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Frontoparietal networks involved in categorization and item working memory

Article in NeuroImage · December 2014

DOI: 10.1016/j.neuroimage.2014.11.051 · Source: PubMed

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Neuroimage. 2015 February 15; 107: 146–162. doi:10.1016/j.neuroimage.2014.11.051.

Frontoparietal networks involved in categorization and item

working memory
Kurt Braunlich,
Colorado State University, Cognitive Psychology and Molecular, Cellular and Integrative
Neurosciences

Javier Gomez-Lavin, and

CUNY, Department of Philosophy

Carol A. Seger*
Colorado State University, Cognitive Psychology and Molecular, Cellular and Integrative
Neurosciences
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Abstract
Categorization and memory for specific items are fundamental processes that allow us to apply
knowledge to novel stimuli. This study directly compares categorization and memory using delay
match to category (DMC) and delay match to sample (DMS) tasks. In DMC participants view and
categorize a stimulus, maintain the category across a delay, and at the probe phase view another
stimulus and indicate whether it is in the same category or not. In DMS, a standard item working
memory task, participants encode and maintain a specific individual item, and at probe decide if
the stimulus is an exact match or not. Constrained Principal Components Analysis was used to
identify and compare activity within neural networks associated with these tasks, and we relate
these networks to those that have been identified with resting state-fMRI. We found that two
frontoparietal networks of particular interest. The first network included regions associated with
the dorsal attention network and frontoparietal salience network; this network showed patterns of
activity consistent with a role in rapid orienting to and processing of complex stimuli. The second
uniquely involved regions of the frontoparietal central-executive network; this network responded
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more slowly following each stimulus and showed a pattern of activity consistent with a general
role in role in decision-making across tasks. Additional components were identified that were
associated with visual, somatomotor and default mode networks.

Keywords
working memory; categorization; connectivity; PCA; fMRI

© 2014 Elsevier Inc. All rights reserved.

*
Corresponding author: Carol A. Seger, Department of Psychology, 1876 Campus Delivery, Colorado State University, Fort Collins,
CO 80523, USA, Tel: +1 970 491-3540, Fax: +1 970 491-1032, [email protected].
Publisher's Disclaimer: This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our
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The authors declare no competing financial interests.
 Braunlich et al. Page 2

1 Introduction
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Categorization and specific-item memory are fundamental processes which allow us to

apply knowledge to novel situations. Categorization requires abstraction from inherent
stimulus features to generalizable latent features, and plays an important role in the flexible
transfer of knowledge and skills across stimuli and tasks (for review, see Seger & Miller,
2010). In contrast, memory for specific items maintains these inherent stimulus features in
order to enable us to make fine distinctions between items. Despite these fundamental
differences, both categorization and specific item memory tasks recruit common cognitive
control systems to support task performance (Cole et al., 2013; Gazzaley & Nobre, 2012;
Seger & Peterson, 2013), raising the question of how the same neural systems can serve
different ends. This study directly compares categorization and specific item memory using
delayed-match-to-category (DMC) and delayed-match-to-sample (DMS) tasks in which
participants encode a stimulus, maintain information across a delay, see a second stimulus
and then decide if it matches the first. The tasks share similar structure, and therefore place
similar demands on perceptual (stimulus encoding), motor (response execution) and some
executive functions (working memory and decision-making). The tasks differ in what is
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encoded at the first stimulus and in the basis of the match-mismatch decision occurring at
probe: specific item identity in the DMS task and category in the DMC task. Our design,
therefore, allows us to isolate differences between processes associated with categorization
and those associated with item-specific memory, and also to identify shared processes.
Below we first discuss proposed shared cognitive control functions across categorization and
specific-item tasks and how they may rely on intrinsically connected frontoparietal neural
networks. We then discuss aspects of cognitive processing specific to categorization and to
item working memory. Finally we describe our task and our predictions.

1.1 Shared cognitive control processes and intrinsic neural systems

Much recent research has focused on how frontoparietal networks can be flexibly recruited
to support cognitive control in diverse task environments (Cole et al., 2013; Dumontheil,
Thompson, & Duncan, 2011; Duncan, 2010). Multiple networks supporting cognitive
control have been identified, and although there is currently little consensus concerning
network nomenclature, we will focus on two networks that show coactivation across a
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variety of cognitive tasks and correlated patterns of intrinsic activity during resting-state
fMRI (Dosenbach et al., 2007; Seeley et al., 2007). The first, the salience network
(abbreviated here as SA), which has important nodes in the anterior insula / frontoinsular
cortex and dorsal anterior cingulate (ACC) / medial frontal gyrus, is thought to play an
important role in bottom-up detection of salient external events, the coordination of
functional networks to meet task demands, and in moderating autonomic arousal (Medford
& Critchley, 2010; Menon & Uddin, 2010; Menon, 2011; Sridharan, Levitin, & Menon,
2008). The second, the central executive network (FP-CEN), which has important nodes in
the dorsolateral prefrontal cortex and posterior parietal cortex / intraparietal sulcus, is
thought to operate on the salient stimuli marked by the SA network (Seeley et al., 2007), and
to play an important role in the manipulation and maintenance of these representations in
working memory and rule-based processes (Miller & Cohen, 2001).

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Intrinsic connectivity has also identified other networks, such as the somatomotor network
(SM; primary motor and somatosensory cortex), dorsal attentional network (DA; premotor
and superior parietal cortex), default mode network (DMN; medial frontal and posterior
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cingulate regions, inferior parietal and medial temporal lobe), and visual network (VS;
occipital and inferior temporal cortex) (Buckner, Krienen, Castellanos, Diaz, & Yeo, 2011;
Choi, Yeo, & Buckner, 2012; Yeo et al., 2011). A focus of recent research has been to
identify how these networks interact, and one particularly important finding is that the FP-
CEN and SA, which show greater activity during cognitively-demanding tasks (Chen et al.,
2013; Dang, Donde, & Madison, 2012; Vanhaudenhuyse & Demertzi, 2011) are
anticorrelated with the DMN. The SA is thought to play an important role in mediating this
anticorrelated relationship, and in switching from the DMN to the FP-CEN in response to
salient external events (Bonnelle et al., 2012; Goulden et al., 2014; Menon, 2011;
Palaniyappan, Simmonite, White, Liddle, & Liddle, 2013; Sridharan et al., 2008).

How these primarily cortical intrinsic connectivity networks interact with subcortical and
cerebellar regions is an active area of research. Buckner and colleagues, for instance,
examined functional connectivity between the cortical intrinsic connectivity networks and
the basal ganglia and the cerebellum (Buckner, Krienen, Castellanos, Diaz, & Yeo, 2011;
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Choi, Yeo, & Buckner, 2012). Both basal ganglia and cerebellum had separate regions that
correlated with each cortical network, consistent with known projections from cortex to
these structures. Particularly relevant for our study are the interconnections with the FP-
CEN, which are primarily correlated with the dorsal head and body of the caudate nucleus
(Choi et al., 2012), and the lateral cerebellar hemisphere (Buckner et al., 2011).

1.2 Categorization
Cognitive neuroscience studies have associated categorization with a large distributed neural
network including the basal ganglia (Seger, 2008), lateral frontal (Muhammad, Wallis, &
Miller, 2006), lateral parietal cortex (Daniel et al., 2011; Freedman & Assad, 2009; Rishel,
Huang, & Freedman, 2013), precuneus (Wenzlaff, Bauer, Maess, & Heekeren, 2011),
premotor and supplementary motor areas (Ashby, Ennis, & Spiering, 2007; Little, Shin,
Sisco, & Thulborn, 2006; Waldschmidt & Ashby, 2011). Although still an active area of
research, clues are emerging as to the individual contributions made by each region. The
basal ganglia has been associated with multiple processes: posterior regions are involved in
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mapping visual stimuli to category, and category to motor response, whereas anterior
regions and the ventral striatum are associated with feedback and reward processing (Seger,
2008). Frontal regions have been associated with maintenance and implementation of
categorization rules (Antzoulatos & Miller, 2011; Buschman, Denovellis, & Diogo, 2012;
Freedman, Riesenhuber, Poggio, & Miller, 2001; Meyers, Freedman, Kreiman, Miller, &
Poggio, 2008; Muhammad et al., 2006; Wallis & Miller, 2003). The parietal cortex
combines category with motor response, and may be responsible for integration of relevant
information for category membership (Freedman & Assad, 2009; Shadlen & Newsome,
2001; Swaminathan & Freedman, 2012). The precuneus and SMA, along with regions of the
basal ganglia they interact with, may be associated with setting a response criterion
(Forstmann et al., 2008; Wenzlaff et al., 2011). In addition, inferotemporal cortex performs
relevant visual processing necessary for categorization, though it is still unclear the degree

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to which this region changes with learning and contributes to the representation of novel
categories.
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Previous categorization studies in humans have typically required participants to view,

categorize and respond to single stimuli in rapid succession. However, like the DMC task,
real life situations often have a delay between the categorization of a stimulus and a
subsequent behavioral response, or require that multiple categorical representations be
integrated in order to determine the correct response. The DMC task is advantageous in that
it allows us to examine category maintenance and integration across stimuli, and also allows
the dissociation of categorization processes from those related to motor preparation. The
DMC task was originally developed for research with non-human primates, which find
category sensitivity independent of motor response for neurons in inferotemporal, parietal,
frontal, and basal ganglia regions (Freedman & Miller, 2008). Electrophysiological
recordings suggest coordination between the frontoparietal network and regions within the
inferior temporal lobe during this task, such that inferior temporal regions tend to be more
sensitive to the visual features of individual exemplars, while prefrontal regions are more
sensitive to features relevant for successful task performance (e.g., categorical-status of the
first stimulus, and match-mismatch status of the second; Freedman et al., 2003; Meyers et
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al., 2008a).

