Cognition 146 (2016) 431438

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Cognition
journal homepage: www.elsevier.com/locate/COGNIT

Believe it or not: Moving non-biological stimuli believed to have human

origin can be represented as human movement
E. Gowen a,, E. Bolton a, E. Poliakoff b
a
Faculty of Life Sciences, University of Manchester, UK
b
School of Psychological Sciences, University of Manchester, UK

a r t i c l e i n f o a b s t r a c t

Article history: Does our brain treat non-biological movements (e.g. moving abstract shapes or robots) in the same way
Received 9 December 2014 as human movements? The current work tested whether the movement of a non-biological rectangular
Revised 28 July 2015 object, believed to be based on a human action is represented within the observers motor system. A
Accepted 13 October 2015
novel visuomotor priming task was designed to pit true imitative compatibility, due to human action
Available online 9 November 2015
representation against more general stimulus response compatibility that has confounded previous belief
experiments. Stimulus response compatibility effects were found for the object. However, imitative
Keywords:
compatibility was found when participants repeated the object task with the belief that the object was
Visuomotor priming
Automatic imitation
based on a human finger movement, and when they performed the task viewing a real human hand.
Mirror neuron system These results provide the first demonstration that non-biological stimuli can be represented as a human
Stimulus response compatibility movement if they are believed to have human agency and have implications for interactions with
Humanrobot interaction technology and robots.
Crown Copyright 2015 Published by Elsevier B.V. All rights reserved.

1. Introduction such as in healthcare, education and entertainment (Andrade et al.,

It is well known that observation of a human action can influ-
ence the observers own motor system. For example, observing
another persons action activates brain areas involved in execution
of that action (Gazzola & Keysers, 2009; Kilner, Neal, Weiskopf,
Friston, & Frith, 2009) and can interfere with or facilitate move-
ment production (Brass, Bekkering, & Prinz, 2001; Sturmer,
Aschersleben, & Prinz, 2000). These effects are thought to be due
to the mirror neuron system (MNS) present within the premotor
cortex and inferior parietal lobe that responds during both obser-
vation and execution of an action (Buccino et al., 2001; Rizzolatti
& Craighero, 2004; Van Overwalle & Baetens, 2009). Recently, there
has been increasing interest in whether non-biological stimuli (e.g.
abstract shapes or robots) are processed in a similar way to human
actions, leading to non-biological movements being represented
within the observers motor system (Gowen & Poliakoff, 2012;
Press, 2011). Measuring whether non-biological movements are
represented in a similar way to human actions could indicate the
success of humanrobot interaction which is particularly relevant
as humanoid robots are likely to increasingly play a role in society,
Corresponding author at: Carys Bannister Building, Dover Street, Manchester
M13 9PL, UK.
E-mail address: emma.gowen@manchester.ac.uk (E. Gowen).
2014; Chaminade & Cheng, 2009; Dautenhahn, 2007; Tapus,
Mataric, & Scassellati, 2007). More controversially, representing
the action of a non-human agent may suggest the attribution of
characteristics associated with humans such as mental states to
non-human stimuli (Chaminade & Cheng, 2009). In this work, we
address whether belief that a non-biological stimulus is based
on a human action produces action representation, using a
behavioural visuomotor priming task.
In visuomotor priming, also termed Automatic Imitation,
observing and performing a compatible action (e.g. lifting ones
index finger while observing another index finger move upwards)
facilitates reaction times, whereas reaction times are slowed when
observing a movement incompatible with a performed action (e.g.
lifting ones index finger while observing a finger press; Brass
et al., 2001). As visuomotor priming is likely to result from activa-
tion of the MNS (Catmur, Walsh, & Heyes, 2009; Heyes, 2011), it
provides a behavioural measure of whether an action is repre-
sented within the observers motor system. Although previous
studies have compared visuomotor priming for human and non-
biological movements (Gowen, Bradshaw, Galpin, Lawrence, &
Poliakoff, 2010; Jansson, Wilson, Williams, & Mon-Williams,
2007; Press, Bird, Flach, & Heyes, 2005) these are confounded by
Stimulus Response Compatibility effects, whereby responses are
faster to spatially or directionally aligned stimuli (Cho & Proctor,

http://dx.doi.org/10.1016/j.cognition.2015.10.010
0010-0277/Crown Copyright 2015 Published by Elsevier B.V. All rights reserved.
