THE VISUAL SYSTEM Wavelength and intensity are two properties

of light that are of particular interest.

Chapter Overview • Wavelength because it plays a role in
the perception of color
The visual system does not produce an • Intensity because it plays an important
accurate internal copy of the external world. role in the perception of brightness.
It does much more.
In everyday language, the concepts of
The visual system creates an accurate, richly wavelength and color are often used
detailed, three-dimensional perception that is interchangeably, as are intensity and
even better than the external reality from brightness.
which it was created.
Pupil and Lens
LIGHT ENTERS THE EYE AND
REACHES THE RETINA The amount of light reaching the retinas is
regulated by the donut-shaped bands of
Some animals have special adaptations that contractile tissue, the irises, which give our
allow them to see under very dim illumination, eyes their characteristic color.
but no animal can see in complete darkness. • Light enters the eye through the pupil,
the hole in the iris.
The light reflected into your eyes from
the objects around you is the basis for your The adjustment of pupil size in response to
ability to see them. If there is no light, there is changes in illumination represents a
no vision. compromise between sensitivity (the ability to
detect the presence of dimly lit objects) and
Light can be thought of in two different ways: acuity (the ability to see the details of objects).
as discrete particles of energy, called photons
or as waves of energy. When the level of illumination is high and
• Both theories are useful as in some sensitivity is not important, the pupils
ways, light behaves like particles; and constrict the regulate light passing. Images
in others, it behaves like waves. become sharper and there is a greater
depth of focus.
Light is sometimes defined as waves of
electromagnetic energy between 380 and 760 However, when the level of illumination is
nanometers in length. too low to adequately activate the
receptors, the pupils dilate to let in more
light, thereby sacrificing acuity and depth
of focus.

Behind each pupil is a lens, which focuses

incoming light on the retina.

When we direct our gaze at something near,

the lens is adjusted by the ciliary muscles, and
assumes its natural cylindrical shape.
• This increases the ability of the lens to
refract (bend) light and thus brings
close objects into sharp focus.
There is nothing special about these wavelengths
But, when we focus on a distant object, the
except that the human visual system responds to
lens is flattened.
them. In fact, some animals can see wavelengths
that we cannot. For example, rattlesnakes can see The process of adjusting the configuration
infrared waves. of the lenses to bring images into focus on
the retina is called accommodation.
 THE RETINA AND TRANSLATION
OF LIGHT INTO NEURAL SIGNALS
After light passes through the pupil and the
lens, it reaches the retina.
The retina converts light to neural signals,
conducts them toward the CNS, and
participates in the processing of the signals.

STRUCTURE OF THE RETINA

Eye Position and Binocular Disparity

One reason vertebrates have two eyes is that

vertebrates have two sides: left and right.

By having one eye on each side, which is by far

the most common arrangement, vertebrates
can see in almost every direction without
moving their heads.

This arrangement sacrifices the ability to see

behind so that what is in front can be viewed
through both eyes simultaneously.

Such arrangement is an important basis for The retina is composed of five different types
our visual system’s ability to create three- of neurons: receptors, horizontal cells, bipolar
dimensional perceptions (to see depth) cells, amacrine cells, and retinal ganglion cells.
from two-dimensional retinal images.
The amacrine cells and the horizontal cells
are specialized for lateral communication
The movements of your eyes are coordinated
(communication across the major channels
so each point in your visual world is projected
of sensory input).
to corresponding points on your two retinas.
Retinal neurons communicate both
To accomplish this, your eyes must chemically via synapses and electrically via
converge (turning inward); convergence is gap junctions.
greatest when you are inspecting things
that are close. The retina is in a sense inside-out:
Light reaches the receptor layer only after
But the positions of the images on your two
passing through the other layers.
retinas can never correspond exactly because
your two eyes do not view the world from
Then, once the receptors have been activated,
exactly the same position.
the neural message is transmitted back out
Binocular disparity—the difference in the through the retinal layers to the retinal
position of the same image on the two ganglion cells, whose axons project across the
retinas—is greater for close objects than outside of the retina before gathering together
for distant objects. in a bundle and exiting the eyeball.

This inside-out arrangement creates two visual

Therefore, your visual system can use the
problems:
degree of binocular disparity to construct
• One is that the incoming light is distorted by
one three-dimensional perception from
the retinal tissue through which it must pass
two two-dimensional retinal images
before reaching the receptors.
• The other is that for the bundle of retinal
ganglion cell axons to leave the eye, there
must be a gap in the receptor layer; this gap
is called the blind spot.
 CONE AND ROD VISION
The first of these two problems are minimized There are two different types of receptors in
by the fovea, an indentation at the center of the human retina: cones and rods.
the retina that is specialized for high-acuity
vision (for seeing fine details). It emerged after it was first noticed that
species active only in the day tend to have
The thinning of the retinal ganglion cell cone-only retinas, and species active only
layer at the fovea reduces the distortion of at night tend to have rod-only retinas.
incoming light.
From this observation emerged the duplexity
The blind spot, the second of the two visual theory of vision—the theory that cones and
problems, requires a more creative solution. rods mediate different kinds of vision.

When the visual system detects a straight bar Photopic vision (cone-mediated vision)
going into one side of the blind spot and predominates in good lighting and provides
another straight bar leaving the other side, it high-acuity (finely detailed) colored
fills in the missing bit and what you see is a perceptions of the world.
continuous straight bar, regardless of what is
actually there. In dim illumination, there is not enough light
to reliably excite the cones, and the more
The visual system uses information sensitive scotopic vision (rod-mediated
provided by the receptors around the vision) predominates.
blind spot to fill in the gaps in your retinal
However, the sensitivity of scotopic vision
images—a process known as Completion.
is not achieved without cost: Scotopic
vision lacks both the detail and the color
It is one compelling demonstration that the
of photopic vision.
visual system does much more than make
a faithful copy of the external world.
The differences between photopic and
It is a mistake to think that completion is scotopic vision result in part from a difference
merely a response to blind spots. Completion in the way the two systems are “wired.”
also plays an important role in normal vision. • In the scotopic system, the output of
several hundred rods converges on a
When you look at an object, your visual single retinal ganglion cell
system does not conduct an image of that • Whereas in the photopic system, only
object from your retina to your cortex. a few cones converge on each retinal
Instead, it extracts key information about ganglion cell.
the object—primarily information about its
edges and their location—and conducts
that information to the cortex, where a
perception of the entire object is created
from that partial information.

For example, the color and brightness of

large unpatterned surfaces are not
perceived directly but are filled in
(completed) by a completion process called
surface interpolation—the process by
which we perceive surfaces.

The visual system extracts information

about edges and from it infers the
appearance of large surfaces.
 As a result, when dim light stimulates Under photopic conditions, notice that the
many rods simultaneously, the outputs of visual system is maximally sensitive to
this stimulation converge and summate wavelengths of about 560 nanometers.
(add) on the retinal ganglion cell. • Thus, under photopic conditions, a
light at 500 nanometers would
On the other hand, the effects of the same have to be much more intense
dim light applied to a sheet of cones than one at 560 nanometers to be
cannot summate to the same degree, and seen as equally bright.
the retinal ganglion cells may not respond
at all to the light. In contrast, under scotopic conditions, the
visual system is maximally sensitive to
wavelengths of about 500 nanometers.
• Thus, under scotopic conditions, a
light of 560 nanometers would
have to be much more intense
than one at 500 nanometers to be
seen as equally bright.

Because of the difference in photopic and

scotopic spectral sensitivity, an interesting
visual effect—the Purkinje Effect—can be
observed during the transition from photopic
to scotopic vision.

Cones and rods differ in their distribution on

the retina. There are no rods at all in the
fovea, only cones.

