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Automatic Imitation: A Meta-Analysis
Article in Psychological Bulletin · March 2018

DOI: 10.1037/bul0000143
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Running head: AUTOMATIC IMITATION: A META-ANALYSIS 1
Automatic Imitation: A Meta-Analysis
Emiel Cracco, Lara Bardi, and Charlotte Desmet
Ghent University
Oliver Genschow
University of Cologne
Davide Rigoni
Ghent University
Lize De Coster
University of California, San Francisco
Ina Radkova, Eliane Deschrijver, and Marcel Brass
Ghent University
AUTOMATIC IMITATION: A META-ANALYSIS 2
Author Note
Emiel Cracco, Lara Bardi, Charlotte Desmet, Davide Rigoni, Ina Radkova, Eliane
Deschrijver, and Marcel Brass, Department of Experimental Psychology, Ghent University;
Oliver Genschow, Social Cognition Center Cologne, University of Cologne; Lize De Coster,
Department of Psychiatry, University of California, San Francisco.
This work was supported by the Research Foundation Flanders Grant
FWO14/ASP/050 awarded to the first author. The second to eight author contributed
equally to the manuscript and the order of authorship for these authors was determined at
random. We are grateful to the researchers who provided us the required data for their
studies. We are especially thankful to the researchers who provided us their unpublished
data: Henryk Bukowski, Emily Butler, Tullio Liuzza, Barbara Müller, Lior Noy, Roland Pfister,
Birgit Rauchbauer, and Federica Riva. Finally, we would like to thank Richard Ramsey for his
insightful comments on a previous draft of this manuscript.
Correspondence concerning this article should be addressed to Emiel Cracco,
Department of Experimental Psychology, Ghent University, Henri Dunantlaan 2, Gent, B-
9000. E-mail: emiel.cracco@ugent.be.
Copyright Statement
The following manuscript is accepted for publication in Psychological Bulletin. ©American
Psychological Association, 2018. This paper is not the copy of record and may not exactly
replicate the final, authoritative version of the article. Please do not copy or cite without
authors permission. The final article will be available, upon publication, via its DOI:
10.1037/bul0000143.
Abstract
Automatic imitation is the finding that movement execution is facilitated by compatible and
impeded by incompatible observed movements. In the past fifteen years, automatic
imitation has been studied to understand the relation between perception and action in
social interaction. Although research on this topic started in cognitive science, interest
quickly spread to related disciplines such as social psychology, clinical psychology, and
neuroscience. However, important theoretical questions have remained unanswered.
Therefore, in the present meta-analysis, we evaluated seven key questions on automatic
imitation. The results, based on 161 studies containing 226 experiments, revealed an overall
effect size of gz = 0.95, 95% CI = [0.88, 1.02]. Moderator analyses identified automatic
imitation as a flexible, largely automatic process that is driven by movement and effector
compatibility, but is also influenced by spatial compatibility. Automatic imitation was found
to be stronger for forced choice tasks than for simple response tasks, for human agents than
for non-human agents, and for goalless actions than for goal-directed actions. However, it
was not modulated by more subtle factors such as animacy beliefs, motion profiles, or visual
perspective. Finally, there was no evidence for a relation between automatic imitation and
either empathy or autism. Among other things, these findings point towards actor-imitator
similarity as a crucial modulator of automatic imitation and challenge the view that imitative
tendencies are an indicator of social functioning. The current meta-analysis has important
theoretical implications and sheds light on longstanding controversies in the literature on
automatic imitation and related domains.

Public Significance Statement
The current meta-analysis indicates that automatic imitation is a flexible, automatic process
that depends on how similar the actor is to the imitator. In contrast to popular belief, we
found no support for the hypothesis that automatic imitation is an indicator of social
functioning, which was studied in the form of empathy and autism spectrum disorder. These
results have important implications for our understanding of perception and action in the
context of social interaction.

The mob, when they are gazing at a dancer on the slack rope, naturally writhe and
twist and balance their own bodies, as they see him do, and as they feel that they
themselves must do if in his situation (Smith, 1759, p. 5)
Why do we imitate others? Is it that we choose to imitate them, or rather that we
cannot control the urge to do so? Interest in these questions dates back to at least the 18th
century when Adam Smith (1759) argued that spontaneous imitation can be seen as a
primitive form of sympathy. In spite of the longstanding theoretical interest, however, it was
not until the 20th century that experimental evidence started to emerge. That is, in an
ingenious study, Hull (1933) was able to demonstrate that subjects had inadvertently copied
the body movements of an experimenter reaching forward and backward during a series of
psychological tests. This finding was later extended to mannerism (Chartrand & Bargh,
1999), postures (Sheflen, 1964), facial expressions (Chartrand & Bargh, 1999), and speech
patterns (Bock, 1986), and has subsequently become known as motor mimicry (Chartrand &
Lakin, 2013; Chartrand & van Baaren, 2009). More precisely, motor mimicry is the finding
that people tend to imitate each other during social interaction (Chartrand & Lakin, 2013;
Chartrand & van Baaren, 2009; Heyes, 2011).
However, the strongest evidence that imitative tendencies cannot be controlled
comes from research on automatic imitation. More specifically, this research has shown
that people imitate others even when it impairs task performance (Brass, Bekkering, &
Prinz, 2001; Brass, Bekkering, Wohlschlager, & Prinz, 2000; Kilner, Paulignan, & Blakemore,
2003; Stürmer et al., 2000). For example, in one of the first studies, Brass et al. (2000)
instructed participants to lift their index finger in response to the number “1” and their
middle finger in response to the number “2”. At the same time, a hand on the screen also
lifted its index finger, also lifted its middle finger, or did not move (Figure 1a). The results
showed that, compared with when the hand did not move, responses were faster and more
accurate when the observed movement matched the instructed response, but slower and
less accurate when the observed movement did not match the instructed response. Given
that participants copied the observed movements regardless of whether they facilitated or
impaired response selection, these results indicate that imitation could not be controlled
and therefore that it was involuntarily (Heyes, 2011; Moors & De Houwer, 2006; Proctor &
Vu, 2006). The finding that performance on a primary task is modulated by the observation
of actions that can either correspond or not correspond with the required response is now
widely known as automatic imitation (Heyes, 2011), and will be how the term is used in the
current paper.
As demonstrated in Figure 2, research on automatic imitation has grown
exponentially since the effect was first demonstrated in the early 2000s (Brass, Bekkering, et
al., 2001; Brass et al., 2000; Kilner et al., 2003; Stürmer et al., 2000). In contrast to motor
mimicry, which typically measures the frequency of imitation over an extended period of
time, automatic imitation measures reaction times (RTs), errors rates (ERs), or kinematics
(KMs) using stimulus-response compatibility paradigms (Heyes, 2011). The use of these
paradigms is rooted in a long tradition of psychological research on stimulus-response
compatibility (Simon, 1969; Stroop, 1935), and has been instrumental in understanding the
mechanisms that connect perception to action (Proctor & Vu, 2006). As part of this
tradition, the study of automatic imitation has provided crucial insights into the interplay
between perception and action in social settings (Heyes, 2011).
Although automatic imitation was first demonstrated in cognitive science, it is now
widely studied across a broad range of psychological disciplines. For example, it is thought
to rely on the same mechanisms as motor mimicry (Heyes, 2011). Moreover, these
mechanism are believed to facilitate social interaction (Stel, van Dijk, & van Baaren, 2016;
Wang & Hamilton, 2012) and to predict both empathy (Butler, Ward, & Ramsey, 2015;
Genschow et al., 2017) and autism spectrum disorder (Spengler, Bird, & Brass, 2010;
Williams, 2008; Williams, Whiten, Suddendorf, & Perrett, 2001). Furthermore, automatic
imitation is widely considered an index of mirror neuron activation (Bien, Roebroeck,
Goebel, & Sack, 2009; Catmur, Walsh, & Heyes, 2009; R. Cook, Bird, Catmur, Press, & Heyes,
2014; Heyes, 2010), serves as a model for social interaction in artificial intelligence
(Chaminade & Cheng, 2009), and has greatly advanced our understanding of the
mechanisms behind imitation in adults (Heyes, 2011), children (Heyes, 2016; Ray & Heyes,
2011), and non-human species (Heyes, 2012).
Following fifteen years of research, it is now well-established that automatic
imitation – the finding that movement execution is facilitated by compatible and impeded
by incompatible observed movements – exists (Heyes, 2011). However, important
theoretical questions remain. Some questions relate to the mechanisms underlying
automatic imitation: What processes contribute to automatic imitation? Is automatic
imitation flexible? When during action preparation does automatic imitation operate? Is
automatic imitation really automatic? Other questions relate to the factors that modulate
automatic imitation: What is the role of action goals in automatic imitation? Is automatic
imitation sensitive to the similarity between actor and imitator? Is automatic imitation
related to social abilities? Each of these questions is essential to understand the connection
between perception and action in social interaction. As such, answering them will not only
have important implications for the domain of automatic imitation but also for the broader
literature on how we represent others’ actions.

Although several narrative reviews on automatic imitation already exist (e.g.,
Chaminade & Cheng, 2009; Gowen, Stanley, & Miall, 2008; Heyes, 2011; Press, 2011),
inconsistent evidence has made it difficult to provide conclusive answers to the
aforementioned questions. To address this issue, the current study, we will use a meta-
analytic approach. A meta-analysis is well-suited to evaluate inconsistent evidence because
it provides a quantitative estimate of the available evidence based on the combined data of
many independent studies (Hedges & Vevea, 1998; Lipsey & Wilson, 2001). Moreover, a
meta-analysis allows us to identify important methodological variables that should be taken
into account in research on automatic imitation to ensure that the effects are unbiased and
reliable. In the remainder of our introduction, we will first describe the paradigms and
theories of automatic imitation and will then provide a brief review of the different
questions that are addressed in the meta-analysis.
--- Insert Figure 1 and 2 about here ---
Paradigms of Automatic Imitation
Automatic imitation in the current meta-analysis will be operationalized as “a type of
stimulus-response compatibility effect in which the topographical features of task-irrelevant
action stimuli facilitate similar, and interfere with dissimilar, responses” (Heyes, 2011, p.
463). There are three paradigms that fit this description: a forced choice RT paradigm, a
simple response RT paradigm, and a simple response KM paradigm. All three paradigms
investigate the interference of an irrelevant observed movement on a relevant executed
movement. In the forced choice RT paradigm, participants are required to select a response
on the basis of a symbolic cue while a hand on the screen makes an irrelevant compatible or
incompatible movement (Brass et al., 2000; Stürmer et al., 2000). In the study by Brass et al.
(2000) described earlier, a compatible trial is when the stimulus hand lifts the same finger as
the finger that has to be lifted by the participant (e.g., see index finger, move index finger)
and an incompatible trial is when the stimulus hand lifts a different finger as the finger that
has to be lifted by the participant (e.g., see index finger, move middle finger). Automatic
imitation in this paradigm is the finding that RTs are slower and ERs higher on incompatible
trials than on compatible trials (e.g., Bertenthal, Longo, & Kosobud, 2006; Brass et al., 2000;
Catmur & Heyes, 2011; Gillmeister, Catmur, Liepelt, Brass, & Heyes, 2008; Otte, Habel,
Schulte-Ruther, Konrad, & Koch, 2011; Stürmer et al., 2000).
In the simple response RT paradigms, participants are no longer required to choose a
response on the basis of a cue but can instead prepare the relevant response before the
onset of the trial. For example, Brass et al. (2001) instructed participants to either lift or tap
their index finger as soon as the stimulus hand lifted or tapped its index finger. In line with
the forced choice RT paradigm, it was found that responses on compatible trials were faster
than responses on incompatible trials (e.g., Bird, Leighton, Press, & Heyes, 2007; Brass,
Bekkering, et al., 2001; Heyes, Bird, Johnson, & Haggard, 2005; Otte, Habel, et al., 2011;
Press, Bird, Walsh, & Heyes, 2008). Finally, the simple response KM paradigm follows a
procedure that is similar to the simple response RT paradigm but focuses on KMs rather
than RTs. In a prominent study, Kilner et al. (2003) asked participants to move their arm
back and forth in the horizontal or vertical plane while facing an experimenter who moved
his arm in the compatible (i.e., parallel) or incompatible (i.e., orthogonal) plane. In line with
the RT paradigms, participants’ movement trajectory contained more variability when the
experimenter performed an incompatible movement than when he performed a compatible

movement (e.g., Bouquet, Gaurier, Shipley, Toussaint, & Blandin, 2007; Kilner, Hamilton, &
Blakemore, 2007; Roberts, Bennett, & Hayes, 2015; Stanley, Gowen, & Miall, 2007).
Theories of Automatic Imitation
In contrast to early views assuming an innate imitation mechanism (Meltzoff &
Moore, 1983, 1989), the current dominant view is that automatic imitation develops as a
consequence of domain-general learning mechanisms (Brass & Heyes, 2005). In particular,
this view is substantiated by two distinct yet related theories. A first theory is the associative
sequence learning (ASL) theory (Brass & Heyes, 2005; Cook et al., 2014; Heyes, 2010; Ray &
Heyes, 2011). This theory argues that the visual representation of an action is initially
unconnected to its motor representation. Instead, these connections develop gradually
because action execution produces perceivable sensory consequences that over time
become associated with the motor command that produces them. For example, when we
grasp an object, we typically see how our hand grasps the object. Likewise, when we express
an emotion, we tend to see the same emotion expressed on the face of others. These
experiences lead to bidirectional connections between perceiving and executing an action,
and this is thought to cause automatic imitation.
A second theory, named ideomotor (IM) theory, extends ASL by assuming an
additional mechanism (Brass & Heyes, 2005; Greenwald, 1970; Hommel, Müsseler,
Aschersleben, & Prinz, 2001; Prinz, 1997; Shin, Proctor, & Capaldi, 2010). Like ASL, IM posits
that associative learning leads to connections between visual and motor representations.
However, IM further predicts that this learning process culminates in the development of
ideomotor representations that represent actions in terms of their anticipated sensory

consequences (Greenwald, 1970). Given that the visual image of an action is part of its
sensory consequences, seeing an action primes the ideomotor representation controlling
that action (Brass & Muhle-Karbe, 2014), and this leads to automatic imitation. In other
words, IM and ASL differ in whether or not they assume ideomotor representations, but
agree in their emphasis on the role of motor learning (ML) in automatic imitation (Brass &
Heyes, 2005). Therefore we will refer to these theories as ML theories of automatic
imitation.
At the neurophysiological level, automatic imitation has mostly been explained in
relation to the mirror neuron system (Bien et al., 2009; Brass & Heyes, 2005; Catmur et al.,
2009; R. Cook et al., 2014; Heyes, 2011; Simpson, Murray, Paukner, & Ferrari, 2014). The
mirror neuron system refers to a network of motor areas in the frontal and parietal cortex
that do not only respond to action execution but also to action observation (Rizzolatti &
Craighero, 2004; Rizzolatti & Sinigaglia, 2010). Although this system was initially discovered
in the monkey brain (Gallese, Fadiga, Fogassi, & Rizzolatti, 1996), there is now converging
evidence that a similar system exists in humans as well (Molenberghs, Cunnington, &
Mattingley, 2012). Importantly, the presence of a mirror neuron system is consistent with
both ML theories of automatic imitation (Brass & Heyes, 2005). More specifically, ASL
interprets mirror neurons in the context of associative networks (Cook et al., 2014; Heyes,
2010) and IM sees mirror neurons as the neurophysiological manifestation of ideomotor
representations (Brass & Muhle-Karbe, 2014).

The Present Meta-Analysis
The current meta-analysis is structured around seven key questions in the literature.
As outlined above, these questions concern the mechanisms underlying automatic imitation
and the conditions under which automatic imitation occurs. In the remainder of the
introduction, we will briefly review extant research on these questions and explain how they
will be addressed in the meta-analysis that follows. Our meta-analysis will focus primarily on
RT data, but will also report ERs as a secondary outcome measure. In general, we expect
that RTs will be more sensitive to modulatory influences than ERs because the number of
errors is typically low (i.e. < 5%) in automatic imitation studies (e.g., Brass et al., 2000;
Stürmer et al., 2000). For KM tasks of automatic imitation, we will restrict our analysis to
questions that have been considered relevant in these tasks because the number of KM
studies (n = 23) is insufficient to explore all moderators.
What Processes Contribute to Automatic Imitation?
Imitative Versus Spatial Compatibility. To understand automatic imitation, it is
important to delimit the processes that do and the processes that do not contribute. For
example, a longstanding discussion concerns the role of spatial processes in automatic
imitation (Aicken, Wilson, Williams, & Mon-Williams, 2007; Bertenthal et al., 2006; Boyer,
Longo, & Bertenthal, 2012; Catmur & Heyes, 2011; Heyes, 2011; Jansson, Wilson, Williams,
& Mon-Williams, 2007; Jimenez et al., 2012; Press et al., 2008). For example, in the study of
Brass et al. (2000), participants observed a left stimulus hand mirroring their right response
hand (Figure 1a). As a result, index finger stimulus movements were not only compatible
with index finger responses in terms of imitation (i.e., see index finger, move index finger)
but also in terms of location (i.e., see left finger, move left finger). A fundamental question is
therefore whether automatic imitation (i.e., imitative compatibility) can be explained as a
mere byproduct of spatial processing (i.e., spatial compatibility).
Three methods have been developed to investigate the extent to which automatic
imitation depends on spatial compatibility. The first method is to position the stimulus hand
orthogonal to the response hand (e.g., J. L. Cook & Bird, 2011, 2012; Heyes et al., 2005;
Press, Bird, Flach, & Heyes, 2005). For example, in the Brass et al. (2000) study, spatial
processes can be neutralized by rotating the stimulus hand 90° counterclockwise so that its
fingers point to the right instead of downwards (Figure 1b). In this case, an observed index
finger movement is still compatible with an index finger response in terms of imitation (i.e.,
see index finger, move index finger), but no longer in terms of location (i.e., see lower
finger, move left finger). However, a potential issue with this method is that there is a
documented tendency to associate “down” with “left” and “up” with “right” (Weeks &
Proctor, 1990). As a result, this method confounds automatic imitation with orthogonal
spatial compatibility (Heyes, 2011).
The second method offers a solution to orthogonal spatial confounds by separately
manipulating imitative and spatial compatibility (e.g., Bertenthal et al., 2006; Boyer et al.,
2012; Catmur & Heyes, 2011). For instance, using the stimuli of Brass et al. (2000), this can
be achieved by presenting a left stimulus hand in one half of the trials and a right stimulus
hand in the other half of the trials. This setup results in a positive relation between imitative
and spatial compatibility in left hand trials and a negative relation in right hand trials, which
makes it possible to calculate a main effect of imitative compatibility that is independent of
spatial compatibility (Figure 1c). Finally, the third method addresses the spatial compatibility
confound by using more complex stimuli such as symbolic gestures that cannot easily be
categorized on a simple spatial dimension (Bortoletto, Mattingley, & Cunnington, 2013;
Cracco, Genschow, Radkova, & Brass, 2018; Liepelt, Prinz, & Brass, 2010). Since all three
methods have been proven to elicit automatic imitation, it is now widely accepted that
imitative compatibility can be influenced but not be reduced to spatial compatibility (Heyes,
2011).
However, it is still an open question how much of automatic imitation can be
explained by spatial processes. To address this question, we will compare RT tasks that
confound imitative and spatial compatibility with RT tasks that control for simple (e.g., left-
right) spatial compatibility by using one of the three methods discussed above. In addition,
we will explore whether orthogonal spatial compatibility has an influence on automatic
imitation when orthogonal stimulus-response compositions are used. While preliminary
research suggests that orthogonal spatial compatibility poses no threat to automatic
imitation (Jimenez et al., 2012; Press et al., 2008), the small sample size of these studies (N =
17 and 18) has made it difficult to draw strong conclusions. In the current meta-analysis, we
will perform a stringent test of the orthogonal spatial compatibility hypothesis by comparing
studies that controlled for simple but not orthogonal spatial compatibility with studies that
controlled for both types of spatial compatibility.
Effector Versus Movement Compatibility. If automatic imitation is not a spatial
effect, then what is it? According to Heyes (2011), it is a combination of effector and
movement compatibility. The term effector in this context is used as a synonym of body
part. In other words, effector compatibility refers to the overlap between the body part
moved by the model and the body part that has to be moved by the participant (e.g., index
or middle finger). In contrast, movement compatibility refers to the overlap between the
type of movement made by the model and the type of movement that has to be made by
the participant (e.g., lifting or tapping). Studies on effector compatibility manipulate effector
overlap while keeping the movement constant (e.g., lift index finger or middle finger),
whereas studies on movement compatibility manipulate movement overlap while keeping
the effector constant (e.g., lift or tap index finger). Research has found reliable imitation in
both effector (e.g., Biermann-Ruben et al., 2008; Brass et al., 2000; Catmur & Heyes, 2011;
Obhi, Hogeveen, Giacomin, & Jordan, 2014) and movement compatibility tasks (e.g., Brass,
Bekkering, et al., 2001; Heyes et al., 2005; Obhi & Hogeveen, 2010; Stürmer et al., 2000).
Moreover, research has demonstrated that effector compatibility can be found in the
absence of a stimulus movement by highlighting the relevant effector (J. L. Cook & Bird,
2011, 2012), and two studies have now shown that movement and effector compatibility
are at least partially independent (Leighton & Heyes, 2010; Press, Gherri, Heyes, & Eimer,
2010).
However, it is currently unknown which type of compatibility contributes most to
automatic imitation. An important distinction between effector and movement
compatibility is that effector incompatible trials activate two movements that can be
executed at the same time (e.g., lift index and middle finger), whereas movement
incompatible trials activate two movements that cannot be executed at the same time (e.g.,
lift and tap index finger). If the mutually exclusive nature of the movements activated on
movement incompatible trials causes more response conflict, then it can be predicted that
movement compatibility should be stronger than effector compatibility. Nevertheless, the
opposite can also be predicted if the two movements activated on effector incompatible
trials are combined into a composite movement (e.g., lift both index and middle finger). In
this case, effector incompatible trials would require participants to inhibit not only the
stimulus movement (e.g., lift middle finger) but also the composite movement (e.g., lift both
index and middle finger) in order to select the correct response (e.g., lift index finger). To
distinguish between these two hypotheses, we will compare RT automatic imitation in
effector compatibility tasks with RT automatic imitation in movement compatibility tasks.
Is Automatic Imitation Flexible?
In addition to the processes that make up automatic imitation, research has also
started to investigate the extent to which these processes are susceptible to additional
learning (R. Cook et al., 2014; Heyes, 2010). That is, both ML theories assume that automatic
imitation develops through associative learning (Brass & Heyes, 2005). As a result, these
theories predict that the acquisition of incompatible sensorimotor associations can overrule
preexisting compatible associations (R. Cook et al., 2014; Heyes, 2010). In support, research
has shown that automatic imitation was reduced after short periods of sensorimotor
training where participants had to counterimitate the actions they observed (e.g., R. Cook,
Press, Dickinson, & Heyes, 2010; Gillmeister et al., 2008; Heyes et al., 2005; Wiggett,
Hudson, Tipper, & Downing, 2011). Furthermore, automatic imitation has been found to
reverse when it was embedded in an action context that required participants to
counterimitate the stimulus movements (e.g., Ocampo, Kritikos, & Cunnington, 2011; van
Schie, van Waterschoot, & Bekkering, 2008). However, not all studies have supported the
flexible nature of automatic imitation (Poljac, van Schie, & Bekkering, 2009). Therefore, we
will investigate whether automatic imitation in RT tasks is sensitive to (in)compatible
training and (in)compatible action contexts. In particular, we expect a reduction of the
imitation effect after incompatible associations are acquired and either an enhancement or
no changes after compatible associations are acquired.