432 E. Gowen et al. / Cognition 146 (2016) 431438
2003). Consequently, visuomotor priming to non-biological stimuli
could result purely from stimulus response compatibility effects in
the absence of action representation (Jansson et al., 2007).
Models of visuomotor priming share the idea that priming
occurs along a visuomotor route, transforming visual input into a
motor response and that priming produced by stimulus response
compatibility and imitative compatibility are dissociated with
the latter involving the MNS (Gowen & Poliakoff, 2012; Heyes,
2011; Wang & Hamilton, 2012). This visuomotor route is modu-
lated by top-down factors such as attention, prior knowledge and
social cognitive processes which can exert influence at the early
sensory input stage or at the later motor, output stage (Gowen &
Poliakoff, 2012; Heyes, 2011). One top-down influence, termed
belief refers to prior knowledge or assumptions that a person has
about the observed stimulus. For example, visuomotor priming is
greater if a person believes (having received explicit instruction)
that the non-biological stimulus is created from a human move-
ment (Liepelt & Brass, 2010; Shen, Kose-Bagci, Saunders, &
Dautenhahn, 2011; Stanley, Gowen, & Miall, 2007), whereas belief
that a hand is virtual can reduce priming (Longo & Bertenthal,
2009). A more spontaneous or implicit form of belief could also
occur for non-biological stimuli that have human characteristics
(e.g. a robot) or for non-biological stimuli that are presented in a
similar context to a previous human stimulus (Stanley et al.,
2007). On the one hand, these results could suggest that belief pro-
duces action representation for non-biological stimuli by activating
the MNS at the input stage or enhancing the MNS at the output
stage. However, it could be that implicit or explicit belief merely
alters attention to the stimulus movement, which either enhances
or reduces stimulus response compatibility effects via the input
route, without activating the MNS (Heyes, 2011; Press, 2011). Con-
sequently, it is still unknown whether a non-biological movement
can produce action representation equivalent to a human
movement.
The aim of this work was to resolve these issues by separating
imitative compatibility, due to action representation, from more
basic stimulus response compatibility effects. We used a modified
version of the visuomotor priming task where participants observe
an index finger or blue rectangular object moving upwards or
Fig. 1. Time course of one trial for the hand (top) and object stimuli (bottom). Trial starts at left of picture in neutral position and shows a downward movement for both
stimulus types. The yellow go signal is presented for 80 ms at 0, 120 or 280 ms following the start frame. A second end frame is presented for the 280 ms go signal. Pictures to
right of dashed line show position of flash on stimuli. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this
article.)
downward and must respond when they observe a go signal in
the form of a yellow flash (Fig. 1). Participants viewed a right hand
rotated 90 degrees counter clockwise (from the participants view-
point), in a thumb up orientation and were required to make a
key press response with their left hand. This stimulus orientation
and response combination separated three stimulus response com-
patibility effects from imitative compatibility (Fig. 2). Directional
stimulus response compatibility effects were removed by rotating
the hand so that up/down index finger movements now became
left/right movements. However, rotating the hand introduces two
further potential stimulus response compatibility effects (i) left-
down and up-right stimulus response pairings are faster (orthogo-
nal stimulus response compatibility; Weeks & Proctor, 1990); (ii)
an advantage when the stimulus and response are on the same side
of space (Simon effect; Simon, 1990).
By using a thumb up orientation together with a left handed
pressing response (Fig. 2) we were able to isolate imitative com-
patibility from both orthogonal stimulus response compatibility
and the Simon effect. Thus, when the finger moves leftward across
the screen, this is compatible in terms of the Simon effect (left
advantage due to using left hand to response) and orthogonal stim-
ulus response compatibility (down-left advantage when pressing
button), but is imitatively incompatible (downward response,
observing upward finger movement). However, when the finger
moves rightward across the screen, this is incompatible in terms
of the Simon effect and orthogonal stimulus response compatibil-
ity, but is imitatively compatible (downward response, observing
downward finger movement).
