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J Cogn Neurosci. 2004 September ; 16(7): 1234–1249.

Development of Brain Mechanisms for Processing Orthographic

and Phonologic Representations

James R. Booth1,2,3, Douglas D. Burman1, Joel R. Meyer2, Darren R. Gitelman3, Todd B.

Parrish3, and M. Marsel Mesulam3
1 Northwestern University,
2 Evanston Northwestern Healthcare,
3 Northwestern University Medical School

Abstract
Developmental differences in the neurocognitive networks for lexical processing were examined in
15 adults and 15 children (9- to 12-year-olds) using functional magnetic resonance imaging (fMRI).
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The lexical tasks involved spelling and rhyming judgments in either the visual or auditory modality.
These lexical tasks were compared with nonlinguistic control tasks involving judgments of line
patterns or tone sequences. The first main finding was that adults showed greater activation than
children during the cross-modal lexical tasks in a region proposed to be involved in mapping between
orthographic and phonologic representations. The visual rhyming task, which required conversion
from orthography to phonology, produced greater activation for adults in the angular gyrus. The
auditory spelling task, which required the conversion from phonology to orthography, also produced
greater activation for adults in the angular gyrus. The greater activation for adults suggests they may
have a more elaborated posterior heteromodal system for mapping between representational systems.
The second main finding was that adults showed greater activation than children during the intra-
modal lexical tasks in the angular gyrus. The visual spelling and auditory rhyming did not require
conversion between orthography and phonology for correct performance but the adults showed
greater activation in a system implicated for this mapping. The greater activation for adults suggests
that they have more interactive convergence between representational systems during lexical
processing.

INTRODUCTION
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In a series of studies, Booth et al. (2002a, 2002b) have developed a neurocognitive model of
lexical processing using functional magnetic resonance imaging (fMRI). This model suggests
that auditory word forms (phonologic representations) involve the superior temporal gyrus,
visual word forms (orthographic representations) involve the fusiform gyrus, and meaning
forms (amodal semantic representations) involve the middle temporal gyrus (Booth, Burman,
Meyer, Zhang, et al., 2003; Booth et al., 2001, 2002a, 2002b). Interactions among these
representations are mediated by posterior heteromodal regions including the supramarginal
and angular gyrus. In cross-modal tasks that require the conversion of orthography to
phonology (rhyming judgments to visually presented words), for example, better performance
in adults is associated with greater supramarginal/angular gyrus and superior temporal gyrus
activations (Booth, Burman, Meyer, Gitelman, et al., 2003). Similarly, in cross-modal tasks

Reprint requests should be sent to James R. Booth, Department of Communication Sciences and Disorders, Northwestern University,
2240 Campus Drive, Evanston, Illinois, 60208, USA, or via e-mail:[email protected]..
The data reported in this experiment have been deposited in the fMRI Data Center (http://www.fmridc.org). The accession number is
2-2004-1165T.
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that require the conversion of phonology to orthography (spelling judgments to auditorily

presented words), better performance is associated with greater supramarginal/angular gyrus
and fusiform gyrus activations (Booth, Burman, Meyer, Gitelman, et al., 2003). However,
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developmental differences in the neurocognitive networks involved in cross-modal tasks have

not been investigated. Because children are less skilled at lexical processing than adults, we
expected that children may show less activation in the supramarginal/angular gyrus for
mapping between orthographic and phonologic representations during cross-modal tasks. This
greater activation in this posterior heteromodal system may reflect greater elaboration of this
system to represent different levels of mapping between orthographic and phonology. Indeed,
several behavioral studies have argued for skilled lexical processing involving a mechanism
that includes grapheme–phoneme, onset–rime, syllabic and word level mappings for both
reading (Ehri, 1995; Marsh, Friedman, Welch, & Desberg, 1981; Gough & Hillinger, 1980)
and spelling (Nunes, Bryant, & Bindman, 1997).

Skilled lexical processing may also be characterized by enhanced interactivity between the
orthographic and phonologic representational systems. VanOrden, Pennington, and Stone
(1990) and VanOrden and Goldinger (1994) have argued that efficient processing is
characterized by resonance between representational systems (VanOrden & Goldinger, 1994;
VanOrden et al., 1990). Resonance between systems occurs when input to the orthographic
system closely matches the information that is fed back from the phonologic system or vice
versa. Several lines of research suggest that skilled readers activate phonologic information
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when reading earlier and more automatically than less skilled readers (Booth, Perfetti,
MacWhinney, & Hunt, 2000; Plaut & Booth, 2000; Booth, Perfetti, & MacWhinney, 1999).
There is also a 25-year history of behavioral research with adults illustrating the influence of
orthographic information on the speed of spoken word recognition (Ziegler & Ferrand, 1998;
Dijkstra, Roelofs, & Fieuws, 1995; Jakimik, Cole, & Rudnicky, 1985; Donnenwerth-Nolan,
Tanenhaus, & Seidenberg, 1981; Seidenberg & Tanenhaus, 1979). Research shows that the
influence of orthography on auditory language tasks is inconsistent in the early elementary
grades and becomes reliable in later elementary grades (Zecker, 1991; Perin, 1983; Tunmer &
Nesdale, 1982; Ehri & Wilce, 1980). Research on dyslexics has also found evidence for a
reduced role of orthographic knowledge on auditory processing in children with reading
disabilities compared to control children (Landerl, Frith, & Wimmer, 1996; Zecker, 1991). All
of this research suggests that there may be more interactivity between orthographic and
phonologic representations in skilled readers. Because adults are more skilled than children,
we expected that adults may show more activation in the supramarginal/angular gyrus even
during intra-modal tasks that do not require conversion between orthographic and phonologic
representations for correct performance.

