Prousali, Evi. 2021.

"Performing Art, Performing Life: A Proposition on the Origins of Art."

The International Journal of Arts Theory and History 16 (2): 17-36.
doi:10.18848/2326-9952/CGP/v16i02/17-36.

The International Journal of Arts Theory and History

Volume 16, Issue 2, 2021, https://artsinsociety.com
© Common Ground Research Networks, Evi Prousali, All Rights Reserved.
Permissions: cgscholar.com/cg_support
ISSN: 2326-9952 (Print), ISSN: 2327-1779 (Online)
https://doi.org/10.18848/2326-9952/CGP/v16i02/17-36 (Article)
 PROUSALI: PERFORMING ART, PERFORMING LIFE

Performing Art, Performing Life:

A Proposition on the Origins of Art
Evi Prousali,1 University of the Peloponnese, Greece

Abstract: How do we experience Art? What does it mean to have an “aesthetic” experience? Philosophers and
psychologists have contemplated on such questions for centuries. Contemporary neuroscientists, from their part, have
posed the question “What happens in the brain when we experience Art?” The use of functional neuroimaging techniques
(fMRI) has revealed a number of significant results among which the most significant are: a) artworks are perceived
through a sensorimotor neural mechanism in the brain, the Mirror Neuron System (MNs); b) art perception is an
“embodied” experience; and c) primary survival neural circuits are involved in aesthetic appreciation. The field of
neuroaesthetics was introduced in the early 1990s when neuroscientists began exploring the neural correlates of aesthetic
experience. An international collaborative platform has since been created which analyzes and studies the emerging
issues at the intersection of cognitive neuroscience and the Arts. In this article, I will present the scientific research data
with relation to art, as well as refer to contemporary evolutionary theories of art and archaeological evidence which
indicates that art originated in the Pleistocene era. Based on these data, I will make a proposition on the origins of art. I
will argue that the “aesthetic modality,” as I call it, is an exaptation of the biological function of the Mirror Neurons’
simulation of the world. Finally, I posit that performing art is a way of performing life.

Keywords: Performing Arts, Mirror Neurons, Embodied Simulation,

Art Perception, Evolutionary Theories of Art, Aesthetics

Introduction

I n the early 1990s, neuroscientists begun investigating the neural mechanisms underpinning
various brain functions. At the same time, the eminent neurobiologist Semir Zeki introduced
the term “neuroaesthetics” (Zeki 1999), in order to associate neuroscientific data with
aesthetic experience. The new field of neuroaesthetics, along with the discovery in the brain of
the Mirror Neuron System (MNs) by neurophysiologist Giacomo Rizzolatti, enhanced the
relations between art and neuroscience (Rizzolatti and Craighero 2004). At the same time, many
neuroscientists addressed the question, “What happens in the brain when we experience Art?”
and employed brain-imaging techniques to investigate aesthetic experience. Specifically,
neuroscientists used artworks as stimuli in their experiments and applied neuroimaging
techniques in order to study the brain’s response during the perception of art.2 In due course, a
large amount of qualitative and quantitative data has been collected leading to the formulation of
a number of key concepts such as: “embodied simulation,” “embodied language,” and
“experimental aesthetics,” which are concerned with understanding the neurophysiologic base of
aesthetic creation and aesthetic experience. Below, I will present the new data which reveal that
aesthetic experience is, primarily, an embodied experience; a fact that revolutionizes the ideas
about the creation and perception of art.

1
Corresponding Author: Evi Prousali, 21 Vassileos Konstantinou & Terzaki, Department of Performing and Digital Arts,
School of Fine Arts, University of the Peloponnese, Nafplion, P.C. 21100, Greece. email: eprousali@uop.gr
2
Techniques that had been used in the research for detecting brain physiology and function, which are also being used
for diagnostic purposes, include EEG (Electroencephalography), MRI (Functional Magnetic Resonance), and PET
(Positron Emission Tomography).
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Neuroaesthetics
Visual Brain and “Beauty”

While investigating the neural underpinnings of vision, neuroscientist Semir Zeki discovered
several functionally-specialized areas of processing, in terms of different attributes of vision.3
The primary visual cortex, which receives the signals from the retina through the thalamus, is the
area known as V1 (visual 1) surrounded by V2. The signals from VI, and V2 project to V4 (color
processing area) and V5 (motion processing), and the processing goes on until it ends to essential
nodes (Figure 1). Zeki also discovered that the visual input, besides being distributed in space, is
also distributed in time, due to an asynchrony in the processing time of the visual attributes. For
example, while viewing a moving object, color is perceived prior to motion by approximately 80
milliseconds (Zeki and Bartels 1998). In short, neuroimaging evidence reveals that, there are
many different visual areas in the brain, each one of which receives visual input in stages; thus,
vision and visual perception is not a momentary perception but a modular and asynchronous
process through which the brain gains knowledge about the world.

Figure 1: Visual Cortex

Source: Muddamsetty, 2014

Zeki also discovered that there is an immanent neural mechanism in the brain which, in cases
of ambiguous images, allows for only one interpretation to be made at any given time, as happens,
for example, with the Cube of Kanizsa or with the painting by Dali (Figures 2 and 3). Zeki also
asserts that, in such cases “it is not ambiguity itself that is aesthetically pleasing…it’s rather the
capacity of multiple experiences, even though we are conscious of only one at any given moment,
that a stimulus can provide” (Zeki 2004, 192). By extension, it seems plausible that this very brain
mechanism is also involved in the interpretation of ambiguous facial expressions, such as for
example with the paintings by Vermeer or Da Vinci (Figures 4 and 5), where multiple
interpretations of the facial expression can be given. The viewer can offer several equally valid
interpretations, but he/she is only conscious of one interpretation at any given moment (Zeki
2001).

