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Mirror neuron research: The past and the future

Article  in  Philosophical Transactions of The Royal Society B Biological Sciences · April 2014

DOI: 10.1098/rstb.2013.0169 · Source: PubMed

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Mirror neuron research: the past and the future

Pier Francesco Ferrari and Giacomo Rizzolatti
Phil. Trans. R. Soc. B 2014 369, 20130169, published 28 April 2014

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Mirror neuron research: the past and

the future
Pier Francesco Ferrari1 and Giacomo Rizzolatti1,2
1
Dipartimento di Neuroscience, Università di Parma, 43125 Parma, Italy
2
Centre for Brain and Motor Cognition, Italian Institute of Technology, Via Volturno 39, Parma, Italy
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This special issue is, in large part, based on the presentations given at a workshop
held at the ‘Ettore Majorana’ Centre in Erice (Sicily) to celebrate the discovery of
mirror neurons, 20 years after the first report on their discovery [1]. All articles in
Introduction this issue have been updated and deal with the present state of the art of mirror
research in different fields.
Cite this article: Ferrari PF, Rizzolatti G. 2014 It was certainly difficult to predict from the note published in 1992 how pro-
Mirror neuron research: the past and foundly the discovery of mirror neurons would influence cognitive neuroscience
the future. Phil. Trans. R. Soc. B 369: in the next 20 years. Even more difficult was to forecast that the discovery of
mirror neurons would have an impact on disciplines outside neurosciences,
20130169.
such as psychology, ethology, sociology and philosophy, or that they would
http://dx.doi.org/10.1098/rstb.2013.0169 interest novelists (e.g. The elegance of the hedgehog [2]) and laymen.
Why has the notion of mirror neurons had such a wide impact on cognitive
One contribution of 19 to a Theme Issue disciplines? Two reasons are the most likely. The first is that their discovery put
‘Mirror neurons: fundamental discoveries, the problem of how we understand others at the forefront of neuroscience. The
second is that, by showing that mirror neurons were basically motor neurons,
theoretical perspectives and clinical
they suggested a rather unexpected solution to this problem: the motor
implications’. system is involved in understanding the actions and intentions of others.
This problem of how we understand others is not new in philosophical
Subject Areas: debates. One of the most accepted views is that the capacity to understand
neuroscience, cognition, behaviour the intentions of others is based on the observer’s capacity to infer others’
internal mental states and to ascribe to them a causal role in generating the
observed behaviour [3,4]. This view is in line with the analytical tradition
Keywords: based on the propositional account of others’ minds. An alternative view, put
epigenetics, motor cognition, social forward by phenomenologists, is that we understand others by comparing an
development, understanding others, empathy action done by others with our own behaviour in a similar situation [5]. The dis-
covery of mirror neurons did not disprove the conventional analytical view, but
demonstrated the validity of the phenomenological stance, at least in most
Author for correspondence: everyday life conditions. It is important to stress that action understanding
Pier Francesco Ferrari through the mirror mechanism is a direct activation of motor representation.
e-mail: [email protected] It does not require a cognitive simulation of others’ behaviour as suggested
by simulation theory of action understanding [6].
Interest in the direct understanding of others’ actions increased enormously
when it was found that the mirror mechanism is also present in the emotional
brain centres. Although there is no doubt that one can understand others’
emotions via inferential mental processes (as during the observation of emotions),
there is clear evidence that brain structures involved in the integration and control
of emotions, like the insula and the anterior cingulate, respond both when one
feels an emotion (e.g. pain or disgust) owing to natural stimuli, or when one
observes that emotion in others [7–10]. This mechanism allows a direct
first-person understanding of others’ emotion. ‘Your pain is my pain’.
Some authors previously questioned the presence of mirror neurons in
humans [11]. This is not, however, a current issue, given the overwhelming evi-
dence for the existence of mirror neurons in humans from hundreds of
experiments [12] carried out with a variety of techniques (positron emission
tomography, functional magnetic resonance imaging, transcranial magnetic
stimulation, magnetoencephalography and electroencephalography), as well as
from a few single neuron studies [13]. It is, nevertheless, of some interest to exam-
ine the reasons why this somehow surprising debate took place. In fact,
traditionally, the mechanisms discovered in primates as well as in other species

& 2014 The Author(s) Published by the Royal Society. All rights reserved.
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of animals are considered, at least prima facie, to exist also in order to develop, require genetic instructions as well as 2
humans. Nobody questioned, for example, the existence of specific environmental inputs.

