Heyes 2022, What Happend To Myrror Neurons

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research-article2021
PPSXXX10.1177/1745691621990638Heyes, CatmurWhat Happened to Mirror Neurons?
ASSOCIATION FOR
PSYCHOLOGICAL SCIENCE
Perspectives on Psychological Science
What Happened to Mirror Neurons? 2022, Vol. 17(1) 153–168

© The Author(s) 2021
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https://doi.org/10.1177/1745691621990638
DOI: 10.1177/1745691621990638
Cecilia Heyes1,2 and Caroline Catmur3 www.psychologicalscience.org/PPS
1
All Souls College, University of Oxford; 2Department of Experimental Psychology, University of Oxford;
and 3Department of Psychology, Institute of Psychiatry, Psychology, and Neuroscience, King’s College
London
Abstract
Ten years ago, Perspectives in Psychological Science published the Mirror Neuron Forum, in which authors debated the
role of mirror neurons in action understanding, speech, imitation, and autism and asked whether mirror neurons are
acquired through visual-motor learning. Subsequent research on these themes has made significant advances, which
should encourage further, more systematic research. For action understanding, multivoxel pattern analysis, patient
studies, and brain stimulation suggest that mirror-neuron brain areas contribute to low-level processing of observed
actions (e.g., distinguishing types of grip) but not to high-level action interpretation (e.g., inferring actors’ intentions).
In the area of speech perception, although it remains unclear whether mirror neurons play a specific, causal role in
speech perception, there is compelling evidence for the involvement of the motor system in the discrimination of
speech in perceptually noisy conditions. For imitation, there is strong evidence from patient, brain-stimulation, and
brain-imaging studies that mirror-neuron brain areas play a causal role in copying of body movement topography.
In the area of autism, studies using behavioral and neurological measures have tried and failed to find evidence
supporting the “broken-mirror theory” of autism. Furthermore, research on the origin of mirror neurons has confirmed
the importance of domain-general visual-motor associative learning rather than canalized visual-motor learning, or
motor learning alone.
Keywords
action understanding, associative learning, autism, mirror neurons, neuroscience, social cognition
Ten years ago, mirror neurons were everywhere. In scientists are no longer working as actively in this field,
2011, when Perspectives on Psychological Science pub- or that the mirror-neuron “brand” is losing its appeal,
lished a forum focused on the functions and origins of or both and therefore raises the question of what hap-
these fascinating cells (Gallese et al., 2011; Glenberg, pened to mirror neurons.
2011a, 2011b), mirror neurons featured in Time maga- Given the extent of public interest in mirror neurons
zine and The New York Times, programs about mirror and the liveliness of the controversy they provoked
neurons were broadcast by CNN and the BBC, and among scientists and philosophers, this question could
more than 200 articles were published in academic be fruitfully interpreted in an historical and sociological
journals implicating mirror neurons in, among other way. We could ask about the currents in Western society
functions, action understanding, alexithymia, autism, and in contemporary cognitive science that first made
business management, empathy, imitation, language mirror neurons “mesmerising” (Heyes, 2010) and then
comprehension, language production, literary mimesis, weakened their appeal. But this article takes a more
posttraumatic stress disorder, and schizophrenia. Mea- straightforward, natural science approach. After a brief
sured by number of academic publications, interest in introduction to mirror neurons, we use the questions
mirror neurons peaked 2 years later in 2013 and then discussed in the Mirror Neuron Forum (Gallese et al.,
began to decline (Fig. 1). Of course, the numbers in 2011), concerning the functions and origins of mirror
Figure 1 are not an infallible measure of scientific inter-
est in mirror neurons. Since 2013, researchers may have Corresponding Author:
begun to use other terms for the same targets of inves- Cecilia Heyes, All Souls College, University of Oxford
tigation. However, Figure 1 suggests that cognitive E-mail: [email protected]
154 Heyes, Catmur
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Fig. 1. Number of articles published per year from 1996 to 2020 that included the words “mirror
neuron” in the title, abstract, or keywords. Data from Scopus, January 8, 2021.
neurons, to structure a succinct survey of research and execute conditions. For example, they fire during
published in the past 10 years. We then consider the observation of an experimenter placing food in
whether these recent findings have taken the shine off front of the monkey and when the monkey grasps the
mirror neurons and, if so, whether that reaction is food to eat it (di Pellegrino et al., 1992). Strictly and
appropriate. We conclude that although the results of broadly congruent mirror neurons were, from the
careful empirical research were bound to be disap- beginning, of primary interest, and they are what we
pointing relative to the more grandiose claims, recent and most other researchers mean when we use the term
work on mirror neurons should encourage further sys- “mirror neuron.” These cells are intriguing because, like
tematic investigation. a mirror, they match observed and executed actions;
they code both “my action” and “your action.”