Participants responded to the go signal under three stimulus
conditions. Firstly, they carried out the task while observing the
object (object condition). Next, they responded while observing
the object following a belief manipulation informing them that
the object was based on the movement of a human index finger
(belief condition). Lastly, they performed the task with the real
human hand (hand condition). We hypothesized that orthogonal
spatial compatibility/Simon effects would be present in the initial
object condition and that imitative compatibility would be present
for the hand stimulus. However, in the belief condition, there were
two possibilities: (1) There would be an increase in orthogonal
 E. Gowen et al. / Cognition 146 (2016) 431438
Left
Imitation=incompatible
Simon effect=compatible
Orthogonal =compatible
Response: press (down) using left hand
Fig. 2. Dissociating imitative and stimulusresponse compatibility. The participants response (downward key press) shares imitative compatibility with the finger when it
moves rightward across the screen (downward finger movement), but is incompatible when the finger moves leftward across the screen (upward finger movement). In
contrast, stimulus response compatibility effects are compatible with a leftward movement: the leftward stimulus movement is compatible with the left hand response
(Simon effect) and the downward response (orthogonal spatial compatibility; right-up left-down).
spatial compatibility/Simon effect if the belief effect produced
increased attention to the stimulus or (2) Imitative compatibility
in the opposite direction, similar to the hand would be present if
the belief effect is caused by representation of a human movement.
In this case, as stimulus response compatibility can be stronger
than imitative compatibility (Longo & Bertenthal, 2009; Jimenez
et al., 2012) we expected the imitative compatibility effects to be
smaller than previously reported, but importantly in the opposite
direction to stimulus response compatibility effects.
2. Methods
2.1. Participants
20 right-handed healthy individuals were recruited for the
study, but the data of two participants was excluded due to failure
to comply with instructions or a high percentage (>20%) of
responses made prior to the go signal. The mean age (SD) of the
remaining 18 participants (9 female) was 22.94 (4.71). As the task
was new, a power calculation was not possible. However, an N of
20 was chosen based on earlier versions of the task that showed
a significant interaction between stimulus condition (object/
hand/belief) and compatibility, with a large effect size of
gp2 = 0.2. The study was approved by the University of Manchester
Research Ethics Committee and carried out in accordance with the
provisions of the World Medical Association Declaration of
Helsinki.
2.2. Stimuli
Participants sat 80 cm from a flat screen monitor, upon which
sequences of stimuli were displayed using Presentation Software
(Neurobehavioral Systems). Stimuli consisted of either a right hand
rotated 90 degrees counter clockwise, in a thumb up orientation
or a blue rectangular object in an equivalent position (Fig. 1). Par-
ticipants observed the object or the index finger of the hand mov-
ing in a leftward or rightward direction across the screen (i.e.
up/down movement of the finger) and were required to make a
press response using their left index finger, when they observed
a yellow flash appear (Fig. 1). Key presses were recorded on a sep-
arate keypad that was positioned centrally in front of the computer
screen, so that the participants left hand was aligned with the cen-
tre of the image on screen. This stimulus orientation and response
combination separated the Simon effect (left advantage due to
using left hand to response) and orthogonal spatial compatibility
effects (down-left advantage when pressing button) from imitative
433
Right
Imitation=compatible
Simon effect=incompatible
Orthogonal =incompatible
compatibility (right stimulus movement, representing downward
movement, imitatively compatible with downward key press)
(Fig. 2).
The hand stimuli were created by converting digital .avi files
into a sequence of 9 still frames consisting of an initial start frame
(finger in a neutral position) and 7 movement frames. The initial
start frame was presented for 1600 ms and was identical for
upward and downward movements, ensuring it was not possible
to predict the movement direction. The 7 movement frames (pre-
sented for 40 ms each) depicted the finger making an upward
(33 pixels or 12.6 mm) or downward (27 pixels or 10.3 mm) move-
ment. The movement of both the hand and object had a biological
profile, accelerating toward the middle and decelerating towards
the end. To plot the trajectory in the object movement condition,
a blue rectangle, was positioned over the moving index finger in
each frame. However, the hand was not visible in the resulting
clips in this condition; instead the rectangle was shown alone over
a background constructed to resemble that used in the hand clips.
The luminance (142 cd/m2) and size (5.5  1.8 cm) of the rectangle
were matched to the finger of the hand. The yellow flash
9.8  1.6 cm and was presented for 80 ms. It was positioned over
the finger or object so that it covered the full extent of the move-
ment (Fig. 1).
