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Chapter 3:
Neural Processing and
Perception

Neural Processing and Perception

Car B Car A
The way electrical signals travel through the nervous system is more like Car B’s
journey. The pathway from receptors to brain is not a nonstop turnpike. Every signal
leaving a receptor travels through a complex network of interconnected neurons, often
meeting, and being affected by, other signals along the way.

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Neural processing is the interaction of signals in many

neurons.

Lateral Inhibition and Perception

• Experiments with eye of Limulus

– Ommatidia allow recordings
from a single receptor.
– Light shown into a single
receptor leads to rapid firing
rate of nerve fiber.
– Adding light into neighboring A Limulus, or horseshoe crab. Its
receptors leads to reduced large eyes are made up of
firing rate of initial nerve fiber. hundreds of ommatidia, each
containing a single receptor.

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A demonstration of lateral
inhibition in the Limulus.
The records show the
response recorded by the
electrode in the nerve fiber
of receptor A: (a) when only
receptor A is stimulated;
(b) when receptor A and the
receptors at B are stimulated
together;
(c) when A and B are
stimulated, with B at an
increased intensity.
Figure 3-3 p55

Lateral Inhibition and Lightness

Perception

• Three lightness perception phenomena

explained by lateral inhibition
– The Hermann Grid: Seeing spots at an
intersection
– Mach Bands: Seeing borders more sharply
– Simultaneous Contrast: Seeing areas of
different brightness due to adjacent areas

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Hermann Grid

• People see an illusion of gray

images in intersections of
white areas.
• Signals from bipolar cells
cause effect
– Receptors responding to
white corridors send
inhibiting signals to receptor
at the intersection
– The lateral inhibition causes
a reduced response which The Hermann grid. Notice the
leads to the perception of gray “ghost images” at the
gray. intersections of the white areas,
which decrease or vanish when
you look directly at an
intersection.

(a) Four squares of the Hermann grid, showing

five of the receptors under the pattern.
Receptor A is located at the intersection, The bipolar cells from the circuit in
and B, C, D, and E have a black square on Figure 3.5. Each bipolar cell has
either side. an initial response of 100. Bipolar
(b) Perspective view of the grid and five cells B, C, D, and E each send 10
receptors, showing how the receptors units of inhibition to bipolar cell A,
(green) connect to bipolar cells (blue). The as indicated by the red arrows.
response of all five receptors is 100. The
Because the total inhibition is 40,
initial response of the bipolars matches the
the final response of bipolar A is
response of the receptors. Lateral inhibition
60.
travels to bipolar cell A along the red
pathways
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(a) Four squares of the Hermann grid, as in

Figure 3.5, but now focusing on receptor D,
which is flanked by two black squares.
Receptor D is surrounded by receptors A, F,
G, and H. Notice that receptors F and H are The bipolar cells from the circuit in
located under the two black squares, so Figure 3.7. Bipolars A and G have an
they receive less light than the other initial response of 100, and F and G
receptors. (b) Perspective view of the grid have an initial response of 20. Bipolars A
and five receptors, showing how the and G each send 10 units of inhibition to
receptors (green) connect to bipolar cells bipolar cell D; Bipolars F and H each
(blue). The response of receptors A and G is send 2 units of inhibition to D. The total
100, and the response of F and H is 20. inhibition is 24, so the final response of
Lateral inhibition travels to bipolar D along
bipolar D is 76.
the red pathways.

Mach Bands

• People see an illusion of enhanced lightness

and darkness at borders of light and dark
areas.
– Actual physical intensities indicate that this
is not in the stimulus itself.
– Receptors responding to low intensity
(dark) area have smallest output.
– Receptors responding to high intensity
(light) area have largest output.

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Shadow-casting technique
for observing Mach bands.
Illuminate a light-colored
surface with your desk lamp
and cast a shadow with a
piece of paper.

Mach bands at a contour between light and dark.

(a) Just to the left of the contour, near B, a faint
light band can be perceived, and just to the right
at C, a faint dark band can be perceived.
(b) The physical intensity distribution of the light,
as measured with a light meter.
(c) A plot showing the perceptual effect described
in (a). The bump in the curve at B indicates the
light Mach band, and the dip in the curve at C
indicates the dark Mach band. The bumps that
represent our perception of the bands are not
present in the physical intensity distribution.

