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Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.
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Hum Brain Mapp. 2010 March ; 31(3): 391397. doi:10.1002/hbm.20873.
Neural Decoding of Goal Locations in Spatial Navigation in

Humans with fMRI
Paul F. Rodriguez
Department of Radiology University of California at San Diego 8950 Villa La Jolla Drive C101 La
Jolla, CA 92037 p4rodriguez@ucsd.edu ph. 858-246-0994
Abstract
We demonstrate that multivoxel pattern analysis can be used to decode place-related information
in fMRI. Subjects performed a working memory version of the Morris water maze task in a virtual
environment with a single wall cue. The voxel data that corresponds to when subjects were located
at the goal was extracted for seven regions implicated in spatial navigation, and then used to train
a pattern classifier based on partial least squares. Using a leave-one-out (LOO) test procedure,
goal locations at E, W, N positions (relative to the cue as S) were predicted significantly better
than a nave classifier for voxels in medial prefrontal cortex, hippocampus, and inferior parietal
cortex. Prediction with voxels from other regions involved in navigation was also better than a
nave classifier, which raises the possibility that goal-location information is widely disseminated
among the navigation network. It turns out that predictive capability of all regions combined
significantly decreases, relative to no change, only when voxel data from the hippocampus is left
out. This implies that the hippocampus contains some unique information that identifies goal
locations, whereas other regions contain information that also identifies goal locations but is more
redundant. Classification of goal locations is an important step toward decoding a variety of place-
related information in spatial navigation with fMRI.
Keywords
hippocampus; prefrontal cortex; multivoxel pattern analysis; place coding; cognitive map
Introduction
FMRI studies of spatial navigation using virtual environments have consistently found a
network of activated regions that is compatible with regions identified in the animal
literature (e.g. Aguirre, et al., 1996; Maguire et al., 1998; Iarria et al. 2003; Hartley et al.,
2003; Voermans et al., 2004; Wolbers & Buchel, 2005; Spiers & Maguire, 2006; Wolbers, et
al., 2007; Shipman & Astur, 2008). This network includes posterior/inferior parietal regions,
retrosplenial cortex, and medial temporal regions, including hippocampus and
parahippocampal regions.
Multivoxel pattern analysis has shown that fMRI activity may contain predictive
information for a variety of sensory and cognitive processes, such as for object categories,
stimulus orientation, and intentions (Haxby, et al., 2001; Hanson, et al, 2004; Kamitani, et
al., 2005; Polyn, et al., 2005; Haynes, et al., 2007). Given the growing fMRI literature on
spatial navigation, and the well-studied properties of neurons in animal spatial navigation,
the ability to predict place-related information would also be significant. Here we use a
visually austere virtual environment to test if goal locations can also be predicted from a
pattern of fMRI activity.
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Single cell recordings in rodents show that information about goal locations are prominently
represented by neurons in the hippocampus and prefrontal cortex. For example, one study
found an accumulation of place fields at goal locations in the hippocampus, even when those
locations are unmarked (Hollup et al., 2001; see also Hok, et al, 2005). Another study found
place fields in the prefrontal cortical regions of rats that prominently represented goal
locations distinct from reward sites (Hok, et al., 2007). In a study with pre-operative
humans, goal information was primarily found prefrontal cortex, hippocampus, and
parahippocampus (Eckstrom, et al., 2003). It is also the case that hippocampal place-fields
do not show a topology and that the reconstruction of location depends on an ensemble of
neurons (Zhang, et al., 1998; Redish, et al., 2001). Based on these studies, we hypothesized
that goal locations can be discriminated from the distributed pattern of voxel activity in
medial temporal lobe and prefrontal cortex. We also plan to test other regions in the spatial
network because encoding goal locations are only one aspect of route planning and
execution in goal-oriented navigation (Poucet, et al., 2004).
The task we used is a version of the Morris water maze that relies on spatial working
memory (Morris, et al., 1986). Subjects freely navigated with a MRI-compatible joystick in
a circular arena with one small red square on the wall providing the sole landmark. In an
encoding phase, starting from a random location and orientation, subjects picked up a coin
by moving to a yellow disk on the arena floor. After a short delay with a blank screen,
subjects were randomly repositioned and reoriented, and they then retrieved the now
invisible coin by returning to the vicinity of its prior location. Successful retrieval requires
integration of joystick motion, optic-flow cues, wall views, and allocentric spatial memory.
Our results show that goal locations can be predicted significantly better than a baseline
model for voxels in medial prefrontal cortex, hippocampus, and inferior parietal cortex. We
also found best performance when data from all regions were combined. We then tried
removing voxel data one region at a time from the combined dataset and found that
prediction performance decreased significantly less than zero only when removing voxel
data from the hippocampus. Thus, it seems that goal location information may be distributed
or broadcast throughout many regions of the navigation network, but the hippocampus
contributes something more unique or specialized. These results are the first step toward
decoding a variety of place-related information in spatial navigation with fMRI.
Materials and Methods

