J Neurosurg 64:693-704, 1986

Review Article

The right hemisphere: neuropsychological functions

KENNETH M. HEILMAN, M.D., DAWN BOWERS, PH.D., EDWARD VALENSTEIN, M.D., AND
ROBERT T. WATSON, M.D.
Department of Neurology, College of Medicine, University of Florida, and Veterans Administration
Medical Center, Gainesville, Florida

v" In the past two to three decades, clinicians and neuroscientists have been studying the functions of the
fight hemisphere. Neither hemisphere seems to be dominant in the absolute sense. Each appears to be
specialized and is dominant for different functions. However, most functions require the cooperation of both
hemispheres. When one is damaged, the other can often compensate for the damaged one. Lesions of the left
hemisphere are associated with language (speech, reading, and writing) and praxic disorders, and lesions of the
fight hemisphere can result in visuospatial, attentional, and emotional disorders.
The authors review some of the major behavioral disorders associated with fight hemisphere dysfunction
and concentrate on three major types of disorders - - visuospatial, attentional, and emotional. Although not
all the behavioral defects associated with fight hemisphere damage can be subgrouped under these three types,
they are the ones most often associated with right hemisphere lesions.

KEY WORDS 9right cerebral hemisphere 9 visuospatial disorder 9

attentional disorder 9 psychological function 9 emotional disorder

A
FTER Broca's description ~8 of eight fight-handed
patients who had aphasia associated with left
hemisphere lesions, the left hemisphere became
known as the dominant (controlling) side. Other 19th
and early 20th century neurologists provided evidence
that the left hemisphere also mediated skilled move-
ment as well as cognitive activities such as reading and
writing. These reports provide additional evidence for
the dominance of the left hemisphere.
Clinicians were well aware that lesions of the fight
hemisphere may induce contralateral sensory and mo-
tor defects. Although there were reports of visuospa-
tial disorders associated with the fight hemisphere dys-
function, 55 the higher functions of the "non-dominant"
fight hemisphere were largely unrecognized. Even today
many clinicians think that, other than sensory or motor
function, the fight hemisphere mediates no important
This contribution is the 20th in a series of review articles
selected jointly by the Committee on Graduate Education of
the American Association of Neurological Surgeons and the
Congress of Neurological Surgeons, and the Journal of Neu-
rosurgery. - - Editor.
behavioral functions. Because the functions of the right
hemisphere are not recognized, surgeons are more likely
to operate on the right hemisphere. For example, ven-
tricular shunts are more often placed through the right
hemisphere. Surgeons also feel more at ease when re-
moving tumors and epileptic foci from the right hemi-
sphere association areas because they are less concerned
about the residual behavioral defects.
In the past two to three decades behavioral neurolo-
gists, neuropsychologists, neurosurgeons, and neuro-
physiologists have been studying the functions of the
right hemisphere. Neither hemisphere seems to be dom-
inant in the absolute sense. Each appears to be special-
ized and is dominant for different functions. However,
most functions require the cooperation of both hemi-
spheres. When one is damaged, the other can often
compensate for the damaged one. Left hemisphere le-
sions are associated with language (speech, reading, and
writing) and praxic disorders, and right hemisphere
lesions with visuospatial, attentional, and emotional
disorders.
In reviewing some of the major behavioral disorders

J. Neurosurg. / Volume 6 4 / M a y , 1986 693


associated with right hemisphere dysfunction, we shall
concentrate on three major types of disorders - - vis-
uospatial, attentional, and emotional. Although not all
behavioral defects associated with right hemisphere
damage can be subgrouped under these three types,
they are the ones most often associated with right
hemisphere lesions.
Visuospatial Disorders
Almost all human behavior has some spatial com-
ponent. For example, the ability to recognize and name
objects, letters, and printed words relies, at least in part,
on spatial discriminations. However, most of these ac-
tivities appear to be mediated by the left hemisphere.
Benton 8 noted important differences between the iden-
tification of objects and their localization in space.
These defects are dissociable; patients with right-sided
damage can often identify words, letters, objects, and
even faces, but many have difficulty in localizing objects
in space. Hughlings Jackson 55 was one of the first to
recognize the significance of visuospatial disorders. He
described a patient who had a tumor in the posterior
portion of the right hemisphere who lost his way and
could not dress himself.
Defective localization in space can be tested using a
large variety of tasks. For example, Warrington and
Rabin 94 presented pairs of cards containing dots in a
two-dimensional spatial position. The subjects were
asked whether the dots were located in the same place
on both cards. The performance of patients with right
parietal disease was poorer than the results of those
with left hemisphere disease or of a control group
without brain damage. Hannay, eta/., 43 also demon-
strated that right hemisphere-damaged patients have
defective localization. In both of these studies the in-
vestigators used relative rather than absolute localiza-
tion, and Benton 8 suggested that the right hemisphere
may be more important for allocentric (location in
relation to external cues) than egocentric (location in
relation to self) localization. Judgment of direction 9
and distance (depth perception) 22 may also be impaired
with right hemisphere damage. It is nevertheless impor-
tant to recognize that there are two types of depth
perception - - global and local stereopsis - - and it is
only the former (which does not depend on binocular
disparity) that is impaired with right hemisphere lesions.
Geographic and topographic orientation may be
tested in various manners. For example, Hfcaen, et
al., 44 had patients identify principal geographic land-
marks such as cities and rivers on a map and found
that those with right hemisphere retrorolandic lesions
performed more poorly than did control subjects with
left hemisphere damage. Although this type of test is
easy to perform in the clinic, Benton 8 ascribes to ne-
glect, rather than to visuospatial disorders, the propen-
sity of patients with right hemisphere damage to make
errors on this test.
Visuospatial skills are also necessary for building
694
K. M. Heilman, et al.
objects and drawing. A defect in spatial-organization
performance is referred to as constructional apraxia.
