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So Mondrian, for example, might trigger, suggested Zeki in his book
Inner Vision
, neuronal cells that have a preference for the orientation of the lines he has painted. Is a liking for Mondrian then a preference that our art-viewing
cells
were having? Would they not respond, like an eager dog, to
any
set of lines in the world? If we did
not
care for Mondrian, was the problem with our line-orienting cells? Could we retrain them to appreciate what Mondrian was doing?
It is not unthinkable: When people have received pulses of transcranial direct current stimulation to the left dorsolateral prefrontal cortex, they seem to like the images they are looking at more.

When I find Zeki in his office—strewn with Mondrian coffee mugs and other paraphernalia—I encounter not a raving rationalist ideologue bent on stripping the pleasure and mystery from artistic encounters but rather a courtly man with an intense twinkle in his eye, someone with a passionate interest in the workings of the human mind, an almost equally intense love of art, and what seems an entirely sensible desire to see what each might have to do with the other. “They say that neuroscience will never explain beauty and art,” he said. “First of all, we never conflate beauty and art. Second of all, we do not seek to explain either art or beauty. The most detailed knowledge of the brain would not improve Beethoven's symphonies.” I felt I was hearing a defensive echo of past slights, someone who had been politely told by fellow neuroscientists that art was beneath him and by artists that art was beyond the ken of neuroscience.

What might be going on in the brain of the average person as he looks at a painting? Whether the response has anything to do with why it is considered art, it seems churlish, even antiscientific, not to consider that knowledge as a part of the appreciation of that work. Consider Francis Bacon, with his portraits of notoriously misaligned faces. “Nobody ever describes Francis Bacon's work as beautiful,” Zeki said. “They may ascribe lots of other qualities to it, painterly qualities, and attribute savage truth to it and so forth, but nobody calls it beautiful. They call it a chamber of horrors.” Part of the reason, Zeki has suggested, may lie in the brain's instinctual response to severely disfigured faces. There is little the brain responds to more readily than human faces, particularly ones it deems attractive. Weeks out of the womb, infants are already voting with their eyes, staring longer at more visually pleasing faces.
We even seem able to judge the attractiveness of a face before we have recognized it
as
a face (it may be
so
instinctive that to even think about it, as research has suggested, seems to reduce the pleasure).
When people look at painted portraits, their liking of the paintings has correlated with the perceived attractiveness of their subjects.

Over time and over multiple exposures, sadly, the hedonic glow of looking at a lovely face begins to subside, at least as a neural signal. But there is one facial register, Zeki told me, to which the brain response never flags: severe disfigurement. “If you expose subjects to images of disfigured faces and images of disfigured objects, they soon get used to
the objects. The activity in the frontal cortex soon dies out,” Zeki said. “It never does so with disfigured faces. You can never accommodate to a disfigured face.” And so Bacon's “
visual shock,” as the artist called it, seems to tap into this innate response in a way that, for example, the Cubists do not (perhaps because, Zeki suggests, the disfigurations are less violent). As Bacon's biographer Michael Peppiatt perceptively notes, the effect of a work like
Head I
is to alert the nerves “
to
something unusual
, something sinisterly unpleasant, before the image has spelled itself out in the brain.”

The neuroscience view supports that. The visual shock may be something acutely felt, much the same way it can be when looking at disfigured images of another sort: perfectly symmetrical faces. “Symmetry” has been taken, in a just-so kind of way, as “equating” with facial beauty. Actually, all human faces have some kind of asymmetry. “
It is unnatural to have a perfectly symmetrical face,” as Dahlia Zaidel, a professor of neuroscience at UCLA, told me. There is no “golden ratio,” a universally preferred ratio of asymmetry. As the other famous Francis Bacon put it, “All beauty hath some strangeness in its proportion.”
The left side of the face, for example, is often more expressive—which is why it has more wrinkles—and “the fact that these facial asymmetries are present,” Zaidel noted, “tells us that the brain of the observer has to pick them up.”
Artists, indeed, might have sensed these imbalances in expressiveness, because European portrait painters from the Renaissance onward have shown a preference for the left profile, particularly among women.

