he ot her vi sual f i el d usi ng el ect r oni c shut t er s at t ached t o
sl i de proj ect ors (see Fi gure 1). As t he pat i ent f i xat es on
a poi nt i n space, i nf or mat i on f l ashed to t he l eft of f i xat i on
is present ed excl usi vel y to t he ri ght hemi spher e. The qui ck
f l ashi ng is necessar y to cont r ol f or unwant ed eye move-
ments, whi ch woul d r edi r ect t he i nf or mat i on i nto t he un-
want ed hemi spher e. Thi s met hod has been repl aced by
usi ng comput er - based st i mul i pr esent at i on ar r angement s.
More i mport ant l y, t he devel opment of an i mage st abi l i zi ng
syst em, used i n concer t wi t h a Purki nj e eye t racker, now
per mi t s sust ai ned pr esent at i on of i nf or mat i on t o ei t her vi -
sual fi el d, and t heref ore, ei t her hemi spher e. Accor di ngl y,
i f a subj ect moves hi s or her eyes away from f i xat i on, t he
st abi l i zi ng syst em moves t he st i mul us wi t h t he eyes and
t her eby pr event s t he i nf or mat i on from bei ng present ed to
t he wr ong hemi spher e. Thi s t echnol ogi cal devel opment
has al l owed f or new f i ndi ngs on bot h t he neur ol ogi cal and
psychol ogi cal aspect s of hemi spher e di sconnect i on.
Fundamental Principles Arising from Initial Studies
The or i gi nal report s on t he Cal i f or ni a seri es of pat i ent s
deal t wi t h a number of f undament al i ssues concer ni ng t he
basi c psychol ogi cal pr oper t i es of t he separ at ed cerebral
hemi spheres, as wel l as basi c i ssues of neur ol ogi cal orga-
ni zat i on (Sperry et al., 1969; Gazzani ga, 1970). In many
respects, t he i ssues rai sed in t he or i gi nal st udi es st i l l dr i ve
cur r ent research efforts. In t he f ol l owi ng, t he neur ol ogi c
consequences wi l l be r evi ewed fi rst, f ol l owed by t he st ud-
i es on t he separat e psychol ogi cal pr oper t i es of t he t wo
cerebral hemi spher es.
Sensory and Motor Studies
The human st udi es wer e car r i ed out i n t he cont ext of
st rong new ani mal evi dence, obt ai ned by Myer s and
Sper r y (1958), t hat di vi di ng t he cerebral commi ssur es pro-
duced a pr of ound def i ci t i n t he i nt er hemi spher i c t r ansf er
of sensor y and mot or i nf ormat i on. In t he cat, monkey, and
chi mpanzee, Myer s (1956) had det er mi ned t hat , f ol l owi ng
mi dl i ne sect i on of t he corpus cal l osum and ant er i or com-
mi ssure, vi sual and t act i l e i nf or mat i on l at er al i zed to one
hemi spher e di d not t r ansf er t o t he opposi t e hemi spher e.
Thi s resul t ed i n t he so-cal l ed "spl i t -brai n" ani mal . Thi s star-
t l i ng di scover y was compl et el y cont r ar y t o earl i er report s
on t he ef f ect s of human commi ssur e sect i on report ed by
Akel ai t i s (1941, 1944). Akel ai t i s report ed no si gni f i cant
neur ol ogi cal or psychol ogi cal ef f ect s f ol l owi ng sect i on of
t he cal l osum.
Neuron
218
. . . . . .
DEFLECTOR
Horizontal
~ e~ . ya
Figure 1. Stimulus Lateralization Technique
To examine differentially hemispheric processing differences, it is nec-
essary to lateralize stimuli within the left and right visual fields (upper
left). However, if the subject makes an eye movement during the stimu-
lus presentation, proper lateralization is no longer maintained (upper
middle). Thus, retinal stabilization is useful to counteract the effects
of such eye movements (upper right). A dual Purkinje image eyetracker
coupled with a mirror stimulus deflector allows such retinal stabilization
(lower left). As eye movements occur, horizontal and vertical deflection
mirrors move to counteract such movements, maintaining proper later-
alization (lower right).
The sur ger i es on t he or i gi nal Cal i f or ni a cases (W J, NG,
and LB) wer e abl e to show t hat humans responded to
f or ebr ai n commi ssur ot omy i n essent i al l y t he same way as
t he monkey and chi mpanzee. Vi sual i nf or mat i on pre-
sent ed to one hal f -brai n was not avai l abl e t o t he ot her
hal f -brai n f or anal ysi s. A si mi l ar pr i nci pl e appl i es to t ouch.
Obj ect s pl aced i n t he ri ght hand wer e nor mal l y named
and descri bed, but obj ect s pr esent ed in t he l eft hand wer e
not. Si mi l arl y, sensor y i nf or mat i on present ed to one hemi -
spher e was usef ul i n gui di ng t he cont ral at eral hand but
ver y i nef f ect i ve i n cont r ol l i ng t he i psi l at eral hand.
From a cogni t i ve poi nt of vi ew, t he fi rst st udi es conf i rmed
l ong- st andi ng neur ol ogi c knowl edge about t he nat ure of
t he t wo cerebral hemi spher es. The l eft brai n was domi nant
f or l anguage, speech, and maj or probl em sol vi ng, whi l e
t he ri ght appear ed speci al i zed f or vi suospat i al t asks such
as dr awi ng cubes and ot her t hr ee- di mensi onal patterns.
Of course, t hi s meant t hat vi sual and t act i l e st i mul i pre-
sent ed to t he r i ght hemi spher e coul d not be named or
descri bed, si nce t he sensor y i nf ormat i on was di scon-
nect ed from t he domi nant l eft speech hemi spher e. Thi s
dr amat i c resul t st ood in cont r ast to t he r i ght hemi sphere' s
abi l i t y to acknowl edge t he pr esence of t hese st i mul i by
al l owi ng i t to respond in a nonver bal manner, such as
poi nt i ng to mat chi ng obj ects and t he like.
In t he subsequent 30 years, t hese st udi es have been
f ol l owed up by a nu mber of i nvest i gat ors. Whi l e t he or i gi nal
st udi es set t he f r amewor k f or subsequent research, t he
huge effort to char act er i ze f ul l y t hese uni que human be-
i ngs has cont i nued to yi el d maj or i nsi ght s i nt o t he or gani -
zat i on of t he human brai n.
Perceptual and Attentional Studies Following
Cerebral Disconnection
The at t ent i onal and per cept ual abi l i t i es of spl i t -brai n pa-
t i ent s have been ext ensi vel y expl ored. Vi sual percept i on
is t he easi est to st udy. Over al l , f ol l owi ng cor t i cal di scon-
nect i on, percept ual i nf or mat i on does not i nt eract bet ween
t he t wo cerebral hemi spheres, wher eas t he suppor t i ng
cogni t i ve processes of at t ent i onal mechani sms do some-
t i mes i nt eract .
Simple P erceptual Interactions Are Not Seen
Spl i t -brai n pat i ent s are not abl e to cross-i nt egrat e vi sual
i nf ormat i on bet ween t hei r t wo vi sual hal f -f i el ds. When vi -
sual i nf ormat i on is l at eral i zed to ei t her t he l eft or ri ght di s-
connect ed hemi spher e, t he unst i mul at ed hemi spher e
cannot use t he i nf ormat i on f or per cept ual anal ysi s (see
Fi gure 2). Thi s is al so t r ue f or st er eognost i c i nf or mat i on
present ed to each hand. Whi l e t he pr esence or absence
of t ouch st i mul at i on is not ed t o any part of t he body by
ei t her hemi spher e, pat t erned somat osensor y i nf or mat i on
is l at eral i zed. Thus, an obj ect hel d in t he l eft hand cannot
hel p t he ri ght hand f i nd an i dent i cal obj ect. Al t hough t her e
have been report s ar gui ng t hat some hi gher or der percep-
t ual i nf ormat i on is i nt egrat ed at some l evel vi a subcor t i cal
st ruct ures (Croni n-Gol omb, 1986; Sergent , 1990), t hese
resul t s have not been repl i cat ed by ot hers (Seymour et
al., 1994; Cor bal l i s et al., 1993; Corbal l i s, 1994; McKeever
et al., 1981).
