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In search of the period eye:

Contributions from neuroscience

MA Renaissance Studies: Dissertation

Pa le Ni ko i


Table of Figures ....................................................................................................................................... 3

Introduction .................................................................................................................................... 4


General Literature Review .............................................................................................................. 6



A Note on Function ............................................................................................................... 13

Contributions from Neuroscience................................................................................................. 16


Mirror Neurons ..................................................................................................................... 16

Case Studies: Empathetic Responses ............................................................................................ 20



Ambiguity .............................................................................................................................. 34


Plasticity ................................................................................................................................ 41

Conclusions ................................................................................................................................... 48

Appendix 1 - Contemporary account of empathetic responses to paintings ....................................... 51

Bibliography .......................................................................................................................................... 52

Table of Figures
Figure 1 - Olga Rozanova, Green Stripe (1918)
Figure 2 - Rogier van der Weyden, Deposition (c.1435)
Figure 3 - The Goddess Parvati
Figure 4 - Bartholomew, James, son of Alphaeus and Andrew, detail from the Last Supper
Figure 5 - Jesus, Thomas, James the Greater and Philip, detail from the Last Supper
Figure 6 - Matthew, Jude Thaddeus and Simon the Zealot, detail from the Last Supper
Figure 7 - Judas Iscariot, Peter and John, detail from the Last Supper
Figure 8 - Stefano Di Giovanni Sassetta, The Last Supper (1423)
Figure 9 - The Crucifixion, miniature from a 13th-century Psalter
Figure 10 - Mary of Cleophas (?), detail from the Deposition
Figure 11 - Mary Magdalene, detail from the Deposition
Figure 12 - Eos and Kephalos (1st half of 5th century before Christ)
Figure 13 - Niobide (1st half of 5th century before Christ)
Figure 14 - Image of Maenad
Figure 15 - Relief of Maenad
Figure 16 - The 'rabbit or duck' illusion
Figure 17 - The Necker cube illusion
Figure 18 - Bi-stable image showing a vase or two faces
Figure 19 - Johannes Vermeer, Girl reading a letter at an open window (c. 1657-1659)
Figure 20 - Johannes Vermeer, The Music Lesson (c. 1662-1665)
Figure 21 - Diagrams representing functional and anatomical changes to a neural network (Kandel
2006, 214)

1. Introduction
One fundamental problem in art history is the difficulty of coming to terms with art produced in
times and cultures remote from our own. How can the paintings of Rogier van der Weyden, for
example, be fully appreciated today, when paintings in 15th-century Netherlands were conceived
under very different economic conditions to those of today and with very different objectives in
mind? I his primer in the social history of pictorial style , Painting and Experience in FifteenthCentury Italy1 (1988), Baxandall proposes several ways of bridging this gap, albeit in the context of
15th-century Italian painting. Partly by being attentive to both the economic context of the
Quattrocento and its proper language for talking about art, and partly by invoking the concept of the
pe iod e e , da a dall (1988, 151)

a ages to sha pe ou pe eptio

of Renaissance painting.

The concept of the period eye refers to those cognitive skills that influence the way a particular
society in a particular period perceives art. The cognitive revolution of the 20 th century most
certainly contributed to the formation of this idea. I believe that the neuroscientific developments of
today have a similar potential for influencing the way art is understood. With this in mind, in this
dissertation I aim to answer the following research question: How can neuroscience contribute to
the appreciation of Renaissance art nowadays?
There are three relevant sets of skill, as Baxandall (1988, 31) explains; a sto k of patte s, atego ies
and methods of inference; training in a range of representational conventions; and experience,
drawn from the environment. Essentially, standing before a work of art, the beholder brings to it a
combination of innate and socially-developed skills, olle ti el also efe ed to as og iti e st le ,
that contribute to the meaning of that work for the beholder (Baxandall 1988, 29). Essentially, this is
a more thorough address of what Gombrich (2002a, 246) called the
projection of the

eholde s sha e , that is the

eholde s own life and experience into the image. And, since the painter

To be referred to as Painting & Experience for the remainder of this essay.

endeavours to

at h the o

o de o i ato of his audie e s og iti e skills, the pe iod e e

influences pictorial style itself (Baxandall 1988, 40).

I po ta tl , the pe iod e e shapes ou po e s of dis i i atio . o a e tai p ofessio , da a dall
(1988, 39) ela o ates, leads a

a to dis i i ate pa ti ula l effi ie tl i ide tifia le a eas. Fo

example, Baxandall (1988) suggests that merchants are capable of discriminating between volumes
of barrels, sacks or bales, and are invited to do so by painters who would place in their works
geometric puzzles as a way of engaging their audience.
In this way, Baxandall places visual culture within a social context. His former teacher, Gombrich, on
the other hand, has been criticized for divorcing art from society, as a subject of study removed from
all other aspects of human life (Langdale 1998). It is its contextualised nature that makes the period
eye a particularly helpful instrument of art historical study, since it permits one to glean what it was
like to experience painting in fifteenth century Italy.
I believe that recent advances in the understanding of the human brain could enrich the concept of
the period eye. Interdisciplinary studies have, thus far, focused on modern art. In this dissertation I
propose to investigate the relevance of these studies for art of the Renaissance in parts of Europe
(for the purposes of this essay this covers mainly c.13th to 16th centuries). To do this, and to provide
the academic context for the remainder of the work, I will begin with an overview of the literature
dealing with the application of neuroscience to art. This shall be followed by a more focused analysis
concerning, in turn, the following three areas: mirror neurons, ambiguity, and neuroplasticity. The
section on mirror neurons will also include a more extensive case study of empathetic response
hi h

ill e ased o Leo a do da Vi i s Last Supper a d ogie

a de We de s Deposition.

Unfortunately, due to limited space, the latter two sections will focus on the theoretical aspects
without detailed case studies, although I will venture to provide examples where appropriate. A
concluding section will summarize the main findings in light of the literature and case studies

It should be noted at the outset that this dissertation will not differentiate between different types
of beholder. Although a limitation of the work, it should not hinder the identification of certain
trends in the analysis. It must be recognised, however, that different types of audience may have
experienced paintings in very different ways. Any further study in this field would certainly warrant
closer attention to discrete categories of audience, based on social class, sex, or profession, for

2. General Literature Review

In this section I shall give an overview of the literature broadly concerned with the contributions of
neuroscience to art history. But firstly, to set the context for the discussion to follow, it is necessary
to define the problem in more detail.
The problem of recreating the social and artistic experience from history is an enduring one in art
history. The works of Heinrich Wlfflin in particular, embedded a historical awareness into art
history as a discipline (Harrison et al. 1998, 682-683). In Renaissance und Barock (1888), in which he
sets about explaining the stylistic change in architecture from one period to the other, Wlfflin (1998
[1888], 723)

ites that To explain a style then can mean nothing other than to place it in its general

histo i al o te t 2. Wlffli s ie
depe ded o histo i all disti t

of the ha ges i

st le, as Kes e (2009, 266) points out,

odes of isio , elated to the Weltanschauung of the pe iod .

Additionally, Wlfflin plays no small part in establishing the concept of the history of seeing in art
history (Gombrich 2002a, 14).
If we accept the premise then that there indeed exists a histo
espo ses, i deed of e lai i g the



of seei g, the

assessi g past

(Freedberg 1989, 22) seems to be a necessary

step towards sharpening our perception of Renaissance painting. Going as far back as the
Renaissance, however, a culture very unlike ours, the problem is exacerbated by the lack of

Ei e til e kl e ka
i hts A de es heisse als ih
ei eihe (Wlfflin 1888, 65)

a h sei e

Ausd u k i die allge ei e eitges hi hte

documentary evidence that might otherwise offer an insight into contemporary responses to
artworks (Baxandall 1988, 24).
Harrison et al. (1998, 683) identify the work of Wilhelm Dilthey as one of the most thorough to
address this problem. While some observers have deemed the problem of historicism
insurmountable (Gombrich 2002a, 54), Dilthey maintained, as did Gombrich after him, a more
optimistic stance that the a eful stud of a t o ks ould i deed i spi e o e s histo i al i agi atio
to t a s e d o e s o

o te t (Harrison et al. 1998, 683, Gombrich 2002a, 54). In this way Dilthey

hopes to e o e the app e iatio , utilizatio , a d e e

a e of the e isti g

ajo fo

s of a t

(Dilthey 1998 [1892], 725-726).

It is important to note that

of da a dall s ideas a e fou ded i pli itl o e pli itl o


and, in particular, psychology. Indeed his chapter on the period eye (1988) begins with a standard
description of the mechanics of vision. Gombrich, perhaps more than any other art historian,
promoted the use of psychology (claiming in one interview that his approach to art history was
always biological (Onians 2007, 159)) to shed light on problems in the discipline. His influential book,
Art and Illusion, egi s ith a uote f o

Ma J. F iedl de s Von Kunst und Kennershaft: A t ei g

a thing of the mind, it follows that any scientific study of art will be psychological. It may be other
things as ell, ut ps hologi al it ill al a s e. (2002a, 1).
However, Gombrich wrote Art and Illusion in the 1960s, and Baxandall wrote Painting & Experience
in the 1970s. Since then, a great deal of progress has been made in understanding of the mind3. The
20th e tu


the a i al of a

s ie e 4 (Kandel 2006, xii). Considerable advances in

technology have exposed the human brain to closer examination than ever before. Although still far
from fully understanding this highly complex organ, techniques such as functional magnetic
resonance imaging (fMRI) and positron-emission tomography (PET) scans have allowed new insights

Fo the pu poses of this disse tatio I shall use the te s i d a d ai i te ha gea l . Ka del (2006,
xii) states that o e of the p i iples of the e s ie e is that i d a d ai a e i sepa a le a d that,
si pl put, i d is a set of ope atio s a ied out the ai .
Else he e efe ed to as eu os ie e , ai studies o
ai s ie es .

into the

ai s inner workings. Not only does the new technology permit the observation of single

cells, but also of neural circuits in action (Kandel 2006, 305). Indeed, by the 1990s, fMRI allowed for

o ito i g of eu al a ti it i

eal ti e (Onians 2007, 3). The ability to investigate the brain

directly at a neural level has allowed scientists gain greater understanding of the mind by direct
observation, thus gradually leading away from a strict behaviourist approach to psychology. Kandel
(2006, xii) su

a ises the

s ie e of

iolog to e a i e the g eat e ai i g

i d as a s ie e that uses the po e of

ole ula

ste ies of life .