When During Action Preparation Does Automatic Imitation Operate?
A third question that will be addressed is the locus of automatic imitation. Action
preparation is a hierarchical process that entails two consecutive stages (Wong, Haith, &
Krakauer, 2015). The first stage consists of determining a motor goal (i.e., “what”) and the
second stage of developing a motor plan that achieves the established goal (i.e., “how”).
From this perspective, action observation can interfere either with the motor goal (e.g., “lift
the right index finger…”), with the motor plan (e.g., “… by activating the relevant muscles”),
or with both. To distinguish between these two possibilities, we will compare automatic
imitation in forced choice and simple response RT tasks. While observed movements in the
former task can interfere both with the motor goal and with the motor plan, observed
movements in the latter task can only interfere with the motor plan because the motor goal
is known before the stimulus movement is observed. Therefore, if automatic imitation is
restricted to forced choice RT tasks, then it influences the motor goal but not the motor
plan. In contrast, if automatic imitation is equally strong in both tasks, then it influences the
motor plan but not the motor goal. Finally, if automatic imitation occurs in both tasks but
more strongly in forced choice RT tasks, then it influences the motor goal as well as the
motor plan.
Is Automatic Imitation Really Automatic?
A fourth important question concerns the degree to which automatic imitation is
automatic. In the current meta-analysis, we will adopt a componential view on automaticity.
This is consistent with recent insights that automaticity is not a binary concept but rather a
multifaceted concept that encompasses a range of related features such as intentionality,
consciousness, speed, and attention (Moors & De Houwer, 2006). The word automatic in
automatic imitation therefore implies that it is a fast process that requires no intention, no
awareness, and no attention (Moors & De Houwer, 2006). Although it is well-established
that automatic imitation is unintentional (Heyes, 2011), less is known about its speed
(Catmur & Heyes, 2011; Cracco et al., 2018; Wiggett, Downing, & Tipper, 2013) and the role
of awareness (Mele, Mattiassi, & Urgesi, 2014) or attention (Bach, Peatfield, & Tipper, 2007;
Catmur, 2016; Chong, Cunnington, Williams, & Mattingley, 2009; Cracco & Brass, 2017). To
obtain a more informed view on the degree to which automatic imitation can be considered
automatic, we will focus on the features speed and attention.
Speed. A process can be considered automatic if it is a fast process (Moors & De
Houwer, 2006). A useful method to measure the speed of a process is to look at the
influence of stimulus-onset asynchronies (SOAs). Applied to automatic imitation, this can be
defined as the delay between the presentation of the stimulus movement and the
presentation of the imperative cue. Following this approach, research has discovered that
automatic imitation is a transient process that first increases with SOA but then decreases
again when the delay exceeds 80 to 150 ms (Catmur & Heyes, 2011; Cracco et al., 2018;
Wiggett et al., 2013). However, it is currently unknown how long after action observation
that automatic imitation can influence responses. Therefore, to further evaluate the extent
to which automatic imitation is a fast process, we will explore in detail how automatic
imitation develops as SOA increases.
Attention. A process is automatic in terms of attention if it requires little or no
attentional processes (Moors & De Houwer, 2006). Although initial research argued that
attention to the relevant features (Chong et al., 2009) of the relevant effector (Bach et al.,
2007) is a necessary condition of automatic imitation, a more recent study found that
directing attention away from the stimulus movements decreased but did not eliminate the
imitation effect (Cracco & Brass, 2017). Similarly, using a different technique, another study
showed that increasing the perceptual load did not make automatic imitation disappear but
instead made it reverse (Catmur, 2016). Since response speed was also slower at high
perceptual load, this reversal was interpreted as evidence that the processing of task-
irrelevant stimuli was inhibited after initially being facilitated (Eimer & Schlaghecken, 2003;
Jonas et al., 2007). In other words, it was argued that stimulus movements were still
processed when attentional load was high but that this led to a negative imitation effect
attributabel to inhibitory processes (Catmur, 2016). To further examine the role of attention
in automatic imitation, we will investigate whether the imitation effect in RT tasks depends
on the position of the imperative cue relative to the stimulus movement. That is, we expect
that automatic imitation will be stronger when the cue is positioned close to the stimulus
movement. In addition, we will also examine whether automatic imitation is stronger when
the cue is visual compared with auditory. This is because auditory cues do not force
participants to focus their attention on the screen while visual cues do.
What Is the Role of Action Goals in Automatic Imitation?
Another debate is whether automatic imitation is driven by action goals or by
movement parameters (Bouquet, Shipley, Capa, & Marshall, 2011; Chiavarino, Bugiani,
Grandi, & Colle, 2013; Liepelt et al., 2010; Liepelt, Von Cramon, & Brass, 2008; Press et al.,
2008). According to an influential view, imitation is a goal-directed process that involves
decomposing observed actions into a hierarchy of features where goals are more important
than how those goals are achieved (Wohlschläger, Gattis, & Bekkering, 2003). In line with
this view, research has shown that children (Bekkering, Wohlschläger, & Gattis, 2000) as
well as adults (Avikainen, Wohlschlager, Liuhanen, Hanninen, & Hari, 2003) prioritize the
ends over the means when asked to imitate (see also Gattis, Bekkering, & Wohlschläger,
2002; Gleissner, Meltzoff, & Bekkering, 2000; Want & Gattis, 2005). Furthermore, the notion
of goal-directed imitation is consistent with the observation that mirror neurons
preferentially respond to actions that are directed towards an object (Caspers, Zilles, Laird,
& Eickhoff, 2010; Cattaneo, Caruana, Jezzini, & Rizzolatti, 2009; Rizzolatti & Sinigaglia, 2010).
Applied to automatic imitation, goal-directed theories predict that imitative responses
should be stronger for goal-directed actions than for goalless actions (Bouquet et al., 2011;
Chiavarino et al., 2013). However, evidence for this theory is currently mixed with one study
supporting this pattern (Bouquet et al., 2011) and two other studies finding either the
opposite pattern (Chiavarino et al., 2013) or no difference (Liepelt et al., 2010).
Interestingly, the latter two studies suggest that movement parameters rather than
action goals might be what drives imitation (Chiavarino et al., 2013; Genschow, Florack, &
Waenke, 2013). In line with this view, it has been shown that the reason why goals are
imitated over movements is that goals tend to be more salient (Bird, Brindley, Leighton, &
Heyes, 2007; Leighton, Bird, & Heyes, 2010). Furthermore, strong evidence that movements
can guide imitation comes from research on cross-contextual imitation, which has
demonstrated that watching someone lift a barbell can trigger an action with similar
movement parameters but a different action goal such as lifting a cup to drink (Genschow &
Florack, 2014; Genschow et al., 2013; Genschow & Schindler, 2016). This is further
supported by the fact that automatic imitation occurs even when the stimulus movements
have no clear goal (e.g., Heyes et al., 2005; Press et al., 2008; Stürmer et al., 2000).
In sum, research on automatic imitation as well as related research has supported
both the view that imitation is driven by goals and the view that it is driven by movements.
Therefore, to further investigate the hypothesis that automatic imitation is a goal-directed
process, we will compare RT studies that used goalless stimulus movements such as finger
lifting (e.g., Brass et al., 2000; Stürmer et al., 2000) with RT studies that used object-directed
stimulus movements such as grasping (Chong et al., 2009; van Schie et al., 2008) or
communicative stimulus movements such as symbolic gestures (Bortoletto et al., 2013;
Cracco et al., 2018; Liepelt et al., 2010) and facial expressions (Otte, Jost, Habel, & Koch,
2011; Press, Richardson, & Bird, 2010). If goals have a stronger influence than movements,
then automatic imitation should be weaker when the stimulus movement is goalless
compared with when it is objected-directed or communicative (Bouquet et al., 2011;
Chiavarino et al., 2013). However, given that action goals are often more salient than
movement parameters (Bird, Brindley, et al., 2007; Leighton, Bird, & Heyes, 2010), an
alternative hypothesis is also that goal-related actions may drive away attention from the
observed movement towards the action goal. In this case, automatic imitation should be
weaker for goal-directed actions, and this should be especially true for object-directed
actions where attention can be drawn towards the object (Chiavarino et al., 2013).
Is Automatic Imitation Sensitive to the Similarity Between Actor and Imitator?
Apart from research on the mechanisms and conditions of automatic imitation, there
is also an extensive literature on whether automatic imitation is sensitive to the similarity
between the actor and the imitator (Chaminade & Cheng, 2009; Gowen & Poliakoff, 2012;
Press, 2011). In particular, this research builds on the assumption that automatic imitation is
a social process that should be moderated by the degree to which the imitator resembles
the actor in terms of perceived or actual appearance. At a theoretical level, both ML
theories consider similarity to be an important modulator of imitative behavior. For
example, according to ASL, imitative tendencies are learned responses that develop as a
result of self-observation and interaction with other, often similar (Efferson, Lalive, & Fehr,
2008), individuals (Brass & Heyes, 2005; Cook et al., 2014; Heyes, 2010; Ray & Heyes, 2011).
Although perception-action links are flexible in the sense that they allow generalization to
comparable stimuli, the strength of the imitative response is thought to depend on the
number of features that the observed movement shares with the stored action
representation (Shanks & Darby, 1998). Similarly, IM postulates that stimuli differ in terms
of ideomotor compatibility – the extent to which they resemble the sensory outcomes of an
action (Brass, Bekkering, et al., 2001; Greenwald, 1970; Shin et al., 2010). As a result, IM
assumes that stimuli with higher ideomotor compatibility can access the corresponding
ideomotor representation more directly and therefore that automatic imitation is stronger
when the model is similar to the participant.
Bottom-Up Animacy. If similarity modulates automatic imitation, then human agents
should elicit a stronger imitative response than nonhuman agents. In support of this
hypothesis, research has found stronger automatic imitation for human agents compared
with nonhuman agents in both RT (e.g., Bird et al., 2007; Brass, Bekkering, et al., 2001;
Gowen, Bolton, & Poliakoff, 2016; Press et al., 2005; Press, Gillmeister, & Heyes, 2006, 2007)
and KM (e.g., Chaminade, Franklin, Oztop, & Cheng, 2005; J. L. Cook, Swapp, Pan, Bianchi-
Berthouze, & Blakemore, 2014; Gowen et al., 2008; Kilner et al., 2003) tasks. However, not
all studies have supported a clear-cut human bias in automatic imitation (Cracco, De Coster,
Andres, & Brass, 2015; Jansson et al., 2007; Kilner et al., 2007; Klapper, Ramsey, Wigboldus,
& Cross, 2014; Kupferberg et al., 2012; Liepelt et al., 2010; Oztop, Franklin, & Chaminade,
2005; Press et al., 2006). A closer look at these studies shows that they mostly used
nonhuman agents with high (e.g., a wooden hand) as opposed to low (e.g., a moving dot)
human resemblance. To investigate whether automatic imitation is reduced for nonhuman
agents, we will compare RT and KM automatic imitation of human models with RT and KM
automatic imitation of nonhuman models such as robots (e.g., Press et al., 2006) and
geometrical models such as moving dots (e.g., Gowen et al., 2016). If automatic imitation is
sensitive to similarity, then imitative responses should become stronger as the stimuli
become more humanlike.
Top-Down Animacy. In addition to bottom-up animacy, beliefs about animacy may
also influence automatic imitation. For example, Liepelt and Brass (2010b) measured
automatic imitation of a gloved hand after showing participants that the hand was human
or wooden. The results revealed a stronger imitation effect in the human condition,
indicating that beliefs about animacy can influence automatic imitation. The influence of
animacy beliefs on automatic imitation has since been supported in some (Gowen et al.,
2016; Klapper et al., 2014; Longo & Bertenthal, 2009; Stanley et al., 2007) but not all studies
(Press et al., 2006). Therefore, we will investigate whether RT automatic imitation is weaker
for human and ambiguous agents when participants are told that the agent is nonhuman
and whether it is stronger for nonhuman and ambiguous agents when participants are told
that the agent is human.
Motion Profile. Apart from model characteristics, the degree to which the stimulus
movement follows a human motion profile might also play a role. For example, in a study on
KM automatic imitation, Kilner et al. (2007) found that participants did not imitate a human
model when its movements followed a non-biological rather than a biological motion
profile. Although some studies have obtained similar results (Chaminade et al., 2005;
Stanley, Gowen, & Miall, 2007), other studies found no difference between biological and
non-biological motion profiles (Bouquet et al., 2007; J. L. Cook et al., 2014; Gowen et al.,
2008). Therefore, we will compare KM automatic imitation of stimulus movements that
followed a biological motion profile with KM automatic imitation of stimulus movements
that did not follow a biological motion profile.
Although movement characteristics have often been studied in KM studies, this
factor has remained relatively unexplored in RT studies. To address this issue, we will look at
the presentation mode of the stimulus movements. In particular, we will compare stimulus
movements presented as short videos (e.g., Brass et al., 2000; Gowen et al., 2016; Heyes et
al., 2005) with stimulus movements presented as a two-frame sequence in which a picture
of the stimulus in neutral position is replaced with a picture of the stimulus in final position
(e.g., Catmur & Heyes, 2011; Cracco et al., 2015; Press et al., 2007) and stimulus movements
presented as a single picture of the stimulus in final position (e.g., Chong et al., 2009; Liepelt
et al., 2010; Otte, Jost, et al., 2011). Given that videos provide a more realistic
representation of human motion than a sequence of two pictures or a single picture, it can
be predicted that RT automatic imitation will be stronger when videos are used instead of
pictures.
Perspective. A more subtle aspect of similarity is the perspective from which the
stimulus movement is seen. That is, at least for non-facial actions, ML theories postulate
that self-observation is the main mechanism behind the formation of perception-action links
(Brass & Heyes, 2005). As a result, it can be predicted from these theories that automatic
imitation should be stronger when a movement is seen from a first-person perspective than
when it is seen from a third-person perspective. In line with this idea, research has found
stronger automatic imitation for actions presented in a first-person perspective than for
actions presented in a third-person perspective (Bortoletto et al., 2013; Genschow et al.,
2013; Vogt, Taylor, & Hopkins, 2003). However, some studies have also suggested that the
presence of a first-person bias might depend on contextual variables such as “cognitive
style” (Bortoletto et al., 2013). To evaluate the role of visual perspective, we will examine
whether automatic imitation in RT tasks is stronger when stimulus movements are
presented in a first-person perspective than when they are presented in a third-person
perspective.
Gender Overlap. Finally, we will look at two similarity features that have not yet
been explored in the literature. The first feature is gender overlap. Given that two persons
tend to be more similar when they are of the same gender, it can be predicted that
automatic imitation should be stronger when there is a match between the gender of the
model and the gender of the participant. In support of this idea, a recent study reported
stronger imitative responses for female than for male participants when a female stimulus
hand was used (Butler et al., 2015). However, this study did not manipulate the gender of
the stimulus hand. As a result, it is unclear whether this effect was a gender effect or a
gender overlap effect. To address this question, we will investigate whether automatic
imitation in RT tasks is stronger when the gender of the model corresponds to the gender of
the majority of the sample.
Response Overlap. The second feature of similarity that has remained unexplored is
response overlap, which refers to how similar the stimulus movement is to the response
movement. For example, studies with finger lifting/tapping stimuli have not always used
finger lifting/tapping responses (e.g., Brass, Bekkering, et al., 2001; Brass et al., 2000;
Liepelt, Stenzel, & Lappe, 2012), but have sometimes also used key release/press responses
(e.g., Butler et al., 2015; Longo & Bertenthal, 2009; Longo, Kosobud, & Bertenthal, 2008).
Although a key press looks similar to a finger tap, these two actions feel different, sound
different, and have a different outcome. Since ML theories assume that perception-action
associations are not only constructed from the visual but also from the sensory and auditory
outcomes of an action (Hommel et al., 2001; Shin et al., 2010), it can be predicted, to put it
in the words of IM, that a key press is represented with a different ideomotor
representation than a finger tap. To test this hypothesis, we will investigate whether
automatic imitation of finger lifting/tapping stimuli is stronger when the response is also
finger lifting/tapping than when the response is releasing/pressing a key.
Is Automatic Imitation Related to Social Abilities?
The finding that automatic imitation is sensitive to the similarity between the actor
and imitator has been interpreted as evidence that imitation is an inherently social process
(Wang & Hamilton, 2012). In extension, a prominent view is that automatic imitation
(Gallese, 2007; Wang & Hamilton, 2012) or how it is controlled (Brass, Ruby, & Spengler,
2009; Sowden & Shah, 2014) can provide insight into people’s social abilities. For example, it
has been proposed that automatic imitation might be related to social functioning because
it relies on embodied simulation, which is assumed to be at the basis of important social
skills such as mentalizing and empathy (Gallese, 2007). In the case of empathy, for example,
it is argued that simulation is necessary to feel the emotional state of someone in distress
(Carr, Iacoboni, Dubeau, Mazziotta, & Lenzi, 2003). In contrast, another view argues that it is
not self-other overlap but self-other control that is important for social cognition (Brass et
al., 2009; Sowden & Shah, 2014). For instance, empathy does not only require individuals to
share others’ emotional state but also to control these emotions in order to provide a
complementary empathic response (Bird & Viding, 2014; Decety & Lamm, 2006). From this
perspective, automatic imitation is an indicator of social abilities because participants have
to distinguish between self and other representations on incongruent trials to overcome the
urge to imitate (Brass et al., 2009; Sowden & Shah, 2014). Although these two theories
differ in how automatic imitation is related to social cognition, they agree that impairments
in automatic imitation should be accompanied by social difficulties, which should be visible
as impaired empathic abilities (Carr et al., 2003; Gallese, 2007; Sowden & Shah, 2014) and
might lead to autism spectrum disorder (ASD) (Spengler, Bird, et al., 2010; Williams, 2008;
Williams et al., 2001).
Empathy. The construct of empathy can be divided into cognitive and emotional
empathy, with the former referring to perspective taking and the latter to emotional
concern (Blair, 2005; Davis, 1983). However, in contrast to the two theories outlined earlier,
research on the relation between automatic imitation and empathy has yielded mixed
results. That is, one study found a relation with empathy only when the model was
attractive (Müller, Van Leeuwen, Van Baaren, Bekkering, & Dijksterhuis, 2013), a second
study only with the social modulation of automatic imitation (Haffey, Press, O’Connell, &
Chakrabarti, 2013), and two final studies found no relation (Butler et al., 2015; Genschow et
al., 2017). In the current meta-analysis, we will investigate the relation between automatic
imitation and both cognitive and emotional empathy across studies. Since multiple
instruments have been used in the literature, we will combine data from the Interpersonal
Reactivity Index (IRI; Davis, 1983; De Corte et al., 2007), the Empathy Assessment Index (EAI;
Gerdes, Lietz, & Segal, 2011; Inzunza, 2014), and the Empathy Quotient (EQ; Baron-cohen &
Wheelwright, 2004), for which validation studies have found high inter-correlations (De
Corte et al., 2007; Gerdes et al., 2011; Inzunza, 2014; Lawrence, Shaw, Baker, Baron-Cohen,
& David, 2004).
Autism Spectrum Disorder. If automatic imitation is a mirror into the social mind,
then it should be impaired in social disorders such as ASD (Spengler, Bird, et al., 2010;
Williams, 2008; Williams et al., 2001). This view is instantiated by two important theories on
ASD, each corresponding to one of the two social cognition accounts outlined earlier.
According to one theory, ASD is the result of a ‘broken mirror neuron system’ and this is
thought to cause hypo-imitation (Williams, 2008; Williams et al., 2001). In particular, it is
assumed that impairments in the perception-action system lead to difficulties in simulating
the mental states of others, which in turn is at the root of social problems in ASD. However,
it is important to note that hypo-imitation can also point to a different mechanism, namely
that participants with ASD do not attend to social stimuli or are less motivated to process
them (Blake, Turner, Smoski, Pozdol, & Stone, 2003; Chevallier, Kohls, Troiani, Brodkin, &
Schultz, 2012; Klin, Lin, Gorrindo, Ramsay, & Jones, 2009). Moreover, there is currently only
one study showing reduced automatic imitation in ASD (J. L. Cook et al., 2014). Therefore, a
second theory has argued that it is not the mirror system but the control over this system
that is impaired in ASD (Spengler, Bird, et al., 2010). More precisely, this theory argues that
individuals with ASD have difficulties to distinguish self from other representations. As a
result, they are slower to inhibit automatic imitation when the observed movement does
not match the instructed response and for that reason are expected to display hyper- rather
than hypo-imitation (Spengler, Bird, et al., 2010).
In support of the self-other control account, two studies have now reported hyper-
imitation in ASD (Deschrijver, Wiersema, & Brass, 2017; Spengler, Bird, et al., 2010), two
additional studies reported context-bound hyper-imitation (Bird, Leighton, et al., 2007;

Schunke et al., 2015), and one final study reported a positive relation between autism
symptom severity and automatic imitation (Sowden, Koehne, Catmur, & Dziobek, 2015).
Nevertheless, a substantial research body also failed to find evidence for hyper-imitation in
both children (Grecucci et al., 2013) and adults (Forbes, Wang, & Hamilton, 2016; Gowen et
al., 2008; Press, Richardson, et al., 2010; Sowden et al., 2015) with ASD. Taken together,
studies on ASD have provided mixed results. An important reason is presumably that
research on this topic has mostly used small samples (for an exception: Sowden et al.,
2015). As a stringent test of the hypo- and hyperimitation hypotheses of ASD, we will
therefore compare RT and KM automatic imitation between ASD and neurotypical samples.
Secondary Moderators
In addition to the moderators discussed above, we will also consider the influence of
participant age, number of trials, and response effector as moderators of secondary
interest.
Age. With regard to participant age, we will explore whether the mean age of the
sample is related to the strength of automatic imitation in RT tasks.
Trials. To investigate whether automatic imitation in RT tasks is influenced by the
length of the experiment, we will look at the total number of trials in the blocks on which
the effect size was based.
Effector. The effect of response effector will be investigated to explore whether
automatic imitation differs for finger movements, hand movements, foot movements, and
facial movements. To this end, we will examine if the imitation effect in RT tasks depends
on the effector that is involved.