Many assembling and drawing tasks, including block
designs, stick constructions, construction of models,
drawing to command, and copying a drawing, have
been used to assess constructional praxis. Kleist 56 ini-
tially thought that constructional apraxia was associated
with left parietal dysfunction. Multiple studies have
since shown that constructional apraxia is more fre-
quent and more severe after right than after left hemi-
sphere disease (for a review seen Benton8). However,
some investigators have not found differences between
patients with right and those with left hemisphere dam-
age. Others have suggested that there may be qualitative
differences in the performance of patients with right
and those with left hemisphere damage. Left hemi-
sphere damage induces an executive-ideation defect,
and right hemisphere damage induces a visuospatial
defect.93 Unfortunately, this hemispheric dichotomy
has not been completely supported.
Patients with isolated right hemisphere disease not
only can name or recognize words, letters, objects, and
familiar people, but they also have no difficulty making
simple visual discriminations. However, when visual
discriminations become difficult and the patient must
rely more on spatial skill, 9'65 patients with right hemi-
sphere disease may have defective performance. Pa-
tients with right hemisphere damage may also have
difficulty discriminating between incomplete or muti-
lated figures. 58'92
Discriminating and matching unfamiliar faces ap-
pears to be different from recognizing familiar faces.
Although prosopagnosia (inability to recognize familar
faces) is associated with bilateral lesions, 64 the inability
to discriminate unfamiliar faces appears to be associated
with right hemisphere lesions. 7 Unlike the patients with
prosopagnosia who have bilateral inferior temporo-oc-
cipital lesions, patients with an inability to discriminate
unfamiliar faces usually have right parietal lesions. That
most patients with prosopagnosia can discriminate un-
familiar faces suggests that the inferior temporo-occip-
ital regions are not critical for this type of visuospatial
function. Most patients with profound visuospatial dis-
orders from right parietal disease also have little diffi-
culty in recognizing familiar faces.
Studies in monkeys led Mishkin and co-workers66 to
conclude that the striate cortex is the source of two
multisynaptic corticocortical pathways. One courses
ventrally, connecting the striate, prestriate, and inferior
temporal areas; the other runs dorsally interconnecting
with the striate, prestriate, and parietal lobe. Although
the ventral system seems important for recognition of
objects, the dorsal system appears to influence spatial
localization. In humans, ventral pathways also appear
important in object recognition. However, the dorsal
system in humans appears to be lateralized, the right
parietal lobe having a critical role in visuospatial proc-
essing.
J. Neurosurg. / Volume 64/May, 1986
The fight hemisphere: neuropsychological functions
Attentional and Intentional Disorders
When patients with fight parietal disease are asked
to draw a picture, they often fail to draw the left side.
This type of defect was thought to reflect a visuospatial
defect and had been termed "visual spatial agnosia. "63
Visuospatial agnosia, or hemispatial neglect, is part of
the neglect syndrome, which is a group of behavioral
disorders in which a subject fails to report, respond to,
or orient toward novel or meaningful stimuli. The
response failure occurs when the patient has to move a
contralateral limb or move a body part toward or in
contralateral hemispace. The report failure occurs when
stimuli are presented contralateral to a brain lesion.
Although elements of the neglect syndrome can follow
left hemisphere lesions, neglect is more frequent and
severe after fight hemisphere lesions. Four major ele-
ments of the neglect syndrome will be discussed sepa-
rately in this review: 1) sensory neglect or hemi-inatten-
tion; 2) extinction to simultaneous stimuli; 3) motor
neglect; and 4) hemispatial neglect.
Sensory Neglect
For hemi-inattention or sensory neglect to be tested,
somatesthetic, visual, and auditory stimuli should be
presented in a random order to the sides ipsilateral and
contralateral to the lesion. Although sensory loss may
be the most common cause of a failure to report or
respond to a stimulus presented contralateral to the
damaged hemisphere, patients and animals with lesions
in locations other than a primary sensory area or sen-
sory projection system may also fail to report or respond
to stimuli. This is termed "sensory neglect." If the lesion
site is unknown, one may be unable to distinguish
hemianesthesia or hemianopsia from severe inatten-
tion. Unilateral central nervous system lesions, how-
ever, do not produce a contralateral hearing loss. Even
patients with a peripheral hearing loss will respond to
unilateral stimuli unless the sound is of low amplitude
and close to the ear. Therefore, when patients do not
respond to unilateral auditory stimuli, they usually have
sensory neglect.
Several theories have been proposed to account for
sensory neglect, which has been postulated to be in-
duced by decreased sensory input. T M Brain 17 believed
that the parietal lobes contain the body schema and
also mediate spatial perception. Parietal lesions, there-
fore, caused the patient to fail to recognize only half his
body and half of space. Neglect in humans and mon-
keys can be induced by lesions in many different brain
regions, including the temporoparietal occipital junc-
tion; 25'52 limbic areas, such as the cingulate gyrus; 52'96
and subcortical areas, such as the thalamus 95 and mes-
encephalic reticular formation. 73'97Heilman and Valen-
stein 49 and Watson, et al., 96"99 postulated that sensory
neglect is an attentional-arousal disorder induced by
dysfunction in a corticolimbic reticular formation loop.
Since the classic work of Moruzzi and Magoun, 68 it
J, Neurosurg. / Volume 64/May, 1986
has been recognized that the reticular activating system
is critical for arousal, a physiological state of heightened
excitability that prepares the organism for sensory and
motor processing. Whereas bilateral lesions of the retic-
ular activating system induce coma, unilateral lesions
induce contralateral neglect. The reticular activating
system does not have direct connections to the cortex.