Although there may well be other aspe ts of the

s ie e ele a t to e aissa e pai ti g,

owing to restrictions of space, this work will limit its scope to mirror neurons, ambiguity, and
neuroplasticity. In the process, I will highlight how each of these complements existing art-historical
thinking about Renaissance painting, using examples where appropriate. In order to establish a
suitable context it will be helpful to begin with a brief overview of the more general trends in the
One of the most comprehensive accounts of the rapprochement of the disciplines of art history and
neuroscience is John O ia s Neuroarthistory (2007). An important feature of this book is its title,
indicative in itself of the rising interest of both the neuroscientific and the art-historic communities
in the bearing of recent advances in neuroscience on understanding art production and reception.
Furthermore, Onians himself taught a postgraduate module in A t a d the d ai , named after the
1999 paper by Zeki, which was the first postgraduate course in an art history department that
applied neuroscientific principles. Freedberg, for his part, founded the Art and Neuroscience
Project at the Italian Academy for Advanced Studies in America in 2001. This project encourages,
a o g othe thi gs, i estigatio of espo ses to o ks of a t [since] developments in the cognitive
eu os ie es ha e g eatl

illu i ated the

eu al su st ate of su h espo ses. (The Art &

Neuroscience Project n.d.).

An interdisciplinary approach however is not without its risks. Considerable criticism can be found of
those advocating o e lo

lai s a d a

itio s of iolog a d the s ie es of the

2009, 268). One of the more high-profile cases involves Go

i d (Kesner

i h s espo se to a a ha d a and

Hi stei s (1999) somewhat provocative article The Science of Art A Neurological Theory of
Aesthetic Experience (Onians 2007, 8). Gombrich, Kesner (2009, 268) explains, objected to the
autho s poor application of biology to culture, based as it was on a simplistic and reductionist
account of art .
Although the potential pitfalls of straddling multiple disciplines are clear, I do not believe that this
confrontational account of the relationship between art history and brain studies is representative
of real trends. Each discipline has much to learn from the other. Indeed, the emphasis seems to be
very much on inter-disciplinary collaboration, as a ui k gla e at O ia s (2007) bibliography
reveals, with combined efforts from the likes of Freedberg (art historian) and Gallese (neuroscientist)
or Hobson (psychiatrist) and Wohl (art historian). Mandelbaum (1970, vii) summarises this point
The paths of art historians, psychologists, and philosophers do not normally converge,
even when they are dealing with a common problem, since their aims and their methods
are in most cases radically different. Yet, questions regarding the nature of
representation in art have recently provided a common meeting ground.
A more particular problem for the present work is that of the applicability of neuroscience to
Renaissance painting. It has been noted (Zeki 1999, 99) that the developments in our understanding
of the mind lend themselves more to analysing art of the 20th century than to previous periods
including the Renaissance. To u de sta d the easo s fo this, it is i po ta t to app e iate )eki s
general approach to the subject. His starting assumption is that the function of art is an extension of
the function of the visual brain, that is to acquire knowledge about the world (Zeki 1999, 1, 8). This

assumption leads Zeki to extrapolate principles of art from principles of vision. As shall be shown,
this method encounters important limitations when applied to Renaissance art.
As a neuroscientist specialising in the human visual system, Zeki in the 1980s and 1990s contributed
much to the discovery and investigation of functionally specialised areas of the visual brain. The level
of specialisation turned out to be greater than scientists suspected. Not only does the modularity of
vision mean that there a e spe ifi g oups of ells spe ialised i a diffe e t att i ute of the isual
s e e, su h as fo

, olou a d


(Zeki 1999, 59), but there are specific groups sensitive to

particular colours but not to others, to vertical lines but not horizontal ones, or to movement in a
particular direction (Zeki 1999, 60-62).
One can begin to understand the way in which neuroscience can help one understand how a
painting such as Olga oza o a s Green Stripe (Figure 1) affects the viewer. A bold green vertical
stripe on a pale background excites localised groups of cells activated by those formal features, and
the contrast itself is reinforcing for the brain since areas of high contrast in nature are usually
information-rich and therefore attract attention(Ramachandran and Hirstein 1999, 25). The act of
activating particular modules of the visual system in a targeted way and then enhancing them is also
known as isolation (Ramachandran and Hirstein 1999, 22). It is this type of rather direct connection
between the painting as a visual stimulus and the excitation of particular brain areas that prompts
Zeki to declare that the artist is in a sense, a neuroscientist, exploring the potentials and capacities
of the


(Zeki n.d.). Where the approach seems to be less effective, however, is with a painting

such as Leo a do s Last Supper or Rogier van der Weyden s Descent from the Cross (Figure 2).


Figure 1 - Olga Rozanova, Green Stripe (1918)


a t s te de

Figure 2 - Rogier van der Weyden, Deposition (c.1435)

to a ds abstraction and simplification means that, as stimuli, modern

paintings arouse more isolated areas of the brain and therefore more specific responses than more
complex figurative or narrative paintings (Zeki 1999, 99). For this reason, Inner Vision (1999) contains
many examples from Impressionism, Cubism, Suprematism, Fauvism, and Kinetic art. Although Zeki
develops a chapter entitled A

eu o iologi al app aisal of Ve

ee a d Mi hela gelo su h

examples are rare. It is interesting to note too, that Ramachandran typically uses Indian art, such as
the sculpture of the Goddess Parvati in Figure 3 for example, to elucidate his laws of aesthetics, but
these too exhibit elements of abstraction in a way that Renaissance paintings do not. Even within
the scope of modern art, it is recognised that science has little to offer in terms of explaining works
such as Ma el Du ha p s Pissoir, implying that responses to this work are primarily socially
constructed (Ramachandran 2000).


Figure 3 - The Goddess Parvati

I believe that the starting proposition that the overriding function of painting is the discovery of the
world is reductionist to the point of offering little if any insight into figurative and narrative art.
Although Zeki may be correct on some meta-level, this is not particularly helpful to art historians.
Co t a

to )eki s u if i g defi itio , a useful and necessary distinction between arts produced in

different periods must be based on function.

According to Gombrich (1984, 20), art should be judged primarily by standards of utility. Throughout
histo ,

o ks of a t

e e o side ed to ha e a defi ite fu tio

(Gombrich 1984, 20). As a

consequence, it is not against our present taste that art should be assessed, but rather against a
easu e of

hethe it

o ks in terms of fulfilling its intended function (Gombrich 1984, 23).

Similarly, in his discussion of the Aims and Limits of Iconology (1978, 5), Gombrich emphasises
Hi s h s p i a

of ge es i esta lishi g the i te tio of the a tists a d the efo e the asi le el

of meaning of the work itself. Baxandall too highlights the importance of reconstructing the a tist s
intention in order to fully appreciate a painting, although it is debatable to what extent this is really
ever achievable (Baxandall 1989, 109).
While in Expressionism the 20th century saw the rise of an entire movement dedicated to exploring
the a tist s i

e e otio al

o ld a d o

u i ati g it to a

ide audie e, in the 15th century, as


Baxandall (Baxandall 1988, 3) put it, painting was still too important to be left to the painters. The
a tist i the e aissa e o ked to a p og a

e p o ided

his pat o (Gombrich 1978, 6). Such

a programme was contractual and could specify not only the subject to be painted, but also which
colours were to be used, what quality pigment was to be used, and what figures were to be
included, for example (Baxandall 1988, 6, 12). This therefore poses a problem for the Renaissance
art historian wishing to explore Renaissance art from a biological perspective, since it appears that
biology played a relatively minor role in shaping the art relative to certain other factors. This point is
central to Freedberg also, who, in his quest for universals in The Power of Images Studies in the
History and Theory of Response (1989, xx), stresses the importance of not fo getti g that espo ses
are forged on the anvil of culture and in the fi e of pa ti ula histo

I believe this problem is not insurmountable, but it does require a more detailed look at the function
of Renaissance paintings.


A Note on Function

Painting in the Renaissance had a fundamentally different function to art of the 20th century. Most
Renaissance painting was religious. This apparently trivial observation, in fact, has certain important
consequences. There existed a tradition in Catholic thinking (Welch 2000, 137), stemming from Pope
Gregory I at the turn of the 6th century, and filtering down to Thomas Aquinas in the 13th century. A
contemporary of Aquinas, Johannes Balbus (John of Genoa), wrote in his Catholicon that the reason
for images in churches was threefold;
First, for the instruction of simple people [...] Second, so that the mystery of the
incarnation and the examples of the Saints may be the more active in our memory
through being presented daily to our eyes. Third, to excite feelings of devotion, these
being aroused more effectively by things seen than by things heard. (Johannes Balbus
cited in Baxandall 1988, 41)


This orthodox stance was reiterated by the Council of Trent in 1563 (Welch 2000, 165). This is not to
suggest, however, that religious art was uniform since the time of Gregory I. Indeed, an important
poi t to ote is that G ego

I s adage o l

efe s to the fi st of the th ee parts that Balbus mentions

above, and the other two were, therefore, added later. This development may be related to the
important shift which occurred between the 13th and 16th e tu ies, he e

the ste

atio alit of

scholastic theologians was gradually modified by warmth of feeling and emotion (Huyghe 1967, 19).
St Bernard, like St Francis, encouraged their followers to contemplate God not only though the mind
but also through the emotions. This shift resulted in a need for a physical and material
In Painting & Experience, Baxandall (1988, 43-47) deals with Quattrocento painting, which
functioned very much as a collaborative creation between the artist and the audience, the former
providing the scene as a generic foundation for the latter to build their personal mediation upon. As
su h, the pai te

as a p ofessio al isualize of the hol sto ies (Baxandall 1988, 43). The painter,

therefore, avoided depicting people or places in too much detail, instead complimenting the
eholde s interior vision (Baxandall 1988, 45-47). Although the centuries that followed saw a shift
towards greater detail, partly influenced by both technological and stylistic developments in the
north countries, the basic function of paintings in Renaissance Italy as aids to religious mediation did
not change.
An important piece of evidence for the role internal representations played in prayer, the book
Zardino de Oration (1454) offers the following instructions:
Moving slowly from episode to episode, meditate on each one, dwelling on each single
stage and step of the story. And if at any point you feel a sensation of piety, stop: do not
pass on as long as that sweet and devout sentiment lasts. (cited in Baxandall 1988, 46)


In his take on the emotive function of images, Dominican Fra Michele da Carcano draws attention to
another important aspect of the tendency towards realism during the 15th century. He writes that
with images

be moved when they see [the Saints as if a tuall p ese t cited in Baxandall

1988, 41). Gombrich (2002b, 84), among others, repeatedly draws attention to the importance of
the beholder s ole as eyewitness for the development of painting in the Renaissance away from the
pi tog aphi


stee pai ti g i

of pi to ial a atio to a ds atu alis . Giotto was one of the key figures to
the di e tio

of d a ati

ep ese tatio , i flue ced partly by friars whose

sermons encouraged the faithful to visualise the events from the Bible and the lives of the Saints as
if they were actually present (Gombrich 1984, 151-152). It became necessary not only to represent
hat happe ed, as i the pi tog aphi st le , ut also
21, 89, Gombrich 1976, 16). It

u iall how it happened (Gombrich 2002b,

as pe haps thus that G ego

I s adage

as modified in the late

middle ages, although one can only speculate.