Method
Literature Search
The literature search was initiated in February 2015 and terminated in January 2016.
We searched for studies published between January 1 2000 and January 1 2016 based on
the publication date of two important papers (Brass et al., 2000; Stürmer et al., 2000). To
collect published studies, we entered the following search terms in Web of Science,
PubMed, and Google Scholar: “automatic imitation” OR “imitation inhibition” OR "imitative
compatibility" OR "imitative congruency" OR "movement compatibility" OR "movement
congruency" OR "action compatibility" OR "action congruency" OR "motor compatibility" OR
"motor congruency" OR "movement interference" OR "action interference" OR "motor
interference" OR "motor facilitation" OR "movement facilitation" OR "action facilitation" OR
"motor mimicry" OR "action mimicry" OR "movement mimicry" OR "motor priming" OR
“action priming" OR "movement priming" OR "visuomotor priming" OR "body part priming"
OR "effector priming". Most of these search terms were identified by Heyes (2011). The
Web of Science and PubMed search was performed on titles, abstracts, and keywords. A
similar strategy was initially used for Google Scholar, but was later restricted to a title search
to limit the number of initial results (i.e., 2870).
In addition to the Boolean search, we also looked for studies that cited at least one
out of four influential papers on automatic imitation (Brass, Bekkering, et al., 2001; Brass et
al., 2000; Heyes, 2011; Stürmer et al., 2000). Furthermore, to include doctoral dissertations
in the database we added AND “dissertation” to the Google Scholar title, abstract, and
keyword search. Finally, unpublished data was collected by sending requests to prominent
authors, the European Social and Affective Neuroscience mailing list, and the Belgian
Association for Psychological Science mailing list. The above search strategy resulted in 1374
unduplicated results consisting of 1326 published articles or books, 10 doctoral
dissertations, and 38 unpublished datasets.
Screening Process
The studies obtained from the literature search were divided among the different
authors and were evaluated on the basis of the criteria described below. At the end of this
process, the first author checked the screening of all authors for errors. We first screened
the abstracts to identify studies that were clearly not eligible. The remaining studies were
then evaluated in more detail, resulting in a database of 161 studies. Together, these
studies contained 251 experiments, of which 25 did not meet the criteria for inclusion.
Importantly, we only included experimental conditions that satisfied all inclusion criteria.
The criteria used to screen the 1374 search results will now be described in detail. The
number of excluded studies is denoted with n and the number of excluded experiments
within otherwise included studies is denoted with m (Figure 3).
1. The study should report original empirical data (n = 238).
2. The study should investigate RT, ER, or KM automatic imitation (n = 939, m = 19). A
task was considered to assess automatic imitation if it measured how task-related
motor responses were influenced by seeing compatible and incompatible
movements (e.g., Brass, Bekkering, et al., 2001; Brass et al., 2000; Kilner et al., 2003).
The dependent variable of these studies should be RTs, ERs, or KMs. However,
studies on KM automatic imitation were only considered if they measured

movement trajectories as in Kilner et al. (2003) to ensure that the effect sizes were
comparable (Lipsey & Wilson, 2001). This means that we did not include studies that
measured alternative KM properties such as force (e.g., Salama, Turner, & Edwards,
2011) or movement duration (e.g., Pfister, Dignath, Hommel, & Kunde, 2013).
Furthermore, studies that measured automatic imitation of tool use were considered
beyond the scope of the meta-analysis and were therefore not included (e.g.,
Massen, 2009). Finally, studies on social attention were not included even if they
reported automatic imitation of movements such as eye gaze or pointing because
imitation in these paradigms cannot be distinguished from attentional processes
(e.g., Crostella, Carducci, & Aglioti, 2009).
3. The compatibility between the observed movement and the instructed response
should be manipulated within-subjects (n = 2). This criterion was included to ensure
that the effect sizes could be compared (Dunlap, Cortina, Vaslow, & Burke, 1996;
Lakens, 2013).
4. Studies should report a measure of imitative compatibility that is independent of
other types of compatibility with the exception of spatial compatibility (n = 10, m =
1). We did not exclude studies in which imitative compatibility was confounded with
spatial compatibility so we could investigate the influence of spatial processes on
automatic imitation. However, we did exclude studies in which imitative
compatibility and simple spatial compatibility were always negatively correlated
because it is impossible to extract a clean imitation effect from such studies (e.g.,
Müller et al., 2013).
5. Each response should occur equally often in the compatible condition as in the
incompatible condition (n = 1, m = 1). Studies that did not meet this criterion were
excluded to ensure that automatic imitation was not confounded with response
execution.
6. Studies reporting a reversed imitation effect were not included when this could be
explained as a complementary response (n = 1). This resulted in the exclusion of one
unpublished study in which it was found that participants were faster to take a
dominant pose when seeing a submissive pose and vice versa (Müller, de Graag, &
van Baaren, 2016). Moreover, studies that predicted a reversed automatic imitation
effect due to inhibitory processes were excluded as well (n = 2). For example, one
study was excluded because it predicted reversed automatic imitation due to social
inhibition of return (Jonas et al., 2007).
7. Stimuli should be presented for more than 100 ms to ensure that they were
processed consciously (n = 1).
8. The dependent measure should be recorded reliably (n = 3). For example, one study
was excluded because RT was measured by counting the number of video frames
with 40 ms duration each between the presentation of the stimulus movement and
the movement onset (Wild, Erb, Eyb, Bartels, & Grodd, 2003).
9. The sample should consist of human participants (n = 8). Moreover, a study was only
considered when the sample consisted of participants that (a) had no actual or
virtual brain damage, (b) had not been diagnosed with a psycho- or
neuropathological disorder except for ASD, and (c) had not been administered a
psychopharmaceutic substance (n = 1). An exception was made for ASD to test the
hypo- and hyper-imitation hypotheses of ASD. If an experiment contained an
ineligible clinical group together with a control group, then only the control group
was included in the analysis. Sham brain stimulation or brain stimulation at control
locations was not considered virtual brain damage.
10. The same dataset should not be included more than once in the meta-analysis (n = 2,
m = 2). If multiple studies used the same dataset, then only the first study was
included.
11. Sufficient statistical information should be available to calculate the necessary effect
sizes (n = 5, m = 2). The required information should be either reported in the paper
or obtained from the authors.
--- Insert Figure 3 about here ---
Coding and Reliability
The eligible studies were coded by the first author according to a coding manual that
was approved by all authors. The coded variables are described in Appendix A. Afterwards,
we added one additional variable that indicated whether studies that controlled for simple
spatial compatibility also controlled for orthogonal spatial compatibility. With regard to
perspective, a stimulus movement was coded as third-person when it was biomechanically
impossible for the observer to position the relevant body part in the same way as the
observed body part. Furthermore, because studies rarely report their results separately for
male and female participants, participant gender was coded as ‘female’ when > 50% of the
participants in the sample were female (MRT = 69%, MER = 69%) and as ‘male’ when ≤ 50% of
the participants were female (MRT = 39%, MER = 42%). Finally, we used the variable response
device to determine response overlap. Regular meetings were organized among eight
authors to discuss problems that were encountered during coding. For example, one
notable problem was that model gender was difficult to code objectively. Therefore, this
variable was coded according to a consensus method in which each of the eight authors
coded this variable individually by indicating whether they considered the model to be male,
female, or ambiguous. If ≥ 75% of the coders agreed on the gender, it was coded as such.
Otherwise, the gender was coded as ambiguous.
To evaluate the reliability of the coding, the ninth co-author was trained to use the
coding manual and subsequently recoded 28 randomly selected studies containing 47
experiments in total. Afterwards, to further identify errors, the same co-author also checked
the coding of the first author. Disagreements and inconsistencies that arose during coding
were resolved through discussion in group. Coding reliability was assessed with intraclass
correlation coefficients (ICC) for continuous variables and with kappa coefficients for
categorical variables. The average reliability for continuous variables (ICC = .98) was high
and varied between .94 (Trials) and 1.00 (Age and SOA). Similarly, high reliability was
observed for categorical variables (κ = .91) and varied between .70 (Action Goals) and 1.00
(Flexibility, Paradigm Type, Cue Modality, Top-Down Animacy, ASD).
Meta-Analytic Procedures
All analyses were performed in R using the metafor (Viechtbauer, 2010) and
robumeta (Fisher & Tipton, 2015) packages.
Mixed Effects Model. The data was analyzed with a mixed effects model:
𝑦𝑖 = 𝛽0 + 𝛽1 𝑥𝑖1 + … + 𝛽𝑝 𝑥𝑖𝑝 + 𝑢𝑖 + 𝑒𝑖 (1)
where yi is the effect size of experiment i, β0 is the weighted average population effect, β1 …
βp are the regression coefficients for moderators xi1 … xip, ui ~ N(0, τ2) is the error between
experiments, and ei ~ N(0, vi) is the error within experiments. More precisely, mixed effects
models assume that variability among effect sizes can arise both within and between
experiments. First, variability within experiments vi is introduced through inaccurate
measurement of the underlying effect size. Second, variability between experiments or
heterogeneity τ2 is introduced through methodological or contextual differences between
experiments. Importantly, τ2 reflects the amount of heterogeneity that remains after
accounting for the moderator variables that are included in the model. Since mixed effects
models allow for heterogeneity among the effect sizes, these models can be used to make
inferences about the population rather than about the m experiments included in the meta-
analysis (Field & Gillett, 2010). However, a crucial assumption of mixed models is that the
effect sizes yi should be statistically independent (Lipsey & Wilson, 2001). This assumption is
violated when multiple effect sizes from the same sample are included and such a violation
is known to inflate the false-positive rate (Tanner-Smith, Tipton, & Polanin, 2016). Since
some of the moderators have mostly been studied in repeated measures designs, we
decided to control for statistical dependency by using robust variance estimation (RVE)
(Fisher & Tipton, 2015; Hedges, Tipton, & Johnson, 2010; Tanner-Smith & Tipton, 2014;
Tanner-Smith et al., 2016).
Robust Variance Estimation. RVE accounts for dependency in the data by calculating
an empirical estimate of the standard error that does not assume independent effect sizes
(Fisher & Tipton, 2015; Hedges et al., 2010; Tanner-Smith & Tipton, 2014; Tanner-Smith et
al., 2016). Simulation studies have shown that RVE provides unbiased estimates of the true
standard error, does not assume a specific distribution, and is robust to misspecification of
the weights (Hedges et al., 2010). However, even though statistical inferences are valid for
any set of weights, inverse variance weights are preferred in terms of statistical efficiency
(Hedges et al., 2010). In particular, two types of weights have been proposed: hierarchical
effects weights and correlated effects weights. Hierarchical effects weights are most
appropriate when multiple experiments are nested within a larger cluster (e.g., research
group), while correlated effects weights are most appropriate when multiple effect sizes
from the same sample are included. Although both types of dependency typically co-occur,
the choice of weights only affects statistical efficiency. It has therefore been recommended
to pick a weighting method based on the main source of dependency (Tanner-Smith &
Tipton, 2014).
To optimally account for the inclusion of multiple effect sizes from the same sample,
the current meta-analysis used correlated effects weights (Fisher & Tipton, 2015):
1
𝑤𝑖 = (2)
𝑘𝑖 (𝑣∙𝑖 + 𝜏 2 )
where v∙i is the average variability within experiment i, τ2 is an estimate of the variability
between experiments, and ki is the number of effect sizes in experiment i. Note that the
total variability in each experiment v∙i + τ2 is divided equally among the ki effect sizes of
experiment i so that each effect size has the same weight. This strategy ensures that the
total weight assigned to experiment i does not depend on the number of effect sizes
provided by this experiment. Moreover, the calculation of τ2 depends on a parameter ρ that
represents the correlation among the effect sizes and is assumed to be the same for all
experiments (Fisher & Tipton, 2015). Although the value of ρ has to be set manually, varying
this parameter typically has little influence on the results. In the current study, the default
value ρ = 0.8 of the robumeta package was used (Fisher & Tipton, 2015). In addition,
sensitivity analyses will be reported to investigate whether the results depend on the value
of ρ.
Although RVE provides an unbiased estimate of the standard error, it is known to
underestimate the true variance when the number of experiments is small. To address this
issue, small sample corrections have been proposed (Tipton, 2015; Tipton & Pustejovsky,
2015). As these corrections also control for unbalanced or skewed moderators, it was
recently recommended that these corrections should always be implemented (Fisher &
Tipton, 2015; Tanner-Smith et al., 2016). In keeping with this recommendation, small
sample adjustments were applied to all tests in the current meta-analysis. Importantly, p
values from t tests with small sample correction are valid when df ≥ 4 but suffer from
inflated Type I error rates when df < 4 (Tanner-Smith et al., 2016). Therefore, we will only
report analyses where all t test contrasts have df ≥ 4. In addition, for continuous
moderators, we will remove outlier values exceeding the weighted mean value by > 3 SD
because including these values reduces the dfs and therefore reduces statistical power
(Tanner-Smith et al., 2016).
Effect Size Computation. Cohen’s d was used as a measure of effect size. In
particular, dz for the difference between incongruent (IC) and congruent (C) trials was
calculated with the following formula:
𝑀𝐼𝐶 − 𝑀𝐶 𝑀𝐼𝐶−𝐶
𝑑𝑧 = = . (3)
√𝑣𝑎𝑟𝐶 + 𝑣𝑎𝑟𝐼𝐶 − 2 × 𝑐𝑜𝑣𝐶,𝐼𝐶 𝑆𝐷𝐼𝐶−𝐶
This effect size controls for inter-individual differences by including the correlation between
C and IC in the denominator (Lakens, 2013). Since the value of dz depends on the strength of
the correlation, it has been argued that dz does not provide a good estimate of the true
effect size (Dunlap et al., 1996). However, this argument rests on the assumption that
between-subject designs are the default design in psychology (Lakens, 2013). Given that
automatic imitation is almost always assessed in a repeated measures design, this argument
does not hold for the current meta-analysis. Indeed, the natural unit of analysis in such
studies is the compatibility effect (IC - C). As this unit is calculated at the intra-individual
level, it is reasonable to rely on effect sizes that control for inter-individual differences
(Lakens, 2013). When dz could not be calculated on the basis of the raw data, it was
calculated on the basis of the relevant t or F test:
𝑡 𝐹
𝑑𝑧 = or √ . (4)
√𝑛 𝑛
Finally, because dz provides a slightly biased estimate of the population effect size, it was
corrected using the “escalc” function of the metafor package (Viechtbauer, 2010) to obtain
the unbiased effect size gz:
𝑛−1
𝛤( 2 )
𝑔𝑧 = 𝑑𝑧 × (5)
√(𝑛 − 1) × 𝛤 (𝑛 − 2)
2 2
With Γ(x) representing the gamma function and with the following variance:
1 𝑔𝑧2
𝑉𝑎𝑟𝑔𝑧 = + . (6)
𝑛 2𝑛
Seventy-three studies did not contain sufficient statistical information to calculate an
effect size in at least one experiment. To maximize the number of effect sizes that could be
included, we therefore sent data requests to the corresponding author of these studies
between November and December 2015. A subsequent reminder was sent to non-
responding authors between January and February 2016. We received all requested
statistics for 46 studies and part of the requested statistics for 1 study. For 22 studies, the
data was no longer available or could not be retrieved in time. Finally, we received no
response to our data requests for 4 studies.
For the 27 studies without sufficient statistical information, we obtained an effect
size estimate using the following procedure. First, we looked for studies where we could
extract MIC-C, SDC, and SDIC from the text (n = 6, m = 13, k = 139). To calculate an effect size
for these studies, we computed the correlation between C and IC for all experiments in the
meta-analysis where this was possible and then imputed the relevant RT (r = .94, n = 61, m =
95), ER (r = .47, n = 23, m = 36), or KM (r = .81, n = 11, m = 12) correlation coefficient in the
dz formula (Higgins, Deeks, & Altman, 2011). Second, we looked for studies where MIC-C and
either SDIC-C or both SDC and SDIC could be extracted from the provided figures using a
computerized measurement tool (n = 8, m = 12, k = 51). When SDC and SDIC were extracted,
we imputed the mean correlation coefficient in the dz formula following the procedure
described above (Higgins et al., 2011). Third, we looked for studies where we could extract
MIC-C but not SDC or SDIC from the text or figures (n = 6, m = 10, k = 27). To get an effect size
for these studies, we calculated SDIC-C for all experiments in the meta-analysis where this
was possible and then imputed the relevant RT (SDIC-C = 30.83 ms, n = 97, m = 138) or ER
(SDIC-C = 4.27%, n = 25, m = 38) SD in the dz formula (Higgins et al., 2011). Finally, we were
able to calculate an effect size for an additional 2 studies (m = 2) and 1 experiment when we
did not code one of the moderators. However, despite these efforts, we could not calculate
an effect size for 5 studies (m = 5) and 2 experiments of studies that were otherwise
included.
Outliers. RT, ER, and KM extreme values were identified by looking for effect sizes
that exceeded the weighted mean effect size by more than 3 SDs. Note that a weighted
mean was used to ensure that the influence of each experiment did not depend on the
number of provided effect sizes. To calculate the weighted mean, a weight of 1 was divided
equally among the different effect sizes belonging to the same experiment. The above
procedure identified 1 RT outlier (gz = 3.99), 2 ER outliers (gz = 2.19, 2.37), and no KM
outliers. These outliers were subsequently replaced by the 3 SD deviation value for RTs (gz =
3.41) and ERs (gz = 2.03) respectively.
Moderator Correlations. A potential issue in meta-analyses is confounded
moderators (Field & Gillett, 2010; Lipsey & Wilson, 2001). To identify such confounds, we
assessed the weighted association between the different RT and KM moderators with r for
continuous-continuous and continuous-dichotomous pairs, with multiple R for continuous-
polytomous pairs, and with Cramér’s V for dichotomous-dichotomous, dichotomous-
polytomous, and polytomous-polytomous pairs (Table 1). For all three statistics, a value of 0
indicates the absence of a relation and a value of 1 indicates a perfect relation. In the
current meta-analysis, an association ≥ .40 was considered substantial. According to Cohen
(1988), a value of .40 can be considered a medium-to-large association. When a substantial
relation between two moderators was identified, we examined the moderator effect while
controlling for the other moderator to rule out that the presence or absence of a moderator
effect was the result of confounding.
Publication Bias. To test for potential publication bias, we will plot each
1
experiment’s mean effect size against √ with a funnel plot. A negative association
𝑁
between the sample size and the effect size across studies is indicative of publication bias
because studies with small samples need large effect sizes to yield a significant result. We
used N rather than SE as a measure of precision because the latter depends on gz and can
therefore result in spurious funnel plot asymmetry, especially when the average effect size
is large (Sterne, Egger, & Moher, 2011; Zwetsloot et al., 2017). Funnel plots allow us to test
for publication bias at the level of the main effect (i.e., the congruency effect).
However, publication bias can also occur at the moderator level. That is, if studies
that did not find a significant moderation effect were not published, then this could lead to
publication bias that cannot be detected with funnel plots. Therefore, to address this issue,
we will report p-curve analyses for three moderators that have mostly been manipulated
within experiment: Flexibility, Bottom-Up Animacy, and Top-Down Animacy. A p-curve
analysis tests if the distribution of significant p-values is consistent with what can be
expected when the effect is true (Simonsohn, Nelson, & Simmons, 2014; Simonsohn,
Simmons, & Nelson, 2015). That is, true effects have a right-skewed distribution in which p <
.025 is more likely than .025 < p < .050. In contrast, nonexistent effects have a uniform
distribution in which each p-value is equally likely. Finally, p-hacked effects have a left-
skewed distribution in which .025 < p < .050 is more likely than p < .025. A p-curve analysis
investigates the evidential value in a set of studies by testing whether the observed
distribution of p-values is significantly right-skewed. Because it only looks at significant p-
values, this method is robust to publication bias and p-hacking (Simonsohn et al., 2014,
2015). Importantly, simulation studies have shown that p-curve analyses have a controlled
5% false positive rate even when the number of included p-values is low, and that their
statistical power is always higher than that of the individual studies included in the analysis
(Simonsohn et al., 2014).
--- Insert Table 1 about here ---

