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A Face to Remember
Once dominated by correlational studies, face-perception research is moving into the realm of
experimentationand gaining tremendous insight.
By Kerry Grens | November 1, 2014
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THOMAS NORTHCUT/GETTY IMAGES
on Blackwell reclined in a hospital bed at Stanford University, bandages from his recent brain
surgery wrapped snugly around his head. Doctors had just removed a piece of his cranium,
implanted electrodes on the surface of his brain, and closed him back up. He waited for a seizure.
Blackwell, 49, had his first seizure when he was 11 and had experienced similar incidents periodically
thereafter. But after he turned 40, the seizures became more frequent. He wanted to feel secure when
caring for his two young children instead of worrying that he might have a seizure while bringing them to
the park. So in 2012 he gave doctors the OK to implant the electrodes, which were designed to pinpoint
the epicenter of his seizures as they came and help determine whether hed be good candidate for
surgery to remove the culprit tissue.
http://www.the-scientist.com/?articles.view/articleNo/41326/title/A-Face-to-Remember/
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Once such electrodes are in place, the wait for a seizure can take days, so to pass the time Blackwell
participated in some cognitive and perceptual tests. On occasion, researchers visited him and presented
him with tasks to perform on a computer, such as clicking a button when a particular object appeared on
a screen. After days with no sign of a seizure, Josef Parvizi, Blackwells neurologist, asked for permission
to stimulate the electrodes on Blackwells brain as part of an experiment. Blackwell agreed.
Curr
Parvizi instructed Blackwell to look at objects in the room: a Mylar balloon, a TV screen. The doctor
clicked a button, but nothing happened. Then he said, Look at my face, and he hit the button, and it
was the most bizarre thing, Blackwell recalls. In that brief moment when Parvizi zapped the electrodes,
Blackwell saw the doctors face metamorphose.
His face just sagged. His eyes drooped; his nose drooped and just shifted. It was very cartoonish, he
says. The face looked somewhat familiar, but it was no longer Parvizisuntil the doctor stopped
stimulating the electrodes on Blackwells brain. As soon as the stimulus was over, the familiar face of the
neurologist returned.
Wanting to learn more, Parvizi asked Blackwell some questions: Could he still tell that Parvizi was a
male? Oh, yeah, Blackwell replied. How did you know? asked Parvizi. Because youre still wearing a
suit and tie. Only your face changed. Everything else was the same. And with that, Blackwell gave to
science the best experimental evidence yet that humans have what researchers call a face-selective
area, or face patch for shorta region of the brain specialized for the perception of faces.1
View the
Previous human studies had relied on imaging techniques to link face perception to face patches by
association; none of them showed that disrupting the face patch could alter face perception. But more
recent research provides evidence that face patches, generally recognized as three chunks of the
temporal lobe, are critical to the everyday observation of faces. Blackwell is now just one of about a
dozen patients in whom Parvizi and Stanford colleague Kalanit Grill-Spector have demonstrated the
electrical disruption of face perception.
Subs
Its just so striking how specific the perceptual distortion is to the face, says Grill-Spector. This is why
its a very important discovery, because it shows the specificity of the cortical region to processing
faces.
Neuroscientists are now capitalizing on this specificity to unpack the fundamental computational
processes that go into identifying a face, a feat most of us perform without thought or effort. The face
patchesand even individual neurons in themappear to do different jobs, such as analyzing the
features of the face, responding to how the head is tilted, and, ultimately, determining someones
identity.
We meet thousands of individuals . . . and we can differentiate them, we can recognize them in different
conditions: when there are shadows on them, when the face is turned at different angles, if they get a
haircut, says Marlene Behrmann, a cognitive neuroscientist at Carnegie Mellon University. Its an
incredibly robust human ability.
Natural experiments
Page 2 of 10
11/13/14 1:33 PM
The neural stimulation Parvizi gave to Blackwell