The early Franciscan tract, the Little book on the meditation on the Passion of Christ divided
according to seven hours of the day by Pseudo-Bede, provides an invaluable glimpse into the method
of meditation required from the faithful:

It is necessary that when you concentrate on these things in your contemplation, you do
so as if you were actually present at the very time when he suffered. And in grieving you
should regard yourself as if you had our Lord suffering before your very eyes. (cited in
Freedberg 1989, 171)

According to Freedberg (1989, 171), the narrative in the book addresses the reader occasionally to

ui e, What the

ould ou do if ou e e to see these thi gs? . Des iptio s of Ch ist s suffe i g

in this, as in other treatises that followed, were very gruesome and vivid (ibid.).

The above discussion shows that when considering art from any period, including the one under
investigation here, from a neuroscientific perspective, the focus should be less on the direct

connection between the mind of the artist and that of the beholder, but rather on the intended
function of the art work and the mind of the beholder. By implementing an integrated approach
which combines the analytical elements in this way, it may be possible to establish the relevance of
neuroscience for Renaissance art. With this in mind, I will not turn to the first of the three
developments from neuroscience to be discussed.

3. Contributions from Neuroscience

The present section will focus in more detail on the three key discoveries to be considered, namely,
mirror neurons, ambiguity, and neuroplasticity. The section on mirror neurons will also include two
paintings Leo a do da Vi i s Last Supper a d ogie

a de We de s Deposition) as case studies

to be analysed in light of the theory. At a high level, the different parts will be organised in terms of
an explanation of the discovery followed by an examination of its relevance for Renaissance
painting. At the end of each section, I will pause to reflect on the contributions offered by each of
the discoveries.


Mirror Neurons

there was speech in their dumbness, language in their very gesture (Shakespeare, The
Winter's Tale Act v Scene 2 Line no 14-15)
The observation by Rizzolatti et al. in 1996 that certain neurons in macaque monkeys fire both when
the monkey performs an action and when it observes that action being performed by a human is
co side ed

a a ha d a as the single most important "unreported" ... sto

(Ramachandran n.d.). The easo

fo a a ha d a s e thusias

behaviour of mirror neurons (dubbed by hi


o ke see

is that the

o ke do eu o s

of the de ade
ha a te isti
ould e

u ial

in understanding previously elusive aspects of human nature such as non-verbal communication,

learning by imitation, the development of language, and, crucially for the present discussion,
empathy. O e s a ilit to put o eself i

a othe s shoes, to e pathise

ith thei

ood, to

understand the intention behind their actions, could all be attributable to a large extent to mirror
It must be recognised that discussion of the role of empathy in art follows in the tradition of the likes
of Friedrich Theodor Vischer, Robert Vischer and Heinrich Wlfflin. As early as the 19 th century,
F ied i h Vis he

ote that the u o s ious p o ess

he e

the su je t p oje ts his o he

feelings into the inanimate object must be investigated on the level of our physical and psychological
o ga izatio

(Harrison et al. 1998, 682). Ulti atel , it

as F ied i h s so , o e t Vis he ,


embedded empathy as a central concern in art history (Harrison, Wood and Gaiger 1998, 682). In
1873, Robert Vischer adopted the term Einfhlung5 to des i e the ph si al espo ses that a e
ge e ated

the o se atio of pai ti gs (Freedberg and Gallese 2007, 198). Building on Robert

Vis he s ideas, Wlffli e plo ed the ps hologi al a d ph si al effe ts of pa ti ula a hite tu al

forms on the beholder (Freedberg and Gallese 2007, 198). At the turn of the 20th century, Aby
Warburg wrote about the Pathosformel: the representation of inner emotional states through
particular physical representations of figures. Bernard Berenson even approached the modern
understanding of empathetic responses, suggesting that the movement of figures depicted in
e aissa e a t as e pe ie ed

the ie e s o

us les, the e

e ha i g thei e pe ie e

of the work itself (Freedberg and Gallese 2007, 198).

Looking back even further, Socrates is purported to have taught the need to represent not only the
ph si al i a t, ut that the ph si al should i tu

e eal the the a tio of the soul Tes psyches

erga) (Gombrich 2002a, xxii). Indeed, Warburg believed that Renaissance masters learned the skill of
achieving Pathosformel from classical Greece (Gombrich 2002a, 20). This seems to agree with
Pa ofsk s theo

that the Italia enaissance of the 14th and 15th centuries was characterised by a

app o he e t of lassi al fo

a d lassi al o te t (Panofsky 1972, 111).

A lite al t a slatio i E glish ould e feeli g i .


All this , Go

i h (1970, 36) su

a ises su i tl , te ds to o o o ate the h pothesis that e

interpret and code the perception of our fellow creatures not so much in visual but in muscular

Since their discovery, the existence of mirror neurons in the human brain could only be suggested
via fMRI scans that indicated increased activity in certain areas of the brain. Very recently, however,
obstacles to observing individual cells within the brain were finally overcome. This allowed for the
existence of mirror neurons in humans to be proven, and for their mechanism to be demonstrated
(Mukamel et al. 2010). In their important article, Motion, emotion and empathy in esthetic
experience, Freedberg and Gallese (2007) specifically focus on mirroring mechanisms and empathy,
describing several different types of empathetic response to visual stimuli. Broadly, three key types
of response are identified. Each shall be described in turn.

The fi st o e s the

odil e gage e t

ith gestu es,

o e e ts a d i te tio s of othe s

(Freedberg and Gallese 2007, 201). The authors offer as an example Mi hela gelo s Prisoners,
explaining that espo ses ofte take the form of felt activation of the muscles that appear to be
a ti ated


the s ulptu e itself (Freedberg and Gallese 2007, 197), thus effectively

communicating to the beholder the sense of exertion the figure must be experiencing as it tries to
free itself from the block of marble (Freedberg and Gallese 2007, 198).

The se o d t pe of

i o

espo se o e s the feeli g of e path fo

odil se satio s

(Freedberg and Gallese 2007, 201-202). Ca a aggio s Incredulity of St. Thomas, for example, is shown
to arouse ie e s so atose so

s ste

as the pe ei e to e tou hed a d p o ed the sel es.

Furthermore, these tactile sensations are localised in that part of the body observed to be affected
in the painting, i this ase the tho a

he e Jesus

ou d is situated. A d i Go a s Desastres de la

Guerra, bodily empathy arises as a result of both the representation of unbalanced figures as well as
from images of bodily harm and mutilation. The latter is seen to activate those areas of the brain


normally activated during the actual experience of pain as well as the ensuing shock (Freedberg and
Gallese 2007, 198). Significantly, the experience of physical empathy often translates into emotional
empathy the thi d t pe of

i o

espo se (Freedberg and Gallese 2007, 197). The latter point

suggests that a beholder is capable of identifying the emotions of others by internally imitating their
actions (Freedberg and Gallese 2007, 201-202).

It is worthwhile introducing, at this point, a consideration of function into the discussion. In The
Power of Images, Freedberg (1989, 1-2) offers two contemporary accounts of empathetic responses
to paintings. The first of these, from 1584, emphasises the ability of a painting to rouse the beholder
to a wide variety of emotions or desires, depending on what is represented in the painting (see
Appendix 1). This a ou t e hoes Al e ti s On Painting (1970 [1435-1436]) where he wrote that we
weep with the weeping, laugh with the laughing, a d g ie e ith the g ie i g . The second account,
from 1587, is worth restating here in full:

[...] since the eye is the most perfect among the exterior senses, it moves the minds to
hatred, love and fear, more than all the other senses ...; and when the beholders see
very grave tortures present and apparently real ... they are moved to true piety, and
thereby drawn to devotion and reverence all of which are remedies and excellent
means for their salvation. (cited in Freedberg 1989, 2)

This account brings to the fore the tradition of empathic meditation (Freedberg 1989, 164). This
tradition ties in firmly with the earlier discussion of function, and it is worth remembering that

e e a effe ti e

a of di e ti g people s atte tio to a ds Ch ist s suffe i gs a d good

deeds by exciting them to empathy (Freedberg 1989, 164).

It is interesting to note that mirror neurons are an inbuilt feature of the human brain. This suggests
that the neural mechanism underlying empathy is universal as it is shared by all humans regardless
of when or where they lived. This is an important point, and one we shall return to, as it addresses

the perennial debate in art history about whether there are any aspects of human cognition that are
universal in this sense, or whether this is always historically specific and moulded by the particular
society or culture in question.