At the boundaries of the foveal indentation,

the proportion of cones declines markedly,
and there is an increase in the number of rods. EYE MOVEMENT
There are more rods in the nasal What we see is determined not just by what is
hemiretina (the half of each retina next to projected on the retina at that instant.
the nose) than in the temporal hemiretina
(the half of each retina next to the temples). Although we are not aware of it, the eyes
continually scan the visual field, and our visual
SPECTRAL SENSITIVITY perception at any instant is a summation of
recent visual information.
Because our visual systems are not equally
sensitive to all wavelengths in the visible It is because of this temporal integration that
spectrum, lights of the same intensity but of the world does not vanish momentarily each
different wavelengths can differ in brightness. time we blink.

Our eyes continuously move even when

we try to keep them fixated. Involuntary
fixational eye movements are of three
kinds: tremor, drifts, and saccades.

Although we are normally unaware of

fixational eye movements, they have a critical
visual function.
 When eye movements or their main effect The figure shows the relationship between
are blocked, visual objects begin to fade the absorption spectrum of rhodopsin and
and disappear. It happens as most visual the human scotopic spectral sensitivity
neurons respond only to changing images. curve. It means that, in dim light, our
sensitivity to various wavelengths is a
If retinal images are artificially stabilized direct consequence of rhodopsin’s ability
(kept from moving on the retina), the to absorb them.
images start to disappear and reappear.
Thus, eye movements enable us to see Rhodopsin is a G-protein–coupled receptor
during fixation by keeping the images that responds to light rather than to
moving on the retina. neurotransmitter molecules.
• Rhodopsin receptors initiate a cascade
VISUAL TRANSDUCTION: CONVERSION OF of intracellular chemical events when
LIGHT TO NEURAL SIGNALS they are activated.

Visual transduction is the conversion of light When rods are in darkness, their sodium
to neural signals by the visual receptors. channels are partially open, thus keeping the
rods slightly depolarized and allowing a steady
When the pigment from the rods—which flow of excitatory glutamate neurotransmitter
became known as rhodopsin—was molecules to emanate from them.
exposed to continuous intense light, it was
bleached (lost its color) and lost its ability However, when rhodopsin receptors are
to absorb light, but when it was returned bleached by light, the resulting cascade of
to the dark, it regained both its redness intracellular chemical events closes the sodium
and its light-absorbing capacity. channels, hyperpolarizes the rods, and reduces
the release of glutamate.
Rhodopsin’s absorption of light (and the
accompanying bleaching) is the first step in The transduction of light by rods
rod-mediated vision. exemplifies an important point: Signals are
• Evidence forwards that the degree to often transmitted through neural systems
which rhodopsin absorbs light in various by decreases in activity.
situations predicts how humans see
under the very same conditions. FROM RETINA TO
PRIMARY VISUAL CORTEX
The degree to which rhodopsin absorbs lights
of different wavelengths is related to the ability Many pathways in the brain carry visual
of humans and other animals with rods to information. The largest and most thoroughly
detect the presence of different wavelengths of studied visual pathways are the retina-
light under scotopic conditions. geniculate-striate pathways, which conduct
signals from each retina to the primary visual
cortex (striate cortex) via the lateral geniculate
nuclei of the thalamus.

RETINA-GENICULATE-STRIATE SYSTEM

About 90 percent of axons of retinal ganglion

cells become part of the retina-geniculate-
striate pathways. No other sensory system has
such a predominant pair (left and right) of
pathways to the cortex.
All signals from the left visual field reach the right
primary visual cortex—either ipsilaterally from the
temporal hemiretina of the right eye or
contralaterally (via the optic chiasm) from the nasal
hemiretina of the left eye—and that the opposite
is true of all signals from the right visual field.
 THE M AND P CHANNELS
Two parallel channels of communication flow
through each lateral geniculate nucleus.

One channel runs through the top four layers.

These layers are called the parvocellular layers
(or P layers) because they are composed of
neurons with small cell bodies.

The other channel runs through the bottom

two layers, which are called the magnocellular
layers (or M layers) because they are
composed of neurons with large cell bodies.

Each lateral geniculate nucleus receives visual The parvocellular neurons are particularly
input only from the contralateral visual field; responsive to color, fine pattern details,
three layers receive input from one eye, and and stationary or slowly moving objects. In
three receive input from the other. contrast, the magnocellular neurons are
particularly responsive to movement.
Most of the lateral geniculate neurons that
project to the primary visual cortex terminate Cones provide the majority of the input to
in the lower part of cortical layer IV, producing the P layers, whereas rods provide the
a characteristic stripe, or striation, when majority of the input to the M layers.
viewed in cross section—hence, primary visual
cortex is often referred to as striate cortex. SEEING EDGES

RETINOPIC ORGANIZATION Edge perception does not sound like a

particularly important topic, but it is. Edges are
The retina-geniculate-striate system is the most informative features of any visual
retinotopic; each level of the system is display because they define the extent and
organized like a map of the retina. position of the various objects in it.
• Two stimuli presented to adjacent areas
of the retina excite adjacent neurons at all Accordingly, the perception of an edge is
levels of the system really the perception of a contrast between
two adjacent areas of the visual field.
A dramatic demonstration of the
retinotopic organization of the primary CONTRAST ENHANCEMENT
visual cortex was provided by Dobelle,
Mladejovsky, and Girvin (1974). Adjacent to each edge, the brighter stripe
looks brighter than it really is and the darker
If electrical current was administered stripe looks darker than it really is.
simultaneously through an array of
electrodes forming a shape, such as a cross, The nonexistent stripes of brightness and
on the surface of a patient’s cortex, the darkness running adjacent to the edges are
patient reported “seeing” a glowing image called Mach bands; they enhance the contrast
of that shape. at each edge and make the edge easier to see.

Every edge we look at is highlighted for us

This finding and recent research on retinal by the contrast-enhancing mechanisms of
implants could be the basis for the our nervous systems.
development of visual prostheses that
could benefit many blind people. In effect, our perception of edges is better
than the real thing.
 RECEPTIVE FIELDS OF
RETINA-GENICULATE-STRIATE SYSTEM

Hubel and Wiesel (1979) began their studies of

visual system neurons by recording from the
three levels of the retina-geniculate-striate
system: first from retinal ganglion cells, then
from lateral geniculate neurons, and finally
from the striate neurons of lower layer IV.

When Hubel and Wiesel compared the receptive

fields recorded from retinal ganglion cells, lateral
geniculate nuclei, and lower layer IV striate
neurons, four commonalties were apparent:
• At each level, the receptive fields in the
RECEPTIVE FIELDS OF VISUAL NEURONS foveal area of the retina were smaller
than those at the periphery; this is
Hubel and Wiesel’s influential method is a consistent with the fact that the fovea
technique for studying single neurons in the mediates high-acuity vision.
visual systems of laboratory animals. • All the neurons had receptive fields that
were circular.
First, the tip of a microelectrode is • All the neurons were monocular; that is,
positioned near a single neuron in the part each neuron had a receptive field in one
of the visual system under investigation. eye but not the other.
• Many neurons at each of the three levels
During testing, eye movements are of the retina-geniculate-striate system
blocked by paralyzing the eye muscles, had receptive fields that comprised an
and the images on a screen in front of the excitatory area and an inhibitory area
subject are focused sharply on the retina separated by a circular boundary.
by an adjustable lens.
When Hubel and Wiesel shone a spot of
The next step in the procedure is to identify achromatic light onto the various parts of the
the receptive field of the neuron. The receptive fields of a neuron in the retina-
receptive field of a visual neuron is the geniculate-striate pathway, they discovered
area of the visual field within which it is two different responses.
possible for a visual stimulus to influence • The neuron responded with either
the firing of that neuron. “on” firing or “off” firing, depending
on the location of the spot of light in
The final step in the method is to record the the receptive field.
responses of the neuron to various simple
stimuli within its receptive field in order to For most of the retinal ganglion cells, lateral
characterize the types of stimuli that most geniculate nuclei, and lower layer IV striate
influence its activity. neurons, the reaction— “on” firing or “off”
firing—to a light in a particular part of the
Then the electrode is advanced slightly, receptive field was quite predictable.
and the entire process of identifying and • It depended on whether they were
characterizing the receptive field on-center cells or off-center cells.
properties is repeated for another neuron,
and then for another, and so on.