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Mach Bands - continued

– All receptors are receiving lateral Mach bands at a contour

between light and dark.
inhibition from neighbors (a) Just to the left of the
– In low and high intensity areas contour, near B, a faint
light band can be
amount of inhibition is equal for all perceived, and just to the
receptors right at C, a faint dark
band can be perceived. (b)
– Receptors on the border receive The physical intensity
differential inhibition distribution of the light, as
– On the low intensity side, there is measured with a light
meter. (c) A plot showing
additional inhibition resulting in an the perceptual effect
enhanced dark band. described in (a). The bump
in the curve at B indicates
– On the high intensity side, there is the light Mach band, and
less inhibition resulting in an the dip in the curve at C
indicates the dark Mach
enhanced light band. band. The bumps that
– The resulting perception gives a represent our perception of
boost for detecting contours of the bands are not present
in the physical intensity
objects. distribution.

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Mach Bands

Table for determining the final output of the bipolar The final receptor output calculated
cells in Figure 3.11, by starting with the initial for the circuit in Figure 3.11. The
response and subtracting inhibition coming from bump at B and the dip at C
the left and right. correspond to the light and dark
Mach bands, respectively.

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Lateral Inhibition and Simultaneous

Contrast

• People see an illusion of changed brightness

or color due to effect of adjacent area
– An area that is of the same physical
intensity appears:
• lighter when surrounded by a dark area.
• darker when surrounded by a light area.

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Lateral Inhibition and Simultaneous

Contrast - continued
– Receptors stimulated by bright
surrounding area send a large
amount of inhibition to cells in
center.
– Resulting perception is of a darker
area than when this stimulus is
viewed alone.
– Receptors stimulated by dark Simultaneous contrast. The two
surrounding area send a small center squares reflect the same
amount of light into your eyes
amount of inhibition to cells in but look different because of
center. simultaneous contrast.
– Resulting perception is of a lighter
area than when this stimulus
viewed alone.

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How lateral inhibition has been used to explain the simultaneous contrast effect. The
size of the arrows indicates the amount of lateral inhibition. Because the square on
the left receives more inhibition, it appears darker.

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A Display That Can’t Be Explained by

Lateral Inhibition

• White’s Illusion
– People see light and dark rectangles even
though lateral inhibition would result in the
opposite effect.

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Figure 3.16 White’s illusion. The rectangles at A and B appear different, even though
they are printed from the same ink and reflect the same amount of light.

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Figure 3.17 When you mask off part of the White’s illusion display, as shown here, you
can see that rectangles A and B are actually the same. (Try it!)
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Figure 3.18 The arrows indicate the amount of lateral inhibition received by parts of
rectangles A and B. Because the part of rectangle B is surrounded by more white, it
receives more lateral inhibition. This would predict that B should appear darker than A
(as in the simultaneous contrast display in Figure 3.14), but the opposite happens. This
means that lateral inhibition cannot explain our perception of White’s illusion.

Figure 3-18 p60

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Explanation of White’s Illusion

• Belongingness
– An area’s appearance is affected by where
we perceive it belongs.
– Effect probably occurs in cortex rather than
retina.
– Exact physiological mechanism is
unknown.

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Processing From Retina to Visual

Cortex and Beyond

• Area of receptors that affects

firing rate of a given neuron in
the circuit
• Receptive fields are
determined by monitoring
single cell responses.
• Research example for vision
– Stimulus is presented to
retina and response of cell The optic nerve, which
is measured by an leaves the back of the eye,
contains about one million
electrode. optic nerve fibers in the
human.
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Receptive field
• Hartline (1938, 1940) isolated a single fiber in the optic nerve by teasing
apart the optic nerve. While recording from this teased-out fiber, Hartline
illuminated different areas of the retina and found that the fiber he was
recording from responded only when a small area of the retina was
illuminated. He called the area that caused the neuron to fire the nerve
fiber’s receptive field.
a) Hartline’s experiment in which he
determined which area of a frog’s
retina caused firing in a single optic
nerve fiber. This area is called the
receptive field of that optic nerve
fiber.
b) Receptive fields of three optic
nerve fibers. These receptive fields
overlap, so stimulating at a
particular point on the retina will
generally activate a number of
fibers in the optic nerve.