caused momentary prosopagnosia?,? or face
blindnessan inability to discern the identity of a
person by his face. There are those who have had
prosopagnosia all their lives. Author and
neurologist Oliver Sacks famously documented his
own condition in his writings. In a 2010 New
Yorker article, Sacks wrote that he often cannot
recognize people he knows well; he has even
bumped into his own reflection, thinking it was
another bearded man walking toward him.2
Given face perceptions importance and ubiquity in
social interactions, prosopagnosia is a stressful
deficiency to live with. I avoid conferences,
parties, and large gatherings as much as I can,
knowing that they will lead to anxiety and
embarrassing situations, since I not only fail to
recognize people that I know well but tend also to
greet strangers as old friends, Sacks wrote. I am
much better at recognizing my neighbors dogs
(they have characteristic shapes and colors) than
my neighbors themselves.
MODIFIED FROM CREATIVEMARC/ISTOCKPHOTO.COM
To investigate the roots of such face blindness, Behrmann is studying the brains of prosopagnosia
patients. In one study, she asked such patients to view faces while they sat in a functional magnetic
resonance imaging (fMRI) scanner. Like people without the perceptual deficiency, the patients showed
normal activity in what Behrmann refers to as the core patches involved in face perception: the
fusiform face area (FFA), the occipital face area (OFA), and the superior temporal sulcus (STS). We
were perplexed, says Behrmann. We knew [the patients brains] were impaired, but we couldnt find
where.
So Behrmanns team, using an MRI approach called diffusion tensor imaging, embarked on a highresolution sleuthing trip through the hills and valleys of the brains morphology to find the differences.
The researchers found a reduction in two white-matter tractsthe myelin-wrapped axons connecting
neurons of different brain regionsbetween these core face patches in the back of the brain and
extended face-processing areas toward the front of the brain.3 (See illustration below.) The frayed
cabling suggests its a failure to propagate signals from the core to the extended regions, says
Behrmann. (For more information on the technique of diffusion tension imaging, see Whites the
Matter, The Scientist, November 2014.)
Intriguingly, the magnitude of the compromise
[in white matter] was correlated with the
magnitude of the prosopagnosia, says
Behrmann. It was a key piece of the puzzle.
More recently, the group has found, through fMRI
comparisons between people with and without
prosopagnosia, that activity in the extended
regions contacted by these white matter tracts is
indeed reduced during face perception.4 It
appears that the prosopagnosics face patches
operate well enough to know that a face is a face,
but the poor connection to the extended regions
prevents them from determining that persons
identity.
It appears that the

prosopagnosics face
patches operate well enough
to determine that a face is a
face, but the poor
connection to the extended
regions prevents them from
establishing that persons
Page 3 of 10
Also using fMRI, Ida Gobbini and James Haxby of

Dartmouth College have shown that the extended
regions are much more active when viewing
11/13/14 1:33 PM
identity.
familiar faces than unfamiliar ones.5 My

interpretation is, when you see someone familiar, you retrieve the person knowledge of that person,
says Gobbini. By recruiting extended brain regions such as the precuneus or amygdala, people may be
able to pull together the memories and emotions that go along with seeing someones face.
Something [cognitive neuroscience] is working really hard on is trying to understand the relative
contribution of these different areas, Behrmann says.
Manipulating humans
In September, Stanfords Parvizi and Grill-Spector
published data from electrode-stimulation
experiments on 10 participants, all of them with
medical situations like Blackwells who were
hoping to find a surgical solution to their maladies
and willing to lend their brains for study. Half the
patients had electrodes implanted on the fusiform
gyrus in their right cerebral hemispheres; the
other half, on the same area on the left side of
their brains. Although both areas were active
during face perception, only when Parvizi tickled
the right hemisphere face patch with electrical
stimulation did the patients report seeing an
altered face.6 Intriguingly, some patients also saw
faces that werent really there. One reported: The
black spot on the top of the TV shows some kind
of face expression. It looked like a human face,
then disappeared. Another patient had an
experience similar to Blackwells, in which the
doctors face morphed: It was almost like you
were a cat.
The results support the idea that face perception
is lateralized, which scientists had suspected since
the first documented cases of prosopagnosia in
patients with damage to the right-hemisphere
fusiform gyrus. Only the right side is important
for changing conscious perception of faces, says
Parvizi. We think the left side might be important
for retrieving names or anything language-related,
but its probably not doing the same thing as the
right hemisphere.
PATCHWORK: In the 1990s, researchers published data