The two possible mechanisms by which this system
may influence arousal are a polysynaptic pathway to
the cortex through the thalamus and a modulating
activity of the thalamic reticular nucleus (for review see
Watson, et a/.99). The thalamic reticular nucleus is a
thin nucleus that envelops the thalamus, projects to the
thalamic relay nuclei, and inhibits relay to the cortex. 81
The reticular activating system projects to the thalamic
reticular nucleus and inhibits this inhibitory nucleus,
allowing increased transmission of sensory information
to the cortex. 84
In the human and the ape, the primary sensory cortex
projects to modality-specific sensory association areas,
which project to polymodal sensory areas, including the
temporoparietal region. 69 The polymodal sensory areas
may be important in modeling (detecting stimulus nov-
elty). When a stimulus is novel, multimodal conver-
gence areas may have a general inhibitory action on the
thalamic reticular nucleus or may activate the mesen-
cephalic reticular formation, thereby providing an
arousal response after cortical analysis.
Unlike stimulus novelty, stimulus significance is de-
termined partly by the needs of the organism. Limbic
system input into the temporoparietal region may pro-
vide information about biological needs. Since the fron-
tal lobes are critical in goal-oriented behavior and de-
veloping sets, the frontal lobes may provide input about
goals that are neither stimulus-dependent nor moti-
vated by an immediate biological need. The frontal
lobes and cingulate cortex have strong input into the
temporoparietal region. These connections may pro-
vide an anatomical substrate by which motivational
states may influence stimulus processing. 4599 Cellular
recordings from the parietal lobe have shown that some
neurons enhance their activity when an animal is pre-
sented with motivationally significant visual stimuli.
The activity of these neurons is spatially selective. Re-
cordings from parietal neurons suggest that the parietal
lobe is subserving attentional processes. 2~
In summary, the temporoparietal region receives
polymodal sensory, limbic, and frontal input; can acti-
vate the reticular activating system; and has cells that
show increased activity when presented with significant
stimuli. Because the temporoparietal region appears to
be mediating attentional processes, it should not be
surprising that sensory neglect occurs when the region
is ablated.
In humans, neglect is more frequent and severe after
fight than after left hemisphere lesions. ~7'25The atten-
tional cells found in the parietal lobe of monkeys by
Lynch62 and Robinson, eta/., 75 usually have contralat-
695

eral receptive fields, but some of these neurons have
bilateral receptive fields. That is, some respond to stim-
uli presented in the right or the left visual half-fields.
To account for a hemispheric asymmetry of attention
in humans, we suggest that the temporoparietal regions
of the human brain also have attentional or compara-
tor neurons, but that the cells in the right hemisphere
are more likely to have bilateral receptive fields than
are cells in the left hemisphere. Thus, cells in the left
hemisphere would be activated predominantly by novel
or significant stimuli in the right hemispace or hemi-
field, but cells in the right hemisphere would be acti-
vated by novel or significant stimuli in either visual
field or either side of the hemispace, or both. If this
were the case, right hemisphere lesions would more
often cause inattention than would left hemisphere
lesions. When the left hemisphere is damaged, the right
can attend to ipsilateral stimuli, but the left cannot
attend to ipsilateral stimuli after right-sided damage.
Support for this hypothesis has been provided by both
physiological5~and imaging studies. TM
Extinction to Simultaneous Stimulation
Most patients improve after initial sensory neglect.
At first they ignore stimuli presented to the side opposite
the lesion, but they eventually become able to detect
and lateralize these stimuli correctly. When given bilat-
eral stimulation, however, they often fail to report the
stimulus presented to the side contralateral to the lesion.
This phenomenon, first noted by Loeb 6~to occur in the
tactile modality, has been termed "extinction to double
simultaneous stimulation. ''6 It may also occur in the
auditory and visual modalities.6
The methods used to test for extinction are similar
to those used to test for sensory neglect. If the patient
responds normally to unilateral stimulation, bilateral
simultaneous stimulation may be used. Unilateral stim-
uli should be randomly interspersed with bilateral si-
multaneous stimuli. The lesions causing extinction are
often in the same areas as lesions that cause sensory
neglect. However, certain forms of extinction may also
occur after lesions of the corpus callosum, 85 and even
left-sided extinction has been reported to follow left
hemisphere lesions. 82
Different mechanisms may underlie extinction in
patients with callosal lesions, sensory defects, or hemi-
spheric lesions. Such mechanisms are poorly under-
stood, but several explanatory hypotheses have been
advanced. Because patients with partial deafferentation
may have extinction, several authors have postulated a
sensory mechanism to explain sensory neglect and sen-
sory extinction from parietal lesions.6 Psychophysical
methods have been used to demonstrate that in normal
subjects sensory threshold increases on one side when
the opposite side is stimulated (obscuration). ~~If this
obscuration phenomenon occurs in a patient with an
elevated threshold caused by an afferent lesion, it would
appear similar to extinction in patients without deaffer-
696
K. M. Heilman, et al.
entation. The mechanism underlying obscuration, how-
ever, is not known.
Several investigators have suggested that extinction
and perhaps obscuration result from suppression or
reciprocal inhibition. In the case of cerebral damage,
the normal hemisphere inhibits the damaged hemi-
sphere more than the damaged hemisphere inhibits the
normal one. Consequently, stimuli contralateral to the
damaged hemisphere are not perceived when the nor-
mal side is stimulated. The physiological mechanisms
that induce this reciprocal inhibition are unknown.
Each association cortex may project not only to the
ipsilateral but also to the contralateral thalamic reticular
nucleus. Unlike the ipsilateral connections, which are
inhibitory, the contralateral projections may be facili-
tatory.
Another explanation for extinction, the limited atten-
tion theory, specifies that under normal circumstances
bilateral simultaneous stimuli are processed simulta-
neously, with each hemisphere processing the contra-
lateral stimulus. However, a damaged hemisphere may
be unable to attend to contralateral stimuli, making the
organism inattentive to those stimuli. As the organism
recovers, it becomes capable of attending to contralat-
eral stimuli. This improvement may be mediated by
the normal ipsilateral hemisphere. The normal hemi-
sphere, however, may have a limited attentional capac-
ity. Therefore, with bilateral simultaneous stimulation,
the normal hemisphere's attentional mechanism is oc-
cupied with the contralateral stimulus and may be
unable to attend to the ipsilateral stimulus.