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Receptive field
• The region of the retina that must receive illumination in order to obtain
a response in any given fiber. (Hartline, 1938, p. 410

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Receptive field
• Researchers studying the responding of optic nerve fibers in the cat discovered a
property of receptive fields. The cat receptive fields are arranged in a center-
surround organization
• The area in the “center” of the receptive field responds differently to light than the
area in the “surround” of the receptive fi eld (Barlow et al., 1957; Hubel & Wiesel,
1965; Kuffler, 1953).

Excitatory-center, Inhibitory-center,
inhibitory- excitatory
surround surround
receptive field. receptive field.

Center-surround receptive fields. (a) Excitatory center, inhibitory surround; (b)

inhibitory center, excitatory surround.
Figure 3-21 p61

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Receptive field
• The discovery that receptive fields can have oppositely responding areas made it
necessary to modify Hartline’s definition of receptive field to the retinal region
over which a cell in the visual system can be influenced (excited or inhibited)
by light (Hubel & Wiesel, 1961). The word “influences” and reference to
excitation and inhibition make it clear that any change in firing—either an increase
or a decrease—needs to be taken into account in determining a neuron’s receptive
field.

Excitatory-center, Inhibitory-center,
inhibitory- excitatory
surround surround
receptive field. receptive field.

Center-surround receptive fields. (a) Excitatory center, inhibitory surround; (b)

inhibitory center, excitatory surround.
Figure 3-21 p61

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Center-Surround Antagonism
• The discovery of center-surround receptive fields was also
important because it showed that neural processing could result in
neurons that respond best to specific patterns of illumination. This is
illustrated by an effect called center-surround antagonism,

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Center-Surround Antagonism
• Output of center-surround receptive fields changes depending on
area stimulated:
– Highest response when only the excitatory area is stimulated
– Lowest response when only the inhibitory area is stimulated
– Intermediate responses when both areas are stimulated

Response of an excitatory-center, inhibitory-surround receptive field as stimulus size is

increased. Shading indicates the area stimulated with light. The response to the stimulus
is indicated below each receptive field. The largest response occurs when the entire
excitatory area is illuminated, as in (b). Increasing stimulus size further causes a
decrease in firing due to center-surround antagonism.

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Center-Surround Antagonism
We can explain center-surround receptive fi elds and center-surround antagonism in
terms of neural processing by describing the operation of a neural circuit, which, as you
will recall from Chapter 2, is a group of interconnected neurons. These neurons, working
together, help create the excitatory center, inhibitory-surround receptive field of neuron
B. Center-surround receptive fi elds are created by the interplay between excitation and
inhibition.

A seven-receptor neural circuit underlying a center- surround receptive field.

Receptors 3, 4, and 5 are in the excitatory center, and receptors 1, 2, 6, and 7
are in the inhibitory surround.
Figure 3-23 p62

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Hubel and Wiesel’s Rational for

Studying Receptive Fields
• How neurons at higher levels of the
visual system become tuned to
respond best to more and more
specific kinds of stimuli. To do this,
Hubel and Wiesel modified Hartline’s
procedure for presenting light to the
retina. Instead of shining light directly
into the animal’s eye, Hubel and
Wiesel had animals look at a screen
on which they projected stimuli.

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Hubel and Wiesel’s Rational for

Studying Receptive Fields

• Signals from the retina travel

through the optic nerve to the
– Lateral geniculate nucleus
(LGN)
– Primary visual receiving area
in the occipital lobe (the striate
cortex or area V1)
– And then through two
pathways to the temporal lobe
and the parietal lobe
– Finally arriving at the frontal
lobe
Recording electrical signals from a fiber
in the optic nerve of an anesthetized cat.
Each point on the screen corresponds to
a point on the cat’s retina.

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Hubel and Wiesel’s Rational for

Studying Receptive Fields

• An important thing to remember

about receptive fields, which is
always true no matter what
method is used, is that the
receptive field is always on the
retina.
• It doesn’t matter where the neuron
is—the neuron can be in the
retina, the visual cortex, or
elsewhere in the brain, but the
receptive field is always on the
retina, because that is where the
stimuli are received. Recording electrical signals from a fiber
in the optic nerve of an anesthetized cat.
Each point on the screen corresponds to
a point on the cat’s retina.