1.3 Working Memory

The DMS task has been used extensively to investigate specific-item working memory in
human and non-human primates. A large body of research finds that frontoparietal regions
are recruited during the performance of DMS tasks (Sala, Rämä, & Courtney, 2003).
However, there are some differences based on task demands; for example, working memory
for objects rather than spatial location is particularly reliant on ventrolateral PFC regions in
the middle and inferior frontal gyri (Sala et al., 2003). There is also evidence that working
memory for objects involves interactions between these frontoparietal networks and higher
order visual cortical regions involved in representing the objects (Gazzaley, Rissman, &
D’Esposito, 2004), and evidence that memory for specific items can lead to interactions with
the hippocampus (Rissman, Gazzaley, & D’Esposito, 2008). Neural networks recruited
during DMS performance vary depending on task demands. Increased working memory
demands have been associated with increased connectivity between regions in the
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inferotemporal cortex and the hippocampus, and decreased connectivity between

inferotemporal regions and the inferior frontal gyrus (Rissman et al., 2008). Similarly,
during the delay epoch, inferotemporal regions associated with task-relevant processes show
increased connectivity with the frontoparietal network while regions associated with task-
irrelevant processes show increased connectivity with the default mode network (Chadick
and Gazzaley, 2011).

1.4 Present Study

In the present study, we directly compared patterns of activity during DMC and DMS tasks
utilizing the same perceptually-similar stimuli (young Caucasian female faces) and the same
timing and responses such that the tasks differed only in the requirement to either categorize
the face or remember the specific face. As in several previous DMS studies, we chose to use

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facial stimuli, as the processing of these stimuli is known to occur with localized regions of
the fusiform cortex (cf., Gazzaley et al., 2004; Rissman et al., 2008). Two versions of the
DMC task were used, which we termed “Category” and “Label.” In both, participants
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viewed a face at encoding, categorized it, and maintained the category membership across a
delay. At probe, the conditions differed: in the Category version participants viewed a
second face, whereas in the Label version they viewed the category label (“A” or “B”). In
both of these conditions, participants decided whether the categories matched. The Label
condition allowed us to discriminate between activity due to comparing category labels and
match-mismatch decision making, versus activity due to viewing and categorizing the face.
Thus, we predicted that regions involved in stimulus categorization should have high
activity at encoding for both Label and Category, but only for Category at probe. However,
regions associated with category match-mismatch decision making should show activity
more broadly in both tasks.

We predicted some common and some differing patterns of activity based on the shared and
individual characteristics of categorization and item working memory. First, because of the
paired task design that equated basic visual and motor demands, we predicted similar
recruitment of the visual and motor systems. Second, as both tasks require weighing
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evidence towards the binary response options, we predicted that the tasks would commonly
elicit activity in regions associated with general decision-making processes (Freedman &
Assad, 2011; Seger & Peterson, 2013). We predicted that the primary differences between
tasks would be found in regions associated with the FP-CEN. Categorization involves
several different strategies that require cognitive control, including evaluating information
with respect to categorization criteria and mapping the stimulus to category membership
(Seger & Peterson, 2013). In contrast, working memory requires different control processes
for encoding and maintenance, potentially via interactions with inferotemporal cortex and
the hippocampus (Rissman et al., 2008).

2 Methods
2.1 Participants
Seventeen participants were recruited from the Colorado State University Community. All
participants were healthy, right-handed adults (11 females, 6 males) with an average age of
27 (range: 20–37). Participants were screened for history of psychiatric or neurological
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disorders, for current use of psychoactive medications, and exclusionary criteria for fMRI
(e.g., claustrophobia, metallic implants).

2.2 Stimuli
Twenty-five similar young adult female Caucasian faces were selected for the stimulus set.
To discourage use of verbalizable memory strategies, all images were cropped so that the
whole face, but no other defining characteristics, was shown. The faces were then warped
and resized to subtend a visual angle of roughly 3.9 degrees horizontally and 6.9 degrees
vertically. For each participant, eight stimuli were randomly assigned to category “A” and
eight were randomly assigned to category “B.” This type of categorization task is sometimes
referred to as arbitrary, or unstructured because the stimuli are randomly assigned to

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category and do not include any intentional within-category similarities. Unstructured tasks
rely on procedural knowledge to a similar degree as structured implicit categorization tasks
(Crossley, Madsen, & Ashby, 2012), and recruit similar cortical and striatal systems (Seger,
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Dennison, Lopez-Paniagua, Peterson, & Roark, 2011; Seger, Peterson, Cincotta, Lopez-
Paniagua, & Anderson, 2010). The remaining stimuli were used in the Item condition.

2.3 Procedure
Prior to scanning, participants performed two tasks on a laptop computer; they first learned
to categorize faces and then trained on a task that was similar to what they would later
perform in the scanner. In the category-learning task, participants learned to categorize each
of the 16 faces into category “A” or category “B” via trial and error. On each trial, a face
was presented in the center of the computer screen, and the category labels were presented at
the bottom left and right of the screen. To encourage participants to learn category labels
rather than specific motor responses, the locations of the labels were randomly determined
on each trial (i.e., “A” appeared at the bottom left of the screen on some trials, but at the
bottom right on others). Participants responded by pressing the “d” key on the laptop
keyboard if the chosen category label was on the bottom left side of the screen, and the “k”
key if it was on the bottom right. Each image remained on the screen until the participant
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made a response. Following each response, auditory and visual feedback was presented for
0.75 seconds. Following correct responses, the word “Correct” was presented in the center
of the computer screen with a pleasant tone. Following incorrect responses, the word
“Wrong” was presented with an unpleasant tone. Every 100 trials, participants were given a
self-paced break. Participants trained until they reached an 85% correct performance
criterion on the final block of 100 trials.

To gain familiarity with the task that would later be used in the scanner, after reaching the
85% performance criterion, participants performed a second training task similar to that they
would perform in the scanner (Figure 1 illustrates the task performed in the scanner). At the
beginning of each trial, a cue was presented for 1.5 seconds, which instructed participants to
either “Match the Specific Face,” or “Match the Category.” After this cue was presented, a
face stimulus was presented for 1.5 seconds in the center of the computer screen. After a
nine-second delay (during which time participants saw only a blank screen), a second face
stimulus (or, in the Label condition, the category label “A” or “B”) was presented for three
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seconds. Two response cues, “Match” and “Mismatch” were then added to the display (at
the bottom left and bottom right of the screen, respectively). On trials in which participants
were cued to remember the specific face, participants had to indicate whether the second
face was the same as the first. On trials in which participants were instructed to remember
the category, they had to indicate whether the second stimulus (face or category label)
belonged to the same category as the first. No feedback was delivered. Trials were separated
by a 1.5 second inter-trial interval (ITI) during training. All participants performed 30 trials
of the second training task. The assignment of condition to trial was random (selected with
replacement), such that there were 10 trials per condition. As in the first training task,
participants made their responses via the index fingers of their right and left hands using the
“d” and “k” keys.

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In the scanner, the task was similar to the second pretraining task described above. Stimuli
were, however, presented via a back-projection mirror positioned above the participant, and
responses were made with fingers of the right and left hands via separate button boxes. The
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ITI was jittered according to a positively-skewed geometric distribution ranging from 1.5 to
9 seconds. Participants performed two 15-minute runs. In order to increase power for
analyses of the Item trials in contrast with Categorical Encoding trials, we presented fewer
Category (14) and Label (14) trials than Item trials (17) during each run. Both correct and
incorrect trials were included in the analyses to maximize statistical power.

2.4 Image Acquisition

Images were obtained with a 3.0 Tesla MRI scanner (Siemens) at the Intermountain
Neuroimaging Consortium (Boulder, CO). The scanner was equipped with a 12-channel
head coil. Structural images were collected using a 3D T1-weighted rapid gradient-echo
(MPRAGE) sequence (256 × 256 matrix; FOV, 256; 192 1-mm sagittal slices). Functional
images were reconstructed from 28 axial oblique slices obtained using a T2* -weighted 2D-
EPI sequence (TR, 1500ms; TE, 25ms; FA, 75; FOV, 220-mm, 96 × 96 matrix; 4.5-mm
thick slices; no inter-slice gap). Each run consisted of 597 volumes. The first three volumes,
which were collected before the magnetic field reached a steady state, were discarded.
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2.4.1 Preprocessing—Image preprocessing was performed using SPM8 (http://

www.fil.ion.ucl.ac.uk/spm/software/spm8). Preprocessing involved correction of slice time
acquisition differences (images were adjusted to the 14th slice), motion correction of each
volume to the first volume of the first run using 3rd degree B spline interpolation,
coregistration of the functional to the structural data, normalization to the MNI template,
smoothing (with a 6 mm Gaussian kernel), and temporal filtering (with a 128 sec. high-pass
filter). One participant had excessive head-movement (defined as greater than 3mm
translational or 2.5° rotational movement). This participant’s data were excluded from
subsequent analyses.

2.5 FMRI Analyses

2.5.1 Univariate General Linear Model—Trial epochs (i.e., encoding, delay, and probe)
were modeled as independent regressors in a univariate whole-brain analysis. All trials were
included in this analysis. The encoding regressor was coded as a 1.5 s. boxcar coinciding
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with the presentation of the first stimulus (1.5–3 sec. after cue onset). The delay period was
modeled as a 2 second boxcar placed halfway through the delay period (6–8 sec. after cue
onset). The probe period was modeled as a 3 second boxcar coinciding with the presentation
of the second stimulus (12–15 sec. after cue onset). As in previous research, the onsets of
these regressors were placed at least 4 seconds apart to minimize the influence of preceding
trial epochs (Barde & Thompson-Schill, 2002; Druzgal & D’Esposito, 2003; Gazzaley et al.,
2007, 2004; Pessoa, Gutierrez, Bandettini, & Ungerleider, 2002; Postle, Zarahn, &
D’Esposito, 2000; Rissman, Gazzaley, & D’Esposito, 2004; Rissman et al., 2008; Zarahn,
Aguirre, & D’Esposito, 1997). We convolved each boxcar with the canonical SPM HRF.
For each contrast, we generated maps at an uncorrected threshold of p < 0.001 and corrected
for multiple comparisons using the topological false-discovery rate (Chumbley & Friston,
2009).