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‘simple’ and ‘complex’ neurons in the visual cortex of One should also consider that neural circuits very rarely
humans, despite the fact that they were first discovered in the evolved for one purpose only. They are often co-opted for
cat and then studied in detail in the monkey [14,15]. other purposes as well; that is, they are exapted to serve
One reason for the opposition of some neuropsychologists additional functions. Thus, the brain networks that are involved
to accept the existence of mirror neurons in humans is probably in sensorimotor transformation can be recruited to support
due to the fact that the mirror mechanism was discovered when additional functions such as action understanding and imitation.
it was not completely clear whether damage to the areas The key role of epigenetic events in driving neuronal devel-
endowed with this mechanism determines deficits in under- opment has recently been emphasized [26]. These epigenetic
standing others’ actions (for evidence for the causative role of events might facilitate and stabilize, in the course of phylogeny,
mirror neurons in action understanding, see [12,16]). Typically, sensorimotor mechanisms involved in the coding of others’

Phil. Trans. R. Soc. B 369: 20130169

the opposite series of events occurs, that is experimental and/ actions (i.e. mirror neurons). The markers of this gene expression
or clinical studies indicate that a certain cortical region is can consistently emerge in each generation, when infants face
involved in a particular function (e.g. occipital cortex in specific and predictable environmental conditions [28], thus
vision) and then single neuron recordings in animals reveal leading to long-lasting changes in the phenotype. In fact, in all
the mechanism underlying this function. individuals of our species, infants experience the vision of
The second reason is that the motor system has been their own hand and benefit from universal forms of face-
traditionally considered not to be involved in cognitive func- to-face interactions with the caregiver. These behavioural
tions [17]. Its role was thought to be limited to the control events affect those brain mechanisms (including the mirror
and production of movements. The discovery of mirror mechanism) that have been referred as to experience-expectant,
neurons as well as of other motor neurons involved in cogni- which actively ‘seek out’ specific environmental inputs. If the
tion, such as neurons coding peripersonal space or neurons elements of this ‘expectable’ environment are either lacking, or
transforming object affordances into potential motor acts else markedly disturbed, then the immature nervous system is
(canonical neurons), radically changed this view. Yet, as likely to be profoundly affected, thus requiring alternative strat-
often occurs in science, there was some resistance to changing egies for rewiring the networks. According to the epigenetic
consolidated points of view. hypothesis, new emergent neuronal responses and the related
An exciting and not fully explored field is the origin of mirror epigenetic mechanisms yielded some advantages for the organ-
neurons and how the development of social-cognitive skills in ism (e.g. faster or more accurate capacity to recognize others’
infancy may rely on the maturation of the motor system. We actions through mapping the actions onto one’s own motor
gave ample space in this special issue to this topic [18–22]. It knowledge), and natural selection would have favoured their
is highly debated whether the mirror system arose as the conse- stabilization and facilitation of expression [26].
quence of association learning or an evolutionary process that An important field for future research studies will be to
endowed a population of neurons with a mechanism necessary assess the sensitivity of mirror neurons to early perturbations
for accomplishing a specific function [20,23]. of the social environment and whether there are critical
According to the associative account, mirror neurons periods necessary for the wiring of the sensorimotor connec-
emerge during development as the result of strengthening sen- tions responsible for mirror neuron construction and
sorimotor connections that originally evolved for visuomotor functioning. Clinical investigations of disturbances of early
control [20]. From this perspective, for example, the motor social interactions, such as in children who experience a
command for reaching/grasping made by the infant is associ- severe lack of early social stimulation or who, for genetic
ated with the sight of his/her own hand. The simultaneous reasons, have deficits in social communication, could rep-
firing of motor and visual neurons can strengthen their connec- resent an interesting challenge to explore the ontogeny of
tions (see also [24,25]). The further step would be to generalize this mechanism and its functional role.
the visual description of the agent’s action to that of others Another important issue that will be fundamental for
and therefore the observation of others’ actions is sufficient to understanding the mirror mechanism more deeply is to eluci-
trigger the observer’s motor neurons. date its neurochemical and molecular bases. This appears
The alternative account maintains that at least part of the very difficult to address with macaques due to the limitation
mirror neuron system is genetically determined and adapted in applying biomolecular techniques in a species with low
to serve one function as the consequence of the natural selec- reproductive efficiency (one infant per year, long gestation
tion process ([20], but also see [26,27]). More recently, models period and delivery interval) and which requires high costs
have been proposed which avoid pitting one view against the in terms of housing and experiments performing. By contrast,
other, thus escaping an old fruitless nature–nurture divide. other animal models, such as marmosets or birds, appear to
Rather, these models integrate learning processes into a be suitable for investigating complex social-cognitive skills
more realistic theoretical framework on how the brain devel- and their neural underpinnings.
ops in the peri- and post-natal period, and on how some Marmosets, for example, have been used with success in
brain mechanisms do not need to be specified by strict gen- biomedical research, not only because of the reduced costs
etic rules. It is known that plastic changes do occur in the involved, but also because of the possibility of using them
brain during ontogeny and involve a complex interaction in neuroscience research both with invasive and non-invasive
between the environment and gene expression in different techniques (to use them in neuroscience research both with
brain networks. Nobody could deny, for example, that the invasive and non-invasive techniques [29 –31]). The results
development of vision and the organization of the visual of these and other studies clearly showed that marmosets dis-
cortex in primates and cats, and of human language and play remarkable similarities with humans in terms of brain
song learning in birds, are examples of adaptations that, in networks, immune response and endocrinological profile.
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The recent studies on transgenic techniques [31] and on the area in speech production and perception [35] and the HVC 3
embryonic stem cells of marmosets indicate that this species in birds. The possibility of investigating the molecular basis

