Behavioural Brain Research xxx (xxxx) xxx–xxx

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Behavioural Brain Research

journal homepage: www.elsevier.com/locate/bbr

Hex Maze: A new virtual maze able to track acquisition and usage of three
navigation strategies
⁎
Meg J. Spriggsa,b, , Ian J. Kirka,b, Ronald W. Skeltonc
a
School of Psychology, University of Auckland, New Zealand
b
Brain Research New Zealand, New Zealand
c
Department of Psychology, University of Victoria, Canada

A R T I C L E I N F O A B S T R A C T

Keywords: Spatial navigation is a complex and multi-faceted skill that, in humans, is understood to encompass two distinct
Allocentric navigational strategies, namely allocentric and egocentric navigation. These differ in the frame of reference used
Egocentric and the brain networks activated. However, egocentric navigation can be further divided into two, equally
Spatial navigation distinct strategies depending on whether the navigator is using subject-to-object relations (egocentric-cue) or
Virtual maze
direction of body turns (egocentric-response) to navigate. To date, there are no experimental paradigms able to
Sex differences
distinguish between participants’ employment of allocentric, egocentric-cue and egocentric-response strategies,
and to track their usage over time. The current study presents the Hex Maze: a novel virtual environment that
can not only distinguish between the three navigational strategies, but can also be used to index aspects of
strategy use such as preference, acquisition, stability and competence. To illustrate this, 32 male and 32 female
participants were presented with the Hex Maze and sex differences in strategy usage were explored. While the
results offer some support for previously identified sex differences in strategy preference, there were no sig-
nificant sex differences in the novel measures of strategy acquisition, stability, or multi-strategy competence.
Additionally, our results suggest that strategy preference does not preclude learning to competently navigate
using other strategies. Importantly, the current study offers validation for the Hex Maze as an unbiased method
of exploring spatial navigation, and it is anticipated that this easy-to-use tool will be valuable across research and
clinical settings.

1. Introduction irrespective of their perspective. Single cell recordings in rodents have

The ability to navigate through one’s environment is a crucial skill
for successful interaction with the external world. Recent technological
developments have allowed a smooth transition from studying spatial
navigation in rodents to studying spatial navigation in humans, and the
integration of findings from both fields grants an in-depth under-
standing of this important cognitive skill [1,2]. With this has come the
understanding that spatial navigation is multi-faceted and employs a
number of distinct brain networks. This therefore raises questions of
how different components of spatial navigation combine, and how they
are differentially utilized in distinctive environments [3,1,4].
Spatial navigation is widely recognized as consisting of two broad
strategies differing in the frame of reference used; allocentric naviga-
tion and egocentric navigation. Allocentric spatial navigation uses a
world-centered frame of reference, and involves encoding object-to-
object relations [1,5,6,7]. The allocentric reference frame is in-
dependent from the navigator as the same information is encoded
identified a group of hippocampal ‘place cells’ that encode location
relative to an external boundary [8,7]. Human neuroimaging studies
support the central role of the hippocampus [9–14] and surrounding
parahippocampal gyrus [13,14] in allocentric navigation. Both rodent
and human studies also indicate a role for the retrosplenial cortex in
landmark identification, and the translation between allocentric and
egocentric navigation [15–19]. However, activation is not limited to
these regions, and in a meta-analysis, Boccia et al. [20] identified a
wider network also frequently activated during allocentric navigation
including regions within the occipital and frontal cortices.
Conversely, egocentric navigation involves spatial representations
that are dependent on the location of the observer. Egocentric naviga-
tion uses the location of the observer as a point of reference, which
changes with a change in the observer’s perspective [1,5]. However,
there remains some ambiguity as to how egocentric information is en-
coded. Traditionally, egocentric reference frames have been defined as
involving subject-to-object relations [5]. This type of egocentric

⁎
Corresponding author at: The University of Auckland, Private Bag 92019, Auckland 1142, New Zealand.
E-mail address: mspr827@aucklanduni.ac.nz (M.J. Spriggs).

https://doi.org/10.1016/j.bbr.2017.11.041
Received 24 June 2017; Received in revised form 17 November 2017; Accepted 30 November 2017
0166-4328/ © 2017 Elsevier B.V. All rights reserved.