2.3. Procedure
The experiment was split into three blocks. Firstly, the object
was presented, followed by a questionnaire asking whether partic-
ipants found that the rectangle made them think of a human finger
movement (Table 2, questions 12). Secondly, participants were
told that the object movement was generated from a human finger
movement. This was demonstrated using the experimenters own
hand, as well as using a printed picture depicting how the hand
had been rotated. Therefore, the way the hand was orientated
and the direction of movement (upwards = rightwards; down-
wards = leftward) was made clear to the participants and the
object was referred to as the finger, moving upwards/down
wards. They then repeated the task. Following the belief condition,
participants completed a second short questionnaire, asking to
what extent they believed that the object movement was based
on a human finger movement (Table 2, questions 35). Thirdly,
participants observed the hand stimulus.
Each trial consisted of the 9 frames detailed above depicting the
moving finger or object (Fig. 1). The yellow flash go signal appeared
at one of three different stimulus onset asynchronies (SOAs)
following the start frame (0, 120, 280 ms). The three SOAs were
434 E. Gowen et al. / Cognition 146 (2016) 431438

chosen to assess the strength of priming at the start, middle and Table 1
end of the movement. As the 280 ms SOA occurred after the end Reaction times for the different conditions.

frame, a second end frame was presented. In the case of no-go tri- Condition SOA Reaction time (ms) [95% Compatibility effect (ms)
als, the flash did not appear. For baseline trials, no movement (ms) confidence intervals] [95% confidence intervals]
frames were presented. Instead, the flash appeared after the start Object 0 355.03 [334.7, 375.4] 5.25 [ 16.89, 6.38]
frame, followed again by the start frame for 40 ms. Both the no- 349.78 [328.9, 370.6]
go and baseline trials decreased predictability of the flash go signal. 120 331.19 [308.2, 354.2] 13.48 [ 24.31, 2.64]
317.77 [298.6, 336.8]
Trials were terminated if the participant made a response or if no 280 322.98 [307.5, 338.5] 11.34 [ 24.86, 2.06]
response occurred within 2000 ms of the appearance of the flash. 311.58 [294.4, 328.8]
Between each trial a blank screen was displayed for 2000 ms. Belief 0 389.64 [353.0, 426.3] 0.99 [ 16.69, 18.67]
Compatibility was determined by whether the direction (press) 390.63 [346.3, 435.0]
was compatible or incompatible with the finger stimulus move- 120 334.33 [301.8, 366.9] 12.3 [ 0.12, 24.72]
ment (upward/downward). As detailed in Fig. 2, upward finger 346.63 [305.7, 387.6]
280 332.21 [305.1, 359.3] 5.32 [ 9.8, 20.46]
movements correspond to leftward movements across the screen
337.53 [303.5, 371.5]
for all three stimuli, while downward movements correspond to
Hand 0 378.24 [343.7, 412.8] 4.48 [ 20.18, 11.22]
rightward movements. If Reaction Times (RTs) are faster when
373.76 [345.4, 402.1]
the stimulus was moving in the rightward (compatible) compared 120 319.93 [299.2, 340.6] 7.6 [ 11.1, 26.3]
to leftward (incompatible) direction, this will result in positive 327.53 [292.4, 339.6]
compatibility effects (IncompatibleCompatible RTs). In this case, 280 316.01 [292.4, 339.6] 9.15 [ 9.18, 27.48]
movements have been facilitated by representation of an upward 325.16 [301.5, 348.8]

moving finger movement and indicate the presence of imitative Note: Bold = compatible responses.
compatibility. However, if RTs are faster when the stimulus was
moving in the leftward (incompatible) compared to rightward
(compatible) direction, this will result in negative compatibility Table 2
Questionnaire results.
effects (IncompatibleCompatible RTs). Here, movements have
been facilitated by the direction/position of the stimulus and indi- Question Mean One
cate the presence of stimulus response compatibility effects. score [95% sample t
CI] test
Therefore, positive compatibility effects represent imitative com-
patibility, whereas negative compatibility effects represent stimu- 1 During the first half of the experiment, did you 0.12 t = 1.46;
lus response compatibility (the left/right movement). at any time think that the moving rectangle [ 0.1, 0.3] p = .16
might represent a human finger movement?
