Professional Documents
Culture Documents
Inferior Parietal Lobule Contributions To 1 PDF
Inferior Parietal Lobule Contributions To 1 PDF
Abstract
■ This study investigated how the left inferior parietal lobule slowed responses in the meaning, but not sound, task, whereas
(IPL) contributes to visual word recognition. We used repetitive stimulation of the left SMG affected responses in the sound,
TMS to temporarily disrupt neural information processing in two but not meaning, task. These results demonstrate that ANG and
anatomical fields of the IPL, namely, the angular (ANG) and supra- SMG doubly dissociate in their contributions to visual word rec-
marginal (SMG) gyri, and observed the effects on reading tasks ognition. We suggest that this functional division of labor may be
that focused attention on either the meaning or sounds of written understood in terms of the distinct patterns of cortico-cortical
words. Relative to no TMS, stimulation of the left ANG selectively connectivity resulting in separable functional circuits. ■
INTRODUCTION
or posterior (ANG) fields of the IPL. Similarly, Philipose
Reading is one of the most important skills humans need and colleagues (2007) found that reading deficits were
to learn to function in modern society, and understand- more commonly because of SMG, rather than ANG, lesions.
ing how this complex process is achieved by the brain is In contrast to Dejerineʼs hypothesis that the IPL stores
an important goal of cognitive neuroscience. On a neuro- visual word forms, others have claimed that this brain re-
logical level, reading involves a set of brain regions that gion performs the procedures necessary for converting
work together to recognize a visual input and link it to its spelling to sound during visual word recognition (Roux
corresponding sound and meaning. For example, extra- et al., 2012; Booth et al., 2004; Joubert et al., 2004; Pugh
striate visual areas are critical for recognizing the visual et al., 2000; Shaywitz et al., 1998; Law et al., 1991). For
form of a word, whereas higher-order association areas instance, Shaywitz and colleagues (1998) used fMRI to
located in the frontal, parietal, and temporal lobes are measure brain activity during a set of silent reading tasks
important for processing its sound and meaning (Price & and observed that normal, skilled readers robustly engaged
Mechelli, 2005; Shaywitz et al., 2002; Pugh et al., 2001). ANG when performing a nonword rhyming task (e.g., “Do
Here we specifically focused on the functional contribu- jete and leat rhyme?”). Unlike words, pronounceable non-
tions of the left inferior parietal lobule (IPL) to visual word words do not have a learned, associated phonology and
recognition. thus require the reader to assemble the phonological code
Joseph Jules Dejerine (1891) was the first to highlight from the orthography (Marshall & Newcombe, 1973). As a
the importance of the IPL in reading. He described the result, the authors associated ANG activation with spelling-
case of a 63-year-old sailor who became unable to read or to-sound conversion rather than with storage of visual
write because of a lesion of the left posterior IPL, more spe- word forms. Consistent with this claim, they found that
cifically the angular gyrus (ANG). Dejerine reasoned that developmental dyslexics, who were by definition poor at
the patientʼs inability to recognize visual words (alexia) phonological assembly, showed abnormally low ANG acti-
coupled with his writing difficulty (agraphia) indicated a vation (see also Pugh et al., 2000). Using a similar task but
central loss of visual word forms, which he argued were a different methodology, Roux et al. (2012) found that
stored in ANG. Subsequent studies of patients with ac- SMG—not ANG—was the critical site for spelling-to-sound
quired reading deficits have confirmed the importance of conversion. In this experiment, they used intracranial stim-
the IPL for reading but have introduced uncertainty regard- ulation in awake neurosurgical patients to temporarily dis-
ing the specific anatomical fields. For instance, Warrington rupt local processing in the stimulated region and measure
and Shallice (1980) reported two patients with profound its effect on behavior. Stimulation of the left anterior SMG
reading impairments subsequent to lesions predominantly preferentially interfered with reading nonwords (e.g.,
affecting either the anterior (supramarginal gyrus [SMG]) “dasul”) but did not affect real words, suggesting that the
SMG was necessary for spelling-to-sound conversion.
Therefore, although these studies agree that the IPL plays
University College London a critical role in spelling-to-sound conversion, there is no
© 2015 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 27:3, pp. 593–604
doi:10.1162/jocn_a_00721
consensus regarding the specific anatomical locus of this during SMG stimulation that interfered with making a
function with some focusing on ANG (Pugh et al., 2000; button press response and therefore could not participate
Horwitz, Rumsey, & Donohue, 1998; Shaywitz et al., in the experiment. In the other four, functional localization
1998), others on SMG (Roux et al., 2012; Jobard, Crivello, failed to identify an appropriate ANG (2) or SMG (2) testing
& Tzourio-Mazoyer, 2003; Law et al., 1991), and still others site. All of the remaining participants were right-handed,
arguing that both fields are important for this process monolingual native English speakers with normal or
(Booth et al., 2002, 2003, 2004; Joubert et al., 2004). corrected-to-normal vision. They reported having no neu-
A third hypothesis associates ANG and SMG with differ- rological conditions and no form of dyslexia. Each person
ent functional properties during visual word recognition. provided informed consent after the experimental proce-
By this account, ANG is involved when processing the dures were explained and was paid for their participation.
meaning of words whereas SMG contributes to processing The experiment was approved by the University College
their sound (Graves, Desai, Humphries, Seidenberg, & Binder, London research ethics committee.