Our study examined developmental differences between 15 adults and 15 children (9- to 12-
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year-olds) in the functional neuroanatomy for lexical processing using visual or auditory lexical
tasks that required spelling and rhyming judgments (see Table 1). These visual word judgment
tasks were compared to a control condition involving line patterns and the auditory word
judgment tasks were compared to a control condition involving pure tones (see Table 2). All
tasks involved the sequential presentation of three stimuli and required the participants to
determine whether the third stimuli matched one of the previous two stimuli based on a
predefined criterion. The goal of this study was to examine whether adults would show greater
activation in the supramarginal/angular gyrus during cross-modal and intra-modal tasks.
Greater activation in this region during cross-modal tasks would indicate a more elaborate
system involved in mapping between orthographic and phonologic representations and greater
activation in this region for the intra-modal tasks would indicate greater interactivity between
the orthographic and phonologic systems.

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RESULTS
Behavioral Performance
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Means (and standard errors) for accuracy and reaction time on the lexical and control tasks are
presented in Table 3. Only data for the fMRI session are presented because statistical analyses
revealed no main effects or interactions involving accuracy or reaction time on the practice
versus fMRI sessions. This indicates that the environment of the scanner did not adversely
affect the behavioral performance of the adults or children. Table 3 shows that both the adults
and children could effectively perform the tasks in the fMRI scanner as all mean accuracy
levels were above 80%.

We calculated age (adults, children) by condition (word, control) ANOVAs on accuracy and
reaction time separately for each of the lexical tasks in each of the modalities. Adults were
more accurate than children on all tasks [visual spelling, F(1,63) = 21.20, p < .001; visual
rhyming, F(1,63) = 17.76, p < .001; auditory spelling, F(1,59) = 12.89, p< .01; auditory
rhyming, F(1,63) = 9.27, p < .01]. Adults were also faster than children on all tasks [visual
spelling, F(1,63) = 78.33, p < .001; visual rhyming, F(l,63) = 69.98, p < .001; auditory spelling,
F(1,59) = 45.11, p < .01; auditory rhyming, F(1,63) = 65.71, p < .01]. There was a main effect
of condition for accuracy on auditory spelling [F(1,59) = 10.11, p < .01], revealing that word
judgment was less accurate than tone judgment for this task. There were main effects of
condition for reaction time on all tasks but auditory rhyming, revealing that the word judgment
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was slower than control judgments [visual spelling, F(1,63) = 15.88, p < .001; visual rhyming,
F(1,63) = 10.83, p < .001; auditory spelling, F(1,59) = 20.16, p < .01].

There were no interactions between age and condition for the visual and auditory rhyming tasks
for either accuracy or reaction time or for visual and auditory spelling for reaction time. The
lack of interactions suggests that any reported fMRI differences may not be attributable to
performance differences on these tasks because the exact same analyses (age by condition)
were done on the behavioral and fMRI data. However, the main effects for accuracy were
qualified by interactions between age and condition for visual spelling [F(l,63) = 4.09, p < .
05] and auditory spelling [F(l,59) = 4.94, p < .05], indicating that children had especially low
accuracy on word judgment for the spelling tasks. This interaction means that part of the
developmental effects in the fMRI data may be due to performance differences. In
developmental work, it is extremely hard to equate different ages in behavioral performance
especially on tasks that tap into explicit judgments of meaningful linguistic contrasts.

Activation for Each Lexical Task

Figure 1 presents significant activation separately for the adults (red) and children (green) in
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the spelling and rhyming tasks for the visual and auditory modality as compared to the
nonlinguistic control tasks (lines or tones). Figure 1 also shows overlap (purple) between the
activation maps for the adults and children. Figure 2 presents significantly greater activation
for the adults than for children (red) or significantly greater activation for children than for
adults (green). The numerical data for Figures 1 and 2 are presented in Tables 4–7. We
concentrate our reporting of the results to our regions of interest that included visual association
regions (fusiform gyrus, BA 37), auditory association regions (superior temporal gyrus, BA
22), posterior heteromodal regions (angular gyrus, BA 39, and middle temporal gyrus, BA 21),
and inferior frontal gyrus (BA 9, 44, 45, 47). All areas of significant activation outside of these
regions of interest are presented in the tables and figures.

Visual Spelling Task—Both adults and children showed more activation in the left than
right inferior frontal gyrus. However, adults showed significantly greater activation than
children in the left (68 voxels) and right (51 voxels) inferior frontal gyrus. Both adults and

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children showed activation in the left angular gyrus, but only adults showed activation in the
right angular gyrus. Adults showed significantly greater activation in the left (89 voxels) and
right angular gyrus (21 voxels). Both adults and children showed more activation in the left
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than in the right fusiform gyrus and there was no significant developmental difference in this
region. Only children showed activation in the left middle temporal gyrus.

Visual Rhyming Task—Both adults and children showed activation in the left inferior
frontal gyrus, but only adults showed activation in the right inferior frontal gyrus. Adults
showed significantly greater activation than children in the bilateral inferior frontal gyrus (43
voxels for left and 20 voxels for right). Both adults and children showed more activation in the
left than in the right fusiform gyrus and there was no significant developmental difference in
this region. Both adults and children also showed activation in the left middle temporal gyrus,
but only adults showed activation in the left angular gyrus and this activation was significantly
greater than for children (34 voxels). Neither the adults nor children showed activation in the
left superior temporal gyrus at the p < .001 level; however, both the children and the adults
showed activation in the left superior temporal gyrus at the p < .01 level.