Spatial Environment and Trial Detail
The virtual environment was coded in C++ with the OpenGL library (www.opengl.org) for
rendering 3-D graphics. The environment consisted of a circular arena with a radius of 100
arbitrary units and a wall 16 units high (Figure 1). A fixation point indicating view and
heading was placed mid-height on the wall, and two parallel horizontal lines were drawn on
the wall to help perspective. Dots were randomly placed on the floor, and shuffled between
trial phases, to provide optic-flow cues.
Subjects moved around freely with a joystick, which in the scanner was an MRI compatible
joystick (Current Designs: www.curdes.com). Movement consisted of forward, backward,
left, and right steps. To simulate natural movement, the forward motion was allowed to be
relatively quick and slightly graded (speed range of about 50-75 units/sec), and the other
lateral directions were limited to 10 units in any one continuous movement at a constant
speed (20 units/sec). The left/right buttons of the joystick controlled heading. One button
press turned 1 degree and holding the button turned 90 degrees/sec. If subjects turned while
simultaneously moving forward, then the rotation would be slow so that they would veer
slightly as they moved.

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Each trial consisted of an encoding and a test phase. In the encoding phase, the subject
started in a random orientation and position, and then moved to a visible yellow coin located
on the floor within 15 seconds (Figure 1). When the subject was within 15 units of the coin
movement was frozen and the word found appeared centrally on the screen for 1.5
seconds. The screen was then blanked out, except for a fixation cross, for a jittered delay
period (6 second mean, range 4-10 seconds). The test phase started with the subject again
randomly repositioned and reoriented. The coin was not visible, and the subject had to move
to the location where the coin was previously. There was a time limit of 20 seconds in the
test phase to limit searching. The subject had to be near the correct location (within 35 units)
for at least 2 seconds for a successful retrieval. If the coin was retrieved (not retrieved) the
word recalled (not recalled) appeared on the screen for 1.5 seconds, the screen was
blanked out and replaced by a fixation cross for another 4-10 second jittered delay time
before the next trial.
There was one red square on the wall (12 units wide) that covered only about 7 degrees of
the wall. The circular arena with one wall cue is patterned after Muller et al., (1987).
Subjects were pre-trained to retrieve coins all over the arena, but during fMRI scanning the
coin was always placed East, West, North (E, W, N) relative to the red square (South), and
10 units in front of the wall. The main reason we limited goals to three locations was so that
we can perform neural decoding of place information and have sufficient number of trials at
each position. Starting locations for each trial phase were limited to seven of the eight
compass positions along the wall circumference (i.e. E, NE, etc , but not S), and they were
counter balanced across phases and goal locations.
There was also a cue-place control condition in which there were eight unique cues of
various colors and polygon shapes evenly spaced out on the wall. The coin was always
located next to one of the cues. The cues were shuffled between trial phases to prohibit
allocentric encoding. (see Supporting documentation: Spatial environment and Trial Detail).
Subjects
The study was approved by UC Irvine Institutional Review Board committee and all
subjects approved and signed consent forms, and were compensated for their participation in
a training ($10/hr) and scanning session ($25/hr). Eleven subjects (6 female, 5 male; 24-39
years of age; mean(std) age 29.8(5.15); all with post-secondary education) were recruited
directly or by referral. All subjects were self-reported right handed and none had any known
neurological disorders.
fMRI Acquisition and Analysis

All subjects were given 1 hour of training one or two days prior to scanning, and were also
given 5 minutes of practice while lying down in the scanner, just before scanning. FMRI
data was acquired with a Phillips 3T Intera Achieva, sense factor 2, 37 interleaved 3mm-
thick slices, TR=2, no gap, flip angle 70 degrees, TE=30ms, FOV=240240, 33mm in
plane resolution. Slices were taken perpendicular to the long axis of the hippocampus, which
has been shown to increase signal in the temporal lobe, accompanied by a decrease in
posterior orbital frontal cortex and rostral anterior cingulate (Weiskopf, et al., 2005). The
most anterior slice reached the genu of the corpus callosum and covered Broadman Area 9,
but cut off orbital frontal cortex.
In the scanning session for each subject, there were three scanning runs. In each run there
were 12 trials with 1 wall cue and 4 trials with the cue-place control condition. Data were
analyzed with SPM2 (http://www.fil.ion.ucl.ac.uk). Images were realigned and normalized
to the EPI template in MNI (Montreal Neurological Institute) space, and smoothed with 8