Motor Neglect (Hemiakinesia)
An aroused animal can selectively attend to a stim-
ulus and either act or not act. The decision is based not
only on the nature of the stimulus but also on the state
of the organism. The physiological state in which one
prepares for an action is called "intention." A disorder
of intentional processes leads to akinesia.
Akinesia is behaviorally defined as the inability to
initiate a movement in an aroused organism in which
the lack of movement cannot be attributed to a defect
in the motor unit or pyramidal system. Akinesia may
also be expressed as less forceful movements, move-
ment of decreased amplitude (hypometria), and slower
movement (bradykinesia). Some patients have in-
creased contralateral limb akinesia when they must use
their ipsilateral extremities simultaneously. This has
been termed "motor extinction. ''s9
Lesions of the dorsolateral frontal lobe, especially the
region of the frontal eye field (Brodmann's area) may
induce contralateral akinesia. 98 The connections of the
frontal eye fields are important in understanding the
possible role of the dorsolateral frontal lobe in inten-
tion. 99 It has reciprocal connections with unimodal and
polymodal posterior sensory association cortices23 and
is an area of sensory convergence. ~2 The dorsolateral
frontal lobe has reciprocal connections with medial
J. Neurosurg. / Volume 6 4 / M a y , 1986
The right hemisphere: neuropsychological functions
(nonspecific) thalamic nuclei. Projections to the mes-
encephalic reticular formation,57 as well as nonrecipro-
cal projections to the caudate nucleus, also exist. Last,
the dorsolateral frontal lobe also receives input from
the limbic system, mainly from the anterior cingulate
gyrus.3
Its connections with neocortical sensory association
and sensory convergence areas may provide the frontal
lobe with information about external stimuli that may
call the organism to action. The limbic connections
(anterior cingulate gyrus) may provide the frontal lobe
with motivational information. Connections with the
mesencephalic reticular formation may be important
in arousal, and connections with the basal ganglia and
premotor regions may be related to motor preparation.
Physiological studies support the hypothesis that the
frontal lobe mediates intentional activity. 38 Lesions in
many of the regions that connect with the frontal lobe
may also induce contralateral akinesia, including poly-
modal sensory cortex (temporoparietal regions)9~ and
supplementary motor area cingulate regions (Meador,
et al., in preparation). Neglect or akinesia may also
accompany lesions of the "nonspecific" intralaminar
thalamic nuclei, 98 which project both to the frontal
lobes and to the neostriatum. Akinesia may also result
from lesions of the basal ganglia and ventral thalamus
(ventralis lateralis and ventralis anterior). 9~ The basal
ganglia project to the ventral thalamus, and this "mo-
tor" portion of the thalamus is also gated by the nucleus
reticularis thalami. The reticular nucleus of the thala-
mus may be inhibited by the mesencephalic reticular
formation during an arousal or orienting response and
be inhibited by the frontal lobes during a motor set.99
Much evidence suggests the importance of dopami-
nergic neurons in the mediation of intention. Marked
defects of intention have long been known to be prom-
inent in patients with Parkinson's disease, which is
characterized pathologically by degeneration of ascend-
ing dopaminergic neurons. In animals, unilateral le-
sions in these pathways cause unilateral motor neglect.
Patients with fight hemisphere lesions have a more
severe contralateral limb akinesia than do patients with
left hemisphere lesions (Coslett and Heilman, in prep-
aration). Howes and Boiler54demonstrated that patients
with fight hemisphere lesions had slower reaction times
of the ipsilateral hand than did patients with left hemi-
sphere lesions. The slowing of reaction time was not as
dependent on lesion size as on location. Right hemi-
sphere lesions appeared to induce the most profound
slowing. These investigators did not mention whether
the patients with ipsilateral slowing had unilateral ne-
glect. In monkeys, no hemispheric asymmetries in the
production of the neglect syndrome have been noted.
However, monkeys with lesions inducing neglect had
slower ipsilateral reaction times than did monkeys with
equal-sized lesions that did not induce neglect (Valen-
stein, et al., in preparation).
It has been shown that warning stimuli may prepare
J. Neurosurg. / Volume 6 4 / M a y , 1986
an organism for action and thereby reduce reaction
times? 9 Because patients with fight hemisphere lesions
have been shown to have reduced behavioral evidence
of activation,54it has been postulated that in the human
the right hemisphere may dominate in mediating the
intention (activation) process? ~ That is, the left hemi-
sphere prepares the fight extremities for action, and the
fight prepares both. Therefore, with left-sided lesions,
left limb akinesia is minimal, but with fight-sided le-
sions, there is severe left limb akinesia. In addition,
because the right hemisphere is more involved than the
left in activating the right extremities with fight hemi-
sphere lesions, there is more ipsilateral hypokinesia than
with left hemisphere lesions.
That the fight hemisphere dominates mediation of
activation or intention (physiological readiness to re-
spond) has been demonstrated in normal subjects. They
show more activation (measured behaviorally by reac-
tion times) with warning stimuli delivered to the fight
hemisphere than with warning stimuli delivered to the
left hemisphere. That is, warning stimuli projected to
the fight hemisphere reduced reaction times of the right
hand more than warning stimuli projected to the left
hemisphere reduced reaction times of the right hand.
Warning stimuli projected to the fight hemisphere re-
duced reaction times even more than did warning stim-
uli projected directly to the left hemisphere. These
results support the hypothesis that the fight hemisphere
dominates intention (activation, preparation)? ~
Hemispatial Neglect
When patients with hemispatial neglect are asked to
perform a variety of behavioral tasks, they neglect the
hemispace contralateral to their lesion. For example,
when asked to draw a picture of a flower, they may
draw only half of the flower. When asked to bisect a
line, they may quarter it instead, and they may fail to
cross out lines distributed over a page.