The experimental conditions were stimulus condition (object, 2 To what extent, if at all did the movement of 0.43 t = 1.58;
belief, hand)  compatibility (compatible, incompatible)  SOA the rectangle remind you of a human finger [ 0.1, 1.0] p = .13
(0, 120, 280 ms). For each block, the 6 compatibility  SOA condi- movement?
tions were presented 12 times, together with 12 baseline and 12 3 Did you think of the rectangle as more of a 7.59 [6.5, t = 15.4;
block representation of a moving finger, 8.6] p < .001
no go trials giving a total of 96 trials. These trials were split into
compared to when you had seen the rectangle
2 mini-blocks of 48 trials and a short break was given after each for the first time?
mini-block. A longer break was given after each block. For the 4 To what extent did you think of the moving 7.0 [5.8, t = 12.5;
entire experiment there were 216 experimental trials, 36 baseline rectangle as being a human finger compared to 8.2] p < .001
trials and 36 no go trials, totalling 288 trials. when you had seen the block for the first time?
5 When you were told the rectangle represented 7.43 [6.3, t = 13.4;
a finger movement, how much did you believe 8.6] p < .001
that this was true?
3. Results
Note: Questions 12 were asked following the object condition, questions 35 fol-
lowing the belief condition. For questions 1, 2, 4, 5, a score of 0 represents not at
RTs for trials were removed if the participant made an incorrect all and a score of 10 represents very much. For question 3, a score of 0 represents
response, did not respond, if the response was longer than 1000 ms disagree and 10 represents agree.
or shorter than 150 ms or lay outside of 2.365 standard deviations
of the participants mean RT (Van Selst & Jolicoeur, 1994). This
resulted in a loss of 0.39% of trials. compatible and incompatible reaction times (across SOA) revealed
A repeated-measures ANOVA with factors of stimulus condition that a significant negative compatibility effect was present for the
(object, belief, hand), SOA and compatibility was conducted on object condition (t(53) = 3.077, p = 0.003; d = 0.24), but that com-
mean RTs (Table 1). There was a main effect of SOA (F(2,34) patibility effects were not significant for the belief or hand condi-
= 51.026, p < .0005, gp2 = .75), indicating that while RTs were sig- tion (t(53) < 1.5, p > 0.14; d < 0.08) (Bonferroni correction = 0.02).
nificantly slower at 0 ms compared to 120 ms (t(17) = 11.202, These results show that while negative compatibility effects, repre-
p < .0005) and 280 ms (t(17) = 5.193, p < .0005), there was no sig- senting stimulus response compatibility and Simon effects were
nificant difference between RTs at 120 ms and 280 ms (t(17) observed for the object, positive (non significant) compatibility
= 1.017, p = .323) (Bonferroni correction = 0.02). Importantly, there effects, representing imitation of a human action were observed
was a significant interaction between stimulus condition and com- for both the belief and hand condition.
patibility (F(2,34) = 5.461, p = .009, gp2 = .24) (Fig. 3). Paired t-tests Finally, there was a significant interaction between stimulus
conducted between mean compatibility effects in each stimulus condition and SOA (F(4,72) = 2.850, p = .049, gp2 = 0.13). Paired t
condition (averaged across SOA) indicated that compatibility tests comparing differences in RTs between the stimulus pairings
effects were significantly smaller for the object ( 10.04 ms) than at each SOA indicated that the difference between belief and hand
both the belief (6.20 ms) (t(17) = 3.024, p = .008; d = 0.6) and (14.4 ms) was significantly smaller than the difference between
the hand condition (4.09 ms) (t(17) = 2.535, p = .021; d = 0.48), the object and hand conditions (23.6 ms) for the 0 ms SOA only
while there was no significant difference between the belief and (t(35) = 3.044, p = 0.004, Bonferroni correction = 0.005; d = 0.71).
hand conditions (t(17) = .416, p = .628; d = 0.06) (Bonferroni Simple main effects comparisons at 0 ms SOA also revealed signif-
correction = 0.02). Simple main effects (paired t tests) comparing icantly faster responses for the object (352.4 ms) than both the
 E. Gowen et al. / Cognition 146 (2016) 431438
Fig. 3. Compatibility effects for each of the three stimulus conditions. Positive compatibility effects indicate imitative compatibility, whereas negative compatibility effects
indicate stimulus response compatibility. Standard error bars are shown (Morey, 2008).