Please cite this article as: Spriggs, M.J., Behavioural Brain Research (2017), https://doi.org/10.1016/j.bbr.2017.11.041
M.J. Spriggs et al.
navigation can be referred to as ‘cue-based’, as it relies of the location of
cues in the environment. However, egocentric navigation also en-
comapsses navigating based on the direction of body-turns, which can
be referred to as ‘non-spatial’ or ‘response-based’ as it does not involve
encoding of environmental information [11,12,7]. Accordingly, ego-
centric navigation is frequently associated with activation across both
the parietal lobe (posterior cingulate, precuneus, inferior parietal lo-
bule) [21,16,17,22] and striatum [11,23,12]. The parietal lobe is un-
derstood to be involved in “body referencing” [1,24] and may therefore
be better suited for cue-based navigation. Conversely, the striatal
system is associated with stimulus-response driven behaviour [25,11]
as well as the encoding of spatial layout [1] and may thus be better
suited for response-based egocentric navigation. Similarly to allocentric
navigation, a wider network of regions has frequently been identified in
neuroimaging studies of egocentric navigation including frontal and
occipital regions, some of which showing some overlap with those ac-
tivated in allocentric tasks [20].
In recent years, human spatial navigation has increasingly been
studied in virtual environments. The use of a virtual environment has
several advantages over real world navigation including easy control
and manipulation of experimental factors, and allowing participants
active exploration, repetitive practice, and feedback [26,27]. There are
however some limitations to virtual environments, such as a narrower
point of view and limited resolution. Regardless, virtual reality, and the
cognitive maps built within, show good translation to real world spatial
navigation and are easy to transfer between clinical and research set-
tings [28,27].
The virtual environments used to study spatial navigation in hu-
mans are often derivatives of well established rodent tasks, such as the
radial arm maze (RAM) and the Morris water maze [29]. The RAM has a
central hub with radiating arms, ranging in number from just four to 17
[30]. The rodent RAM is usually constructed so that distal cues are
visible, and a subset of the arms are baited with food. In general, testing
takes two forms: i) working memory, in which within-trial arm re-visits
are considered errors and; ii) reference memory, in which maximally
successful performance requires the animal to only visit the arms that
have been baited on previous trials. Virtual versions of the RAM also
require human participants to navigate to ‘goals’ hidden at the end of a
selection of the arms, and consequently remember which arms they
have and have not visited [31–34,10,12]. Performance is measured in
number of errors (wrong arms visited), and latency. Navigational
strategy use is determined using a small number (1 or 2) of probe trials.
Hidden goals can be located allocentrically using the extra-maze en-
vironment, or egocentrically using the spatial arrangement of arms of
the maze (e.g., “the first goal is in the second arm to the left of the start
position”). This egocentric navigation corresponds to ‘response-based’
navigation.
One of the limitations of the rodent RAM is the potential use
proximal visual or olfactory cues [29]. The rodent MWM [35,36] was
designed specifically to address this issue. The MWM consists of a cir-
cular pool of opaque water, featureless on the inside, and surrounded
by curtains so that distal cues can be manipulated (or removed). A ty-
pical paradigm might involve learning the position of a hidden platform
over repeated trials with varying start position (i.e., in the N,S,E, or W
quadrants of the maze). Subsequent to learning the position of the
platform (a day later than the last hidden platform trial for example),
probe trials are run in which the platform is removed altogether. Spatial
memory can be assessed by the amount of time spent in, or distance
swam within the previous target quadrant. Virtual versions of the MWM
also require human participants to locate a single hidden goal or plat-
form and performance is measured via accuracy, latency, and path
length [32,37,11,38,39,40–42]. A single probe trial is used to de-
termine whether the platform was located using an allocentric (extra-
maze environmental cues) or egocentric-cue (an intramaze landmark)
strategy. Because the start location is varied, an egocentric-response
strategy cannot be used.
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As can be seen from the above discussion, virtual versions of the
RAM and the MWM only distinguish between allocentric navigation,
and one of the types of egocentric navigation (egocentric-response and
egocentric-cue respectively). Additionally, measures of performance are
largely limited to accuracy, latency and performance on a single probe
trial. While such measures indicate strategy choice and task proficiency,
they do not provide more detailed information about strategy usage,
such as acquisition, stability, and competence. These limitations may
have implications for how we interpret individual, or group based
differences in spatial navigation, for example, the widely cited sex
differences in strategy preference and navigational efficiency. Both
subjective (self-report) and objective (providing navigational informa-
tion) measures have previously indicated that males are more likely to
use allocentric strategies while females are more likely to use egocentric
strategies [43–46]. Additionally, while males tend to perform better in
tasks requiring allocentric navigation, this male-advantage is reduced
or eliminated when egocentric strategies can be employed
[32,37,47–49]. However, whether these sex differences are due to
strategy preference (the strategy a participant will chosen most often
over all others) and/or competence (accuracy and efficiency using a
particular strategy), and whether there are sex differences in strategy
acquisition, or stability remains unclear. Furthermore, it is uncertain
whether these sex differences would persevere in an environment in
which navigators have the choice of using all three navigation strategies
(allocentric, egocentric-cue, and egocentric-response).
Here we present a new paradigm for studying spatial navigation; the
Hex Maze. The Hex Maze is a 6 arm RAM, but follows the paradigmatic
procedures of a MWM. This means that while it has a countable number
of pathways, and thus a limited number of choices as to where the goal
might be, there is only one goal in a fixed location, and trials begin from
variable start locations (always at the end of an arm, rather than in the
centre of the maze). Participants are presented with alternating
learning and probe trials. In learning trials (called “Find-It” trials),
participants must locate an invisible platform. On probe trials (called
“Show-Me” trials), they must then return to the location of the platform
on the previous Find-It trial. Importantly, a shift in maze configuration
between Find-It and Show-Me trials means that participants will give
different responses on probe trials depending on which of the three
navigational strategies they use (allocentric, egocentric-cue and ego-
centric-response). This not only allows an examination of strategy
preference, but also provides more detailed information on strategy use,
such as acquisition, stability, and competence. The aims of the current
study were to 1) establish the validity and virtues of the paradigm, 2)
provide a baseline from young, healthy university students and 3) ex-
plore sex differences in the various aspects of strategy use elucidated by
the paradigm.
2. Materials and methods
2.1. Participants
Participants were 64 undergraduates recruited primarily from a first
year Psychology class (32 Men and 32 Women, M Age = 21.0,
SD = 3.8) who received partial course credit for participating. The
study was approved by the University of Victoria’s Human Research
Ethics Board. After consent was obtained, participants were given a
brief survey to collect demographic information (age, handedness, etc.),
to check exclusion criteria (English not first language, brain injury,
psychiatric and neurological disorders) and some potential influences
(exhaustion, dizziness, and recent food, alcohol or tobacco).
2.2. Apparatus
All navigational testing was completed in the Hex Maze. The maze
was built using the UnReal Development Kit (UDK® Epic Megagames),
run on a Window 7 desktop computer (Dell, 2.83 GHz) using a 23”
M.J. Spriggs et al.
Fig. 1. Hex maze design. a) A view of the Hex Maze from above (Note that participants do not see the maze from this angle). b) Four views of Hex Maze looking north (i), south (ii), east