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2.5.2 Constrained Principal Component Analyses (CPCA)—To investigate task-

related differences across functional networks, we used Constrained Principal Component
Analyses (CPCA) using a finite-impulse response (FIR) model, as implemented in the
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fMRI-CPCA toolbox (www.nitrc.org/projects/fmricpca). CPCA combines multivariate

regression and principal component analysis to identify multiple functional networks
involved in a given task, and has been used successfully with similar experimental
paradigms (Metzak et al., 2012; Metzak et al., 2011; Woodward, Feredoes, Metzak, Takane,
& Manoach, 2013). This approach is mathematically similar to Partial Least Squares
analysis (McIntosh, Bookstein, Haxby, & Grady, 1996), and is attractive, as it allows
estimation of changes in the BOLD response across peristimulus time within each functional
network, and also allows statistical inference concerning the importance of each column of
the design matrix for each component.

CPCA involves preparation of two matrices: Z, and G. Z contains the BOLD time course of
each voxel, with one column per voxel and one row per scan. The design matrix, G contains
a FIR model of the BOLD response related to the event onsets. The BOLD time-series in Z
is regressed onto the design matrix, G, yielding a matrix, C, of regression weights. GC thus
contains the variance in Z, that is accounted for by the design matrix, G. Components are
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then extracted from the variance in GC via singular value decomposition, yielding U, a
matrix of left singular vectors, D a diagonal matrix of singular values, and V, a matrix of
right singular vectors. The columns of VD, which reflect component loadings, can be
overlaid on a structural image to visualize the functional networks. To maximize the
variance of the squared loadings, we orthogonally rotated VD prior to display. The top 5% of
these rotated loadings for each component are illustrated in Figures 3B, 4B, 5B, 6B and 7A.
Several previous studies (e.g., Metzak et al., 2012; Metzak et al., 2011) have used a similar
threshold. For each combination of peristimulus time-point, condition and participant,
CPCA estimates a set of predictor weights (P), which are the values that relate the design
matrix, G, to the networks associated with each component, such that U = G × P. All trials
were included in this analysis.

We conducted a repeated measures ANOVA using SPSS, which allowed us to investigate

the consecutive scans where the slope of the predictor weight time course differed between
conditions. This allows investigation of differences between conditions or contiguous time
points, without considering the complex hemodynamic shape (cf., Metzak et al., 2012). For
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these analyses, and in Figures 3B, 4B, 5B, 6B and 7A, we adjusted the time-series so that
the first observation was zero for all conditions (cf., Metzak et al., 2011, 2012; Woodward et
al., 2013). We tested assumptions of sphericity, and controlled for violations using
Greenhouse-Geisser adjusted degrees of freedom.

Readers more familiar with the interpretation of statistical maps derived from univariate
analyses should take care when interpreting the multivariate results shown in Figures 3, 4, 5,
6 and 7. Whereas statistical maps derived from univariate analyses provide information
about activity occurring within specific regions, each CPCA component reflects a pattern of
task-related variance derived from all voxels in the brain. The maps derived from CPCA
analyses, therefore, provide information about regions that cooperate to subserve a particular
function. Whereas most univariate analyses assume a HDR shape by using a standard GLM

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with a canonical HRF, CPCA uses a FIR model which uncovers network-specific HDR
shapes of task-related variance in a data-driven manner. This is valuable, as it can help
segregate and characterize task-related processes that might not have been predicted by the
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experimenter.

In the present paper, we use univariate analyses to characterize activity occurring within
specific regions of the brain, and we use CPCA to investigate how distributed regions
coordinate to subserve different processes. In the tables provided, we label cluster peaks
according to a 7-network parcellation identified in previous research (Buckner et al., 2011;
Choi et al., 2012; Yeo et al., 2011), but refer to some regions that these papers term the
ventral attentional network as the salience network (SA), in line with current usage
(Buckner, Krienen, & Yeo, 2013).

3 Results
3.1 Behavioral
All trials for which participants made a behavioral response were included in all analyses.
Accuracy was highest for the Item conditions (M = 94.87% correct, SD = 5.83), lower for
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the Label condition (M= 81.11%, SD=12.36) and lowest for the Category condition (M =
75.46%, SD = 16.71), F(2,45) = 10.25, p < .001, η2 = 0.31. The accuracy difference between
the Label and Category conditions is likely related to the different number of categorization
decisions required for each condition: the Category trials required participants to categorize
stimuli at encoding and probe, and an error on either decision could lead to an incorrect
response, whereas the Label trials required participants to categorize stimuli only at
encoding. Performance in the Label condition was close to that of the 85% accuracy
criterion from the learning phase. We did not collect reaction time data because it was
unlikely to be of interest due to the requirement that participants delay their response until
the response cue was presented.

3.2 Neuroimaging
3.2.1 Univariate GLM Analyses—Whole brain GLM analyses were used to examine
regions of activity during each trial epoch: encoding, delay and probe (cf., Gazzaley et al.,
2007, 2004). Because the Category and Label trials were identical until the onset of the
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second stimulus, these conditions were combined as the “categorical-encoding” condition

for examination of activity during the encoding and delay epochs. In this section we present
univariate regions of activity across the whole brain; we also discuss these results masked by
each component later following the CPCA results.

As can be seen in Table A.1 and Figure 2, during encoding, the Categorical-Encoding trials
elicited greater activity than Item trials primarily within frontal lobe regions (middle
cingulate, superior medial gyrus, inferior frontal / anterior insula) and subcortical regions
that are known to interact with the frontal lobe (right caudate, left cerebellar lobule VI).
These regions participate in the frontoparietal intrinsic connectivity network (Yeo et al.,
2011). In addition, categorical encoding recruited visual regions including the bilateral
calcarine gyri. The Item condition elicited greater activity than Categorical-Encoding trials

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in the left inferior frontal gyrus, regions of the temporal and occipital lobe associated with
high level visual processing, and the bilateral hippocampus. All of these regions have been
identified in previous studies as being recruited during visual working memory encoding
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(Gazzaley et al., 2007; Sala et al., 2003). The only region showing significant activation in
response to both conditions (conjunction analysis, contrast with implicit baseline) at
encoding was the crus I region of the right cerebellum.

During the delay period, the majority of regions sensitive to differences between conditions
showed patterns of activity suppressed below implicit baseline (cf. Gazzaley, Rissman, &
D’Esposito, 2004). To avoid difficulties in interpreting deactivation, we conducted a
conjunction analysis, and have reported only regions that showed activity greater than
implicit baseline and were also sensitive to direct contrasts between conditions. We found
that regions in the right superior temporal lobe and middle cingulate gyrus showed
significantly greater activity during Item trials than during Categorical-Encoding trials. No
regions showed greater activity during Categorical-Encoding trials than during Item trials.

During the probe epoch, Category and Label trials were analyzed separately, and compared
with each other and with Item trials. Not surprisingly, during the probe epoch, conditions in
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which participants viewed faces (Category and Item) had greater activity in higher order
visual processing regions than the Label condition (in which participants viewed the
Category Label, “A” or “B”). As shown in Figure 2, these included bilateral inferior
occipital and bilateral fusiform gyri. Conversely, Label trials led to greater activity than
Category and Item trials in other visual processing regions including the right cuneus /
superior occipital gyrus, and a region of the left fusiform (Label > Category only). These
differences are likely due to visual processing differences between faces and letters. In
addition to visual regions, the Category > Label contrast during the probe epoch revealed
recruitment of frontoparietal regions including the right superior medial gyrus, and the
middle and posterior cingulate, along with a region of the cerebellum (left cerebellar crus
II).

The Category versus Item contrast (see Figure 2, bottom row) was the most direct
comparison between categorization and item recognition; both conditions had similar
requirements for viewing and processing face stimuli and making same-different judgments.
The only region showing greater activity during Item trials than during Category trials was a
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region in the left middle temporal gyrus. Category trials elicited greater activity than Item
trials in executive regions of the cerebellum, frontal (middle frontal, anterior insula/inferior
frontal, and superior medial gyrus) and parietal regions (inferior parietal, angular gyrus, and
precuneus), including the salience network. Finally a conjunction analysis revealed that
motor planning regions of the SMA were recruited in all three conditions, consistent with
the similar motor response demands across the conditions.

3.2.2 CPCA—The GLM model, GC, accounted for 36.31% of the variance in the BOLD
signal. Based on inspection of the scree-plot, we extracted five components. After varimax
rotation, the first through fifth components accounted for 14.15%, 8.13%, 4.95%, 4.90%,
and 4.16% of the task-related variance, respectively.