belief (390.14 ms; t(35) = 3.044, p = .004; d = 0.62) and the hand
(376.0 ms; t(35) = 2.98, p = .005; d = 0.46), but no difference
between the belief and hand condition (t(35) = 1.46, p = .15;
d = 0.2) (Bonferroni correction = 0.02). There were no significant
interactions between compatibility and SOA (F(2,34) = 0.37,
p = .69, gp2 = .02) and condition, compatibility and SOA (F(4,68)
= 1.02, p = .40, gp2 = .0.06).
Prior to the belief instruction, participants did not associate the
object with a finger movement (Table 2). However, following the
belief instruction, scores were significantly different from 0, indi-
cating that participants considered that the object represented a
human movement.
3.1. Control conditions
3.1.1. Control Experiment 1: Order effects
To test for possible practice/order effects when the object was
viewed for a second time in the belief condition, compatibility
effects for the first and last six trials of the object condition were
calculated and averaged across participants. Compatibility effects
were negative for both first (0 ms = 2.65, 120 ms = 10.7,
280 ms = 8.52) and last (0 ms = 8.42, 120 ms = 16.8,
280 ms = 17.7) six trials. This suggests that the positive compat-
ibility found in the belief condition was not due to a practice effect.
This was further confirmed by presenting two separate blocks of
the object condition to a separate group of naive participants
(n = 22, 15 females, mean age (SD) = 21.7 3.8 years). Compatibil-
ity effects for the second object block (mean across
SOA = 4.59 ms, standard error = 2.89) were significantly lower
than the belief condition in the main experiment (mean = 6.2 ms,
standard error = 4.13; t = 2.8, p = 0.04).
3.1.2. Control Experiment 2: Weaker positive compatibility effects
Although the negative compatibility effects for the object were
significantly smaller than the positive compatibility effects found
for the belief and hand conditions, these positive compatibility
effects were non-significant. This is likely to be due to the require-
435
ment to pit orthogonal spatial compatibility and Simon effects
against imitative compatibility. To test this we presented a new
group of 19 participants (9 female, mean age (SD): 22.1 (3.01)
with identical hand stimuli to previously described except that
the hand was a left rather than right hand in thumb down position
(Fig. 4a). Participants responded to the appearance of the yellow
flash by pressing down their right index finger. Using this setup,
when the finger moves downward (rightwards), the participants
pressing response has both imitative and stimulus response
(Simon effect) compatibility, whereas when the finger moves
upward (leftward), the pressing response is incompatible for both
imitation and the Simon effect. As imitative and stimulus response
compatibility effects are combined, we would expect larger, signif-
icant positive compatibility effects. In line with this, a compatibil-
ity  SOA ANOVA revealed a main effect of compatibility (F(1,18)
= 11.87, p = .0003, gp2 = 0.4) and an interaction between compati-
bility and SOA (F(2,36) = 4.08, p = .03, gp2 = 0.19) (Fig. 4b). Simple
main effects (paired t tests) revealed significant positive compati-
bility effects for the 280 ms SOA only (t = 4.6, p < 0.001;
d = 0.67, Bonferroni correction = 0.02). This compatibility effect
was twice as large as the compatibility effects for the finger in
our main experiment (Fig. 3), highlighting that imitative and stim-
ulus response compatibility effects are likely to be combined,
resulting in strong compatibility effects when in the same direction
(Control Experiment 2), but weaker compatibility effects when in
opposite directions (Main Experiment).
4. Discussion
Our results provide the first behavioural demonstration that
observation of a non-biological stimulus, believed to have human
agency, leads to imitative compatibility as opposed to stimulus
response compatibility. More generally, by separating imitative
compatibility from stimulus response compatibility, this is the first
evidence to show that priming caused by non-biological stimuli
can be generated by imitative compatibility. When participants
observed the object, prior to the belief manipulation, negative
436 E. Gowen et al. / Cognition 146 (2016) 431438
(a) Le Right
Imitaon=incompable
Simon eect=incompable
Orthogonal =compable
Response: press (down) using right hand
Fig. 4. (a) Left hand in thumb down position, showing co-occurrence of imitation and Simon effect compatibility when responding with right finger press. (b) Compatibility
effects across the three different SOAs, showing large compatibility effects at the 280 ms SOA. Standard error bars are shown.
compatibility effects were present due to stimulus response com-
patibility produced by the direction and position of the stimulus.