Monkey mirror neurons are responsive to the obser-
Mirror Neuron Basics vation and execution of hand and mouth actions. The
Mirror neurons were discovered by chance in monkeys hand actions include grasping, placing, manipulating
in 1992 and given their evocative name 4 years later (di with the fingers, and holding (di Pellegrino et al., 1992;
Pellegrino et al., 1992; Gallese et al., 1996). Early studies Gallese et al., 1996). The mouth actions include inges-
of the field properties of mirror neurons—the sensory tive behaviors such as breaking food items, chewing,
and motoric conditions in which they fire—revealed and sucking and communicative gestures such as lip
three basic types. Strictly congruent mirror neurons dis- smacking, lip protrusion, and tongue protrusion (Ferrari
charge during execution and observation of the same et al., 2003).
action, for example, when the monkey performs a pre- Mirror neurons were originally found using single-
cision grip and when it passively observes a precision cell recording in area F5 of the ventral premotor cortex
grip performed by another agent. Broadly congruent (di Pellegrino et al., 1992; Gallese et al., 1996) and the
mirror neurons are typically active during the execution inferior parietal lobule (Bonini et al., 2010; Fogassi
of one action (e.g., precision grip) and during the et al., 2005) of the monkey brain. Subsequently, they
observation of one or more similar, but not identical, were found not only in these “classical” areas but also
actions (e.g., power grip alone, or precision grip, power in nonclassical areas, including primary motor cortex
grip, and grasping with the mouth). Logically related (Dushanova & Donoghue, 2010; Tkach et al., 2007) and
mirror neurons respond to different actions in observe dorsal premotor cortex (Tkach et al., 2007).
What Happened to Mirror Neurons? 155
Early research with human participants used func- From their discovery in 1992, theorizing about the
tional MRI (fMRI) to show spatial overlap in the areas function of mirror neurons was dominated not by com-
of ventral premotor cortex and inferior parietal lobule putational modeling and experimental intervention but
that are active when people observe and execute move- by consideration of their field properties. Defining
ments (Buccino et al., 2001; Decety et al., 1997; Grezes functions broadly and in everyday language, research-
& Decety, 2001; Rizzolatti et al., 1996). This was not ers reflected on what neurons responsive to similar
conclusive evidence of the existence of human mirror observed and executed actions would be “good for,”
neurons because the spatial overlap could have been what kinds of psychological tasks they might be able
due not to neurons that each respond both to observa- to fulfill. In some cases, this strategy produced hyper-
tion and execution of action (mirror neurons) but to bole. Mirror neurons were hailed as “cells that read
clusters of neurons each responding either to action minds” (Blakesee, 2006), “the neurons that shaped civi-
observation or to action execution (Dinstein, 2008; lization” (Ramachandran, 2009), and a “revolution” in
Dinstein et al., 2007). By 2011, doubts about the pres- understanding social behavior (Iacoboni, 2008). But
ence of mirror neurons in the human brain had been most researchers, including the group in Parma that
assuaged by studies using single-cell recording in pre- discovered mirror neurons, focused on four realistic
surgical patients (Mukamel et al., 2010) and the repeti- possibilities: action understanding, speech perception,
tion suppression fMRI procedure in healthy volunteers imitation, and their potential dysfunction in autism.
(Kilner et al., 2009). Like monkey mirror neurons, evi- Each of these four hypotheses about the function of
dence consistent with the existence of human mirror mirror neurons was debated in the Mirror Neuron
neurons has been found in both classical areas—ventral Forum (Gallese et al., 2011; referred to hereafter as the
premotor cortex and inferior parietal lobule—and non- forum) alongside a key question about their origins:
classical areas, including dorsal premotor cortex, supe- Do mirror neurons get their characteristic visual-motor
rior parietal lobule, cerebellum (Molenberghs et al., matching properties from learning?
2012), supplementary motor area, and medial temporal
lobe (Mukamel et al., 2010).
Research using single-cell recording (and to some Functions
extent, repetition suppression) suggests that mirror
neurons are typically present in adult human brains.
Action understanding
However, this research does not license the inference In his lucid summary analysis of the forum, Glenberg
that mirror neurons are always or usually responsible (2011b) concluded in relation to action understanding
for spatial overlap in fMRI responses during the obser- that there was broad agreement that mirror neurons, or
vation and execution of action. Consequently, it has a mirror-neuron system, “plays some role in action pro-
become common to use terms such as “the mirror- cessing” (p. 408) but no consensus about what that role
neuron system” and “mirror-neuron brain areas” to refer might be. It could be relatively low level; mirror neu-
to regions of the brain that are active during action rons may contribute to action selection or to action
observation and execution and/or for which there is recognition, helping to distinguish one type of action
evidence of the presence of mirror neurons. As review- from another (e.g., precision grip from power grip).
ers, we have adopted the latter of these conventions, Alternatively, mirror neurons may have a high-level
but note that these terms are unsatisfactory in at least function in action processing, enabling “understanding
two respects. First, it is not clear in what sense the areas from within” (Rizzolatti & Sinigaglia, 2010) or inferences
containing mirror neurons constitute a system. Second, about actors’ mental states. Since 2011, advances in
it is likely that only a few of the neurons in each of addressing this issue have come mainly from two broad
these areas have mirror properties. For example, fewer lines of evidence: the use of multivoxel pattern analysis
than 10% of the neurons studied in di Pellegrino et al.’s in fMRI to “decode” the information represented within
(1992) seminal article showed “strict” or “broad” con- and across brain areas and the use of patient studies
gruence in their firing patterns to observed and exe- and neurostimulation to investigate the causal role of
cuted actions, and although some single-unit studies mirror-neuron brain areas for action understanding.
have reported higher proportions of mirror neurons, Multivoxel pattern analysis has revealed that “mirror”
the nature of the single-unit technique makes it prob- areas including premotor cortex encode concrete repre-
lematic to estimate the true prevalence of mirror neu- sentations of observed actions (e.g., the action involved
rons in any brain area (see also Kilner & Lemon, 2013). in opening a particular bottle) rather than abstract,
These issues should be borne in mind when consider- higher level representations (e.g., the goal “to open”;
ing the results of neuroimaging and neurostimulation Wurm & Caramazza, 2019; Wurm & Lingnau, 2015).
studies discussed in this review. These findings are consistent with the involvement of
156 Heyes, Catmur
mirror neurons in lower level processing of observed reduced, suggesting that such responses may improve
actions. speech discrimination in perceptually noisy conditions.
