The Special Senses

8
The Special Senses

LEARNING OBJECTIVES images, nor do they contribute to color vision. Rods operate
Objectives Heading best under conditions of reduced lighting (scotopic vision).
Upon completion of this chapter, the student should be able to
Cones, in contrast, are not as sensitive to light as rods are
answer the following questions: and thus operate best under daylight conditions (photopic
vision). Cones are responsible for high visual acuity and
1. What is the dark current, and how does the absorption of
color vision.
a photon change it?
2. What are the synaptic pathways for the central and
The retina is an outgrowth of the thalamus. Thus infor-
surround portions of the receptive field of an on-center mation processing within the retina is performed by retinal
bipolar cell? Of an off-center bipolar cell? neurons, and the output signals are carried to the brain by
3. What are the receptive field properties of simple and the axons of retinal ganglion cells in the optic nerves.
complex cells in the visual cortex? There is a partial crossing of these axons in the optic chiasm
4. What is the frequency theory of sound encoding? Why is that causes all input from one side of the visual space to
the place theory also required? pass to the opposite side of the brain. Posterior to the
5. What are the stimuli that are normally transduced by the optic chiasm, the axons of retinal ganglion cells form the
hair cells in the semicircular canals and otolith organs? optic tracts and synapse in nuclei of the brain. The main
6. What are the functional consequences of the differing visual pathway in humans targets the lateral geniculate
numbers of different receptor molecules between
nucleus (LGN) of the thalamus and this nucleus, through
olfactory and gustatory receptor cells?
the visual radiations, projects the visual information to the
visual cortex. Other visual pathways project to the superior
colliculus, pretectum, and hypothalamus, structures that
participate in orientation of the eyes, control of pupil size,
T
he evolution of vertebrates shows a trend called and circadian rhythms, respectively.
cephalization in which special sensory organs
develop in the heads of animals, along with the cor- Structure of the Eye
responding development of the brain. These special sensory
systems, which include the visual, auditory, vestibular, The wall of the eye is composed of three concentric layers
olfactory, and gustatory systems, detect and analyze light, (Fig. 8.1). The outer layer, or the fibrous coat, includes the
sound, and chemical signals in the environment, as well as transparent cornea, with its epithelium, and the opaque
signal the position and movement of the head. The stimuli sclera. The middle layer, or vascular coat, includes the iris
transduced by these systems are most familiar to humans and the choroid. The iris contains both radially and cir-
when they provide conscious awareness of the environment, cularly oriented smooth muscle fibers, which make up the
but they are equally important as the sensory basis for pupillary dilator and constricter muscles, respectively. The
reflexive and subconscious behavior. choroid is rich in blood vessels that support the outer layers
of the retina, and it also contains pigment. The innermost
The Visual System layer of the eye, the retina, is embryologically derived from
the diencephalon and therefore is part of the central nervous
Vision is one of the most important special senses in humans system (CNS). The functional part of the retina covers
and, along with audition, is the basis for most human the entire posterior aspect of the eye except for the optic
communication. The visual system detects electromagnetic nerve head, or optic disc, which is where the optic nerve
waves between 400 and 750nm long as visible light, which axons leave the retina. Because there are no receptors at
enters the eye and impinges on photoreceptors in a special- this location, it is often referred to as the anatomical blind
ized sensory epithelium, the retina. spot (see Fig. 8.1).
The photoreceptors, rods and cones, can distinguish two A number of functions of the eyes are under muscular
aspects of light: its brightness (or luminance) and its wave- control. Externally attached extraocular muscles aim the
length (or color). Rods have high sensitivity for detecting eyes toward an appropriate visual target (see Chapter 9).
low light intensities but do not provide well-defined visual These muscles are innervated by the oculomotor nerve
127
128 S E C T I O N 2 Berne & Levy Physiology
Visual axis Optic axis
Iris Aqueous humor