The lesions associated with hemispatial neglect are
similar to those associated with motor and sensory
neglect. Although hemianopsia may enhance the symp-
toms of hemispatial neglect, hemianopsia alone cannot
entirely account for the deficit. First, some patients with
hemispatial neglect are not hemianopsic.63 Second,
hemianopsic patients without neglect do not perform
differently from nonhemianopsic patients in the visual
discrimination of bisected lines. TM
The defective performance cannot be explained by a
visual half-field defect, but it may be a hemispatial
defect. Hemispace is a complex concept because it can
be defined according to eye, head, or trunk position.
With the eyes and head facing directly forward, the
hemispaces defined by these three standards are con-
gruent. But if the eyes are directed to the far right, for
example, the left visual field falls in large part in the
fight hemispace, as defined by the head and body
midline. Similarly, if the head and eyes are turned far
to the right, left head and eye hemispace can both be
697

in the right hemispace of the body. There is evidence
to suggest that head or body hemispace, or both, is
important in determining the symptoms of hemispatial
neglect45 (Coslett, et al., in preparation). For example,
when patients with left hemispatial neglect are asked to
bisect a line, their performance is poorer in left body
hemispace than it is in right body hemispace even when
using a strategy that ensures that the line is seen in the
normal right visual field. Similarly, independent of
visual field and body hemispace, lines are more poorly
bisected in left head hemispace than in right head
hemispace.
The abnormal performance of patients in contralat-
eral space suggests that brain mechanisms relating to
the opposite hemispace have been disturbed. It suggests
that each hemisphere is responsible not only for receiv-
ing stimuli from the contralateral visual field and for
controlling the contralateral limbs, but also for atten-
tion and intention in contralateral hemispace independ-
ent of which visual field the stimulus enters or which
hand is used. 45 The postulate that each hemisphere
attends to and intends in contralateral hemispace has
been supported by studies of normal subjectsJ 6
Several neuropsychological mechanisms could ac-
count for the hemispatial defect associated with neglect.
Bisiach and Luzzatti ~3proposed a representational map
hypothesis similar to Brain's body schema hypothesis.17
They asked two patients with fight hemisphere damage
to describe from memory a familar scene, the main
square in Milan, from two different spatial perspectives
(one facing the cathedral and the other facing away
from the cathedral). Regardless of the patients' orienta-
tion, left-sided details were omitted. On the basis of the
findings from these two patients and from a second
study, TM these investigators postulated that the mental
representation of the environment is structured topo-
graphically and is mapped across the brain. That is, the
visual image conjured up in the mind may be split
between the two hemispheres (like the projection of a
real scene). With right hemisphere damage, there is a
representational disorder for the left half of this image.
The representation map postulated by Bisiach and co-
workers ~3'~4may be hemispatially organized so that left
hemispace is represented in the right hemisphere and
right hemispace is represented in the left hemisphere.
Hemispatial neglect may be associated with a gaze
defect which might prevent patients with hemispatial
neglect from fully exploring the neglected side of the
line. Partial support for the gaze hypothesis is provided
by De Renzi, et al., 28 who demonstrated that patients
with right hemisphere lesions and neglect have eye
deviation to the right.
Patients with hemispatial neglect may have a hemi-
spatial attentional deficit. Although the line is "seen"
in the normal visual field, the line is in neglected hemi-
space and therefore is not attended to. Patients with
neglect may also have a hemispatial or directional aki-
nesia. 53 When patients were asked to point with their
698
K. M. Heilman, et al.
normal pointing hand with their eyes closed to an
imaginary point directly in front of them, they pointed
toward right hemispace. 46 In addition, when directional
reaction times were measured, patients with hemispatial
neglect were slower initiating a response toward left
than right hemispace) 3 These results cannot be ac-
counted for by inattention, representation disorders, or
gaze defects; therefore, they provide evidence for a
hemispatial or directional akinesia. Although these stud-
ies support a directional or hemispatial akinesia, or
both, they do not refute alternative hypotheses. Atten-
tion, intention, gaze, and memory (representations) are
all closely linked functions.
Lesions in the right hemisphere induce hemispatial
neglect more often than do those in the left hemisphere.
The neglect induced by right hemisphere lesions is also
more severe, t.33 The mechanism underlying this asym-
metry is unknown; however, the right hemisphere may
be able to intend and attend in and toward both hemi-
spatial fields, but the left may be able to attend and
intend only in and toward the right hemispatial field.
With left hemisphere lesions, therefore, the right hemi-
sphere will attend and intend in and toward ipsilateral
(fight) hemispace. However, with right hemisphere le-
sions, the left hemisphere attends and intends in and
toward the right hemispace; the left hemispace is ne-
glected.
Affeetive Disorders
Babinski 2 noted that patients with right hemisphere
disease often appear indifferent or euphoric. Goldstein 4~
noted that patients with left hemisphere lesions and
aphasia often showed a profound depression, which he
termed the "catastrophic reaction." These clinical ob-
servations were also seen by Gainotti 3~who studied 160
patients with lateralized brain damage.
Affective Comprehension
Auditory-Verbal Comprehension. Patients with
right hemisphere lesions might have a defect in the
comprehension of affect. Developing an appropriate
emotion depends, in part, on comprehending speech.
Speech can simultaneously carry at least two different
types of information - - the linguistic content (what is
said) and the affective content (how it is said). The
linguistic content is conveyed by a complex code that
requires semantic and phonemic decoding. In contrast,
the affective content is conveyed by the pitch, tempo,
and tonal contours of speech 7~ and consequently re-
quires a different type of processing.
