Neurobiology and the Humanities


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Neurobiology and the Humanities
Semir Zeki1,*
1University College London, London, WC1E 6BT, UK
*Correspondence: s.zeki@ucl.ac.uk
http://dx.doi.org/10.1016/j.neuron.2014.09.016
This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/3.0/).

Can the arts and humanities contribute significantly to the study of the brain? Similar brain processes are
involved in humanistic and scientific inference, and in this essay, I argue that conclusions reached by one
are relevant to the other.
Can the arts and humanities contribute

significantly to brain studies? Do they

frame questions regarding human experi-

ence that can be tested experimentally

and are these fundamentally different

from those posed by neuroscience? Is

there any present need or imperative to

appropriate questions from them in

neurobiological studies, or should that

be deferred until more is known about

the functions and functioning of the brain?

These questions impose themselves

forcefully at a time when a significant pro-

portion of human brain studies are ad-

dressing questions that are of importance

to human experience.

Common Questions
Science and the humanities have much to

separate them but much to unite them

too. Artistic and scientific questions are

commonly the same, though addressed

differently, and hence, the former provide

hints and guesses for scientific experi-

mentation. I have written of artists and

humanists as being, in a sense, neurobiol-

ogists who explore the brain, though with

techniques that are unique to them (Zeki,

1993). Paul Cézanne’s preoccupation,

and artistic experimentation, with how

color modulates form is but a variant of

the neurobiological question of how the

separate representations of form and

color are integrated in the brain to give

us a unitary percept of both (Zeki, 1978;

Livingstone and Hubel, 1988). The exper-

iments of Picasso and Braque in the early,

analytic, phase of cubism—of how a

form maintains its identity in spite of

wide variations in the context in which it

is viewed—resolves itself scientifically

into the neurobiological problem of form

constancy. The quest of Piet Mondrian

for the ‘‘constant truths concerning
12 Neuron 84, October 1, 2014 ª2014 The Au
forms’’ (Mondrian, 1941) is an artistic

version of the question of what the neural

building blocks of all forms are (often pre-

sumed to be the orientation-selective

cells of the visual cortex), while kinetic

art, which sought to represent motion

artistically, reached conclusions that are

consistent with conclusions reached later

by neurobiology (Zeki and Lamb, 1994).

All Truths Are Subjective
These are, in a sense, facile rallying points

that merely serve to emphasize different

approaches to what are, at heart, com-

mon questions. More difficult to address

are shared questions regarding human

experience and what they signify about

brain operations and the world in which

it has developed. Here the boundary

between neurobiological and humanistic

questions is faint and separating the

two, I believe, does both a disservice

even if, at present, the relationship be-

tween neuroesthetics and the humanities

is asymmetric, in that neuroesthetics

has a good deal more to gain from the

humanities than the latter from us. Many

of the critical questions now addressed

experimentally by neuroesthetics have

been addressed in philosophical dis-

course for centuries. Prominent among

these is the problem of knowledge, a pri-

mordial function of the brain and a central

issue in philosophy. Using color vision as

an example, Arthur Schopenhauer argued

that ‘‘a more precise knowledge and

firmer conviction of the wholly subjective

nature of color contributes to a more pro-

found comprehension of the Kantian doc-

trine of the likewise subjective, intellectual

forms of all knowledge’’ (Schopenhauer,

1854), since color is a subjective experi-

ence that is the result of a transformation

of the objective reality of the outside world
thor
by rules that govern the operations of the

mind (brain). The only knowledge we can

therefore have of color is ‘‘brain knowl-

edge’’. The brain, far from representing

colors (or indeed the sensory world)

passively and veridically, constructs

them through inherited programs (algo-

rithms) (Zeki, 1993). Neurobiology has

yet to unravel the details of these opera-

tions, but their purpose is to stabilize the

colors of surfaces in spite of continual

fluctuations in the wavelength-energy

composition of the light reflected from

them, leading to a constancy of colors.