(iii), and west (iv) from the centre of the maze.
1680 × 1050 screen resolution placed on a desktop ∼80 cm from the
participant.
Participants moved using an Xbox 360-type controller with buttons
programmed to allow only left, right and forwards movements (both
joysticks were disabled).
All statistical analyses were conducted in Microsoft Excel.
Fig. 1a provides an aerial view of the maze environment. The maze
was contained in a large circular arena with an apparent diameter
∼60m. It was bounded by a low stone wall and had a tiled floor. The
floor of the maze was trisected by three recessed alleys each 1/10th the
diameter of the arena, at 60° angles to each other with one alley or-
iented East-West, and a hexagonal plate in the centre so that the floor
tiling was not a clue to orientation. The entire arena was enclosed in a
round room that was twice the diameter of the arena. Two large win-
dows to the arbitrarily labelled north and south afforded views to a
mountain range that peaked at the north, and a large body of water to
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the south with a large island (Fig. 1b). The room also had two sets of
narrower but equally high windows to the east and west which afforded
views of the mountain ranges coming down to meet the large body of
water almost exactly at the east and west. At the end of each arm,
floating above the low wall bounding the arena, was a whitish/grey
sphere.
2.3. Navigation training and testing
2.3.1. Training
Throughout training and testing, participants were instructed as to
the nature of the upcoming trial by the experimenter reading from a
script. Training consisted of three phases: 1) exploration, 2) visible
platform trials, and 3) an explicit probe.
Firstly, the exploration phase was used to familiarize participants
with the environment and how to move using the controller. At the start
M.J. Spriggs et al.
of the single “explore” trial, participants were teleported to the eastern
extreme of the room outside of the area (grassy area visible in Fig. 1a),
facing out the window. They were instructed to explore the room, look
out the windows, and to let the experimenter know when they were
satisfied with their knowledge of the environment and the use of the
controller.
The exploration phase was followed by four visible platform trials to
familiarize the participant with the platform-finding procedure.
Participants were teleported to the end of one arm of the area and were
told to move to a visible platform within the arena as quickly and di-
rectly as possible, and step on it. The visible platform locations were, in
order: 1) the centre of the arena, 2) the far end of the alley in line with
the starting alley, 3) the second alley on the right and 4) the first alley
on the left. Once participants reached the platform, it made an audible
sound and they were instructed that they could look around as much as
they liked, but would be prevented from moving off the platform by an
invisible barrier. For these four trials, an inner wall was raised to mask
the outside landscape and the spheres marking the end of each arm
were removed.
Finally, participants were presented with an explicit probe.
Participants were told that this was the first “Show-Me” trial and that
the experimenter wanted to see how good their “intuition” was by
having them go to where they thought the invisible platform might be
hidden on the next trial. The trial started with the participant in the
centre of the maze, facing east. One of the six white spheres was re-
placed with a dark grey sphere. The purpose of this trial was to see how
many participants could guess that the platform would be hidden in the
arm marked by the coloured sphere, or whether participants were un-
intentionally biased by any other aspects of the maze environment.
Participants were told to go to the end of the arm where they thought
the platform would be hidden. Once the participant reached the end of
an arm, there was an audible beep and they were trapped by an in-
visible barrier.
2.3.2. Testing
In order to measure acquisition of navigational knowledge and
strategy preference, participants were given 10 pairs of trials in sets of
five across two maze configurations (Fig. 2a). Each pair of trials con-
sisted of a learning trial followed by an explicit probe trial, labelled
“Find-It” trials and “Show-Me” trials respectively. Prior to the first
learning trial, participants were informed that they would be alter-
nating between two types of trials across the next two blocks of trials,
and the two types of trials were described. On “Find-It” trials, their task
was find an invisible platform at the end of one arm, which would
visibly rise-up below them and make a sound when it was found. Like
on the visible platform trials, they were not able to get back off the
platform once on it, but could look around as much as they liked and
were asked to inform the experimenter when ready to move on. “Find-
It” trials were always followed by “Show-Me” trials, in which there was
no platform. Instead, participants were tasked with returning to the
location of the platform on the previous “Find-It” trial. Once they
reached the end of their chosen arm, they would hear a beep and an
invisible barrier would keep them from leaving the arm. Participants
were also told that for each configuration, the platform would remain in
the same place, and that they would be informed on each trial whether
it was a “Find-It” or “Show-Me” trial.
In the first set of five trial pairs, the “Find-It” trials started at the end
of the east arm and the goal (platform) was at the end of the southwest
arm (see Fig. 2a), which was the second arm to the left and was marked
by a dark grey sphere. The “Show-Me” trials started at the end of the
northwest arm, and the grey sphere was move to the end of the west
arm. Shifting the configuration of the maze on the “Show-Me” trial
enables a distinction between three navigational strategies based on
participants’ chosen arm. If participants went straight to the southwest
arm, they were scored as navigating by an allocentric “place” strategy
as this is the original location of the platform as indicated by the arena
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windows (but is no longer marked by the dark grey sphere and is no
longer the second arm to the left). If they went to the end of the west
arm, they were scored as navigating by an egocentric “cue” strategy as
this is now marked by the dark grey sphere. If they took the second left
turn and went to the southeast arm, they were scored as navigating by
an egocentric “response” strategy, as this corresponds to the original
turning direction. If they went to either of the other two arms, or re-
turned to the original start arm, they were scored as not having learned
the goal location or any strategy.
On the second set of 5 trial pairs, the “Find-It” trials started in the
southwest, and the platform was at the end of the east arm (the second
right) and was marked by a blue sphere. “Show-Me” trials started at the
end of the west arm and the blue sphere was moved to the end of the
northeast arm. Once again, participants were scored by their arm
choice: to the same place (east), to the same cue (blue-sphere, north-
east) or by the same response pattern (taking the second right, south-
east).
In sum, on learning trials, the three navigational strategies were
convergent, while on the probe trials they were divergent and revealed
strategy selection. At no point were participants informed of this di-
vergence on the Show-Me trials or that the start position would be
different. The most important measure of performance was arm choice
on the probe (“Show-Me”) trials, however distance and latency were
also recorded. The use of two different maze configurations (with dif-
ferent places, cues and responses) reduces the effects of overtraining
and allows additional measurements such the effects of experience on
acquisition of a new goal location and the stability of strategy selection.
2.3.3. Ancillary tasks
Three tasks were administered to assess the extent of each partici-
pant’s navigational knowledge. First, three “forced-strategy” probes
were administered (Fig. 2b). Participants were instructed to go to the
location of the platform on the last set of trial pairs (configuration 2).
“Place” probe trials started in the centre of the Hex maze (to obviate
response navigation) and there were no cue spheres. The landscape was
visible through the windows. The “Cue” probe also started in the centre
of the maze, but the windows were blocked by a blank wall (to obviate
allocentric navigation). Each arm was marked with one of three white
spheres and three coloured spheres corresponding to the cues of the