Humans Show Visual Midline
Overlap Phenomenon
Ther e is some nasot empor al over l ap at t he ret i nal ver t i cal
meri di an in cat and monkey (Stone, t 966; St one et al.,
1973). In a st ri pe 1- 2 degr ees of vi sual angl e wi de t hat
st raddl es t he t wo vi sual hal f -f i el ds, vi sual i nf or mat i on i s
sent to both t he l eft and r i ght vi sual corti ces. Whet her or
not t he anat omi cal proj ect i ons have any f unct i onal si gni f i -
cance has never been exami ned i n t he ani mal . Fendr i ch et
al. (1989) have exami ned t hi s i ssue i n spl i t - br ai n pat i ent s.
Usi ng an i mage st abi l i zer combi ned wi t h a Purki nj e eye
tracker, caref ul assessment of t he vi sual mi dl i ne of t wo
spl i t -brai n pat i ent s has r eveal ed an area of no more t han
2 degrees in wi dt h at t he ver t i cal mi dl i ne wher e vi sual
i nf ormat i on appear s avai l abl e t o each hal f -brai n. Thi s con-
trasts wi t h t he f i ndi ngs of Sugi shi t a et al. (1994), who f ound
no overl ap but who wer e di sadvant aged by not havi ng t he
i mage st abi l i zer to car r y out t hei r mi dl i ne st udi es. Sti l l ,
wi t hi n t hi s st r i p of overl ap, t he si gnal s conveyed to each
hemi spher e from t he cont r al at er al hemi r et i na appear to
be weak or degraded. St i mul i coul d not be compar ed
across t he ver t i cal meri di an i f t hese compar i son requi red
det ai l ed shape i nformati on, or i f t he sti mul i wer e presented
f or onl y 200 ms.
Review: Human Split-Brain Studies
219
I I L V F R V F
-~
x \ .
N
/ / B
: \
Figure 2. Interhemispheric Transfer Tests
The upper example depicts spatial tests. On within-field trials (A), the
eyes moved to the stimulus that was surrounded by the probe. On
between-field trials (B), the eyes also moved to the corresponding
stimulus in the other hemifield. In contrast, when perceptual informa-
tion was used as the cue (lower examples), the capacity to find the
patterned cue was only possible on the within-field trials (A). On the
between-field trials (B), the response to the opposite field was only at
chance.
I nter hemi s pher i c T rans f er I s S een f o r C rude
S pati al L o c ati o n I nf o r mati o n
Unlike visual and somatosensory cues, crude i nformati on
concerni ng spatial l ocati ons can be cross-integrated
(Trevarthen, 1968; Trevarthen and Sperry, 1973; Holtz-
man, 1984). In one set of experi ments, a four-point grid
was presented to each vi sual field. On a given trial, one
of the positions on the grid was highlighted, and one con-
dition of the task required the subject to move his eyes
to the hi ghl i ghted poi nt within the visual fi el d stimulated.
In the second condition, the subject was required to
move the eyes to the relative poi nt in the opposi te visual
field. Split-brain subjects were easily abl e to do this,
thereby suggesting some crude cross-integration of spa-
tial information. This was true even if the grid was randoml y
posi ti oned in the tested field.
S pati al A ttenti o n C an B e Di r ec ted but No t Divided
between the Hemi s pher es
The finding that some type of spatial i nformati on remains
integrated between the two half-brains raises the question
of whether the attenti onal processes associated with spa-
tial i nformati on are affected by cortical di sconnecti on sur-
gery. Using a modi fi cati on of a paradi gm devel oped by
Posner et al. (1980) that capi tal i zes on spatial priming phe-
nomena, Hol tzman et al. (1981) found that ei ther hemi-
sphere can direct attention to a poi nt in ei ther the left or
right visual field. Posner first showed that the response
latency to a peri pheral visual target is reduced when ob-
servers have pri or i nformati on regardi ng its spatial locus,
even when eye movements are prevented. Presumably,
the spatial cue allows observers to direct thei r attention
to the appropri ate location pri or to the onset of the target.
When this paradi gm was used in split-brain patients to
measure the extent to whi ch such attenti onal cues affect
performance, the separated hemi spheres were found not
to be strictly i ndependent in thei r control of spati al orienta-
tion. Rather, the two hemi spheres rely on a common ori-
enting system to mai ntai n a si ngl e focus of attention. Thus,
as with normal peopl e, a cue to direct attenti on to a particu-
lar poi nt in the vi sual field is used no matter which hemi-
sphere is presented with the spatial cue.
The discovery that spatial attenti on coul d be di rected
with ease to ei ther visual fi el d raised the question of
whether each separate cogni ti ve system in the split-brain
patient could, if instructed to do so, i ndependentl y direct
attention to a particular part of its own vi sual fi el d (Holtz-
man et al., 1984). Can the right hemi sphere direct attenti on
to a point in the left visual fi el d whi l e the left brain simulta-
neously directs attention to a poi nt in the right visual field?
Normal subjects cannot so di vi de thei r attention. Can split-
brain patients with evi dent cogni ti on in both hemi spheres
do so?
Results from several studies show that the split-brain
pati ent is unable to di vi de spatial attenti on between the two
half-brains. There woul d appear to be onl y one integrated
spatial attention system that remains intact fol l owi ng corti-
cal disconnection. Thus, like neurol ogi cal l y intact observ-
ers, the attenti onal system of split-brain pati ents is unifo-
cal. They are unabl e to prepare for events in two spati al l y
disparate locations.
A ttenti o nal R es o urc es A re S har ed
The dramati c effects of di sconnecti ng the cerebral hemi-
spheres on percepti on and cogni ti on might suggest each
half-brain possessed its own attenti onal resources. If true,
one woul d predict the cogni ti ve operati ons of one half-
brain, no matter what the difficulty, woul d little influence
the cogni ti ve activities of the other. The competi ng vi ew
is that the brain has a set of limited resources that manage
such processes, and if they are bei ng appl i ed to task A,
there are fewer avai l abl e for task B. This model woul d
predict that the harder hemi sphere A worked on a task,
the worse hemi sphere B woul d do on a task of constant
complexity.
Many studies have been carried out on this issue, and
all confirm the notion that t he central resources are limited
(Holtzman and Gazzaniga, 1982). In the ori gi nal experi -
ment, there were two conditions. In what was cal l ed the
"mi xed" or hard condition, two series of three different geo-
metric shapes were di spl ayed concurrentl y to the left and
right of central fi xati on and thus were l ateral i zed to the
right and left hem ispheres, respectively. In t he"redundant "
or easy condition, the series of three di fferent geometri c
shapes was presented to onl y one hemi sphere, whi l e the
other vi ewed three identical geometri c shapes. A unilateral
probe figure subsequentl y appeared, and the observer in-
dicated with a forced-choice key press whet her it matched
any of the probed field's items. The results cl earl y showed
that, when one half-brain was working on processing onl y
Neuron
220
one repeat ed st i mul us, t he opposi t e hemi spher e per-
f ormed bet t er at recal l i ng whet her t he probed st i mul us
was part of t he or i gi nal set of t hr ee st i mul i . When both
hemi spher es wer e t r yi ng to pr ocess t hree st i mul i , t he per-
f or mance of each was i mpai red. These overal l f i ndi ngs
have r ecent l y been repl i cat ed i n a monkey model of t hese
t asks ( Lewi ne et al ., 1994).
The f or egoi ng concept of resources as used here is to
be di st i ngui shed from ot her propert i es of sensor y syst ems
associ at ed wi t h at t ent i onal i nf or mat i on processi ng. The
l i mi t on t he pr ocessi ng of i nf or mat i on t hat is capt ured by
t he concept of r esour ce l i mi t at i ons i n cui ng t asks or pro-
cessi ng l oad t asks is more gener al and di st i nct from t he
l i mi t s and mechani sms now st udi ed i n such phenomena
as sear chi ng a vi sual scene f or i nf or mat i on. The concept of
resources, as concept ual i zed from t he r esul t s i n spl i t -brai n
pat i ent s, l ar gel y refers to processes t hat are engaged
when t here is a vol unt ar y al l ocat i on of at t ent i on to an i nfor-
mat i on pr ocessi ng task. Sear chi ng a vi sual scene, how-
ever, cal l s upon more aut omat i c processes t hat may wel l
be bui l t - i n propert i es of t he vi sual syst em i t sel f ( Ol shausen
et al ., 1993). That t hese syst ems ar e di st i nct is ref l ect ed i n
t he resul t s of t he f ol l owi ng st udi es, i n whi ch i t is di scovered
t hat spl i t t i ng t he brai n has a di f f er ent ef f ect on such pro-
cesses.