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mm FWHM Gaussian kernels. We applied the general linear model to whiten data, with
regressors for each cell of 22 matrix for successful trials (allocentric vs. cue-place X
encoding vs. test), as well as the allocentric and cue-place delay periods. Regressors were
convolved with the canonical hemodynamic response. Additionally, unsuccessful trials were
coded separately.
Voxel Extraction and Dataset Construction

Overall, there was an average of 67.4 (sd 15.0) percent successful trials. We created separate
datasets for each of the 7 (4 females) out of 11 subjects who had at least 7 correct N trials,
which provided a nearly balanced number of E/W/N trials. The average number of
successful trials for the 7 subjects used for decoding, at E, W, and N locations were 9.36 (sd
1.18), 8.57 (sd 1.9), 8.71(1.6). Because each trial contributes two observations to the data set
(for encoding and test phase), the average total number of observations for subjects was 53.3
(sd 7.0).
In post-scan interviews, subjects described locations as generally left/right of the cue, except
for 3 subjects that described the arena as a clock face. No subject noticed that coin locations
were repeated exactly at the E, W, N (or 3, 6, 9 oclock) positions. (As mentioned in
methods section on Trial Details, goal locations were spread out all over the arena during
training, but locations were limited to three positions during scanning.) Those that projected
a clock face thought locations were scattered around 3, 6 or 9 oclock. It is worth pointing
out that the three subjects who used clock face descriptions were the 2nd, 4th, and 6th best
performing subjects, which implies they were above average but not outliers. Moreover,
only two of those three subjects had at least seven successful N trials and were included in
the neural decoding test.
Each data point in each dataset consisted of a row vector of individual voxel activity. The
voxel activity in each row was the average at each voxel over the third and fourth volumes
after the end of successful trial phases. The third and fourth volumes correspond to the end
of trial phase plus a 5-9 second lag time, which covers the peak of the canonical
hemodynamic response to the moment that subjects were located at the goal. We included
both encoding and test phases in one dataset so that neural decoding represents location
information irrespective of differences in specific trial phase processing.
Voxels were extracted using the SPM2 extraction tool to whiten data and regress out motion
parameters. Voxels were extract from seven bilateral regions implicated in spatial navigation
(e.g. Maguire, et al., 1998). The regions were defined anatomically by predefined templates
in the WFU Pickatlas (Maldjian, et al. 2003) or by spheres centered at coordinates defined
functionally, as detailed in Table 1 (see also Supporting documentation: fMRI Results and
Data Extraction). In addition, all seven regions were combined as another data matrix,
thereby giving 8 datasets for each subject.
Neural Decoding
We applied linear partial least squares (PLS) regression combined with taking the maximum
output to classify locations (E, W, or N) for predictions. Briefly, PLS uses singular value
decomposition to find components with maximum variance, and then project data onto those
components prior to regression (e.g. Shawe-Taylor & Cristianini, 2004). Projecting data
onto components reduces the dimensions of the input space and alleviates the need to select
voxels, thereby making it more feasible to operate over all voxels in a regional analysis. This
is especially relevant for spatial navigation because cellular recordings show that an
ensemble of place-cells is most important for place reconstruction (Zhang, et al., 1998;
Redish, et al., 2001).