However, any such involvement does not entail that Measurement of motor cortex excitability has also been
mirror neurons play a causal role in action processing. used to investigate the involvement of motor areas in
The evidence in this case is still rather mixed. Studies speech perception. Motor cortical representations of
of individuals born without upper limbs indicate that speech effectors (e.g., lips or tongue) are enhanced
action recognition can take place without motor repre- during the perception of speech in noise (Nuttall et al.,
sentations of the relevant effectors (Vannuscorps & 2016, 2017; but see Panouilleres et al., 2018, for a con-
Caramazza, 2016), and some stroke patients can identify trasting result). This enhancement may have functional
actions despite damage to mirror-neuron brain areas implications for speech perception ability: Participants
(Tarhan et al., 2015), although a meta-analysis of previ- with greater motor mirroring of perceived speech
ous findings indicates that such patients do show showed better ability to discriminate speech in noise
impairments in action identification (Urgesi et al., 2014). (d’Ausilio et al., 2014).
Patient studies can be hard to interpret, however, However, evidence from patient studies casts doubt
because of heterogeneity in the damage incurred and on whether the motor system is causally involved in
the acquisition over time of compensatory strategies. speech perception. If speech perception requires per-
Temporary disruption of brain function using neuro- ceived speech to be matched with motor commands
stimulation can therefore provide convergent evidence for the production of speech, then one should find
that a particular brain area plays a causal role in relation speech perception impairments in patients who have
to a particular cognitive function. One prominent impairments in speech production as a result of brain
neurostimulation study indicated that premotor cortex lesions. In contrast to this prediction, a series of studies
was necessary for identification of the intentions under- has demonstrated intact speech sound discrimination
lying observed actions (Michael et al., 2014). However, in patients with speech production difficulties (Hickok
in that study, premotor cortex stimulation also disrupted et al., 2011; Rogalsky et al., 2011; Stasenko et al., 2015).
perceptual matching of the observed actions. It is pos- A final line of evidence comes from brain-stimulation
sible, therefore, that the disruption to intention identi- studies. Restle and colleagues (2012) demonstrated that
fication was the result of disruption to low-level action facilitatory brain stimulation to the inferior frontal
processing and that premotor cortex does not play a gyrus—a classical mirror-neuron brain area—improved
direct, causal role in intention reading (Catmur, 2014). participants’ accuracy in repeating unfamiliar foreign
On the basis of evidence including that summarized speech sounds. They argued that this indicates the role
above, Thompson and colleagues’ (2019) recent review of the inferior frontal gyrus in matching perceived
of the putative contribution of mirror neurons to action speech to produced speech, but this result cannot reveal
understanding concluded that any involvement of mir- whether the crucial role of this brain area is in speech
ror neurons appears to be confined to lower level pro- perception, speech production, or the matching process
cessing of observed actions (e.g., aiding action itself. Furthermore, because this study did not include
discrimination or recognition). In particular, they found a control task, it is unclear whether stimulation of this
no compelling evidence for the involvement of mirror brain area had a specific effect on speech processing
neurons, or mirror-neuron brain areas, in higher level or whether any complex sensorimotor task might have
processes such as inferring other people’s intentions been improved by such stimulation. However, a series
from their observed actions. of subsequent studies has shown that stimulation of
motor (Rogers et al., 2014; Smalle et al., 2015) or pre-
motor cortex (Nuttall et al., 2018) affects speech per-
Speech perception
ception ability, in particular for distorted speech (Nuttall
Four contributors to the forum (Gallese et al., 2011)— et al., 2018), and one of these studies included a control
Gernsbacher, Gallese, Hickok, and Iacoboni—agreed task, which permits the conclusion that the stimulation
that the motor system has some role in speech percep- had a specific effect on perception of speech but not
tion, but they disagreed about the type and magnitude nonspeech sounds (Rogers et al., 2014).
of the motor system’s role and about whether mirror In summary, there appears to be reasonably strong
neurons in particular are important (Glenberg, 2011b). evidence for the involvement of the motor system
Neuroimaging data indicate that mirror-neuron brain (including premotor mirror-neuron brain areas as well
areas respond during speech perception. For example, as motor cortex) in the discrimination of speech in
Callan and colleagues (2014) demonstrated that responses perceptually noisy conditions. However, this conclusion
in ventral premotor cortex during a vowel-identification is not yet supported by the patient data. A priority for
task were enhanced when the signal-to-noise ratio was future research, therefore, is to test whether patients
with premotor lesions are impaired at discrimination of motor cognition,” rather than mirror neurons, contrib-
speech from noise. utes to the autistic1 phenotype, Iacoboni stood by the
more specific “broken-mirror theory” (Iacoboni &
Dapretto, 2006; Oberman & Ramachandran, 2007;
Imitation Williams et al., 2001; but see Southgate & Hamilton,
Of the functions discussed in the forum, imitation 2008), citing a range of brain-imaging studies suggest-
attracted the strongest consensus. It was agreed that ing that people with autism have abnormal activity in
although early work on the relationship between mirror mirror-neuron areas of the brain. Gernsbacher con-
neurons and imitation had involved some dubious defi- tested all of this evidence, highlighting methodological
nitions and inferences, “when imitation [is] defined in problems and replication failures, whereas Heyes, also
terms of action topography [how body parts move rela- skeptical about the broken-mirror theory, focused on
tive to one another], most agree mirror neurons con- evidence that people with autism show intact (Bird
tribute” (Glenberg, 2011b, p. 409). This consensus was et al., 2007; J. L. Cook & Bird, 2012; Gowen et al., 2008;
due in large measure to two studies showing that repeti- Press et al., 2010) and sometimes exaggerated (Spengler
tive transcranial magnetic stimulation, a disruptive et al., 2010) automatic imitation.