3
Semir Zeki had previously conducted research on the visual brain of macaque monkeys with congruent results.
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Figure 2: Cube of Kanizsa Figure 3: Salvador Dali, L’Image disparaît, 1938

Source: oxfordre.com Source: Dali 1998

Figure 4: Girl with a Pearl Earring, Jan Vermeer, 1665 Figure 5: Leonardo Da Vinci, La Gioconda, 1503
Source: Vermeer 1665 Source: Da Vinci 1503

Brain neural limitations can also alter the way we see the world as can be vividly
demonstrated by the cases of artists who have suffered partial or temporary brain damage. The
following examples demonstrate the need to resort to an analysis of brain functions as they can
specify the impact which the latter have on artistic creation and perception. The painter Anton
Räderscheidt suffered a right cerebral stroke due to which he was left with hemiplegia and loss
of the left visual field. Prior to the stroke, the painter had painted a portrait of himself in near-
realistic manner and with great accuracy (Figure 6). The artist’s cerebral damage deprived him of
the ability to perceive the left half of his face, and as a consequence of that, in the immediate post-
stroke period, he painted his portrait by depicting only the right side, as this was the way he
perceived it (Figure 7a). As he was recovering, his painting gradually ameliorated (Figure 7b–
7d). Thus, the brain dysfunction affected the way the artist perceived the world and, consequently,
his artistic creation (Petcu et al. 2016).
Brain damage accounts for the change of style in another painter, Katherine Sherwood, who
also suffered from a left hemispheric stroke and right hemiparesis. Her artistic skill was
subsequently recovered but with a totally different style of painting—more vivid and energetic—
probably resonating the new way she perceived the world (Figures 8 and 9).

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Figure 6: A. Räderscheidt: Painter and Model (pre-stroke) Figure 7a–d: Räderscheidt: Self-portraits (post-stroke)
Source: Petcu et al. 2016 Source: Petcu et al. 2016

Figure. 8: Katherine Sherwood: Voyager’s Constant Figure. 9: Katherine Sherwood: Cajal’s Revenge
(pre-stroke) (post-stroke)
Source: Petcu et al. 2016 Source: Petcu et al. 2016

Thus, neuroaesthetic data indicate that, in order to understand art creation and perception, scholars
should enquire not only into the nature of the signs residing in a painting but also into the
limitations that brain neural mechanisms impose upon what we actually see and how we interpret
it.
Another objective of neuroaesthetic research was to investigate the beholder’s appraisal of
“beauty.” Philosophers and scholars have speculated on the issue of beauty through the ages. The
main question posed is whether beauty resides within the apprehended object or in the perceiving
subject. Plato, whose writings dominated aesthetic theories and discourse for much of the last
2,000 years, believed that beauty has an existence of its own, that is, independent of the subject
apprehending it.4 It is with the publication of Kant’s work on aesthetic judgment that the emphasis
shifted to the perceiver’s interpretation in the search for the attributes of beauty and aesthetic
value ([1790] 1952).5
Neuroscientists Ishizu and Zeki were among the first to investigate the brain’s neural
correlates underpinning the appraisal of beauty in connection with different sources, such as

4
Even for Plato, participation by the individual was critical. His lofty discourses in Phaedrus and The Symposium, which
emphasize beauty as something with an eternal presence outside the individual are, nevertheless, counterbalanced by the
concession in Hippias Major that the beautiful is that which “is pleasing to the eye and ear,” that is, by participation.
5
In The Critique of Judgment, Kant supposed the existence of a sensus communis, that is to say a brain organization that
is similar across individuals and cultures.
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painting and music (Ishizu and Zeki 2013). Certain fMRI experiments, on art perception, which
used a) paintings (Yue, Vessel, and Biederman 2007), b) sculptures (Gallese 2017), and c) music
(Brattico, Brattico and Jacobsen 2009) proved that, when subjects are asked to appraise “beauty,”
one common cortical area is activated, that of the medial Orbitofrontal Corte (mOFC). It is worth
stressing that activation of the mOFC has also been detected with connection to neural circuits
underpinning the mechanisms of reward, pleasure and judgment which constitute principal
survival mechanisms for the human species (Grabenhorst and Rolls 2011). Other experiments
revealed that art appraisal evokes almost the same reward neural circuits as do food and sex, and
that art evaluation is also biased by expertise and personality (Chatterjee 2014). The same cortical
area is involved in decision-making situations which are, in turn, related to the reward neural
system (Ishizu and Zeki 2013). In addition, a quantitative meta-analysis aiming to study the neural
correlates of viewing paintings also revealed that reward processing areas are involved (Vartanian
and Skov 2014). According to current neuroscientific data, the appraisal of “beauty” seems to be
connected with the primary survival neural circuits, mainly the reward neural circuits (Grassi
2020).
This multifaceted and complex neural circuit (reward and decision-making neural systems),
which is involved in artworks’ perception, is not simply a mechanistic process. Rather, it
constitutes, and should be considered as being, the dynamic and flexible primal mode of
perception that our species has attained through evolution in order to gain an understanding of
the challenges of living in the world. The fact that aesthetic perception is founded, though not
exhausted, on this neural structure, should be taken into consideration by modern aesthetic
theories, since it may indicate the important role that “art” has played in human evolution.