Results
Overall Analysis
The overall analysis revealed a large imitation effect, gz = 0.95, SE = 0.04, CI 95% =
[0.88, 1.02], m = 268, k = 7561. However, the I² coefficient indicated that 83% of the
variability was due to heterogeneity of the effect sizes2. Adding dependent variable to the
model as a moderator revealed a significant modulation, F(2, 69.3) = 20.51, p < .001, with a
larger effect size for RTs, gz = 1.03, SE = 0.04, CI 95% = [0.95, 1.12], m = 242, k = 546, than for
ERs, gz = 0.69, SE = 0.04, CI 95% = [0.60, 0.77], m = 96, k = 142, and KMs, gz = 0.77, SE = 0.10,
CI 95% = [0.56, 0.97], m = 26, k = 68.
Publication Bias
An inspection of the funnel plots revealed that the effect size distribution was
symmetrical for all three dependent variables (Figures 4-6). In line with this visual analysis,
Egger’s regression test found no relation between sample size and effect size for either RTs,
z = -1.55, p = .121, ERs, z = -1.86, p = .062, or KMs, z = 0.64, p = .521 (Sterne & Egger, 2005).
Although the ER regression test approached significance, the funnel plot suggested that this
was due to a single study with a large effect size and a large sample. Indeed, when this study
was removed, there was no longer evidence for ER funnel plot asymmetry, z = -1.00, p =
.317. In conclusion, the funnel plot analyses suggest it is unlikely that publication bias had a
strong influence on the main effect of automatic imitation. P-curve analyses testing for
publication bias at the moderator level will be reported together with the corresponding
moderator effects.
--- Insert Figures 4-6 about here ---
Spatial Compatibility. We first explored the effect of simple spatial compatibility on
RT/ER automatic imitation by comparing studies that confounded imitative and simple
spatial compatibility with studies that controlled for simple spatial compatibility either by
using orthogonal stimulus-response compositions, by separately manipulating imitative and
spatial compatibility, or by using stimuli that could not be classified on a simple spatial
dimension3. This revealed smaller effect sizes for both RTs, t(231.0) = -4.15, p < .001, and
ERs, t(90.8) = -3.51, p < .001, when simple spatial compatibility was controlled. However,
robust automatic imitation was still present even when simple spatial compatibility did not
contribute (Table 2 and 3).
Second, we investigated the effect of orthogonal spatial compatibility on RT/ER
automatic imitation by comparing studies that used orthogonal stimulus-response
compositions (i.e., orthogonal) with studies that separately manipulated imitative and
spatial compatibility (i.e., independent). Importantly, this analysis was restricted to studies
using finger lifting/tapping or hand opening/closing stimuli because other studies have not
yet used orthogonal stimulus-response compositions (Heyes, 2011). The finger analysis
indicated smaller effect sizes for RTs, t(38.5) = -4.95, p < .001, gdiff = 0.48, and ERs, t(11.8) = -
2.23, p = .046, gdiff = 0.23, in independent studies than in orthogonal studies. However, even
in independent studies, there was robust RT, gz = 0.78, SE = 0.08, CI 95% = [0.63, 0.93], and
ER, gz = 0.48, SE = 0.09, CI 95% = [0.30, 0.66] automatic imitation. The RT hand analysis also
indicated smaller effect sizes in independent studies, t(24.7) = -4.07, p < .001, gdiff = 0.38.
However, this analysis should be interpreted with care because it included only a single
independent study (Press et al., 2008). The ER hand analysis did not have sufficient dfs and
could therefore not be interpreted. Taken together, the spatial compatibility analysis thus
indicates that both simple and orthogonal spatial compatibility inflated automatic imitation,
but could not explain it.
Compatibility Type. The difference between movement and effector compatibility
was explored for RTs and ERs. The RT analysis found a smaller imitation effect for
movement compatibility than for effector compatibility, t(227.1) = 6.35, p < .001 (Table 2).
However, compatibility type was confounded with paradigm type and response effector
(Table 1). We therefore controlled for these variables in a second analysis, but the effect of
compatibility type remained significant, both p ≤ 0.024. The ER analysis mirrored the RT
pattern but did not reach significance, t(31.3) = 1.73, p = 0.093 (Table 3). This indicates that
automatic imitation was stronger in effector compatibility tasks than in movement
compatibility tasks.
The flexibility of RT and ER automatic imitation was investigated by comparing
conditions in which compatible sensorimotor association were formed with conditions in
which incompatible sensorimotor association were formed, and conditions in which no or
non-contingent sensorimotor associations were formed. However, the ER analysis could not
be interpreted because it did not have sufficient dfs for all relevant t contrasts. The RT
analysis revealed a marginally significant main effect, F(2, 17.8) = 2.76, p = 0.090, with a
smaller effect size for incompatible associations than for compatible associations, t(20.0) = -
2.40, p = 0.027, and a trend towards a smaller effect size for incompatible associations than
for no associations, t(16.3) = -1.99, p = 0.064. No difference was found between compatible
associations and no associations, t(11.9) = 1.04, p = 0.317 (Table 2).
In addition to the main analysis, we also performed a p-curve analysis on the RT data
(Supplementary Material 1). The half p-curve test revealed a significant right-skewed
distribution, z = -4.64, p < .001 (Figure 7). The robustness of this analysis was checked in
three steps. In a first step, we excluded one experiment with unclear hypotheses and one
experiment that used non-standard analyses. In a second step, we additionally excluded two
experiments in which non-contingent incompatible associations were used as a baseline
instead of no associations. Finally, in a third step, we explored the influence of removing the
lowest p-values from each of the analyses reported above. The results revealed a significant
half p-curve for the first two robustness analyses, both z ≤ -4.2, both p < .001, that was
resistant to removal of the 5 lowest p-values. In other words, the p-curve analysis showed
robust evidence for the flexibility of automatic imitation and thus strengthened the
preliminary evidence provided by the main analysis.
To investigate whether automatic imitation interferes with the motor goal or with
the motor plan, we compared forced choice and simple response tasks using the factor
paradigm type. This analysis was restricted to RTs because there are no meaningful errors in
simple response tasks. The RT analysis revealed a larger effect size in forced choice tasks
than in simple response tasks, t(148.6) = 7.31, p < .001, and this effect was still present
when the confounding influence of compatibility type was controlled, p < .001 (Table 1).
Importantly, however, a robust imitation effect was obtained in both tasks (Table 2). This
suggests that automatic imitation influenced both the motor goal and the motor plan.
Speed. To investigate the speed of automatic imitation, we investigated the
influence of SOA on automatic imitation. The SOA analysis was restricted to conditions
where the imperative cue was presented either together with the stimulus movement (i.e.,
SOA = 0) or after the stimulus movement (i.e., SOA > 0). Given that SOA varied not only
between but also within experiments, we furthermore tested both its between- and within-
experiment effect (Tanner-Smith & Tipton, 2014). The between-experiment analysis
revealed a negative effect of SOA on automatic imitation for both RTs, t(13.9) = -7.44, p <
.001, and ERs, t(7.0) = -3.83, p = .007. The within-experiment analysis confirmed this pattern
for RTs, t(13.2) = -2.55, p = .024, but not for ERs, t(4.9) = -0.91, p = .408. An inspection of
Figure 8 revealed that the imitation effect could be observed until SOAs of around 1000 to
1200 ms. Together, these results suggest that automatic imitation became weaker as SOA
increased but did not disappear unless very long SOAs were used.

Spatial Attention. The role of spatial attention on RT and ER automatic imitation was
investigated by looking at the effect of cue location and cue modality. The RT analysis
revealed a significant effect of cue location, t(39.8) = 3.81, p < .001, with a stronger
imitation effect when the cue was positioned near the stimulus movement compared to
when it was positioned away from the stimulus movement. The ER analysis did not find an
effect of cue location, t(21.4) = 1.18, p = .252, but the numerical pattern mirrored the RT
results (Table 3). The cue modality analysis was restricted to RTs because the ER analysis did
not have sufficient dfs to be interpreted. This revealed a marginally stronger effect size
when visual cues rather than auditory cues were used, t(5.3) = 2.32, p = 0.065 (Table 2).
However, there was also a substantial correlation between cue location and cue modality
(Table 1). When we included both moderators in the same model, a significant effect was
found for cue location, t(30.2) = 3.42, p = .002, but not for cue modality, t(7.5) = -0.64, p =
.542. Taken together, these analyses indicate that automatic imitation was weaker when
spatial attention was not directed to the stimulus movement.
To investigate the effect of action goals on automatic imitation, we compared
goalless stimulus movements with object-directed and communicative stimulus movements
for RTs and ERs. The RT analysis revealed a significant main effect of action goal, F(2, 33.5) =
11.19, p < .001, with a stronger imitation effect for goalless stimulus movements than for
both communicative stimulus movements, t(17.3) = 2.60, p = .019, and objected-directed
stimulus movements, t(37.5) = 4.5, p < .001 (Table 2). There was no significant difference
between communicative and object-directed stimulus movements, t(30.6) = 1.29, p = 0.205.

Although there was a substantial correlation between action goal and response effector, the
main effect of action goal did not change when response effector was included as a control
variable, F(2, 23.4) = 6.85, p = .005 (Table 1). In line with the RT analysis, the ER analysis also
demonstrated a main effect of action goal, F(2, 12.4) = 4.89, p = .027. This effect was
characterized by a stronger imitation effect for goalless stimulus movements than for
object-directed stimulus movements, t(8.0) = 3.12, p = .014, and a trend towards a stronger
imitation effect for goalless stimulus movements than for communicative stimulus
movements, t(11.1) = 2.14, p = .056. No difference was found between object-directed and
communicative stimulus movements, t(13.6) = 1.34, p = .202 (Table 3).
These findings speak against the hypothesis that goals have a stronger influence on
automatic imitation than movements. Furthermore, because the imitation effect was not
only weaker for object-directed stimulus movements but also for communicative stimulus
movements, they also speak against the hypothesis that objects may drive attention away
from the stimulus movement. Nevertheless, to further examine the attentional hypothesis,
we additionally tested whether RT automatic imitation differed between object-directed
stimulus movements that were performed towards an actual object and object-directed
stimulus movements that were performed towards an implied object (i.e., pantomimes). In
contrast with the attentional hypothesis, the effect sizes for pantomimes, gz = 0.66, SE =
0.09, CI 95% = [0.45, 0.87], and object-directed stimulus movements, gz = 0.71, SE = 0.13, CI
95% = [0.45, 0.97], were highly similar. Moreover, the action goal analysis confirmed the
significant main effect of action goal, F(3, 25.4) = 9.34, p < .001, with a stronger imitation
effect for goalless stimulus movements than for object-directed stimulus movements,
t(23.6) = 3.27, p = .003, pantomimes, t(8.8) = 4.71, p = .001, and communicative stimulus
movements, t(17.3) = 2.60, p = .019. None of the other comparisons reached significance, all
t(16.8) ≥ 1.52, all p ≥ .147.
Bottom-Up Animacy. The role of bottom-up animacy was tested by comparing RT,
ER, and KM automatic imitation of human, nonhuman, and geometrical models. However,
the ER analysis could not be interpreted because it did not have sufficient dfs. The RT
analysis revealed a significant main effect of animacy, F(2, 23.2) = 26.90, p < .001. Follow-up
tests indicated the presence of a linear trend with stronger automatic imitation for human
models than for nonhuman models, t(11.9) = 2.89, p = .014, and stronger automatic
imitation for nonhuman models than for geometrical models, t(21.7) = 2.97, p = .007 (Table
2). The KM analysis likewise found a main effect of animacy, F(2, 12.5) = 7.23, p = .008.
However, in contrast to our expectations, a weaker imitation effect was obtained for
nonhuman models than for both human models, t(8.6) = -3.46, p = .008, and geometrical
models, t(10.7) = -3.48, p = .005, and no difference was observed between human models
and geometrical models, t(15.6) = 0.89, p = 0.386 (Table 4). Although bottom-up animacy
was highly correlated with motion profile in KM studies (V = .56), including both moderators
in the same model did not change the animacy main effect, F(2, 9.9) = 8.13, p = .008.
In addition to the main analysis, we also conducted a p-curve analysis on the
difference between human and nonhuman models. Comparisons between human and
geometrical models were not included. To maximize the number of data points in the
analysis, we included both eligible RT (n = 3) and KM (n = 1) studies (Supplementary
Material 2). The half p-curve analysis revealed a significantly right-skewed distribution, z = -
3.60, p < .001 (Figure 8). We then repeated the p-curve analysis including only RT studies.
However, this also indicated significant right skew, z = -4.22, p < .001. Taken together, the
RT analysis showed that automatic imitation became stronger when the similarity between
the model and the participant increased. In contrast, the KM analysis revealed an
unexpected pattern with reduced automatic imitation for nonhuman models but not for
geometrical models.
Top-Down Animacy. The influence of top-down animacy was investigated for RTs
and ERs. That is, we examined whether automatic imitation of human/ambiguous models
was reduced when participants were told that these models were nonhuman and whether
automatic imitation of nonhuman/ambiguous models was increased when participants were
told that these models were human. However, we will report only the RT analysis for
nonhuman/ambiguous models because the other analyses did not have sufficient dfs to be
interpreted. In contrast to our expectations, we found no increased automatic imitation in
the human belief condition, t(6.8) = 0.60, p = 0.566 (Table 2).
We followed up on these results with a p-curve analysis. Similar to bottom-up
animacy, both RT (n = 3) and KM (n = 1) studies were included to maximize the number of
data points in the analysis (Supplementary Material 3). In line with the main analysis, the
half p-curve analysis found no significant right skew, z = -0.9, p = .185. The flat test
furthermore indicated that the obtained curve was not significantly flatter than the curve
that is expected when studies are powered at 33%, z =-0.67, p =.251. This indicates that
there is currently no evidence for either the presence or absence of a top-down animacy
effect (Figure 9). Similar to bottom-up animacy, including only RT studies did not change the
results.
Motion Profile. To explore the influence of motion profile on KM automatic
imitation, we compared biological and non-biological motion profiles. However, there was
no difference in effect size between the two motion profiles, t(7.5) = 0.02, p = .986, and
controlling for the relation between motion profile and bottom-up animacy (V = .56) did not
change this, t(7.4) = -0.62, p = .554 (Table 4).
For RT and ER automatic imitation, we instead compared one-frame stimuli with
two-frame stimuli and multi-frame stimuli. The RT analysis revealed a main effect of number
of frames, F(2, 44.9) = 3.52, p = .038. This effect was characterized by a smaller effect size
for one-frame stimuli than for two-frame stimuli, t(19.3) = -2.68, p = .015, and a marginally
smaller effect size for one-frame stimuli than for multi-frame stimuli, t(18.6) = -1.79, p =
.090 (Table 2). No difference was found between two-frame stimuli and multi-frame stimuli,
t(207.4) = -1.10, p = .273. The ER analysis revealed a marginally significant main effect of
number of frames, F(2, 16.9) = 3.54, p = .052. Similar to RTs, this was due to a smaller effect
size for one-frame stimuli than for two-frame stimuli, t(7.3) = -2.72, p = .028. However,
there was no difference between multi-frame stimuli and one-frame stimuli, t(7.5) = -1.80, p
= .113, or two-frame stimuli, t(81.6) = -1.41, p = .162 (Table 3). In sum, the KM analysis
found no effect of motion profile and the RT/ER analysis found no evidence that automatic
imitation was stronger when the stimulus movement was presented with a short video
rather than with a sequence of two frames.