While we can be (subjectively) sure that

a leaf is green even when it reflects more

long-wave (red) light (as is common at

sunset or sunrise), we can never be sure,

unless armed with light-measuring de-

vices, of the ‘‘objective’’ reality in terms

of the precise wavelength-energy com-

position of the light reflected from a sur-

face and from its surrounds. Generally

speaking, the only truths that we can be

certain of are those that we experience,

namely subjective truths. This is but one

example of a shared general question in

neurobiology and the humanities—of

how objects and situations maintain their

identity in spite of continual changes in

the signals reaching the brain from them,

summarized for Western philosophy in

the Heraclitan doctrine of flux and for

Eastern (Buddhist) philosophy in the

statement that ‘‘nothing is permanent

except change.’’

Similar Inferential Processes in
Scientific and Humanistic
Approaches
The primacy of subjective truths extends

from an apparently elementary process

such as color to much more complex

experiences, such as those of beauty,

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desire, and love as well as to abstract con-

cepts such as the experience of mathe-

matical beauty. The path to acquiring

knowledge—whether grounded in scien-

tific experimentation or through philo-

sophical (Cellucci, 2013) or humanistic

speculation—must use similar mental

processes. There is no reason to suppose

that thebrain processes leading to subjec-

tive truths—in terms of inference, which is

the result of observation and of inductive,

deductive, and analogic reasoning—are

different for the sciences and the human-

ities. Indeed, the similarity may extend to

metaphoric and metonymic reasoning.

The humanistic approach—be it in art

or philosophy—is equally grounded in

experimentation, of a different, more

speculative kind but one that is neverthe-

less also subject to the logic of the brain.

Its results, significantly, lend themselves

to scientific experimentation. Hence, in

seeking to understand human nature

and the human condition, conclusions

reached by humanistic debate and dis-

cussion are no less or more valid than

those reached by scientific experimenta-

tion, even if translation from humanistic

achievements to scientific experimenta-

tion is neither straightforward nor easy. A

major difference is that, to attain scientific

status as valid for populations instead

of individuals, subjective truths require

scientific validation, usually through sta-

tistical inference. Indeed, given their

longevity and the similarity in brain pro-

cesses leading to inferences in both the

sciences and humanities, subjective

truths revealed by humanistic discourse

can in fact be said to have also been sub-

ject to scientific experimentation and sta-

tistical validation and hence provide rich

material for scientific experimentation.

The works of Plato, Sophocles, Kant,

Shakespeare, Dostoevsky, and Balzac,

among others, have a longevity even sur-

passing those of scientific works because

they reveal subjective truths that are

generally applicable to all humans. One

is likely to acquire as much experimentally

testable knowledge, for example, from

reading Kant on aesthetics or Balzac and

Zola on creativity than one would from

any presently available scientific text.

The Experience of Beauty
Perhaps nowhere is the interdependence

of humanistic enquiry and experimental
investigation more intertwined than in

the study of one of the most ubiquitous

of (subjective) human experiences—that

of beauty; it serves as a powerful ground,

as well as an example, for uniting

the humanistic and neurobiological ap-

proaches. Neuroesthetics does not

enquire into what beauty is and does

not (contrary to common belief) confound

it with art. It also acknowledges the

importance of culture and learning in

shaping aesthetic experience. But its pri-

mary concern at present is to understand

the neural mechanisms that allow all

humans, regardless of race or culture, to

experience beauty. Since an aesthetic

experience implies having made a judg-

ment, it also aims to unravel the neural

systems underlying aesthetic judgments

and address the question, first posed by

Kant, of whether aesthetic judgments

precede or succeed aesthetic experi-

ences. In short, like the art critic Clive

Bell, neuroesthetics seeks to understand

what, in aesthetic experience, is ‘‘com-

mon to all and peculiar to none’’ (Bell,

1914), which is not to deny that, superim-

posed upon the commonality, there are

subjective differences in experiences

that science must account for.