Visual Search Is Independent and Different
in the Half.Brains
The more i t ems present ed to be anal yzed i n a vi sual array,
t he l onger it t akes to search t he array f or t he i nf or mat i on
bei ng sought i n a par t i cul ar t ask (St ernberg, 1975). Thus,
af t er a basel i ne react i on t i me is est abl i shed, it t akes nor-
mal cont r ol s an addi t i onal 70 ms t o det ect a t arget i f t here
ar e t wo more items, anot her 70 ms f or an addi t i onal t wo
i tems, and so on. Thi s l i near i ncr ease i n response t i me
wi t h number s of el ement s i n a di spl ay is char act er i st i c
onl y of "conjunction search, " i.e., wher e t ar get s cannot be
di st i ngui shed f rom nont arget s by a si mpl e f eat ur al di ffer-
ence. In spl i t - br ai n pat i ent s, it has been shown t hat , when
t he i t ems are di st r i but ed across t he mi dl i ne of t he vi sual
f i el d as opposed to al l bei ng i n one vi sual fi el d, t he react i on
t i me to added st i mul i is cut al most i n hal f (Luck et al., 1989,
1994). These resul t s i ndi cat e t hat t he t wo di sconnect ed
hemi spher es can search t he arrays i n paral l el , somet hi ng
t hat normal subj ect s cannot do.
Thi s f i ndi ng has r ecent l y been ext ended by Ki ngst one
et al. (1995) to show t hat , al t hough t he over al l resources
a br ai n commi t s to a t ask appear const ant , t he st r at egy
by whi ch t hey are used can var y as a f unct i on of t he hemi -
spher e act i vat ed f or t he task. It now appear s t he f i xed
propert i es seen i n vi sual search exper i ment s can be i nf l u-
enced by what are cal l ed "t op-down" propert i es. In t hi s
st udy i t was di scover ed t hat t he st r at egy by whi ch each
hemi spher e car r i ed out t he exami nat i on of t he cont ent s
of i t s vi sual f i el d di f f ered. The left, domi nant hemi spher e
ut i l i zed a "gui ded" or "smart" st rat egy, wher eas t he r i ght
hemi spher e di d not. Thi s means t he l eft hemi spher e
adopt ed a hel pf ul cogni t i ve st r at egy in sol vi ng t he prob-
l em, whi l e t he r i ght hemi spher e does not possess t hose
ki nds of ext r a cogni t i ve ski l l s t o assi st i t s vi sual search
mechani sm.
Sensorimotor Control Mechanisms Differ
for Eye and Hand
Wi t h t he remarkabl e separat i on of sensor y i nf ormat i on
and wi t h t he wel l known l at er al i zat i on of cor t i cospi nal mo-
t or syst ems, t he spl i t -brai n ani mal and human rai se i nter-
est i ng quest i ons concer ni ng t he neural mechani sm by
whi ch a mot or act i vi t y can t ake pl ace. In recent years,
t hese pat i ent s have al so of f ered t he oppor t uni t y to test
t heor i es about t he nat ur e of t he neur al pat hways act i ve
i n vi suo- ocul ar cont rol .
A Disconnected Hemisphere Can Control Both
Arms but Only the Opposite Hand
One of t he endur i ng f i ndi ngs of spl i t - br ai n research has
been t he di st i nct i on bet ween t he capaci t y a di sconnect ed
hemi spher e has f or cont r ol l i ng pr oxi mal ver sus di st al mus-
cl es. Sect i oni ng of t he cal l osum cl ear l y i mpai r s t he capac-
i t y of t he l eft hemi spher e t o cont r ol t he l eft hand and of
t he ri ght hemi spher e to cont r ol t he ri ght hand. These i psi -
l ateral sensor i mot or combi nat i ons need t he i nt act cal l o-
sum to i nt egrat e i nf ormat i on from t he cor t i cal sensor y
areas to t he mot or cor t ex t hat cont r ol s di st al hand move-
ment. At t he same ti me, ei t her hemi spher e can gui de and
cont rol more pr oxi mal movement s of each arm, and of
course, t he l egs. Thi s abi l i t y of each separ at ed hemi -
spher e to cont r ol each arm al l ows f or t he appear ance of
cent r al l y i nt egrat ed i nf ormat i on when, i n fact, t he i nt egra-
t i on seen is somat i c and peri pheral .
In a recent st udy by Ki ngst one et al. (1995), i t was shown
t hat a t ask whi ch appeared to i nvol ve t he cent r al i nt egra-
t i on of i nf or mat i on i n fact ref l ect ed onl y t he peri pheral i nte-
grat i on of i nf or mat i on. In t hese exper i ment s, one wor d was
f l ashed to one hemi spher e and anot her to t he opposi t e
hemi spher e. The t ask requi red t he pat i ent t o dr aw what t he
message sai d; i .e., f or wor ds such as "bow" and "arrow,"
wher e "bow" was f l ashed to t he ri ght hemi spher e and
"arrow" was f l ashed to t he l eft hemi spher e, t he subj ect
woul d dr aw a bow wi t h an ar r ow i n posi t i on. At f i rst gl ance
t hi s appears to suggest t he i nf ormat i on pr esent ed t o each
hal f -brai n was somehow i nt egrat ed t hr ough subcor t i cal
syst ems t hat i n t ur ned managed t he mot or response. How-
ever, t he real st r at egy was unear t hed when wor d pai rs
wer e pr esent ed that, i f t r ul y i nt egrat ed, woul d resul t i n one
ki nd of dr awi ng ver sus anot her. In t hese t ri al s, t he wor d
pai r "hot-dog" was present ed, wher e "hot" was f l ashed to
t he ri ght hemi spher e and "dog" was f l ashed t o t he l eft
hemi sphere. In t hi s case t he subj ect dr ew a pi ct ur e of a
dog and t hen added l i nes i ndi cat i ng i t was pant i ng from
heat. If, of course, t he ent i re wor d had been f l ashed t o one
or t he ot her hemi sphere, t he subj ect woul d have dr awn a
hot dog, whi ch is to say a f r ankf ur t er i n a bun. Thus, what
mi ght appear t o be cent r al l y i nt egrat ed sensor i mot or re-
sponse was i n f act a case i n whi ch each hemi spher e con-
t rol l ed and rel eased t he arm used i n dr awi ng i n an ef-
f ort l ess and ef f i ci ent manner.
Either Hemisphere Can Control Saccadic
Eye Movements
In cont r ast to t he i nabi l i t y of a di sconnect ed hemi spher e
to cont r ol t he i psi l at eral hand wi t h accur acy, each hemi -
spher e is abl e t o di rect t he eyes ei t her cont r aver si vel y or
i psi ver si vel y ( Hughes et al., 1992). Thi s capaci t y woul d
Review: Human Split-Brain Studies
221
Figure 3. Postoperative Mid-Saggital MR Images
Magnetic resonance images (MRIs) of four patients with varying degrees of callosal section.
(A) The entire corpus callosum has been sectioned, resulting in no transfer of information between the cerebral hemispheres.
(B) The posterior or splenia! regions have some sparing in a particular area, with the result that visual information could be transferred between
the cerebral hemispheres.
(C) The posterior callosum was sectioned, while the anterior half remained intact. This patient had difficulty with sensorimotor integration of one
hand but not the other.
(D) Two remnants were inadvertently left intact, one in the splenial region and one in anterior genu region (see Gazzaniga, 1989).
not be predicted by dozens of studies showi ng that in each
hemi sphere the frontal eye fields control contraversive eye
movements (Bruce and Gol dberg, 1984; Wurtz and AI-
bane, 1980). This particular result reveals how psycho-
physical studies on human patients with discrete lesions
suggest that alternate neural pathways are involved, path-
ways different from those that mi ght otherwise be evident
from the ani mal studies.