identifying a face patch, dubbed the fusiform face
area (FFA), which lit up more in response to faces than
to other objects.1 Soon, other researchers identified
two additional face patches, the occipital face area
(OFA) and the superior temporal sulcus (STS).
Together, the FFA, OFA, and STS comprise what some
call the core regions of face perception.
See full infographic: JPG | PDF
CATHERINE DELPHIA
Parvizi says additional studies of this sort could help to determine how the patches are connected and
what jobs they perform, as well as the precise brain regions where the visual decoding involved in face
perception takes place. Intracranial recordings could also help resolve questions about the specificity of
the face patches and the role of neighboring cells. Of course, patients who require brain electrode
implantation and are willing to participate in such neuroscience studies are few and far between, making
it difficult to amass data. To achieve bigger sample sizes, Brad Duchaine of Dartmouth College and David
Pitcher of the National Institute of Mental Health have used transcranial magnetic stimulation (TMS) to
noninvasively excite brain regions of healthy volunteers. (See Brain Massage, The Scientist, November
2014.) A magnetic coil delivers short bursts of electrical stimulation, which interrupts normal brain
activity for about 20 minutes. By placing the coil close to a face patch, we can temporarily make you
bad at face perception, says Pitcher.
Page 4 of 10
11/13/14 1:33 PM
Two patchesthe OFA and the posterior STS (pSTS)lie at the surface of the brain. With fMRI, the
researchers can determine the precise location of the patches in an individual, then use TMS at a
particularly high frequency (called theta burst TMS, or TBS) to scramble the neurons normal function.
Volunteers then reenter the brain scanner to look at various images as the researchers observe the
activity in the patches.
A prevailing model has held that visual information about faces first comes into the OFA from the early
visual cortex, then branches out to the other face patches. According to this view, the OFA acts kind of
like a gatekeeper, says Pitcher. But his latest work shows that, although disrupting the OFA reduces
brain activity in the pSTS when the participants view photographs of faces, pSTS activity remains normal
when they view videos.7 These results suggest that facial information reaches more than just the OFA
initially and does not travel linearly through the patches. What people have been arguing more and
more so, and my paper is one of the first to show it experimentally, is that all the patches are connected
to each other, says Pitcher. The face patches all get information round about the same time and share
that information. (See illustration above.)
For all thats been gleaned from the human experiments, there is still much left to be discovered. Pitcher
points out that the neuronal connections between patches are still largely unmapped in humans, for
instance. Additionally, we dont have a good feel for how the division of labor is being set up among the
various face patches, says Duchaine. Such work would require invasiveness so far impossible in humans,
he notes, but research on face perception in other animals is beginning to yield clues.
Monkey business
Much of what scientists can only dream of doing in human brains, Doris Tsao and Winrich Freiwald have
accomplished in macaques. As a graduate student in Margaret Livingstones lab at Harvard, Tsao read
about the discovery of the FFA, finding it astonishing that there would be a region specialized for faces
when there are so many other objects humans have to identify. But the fMRI data that revealed the
existence of the FFA couldnt explain what the cells themselves were doing. To get at function on the
cellular level, Tsao says, it seemed easy to test in a monkey.
So, in the early 2000s, she teamed up with Freiwald, then a postdoc in Nancy Kanwishers lab at MIT.
The duo would insert electrodes into the brain region that responded to faces as the monkeys viewed a
slide show of a variety of objects and human faces. Freiwald, now at Rockefeller University, vividly
recalls those initial trials as the researchers guided electrodes slowly through the brain toward what they
call the middle face patch while pictures flashed before the monkey. A crackling sound heard over
speakers connected to the electrophysiology rig would alert the researchers to a neuron firing. On
occasion, a little rumbling would sound and then fade away. Then, just as a face popped up on the
screen, they heard a loud kkkrrrr, and made a note of it as a cell that likes faces.
Then they got another. Kkkrrr. And another. And then we realized, Wow, every time we stuck our
electrode into that patch we got face cell after face cell after face cell,? says Tsao.8 It was
tremendously exciting. It meant we could now have hope to understand the brains vocabulary for how
the brain codes objects.
The study also provided strong support for the specificity of face patches. The response to the other
objects the macaques viewed was much smaller, sometimes silence altogether, compared to the neural
activity triggered by faces. (See Just for Faces? at bottom.)
Freiwald and Tsao spent hours Photoshopping
faces for new experiments. To see whether
individual neurons respond to particular features
in the face, they created a simple cartoon in
which the parts could change independently of
one anotherthe eyebrows could be removed,
the eyes spaced far apart, and so on. They found
that within the same face patch, individual
'Pretty much everybody you