Mirror Neurons and Action Understanding

In the last decades, neuroscientists proved that body, brain, and experience, constitute a
multifaceted and diversified unity. A wide range of neuroscientific experiments have
demonstrated the mechanisms of this body-brain system. Neuroimaging techniques have shown
that there is a basic brain mechanism called “mirror mechanism” which underpins the performing
or the perception of an action. For example, every movement or action that our body performs
toward someone or something, or every movement or action that we observe, activates the mirror
neuron mechanism in our brain.
Specifically, when an individual observes another individual performing an action (e.g.,
grasping an object or drinking water etc.), specific neural regions are activated in the spectator’s
brain, as if the spectator was performing the action himself/herself, but without any overt motor
action taking place (Buccino et al. 2001). The observer’s cortical motor network is activated,
though not in all of its components, hence action is not produced but only simulated in the
spectator’s brain. Neuroscientists call this integral brain mechanism the Mirror Neuron System
(MNs) (Rizzolatti and Sinigaglia 2008).6 In addition, it is confirmed that transitive (i.e., goal-
directed) actions and intransitive (i.e., non-goal-directed) actions activate in humans, as well as
in monkeys, the MNs which means that even intentions of actions are simulated in the observer’s
brain (Rizzolatti and Sinigaglia 2010). Another important property of the mirror-neuron system
is that it encodes each of the movements forming an action, that is, the syntax of an action or the
stages of an action and not only the action itself (Gangitano, Mottaghy and Pascual-Leone 2001).
This mirror neuron mechanism is activated by action stimuli ranging from simple finger, to whole
body movements (Springer, Hamilton, and Cross 2012).

6
But what are mirror neurons? Using the fMRI procedure (functional magnetic resonance imaging), in area F5 part of
frontal motor areas, Rizzolatti and his team discovered, first in monkeys, a set of neurons that became active both when
the animal itself executed a motor act (for example, when it grasped food) and when it observed the experimenter doing
it. These neurons were recorded in the cortical convexity of F5 and were named mirror neurons.
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Mirror neurons were first discovered in the premotor cortex of the macaque monkey and,
subsequently, neurons with mirror properties have been found in many brain cortical areas of
monkeys and other species such as birds (Tramacere and Ferrari 2016; Keysers and Gazzola
2009). Research has shown that newborns as young as eighteen hours are capable of reproducing
mouth and face movements displayed by the adult they are facing (Meltzoff and Moore 1997)
and that thirty-six-hour newborns can discriminate and reproduce three different emotional facial
gestures, i.e., happiness, sadness, and surprise (Simpson et al. 2016). The capacity of a young
infant to reproduce mouth and face movements of adults is surely not based on any inferential
process. It means that newborns set into motion, and in the “correct” way, a part of their body
they have no visual access to, but that nevertheless matches an observed behavior. It means that
visual information is transformed into motor information. Also, there is evidence that infants
succeed in ascribing goals by capitalizing on their own motor processes and representations from
very early in life, with the involvement of the Mirror Neuron system (Southgate et al. 2010).
Thus, what is proven so far is that the MNs is instantly and non-consciously activated during
the observation of an action or even when detecting an intention for an action. Thus, the
observation of actions performed by others, or even the detection of an intention in others to act,
activates the mirror neural network in the observer’s brain which creates an internal
(re)presentation by which the observer simulates bodily the performing/intended action
himself/herself.

Empathy and Social Cognition: An “Embodied Simulation”

In our daily life, we constantly observe the actions and behaviors of others and, as demonstrated
above, we interpret them in terms of goals and intentions. But, apart from others’ actions and
intentions, individuals understand emotions as well.
Neurocientific evidence shows that the Mirror Neuron System (MNs) is also involved in the
understanding of another’s mind and emotion (Gallese 2003). Further research also verified that
the recognition of the emotions and the inference about the feelings of others involve the
activation of the sensorimotor mirroring mechanism in the observer’s brain (Moore, Gorodnitsky,
and Pineda 2012). The ability of people to recognize and respond to the emotions of others is
called empathy and is fundamental to our emotional and social lives.
Empathy is connected to core mechanisms associated with affective communication, social
attachment, and parental care. Social neuroscience is the discipline which examines the
neurobiological mechanisms that instantiate empathy (Batson 2009). As far as emotional empathy
is concerned, it has been proved that, apart from the thalamus and the limbic areas which are
involved in the processing of emotions, the mirror neuron system is also involved (Buck, Powers,
Hull 2017). Moreover, empathy in humans is assisted by other domain-general high-level
cognitive abilities, such as executive functions, mentalizing, and language, which expand the
range of behaviors that can be driven by empathy (Decety 2011). In general, researchers of social
cognition argue that “our brains, and those of other primates, appear to have developed a basic
functional mechanism, the mirror mechanism, which gives us an experiential insight into other
minds. This mechanism could provide the first unifying perspective of the neural basis of social
cognition” (Gallese, Keysers, Rizzolatti 2004, 401).
Specifically, neuroscientist Vittorio Gallese argues that the MNs underpins the action and
emotion “embodiment” sited in the brain which allows basic forms of intersubjective
communication and mutual, implicit understanding and emotional exchange. He calls this
functional neural mechanism “embodied simulation” and posits that “in this context, simulation
is conceived of as a non-conscious, pre-reflective functional mechanism of the brain-body system
whose function is to model objects, agents and events” (Gallese 2014, 3-4). It is a corporeal
procedure through which we are engaged with others and connected to the world: “a world
populated by natural objects, man-made objects and other individuals, a world in which most of
the time we feel at home” (Gallese 2017, 44–45).
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Thus, the MNs are an integral part of our cognitive system because its neurofunctional
architecture supports not only action execution and action perception but interpersonal
understanding as well. Far from being a specific characteristic of the premotor system, the MNs
is a basic, multifaceted mechanism of brain functioning which enables us to map the acting and
emotional world in our brain.