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3.2.2.1 Component 1: Component 1 is illustrated in Figure 3; cluster coordinates are given

in Table A.2. This component was characterized by activity in the bilateral fusiform gyrus
and occipital regions, bilateral thalamus, bilateral parietal regions, bilateral inferior frontal
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gyrus and anterior insula, and bilateral medial frontal gyrus. Overall this component
overlapped with the visual, dorsal attentional, FP-CEN, and SA networks (Buckner et al.,
2011; Dosenbach, Visscher, & Palmer, 2006; Seeley et al., 2007). Inspection of the predictor
weights (Figure 3b) revealed a bimodal shape, and an ANOVA revealed a significant main
effect of condition, F(2,30) = 6.1, p < .05, η2= .01, a significant main effect of time-point,
F(17,255) = 27.29, p < .001, η2 = .56, and an interaction between condition and time-point,
F(34,510) = 9.88, p < .001, η2 = .04. This interaction was driven by differences following
the second stimulus. The slope of the predictor weight timecourse differed between the
Category and Label trials for the two consecutive time points between 2.25 and 5.25 seconds
(p<.001 for both time points), and the time points between 6.75 and 11.25 seconds (p<.05;
p<.001; p<.001) following the second stimulus. The slope also differed between the
Category and Item conditions (2.25 to 3.75, 6.75 to 8.25, 9.75 to 11.25, and 14.25 to 15.75
seconds; p<.05 for all time-points), and between the Item and Label trials (2.25 to 5.25
seconds and 6.75 to 12.75 seconds; p<.05 for all time points). Following the second
stimulus, the predictor weights associated with the Category and Item conditions showed
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greater amplitude than those of the Label condition, suggesting that the Category and Item
conditions placed greater demands on the Component 1 network than did the Label
condition. This might have been due to the visual differences between faces and labels, or
due to differences in higher-order cognitive demands.

3.2.2.2 Component 2: Component 2 loadings (Figure 4 and Table A.3) were associated with
regions largely involved in motor and visual processing, including motor and premotor
cortex (e.g., precentral and postcentral gyri, and the SMA) and visual regions (superior
occipital, lateral occipital and lingual gyri). This component largely overlaps with the visual
and somatomotor networks, along with some adjoining areas of the salience network and
dorsal and ventral attentional networks (Yeo et al., 2011). Inspection of the predictor
weights revealed a unimodal peak occurring roughly 8 seconds after the onset of the second
stimulus. An ANOVA on the predictor weights revealed a significant main effect of
condition, F(2,30) = 19.23, p< .001, η2=.03, a main effect of time-point F(17,255) = 5.00, p
< .001, η2=.18, and an interaction between condition and time-point F(34,510) = 2.9, p < .
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001, η2=.04. There was a significant difference in the slope of the predictor weight time
course between the Category and Label condition following the first peak (6.75–8.25
seconds after the first stimulus; p< .05), but the effects predominantly followed the second
stimulus. The slope between the Category and Label Trials differed between 2.25 and 3.75
seconds and 12.75 to 14.25 seconds following the second stimulus (p<.05 for both time
points). The slopes associated with the Label and Item conditions differed for the
consecutive time points between .75 and 3.75 seconds (p<.05); and the slopes of the
Category and Item conditions differed between 3.75 and 5.25 seconds (p.<.05). Overall,
following the second stimulus, the predictor weight peak was greatest for the Label
condition followed by the Item condition, and then the Category Condition.

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3.2.2.3 Component 3: The top 5% of loadings on Component 3 (see Figure 5 and Table A.
4) were associated primarily with regions within the FP-CEN network (Buckner et al., 2011;
Choi et al., 2012; Yeo et al., 2011), including extensive regions of the inferior and medial
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parietal lobe, regions of the inferior, middle, and superior frontal gyri, and subcortical
regions including the caudate nucleus and cerebellar regions. In addition, this component
included primary visual cortex and regions of the medial frontal gyrus, inferior parietal lobe,
and cerebellum associated with the default network. Inspection of the predictor weights
associated with Component 3 revealed a bimodal shape similar to that of Component 1, but
with peaks delayed several seconds in time. An ANOVA on the predictor weights (shown in
Figure 5B) revealed a significant main effect of condition, F(2,30) = 13.24, p < .001, η2=.
04, a main effect of time-point F(17,255) = 4.67, p < .001, η2=.16, and an interaction
between condition and time-point F(34,510) = 2.34, p < .001, η2=.03. This interaction was
driven by effects following the second stimulus. The predictor weight slopes were
significantly steeper in the Category condition than in the Label conditions from 6.75 to 8.25
seconds following the second stimulus (p<.01). The slope was steeper in the item condition
than the Label condition from 2.25 to 3.75 seconds (p<.05) and during the consecutive time
points from 6.75 to 9.75 seconds (p<.01; p<.05). From 8.25 to 9.75 seconds, the slope
associated with the Item condition was steeper than that of the Category condition (p < .05),
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an effect driven by a slightly shorter time-to-peak in the Category condition.

3.2.2.4 Component 4: Component 4 (see Figure 6 and Table A.5) was associated
exclusively with regions of the visual network (Yeo et al., 2011), and included regions
extending from the primary visual cortex superiorly to the precuneus and anteriorly to the
inferior temporal lobe. An ANOVA on the predictor weight timecourse indicated a
significant main effect of condition, F(1.44,21.63) = 4.55, p < .05, η2=.01, and a main effect
of time-point F(4.25,63.78) = 7.76, p < .001, η2=.26. The interaction between condition and
time-point was not significant, F(7.78,116.69) = 1.63, p >.05, η2=.02.

3.2.2.5 Component 5: The top 5% of loadings (Table A.6 & Figure 7) were associated
primarily with regions within the default mode network, including medial parietal, medial
prefrontal, middle temporal gyrus, and the hippocampus (Buckner et al., 2011; Choi et al.,
2012; Yeo et al., 2011). An ANOVA on the predictor weight time course revealed a
significant main effect of condition, F(2,30) = 12.56, p < .01, η2= .46, a significant main
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effect of time-point, F(17,255) = 19.24, p < .001, η2 = .41, and an interaction between
condition and time-point, F(34,510) = 2.5, p < .001, η2 = .14.

The interaction was driven by differences following the second stimulus: the slopes differed
between the Category and Label conditions from 2.25 to 3.75 seconds (p<.05), from 5.25
8.25 (p<.01; p<.01), and between the Category and Item conditions from .75 to 2.25 (p<.05),
and from 11.25 to 14.25 (p<.05; p<.05) seconds following the second stimulus. As predicted
based on task difficulty, we found that Category (M = −0.11, SD= .08, t(15)=3.04, p < .01,
g= .61) and Label (M= −.13, SD= .09, t(15)= 4.97, p < .001, g= 0.7) trials elicited greater
suppression across the predictor weight time series than Item trials (M= −.05, SD= .11).

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3.2.3 CPCA-masked Univariate GLM Analyses—In order to further explore how

regions within CPCA components were affected by task differences, we examined our
univariate results within masks formed by each individual component. However, it is
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important to understand the limitations of this approach. Whereas the CPCA results
highlight patterns of task-related variance shared between brain regions, univariate analyses
ignore these patterns of shared variance and consider only variance within specific clusters.
As a result, univariate results can represent several sources of overlapping variance, and
may, therefore, not closely resemble the patterns revealed by CPCA (e.g., Components 1
and 3 overlapped in regions of the bilateral precuneus, and Components 3 and 4 overlapped
in bilateral regions of the lingual gyrus). While univariate analyses can be conceptualized as
providing a view of task-related variance that slices across network variance, CPCA can be
conceptualized as providing a view that slices across the variance within specific brain
regions. The evaluation of the statistical reliability of the results from each analysis reflects
this distinction; univariate results are evaluated for each cluster separately, while CPCA
results are evaluated at the level of the hemodynamic response of the entire network.
Additionally, whereas the univariate analyses assume a canonical hemodynamic response
associated with each task epoch, CPCA uses a flexible FIR model, and is capable of
uncovering patterns of task-related variance in a data-driven manner. Because the univariate
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analyses are sensitive to matches between the network-unspecific hemodynamic response

and the canonical hemodynamic response function modeling each task epoch separately,
whereas the multivariate analysis reflect network-specific, data-driven, hemodynamic
response shapes during overall task performance, we consider the pattern of univariate
results to convey a broader, network-unspecific view of task related activity, although this
activity is restricted to matches to the canonical HRFs.

3.2.3.1 Component 1: As shown in Table A.7, consistent with the predictor weight time
course, we found that visual regions within Component 1 tended to respond preferentially to
visual stimulus features at probe; regions in the bilateral inferior occipital gyrus, bilateral
fusiform, and the left calcarine showed greater activity when a face was presented (i.e.,
during both Category and Item trials) than when a Label was presented. Interestingly,
neighboring regions within the fusiform showed greater activity during the encoding epoch
when it was necessary to encode a specific stimulus (Item trials) than when it was necessary
to categorize it (Category and Label trials), indicating that this region was sensitive to
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specific-item encoding demands. Taken together, these results indicate that these visual
regions have both feature specific processing roles, and functional roles within the task-
related salience network in responding to stimuli.

A second pattern was that frontal and parietal regions associated with FP-CEN and SA
tended to show sensitivity to categorization demands. Regions in the superior medial frontal
cortex, the left cerebellar crus 1, and the bilateral precuneus showed greater activity for
categorical encoding trials than for Item trials during encoding, and also showed greater
activity for Category trials than for Item trials during probe. Interestingly, only the left
precuneus, a region thought to be a hub in the FP-CEN (Niendam, Laird, & Ray, 2012;
Spreng, Sepulcre, Turner, Stevens, & Schacter, 2013) also showed greater activity for the
Label condition than the Item condition at probe. The right anterior insula, a key node in the

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salience network (Seeley et al., 2007), and a region of the thalamus known to be connected
with the prefrontal cortex (Behrens et al., 2003), showed greater activity for the categorical-
encoding trials than the Item trials during encoding, but showed no differences between
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conditions at probe.

3.2.3.2 Component 2: Mirroring the predictor weight timecourse, we found that visual
cortical regions of Component 2 were sensitive to visual stimulus features, as detailed in
Table A.8. Regions of the left lingual and superior occipital gyrus showed greater activity
during categorical encoding than during item trials at encoding, while the right superior
temporal lobe showed the opposite pattern (item > categorical encoding). The primary
effect, however, was that regions of the bilateral lingual gyrus, bilateral occipital gyri, and
left supramarginal gyrus were more active for Label than for Category and/or Item trials.
The univariate results, however, did not reveal similar patterns of activity within the motor
regions associated with Component 2.