speak to has the total
intuition that face recognition
is seamless, rapid, effortless.
Page 5 of 10
neurons were tuned to respond more readily to

particular features, such as the eyes or hair, and
that many of the cells were especially keen on
facial geometry, such as the roundness of the
face, the size of the irises, or the space between
the eyes.9 Here you can have neighboring cells,
cells separated by microns, doing totally separate
things, says Tsao, who pursued this work as a
postdoc with Livingstone and, later, Roger Tootell
at Massachusetts General Hospital.
11/13/14 1:33 PM
And yet its probably the

most difficult problem the
visual system has to solve.
Marlene Behrmann,
Carnegie Mellon University
Jim DiCarlo and his colleagues at MIT have used optogenetics to actually turn off some of those neurons
and see what happens. In a recent study demonstrating the technique, they asked what would happen if
they silenced the neurons in and around a particular face patch in macaques that had been trained to
discriminate the gender of human faces. The fields hypothesis would be that silencing the face patch
should produce deficits [in discrimination], and silencing other tissue should affect [perception of
nonface] objects, says DiCarlo. Sure enough, with suppressed face-patch activity, the monkeys were
less able to distinguish men from women. Inhibition of neighboring regions, on the other hand, had no
such effect.10
DiCarlo says this study, presented by lead author Arash Afraz at the Vision Sciences Society meeting
earlier this year, is just the beginning of what optogenetics can bring to the study of object and face
perception. DiCarlos group is now conducting extensive face and object discrimination tests, with plans
to silence bits of neural tissue in one or more brain regions.
Tsao and Freiwalds studies have also supported the idea that faces are processed across a network of
patches. After working with the middle face patch in macaques, the researchers identified a number of
other regions active during face perception. They found that these regions are tightly coupled
anatomically; one patch communicates with multiple other patches, each of which appears to perform a
distinct task. In one region, for instance, the cells responded only to faces in particular orientationssay,
profile or straight-on. A different patch might respond to just one individual face, but will do so
regardless of the faces orientation.
Many consider Tsao and Freiwalds work the best evidence to date that face perception operates like an
orchestra, with units cooperating, communicating, and building upon one another to provide a
harmonious picture of facial identity. Ive learned more from one of their papers than from 10 to 20
human papers because you can get in there and record from single neurons, says Duchaine. They get
so much interesting evidence out of their recordings, it blows me away.
Points of view
Page 6 of 10
11/13/14 1:33 PM
Comprehending how the brain computes faces has