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In PLS regression the decomposition is applied to the covariance of the data with target
vectors, which is in contrast to the better known PCA (Principle Component Analysis)
method that uses the covariance of the data itself. Using the covariance with the target
vectors implies that components are directly related to the target. This is especially
important for fMRI data, which is known to have many noise sources that could dominate a
PCA application. Moreover, in PLS regression the decomposition of the covariance matrix
can be iterated with the residuals as a new target, as we detail below. Iterating over the
residuals makes this a very different PLS procedure than used for statistical inferencing in
fMRI (McIntosh, et al., 1996), one that is more appropriate for prediction. PLS also
compares quite favorably to support vector machines in a variety of benchmark datasets
(Bennett & Embrechts, 2003), and was previously shown to have good performance with
fMRI data (Rodriguez, 2006).
In detail, the procedure starts with a standard multivariate regression equation, Y=XW,
where in our case Y is a matrix of t3 target vectors, where there are t observations, and
labels in each row of Y were set either 1 or 0 to indicate one of three goal locations. X is a
mean-centered data matrix of size t v, where the v columns are voxels, and W is a vector of
weights. In the first step, singular value decomposition is used to find the one component U1
that can best account for the variance of XY. Then, the data matrix X in the regression
equation is replaced by XU1, giving Y= XU1W. The weight vector is determined by solving
the normal equations ([XU1][XU1])W = [XU1]Y. Then, U1 is removed from the space of X
(i.e. take the residual of the equation X=b U1, where b is another weight vector), thereby
producing a new matrix X2, that is orthogonal to U1. Likewise, Y is replaced by the residual
Y XU1W =Y2, and the process is repeated using the covariance of X2Y2. After iterating K
times, the above regression equation becomes Y=[XU1 X2U2 X3U3 XKUK]W, where W
=[ W1 W2 WK]. For making a prediction, a test data vector is used instead of X (for
details, see Bennet & Embrechts, 2003; Shawe-Taylor & Cristianini, 2004). In pilot testing,
K~30 was found to be optimal, and for 27<=K<=31 there are only small changes in
classification performance (i.e. less than 0.5 percent change) that do not affect the
significance of p-values (see Supporting documentation: Neural Decoding).
Leave-one out cross-validation was used to calculate the percentage of correct predictions
for each of the eight datasets, for each of the seven subjects. This performance was then
compared to a nave classifier as a baseline null model. The nave classifier ignores voxel
data and merely selects the most likely class according to prior probabilities for each subject.
It performs slightly above a 33 percent chance level because subjects do not have an equal
number of E/W/N successful trials. Significant performance for each region was inferred by
a within-subject, paired (PLS vs. nave classifier), right-tailed t-test using p-value threshold
of 0.0063, which represents a p-value of 0.05 with a conservative Bonferroni correction for
eight datasets.
Results
The average percent correct across subjects was significantly better than the nave classifier
for medial frontal gyrus, hippocampus and inferior parietal regions (51.4 percent, p~.0006;
47.7 percent, p~0.0056; 49.2 percent, p~0.0026, respectively; Figure 2a). Performance for
all other regions was better than the nave classifier at p<0.03. The maximum performance
of 60.7 percent correct (p<0.0001) was attained using the combined regions dataset. Figure 3
shows the percent correct for each region across time windows. There is a clear peak for the
time window that corresponds to the peak of a canonical hemodynamic response for when
the subject was located at the goal.

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As a confirmation that classification pertains to navigational processes, we compared

prediction performance with other test datasets (Supporting documentation: Control Tests).
In one test, we split classification performance for each goal location separately to see if
perhaps any location had a large impact on the total percent correct. One might for example
suppose that N goal locations are classified very well because they are more difficult or
perhaps more salient in some regard. We tallied the percent correct classification for each
goal location separately for each subject, and then compared locations by paired t-tests of N
minus E, N minus W, etc. . No region showed a significantly greater performance for one
location over both of the other two (Figure S5). Thus, there is no evidence that neural
decoding is only picking up on one class for making predictions about goal locations.
Instead it seems that each goal location is predicted correctly with similar percentages.
We also considered data from the test phase of unsuccessful trials, which likely represent
navigational errors. Given the small number of unsuccessful trials and lack of consistency in
subjects actual locations at the end of the trial, we use the desired goal location as the class
label. Performance was poor for all regions (range 29.3 to 42.5, see Table S3.), which shows
that error-trial data are unclassifiable, as one should expect. In another control test, we found
that data from the cue-place control condition are not classified as belonging to any goal
location. These tests help rule out the possibility that classification performance in
successful trials is based on some non-spatial factor.
The fact that many regions of the spatial navigation network carry predictive information
suggests that specific goal identity is disseminated throughout the network, and/or each
region carries some distinguishing information about goal locations. Indeed, many regions
of the navigation network have cells with a coarse representation of location, which likely
contributes to goal-oriented, allocentric navigation (Knierem, 2006). To address this issue,
we performed an exploratory analysis to test which regions have the biggest impact on
performance of the combined regions dataset. For this we applied a paired t-test (within
subject, uncorrected) to the difference of percent correct with the combined dataset versus
percent correct when data from each region was taken out separately. It turns out that only
when removing the hippocampus data did we find a significant decrease in performance
relative to the null hypothesis of zero change (60.7 percent for all seven regions combined
versus 58.4 percent for all regions except hippocampus, within subject mean difference
0.021, sd 0.027, p~0.039; all other regions were in a range of mean differences 0.003 to
0.029, and p~ 0.086 to 0.67). Consequently, it seems that voxel data from the hippocampus
adds unique information to the combined regions data set, whereas information in other
regions are more redundant. The presence of unique information does not guarantee the best
prediction of goal locations because other factors, such as MRI signal noise, or noise related
to other hippocampal coding functions, could interfere with classification. Nonetheless, this
finding is consistent with the cellular recordings that show the hippocampus is specialized
for place representations, and other regions are involved in coding coarser aspects of goal
locations, as part of route planning and execution (e.g. Poucet et al., 2004; Knierem, 2006).
Although speculative at the spatial resolution of fMRI, our data suggest that those other
representations are somehow more shared among regions of the spatial navigation network.
Discussion
In summary, we have demonstrated that goal locations in virtual navigation can be decoded
from the pattern of voxels that corresponds to the moment subjects were located there.
Prediction performance was most significant in the medial prefrontal cortex, hippocampus,
and inferior parietal cortex. Moreover, we used pattern analysis with all regions combined to
show that information about goals seems to be disseminated in the navigation network,
except that hippocampus may carry unique information.