In o side i g the od a d its la guage i


-century painting, Baxandall (1988) focuses on the

highly codified and formalised movements and gestures exhibited. This method of interpretation can
be revealing, although highly dependent on the scarce literary evidence from the time. Thus, the
Mirror of the World from the 1520s provides a rare insight into the meaning of gestures by matching
them to particular emotional states or particular circumstances. This approach does, however,
overlook those more spontaneous gestures, of which the interpretation is none the less important. I
believe that complementing the contemporary literary evidence from the period with an
understanding of mirror neurons may help the modern audience view Renaissance art more like a
Renaissance audience. I will now look at two specific examples of paintings where I believe
knowledge of mirror neurons and their role can provide toda s viewer with an augmented
experience of the works.
Case Studies: Empathetic Responses
The t o pai ti gs I ill o e plo e i

o e detail, i

ie of the dis ussio a o e, a e Leo a do s

Last Supper a d ogie s Deposition. Although the paintings are from different decades and
locations, I believe each one can demonstrate different aspects of the empathetic response. It is the
i te tio he e to use the t o ase studies to de o st ate ho the p oposed i teg ated app oa h ,
combining neuroscience and a consideration of function, can contribute to the appreciation of
Renaissance art and also highlight any limitations of the approach.
Leonardo da Vincis Last Supper
The istoria will move the soul of the beholder when each man painted there clearly
shows the movement of his own soul. [...] These movements of the soul are made
known by movements of the body. (Alberti 1970 [1435-1436])

The most important consideration in painting is that the movements of the figure
expresses its mental state, such as desire, scorn, anger, pity, and the like. (da Vinci 2008,
Leonardo da Vinci painted the Last Supper in 14951497 in what was at the time a mausoleum of
the monastery of Santa Maria delle Grazie in Milan (Zllner 2007, 122). The painting depicts the
moment that Jesus tells his apostles that one of them would betray him: When Jesus had thus said,
he was troubled in spirit, and testified, and said, Verily, verily, I say unto you, that one of you shall
betray me. (John 13.21 . The Gospel of Joh fu the e ou ts the dis iples ea tio s as the looked
one on another, doubting of whom he spake Joh

. These reactions of surprise, anger and

questioning are captured in the group of three to the left of the painting (Saints Bartholomew,
James, son of Alphaeus and Andrew) and another to the right of Christ (Saints Thomas, James the
Greater and Philip).

Figure 4 - Bartholomew, James, son of Alphaeus and

Figure 5 - Jesus, Thomas, James the Greater and Philip,

Andrew, detail from the Last Supper

detail from the Last Supper

The a ou t o ti ues to des i e the s e e: Simon Peter therefore beckoned to him, that he
should ask who it should be of whom he spake. (John 13.24). This is captured in Figure 6. Peter then
lea s i to a ds Jesus

east a d asks hi , hile Jesus a s e s: He it is, to whom I shall give a sop,


when I have dipped it. And when he had dipped the sop, he gave it to Judas Iscariot, the son of
i o . (John 13.26). Figure 7 shows Judas and Jesus reaching for the same piece of bread.

Figure 6 - Matthew, Jude Thaddeus and Simon the Zealot,

Figure 7 - Judas Iscariot, Peter and John, detail from the

detail from the Last Supper

Last Supper

The story is infused with drama. Leonardo manages to convey a sense of this through the gestures
and attitudes of the apostles. Both James, son of Alphaeus, and James the Greater raise their hands
in astonishment, for example, the former raising his eyebrows and the latter with mouth wide open.
Bartholomew, meanwhile, stands up to lean across the table with great intensity, both hands on the
table, and his powerful gaze adding to the spectacle. It is interesting to note an account of
Leo a do s f iend Luca Pacioli, who wrote that it was
not possible to imagine the apostles more agitated upon hearing the voice of unfailing
truth [...] In their poses and their expressions, they seem to be speaking one to another
and this one to that with vigorous astonishment and dismay. (cited in Zllner 2007, 122)
Even Darwin (2006 [1872], 1430) efe s to the Leo a do s Last Supper to e e plif asto ish e t i
his writings on the gestures of the human body. Just as Alberti (1970 [1435-1436]) advocates the use
of many and varied gestures in a painting to render the picture more compelling, Leonardo here
deploys these gestures with a specific aim. The image conveys

hat it

ust ha e ee like


Christ uttered those words (Gombrich 1984, 226).


A consideration of the eholde s

i o espo se permits the art historian to place the beholder at

the centre of the analysis. To that end, it is possi le to u de sta d Leo a do s use of e p essi e
gesti ulatio as i ed to the ha a te s as a

ethod of heighte i g the eholde s e pe ie e of the

scene. In beholding the Last Supper, in accordance with the taught meditation techniques, the

ust o e t ate o ea h of the figu es i tu . A d, i a o da e ith Leo a do s o

intention, and thanks to the mirror system, he should feel angered with Andrew, astonished with
James the Greater, and perhaps saddened with John. Not only does the beholder reflect
intellectually on the parts of the story as he concentrates on each character or group of characters in
turn (as he would have done presented with an earlier, pictographic version of the story (Figure 8)),
but reacts empathically to each character also, based on a mirror response to each gesture, thus
being further implicated as a witness to the scene.

Figure 8 - Stefano Di Giovanni Sassetta, The Last Supper (1423)

There is the danger here of overstating the case. The gesticulations of the Apostles would be difficult
to interpret to this level of meaning if the beholder was unfamiliar with the story (Gombrich 2002b,
69). U dou tedl ,

othe fa to s ould ha e fed i to the eholde s e pe ie e of this

u al.

There is, simultaneously however, the danger of dismissing altogether the part played by empathy
(and by the mirror mechanism). An experiment cited by Changeux (1994, 192) provides considerable

insight. The neurologist Alexandre Luria, while investigating the impact of lesions of the frontal lobe
one of the areas where mirror neurons occur (Rizzolatti and Craighero 2004) found that when
patients were presented with a painting by Baron Klodt, the Last Spring, they were incapable of
understanding its content. The scene depicted a d i g gi l sitti g i a a


hile he elde l

parents watch her sadly and her sister stands by the window in an attitude of profound sorrow
(Luria in Changeux (1994, 192) . Lu ia s su je ts, u a le to establish a coherent view of the picture,
guessed ased o the gi l s go

that the

e e looki g at a

eddi g s e e. Cha geu i tuiti el

suggests that the frontal cortex pla s a esse tial ole i dis e i g the

e tal states, affects,

beliefs, desires and intentions of othe s, whether real or represented in a painting (Changeux 1994,
192). It may be said then, without exaggeration, that it is thanks to the mirror neuron system that
the gestures can be interpreted at all. I believe the knowledge of the story and the understanding of
the gestures in terms of emotional states combine in the painting and reinforce one another to
produce a powerful holistic effect.

The e is a eed, ho e e , fo

ultu al se siti it . Go

i h s (2002b, 69) perceptive analysis of the

mural reveals that in the Last Supper, Leonardo combines spontaneous and conventionalised
gesticulation, reporting that the gestures themselves are often taken as representative of a
Mediterranean culture. Having said that, both Gombrich (1976, 33) and Baxandall (1988, 48-49)
emphasise the trans-cultural influence of popular preachers in the 13th century. As Baxandall (1988,
64) points out, these e e skilled isual pe fo

e s ith a odified a ge of gesti ulatio

ot spe ial

to Ital . It has been demonstrated, in addition, that the expression and interpretation of certain
basic emotions (such as happiness, sadness, fear, and surprise) exhibit remarkable consistency
across cultures (Feldman 1993, 368-369, Ekman and O'Sullivan 1991).


ust e oted that,

all a ou ts, Leo a do s Last Supper was an exceptional rendition of this

traditional theme (Zllner 2007, Gombrich 1984, 224). It is notable precisely for the animated nature
of the characters and the diversity of their attitudes, as the contemporary accounts demonstrate.

Pe haps this the pla es Leo a do s Last Supper among those works that traversed the tradition of
pictographic representation and ushered in the new, dramatic, style of painting. One reason why this
depiction was unusual was due to the elati e diffi ult of t a slati g Jesus ps hologi al suffe i g
into a visual representation. The dis ussio a o e has sho


o side i g the eholde s

mirror response, the audience is brought to the fore of the analysis, and the intended experience of
the work can be reconstructed somewhat more accurately than is possible otherwise. Furthermore,
the analysis has shown that consideration of the mirror response only makes sense and adds value
to the analysis if considered in conjunction with the art-historical context. As a theme, the Last
Supper seems to have been less open to the imparting of the type of emotional intensity discussed
above than certain other themes from the Passion. Other more physical aspects of the Passion offer
the painter a more straightforward message to convey. The following section addresses this
hypothesis in more detail by examining Rogier a de We de s Deposition6.

Rogier van der Weydens Deposition

One approach to determining the meaning of medieval painting, pioneered by Panofsky in his
influential article on the Arnolfini Portrait (1934), is that of disguised s



espo se to the

ensuing pursuit of coded messages in pictures, Marrow (1986) tries to re-focus attention on two
points that he deems more central to the task of understanding pictures. The first of these is a
greater emphasis on the eholde s espo se i.e.
152)), and the second is the ole of the

hat o e "does" i f o t of it, o

o ks i sti ulati g e

ith it (1986,

states of o s ious ess Marrow

uses Rogier s Deposition to elaborate on these points (Figure 2).

i ila l to Leo a do s Last Supper, Rogier s Deposition stands at the crossroads between the 13thcentury pictographic tradition and the increasingly prominent dramatic representation in the 15th
century. The 13th-century Crucifixion below (Figure 9), for example, converted scripture into image
form in such a way as to conceptualise the key theme; the idea of compassion. This was embodied,

Also known as The Descent from the Cross, or simply Descent.


t pi all ,
othe ,

the Vi gi o the o e ha d,

ho e pathised


it essed he so s death, a d

t. F a is o the

ith Ch ist s suffe i g to such a degree that he experienced the stomata

himself (Marrow 1986, 153). The Virgin is shown, furthermore, with a sword piercing her heart, as a
lite al ep ese tatio of i eo s p ophe

Figure 9 - The Crucifixion, miniature from a 13th-century Psalter

The 14th century saw the invention of the Andachtsbild (the devotional image). These images were
characterised by the gradual isolation of holy figures from recognisable settings (Marrow 1986, 153).
Rather, artists distilled the

essage of Ma

s a d Ch ist s o passio ate lo e i to its

o e esse tial

elements (Marrow 1986, 153-154). This development coincides with the rise of personal meditative
hat Pa ofsk

devotion in the northern countries,

efe ed to as

o te plati e i

e sio

(Marrow 1986, 154). These images had the function, therefore, of stimulating the appropriate
emotions in viewers (Marrow 1986, 155).