Auditory Spelling Task—Both adults and children showed activation in the left inferior
frontal gyrus, but only adults showed activation in the right inferior frontal gyrus. Adults
showed significantly greater activation than children in the bilateral inferior frontal gyrus (110
voxels for left and 189 voxels for right). Only adults showed activation in the bilateral angular
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gyrus and they showed significantly greater activation than children in the left angular gyrus
(35 voxels). Both adults and children showed more activation in the left than in the right
superior/middle temporal gyrus, but adults showed significantly greater activation in the left
superior temporal gyrus (24 voxels) and right middle temporal gyrus (30 voxels). Both adults
and children showed activation in the left fusiform gyrus, but children showed significantly
greater activation than adults in this region (74 voxels).

Auditory Rhyming Task—Both adults and children showed activation in the left inferior
frontal gyrus, but adults showed significantly greater activation in this region (213 voxels).
Both adults and children showed more activation in the left than in the right superior/middle
temporal gyrus, but adults showed significantly greater activation in the left superior temporal
gyrus (23 voxels). Both adults and children showed activation in the left fusiform gyrus and
there was no significant developmental difference in this region.

Additional Activation Clusters—The medial frontal gyrus/anterior cingulate was

activated for both adults and children for the spelling tasks in both modalities. Only the adults
showed activation in the medial frontal gyrus during the visual rhyming task and neither group
showed activation in these regions during the auditory rhyming task. In addition, the adults
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showed activation in the cerebellum for all tasks, but the children only showed activation in
the cerebellum in the auditory spelling task.

DISCUSSION
The purpose of this study was to examine developmental differences between adults and 9- to
12-year-old children in the neural substrate for lexical processing using word judgment tasks
that required explicit manipulation of orthographic (spelling tasks) and phonologic (rhyming
tasks) representations. The left inferior frontal gyrus was activated during all of these tasks for
both the adults and children. However, the adults showed significantly greater activation in the
left inferior frontal gyrus on the spelling and rhyming tasks in both modalities. This is consistent
with previous research that shows developmental increases in the magnitude of activation in
the left inferior frontal gyrus during covert verb generation tasks (Holland et al., 2001), overt
word generation tasks (Schlaggar et al., 2002), covert verbal fluency tasks (Gaillard et al.,

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2003), category judgment tasks (Shaywitz et al., 2002), and implicit reading tasks (Turkeltaub,
Garaeu, Flowers, Zefirro, & Eden, 2003). The right inferior frontal gyrus was activated in the
adults for all tasks but the auditory rhyming task. The children only activated the right inferior
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frontal gyrus during the visual spelling task. In a direct statistical comparison, the adults showed
significantly greater activation than the children in the right inferior frontal gyrus for the visual
spelling, visual rhyming, and auditory spelling tasks. The developmental finding for the
spelling task is consistent with a previous study that found greater right inferior frontal gyrus
activation in adults than in children (Booth, Burman, Meyer, Zhang, et al., 2003). The finding
of bilateral developmental differences during spelling is also consistent with adult research
that shows both right and left hemisphere involvement during spelling tasks (Flowers, Wood,
& Naylor, 1991).

Several lines of research suggest that the left fusiform gyrus is important for processing
orthographic information (Fujimaki et al., 1999; Herbster, Mintun, Nebes, & Becker, 1997;
Nobre, Allison, & McCarthy, 1994; Petersen, Fox, Snyder, & Raichle, 1990) and that the
superior temporal gyrus is important for processing phonologic information (Giraud & Price,
2001; Binder et al., 1994; Howard et al., 1992). Consistent with this, both adults and children
in our study showed greater activation in the left than in the right fusiform gyrus for the visual
tasks and showed greater activation in the left than in the right superior temporal gyrus for the
auditory tasks. Our study included cross-modal tasks that required the mapping between
orthographic and phonologic representations. Both adults and children showed activation in
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the left fusiform gyrus during the auditory spelling task that required access to orthographic
representations and both also showed activation in the left superior temporal gyrus during the
visual rhyming task that required the access to phonologic representations. However, the adults
showed significantly greater activation in the angular gyrus for both of the cross-modal tasks,
suggesting that they may have a more elaborated system that allows for more efficient mapping
between representations (Booth et al., 1999, 2000). The developmental differences in our study
are similar to a previous study that has reported a positive correlation between skill and brain
activation during cross-modal tasks within a group of adults (Booth, Burman, Meyer, Gitelman,
et al., 2003). These differences are also consistent with behavioral studies which suggest that
acquisition is marked by the elaboration of a single mechanism for mapping between
orthography and phonology that includes grapheme–phoneme, onset–rime, syllabic, and word
level mappings (Nunes et al., 1997; Ehri, 1995; Marsh et al., 1981; Gough & Hillinger,
1980).

Some models of skilled reading suggest that there are two separate mechanisms for mapping
between orthography and phonology—one system directly maps between whole word
representations and the other involves a grapheme–phoneme correspondence rule system
(Coltheart, Rastle, Perry, Langdon, & Ziegler, 2001; Coltheart, Curtis, Atkins, & Haller,
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1993). Some developmental models argue that reading and spelling acquisition is marked by
a shift from reliance on the grapheme–phoneme (alphabetical) system to the direct mapping
between whole word representations (Frith, 1985). Our neurocognitive model of lexical
processing argues that the supramarginal/angular gyrus is involved in extracting statistical
regularities (or probabilistic rules) between orthography and phonology, so dual mechanism
models predict a developmental decrease of activation in this region due to an increasing
reliance on direct mapping between whole word representations. Because we show a
developmental increase of activation in the angular gyrus, this suggests that acquisition may
be marked by the greater role of a system for abstracting regularities between orthography and
phonology. Therefore, our results seem to be more consistent with models of reading which
argue for one mechanism that maps between orthography and phonology (Harm & Seidenberg,
1999; Plaut, McClelland, Seidenberg, & Patterson, 1996; Seidenberg & McClelland, 1989).