Mirror Neurons and Art

The Mirror Neuron System is also activated in an individual’s brain during art perception
(Freedberg and Gallese 2007). Scientific data show that the viewer of a painting also establishes
an embodied relation with the content of the observed artwork. The question, then, is: What
actually happens in the spectator’s brain during art perception?
Neuroimaging results show that perception of artworks depicting scenes of unpleasant
situations, physical exertion or torture, as in the following paintings (Figures 10 and 11),
automatically activates, in the viewer’s brain, the very sensorimotor cortical areas which would
normally be activated if the viewers were subjected to the same conditions themselves (Gallese
2005; Damasio 2003). Moreover, the MNs is also activated when viewers perceive abstract art,
as e.g., with a painting by Franz Kline or Lucio Fontana (Figures 12 and 13), in which cases
the viewer’s brain simulates the creative actions of the artists while painting. The above data
show that, humans perceive actions, emotions or sensations, in both everyday life as well as in
art, through the same neural mechanism, that of the Mirror Neuron System (Sbriscia-Fioretti et
al. 2013).

Figure 10: Michelangelo Buonarotti, Slave called Atlas, Figure 11: Michelangelo Merisi da Caravaggio, Boy
Florence, Academia (ca. 1520–1523) bitten by a lizard (ca. 1593-1594), National Gallery,
Source: Michelangelo.net London
Source: https://www.nationalgallery.org.uk]

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Figure 12: Franz Kline, Painting Number 2, 1954 Figure 13: Lucio Fontana, Concetto Spaziale
Source: moma.org “Atezza” (Spatial Concept “Waiting”), 1960
Source: trouerlereel.wordpress.com

This data led Gallese to extend the “embodied simulation” theory into the field of art,
introducing the term “experimental aesthetics” as the new discipline which deploys
neuroscientific data in order to analyze experiences in the artistic domain: the creation and
perception of art. In “experimental aesthetics,” Gallese emphasizes the inter-corporeality
involved in the perception of art and the social nature of human creative experience (Gallese
2017). Apart from Gallese, other neuroscientists introduce similar terms in order to delineate art
appreciation, such as the term “embodied aesthetics” (Kirsch, Urgesi, and Cross 2016), while
others have also shown that aesthetic experience is grounded in the embodied simulation of
actions, emotions and corporeal sensations represented in artworks (Ticini, Urgesi, and Calvo-
Merino 2015).

Language and Mirror Neurons: “Embodied Language”

Scientific data, so far, have shown that actions depicted in paintings and sculptures are perceived
by viewers through the MNs. Complementary results exist for the actions perceived in the context
of Performing Arts, such as theatre, performance art, cinema etc. (McConachie 2008; Blair 2008).
Since, the Performing Arts involve language it becomes mandatory that we refer to language-
perception, as well.
The available neuroimaging evidence concerning language (De Vega, et al. 2014) seems to
suggest that even “action words” can activate specific sensorimotor areas in the brain. A relatively
recent review concerning linguistic understanding (Buccino et al. 2016) showed that sentences
expressing a transfer action either in a factual sense, e.g. “I give you some pizza,” or in a
metaphorical one, e.g. “I give you my opinion.” activated in the listener’s brain the neural area
correlated with hand motor processing – due to the verb “to give” (Glenberg et al. 2008).
Neuroscientists call this phenomenon “embodied language” and stress the fundamental role
which “embodiment” plays in speaker-listener’s communication. Thus, it seems that even word
articulation can, primarily, provoke an “embodied” re-action in the listener; an activation of the
listener’s sensorimotor neural circuits in order to ascribe meaning. Thus, apart from observed
actions, intentions and emotions, certain classes of words—namely, words describing or implying
motion realistically or metaphorically—are also being interpreted and understood through
sensorimotor processes, mainly on “embodied simulation” processes. In his review article
Buccino asserts that “grounding meaning in experience requires both that the activation of
sensorimotor circuits has a causal role in attributing content to words, and that personal
sensorimotor experiences are already intersubjective to some extent” (Buccino et al. 2016, 77).

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Contemporary neuroscience reveals that what we see or listen to are not only “visual” or
“auditory” information that are cognitively processed in the brain but rather “embodied
simulation” processes that construct a dynamic (re)presentation of the world around us. Thus, a
direct neural link has been located between auditory information, action-perception, and
language, which is further supported by the discovery of the mirror neuron system (Arbib 2013).
As a conclusion, we point out the following:

a) The fact that language –an abstract communicational construct– is also being perceived
and understood in an embodied manner strengthens the argument that meaning is rooted
in bodily experiences and underpinned by brain circuits, rather than being extracted
purely from word content or cultural content.
b) Since the performing arts comprise a world on their own, as each artwork does according
to W. Benjamin, one that includes actions, intentions, emotions and words that are
perceived through the unconscious mechanism of the MNs, it is crucial that we look
more into brain mechanisms in order to elucidate fields of human creativity which for
long have been considered to be purely cultural or/and idiosyncratic.
c) I will elaborate this argument in the last section titled "Performing Art, Performing Life".