Yea, a s o d shall pie e th ough th o

(Luke 2.35)

soul also, that the thoughts of

hea ts

e e ealed.


Rogier s Deposition shares some of the Andachtsbilds characteristics. As von Simson (1953, 9) points
out, the s e e is st i tl li ited to the hu a sphe e ,

ithout the pa ti ula s of a kg ou d o

landscapes to distract the viewer from the unfolding drama. The intended effect of the composition
is to pu sue the si gle the e of la e t i to its fi est e otio al a ifi atio s (von Simson 1953,
10). Indeed, the subject of the painting can be specified as the juxtaposition of the passio and
compassion (von Simson 1953, 11). This is ep ese ted isuall


s postu e

i o i g that of

Jesus. It is worth pausing here to describe briefly the historical development of this theme.
The 12th and 13th centuries saw an increasing intimacy in the relationship between Mary and her
dead son. The asso iatio of Ma

s suffe i g ith that of Ch ist s is a pa t of Ch istia do t i e that

stems from the writings of Bernard of Clairvaux in the 12th century (von Simson 1953, 11). The
association grows stronger throughout the 13th century, inspired partly by the increased emphasis
laid by St. Francis and his followers on the affect. Whereas in previous times Mary was shown to
possess a p iestl dig it

i fa e of he so s sa ifi e, the forming attitude now held the Virgin s

suffering in such reverence that Bonaventure was inspired to suggest that at the Crucifixion Mary
as t a sfo

ed i to the like ess of Ch ist (von Simson 1953, 12-13).

These tendencies reached a peak in the 14th and 15th centuries, giving rise to what von Simson (1953,
13) efe s to as the age of Co passio . The Piet as a devotional image, for example, now
becomes second in popularity only to the Man of Sorrows (von Simson 1953, 14). Indeed, the
Speculum humanae salvationis, published in 1324, did much to popularise the association by
following the e e tatio s of Ch ist s Passio

ith the e e o o s of Mary (von Simson 1953,

12-13). By the mid-15th century, then, the passio and compassio are perceived as inseparable by
writers such as Bernardine of Siena and Denis the Carthusian (von Simson 1953, 14). As von Simson

a ises, The

aste the e

adopted the mariology of the most famous theologian of his time

and country [Denis], i a i age hi h popula de otio s e de ed i telligi le to e e o e. (1953,



Although Marrow (1986, 156) criticises von Simson for over-emphasising the importance of liturgical
and doctrinal content of the painting, both authors do, in fact, insist on the primacy of the
eholde s espo se. Von Simson (1953, 16) e plai s that it is th ough esta lishi g the

o d of

si ilitude between the beholder and Jesus that the artist brings the doctrine to life. Indeed,
unlocking the meaning of Rogier s Deposition lies on the level of human experience or, in other
o ds, the do t i e had to e li ed to e o e effe ti e (von Simson 1953, 15).
Marrow (1986) claims that the ie e s espo se is not only inspired towards the appropriate
response by contemplating Christ and the Virgin, but in the s e e s witnesses, and not least the
Virgin herself, these responses are actually exemplified (Marrow 1986, 155). This, according to
Marrow (1986, 155), represents a shift in emphasis unique to the Renaissance, away from the
traditional subject matter, that is the story or idea depicted, and towards the beholder. In some
se se, Ma

i ogie s Deposition is an aspirational representation of the beholder. In summary

then, Marrow (1986, 157) argues that works such as Rogier s Deposition
surpassed the symbolic images of compassion of the thirteenth century, and the
Andachtsbilder of the fourteenth, both of which espoused compassion only conceptually
or by inference, by visualizing and thus explicitly extolling diverse forms of emotional
response to traditional Christian subjects.
Subsequently, Ma o s (1986, 163) argument turns to establishing the relationship between artistic
stimuli and consciousness itself. At a high level, he explains that the new method of representation
required a different engagement from the viewer. In particular, the meaning of the work now
depended to a much greater degree on the viewer experiencing the appropriate emotional
response. An important change, therefore, from Middle Ages was characterised by a work of art no
lo ge

ei g defi a le

hi h a e o e tl

e el

hat is ep ese ted

ut its

ea i g ei g vested also in effects

ulti ated i the eholde (Marrow 1986, 169).


Although I ag ee ith the ke ideas i

oth Ma o s (1986) and Simson s (1953) analyses, I do not

think that these fully address the question of the beholder s espo se to the
Ma o

ok i

uestio .

does t uite e plai why the image of Mary should affect the viewer in the first place.

Although her image certainly functions as a reminder that hers is the appropriate emotional
response that the beholder should aspire to, I would like to suggest that this works in combination
with the mirror response aroused by the image. The image of Mary involves the viewer in a direct
bodily engagement with the painting. As methods of engaging the ie e s o s ious ess, Ma o
(1986) cites instances of artists using mirrors that face the viewer or outwardly gazes from
protagonists. I think that the evocation of the different types of mirror response could be added to
this list of tools that the artist can deploy to achieve his aim. The recognition and understanding of
this tool seems critical, therefore, to recreating the intended 15th-century experience of the painting.
Practical experiments investigating the viewer response to different paintings are not within scope
of this essay. One might only hypothesise here that results of such tests on the effects of ogie s
Deposition would reveal the different characters in the painting to arouse different mirror
responses. Aside from Christ and Mary, already discussed, one would expect viewers to empathise
with the distressed woman on the left of the panel (Figure 10) (perhaps Mary of Cleophas, Mary s
sister), symptomatic of an emotional mirror response. Viewers may also feel unbalanced, as they
ha e ee sho

to do fa ed

ith Go a s Desastres de la Guerra (see Section 3.1), by the sight of

Mary Magdalene, t isted as she sta ds at Ch ist s feet (Figure 11).


Figure 10 - Mary of Cleophas (?), detail from the

Figure 11 - Mary Magdalene, detail from the Deposition


What may be referred to as the mechanics of a painting - that is its composition, colours, and other
formal aspects - are largely discernible. If the stated hypotheses are correct, however, this would
cast fresh light onto the mechanics of the beholder s espo se, all the more important if this is the
true (or at least other significant) subject of the painting. Again, I would argue that the power of the
painting is derived from the successful integration of the narrative, the formal aspects of the
painting, and the manipulation of the audience participation by the evocation of empathetic
response. Thus, if indeed the do t i e had to e li ed to e o e effe ti e (von Simson 1953, 15), it
is now possible to see the important roles played by the artist, the painting, and the audience in
bringing it to life.
To eite ate, p ese ted ith ogie s Deposition the modern viewer is faced with the usual problem
of bridging his time and culture and that of Rogier and his audience in order to appreciate the
painting more fully. As von Simson (1953, 15) poi ts out, The e te t to hi h the age as
o te platio of these e e ts su passes


o ed

u de sta di g. An awareness of mirror responses

can perhaps reveal an overlap between the modern viewer and the 15th-century viewer s espo se.



ith Leo a do s Last Supper, the analysis demonstrates how the investigation into mirror

responses could be combined with an appreciation of the iconography to contribute to recreating

the original intended experience of the work.
Perhaps the new light shed on this work ca also help add ess Ma o s (1986) concern with the
relationship between the consciousness of the audience and the artwork. While the inversion of the
subject matter described above is certainly significant, it may not be as unique as Marrow suggests
(1986). He writes that, Ne e
ee so e pli itl

efo e [...] had the situations of the observed in art and its observer

e e sed. (Marrow 1986, 165). It

e app op iate he e to e all Pa ofsk s

theory about the rapprochement of classical form and classi al o te t, a d to suggest that ogie s
Deposition i fa t o tai s ele e ts of Wa u g s Pathosformel. What I would like to suggest here is
that ogie s Deposition adopts not only classical forms, but classical aims too, in order to achieve
the highly emotive effect described above, albeit in a different context, namely a Christian one.
Examples of possible classical forms prefiguring images of the dead Christ are shown in the figures
below. Figure 12 is reminiscent of images of the Piet, while Figure 13 of the deposition itself.



Figure 12 - Eos and Kephalos (1 half of 5 century before



Figure 13 - Niobide (1 half of 5 century before Christ)



Parallels between the characters in these mythological scenes and Jesus and Mary are not the only
ones that can be drawn. Indeed, Warburg exemplified his concept of Pathosformel with reference to
the Maenads as the types borrowed in Renaissance scenes of the Crucifixion to depict the mourning
women at the foot of the cross (Wind 1937). Indeed, Figure 15 offers a striking comparison with

Magdale e i ogie s Deposition.

Figure 14 - Image of Maenad

Figure 15 - Relief of Maenad

What is more, I believe it is the universality of the mirror system mechanism that permits certain
forms to be reused in different contexts while maintaining particular effects. If this is true, it would
be testable experimentally, and would be supported if toda s ie e s were shown to exhibit the
predicted espo ses. This

a also pa tl e plai

h so e pai ti gs, su h as ogie s Deposition,

receive as much attention today as they do despite the cultural and temporal distance. This point
shall be developed further in Section 3.3.
To conclude, with reference to the research question stated in Section 1, the discussion and analyses
above have gone some way to answering how neuroscience can contribute to the appreciation of
Renaissance art nowadays. Crucially, the analyses of the case studies are demonstrative of an
integrative approach to understanding art works. Zeki (1999, 28-29) argues that what is important


for a general appreciation of a work of art can be gained through a direct experience of it thanks to
the universality of perception. The rest, he claims, should be left to the connoisseurs. I do not think
that a strict distinction between these different kinds of knowledge about a painting is a helpful one.
I believe it is precisely the marriage of these different elements that can add the most value to
toda s appreciation of the works. I believe that the analysis here has shown that any universal
aspects of brain mechanisms, in this case empathetic response, cannot be isolated from the rest of
iconography and can only be truly helpful in revealing the meaning of paintings when integrated
with iconographic and other art-historical knowledge.
This approach has revealed several specific insights into Renaissance art. Whilst lending valuable
suppo t to da a dall s ideas about how gestures and movements in paintings were read by a
contemporary audience, the consideration of mirror responses has allowed for further
reconstruction of the intended experience of these works. In particular, it has allowed for the
consideration of the effect of the less codified and formalised gestures that Baxandall (1988)
explores in Painting & Experience. A key to this is the emphasis given in the analysis to the
eholde s esponse itself. This seems particularly relevant given the increasing consideration in the
pe iod gi e to the eholde s affe ti e i ol e e t i the o ks. Highlighting these aspects in the
analysis, it has also been possible to suggest broader implications for the persistence of certain
evocative forms (Pathosformel a d thei elatio to the eholde s o s ious ess. This section has
also elaborated on some key themes of the discussion. Namely, it has explored the roles of nature
and nurture in the production and consumption or art works, as well as the corresponding status of
the universal versus the historical in art history. These themes shall be explored further in the
following Sections.
In spite of the limitations of )eki s app oa h discussed above, he does nevertheless make some
significant contributions, not least through the study of ambiguity to which I shall now turn.