However, following the belief manipulation, compatibility effects
became numerically positive and of a similar magnitude to the
subsequently viewed hand stimulus. Importantly, compatibility
effects for both the belief and hand condition were numerically
positive and significantly different to the object condition. If the
belief effect was simply caused by increased attention to the stim-
ulus enhancing bottom-up stimulus response compatibility effects,
the belief condition should have produced larger negative compat-
ibility effects, similar to the object condition. Consequently, our
results indicate that belief that a non-biological stimulus is
human-generated causes action representation, validating previ-
ous belief studies unable to conclusively rule out the influence of
attention (Liepelt & Brass, 2010; Shen et al., 2011; Stanley et al.,
2007). More generally, our hand condition adds to accumulating
data that imitative compatibility cannot be explained by stimulus
response compatibility effects (Catmur & Heyes, 2011; Jimenez
et al., 2012).
While significant negative compatibility effects were observed
for the object, it should be noted that the positive compatibility
effects for the belief and hand condition were non-significant.
However, it is unsurprising that we obtained smaller imitative
compatibility effects since imitative compatibility was pitted
against a number of stronger stimulus response compatibility
effects (Longo & Bertenthal, 2009; Jimenez et al., 2012). Indeed,
when the direction of stimulus response compatibility and imita-
tive compatibility were complementary as in our Control Experi-
ment 2, positive compatibility effects were significant and more
than twice as large as the finger stimulus in the main experiment.
As with previous work, this highlights that stimulus response com-
patibility can mask imitative compatibility when pitted against
one another (Longo & Bertenthal, 2009; Jimenez et al., 2012) and
that significant positive compatibility effects are more likely when
imitative compatibility is confounded with stimulus response
compatibility. Furthermore, the overall pattern of our results sup-
ports the theory that belief causes action representation as
opposed to increased attention to stimulus properties. Firstly, the
direction of the compatibility effects was positive for both the
belief and hand condition and differed significantly from the neg-
ative compatibility effects for the object. In addition, reaction times
were similarly slower in the belief and hand conditions compared
to the object conditions, raising the possibility that although they
were not instructed to do so participants were mentally rotating
the belief and hand stimuli to match their own posture. Previous
work indicates that making explicit 90 mental hand rotations pro-
duces longer reaction times (Parsons, 1994) than the difference
between the belief and hand compared to object conditions. How-
ever, these smaller differences may result from the rotation not
being necessary for the task, as well as viewing a consistent hand
(b)
Imitaon=compable
Simon eect=compable
Orthogonal =incompable
rotation on every trial. As the belief condition closely follows the
pattern of the hand condition, this supports the notion that the
action is represented in the belief condition. Secondly, the finding
of similar compatibility effects for the first and second half of the
trials in the object condition argues against practice effects or
decreased attention to spatial/directional (stimulus response com-
patibility) properties leading to a reduction in stimulus response
compatibility effects in the belief condition. This conclusion is fur-
ther supported by significantly different compatibility effects
between the belief condition and the second object block of Con-
trol Experiment 1. Previous belief experiments that have not sepa-
rated stimulus response compatibility and imitative compatibility
(Liepelt & Brass, 2010; Shen et al., 2011; Stanley et al., 2007) also
indicate that this possibility is unlikely. These studies show
increased compatibility effects for the belief condition, whereas a
reduction in the influence of stimulus response compatibility
effects would have resulted in smaller compatibility effects.