Look at computer-generated portraits of perfectly symmetrical faces, and you feel a tickle in the brain, not unlike when you make the “flip” on an ambiguous figure reversal, like the Necker cube. “The symmetry is monster-like,” Zaidel suggested. “It does not exist in the real world.” The fusiform gyrus, that right-hemispheric area largely devoted to processing human faces, is finding its hypothesis about the world violated. “The brain is very specific about what it considers to be a face,” she said. “Your brain has to do a double take.” Probably, as with Bacon's work, before you even know what you are looking at. None of this explains why we might think of Bacon as a great artist—or an artist at all. We could simply look at severely disfigured faces in an obscure medical journal. But does the knowledge that his work may affect us,
at an irrevocably fundamental level, not add something to its appreciation, the way Smith began to like his Delacroix more by learning the life story of its subject? The pleasure of art and the pleasure of ice cream were different, the philosopher Rudolf Arnheim once protested. Might neuroscience not provide some way of demonstrating that was not merely a comforting fiction told by art lovers?

Later that day, Semir Zeki and I went out walking, to look at paintings. At the National Gallery, we looked at what is reputed to be Titian's self-portrait. “
He uses the technique where his face was partly turned away,” Zeki said. “There was a way in Venice in that time of showing some contempt, haughtiness, for everyone else. And the fact that the kind of half-turned-away face is a sign of contempt—and still is—tells you that our brains are similar in interpreting these sorts of things.”

Later, over lunch at the Garrick Club—a ridiculously genteel place where members, instead of sending each other text messages, leave notes in creamy white envelopes in the lobby—Zeki told me, “I believe there's a basic skeleton in the brain that allows you to experience beauty. What the characteristics of the objects are, I would leave that to the art historian.”

Art, rather than responding to or reflecting innate preferences, may actually succeed by tweaking them. Humans seem to overwhelmingly favor fractal patterns—those geometrically recurring forms seen in snowflakes and tree branches. The painter Jackson Pollock, as analysis revealed, generated fractal patterns in his monumental works of Abstract Expressionism. This alone would hardly explain his success as an artist, given the initial hostility to his work.

What is more interesting, however, as the physicist Richard Taylor and colleagues have found, is that Pollock's later, most developed, and eventually most famous work actually
departed
from the rather narrow bandwidth of the most preferred fractal patterns. As if, they suggested, he were trying to challenge or even confound viewers, pushing the limits of what they might like. After all, if the brain is primed for symmetry, what better way to capture attention than to violate it? With typefaces, we prefer fonts that are easier to read. No surprise there—we like fluency.
But research has shown that when you make fonts more
disfluent
,
harder to read, people seem to better remember the information conveyed by the words. Similarly, art that goes down too easy may be art that is easier to forget.

One critique of neuroaesthetics is that it is of little use if one could not, neurologically, distinguish between the urinal that Duchamp famously rendered as art and that same urinal in a hardware store. Of course, no one
else
could until Duchamp went ahead and claimed the distinction; he later admitted to being “horrified” that his readymades had come to be admired for “
their aesthetic beauty.” Art itself has not been very good at deciding what is art or what is good art. When Andy Warhol created his famous
Brillo Boxes
sculptures, as the critic Arthur Danto has noted, the more interesting question than whether they were art was why the actual Brillo box they faithfully replicated (designed by the artist James Harvey) was
not
. Those reasons, Danto notes, were “
nothing that meets the eye.” But they were certainly met somewhere in the brain. Perhaps, Danto averred, we could call them “commercial art.”