Ambiguity rabbit or duck is clearly the key to the whole problem of image reading.
(Gombrich 2002a, 198)

i h s u ha a te isti all

hi si al o

e t does i t odu e a o etheless i po ta t the e

to the present discussion. Ambiguity can be understood at various levels of complexity. Simple
perceptual ambiguity, at one end of the scale, is well exemplified by such bi-stable illusions as the
rabbit/duck illusion (Figure 16) or the Necker cube illusion (Figure 17). In both cases, the stimulus on
the page is constant while the brain switches between two alternative views (between a rabbit and a
duck in the former and between two alternative perspectives on the cube in the latter). These
illusions present the brain with too little information to be able to resolve the ambiguity. As a result

ai s it hes et ee t o e uall plausi le i te p etatio s, although at a

given time, only

o e i te p etatio is possi le (Zeki 2004, 183).

Figure 16 - The 'rabbit or duck' illusion

Figure 17 - The Necker cube illusion

To understand this phenomenon at a neural level, it is necessary to distinguish, in the first place,
et ee o di a

odes a d esse tial odes . As

e tio ed i Section 1, the brain is subject to a

high degree of functional specification. In the human visual system, some groups of cells are
responsible solely for detecting motion, whilst others react only to certain colours, and others


respond to orientation (i.e. vertical or horizontal forms). A neural node describes one collective of
su h spe ialised ells. A ode is said to e esse tial if its a ti atio

esults i the o s ious

perception of the attribute that that node is responsible for, without further processing by the brain.
To account for the register of discrete attributes due to the activity of essential nodes, Zeki (2004)
has oi ed the te

' i o o s ious ess . O e pa t of the isual s ste

i the


olou pe eptio , a ed V , is a good e a ple of a esse tial ode. A ti it the ei

o s ious pe eptio of olou

esponsible for
esults i


ithout the eed fo fu the p o essi g (Zeki 2004, 177). Essential

nodes can, therefore, be thought of as perceptual sites in the brain that function simultaneously as
processing sites. Intervention from a higher cognitive area such as learning or memory (also
efe ed to as the top-do

i flue e is not required to process and interpret the stimulus.

Consequently, the illusions discussed above imply that each essential node, and therefore each
microconsciousness, can have multiple states, with each state resulting in a different interpretation
of the stimulus (i.e. rabbit or duck). Zeki (ibid.) conjectures that these states must be determined by
the strengthening of one set of cells within the node at the expense of another.
These ambiguous images demonstrate another key principle of perception. At least since Plato, a
psychological hypothesis has existed that human perception is concerned with the extraction of
stable universals (the essential) from the ever-changing world around us (Gombrich et al. 1970, 3).
This particular role of the human visual system is upheld
k o ledge that is

o th a ui i g,


studies i this field. The o l

ites Zeki (1999, 5), is k o ledge a out the e du i g a d

ha a te isti p ope ties of the o ld . As su h, under normal circumstances (for the above illusions
are contrived), the

ai is a

iguit - esol i g

a hi e pa e elle e (Kandel 2006, 297),

processing, as it does, incomplete and changing inputs to create a logi all

ohe e t output. It

should be noted, furthermore, that as an incentive for the brain to discover hidden objects (an
important skill in nature), the brain has evolved to enjoy the exercise (Ramachandran 2000).
Following from this evidence, ambiguity can be defined as an unstable state characterised by a
certainty of several logically coherent interpretations a plurality of truths (Zeki 1999, 25).

Psychologists have termed the

t a sie t as

o sta

efe ed to as olou

ai s a ilit to e de sta le a d lo g-lasting what is dynamic and

. A o di gl , the a ilit to pe ei e a olou i

o sta

a d dista es is alled fo

a i g light o ditio s is

, hile the a ilit to ide tif a pa ti ula fo m from different angles

o sta

. (Zeki 1999, 22). In practical terms, this is the capacity, for

example, to identify a particular face not only as the lighting and angles vary, but even as its
expressions are animated.
Zeki (1999), building on his assumption that the function of art is the same as that of the visual
system, offers a neurological appraisal of Michelangelo. The following is an example of the potential
effect of formal ambiguity. Zeki (1999) claims that Michelangelo, in pursuit of the Platonic forms,
struggled with the problem of depicting something as ethereal as spiritual beauty and divine love.
Being subjects not of this world, it is only through the imagination that mortal men can hope to
attain them. Michelangelo thus leaves the Rondanini Piet in a comparatively unfinished state,
unlike his earlier more idealised works, to capture the imagination of the beholder (Zeki 1999, 31).
d lea i g the

o finite, Michelangelo invites the spectator to be imaginatively involved, and the

spe tato s ie

a fit

of the Co epts, the sto ed ep ese tatio s, i his


(Zeki 1999,

Interestingly, a comparison between the Rondanini Piet and the Atlas from the Prisoners series,
which Freedberg (2007, 197) noted tends to evoke a muscle response from beholders, as discussed
in the previous section, reveals a shared unfinished quality. The particular example of the Rondanini
Piet, I am tempted to speculate, thus combines effects from embodied empathetic responses with
those of ambiguous form to stir both mind and body of the beholder towards the intended
In terms of brain mechanisms, the more complex an ambiguous image the greater the number of
areas that are activated in the brain. It has been reported (Zeki 2004, 186) that the bi-stable image in
Figure 18 activates separately the areas involved in object recognition and those involved in face

recognition. Experiments, however, also indicate that the fronto-parietal cortex (involved in higher
cognitive function) is engaged as the brain switches between one percept (i.e. interpretation) and
the other (Zeki 2004, 186).

Figure 18 - Bi-stable image showing a vase or two faces

This has led Zeki (1999, 29) to speculate that highly complex ambiguous images such as the paintings
of Vermeer, while subject to the same principles as simpler ambiguous images, engage many more
areas of the brain, including its memory. The brain is still tasked with attributing meaning to the
image by establishing the most likely state (or situation in the ase of Ve

ee s pai ti gs that

befits the visual clues. Typically, however, Vermeer does not provide sufficient clues to allow for the
determination of just one stable state, inviting more questions than offering answers. Aside from
physical aspects, like colour and form, ambiguity can thus apply equally to less tangible aspects of art
(1999, 22). Zeki terms this notion situatio al o sta

. What is written in the note the woman in

Figure 19 is reading? What is the relationship between the man and woman in Figure 20? The
paintings themselves do not provide the answers to these questions. Hence, according to Zeki (1999,


24-25), the ps hologi al po e


of Ve

ee s

o ks o es f o

thei a ilit

to ge e ate

iguit .

Figure 19 - Johannes Vermeer, Girl reading a letter at

Figure 20 - Johannes Vermeer, The Music Lesson (c.

an open window (c. 1657-1659)


As a practical example, I would now like to consider the role of ambiguity as discussed above with
spe ial efe e e to Velz uez s Las Meninas. This large format (318cm 276cm) group portrait
depicts the King and Queen of Spain, their daughter, Velazquez himself, the court dwarfs, a
chaperone, a guard, and an attendant. Art historians, philosophers, and indeed other artists have
produced voluminous analyses attempting to decipher this highly complex painting.
The painting contains three focal points: the I fa ta Ma ga ita, Velaz uez s self-portrait, and the
reflection of the royal couple. Velzquez is one of five characters that look out at the viewer from
the painting. These gazes have the effect of implicating the viewer in the scene, effectively rendering
them a fourth focal point of the painting. This is enhanced by both the mirror where the beholder
might expect to see himself, a d the pai te s o

attitude as he sta ds efo e his a as. Perhaps

he is pausing to study his subject, before adding another brush-stroke. Thus the spectator becomes
that subject.

But the subject of the canvas that Velzquez so tantalizingly obscures from view could equally be the
king and queen or the Infanta Margarita or even the whole scene from the point of view of the
couple i.e. Las Meninas itself. Although previous studies have attempted to decipher this enigma
with certainty, I believe it is precisely the ambiguity between this multitude of certainties that lends
the painting its dynamic. It is thanks to this that Each focal point involves us in a new set of
elatio s (Clark 1960, 36). Foucault (2004, 5) also acknowledges the complex relationships between
the different roles, writing: No gaze is sta le ... su je t a d o je t, the spe tato a d the


e e se thei oles to i fi it . As with the simple illusions seen at the beginning of this section, the
compositional balance of Las Meninas means that at any one time the viewer is capable of
contemplating only one of these focal points as the subject matter, and the instability of each
perpetuates the ie e s interaction with the painting.
In the case of Las Meninas, the ultimate purpose of this illusionistic game is open to speculation.
One might argue that it addresses concerns particular to the 17th-century Spanish empire on the one
hand, and those of Velazquez in improving his social standing on the other. Considering first the
latter, the painting in which Velazquez had included his self-portrait was originally entitled The
Family (Levey 1971, 147, Portus 2004, 171), thus indicating the aspirational nature of his inclusion in
the group. With espe t to the fo

e , i pai pai te s e e ot e e pted f o

the a tisa s ta

the alcabala sales tax imposed on manufactured goods until 1677 (Shiner 2001, 67, Taggard 1996,
80). In both Florence and Rome, however, it was already recognised as a liberal art. Velzquez may
thus be making an appeal to his audience to reconsider both the intellectual underpinnings of his
art, and his own social status as an artist. Seen in this way, the ambiguity may function as a powerful
heto i al tool that poses the uestio

athe tha offe i g a

a s e s since these, ultimately,

must come from the audience. In this case, then, an understanding of ambiguity does help explain
how the painting works, but, for lack of material evidence, one can only speculate as to the aims of
the painter, or why he chose to paint the work the way that he did.