intervention, of the inferior frontal gyrus, a mirror- A systematic review in 2013 of neuroscientific evi-
neuron brain area, selectively impaired imitative behav- dence concluded that there was “little evidence for a
ior (Catmur et al., 2009; Heiser et al., 2003). global dysfunction of the mirror system in autism”
Causal methodologies, including brain-stimulation (Hamilton, 2013, p. 91). Studies published since that
and patient studies, have continued to support the con- review, in which a range of techniques were used to
sensus that mirror-neuron brain areas contribute to investigate neural responses to observed actions and
imitation. Two studies in which facilitatory brain stimu- imitation (the cognitive function thought to rely most
lation to inferior frontal gyrus were used demonstrated strongly on such responses), support this conclusion.
improvements in vocal imitation and naturalistic mim- A popular technique to measure motor system
icry (Hogeveen et al., 2015; Restle et al., 2012), whereas responses during action observation is mu suppression,
inhibitory stimulation to the inferior parietal lobule an electroencephalographic (EEG) measure of the
slowed participants in an instructed imitation task reduction in coherence in certain frequency bands that
(Reader et al., 2018). Inhibitory stimulation of the infe- occurs both when performing and when observing
rior frontal gyrus also disrupted automatic imitation actions. However, the use of this technique to index
(Newman-Norlund et al., 2010), but this effect was not mirror-neuron responses has been criticized on the
specific to human body movements, and similar disrup- grounds that it measures attentional (Hobson & Bishop,
tion was found for nonbiological stimuli. 2016, 2017) and somatosensory (Coll et al., 2015, 2017),
Binder and colleagues (2017) demonstrated that rather than motor, responses. Notwithstanding these
patients with apraxia were impaired on an instructed critiques, recent studies of mu suppression during
imitation task and that this impairment was associated action observation in autism have shown no differ-
with lesions to a set of brain areas thought to contain ences from neurotypical controls (Bernier et al., 2013;
mirror neurons, including the left postcentral gyrus, Ruysschaert et al., 2014).
intraparietal sulcus, and inferior frontal cortex. A similar In contrast, fMRI of neural responses during action
result was reported by Frenkel-Toledo et al. (2016), observation indicates some differences between autistic
who found imitation impairments to be associated with and neurotypical participants. A recent meta-analysis
lesions to the left inferior and superior parietal lobules of six fMRI studies of action observation and imitation
and postcentral gyrus. indicated that participants with autism showed greater
Data from causal studies such as these have been com- responses in bilateral fronto-parietal regions than par-
plemented by a series of fMRI studies over the past ticipants without autism (Yang & Hofmann, 2016),
decade, which demonstrate greater responses in mirror- although another study showed no differences in neural
neuron brain areas during imitation than during other responses during action observation (Pokorny et al.,
closely matched tasks (e.g., Campbell et al., 2018; Mainieri 2015). These data suggest some differences in responses
et al., 2013; Mengotti et al., 2012; Ocampo et al., 2011). during action observation, but they are not consistent
with a broken-mirror account of autism; if anything,
they point to greater neural responses in mirror-neuron
Autism
brain areas during action observation in people with
In marked contrast with imitation, the forum unearthed autism compared with those without autism. Further-
“huge disagreement on autism” (Glenberg, 2011b). more, although these responses are in brain areas
Although Gallese defended the view that “impaired thought to contain mirror neurons, none of these
158 Heyes, Catmur
researchers used techniques that permit the definitive neuron—responsive to both the sight and performance
conclusion that such responses are due to mirror neu- of an action. Iacoboni agreed that visual-motor learning
rons instead of other neurons colocated in these brain is likely to be important but saw signs that it is “cana-
areas. lized” by a genetic predisposition to develop mirror
Techniques that measure motor representations of neurons (Del Giudice et al., 2009). Gallese went further,
specific actions (e.g., grasping) are potentially more arguing that there is an “innate” or “genetically pre-
informative in this respect. Recent studies in which determined” (Gallese et al., 2011, p. 384) propensity to
motor-evoked potentials were used to measure motor develop mirror neurons that is facilitated not primarily
cortical excitability during action observation provide by visual-motor learning but by motor experience
mixed evidence for differences between autistic and before and after birth (Gallese et al., 2009). Summariz-
neurotypical participants. Enticott and colleagues (2012) ing the debate about origins, Glenberg (2011b, p. 409)
reported a smaller increase in motor cortical excitability identified as a key question “the degree to which neo-
in participants with autism compared with neurotypical natal imitation is a reliable phenomenon.”