The general strategy is to begin by

studying neurons near the receptors and
gradually work up through “higher” and
“higher” levels of the system in an effort to
understand the increasing complexity of
the neural responses at each level.
 On-center cells respond to lights shone in
RECEPTIVE FIELDS OF
the central region of their receptive fields
PRIMARY VISUAL CORTEX
with “on” firing and to lights shone in the
periphery of their receptive fields with The receptive fields of most primary visual
inhibition, followed by “off” firing when cortex neurons fall into one of two classes:
light is turned off. simple or complex.

SIMPLE STRIATE CELLS

• They have receptive fields that can be

divided into antagonistic “on” and “off”
regions and are thus unresponsive to
diffuse light.
• They are all monocular.
• Borders between the “on” and “off” regions
of the cortical receptive fields of simple
cells are straight lines rather than circles.
Off-center cells display the opposite • They respond best to bars of light in a dark
pattern: They respond with inhibition and field, dark bars in a light field, or straight
“off” firing in response to lights in the edges between dark and light areas.
center of their receptive fields and with • They respond maximally only when its
“on” firing to lights in the periphery of their preferred straight-edge stimulus is in a
receptive fields. particular position and in a particular
orientation.
• The receptive fields of simple cortical cells
In effect, on-center and off-center cells
are rectangular rather than circular.
respond best to contrast. The most effective
way to influence the firing rate of an on-center
or off-center cell is to maximize the contrast COMPLEX STRIATE CELLS
between the center and the periphery of its
• They are more numerous than simple cells.
receptive field by illuminating either the entire
center or the entire surround (periphery) while • They have rectangular receptive fields,
leaving the other region completely dark. respond best to straight-line stimuli in a
specific orientation, and are unresponsive
Hubel and Wiesel thus concluded that one to diffuse light.
function of many of the neurons in the
retina-geniculate-striate system is to However, they differ from simple cells in three
respond to the degree of brightness important ways.
contrast between the two areas of their • First, they have larger receptive fields.
receptive fields. • Second, it is not possible to divide the
receptive fields of complex cells into static
“on” and “off” regions: A complex cell
responds to a particular straight-edge
stimulus of a particular orientation
regardless of its position within the
receptive field of that cell.

Thus, if a stimulus that produces “on” firing

in a particular complex cell is swept across
that cell’s receptive field, the cell will
respond continuously to it as it moves
across the field.

Many complex cells respond more robustly

to the movement of a straight line across
their receptive fields in a particular
direction.
 • Third, many complex cells are binocular • The “preferences” of the neurons became
(respond to stimulation of either eye). more complex. It occurred because neurons
with simpler preferences converged on
BINOCULAR COMPLEX STRIATE CELLS neurons with more complex preferences.

What you learn about the cell by stimulating CHANGING CONCEPT OF THE
one eye is confirmed by stimulating the other. CHARACTERISTICS OF THE VISUAL
RECEPTIVE FIELDS
What is more, if the appropriate stimulation is
applied through both eyes simultaneously, a Numerous studies after Hubel and Wiesel have
binocular cell usually fires more robustly than discovered that receptive fields are much more
if only one eye is stimulated. complex than was originally recognized.

Most of the binocular cells in the primary visual RETINAL GANGLION CELLS
cortex display some degree of ocular
dominance; that is, they respond more Primates have about 20 and 40 distinct sorts of
robustly to stimulation of one eye than they retinal ganglion cells, respectively—each with
do to the same stimulation of the other. its own sort of receptive field.

In addition, some binocular cells fire best when In addition to the on-center and off-center
the preferred stimulus is presented to both receptive fields, there are also retinal ganglion
eyes at the same time but in slightly different cells with receptive fields that are selective to
positions on the two retinas. These cells one or more of the following: (1) uniform
respond best to retinal disparity and thus are illumination, (2) orientation, (3) motion, and
likely to play a role in depth perception. (4) direction of motion.

LATERAL GENICULATE CELLS

ORGANIZATION OF
PRIMARY VISUAL CORTEX Analyses have shown that some cells in the
Hubel and Wiesel reached three important lateral geniculate nucleus have receptive
conclusions about the organization of primate fields that are sensitive to more than contrast.
visual cortex:
• The primary visual cortex was organized These cells have receptive fields that are
into functional vertical columns: All of the sensitive to one or more of the following: (1)
neurons in the same vertical column orientation, (2) motion, and (3) direction of
respond to stimuli applied to the same area motion. These receptive fields are similar to
of the retina, are dominated by the same those of retinal ganglion cell.
eye and “prefer” the same straight-line
angles (if they display a preference for CHANGING CONCEPT OF THE VISUAL
straight-line stimuli). RECEPTIVE FIELDS: CONTEXTUAL
• The location of various functional columns INFLUENCES IN VISUAL PROCESSING
in primary visual cortex is influenced by the
Most investigations of the responsiveness of
location on the retina of the column’s visual
visual system neurons have been based on two
fields, by the dominant eye of the column,
implicit assumptions.
and by the column’s straight-line angle.
• The first is that the mechanisms of visual
All of the functional columns in the primary visual processing can be best identified by
cortex that analyze input from one area of the studies using simplified, controllable,
retina are clustered together. artificial stimuli.
• The second is that the receptive field
Half of a cluster receives input from the left eye and properties of each neuron are static,
the other half receives input from the right eye, and unchanging properties of that neuron.
that each cluster includes neurons with preferences
for straight-line stimuli of various orientations.
 Research that has employed video clips of Sunlight and most sources of artificial light
real scenes involving natural movement contain complex mixtures of most visible
suggests that neither of these assumptions wavelengths. The mixture of wavelengths that
is correct at any of the three levels of the objects reflect influences our perception of
retina-geniculate-striate system. their color.