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The work reported here supports another recent fMRI study on neural decoding for place-
related information by Hassabis and colleagues (Hassabis, et al., 2009). In that study,
subjects navigated to one of four corners of a room, which were identified by wall cues
between the corners. In a search light procedure limited to hippocampus and

parahippocampus, neural decoding could discriminate activation patterns that correspond to
the four corners for voxels in both regions. However, there were significantly more voxels in
the hippocampus that could discriminate above chance than in the parahippocampus. Thus,
our results and that of Hassabis and colleagues show that goal location can be decoded and
that the hippocampus may be more specialized. An important difference is that the fMRI
acquisition in that study used a high resolution protocol and only covered slices that
encompass the hippocampal axis, which precludes a complete comparison to the work here.
Nonetheless, both studies raise an issue whether neural decoding of voxel patterns actually
reflects place-cell activity, or some other undetermined spatial representations. Clearly,
more studies are needed to fully establish a connection to place-cells.
It is also worthwhile to point out other fMRI studies related to goal processing. In a recent
spatial navigation study, a region of medial prefrontal cortex was found to correlate with the
proximity to goal locations (Spiers & Maguire, 2007). In that study, goals were destinations
within a virtual depiction of the city of London. The correlation pertained to all destinations
and did not involve neural decoding of specific locations. Thus it is not clear if voxel
activity would also track proximity to specific goal locations. In our case, where there are no
barriers or corners that have to be negotiated, the proximity to the goal is confounded with
the time to a trial end. Future work could try to separate this more clearly by using obstacles
to force the subject to be close to the goal in virtual space, but still far in time from reaching
it.
Prefrontal cortex has also been implicated in non-spatial goal processing functions, such as
behavioral planning, decision making/intentions, motor planning and task-set maintenance.
Haynes et al. (2007) have shown that neural decoding can dissociate motor execution/
planning from the intentional aspects of behavior. They found that voxel activity could
predict motor execution/planning best in a dorsal-medial frontal region, whereas intentions
were predicted in more anterior and lateral frontal regions. In our task set up, we have
randomized and counterbalanced starting positions and orientations with respect to goal
locations, so that we do not expect motor movements to be correlated with specific goals.
However, we do not claim to separate goal locations from all motor-related activity because
navigational planning may use heading vectors that are egocentrically defined motion plans
(e.g. Knierem, 2006).
We also do not have strong claims about what kind of task-set information subjects must
maintain during navigation. One possibility is that the goal-location as defined by a

cognitive map is maintained and used in navigation. However, as partially shown in Figure
2b, we did not find strong support for any region that maintains information about specific
goals locations throughout the navigation process. Another possibility is that once a subject
is oriented and a route is planned, a heading vector is all that is maintained (e.g. Pearce et
al., 1998; Kubie & Fenton, 2009).
One potential caveat in the study is that visual input could be driving prediction performance
if subjects keep the wall cue in view while searching for the coin. However, within the last
four seconds, subjects have the wall cue in view only 7.7 (sd 6.9) percent of trial phase
endings, which implies that direct views of the cue have limited influence. Moreover,
classification performance with voxels from early visual cortex (39.1 percent) showed no
evidence of prediction greater than the nave classifier (p~0.339) or chance level (p~0.096;
Supporting documentation: Control Tests). Another potential caveat is that attention,