control participants when viewing hand grasps. How-
ever, a later study found no differences on the same
Sensorimotor learning
measure when observing socially relevant hand actions
(Enticott et al., 2013). Previous claims of disrupted mir- In the past decade, more evidence has emerged that
ror responses in autism based on electromyographic evi- learning plays an important role in the development of
dence (Cattaneo et al., 2007) have also come under mirror neurons (Brunsdon et al., 2020; Catmur et al.,
scrutiny, and a series of methodological critiques has cast 2011; Copete et al., 2016; de Klerk et al., 2015;
doubt on the interpretation of previous data (Pascolo & Fitzgibbon et al., 2016; Furukawa et al., 2017; Guidali
Cattarinussi, 2012; Ruggiero & Catmur, 2018). et al., 2020; Hou et al., 2017; McKyton et al., 2018;
Finally, behavioral measures of imitation have also Orlandi et al., 2017; Press et al., 2012; Wiggett et al.,
provided little evidence for a broken-mirror account of 2012; Zazio et al., 2019). Some of the recent studies
autism. Although one study reported some differences have reported greater activity in mirror-neuron brain
in instructed imitation (Cossu et al., 2012), a task with areas in pianists and dancers than in people who lack
many demands unrelated to mirror neurons, the major- such expertise during observation of musical perfor-
ity of recent studies have found either no difference in mance and dance, respectively (Furukawa et al., 2017;
imitation between autistic and neurotypical participants Hou et al., 2017; Orlandi et al., 2017). These studies are
or greater imitation in people with autism (Gordon of interest because they indicate that activity in mirror-
et al., 2020; Schulte-Ruther et al., 2017; Schunke et al., neuron brain areas is affected by long-term learning
2016; Sowden et al., 2016). Overall, therefore, the past under naturalistic conditions, but they do not indicate
10 years of research have produced no compelling evi- what kind of learning is important. For example, danc-
dence for the claim that autism is associated with ers may show greater mirror-neuron brain area activity
mirror-neuron dysfunction. than control participants during dance observation
because the dancers have watched more dance move-
ments (sensory learning), performed more dance move-
Origins ments (motor learning), and/or watched more dance
Contributors to the forum also debated the origins of movements while performing similar dance movements
mirror neurons, addressing the question of whether (sensorimotor learning).
mirror neurons get their characteristic visual-motor Experiments that were designed to isolate the kind
matching properties from learning. Heyes argued that of learning involved in mirror-neuron development
mirror neurons get their matching properties via stan- have suggested that sensorimotor learning is crucial
dard mechanisms of sensorimotor associative learning. (Catmur et al., 2011; de Klerk et al., 2015; Fitzgibbon
They start out as motor neurons, active only during the et al., 2016; Guidali et al., 2020; Press et al., 2012;
performance of action. Then, through correlated experi- Wiggett et al., 2012). For example, replicating and
ence of seeing and doing the same actions—in the extending earlier work with a similar design (e.g.,
context of self-observation (e.g., an infant watches his Catmur et al., 2007, 2008), Wiggett et al. (2012; see also
or her hand own in motion) and social interactions in Brunsdon et al., 2020) found using fMRI that mirror-
which the same movements are repeatedly both neuron brain areas were more strongly activated by
observed and executed (e.g., pat-a-cake; Heyes, 2001)— observation of hand movement sequences in partici-
these motor neurons become strongly connected to pants who had simultaneously observed and executed
visual neurons tuned to similar actions. Consequently, the movements (sensorimotor learning) than in partici-
what was once a motor neuron becomes a mirror pants who had either observed the movements without
performing them (sensory learning) or performed the The idea that motor learning alone is sufficient to
movements without observing them (motor learning). change the properties of mirror neurons (Gallese et al.,
Furthermore, using paired-pulse transcranial magnetic 2009) has not been supported. The study by de Klerk
stimulation (TMS), Catmur et al. (2011) confirmed that et al. (2015) failed to find a relationship between the
novel sensorimotor experience of the kind given by frequency with which infants performed stepping
Wiggett et al. acts on mirror responses. They showed movements during training and the extent of senso-
that “counter-mirror” training (in which participants rimotor α suppression during observation of stepping
performed index-finger movements while observing movements after training. Yet more striking, using imita-
little-finger movements and vice versa) reversed mirror tion as a behavioral index of mirror-neuron activity,
responses (e.g., resulted in greater activation of an McKyton et al. (2018) found reduced automatic imita-
index-finger muscle during observation of little-finger tion in newly sighted children who suffered from dense
movement than during observation of index-finger bilateral cataracts from early infancy and were surgi-
movement) via the same connections between premo- cally treated only years later. These children, who had
tor and motor cortex that were responsible for mirror been deprived of sensory and sensorimotor experience
effects before training (e.g., greater activation of an of action but not of motor experience, were less inclined
index-finger muscle during observation of index-finger than control children to imitate task-irrelevant hand
movement than little-finger movement). actions.
Responding to a study in which counter-mirror train- In addition to showing that sensorimotor learning
ing yielded later effects on motor excitability than mir- is important for mirror-neuron development, recent
ror training (Barchiesi & Cattaneo, 2013; see also Ubaldi research suggests that in everyday life, much of this
et al., 2015), Cavallo et al. (2014), like Catmur et al. learning occurs in the context of social interactions
(2011), found that mirror responses and counter-mirror between infants and their caregivers. The extent to
responses followed the same time course. Another TMS which mothers imitate infant facial expressions at 2
study found that counter-mirror responses can be months postpartum predicts EEG α suppression at 9
induced by instruction alone (Bardi et al., 2015). How- months during observation of the same facial expres-
ever, it is unlikely that instructional learning, rather than sions (Rayson et al., 2017; see also Markodimitraki &
sensorimotor learning, was responsible for the training Kalpidou, 2019; Murray et al., 2018). Furthermore, this
effects reviewed above because (a) in the study by relationship is action specific. De Klerk et al. (2019)
Bardi et al. (2015), participants were tested immediately found that parental imitation of facial expressions pre-
after instruction, whereas in the earlier studies they dicted infant imitation of facial but not hand move-
were tested 24 hr after both instruction and sensorimo- ments, implying that parental imitation supports
tor training, and (b) training effects have been observed mirror-neuron development through learning of spe-
in uninstructed infants (de Klerk et al., 2015). cific sensorimotor associations (e.g., between the sight
The training studies described above involved adult and performance of mouth opening) rather than by
participants, but there is also now evidence that sen- enhancing attention to body movements or social
sorimotor learning is important in the early develop- motivation.