The foregoi ng group of studies reveals how discon-
necting the cerebral hemi spheres at the cortical level inter-
rupts some kinds of attentional, perceptual, and sensori-
motor processes, but not others. Although there is little
evi dence that any perceptual information can be trans-
ferred between the hemispheres, there are some kinds of
attentional and sensori motor processes that remain un-
changed with cortical disconnection. Just as formal genet-
ics helps formul ate the concept of a gene that later be-
comes understood at the mol ecul ar level, identifying these
differences in brain organi zati on will help neuroscientists
understand the systems basis of these mental activities.
Partial Callosal Sections Reveal Specificity
of Commissure Function
In animal studies, it was quickly determi ned that sectioning
the entire corpus callosum and anterior commi ssure pre-
vented the i nterhemi spheri c transfer of a wi de range of
modal and motor information. It was also shown that partial
sectioning of the commi ssures could block specific func-
tions from transferring across the callosum (Black and My-
ers, 1966; Sullivan and Hamilton, 1973; Hamilton and Ver-
meire, 1986). In humans, it was only after other cases
appeared who had not undergone full callosal section that
comparabl e studies could be done, and it became appar-
ent that specific regions of the callosum were responsible
for the transfer of specific modalities. This work was en-
hanced when MRI allowed for the accurate description of
cut and uncut fiber systems.
MRI.Verified Lesions of Partial Sections Reveal
Specific Modal Functions
When the corpus callosum is fully sectioned, there is little
or no perceptual or cognitive interaction between the hemi-
Neuron
222
SPLIT BRAIN
Right Hemisphere
S T I M U L U S
< >
COMPARISONS
Left Hemisphere
VERBALRESPONSE
LW t
K n i g h t
RVF
N o r m a l B r a i n
" K n i g h t "
LVF t RVF
K n i g h t
P a r t i a l S p l i t "1 h a v e o p i c t u r e i n
m i n d b u l c a n ' t soy
it.,.. T w o f i g h t e r s i n
a r i n g . . , , A n c i e n t ....
w e a r i n g u n i f o r m s
ond h e l m e t s . . . , o n
h o r s e s , t r y i n g t o
k n o c k e a c h o t h e r
o f f . . . K n i g h t s ? "
I LVF t RVF
K n i g h t
C o m p l e t e S p l i t
"\;
\ l l
"1 d i d n ' t s e e a n y t h i n g "
Figure 4. Visual Transfer Fallowing Staged Callosal Section
Case JW underwent a staged callosal section in which the posterior
half of the callosum was first sectioned. It has been shown that the
posterior disconnecting surgery prevents the transfer of information
related to the sensory dimensions of the stimulus, but not the semantic.
The result was consistent with the overall knowledge that more anterior
regions of the callosum are committed to the transfer of higher order
information (from Sidtis, 1981).
spheres. Surgi cal cases in whi ch cal l osal section was ei-
ther limited or where there was i nadvertent sparing of a
part of the cal l osum al l ow for the exami nat i on of speci fi c
functi ons of the caUosum by region. For exampl e, when
the splenial region (posterior area of t he cal l osum that
interconnects the occipital lobe) is spared, there is normal
transfer of vi sual i nformati on between the two cerebral
hemi spheres (Figure 3). In these cases, pattern, color, and
linguistic i nformati on presented anywhere in ei ther vi sual
fi el d can be matched with i nformati on presented to the
other half-brain. However, such patients show no transfer
of stereognosti c information and also show a left ear sup-
pression to di choti cal l y presented audi tory stimuli. These
kind of observati ons are consistent with other human and
ani mal data that show the maj or subdi vi si ons of the calla-
sum are organi zed into functi onal zones, wi th the posteri or
regions more concerned with vi sual i nformati on and the
more anteri or regi ons transferring audi tory and tacti l e in-
formati on (Hamilton, 1982; Gaz.zaniga, 1989).
Anterior Callosum Is Involved in Higher Order
Transfer of Semantic Information
Patients who have undergone staged cal l osal section
have also provi ded gl i mpses into what anteri or cal l osal
regions are i nvol ved in transferring between the cerebral
hemispheres. When the posteri or half of the cal l osum is
sectioned, transfer of visual, tactile, and audi tory sensory
information is severel y disrupted. However, the remai ni ng
intact anteri or cal l osum is abl e to transfer hi gher order
i nformati on (Figure 4). Hi gher order i nformati on is a term
used when it is cl ear the i nformati on transferred is not
si mpl y sensory in nature. This distinction was shown in
one study (Sidtis et al., 1981) in whi ch the corpus cal l osum
was secti oned in two stages. Fol l owi ng the first stage of
sectioning, t he pati ent was unabl e to name stimuli pre-
sented to the right hemi sphere. However, over a 10 week
period, he gradual l y began to name some of the stimuli.
Cl ose exami nat i on of this capaci ty reveal ed that the right
hemi sphere was transmitting to the left hemi sphere hi gher
order or gnostic cues about the stimulus, but not the actual
stimulus itself (Figure 4). In short, the anteri or cal l osum
transfers gnosti c representati ons of the stimulus, but not
the stimulus itself. Following section of the anteri or calla-
sum, this behavi or ceased.
Memory Studi es Following Cerebral Disconnection
The most powerful impression one has observi ng patients
who have had thei r hemi spheres di vi ded is how unaffected
they appear to be in terms of thei r general cogni ti ve aware-
ness, affect, and sense of self (Gazzaniga, 1970). At a
superficial l evel of observation, separati ng half of the neo-
cortex from t he other half appears to have little i mpact on
overal l cognition. Verbal IQ remains intact, as do within-
hemi sphere reacti on ti mes to perceptual stimuli and gen-
eral problem sol vi ng capacities. On the ot her hand, stan-
dardi zed memory tests admi ni stered postoperati vel y have
suggested that there is an i mpai rment in short-term mem-
ory capaci ty (Zaidel and Sperry, 1974). Recent studies
have extended these observations.
Free Recall but Not Recognition Memory Is
Impaired in Each Cerebral Hemisphere
We have recentl y carried out a vari ety of studies on overal l
i nformati on processing capaci ti es and, in some instances,
have been abl e to compare postoperati ve performance
with preoperati ve capacity. In these new tests, an interest-
ing picture emerges that suggests commi ssurotomy im-
pacts on free recall mechani sms but that recogni ti on mem-
ory remains l argel y unchanged. Free recall requires a
subject, with no cuing, to recall prior i nformati on such as
a previously studi ed word list. Recogni ti on tasks merel y
require a subject to j udge whet her a sti mul us such as a
printed word has been seen before on a previ ousl y studied
list. The overall pattern of results is consi stent with the
i dea that di sconnecti ng the hemi spheres has a powerful
effect on free recall capacity, whi l e recogni ti on capaci ty
remains normal (Phelps et al., 1991). Moreover, it was
shown that onl y posterior cal l osal -sect i oned patients re-
veal ed the free recall deficit. Anteri or cal l osum-sect i oned
pati ents behaved normally. Si nce secti oni ng the posteri or
cal l osum i nevi tabl y i nvol ves sectioning t he hi ppocampal
commissure, i t is suggested this structure may pl ay a cru-
cial role in this speci fi c memory deficit. Thi s woul d be con-
sistent with studies suggesti ng even uni l ateral hi ppocam-
pal damage can produce probems in free recall (Goldstein
and Polkey, 1992; Ott and Saver, 1993). Disconnection
Review: Human Split-Brain Studies
223
RI~DUNIItJ(I" CONDfIION MI X [ D CONOI TI ON
T 5 T robe 'T 5 T F~obe
san~ d;~. ~ d~4
i I ~i ~
/
I 1~
LVF RVF L,A: RV~
Re~J,~an~ Mixed
Figure 5. Memory Performance Following Callosal Section
On each "redundant" trial (top left), a bilaterally displayed target (X)
moved among four homologous cells in the two matrices. A tone
sounded and a unilateral probe sequence was presented. The ob-
server indicated whether the probe sequence matched the probed
field's target sequence. In this example, the bilateral target sequences
began at time T~ in the upper left corners of the two matrices and
concluded at -14 in the upper right corners, and the unilateral probe
sequence was presented from Ts-Ts. All Xs appeared for 150 ms. The
diagram on the right is an example of a mixed condition trial. Mixed
conditions trials differed from redundant condition trials only in that
different target sequences were presented in the two visual fields.