obvious implications for understanding
prosopagnosia. Working with Behrmann, Adrian
Nestor of the University of Toronto Scarborough
aims to build a face from the fMRI data of a
viewer. The researchers have built an algorithm
that attempts to link the objective pictorial
qualities of a face with the corresponding neural
code. Then, starting with the neural code, the
software can work backward to construct the face.
By analyzing the neural codes of healthy
participants, researchers could uncover new
insights into the computation of face perception,
and the scans of prosopagnosia patients may
prove even more interesting.
If we can approximate the face [seen by]
prosopagnosics, we can finally not only theorize
and hypothesize as to why they cannot recognize
faces at the individual level, we can see what they
see, says Nestor. Thats a very powerful
demonstration of whats wrongwhats not
functioning at the neural level.
MODIFIED FROM FRANCKREPORTER/ISTOCKPHOTO.COM
The early results show that the reconstructed

faces formed by the data from congenital prosopagnosics are remarkably alike. The neural patterns are
so similar, Nestor says. Were still trying to figure out why. In a person without face blindness, on the
other hand, the neural responses to viewing two different faces would be measurably different.
For Freiwald and many others, face perception
also offers a window into the mechanism
underlying our nature as social beings. Over the
last [few] years, I realized that faces are such
potent stimuli to get us into the emotional and
social brain in ways that escape cognitive control
in the first pass, he says. Take, for instance, the
way humans ooh and aah and feel warmth for
the face of a baby animal. Were just showing
pixels of colors, says Freiwald, but by the way
theyre arranged were getting into the emotional
brain.
Faces are such potent stimuli

to get us into the emotional
and social brain in ways that
escape cognitive control in
Winrich Freiwald,
the first pass.
Rockefeller University
Could this social nature be at the heart of how we so easily identify facesthree-dimensional objects that
follow the same basic pattern, yet carry so much significance individually? The face is what one goes by,
generally, Alice tells Humpty Dumpty in Lewis Carrolls Through the Looking Glass. Yet, as Humpty
Dumpty rightly responds, Your face is the same as everybody hasthe two eyes, so . . . nose in the
middle, mouth under.
Thats exactly why this is such an interesting domain to be in, says Behrmann. Pretty much everybody
you speak to has the total intuition that face recognition is seamless, rapid, effortless. And yet . . . its
probably the most difficult problem the visual system has to solve. Theres a real disconnect between our
introspection and the nature of the computation.
JUST FOR FACES?
Page 7 of 10
11/13/14 1:33 PM
While plenty of imaging data support the

involvement of several brain regions in
perceiving faces, there remains considerable
debate about each areas precise function,
especially regarding whether the neurons are
exclusively devoted to facial recognition. Isabel
Gauthier of Vanderbilt University makes the
argument that the patches may not be facespecific at all, but might simply reflect our
expertise in facial recognition.
She and her colleagues have asked experts in
recognizing particular objects, say, car
enthusiasts or bird watchers, to view faces,
cars, birds, and other items while in an fMRI
scanner. And over and over again, we find the THOMAS NORTHCUT/GETTY IMAGES
same relationship: the response to nonface
objects in the fusiform face area [FFA] predicts
your performance in recognizing objects, she says. In other words, experts use the FFA to
discriminate the objects of their expertise, just as they do when perceiving faces.1 Its not surprising
faces would activate this region, because were experts at [recognizing faces], Gauthier says.
Other researchers have countered Gauthiers argument with their own data. Earlier this year, Brad
Duchaine of Dartmouth College and his colleagues showed that prosopagnosics can still become
experts in identifying particular objects when theyre trained to do so.2 They were able to learn [the
task] even though they dont have [a fully functioning] FFA, says David Pitcher, a research fellow at
the National Institute of Mental Health who was a coauthor of the study. If the FFA is an expertise
brain region, the prosopagnosics should have failed to learn, just as they cannot be trained to
recognize faces.
But others have also raised doubts about the idea that face patches are solely responsible for
processing faces. Its also possible that neurons neighboring the face patches contribute to detecting
someones identity. To get to the bottom of face perception, Jim DiCarlo and Arash Afraz of MIT and
other researchers are moving the field from one of observation and correlation to one of
experimentation and causation. If we can do causality studies, I think we will find [the so-called
face] neurons are doing other things not face-related, he says. So far though, their data seem to
support the idea that face patches are specialized for recognizing faces, DiCarlo notes. That
hypothesis is too simple, but we dont have evidence to refute it. So it might be true.
1. I. Gauthier et al., Expertise for cars and birds recruits brain areas involved in face
recognition, Nat Neurosci, 3:191-97, 2000.
2. C. Rezlescu et al., Normal acquisition of expertise with greebles in two cases of acquired
prosopagnosia, PNAS, 111:5123-28, 2014.
References
1. J. Parvizi et al., Electrical stimulation of human fusiform face-selective regions distorts face
perception, J Neurosci, 32:14915-20, 2012.
2. O. Sacks, Face-blind, The New Yorker, August 30, 2010.
3. C. Thomas et al., Reduced structural connectivity in ventral visual cortex in congenital
prosopagnosia, Nat Neurosci, 12:29-31, 2008.
4. G. Avidan et al., Selective dissociation between core and extended regions of the face processing
network in congenital prosopagnosia, Cereb Cortex, doi:10.1093/cercor/bht007, 2013.
5. M.I. Gobbini, J.V. Haxby, Neural systems for recognition of familiar faces, Neuropsychologia,
45:32-41, 2007.
6. V. Rangarajan et al., Electrical stimulation of the left and right human fusiform gyrus causes
Page 8 of 10
11/13/14 1:33 PM
different effects in conscious face perception, J Neurosci, 34:12828-36, 2014.