In the vast majority of persons (particularly fight-
handed individuals), the left hemisphere is clearly more
adept than the right in decoding the linguistic content
of speech. Until about a decade ago little was known
about how affective intonations were being mediated.
To determine whether the fight hemisphere is more
adept than the left in decoding the affective components
of speech, we read sentences with semantically neutral
J. Neurosurg. / Volume 64/May, 1986
The right hemisphere: neuropsychological functions
content (such as "The boy went to the store.") in four
different emotional intonations (happy, sad, angry, in-
different) to patients with right temporoparietal infarc-
tions and to those with aphasia and left temporoparietal
infarctions. The patients' task was to identify the emo-
tional tone of the speaker. Patients with fight hemi-
sphere lesions performed worse on this task than did
those with left hemisphere lesions; suggesting that the
fight hemisphere is more critically involved in process-
ing the affective intonations of speech. 47 Further sup-
port for the hypothesis that the two hemispheres process
different aspects of speech (semantic content vs affec-
tive intonations) has been drawn from dichotic listening
studies with normal adults. 42
The neuropsychological defect underlying the im-
paired ability of patients with right hemisphere disease
to identify affective intonations in speech is not entirely
clear. The defect may be related to a cognitive inability
to denote and classify affective stimuli. It could also
be related to an inability to discriminate perceptually
between affective intonations in speech. In a subse-
quent study, we88 replicated our previous finding and
attempted to determine whether patients with fight
hemisphere disease could, in fact, discriminate between
affective intonations of speech without having to clas-
sify or denote these intonations. Patients were required
to listen to identical pairs of sentences spoken in either
the same or different emotional tones. The patients did
not have to identify the emotional intonation, but had
to tell whether the intonations associated with the sen-
tences sounded the same or different. Patients with right
hemisphere disease performed more poorly on this task
than did patients with left hemisphere disease, which
suggests that the perceptual discrimination between
affectively intoned stimuli was impaired in the patients
with fight hemisphere damage.
Emotional messages may be conveyed by proposi-
tional speech. Graves, et al., 4~ visually presented emo-
tional and nonemotional words to aphasic patients with
left hemisphere lesions. Their task was to read the
words. Unlike speech, which can be affectively intoned,
the written word carries no prosody. Nevertheless,
Graves and co-workers found that emotional words
were read better than were nonemotional words. These
authors also studied normal adults with emotional and
nonemotional words tachistoscopically presented to the
left and right visual half-fields. In the left visual half-
field (right hemisphere) there was a relative superiority
for the recognition of emotional words over nonemo-
tional words.
Visual-Nonverbal Comprehension. The develop-
ment of an appropriate emotional state also depends
on perceiving and comprehending visual stimuli, such
as facial expression and gesture. Gardner, et al.,35 found
that patients with right hemisphere disease and those
with left hemisphere disease were equally impaired in
selecting the most humorous of a group of cartoons
J. Neurosurg. / Volume 6 4 / M a y . 1986
without captions. Patients with right hemisphere disease
performed better with captions.
We gave a variety of affective visual tasks to patients
with left or fight hemisphere lesions as well as to neu-
rological patients without hemispheric disease. 26 We
asked our subjects to discriminate between a pair of
neutral faces (saying, for instance, "Are these two faces
the same person or two different people?"); to name
the emotion expressed by a face (happy, sad, angry,
indifferent); to select a "target" emotion from a multi-
ple-choice array of faces (saying, for instance, "Point to
the happy face."); and to determine whether two pic-
tures of the same person's face expressed the same or a
different emotion.
Compared with other groups, patients with right
hemisphere disease were markedly impaired in their
ability to discriminate between pairs of neutral faces, as
has been previously discussed. Although both left and
right hemisphere-damaged groups had difficulty nam-
ing and selecting emotional faces, there was a trend for
patients with right hemisphere disease to perform more
poorly on these two tasks than patients with left hemi-
sphere disease. In addition, patients with right hemi-
sphere disease were more impaired in making same/
different discriminations between emotional faces.
When performance across these various affective
tasks was covaried for neutral facial discrimination (a
visuospatial nonemotional task), differences between
the two groups disappeared. This finding suggests that
the defect underlying poor discrimination of neutral
faces observed in the group with right hemisphere dis-
ease might also cause their inability to recognize and
discriminate between emotional faces. However, the
poor facial discrimination by the group with right hemi-
sphere disease did not entirely correlate with their abil-
ity to recognize and discriminate between emotional
faces. Retrospective review of the data revealed that
about one-third of the patients with right hemisphere
disease performed poorly on both the neutral facial
discrimination task and the emotional faces task,
whereas about one-third performed well on both. The
remaining patients with right hemisphere disease, how-
ever, performed relatively well on neutral facial discrim-
ination but poorly on the emotional faces task.
We therefore believe that the right hemisphere might
be important for perceiving faces and facial expressions.
These processes may be either interdependent (one
must perceive faces before perceiving the emotions
expressed by the faces), or they may be independent
but share the same or a contiguous anatomical locus.
Cicone, et al., 24 also presented emotional faces to pa-
tients with right and left hemisphere damage and again
found that right hemisphere-damaged patients had im-
paired ability to recognize emotional faces. We 15 have
examined a patient who had a large right hemisphere
tumor in the posterior region that did not appear to
involve cortical structures. It did involve subcortical
association fibers, some of which may have been com-
699

missural fibers that course to and from the left hemi-
sphere. This patient performed flawlessly on the neutral
and emotional facial discrimination task, but had great
difficulty in naming and selecting named emotional
faces. His test performance suggested that the ability to
recognize or match faces can be dissociated from the
ability to recognize emotional faces. A right hemisphere
disconnection from the left hemisphere may have
caused his inability to name emotional faces or to point
to a face named by the examiner.