Perspective. The role of perspective was examined for RTs and ERs. Studies in which
the perspective of the model could not be determined (e.g., a moving dot) were not
included in the analysis. Neither the RT analysis, t(113.3) = 1.53, p = .130, nor the ER
analysis, t(40.8) = 1.24, p = .220, found a difference between first- and third-person
perspective (Table 2 and 3). Importantly, as facial actions are assumed to be learned mainly
through social interaction rather than through self-observation, we also looked at the
influence of perspective while excluding studies on facial automatic imitation, but this
likewise revealed no difference in either the RT, t(110.3) = 1.48, p = .142, or ER, t(34.4) =
1.29, p = .207, analysis. However, Table 1 indicated a substantial correlation between
perspective and response effector for RTs. In a second step, we therefore repeated the RT
analysis while controlling for effector, but the effect of perspective remained non-
significant, t(75.4) = 0.01, p = .996. In other words, there was no support for the hypothesis
that automatic imitation depends on the perspective in which the stimulus movement is
presented.
Gender Overlap. To investigate the hypothesis that automatic imitation depends on
whether the gender of the participant and the model overlap, we looked at the interaction
between participant gender and model gender for RTs and ERs. However, prior to this
analysis, we first explored whether there was a main effect of these two variables. It is
important to note here that non-human agents were not included in any of the model
gender analyses because these models have no gender. The participant gender analysis
found no difference between predominantly male and predominantly female samples for
RTs, t(111.1) = 0.39, p = .695, but a stronger effect in predominantly female samples for ERs,
t(41.8) = 2.20, p = .033. The model gender analysis revealed a main effect for both RTs, F(2,
64.8) = 6.75, p = .002, and ERs, F(2, 28.4) = 3.79, p = .035. The RT main effect was
characterized by stronger automatic imitation for female models than for ambiguous
models, t(32.7) = 2.12, p = .042, and stronger automatic imitation for ambiguous models
than for male models, t(90.9) = 2.49, p = .015 (Table 2). The ER main effect followed a
similar pattern with stronger automatic imitation for female models than for male models,
t(17.3) = 2.75, p = .014, and marginally stronger automatic imitation for ambiguous models
than for male models, t(55.7) = 1.77, p = .082, but no significant difference between female
models and ambiguous models, t(14.3) = 1.30, p = .214 (Table 3).
Given that most of the participants in the dataset were female (RT: 60%, ER: 61%), an
interesting hypothesis is that the main effect of model gender was driven by a participant
gender (male vs. female) x model gender (male vs. ambiguous) interaction. Note that the
interaction analysis did not include female models because only 3 experiments in the
dataset had used a female model in a predominantly male sample. As a result, the analysis
had insufficient dfs to be interpreted if female models were included. In line with the gender
overlap hypothesis, the RT analysis revealed a significant participant gender x model gender
interaction, t(40.7) = -2.33, p = .025. Follow-up tests found no difference between
predominantly male and predominantly female samples when the model was ambiguous,
t(62.9) = -0.54, p = .592, but a larger effect size in predominantly male samples when the
model was male, t(21.8) = 2.33, p = .029 (Figure 10). The participant gender x model gender
interaction did not reach significance for ERs, t(31.9) = -0.56, p = .580, but the numerical
pattern mirrored the RT data.
Nevertheless, an important caveat is that we coded samples as “predominantly
female” even if only 51% of the participants were female. Instead, a more suitable analysis
may be to analyze the proportion of female participants as a continuous variable. However,
this analysis likewise revealed a significant participant gender x model gender interaction,
t(19.1) = 2.11, p = .048, with a significant negative relation between proportion of females
and automatic imitation when the model was male, t(13.2) = -2.63, p = .021, β = -0.95, CI
95% = [-1.72,- 0.17], but no relation when the model was ambiguous, t(28.7) = -0.40, p =
.696, β = -0.10, CI 95% = [-0.59, 0.40]. Taken together, the gender overlap analysis thus
indicates that automatic imitation was weaker when there was a discrepancy between the
gender of the model and the gender of the participant.
Response Overlap. The response overlap analysis was restricted to studies using
finger/lifting tapping stimuli because these were the only stimuli that differed systematically
with respect to response overlap. More precisely, we compared RTs and ERs of studies in
which the required response was a finger lift/tap (i.e., response overlap) with RTs and ERs of
studies in which the required response was a key release/press (i.e., no response overlap).
In support of the response overlap hypothesis, the RT analysis revealed stronger automatic
imitation, t(69.9) = 3.88, p < .001, and the ER analysis revealed marginally stronger
automatic imitation, t(43.5) = 1.77, p = .084, for finger lift/tap responses than for key
release/press responses (Table 2 and 3).
Empathy. The relation between empathy and automatic imitation was assessed for
RTs and ERs, but the ER analysis did not have sufficient dfs to be interpreted. As a result, we
will only report the RT analysis (N = 975). As noted before, we included data from the IRI,
EAI, and EQ. The cognitive empathy analysis was conducted on the IRI Perspective Taking
and EAI Perspective Taking scales. The emotional empathy analysis was conducted on the
IRI Empathic Concern and EAI Affective Responses scales. Although our analysis focuses on
cognitive and emotional empathy, we also report correlations with IRI Fantasy and IRI
Personal Distress because the IRI has often been used in the automatic imitation literature.
In contrast to the IRI and EAI, the EQ is a unidimensional instrument that combines cognitive
and emotional empathy. Therefore, as a final analysis, we also investigated whether
automatic imitation was correlated with a composite measure of cognitive and emotional
empathy obtained from the IRI, EAI, and EQ data.
To estimate the relation between automatic imitation and empathy, we entered the
mean Fisher transformed correlation of each relevant experiment into the analysis. We then
used hierarchical effects weights instead of random effects weights to account for the fact
that some experiments were part of the same study. However, the results revealed no
significant correlation between automatic imitation and either cognitive empathy, t(9.6) =
0.71, p = 0.497, or emotional empathy, t(9.6) = 1.49, p = 0.168 (Figures 12-13). To
determine whether this non-significant correlation could be interpreted as evidence of
absence, we then tested whether the 90% confidence interval around the obtained
correlation was captured by the equivalence range of r = [-.20, .20]. More specifically, this
would show that the correlation was significantly smaller than r = .20 at α = .05 (Rogers,
Howard, & Vessey, 1993). A r = .20 explains 4% of the variance and can be considered a
small-to-medium effect size according to Cohen (1988). We chose r = .20 as the smallest
effect size of interest because an inspection of the literature revealed that this was the
smallest effect size that could be detected with reasonable power by the best powered
study (i.e., N = 193 for 80% power). The fact that none of the studies were powered to test
smaller effect sizes indicates that such effects were considered practically or theoretically
irrelevant. The 90% confidence interval of neither cognitive empathy, CI 90% = [-.06, .13],
nor emotional empathy, CI 90% = [-.01, .10], exceeded the equivalence range. This indicates
that the correlation was significantly smaller than r = .20.
In line with the above analyses, the analysis of IRI Fantasy, t(8.7) = 1.15, p = 0.279, CI
90% = [-.03, .12], and IRI Personal Distress, t(8.5) = 1.62, p = 0.141, CI 90% = [-.01, .20],
likewise showed that the correlations with automatic imitation did not differ significantly
from zero, and that they were significantly smaller than r = .20 (Table 2). Finally, no
significant correlation with automatic imitation was found when we calculated a composite
measure of cognitive and emotional empathy, t(10.0) = 1.29, p = 0.225, and this correlation
was again significantly smaller than r = .20, CI 90% = [-.02, .09].
--- Insert Figures 12-13 about here ---
Autism Spectrum Disorder. To test the hypothesis that automatic imitation is
impaired in individuals with ASD, we first tested for differences in RT automatic imitation
between control samples and ASD samples (Table 2). The results of this analysis revealed
that the imitation effect between the two groups was indistinguishable, t(9.7) = 0.24, p =
0.819. Similar to the empathy analyses, we then performed an equivalence test to
determine whether the non-significant result indicated that the effect was absent (Rogers et
al., 1993). The equivalence range was set to gz = [-0.30, 0.30], which corresponds to a small-
to-medium effect size according to Cohen (1988). The approach described for empathy
revealed that d = 0.50 was the smallest effect size that could be detected with reasonable
power by the best powered study (i.e., N = 64 per group for 80% power). However, because
we did not consider effect sizes below d = 0.50 to be theoretically irrelevant, we decided to
use a similar threshold as in the empathy analysis, namely a small-to-medium effect size.
The 90% confidence interval [-0.22, 0.28] was captured by the equivalence range, indicating
that the effect was significantly smaller than gz = 0.30.
Nevertheless, it could be argued that there were demographic differences between
the two groups (e.g. age, gender, IQ, …) that might have concealed a potential effect. To
address this issue, we performed a second analysis including only the studies that used both
a control group (N = 192) and an ASD group (N = 201). Given that the groups were matched
on demographic variables in all of these studies, this approach controls for a potential
influence of these variables. Notwithstanding, there was again no significant difference
between the two groups, t(16.5) = 0.07, p = .947, and the difference was significantly
smaller than gz = 0.30, CI 90% = [-0.27, 0.30]. Finally, because the previous analyses did not
include ER data or two KM studies on ASD, we repeated the second analysis using not only
RTs but also ERs and KMs. This resulted in N = 213 in the control sample and N = 220 in the
ASD sample. However, in line with the previous analyses, there was again no significant
difference, t(21.0) = -0.06, p = .952, and the difference was significantly smaller than gz =
0.30, CI 90% = [-0.28, 0.26]4. Taken together, the ASD analysis thus strongly suggests that
there was no difference in automatic imitation between the control samples and the ASD
samples.
Secondary Moderators
Age. The influence of age on automatic imitation was explored for RTs and ERs.
However, neither the RT analysis, t(34.7) = 0.85, p = .404, nor the ER analysis, t(16.1) = -1.64,
p = .121, revealed a significant effect (Table 2 and 3).

Trials. The relation between the number of trials and automatic imitation was
investigated for RTs and ERs. The RT analysis indicated that the imitation effect decreased
when the number of trials increased, t(53.6) = -2.01, p = .050. The ER analysis mirrored this
pattern but did not reach significance, t(22.7) = -1.70, p = .103.
Response Effector. The influence of effector was investigated for RTs and ERs.
However, the ER analysis did not have sufficient dfs to be interpreted. The RT analysis
revealed a significant main effect of effector, F(3, 13.8) = 4.00, p = .030 (Table 2). However,
there was a substantial correlation between effector and perspective, action goal, response
device, and compatibility type. Although the main effect of effector was still present when
controlling for response device, p = .003, it disappeared when controlling for perspective,
action goal, or compatibility type, p ≥ .141. Since the response effector is determined by the
stimulus movement, this suggests – all else being equal – that stimulus differences do not
have a discernible influence on automatic imitation. In line with this view, none of the
effects that were confounded with response effector disappeared when its influence was
controlled.
--- Insert Table 2, 3, and 4 about here ---
Sensitivity Analyses
To test the robustness of the RT and KM findings reported in Tables 2 and 4, we
performed four sensitivity analyses. In a first sensitivity analysis, we varied the ρ parameter
of the RVE models (Fisher & Tipton, 2015). However, none of the results changed when the
default ρ = .80 setting was replaced by ρ = .00, ρ = .20, ρ = .40, ρ = .60, ρ = .80, or ρ = 1.00. In
a second sensitivity analysis, we used a different method to estimate the effect size for
experiments that did not provide sufficient statistical information. More precisely, we
imputed the median correlation and standard deviation instead of the mean correlation and
standard deviation. This procedure resulted in a median correlation of r = .92 for RTs and r =
.71 for KMs and in a median standard deviation of SD = 26.72 for RTs. In line with the first
analysis, the second sensitivity analysis did not change any of the results.
In a third sensitivity analysis, we removed effect sizes corresponding to geometrical
models from the meta-analysis. Although geometrical objects have been interpreted as
inanimate models in the literature (Gowen & Poliakoff, 2012), it could be argued that these
studies can no longer be seen as automatic imitation studies. Removing geometrical models
caused the marginally significant effect of cue modality to disappear, t(5.3) = 1.54, p = .182,
but no other changes were observed. Finally, in a fourth sensitivity analysis, we added SOA
as a covariate to the RT analyses. Since we did not exclude studies on the basis of SOA in the
present meta-analysis, it could be argued that some of the studies used SOAs that were too
long to obtain a reliable imitation effect5. Even though Table 1 did not reveal substantial
correlations between SOA and the other moderators, we decided to explicitly address
potential SOA confounds by controlling for its influence in a sensitivity analysis. Adding SOA
to the model removed the marginally significant effect of cue modality, t(5.4) = 1.17, p =
.290, the main effect of number of frames, F(2, 45.9) = .69, p = .509, and the linear effect of
number of trials, t(47.8) = -1.20, p = .235.
In summary, the sensitivity analyses indicated that the results for cue modality,
number of frames, and number of trials were sensitive to methodological decisions.
However, an inspection of Table 2 shows that the evidence for these effects was already
relatively weak in the main analysis. Taken together, it can therefore be concluded that
three small effects proved to be unstable but that none of the other main findings were
sensitive to four important methodological decisions.
Discussion
Automatic imitation is widely studied in cognitive (Heyes, 2011), social (Wang &
Hamilton, 2012), and developmental psychology (Ray & Heyes, 2011), and in animal
cognition (Heyes, 2012), robotics (Chaminade & Cheng, 2009), and neuroscience (Bien et al.,
2009; Brass & Heyes, 2005; Hamilton, 2015). Although the moderators of automatic
imitation have already been reviewed extensively in previous work (Chaminade & Cheng,
2009; R. Cook et al., 2014; Gowen & Poliakoff, 2012; Heyes, 2011; Press, 2011; Sowden &
Shah, 2014; Wang & Hamilton, 2012), the evidence has often been inconsistent and
important questions remain. Therefore, the goal of the current study was to answer seven
key theoretical questions on automatic imitation: What processes contribute to automatic
imitation? Is automatic imitation flexible? When during action preparation does automatic
imitation operate? Is automatic imitation really automatic? What is the role of action goals
in automatic imitation? Is automatic imitation sensitive to the similarity between actor and
imitator? Is automatic imitation related to social abilities? In what follows, we will discuss
the answers obtained to each of these questions, and their implications for related research
fields. We will furthermore elaborate on whether KM automatic imitation is a valid measure
of automatic imitation, and go into the limitations of the present meta-analysis.

What Is The Role of Spatial Compatibility? In line with previous research, we found
that automatic imitation was weaker but not absent when spatial processes were controlled
(Heyes, 2011). The results furthermore showed that both simple and orthogonal spatial
compatibility influenced automatic imitation. This goes against earlier work in which
automatic imitation was found to be insensitive to orthogonal spatial effects (Jimenez et al.,
2012; Press et al., 2008) and suggests that researchers should opt for paradigms that allow
spatial compatibility to be separated from imitative compatibility (e.g., Bertenthal et al.,
2006; Catmur & Heyes, 2011) or use gestures that are matched on spatial properties
(Bortoletto et al., 2013; Cracco et al., 2018). The finding that spatial processes can interfere
with automatic imitation furthermore highlights the need to explore potential non-social
confounds in social paradigms. This issue is not only relevant for research on automatic
imitation (Heyes, 2011), but also for research on related topics such as joint action that
suffers from similar confounds (Dolk, Hommel, Prinz, & Liepelt, 2013). As shown in the
current meta-analysis, a valuable approach is to show that the explanatory power of social
processes exceeds that of non-social processes.
However, in addition to spatial compatibility, it is important to also consider other
non-social processes that might contribute to automatic imitation. One such process is
emulation, which is the phenomenon that task-irrelevant object properties modulate
responses to task-relevant stimuli (Edwards, Humphreys, & Castiello, 2003; Symes, Ellis, &
Tucker, 2005; Tucker & Ellis, 1998; Wilf, Holmes, Schwartz, & Makin, 2013). When automatic
imitation studies have an object in the stimulus display (e.g., Ocampo & Kritikos, 2010; van
Schie et al., 2008), emulation can act as a confound (Heyes, 2011). However, most studies
do not include objects (e.g., Brass, Bekkering, et al., 2001; Brass et al., 2000; Heyes et al.,
2005; Stürmer et al., 2000). Given that the current meta-analysis revealed a robust imitation
effect also in these studies, it is clear that emulation cannot explain automatic imitation.
Is There a Difference Between Effector and Movement Compatibility? The analysis
of effector and movement compatibility revealed that automatic imitation was stronger
when it was measured in terms of effector compatibility. A potential explanation for this
finding is that, on incompatible trials of effector compatibility paradigms, the stimulus
movement (e.g., lift index finger) is integrated with the response movement (e.g., lift middle
finger) into a composite movement (e.g., lift both index and middle finger). More
specifically, such a mechanism makes it more difficult to select the correct response on
effector incompatible trials because participants then have to inhibit not only the stimulus
movement but also the composite movement. Given that automatic imitation depends on
the degree of response conflict, such a mechanism can explain why effector compatibility
was stronger than movement compatibility.
The idea that multiple actions can be integrated into a composite action is consistent
with evidence that the actions of multiple agents can be co-represented in the motor
system, as shown in studies on automatic imitation (Cracco & Brass, 2017; Cracco et al.,
2015), the mirror system (Cracco, De Coster, Andres, & Brass, 2016), and joint action
(Ramenzoni, Sebanz, & Knoblich, 2014; Tsai, Sebanz, & Knoblich, 2011). Being able to
integrate planned actions with observed actions into a composite action may furthermore
be an important tool to achieve interpersonal coordination in joint action tasks that require
the cooperation of multiple persons (e.g., rowing). In particular, it allows each person to
represent the actions of the others as a “group action”. This, in turn, enables the use of
motor prediction mechanisms to achieve synchronized group behavior (Colling, Knoblich, &
Sebanz, 2013; Kourtis, Sebanz, & Knoblich, 2013).

Consistent with the notion that automatic imitation is a flexible process, we found
weaker imitative responses when incompatible sensorimotor associations were established
(Brass & Heyes, 2005; Brass & Muhle-Karbe, 2014; R. Cook et al., 2014). However, there was
no increase in automatic imitation when compatible associations were established instead.
This suggests that perception-action associations are overlearned due to years of experience
and for that reason are difficult to strengthen. The finding that imitative tendencies are
flexible accords well with neural evidence that mirror activation is likewise modulated by
incompatible sensorimotor training (Catmur, Walsh, & Heyes, 2007; R. Cook et al., 2014;
Heyes, 2010). From the perspective of ML theories, this shows that associations between
perception and action are not inborn but acquired through associative learning (R. Cook et
al., 2014; Heyes, 2010, 2016). Interestingly, the claim that imitation is learned was recently
supported by a comprehensive longitudinal study that followed 106 infants until the age of
9 weeks old and found no evidence of neonatal imitation (Oostenbroek et al., 2016) in
contrast with what has been reported before (Meltzoff & Moore, 1983, 1989; Simpson et
al., 2014). Together with our results, this indicates that “nature” does not provide humans
with an inborn capacity to imitate but rather with an inborn capacity to learn imitation
through “nurture”.
Furthermore, flexible perception-action links offer an elegant solution to the
paradoxical finding that action observation does not always trigger an imitative response
but sometimes also a complementary response (Ocampo & Kritikos, 2010; Ocampo et al.,
2011; Sartori, Betti, & Castiello, 2013; van Schie et al., 2008). For instance, when someone
throws you a ball, your initial response is not to imitate but to open your hands so that you
can catch the ball. In the associative learning framework, complementary actions can be
explained as contextual modulations of the dominant imitative association (R. Cook,
Dickinson, & Heyes, 2012). That is, it can be learned through experience or instruction that
in specific contexts certain actions should be followed by a complementary rather than an
imitative response (R. Cook et al., 2012; Ocampo & Kritikos, 2010; Ocampo et al., 2011; van
Schie et al., 2008). As such, associative learning mechanisms allow perception-action
associations to support social interaction well beyond mere imitation (Sebanz, Bekkering, &
Knoblich, 2006).
The analysis of forced choice and simple response tasks revealed that both tasks
produced reliable automatic imitation, but also that imitative responses were stronger in
forced choice tasks. The finding that automatic imitation was stronger in forced choice tasks
shows that it interfered with the motor goal (“what finger should I lift?”), whereas the
finding that it persisted in simple response tasks shows that it influenced the motor plan
(“what muscles will achieve this?”) as well (Wong et al., 2015). This interpretation is
consistent with neuroscientific evidence that seeing an action with the intention to imitate
does not only activate brain regions involved in action planning (Bien et al., 2009;
Molenberghs et al., 2012) but also activates primary motor regions that execute planned
actions (Caspers et al., 2010), and is further supported by single cell evidence that mirror
neurons can also be found in the primary motor cortex of the macaque brain (Vigneswaran,
Philipp, Lemon, & Kraskov, 2013).

With regard to simple response tasks, it is interesting to note that stimulus-response
compatibility research outside the domain of automatic imitation has found little support
for the idea that task-irrelevant stimuli can interfere with prepared responses (Brass,
Bekkering, et al., 2001; Hommel, 1996). A potential explanation for this pattern can be
found in the IM assumption that observed and planned movements are represented in the
form of shared perception-action representations (Brass & Heyes, 2005; Prinz, 1997). In
particular, IM argues that the visual image of an action is part of its motor representation
and therefore that observed actions have direct access to the corresponding motor code. As
a consequence, observed actions can influence behavior even after the response has been
selected (Brass, Bekkering, et al., 2001). In contrast, abstract stimuli with low ideomotor
compatibility are not able to directly activate the motor code and are instead assumed to
rely on a translation process (Boyer et al., 2012; Sauser & Billard, 2006). Therefore, task-
irrelevant stimuli in non-imitative tasks can only access the motor code if the response has
not yet been selected (Brass, Bekkering, et al., 2001).
The extent to which automatic imitation is automatic was evaluated by looking at
two components of automaticity, namely speed and attention. The speed analysis revealed
that automatic imitation became weaker when the duration between the stimulus
movement and the imperative cue increased. This suggests that the processes involved in
automatic imitation are fast, and as such that it is an automatic process (Moors & De
Houwer, 2006). Nevertheless, it is of note that more subtle SOA analyses have shown that
automatic imitation is strongest when there is a short interval of 80 to 150 ms between the
presentation of the stimulus movement and the presentation of the imperative cue (Catmur
& Heyes, 2011; Cracco et al., 2018; Wiggett et al., 2013). Together with the results of the
current study, this suggests that automatic imitation is transient, but also that a short period
of time is needed before the influence of observed actions on behavior reaches its maximal
potential. Given that motor preparation processes are known to operate at a nearly
instantaneous time scale (Wong et al., 2015), a plausible hypothesis is that the delay reflects
the visual processing of observed actions rather than actual imitation.
With regard to attention, the results showed that automatic imitation was reduced
but not eliminated when participants’ attention was directed away from the stimulus
movement. This is consistent with evidence that automatic imitation is weaker (Cracco &
Brass, 2017) or absent (Bach et al., 2007; Chong et al., 2009) when attention is directed
away from the stimulus movement, but also with evidence that imitative responses do not
disappear under conditions of high attentional load (Catmur, 2016). In particular, it indicates
that automatic imitation is sensitive to variations in attention but persists in its absence.
Together with the fact that automatic imitation is an unintentional process (Heyes, 2011),
the present automaticity analysis thus indicates that its classification as “automatic” is
justified (Moors & De Houwer, 2006).
Nevertheless, a full automaticity investigation also requires an evaluation in terms
consciousness (Moors & De Houwer, 2006). Specifically, automatic imitation can be
considered subconscious at the input level if it does not require conscious processing of the
stimulus movement and subconscious at the process level if subjects are unaware of their
imitative behavior (Moors & De Houwer, 2006). An inspection of the literature indicates that
there is currently no evidence that automatic imitation is unconscious at the input level
because it was shown to be absent when the stimulus movement was presented
subliminally (Mele et al., 2014). The same is true at the process level because participants
typically “feel” the urge to imitate the stimulus movement. This is in sharp contrast with
studies on motor mimicry where subjects are often unaware of their imitative tendencies
(Chartrand & Bargh, 1999). Taken together, it can thus be concluded that automatic
imitation is automatic with respect to some but not all components of automaticity (Moors
& De Houwer, 2006). However, additional research will be needed to understand how
automatic imitation varies with different degrees of attention or awareness, whether the
role of attention depends on the type of attention under consideration (e.g., spatial vs. non-
spatial), and at what stage during processing the delay in automatic imitation occurs (i.e.,
visual, visuomotor, or motor).
In contrast to the hypothesis that action goals have a stronger influence on
automatic imitation than movement parameters (Bouquet et al., 2011; Chiavarino et al.,
2013), goalless stimulus movements were found to produce stronger imitative responses
than object-directed or communicative stimulus movements. Furthermore, against the view
that attentional processes might drive attention away from the stimulus movement and
towards the action object (Bird, Brindley, et al., 2007; Leighton, Bird, & Heyes, 2010), no
difference was observed between object-directed movements, pantomimes, and
communicative movements. These findings go against previous research on intentional
imitation (Avikainen et al., 2003; Bekkering et al., 2000) and the mirror neuron system
(Caspers et al., 2010; Rizzolatti & Sinigaglia, 2010), and are instead more consistent with
reports of cross-contextual imitation (Genschow & Florack, 2014; Genschow et al., 2013;
Genschow & Schindler, 2016). However, does this mean that, despite popular belief (Bien et
al., 2009; Brass & Heyes, 2005; Catmur et al., 2009; R. Cook et al., 2014; Hamilton, 2015;
Heyes, 2011; Simpson et al., 2014), automatic imitation recruits different processes than
intentional imitation and does not activate the mirror neuron system?
To answer this question, it is interesting to consider the study of Chiavarino et al.
(2013) in which automatic imitation was likewise found to be weaker for goalless stimulus
movements than for goal-directed stimulus movements. In line with goal-directed theories
of imitation, these authors argued that goalless actions were coded in terms of movement
characteristics and goal-directed actions in terms of action goals. This can lead to stronger
automatic imitation for goalless actions if movement coding is stronger than goal coding
(Chiavarino et al., 2013), but also if goal coding is slower than movement coding. The latter
is consistent with evidence that it is the motor goal and not the motor plan that accounts
for most of the response time in choice tasks (Wong et al., 2015). Constructing a motor goal
in terms of goal coding is more abstract and may therefore be more difficult than in terms of
movement coding. This, in turn, might make goal coding slower and could explain why it
does not have strong influence in fast-paced automatic imitation paradigms. In support of
this idea, increased automatic imitation for goal-directed stimulus movements was reported
in a KM study that measured automatic imitation over an extended period of time (Bouquet
et al., 2011). This suggests that variations in time course could explain why action goals have
a different effect on automatic imitation than on intentional imitation or mirror activation.
From this perspective, an interesting avenue for future work is to explore whether the
influence of action goals on automatic imitation depends on the time available to process
the stimulus movement (i.e., SOA).