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There was also evidence for access of orthographic information during auditory processing as
both adults and children also showed activation in the left fusiform for the auditory rhyming
task. This is consistent with previous literature which has shown the influence of orthographic
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information on spoken word recognition (Ziegler & Ferrand, 1998; Dijkstra et al., 1995;
Jakimik et al., 1985; Donnenwerth-Nolan et al., 1981; Seidenberg & Tanenhaus, 1979). We
also found that the adults showed greater activation than children in the angular gyrus/superior
parietal lobule during the auditory rhyming task, suggesting that the adults showed greater
interactivity between orthography and phonology during spoken word processing. This is
consistent with research which shows that the influence of orthography on auditory language
tasks is weak in the early elementary grades and becomes reliable in later elementary grades
(Zecker, 1991; Perin, 1983; Tunmer & Nesdale, 1982; Ehri & Wilce, 1980).

Skilled lexical processing seems to involve increasing resonance between the orthographic and
phonologic systems (VanOrden & Goldinger, 1994; VanOrden et al., 1990). This would
explain the greater activation for the adults in the angular gyrus during the auditory rhyming
task (as discussed above) and also during the visual rhyming task. Previous research clearly
shows that older and more skilled readers have faster and more automatic access of phonologic
information when reading (Booth et al., 1999, 2000). Although both adults and children showed
reliable activation in the angular gyrus during the visual spelling task, the greater activation
for the adults in this region may be related to their more automatic access of phonology. Further
research should confirm this finding by manipulating the orthographic and phonologic
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consistency between word pairs. Research clearly shows that orthographically (e.g., grade,
laid) and phonologically (e.g., pint, mint) inconsistent pairs are more difficult during spelling
and rhyming judgments (McPherson, Ackerman, & Dykman, 1997; Levinthal & Hornung,
1992; Kramer & Donchin, 1987; Rugg & Barrett, 1987; Johnston & McDermott, 1986; Rack,
1985; Polich, McCarthy, Wang, & Donchin, 1983; Seidenberg & Tanenhaus, 1979). If there
is more automatic access of orthography for the adults during auditory rhyming tasks then one
may expect greater interference (more activation in the angular and fusiform gyrus) for the
orthographically inconsistent pairs. Similarly, if there is more automatic access of phonology
for the adults during visual spelling tasks, then one may expect greater interference (more
activation in the angular and superior temporal gyrus) for the phonologically inconsistent pairs.
We are currently investigating these orthographically and phonologically inconsistent word
pairs during intra-modal and cross-modal tasks using an event-related fMRI design.

The greater activation in the angular gyrus during the intra-modal and cross-modal tasks is
likely not due to developmental differences in the involvement of semantics. Research shows
that older and more skilled readers rely less on semantics than younger and less skilled readers
during rapid word recognition (Schwantes, 1981, 1985; Briggs, Austin, & Underwood, 1984;
Simpson & Lorsbach, 1983; Simpson, Lorsbach, & Whitehouse, 1983; West, Stanovich,
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Freeman, & Cunningham, 1983; Stanovich, West, & Freeman, 1981; West & Stanovich,
1978). Studies have generally shown that the middle temporal gyrus is involved in a variety
of semantic tasks including action, abstract or concrete, living or nonliving, and category
judgments (Phillips, Noppeney, Humphreys, & Price, 2002; Devlin et al., 2002; Friederici,
Opitz, & von Cramon, 2000; Price, Moore, Humphreys, & Wise, 1997; Pugh et al., 1996).
Another way to examine the neural representation of semantics is to compare activation
patterns between words and pseudowords, because the latter do not have meaning
representations. Most studies have found that the left inferior frontal gyrus, the left posterior
superior temporal gyrus, the left inferior parietal lobule, and the left fusiform gyrus show
greater activation for pseudowords than for words (Mechelli, Gorno-Tempini, & Price, 2003;
Fiebach, Friederici, Mueller, & von Cramon, 2002; Simos et al., 2002; Xu et al., 2001;
Mechelli, 2000; Fiez, Balota, Raichle, & Petersen, 1999; Hagoort et al., 1999; Herbster et al.,
1997). The only brain region that seems to consistently show greater activation for words than
for pseudowords is the left posterior middle temporal gyrus (Fiebach et al., 2002; Simos et al.,

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2002; Hagoort et al., 1999). Taken together, these behavioral and neuroimaging results seem
to predict a developmental decrease in the involvement of the left middle temporal gyrus during
our spelling and rhyming tasks. In contrast, we found developmental increases in activation in
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the angular gyrus, so this region is probably not associated with age-related differences in
semantic involvement but rather with extracting statistical regularities between orthography
and phonology. Although some studies have suggested that the angular or supramarginal gyrus
are involved in semantic processing (Devlin, Matthews, & Rushworth, 2003; Rossell, Price,
& Nobre, 2003), if the developmental differences reported for the angular gyrus in our study
were due to semantic effects, we would have expected to find developmental decreases in
activation for this region. Further research will have to be done to confirm our prediction that
development should be marked by developmental decreases in the involvement of the middle
temporal gyrus in nonsemantic lexical processing.

The central finding of this study is an age-related increase in activation in the angular gyrus
during both cross-modal and intra-modal tasks. Previous reports on adults have shown greater
activation in the angular gyrus during cross-modal than intra-modal lexical tasks suggesting
that cross-modal tasks require mapping between orthography and phonology, whereas intra-
modal tasks do not (Booth et al., 2002a). These previous results, together with the present
results, show that there is more robust activation in the angular gyrus during cross-modal tasks
than intra-modal tasks, but that reading skill may be associated with a more elaborated system
for mapping and increasing interactivity between orthographic and phonologic representations.
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METHODS
Participants
Sixteen adults (M age = 25.2 years, range = 20.7–35.7 years) and 16 children (M age = 10.7
years, range = 9.4–11.9 years) participated in this study. There were 6 men and 10 women in
the adult group and there were 8 boys and 8 girls in the child group. One child was removed
from the auditory spelling task due to low accuracy performance. One adult had missing
behavioral data on the auditory spelling task due to computer malfunction. All adults were
undergraduate or graduate students at Northwestern University. All children were recruited
from private and public schools in the Evanston, Illinois, area.