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The Visual System

(a) Side view of the visual system,
showing the major sites along the
primary visual pathway where
processing takes place: the eye, the
optic nerve, the lateral geniculate
nucleus, and the visual receiving
area of the cortex.
(b) Visual system seen from underneath
the brain, showing the superior
colliculus, which receives some of
the signals from the eye.

Figure 3-25 p63

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Hubel and Wiesel: LGN

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Hubel and Wiesel: LGN

• Recording from the LGN, receptive fields were similar to that of
ganglion cells.
• The fact that little change occurred in receptive fields when
moving from the optic nerve fibers to neurons in the LGN

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Hubel and Wiesel: LGN

• Major function of LGN is to
regulate neural information
from the retina to the visual
cortex.
– Signals are received from
the retina, the cortex,
the brain stem, and the
thalamus.
– Signals are organized by
eye, receptor type, and
type of environmental
information.
Information flow into and out of the LGN.
The sizes of the arrows indicate the sizes
of the signals
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Hubel and Wiesel: Receptive Fields of

Neurons in the Visual Cortex
• Neurons that fire to specific
features of a stimulus
• Pathway away from retina
shows neurons that fire to
more complex stimuli
• Cells that are feature
detectors:
– Simple cortical cell
Figure 3.28 When Hubel and Wiesel
– Complex cortical cell dropped a slide into their slide projector,
– End-stopped cortical cell the image of the edge of the slide moving
down unexpectedly triggered activity in a
cortical neuron

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Hubel and Wiesel: Receptive Fields of

Neurons in the Visual Cortex

The relationship between orientation and fi

ring is indicated by a neuron’s orientation
tuning curve, which is determined by
measuring the responses of a simple
cortical cell to bars with different
orientations.

Figure 3-27 p65

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Hubel and Wiesel: Receptive Fields of

Neurons in the Visual Cortex

• Hubel and Wiesel found cells in the

striate cortex with receptive fields
that, like center-surround receptive
fields of neurons in the retina and
LGN, have excitatory and inhibitory
areas. However, these areas are
arranged side by side rather than in
the center-surround configuration.
• Cells with these side-by-side
receptive fields are called simple
cortical cells.

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Hubel and Wiesel: Receptive Fields of

Neurons in the Visual Cortex

Response of a
complex cell recorded
from the visual cortex
of the cat. The stimulus
bar is moved back and
forth across the
receptive field. The cell
fires best when the bar
is positioned with a
specific orientation and
is moved in a specific
direction.

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Hubel and Wiesel: Receptive Fields of

Neurons in the Visual Cortex

• End-stopped cells
• Response of an
end-stopped cell
recorded from the
visual cortex of the
cat.
• The stimulus is
indicated by the
light area on the left.
This cell responds
best to a medium-
sized corner that is
moving up.

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Table 3-1 p66

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Do Feature Detectors Play a Role in

Perception?
• Neural processing endows neurons with properties that make them feature
detectors that respond best to a specific type of stimulus. When
researchers show that neurons respond to oriented lines, they are
measuring the stimulus–physiology relationship.

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Do Feature Detectors Play a Role in

Perception?
• But just measuring this relationship does not prove that these neurons
have anything to do with the perception of oriented lines. To demonstrate a
link between physiology and perception, it is necessary to measure the
physiology–perception relationship. One way this has been accomplished
is by using a psychophysical procedure called selective adaptation.

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Selective Adaptation

• Neurons tuned to specific stimuli fatigue

when exposure is long.
• Fatigue or adaptation to stimulus causes
– Neural firing rate to decrease
– Neuron to fire less when stimulus
immediately presented again
• Selective means that only those neurons that
respond to the specific stimulus adapt.

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Selective Adaptation - continued

• Gratings are used as stimuli

– Made of alternating light
and dark bars
– Angle relative to vertical
can be changed to test for
sensitivity to orientation
– Difference in intensity can
be changed to test for
sensitivity to contrast

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Selective Adaptation - continued

• Measure sensitivity to range of one stimulus

characteristic
• Adapt to that characteristic by extended
exposure
• Re-measure the sensitivity to range of the
stimulus characteristic

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Procedure for carrying out a selective adaptation experiment.

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Selective Adaptation - continued

A grating’s contrast threshold is the minimum intensity difference between

two adjacent bars that can just be detected. The contrast threshold for seeing
a grating is measured by changing the intensity difference between the light
and dark bars until the bars can just barely be seen.