first set (grey), the second set (blue), and an unused option (green). The
Response probe began at the end of one arm. There were no spheres
present and the windows were blocked by a blank wall. In all trials,
participants were instructed to go to the end of an arm to make their
choice and responses were counted as to whether they were correct
relative to the first set or second set of trial pairs.
The second ancillary task was a “Sense of Direction” task [50,40,41]
to test for the ability to point to landscape features that were not in
sight (and sense of immersion or presence) and cognitive map. Parti-
cipants were teleported to the north edge of the room, looking north-
wards (towards the mountains). They were asked to use their hands in
their air above them to point to where they thought the water was.
Their responses were recorded on paper and later scored as follows: 0-at
the display screen, 1-to one side of the screen, 2-straight to left or right,
3-behind the midline, to the side, 4-straight behind.
The third ancillary task was a Room Reconstruction task
[38,50,40,41]. Participants were presented with a template for placing
pictures representing the landscape, the walls, and the radial maze
floor. First they were asked to pick 4 cardinal landscapes from a random
array of 8 cardinal and semi-cardinal views and were awarded a point
for each cardinal landscape in the correct position relative to the first
one placed (maximum − 4). Then they were asked to place the four
walls, and given 1 point if all were correct relative to each other and
another point if at least one wall was correct relative to a cardinal
landscape. Then they were asked to place a representation of the floor
(with 6 arms) with a blue circle representing the cue sphere of the
second set, and were given 1 point if the blue sphere was oriented
M.J. Spriggs et al.
towards the east landscape picture. Finally, they were asked to place on
the “floor” a small red circle representing the platform and were given 1
point if it was correct relative to the first cardinal landscape placed. The
maximum score on this task was 8.
At the conclusion of the test session, participants were asked a series
of 10 questions about the strategies they used in the Hex maze and a
series of 7 questions about their experience with videogames, maps and
compasses and other experiences that might have affected how they
navigated in the Hex maze. Three separate measures of gaming ex-
perience (experience with videogames, 3D games, and 2D games), each
measured on a Likert scale from 0 (never) to 5 (daily) were combined
into a composite “gaming experience” score (out of 15). Controller
experience was also measured on Likert scales from 0 (no experience) to
5 (daily). Finally, participants were asked whether they had previous
map/compass experience and if so, how frequently (1 = occasionally to
5 = daily).
3. Results
First, it was important to validate the Show-Me trials as a means of
evaluating navigation strategy. As part of the post-maze questionnaire,
participants were asked a direct questions regarding strategy con-
sistency across Find-It and Show-Me trials. 82.3% whose responses
could be evaluated (25/30 males, 26/32 females) used the same
strategy on Show-Me trials as they had on the preceding Find-It trial.
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Fig. 2. a) Pathways to the platform on “Find-It” trials
and “Show-Me” trials for configurations 1 and 2.
Stars indicate starting positions and dotted circles
indicate the location of the hidden platforms on
“Find-It” trials. Solid circles represent the spheres;
the coloured circles (dark grey in configuration 1 and
blue in configuration 2) indicate the position of the
single coloured sphere (Cue) in the maze. Coloured
arrows indicate pathways taken on all trials (red), by
those using an egocentric-cue strategy (to C1 or C2,
purple), an egocentric-response strategy (to R1 or R2,
green) or an allocentric-place strategy (to P1 or P2,
yellow). b) Configuration of the Forced Strategy
Probes. Red stars represent start locations and the
arrows represent start facing direction. Large grey
circle surrounding the arena for Cue and Response
probes represent a wall occluding the windows. P1
and P2 indicate the place location for configurations
1 and 2 (determined by windows). C1 and C2 re-
present the location of the coloured spheres used for
configurations 1 and 2. R1 and R2 indicate the di-
rection of turn to reach the platform on configuration
1 and 2, while RP is the response that would have
been employed to get to the Place location for either
set. (For interpretation of the references to colour in
this figure legend, the reader is referred to the web
version of this article.)
This thus provided the groundwork to use the Show-Me trials to probe
strategy prevalence, rate of learning and strategy acquisition.
It was also important to examine participants previous navigational
experience so as to assess the impact of this on measures of Hex Maze
performance. The composite gaming experience score revealed that
most participants had limited previous gaming experience (M = 5.08,
SD = 3.22) and was used to group participants into low/no gaming
experience (score < 3, N = 21), moderate gaming experience (score
3–6, N = 21) and high gaming experience (score > 6 N = 22). The
controller experience score also revealed a relatively low level of pre-
vious experience (M = 1.97, SD = 1.36) and was used to split partici-
pants into groups of little/no controller experience (score < 2 N = 34)
and high control experience (score > 1 N = 30). Interestingly, less
than half of the participants (N = 30) reported any map/compass ex-
perience. As such, participants were grouped by whether or not they
had previous map/compass experience, rather than by frequency.
3.1. Strategy prevalence in the maze
A substantial proportion of participants acquired and used each of
the three possible navigational strategies (allocentric, egocentric-cue,
and egocentric-response) regardless of sex (Fig. 3). One-sample t-tests
against zero indicated that all three strategies were used a significant
proportion of the time (t(63) = 6.60, p < 0.0001). One sample t-tests
against chance indicated that the allocentric (place) strategy was
M.J. Spriggs et al.
chosen more often than chance (1 in 5) (t(63) = 4.43, p < 0.0001,
d = 0.554), and when combined, the two egocentric strategies (cue and
response) were chosen more often than chance (2 in 5) (t(63) = 2.30,
p =0.024, d = 0.287). Pairwise comparisons indicated that the allo-
centric strategy was chosen on more trials that the egocentric-cue
strategy (t(30) = 2.4, p = 0.02, d = 0.3). No other pairwise compar-
isons of strategies were significant (p > 0.05) and there were no sex
differences within strategies (pairwise t-tests p > 0.05).
3.2. Rate of learning
Participants navigated the maze quickly. All latencies on the visible
platform trials were low, reaching an asymptote of between 10 and
15 seconds (M = 10s, SD = 3.91s). After the first Find-It trial, 80% of
the latencies were below 15 seconds (Trials 2–10 M = 14s, SD = 6s)
and 80% of both men and women reached this level of performance by
the fifth trial of the first set, and the second trial of the second set.
Strategy preference did not appear to be a factor in how quickly
participants learned the maze nor how quickly they could navigate to
the hidden platform on Find-It trials (Fig. 4a). A 3 (strategy) x 2 (sex)
ANOVA on average latencies (trials 2–10) revealed no effect of strategy
(F(3.56) = 0.076, p > 0.05, ηp2 = 0.004) or sex (F(1.56) = 2.021,
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Fig. 3. Prevalence of strategy use on 10 Show-Me
trials split by sex. Error bars depict standard error.
p > 0.05, ηp2 = 0.035) and no interaction (F(3.56) = 1.177,
p > 0.05). Additionally, a 3(game experience) x 2(control experience)
x 3(map/compass experience) ANOVA on average latencies revealed no
significant effect of previous game experience (F(2.58) = 0.13,
p > 0.05, ƞp2 = 0.005), previous controller experience (F(1.58)
= 1.55, p > 0.05, d = 0.028) or map/compass experience (F(2.58)
= 0.07, p > 0.05, ƞp2 = 0.001) and no significant interactions (all
p > 0.05). While men were, on average, faster regardless of strategy,
planned post-hoc t-tests showed that they were significantly so only
when their strategy preference was for the egocentric-cue strategy (t(9)
= 2.36, p = 0.04, d = −1.2) (Fig. 4b).
3.3. Strategy use
The ability of the Hex Maze to track the use of the three strategies
over trials made it possible to examine four novel aspects of strategy use
and sex differences within these areas: 1) distribution of preferred
strategies, 2) acquisition of preferred strategies, 3) stability and origin