7. D. Pitcher et al., Combined TMS and fMRI reveal dissociable cortical pathways for dynamic and
static face perception, Curr Biol, 22:2066-70, 2014.
8. D.Y. Tsao et al., A cortical region consisting entirely of face-selective cells, Science, 311:670-74,
2006.
9. W.A. Freiwald et al., A face feature space in the macaque temporal lobe, Nature Neurosci,
12:1187-96, 2009.
10. A. Afraz et al., Optogenetic and pharmacological suppression of face-selective neurons reveal
their causal role in face discrimination behavior, J Vision, 14:600, 2014.
Tags
white matter, neuroscience, faces, face perception, brain map, brain anatomy and brain
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Comments
November 4, 2014
Dr. S
Posts: 1
This is all very interesting--that there is a section of the brain specific for
face recognition. But, although scientists have narrowed the search, the
results say nothing about how we learn to recognize faces. And, I doubt
that face recognition is so special. After all, we learn to recognize slight
and subtle differences between an almost infinite number of other stimuli,
not just faces. Faces get a lot of attention because of their social value,
but I doubt that they are special otherwise.
Also, cognitive science offers no cogent theory of face perception. And
saying that the brain perceives or organizes is misleading. To perceive is
to behave and individual organisms, not brains, behave. The brain is
programmed by a combination of genes and learning experiences, with
the latter surely more important, especially in humans. When we first
come into the world there is little to no perception (i.e., reacting
differentially to stimuli) of any kind; we have to learn it. And we do so
thorugh operant conditioning from interactions with an ever increasing
complex environment. In fact, no other theory besides an operant one can
explain why we cannot recognize someone when seeing them but can do
so when we hear their voice, or see some other visual stimulus associated
with that person. This is because we have been reinforced to call people
by name in the presence of a variety of different stimuli associated with
them, faces being only one. I'm sure neuroscientists could look for the
neurons that mediate voice recognition too. All of these efforts are a little
like a more sophisticated phrenology. We may ultimately be able to point
to exact locations in the brain responsible for mediating thousands of
interesting behaviors; but that will tell us nothing about the genesis or
function of those behaviors.
Neurophysiology without an experimentally based theory of behavior--not
Page 9 of 10
11/13/14 1:33 PM
mind--will only tell a very small part of the story. The decade of the brain
has surely eclipsed the decade of behavior.
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