Studies of the responses of visual cortex to COMPONENT AND

natural scenes indicate that the response of a OPPONENT PROCESSING
visual cortex neuron depends not only on the
stimuli in its receptive field but also on the COMPONENT THEORY
larger scene in which these stimuli are
According to this theory, there are three
embedded.
different kinds of color receptors (cones),
each with a different spectral sensitivity. The
The influences on a visual neuron’s activity that
color of a particular stimulus is presumed to
are caused by stimuli outside the neuron’s
be encoded by the ratio of activity in the three
receptive field are generally referred to as
kinds of receptors.
contextual influences.
• This theory is derived from the
• They can take many forms depending
observation that any color of the visible
on the exact timing, location, and shape
spectrum can be matched by a mixing
of the visual stimuli under investigation
together of three different wavelengths
and on the ambient light levels.
of light in different proportions.
• The nature of contextual influences is
• The fact that three is normally the
dependent on prior exposure to them.
minimum number of different
• Contextual signals associated with
wavelengths necessary to match every
particular actions/states can help shape
color suggested that there were three
the properties of a receptive field. types of receptors.
A visual neuron’s receptive field was
initially assumed to be a property of the OPPONENT-PROCESSING THEORY
neuron resulting from the hardwired
convergence of neural circuits. According to this theory, there are two
different classes of cells in the visual system
Now, a neuron’s receptive field is viewed for encoding color and another class for
as a plastic property of the neuron that is encoding brightness.
continually fine-tuned on the basis of prior
experience and current signals from the Each of the three classes of cells encoded two
animal’s environment. complementary color perceptions.
• One class of color-coding cells signaled
red by changing its activity in one
SEEING COLOR direction and signaled red’s color
complementary, green, by changing its
Color is one of the most obvious qualities of
activity in the other direction.
human visual experience. The correct term for
• Another class of color-coding cells was
colors is hues, but in everyday language they
hypothesized to signal blue and its
are referred to as colors.
complement, yellow, in the same
opponent fashion.
What is there about a visual stimulus that
• A class of brightness-coding cells was
determines the color we perceive?
hypothesized to similarly signal both
• To a large degree, the perception of an
black and white.
object’s color depends on the
wavelengths of light that it reflects
into the eye.
 The opponent-process theory of color COLOR CONSTANCY
vision was based on several behavioral AND RETINEX THEORY
observations.
• One was that complementary Neither component nor opponent processing
colors cannot exist together: can account for the single most important
There is no such thing as bluish characteristic of color vision: color constancy.
yellow or reddish green • It means that the perceived color of an
• Another was that the afterimage object is not a simple function of the
produced by staring at red is green wavelengths reflected by it.
and vice versa, and the afterimage
produced by staring at yellow is Color constancy is an important—but much
blue and vice versa. misunderstood—concept. Color constancy is
the tendency for an object to stay the same
Research subsequently proved that both color- color despite major changes in the
coding mechanisms coexist in our visual wavelengths of light that it reflects.
systems.
Although the phenomenon of color
A technique for measuring the absorption constancy is counterintuitive, its advantage
spectrum of the photopigment contained in a is obvious. It improves our ability to tell
single cone that allowed researchers to objects apart in a memorable way so that
confirm the conclusion that: we can respond appropriately to them.
• There are indeed three different kinds
of cones in the retinas of those Our ability to recognize objects would be
vertebrates with good color vision greatly lessened if their color changed
• Each of the three has a different every time there was a change in
photopigment with its own illumination. In essence, if it were not for
characteristic absorption spectrum. color constancy, color vision would have
little survival value.
•
•
Although color constancy is an important
feature of our vision, we are normally unaware
of it. It is only in the controlled environment of
the laboratory that one can fully appreciate
that color constancy is more than an
important factor in color vision: It is the
essence of color vision.

According to Land’s retinex theory of color

vision, the color of an object is determined by
its reflectance—the proportion of light of
different wavelengths that a surface reflects.

Although the wavelengths of light reflected by

Most primates are trichromats (possessing a surface change dramatically with changes in
three color vision photopigments). illumination, the efficiency with which a
surface absorbs each wavelength and reflects
Most other mammals are dichromats the unabsorbed portion does not change
(possessing two color vision photo
pigments)—they lack the photopigment According to the retinex theory, the visual
sensitive to long wavelengths and thus system calculates the reflectance of
have difficulty seeing light at the red end of surfaces, and thus, perceives their colors,
the visible spectrum by comparing the light reflected by
adjacent surfaces in at least three
In contrast, some birds, fish, and reptiles different wavelength bands (short,
have four photopigments, and some medium, and long).
insects have five or more photopigments—
the dragonfly wins first place with ten.
 CORTICAL MECHANISM OF VISION AND
CONSCIOUS AWARENESS

There is much more to the human visual the corresponding area of the contralateral
system—we are visual animals. The entire visual field of both eyes.
occipital cortex as well as large areas of
temporal cortex and parietal cortex are Neurological patients with suspected damage
involved in vision. to the primary visual cortex are usually given a
perimetry test.
THREE DIFFERENT CLASSES OF
VISUAL CORTEX Many patients with scotomas are not
consciously aware of their deficits.
Visual cortex is often considered to be of three
different classes. One factor that contributes to this lack of
• Primary visual cortex is that area of cortex awareness is completion. A patient with a
that receives most of its input from the scotoma who looks at a complex figure, part of
visual relay nuclei of the thalamus. which lies in the scotoma, often reports seeing
• Areas of secondary visual cortex are a complete image.
those that receive most of their input
from the primary visual cortex. Blindsight is the ability to respond to visual
• Areas of visual association cortex are stimuli in a scotoma with no conscious
those that receive input from areas of awareness of them. For example, a patient
secondary visual cortex as well as from with blindsight might reach out and grab a
the secondary areas of other sensory moving object in her scotoma, all the while
systems. claiming not to see the object.

The primary visual cortex is located in the Functional Areas of Secondary and
posterior region of the occipital lobes. Association Visual Cortex

Areas of secondary visual cortex are located in Secondary visual cortex and the portions of
two general regions: in the prestriate cortex association cortex involved in visual analysis
and in the inferotemporal cortex. are both composed of many different areas,
• The prestriate cortex is the band of each specialized for a type of visual analysis.
tissue in the occipital lobe that
surrounds the primary visual cortex.
• The inferotemporal cortex is the
cortex of the inferior temporal lobe.

Areas of association cortex that receive visual

input are located in several parts of the
cerebral cortex, but the largest single area is in
the posterior parietal cortex.

The major flow of visual information in the

cortex is from primary visual cortex to the
various areas of secondary visual cortex to
the areas of association cortex.
PET, fMRI, and evoked potentials have been used to
Damage to Primary Visual Cortex: identify various areas of visual cortex in humans. The
Scotomas and Completion activity of volunteers’ brains has been monitored
while they inspect various types of visual stimuli.
Damage to an area of the primary visual cortex
produces a scotoma—an area of blindness—in By identifying the areas of activation associated with
various visual properties, researchers have so far
delineated about a dozen different functional areas of
human visual cortex.
 Dorsal and Ventral Streams

Many pathways that conduct information from While, patients with damage to the
the primary visual cortex through various inferotemporal cortex often have no
specialized areas of secondary and association difficulty reaching accurately for objects
cortex can be thought of as components of two they have difficulty describing.
major streams: the dorsal stream and the
ventral stream
• The dorsal stream flows from the primary There is an alternative interpretation for the
visual cortex to the dorsal prestriate same evidence.
cortex to the posterior parietal cortex
• The ventral stream flows from the Goodale and Milner argued that the
primary visual cortex to ventral prestriate primary difference between the dorsal
cortex to inferotemporal cortex. and ventral streams is not the kinds of
information they carry but the use to
which that information is put.

They suggested that the primary function

of the dorsal stream is to direct behavioral
interactions with objects, whereas the
primary function of the ventral stream is
to mediate the conscious perception of
objects.

They termed this the “control of behavior”

versus “conscious perception” theory.

STIMULUS PREFERENCE The major support for the “control of

Most visual cortex neurons in the dorsal behavior” versus “conscious perception”
stream respond most robustly to spatial theory is the confirmation of its two primary
stimuli, such as those indicating the location of predictions:
objects or their direction of movement. (1) that some patients with bilateral lesions
to the ventral stream may have no
In contrast, most neurons in the ventral conscious experience of seeing and yet
stream respond to the characteristics of be able to interact with objects under
objects, such as color and shape. visual guidance
(2) that some patients with bilateral lesions
Ungerleider and Mishkin proposed that the
to the dorsal stream may consciously
dorsal and ventral visual streams perform
see objects but be unable to interact
different visual functions.
with them under visual guidance
The dorsal stream is involved in the
perception of “where” objects are while Prosopagnosia
the ventral stream is involved in the
Prosopagnosia, briefly put, is a visual agnosia
perception of “what” objects are.
for faces that can be acquired either during
development (developmental prosopagnosia)
A major implication of the “where” versus or as a result of brain injury (acquired
“what” theory of vision is that damage to some prosopagnosia).
areas of cortex may abolish certain aspects of
vision while leaving others unaffected. A visual agnosia is a specific agnosia for visual
stimuli. In other words, visual agnosics can see
Patients with damage to the posterior things, but they don’t know what they are.
parietal cortex often have difficulty
reaching accurately for objects but they
have no difficulty describing.
 THE SENSORIMOTOR SYSTEM
The diagnosis of acquired prosopagnosia is
usually associated with damage to either or Chapter Overview
both of the fusiform face area and the occipital The three principles of sensorimotor control
face area. that are the foundations of this chapter:
(1) The sensorimotor system is
The fusiform face area (FFA) is located on the hierarchically organized.
ventral surface of the boundary between the (2) Motor output is guided by sensory input.
occipital and temporal lobes. It has been (3) Learning can change the nature and the
implicated in face identification because parts locus of sensorimotor control
of it are selectively activated by human faces
and because electrical stimulation of this brain
area in humans can metamorphose a viewed THREE PRINCIPLES OF
face into a completely different face. SENSORIMOTOR FUNCTION

The occipital face area (OFA) is located on the The Sensorimotor System
ventral surface of the occipital lobe. Is Hierarchically Organized
Reversible inactivation of the OFA by The operation of the sensorimotor system is
transcranial magnetic stimulation disrupts the directed by commands that cascade down
ability to discriminate between faces/ through the levels of a hierarchy—from the
association cortex (the highest levels) to the
Akinetopsia muscles (the lowest levels).