of strategy preference, and 4) strategy preference vs strategy compe-
tence. These results are discussed below.
Fig. 4. Latencies on Find-It trials by strategy. a)
Latencies over Find It trials 1-10. Dotted line depicts
approximate asymptotic latency. b) Average la-
tencies on trials 2–10 for males and females. Error
bars depict standard errors. * sex difference
p < 0.05.
M.J. Spriggs et al.
3.3.1. Distribution of preferred strategies
Preferred strategy was defined simply as the strategy that an in-
dividual used most often over the 10 trial pairs. The Place strategy was
used most often by both men and women, however the other two
strategies were also preferred by a reasonable proportion of both sexes
(Table 1). Sex differences in strategy preferences were small, with 56%
of men demonstrating a preference for allocentric (Place) navigation
and 56% of women demonstrating a preference for egocentric (Cue and
Response) navigation. A chi-squared analysis of the distribution of
preferred strategies for the 60 participant who had a preferred strategy
showed that not all strategies were preferred equally, X2(2) = 10.0,
p = 0.006, with the distribution for Place/Cue/Response closer to 50/
20/30. A Chi-Square goodness of fit test to these proportions showed a
close fit to this model, X2(2) = 0.117, p = 0.943. Finally, three addi-
tional Pearson chi square analyses revealed no significant relationship
between strategy preference and gaming experience (X2(6) = 4.82,
p = 0.567), previous controller experience (X2(3) = 5.27, p = 0.154),
or previous map/compass experience (X2(3) = 1.88, p = 0.598).
3.3.2. Strategy acquisition
84% of participants chose a strategy on the very first Show-Me trial
(i.e., after only one Find-It trial). 75% of all participants chose a
strategy on all 10 trials, 93.8% chose a strategy on at least nine trials,
and only one participant failed to choose a strategy on 8 or more trials.
Due to the rapidity of strategy learning, it was important to ensure
that the design of the maze did not allow participants to guess the lo-
cation of the platform. On the “Guess” trial given before the location of
the platform was revealed, only 11 males (33%) and six females (19%)
guessed that the platform would be located at the end of the arm
marked by the grey sphere, with only two of the male participants
subsequently demonstrated a preference for the egocentric-cue strategy.
This indicates that participants were not automatically using the co-
loured sphere, and that there was nothing else about the maze that
“gave away” the future location of the platform.1
Analysis of strategy choices on individual trials showed that no one
strategy was adopted first, or before any other strategy. On the very first
Show-Me trial, male participants distributed their strategy choices al-
most equally among the three strategies (Fig. 5a), whereas female
participants were most likely to choose the Response strategy (Fig. 5b).
Over the remaining trials in the first configuration, the Place strategy
was chosen with increasing frequency by both sexes, with a commen-
surate decline in the frequency of choosing the Cue strategy. The rate of
choosing the Response strategy remained stable. By the end of the first
set, all but one participant had acquired at least one of the strategies.
Within the second configuration, the Place strategy was chosen more
often than any other strategy by both sexes. The Cue strategy was
chosen by about a quarter of male and female participants. The Re-
sponse strategy was chosen by a comparable number of female parti-
cipants but a much lower number of male participants.
3.3.3. Stability and origin of strategy preference
Strategy preference tended to remain stable over trials. 31% of
participants chose the same strategy from the first to the last Show-Me
trial, though this tendency was stronger in men than women (15/32
men, 5/32 women, X2(2) = 0.001, p = 0.025). Additionally, there was
stability across maze configurations. Among those who had a preferred
strategy in both sets, 80% (47/59) preferred the same strategy in both
sets (Males: 84%, 26/31; Females: 75%, 21/28). Preference for the
place strategy was the most stable (Table 2).
There was no indication that the distribution was effected by the
maze design via variations in the efficiency or accuracy of the different
1
Although the rate of guessing in males might seem high (compared to chance at 1 in 6
or 16.7%), it is low compared to the number of participants who chose a strategy (i.e. one
of the 3 correct arms) after just one trial: Men 29/32 (91%), women 25/32 (78%).
7
Behavioural Brain Research xxx (xxxx) xxx–xxx
Table 1
Distribution of strategy preferences for males and females.
N Males (%) N Females (%) N All (%)
Place 18 (56) 13 (41) 31 (48)
Cue 5 (16) 6 (19) 11 (17)
Response 6 (19) 12 (38) 18 (28)
Cue + Response 11(34) 18 (56) 29 (45)
Nonea 3 (9) 1 (3) 4 (6)
a
Note: “None” indicates individuals who had no clear strategy preference, usually
because they chose 2 strategies with equal frequency. One female and two males chose
cue and response strategies on 5 trials each, and one male chose place, cue and response
strategies on 3 trials each.
strategies. Additionally, as described above, no strategy was faster than
any other and there was no association between strategy and latency.
This suggests that these results are not idiosyncratic to this maze.2
3.3.4. Strategy preference versus strategy competence
Strategy competence was assessed using the forced strategy probes.
Performance on these probes was scored as perfect (2 points) if the
participant went to the correct arm according to the second (most re-
cent) set of trials or partially correct (1 point) if they went to the arm
that was correct on the first set of trials. Pairwise 2-tailed t-tests re-
vealed no significant differences between the scores of participant on
the 3 different probes (p > 0.05). However, over the 3 forced strategy
probes combined, those who preferred the ego-response strategy per-
formed significantly worse than those preferring the egocentric-cue
strategy (2-tailed t-test, p < 0.05, Fig. 6a), though they were not sig-
nificantly different from those preferring an allocentric strategy (2-
tailed t-test, p > 0.05).
There was a bidirectional association between strategy preference
(on Show-Me trials) and strategy competence (on forced strategy
probes; Fig. 6b). Participants who preferred a particular strategy per-
formed better on the corresponding probe than the other two probes,
and also scored better on that probe than the participants who preferred
the other two strategies. This was true for all three strategies and all
three probes. As such, the poorer overall score of those who preferred
the ego-response strategy was likely due not to poorer scores on the
response probe but rather to poorer performance on the non-congruent
probes. There were no significant sex differences in performance on the
probe trials (Fig. 6c; 2-tailed t-test, p > 0.05), nor were there sig-
nificant sex differences among those of a given strategy preference (2-
tailed t-test, p > 0.05).
Participants who preferred to navigate allocentrically were sig-
nificantly better than chance in the cue probe, and those who preferred
the egocentric-cue strategy were significantly better than chance in the
response probe (1-sample-t-test, p < 0.05; Fig. 6b).3 Additionally,
when the data were analysed in terms of whether the response on each
probe was perfect (i.e., excluding responses that were consistent with
the first configuration) 47% of males and 56% of females received a
perfect score on more than one strategy (Fig. 7d). However, the pro-
portion of participants demonstrating competence across all three
strategies was low. Therefore, many participants were not only con-
currently learning multiple strategies, but demonstrated competence in
using a non-preferred strategy when required.
2
Similarly, there was little or no association between strategy preference and the
ability to find the correct location on the corresponding forced strategy probe. All the
probes were about the same difficulty and there was no relation between frequency of
probe success and frequency of strategy preference.
3
“Chance” was calculated for Place and Cue probes (which started in the centre) as 1/6
chance of getting 2 out of 6 points and 1/6 chance of getting 1 point, for a total of 3 out of
6, or 0.50 points. For Response probes, "chance" was calculated out of 5 because they
started at the end of one arm. Specifically, there were 5 options, so there was a 1/5 chance
of getting 2 points, and a 1/5 chance of getting 1 point, for a total of 3 out of 5, or 0.60
points.
M.J. Spriggs et al. Behavioural Brain Research xxx (xxxx) xxx–xxx