The importance of ambiguity in art has been highlighted by art historians and neuroscientists alike,
both oti g its a ilit to e gage

ith the eholde s i agi atio (Gombrich 1984, 228, Zeki 2004,

189). o ethi g , )eki (2004, 189) quotes Schopenhaue , i deed the ulti ate thi g,
o e fo the
a t , Quat e

i d to do. Whilst )eki (1999, 26) lai s that a

ust e left

iguit is a ha a te isti of all g eat

e (1998, 121) in the early 19th e tu , stipulates that It is the fi tious a d the

incomplete in every art, and these alone, which constitute art, and become moreover the sources of
the pleasu e of i itatio . Co t a

to )eki s alue judge e t, Quatremre rather introduces a

principle; that all art can only ever be incomplete, and the beholder is inevitably absorbed in the
exercise of bridging the gap between it and reality. Whilst there is a danger here of abstracting the
problem to a point where it no longer bears any relevance to the task at hand, I aim to show how
these theories have, in fact, very concrete implications for understanding Renaissance painting.
It is important to note first, that the exercise above in using ambiguity to explain Renaissance
painting is limited. Las Meninas stands out in Renaissance painting, if not in the history of art, as an
exceptional painting. The ability to shed more light on such a unique painting, as welcome as it is,
does not necessarily help improve the art historia s u de sta di g of a pe iod o a st le i a t
I think it may be possible, nonetheless, to broaden the relevance of ambiguity for Renaissance art,
again, by considering the proper context. A paradox of realism during the Renaissance, Gombrich
(1984, 228) explains, was that by attempting to replicate nature in portraits, the sitters depicted
actually came across as lifeless. In an effort to make his characters seem less rigid, Botticelli depicted
the movement of hair and garments. Leo a do s Mona Lisa o the othe ha d, used the a tist s o
i e tio of sfumato to pu posefull disto t a eas a ou d the o e s of the su je t s

outh a d

eyes. The overall effect of using the technique in this way is to introduce an element of uncertainty
a out the sitte s

ood: He e p essio al a s see s just to elude us (Gombrich 1984, 228). An


important element of the painting is thus left to the eholde s i agi atio , a d it is this i agi ati e
engagement borne out of built-in ambiguity that brings the Mona Lisa to life for the beholder.
While Gombrich associates this heightening of the psychological power of painting with Leonardo
and his important invention of distorting outlines of forms, I believe that an important parallel exists
in earlier painting, namely the religious paintings of the 15th century. As discussed in Section 2.1, the
eligious pai ti g f o

this pe iod

as t pi all i te ded to o ple e t the eholde s i te al

vision of a particular scene. Another way of looking at it, in light of the above discussion, is that the
pai te i fa t uilt a

iguit i to the pi tu e i o de to pe

it the i di idual s i agi atio to

create its own version, and become witness to the events. Thus, not by obscuring forms as Leonardo
did, but rather by the omission of specific descriptive detail, artists left open the possibility of many
variations of the scene, engaging the eholde s

i d as he t ies to fi d his. I this

a , the gap

between painting and reality becomes instrumental in the function of the painting.
In this section I have shown how recent developments in understanding the neurology of ambiguity
to o o

)eki s (2004) ph ase

a illu i ate see i gl

o ple

o ks su h as Mi hela gelo s

Rondanini Piet a d Velaz uez s Las Meninas. Crucially, however, again by considering the problem
in context and in terms of the function of the works, it has been possible to add another layer of
u de sta di g to da a dall s (1988) explanation of 15th-century religious paintings.



It seems there is nothing to exceed the plasticity of man, except, of course, the plasticity
of woman. (Gombrich et al. 1970, 11)
The last of the major developments in neuroscience that I wish to discuss is neuroplasticity. In
Painting & Experience, there are two aspects of his social history of pictorial style that Baxandall
does not dedicate much attention to. Firstly, whilst addressing in detail the impact that the period
eye has on style, Baxandall (1988) only hints at the potential reciprocity of this relationship, that is,


the influence that art could have on the period eye. And secondly, although Baxandall (1988)
suggests that experience of art is influenced by both innate and conditioned factors, Painting &
Experience focuses mainly on the latter. It has been pointed out (Onians 2007, 179-180) that,
ultimately, he gives little consideration to the passive structuring of the brain, concentrating instead
on formal training. I think that neuroplasticity, the concept that the neural networks comprising the
brain are malleable, offers fresh insight into the relationship between experience and the brain, and
into the broader but related relationship between society and art.
Since the 1960s, evidence has been growing in support of the idea that, contrary to previous belief,
the human brain does not become ha d- i ed during childhood and adolescence, but continues to
change in several important ways throughout adulthood (Doidge 2007, xviii). Although much
research is being done to determine exactly the extent of the mutability of the human brain (Rakic
2002, 614), already the discovery that brain neural circuits are not permanent or necessarily
function-spe ifi , is dee ed
e tu

so e as o e of the

ost i po ta t dis o e ies of the t e tieth

(Doidge 2007, xix).

The neural network can change in response to experience in several different ways (Kandel 2006,
204). Firstly, synaptic connections between neurons can be made weaker or stronger. Weaker
connections can be observed when a particular stimulus is repeated over time resulting in a
decreased psychological response, a process known as habituation. Alternately, the synaptic
connection can be strengthened (diagram on left of Figure 21) by other types of learning such as
sensitisation or classical conditioning. Sensitisation is the occurrence of a synaptic amplification as a
result of repeated stimulation, while classical conditioning is an associative type of learning where a
behavioural response is elicited by a previously unrelated stimulus through repeated exposure to the
stimulus in synchronisation with a stimulus that normally does evoke the response (famously studied
by Ivan Pavlov on dogs). I the
suggested that the flo of i fo

oadest te

s , Ka del (2006, 171) explains, these observations

atio i the a ious eu al i uits of the

ain could be modified


by learning . Even more astonishingly, experiments have demonstrated that learning can also cause
the a tual

e of s apses i


ai s

connections are established additi e plasti it

et o k to

ha ge (Kandel 2006, 213). New

as a result of learning (diagram on right of Figure

21), and existing connections abandoned su t a ti e plasti it

when they are not used (Doidge

2007, 298).

Figure 21 - Diagrams representing functional and anatomical changes to a neural network (Kandel 2006, 214)

The e te t of the

ai s plasti it has


de o st ated i

d a ati fashio

u e ous

practical experiments. Ramachandran and Blakeslee (Ramachandran and Blakeslee 2005, 59)
describe several methods of quickly altering a pe so s od -image, giving them the impression that
parts of their body have either changed shape or become disembodied. This, the authors explain,
de o st ates that the ight ki d of se so

sti ulatio

o pletel alte o e s appa e tl


certain knowledge developed over a lifetime in just a few seconds (Ramachandran and Blakeslee
2005, 59).
Thus the brain was shown to be undergoing constant change as a direct result of experience,
pe fe ti g its i uits so it

as ette suited to the task at ha d. (Doidge 2007, xviii-xix). It is

important to note that the changes described show a tendency towards greater efficiency. It has
been reported that, under some circumstances, healthy working parts of the brain would
compensate for parts that were not working, either due to genetic mutation or damage caused by
accidents, physically extending their influence in the brain (Doidge 2007, xviii-xix).
Another experiment described by Ramachandran (2000), involving the conditioning of rats to
distinguish a rectangle from a square, reveals a feature of neuroplasticity particularly pertinent to
the present discussion. Conditioning the rats by feeding them cheese every time they are presented
with the rectangle but not the square leads the rats to exhibit a preference for the rectangle by
moving towards it. Once the rats are then presented with the original rectangle and another,
skinnier and longer, rectangle, they exhibit preference for the new rectangle even though this is not
what they were taught to associate with food. As Ramachandran (2000) explains, what the rats
lea ed

as a ule: e ta gula it . I othe

o ds, the g eate the aspe t atio of the shape, the

g eate is the ats p efe e e fo it. This is efe ed to as the peak shift effe t a d oth a t histo ia s
(Gombrich et al. 1970) and neuroscientists (Ramachandran and Hirstein 1999) have explained the
power of caricature in these terms, as well as its bearing on art more generally. What is more, the
effect can be cumulative, evoking a stronger response every time the stimulus is presented.
These discoveries certainly bear importance for the nature/nurture debate. Although DNA

i es

hi h eu o s a e a le to o

e t


hi h othe

eu o s, uilt i to the

ai s

neural network is the potential for a multitude of behaviours. Which particular circuits are then
favoured over others, and therefore which particular responses and behaviours are
generated/produced is down to experience (Kandel 2006, 202). This radical new view of the human


as ot edu tio ist, as some feared, but more open and dynamic than could have been

d ea ed e e a fe

ea s ea lie . (Onians 2007, 4).