Akinetopsia is a deficiency in the ability to see The commands that emerge from the
movement progress in a normal smooth association cortex specify general goals rather
fashion—individuals affected by it only see than specific plans of action. The association
periodic snapshots of the world. cortex does not routinely gets involved in the
details.
It can be either a permanent result of brain
damage, or it can be the transient result of The main advantage of this hierarchical
taking high doses of certain antidepressants. organization is that the higher levels of the
hierarchy are left free to perform more
When akinetopsia is the result of an acquired complex functions.
brain injury, it is often associated with damage
to area MT (middle temporal area) of the The sensorimotor system are parallel
cortex, near the junction of the temporal, hierarchical systems; that is, they are
parietal, and occipital lobe. The function of MT hierarchical systems in which signals flow
appears to be the perception of motion. between levels over multiple paths.
• This parallel structure enables the
END OF THE VISUAL SYSTEM (#) association cortex to exert control
over the lower levels of the hierarchy
in more than one way.
• For example, the association cortex
can directly inhibit an eye blink reflex
to allow the insertion of a contact lens.

The sensorimotor are also characterized by

functional segregation.
• Each level of the sensorimotor
hierarchies tends to be composed of
different neural structures, each of
which performs a different function.
 The organization of individual responses
into continuous motor programs and
the transfer of their control to lower
In summary, the sensorimotor system is a levels of the CNS characterize most
parallel, functionally segregated, sensorimotor learning.
hierarchical system.
General Model of Sensorimotor
The main difference between the sensory
System Function
systems and the sensorimotor system is
the primary direction of information flow.

In sensory systems, information mainly

flows up through the hierarchy; in the
sensorimotor system, information mainly
flows down.

Motor Output Is Guided by Sensory Input

The sensorimotor system is flexible. They

continuously monitor the effects of their own
activities, and they use this information to fine-
tune their activities.
The figure illustrates several principles of
The sensory organs all monitor the body’s sensorimotor system organization.
responses, and they feed their information
back into sensorimotor circuits. Notice its hierarchical structure, the
• In most instances, this sensory functional segregation of the levels (e.g., of
feedback plays an important role in secondary motor cortex), the parallel
directing the continuation of the connections between levels, and the
responses that produced it. numerous feedback pathways.

The only responses that are not normally

influenced by sensory feedback are ballistic Sensorimotor Association Cortex
movements—brief, all-or-none, high-speed
movements. Association cortex is at the top of your
sensorimotor hierarchy. There are two major
Many adjustments in motor output that areas of sensorimotor association cortex: the
occur in response to sensory feedback are posterior parietal association cortex and the
controlled unconsciously by the lower dorsolateral prefrontal association cortex.
levels of sensorimotor hierarchy without
the involvement of the higher levels. Posterior parietal cortex and the dorsolateral
prefrontal cortex are each composed of several
Learning Changes the Nature and different areas, each with different functions.
Locus of Sensorimotor Control
Posterior Parietal Association Cortex
During the initial stages of motor learning,
each individual response is performed under Before an effective movement can be initiated,
conscious control. certain information is required.

Then, after much practice, individual The nervous system must know the original
responses become organized into continuous positions of the parts of the body that are to
integrated sequences of action that flow be moved, and it must know the positions of
smoothly and are adjusted by sensory any external objects with which the body is
feedback without conscious regulation. going to interact.
 The posterior parietal association cortex Apraxia is a disorder of voluntary
plays an important role in integrating movement that is not attributable to a
these two kinds of information, in simple motor deficit or to any deficit in
directing behavior by providing spatial comprehension or motivation.
information, and in directing attention.
Remarkably, patients with apraxia have
The posterior parietal cortex is classified as difficulty making specific movements
association cortex because it receives input when they are requested to do so,
from more than one sensory system. particularly when the movements are out
of context; however, they can often readily
It receives information from the three sensory perform the very same movements under
systems that play roles in the localization of natural conditions when they are not
the body and external objects in space: the thinking about what they are doing.
visual system, the auditory system, and the
somatosensory system. Although its symptoms are bilateral,
apraxia is often caused by unilateral
damage to the left posterior parietal
cortex or its connections

Contralateral neglect, the other striking

consequence of posterior parietal cortex
damage, is a disturbance of a patient’s
ability to respond to stimuli on the side of
the body opposite (contralateral) to the
side of a brain lesion in the absence of
simple sensory or motor deficits.

Most patients with contralateral neglect

often behave as if the left side of their
world does not exist, and they often fail to
In turn, much of the output of the posterior
appreciate that they have a problem.
parietal cortex goes to areas of motor cortex,
to the dorsolateral prefrontal association
The disturbance is often associated with
cortex, to the various areas of secondary
large lesions of the right posterior parietal
motor cortex, and to the frontal eye field—a
cortex, though damage to other brain
small area of prefrontal cortex that controls
regions has also been implicated.
both eye movements and shifts in attention.

Studies in humans indicate that the Dorsolateral Prefrontal

posterior parietal cortex contains a mosaic Association Cortex
of small areas, each specialized for guiding
particular movements of eyes, head, arms, The other large area of association cortex that
or hands. has important sensorimotor functions is the
dorsolateral prefrontal association cortex.
Damage to the posterior parietal cortex can
It receives projections from the posterior
produce a variety of deficits, including deficits
parietal cortex, and it sends projections to
in the perception and memory of spatial
areas of secondary motor cortex, to primary
relationships, in accurate reaching and
motor cortex, and to the frontal eye field.
grasping, in the control of eye movement, and
in attention.
Activity of other dorsolateral prefrontal
However, apraxia and contralateral neglect are neurons is related to the response rather
the two most striking consequences of than to the object.
posterior parietal cortex damage.
These neurons typically begin to fire
before the response and continue to fire
until the response is complete.
 Neurons in many cortical motor areas
begin to fire in anticipation of a motor
activity, but those neurons in the
dorsolateral prefrontal association cortex
tend to fire first.

The response properties of dorsolateral

prefrontal neurons suggest that decisions to
initiate voluntary movements may be made in
this area of cortex, but these decisions depend
on critical interactions with posterior parietal
cortex and other areas of frontal cortex.
To qualify as secondary motor cortex, an
area must be appropriately connected
with association & secondary motor areas.

Electrical stimulation of an area of

secondary motor cortex typically elicits
complex movements, often involving both
sides of the body.