Fig. 5. Number of participants selecting each

strategy on each trial of acquisition, over the two sets
of 5 Show-Me trials for males (a) and females (b).

Table 2 3.4. Cognitive map development

Consistency of strategy preference in first 5 trials (Set 1) versus the second 5 trials (Set 2).
Performance on both tasks of cognitive map development was poor.
Set 1 Set 2
On the Sense of Direction test, six male and five female participants
Place Cue Response No Preference Stable Preference (19% and 15%) pointed behind themselves (Fig. 7a). Conversely, eight
males and three females (25% and 9%) pointed directly at the screen,
a
Place 25 0 0 0 25/25 (100%) suggesting that they did not feel immersed in the environment. The
Cue 2 7a 1 1 7/11 (64%)
Response 3 5 15a 0 15/23 (65%)
average score on the Room Reconstruction task was less than 3, in-
No Preference 2 2 0 1 1/5 (20%) dicating that most participants were not capable of identifying the
views representing all four cardinal directions, and were much less
a
Participants who preferred the same strategy during both sets.

Fig. 6. Results from probe trials. a) Average probe

scores across all probe types split by strategy pre-
ference on previous Show-Me trials (Allo– allo-
centric, Ego-Cue–egocentric cue-based, Ego-
Resp–egocentric response-based). b) Probe scores for
each strategy preference split by probe type (Place,
Cue or Response). Error bars depicting standard
error c) Performance on probe trials split by sex.
Error bars depicting standard error. In these three
graphs (a, b, and c), the maximum score in any ca-
tegory is 2. d) Percentage of males and females able
to respond perfectly on single or multiple place, cue
and response probes. * above bars = two-tailed t-
test, p < 0.05. * within the bars = 1-sample, two-
tailed t-test, p < 0.05 (significantly different from
chance).

8
M.J. Spriggs et al. Behavioural Brain Research xxx (xxxx) xxx–xxx

Fig. 7. Results from 2 cognitive map tests plotted by

sex and preferred strategy. a) Sense of Direction task,
scored out of 4. b) Room Reconstruction, scored out
of 8. * sex difference p < 0.05.

capable of identifying the relative positions of the windows and plat- presented once, or presented only a small number of times over large
form position relative to the landscape outside the windows (Fig. 7b). delays. The Hex Maze is novel in that ‘Show-Me’ trials are presented
Two 2 × 4 ANOVAs, revealed no significant effects of sex or after every learning trial throughout testing. This not only improves
strategy preference on cognitive map development (all p > 0.05). The reliability and applicability to studies requiring averaging over a large
only significant effect present was that male participants who preferred number of trials (i.e., neuroimaging studies), but also allows for
allocentric or cue-based egocentric navigation were better than the strategy usage to be tracked throughout testing. van Gerven et al. [42]
female participants with the same strategy preference (post-hoc pair- presented participants with alternating standard trials (similar to Hex
wise t-test; allocentric t(29) = 2.637, p = 0.01 d = 0.98; egocentric- Maze Find-It trials) and inter-trial probes (similar to Hex Maze Show-
cue t(9) = 2.657, p = 0.03, d = 1.85). Me trials), but were limited to distinguishing between egocentric and
allocentric navigation. The Hex Maze is the first virtual maze to include
repeated ‘probe’ trials that distinguish between three navigational
4. Discussion
strategies.
The current results also demonstrate that the Hex Maze is easy to
The current paper presents a novel method for assessing three dis-
learn and does not bias strategy choice. All strategies were used on a
tinct spatial navigation strategies in a virtual environment. The Hex
considerable proportion of trials. For both men and women, the
Maze is configured as a 6 arm RAM, and is run procedurally as a MWM
strategy used most often was the allocentric strategy, with the ego-
with one platform location, and varied start locations. The paradigm
centric-cue and egocentric-response strategies used for a similar
involves participants repeatedly alternating between locating an in-
number of trials. When grouped together, the egocentric strategies were
visible platform (learning, or ‘Find-It’ trials), and returning to the pre-
used on an equivalent number of trials as the allocentric strategy. This
vious location of that platform (probe, or ‘Show-Me’ trials).
indicates that the maze is not heavily biased towards or against a
Importantly, the Hex Maze not only distinguishes between three navi-
particular strategy (or sex). Additionally, trial latencies indicate that
gational strategies (allocentric, egocentric-cue, and egocentric-re-
the task was easy to learn, with male and female participants reaching
sponse) but also allows further examination of aspects of strategy usage,
asymptote (15 s) after an average of 2 and 4 trials respectively.
such as acquisition, stability, and competence.
Latencies were not significantly affected by strategy choice or the in-
The Hex Maze is easily accessible and easy to use. The maze was
teraction between sex and strategy choice. This is contrary to the notion
constructed in the Unreal Development Kit virtual environment (www.
that males are more efficient allocentric navigators
unrealengine.com) and has been designed so that different key strokes
[31,32,37,48,49,40,41], and suggests that females who prefer to navi-
call different trials, meaning that trial order and several room features
gate allocentrically are equally as efficient.
can be modified without programming. Additionally, the maze creates a
The design of the Hex Maze also allows for an exploration of a
log file with each run, which not only records user locations over time,
number of novel aspects of strategy usage including strategy preference,
but also records the outcome of each trial (e.g., when the platform is
acquisition, stability and competence. Firstly, strategy ‘preference’ was
reached on Find-It trials and which arm is chosen on Show-Me trials).
defined as the strategy used for the most trials by a participant. The
The results of the current study offer validation for the use of the
majority of participants (male and female) demonstrated a preference
Hex Maze as a tool for assessing various features of spatial navigation.
for one strategy, with an approximate 50/50 split between those who
Firstly, the results demonstrate the novel use of “Show-Me” trials to
preferred the allocentric strategy, and those who preferred one of the
assess the strategies participants employ. Show-Me trials require par-
egocentric strategies. Interestingly, previous studies using the RAM
ticipants to move to the position where the invisible platform was
have generally found that a much larger proportion of their participants
hidden on the preceding Find-It trial. Due to a shift in the configuration
use an egocentric-response strategy than an allocentric strategy
of the maze, their response will differ depending on the navigation
[33,10,12]. As previous studies employing alternative maze designs
strategy they used. In the current study, 82.3% of participants reported
have identified similarly high proportions of allocentric navigators
using the same strategy on Find-It and Show-Me trials, indicating that
[56,57,40,41], this suggests a potential bias in the standard RAM. The
Show-Me trials are a credible index of strategy employment.
Hex Maze does not appear to be affected by bias for any one strategy.
The use of a probe trial to determine strategy usage is common
There was a trend for more males to prefer the allocentric strategy,
practice in ‘dual-solution’ rodent mazes [51–54]. Additionally, previous
and more females to prefer the egocentric strategies (primarily the ego-
virtual MWM studies conducted by our group have used similar ‘drop-
response strategy) which is consistent with the premise that males and
the-seed’ trials to distinguish between egocentric and allocentric navi-
females have a tendency to use allocentric and egocentric strategies
gation [55,40,41]. However, such probe trials are usually only