In line with the hypothesis that self-other overlap is an important modulator of
automatic imitation (Chaminade & Cheng, 2009; Gowen & Poliakoff, 2012; Press, 2011), the
results indicated that RT automatic imitation increased linearly from geometrical models via
nonhuman models to human models. This finding accords well with previous research that
likewise found stronger automatic imitation for human models than for nonhuman models
(e.g., Press et al., 2005, 2006, 2007) and suggests that the absence of a human bias in some
studies might have been caused by the fact that they used models that were too humanlike
to produce an effect (Cracco et al., 2015). In contrast to the RT analysis, the KM analysis
revealed an unexpected pattern with reduced automatic imitation for nonhuman models,
but not for geometrical models. While this pattern is consistent with evidence that dot
movements (e.g., Bouquet et al., 2007; Gowen et al., 2008; Jansson et al., 2007) but not
robotic movements (e.g., Kilner et al., 2003) produce KM automatic imitation, it is unclear
how these findings can be explained within the framework of ML theories. As discussed in
detail later on, a potential explanation might be that KM tasks of automatic imitation
measure something else than imitation.
In contrast to bottom-up animacy, we found no evidence that automatic imitation
was sensitive to animacy beliefs. This speaks against previous research reporting such
effects (Gowen et al., 2016; Klapper et al., 2014; Liepelt & Brass, 2010b; Stanley et al., 2007)
and instead is more consistent with the study of Press et al. (2006) where similar results
were obtained (see also Press, 2011). Notwithstanding, it is of note that the number of
studies investigating animacy beliefs is currently limited. For example, the p-curve analysis
indicated that there was no evidence either in support or against a top-down animacy
effect. It will therefore be important to further investigate this factor in future studies.
Indeed, evidence suggests that psychological beliefs on topics such as free will can have
pervasive effects on human performance (e.g., Rigoni, Kühn, Gaudino, Sartori, & Brass,
2012; Rigoni, Wilquin, Brass, & Burle, 2013). In addition to top-down animacy, there was
also no evidence that RT or KM automatic imitation was stronger when the stimulus
movements followed a biologically plausible motion profile (Kilner et al., 2007). However,
similar to top-down animacy, the KM motion profile analysis was based on just a limited set
of studies, highlighting the need for further research. Moreover, the difference between
biological and non-biological motion profiles is often subtle. It will thus be interesting to see
if an effect can be found when stronger manipulations are used. In support of this notion,
research has shown that automatic imitation is reduced when the stimulus movements are
impossible to execute (Liepelt & Brass, 2010a; Longo et al., 2008).
In addition to animacy, we also considered the visual perspective in which the
stimulus movement was presented. Given that ML theories assume that automatic
imitation, especially of non-facial actions, is learned through self-observation (Brass &
Heyes, 2005), it can be predicted from these theories that imitative responses should be
stronger when the stimulus movements are shown in a first-person perspective. However,
in contrast to this hypothesis, we found no influence of perspective on automatic imitation.
A potential explanation is that the effect of perspective depends on contextual moderators
(Bortoletto et al., 2013; Vogt et al., 2003), but there are currently no theories that describe
which variables should and which variables should not interact with visual perspective.
Therefore, a more parsimonious account is that perception-action links gradually generalize
over different perspectives as a result of continual exposure to the actions of others
(Burgess, van Baaren, Fitzgibbon, Fitzgerald, & Enticott, 2013). Indeed, in addition to self-
observation, social exposure is considered to be an important mechanism through which
automatic imitation develops (Brass & Heyes, 2005). Therefore, it is conceivable that with
sufficient social experience a third-person bias rather than a first-person bias starts to
emerge. In line with this view, a study using the rubber hand illusion found that the motor
system responds more strongly to observed actions that are attributed to another person
than to observed actions that are attributed to oneself (Schütz-Bosbach et al., 2006).
Finally, we looked at two factors that have so far not been studied in the literature,
namely gender overlap and response overlap. First, the gender analysis indicated that ER
automatic imitation was stronger in predominantly female samples than in predominantly
male samples and that RT automatic imitation was stronger when the model’s gender
matched the gender of the majority of the participants. This is consistent with the study of
Butler et al. (2015) in which automatic imitation of a female model was found to be stronger
for female participants than for male participants (see also Genschow et al., 2017).
However, this study did not include a male model and was thus unable to determine
whether the gender effect was driven by participant gender or by gender overlap. The
current results provide preliminary evidence that both factors may play a role.
Second, the response overlap analyses revealed that automatic imitation was
stronger when the response matched the stimulus movement not only in terms of visual
characteristics but also in terms of the other outcomes produced by these actions. This
finding supports the theoretical assumption that actions are represented in the motor
system as the combination of their sensory outcomes (Brass & Heyes, 2005). It furthermore
suggests that researchers should strive to maximize stimulus and response overlap in
automatic imitation studies. That is to say, our results indicate that finger tapping/lifting
stimuli may produce stronger automatic imitation in studies measuring finger tap/lift
responses than in studies measuring key press/release responses.
To conclude, the similarity analysis revealed that automatic imitation depends on
how similar the model is to the participant, but not on more subtle factors such as animacy
beliefs or motion profile. These results are consistent with the literature on mirror neurons
(Press, 2011) and motor mimicry (Chartrand & van Baaren, 2009). However, in contrast to
automatic imitation, mimicry research has focused more on overlap of opinions or social
groups than on overlap of physical characteristics (e.g., Castelli, Pavan, Ferrari, & Kashima,
2009; Yabar, Johnston, Miles, & Peace, 2006). Rather than feature overlap (Press, 2011), this
research has argued that similar others are mimicked more often because individuals use
mimicry to foster social relations and are more motivated to affiliate with persons that are
more like themselves (Lakin & Chartrand, 2003; Stel et al., 2016; Wang & Hamilton, 2012).
Interestingly, the idea that affiliation goals shape who we do and who we don’t imitate has
now also received support from studies on automatic imitation (e.g., J. L. Cook & Bird, 2011,
2012; Genschow & Schindler, 2016; Gleibs, Wilson, Reddy, & Catmur, 2016; Leighton, Bird,
Orsini, & Heyes, 2010; Rauchbauer, Majdandzic, Hummer, Windischberger, & Lamm, 2015).
However, the role of feature overlap in motor mimicry so far remains unclear.
Is Automatic Imitation Related to Empathy? The final question addressed in our
meta-analysis was the relation between automatic imitation and social abilities. According
to prominent accounts of social cognition, automatic imitation should be related to
empathy because embodied simulation and self-other control are essential components of
both constructs (Bird & Viding, 2014; Brass et al., 2009; Decety & Lamm, 2006; Sowden &
Shah, 2014). However, in contrast to this prediction, we found no association between
automatic imitation and empathy in a combined sample of almost 1000 participants (Butler
et al., 2015; Genschow et al., 2017). This finding highlights the need for a more nuanced
view on the role of self-other processes in social cognition. For example, self-other theories
could be updated to take into account the representational content on which a process
operates. That is, automatic imitation might be unrelated to empathy because the former
operates on motor representations and the latter on emotional representations even
though the performed operations are functionally equivalent. As would be expected from
such a hypothesis, research that looked at facial feedback in response to emotional
expressions has so far found consistent relations with empathy (Sonnby-Borgström, 2002;
Sonnby-Borgström, Jönsson, & Svensson, 2003).
Alternatively, it could be that a correlational approach is not suitable to investigate
the role of self-other processes in empathy, and that experimental approaches might be
more appropriate instead. In line with this view, it was recently demonstrated that short
periods of imitation or counter-imitation training can improve empathic abilities in a
subsequent unrelated task (De Coster, Andres, & Brass, 2014; De Coster, Verschuere,
Goubert, Tsakiris, & Brass, 2013; De Coster, Wiersema, Deschrijver, & Brass, 2017; de
Guzman, Bird, Banissy, & Catmur, 2015; Santiesteban, White, et al., 2012). Taken together,
the absence of a relation with empathy indicates that theories of social cognition should
consider the content of self-other representations and suggests that experimental
approaches might be more sensitive than correlational approaches.
Finally, although a relation with empathy has been reported in the domain of motor
mimicry, it is notable that the support for this relation is restricted to a single study
(Chartrand & Bargh, 1999) and could not be replicated in more recent work with higher
power (Genschow et al., 2017). Together with the current meta-analysis, this strongly
suggests that the assumed relation between measures of imitation and measures of
empathy should be reconsidered. More generally, our results call for caution when
interpreting correlations with personality measures in small sample studies. For example,
while preliminary studies has also reported relations between automatic imitation and
measures such as narcissism (Hogeveen & Obhi, 2013; Obhi et al., 2014), recent research
with larger samples has revealed that automatic imitation, but also motor mimicry, is largely
uncorrelated with personality traits (Butler et al., 2015; Genschow et al., 2017).
Is Automatic Imitation Impaired in Autism? Theories of embodied simulation and
self-other control furthermore have a prominent position in the literature on ASD (Spengler,
Bird, et al., 2010; Williams, 2008; Williams et al., 2001). The central tenet of simulation
theory is that social difficulties in ASD stem from a broken mirror neuron system (Williams,
2008; Williams et al., 2001). This hypothesis is challenged by proponents of the self-other
control theory who argue that it is not the mirror neuron system but the control over this
system that is abnormal in ASD (Spengler, Bird, et al., 2010). Although both theories assume
that automatic imitation is impaired in ASD, the former predicts hypo-imitation (Williams,
2008; Williams et al., 2001) and the latter predicts hyper-imitation (Spengler, Bird, et al.,
2010). However, In contrast to both hypotheses, we found no difference in automatic
imitation between ASD samples and control samples even after we controlled for
demographic variables. Given that our analysis contained 220 participants with ASD, this
clearly indicates that ASD is not associated with deficits in imitation or imitative control and
as such casts doubt on the idea that self-other processes are impaired in autism (Spengler,
Bird, et al., 2010; Williams, 2008; Williams et al., 2001).

Nevertheless, the absence of hypo- and hyper-imitation does not necessarily indicate
that imitation is normal in ASD. For example, the social top-down response modulation
(STORM) theory proposes that it is not imitation as such but the ability to adapt imitation to
the social context that is impaired in individuals with ASD (Forbes et al., 2016; Hamilton,
2013). This theory originated from evidence that healthy participants do while ASD
participants do not display stronger automatic imitation in prosocial contexts compared
with antisocial contexts (J. L. Cook & Bird, 2012) and is supported by related evidence that
the influence of eye contact on automatic imitation, which is present in control participants,
is absent in ASD participants (Forbes et al., 2016). Importantly, however, our results suggest
that STORM deficits should not be interpreted in terms of imitative control because such
problems should be visible as hyper-imitation (Spengler, Bird, et al., 2010). It instead seems
more likely that individuals with ASD fail to act on social cues because they process or
interpret these cues in an atypical manner (Chevallier et al., 2012; Hamilton, 2013).
However, from this perspective, deficits in imitation should no longer be considered a key
component of ASD and should instead be seen as a byproduct of abnormal social processing
(Blake et al., 2003; Chevallier et al., 2012; Klin et al., 2009).
To conclude, we found little evidence for the hypothesis that automatic imitation is
impaired in ASD. This finding is in strong disagreement with self-other theories of autism
(Spengler, Bird, et al., 2010; Williams, 2008; Williams et al., 2001). In particular, it suggests
that self-other processes are intact in individuals with ASD. Nevertheless, similar to
empathy, it remains possible that self-other impairments in ASD do not generalize across
domains but are instead restricted to a specific set of representations. In support of this
hypothesis, there is preliminary evidence that difficulties to distinguish self from other in
ASD might be restricted to the somatosensory domain (Deschrijver, Wiersema, & Brass,
2016; Deschrijver et al., 2017). Moreover, the STORM hypothesis that social regulation of
imitation is impaired in ASD is a promising avenue for further research (J. L. Cook & Bird,
2012; Forbes et al., 2016; Hamilton, 2013). Nevertheless, research on both theories is only
just developing and further investigation will be required to evaluate the predictions they
put forward. For example, recent work suggests that the effect of social cues on automatic
imitation may be limited to its spatial component (Marsh, Bird, & Catmur, 2016),
highlighting the need to differentiate between spatial and imitative compatibility in studies
of STORM. Finally, it is important to consider the implications of our results for imitative
behavior in related psychiatric disorders. In particular, considering that schizophrenia can be
seen as part of the autism spectrum (King & Lord, 2011), an interesting question is how
intact automatic imitation in ASD can be reconciled with recent reports of disturbed
automatic imitation in schizophrenic samples (Dankinsas, Melynyte, Siurkute, & Dapsys,
2017; Nakagawa & Hoshiyama, 2015).
Is Kinematic Imitation Really Imitation?
An important question that arises from our results is whether KM automatic
imitation actually measures imitation. In contrast to RT automatic imitation, we found no
evidence that KM automatic imitation became stronger when the model looked more like a
human. Instead, we found a reduction of KM automatic imitation when the model was a
robot but not when it was a moving dot. This is consistent with a large number of studies
that likewise found reliable KM automatic imitation in response to moving dots (Bouquet et
al., 2007; Gowen et al., 2008; Jansson et al., 2007; Roberts, Hayes, Uji, & Bennett, 2015;
Romero, Coey, Schmidt, & Richardson, 2012; Stanley et al., 2007). There are at least two
hypotheses that could explain the KM animacy pattern. A first hypothesis is that participants
interpret moving dots as human movements because moving dots, as opposed to robots,
can be ambiguous in terms of animacy. In line with this view, research has found that
human participants are skilled at recognizing biological motion from humanlike point-light
displays (e.g., Ulloa & Pineda, 2007). However, this hypothesis assumes that automatic
imitation is modulated by animacy beliefs, an assumption that could not be supported by
the data. Moreover, it cannot explain why KM automatic imitation was not reduced when
the stimulus movements followed a non-biological motion profile. Therefore, a second
hypothesis is that KM automatic imitation does not measure imitative processes, but
instead measures something else such as spatial processes.
The spatial hypothesis was recently tested by Hardwick and Edwards (2012) who
separately manipulated imitative and spatial compatibility. Participants in their task had to
make forward-to-backward (i.e., ↕) or left-to-right (i.e., ↔) arm movements while seeing
compatible or incompatible movements of an actor who either looked at the participant’s
face (i.e., ↑↓) or at the participant’s right side (i.e., ↑←). The manipulation of position
caused the relation between imitative and spatial compatibility to change from positive in
the face-to-face condition to negative in the face-to-side condition. This, in turn, made it
possible to calculate a main effect of imitative compatibility that was independent of spatial
compatibility. However, in contrast to RT automatic imitation (Heyes, 2011), the results
revealed that spatial compatibility did while imitative compatibility did not contribute to KM
automatic imitation. Although it is too soon to conclude that KM automatic imitation is
driven solely by spatial processes, these results clearly indicate that additional work is
needed to understand the different processes that are involved. Therefore, future research
may want to consider alternative KM measures that are typically independent of spatial
processes such as force (Salama et al., 2011), movement duration (Pfister et al., 2013) or
movement speed (Eaves, Haythornthwaite, & Vogt, 2014; Eaves, Turgeon, & Vogt, 2012).
Limitations of the Current Meta-Analysis
To finish, the present meta-analysis also has a number of limitations. A first
limitation is that we focused on automatic imitation and as such did not consider the closely
related phenomenon of motor mimicry. As noted throughout the paper, research on motor
mimicry has often addressed similar questions as research on automatic imitation
(Chartrand & Lakin, 2013; Chartrand & van Baaren, 2009; Heyes, 2011). There is
furthermore a strong overlap between the assumed mechanisms of the two constructs
(Genschow et al., 2017; Heyes, 2011). In consequence, it could be argued that studies on
motor mimicry should have been included in the meta-analysis. However, there are two
important distinctions between automatic imitation and motor mimicry that make this
difficult. First, automatic imitation differs from motor mimicry in the sense that it measures
imitative responses in situations where imitation can interfere with task performance
(Heyes, 2011). Second, there are substantial differences in the experimental procedure of
both constructs that lead to large differences in effect size (Genschow et al., 2017). A meta-
analysis combining multiple constructs is impossible to interpret because it compares
‘apples with oranges’ (Field & Gillett, 2010; Lipsey & Wilson, 2001). Therefore, even though
the present meta-analysis has implications for motor mimicry, it is important to keep in
mind that more research is needed to understand how automatic imitation and motor
mimicry are related (Genschow et al., 2017), and that the results obtained here might but
do not necessarily generalize to the field of motor mimicry.

A second limitation is that most data points in the meta-analysis were obtained from
published studies. As a consequence, it could be argued that our results were at least partly
driven by publication bias. However, it is notable that we also included a substantial number
of unpublished studies. Moreover, there was no evidence for publication bias in the funnel
plots of RT, ER, or KM automatic imitation. This should not be surprising considering that
stimulus-response compatibility effects such as automatic imitation are known to be highly
reliable (Borgmann, Risko, Stolz, & Besner, 2007; Genschow et al., 2017). In addition to the
funnel plots, we also conducted p-curve analyses of three moderators that have mostly
been manipulated within experiments, but these analyses likewise showed no sign of
publication bias (Simonsohn et al., 2014, 2015). Taken together, it thus seems unlikely that
our results can be explained by publication bias.
A final limitation is that not all the effects reported in the meta-analysis can be
interpreted with the same level of confidence. As a rule of thumb, readers should treat the
analyses based on a larger number of studies (e.g., spatial compatibility) with more
confidence than the analyses based on a smaller number of studies (e.g., animacy beliefs).
For example, the fact that we found no significant effect of animacy beliefs on automatic
imitation does not necessarily mean that there is no effect, but could also mean that there
is insufficient data to detect an effect. This interpretation is consistent with the p-curve
analysis, which, due to the low number of studies, was unable to determine whether the
effect of top-down animacy was supported by the data or not. As a result, it will be
important for future research to follow up on the hypotheses that, according to the present
meta-analysis, have been understudied.
In the same vein, further research will be needed to confirm or disprove the effects
of gender overlap and response overlap. That is, neither effect has so far been studied in the
literature. While the present work provides preliminary evidence for their existence, this
evidence should be interpreted with care. For example, the proportion of female
participants was taken as a proxy for participant gender in the gender overlap analysis.
Likewise, the response overlap effect could also reflect weaker automatic imitation in
studies using key release/press responses regardless of the stimulus movement because
there are currently almost no studies with key release/press stimuli. In other words, we urge
not to interpret the data on gender overlap and response overlap as strong evidence for
these effects but rather as a first step towards a consistent research program that
investigates their influence by manipulating both participant gender and model gender, and
by manipulating the correspondence between the stimulus movement and the response
movement.
Conclusion
Despite the more than 200 experiments on automatic imitation, important
theoretical questions regarding its mechanisms and moderators have proven difficult to
answer. In the current meta-analysis, we evaluated seven such questions. The results
revealed that automatic imitation is a flexible, largely automatic process that is driven
mainly by movement and effector compatibility, but is also modulated by spatial
compatibility. They furthermore identified actor-imitator similarity as an important
modulator and showed that automatic imitation is related to neither empathy nor autism.
Overall, this indicates that automatic imitation is best explained in terms of domain-general
mechanisms such as movement, effector, and spatial compatibility (Heyes, 2011) that
develop through associative learning (Brass & Heyes, 2005; R. Cook et al., 2014).
More generally, our results provide important insights on the relation between
perception and action in social settings. In addition to automatic imitation, this has
important implications for research on motor mimicry (Chartrand & Lakin, 2013), mirror
neurons (Rizzolatti & Sinigaglia, 2010), and joint action (Sebanz et al., 2006). The current
work furthermore sheds light on the mechanisms through which imitation develops (Ray &
Heyes, 2011) and can be used to build robot models of human social behavior (Chaminade
& Cheng, 2009). Finally, our findings highlight the need to reconsider current theories on
self-other processing in empathy and autism, as well as in social cognition as a whole (Brass
et al., 2009; Sowden & Shah, 2014; Spengler, Bird, et al., 2010; Williams, 2008; Williams et
al., 2001).
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Footnotes
1
Note that m does not correspond to the number of experiments as stated in figure 2
because between-subject groups were coded as different experiments for the statistical
analysis. The reasoning behind this is that effect sizes corresponding to different between-
subject groups are not statistically dependent.