It was, after all, a philosopher, Edmund

Burke, who defined beauty in significantly

neurobiological terms, as being ‘‘largely

a property of objects acting upon the

human mind through the intervention of

the senses’’ (Burke, 1757, my emphasis).

Today, much of the inspiration for the

paradigms used to study the neurobi-

ology of aesthetic experience, whether

acknowledged or not, comes from philo-

sophical studies.

Though Bell thought of aesthetic expe-

rience as a ‘‘purely subjective business,’’

he, like others before and after him,

sought for ‘‘objective’’ characteristics

that constitute an essential ingredient of

beauty. Whether such a characteristic

exists has been debated but without a

consensus. This is not surprising. Sym-

metry, for example, is not considered to

be characteristic of beauty in all cultures;

it does not therefore qualify as a charac-

teristic that is ‘‘common to all and pecu-

liar to none.’’ Characteristics such as

proportion or size, though of importance

in domains such as architecture, are

meaningless when applied to the aes-

thetics of, for example, color. As well,
Neuron
there is the functional specialization in

the brain and in vision, for example,

different areas of the (visual) brain are

specialized to process different attributes

such as color, motion, and form (Zeki,

1978; Zeki et al., 1991). This suggests

that, in the visual domain, there may be

many different visual characteristics

(which I have termed ‘‘significant configu-

rations’’; Zeki, 2013), belonging to dif-

ferent visual domains, each one capable

of activating the relevant visual area in

a manner that arouses the ‘‘aesthetic

emotion’’ in that domain, as appears to

be the case for kinetic stimuli (Zeki and

Stutters, 2012). But there is a common

characteristic that is independent of

learning, culture, and ethnic origin to all

that is experienced as beautiful, one

that is ‘‘common to all and peculiar to

none.’’ It lies in a simple neurobiological

fact—that whenever an individual experi-

ences beauty, regardless of whether the

source is visual, musical, moral, or math-

ematical, there is a correlate in the form

of activity in a part of the emotional brain,

namely field A1 of medial orbitofrontal

cortex (A1 mOFC) (Ishizu and Zeki,

2011). Interestingly, this area is also

active when subjects have pleasant or

rewarding experiences—both of which

have been strongly linked to beauty in

the philosophy of aesthetics (Gordon,

1997), providing a good area for future

experimentation designed to reveal the

relationship, in neural terms, between

these subjective experiences. This raises

the question of what role the sensory

areas of the brain play in translating

significant visual configurations into

an aesthetic emotion, a neurobiological

problem of importance that extends well

beyond neuroesthetics. Whether stimuli

such as faces, for example, are perceived

as ugly or beautiful, they activate com-

mon areas critical for the perception of

faces (Kanwisher et al., 1997; Haxby

et al., 2000). But faces that are perceived

as beautiful correlate as well with activity

in mOFC (O’Doherty et al., 2003), while

those experienced as fearful correlate

with activity in amygdala (Morris et al.,

1996). Some feature of these stimuli

must activate the common areas differ-

entially, leading to different outputs from

them. Neurobiologically, the question

resolves itself into the broader one of

the pattern of activity within a common
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area that dictates the selective output

from it to one destination or another.