Our results advance existing models detailing how priming is
influenced by top-down factors (Gowen & Poliakoff, 2012; Heyes,
2011; Wang & Hamilton, 2012). These models share the idea that
priming occurs along a visuomotor route, involving brain areas
such as the Superior Temporal Sulcus (STS), parietal and premotor
areas such as the IFG. The visuomotor route transforms visual
input into a motor response and priming produced by stimulus
response compatibility and imitative compatibility are dissociated
(Heyes, 2011), with the latter involving the MNS. Priming within
the visuomotor route is modulated by top-down factors such as
attention, prior knowledge and social cognitive processes, which
can exert influence at the early sensory input stage or at the later
motor, output stage (Heyes, 2011). There is increasing evidence
that one brain area, the Medial Prefrontal Cortex (MPFC) is likely
to modulate priming (Cross, Torrisi, Losin, & Iacoboni, 2013;
Spengler, von Cramon, & Brass, 2009; Wang, Ramsey, &
Hamilton, 2011, 2010), including during belief manipulations
(Stanley, Gowen, & Miall, 2010). As priming changed from stimulus
response compatibility during the object condition to imitative
compatibility for the belief condition, mechanisms modulating
the level of motor output are unlikely to be responsible for the
effect of belief on priming. Instead, it is more likely that belief
causes the visual input to be processed by the MNS pathway,
potentially through connections between the MPFC and STS, simi-
lar to a gating mechanism (Liepelt & Brass, 2010). In contrast, when
belief is drawn to inanimacy (Longo & Bertenthal, 2009) priming
may be channelled along non-mirror, stimulus response compati-
bility routes only.
Our behavioural results complement fMRI studies showing that
non-biological stimuli activate brain areas associated with the
MNS (Cross, Hamilton, Kraemer, Kelley, & Grafton, 2009; Engel,
Burke, Fiehler, Bien, & Rosler, 2008; Gazzola, Rizzolatti, Wicker, &
Keysers, 2007; Press et al., 2012). However, these studies are
 E. Gowen et al. / Cognition 146 (2016) 431438
unable to identify whether non-biological movement activates the
MNS as they have either not separated stimulus response compat-
ibility from imitative compatibility or do not use both observation
and execution conditions to identify the MNS. Furthermore, some
have used training paradigms to associate non-biological stimuli
with actions, whereas our study shows that action representation
of non-biological movement can occur without prior training.
Combining our task with fMRI could be used to separate brain
activity relating to MNS and stimulus response compatibility.
As imitation can relate to social affiliation (Chartrand & Lakin,
2013), it is possible that imitation could be used to measure the
success of humanrobot interaction, assessing the effect of modifi-
cations in robot design aimed at optimising these interactions
(Hofree, Ruvolo, Bartlett, & Winkielman, 2014). Our findings indi-
cate that providing stimulus response compatibility factors are
removed, measuring the level of imitative compatibility would be
a useful and possible marker for whether a robot elicits action rep-
resentation. In addition, instructions and prior knowledge may be a
useful strategy for encouraging representation of robot actions and
facilitating human acceptance of robots (Chaminade & Cheng,
2009). Supporting this, visuomotor priming has been found during
observation of a robot movement only if participants are informed
that the robot is observing their actions and consequently is en-
gaged in the interaction (Shen et al., 2011). Importantly, the pre-
sent results suggest that this belief effect is likely to be due to
representation of the robot as a human-like agent as opposed to
increased attention to the robot. In our experiment, as the object
moved using a human trajectory, it is unclear whether belief allows
access of this movement to the MNS or whether the object is pro-
cessed as human. Although this question requires testing using a
non-human trajectory, the results of Shen et al. (2011) support
the latter possibility as their robot stimulus moved with a non-
human trajectory. Together, these results suggest that a robot
can trigger human associated imitative behaviours that could lead
to higher level social representations (Chaminade & Cheng, 2009).
These implications would also apply to the situation where a robot
triggers implicit belief that they are human-like. A crucial next step
is to assess whether those humanrobot interactions that produce
imitation lead to better humanrobot acceptance. Furthermore, as
our object stimulus moved with a human trajectory, it will be
important to understand whether the belief effect is apparent for
non-biological stimuli moving with a non-human trajectory.
In summary, the current study has demonstrated that abstract,
non-biological stimuli can elicit imitative compatibility effects
equivalent to those produced for a human movement provided
that the non-biological movement is believed to be human-
generated. These findings imply that non-human movement can
be processed by the MNS, as if it were a human movement. This
has implications for understanding the control of imitation, as well
as developing successful humanrobot interaction.
Acknowledgements
The authors would like to thank Shannon Atkinson and Robyn
Dowlen for their help with data collection.
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