9
M.J. Spriggs et al.
respectively [18,42,50,67–69]. However, these sex differences were
small with only 56% of male and 56% of female participants preferring
the allocentric and egocentric strategies respectively. This suggests that
the option to use any one of the three navigation strategies may miti-
gate this difference. It is however important to acknowledge that there
are a number of other factors that influence strategy preference such as
experience [55] or stress [58,54,42] that were not measured in the
current study.
It may appear that the working definition of ‘preference’ is fairly
liberal, as the strategy used more often than any other is considered a
participant’s preference (and thus can still be a preference even if it is
not used on the majority of trials). However, when focusing instead on
strategy ‘dominance’ (the strategy used on > 50% of trials), the result
changes for only two participants. In both cases, the participants de-
monstrated a preference on 4/10 trials, and had one trial in which their
response did not match any strategy (i.e., they chose one of the 3 other
arms). These erroneous responses may represent an unsuccessful at-
tempt to use their preferred strategy, or a lapse in the participant’s
attention. Therefore, the more liberal threshold does seem to capture
strategy preference, while also potentially accounting for small errors.
Strategy acquisition in the current study was quick, with 75% of
participants using their preferred strategy from the first trial. Data from
the ‘guess’ trials, which were presented before testing began, indicate
that most participants did not immediately identify the cue (coloured
sphere) and were not biased by any particular components of the maze
(e.g., drawn towards a particular window), suggesting that preference is
an automatic response that participants bring to the experiment. Not
only was acquisition fast, but strategy preference was relatively stable
for all three strategies. Even with a change in start location, and plat-
form location, all of the participants who demonstrates an allocentric
preference in the first configuration maintained this preference into the
second configuration. Furthermore, 65% of participants remained
stable for the two egocentric strategies even with a change in cue colour
and direction of turn. Conversely, Iaria et al. [12] found that a large
proportion of participants shifted from a ‘spatial strategy’ (allocentric)
to a ‘non-spatial strategy’ (egocentric-response) with practice in a vir-
tual RAM. A similar shift has also been noted in rodents [59,60], and
has been interpreted as evidence for a difference in the relative pro-
cessing speed of the two systems (hippocampal-based allocentric navi-
gation being acquired faster than striatal-based egocentric-response
based navigation). However, the current results demonstrate no such
differences in the acquisition times for the two strategies. It may
therefore be that this ‘shifting’ is a further bias of the RAM, which is
overcome in the Hex Maze. Together, these results provide further
support that the configuration of the maze does not bias participants
towards a particular strategy but that strategy preference is idiosyn-
cratic to the individual.
Finally, the Hex Maze allows a direct comparison between strategy
preference and strategy competence. While strategy preference refers to
the strategy used most often by a participant, strategy competence re-
fers to accuracy and efficiency using a particular strategy. Previously,
these two constructs have been difficult to isolate, which has influenced
how individual differences in spatial navigation are interpreted. For
example, it is difficult to determine whether the male tendency towards
allocentric navigation and a female tendency towards egocentric navi-
gation is due to differences in strategy preference, competence or both
[61,43,44,46]. In the Hex Maze, strategy competence can be measured
using the forced choice probes, allowing for a distinction between
preference and competence.
The current results indicate that participants performed better on
the forced choice probe for their preferred strategy than both their non-
preferred strategies, and better than participants who preferred another
strategy. This association suggests that strategy preference may be
guided by strategy competence or alternatively, that strategy compe-
tence may be greatest in the preferred strategy due to greater practice
with that strategy. There were no sex differences in this bidirectional
10
Behavioural Brain Research xxx (xxxx) xxx–xxx
association.
Finally, an intriguing finding from the independent measure of
strategy competence is that a large proportion of participants not only
appear to be concurrently learning, but demonstrate proficiency in
more than one strategy. Latent spatial learning has previously been
difficult to measure in rodents [62–66], However, the current study
demonstrates that, in the same experimental environment, many par-
ticipants who have a preference for one strategy are concurrently ac-
quiring information regarding a second, and maybe a third strategy
with no additional training. Thus, preference did not preclude learning
another strategy, as both allocentric navigators and egocentric-cue
navigators demonstrated greater than chance performance in the forced
choice probe for another strategy. Additionally, almost half of the
participants received a perfect score for the forced choice probes of
their preferred strategy and at least one other. Neither of these findings
were significantly affected by sex. This is consistent with dual-solution
mazes that have previously been used to demonstrate that rodents are
able to acquire both ‘cognitive’ (allocentric) and ‘stimulus-response’
(egocentric-response) strategies, and that manipulating experimental
factors such as stress and the amount of training can influence the
dominant strategy of use [67,68]. However, it is difficult to comment
on the distinction between preference and competence based on rodent
studies. To the best of the authors knowledge, the Hex Maze is the first
human spatial navigation task that allows for this distinction to be
made for three navigational strategies.
Interestingly, participants who preferred the egocentric-response
strategy performed worse overall on the probes than the allocentric
strategy users, and were the only group not to perform better than
chance on one of the strategy probes other than their preference. This is
somewhat consistent with the premise that this strategy is ‘non-spatial’
[12], and is the only strategy that does not depend on spatial in-
formation gained from specific environmental features. Both the ego-
centric-cue and allocentric strategies require an awareness of environ-
mental features (albeit, different environmental features), and therefore
navigators using these strategies may be more likely to concurrently
‘pick up’ different types of spatial information from other features. In
light of this, it is difficult to determine whether participants were aware
that they were picking up information useful for another strategy, and
actively choosing not to use it, or whether they were unaware and ig-