ment of mirror responses. Using EEG recordings of
sensorimotor α suppression as an index of mirror- Is the sensorimotor learning
neuron activity, de Klerk et al. (2015) found in 7-month-
old infants, who could not yet walk, that mirror
genetically canalized?
responses during observation of stepping movements The importance of sensorimotor learning for the devel-
increased with the amount of sensorimotor experience opment of mirror neurons is now well established, but
they had received in earlier training sessions. Infants questions remain about the character of this learning.
who had frequently seen their own stepping move- The “associative account” maintains that the sensorimo-
ments while performing those movements showed tor learning that builds mirror neurons is of exactly the
greater α suppression than infants who had relatively same kind as the learning that produces Pavlovian and
little correlated experience of seeing and doing the instrumental conditioning; it is a computationally unde-
stepping movements. De Klerk et al. did not find similar manding, domain-general process that forges excitatory
effects of sensory experience (observing stepping) or and inhibitory links between simple event representa-
motor experience (making stepping movements) on α tions (R. Cook et al., 2014; Heyes, 2001; Keysers &
suppression, suggesting that at least in this study, α Perrett, 2004). In contrast, the “canalization account”
suppression indexed the sensorimotor matching func- (Del Giudice et al., 2009; Gallese et al., 2009), sup-
tion of mirror neurons rather than purely attention or ported by Iacoboni and Gallese in the forum, suggests
arousal. that monkeys and humans genetically inherit a specific
160 Heyes, Catmur
propensity to acquire mirror neurons. On this view, the Recent work with human neonates points in the
sensorimotor learning that contributes to mirror-neuron same direction. In a study with unprecedented power,
development is domain specific—it involves computations conducted in Brisbane, Oostenbroek et al. (2016) tested
distinct from those involved in standard conditioning— more than 100 infants longitudinally at 1, 3, 6, and 9
and/or the learning is primed for the development of weeks of age in a cross-target procedure involving a
mirror neurons (given a head start by genetically inher- wide range of targets. They recorded the frequencies
ited behavioral mechanisms). of nine target actions—tongue protrusion, mouth open-
A few studies in the past 10 years have addressed ing, happy expressions, sad expressions, index-finger
the domain generality of the sensorimotor learning protrusion, grasping, MMM sound, EEE sound, and
involved in mirror-neuron development. Consistent tongue click—while infants observed 11 movement
with the associative account, these have indicated that stimuli—an adult performing each of the nine actions
like standard conditioning, mirror-neuron learning and two object movements (spoon protruding through
depends on contingency as well as contiguity (Cooper a tube and box opening). The results of the Brisbane
et al., 2013) and shows a distinctive pattern of contex- study were wholly negative: In no case did the infants
tual modulation (R. Cook et al., 2012). However, as consistently perform a target action more often while
Glenberg (2011b) predicted, the majority of research observing the same action than while observing all of
bearing on the associative and canalization accounts the alternative actions.
has focused on neonatal imitation. Given the evidence Previous failures to find neonatal imitation have been
that mirror neurons contribute to imitation (see above), attributed to methodological factors—for example, the
reliable evidence that newborns can imitate before they use of an inappropriate model, an inadequate response
have had the opportunity for relevant sensorimotor interval, or suboptimal statistical procedures. In a recent
learning would suggest that the development of mirror meta-analysis of neonatal imitation research by the Bris-
neurons is canalized or genetically predetermined. bane group, encompassing 336 effect sizes dating back
A set of 10 studies from one research group, eight to 1977, researchers sought and did not find a modulat-
of them published since the forum, claimed to provide ing influence of 13 methodological factors previously
evidence of imitation in newborn monkeys (Ferrari cited as reasons for replication failure (Davis et al.,
et al., 2006, 2009; Kaburu et al., 2016; Paukner et al., 2021). However, the meta-analysis did find a modulat-
2011, 2014, 2017; Simpson et al., 2013, 2014, 2016; ing effect of “researcher affiliation,” in which a small
Wooddell et al., 2019). These studies did not use the number of laboratories are more likely than others to
“cross-target” procedure, which both enthusiasts and find large positive effects. Furthermore, across the
skeptics have agreed is necessary to detect imitation in whole data set, there was a relationship between stan-
newborns (e.g., Meltzoff, 1996; Meltzoff & Moore, 1977; dard error and effect size indicative of publication bias
Oostenbroek et al., 2016; Ray & Heyes, 2011; Redshaw, (i.e., suggesting that smaller studies have been con-
2019; Whiten, 2002). For example, when testing for ducted, found no evidence of neonatal imitation, and
imitation of tongue protrusion and lip smacking, they not been published).
did not use these behaviors as controls for one another. Reanalyzing the data from the Brisbane study using
Instead of looking for a higher frequency of tongue a more liberal statistical method, Meltzoff et al. (2018;
protrusion than of lip smacking in infants who had just see also Oostenbroek et al., 2018) found evidence of
observed tongue protrusion and a higher frequency of imitation for one of the nine target actions—tongue
lip smacking than of tongue protrusion in infants who protrusion. Although consistent with other reviews and
had just observed lip smacking, they reported, for meta-analyses of neonatal imitation data (e.g., Anisfeld,
example, a higher frequency of tongue protrusion after 1996; Jones, 2006; Ray & Heyes, 2011), this result does
observation of tongue protrusion than after observation not uphold the historical claim that newborns are capa-
of a rotating disk. An effect of this kind could be due ble of voluntary imitation of a range of actions (Meltzoff
not to imitation of tongue protrusion but to a biological, & Moore, 1977; see also Keven & Akins, 2017) or sup-
social stimulus eliciting more behavior of all kinds than port the view that mirror neurons are learned via a
a nonbiological, asocial stimulus. Pointing out this canalized or genetically predetermined process.