Criticism of Mirror Neuron Theory and Neuroaesthetics

It is indisputable that Mirror Neurons are activated in the observer’s brain every time an action is
performed before him/her, whether this is performed in real life or depicted in artworks.
Neuroscientists, in their majority, agree that the activation of MNs underpins action-
understanding; however, a number of neuroscientists remain skeptical about this theory; the latter
claim, in short, that: a) the mirror neurons’ simulation of an action, per se, cannot be the
mechanism for action-understanding, b) action-execution and action-perception co-occur and
become associated via sensory-motor learning in the response pattern of mirror neurons, and c)
mirror neurons are activated after an action is understood by other mechanisms as a means to
make predictions about the future (Hickok 2013). Csibra eloquently expresses their criticism in
the following sentence: Mirror neurons “reflect action understanding rather than contribute to it”
(Csibra 2006, 457). It is an ongoing discussion about the nature of MNs’ involvement in action-
understanding and is regarded as an invitation to deepen and refine its role. Nevertheless, neither
aspect of MNs criticism does affect the purposes of this article.
Scholars of art and cultural studies have also made intense criticism of Neuroaesthetics.
Skeptical scholars tend to doubt and question the progressing integration of multifarious
neuroaesthetic data with humanistic theories accusing neuroscientists either for a lack of sustained
engagement with the theory of art, as well as with art, per se, or for having reductionist tendencies.
Moreover, they argue that the prefix “neuro” is being used as an indisputable “objective” evidence
for otherwise subjective arguments. In that sense, art theorists and scholars of cultural studies
claim that they are “forced” to defer to neuroscience. Since neuroaesthetics is based on
neuroimaging data, another critical argument is that neuroimaging technologies are not equipped
to answer complex questions about behavior, emotion, artistic taste or art perception (Mondloch
2016). In general, philosophers of art are skeptical about “whether neuroaesthetics has anything
significant to offer to theories of art or aesthetic experience” (Holt 2013, 2).

Toward an Applicable Interdisciplinary Model

Notwithstanding the above skepticism, a number of new transdisciplinary fields have emerged:
neuroarthistory, neuroanthropology, neuroarcheology, and neuroaesthetics. Working at the
intersections of neuroscience, the humanities and various areas of social practice is nothing but
straightforward. For example, long-lasting dichotomies, such as body/brain or nature/culture have
often been criticized by scientists on conceptual grounds. Nonetheless, a successful bridging cannot
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occur without adherence to the premise that the findings of one discipline must be evaluated with
relation to the concepts of the others. On this ground, scholars should provide concrete suggestions
on how integrative experimental work, in the practical sense, could clarify theoretical concepts. In
this way, the interplay between a branch of scientists (such as, neuroscientists, cognitive scientists,
etc.), theorists (such as, art theorists, philosophers of art, etc.) and artists could productively lead to
comprehensive approaches for defining artistic behaviors.
Whenever this has been the case, the “neuroscientific turn” seems to be fruitful and
generative, so far. An interdisciplinary platform has been formed by both scholars and scientists
who are interested in studying the intersection of art and neuroscience, expecting that such
collaboration would be mutually beneficial. Without overlooking their theoretical discrepancies,
they focus on applying new research methodologies and fresh transversal approaches in order to
elucidate artistic creation and perception. In this context, art, art-history and theory, and
neuroscience are equal partners towards advancing our understanding of the human nature itself
(Leder 2014). Early enough, neuroscientist Ramachandran, foreseeing the future, overall
concluded that “mirror neurons will do for psychology what DNA did for biology: they will
provide a unifying framework and help to explain a host of mental abilities that have hitherto
remained mysterious and inaccessible to experiments” (2000, 1).

Evolutionary Theories of Art

Since, both, everyday perception and art perception are, primarily, embodied processes based on
the Mirror Neuron system—a fundamental mechanism of brain function—we move on to the
most crucial question: “Why do we create art?”
Taking into account that: a) the MNs is also found in the chimpanzee brain, b) the fact that
art and aesthetic experience are spontaneous phenomena present in all human cultures and, c)
“art” originated in the earliest times of humanity, tracing back to homo erectus or homo ergaster
as archaeological records indicate (Bednarik 1998), the artistic phenomenon may well comprise
“a behavioural complex, an inherited tendency to act in a certain way” as Dissanayake states
(Dissanayake 1992, 224). Thus, we are led to assume that art should somehow contribute to the
vital everyday functions of human survival, meaning that art is not apart from nature but a part
of nature. It is plausible that the biological origins of art play a significant role in defining the
aesthetic phenomenon as a whole. Under this perspective, it seems crucial for the theory of
aesthetics to shift its focus from the long-standing philosophical question of “what is art?” to a
more palpable one, such as “what people do when they are engaged with art?”
The above chain of inquiry forms an interdisciplinary basis which integrates neuroaesthetics,
aesthetic theories and evolutionary theories of art, in order to study the aesthetic phenomenon.
Human artistic history goes back more than 300,000 years when people painted cave walls and
created human-like figures, animals, palms etc. Art is a worldwide phenomenon among nations,
ethnicities and cultures. But why and how did art become so universal? Is art rooted in culture,
learned and passed down from generation to generation or is it rooted in genetic adaptation,
providing us with an evolutionary advantage over our ancestral competitors?
The discovery of various artifacts coming from the earliest phase in the history of our species
(Figures 14 and 15),7 presupposes that an artistic behavior had developed early in the evolutionary
history of mankind (Bednarik 2014). This assumption has given rise to several evolutionary
theories of art concerning the creation as well as the perception of art (Boyd 2005). The debate
as to whether the arts are biologically adaptive, culturally derived or both, has intensified. Various
approaches to the evolutionary theory of art have appeared, and at this point we shall briefly refer
to the most significant ones.

