276 VOLUME 8 | NUMBER 3 | MARCH 2005 NATURE NEUROSCIENCE
BRIEF COMMUNICATIONS
Attending to local form while
ignoring global aspects depends
on handedness: evidence from
TMS
Carmel Mevorach
1
, Glyn W Humphreys
1
& Lilach Shalev
2
Our perceptions of the whole and of the parts of a visual
stimulus are mediated by different brain regions. We used low-
frequency transcranial magnetic stimulation (TMS) to show
for the first time that opposite, homologous regions in the two
hemispheres are involved in attending to local parts for left-
and right-handed individuals. The brain regions that focus on
the ‘trees’ while ignoring the ‘forest’ are switched as a function
of handedness.
The ability to attend to and identify global (the ‘forest’) or local (the
‘trees’) aspects of a hierarchical object
1
is lateralized between the two
hemispheres in the human brain; the left hemisphere is biased toward
the local level, whereas the right hemisphere is biased toward the global
level
2
. Hence, patients with unilateral damage may have difficulty in
identifying local or global information according to the site of lesion
3
.
However, this assignment is not absolute, as damage to the right
hemisphere can disrupt global
4
as well as local
5
identification.
No previous studies have assessed whether the brain regions that
focus on parts and wholes differ in left- and right-handers, though
handedness is known to influence language lateralization
6
and some
aspects of spatial perception
7
. Here we show differential lateralization
of function for attention to local aspects of form in homologous areas
in the right and left cerebral hemispheres.
Transcranial magnetic stimulation (TMS) is a technique in which
transient disruption of normal brain activity is induced by the
application of focal magnetic pulses to specific regions on the scalp
8
. We
used TMS to assess the role of the left and right posterior parietal lobes
(PPL; electrode sites P3 and P4) for right- (n = 11, mean age = 29.18)
and left-handed participants (n = 11, mean age = 25.36; handedness was
assessed using the Edinburgh Handedness Inventory) when they were
required to identify global and local forms in two stimulus categories
(letters and shapes) under conditions of focal attention (see Fig. 1).
Statistical analysis showed contrasting effects of TMS for right-
and left-handed participants specifically when identifying local
congruent displays as compared with incongruent displays (the global
interference effect; see Fig. 2). This effect did not interact with stimulus
category (letters or shapes). For right-handers, TMS over the left PPL
increased global interference as compared with TMS over the right
PPL (Fig. 2a,c). For left-handers, TMS over the right PPL resulted
in larger global-to-local interference than did TMS over the left PPL
(Fig. 2b,c). There were no effects on the global identification task.
Notably, TMS affected performance primarily when attentional con-
trol was required in order to ignore an irrelevant level (on incongruent
relative to congruent trials).
These findings fit with previous studies pointing to an attentional
involvement in global and local identification and in the apparent lat-
eralization of these processes in right-handed individuals
4,9,10
. More
than this, though, our data show that opposite, homologous regions in
the left and right hemispheres control attention to local form in left-
and right-handed individuals. In right-handers, the left PPL controls
1
Behavioural Brain Sciences Centre, School of Psychology, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK.
2
Department of Education and
Psychology, The Open University Israel, 108 Ravutski St., Ra'anana, Israel. Correspondence should be addressed to C.M. (c.mevorach@bham.ac.uk).
Published online 6 February 2005; doi:10.1038/nn1400
Figure 1 Typical display sequence showing congruent and incongruent
compound stimuli. A red compound figure was presented either to the
left, center or right of fixation and was composed of either ‘H’ and ‘D’
(letter condition) or ‘crosses’ and ‘boxes’ (shape condition). On different
blocks of trials, participants were asked to identify the global or the
local elements while ignoring information on the other level. In half of
the trials, the compound figures consisted of the same global and local
elements (congruent trials), and in the other half, of different global and
local elements (incongruent trials). For TMS, a 70-mm figure-eight coil
connected to a MagStim Rapid stimulator (MagStim) was positioned
over the left or right posterior parietal lobe (P3 or P4 on the 10–20 EEG
coordinate system). TMS trains consisted of 600 1-Hz pulses with intensity
set to 90% of the participants’ motor threshold (this can produce prolonged
inhibition of the stimulated cortical site
11
including P3 and P4; ref.
12). Each block consisted of 96 trials and was given six times to each
participant (three blocks per category: letters or shapes). In a first session,
a pre-TMS run was followed by one of the two TMS trains, which was
immediately followed by another run of the task; in a second session there
was only the TMS train followed by the task. The order of both the stimulus
category and the TMS site was counterbalanced across participants. As
participants initially performed the pre-TMS condition, differences between
pre- and post-TMS could be masked by practice. Consequently our focus
was on the two post-TMS conditions. The experiment was approved by the
Ethics Committee of the School of Psychology, Birmingham University.
Written informed consent was obtained from all participants.
© 2005 Nature Publishing Group http://www.nature.com/natureneuroscience