Neurons in an area of secondary motor

cortex often become more active just prior
to the initiation of a voluntary movement
and continue to be active throughout the
movement.
Secondary Motor Cortex
In general, areas of secondary motor cortex
Areas of secondary motor cortex are those that are thought to be involved in the
receive their input from association cortex. programming of specific patterns of
movements after taking general instructions
For many years, only two areas of secondary from dorsolateral prefrontal cortex.
motor cortex were known: the supplementary
motor area and the premotor cortex. Identifying the Areas of
The supplementary motor area wraps over Secondary Motor Cortex
the top of the frontal lobe and extends
Few discoveries have captured the interest of
down its medial surface into the
neuroscientists as much as the discovery of
longitudinal fissure.
mirror neurons.
Meanwhile, the premotor cortex runs in a
Mirror neurons are neurons that fire when an
strip from the supplementary motor area
individual performs a particular goal-directed
to the lateral fissure.
movement or when they observe the same
movement performed by another.
Identifying the Areas of
Secondary Motor Cortex Why did the discovery of mirror neurons in the
The simple two-area conception of secondary ventral premotor area create such a stir?
motor cortex has become more complex. The reason is that they provide a possible
mechanism for social cognition
Neuroanatomical and neurophysiological (knowledge of the perceptions, ideas, and
research has made a case for at least eight intentions of others). Mapping the actions
areas of secondary motor cortex in each of others onto one’s own action repertoire
hemisphere, each with its own subdivisions. might facilitate social understanding.
Support for the idea that mirror neurons play When they mapped out the relation between
a role in social cognition has come from each cortical site and the muscle that was
demonstrations that these neurons respond activated by its stimulation, they found that
to the understanding of the purpose of an the primary motor cortex is organized
action, not to some superficial characteristic of somatotopically—that is, according to a map
the action itself. of the body.

Most of the research on human mirror neuron The somatotopic layout of the human primary
mechanisms have been functional MRI studies. motor cortex is commonly referred to as the
Many of these studies have found areas of motor homunculus.
human motor cortex that are active when a
person performs, watches, or imagines a As in the left figure, each site in the primary
particular action. motor cortex receives sensory feedback from
receptors in the muscles and joints that the site
Primary Motor Cortex influences.
• Presumably, this latter adaptation
The primary motor cortex is located in the facilitates stereognosis—the process
precentral gyrus of the frontal lobe. of identifying objects by touch.

It is the major point of convergence of cortical What is the function of each primary motor
sensorimotor signals, and it is the major, but cortex neuron?
not the only, point of departure of Until recently, each neuron was thought to
sensorimotor signals from the cerebral cortex. encode the direction of movement.

The main evidence for this was the finding

that each neuron in the arm area of the
primary motor cortex fires maximally
when the arm reaches in a particular
direction and that each neuron has a
different preferred direction.

Current View of
Primary Motor Cortex Function

Recent efforts to map the primary motor

cortex have used a new stimulation technique
using longer pulses of current.

The results were amazing: Rather than

eliciting the contractions of individual muscles,
these currents elicited complex natural-looking
response sequences.

Conventional View of For example, stimulation at one site

Primary Motor Cortex Function reliably produced a feeding response: The
arm reached forward, the hand closed as if
In 1937, Penfield and Boldrey mapped the clasping some food, the closed hand was
primary motor cortex of conscious human brought to the mouth, and finally the
patients during neurosurgery. They found that mouth opened.
the stimulation of each particular cortical site
activated a particular contralateral muscle
and produced a simple movement.
These recent studies have revealed a looser Consider the implications of this finding—they
somatotopic organization than was are as important as they are counterintuitive.
previously thought.
• For example, although stimulations to First, the finding means that the signals from
the face area do tend to elicit facial every site in the primary motor cortex diverge
movements, those movements are greatly, so each particular site has the ability
complex species-typical movements. to get a body part to a target location
• Also, sites that move a particular body regardless of the starting position.
part overlap greatly with sites that
move other body parts. Second, it means that the sensorimotor
system is inherently plastic.
The conventional view that many primary • Apparently, each location in the
motor cortex neurons are tuned to primary motor cortex can produce the
movement in a particular direction has innumerable patterns of muscle
also been challenged. contraction and relaxation required to
get a body part from any starting point
An alternative to the idea that motor to a specific target location.
neurons are coded to particular angles of • Accordingly, it has been suggested
movement has come from the findings of that the primary motor cortex
studies in which the activity of individual contains an action map in addition to
primary motor cortex neurons is recorded a topographic map.
as monkeys moved about freely—rather
than as they performed simple, learned In the case of Belle and the Robotic arm,
arm movements. Belle’s remarkable feat raised a possibility
that is starting to be realized.
The firing of many primary motor cortex
neurons in freely moving monkeys was Indeed, there has been a recent flurry of
often related to the particular end point of technological advances involving brain–
a movement, not to the direction of the computer interfaces brain—usually via an
movement. array of electrodes placed in the brain).
• For example, paralyzed patients have
That is, if a monkey reached toward a learned to control robotic arms with
particular location, primary motor cortex neural signals collected via multi-
neurons sensitive to that target location electrode arrays implanted in the
tended to become active regardless of the primary motor cortex
direction of the movement.
Brain-computer interfaces have also been
The importance of the target of a movement, used to mitigate the effects of spinal-cord
rather than the direction of a movement, for damage.
the function of primary motor cortex was also
apparent in stimulation studies. In a study by Capogrosso, monkeys with
transected spinal cords each had a wireless
For example, if stimulation of a particular
transmitter implanted in their motor cortex to
motor cortex site caused a straight arm to
record and transmit motor cortex activity.
bend at the elbow to a 90-degree angle,
Another wireless receiver positioned on their
stimulation of the same site caused a
spinal cord just below the site of transection
tightly bent arm to straighten to the same converted that transmitted message into a
90-degree angle.
pattern of stimulation that elicited movement
of their paralyzed limb.
In other words, the same stimulation of
motor cortex can produce opposite Such brain–spine-computer interfaces might
movements depending on the starting allow for significant recovery from the effects
position, but the end position of the
of spinal cord damage
movements remains the same.
 EFFECTS OF PRIMARY MOTOR By performing this function, it is believed to
CORTEX LESIONS play a major role in motor learning, particularly
in the learning of sequences of movements in
Extensive damage to the human primary which timing is a critical factor.
motor cortex has less effect expected, given
that this cortex is the major point of departure The effects of diffuse cerebellar damage on
of motor fibers from the cerebral cortex. motor function are devastating.
• The patient loses the ability to
Large lesions to the primary motor cortex may accurately control the direction, force,
disrupt a patient’s ability to move one body velocity, and amplitude of movements
part independently of others, may produce and the ability to adapt patterns of
astereognosia (deficits in stereognosis), and motor output to changing conditions.
may reduce the speed, accuracy, and force of a • It is particularly difficult to maintain
patient’s movements. steady postures, and attempts to do so
frequently lead to tremor.
Such lesions do not, however, eliminate • There are also severe disturbances in
voluntary movement, presumably because balance, gait, speech, and the control
there are parallel pathways that descend of eye movement.
directly from secondary and association motor
areas to subcortical motor circuits without These effects of cerebellar damage
passing through primary motor cortex. suggest that the cerebellum plays a major
role in monitoring and adapting ongoing
Cerebellum and Basal Ganglia patterns of movement.