Data are presented separately for commissurotomy patient JW and
the average performance of the control observers (after Holtzman and
Gazzaniga, 1985).
of t he hemi spheres in thi s commi ssural zone mi ght wel l
mi mi c t he uni l ateral l esi on effects. Overal l , t he dat a sug-
gest t hat t he i nt act neocort i cal syst ems t hat are i ntercon-
nected t hrough t he post eri or cal l osal syst em cont ri but e t o
free recal l mechani sms. It is as if t he neural mechani sms
requi red for ri chl y encodi ng a sti mul us, whi ch cont ri but e t o
f ree recal l , are less avai l abl e t o each hemi spher e f ol l owi ng
post eri or cort i cal di sconnect i on.
Increase in Short-Term Memory Capacity Is More
Apparent Than Real
Al t hough t he dat a suggest t here are di ffi cul ti es in free re-
cal l , t he cort i cal l y di sconnect ed pat i ent can perf orm ot her
memor y t est s in a manner super i or t o t hat of a normal .
Consi der t he f ol l owi ng.
As al ready revi ewed, t he cal l osum-sect i oned pat i ent
has no measur abl e i nt eract i ons bet ween t he t wo hemi -
spheres in t he processi ng of percept ual i nf ormat i on. Iden-
t i cal and si mpl e vi sual pat t erns of all ki nds can be pre-
sent ed t o each separat e hal f-brai n, and t he pat i ent is
unabl e t o say whet her t he st i mul i are t he same or di fferent.
Thi s f act rai ses t he possi bi l i ty t hat , in a memor y t ask involv-
i ng vi sual retenti on, a spl i t-brai n subj ect mi ght perf orm at
a hi gher level t han a normal i ntact cont rol subj ect if t he
percept ual i nf ormat i on wer e di st ri but ed bet ween t he t wo
visual half-fields. In one st udy on this issue, a compl ex
spati al memor y t ask was admi ni st ered t o both a spl i t-brai n
pat i ent and normal cont rol s in whi ch cri ti cal i nformati on
was present ed in each vi sual hal f.fi el d (Hol t zman and Gaz-
zani ga, 1985). For t he control s, t he vi sual i nformati on was
aut omat i cal l y combi ned and percei ved as one l arge prob-
l em (Fi gure 5). For t he spl i t-brai n pati ent, each hemi spher e
percei ved a pr obl em t hat remai ned separat e f rom t he per-
cept ual i nf ormat i on present ed to t he ot her half-brain; thus,
each hemi spher e percei ved a much si mpl er task. The re-
sults were cl ear. The spl i t-brai n pat i ent out -perf ormed t he
normal cont rol s on cri ti cal t est condi t i ons. The cal l osum-
sect i oned pat i ent benef i t ed f rom t he fact t hat t he percep-
tual array under one of t he t est condi t i ons di d not appear
more di ffi cul t because t he wor k was di st ri but ed t o each
separat e hemi sphere, even t hough t he actual sensory
array was i denti cal t o that exper i enced by t he i ntact normal
control s.
Di sconnect i on of t he cerebral hemi spheres cl earl y
al l ows for a uni que cogni t i ve state. In a sense it turns a
seri al , uni fi ed percept ual syst em i nto t wo si mpl er percep-
tual syst ems t hat do not i nteract, and t heref ore do not
i nterfere wi t h each other. It al l ows for t he breaki ng down of
a l arge percept ual pr obl em into smal l er, more manageabl e
pr obl ems t hat a hal f-brai n is capabl e of sol vi ng. From t he
observer' s poi nt of vi ew, however, it l ooks l i ke t he t ot al
i nformati on processi ng capaci t y of t he pat i ent has been
i ncreased and is superi or t o t hat of normal control s. Yet ,
as we saw in t he st udi es on at t ent i on al r eady descri bed,
cl ose i nspecti on of t he pr obl em i ndi cat es spl i t-brai n pa-
ti ents have not i ncreased t he amount of resources t hey
can call upon t o sol ve probl ems. In short, it appear s t hat
t he human brai n has a set amount of resources it can
al l ocat e t o cogni t i ve tasks and t hat t hese resources remai n
const ant f ol l owi ng commi ssur ot omy. How t hen do we ex-
pl ai n t hese t wo seemi ngl y di f f erent sets of resul ts? On t he
one hand, perf ormance seems bet t er than normal , whi l e
on t he ot her hand we have seen t here are resource l i mi t s
for percept ual and cogni t i ve tasks.
The conundrum f orces t he i ssue of wher e in a percep-
tual mot or t ask resource l i mi t at i ons are appl i ed. Are they,
f or exampl e, appl i ed duri ng t he earl y phases of i nf ormat i on
processi ng t hat deal wi t h t he compl exi t y of t he vi sual sti m-
ulus itself? Or, are t he resources appl i ed at l at er loci of
t he i nf ormat i on processi ng sequence t hat deal wi t h t he
mor e cogni t i ve aspect s of t he task? It woul d appear t hat
i nteracti ons bet ween t he hemi spheres on resource l i mi t s
occur when t he t ask is mor e cogni t i ve and i nvol ves work-
i ng memory. Lewi ne et al. (1994) have pr oposed a si mi l ar
scheme and suggest t he si te of subcort i cal i nt eract i on may
be t he brai n st em.
Language and Speech Processes of the Left
and Right Hemispheres
A di chot omy t hat is useful when tryi ng t o underst and lan-
guage is t he di st i nct i on bet ween how t he brai n enabl es
gr ammar and how it enabl es a l exi con. The gr ammar -
l exi con di st i nct i on (Pinker, t 993) is di f f erent f rom t he mor e
Neuron
224
t r adi t i onal synt ax- semant i c di st i nct i on t hat is commonl y
i nvoked to under st and t he di f f er ent i al ef f ect s of brai n le-
si ons on l anguage processes. In gener al t erms, gr ammar
refers to t he rul e-based system humans have f or or der i ng
wor ds to al l ow f or ef f ect i ve communi cat i on. The l exi con,
on t he ot her hand, i s t he mi nd' s di ct i onar y, wher e speci f i c
wor ds are associ at ed wi t h par t i cul ar meani ngs. The rea-
son f or usi ng t he f ormer di st i nct i on is t hat i t t akes i nt o
account such f act ors as memory, i n t hat wi t h memor y such
wor d st r i ngs as i di oms can be l earned by rote. Whi l e i t is
cl ear l y t he case t hat t he l exi con (memory) cannot under l i e
most phrases and sent ences, si nce t her e are endl ess
uni que sent ences such as t he one I am cur r ent l y wr i t i ng,
memory does pl ay a rol e i n many shor t phrases. Thus,
when uttered, such wor d st r i ngs do not ref l ect an under l y-
i ng i nt eract i on of synt ax and semant i c syst ems. They are,
i nst ead, essent i al l y an ent r y f r om t he l exi con. In more gen-
eral terms, a moder n vi ew woul d predi ct t her e ought t o
be brai n areas whol l y r esponsi bl e f or grammar, whi l e evi -
dence f or l ocal i zat i on of t he l exi con ought to be more el u-
si ve, si nce i t woul d ref l ect l ear ned i nf or mat i on and t hus
be part of t he gener al memor y/ knowl edge syst ems of t he
brai n. The gr ammar system ought to be di scr et e and t here-
f ore l ocal i zabl e. The l exi con shoul d be di st r i but ed and
t her ef or e more di f f i cul t to damage compl et el y.
Language and Speech Processes Can Rarely Be
Present in Both Hemispheres: Right Hemisphere
Language Has a Different Organizational
Structure Than Left
Whi l e t he separat ed l eft hemi spher e is f ul l y capabl e of
normal compr ehensi on of al l aspect s of l anguage, t he r i ght
hemi spher e can possess l anguage. When t here was evi -
dence of ri ght hemi spher e l anguage, t hese di sconnect ed
r i ght hemi spher es wer e sever el y l i mi t ed i n t he ki nds of
l i ngui st i c propert i es t hey possessed (Gazzani ga, 1970;
Zai del , 1990).