7
The cobble was found in the cave of Makapansgat in South Africa and it was described as a manuport of 2.5 to 3 million
years ago. The archaeologists concur that the object was collected by australopithecines for its visual qualities, and that
its iconographic properties were recognized by these creatures.
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Figure 14: The Makapansgat cobble, South Africa, a Figure 15: The Tan-Tan proto-figurine, Morocco,
natural object that was carried to a cave and deposited modified to emphasize its human form during the
almost 3 million years ago. middle Acheulian
Source: Bednarik 2013 Source: Bednarik 2013

The substantive intellectual connections between aesthetics and evolutionary biology date
back at least to Darwin’s Descent of Man and Selection in Relation to Sex (1871), in which he
elaborated the theory of sexual selection by mate choice. Darwin hypothesized that a female’s
“taste for the beautiful”—metaphorically, the “aesthetic faculty”—has evolved during mate
choice and constitutes a distinct evolutionary force that leads to the evolution of ornamental traits
in animals; this was the sexual selection hypothesis of the evolutionary theory of art. Darwin,
using aesthetic language, described that these “aesthetic” choices were based on the subjective
sensory and cognitive experiences and could have led to the evolution of arbitrarily attractive
traits that are unrelated to any naturally selected advantage to the individual making the choice
(Cronin 1991). The “sexual selection hypothesis”, as an evolutionary theory of art, was largely
abandoned for more than a century, but nowadays it is brought to the fore again by several
scholars and scientists.
Along the same lines, a contemporary evolutionary theory of art is the “coevolutionary
aesthetic” theory which provides evidence from a heuristic account of aesthetic taste in both
human and non-human biotic artworlds. This theory proposes that the restructure of aesthetics
without placing human beings at the organizing center will stimulate new progress in our
understanding of art. According to R. Prum many biotic/animal advertisements and behaviors are
forms of art, and the inclusion of these artifacts would provide “an intellectual framework for
understanding the nature of the aesthetic processes that give rise to human art forms” (Prum 2013,
830).
Ellen Dissanayake also goes back to about 1.7 million years, in order to refer to the mother-
infant bonding as the primitive reservoir of affective mechanisms from which a primitive form of
art could have arisen. The mother-infant interaction is adaptive during human evolution and is
universally acquainted. During mother-infant interaction certain affect-laden elements of
maternal communication appear -such as vocalizations, gestures, movements etc.—that can be
called “proto-aesthetic.” These “proto-aesthetic” behaviors, on the one hand, strengthen their
bond, giving healthier and/or happier children, and on the other, ancestral humans subsequently
use them as means to gain the same advantages. In that way, “artification” emerges. Dissanayake
accepts that some animals perform art-like behaviors but she claims that “only humans
deliberately use these operations to make their bodies, movements and surroundings
extraordinary or special” (2009, 165). Dissanayake introduces “aesthetic primitives” and
describes that the somatic and behavioral modalities can add further to “aesthetic” fundamentals
inherent in early Homo sapiens. She finally argues that these “universal aesthetic primitives,
along with predisposed aesthetic preferences, contribute to establishing an evolved, adaptive,

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‘bottom-up’ or ‘naturalistic’ aesthetics and demonstrate that, as we became Homo sapiens, we

were at the same time Homo Aestheticus” (Dissanayake 2015, 19).
Brian Boyd develops his own evolutionary account of art arguing that art is an adaptation,
inherent in the biology of the human species, which derived, in the phylogenetic sense, from
adaptive animal play behavior. Boyd claims that art has retained characteristics of nonhuman
play, such as the practice of actions in the secure environment of a community, or the repeated
engaging in play which aids the development of useful skills and relevant sensitivities crucial for
creating adaptive behaviors within the group; something that leads to strengthened synaptic
connections on the neurological level (Boyd 2009).
Jan Verpooten argues for quite the opposite: the arts have evolved culturally rather than
biologically, exploiting biological adaptations rather than extending them. Verpooten does not
reject the evolutionary parameter; rather he confines it in the survival advantage gained by the
“artist” as being the most preferable male/female of the group. In specific, “the arts can be defined
as arising from the interactions between cultural evolution, involving the capacities to learn from
others, and biological evolution, i.e., the capacity for a cognitive stance that accords significance
to an object whether this is expressed explicitly in an artificially contrived artefact or activity, or
implied in a mundane item within a ritualistic or animistic context” (Hodgson and Verpooten
2014, 76). In that sense, he proposes that most of the arts evolved culturally, building on pre-
existing biological traits.
Anjan Chatterjee, in his book The Aesthetic Brain argues that it does not make sense to see
art as purely genetic or cultural (Chaterjee 2014). He uses the brilliant metaphor of the Bengalese
finch’s song to illustrate a more nuanced explanation for the origin of art. According to Chatterjee,
art is like the Bengalese finch’s song. While there is little question of art’s adaptive roots, art no
longer holds an adaptive purpose. Like the Bengalese finch’s singing ability, the selective
pressures on our artistic abilities have relaxed, enabling us to diversify our artistic behaviors for
nearly infinite purposes. “Art can be both the expression of an instinct and the relaxation from
this instinct” (Chaterjee 2014, 179).
Mark Changizi argues that “the arts have been culturally selected over time to be a ‘good fit’
for our brain, and our brain has been naturally selected over time to be a good fit to nature …so,
perhaps the arts have come to be shaped like nature, exactly the shape our brain came to be highly
efficient at processing,” to conclude with the statement that “with the brain put on the shelf, the
goal is, instead, to analyze nature” (Changizi 2010). In fact, he searches for regularities in nature
because he assumes that our brains evolved to fit these regularities (Changizi 2011). As a
consequence, hundreds of articles and many books, representative of the ongoing inquiry on the
biological origin of art, have been written, pointing towards an interdisciplinary perspective that
will bring together evolutionary aesthetics and aesthetic theories, in order to study the aesthetic
phenomenon as a whole.