2
It is noteworthy that RVE models use a simplistic method to estimate heterogeneity.
Consequently, the I² parameter should not be seen as an accurate representation of the
true heterogeneity but rather as a rough indicator (Tanner-Smith et al., 2016).

3
Studies using facial stimuli were coded as controlled in terms of spatial compatibility
because we considered faces too complex to be categorized on a simple spatial dimension.
Nevertheless, it could be argued that some facial expressions do have a clear spatial
component (e.g., smile vs. frown). To address this issue, we also performed the spatial
compatibility analysis without the studies using facial stimuli. In line with the main analysis,
this revealed that automatic imitation was weaker for both RTs, t(222.0) = -4.43, p < .001,
and ERs, t(85.2) = -3.53, p < .001, when simple spatial compatibility was controlled.
4
There is one additional ASD study that was published after data collection had
already been completed (Forbes et al., 2016). Although this study is not included in the
meta-analysis, we did explore whether the results of the ASD analysis changed if this study
was included but this was not the case.

5
The reason why we did not exclude studies on the basis of SOA is that such a
selection would necessarily be arbitrary. That is, it is difficult to determine an objective
criterion to differentiate between SOAs that are acceptable and SOAs that are too long. This
is further complicated by the fact that a cut-off criterion for SOA should take into account
whether the stimulus movement is presented as a video or not because a positive SOA may
coincide with the development of the stimulus movement when it is presented as a video
but always occurs after the stimulus movement if it is presented as a single frame or as a
sequence of two frames. Therefore, we decided to take into account SOA in a sensitivity
analysis rather than using it to exclude studies.

Figures and Tables
Figure 1. Examples of the stimuli used in RT tasks of automatic imitation. In all three panels,
participants have to lift their right index finger when the number 1 is presented and their
right middle finger when the number 2 is presented. The first panel presents a case where
imitative compatibility is confounded with spatial compatibility. The second panel presents
a case where imitative compatibility is orthogonal to spatial compatibility. The third panel
presents a case where the average performance across both trials provides a measure of
imitative compatibility that is independent of spatial compatibility. See text for additional
detail.
Figure 2. The evolution of Google Scholar hits for “automatic imitation”. The y-axis displays
the percentage of hits relative to the total number of entries in Google Scholar for the
respective period. The absolute number of hits is displayed above each bar.
Figure 3. Flowchart of the screening process. The number of studies (n) and experiments (m)
that are included or excluded are denoted. See text for additional detail.
Table 1
Weighted Correlations Among Moderator Variables
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19
1. Spatial Compatibility –
**
2. Compatibility Type .20 –
3. Flexibility .14 .10 –
** *** *
4. Paradigm Type .18 .69 .20 –
***
5. SOA -.06 .20 .07 .02 –
* ***
6. Cue Location .14 .04 .05 .05 -.20 –
* ** ***
7. Cue Modality .11 .09 .05 .11 -.13 .42 –
*** *** ***
8. Action Goal .27 .39 .12 .08 .21 .11 .09 –
*** *** *** *
9. Bottom-Up Animacy .15 .15 .06 .27 .19 .31 .25 .09 –
***
10. Top-Down Animacy .03 .05 .04 .08 .04 .05 .03 .06 .23 –
** *** * *** **
11. Frames .09 .22 .12 .13 .26 .07 .14 .32 .17 .04 –
*** *** ** ***
12. Perspective .27 .33 .13 .20 -.09 .03 .08 .07 .09 .13 .27 –
*
13. Participant Gender .07 .02 .15 .01 .08 .05 .08 .04 .06 .09 .18 .07 –
* *** * ***
14. Model Gender .13 .14 .14 .11 .20 .09 .20 .33 NA .06 .11 .06 .12 –
*** ** ** ** *** **
15. Response Device .13 .22 .07 .05 .01 .06 .15 .22 .19 .03 .11 .31 .09 .23 –
**
16. ASD .03 .06 .07 .01 -.04 .02 .10 .08 .03 .04 .09 .03 .23 .13 .00 –
* ** ** *** *** ***
17. Age .00 -.10 .05 -.04 .07 -.01 -.13 .11 .06 .03 .15 -.03 .34 .08 .18 .27 –
*** * *** *** *** *** *** * *** ** *
18. Trials -.03 .20 .11 .01 .27 -27 -27 .09 .19 .02 .02 -.27 -.12 .25 -.12 -.03 -.09 –
*** *** ** *** ** *** * *** *** *** *** **
19. Effector .32 .67 .22 .36 .17 .20 .16 .58 .17 .03 .37 .46 .05 .16 .43 .12 .23 .17 –
Note. Relations between continuous-continuous variables and between continuous-dichotomous variables were assessed with Pearson’s r.
Relations between continuous-polytomous variables were assessed with multiple R. Relations between dichotomous-dichotomous,
dichotomous-polytomous, and polytomous-polytomous variables were assessed with Cramér’s V. Correlations that could not be calculated are
indicated with NA. We controlled for correlations > .40 (in bold) by analyzing both moderators together in the same model. ASD = autism
spectrum disorder. SOA = stimulus-onset asynchrony. * p < .05, ** p < .01, *** p < .001.
Figure 4. Funnel plot for RTs.

Figure 5. Funnel plot for ERs.

Figure 6. Funnel plot for KMs.

Figure 7. P-curve on the flexibility of RT automatic imitation. See supplementary material 1

for the p-curve disclosure table.
Figure 8. The relation between stimulus-onset asynchrony and automatic imitation. Effect
sizes from the same experiment are represented with the same symbol in the same color.
The fit line of a RVE mixed effects model with stimulus-onset asynchrony as predictor is
shown.
Figure 9. P-curve representing the influence of bottom-up animacy on automatic imitation.
See supplementary material 2 for the p-curve disclosure table.

Figure 10. P-curve representing the influence of top-down animacy on automatic imitation.
See supplementary material 3 for the p-curve disclosure table.

Figure 11. Model gender x participant gender interaction. Values on the y-axis represent the
weighted effect size. The errors bars are standard errors.

Figure 12. Forest plot for the relation between RT automatic imitation and cognitive
empathy. Published studies are annotated with “+” and unpublished studies with “-“.
Genschow et al. (2017) was included in the meta-analysis as an unpublished study but was
published later after data-analysis had already been completed.

Figure 13. Forest plot for the relation between RT automatic imitation and emotional
empathy. Published studies are annotated with “+” and unpublished studies with “-“.
Genschow et al. (2017) was included in the meta-analysis as an unpublished study but was
published later after data-analysis had already been completed.

Table 2
RT Moderator Statistics
Moderator β SE 95% CI m K F/t (df) p

Spatial Compatibility 4.15 (231.0) <.001
Controlled 0.91a 0.05 [0.82, 1.00] 126 258
Confounded 1.26b 0.07 [1.12, 1.40] 116 286
Compatibility Type 6.35 (227.1) <.001
Movement 0.80a 0.05 [0.70, 0.90] 112 353
Effector 1.30b 0.06 [1.18, 1.41] 134 193
Flexibility 2.76 (2, 17.8) .090
Compatible 1.26a 0.16 [0.91, 1.61] 13 16
No 1.09a,b 0.05 [1.00, 1.18] 221 510
Incompatible 0.69b 0.20 [0.26, 1.11] 17 20
Paradigm Type 7.31 (148.6) <.001
Simple 0.69a 0.06 [0.57, 0.80] 78 290
Choice 1.25b 0.05 [1.15, 1.35] 164 256
SOA
Between -0.34 0.05 [-0.44, -0.24] 231 527 -7.44 (13.9) <.001
Within -0.06 0.02 [-0.10, -0.01] 231 523 -2.55 (13.2) .024
Cue Location 3.81 (39.8) <.001
Off 0.74a 0.09 [0.55, 0.93] 31 170
On 1.13b 0.05 [1.04, 1.23] 211 373
Cue Modality 2.32 (5.3) .065
Auditory 0.82a 0.11 [0.55, 1.09] 6 21
Visual 1.08a 0.04 [1.00, 1.17] 236 525
Action Goal 11.19 (2, 33.5) <.001
No Goal 1.15a 0.05 [1.06, 1.25] 198 452
Communicative 0.87b 0.10 [0.66, 1.07] 16 37
Object-Directed 0.70b 0.09 [0.51, 0.88] 30 57
Bottom-up Animacy 26.90 (2, 23.2) <.001
Geometrical 0.40a 0.10 [0.20, 0.60] 25 117
Nonhuman 0.81b 0.10 [0.59, 1.03] 13 18
Human 1.12c 0.04 [1.04, 1.21] 238 409
Top-down Animacy (Nonhuman/Neutral Models) 0.60 (6.8) .566
No 0.56a 0.08 [0.40, 0.71] 38 118
Human 0.78a 0.39 [-0.17, 1.74] 7 18
Frames 3.52 (2, 44.9) .038
One 0.84a 0.10 [0.64, 1.05] 16 37
Two 1.15b 0.06 [1.03, 1.28] 101 166
Multiple 1.05a,b 0.07 [0.93, 1.18] 119 334
Perspective 1.53 (113.3) .130
First 1.00a 0.07 [0.86, 1.14] 68 128
Third 1.13a 0.05 [1.03, 1.24] 169 282
Participant Gender 0.39 (111.1) .695
Male 1.05a 0.08 [0.88, 1.21] 61 151
Moderator β SE 95% CI m K F/t (df) p

Female 1.09a 0.06 [0.97, 1.20] 142 346
Model Gender 6.75 (2, 64.8) .002
Male 0.87a 0.08 [0.71, 1.02] 52 93
Ambiguous 1.11b 0.06 [1.00, 1.22] 139 238
Female 1.43c 0.14 [1.14, 1.72] 25 35
Response Overlap 3.88 (69.9) <.001
No 0.99a 0.07 [0.85, 1.13] 97 287
Yes 1.46b 0.10 [1.26, 1.66] 41 66
Empathy
Cognitive 0.04 0.05 [-0.08, 0.16] 12 15 0.71 (9.6) .497
Emotional 0.05 0.03 [-0.02, 0.12] 12 15 1.49 (9.6) .168
Fantasy 0.05 0.04 [-0.04, 0.13] 11 14 1.15 (8.7) .279
Personal Distress 0.09 0.06 [-0.04, 0.22] 11 14 1.62 (8.5) .141
ASD 0.24 (9.7) .819
Control 1.08a 0.04 [0.99, 1.16] 232 532
ASD 1.11a 0.13 [0.82, 1.40] 10 14
Age 0.13 0.15 [-0.18, 0.44] 198 477 0.85 (34.7) .404
Trials -0.20 0.10 [-0.40, 0.00] 232 525 -2.01 (53.6) .050
Effector 4.00 (3, 13.8) .030
Finger 1.16a 0.06 [1.05, 1.28] 158 380
Face 1.11a,c 0.17 [0.70, 1.53] 7 12
Foot 0.96a,c 0.29 [0.21, 1.71] 6 8
Hand 0.87b,c 0.06 [0.76, 0.98] 72 138
Note. Different subscripts indicate p < .05. For non-continuous variables, the beta
coefficients correspond to Hedge’s g. For empathy, the beta coefficients correspond to
Fisher transformed correlation values. β = beta coefficient; SE = standard error; 95% CI =
95% confidence interval; m = number of experiments; k = number of effect sizes.

Table 3
ER Moderator Statistics
Moderator β SE 95% CI m k F/t (df) p

Spatial Compatibility 3.51 (90.8) <.001
Controlled 0.55a 0.04 [0.46, 0.63] 47 77
Confounded 0.83b 0.07 [0.69, 0.96] 49 65
Compatibility Type 1.73 (31.3) .093
Movement 0.59a 0.05 [0.49, 0.69] 21 41
Effector 0.71a 0.05 [0.61, 0.82] 75 101
SOA
Between -0.15 0.04 [-0.24, -0.06] 92 136 -3.83 (7.0) .007
Within -0.02 0.03 [-0.09, 0.04] 92 136 -0.91 (4.9) .408
Cue Location 1.18 (21.4) .252
Off 0.59a 0.08 [0.42, 0.77] 16 31
On 0.71a 0.05 [0.61, 0.80] 80 109
Action Goal 4.89 (2, 12.4) .027
No Goal 0.73a 0.05 [0.63, 0.82] 80 108
Communicative 0.56a,b 0.06 [0.43, 0.70] 10 23
Object-Directed 0.44b 0.08 [0.25, 0.63] 8 11
Frames 3.54 (2, 16.9) .052
One 0.43a 0.10 [0.18, 0.68] 7 18
Two 0.76b 0.07 [0.63, 0.90] 43 65
Multiple 0.64a,b 0.06 [0.52, 0.75] 43 56
Perspective 1.24 (40.8) .220
First 0.62a 0.06 [0.50, 0.74] 26 48
Third 0.72a 0.05 [0.61, 0.83] 68 82
Participant Gender 2.20 (41.8) .033
Male 0.54a 0.07 [0.40, 0.68] 24 40
Female 0.74b 0.06 [0.62, 0.85] 56 85
Model Gender 3.79 (2, 28.4) .035
Male 0.58a 0.07 [0.43, 0.73] 28 43
Ambiguous 0.75a,b 0.06 [0.62, 0.88] 46 51
Female 0.89b 0.09 [0.69, 1.08] 11 15
Response Overlap 1.77 (43.5) .084
No 0.66a 0.09 [0.49, 0.84] 35 40
Yes 0.86a 0.07 [0.72, 1.01] 22 30
Age -0.43 0.26 [-0.98, 0.12] 77 116 -1.64 (16.1) .121
Trials -0.15 0.09 [-0.33, 0.03] 93 139 -1.70 (22.7) .103
Note. Different subscripts indicate p < .05. For non-continuous variables, the beta
coefficients correspond to Hedge’s g. β = beta coefficient; SE = standard error; 95% CI = 95%
confidence interval; m = number of experiments; k = number of effect sizes.

Table 4
KM Moderator Statistics
Moderator β SE 95% CI M k F/t (df) p

Bottom-up Animacy 7.23 (12.5) .008
Geometrical 0.92a 0.13 [0.64, 1.21] 10 19
Nonhuman 0.24b 0.14 [-0.09, 0.57] 8 13
Human 0.77a 0.12 [0.51, 1.03] 21 36
Motion Profile -0.02 (7.5) .986
Non-biological 0.72a 0.19 [0.23, 1.21] 9 12
Biological 0.72a 0.11 [0.48, 0.95] 23 53
Note. Different subscripts indicate p < .05. Beta coefficients correspond to Hedge’s g. β =
beta coefficient; SE = standard error; 95% CI = 95% confidence interval; m = number of
experiments; k = number of effect sizes.