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may be a more suitable model for genome research and bio- of song production and learning in different species of
molecular investigation than the macaque. These new areas birds makes this model extremely promising in investigating
of research, together with the recent discovery of mirror neur- the development and functional role of the mirror mechanism
ons in the ventral premotor cortex of marmosets [32], and its molecular basis.
represent therefore a first critical step in paving the way to Finally, this new information should be extremely useful
new investigations aimed at clarifying the molecular basis not only for a better comprehension of the mirror mechanism
and the evolutionary origins of this mechanism in primates. itself, but also for its practical application in psychiatric and
Studies in songbirds are another example of how mirror neurological disorders. There is indeed evidence, although
neuron research has expanded and stimulated the field. still preliminary, of possible underdevelopment or impairment
Given the complex interactions occurring during develop- of the mirror system in autism [12] and in those psychiatric dis-

Phil. Trans. R. Soc. B 369: 20130169

ment between parents and their offspring which are so orders [36] in which social competence is compromised [12].
critical for song learning, it is not surprising that in song- A better knowledge of the molecular and neurochemical
birds, neurons with mirror properties (audio –motor basis of this mechanism will represent a breakthrough for
mirroring) have been found. The work by Mooney and co- establishing new therapies and interventions based on solid
workers ([33], see also [34]) clearly demonstrates that, in experimental findings.
songbirds, areas involved in song production and learning
(i.e. area HVC) do have neurons with mirror properties for Acknowledgements. We thank the ‘Ettore Majorana’ Centre in Erice for
audio –motor matching. Lesions in the HVC impair song supporting the workshop on mirror neurons, which stimulated
production and song recognition, as well as the capacity to lively scientific exchanges among scholars and young researchers,
learn new contingencies. These authors suggest that audio – and that inspired the realization of this special issue. We also thank
Lynne Murray for her comments on an early draft of this manuscript.
motor mirror neurons in songbirds may be involved in
song learning and communication [34], thus making a strik- Funding statement. This manuscript was supported by NICHD
ing parallel between the functional role of humans’ Broca P01HD064653 grant to P.F.F.

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