The cerebellum and the basal ganglia are both The functions of the cerebellum were once
important and highly interconnected thought to be entirely sensorimotor, but this
sensorimotor structures but neither is a major conventional view is no longer tenable as
part of the pathway by which signals descend patients with cerebellar damage often display
through the sensorimotor hierarchy. diverse sensory, cognitive, emotional, and
memory deficits.
Instead, both the cerebellum and the basal
ganglia interact with different levels of the There are several competing theories of
sensorimotor hierarchy and, in so doing, cerebellar function, but a popular one is that
coordinate and modulate its activities. the cerebellum plays an important role in
learning from one’s errors and in the
prediction of errors.
CEREBELLUM

The cerebellum’s structure and complex BASAL ANGLIA

connectivity with other brain structures
suggest its functional complexity. It contains The basal ganglia do not contain as many
more than half of the brain’s neurons. neurons as the cerebellum, but in one sense
they are more complex.
It receives information from primary and • It is a complex heterogeneous
secondary motor cortex, information collection of interconnected nuclei.
about descending motor signals from
brain-stem motor nuclei, and feedback The anatomy of the basal ganglia suggests
from motor responses via the that, like the cerebellum, they perform a
somatosensory and vestibular systems. modulatory function.
• They contribute few fibers to
It is thought to compare these three descending motor pathways; instead,
sources of input and correct ongoing they form neural loops via their
movements that deviate from their numerous reciprocal connections with
intended course. cortical areas and the cerebellum.
The traditional view of the basal ganglia was • The ventromedial tracts are much more
that they, like the cerebellum, play a role in diffuse. Many of their axons innervate
the modulation of motor output. interneurons on both sides of the spinal gray
• Now, the basal ganglia are thought to matter and in several different segments,
also be involved in a variety of whereas the axons of the dorsolateral tracts
cognitive functions and in many terminate in the contralateral half of one
aspects of motivation. spinal cord segment, sometimes directly on
a motor neuron.
The basal ganglia have also been shown to • The motor neurons activated by the
participate in learning. ventromedial tracts project to proximal
muscles of the trunk and limbs, whereas the
For example, they play a role in habit motor neurons activated by the
learning, a type of learning that is usually dorsolateral tracts project to distal muscles.
acquired gradually, trial-by-trial and in
classical conditioning. Because all four of the descending motor
tracts originate in the cerebral cortex, all
One theory of basal ganglia sensorimotor are presumed to mediate voluntary
function is based on its known roles in both movement. However, major differences in
movement and motivation. This theory their routes and destinations suggest that
comprises two major assertions: they have different functions.

The first states that the basal ganglia are The experiment by Lawrence and Kuypers
responsible for movement vigor: the control suggest that:
of the speed and amplitude of movement • The ventromedial tracts are involved
based on motivational factors. in the control of posture and whole-
body movements (e.g., walking,
The second assertion is that movement not climbing) and that they can exert
only involves the execution of actions but also control over the limb movements
requires that we actively suppress motor involved in such activities.
activity that is inappropriate or unhealthy. • In contrast, the dorsolateral tracts
control the movements of the limbs.
Descending Motor Pathways
Sensorimotor Spinal Circuits
Neural signals are conducted from the primary
motor cortex to the motor neurons of the We have descended the sensorimotor
spinal cord over four different pathways. hierarchy to its lowest level: the spinal circuits
and the muscles they control.
Two pathways descend in the dorsolateral
region of the spinal cord—collectively known Psychologists often think of the spinal cord
as the dorsolateral motor pathways, and two motor circuits as mere cables that carry
descend in the ventromedial region of the instructions from the brain to the muscles.
spinal cord—collectively known as the
ventromedial motor pathways. But, the motor circuits of the spinal cord show
considerable complexity in their functioning,
Signals conducted over these pathways act independent of signals from the brain.
together in the control of voluntary movement
Muscles
The Two Dorsolateral Motor Pathways and
the Two Ventromedial Motor Pathways Motor units are the smallest units of motor
activity.
When theEach motor
motor unit fires,
neuron comprises a single
all the muscle
The descending dorsolateral and motor
fibers neuron
of its and alltogether.
unit contract of the individual
ventromedial motor pathways are quite skeletal muscle fibers that it innervates.
similar. However, the dorsolateral tracts
differ from the ventromedial tracts in two When the motor neuron fires, all the
major respects: muscle fibers of its unit contract together.
 Any two muscles whose contraction
produces the same movement, be it
Motor units differ appreciably in the number flexion or extension, are said to be
of muscle fibers they contain. The units with synergistic muscles.
the fewest fibers permit the highest degree of
selective motor control. Whereas, those that act in opposition, like
the biceps and the triceps, are said to be
Acetylcholine activates the motor end-plate antagonistic muscles.
on each muscle fiber and causes the fiber to
contract. Muscles are elastic than inflexible and
• Contraction is the only method that cablelike.
muscles have for generating force, • Activation of a muscle can increase the
thus, any muscle can generate force in tension that it exerts on two bones
only one direction. without shortening and pulling them
• All of the motor neurons that together; this is termed isometric
innervate the fibers of a single muscle contraction.
are called its motor pool. • Or it can shorten and pull them
together; this is termed dynamic
Although it is an oversimplification, skeletal contraction.
muscle fibers are often considered to be of two
basic types: fast and slow. Tension in a muscle can be increased by
• Fast muscle fibers are those that increasing the number of neurons in its
contract and relax quickly. They are motor pool that are firing, by increasing
capable of generating great force but the firing rates of those already firing, or
they fatigue quickly because they are more commonly by a combination of these
poorly vascularized. two changes.
• In contrast, slow muscle fibers,
although slower and weaker, are Receptor Organs of Tendons and Muscles
capable of more sustained
contraction because they are more The activity of skeletal muscles is monitored by
richly vascularized. two kinds of receptors: Golgi tendon organs
and muscle spindles.
Each muscle has both fast and slow • Golgi tendon organs are embedded in
fibers—the fast muscle fibers participate in the tendons, which connect each
quick movements such as jumping, whereas skeletal muscle to bones.
the slow muscle fibers participate in • Muscle spindles are embedded in the
gradual movements such as walking. muscle tissue itself.

Because each muscle can apply force in Because of their different locations, Golgi
only one direction, joints that move in tendon organs and muscle spindles respond to
more than one direction must be controlled different aspects of muscle contraction.
by more than one muscle. • Golgi tendon organs respond to increases
in muscle tension but they are completely
Many skeletal muscles belong unambiguously insensitive to changes in muscle length.
to one of two categories: flexors or extensors. • In contrast, muscle spindles respond to
• Flexors act to bend or flex a joint, and changes in muscle length, but they do not
extensors act to straighten or extend it respond to changes in muscle tension.

The function of Golgi tendon organs is to provide the

central nervous system with information about muscle
tension, but they also serve a protective function.

When the contraction of a muscle is so extreme that

there is a risk of damage, the Golgi tendon organs
excite inhibitory interneurons in the spinal cord that
cause the muscle to relax.
 Stretch Reflex
The resulting leg extension after having their
knees rapped with a little rubber-headed
hammer is called the patellar tendon reflex
(patella means “knee”).
• This reflex is a stretch reflex—a reflex
elicited by a sudden external stretching
force on a muscle

Notice that each muscle spindle has its own

threadlike intrafusal muscle, which is
innervated by its own intrafusal motor neuron.

Why so?
Without its intrafusal motor input, a
muscle spindle would fall slack each time
its skeletal muscle (extrafusal) contracted.

In this slack state, the muscle spindle could

not do its job, which is to respond to slight
changes in extrafusal muscle length.

The intrafusal
motor neuron
solves this problem
by shortening the The sudden stretch of the thigh muscle
intrafusal muscle stretches its muscle-spindle stretch receptors,
each time the which in turn initiate a volley of action
extrafusal muscle potentials carried from the stretch receptors
becomes shorter. into the spinal cord by spindle afferent
neurons via the dorsal root.
Thus, keeping
enough tension on This volley of action potentials excites motor
the middle, stretch- neurons in the ventral horn of the spinal cord,
sensitive portion of which respond by sending action potentials
the muscle spindle back to the muscle whose stretch originally
to keep it excited them.
responsive to slight
changes in the The arrival of these impulses back at the
length of the starting point results in a compensatory
extrafusal muscle. muscle contraction and a sudden leg
extension.
In real-life situations, the function of the
stretch reflex is to keep external forces from
altering intended position of the body.

When an external force, such as a push on

your arm while you are holding a cup of
coffee, causes an unanticipated extrafusal
muscle stretch, the muscle-spindle feedback
circuit produces an immediate
compensatory contraction of the muscle that
counteracts the force and keeps you from
spilling the coffee.