The relationship between art and the period eye can now be reconsidered in light of the scientific
advances described above. For one, the le d alua le suppo t to da a dall s original conception of
the period eye. A o di g to the e ide e dis ussed a o e ega di g eu oplasti it , people s dail
a ti ities do i deed alte thei

og iti e st le a d potentially their particular way of creating and

looki g at a t. The eu al effe ts of lea i g a d of ei fo e e t see

to o o o ate da a dall s

thesis that training in a particular kind of geometry will lead people to discriminate between
particular kinds of shape and lead to a preference for looking at objects that share similar (if not
exaggerated) characteristics (Onians 2009, Ramachandran 2000). The mathematical training that
many painters and members of their audience underwent in the 15th century, as Baxandall (1988)
suggests, endowed them with a sense of geometrical proportion and enhanced their powers of
discrimination between particular types of form. It seems, therefore, entirely plausible that artists
incorporated visual prompts into their paintings that would engage their audience by participating in
the exercise of these skills as well as by reinforcing them.
Importantly, neuroplasticity emphasises equally othe fo

ati e fa to s i a tists a d audie es

development besides formal training, in particular the passive structuring of the brain through
experience. In addition, neuroplasticity highlights the reactive nature of the beholder. Neuroscience
makes it possible to venture further than Baxandall could, and suggest that by merely being exposed
to works of art, eholde s espo ses, dynamic as they are, are continuously being refined, whether
they are becoming stronger, weaker, or whether new responses are being fashioned. The reciprocity
of the relationship between art and the period eye is thus brought to the fore. The potential
implications for art history are far from banal. Questions persistent in the discipline concerning, for
example, the development of taste and style over time as well as differences across geographies, can
be approached afresh. A novel explanation has been offered, for example, as to the different


emphasis placed on colour (colorito) and drawing (disegno) in Venetian and in Florentine painting
respectively during the Renaissance. This states that visual preferences developed in each city
according to the influence of their distinct environments on the populatio s

ai s (University of

East Anglia 2006). Alternatively, it may be possible to re-examine the question of what impact the
accurate presentation of perspective on the canvas may have had on the public of the 14 th and 15th
centuries. Doidge (2007, 299-300), explains that perceptual learning occurs as the

ai s a ilit to

discriminate between visual stimuli becomes greater, or when the brain learns a new way of
perceiving altogether, resulting in structural changes to the brain. This seems to support the theory
that Al e ti s On Painting (1970 [1435-1436]) was not only an instructive manual on how to execute

ethod of painting, but also crucially an instructive manual, dedicated originally to the

Duke of Mantua, on how to look at these paintings (Maniura 2010). Al e ti s ai

i this t eatise

Spencer (1970) explains, is one of making the new humanist art of Florence understandable and
desirable for a larger group of artists and patrons. (my emphasis). It seems appropriate to recall
Ma o s (1986, 163) concern with the alternative state of consciousness demanded of the beholder
by Renaissance art. Although further research would be required to test these ideas, it is significant
in itself that such questions should be reopened for consideration by developments in brain studies.

ai s p ope sit to ha ge i

oi the te

e tal Da

i is

espo se to e pe ie e has led so e (Changeux 1994, 193) to

. Plasti it is competitive , Doidge (2007, 298) writes, and Kandel

(2006, 202) discusses plasticity in terms of the selection of preferred connections from a set of
existing connections. Given that the selection process is based on the actual use of synapses and
neural pathways, I wonder whether in fact Lamarckian evolution is not a better analogy. In any case,
the relevance for art is the same, as summarised by Gombrich (cited in Changeux 1994, 189 : In art
history, the word evolution is a good deal more than a metaphor addi g that form adapts to
function through a process of mutation and selection, and then of survival of the fittest forms.


duildi g o Da ki s (2009, 192) idea that memes are the replicators of culture, just as genes are
the replicators of life (in the biological sense), Changeux (1994, 196) suggests that paintings can be
seen as a s thesis of

e es . It should be noted that here too the term is more than a metaphor.

Me es a e actually realized physically, Hu ph e s (cited in Dawkins 2009, 192) claims, [...] as a

st u tu e i the e ous s ste

. Although, plasticity is not mentioned explicitly, I think this is

precisely what Humphreys is referring to. And, continuing the analogy, as evolution of living things
occurs through genes, so the evolution of culture occurs through memes, albeit at a much greater
pace. This means that a meme, far from being an abstract concept, can be understood in terms of a
preference for a particular neural pathway, that is, a particular response to a stimulus. Where
Dawkins (2009, 197) discusses the competition among memes, one might refer back to the
competition between pathways.
In his well-balanced article, Changeux (1994) employs The Lamentation upon the Dead Christ,
attributed to Jacques-Charles de Bellange, as a case study to trace its hypothetical evolutionary
history. Changeux suggests that this work borrows memes from Greco-Roman representations of
the death of a hero. This fo

e e

as adopted i a Ch istia context, in representations of the

Piet scene, in particular that of Villeneuve-les-Avignon, as well as Do atello s The Dead Christ
Supported by Angels. Cha geu sho s ho
e o

i ed

o o i g the te

these fo

e es a d

ea i g

e es

a ha e

from biology) repeatedly over several generations of artists to

ulti atel p ese t the sel es i a u i ue o

i atio i della ge s

197) a gues, illust ates both the remarkable sta ilit of p eg a t

o k. This, Cha geu (1994,

e es of fo

strong emotional power, and their evolution . I i te p et the use of the te

ith thei

p eg a t i this

context to mean the ability of a meme to produce particularly strong effects in the brain. The
Renaissance saw the adaptation of these memes from the ancient world to its own particular
context. This echoes strongly the examples given in Section 3.1 of possible Greco-Roman precursors
to the Renaissance depictions of the Deposition or the Crucifixion. Again, this resonates closely with


Pa ofsk s idea that a distinguishing characteristic of the Renaissance was the re-alignment of
Greco-Roman form and content.
In this Section I have discussed recent discoveries about the plasticity of the human brain, and how
these contribute to the problem of bridging the temporal and cultural gap between the present
context and the particular context of the Renaissance, with a view to sharpening the modern
perception of Renaissance art. Firstly, the extent of the influence of every-day experience on the
workings and even the structure of the human brain has been exposed. This lends more scientific
suppo t to da a dall s o iginal concept of the period eye as the culturally-bound cognitive style of
humans. It has also been shown how brain plasticity can account for the inverse of the relationship
explored by Baxandall, namely the impact that art can have on the cognitive style and on society
itself. The findings, perhaps comforting to the modern observer, imply that even the first witnesses
to Renaissance art may have had to learn to look at these works in a new way. Furthermore, the
o ept of

e tal Da

i is

o La a kis

p o ides a fas i ati g i te p etatio of the e olutio

of visual culture over time. Again alluding to the defining feature of Renaissance art as the meeting
of classical forms with classical content for the first time since classical times, it goes some way to
explain how the images painted in the Renaissance came to take their form they did.

4. Conclusions
The story of the history of art is a cumulative one. Although some of the recent discoveries in brain
sciences are seen as revolutionary, the questions that they contribute to answering are not new.
This dissertation set out to explore specifically how these discoveries could contribute to the
appreciation of Renaissance art today. In particular I have looked at developments in mirror
neurons, ambiguity and neuroplasticity.
To summarise briefly, this work built on the theories of Robert Vischer, Wlfflin and others by
considering the role of mirror neurons in empathetic response. Not only is their activity crucial in


understanding and interpreting those less conventionalised gestures that appear in Renaissance
paintings, but they also illuminate the mechanisms behind personal devotional practice. This was
de o st ated

the dis ussio of ogie s Deposition.

Secondly, I explored the potential contributions a better understanding of the neural basis of
ambiguity could bring to the appreciation of Renaissance paintings. This was brought to bear not
only on the level of individual works, such as Las Meninas, but also on the level of style. Baxandall
(1988) noted the avoidance of overly-detailed descriptions of people or places in 15th-century
religious paintings, and the reasons for this. I have argued that these images functioned not merely
as a static framework onto which the beholder would superimpose his own existing vision of the
scene, but as a dynamic prompt for him to actively search and define this vision.
Lastly, neuroplasticity has provided important support to da a dall s (1988) idea that perception,
and, therefore, the production and consumption of art is influenced by active training. It has, in
addition, revealed the important influence of passive exposure to the environment. It has also been
possible to suggest that the experience of looking at art works could in itself alter people s visual
preferences, and consequently style itself.
Moreover, several important themes have emerged in this work. The debate in art history as to the
relative influence of nature and nurture and the corresponding tension between the universal and
the historical has been considered in light of the discoveries discussed. The level of understanding
today suggests that the relationship between the two sides is dynamic. It appears that certain
universal aspects of brain mechanisms are capable of exhibiting quite different manifestations and
producing varied behaviour in response to the historically-specific tapestry of environmental,
cultural and other external factors. Importantly, it has been demonstrated that bringing brain
sciences to bear on Renaissance art is not necessarily reductionist.


Furthermore, a discussion of cultural memes in terms of neural pathways has produced a possible
explanation for the presence of certain forms and means of expression in the Renaissance. Although
this explanation can only be partial, it does nevertheless cast new light on the relationship between
the Renaissance and the classical world. In particular, it le ds suppo t to Pa ofsk s (1972) theory
that the period saw a return to classical form coupled with classical content, though be it adapted to
a Christian conte t. I po ta tl , the sa e ideas

eso ate

ith Wa u g s theo

of the

The key issue to present itself in pursuing the above investigation is a methodological one. Whilst
the approach suggested by Zeki (1999) and Ramachandran (2005) of matching principles of vision to
principles of art has been moderately successful in relation to 20th century art, it has proved
insufficient in approaching narrative art of previous periods. This dissertation suggests a way of
making neuroscientific knowledge relevant to Renaissance art by contextualising it and relating it to
the definite function of art works between the 13th and 16th centuries. This integrated approach has
emphasised the ways in which each of the key developments complement existing art historical
theory. In this way it has been possible to isolate modest yet important contributions to the field.
Based on this discussion, I believe that neuroscience will continue to play an important role in the
ongoing search for the period eye.

Word count: 14,533


Appendix 1 - Contemporary account of empathetic responses to

Account from 1584, cited in Freedberg (1989, 1):

[A painting] will cause the beholder to wonder when it wondreth, to desire a beautifull young
woman for his wife when he seeth her painted naked; to have a fello-feeling when it is
afflicted; to have an appetite when he seeth it eating of dainties; to fall asleep at the sight of a
sweete sleeping picture; to be be moved and waxe furious when he beholdeth a battel most
lively described; and to be stirred with disdaine and wrath, at the sight of shameful and
dishonest action.


Al e ti, Leo dattista. O Pai ti g. Alberti - 'On Painting'. Translated by John R. Spencer. New
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Baxandall, Michael. Painting And Experience In Fifteenth-Century Italy. 2nd. Oxford: Oxford
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