The Larger Significance of Beauty
If all truths, whether sensory, aesthetic,

or derived from higher cognitive and intel-

lectual sources such as mathematics are

subjective, it becomes interesting to ask

whether the (subjective) experience of

beauty in general is a pointer to universal

truths about ourselves and the Universe

in which we have evolved, just as sensory

experiences such as those of color are

pointers to truths about ourselves and

the ever fluctuating world in which we

have evolved. The experience of color,

derived from a sensory source, reveals

a truth about how our brain obtains

knowledge by stabilizing the continually

changing world in which it has evolved

sensorially. That the experience of mathe-

matical beauty, just like the experience

of musical and visual beauty, correlates

with activity in field A1 of mOFC not only

shows the abstract nature of beauty

but also raises the question of whether

beauty, regardless of its source, is also a

pointer to deeper truths, a sort of yard-

stick for determining the truthfulness of

what that experience reveals. Put simply,

to what extent is the structured order,

or the ordered structure, of the Universe

in which we have evolved reflected in

the organization of our brains and to

what extent is the experience of beauty

a pointer to that structure? Beauty, which

lies at the heart of these questions, is a

topic that has traditionally been more

speculated on in humanistic debate,

though one that is becoming of increasing

interest to science and especially neuro-

biology. A neurobiological quest for what

enables us to experience beauty and

what that experience signifies is vastly

impoverished without significant reliance

on speculations in the humanities.

The Many Uses of Beauty
In The Descent of Man, Charles Darwin

made sexual selection the centerpiece

of his views on beauty and there seems

little doubt that, for example, plumage

on male birds, often perceived as beauti-

ful by humans, reveals a (subjective) truth

in the females about desirable male char-

acteristics in that species, making the

bearer a suitable mate for reproductive

selection. But, as Rothenberg (2011) has
14 Neuron 84, October 1, 2014 ª2014 The Au
emphasized, this raises the question of

why a particular combination of colors

is chosen, and why particular structural

patterns are used by, for example, bow-

birds to create their bowers to attract

females. Basing beauty on sexual selec-

tion alone also leaves out of account other

examples of beauty such as camouflage,

which have functions the opposite of

attracting sexual attention (Rothenberg,

2011). Hence, an enquiry into why partic-

ular patterns or colors are chosen to act

as sexual attractors also constitutes an

enquiry into whether what is experienced

as beautiful is related as well to what

coincides with patterns in our brain, which

has evolved to construct a picture of the

external world.

That fundamental laws governing the

structure of our Universe can be ex-

pressed in mathematical formulations

that arouse the ‘‘aesthetic emotion’’ has

long been emphasized by mathemati-

cians, who in general place a high pre-

mium on beauty. Plato and the Platonic

tradition suppose that mathematical for-

mulations are experienced as beautiful

because they give insights into the funda-

mental structure of the Universe and

hence its beauty. Kant went beyond and

supposed that such formulations arouse

the aesthetic emotion because of the

feeling that ‘‘they make sense’’ (Breiten-

bach, 2013). What ‘‘makes sense’’ is of

course what corresponds to the workings

and above all the logic of the brain. Hence

the aesthetic emotion, even in the ‘‘queen

of sciences,’’ may be a pointer as much

toward truths about both the Universe as

about the workings of the brain. It leads

one to enquire, for example, whether

humans would have developed string

theory, for which there is little if any exper-

imental evidence, if we did not possess

the kind of brain organization that we

have. It is a fascinating question.

In summary, once we acknowledge that

all knowledge is mediated through the

operations of the brain and its cognitive

apparatus, and is therefore subjective,

and that similar brain processes are

involved in humanistic and scientific

inference, we are led ineluctably to the

view that conclusions reached by one

are relevant to those reached by the other

and that the humanities provide a rich

source of hints about the operations of

the brain, which neurobiology and more
thor
particularly neuroesthetics should be

ready to exploit and is doing so.

ACKNOWLEDGMENTS

The work of this Laboratory is supported by
the Wellcome Trust, London. I am grateful to
Carlo Cellucci, Marco Federighi, Tomohiro Ishizu,
Konstantinos Moutoussis, and Dragan Rangelov
for their comments on earlier versions of the
manuscript.

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http://dx.doi.org/10.1007/s11406-013-9497-3
http://dx.doi.org/10.1007/s11406-013-9497-3
http://dx.doi.org/10.1371/journal.pone.0021852
http://dx.doi.org/10.1371/journal.pone.0021852

	Neurobiology and the Humanities
	Common Questions
	All Truths Are Subjective
	Similar Inferential Processes in Scientific and Humanistic Approaches
	The Experience of Beauty
	The Larger Significance of Beauty
	The Many Uses of Beauty
	Acknowledgments
	References