3.2.3.3 Component 3: Overall, many regions within the component 3 mask in the univariate
analysis (Table A.9) showed a pattern of higher activity for both Label and Category than
Item at encoding, along with greater activity for Category than both Label and Item at probe.
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Specifically, at encoding, frontoparietal (right precuneus, left cerebellar lobule VI, and
middle cingulate) and visual (left calcarine gyrus) regions showed greater activity during
categorical-encoding trials than during Item-trials. At probe there was greater activity in FP-
CEN regions during Category trials than during Label trials (right superior medial gyrus,
posterior cingulate, bilateral cerebellar crus I), or Item trials (bilateral precuneus, left inferior
parietal lobe, right angular gyrus, bilateral superior medial gyrus, and posterior cingulate).
Overall, the univariate results were similar to the multivariate results for these regions,
despite the fact that each analysis approach captures different sources of variance. However,
the analysis approaches differed with respect to subcortical regions. Although the basal
ganglia was involved in this component overall, its activity did not differ significantly across
conditions in the univariate analysis, indicating that these regions likely played an important
role across tasks. Similarly, there was widespread cerebellar activity in the component in the
multivariate analysis, indicating cerebellar contributions to the functional network, but only
small regions of cerebellum were present in the univariate analysis.

3.2.3.4 Components 4 & 5: As was the case for visual regions within Components 1 and 2,
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visual regions associated with Component 4 were sensitive to visual stimulus features,
(Table A.10). Interestingly, there were differences in recruitment during encoding between
the working memory and categorization conditions despite both conditions sharing the same
stimulus types (faces). Activity within regions associated with Component 5 tended to be
anticorrelated with task difficulty, mirroring results based on the predictor weight
timecourse. Given the tangential nature of this component to our primary hypotheses, we do
not provide a table of these results.

4 Discussion
We compared delayed match-to-sample and delayed match-to-category tasks to investigate
how neural systems were recruited for categorization and item-specific processes across

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encoding, maintenance across a short delay, and match-mismatch decisions. In the match-to-
sample task, optimal behavior could be subserved by a strategy wherein participants
considered only the intrinsic visual features of the stimuli. The categorization task, however,
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required that participants make judgments based on latent categorical features (the category
labels). We found that categorization and item specific memory recruited five neural
networks (identified as CPCA components). Two of the components are of particular note:
Component 1, which recruited key nodes of the salience network involved in immediate
stimulus processing, and Component 3, which recruited fronto-parietal-striatal regions
linked to executive function.

The first CPCA component had three important characteristics. First, it included regions
associated with frontoparietal networks, especially the salience network (i.e., bilateral
inferior frontal and anterior insula, and dorsal anterior cingulate/ SMA; Chiong et al., 2013;
Ham, Leff, de Boissezon, Joffe, & Sharp, 2013; Menon & Uddin, 2010; Sridharan et al.,
2008) along with visual processing regions. Second, it displayed hemodynamic response
peaks occurring rapidly after stimulus onset. Third, activity in this component was
significantly higher for both conditions that required face processing (Item and Category)
than the Label conditions. These characteristics support the interpretation that this
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component was associated with the detection of behaviourally-salient events and the rapid
allocation of cognitive resources to support task demands. The salience network has been
previously associated with the coordination of large-scale brain networks to support
advantageous behavioral responding (Eckert, Menon, & Walczak, 2009; Ham et al., 2013;
Menon & Uddin, 2010; Sridharan et al., 2008). For instance, damage to the salience network
has been linked to default mode network dysfunction (Bonnelle et al., 2012), and functional
connectivity analyses have provided evidence that the salience network mediates the anti-
correlated relationship between the frontoparietal network and the default mode network
(Goulden et al., 2014; Menon, 2011; Palaniyappan et al., 2013; Sridharan et al., 2008).

Like Component 1, Component 3 was primarily associated with regions of the frontoparietal
network (Buckner et al., 2011; Choi et al., 2012; Yeo et al., 2011), but unlike Component 1,
these regions were primarily associated with the central executive network rather than the
salience network (Goulden et al., 2014; Sridharan et al., 2008). The bimodal predictor-
weight timecourse associated with Component 3 was similar to that of Component 1, but the
peaks occurred later in time following each stimulus, consistent with a greater role in more
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time-demanding processes such as those involved in decision making, rather than rapid
attentional orienting as in Component 1. Interestingly, univariate analyses indicated that
subregions within this network tended to show greater activity when categorization was
required than when it was not.

Many of the regions involved in Component 3 have been associated with categorization in
previous studies. Notably Component 3 recruited frontal lobe regions known to play an
important role in rule and category learning tasks in both monkeys (Antzoulatos & Miller,
2011; Freedman, Riesenhuber, Poggio, & Miller, 2001; Meyers et al., 2008; Wallis &
Miller, 2003) and humans (Seger & Cincotta, 2006). Additionally, there was broad activity
extending medially to laterally across the intraparietal sulcus region, which is thought to
include the human homolog of LIP, shown to be category sensitive in monkey (Fitzgerald,

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 Braunlich et al. Page 16

Freedman, & Assad, 2011; Freedman & Assad, 2006, 2009). This region is also implicated
in perceptual decision making more broadly, and is thought to subserve processes of
accumulation of information from perceptual regions that can serve as input to regions
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involved in response selection (Ploran et al., 2007; Roitman & Shadlen, 2002; Shadlen &
Newsome, 2001). Although univariate analyses indicated that subregions within this
network were preferentially activated during conditions requiring categorization, the
multivariate analyses indicated that this network was similarly recruited across tasks.

Component 3 was also the only component associated with widespread activity in the basal
ganglia, specifically in the body of the caudate, a region associated with visual
categorization in several previous studies (Lopez-Paniagua & Seger, 2011; Nomura et al.,
2007; Seger & Cincotta, 2005, 2006; Seger, Peterson, Cincotta, Lopez-Paniagua, &
Anderson, 2010). Univariate analyses, however, did not indicate that there were significant
differences in basal ganglia activity between tasks; suggesting that this region may play a
similar role across these tasks. An unexpected finding was that widespread regions of the
cerebellum were also included in Component 3. Although these regions of the cerebellum
are associated with frontal cognitive control system (Buckner et al., 2011), and are known to
contribute to higher-order cognitive processes (Balsters, Whelan, Robertson, & Ramnani,
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2012), the cerebellum is not commonly a focus of categorization and decision making
research.

Component 5 closely resembled the DMN, and showed suppressed activity during the more
cognitively demanding Category trials relative to the Item trials. This finding is in
accordance with the known anti-correlated relationship between the default mode and
frontoparietal networks (e.g., Menon & Uddin, 2010; Sridharan et al., 2008). Component 2
resembled the sensorimotor intrinsic connectivity network and showed a single peak
corresponding to the behavioral response. However, this network additionally recruited
regions within the salience and dorsal attention intrinsic connectivity networks. Many of
these regions have been previously associated with abstract motor representation and motor
preparation in functional tasks (Noppeney, Josephs, Kiebel, Friston, & Price, 2005; Rowe,
Hughes, & Nimmo-Smith, 2010), and may therefore have activity patterns that correlated
with the sensorimotor network during our task.

Component 4 was limited to regions within the visual intrinsic connectivity network, and as
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in resting state fcMRI, displayed a pattern of strong local connectivity (Yeo et al., 2011).
Our multivariate analyses, however, suggested that these regions also interacted with
different CPCA components. The bilateral lingual gyrus and cuneus, for instance, interacted
with somatomotor regions in Component 2, while posterior occipital regions were associated
with Component 4, and regions extending from the lateral occipital lobe down through the
fusiform gyri interacted with Component 1. Univariate analyses provided evidence that
visual regions were driven by stimulus type, but were insensitive to categorization demands;
fusiform regions showed greater activity when a face was presented, while medial occipital
regions showed greater activity when the category label (a single letter) was presented.

In this paper we report, for the first time, functional networks involved in the performance of
a delayed matching task that were recruited during categorization and during the processing

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 Braunlich et al. Page 17

of specific items. Most importantly, we found two different frontoparietal networks, one of
which (Component #1) acted on a faster time course, was sensitive to differences between
conditions, and included regions of the salience network in conjunction with regions
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involved in higher level visual processing. The second, (Component #3) operated on a
slower time course, and involved lateral parietal and lateral frontal regions, as well as the
basal ganglia, all regions previously individually associated with categorization and
decision-making.

Acknowledgements
This research was supported in part by NIH Grant R01 MH079182 to CAS, and by NSF REU to Edward DeLosh.
We would like to thank Matthew Rhodes and Danielle Sitzman for providing the face stimuli used in this study.

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Highlights
- Using fMRI, we compared delayed match-to-sample and delayed match-to-
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category tasks

- We identified five intrinsic networks active during the tasks

- The frontoparietal central-executive network played a key role in

categorization

- The frontoparietal salience network was sensitive to rapid visual feature

processing in both tasks

- Visual, motor, and default mode networks were activated similarly for both
tasks
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Figure 1.
During each trial, participants first saw a cue (Item condition: “Match the Specific Face”;
Category and Label conditions: “Match the Category”) for 1.5 sec., they then saw the first
stimulus (1.5 sec). After a brief delay (9 sec) they saw a second stimulus (3 sec). In the
Category and Item conditions, the second stimulus was a face. In the Label condition, the
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second stimulus was the category label (“A” or “B”). After three seconds, a match mismatch
cue was presented, and participants had to indicate whether the second stimulus matched the
first. Trials were separated by a jittered ITI (1.5–9 sec).