Over t he l ast 30 years, ver y f ew cases have been added
t o t he gr oup t hat demonst r at e some ki nd of l anguage i n
t he ri ght hemi spher e (revi ewed i n Baynes, 1990). In t he
ear l y ei ght i es, onl y f i ve cases i n al l seri es of spl i t -brai n
pat i ent s had pr oven to have a l exi con i n t he ri ght hemi -
sphere. Si nce t hat t i me we have been successf ul i n f i ndi ng
onl y one more nor mal l y r i ght - handed pat i ent wi t h a l exi con
i n bot h t he l eft and t he ri ght hemi spher es ( Baynes et al.,
1992).
The l eft and t he ri ght l exi cons of t hese speci al pat i ent s
can be nearl y equal i n t hei r capaci t y, but t hey are orga-
ni zed qui t e di f f erent l y. For exampl e, bot h hemi spher es
show a phenomenon cal l ed t he "word super i or i t y ef f ect "
(WSE). Normal readers are abl e to i dent i f y l etters bet t er
when t hey occur i n t he cont ext of real Engl i sh words, rat her
t han when t he same l etters occur i n pseudo- wor d (korad)
or nonwor d (t whx) st ri ngs. Si nce pseudo- and nonwor ds
do not have l exi cal entri es, l et t ers occur r i ng i n such st r i ngs
do not recei ve t he addi t i onal pr ocessi ng benef i t best owed
on words, and t hus t he WSE emerges. By t hi s vi ew, t he
WSE may be a usef ul measure of t he i nt egr i t y of t he vi sual
l exi con. When t he appr opr i at e st udi es assessi ng t hese
ef f ect s wer e car r i ed out wi t h each hemi spher e, i t was ap-
parent each hemi spher e i n t hese speci al pat i ent s has a
vi sual l exi con t hat pr ovi des an advant age f or r ecogni zi ng
l et t ers t hat occur i n real words. At t he same t i me, a cl oser
anal ysi s of t he dat a reveal ed each hemi spher e was usi ng
a di f f erent st r at egy i n pr ocessi ng t he wor ds. Exami nat i on
of t he react i on t i me data, f or exampl e, suggest ed t hat t he
r i ght hemi spher e t ook more t i me t o r espond as t he wor ds
became l onger. The l eft hemi spher e di d not , j ust as i s
t he case f or normal readers. In addi t i on, i n t he WSE t ask
descri bed above, t he error dat a reveal ed a di f f er ent pat-
t ern f or each hemi sphere. The l eft hemi spher e made more
errors on t he mi ddl e l etters of words, j ust as i n cont rol s,
whi l e t he ri ght hemi spher e made more errors on t he l ast
l etters of words. Fi nal l y, achi evi ng si mi l ar l evel s of accu-
r acy i n each hemi spher e requi red exposur e dur at i ons al-
most 10 t i mes as l ong i n t he nondomi nant hemi spher es.
Thi s resul t suggest s l et t er pr ocessi ng f or t he r i ght hemi -
spher e is seri al and not paral l el i n nat ur e, as is nor mal l y
t he case (Reut er-Lorenz and Baynes, 1992).
To l ocate wher e i n t he processi ng chai n t he hemi spher e
di f f er ences begi n, we eval uat ed each hemi spher e' s per-
f or mance on a l et t er pr i mi ng t ask. The t ask was si mpl y t o
i ndi cat e whet her a br i ef l y f l ashed upper case l et t er was
an "H" or a "T." On each t ri al t he upper case l et t er was
preceded by a l ower case l et t er t hat was ei t her an "h" or
a "t." Normal l y, subj ect s are si gni f i cant l y f ast er when an
upper case" H" is preceded by a l ower case "h," r at her t han
a l ower case "t." Thi s is a ver si on of t he cl assi c " pr i mi ng"
phenomenon f r equent l y used i n cogni t i ve sci ence t o get
at mechani sms of ment al processes. When usi ng l et t ers
i nst ead of wor ds i n a pr i mi ng task, i t is assumed ver y ear l y
st ages i n t he per cept ual - cogni t i ve cont i nuum are bei ng
exami ned.
The di f f er ence bet ween r esponse l at ency on compat i bl e
( h- H) ver sus i ncompat i bl e ( t - H) t ri al s is t aken t o be a mea-
sur e of l etter pr i mi ng. JW per f or med a l at er al i zed ver si on
of t hi s t ask i n whi ch t he pr i me was exposed f or 100 ms
t o ei t her t he r i ght or l ef t vi sual f i el d, and 400 ms l ater, t he
t ar get l et t er appear ed i n ei t her t he ri ght or l ef t vi sual fi el d.
Hi s per f or mance pr ovi des no evi dence of l et t er pr i mi ng
f or l eft vi sual f i el d t ri al s, but cl ear evi dence of pr i mi ng f or
r i ght vi sual f i el d t ri al s. Thus, t her e ar e no pr i mi ng phenom-
ena i n t he di sconnect ed r i ght hemi spher e. In summar y,
t her e can be t wo l exi cons, one i n each hemi spher e, but
t hi s l exi cal or gani zat i on is rare. When present , t he r i ght
hemi spher e l exi con seems t o be or gani zed di f f er ent l y t han
t he l exi con of t he l eft hemi spher e. These obser vat i ons
woul d be consi st ent wi t h t he vi ew t hat l exi cons ref l ect
l ear ni ng processes and as such woul d have a wi der di st ri -
but i on i n t he cerebral cort ex. Sti l l , i t woul d be f ol l y not to
not e t hat i n t he gener al popul at i on t he l exi con seems t o
be i n t he l eft hemi spher e. A r i ght hemi spher e l exi con i s
rarely present, and when i t is, it stores i nformati on di fferentl y.
Generative Syntax Is Present
in Only One Hemisphere
Whi l e t he r i ght hemi spher es of some pat i ent s cl ear l y have
l exi cons, t hese r i ght hemi spher es have shown errat i c per-
f or mance on ot her aspect s of l anguage such as under-
st andi ng verbs, pl ur al i zat i ons, t he possessi ve, act i ve/
Review: Human Split-Brain Studies
225
passi ve di f f er ences, etc. (Gazzani ga, 1970). The ri ght
hemi spher e i n t he pat i ent s t hat possess some l anguage
al so l acks a capaci t y to use wor d or der to di sambi guat e
st i mul i f or cor r ect gr ammat i cal meani ng. Nonet hel ess,
t hese ri ght hemi spher es can i ndi cat e when a sent ence
ends wi t h a semant i cal l y odd wor d (Kut as et al ., 1988).
Addi t i onal l y, r i ght hemi spher es t hat reveal l anguage ca-
paci t i es are abl e to make j udgment s of gr ammat i cal i t y
( Baynes and Gazzani ga, 1988; Baynes, 1990). Thus, even
t hough t hey cannot use synt ax t o di sambi guat e st i mul i or
to gui de compr ehensi on j udgment s, t hey can r ecogni ze
t hat one set of ut t erances is gr ammat i cal whi l e anot her
is not. Thi s f i ndi ng suggest s t hat pat t erns of speech are
l ear ned by rote. Yet, r ecogni zi ng t he pat t ern of accept abl e
ut t erances does not mean a neural syst em can use t hi s
i nf or mat i on t o assi st i n t he under st andi ng of wor d stri ngs
Some Right Hemispheres Can Develop Speech
One of t he hal l marks of most spl i t - br ai n pat i ent s is t hat
t hey speak out of t he l eft hemi spher e and not t he ri ght.
Thi s obser vat i on has been consi st ent wi t h t he neur ol ogi c
l i t erat ure and amyt al st udi es whi ch have shown t hat t he
l eft hemi spher e is t he domi nant hemi spher e f or bot h l an-
guage and speech ( Lennenber g, 1967).
Ther e are now t hr ee and possi bl y f our spl i t - br ai n pa-
t i ent s who seem abl e to speak out of each hemi spher e.
Whi l e t here is al ways an i ni t i al l y domi nant hemi spher e
f ol l owi ng brai n bi sect i on, some pat i ent s have devel oped
t he capaci t y to make one wor d ut t er ances f rom t he di scon-
nect ed r i ght hemi spher e ( Gazzani ga et al., 1979). In t hese
exper i ment s, i t became appar ent t hat st i mul i pr esent ed t o
t he l eft vi sual f i el d (ri ght hemi sphere) began to be named.