Figure 3-32 p67

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Selective Adaptation - continued

• Measure contrast threshold by decreasing

intensity of grating until person can just see it.
• Calculate the contrast sensitivity by taking
1/threshold.
• If threshold is low, person has high contrast
sensitivity.

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Figure 3.33
(a) Results of a psychophysical selective adaptation experiment. This graph shows
that the person’s adaptation to the vertical grating causes a large decrease in her
ability to detect the vertical grating when it is presented again but has less effect on
gratings that are tilted to either side of the vertical.
(b) Orientation tuning curve of the simple cortical neuron from Figure 3.27.
Figure 3-33 p68

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Hubel and Wiesel: Receptive Fields of

Neurons in the Visual Cortex

The relationship between orientation and fi

ring is indicated by a neuron’s orientation
tuning curve, which is determined by
measuring the responses of a simple
cortical cell to bars with different
orientations.

Figure 3-27 p65

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The psychophysical curve is slightly wider because the adapting stimulus

affects some neurons that respond to orientations near the adapting
orientation.

Figure 3-33 p68

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Selective Rearing

• Further evidence that feature detectors are involved in

perception is provided by selective rearing experiments.
• The idea behind selective rearing is that if an animal is reared in
an environment that contains only certain types of stimuli, then
neurons that respond to these stimuli will become more
prevalent.
• This follows from a phenomenon called neural plasticity or
experience-dependent plasticity—the idea that the response
properties of neurons can be shaped by perceptual experience.
• According to this idea, rearing an animal in an environment that
contains only vertical lines should result in the animal’s visual
system having neurons that respond predominantly to verticals.

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Selective Rearing

• Blakemore and Cooper (1970) showed this by rearing

kittens in tubes with either horizontal or vertical lines.
• Both behavioral and neural responses showed the
development of neurons for the environmental
stimuli.
• One way to describe the results of selective rearing
experiments is “Use it or lose it.”

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Selective Rearing

• Striped tube used in

Blakemore and Cooper’s
(1970) selective rearing
experiments.
• The kittens were kept in the
dark from birth to 2 weeks of
age, at which time they were
placed in the tube for 5 hours
a day; the rest of the time
they remained in the dark.

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Selective Rearing
• When the kittens’ behavior was
tested after 5 months of selective
rearing, they seemed blind to the
orientations that they hadn’t seen in
the tube
• Each line indicates the orientation
preferred by a single neuron in the
cat’s cortex.
• This cat, which was reared in a
vertical environment, has many
neurons that respond best to
vertical or near-vertical stimuli, but
none that respond to horizontal
stimuli. The horizontally responding
neurons were apparently lost
because they hadn’t been used.
The opposite result occurred for the
horizontally reared cats.

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Selective Rearing

• This result may seem to contradict the results

of the selective adaptation experiment just
described, in which exposure to verticals
decreases the response to verticals.
However, adaptation is a short-term effect.
Presenting the adapting orientation for a few
minutes decreases responding to that
orientation. In contrast, selective rearing is a
longer-term effect.

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Higher Level Neurons

• The idea that perception can be explained in terms of

feature detectors that respond to straight lines or corners
was popular in the 1970s because, as anyone who has
played with building blocks or Legos knows, many objects
can be created from rectangular shapes.
• Researchers continued to study feature detectors in the
striate cortex and nearby areas, vision researchers were
beginning to pay attention to brain areas far outside of the
striate cortex.

• Inferotemporal (IT) cortex

• Prosopagnosia
• Fusiform face area

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Higher Level Neurons

One of these researchers was psychologist Charles Gross, who decided
that the inferotemporal (IT) cortex in the temporal lobe was ripe for study.
Removing parts of the IT cortex in monkeys affected the monkeys’ ability to
recognize objects, as well as on research on a human condition called
prosopagnosia, in which people with temporal lobe damage were unable to
recognize faces.

Figure 3.35 (a) Location of the inferotemporal (IT) cortex in the monkey. (b)
Location of the fusiform face area (FFA) in the human, just under the temporal
lobe. Both of these areas are rich in neurons that respond to faces.
Figure 3-35 p69

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Higher Level Neurons

• The discovery that neurons in the IT cortex respond to complex stimuli came a few
days into one of their experiments, when they found a neuron that refused to respond
to any of the standard stimuli like oriented lines or circles or squares.
• Nothing worked, until one of the experimenters pointed at something in the room,
casting a shadow of his hand on the screen. When this hand shadow caused a burst
of firing, the experimenters knew they were on to something and began testing the
neuron with a variety of stimuli, including cutouts of a monkey’s hand. After a great
deal of testing, they determined that this neuron responded to a handlike shape with
fingers pointing up (Rocha-Miranda, 2011; also see Gross, 2002). After expanding
the types of stimuli presented, they also found some neurons that responded best to
faces.