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Figure 2.
Whole brain univariate analyses: activity differing between conditions within individual trial
epochs (Encoding and Probe). TOP FIGURE: Encoding epoch. Red: Categorical Encoding
(Category and Label trials) greater than Item; Blue: Item greater than Categorization
Encoding. BOTTOM FIGURES: Probe epoch. TOP: Green: Label greater than Item; Blue:
Item greater than Label. MIDDLE: Green: Label greater than Category; Red: Category
greater than Label. BOTTOM: Red: Category greater than Item. Blue: Item greater than
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Category. Regions of activity are overlaid on the average normalized anatomical image
across subjects. For each contrast, we generated maps at an uncorrected threshold of p <
0.001 and corrected for multiple comparisons using the topological false-discovery rate (q
< .05; Chumbley & Friston, 2009).

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Figure 3.
Component 1. Note the recruitment of regions involved in the salience network (inferior
frontal/anterior insula and anterior cingulate) along with visual processing regions (fusiform
gyrus and occipital lobe). A) The top 5% of component loadings overlaid on the MNI
template provided by MRIcron (3d renderings, top) and the average structural image (slices,
bottom). B) Predictor weight timecourse across peristimulus time. Error bars represent the
standard error of the mean. Vertical lines indicate onsets of visual stimuli.
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Figure 4.
Component 2. Note the recruitment of sensorimotor and premotor regions. A) The top 5% of
component loadings overlaid on the MNI template provided by MRIcron (3d renderings,
top) and an averaged structural image (slices, bottom). B) Predictor weight timecourse. Error
bars represent the standard error of the mean. Vertical lines indicate onsets of visual stimuli.
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Figure 5.
Component 3. Note the recruitment of FP-CEN regions including the lateral prefrontal
cortex and intraparietal sulcus, along with the cerebellum and caudate. A) The top 5% of
component loadings overlaid on the MNI template provided by MRIcron (3d renderings,
top) and an averaged structural image (slices, bottom). B) Predictor weight timecourse. Error
bars represent the standard error of the mean. Vertical lines indicate onsets of visual stimuli.
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Figure 6.
Component 4. Note the recruitment of visual processing regions. A) The top 5% of
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component loadings overlaid on the MNI template provided by MRIcron (3d renderings,
top) and an averaged structural image (slices, bottom).. B) Predictor weight timecourse.
Error bars represent the standard error of the mean. Vertical lines indicate onsets of visual
stimuli.
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Figure 7.
Component 5. Note the recruitment of default mode network regions. A) The predictor
weight timecourse. B) The top 5% of component loadings overlaid on the MNI template
provided by MRIcron (3d rendering, left) and an averaged structural image (slices on the
right). We changed the color map to cool colors to emphasize that Component 5 was
anticorrelated with cognitive demands.
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Table A.1

Univariate results for the encoding, delay and probe epochs. As Category and Label trials were methodologically identical during encoding and delay,
they were combined into a single Categorical-Encoding condition.

Peak MNI
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Contrast Coordinates
Voxels mm^3 x y z Region BA Network

Encoding
  Categorical Encoding >item
1185 9480
−2 −90 −4 Left Calcarine Gyrus 17 VS
2 −76 8 Right Calcarine Gyrus 18 VS
731 5848
12 −64 40 Right Precuneus 7 FP-CEN
−4 −64 44 Left Precuneus 7 DMN
357 2856 −4 −28 28 Middle Cingulate 23 SA
152 1216 12 6 14 Right Caudate Nucleus FP-CEN
106 848 −6 −18 10 Left Thalamus
99 792 −12 −80 −22 Left Cerebelar Lobule VI (hem) 18 FP-CEN
79 632 4 28 42 Right Superior Medial Gyrus 6 SA
75 600 30 22 −4 Right Inferior Frontal / Anterior Insula 47 SA
  Item> Categorical Encoding
451 3608 −42 −66 −12 Left Inferior Occipital Gyrus 19 VS
284 2272

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−46 −8 −30 Left Inferior Temporal Gyrus 20 DMN
−24 −2 −22 Left Amygdala 28
−36 −2 −20 Superior Temporal Pole 36
−22 −12 −18 Left Hippocampus 35
177 1416 −4 52 −16 Left Rectal Gyrus 11 DMN
124 992 22 −10 −14 Right Hippocampus
111 888 44 −62 −6 Right Inferior Temporal Gyrus 36 VS/DA
91 728 −42 28 −16 Left Inferior Frontal Gyrus 38 DMN
87 696 −44 −46 −14 Left Inferior Temporal Gyrus 37 DA
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Peak MNI
Contrast Coordinates
Voxels mm^3 x y z Region BA Network

  Conjunction (Item> Implicit Baseline & Categorical Encoding Trials > Implicit Baseline)
100 800 40 −73 −29 Right Cerebellar Crus I FP-CEN/DMN
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Delay
  Conjunction (Item>Categorical Encoding & Item > Implicit Baseline)
131 1048 56 −18 10 Right Superior Temporal SM
52 416 −6 −18 44 Middle Cingulate SM/VA
Probe
  Item > Label
1325 10600
42 −82 −2 Right Inferior Occipital Gyrus 19 VS
42 −74 −10 Right Inferior Occipital Gyrus 19 VS
42 −46 −16 Right Fusiform Gyrus 37 DA
937 7496
−26 −94 −4 Left Inferior Occipital Gyrus 18 VS
−38 −84 −14 Left Fusiform Gyrus 19 VS
  Label > Item
191 1528 16 −76 24 Right Cuneus 18 VS
135 1080 −50 −50 6 Left Middle Temporal Gyrus/ Lateral Occipital 21 VA
103 824 −14 −88 24 Left Superior Occipital Gyrus 18 VS
84 672 −4 −68 46 Left Precuneus 7 FP-CEN
  Label > Category

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513 4104
18 −88 18 Right Superior Occipital Gyrus 18 VS
18 −78 24 Right Cuneus 19 VS
325 2600 −56 −56 10 Left Middle Temporal Gyrus 37 VA
298 2384 −12 −90 24 Left Superior Occipital Gyrus 18 VS
156 1248 52 −32 16 Right Superior Temporal Gyrus 41 SM
120 960 −38 −18 −8 Left Insula (Id1) 20 VA
112 896 −54 −2 −10 Left Superior Temporal Gyrus 22 DMN
102 816 10 −70 −6 Right Lingual Gyrus 18 VS
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Peak MNI
Contrast Coordinates
Voxels mm^3 x y z Region BA Network
81 648 0 −24 60 Left Paracentral Lobule 6 SM
80 640 68 −30 30 Right SupraMarginal Gyrus 2 VA
Braunlich et al.

77 616 −6 −70 −2 Left Lingual Gyrus 18 VS

71 568 −32 0 12 Left Insula 48 VA
67 536 −62 −24 16 Left SupraMarginal Gyrus 42 VA
63 504 60 −58 20 Right Superior Temporal Gyrus 21 DMN
62 496 −32 −38 −14 Left Fusiform Gyrus 37 VS
  Category > Label
1629 13032
42 −88 −2 Right Inferior Occipital Gyrus 19 VS
42 −74 −10 Right Inferior Occipital Gyrus 19 VS
42 −52 −16 Right Fusiform Gyrus 37 VS
32 −92 −2 Right Inferior Occipital Gyrus 18 VS
1114 8912
−26 −94 −4 Left Inferior Occipital Gyrus 18 VS
−38 −82 −12 Left Inferior Occipital Gyrus 19 VS
211 1688
2 28 46 Right Superior Medial Gyrus 8 SA
8 30 30 Right Middle Cingulate Cortex 32 SA
148 1184 0 −44 20 Posterior Cingulate DMN
136 1088 −8 −74 −38 Left Cerebellar Lobule VIIb (Hem) DA

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97 776 −6 −32 28 Posterior Cingulate 23 FP-CEN
97 776 −2 −34 −6 Superior Colliculus
  Item > Category
122 976 −58 −2 −18 Left Middle Temporal Gyrus 21 DMN
  Category > Item
863 6904
−6 −72 44 Left Precuneus 7 FP-CEN
8 −70 44 Right Precuneus 7 FP-CEN
758 6064 −34 −56 46 Left Inferior Parietal Lobule 7 FP-CEN
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Peak MNI
Contrast Coordinates
Voxels mm^3 x y z Region BA Network
555 4440
0 28 42 Left Superior Medial Gyrus 6 SA
Braunlich et al.

−2 14 46 Left preSMA 6 SA
312 2496
−30 6 60 Left Middle Frontal Gyrus 6 SA
−48 6 44 Left Precentral Gyrus 44 FP-CEN
252 2016 34 −60 48 Right Angular Gyrus 7 FP-CEN
91 728 −4 −38 24 Left Posterior Cingulate Cortex 26 FP-CEN
76 608 −44 30 32 Left Middle Frontal Gyrus 45 FP
71 568 −36 18 −2 Left Inferior Frontal / Anterior Insula 47 SA
59 472 −8 −74 −26 Left Cerebellar Lobule VI (Hem) FP-CEN
  Conjunction(Category> Implicit, Label > Implicit, & Item> Implicit)
121 968 −1 2 50 Bilateral preSMA 6 SA
39 312 −16 −14 15 Left Thalamus
26 208 19 −13 16 Right Thalamus

VS=Visual; SM=Somatomotor; DA=Dorsal Attention; FP-CEN=Central Executive; SA=Salience; DMN=Default Mode Network. Cluster volumes smaller than 10 voxels have been omitted.

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Table A.2

Cluster volumes and peak coordinates for the rotated top 5% of Component 1 loadings.

Peak MNI
Coordinates
Braunlich et al.