2010; Price & Mechelli, 2005; Price, Moore, Humphreys, &
Wise, 1997; Demonet, Price, Wise, & Frackowiak, 1994).
Experimental Procedures
This hypothesis builds on the findings that ANG is consid-
ered a key node in the cortical semantic system (Binder, The experiment consisted of three separate testing ses-
Desai, Graves, & Conant, 2009) and that SMG is important sions for each participant. The first lasted approximately
for phonological processes associated with verbal working 30 min and involved acquisition of a T1-weighted MRI
memory (Buchsbaum & DʼEsposito, 2008). Several neuro- scan (FLASH sequence, repetition time = 12 msec, echo
imaging studies have confirmed a double dissociation time = 5.6 msec, flip angle = 19°, resolution = 1 mm ×
within IPL when processing the sound and meaning of 1 mm × 1 mm) at the Birkbeck–UCL Centre for Neuro-
written words (Vigneau et al., 2006; Devlin, Matthews, & imaging. The structural images were used for anatomical
Rushworth, 2003; Mummery, Patterson, Hodges, & Price, identification of left ANG and SMG in each participant.
1998; Price et al., 1997). It is unclear, however, whether Scanning was followed by two TMS sessions in which
the differential contribution of ANG and SMG to semantic either ANG or SMG was tested, with the order counter-
and phonological processing is necessary because of the balanced over participants. The TMS sessions were sepa-
nature of the neuroimaging techniques, which can only rated by at least 2 days and lasted approximately 1 hr
indicate correlations between brain and behavior, but do each. Each testing session consisted of a TMS-guided
not allow causal relations to be drawn. Moreover, in pa- functional localization and then the main experiment.
tients with IPL lesions, this double dissociation is not read- The aim of the localization procedure was to identify
ily apparent, in part because focal lesions selectively specific testing sites within ANG and SMG. In other
affecting either ANG or SMG are rare. words, the testing sites used in the main experiment
In summary, then, there are three hypotheses con- were determined using a TMS-based functional locali-
cerning IPL contributions to visual word recognition. zation procedure (Pattamadilok, Knierim, Duncan, &
The first claims that the IPL is the site of stored visual Devlin, 2010; Ellison, Lane, & Schenk, 2007; Taylor,
forms of written words although it remains unclear pre- Nobre, & Rushworth, 2007), similar to “functional locali-
cisely where within the IPL these are stored. The second zer” scans commonly used in fMRI experiments (Kraft
hypothesis argues that the procedures for converting et al., 2005; Kanwisher, McDermott, & Chun, 1997). The
spelling-to-sound are a function of the IPL, but it is un- aim of this functional localization procedure was to cus-
clear whether these are specifically located in ANG or tomize the stimulation site in each individual taking into
SMG, or both. Finally, a third hypothesis suggests that account intersubject functional–anatomical variability.
the angular and supramarginal fields of the IPL preferen- To identify appropriate testing sites, we chose localiza-
tially contribute to semantic and phonological processing tion tasks that optimized the constraints placed by the
of written words, respectively. The aim of the current three hypotheses under investigation. According to the
study was to evaluate these hypotheses using repetitive first hypothesis, left IPL stores visual word forms, and
TMS (rTMS) to temporarily and selectively disrupt pro- therefore, the only constraint was that the task used real
cessing in left ANG and SMG during visual word recogni- words (i.e., as opposed to pseudowords). The second
tion and measure the effect on reading behavior. hypothesis suggests that IPL is involved in orthography-
to-phonology conversion. In this case, localization re-
quired a task that involved letter-to-sound conversion, a
METHODS procedure that is thought to occur automatically in virtu-
ally all reading tasks (Coltheart, Rastle, Perry, Langdon, &
Participants Ziegler, 2001; Frost, 1998; Plaut, McClelland, Seidenberg,
Seventeen people volunteered for this study and 12 & Patterson, 1996). Finally, the third hypothesis claims
(seven women, five men; aged 18–42 years, mean = 26 years) that ANG and SMG are required in semantic and phono-
participated in the main experiment. One of the five logical processing of written words, respectively, and
excluded participants experienced right-hand twitching thus, localization required separate tasks that were either