The important role played by sensory feedback

in the regulation of motor output and the “Bad news” of a sudden painful event in the
ability of lower circuits in the motor hierarchy hand arrives in the dorsal horn of the spinal
to take care of “business details” without the cord and has two effects:
involvement of higher levels. • The signals excite both excitatory and
inhibitory interneurons. The excitatory
interneurons excite the motor
neurons of the elbow flexor; the
inhibitory interneurons inhibit the
motor neurons of the elbow extensor.
• Thus, a single sensory input produces a
coordinated pattern of motor output;
the activities of agonists and
antagonists are automatically
coordinated by the internal circuitry
of the spinal cord.
Movements are quickest when there is
Withdrawal Reflex simultaneous excitation of all agonists and
complete inhibition of all antagonists;
Unlike the stretch reflex, the withdrawal reflex
however, this is not the way voluntary
is not monosynaptic.
movement is normally produced.
• When a painful stimulus is applied to
the hand, the first responses are
Most muscles are always contracted to
recorded in the motor neurons of the
some degree, and movements are produced
arm flexor muscles about the time it
by adjustment in the level of relative
takes a neural signal to cross two
cocontraction between antagonists.
synapses.
Movements produced by cocontraction are
Other responses are recorded in the motor
smooth, and they can be stopped with
neurons of the arm flexor muscles after the
precision by a slight increase in the
initial volley; these responses are triggered by
contraction of the antagonistic muscles.
signals that have traveled over multisynaptic
Moreover, cocontraction insulates us from
pathways—some involving the cortex.
the effects of unexpected external forces.

Reciprocal Innervation
Recurrent Collateral Inhibition
Reciprocal innervation is an important
principle of spinal cord circuitry.
Like most workers, muscle fibers and the
It means that antagonistic muscles are motor neurons that innervate them need an
innervated in a way that permits a smooth, occasional break, and inhibitory neurons in
unimpeded motor response: When one is the spinal cord make sure they get it.
contracted, the other relaxes.
 Each motor neuron branches just before it That program must produce, on the basis of all
leaves the spinal cord, and the branch this information, an integrated series of
synapses on a small inhibitory interneuron, movements that involves the muscles of the
which inhibits the very motor neuron from trunk, legs, feet, and upper arms.
which it receives its input.
• The inhibition produced by these local This program of reflexes must also be
feedback circuits is called recurrent incredibly plastic; it must be able to adjust its
collateral inhibition output immediately to changes in the slope of
• The small inhibitory interneurons that the terrain, to instructions from the brain, or to
mediate recurrent collateral inhibition sudden external forces.
are called Renshaw cells.
Central Sensorimotor Programs
As a consequence of recurrent collateral and Learning
inhibition, each time a motor neuron fires,
it momentarily inhibits itself and shifts the A Hierarchy of Central
responsibility for the contraction of a Sensorimotor Programs
particular muscle to other members of the
muscle’s motor pool. One view of sensorimotor function is that the
sensorimotor system comprises a hierarchy of
central sensorimotor programs.

According to this view, all but the highest

levels of the sensorimotor system have
certain patterns of activity programmed into
them, and complex movements are produced
by activating the appropriate combinations of
these programs.

Once activated, each level of the sensorimotor

system is capable of operating on the basis of
current sensory feedback without the direct
control of higher levels.

Thus, although the highest levels of your

sensorimotor system retain the option of
directly controlling your activities, most of the
Walking: A Complex Sensorimotor Reflex
individual responses that you make are
Most reflexes are much more complex than performed without direct cortical
withdrawal and stretch reflexes. involvement.

A walking program must integrate: Characteristics of Central

• visual information from the eyes; Sensorimotor Programs
• somatosensory information from the
feet, knees, hips, arms CENTRAL SENSORIMOTOR PROGRAMS
• information about balance from the ARE CAPABLE OF MOTOR EQUIVALENCE
semicircular canals of the inner ears;
The sensorimotor system does not always
• information from newly discovered
accomplish a particular task in exactly the
sensory receptors in the spinal cord
same way.
that detect mechanical and chemical
information from the cerebrospinal The same basic movement can be carried
fluid in the central canal. out in different ways involving different
muscles is called motor equivalence.
 CENTRAL SENSORIMOTOR PROGRAMS
Motor equivalence illustrates the inherent CAN DEVELOP WITHOUT PRACTICE
plasticity of the sensorimotor system.
• It suggests that specific central Although central sensorimotor programs for
sensorimotor programs are not some behaviors can be established by
stored in the neural circuits. practicing the behaviors, the central
• General programs are stored higher in sensorimotor programs for many species-
your sensorimotor hierarchy and then typical behaviors are established without
are adapted to the situation as explicit practice of the behaviors.
required.
PRACTICE CAN CREATE CENTRAL
SENSORY INFORMATION THAT CONTROLS SENSORIMOTOR PROGRAMS
CENTRAL SENSORIMOTOR PROGRAMS IS
NOT NECESSARILY CONSCIOUS Although central sensorimotor programs for
many species-typical behaviors develop
Is there evidence for the separation of without practice, practice can generate or
conscious perception and sensory control of modify them.
behavior in intact humans?
Theories of sensorimotor learning emphasize
Haffenden and Goodale (1998) supplied such two kinds of processes that influence the
evidence. They showed healthy volunteers a learning of central sensorimotor programs:
three-dimensional version of the visual response chunking and shifting control to
illusion. lower levels of the sensorimotor system

Remarkably, when the volunteers were asked RESPONSE CHUNKING

to indicate the size of each central disk with According to the response-chunking
their right thumb and pointing finger, they hypothesis, practice combines the central
judged the disk on the left to be bigger than the sensorimotor programs that control
one on the right. individual responses into programs that
control sequences (chunks) of behavior.
However, when they were asked to reach out
and pick up the disks with the same two digits, An important principle of chunking is that
the preparatory gap between the digits was a chunks can themselves be combined into
function of the actual size of each disk rather higher-order chunks.
than its perceived size.
SHIFTING CONTROL TO LOWER LEVELS
In the process of learning a central
sensorimotor program, control is shifted from
higher levels of the sensorimotor hierarchy to
lower levels.

Shifting the level of control to lower levels of

the sensorimotor system during training has
two advantages.
• One is that it frees up the higher levels of
the system to deal with more esoteric
aspects of performance.
• The other advantage is that it permits great
speed because different circuits at the
lower levels of the hierarchy can act
simultaneously, without interfering with
one another.
 Functional Brain Imaging of
Sensorimotor Learning The Jenkins and colleagues brain-imaging
study of sensorimotor learning and
Functional brain-imaging techniques have subsequent studies like it have made
provided opportunities for studying gross important contributions by identifying
neural correlates of sensorimotor learning. where changes occur in the brain while
volunteer learn sensorimotor tasks.
By recording the brain activity of human
volunteers as they learn to perform new motor Neuroplasticity Associated with
sequences, researchers can develop Sensorimotor Learning
hypotheses about the roles of various
structures in sensorimotor learning. The learning of new sensorimotor tasks is
accompanied by both cortical and subcortical
One of the first studies of this type was the PET changes.
study of Jenkins. The results are summarized in
the figure below. First, at the level of the motor cortex, there is
a strengthening of the inputs from the
thalamus and from other areas of motor
cortex with learning. Such strengthening is
related to an increase in the number of
dendritic spines—which suggests an increase in
the number of synapses.

Second, there is a large increase in the

number of oligodendrocytes in subcortical
white matter just after sensorimotor learning.
This increase in the number of
oligodendrocytes is presumably due to an
increased demand for myelination of new
and/or existing axonal connections.

END OF THE SENSORIMOTOR SYSTEM (#)

Notice two things.

• First, notice the involvement of the
cortical sensorimotor areas
• Second, notice how the involvement of
association areas and the cerebellum
diminished when sequences were well
practiced.