Figure 3.36 Some of the shapes used by Gross et al. (1972) to study the responses of
neurons in the monkey’s inferotemporal cortex. The shapes are arranged in order of
their ability to cause the neuron to fire, from none (1) to little (2 and 3) to maximum (6).

Figure 3-36 p70

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Higher Level Neurons

Finally, in the 1980s, other experimenters began recording from neurons in
the IT cortex of the monkey that responded to faces and other complex
objects (Rolls, 1981; Perrett et al., 1982), and in the 1990s, researchers
discovered an area on the underside of the temporal lobe of the human
cortex that was named the fusiform face area because it responded
strongly to faces (Kanwisher et al., 1997; McCarthy et al., 1997) (Figure
3.35b).

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The Sensory Code

• Sensory code - representation of
perceived objects through neural
firing
– Specificity coding - specific
neurons responding to
specific stimuli
• Leads to the
“grandmother cell”
hypothesis
• Recent research shows
cells in the hippocampus
that respond to concepts
such as Halle Berry. Figure 3.37 Specificity coding, in which
each face causes a different neuron to fire.

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The Sensory Code

• Problems with specificity coding:
• Too many different stimuli to
assign specific neurons
• Most neurons respond to a
number of different stimuli.

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The Sensory Code continued

• Distributed coding -
pattern of firing across
many neurons codes
specific objects
– Large number of
stimuli can be coded
by a few neurons.

Figure 3.38 Distributed coding, in which the

face’s identity is indicated by the pattern of
firing of a large number of neurons.

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Sensory Code The Sensory Code -

continued
• How many neurons are
needed for an object in
distributed coding?
– Sparse coding - only a
relatively small number
of neurons are necessary
• This theory can be
viewed as a midpoint
between specificity
and distributed
coding.
Figure 3.39 Sparse coding, in which the face’s
identity is indicated by the pattern of firing of a
small group of neurons.

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Sensory Code The Sensory Code -

continued
• Recently, neurons were discovered when
recording from the temporal lobe of
patients undergoing brain surgery for
epilepsy. (Stimulating and recording from
neurons is a common procedure before
and during brain surgery, because it
makes it possible to determine the exact
layout of a particular person’s brain.)

• These neurons responded to very specific

stimuli. Figure 3.40 shows the records for
a neuron that responded to pictures of the
actor Steve Carell and not to other
people’s faces (Quiroga et al., 2008).
• Given the likelihood that even these
special neurons are likely to fi re to more Figure 3.40 Records from a neuron in the
than onestimulus, Quiroga and coworkers temporal lobe that responded to different views
of Steve Carell (top records) but did not respond
(2008) suggested that their neurons are
to pictures of other well-known people (bottom
probably an example of sparse coding. records).

Figure 3-40 p72

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Sensory Code The Sensory Code -

continued
• How neural firing can represent various features in the
environment?
• We can state that part of the answer is that features or objects
are represented by the pattern of firing of groups of neurons.
• Sometimes the groups are small (sparse coding), sometimes
large (distributed coding).
• As we will see in the next chapter, another part of the answer
involves considering how neurons in sensory systems are
organized.

Figure 3-40 p72

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The Mind-body Problem

• How do physiological processes become

transformed into perceptual experience?
– Easy problem of consciousness
• Neural correlate of consciousness
(NCC) - how physiological responses
correlate with experience
– Hard problem of consciousness
• How do physiological responses cause
experience?

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The Mind-body Problem

Figure 3.41
(a) Solving the “easy” problem of
consciousness involves
looking for connections
between physiological
responding and experiences
such as perceiving “Susan’s
face” or “red.” This is also
called the search for the
neural correlate of
consciousness.

(b) Solving the “hard” problem of

consciousness involves
determining how physiological
processes, such as ions
flowing across the nerve
membrane, cause us to have
experiences. Figure 3-41 p73

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