All participants were given an interview to ensure that they did not have a history of
intelligence, reading, or oral-language deficits. All participants were native English speakers
and had normal hearing and normal or corrected-to-normal vision. All participants were free
of neurological diseases or psychiatric disorders and were not taking medication affecting the
central nervous system. The Institutional Review Board at Northwestern University and
Evanston Northwestern Healthcare Research Institute approved the informed consent
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procedures.

Functional Activation Tasks

Word Judgment Tasks—In the word judgment tasks, three words were presented
sequentially and the participant had to determine whether the final word matched either of the
two previous words according to a predefined rule. In the spelling task, participants determined
whether the final word had the same “rime” spelling as either of the first two words. The rime
included all letters after the first consonant or consonant cluster (Bowey, 1990). In the rhyming
task, participants determined whether the final word “rhymed” with either of the first two
words. Table 1 presents the stimuli for the spelling and rhyming tasks.

For both the spelling and rhyming tasks, half of the target trials contained a target word that
rhymed and was orthographically similar to one of the two preceding words (i.e., had the same

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rime). The other half contained a target word that rhymed but was orthographically dissimilar
to one of the two preceding words. In addition, half of the correct trials involved a match to
the first stimulus (first match) and half involved a match to the second stimulus (second match).
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The unrelated trials (40%) involved three orthographically different words that were
nonrhyming and semantically unrelated. If there was a match according to the criterion, the
participant pressed a button with the index finger; if there was no match, the participant pressed
a different button with the middle finger.

Stimulus Characteristics—All tasks were structured in a similar way so patterns of brain

activation could be directly compared across tasks (Crosson et al., 1999). First, the tasks
consisted of words with similar written word frequency for children and adults (The Educator’s
Word Frequency Guide, 1996) and similar adult word frequency for written and spoken
language (Baayen, Piepenbrock, & Gulikers, 1995). Second, no homophones were included in
the experimental lists. Third, the tasks contained about the same number of nouns (55–65%),
verbs (25–35%), and adjectives (10–20%) based on their most frequent usage in the Oxford
English Dictionary.

Visual Word Judgment Tasks—Each word reading task lasted 9 min consisting of 10
blocks of 54 sec. This included a 4-sec introduction screen to each block: “Spelling” for
orthographic task and “Rhyming” for phonologic task. The five experimental blocks alternated
with the five control blocks. In each trial for the experimental blocks, three consecutive words
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were presented in lowercase letters with each word presented for 800 msec followed by a 200-
msec blank interval. A yellow fixation cross (+) appeared on the screen after the third stimulus
was removed, indicating the need to make a response during the subsequent 2000-msec
interval. Participants were told that they could respond before the yellow cross (+) appeared
on the screen. Participants were encouraged to respond as quickly as possible without making
errors. Each trial lasted a total of 5000 msec and there were 10 trials in each block.

Auditory Word Judgment Tasks—The timing for the auditory tasks was the same as for
the visual tasks, but the auditory tasks employed a different list of stimuli. All stimuli for this
task were recorded in a soundproof booth using a digital recorder and a high-quality stereo
microphone. A native Chicagoan woman spoke each word in isolation so that there would be
no contextual effects. All words longer than 800 msec were shortened to this duration (less
than 1% of the words). All words were then normalized so that they were of equal amplitude.
The stimuli were easily heard through the headphones in the 1.5 Tesla scanner.

During the auditory tasks, a white fixation cross (+) was presented during the presentation of
the auditory stimuli. As in the visual word tasks, a yellow fixation cross (+) appeared on the
screen after the third stimulus was presented, indicating the need to make a response.
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Participants were asked to fixate on the cross during the entire trial.

Control Conditions—The control blocks for the visual and auditory tasks were designed to
equate the experimental and control blocks in terms of their memory demands and response
characteristics. The experimental setup and timing for the control blocks was exactly the same
as for the word blocks. For control blocks in the visual tasks, the three stimuli were abstract,
“nonlinguistic” symbols consisting of straight lines (see Table 2). Participants determined
whether the third stimulus was the same as one of the first two stimuli. Half of the correct trials
involved a match to the first stimulus (first match) and half involved a match to the second
stimulus (second match). The nonmatching trials involved three different stimuli. As with the
experimental blocks, 60% of the trials involved a match and 40% involved a nonmatch. For
control blocks in the auditory tasks, the three stimuli were high (700 Hz), medium (500 Hz),
and low frequency (300 Hz) “nonlinguistic” pure tones (see Table 2). The tones were 600 msec

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in duration and contained a 100-msec linear fade in and a 100-msec linear fade out. Otherwise,
the auditory control task was structured exactly like the visual control.
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Experimental Procedure
After informed consent was obtained, participants were administered the informal interview
(see above) and the first practice session in a simulator in order to acclimate the participant to
the scanner environment (Rosenberg et al., 1997). The participant practiced a full-length
version of each experimental task in the simulator. Different stimuli (matched in their stimulus
characteristics) were used in the practice and fMRI sessions. Within 3 days, the participant was
administered the first MRI session. Within 2 months, the participant was administered the
second practice and MRI session. The auditory and visual tasks were run on separate days with
an approximately equal number of participants receiving the visual and auditory modality first.