problem alongside a number of others (e.g., multiple Rather than supporting canalization, one study of
comparisons without correction), Redshaw (2019) rean- young human infants provided evidence that the devel-
alyzed the data from the full corpus of 10 neonatal opment of imitation (and by inference, mirror neurons)
monkey studies. Applying the cross-target methodol- depends on unspecialized, unconstrained associative
ogy, the reanalysis found no evidence whatever of imi- learning. Reeb-Sutherland et al. (2012) found that associa-
tation in newborn monkeys. tive learning ability at 1-month postpartum, measured
using a delay- eyeblink-conditioning paradigm, pre- unit. From ancient Greece to particle physics, there is
dicted performance on a range of imitation tasks at 9 a long tradition in which atoms of this kind are under-
months of age. stood to be the building blocks of reality. Immersed in
this tradition, people may be captivated by the idea that
simple, tidy mirror neurons explain the distressing com-
Reflection
plexities of the social world—including political strife,
In the past 10 years, there has been significant progress drug addiction, pornography, and responses to media
in resolving the questions debated in the Mirror N euron violence (Bocher et al., 2001; Iacoboni, 2008). Second,
Forum (Gallese et al., 2011). Regarding action under- some descriptions of mirror neurons imply telepathy.
standing, multivoxel pattern analysis, patient studies, If mirror neurons mediate “understanding from within”
and research using TMS now suggest that mirror- (Rizzolatti & Sinigaglia, 2010), a “pre-conceptual and
neuron brain areas contribute to low-level processing pre-linguistic form of understanding,” which can “over-
of observed actions (e.g., distinguishing types of grip) come all linguistic and cultural barriers” (Rizzolatti &
but not directly to high-level action interpretation (e.g., Sinigaglia, 2008; p. xiii), they allow an effortless, word-
inferring actors’ intentions). In terms of speech percep- less form of communication that is a lot like telepathy.
tion, although it remains unclear whether mirror neu- Given ancient links between mirrors, oracles, and divi-
rons play a specific, causal role in speech perception, nation (e.g., Orofino, 1994), even the name mirror neu-
there is now compelling evidence for the involvement rons may pump the intuition (Dennett, 1984) that these
of the motor system (including premotor mirror-neuron cells give direct, transparent access to other minds
brain areas as well as primary motor cortex) in the (Heyes, 2010).
discrimination of speech in perceptually noisy condi- Figure 1 suggests that even if public enthusiasm for
tions. Regarding imitation, building on research pub- mirror neurons has been sustained, scientific interest
lished before 2011, researchers conducting patient, began to decline in 2014. Two high-profile reviews
TMS, and fMRI studies have found strong evidence that were published in that year. One of them, a target
mirror-neuron brain areas play a causal role in behav- article with commentaries in Behavioral and Brain Sci-
ioral copying of body movement topography. Finally, ences, did not contest the existence of mirror neurons,
concerning autism, researchers using behavioral and or that they contribute to social behavior, but marshaled
neurological measures have tried and failed to find evidence that they are forged by sensorimotor associa-
evidence for the broken-mirror theory of autism. tive learning (R. Cook et al., 2014). This evidence chal-
Instead, there are intriguing signs that under some lenged the view that mirror neurons are a biological
conditions, people with autism have stronger mirror adaptation—that they evolved via genetic mechanisms
responses than neurotypical control participants. for action understanding, speech, imitation, or any
Alongside these developments, research on the origin other function. The other review, a book titled The Myth
of mirror neurons has confirmed the importance of of Mirror Neurons, was more skeptical. It argued that
domain-general visual-motor associative learning because mirror neurons are products of associative
rather than canalized visual-motor learning or motor learning, they could not mediate action understanding
learning alone. Specifically, major studies assessing or any other cognitive function (Hickok, 2014). The
“the degree to which neonatal imitation is a reliable impact of these two publications should not be over-
phenomenon” (Glenberg, 2011b) have shown that it is estimated. They distilled and developed insights and
not reliable at all. concerns that had been emerging over the preceding
These findings are disappointing relative to early years. But it is plausible that they precipitated the sub-
newspaper headlines. It turns out that mirror neurons sequent decline of mirror-neuron research.
are not “Cells That Read Minds” (Blakesee, 2006), they Should the decline be resisted, or should it be hoped
do not alone explain “what makes humans social” that the trend apparent in Figure 1 continues until mir-
(Lehrer, 2008), and they have not been able to “do for ror neurons are a thing of the past? We support resis-
psychology what DNA did for biology” (Ramachandran, tance for two reasons. First, even if the mirror-neuron
2009). But it is unlikely that any serious cognitive sci- boom was fueled by less than rational currents of
entist or neuroscientist would be surprised that mirror thought (e.g., atomism and telepathy) inside as well as
neurons have not lived up to these sensational claims. outside science, it would not be rational to allow mirror
The puzzle is why mirror neurons have so inflamed the neurons to go bust. This field has produced substantial
popular imagination. We speculate that two factors are findings, many summarized in this article, which should
important. First is the deep historical pull of atomism. not be dismissed just because mirror neurons are “the
Mirror neurons are small and apparently indivisible; most hyped concept in neuroscience” ( Jarrett, 2012).