A Proposition on the Origins of Art

The definition of Art is undoubtedly controversial. For centuries, various cultural and social
constructions and processes have been proposed and prevailed as constituting art. But if we want
to unravel the essence and the origin of art, it would be beneficial to follow Dewey’s proposal,
“In order to understand the meaning of artistic products, we have to forget them for a time, to turn
aside from them and have recourse to the ordinary forces and conditions of experience that we do
not usually regard as aesthetic”8 (Dewey 1934, 4).
Approaching art under this perspective entails a synergy and cross-fertilization of disciplines
that, until recently, were considered independently, such as, evolutionary biology, neuroscience
and archaeology. In specific, old questions and definitions can be addressed and answered
through recent developmental studies. In particular, we should resort to: a) evolution, adaptation
8
Emphasis is mine
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and natural selection, as the process through which the human species evolved; b) the hunter-
gatherer’s society, where the artistic behavior first appeared; and c) the neuroscientific data
concerning artistic behavior. Multifarious evolutionary approaches to art have been proposed in
consistency with this demand. The fact that, so far, no brain mechanism has been discovered
exclusively connected with art, as for example, has been the case with language, does not rule out
other possibilities.
In this article, I combine the above neuroscientific data with the evolutionary perspective
concerning the genesis and development of art, along with the archeological findings, in order to
propose a new hypothesis for the origins and the evolutionary advantage the artistic phenomenon
provides to human species. Therefore, I argue that the artistic phenomenon may well have
appeared as an exaptation (Gould and Vrba 1982) of the Mirror Neuron System and become
beneficially adapted by way of its utility as an “experimental laboratory” of life where multiple
artworlds are (re)presented and tested for their efficacy.
Thus, according to my proposition, in the course of time the MNs came to function
bidirectionally: besides mirroring external actions through embodied simulation in the
individual’s brain, it came to serve the mirroring mechanism in an inverse way by projecting the
individual’s internal state concerning actions in the world through creative acts. Βy consequence,
the individual’s thoughts about actions and/or intentions in the world are being embodied in an
artistic (re)presentational framework. If such a proposition is founded, humans became capable
of externalizing and expressing their internal thoughts about actions and intentions in the form of
(re)presentational acts through the reverse function of the Mirror Neuron System. Τhe world
presents itself to the individual via the mechanism of MNs-embodied simulation, while the
individual (re)presents himself/herself to and in the world via the inverse function of the MNs,
leading to the artistic creation.
The above proposition is sustained by the scientific data which show that the MNs unifies
the “execution and perception of an action, with a set of neurons, ranging from premotor and
supplementary motor areas to primary somatosensory and inferior parietal cortices, coding for a
precise action and activated also in the observers’ motor system” and that the “MNs is crucial for
the study of the self” and self-awareness” (Sandrone 2013, 1). In addition, it has been shown that,
in order for the individual to present itself in the world, the imagery and imagination mechanisms
should be employed. Scientists, defining imagery as the production of mental images associated
with previous percepts and imagination as the faculty of forming mental images of a novel
character or situation, have found that the MNs is also involved in imagery and in imagination.
In particular, they argue that “the mirror neurons could be part of a broader system, the imagery
neuronal system, existing in the human brain” (Agnati et al. 2013, 6). But imagery and
imagination are constitutive parameters of the artistic phenomenon.
In this case, the internal thoughts of prehistoric humans concerning their lives and their
existence in the world were mirrored through the MNs into artistic (re)presentations. These
internal thoughts, primitive as though they might have been, derived from experiences or
knowledge gained from their living in the world. Embodied simulation, now unsuppressed by the
brain mechanisms, was externalized and “realized” as (re)presentations in, of, and on the world.
These (re)presentations led to creations such as the well-known cobble conformations and, in due
course, cave images which in modern terms are usually referred to as “art.” But Tim Ingold wrote
“Hunters and gatherers of the past were painting and carving, but they were not ‘producing
art.’…We must cease thinking of painting and carving as modalities of the production of art, and
view art instead as one rather peculiar, and historically very specific objectification of the
activities of painting and carving” (Ingold 2000, 131).
It is my view that for the primitive hunter-gatherers these creations denote a concern for
communicating the precariousness of everyday life. In fact, these (re)presentations depicted
different states of relations that the hunter-gatherers had in and with the world, and various
potential (re)actions as if the “aesthetic” practice was an “experimental laboratory” where