Or di nar i l y, one st i mul us present ed t o t he l eft hemi spher e
coul d be named. Thi s rat her st ar t l i ng devel opment rai sed
t he quest i on of whet her t he i nf or mat i on was somehow
t r ansf er r i ng to t he domi nant hemi spher e f or speech out -
put, or whet her t he r i ght hemi spher e i t sel f had devel oped
speech= Af t er a seri es of tests, i t was appar ent t hat t he
l atter opt i on was correct . The pat i ent s, f or exampl e, whi l e
abl e t o name an obj ect pr esent ed i n t he l ef t fi el d, i n t he
r i ght f i el d coul d not j udge i f t hey wer e t he same obj ect s
or not. Or, i f wor ds l i ke "f at her" wer e pr esent ed such t hat
t he f i xat i on poi nt fel l bet ween t he "t" and t he "h," t he pa-
t i ent s woul d say ei t her "fat" or "her," dependi ng on whi ch
hemi spher e spoke fi rst. Thus, i n t hese pat i ent s t wo of t he
t hr ee maj or syst ems i n human l anguage can be managed
to some ext ent by ei t her hemi spher e. It al so i l l ust r at es
an ext r aor di nar y pl ast i ci t y, occur r i ng somet i mes over 10
years af t er cal l osal surgery.
Hemisphere Specialization
One of t he car di nal f eat ur es of behavi or al st udi es on t he
brai n, bot h human and ani mal , has been t he r eal i zat i on
t hat speci f i c br ai n areas seem to be i nvol ved wi t h r at her
speci f i c per cept ual and cogni t i ve f unct i ons. The spl i t - br ai n
appr oach to t hi s i ssue has been st r ai ght f or war d. By t est i ng
each di sconnect ed hemi spher e, one can assess t he di ffer-
ent capaci t i es each mi ght possess. Whi l e some of t he
cl ai ms i n t hi s regard have become exagger at ed, t her e are
marked di f f er ences bet ween t he t wo hal f -brai ns (see Hel-
l i ge, 1993).
The Left Hemisphere P ossesses a Unique Capacity
to Interpret Behavior and Unconsciously Driven
Emotional States
A number of year s ago we obser ved how t he left, domi nant
speaki ng hemi spher e deal t wi t h t he behavi or s we knew
we had el i ci t ed from t he di sconnect ed r i ght hemi spher e
(Gazzani ga, 1995). We fi rst reveal ed t he phenomenon us-
i ng a si mul t aneous concept t est . The pat i ent is shown t wo
pi ct ures, one excl usi vel y to t he l eft hemi spher e and one
excl usi vel y to t he ri ght, and i s asked to choose from an
array of pi ct ures pl aced in f ul l vi ew i n f ront of hi m t he ones
associ at ed wi t h t he pi ct ures l at er al i zed to t he l eft and ri ght
brai n. In one exampl e of t hi s ki nd of test, a pi ct ur e of a
chi cken cl aw was f l ashed to t he l eft hemi spher e, and a
pi ct ur e of a snow scene to t he ri ght hemi spher e. Of t he
array of pi ct ur es pl aced i n f r ont of t he subj ect , t he obvi -
ousl y correct associ at i on is a chi cken f or t he chi cken cl aw
and a shovel f or t he snow scene. Case PS r esponded by
choosi ng t he shovel wi t h t he l eft hand and t he chi cken
wi t h t he right. When asked why he chose t hese i tems, hi s
l eft hemi spher e repl i ed, "Oh, t hat ' s si mpl e. The chi cken
cl aw goes wi t h t he chi cken, and you need a shovel t o cl ean
out t he chi cken shed." Here, t he l eft brai n, obser vi ng t he
l eft hand' s response, i nt erpret s t hat response i nt o a con-
t ext consi st ent wi t h i ts sphere of knowl edge, one t hat does
not i ncl ude i nf or mat i on about t he l eft hemi f i el d snow
scene. We cal l ed t hi s l eft hemi spher e pr ocess t he "i nter-
preter."
Thi s same gener al i dea has been obser ved when t he
l eft brai n i nt er pr et er st r uggl es to deal wi t h mood shi f t s,
produced i n t he exper i ment al si t uat i on by mani pul at i ng
t he di sconnect ed ri ght hemi spher e. A posi t i ve mood shi f t
t ri ggered by t he ri ght hemi spher e f i nds t he l eft i nt er pr et i ng
i ts cur r ent exper i ence i n a posi t i ve way. Si mi l arl y, when
t he ri ght t r i gger s a negat i ve mood state, t he l eft i nt erpret s
a pr evi ousl y neut r al si t uat i on i n negat i ve terms.
In recent st udi es we have been abl e to show one of t he
many i mpl i cat i ons f or havi ng a l eft hemi spher e i nt erpret er.
It has l ong been known t hat i nf er ence and i nt er pr et at i on
are i mport ant aspect s of normal memory f unct i oni ng (Bart-
lett, 1932), and one woul d predi ct t he t wo hemi spher es
mi ght respond di f f er ent l y i n some mnemoni c tasks. Phel ps
and Gazzani ga (1992) showed that, when spl i t -brai n pa-
t i ent s wer e shown pi ct ures r epr esent i ng a common scene,
t he t wo hemi spher es r esponded di f f er ent l y when tested
2 hr later. Thei r memory was t est ed wi t h a l at eral i zed y e s -
no recogni t i on test i n whi ch t he di st r act er pi ct ures wer e
ei t her consi st ent or i nconsi st ent wi t h t he or i gi nal scene.
The l eft hemi spher e perf ormed bel ow chance on consi s-
t ent di st ract er pi ct ures, wher eas t he ri ght hemi spher e was
above chance on t hese pi ct ur es and perf ormed at t he
same l evel of accuracy as t he pi ct ur es or i gi nal l y pre-
sent ed. In short, t he ri ght hemi spher e, whi ch has no i nter-
pret i ve mechani sm, rej ected as seen before pi ct ur es t hat
coul d have been part of t he story. The l eft hemi sphere,
on t he ot her hand, wi t h i ts capaci t y f or maki ng i nf er ences
and i nt erpret at i ons, was more st r ongl y i nf l uenced by t he
Neuron
226
expect at i ons f or act i ons common t o a scene and f al sel y
r ecogni zed pi ct ur es consi st ent wi t h t he obser ved scene.
Si mi l ar resul t s have r ecent l y been report ed by Met cal f e
et al. (1995).
Moni tori ng and P roduction of F ac i al Expressions
Are Managed by Di fferent Hemi s pheres
In t he per cept ual domai n, it appear s t hat t he ri ght hemi -
spher e has speci al processes devot ed to t he ef f i ci ent de-
t ect i on of upr i ght f aces ( Gazzani ga, 1989). Al t hough t he
l eft hemi spher e can al so per cei ve and r ecogni ze f aces
as wel l as reveal super i or capaci t i es when t he f aces are
f ami l i ar, t he r i ght hemi spher e appear s speci al i zed f or un-
f ami l i ar faci al st i mul i (Levy et al., 1972; Gazzani ga and
Smyl i e, 1983). Thi s pat t ern of asymmet r y has al so been
shown f or t he rhesus monkey (Hami l t on and Vermi ere,
1988).
Si nce t he r i ght hemi spher e is super i or f or percept i on of
faces, i t woul d be r easonabl e to suppose i t is al so speci al -
i zed f or t he management of f aci al expr essi ons. Recent
st udi es, however , have shown t hat , whi l e bot h hemi -
spher es can gener at e spont aneous f aci al expr essi ons,
onl y t he domi nant l eft hemi spher e can gener at e vol unt ar y
f aci al expr essi ons ( Gazzani ga and Smyl i e, 1990). In t hese
st udi es, si ngl e wor d commands such as "sm ile" or "f rown"
wer e gi ven to ei t her t he l eft or r i ght hemi spher e. Mi xed
i n wi t h t hese commands t hat woul d r equi r e cont r ol of l ower
f aci al muscl es woul d be ot her commands t hat cal l ed upon
t he upper f aci al muscl es, such as "bl ow, " "wi nk," and
"bl i nk. " Whi l e bot h hemi spher es wer e abl e to respond to
t he l atter commands, onl y t he l eft hemi spher e coul d re-
spond to t he commands of "smi l e" and "f rown. " The ri ght
hemi spher e responded at chance, t her eby suggest i ng t he
l ef t was speci al i zed f or t he pr oduct i on of vol unt ar y faci al
expr essi ons.