Voxels mm^3 x y z Region BA Network

3755 30040
32 −90 0 Right Middle Occipital Gyrus 18 VS
42 −56 −20 Right Fusiform Gyrus 37 VS
40 −78 −12 Right Inferior Occipital Gyrus 19 VS
32 −60 46 Right Angular Gyrus 7 DA/FP-CEN
30 −52 46 Right Inferior Parietal Lobule 7 DA/FP-CEN
2916 23328
−32 −90 −6 Left Middle Occipital Gyrus 19 VS
−38 −82 −12 Left Inferior Occipital Gyrus 19 VS
−40 −72 −14 Left Fusiform Gyrus 19 VS
−38 −56 −20 Left Fusiform Gyrus 37 VS
1021 8168
−28 −70 26 Left Middle Occipital Gyrus 19 VS/DA
−28 −62 48 Left Superior Parietal Lobule 7 DA
−30 −48 42 Left Inferior Parietal Lobule 40 DA
627 5016
−4 10 50 Left preSMA / Medial Frontal Gyrus 6 SA
4 16 46 Right preSMA / Medial Frontal Gyrus 6 SA

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525 4200
48 12 30 Right Inferior Frontal Gyrus (p. Opercularis)) 44 DA
44 6 34 RightPrecentral Gyrus 44 DA
48 32 18 Right Inferior Frontal Gyrus (p. Triangularis) 45 SA
113 904
12 −64 42 Right Precuneus 7 FP-CEN
110 880
−46 4 34 Left Precentral Gyrus 44 DA
−46 12 26 Left Inferior Frontal Gyrus (p. Triangularis) 44 SA
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Peak MNI
Coordinates
Voxels mm^3 x y z Region BA Network
95 760 32 24 −4 Right Anterior Insula 47 SA
89 712 −10 −72 44 Left Precuneus 7 FP-CEN
Braunlich et al.

71 568 −6 −76 −24 Left Cerebellar Lobule VI FP-CEN

68 544 10 −14 8 Right Thalamus SA
63 504 −2 −30 26 Middle/Posterior Cingulate 23 FP-CEN
60 480 −10 −16 8 Left Thalamus SA
49 392 −30 24 −2 Left Anterior Insula 47 SA
35 280 −48 0 50 Left Precentral Gyrus 6 DA/SM

VS=Visual; SM=Somatomotor; DA=Dorsal Attention; FP-CEN=Central Executive; SA=Salience. Cluster volumes smaller than 10 voxels have been omitted.

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Table A.3

Table A.3. Cluster volumes and peak coordinates for the rotated top 5% of Component 2 loadings.

Peak MNI
Coordinates
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Voxels mm^3 x y z Region BA Network

3170 25360
−56 −22 40 Left Inferior Parietal Lobule 3 DA
−58 −20 18 Left Postcentral Gyrus 48 SM
−38 −40 58 Left Postcentral Gyrus 2 SM
−62 −36 24 Left SupraMarginal Gyrus 48 SA
−46 −36 22 Left Superior Temporal Gyrus 48 SM
−42 −12 52 Left Precentral Gyrus 6 SM
2679 21432
56 −18 46 Right Postcentral Gyrus 1 SM
48 −20 18 Right Rolandic Operculum 48 SM
46 −26 58 Right Postcentral Gyrus 1 SM
64 −40 22 Right Superior Temporal Gyrus 22 SA
64 −24 24 Right SupraMarginal Gyrus 48 SA
38 −26 62 Right Precentral Gyrus 4 SM
1243 9944
2 −4 52 Right preSMA 6 SM
−2 6 40 Left Middle Cingulate Cortex 24 SA
12 −28 44 Right Middle Cingulate Cortex SA

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1085 8680
−6 −72 −2 Left Lingual Gyrus 18 VS
8 −68 −2 Right Lingual Gyrus 18 VS
−20 −60 −14 Left Cerebellum Lobule VI 19
20 −60 −12 Right Fusiform Gyrus 19 VS
406 3248
−50 −2 0 Left Insula 48 VA/SM
−44 −4 10 Left Insula 48 VA/SM
−56 8 20 Left Inferior Frontal Gyrus 44 SA
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Peak MNI
Coordinates
Voxels mm^3 x y z Region BA Network
306 2448 −8 −94 14 Left Cuneus 18 VS
277 2216
Braunlich et al.

44 2 6 Right Insula 48 SA
58 10 8 Right Inferior Frontal Gyrus 44 SA
243 1944 14 −90 20 Right Cuneus 18 VS
54 432 20 −4 68 Right Superior Frontal Gyrus 6 VA/SM
45 360 50 −66 8 Right Lateral Occipital Gyrus 37 VS
36 288 −12 −26 40 Left Middle Cingulate Cortex SA
32 256 −52 −74 8 Left Lateral Occipital Gyrus 19 VS
18 144 12 −18 6 Right Thalamus

VS=Visual; SM=Somatomotor; DA=Dorsal Attention; SA=Salience. Cluster volumes smaller than 10 voxels have been omitted.

Neuroimage. Author manuscript; available in PMC 2016 February 15.

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Table A.4

Cluster volumes and peak coordinates for the rotated top 5% of Component 3 loadings.

Peak MNI
Coordinates
Braunlich et al.

Voxels mm^3 x y z Region BA Network

5040 40320
4 −72 46 Right Precuneus 7 FP-CEN
−4 −66 58 Left Precuneus 7 DA
−38 −58 38 Left Angular Gyrus 40 FP-CEN
−32 −72 42 Left Inferior Parietal Lobule 7 FP-CEN
38 −64 48 Right Angular Gyrus 7 FP-CEN
−44 −58 48 Left Inferior Parietal Lobule 39 FP-CEN
40 −56 42 Right Inferior Parietal Lobule 40 FP-CEN
0 −34 24 Bilateral Posterior Cingulate 23 FP-CEN
2774 22192
12 −88 −26 Right Cerebellum Crus I DMN
−4 −82 −22 Left Cerebellum Lobule VI FP-CEN
−8 −88 −26 Left Cerebellum Crus I FP-CEN
44 −72 −30 Right Cerebellum Crus I DMN
6 −94 −10 Right Calcarine Gyrus VS
−40 −78 −30 Left Cerebellum Crus I DMN
1061 8488
0 −8 10 Thalamus

Neuroimage. Author manuscript; available in PMC 2016 February 15.

18 4 22 Right Caudate Nucleus FP-CEN
−10 −6 12 Thalamus
−18 −14 24 Left Caudate Nucleus FP-CEN
174 1392 46 38 28 Right Middle Frontal Gyrus 45 FP-CEN
165 1320 −38 58 2 Left Superior Frontal Gyrus 10 FP-CEN
158 1264
0 34 38 Bilateral Superior Medial Gyrus 32 FP-CEN/DMN
0 24 48 Bilateral preSMA 6 FP-SA
77 616 −12 −92 −2 Left Calcarine Gyrus 17 VS
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Peak MNI
Coordinates
Voxels mm^3 x y z Region BA Network
71 568 −50 18 34 Left Middle Frontal Gyrus 45 FP-CEN
32 256 34 62 10 Right Superior Frontal Gyrus 10 FP-CEN
Braunlich et al.

VS= Visual; SM=Somatomotor; DA=Dorsal Attention; FP-CEN=Central Executive; SA=Salience; = DMN=Default Mode Network. Cluster volumes smaller than 10 voxels have been omitted.

Neuroimage. Author manuscript; available in PMC 2016 February 15.

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Table A 5

Cluster volumes and peak coordinates for the rotated top 5% of Component 4 loadings.

Peak MNI
Coordinates
Braunlich et al.

Voxels mm^3 x y z Region BA Network

9600 76800
0 −80 8 Left Calcarine Gyrus 17 VS
−40 −88 −2 Left Middle Occipital Gyrus 19 VS
−38 −80 −14 Left Fusiform Gyrus 19 VS
16 −94 −2 Right Calcarine Gyrus 17 VS
0 −84 26 Left Cuneus 18 VS
10 −64 8 Right Lingual Gyrus 17 VS
−12 −62 4 Left Calcarine Gyrus 17 VS
−36 −88 −12 Left Inferior Occipital Gyrus 19 VS
−28 −68 −18 Left Cerebellum 19 VS
6 −80 42 Right Cuneus 7 VS
−4 −82 42 Left Superior Occipital Gyrus 19 VS
36 −68 −20 Right Fusiform Gyrus 19 VS

VS=Visual. Cluster volumes smaller than 10 voxels have been omitted.

Neuroimage. Author manuscript; available in PMC 2016 February 15.

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Table A.6

Cluster volumes and peak coordinates for the rotated top 5% of Component 5 loadings.

Peak MNI
Coordinates
Braunlich et al.

Voxels mm^3 x y z Region BA Network

4231 33848
2 54 2 Right Superior Medial Gyrus 10 DMN
−2 52 −12 Left Mid Orbital Gyrus 11 DMN
0 62 6 Left Superior Medial Gyrus DMN
6 52 16 Right Superior Medial Gyrus 32 DMN
−8 42 −8 Left Mid Orbital Gyrus 11 DMN
−12 66 22 Left Superior Frontal Gyrus 10 DMN
1770 14160
−4 −56 18 Left Precuneus 7 DMN
8 −52 8 Right Precuneus 17 DMN
1301 10408
−50 −68 24 Left Angular Gyrus 39 DMN
−62 −50 4 Left Middle Temporal Gyrus 21 DMN
523 4184 50 −10 −16 Right Middle Temporal Gyrus 20 DMN
514 4112 52 −56 18 Right Middle Temporal Gyrus 21 DA/DMN
424 3392
−58 −2 −18 Left Middle Temporal Gyrus 21 DMN
−54 −14 −14 Left Middle Temporal Gyrus 20 DMN

Neuroimage. Author manuscript; available in PMC 2016 February 15.