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multiple human “performances” could be tested. Art, in that way, functions as a “trial and error”
practice where predictions or imaginary human states and situations can be manifested and
communicated among the group. In fact, with these (re)presentations humans “mirror”
themselves—both, presently as well as forward in time—in alternative world-performances, in
the same way that world-performances “mirror” themselves in their brain through the MN system.
It is in such terms that I describe the “aesthetic modality” of the MNs.
According to my argument, therefore, the “aesthetic modality” is the exaptation of the reverse
biological function of the Mirror Neurons’ simulation of actions in the world. The present
proposition, that art emerged as an exaptation through a reverse function of the MNs, is not to be
found in any bibliography to date, though similar approaches have been expressed.9 For example,
J. Onians argues that the MNs is the neural mechanism underpinning the arts, basing his argument
on the mirror neurons’ “mimetic” force (Onians 2007), whereas Tooby and Cosmides argue that
the artistic faculty provides humans with the ability to construct imagined worlds where learning
can take place through vicarious experiences (Tooby and Cosmides 2001).
According to my proposition, the evolutionary advantage of the acquisition of this “aesthetic
modality” is that it allows the individual to “experiment” under the conditions of, what I call, a
“(re)presentational laboratory,” in order to become informed and purposeful in its estimation of the
prospects and the planning-out of actions in similar future situations. In this context, the prefix “re”,
in artistic (re)presentation, does not signify repetition, which refers to the past, but (re)creation,
which refers to the future. In this sense, primitive art, performing arts, as well as art in general, aim
dynamically and purposefully towards the future, investigating the possibilities for alternative ways
of living and existing, rendering the arts ontologically performative.
To some evolutionists, the artistic behaviors seem costly and highly-energetic activities to
the human species because their ultimate benefits are not immediately apparent. The experiential
and cognitive “preparedness” which is acquired through the development of the aesthetic
modality, equips the human being with higher prospects of survival and enhanced adaptability,
as it helps him/her comprehend the situation at present, communicate it in the group, as well as
to anticipate their place in similar future situations with better efficiency and flexibility. Thus, the
aesthetic modality endowed humans with the advantage of knowledge gained by “experimenting”
in real life under the (re)presentational conditions of aesthetic performance. These factors
propelled the survival of societies that had developed artistic faculties.
To primitive communities, engagement in artistic practices promoted social cohesion since it
referred to everyday activity commonly shared by everyone. In that way, artistic behavior enhanced
human communication and cooperation and, due to its potential efficacy, it was passed on through
social learning. I, thus, argue that, in the course of time, artistic practices were also culturally
transmitted, adopted and evolved. The enforcement of cultural activity, which was induced by the
artistic practices, modified, in turn, “the chemistry and structure of the humans’ brain through
affecting the flow of neurotransmitters and hormones” (Bednarik 2014, 51). The use and
proliferation of the products of artistic practices must have profoundly altered the hominid’s brain.
In that sense the artistic trait is both an exaptation and a culturally acquired behavior.

9
To elucidate my proposition further, I am presently planning a number of experiments.
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Performing Art, Performing Life

A great number of manuports were found in caves of the Pleistocene era (2.5 to 3 million years
BT). Among them there were cobbles which, in a rough way, resemble hominid faces or figures
(Figures 14 and 15). What justification can be given to a “costly” transportation of such items if
they were not useful in some way to their carrier and beholder? I argue that such cobbles instigated
an internal self-recognition of the beholder’s image through the mirroring function of the MNs.
The cobbles were seen as an external “advertisement” of the beholder being mirrored in the world.
Due to their “mirroring” qualities these cobbles were deliberately “taken out” of their natural
context, they became “meaningful” for their beholder and were invested with a “content” that
enabled them to stand for something else—i.e., the gatherer’s face.
in my view, it was the fortuitous association with such cobbles which induced in hominids
thoughts about a primordial process of self-projection in the world; a process, which, may have
triggered the bidirectional functioning of the MNs which, in the course of millennia, shaped a
behavior that approximates “art creation.” Hominids eventually started to perform actions on the
world, thus creating an “artworld.”
These cobbles were, in a way, the first “readymade” artworks.10 Beads and pendants, proto-
sculpture, engravings and notches, petroglyphs and manuports were the earliest vestiges of
hominid palaeoart, marking their everyday “artistic” activity which, along with cave paintings
depicting animals in different postures as well as scenes with hominids in rituals and ceremonies,
delineated specific perspectives of life-performances in their community. “Yet no practice of art
could carry force that was not already grounded in careful and attentive observation of the lived
world,” as Ingold puts it (Ingold 2019, 660). These were the earliest manifestations of “artistic
behavior.” It was the initiation of a process of expressing inquiries about different possible modes
of living and imagining different relations with the world, which from that point onwards were
depicted in the form of primitive “artworks.” The creation of alternative modes of living depicted
in “artworks” was the hominids’ performative act on their own daily lives, a behavior which
nowadays is called “artistic.” For our early ancestors, “performing art” was a way of being
performative in their life, testing different possibilities with a view to being more effective in the
future. In that sense, “art” is ontologically performative; hence the claim of my proposal that
performing art is, essentially, a way of performing life.
There is no evidence that for our earliest ancestors “art objects”, such as the manuports referred
to, were unconnected from their everyday activity or that they constituted a natural category that
was distinct from artifacts. On the contrary, the scenes depicted in caves and the engraves in the
open air, or the variety and abundance of “artworks” found all over the world, seem to be deeply
embedded in their everyday practices in a performative way. The interdisciplinary approach to the
Arts may provide additional premises and produce new theoretical advancements for the theorists,
critics of art and artists to elaborate further into “art’s” performativity in life. That art performance
is essentially a performance of life had come to the fore in the so-called ‘performative turn’ in the
arts—during the last decades of the twentieth century and in the twenty-first century—when the
boundaries between life and art were crossed over, becoming almost imperceptible. By adopting
this stance, we could say, contemporary performing arts seem to resonate with the original necessity
out of which artistic modality emerged.

10
The concept of “readymades” which I introduce here in relation to the hominids’ manuports, is made with reference to
Marcel Duchamp’s “readymades.” Both have marked important turns in the performing arts, which I will elaborate further
in a future article.
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ABOUT THE AUTHOR
Evi Prousali: Adjunct Lecturer, Department of Performing and Digital Arts, School of Fine Arts,
University of the Peloponnese, Nafplion, Greece

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