Appendix A
Description of the Moderator Variables
Variable Description Options

1. Spatial Is spatial compatibility controlled or 1: controlled, 2: confounded
Compatibility confounded with imitative compatibility?
2. Compatibility Does the task manipulate movement 1: movement, 2: effector
Type compatibility or effector compatibility?
3. Flexibility Is there a compatible or incompatible action 1: no or non-contingent, 2: compatible,
context or training phase? 3: incompatible
4. Paradigm Type Do participants perform a simple response 1: simple, 2: choice
task or a forced choice task?
5. SOA What is the SOA between the imperative continuous
cue and the stimulus movement?
6. Cue Location What is the position of the cue with respect 1: on, 2: off
to the stimulus movement?
7. Cue Modality Is the imperative cue visual or auditory? 1: visual, 2: auditory
8. Action Goal Does the stimulus movement have a goal? If 1: no goal, 2: communicative, 3:
so, what kind of goal? object-directed
9. Bottom-Up Is the model human? If not, what type of 1: human, 2: nonhuman, 3:
Animacy nonhuman model is used? geometrical
10. Top-Down Is there a belief manipulation? If so, what 1: no, 2: human, 3: nonhuman
Animacy type of belief manipulation?
11. Motion Profile Is the motion profile of the stimulus 1: biological, 2: non-biological
movement biologically plausible?
12. Frames How many frames are used to display the 1: one, 2: two, 3: multiple
stimulus movement?
13. Perspective From what perspective is the model 1: first, 2: third
observed?
14. Participant What is the percentage of females in the continuous
Gender sample?
15. Model Gender What is the gender of the model? 1: ambiguous, 2: male, 3: female
16. Response Device Does the response device have keys? 1: keys, 2: no keys
17. Disorder Do the participants suffer from a clinical 1: no, 2: ASD, 3: other
disorder?
18. Age What is the mean age of the sample? continuous
19. Trials On how many trials (RT/ER) or seconds (KM) continuous
is the effect size based?
20. Effector What effector do participants use to 1: finger, 2: hand, 3: arm, 4: face, 5:
respond? foot
21. Lesion Do subjects suffer from a virtual or actual 1: no, 2: tdcs anodal, 3: tdcs cathodal,
brain lesion? 4: tms, 5: brain damage
Appendix B
Evidence Table for the Published Studies in the Meta-Analysis
Study Exp N gz Age Sample DV Animacy Motion Belief Persp Goal Paradigm Spatial Type
Brass et al. (2000) 1 8 1.58 Adult Control RT HM BP NB Third GL Choice CFD EFF
Stürmer et al. (2000) 1 10 0.84 Adult Control RT HM BP NB First GL Choice CFD MOV
Brass, Bekkering, et al. 1 8 0.90 Adult Control RT HM BP NB Third GL Simple CFD MOV
(2001)
Brass, Bekkering, et al. 2 10 0.85 Adult Control RT HM BP NB Third GL Simple CFD MOV
(2001) GM NA
Brass, Bekkering, et al. 3 8 1.66 Adult Control RT HM BP NB Third GL Simple IND MOV
(2001)
Brass, Zysset, et al. 1 10 1.72 Adult Control RT HM BP NB Third GL Simple CFD MOV
(2001)
Craighero et al. (2002) 1 12 0.59 Adult Control RT HM BP NB Third OBJ Simple CFD MOV
Craighero et al. (2002) 2 15 1.06 Adult Control RT HM BP NB Third OBJ Simple CFD MOV
Kilner et al. (2003) 1 8 0.83 Adult Control KM HM BP NB Third GL Simple CFD MOV
NHM NBP
Vogt et al. (2003) 1 24 0.71 Adult Control RT HM BP NB First OBJ Simple CFD MOV
Third
Vogt et al. (2003) 2 23 1.07 Adult Control RT HM BP NB First OBJ Simple CFD MOV
Third
Heyes et al. (2005) 1 10 1.41 Adult Control RT HM BP NB First GL Simple ORT MOV
Heyes et al. (2005) 2 20 1.03 Adult Control RT HM BP NB First GL Simple ORT MOV
Oztop et al. (2005) 1 8 0.27 Adult Control KM HM BP NB Third GL Simple CFD MOV
NHM
Press et al. (2005) 1 16 1.10 Adult Control RT HM BP NB First GL Simple ORT MOV
NHM NA
Alegre et al. (2006) 1 7 1.24 Adult Control RT HM BP NB Third GL Choice CFD MOV
Bertenthal et al. (2006) 1 12 1.66 Adult Control RT HM BP NB Third GL Choice CFD EFF
Bertenthal et al. (2006) 3b 12 0.78 Adult Control RT HM BP NB Third GL Choice IND EFF
Jackson et al. (2006) 1 8 -0.51 Adult Control KM HM BP NB Third GL Simple CFD MOV
NHM NA
NHM
Press et al. (2006) 2 24 0.90 Adult Control RT HM BP HB First GL Simple ORT MOV
NHM NHB NA
Aicken et al. (2007) 1 8 1.35 Adult Control RT HM BP NB Third OBJ Choice CFD EFF
Aicken et al. (2007) 2 8 0.44 Adult Control RT HM BP NB Third OBJ Choice CFD EFF
Bouquet et al. (2007) 1 16 1.16 Adult Control KM HM BP NB Third GL Simple CFD MOV
Bouquet et al. (2007) 2 12 0.79 Adult Control KM GM BP NB NA GL Simple CFD MOV
NBP
Bird et al. (2007) 1 32 1.53 Adult Control RT HM BP NB First GL Simple ORT MOV
ASD NHM
Jansson et al. (2007) 1 8 0.87 Adult Control RT HM BP NB First GL Simple CFD MOV
NHM
Jansson et al. (2007) 2 16 0.57 Adult Control RT HM BP NB First GL Simple ORT MOV
GM NA
Jansson et al. (2007) 3 8 1.46 Adult Control KM GM BP NB NA GL Simple CFD MOV
NBP
Kilner et al. (2007) 1 15 0.87 Adult Control KM HM BP NB Third GL Simple CFD MOV
GM NBP NA
NHM NA
Poliakoff et al. (2007) 2 24 0.69 Senior Control RT HM BP NB Third GL Simple CFD MOV
PD GM NA
Stanley et al. (2007) 1 10 1.13 Adult Control KM HM BP NB Third GL Simple CFD MOV
GM NBP NA
Stanley et al. (2007) 2 20 0.81 Adult Control KM HM BP NHB Third GL Simple CFD MOV
GM NBP HB NA
Schroeter et al. (2007) 1 10 2.23 Adult Control RT HM BP NB Third GL Choice CFD EFF
Lesion
Vaino et al. (2007) 1 31 1.01 Adult Control RT HM BP NB First OBJ Choice IND MOV
Biermann-Ruben et al. 1 15 0.57 Adult Control RT HM BP NB Third GL Choice CFD EFF
(2008) GM NA
Biermann-Ruben et al. 2 16 1.94 Adult Control RT HM BP NB Third GL Choice CFD EFF
(2008) GM NA
Gowen et al. (2008) 1 18 0.98 Adult Control KM HM BP NB Third GL Simple CFD MOV
ASD GM NBP NA
Gillmeister et al. (2008) 1 25 0.56 Adult Control RT HM BP NB First GL Choice IND EFF
Third
Gillmeister et al. (2008) 2 32 0.89 Adult Control RT HM BP NB First GL Choice IND EFF
Third
Jarick & Jones (2008) 1 29 1.45 Adult Control RT HM BP NB Third COM Choice IND MOV
Longo et al. (2008) 1 24 0.62 Adult Control RT HM NBP NB Third GL Choice IND EFF
Longo et al. (2008) 2 24 0.45 Adult Control RT HM NBP NB Third GL Choice IND EFF
Liepelt et al. (2008) 1 18 1.84 Adult Control RT HM BP NB First GL Choice CFD EFF
Press et al. (2008) 1 16 0.78 Adult Control RT HM BP NB First GL Simple IND MOV
Van Schie et al. (2008) 1 16 0.70 Adult Control RT HM BP NB Third OBJ Simple CFD MOV
Catmur et al. (2009) 1 8 1.06 Adult Control RT HM BP NB First GL Choice IND EFF
TMS
Catmur et al. (2009) 2 8 1.34 Adult Control RT HM BP NB First GL Choice IND EFF
Chong et al. (2009) 1 16 0.80 Adult Control RT HM BP NB Third OBJ Simple IND MOV
Jarick & Jones (2009) 1 42 0.86 Adult Control RT HM BP NB Third COM Choice IND MOV
Kupferberg et al. (2009) 1 10 1.35 Adult Control KM HM BP NB Third GL Simple CFD MOV
NHM
Longo & Bertenthal 1 120 0.59 Adult Control RT HM BP NB Third GL Choice IND EFF
(2009) NHB
Van Leeuwen et al. 1 48 1.61 Adult Control RT HM BP NB Third GL Choice CFD EFF
(2009)
Poljac et al. (2009) 1 20 1.00 Adult Control RT HM BP NB Third OBJ Choice CFD MOV
Richardson et al. (2009) 1 9 1.24 Adult Control KM HM BP NB Third GL Simple CFD MOV
Spengler et al. (2009) 1 17 1.94 Adult Control RT HM BP NB Third GL Choice CFD EFF
R. Cook et al. (2010) 1 24 2.50 Adult Control RT HM BP NB First GL Simple ORT MOV
Gowen et al. (2010) 1 29 0.44 Adult Control RT HM BP NB Third GL Simple CFD MOV
GM NA
Gowen et al. (2010) 2a 20 0.71 Adult Control RT HM BP NB Third GL Simple CFD MOV
GM NA
Gowen et al. (2010) 2b 20 0.20 Adult Control RT HM BP NB Third GL Simple CFD MOV
GM NA
Jonas et al. (2010) 2 16 1.38 Adult Control RT HM BP NB Third GL Choice CFD EFF
GTS GM NA
Leighton et al. (2010) 1 36 0.83 Adult Control RT HM BP NB First GL Simple ORT MOV
Leighton & Heyes (2010) 1 12 2.15 Adult Control RT HM BP NB First GL Choice CFD MOV
Third IND EFF
Leighton & Heyes (2010) 2 12 1.86 Adult Control RT HM BP NB First GL Choice ORT MOV
Third IND EFF
Leighton & Heyes (2010) 3 12 0.81 Adult Control RT HM BP NB First GL Choice ORT MOV
Third IND EFF
Liepelt & Brass (2010a) 1 20 2.82 Adult Control RT HM NBP NB Third GL Choice CFD MOV
Liepelt & Brass (2010a) 2 25 2.89 Adult Control RT HM NBP NB Third GL Choice CFD EFF
Liepelt & Brass (2010b) 1 19 3.12 Adult Control RT NA BP HB First GL Choice CFD EFF
NHB
Liepelt et al. (2010) 1 27 0.68 Adult Control RT HM BP NB Third GL Choice IND EFF
NHM COM
OBJ
Liepelt et al. (2010) 2 32 0.95 Adult Control RT HM BP NB Third COM Choice IND EFF
NHM OBJ
Marshall et al. (2010) 1 25 0.75 Child Control KM HM BP NB Third OBJ Simple CFD MOV
Newman-Norlund et al. 1 16 0.55 Adult Control RT HM BP NB Third OBJ Choice CFD MOV
(2010) TMS
Ocampo & Kritikos 1 26 -0.16 Adult Control RT HM BP NB Third OBJ Simple CFD MOV
(2010)
Obhi & Hogeveen (2010) 1 16 0.36 Adult Control RT HM BP NB NA OBJ Choice IND MOV
Press, Gherri, et al. 1 12 1.25 Adult Control RT HM BP NB First GL Simple CFD EFF
(2010) ORT MOV
Press, Gherri, et al. 2 12 1.39 Adult Control RT HM BP NB First GL Simple ORT MOV
(2010)
Press, Richardson, et al. 1 28 1.21 Adult Control RT HM BP NB Third COM Simple IND MOV
(2010) ASD
Spengler, Bird, et al. 1 36 1.84 Adult Control RT HM BP NB Third GL Choice CFD EFF
(2010) ASD
Spengler, Brass, et al. 1 20 2.48 Adult Control RT HM BP NB Third GL Choice CFD EFF
(2010)
Spengler, Brass, et al. 2 19 1.58 Adult Control RT HM BP NB Third GL Choice CFD EFF
(2010)
Spengler, von Cramon, et 1 28 2.67 Adult Control RT HM BP NB Third GL Choice CFD EFF
al. (2010) Lesion
Bouquet et al. (2011) 1 24 0.75 Adult Control KM HM BP NB Third OBJ Simple CFD MOV
GL
Catmur & Heyes (2011) 1 16 0.86 Adult Control RT HM BP NB First GL Choice IND EFF
Catmur & Heyes (2011) 2 8 1.56 Adult Control RT HM BP NB First GL Choice IND EFF
Crescentini et al. (2011) 1 19 0.77 Adult Control RT HM BP NB Third GL Simple CFD MOV
GM NA
J. L. Cook & Bird (2011) 1 45 1.02 Adult Control RT HM BP NB Third GL Choice ORT EFF
Child
Grecucci et al. (2011) 1 21 0.87 Adult Control RT HM BP NB Third GL Simple CFD MOV
Grecucci et al. (2011) 2 21 0.75 Adult Control RT HM BP NB Third GL Simple CFD MOV
Hardwick & Edwards 1 8 0.95 Adult Control KM HM BP NB Third OBJ Simple CFD MOV
(2011)
Hogeveen & Obhi (2011) 1 12 1.45 Adult Control RT HM BP NB First OBJ Choice IND MOV
Ocampo et al. (2011) 1 12 -0.04 Adult Control RT HM BP NB Third OBJ Simple CFD MOV
Otte, Habel, et al. (2011) 1 16 1.98 Adult Control RT HM BP NB Third COM Choice IND MOV
Otte, Habel, et al. (2011) 2 16 0.72 Adult Control RT HM BP NB Third COM Simple IND MOV
Otte, Jost, et al. (2011) 1 14 0.62 Adult Control RT HM BP NB Third COM Choice IND MOV
Shen et al. (2011) 1 24 0.57 Adult Control KM NHM NBP NB Third GL Simple CFD MOV
GM NA
Van Elk et al. (2011) 1 19 0.54 Adult Control RT HM BP NB Third OBJ Choice CFD MOV
Wiggett et al. (2011) 1 96 1.25 Adult Control RT HM BP NB First GL Choice IND EFF
Wang et al. (2011) 1 20 1.37 Adult Control RT HM BP NB Third GL Simple ORT MOV
Wang et al. (2011) 2 23 1.09 Adult Control RT HM BP NB Third GL Simple ORT MOV
Boyer et al. (2012) 2 24 0.59 Adult Control RT HM BP NB Third GL Choice IND EFF
J. L. Cook & Bird (2012) 1 41 1.21 Adult Control RT HM BP NB Third GL Choice ORT EFF
ASD
Hardwick et al. (2012) 1 12 0.25 Adult Control KM HM BP NB Third GL Simple IND MOV
Jiménez et al. (2012) 1 18 -0.19 Adult Control RT HM BP NB Third GL Choice IND EFF
Jiménez et al. (2012) 2 17 0.97 Adult Control RT HM BP NB First GL Choice IND EFF
Third
Kupferberg et al. (2012) 1 43 0.67 Adult Control KM HM BP NB Third GL Simple CFD MOV
NHM
Killingsworth (2012) 1 40 0.44 Adult Control RT HM BP NB Third GL Simple IND MOV
Liepelt et al. (2012) 1 32 1.39 Adult Control RT HM BP NB Third GL Choice CFD EFF
Mengotti et al. (2012) 1 22 0.64 Adult Control RT HM BP NB Third GL Choice IND EFF
Romero et al. (2012) 1 12 1.19 Adult Control KM GM NBP NB NA GL Simple CFD MOV
Santiesteban, Banissy, et 1 15 0.97 Adult Control RT HM BP NB Third GL Choice ORT EFF
al. (2012) TDCS
Santiesteban, White, et 1 36 1.78 Adult Control RT HM BP NB Third GL Choice ORT EFF
al. (2012)
Silas et al. (2012) 1 22 1.90 Adult Control RT HM BP NB First OBJ Choice CFD MOV
Vlaskamp & Schübo 1b 8 0.81 Adult Control RT HM BP NB Third OBJ Choice CFD MOV
(2012)
Vlaskamp & Schübo 2 9 0.20 Adult Control RT HM BP NB Third OBJ Choice CFD MOV
(2012)
Badets et al. (2013) 1 36 1.68 Adult Control RT HM BP NB First GL Choice NA MOV
Bortoletto et al. (2013) 1 28 0.70 Adult Control RT HM BP NB First COM Simple IND MOV
Third
Chiavarino et al. (2013) 1 48 0.84 Adult Control RT HM BP NB Third GL Choice IND EFF
OBJ
Chiavarino et al. (2013) 2 40 1.16 Adult Control RT HM BP NB Third GL Choice IND EFF
OBJ
Cross & Iacoboni (2013) 1 24 1.55 Adult Control RT HM BP NB Third GL Choice CFD EFF
GM NA
Cross et al. (2013) 1 20 1.32 Adult Control RT HM BP NB Third GL Simple CFD EFF
GM NA
Fine et al. (2013) 1 8 1.86 Adult Control KM HM BP NB Third GL Simple CFD MOV
Grecucci et al. (2013) 1 30 0.77 Child Control RT HM BP NB Third GL Simple CFD MOV
ASD
Haffey et al. (2013) 1 36 0.81 Adult Control RT HM BP NB First GL Simple ORT MOV
NHM
Hogeveen & Obhi (2013) 1 18 1.69 Adult Control RT HM BP NB Third GL Choice ORT EFF
Mengotti et al. (2013) 1 21 0.85 Adult Control RT HM BP NB Third GL Choice IND EFF
TMS
Shaw et al. (2013) 1 68 1.86 Adult Control RT HM BP NB Third GL Choice ORT MOV
Wiggett et al. (2013) 1 27 1.45 Adult Control RT HM BP NB First GL Choice IND EFF
Wang & Hamilton (2013) 1 19 1.10 Adult Control RT HM BP NB Third GL Choice CFD EFF
Ainley et al. (2014) 1 43 2.41 Adult Control RT HM BP NB Third GL Choice CFD EFF
J. L. Cook et al. (2014) 1 22 0.04 Adult Control KM HM BP NB Third GL Simple CFD MOV
ASD NHM NBP
Cross & Iacoboni (2014a) 1 32 2.47 Adult Control RT HM BP NB Third GL Choice CFD EFF
GM NA
Cross & Iacoboni (2014b) 1 10 2.15 Adult Control RT HM BP NB Third GL Choice CFD MOV
De Coster et al. (2014) 1 24 1.05 Adult Control RT HM BP NB Third GL Choice CFD EFF
Oxytocin
Evans (2014) 1a 20 0.04 Adult Control RT HM BP NB Third GL Simple CFD MOV
GM NA
Evans (2014) 1b 20 0.16 Adult Control RT HM BP NB Third GL Simple CFD MOV
GM NA
Evans (2014) 2 20 0.16 Adult Control RT HM BP NB Third GL Simple CFD MOV
GM NA
Evans (2014) 3 24 0.19 Adult Control RT HM BP NB Third GL Simple CFD MOV
GM NA
Evans (2014) 4 48 0.03 Adult Control RT HM BP NB First GL Simple ORT MOV
GM NA
GM HB NA
Evans (2014) 6 30 0.15 Adult Control RT GM BP NB NA GL Simple ORT MOV
HB
GM HB NA
Hogeveen et al. (2014) 1 16 1.33 Adult Control RT HM BP NB Third GL Choice ORT EFF
TDCS
Klapper et al. (2014) 1 19 2.45 Adult Control RT HM BP HB Third GL Choice CFD EFF
NHM NHB
Obhi et al. (2014) 1 24 3.41 Adult Control RT HM BP NB Third GL Choice CFD EFF
Verrel et al. (2014) 1 16 1.46 Adult Control RT HM BP NB Third GL Choice CFD EFF
Wang & Hamilton (2014) 1 20 1.83 Adult Control RT HM BP NB Third GL Simple ORT MOV
Wang & Hamilton (2014) 2 19 1.13 Adult Control RT HM BP NB Third GL Simple ORT MOV
Butler et al. (2015) 1 230 1.11 Adult Control RT HM BP NB Third GL Choice CFD EFF
Cracco et al. (2015) 1 38 1.29 Adult Control RT HM BP NB Third GL Choice IND EFF
Cracco et al. (2015) 2 40 1.51 Adult Control RT HM BP NB Third GL Choice IND EFF
NHM
de Guzman et al. (2015) 1 25 1.98 Adult Control RT HM BP NB Third GL Choice ORT EFF
de Guzman et al. (2015) 2 38 -0.08 Adult Control RT HM BP NB Third GL Simple ORT EFF
Rauchbauer et al. (2015) 1 27 2.75 Adult Control RT HM BP NB Third GL Choice CFD EFF
Roberts, Bennett, et al. 1 18 0.74 Adult Control KM HM BP NB Third GL Simple CFD MOV
(2015)
Roberts, Hayes, et al. 1 17 0.97 Adult Control KM GM BP NB NA GL Simple CFD MOV
(2015)
Santiesteban, Bird, et al. 1 16 1.47 Adult Control RT HM BP NB Third GL Choice ORT EFF
(2015) MTS
Santiesteban, Banissy, et 1 14 0.44 Adult Control RT HM BP NB Third GL Choice ORT EFF
al. (2015) TDCS
Schunke et al. (2015) 3 40 1.41 Adult Control RT HM BP NB Third GL Choice CFD EFF
ASD GM NA
Sowden & Catmur (2015) 1 16 0.31 Adult Control RT HM BP NB First GL Choice IND EFF
TMS
Sowden et al. (2015) 1 105 1.06 Adult Control RT HM BP NB Third GL Choice ORT EFF
ASD
Sowden et al. (2015) 2 36 0.65 Adult Control RT HM BP NB First GL Choice IND EFF
ASD
Butler et al. (2016)* 1 31 2.45 Adult Control RT HM BP NB Third GL Choice CFD EFF
*
Butler et al. (2016) 4 45 1.66 Adult Control RT HM BP NB Third GL Choice CFD EFF
Catmur (2016) 1 22 0.67 Adult Control RT HM BP NB Third GL Choice CFD EFF
Catmur (2016) 4 71 0.37 Adult Control RT HM BP NB Third GL Choice CFD EFF
Gowen et al. (2016) 1 18 -0.04 Adult Control RT HM BP NB Third GL Simple ORT MOV
GM HB NA
Gowen et al. (2016) 2 19 0.76 Adult Control RT HM BP NB First GL Simple ORT MOV
Rauchbauer et al. (2016)* 1 62 1.82 Adult Control RT HM BP NB Third GL Choice CFD EFF
Verrel et al. (2016) 1 16 1.65 Adult Control RT HM BP NB Third GL Choice CFD EFF
Senior
Deschrijver et al. (2017)* 1 45 1.46 Adult Control RT HM BP NB Third GL Choice CFD EFF
ASD
Genschow et al. (2017)* 1 199 1.37 Adult Control RT HM BP NB Third GL Choice ORT EFF
Cracco et al. (2018)* 1 40 0.99 Adult Control RT HM BP NB First COM Choice IND MOV
Note. The evidence table includes all published research that was used in the meta-analysis. Unpublished studies that were eventually published are also
included and are annotated with *. If a moderator was manipulated in a particular experiment, then all moderator levels that were present in the experiment
are listed. Participant groups that were not included in the meta-analysis are in italics. If this was reported, gz is the effect size corresponding to the main
effect of congruency. Otherwise, it is the mean of all effect sizes included in the meta-analysis. N: number of participants included in the meta-analysis; gz:
overall effect size; NA: not available or not applicable; Child: < 18 years old; Adult: 18-60 years old; Senior: > 60 years old; ASD: autism spectrum disorder; PD:
Parkinson disease; Lesion: brain lesion; GTS: Gilles de la Tourette syndrome; MTS: mirror-touch synaesthesia; tDCS: non-sham tDCS stimulation; TMS: non-
sham TMS stimulation; RT: response time task; KM: kinematics task; HM: human model; NHM: nonhuman model; GM: geometrical model; BP: biological
motion profile; NBP: non-biological motion profile; NB: no belief; HB: human belief; NHB: nonhuman belief; GL: goalless stimulus movement; OBJ: object-
directed stimulus movement; COM: communicative stimulus movement; Choice: forced choice task; Simple: simple response task; CFD: spatial compatibility
is confounded with imitative compatibility; ORTH: spatial compatibility is orthogonal to imitative compatibility; IND: spatial compatibility is independent of
imitative compatibility; EFF: effector compatibility paradigm; MOV: movement compatibility paradigm.
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Automatic Imitation: A Meta-Analysis

Article in Psychological Bulletin · March 2018

Emiel Cracco Lara Bardi

SEE PROFILE SEE PROFILE

Charlotte Desmet Oliver Genschow

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Validation study using fNIRS View project

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Automatic Imitation: A Meta-Analysis

Emiel Cracco, Lara Bardi, and Charlotte Desmet

University of California, San Francisco

Ina Radkova, Eliane Deschrijver, and Marcel Brass

Deschrijver, and Marcel Brass, Department of Experimental Psychology, Ghent University;

Department of Psychiatry, University of California, San Francisco.

This work was supported by the Research Foundation Flanders Grant

insightful comments on a previous draft of this manuscript.

Correspondence concerning this article should be addressed to Emiel Cracco,

Department of Experimental Psychology, Ghent University, Henri Dunantlaan 2, Gent, B-

9000. E-mail: emiel.cracco@ugent.be.

The following manuscript is accepted for publication in Psychological Bulletin. ©American

impeded by incompatible observed movements. In the past fifteen years, automatic

neuroscience. However, important theoretical questions have remained unanswered.

Therefore, in the present meta-analysis, we evaluated seven key questions on automatic

theoretical implications and sheds light on longstanding controversies in the literature on

automatic imitation and related domains.

Public Significance Statement

context of social interaction.

themselves must do if in his situation (Smith, 1759, p. 5)

Why do we imitate others? Is it that we choose to imitate them, or rather that we

Chartrand & van Baaren, 2009; Heyes, 2011).

However, the strongest evidence that imitative tendencies cannot be controlled

As demonstrated in Figure 2, research on automatic imitation has grown

paradigms is rooted in a long tradition of psychological research on stimulus-response

between perception and action in social settings (Heyes, 2011).

Although automatic imitation was first demonstrated in cognitive science, it is now

imitation is widely considered an index of mirror neuron activation (Bien, Roebroeck,

2011), and non-human species (Heyes, 2012).

Following fifteen years of research, it is now well-established that automatic

by incompatible observed movements – exists (Heyes, 2011). However, important

theoretical questions remain. Some questions relate to the mechanisms underlying

automatic imitation: What processes contribute to automatic imitation? Is automatic

literature on how we represent others’ actions.

Although several narrative reviews on automatic imitation already exist (e.g.,

inconsistent evidence has made it difficult to provide conclusive answers to the

analytic approach. A meta-analysis is well-suited to evaluate inconsistent evidence because

meta-analysis allows us to identify important methodological variables that should be taken

questions that are addressed in the meta-analysis.

--- Insert Figure 1 and 2 about here ---

Paradigms of Automatic Imitation

Automatic imitation in the current meta-analysis will be operationalized as “a type of

stimulus-response compatibility effect in which the topographical features of task-irrelevant

investigate the interference of an irrelevant observed movement on a relevant executed

Schulte-Ruther, Konrad, & Koch, 2011; Stürmer et al., 2000).

In the simple response RT paradigms, participants are no longer required to choose a

experimenter performed an incompatible movement than when he performed a compatible

Theories of Automatic Imitation

In contrast to early views assuming an innate imitation mechanism (Meltzoff &