You can parse the validity of that distinction, but at least one experiment has found that when you ask people to look at images with an aesthetic goal in mind, versus simply to look at them, different brain activity is observed with each exercise. This implies that even if not everything is art, we can try to
look
at everything as if it were. (Warhol, after all, said, “
I just paint things I always thought were beautiful.”) What would make it art? Danto, famously, summed it up:
Art is what the art world says is art.
This flip tautology raises the question, returning to the Tuymans experiment: How much of our response to art is being told that something
is
art? (Answer: A
lot
.)

Most people, even devoted lovers of visual arts, do not visit galleries and museums as philosophers of art. They are not trying to place works in a tradition, to confirm that these works embody meaning, or to tick some box of whatever the reigning art discourse may be. They look at what catches their eye—metaphorically or actually. And they look longer at what they like.

Looking can be a pleasure in itself.
As Edward Vessel, a neuroscientist who directs New York University's ArtLab, told me in his office in lower Manhattan, he was surprised, during a study a few years ago with
his colleague Irving Biederman, to find that there seemed to be opioid receptors in the brain's ventral visual pathway. The region is normally involved in the recognition of forms. Opioids, up to that point, had been associated with synthetic opiates (like heroin) in “reward” or “pain” pathways. So what were they doing in this “visual” area? He wondered if they might “mediate” some connection between the way we form visual representations and meaning in the brain and the way we get pleasure. As you moved up in the brain's sensory processing hierarchies, he theorized, the rewards began to kick in.

From here, it seemed a natural progression to ask, what is it about a scene that we find pleasing? “If you and I share a preference,” he told me, “maybe it's because there's something out there in the world that is driving both of our preferences. Or maybe it's because our internal representations of the world are quite similar and it's those internal simulations that are really driving our preferences.”

In one study, Vessel and a colleague asked people to rate their preferences for real-world scenes and a set of “abstract stimuli.” People's preferences for real-world scenes were quite stable: Almost everyone liked an image of nature over an image of a parking lot. But when they were asked to describe how they felt about the abstract images, hardly anyone liked the same thing. Vessel suggests that with representational images, we are able to extract more “semantic meaning.” That meaning is what subjects agreed upon. With the abstract images, subjects (and their brains) had to sort it out for themselves—much like the hapless viewer in front of an unlabeled piece of nonrepresentational art—and so preferences diverged.

At the height of mid-century Abstract Expressionism, it was suggested that nonrepresentational art could function as a kind of “
universal language,” typically stripped as it was of cultural references. A number of studies have shown, however, that abstract art is a much tougher sell.
People seem to prefer things with meaning, and if you give them some context to the abstraction—even a title—they seem to like it more.

Neuroscience, far from reducing the mystery and power of our response to art, may only affirm them.
After people are shown photographs of faces, algorithms can now predict, at a level higher than chance, which face someone saw, purely based on the subject's neural signature. But such neural “decoders” only work for
that person
. Similarly,
the pattern of your neural response to a work of art will not be the same as mine; it may not even be the same as yours was a moment ago.

And yet, while our preferences may radically diverge, some of our brain activity may be strikingly similar, even while we are looking at different works. Vessel and his colleagues have done some tantalizingly suggestive research about what art—the art that we like—seems to do to us. In one study, while subjects were lying in a scanner (admittedly, not the way most of us view art), they were asked to rate, on a scale of one to four, how strongly they felt a painting moved them. As with any single scale, being “moved” may not be the appropriate measure, but it is a
response
. Subjects saw a wide range of artworks: new, old, familiar, unfamiliar. There was little similarity across what people rated. But something interesting happened to those people when they looked at paintings they rated a four. Those highest-rated paintings alone activated a set of regions, including the medial frontal cortex, known as the default mode network (DMN).

The DMN was discovered, more or less accidentally, in early brain-imaging studies. People were asked to do some task, and the brain's response was observed. In between tasks, however, researchers observed a number of regions that flickered to life as people, in essence, did nothing. The exact role of the DMN is unclear. It could be some sort of background monitoring system; it could be mind wandering. When people are asked to do things, the DMN is suppressed.