MRI Data Acquisition—After screening, the participant was asked to lie down on the
scanner bed. The head position was secured with a specially designed vacuum pillow (Bionix,
Toledo, OH) that allowed for the insertion of two earphones (for the auditory sessions). An
optical response box (Lightwave Medical, Burnaby, Canada) was placed in the participant’s
right hand and a compression alarm ball was placed in the left hand. The head coil was
positioned over the participant’s head and a goggle system for the visual presentation of stimuli
(Avotec, Jensen Beach, FL) was secured to the head coil. Each imaging session took less than
one hour.
NIH-PA Author Manuscript

All images were acquired using a 1.5 Tesla GE scanner. Gradient-echo localizer images were
acquired to determine the placement of the functional slices. For the functional imaging studies,
a susceptibility weighted single-shot EPI (echo-planar imaging) method with BOLD (blood
oxygenation level dependent) was used. The following scan parameters were used: TE = 40
msec, flip angle = 90°, matrix size = 64 × 64, field of view = 22 cm, slice thickness = 4 mm,
number of slices = 32. These scanning parameters resulted in a 3.437 × 3.437 × 4 mm voxel
size. The acquisition of this volume was repeated every 3 sec (TR = 3000 msec) for a total of
9 min per run.

At the end of the functional imaging session, a high-resolution, T1-weighted 3-D image was
acquired (SPGR, TR = 21 msec, TE = 8 msec, flip angle = 20°, matrix size = 256 × 256, field
of view = 22 cm, slice thickness = 1 mm). These scanning parameters resulted in a 0.86 × 0.86
× 1 mm voxel size. The orientation of this 3-D volume was identical to the functional slices.

Image Analysis—Analysis was performed using SPM-99 (Statistical Parametric Mapping)

for motion correction and statistical inference (Friston, Ashburner, et al., 1995; Friston,
Holmes, et al., 1995; Friston, Jezzard, & Turner, 1994). Application Visualization System
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(AVS) software with customized modules was used for visualization.

The functional images were realigned to the last functional volume in the scanning session
using affine transformations. All statistical analyses were conducted on these movement-
corrected images. No individual runs had more than 3 mm of movement. We calculated 2 Age
(children, adults) × 2 Mode (visual, auditory) × 2 Task (spelling, rhyming) ANOVAs separately
on the x-plane, y-plane, and z-plane motion estimates. This analyses revealed no significant
main effects or interactions involving age indicating that movement was not reliably different
between the adults and children for the x-plane (M = 0.18; range = 0.02 to 0.70 vs. M = 0.21;
range = 0.02 to 1.60), y-plane (M = 0.31; range = 0.06 to 1.66 vs. M= 0.30; range = 0.05 to
1.41), or z-plane (M = 0.54; range = 0.10 to 2.48 vs. M = .66; range = 0.12 to 2.64).

Images were then segmented and the gray–white matter information was used to coregister the
structural and functional images. The coregistered images were normalized to the MNI

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 Booth et al. Page 10

stereotaxic template (12 linear affine parameters for brain size and position, 8 nonlinear
iterations and 2 × 2 × 2 nonlinear basis functions for subtle morphological differences). The
MNI template used for normalization by SPM-99 is similar to the Talairach and Tournoux
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(1988) stereotaxic atlas. The major difference between these two atlases is in the inferior
portion of the temporal lobes (Calder, Lawrence, & Young, 2001; Duncan et al., 2000).
Considering the age of our participants and our voxel size, it is reasonable to normalize all
participants into the standard MNI template (Kang, Burgund, Lugar, Petersen, & Schlaggar,
2003; Burgund et al., 2002; Wilke, Schmithorst, & Holland, 2002; Muzik, Chugani, Juhasz,
Shen, & Chugani, 2000).

Statistical analyses were calculated on the smoothed data (7 mm isotropic gaussian kernel)
using a delayed boxcar design with a 6-sec delay from onset of block in order to account for
the lag in hemodynamic response. A high pass filter was applied equal to two cycles of the
experimental and control conditions (216 sec) in order to remove low-frequency effects such
as signal drift, cardiac and respiratory pulsations. We used global normalization to scale the
mean of each scan to a common value in order to correct for whole brain differences over time.

We calculated contrasts (word-control) in order to analyze the two word judgment tasks
(spelling, rhyming) in the two modalities (visual, auditory). A one-sample t test compared each
voxel across all participants within a group (children or adults) to determine whether the
activation during a condition was significant (i.e., greater than 0). A two-sample t test was used
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to determine whether the magnitude of activation across groups was significantly different.
Reported areas of activation were significant using p < .001 corrected at the voxel level for the
one-sample tests and p < .01 corrected at the voxel level for the two-sample tests. All clusters
were greater or equal to 15 voxels.

Acknowledgements
We thank Sarah Brennan, Yasu Harasaki, and Frank Van Santen for their assistance in stimulus development, and
Kristin Bettenhausen, Jean Rex, and Cheryl Wolf for their assistance in conducting the behavioral study. We thank
Nirmal Christian, Wei Li, Paul Springer, and Robert Salzman for their operation of the MRI. We also thank the students,
teachers, and administrators at Pope John XXIII School, Saint Athanasius School, and Saint Peter’s Catholic School
for their participation in the behavioral study. Finally, we thank the participants in the fMRI study for their time and
commitment to research. This research was supported by grants from the National Institute of Child Health and Human
Development (HD042049), the National Institute of Deafness and Other Communication Disorders (DC06149), and
the National Institute of Aging (P50 AG06882 and P30 AG097761).

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Figure 1.
Significant activation for the adults and children on the spelling (S) and rhyming (R) tasks in
the visual (V) and auditory (A) modalities. These slices show activation for the adults (red),
for the children (green), and for both adults and children (purple).
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 Booth et al. Page 16
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Figure 2.
Significant differences between the adults and children activation on the spelling (S) and
rhyming (R) tasks in the visual (V) and auditory (A) modalities. These slices show significantly
greater activation for the adults (red) or for the children (green). Only clusters in our regions
of interest are labeled (see Tables 4, 5, 6, and 7 for all significant regions). Slices were chosen
to maximize visualization of clusters while selecting similar z-coordinates across tasks (AG =
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angular gyrus; FG = fusiform gyrus; IFG = inferior frontal gyrus; MTG = middle temporal
gyrus; STG = superior temporal gyrus).

