Neuropsychologia 40 (2002) 1300–1312
Interhemispheric neural summation in the split brain
with symmetrical and asymmetrical displays
Matthew Roser
∗
, Michael C. Corballis
Department of Psychology, Research Centre for Cognitive Neuroscience, University of Auckland, Private Bag 92019, Auckland, New Zealand
Received 20 March 2001; received in revised form 6 November 2001; accepted 6 November 2001
Abstract
The present study, investigates interhemispheric integration in the split brain. Four split-brained, two acallosal and 14 normal subjects
carried out a simple reaction time task in which they responded to stimuli presented either singly in the left visual field, singly in the
right visual field, or simultaneously in both visual fields. Stimuli were white against a black background and bilateral stimuli were either
symmetrical or asymmetrical around the central vertical meridian. For unilateral stimuli, the difference in response time (RT) between
crossed and uncrossed hand-field combinations (crossed–uncrossed difference, or CUD) measured interhemispheric transfer time. RTs to
bilateral and unilateral stimulus displays were compared to provide a measure of redundancy gain (RG). Normal subjects exhibited small
CUDs and RGs. Split-brained and acallosal subjects were found to have much longer CUDs, and to show enhanced RGs which could not
be explained by a probability (race) model, implying subcortical neural summation. This summation did not depend on the preservation
of symmetry, suggesting that it may not occur at the retinotopically organized superior colliculus, but at another site such as the pons or
reticular formation. © 2002 Elsevier Science Ltd. All rights reserved.
Keywords: Redundancy gain; Commissurotomy; Callosotomy; Callosal agenesis
1. Introduction
A well established finding in studies of simple reaction
time is that response times (RTs) to stimuli such as lumi-
nance onsets are faster if there is more than one stimulus
presented. This is known as the redundancy gain (RG) and
can often be explained as the result of probability summa-
tion [25,35]. If the two stimuli trigger independent response
preparation processes and a response is made as soon as the
first process is completed, then on average, responses will
be made faster than when there is only one process in oper-
ation, as long as the two RT distributions overlap. As RTs
to redundant signals are determined by the first process to
be completed, this is known as the “race model”. If however
there is an enhancement of RT greater than that which can
be explained by probability summation, then coactivation
or neural summation is implied [7,36]. Coactivation mod-
els involve activation from redundant processes combining
towards reaching a single criterion for response initiation
[25,30]. Thus, activity on otherwise independent channels
is summed together to produce a speeded response [36].
∗
Corresponding author. Tel.: +64-9-373-7599; fax: +64-9-373-7450.
E-mail address: m.roser@auckland.ac.nz (M. Roser).
The race model seems sufficient to explain RG data in
most normal subjects [29,36], although there are excep-
tions [28]. However split-brained and acallosal subjects often
show RGs which exceed the predictions of the race model
by a far greater extent than the small violations occasion-
ally observed in normal subjects [7,18,33,36]. This is despite
the fact that redundant stimuli, displayed one in each visual
field, are presented separately to hemispheres which have
been disconnected at the cortical level by commissurotomy
or callosotomy, or by congenital failure of callosal fibers to
develop, as in callosal agenesis. Indeed bilateral presentation
seems to be a necessary condition for race-model violations
in at least one split-brained subject, since there was no viola-
tion when redundant stimuli were presented to a single hemi-
sphere and the split-brained subject showed similar mean RT
effects from redundant targets as normal subjects [36]. In ad-
dition to this, crossed responses, or those made by the hand
contralateral to the visual field in which the stimulus was dis-
played, are usually slowed relative to uncrossed responses.
In normals this crossed–uncrossed difference (CUD) is typ-
ically in the range of 2–6 ms [2,32] and has been used as an
index of callosal transmission times. In split-brained sub-
jects it is much longer, commonly around 70 ms, although
there is considerable variation [7,14,36]. The CUD is also
prolonged in acallosal subjects for whom CUDs of around
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