−64 −18 −8 Left Middle Temporal Gyrus 21 DMN
258 2064 26 −20 −18 Hipp (SUB) 20
186 1488 −42 22 −20 Left Temporal Pole) 38 DMN
134 1072 −24 −22 −18 Hipp (SUB) 30
36 288 −20 −86 −10 Left Fusiform Gyrus 18 VS

VS=Visual; DA=Dorsal Attention; DMN=Default Mode Network. Cluster volumes smaller than 10 voxels have been omitted.
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Table A.7

Cluster volumes and MNI peak coordinates regions within Component 1 that showed significant differences between conditions, based on the full-brain
univariate statistics.

Peak MNI
Braunlich et al.

Coordinates
Voxels mm^3 x y z Region BA Network

Encoding
  Categorical Encoding > Item
103 824 12 −64 40 Right Precuneus 7 FP-CEN
55 440 −4 −28 28 Bilateral Posterior Cingulate 23 FP-CEN
54 432 32 23 −4 Right Anterior Insula 47 SA
51 408 4 28 42 Right Superior Medial Gyrus 6 SA
35 280 −10 −78 −24 Left Cerebellar Lobule VI FP-CEN
35 280 −4 −70 46 Left Precuneus 7 FP-CEN
32 256 −6 −18 10 Left Thalamus
  Item > Categorical Encoding
275 2200 −42 −66 −12 Left Inferior Occipital Gyrus 19 VS
50 400 44 −62 −6 Right Inferior Temporal Gyrus 37 VS
24 192 −42 −46 −16 Left Inferior Temporal Gyrus 37 DA
Probe
  Item > Label
1271 10168
42 −82 −2 Right Inferior Occipital Gyrus 19 VS

Neuroimage. Author manuscript; available in PMC 2016 February 15.

42 −74 −10 Right Inferior Occipital Gyrus 19 VS
42 −46 −16 Right Fusiform Gyrus 37 DA/VS
851 6808
−26 −94 −4 Left Inferior Occipital Gyrus 18 VS
−38 −84 −14 Left Fusiform Gyrus 19 VS
  Label > Item
11 88 −6 −68 46 Left Precuneus 7 FP-CEN
  Category > Label
1471 11768
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Peak MNI
Coordinates
Voxels mm^3 x y z Region BA Network
42 −88 −2 Right Inferior Occipital Gyrus 19 VS
42 −52 −16 Right Fusiform Gyrus 37 VS/DA
Braunlich et al.

32 −92 −2 Right Inferior Occipital Gyrus 18 VS

42 −50 −28 Right Cerebellar Lobule VI (Hem) 37 SA
999 7992
−26 −94 −4 Left Inferior Occipital Gyrus 18 VS
−38 −82 −12 Left Inferior Occipital Gyrus 19 VS
−40 −58 −24 Left Cerebellar Lobule VI 37 SA
96 768 2 28 46 Bilateral Superior Medial Gyrus 8 SA
24 192 −4 −32 28 Bilateral Posterior Cingulate 23 FP-CEN
  Category > Item
412 3296 −34 −56 46 Left Inferior Parietal Lobule 7 FP-CEN
309 2472
0 28 42 BIlateral Superior Medial Gyrus 32 SA
−2 14 46 Left preSMA 6 SA
−2 6 58 Left preSMA 6 SA
190 1520 34 −60 48 Right Angular Gyrus 7 FP-CEN
103 824 8 −70 44 Right Precuneus 7 FP-CEN
75 600 −6 −72 44 Left Precuneus 7 FP-CEN
30 240 −2 −36 24 Posterior Cingulate 23 FP-CEN

VS=Visual; SM=Somatomotor; DA=Dorsal Attention; FP-CEN=Central Executive; SA=Salience; DMN=Default Mode Network. Cluster volumes smaller than 10 voxels have been omitted.

Neuroimage. Author manuscript; available in PMC 2016 February 15.

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Table A 8

Cluster volumes and peak coordinates for the rotated top 5% of Component 4 loadings.

Peak MNI
Coordinates
Braunlich et al.

Voxels mm^3 x y z Region BA Network

9600 76800
0 −80 8 Left Calcarine Gyrus 17 VS
−40 −88 −2 Left Middle Occipital Gyrus 19 VS
−38 −80 −14 Left Fusiform Gyrus 19 VS
16 −94 −2 Right Calcarine Gyrus 17 VS
0 −84 26 Left Cuneus 18 VS
10 −64 8 Right Lingual Gyrus 17 VS
−12 −62 4 Left Calcarine Gyrus 17 VS
−36 −88 −12 Left Inferior Occipital Gyrus 19 VS
−28 −68 −18 Left Cerebellum 19 VS
6 −80 42 Right Cuneus 7 VS
−4 −82 42 Left Superior Occipital Gyrus 19 VS
36 −68 −20 Right Fusiform Gyrus 19 VS

VS=Visual. Cluster volumes smaller than 10 voxels have been omitted.

Neuroimage. Author manuscript; available in PMC 2016 February 15.

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Table A.9

Cluster volumes and peak MNI coordinates regions within Component 3 that showed significant differences between conditions, based on the full-brain
univariate statistics.

MNI Peak
Braunlich et al.

Coordinates
Voxels mm^3 x y z Region BA Network

Encoding
  Categorical Encoding > Item
599 4792
12 −64 40 Right Precuneus 7 FP-CEN
−4 −64 44 Left Precuneus 7 DMN
161 1288 −4 −28 28 Left Middle Cingulate/Medial Frontal Gyrus 23 SA
65 520 −12 −80 −22 Left Cerebellar Lobule VI (Hem) 18 FP-CEN
47 376 4 −88 −4 Left Calcarine Gyrus 17 VS
16 128 −8 −88 −6 Left Calcarine Gyrus 18 VS
14 112 −6 −8 8 Left Thalamus
Probe
  Label > Item
67 536 −4 −68 46 Left Precuneus 7 FP-CEN
Category> Label
40 320 2 26 46 Right Superior Medial Gyrus SA
33 264 −4 −34 28 Bilateral Posterior Cingulate 23 FP-CEN
33 264 36 −62 −28 Right Cerebellar Lobule VIIa Crus I FP-CEN

Neuroimage. Author manuscript; available in PMC 2016 February 15.

30 240 2 −42 22 Bilateral Posterior Cingulate Cortex 26 DMN
24 192 −8 −80 −30 Left Cerebellar Lobule VIIa Crus I FP-CEN
  Category > Item
743 5944
−6 −72 44 Left Precuneus 7 FP-CEN
8 −70 44 Right Precuneus 7 FP-CEN
556 4448 −34 −56 46 Left Inferior Parietal Lobule 7 FP-CEN
168 1344 34 −60 46 Right Angular Gyrus 7 FP-CEN
85 680 0 28 42 Bilateral Superior Medial Gyrus 32 SA
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MNI Peak
Coordinates
Voxels mm^3 x y z Region BA Network
68 544 −4 −38 24 Posterior Cingulate Cortex 26 FP-CEN
14 112 −8 −84 −34 Left Cerebellar Lobule VIIa Crus II (Hem) FP-CEN
Braunlich et al.

11 88 −50 22 34 Left Middle Frontal Gyrus 45 FP-CEN

VS=Visual; SM=Somatomotor; DA=Dorsal Attention; FP-CEN=Central Executive; SA=Salience; DMN=Default Mode Network. Cluster volumes smaller than 10 voxels have been omitted.

Neuroimage. Author manuscript; available in PMC 2016 February 15.

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Table A.10

Cluster volumes and peak MNI coordinates regions within Component 4 that showed significant differences between conditions, based on the full-brain
univariate statistics.

MNI Peak
Braunlich et al.

Coordinates
Voxels mm^3 x y z Region BA Network

Encoding
  Categorical Encoding > Item
1040 8320
−2 −90 −4 Left Calcarine Gyrus 17 VS
2 −76 8 Right Calcarine Gyrus 18 VS
16 −76 10 Right Calcarine Gyrus 17 VS
−14 −62 2 Left Lingual Gyrus 17 VS
20 160 −12 −74 −10 Left Lingual Gyrus 18 VS
  Item > Categorical Encoding
138 1104
−42 −66 −12 Left Inferior Occipital Gyrus 19 VS
−46 −74 −8 Left Inferior Occipital Gyrus 19 VS
Probe
  Item>Label
764 6112
−26 −94 −4 Left Inferior Occipital Gyrus 18 VS
−38 −84 −14 Left Fusiform Gyrus 19 VS

Neuroimage. Author manuscript; available in PMC 2016 February 15.

−14 −96 −12 Left Lingual Gyrus 18 VS
436 3488 42 −82 −2 Right Inferior Occipital Gyrus 19 VS
  Label > Item
56 448 −14 −88 24 Left Superior Occipital Gyrus 18 VS
22 176 12 −86 16 Right Cuneus 18 VS
  Category > Label
861 6888
−26 −94 −4 Left Inferior Occipital Gyrus 18 VS
−24 −82 −16 Left Lingual Gyrus 18 VS
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MNI Peak
Coordinates

View publication stats

Voxels mm^3 x y z Region BA Network
591 4728
42 −88 −2 Right Inferior Occipital Gyrus 19 VS
Braunlich et al.

22 −96 −4 Right Calcarine Gyrus 17 VS

  Label > Category
125 1000 −12 −90 26 Left Cuneus 18 VS
165 1320 6 −86 18 Right Cuneus 18 VS
82 656 10 −70 −6 Right Lingual Gyrus 18 VS
59 472 −6 −70 −2 Left Lingual Gyrus 18 VS

VS=Visual; SM=Somatomotor; DA=Dorsal Attention; FP-CEN=Central Executive; SA=Salience; DMN=Default Mode Network. Cluster volumes smaller than 10 voxels have been omitted.

Neuroimage. Author manuscript; available in PMC 2016 February 15.

Page 48