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Table 1
Examples of Stimuli for the Word Judgment Tasks

Parametric Manipulation
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Similar Orthography Dissimilar Orthography

Spelling
First match hold–plant–cold Hope–colt–soap
Second match built–vote–note slid–lane–strain
Rhyming
First match seat–fresh–heat Jazz–last–has
Second match wish–fall–wall myth–home–foam

For the spelling and rhyming task, there was a parametric manipulation of orthographic similarity.
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Table 2
Examples of Stimuli for Control Tasks in the Visual and Auditory Modality

Modality
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Visual Auditory

First match / \–\\–/ \ 300–500–300 Hz

Second match //–\\–\\ 500–700–700 Hz
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Table 3
Means (M) and Standard Errors (SE) for Accuracy (%) and Reaction (RT) for the Word Judgment and Control Tasks

Word Control

% RT % RT

M SE M SE M SE M SE
Booth et al.

ADULT—VIS
Spelling 97.3 4.1 931 80 96.5 4.6 797 69
Rhyming 96.9 4.4 947 79 97.5 3.9 804 70
CHILD—VIS
Spelling 89.3 7.7 1545 105 93.4 6.2 1214 93
Rhyming 91.6 6.9 1444 89 94.2 5.8 1229 82
ADULT—AUD
Spelling 91.4 7.0 1265 107 93.0 6.4 1003 73
Rhyming 97.4 4.0 993 68 95.9 5.0 977 68
CHILD—AUD
Spelling 81.8 9.6 1660 123 90.7 7.3 1391 94
Rhyming 94.8 5.6 1350 84 90.5 7.3 1376 92

Data are presented separately for the adults and children for each task (spelling, rhyming, and meaning) in each modality (visual and auditory).

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Table 4
Significant Activation Separately for the Adults and Children and Significant Differences between Adults and Children for the Visual Spelling Task

Location Significance Coordinate

Area H BA z-test Voxels x y z

Adult
Booth et al.

Angular gyrus/Precuneus/Superior parietal L 39/19/7 15.13 372 −30 −66 42

lobule
Angular gyrus/Precuneus/Superior parietal R 39/19/7 12.61 236 30 −54 42
lobule
Medial frontal gyrus/Anterior cingulate B 6/8/32 12.44 257 −3 33 42
Inferior frontal gyrus R 9/44/45 11.78 302 51 30 21
Inferior frontal gyrus L 9/44/45/47 21.22 1073 −48 36 6
Middle occipital gyrus/Fusiform gyrus R 18/19/37 17.61 199 24 −90 −6
Middle occipital gyrus/Fusiform gyrus L 18/19/37 22.51 639 −36 −51 −21
Cerebellum L * 8.85 20 −9 −81 −30
Cerebellum R * 8.63 37 12 −78 −33
Child
Medial frontal gyrus/Anterior cingulate L 6/32 10.09 283 −6 15 45
Angular gyrus/Precuneus/Superior parietal L 39/19/7 12.59 205 −24 −57 39
lobule
Inferior frontal gyrus L 9/44/45/47 18.20 1140 −42 9 24
Inferior frontal gyrus R 44/45 7.11 37 51 33 18
Middle temporal gyrus L 21 8.83 41 −51 −36 0
Inferior frontal gyrus/Insula R 47/13 8.35 83 30 27 −3
Putamen L * 7.62 51 −15 6 −3
Middle occipital gyrus/Fusiform gyrus L 18/19/37 14.52 392 −36 −72 −12
Middle occipital gyrus/Fusiform gyrus R 19/37 10.39 131 39 −69 −18
AD–CH
Angular gyrus/Superior parietal lobule L 39/7 8.09 89 −30 −69 45
Inferior frontal gyrus L 9 7.34 68 −48 15 39
Middle cingulate L 31 6.44 18 −6 −36 36
Medial frontal gyrus L 9 6.09 18 −3 42 33
Inferior frontal gyrus R 9/44 8.38 51 51 18 30
Angular gyrus R 39 6.26 21 45 −54 30
CH–AD
Medial frontal gyrus L 6 6.40 40 −6 3 54
Precuneus L 7 8.43 54 −21 −57 51
Postcentral gyrus R 2 5.85 30 48 −24 45
Precentral gyrus L 6 6.48 49 −42 −3 42

J Cogn Neurosci. Author manuscript; available in PMC 2006 February 27.

Anterior cingulate L 32 6.41 25 −6 27 30
Insula R 13 7.01 25 33 21 6
Lingual gyrus L 18 6.60 35 −6 −75 0
Inferior temporal gyrus R 37 6.52 15 57 −54 −3

H = left (L), right (R), or bilateral (B) hemispheres; BA = Brodmann’s area of activation; voxels = number of voxels in cluster, only clusters 15 or greater are presented; coordinates = −X left
hemisphere, +X right hemisphere, −Y behind anterior commisure, +Y in front of anterior commisure, −Z below anterior–posterior commisure plane, +Z above anterior–posterior commisure plane.
Regions are sorted by z-coordinate within a group.
*
No Brodmann’s area for this brain region.
Page 20
 Booth et al. Page 21

Table 5
Significant Activation Separately for the Adults and Children and Significant Differences between Adults and
Children for the Visual Rhyming Task
NIH-PA Author Manuscript