Affective Expression
Verbal Expression. Previous research with normal
and brain-damaged subjects has suggested that there are
hemispheric differences in the ability to comprehend
affectively intoned speech. A hemispheric difference
may also exist in the ability to express various emotions
in speech. Patients with left hemisphere disease are
often aphasic and thus have difficulty using proposi-
tional speech. They are impaired in the ability to convey
emotional content with a semantic-propositional con-
text. However, as noted by Hughlings Jackson (see
Taylor87), some aphasic patients can, by varying tempo,
pitch, and timbre, convey a rich variety of emotional
feeling, despite their inability to produce propositional
speech. Patients with left hemisphere lesions who have
severe nonfluent aphasia and limited ability to express
propositional speech not only can affectively intone
words and sounds but also may be very fluent when
using emotional words, especially expletives.
We attempted to determine whether patients with
right hemisphere disease can express emotionally in-
toned speech 88 by asking them to say semantically neu-
tral sentences (such as "The boy went to the store."),
and to use a happy, sad, angry, or indifferent tone of
voice. These patients were severely impaired. Typically,
they spoke sentences in a flat monotone and often
denoted the target affect (for example, "The boy went
to the store, and was sad."). Ross and MesulamTM de-
scribed two patients who could not express affectively
intoned speech but could comprehend affective speech.
R o s s 77 has also described patients who could not com-
prehend affective intonations but could repeat affec-
tively intoned speech. He has postulated that right
hemisphere lesions may disrupt affective speech (in
terms of comprehension, repetition, and production) in
the same manner that left hemisphere lesions disrupt
propositional speech.
Facial Expression. Buck and D u f f y 19 studied the
ability of fight and of left hemisphere-damaged patients
to show emotional facial expressions in response to
viewing slides such as those of familiar people, an
unpleasant scene (such as a starving child or a crying
woman), and unusual pictures (such as double expo-
sures). The patients' faces were videotaped as they
viewed the slides and were later judged. The results
showed that fight hemisphere-damaged patients were
700
K. M. Heilman, et al.
less facially expressive than left hemisphere-injured sub-
jects.
Occasionally, patients who cannot move their face
normally on command can have pathological outbursts
of crying or laughing. Usually these emotional outbursts
are not under volitional control, and the patients will
often tell the examiner that their facial expression is not
always consistent with their mood. Most patients with
pathological laughing or crying have bilateral subcorti-
cal lesions. 72 Most observers have proposed that this
laughing or crying may be induced by disinhibition of
a subcortical region where these behaviors are pro-
grammed. Sackeim, eta/., TM retrospectively analyzed
such cases and noted that, although most of these
patients have bilateral lesions, those with pathologi-
cal laughing have larger lesions on the right side and
those with pathological crying have larger lesions on the
left side.
Affective Memory
In one of the few studies of affective memory,
Wechsler ~~176presented right and left hemisphere-dam-
aged patients with two stories - - one designed to elicit
an affective response, the other a neutral response. For
example, the emotional story told of a kind man "who
was very sick and his doctors were unable to cure him.
He sent for his wise men who told him he would get
well if he wore the shirt of a truly happy man." Im-
mediately after a story was read to the patients, they
were asked to reproduce it verbally. The score was based
on the total units recalled (similar to logical memory
subtest of the Wechsler Memory Scale). The patients
with left hemisphere lesions made fewer errors on the
emotionally charged story than did patients with right
hemisphere disease; there were no differences in their
performance on the neutral story. However, it was also
noted that even left hemisphere-damaged subjects,
compared with normal controls, had more difficulty
with the emotionally charged stories than with neutral
stories. In normal subjects, affectively charged facts are
recalled better than neutral stories.
Underlying Affect
The studies thus far have suggested that patients
with fight hemisphere disease have more difficulty than
those with left hemisphere disease in comprehending
and expressing affectively intoned speech as well as in
comprehending emotional facial expressions. Patients
with right hemisphere disease may also have more diffi-
culty in comprehending or remembering emotionally
charged speech. These perceptual, cognitive, and ex-
pressive deficits must underlie and account for the
different emotional reactions of patients with fight ver-
sus left hemisphere lesions (indifference vs catastrophic
reaction), as previously described by clinical investi-
gators. 32,4~ Alternatively, these perceptual, cognitive,
and expressive deficits may be independent of and not
J. Neurosurg. / Volume 64/May, 1986
The fight hemisphere: neuropsychological functions
contribute to the different patterns of emotional reac-
tivity after unilateral hemispheric lesions.
To learn more about mood, we36 administered the
Minnesota Multiphasic Personality Inventory (MMPI)
to patients with unilateral hemisphere lesions. The pa-
tients were matched for severity of cognitive (such as
intelligence quotient) and motor defects (such as motor
tapping). The MMPI has been widely used as an index
of underlying affective experiences, and the completion
of this inventory does not require the perception or
expression of affectively intoned speech or the percep-
tion of facial expression. Patients with left hemisphere
disease showed a marked increase on the depression
scale of this inventory, whereas patients with right
hemisphere disease did not. This finding suggests that
the differences in emotional reactions of patients after
right versus left hemisphere disease cannot be attributed
entirely to difficulties in perceiving or expressing affec-
tive stimuli. Equally important, the difference in "de-
pression" between the left and right hemisphere-dam-
aged groups appears unrelated to differences in the
severity of cognitive or motor defects.
Pathophysiology Underlying the Indifference
Response
Cannon 21 regarded the thalamus as an important
central structure responsible for mediating emotions. It
can be stimulated either by peripheral sensory input
or by cortical impulses. He proposed that thalamic
activity could excite both the cortex and the viscera.
Cortical activation induces the conscious emotional
state; the visceral changes occur simultaneously and
serve adaptive purposes. Bard 4 suggested that the hy-
pothalamus was, in fact, the major effector of emotional
expression, since it regulates both the endocrine system
and the autonomic nervous system. Papez, TM in turn,
proposed that the limbic system, which has important
connections with the hypothalamus, cortex, and thala-
mus, is important in regulating emotion.