they combine sensory and motor properties in a single Second, in our view, much of the skepticism about
162 Heyes, Catmur
mirror neurons is based on a misunderstanding. The sources and developmental timing of the sensorimotor
discovery that mirror neurons are forged by associative experience that builds mirror neurons, how this varies
learning does not imply that they are without function. across cultures, and how this information might be used
It suggests they are by-products with respect to genetic for clinical and educational interventions. More broadly,
evolution, but by-products can be very useful indeed— investigation of how mirror neurons are woven into
in the oil business and in the brain. Consider the area computational-neurological systems could provide
of the left fusiform gyrus that mediates identification valuable clues about how other cognitive systems are
of visual word forms. Literacy emerged late in human assembled through learning. When a function is “hard-
history, only 5,000 to 6,000 years ago, making it clear wired” or “innate,” the construction work is done by
that the visual word form area was not designed by evolution acting on genetic variants and lost in the mists
genetic evolution to enable reading. It is a by-product of time, but when the construction is done by learning,
of genetic adaptation for the discrimination of visually it can be studied in creatures alive today (Heyes, 2018).
complex objects—just as mirror neurons are a by- Turning from targets of study to methods, it is clear
product of genetic adaptation for learning about predic- that causal methodologies, such as TMS, have more
tive relationships—but literacy is hugely important as power to illuminate functions than correlational meth-
a means of relating to and learning from other people; odologies, and if it is concluded that the unitary char-
literacy is a social cognitive function (Heyes, 2018). acter of mirror neurons is of crucial importance (see
Our review of mirror-neuron research in the past 10 below), the use of more single-unit recording would
years and the critiques discussed in this section have be desirable. The potential for such recording would
several implications for future research. In relation to be greatly expanded by the development of rodent
mirror-neuron functions, research on low-level action models using sensorimotor training (Heyes, 2013).
discrimination is more likely to make progress than Thinking more broadly about methodology, research
continuing effort to find a direct role for mirror neurons on mirror neurons would benefit greatly from extension
in high-level action interpretation. Likewise, imitation of a system-level, computational approach of the type
looks more promising than speech perception, and— advanced by Hickok and Poeppel (2015) for language
given the strength of the evidence that mirror neurons and by Rushworth and colleagues (e.g., Apps et al.,
contribute to imitation and the lack of evidence that 2016) for other aspects of social cognition. Predictive
autism is due to a broken mirror—dyspraxias are a coding is one potential example of a system-level com-
more promising target than autism for research with putational approach to investigating mirror-neuron con-
clinical applications. The idea that mirror neurons contributions to social cognition (Kilner et al., 2007; Press
tribute to imitation was dismissed at an early stage on et al., 2011). Ideally, in any such approach, each
the grounds that monkeys, the first species in which hypothesis about function would specify a part in a
mirror neurons were identified, cannot imitate (Rizzolatti psychological process—a process going all the way
& Craighero, 2004). This argument assumed that imita- from peripheral sensory input to overt motor output—
tion involves copying an entirely novel action guided that mirror neurons are thought to fulfill and do this in
by understanding of the model’s intentions, a definition a way that is testable using behavioral and neurophysi-
so rich that it implies imitation is rare even in adult ological methods. The name given to this part is not
humans (Heyes, as cited in Gallese et al., 2011). Defin- important in itself. What is important is that the name
ing imitation in a way that is more common in cognitive does not derive its meaning purely from folk psychol-
science, as copying the topography of body movement, ogy and that the hypothetical function of mirror neu-
the associative account of the origin of mirror neurons rons is distinguished clearly from other components
implies that any animal capable of sensorimotor asso- and from the overall process. For example, movement
ciative learning has the potential to develop mirror discrimination or action recognition (component) needs
neurons and to imitate. Humans are better imitators to be distinguished from mentalizing (whole process).
than other animals, including monkeys, because socio- Likewise, on the neurological side, we need more spe-
cultural experience (e.g., synchronous dance and sport- cific, testable theories of how mirror neurons work with
ing activities, being imitated by others) provides other types of neurons and the kinds of networks in
humans with matching sensorimotor experience for a which they are embedded (including distinguishing
broader range of actions (R. Cook et al., 2014). In ret- whether any differences in function are due to region-
rospect, it seems that the rich definition of imitation or species-specific differences in mirror neurons them-
diverted attention from one of the most important func- selves or the properties of the region or organism in
tions of mirror neurons. which they are located). The term “mirror-neuron sys-
As for future research on the origins of mirror neu- tem” is commonly used, but proponents of this term
rons, there is still much to be discovered about the need to specify the sense in which mirror neurons
constitute a system rather than just cells with similar Funding

properties found in interconnected areas of the brain. This work was supported by All Souls College, Oxford,
At the broadest, conceptual level, we need to think and the Leverhulme Trust (Grant PLP-2015-019 to C.
hard about why, if at all, it is important that single Catmur).
neurons have mirror properties. Setting aside the his-
torical appeal of atomism, does it really matter in the ORCID iD
context of contemporary psychology and neuroscience Cecilia Heyes https://orcid.org/0000-0001-9119-9913
whether individual neurons or small networks of neu-
rons match observed and executed actions? Indeed, in Note
a potentially welcome development, recent single-unit 1. The use of the term autistic is endorsed by many individuals
recording studies indicate a move away from consider- with autism (see Kenny et al., 2016). We therefore use this term
ing the properties of individual neurons and instead as well as “person-first” language (e.g., people with autism) to
focus on population-level encoding. For example, indi- respect the wishes of autistic individuals.
vidual neurons responsive to the observation of others’
manipulative actions (grasping, dragging, etc.) in ante- References
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