Location Significance Coordinate

Area H BA z-test Voxels x y z

Adult
Medial frontal gyrus L 6/8 9.84 161 −6 33 42
Precuneus L 7 7.55 27 −24 −54 39
Inferior frontal gyrus R 44 7.26 19 54 33 12
Inferior frontal gyrus L 9/44/45/47 19.69 1070 −48 33 6
Middle temporal gyrus/Angular L 21 8.01 74 −48 −42 6
gyrus
Middle temporal gyrus L 21 8.93 58 −63 −33 3
Middle occipital gyrus/Fusiform R 18/37 13.69 224 24 −90 −6
gyrus
Middle occipital gyrus/Fusiform L 18/19/37 21.31 485 −39 −54 −21
gyrus
Cerebellum R * 9.57 87 12 −75 −30
Child
Inferior frontal gyrus L 9 8.57 33 −51 −3 42
Inferior frontal gyrus L 44/45/47 14.30 707 −51 30 9
Cuneus R 17 7.36 16 21 −81 3
Middle temporal gyrus L 21 11.49 128 −51 −36 0
Parahippocampus L 35 7.21 47 −24 −27 −12
Middle occipital gyrus/Fusiform L 18/19/37 14.99 355 −39 −48 −21
gyrus
Fusiform gyrus R 37 8.27 58 36 −63 −21
AD–CH
NIH-PA Author Manuscript

Inferior frontal gyrus L 9 7.73 43 −48 21 33

Angular gyrus/Middle temporal L 39/37 7.47 34 −51 −57 12
gyrus
Inferior frontal gyrus R 44 7.04 20 57 6 12
CH–AD
 No significant differences

See Table 4 note.

NIH-PA Author Manuscript

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 Booth et al. Page 22

Table 6
Significant Activation Separately for the Adults and Children and Significant Differences between Adults and
Children for the Auditory Spelling Task
NIH-PA Author Manuscript

Location Significance Coordinate

Area H BA z-test Voxels x y z

Adult
Medial frontal gyrus/Anterior L 6/8/32 14.72 296 −6 24 42
cingulate
Angular gyrus/Superior parietal R 39/7 7.23 38 33 −51 42
lobule
Angular gyrus/Precuneus/ L 39/19/7 13.05 343 −30 −63 36
Superior parietal lobule
Inferior frontal gyrus L 9/44/45/47 22.4 1101 −45 12 21
Inferior frontal gyrus/Middle R 45/46 9.58 253 54 33 18
frontal gyrus
Putamen L * 7.43 68 −12 0 9
Superior temporal gyrus/Middle L 22/21/37 25.89 948 −60 −12 −3
temporal gyrus/Fusiform gyrus
Superior temporal gyrus/Middle R 22/21 21.5 539 60 −12 −6
temporal gyrus
Inferior frontal gyrus/Insula R 47/13 9.84 79 33 27 −9
Cerebellum R * 7.16 29 30 −66 −30
Cerebellum R * 11.56 72 12 −81 −33
Child
Medial frontal gyrus L 6 7.28 48 −6 27 39
Inferior frontal gyrus L 9/44/45/47 12.25 516 −48 12 21
Posterior cingulate L 30 6.90 61 −6 −78 6
Superior temporal gyrus/Middle L 22/21/37 18.91 765 −60 −18 −3
NIH-PA Author Manuscript

temporal gyrus/Fusiform gyrus

Superior temporal gyrus/Middle R 22/21 13.6 317 60 −12 −3
temporal gyrus
Inferior frontal gyrus L 47 6.93 15 −30 21 −9
Cerebellum R * 7.50 20 12 −75 −27
AD–CH
Superior parietal lobule L 7 7.46 40 −45 −42 48
Inferior frontal gyrus L 9 6.25 15 −45 12 42
Angular gyrus L 39 7.50 35 −30 −63 36
Inferior frontal gyrus R 9/44 8.12 138 48 12 27
Inferior frontal gyrus L 9/44 7.06 70 −42 36 9
Inferior frontal gyrus L 45 6.95 25 −45 36 9
Inferior frontal gyrus R 45 7.58 51 42 36 6
Superior temporal gyrus L 22 7.35 24 −60 −9 0
Middle temporal gyrus R 21 6.53 30 54 −15 −12
CH–AD
Fusiform gyrus L 37 7.55 74 −33 −54 −15

See Table 4 note.

NIH-PA Author Manuscript

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 Booth et al. Page 23

Table 7
Significant Activation Separately for the Adults and Children and Significant Differences between Adults and
Children for the Auditory Rhyming Task
NIH-PA Author Manuscript

Location Significance Coordinate

Area H BA z-test Voxels x y z

Adult
Precentral gyrus L 6 8.00 27 −54 −6 42
Inferior frontal gyrus L 9/44/45/47 13.51 489 −48 36 6
Superior temporal gyrus/Middle L 22/21/37 24.69 660 −63 −12 −3
temporal gyrus/Fusiform gyrus
Superior temporal gyrus/Middle R 22/21 20.48 309 63 −9 −6
temporal gyrus
Cerebellum R * 9.40 58 24 −63 −24
Child
Inferior frontal gyrus L 44/45/47 10.11 61 −54 27 3
Superior temporal gyrus/Middle L 22/21 17.74 442 −60 −18 −3
temporal gyrus
Superior temporal gyrus/Middle R 22/21 15.81 270 60 −9 −6
temporal gyrus
Fusiform gyrus L 37 7.43 21 −42 −51 −15
AD–CH
Angular gyrus/Superior parietal L 39/7 6.51 27 −42 −45 48
lobule
Precentral gyrus L 6 6.57 17 −48 −9 39
Inferior frontal gyrus L 9/44/45 8.37 213 −45 36 9
Superior temporal gyrus L 22 7.74 23 −60 −9 −3
CH–AD
 No significant differences
NIH-PA Author Manuscript

See Table 4 note.

NIH-PA Author Manuscript

J Cogn Neurosci. Author manuscript; available in PMC 2006 February 27.