The concept of cortical activation or arousal was
central to Cannon's theory? ~ Subsequent research led
to a better understanding of the physiology of arousal.
Berger ~l noted that the electroencephalographic pattern
of behavioral arousal is decreased amplitude and in-
creased frequency. This was termed "electroencepha-
lographic desynchronization." Desynchronization also
occurs during emotional states. 6~As we discussed, stim-
ulation of nonspecific thalamic nuclei or meseneephalic
reticular formation induces behavioral arousal and elec-
troencephalographic desynchronization. 68 In addition,
stimulation of certain cortical areas like the frontal or
temporoparietal regions activates the mesencephalic re-
ticular formation 31 and elicits an arousal response. 83
The limbic system, which has strong input into the
reticular formation, is another pathway by which cor-
tical stimulation can produce arousal. 49'96
The relationship between arousal and the "experi-
ence" of emotion has been studied in normal human
J. Neurosurg. / Volume 64/May, 1986
subjects. In 1924, Maranon (see Fehr and Stern 3~ in-
duced physiological arousal by administering sympath-
omimetic drugs to normal subjects. He stated that
most of his subjects reported feeling "no emotions,"
although many reported that they experienced "as if"
feelings. When Maranon induced an affective memory
that was not strong enough to produce an emotion in
the normal state, emotional reactions then occurred if
there was concomitant pharmacological arousal. Simi-
larly, Schachter 8~ aroused normal subjects pharmaco-
logically, placed them in a stressful situation, and found
both subjective and objective evidence of emotional
states. Pharmacological arousal alone did not produce
such emotional states. Likewise, the stressful situation
alone produced less emotion when the subjects were
not pharmacologically aroused. Maranon's and Schach-
ter's studies both support the hypothesis that, in order
to experience an emotion, one must have the appropri-
ate cognitive state plus a certain degree of arousal.
Arousal depends on the brain-stem reticular formation,
nonspecific thalamic nuclei, and certain changes me-
diated by the hypothalamus. The hypothalamus is
strongly influenced by the limbic system which, in turn,
has considerable input from the neocortex. An emotion
thus depends on varied anatomical structures, including
cortical systems for producing the appropriate cognitive
set, limbic structures for activating the brain stem and
thalamic activating centers as well as for controlling
hypothalamic output, the hypothalamus for regulating
endocrine and autonomic responses, and the brain stem
and thalamic activating systems for producing cortical
arousal.
Studies in our laboratory have suggested that patients
with fight hemisphere disease have difficulty in com-
prehending affectively intoned speech and affective fa-
cial expressions. Such deficits might therefore interfere
with the development of an appropriate cognitive set
(thought to be an essential component of emotion),
thereby resulting in emotional flattening in patients
with fight hemisphere disease. In addition, patients with
fight hemisphere disease and the indifference reaction
might be inadequately aroused. To determine whether
such patients had normal arousal, we48 stimulated the
normal side of patients with right or left hemisphere
disease with an electrical stimulus and simultaneously
recorded galvanic skin responses. The galvanic skin
response is a measure of peripheral sympathetic activity
which correlates well with other central measures of
arousal, such as the electroencephalogram. We found
that patients with fight hemisphere disease and the
indifference reaction had dramatically smaller arousal
responses (galvanic skin response) than aphasic patients
with left hemisphere disease or control patients without
hemispheric lesions.
Morrow, et al., 67 have presented neutral and emo-
tionally loaded stimuli to fight or left hemisphere-
damaged patients. These investigators found that the
patients with right hemisphere disease had reduced
701

galvanic skin responses to both neutral and emotional
stimuli. These findings provide further evidence that
the indifference reaction is associated with hypoarousal.
Patients with the indifference reaction resulting from
right hemisphere disease typically have unilateral
neglect. 27'32'49 As we have discussed, unilateral neglect
is an attention-arousal-activation defect caused by dys-
function in a corticolimbic-reticular loop. 45 Since ne-
glect occurs more often after fight hemisphere lesions,
we have also proposed and provided evidence for the
postulate that the right hemisphere may be dominant
for mediating an attention-arousal response. 48'5~
It is therefore possible that the fight hemisphere may
have a special role in mediating emotional behavior.
Not only can the fight hemisphere process affective
stimuli and program emotional behavior, but it also
appears to have a special relationship to those subcor-
tical structures important for mediating cerebral arousal
and activation.
Conclusions
In this review, we have presented evidence that the
right hemisphere has an important role in mediat-
ing visuospatial skills, attention, and emotions. Several
theories have been advanced as to anatomical and
physiological substrates underlying right versus left
hemisphere dominance. For example, functions like
attention and emotion rely in part on cortical-sub-
cortical connections, and it has been proposed that the
right hemisphere has more cortical-subcortical and cor-
ticolimbic projections than does the left hemisphere,
which has more corticocortical projections. 29 Whereas
the left hemisphere has been thought of as an analyti-
cal or serial processor, the right has been thought of as
a Gestalt (holistic) simultaneous and parallel processor.
Based on these psychological constructs, it has been
postulated that corticocortical connections of the two
hemispheres are different, the right being more diffusely
organized than the left.
Unfortunately, the anatomical and physiological
basis for dominance is not clearly understood. The
anatomical asymmetries demonstrated by Geschwind
and Levitsky 37 and Galaburda, et al., 34 are an exciting
beginning, but further studies are needed.
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Manuscript received July 5, 1985.
Accepted in final form November 4, 1985.
Address reprint requests to: Kenneth M. Heilman, M.D.,
Box J-236, Department of Neurology, J. Hillis Miller Health
Center, University of Florida, Gainesville, Florida 32610.
J. Neurosurg. / Volume 6 4 / M a y , 1986