nored the information until specifically probed. It will be of interest for
future studies to assess this further. Nevertheless, these findings de-
monstrate that the forced choice probes are able to measure concurrent
or incidental learning which is latent in participants, and that, im-
portantly, strategy preference does not characterize the limits of
strategy competence.
An unexpected finding from the current study was that there was
little evidence for the development of a cognitive map of the environ-
ment outside the arena. The Room Reconstruction and Sense of
Direction tasks have been used in previous studies employing a virtual
MWM [38,50,40,41] and have largely demonstrated good development
of a cognitive map in healthy participants. The cause of this discrepancy
between the Hex Maze and the MWM is unclear. Nevertheless, it ap-
pears that while the Hex Maze may be more sensitive to strategy ac-
quisition, preference, and competence, the MWM may encourage the
development of a more holistic cognitive map.
It could be argued that in the absence of explicit reports from par-
ticipants on strategy use, we cannot determine whether a response on a
given trial was a correct use of a particular strategy, or an error using an
alternative strategy. However, it would be expected that if the errors
were random, then they should not manifest consistently. Participants
in the Hex Maze were highly consistent in strategy use within each
maze configuration. If the errors were systematic, then they would not
be expected to be consistent across maze configurations. As the spatial
relationship between the correct arms for each strategy changes be-
tween configurations, it seems unlikely that a participant’s response
would erroneously align with the same strategy across both
M.J. Spriggs et al.
configurations. Additionally, participants performed better on the
forced choice probe for their preferred strategy than either of the other
strategies. If the responses of participants were due to errors, their
performance would have been poor on the corresponding probe. As
such, the argument of errors in strategy uses seems non-parsimonious.
The primary contribution of the Hex Maze to future research is that
it is able to index and characterize three different spatial navigation
strategies. Previous research has only focused on the distinction be-
tween allocentric and egocentric navigation, however these results
demonstrate cue-based egocentric navigation and response-based ego-
centric navigation can (and should) be differentiated. As these strate-
gies are understood to have different neural correlates, the ability of the
Hex Maze to distinguish between the strategies will be important when
studying factors that may influence these brain areas, such as stress
[58,54,42], genetics [33,69,70] brain injury [71–73], ageing [74,75]
and neurodegenerative disease [76,28,77,78]. It will also be of interest
to future studies to explore the relationship between the measures of
strategy usage afforded by the Hex Maze and navigational ability, using
a measure such as the Santa Barbara Sense of Direction Scale [79]. As
many aspects of the task can be modified with simple key strokes (e.g.,
trial order, configuration, available cues), the Hex Maze is well suited
for use across different populations. Additionally, the repetitive nature
of the ‘probe’ (Show-Me) trials renders this task favorable for neuroi-
maging and neurophysiology studies that require averaging over a large
number of trials.
It is important to note a couple of limitations of the current study.
Firstly, the sample population was comprised of English first-language
university students in Canada, which limits the generalizability of the
results. Secondly, it could be argued that the sample size was small for a
methodological study, and that the lack of significant sex effects could
be due to insufficient power. However, previous research from our lab
has established a sample size of 15–20 per group is sufficient for de-
tecting sex differences [80,40,41]. Additionally, with the potential for
clinical applications, it was important to demonstrate the sensitivity of
the Hex Maze with a sample size close to what would be seen in such
studies. This sensitivity is demonstrated by the included effect sizes for
significant effects. Nevertheless, it is important to acknowledge that,
based on the current results, we cannot claim an absence of sex dif-
ference (i.e., one cannot claim support for the null hypothesis) and it
will be important for future research to substantiate the current study
with a larger cohort. Third, we did not include a measure of naviga-
tional ability in the current study. Previous studies using our virtual
MWM have found no relationship between maze performance and
questionnaires of navigational ability [50,81]. As the relationship be-
tween navigational ability and strategy usage remains elusive [82], it
was not a pertinent to the main goals of the current study to include
such a measure. However, as there may be some influence of naviga-
tional ability on strategy use, this is an important avenue for future
studies. Finally, the Hex Maze did not appear to encourage develop-
ment of a cognitive map of the environment outside the arena. This
could suggest that the allocentric cues provided by the Hex Maze are
not actually being used within an allocentric cognitive map. However,
this seems unlikely as the allocentric environment is similar to that used
in previous MWM studies where participants have demonstrated good
cognitive map development [38,50,40,41]. It is unclear why there is
such a discrepancy between the Hex Maze and the MWM in this regard,
and it will be important for future studies to examine this further.
Here, we present a novel paradigm for exploring human spatial
navigation; the Hex Maze. In the current study we were able to char-
acterize not only strategy preference and rates of learning, but also the
stability and acquisition of strategy use. Together our results show that
across all three strategies, acquisition was quick and relatively stable,
and while there was some indication of sex differences in strategy
preference, there appear to be no evidence for significant sex differ-
ences in any of our novel measures of acquisition, stability or multi
strategy competence. Additionally, we have demonstrated that strategy
11
Behavioural Brain Research xxx (xxxx) xxx–xxx
preference and competence are not analogous, and that multiple stra-
tegies can be learnt concurrently and with proficiency. It is envisaged
that this paradigm will be widely applicable and will be beneficial in
furthering our understanding on the cognitive and neural basis of
human navigation, and the factors that may influence it.
Funding
MJS is supported by Brain Research New Zealand PHD scholarship.
Conflicts of interest
There are no conflicts of interest to report.
Acknowledgements
The authors would like to thank Maureen Krulak for data collection.
Additionally, the authors would like to thank Thomas Ferguson and
Dustin Van Gerven for assisting in the development of the virtual en-
vironment.
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