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working memory, and/or verbal processes about broadly defined goal locations are driving
performance, as opposed to navigational representations related to place. However, if that
were the case we would not expect there to be a well defined peak in Figure 3, because
subjects determine their general destination well before reaching the goal location. This
argument could be supported by neural decoding of more varied goal locations or other
kinds of spatial information.
A potential limitation of the study is that during the test phase subjects might wander around
the goal location and retrieve the coin only by chance. However, subjects are required to be
in the goal vicinity for at least 2 seconds so that they cannot achieve good performance
merely by a random walk. They must have at least a general idea of where the coin is
located. As shown in Table S1, subjects on average take about 2 seconds more to find a coin
during retrieval than during encoding, which implies they are only wandering around at
most a few spots near the goal location. Moreover, if subjects were actually retrieving coins
by a random search on occasion, then one would expect that to impair classification
performance. Nonetheless, a follow-up study could vary the goal locations around some
general areas and demand that subjects indicate one spot where they believe the coin to be in
the retrieval phase.
Future work could also extend the findings here by looking at a variety of place-related
information with multivoxel pattern analysis that reflects, and potentially extends, data from
cellular recordings. For example, one could consider salient non-goal locations, such as
decision points in a T-maze, or heading vectors and head directions. A primary concern is
that there would be enough data points. As we show in the control test with smaller samples
(e.g. Table S5), it is important to have a sufficient number of data points for good
classification performance. Exactly how many is difficult to say, but for a freely navigated
environment, there are trade offs between speed of navigation, the ability to acquire fMRI
volumes while the subject is at some location, the availability of landmarks and/or visual
cues, total number of discernible locations, and of course, total number of trials.
Supplementary Material
Refer to Web version on PubMed Central for supplementary material.
Acknowledgments
This work was supported by the Research Imaging Center at University California, Irvine, by a faculty development
award from University of California, Irvine, and by the General Clinical Research Council, UC Irvine School of
Medicine through a National Center for Research Resources health service research grant M01 RR00827.
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Rodriguez Page 11
Figure 1.
Snapshots of the virtual environment are shown for the encoding phase. In the retrieval test
phase (not shown) the coin is not visible (see also Figure S1).

Rodriguez Page 12
Figure 2.
Prediction performance across regions for the PLS procedure are well above that of a nave
classifier. a. Percent correct prediction performance across seven bilateral regions and the
combination of those regions. (HPC= hippocampus, PHPC=parahippocampus,
InfPar=inferior parietal, SupPar=superior parietal, OP/Ret, occipito-parietal/retrosplenial
cortex, m/vStr=medial/ventral striatum, mPFC=medial prefrontal cortex, Comb=regions
combined; * indicates significance at p-values < 0.0063, which is 0.05 corrected for multiple
comparisons of 8 regions). b. Prediction performance for each subject for the combined
regions dataset.

Rodriguez Page 13
Figure 3.
Prediction performance for each region peaks around the same time as the peak of a
canonical hemodynamic response to the moment when subjects are located at the goal (o
combined regions; -- m/vStr regions; all other regions are unmarked because their
performance is highly similar).

Rodriguez Page 14
Table 1
Datasets were created from 7 regions defined functionally or from the WFU Pickatlas, as well as their
combination.
Region Whole region or Number of

Center/Radius Voxels
Hippocampus Whole 523

Parahippocampus Whole 585
Inferior Parietal Whole 1065
Superior Parietal Whole 977
Occipito- -15 -66 18; 16mm 2342
Parietal/Retrosplenial
Striatum -12 10 0; 12 mm 531
Medial PFC Whole 1616
Combined 7639

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Hum Brain Mapp. 2010 March ; 31(3): 391397. doi:10.1002/hbm.20873.

Neural Decoding of Goal Locations in Spatial Navigation in

Materials and Methods

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

fMRI Acquisition and Analysis

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Voxel Extraction and Dataset Construction

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

As a confirmation that classification pertains to navigational processes, we compared

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

between the corners. In a search light procedure limited to hippocampus and

maintain during navigation. One possibility is that the goal-location as defined by a

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

Region Whole region or Number of

Hippocampus Whole 523

Hum Brain Mapp. Author manuscript; available in PMC 2011 March 1.

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