Hemi s phere S peci al i zati on S ens ori motor Tasks
Ther e are some t est s t hat br i ng out hemi spher e superi ori -
t i es i n some of t he pat i ent s. The bl ock desi gn test from
t he Weschl er Adul t I nt el l i gence Scal e is one such test.
Here, t he si mpl e task of ar r angi ng some red and whi t e
bl ocks t o mat ch t hose of a gi ven pat t ern can f i nd t he l eft
hemi spher e per f or mi ng poorl y whi l e t he r i ght t r i umphs
(Bogen and Gazzani ga, 1965). However, i n ot her pat i ent s
bot h hemi spher es when separat ed appear i mpai red, and
in sti l l ot hers t he l eft hemi spher e, i n addi t i on to speaki ng
and t hi nki ng, perf orms t hi s t ask wel l .
The same pat t er n of r esul t s as seen wi t h t he bl ock de-
si gn test is al so seen f or ot her t est s such as t he nonsense
wi r e f i gur e t est ( Mi l ner and Tayl or, 1972). Yet, f or both
t est s t he pr eval ent pi ct ur e woul d be t hat t he ski l l f r equent l y
seems l ocal i zed to t he ri ght hemi spher e. When t he capac-
i t y happens to be l at eral i zed i n t hi s way, it shoul d be easi er
to anal yze t han when t he processes i nvol ved are shared
bet ween t he t wo hal f -brai ns. The quest i on is, what is it?
The component s of t he bl ock desi gn t ask have not yet
been i dent i f i ed. We do know t hat a pat i ent who demon-
st rat es a ri ght hemi spher e super i or i t y f or t hi s ki nd of t ask
can show no super i or i t y on t he percept ual aspect s of t he
task. If a pi ct ure of t he bl ock desi gn pat t ern is f l ashed to
ei t her hemi spher e, each can easi l y f i nd t he match from
a seri es of pi ct ures. And, si nce each hand is demonst r abl y
dext erous, t he r i ght f or wr i t i ng and t he l ef t f or t hi s ki nd of
task, t he cr uci al l i nk must be i n t he mappi ng of t he sensor y
message ont o t he capabl e mot or syst em. It r emai ns f or
f ut ur e research t o under st and t hi s super i or i t y i n perfor-
mance when i t is seen i n one hemi spher e.
The L eft Hemi s phere Is S peci al i zed
for Intel l i gent Behavior
Fol l owi ng di sconnect i on of t he human cerebral hemi -
spheres, t he ver bal IQ of t he pat i ent r emai ns i nt act (Nass
and Gazzani ga, 1987; Zai del , 1990), and t he probl em sol v-
i ng capaci t y, such as seen i n hypot hesi s f ormat i on tasks,
r emai ns unchanged f or t he l eft hemi spher e ( LeDoux et
al., 1977a, 1977b). Whi l e t her e can be def i ci t s i n free recal l
capaci t y (i.e., to recal l ei t her ver bal l y or vi sual l y pr evi ousl y
l ear ned i t ems wi t hout t he benef i t of any cues or r ecogni t i on
ai ds) and i n some ot her per f or mance measures, t he over-
al l capaci t y f or pr obl em sol vi ng seems unaf f ect ed. In ot her
words, i sol at i ng essent i al l y hal f of t he cor t ex from t he dom-
i nant l eft hemi spher e causes no maj or change i n cogni t i ve
f unct i ons. The l eft r emai ns unchanged f rom i ts preopera-
t i ve capaci t y, whi l e t he l ar gel y di sconnect ed, same-si ze
r i ght hemi spher e is ser i ousl y i mpover i shed on a var i et y
of cogni t i ve tasks. Whi l e t he l ar gel y i sol at ed ri ght hemi -
spher e r emai ns super i or to t he i sol at ed l ef t hemi spher e
f or some act i vi t i es, such as t he r ecogni t i on of upr i ght
f ac es - - as wel l as ot her f unct i ons not cover ed i n t hi s re-
vi ew, such as some at t ent i onal ski l l s ( Mangun et al., 1994)
and per haps al so emot i onal processes (Gai not t i et al.,
1 9 9 3 ) - i t is poor at probl em sol vi ng and many ot her mental
act i vi t i es. A brai n system (t he r i ght hemi spher e) wi t h
r oughl y t he same number of neur ons as one t hat easi l y
cogi t at es (the l eft hemi spher e) is not capabl e of hi gher
or der cogni t i on. Thi s r epr esent s st rong evi dence t hat si m-
pl e cort i cal cel l number by i t sel f cannot f ul l y expl ai n human
i nt el l i gence ( Gazzani ga, 1994).
Concl usi ons
O v e r 30 years of spl i t -brai n research has pr ovi ded ext en-
si ve i nsi ght s i nt o t he or gani zat i on of t he human brai n. The
sur gi cal di sconnect i on of t he cerebral hemi spher es pro-
duces an ext r aor di nar y oppor t uni t y to st udy whi ch percep-
t ual and cogni t i ve processes are cort i cal i n nat ure and
whi ch are subcor t i cal . Vi sual percept ual i nf or mat i on, f or
exampl e, r emai ns st ri ct l y l at er al i zed to one hemi spher e
f ol l owi ng cal l osal sect i on. Tact i l e pat t er ned i nf ormat i on
r emai ns l at eral i zed. At t ent i onal mechani sms, however,
can i nvol ve subcor t i cal syst ems. Taken t oget her, cort i cal
di sconnect i on pr oduces t wo i ndependent sensor y i nfor-
mat i on pr ocessi ng syst ems t hat cal l upon a common atten-
t i onal r esour ce system i n t he car r yi ng out of percept ual
tasks.
Cort i cal di sconnect i on al so provi des cl ues to t he nat ure
of human memory. Fol l owi ng post eri or cal l osal sur ger y
t hat i nvol ves sect i oni ng t he hi ppocampal commi ssur e as
wel l , spl i t -brai n pat i ent s retai n normal r ecogni t i on memory
but are i mpai red on free recal l tasks. Thi s f i ndi ng not onl y
demonst r at es t hat t hese t wo processes are di ssoci abl e,
it al so suggest s t hat one rol e of t he hi ppocampal commi s-
sur es is to al l ow f or t he mul t i pl e r epr esent at i on of event s
t hat is known to be i mport ant in free recal l memory.
Review: Human Split-Brain Studies
227
Spl i t - br ai n st udi es have al so r eveal ed t he c ompl ex mo-
sai c of ment al pr ocesses t hat go i nt o human cogni t i on.
On t he one hand, each cer ebr al hemi spher e has i ts own
set of speci al i zed capaci t i es. The l eft hemi spher e, f or ex-
ampl e, is speci al i zed f or l anguage and speech, and maj or
pr obl em sol vi ng capaci t i es cr uci al f or i nt el l i gent behavi or .
It al so possesses a uni quel y human capaci t y t o i nt er pr et
behavi or and t o const r uct t heor i es about t he r el at i onshi ps
bet ween per cei ved event s and f eel i ngs. The ri ght hemi -
spher e, on t he ot her hand, is speci al i zed f or speci f i c t asks
such as f aci al r ecogni t i on, at t ent i onal moni t or i ng, and pos-
si bl y ot her ment al trai ts. It does not possess t he over al l
cogni t i ve capaci t i es of t he l eft brai n, whi ch f i nds it r eact i ng
mor e di r ect l y and si mpl y t o per cept ual i nf or mat i on.
Fi nal l y, spl i t - br ai n st udi es have demonst r at ed how spe-
ci fi c per cept ual and cogni t i ve ski l l s can be i sol at ed,
t hr ough di sconnect i on sur ger y, t o par t i cul ar cer ebr al ar-
eas. These ski l l s t hen ar e not r eveal ed by t hei r absence as
a resul t of a cor t i cal l esi on, whi ch in turn makes st r uct ur e-
f unct i on cor r el at i ons di f f i cul t t o make. The ski l l s ar e pres-
ent and can be demonst r at ed, but onl y when t est ed
t hr ough ei t her t he l eft or r i ght hemi spher e.
Acknowledgments
This work was aided by NIH grants NINDS 5 R01, NS22626-09, and
NINDS 5 P01 NS17778-012 and by the James S. McDonnell Foun-
dation.
January 12, 1995.
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