Posterior Cornea
chamber
Anterior chamber
Ciliary body Limbus
Lens Sclera
Zonule Nodal
fibers point
Choroid
Vitreous
humor
Retina
Nasal
retina
Temporal
Optic retina
disc Fovea
Optic
nerve
Fig. 8.1 Illustration of a view of a horizontal section of the right eye. (Redrawn from Wall GL. The
Vertebrate Eye and Its Adaptive Radiation. Bloomfield Hills, MI: Cranbrook Institute of Science; 1942.)
(cranial nerve [CN] III), the trochlear nerve (CN IV), Normally, light from a visual target is focused sharply
and the abducens nerve (CN VI). Several muscles are also on the retina by the cornea and lens, which bend or refract
found within the eye (intraocular muscles). The muscles in the light. The cornea is the major refractive element of the
the ciliary body control lens shape and thereby the focus of eye, with a refractive power of 43 dioptersa (D). However,
images on the retina. The pupillary dilator and sphincter unlike the cornea, the lens can change shape and vary
muscles in the iris control the amount of light entering the its refractive power between 13 and 26D. Thus the lens
eye, in a way similar to that of the diaphragm of a camera. is responsible for adjusting the optical focus of the eye.
The dilator is activated by the sympathetic nervous system, Suspensory ligaments (or zonule fibers) attach to the
whereas the sphincter and ciliary muscles are controlled by wall of the eye at the ciliary body (see Fig. 8.1) and hold
the parasympathetic nervous system (through the oculomo- the lens in place. When the muscles in the ciliary body
tor nerve; see Chapter 11). are relaxed, the tension exerted by the suspensory liga-
Light enters the eye through the cornea and passes ments flattens the lens. When the ciliary muscles contract,
through a series of transparent fluids and structures that the tension on the suspensory ligaments is reduced; this
are collectively called the dioptric media. These fluids and process allows the somewhat elastic lens to assume a more
structures consist of the cornea, aqueous humor, lens, and spherical shape. The ciliary muscles are activated by the
vitreous humor (see Fig. 8.1). The aqueous humor (located parasympathetic nervous system via the oculomotor nerve.
in the anterior and posterior chambers) and the vitreous In this way, the lens allows the eye to focus on, or accom-
humor (located in the space behind the lens) help maintain modate to, either near or distant objects. For instance, when
the shape of the eye. light from a distant visual target enters a normal eye (one
Although the geometrical optic axis of the human eye with a relaxed ciliary muscle), the target image is in focus on
passes through the nodal point of the lens and reaches the the retina. However, if the eye is directed at a nearby visual
retina at a point between the fovea and the optic disc (see target, the light is initially focused behind the retina (i.e.,
Fig. 8.1), the eyes are oriented by the oculomotor system the image at the retina is blurred) until accommodation
to a point, called the fixation point, on the visual target.
Light from the fixation point passes through the nodal point a
A diopter is a unit of measurement of optical power that is equal to the
of the lens and is focused on the fovea. Light from the reciprocal of the focal length measured in meters. Thus it is a unit of
remainder of the visual target falls on the retina surrounding reciprocal length, and a 2-D lens would bring parallel rays of light into
the fovea. focus at a distance of 0.5m.
CHAPTER 8 The Special Senses 129
occurs; that is, until the ciliary muscle contracts, causing

the lens to become more spherical, the increased convexity IN THE CLINIC
causes the lens to refract the light waves more strongly, As an individual ages, the elasticity of the lens gradually
bringing the image into focus on the retina. declines. As a result, accommodation of the lens for near
Proper imaging of light on the retina depends not only vision becomes progressively less effective, a condition
on the lens and cornea but also on the iris, which adjusts called presbyopia. A young person can change the power
the amount of light that can enter the eye through the of the lens by as much as 14D. However, by the time
that a person reaches 40 years of age, the amount of
pupil. In this regard, the pupil is analogous to the aperture accommodation halves, and after 50 years, it can decrease
in a camera, which also controls the depth of field of the to 2D or less. Presbyopia can be corrected by convex
image and the amount of spherical aberration produced by lenses.
the lens. When the pupil is constricted, the depth of field is Defects in focus can also be caused by a discrepancy
increased, and the light is directed through the central part between the size of the eye and the refractive power of the
dioptric media. For example, in myopia (near-sightedness),
of the lens, where spherical aberration is minimal. Pupillary the images of distant objects are focused in front of the
constriction occurs reflexively when the eye accommodates retina. Concave lenses correct this problem. Conversely,
for near vision or adapts to bright light, or both. Thus when in hypermetropia (far-sightedness), the images of distant
a person reads or does other fine visual work, the quality of objects are focused behind the retina; this problem can
the image is improved by adequate light. be corrected with convex lenses. In astigmatism, an
asymmetry exists in the radii of curvature of different
meridians of the cornea or lens (or sometimes of the retina).
Retina Astigmatism can often be corrected with lenses that possess
complementary radii of curvature.
Layers of the Retina
The 10 layers of the retina are shown in Fig. 8.2. The
outermost portion is the pigmented epithelium (layer 1),
which is just inside the choroid. The pigment cells have
Retinal layers Components

10 Inner limiting membrane
9 Nerve fiber layer
Axons at surface of
retina passing via
8 Ganglion cell layer
optic nerve, chiasm
and tract to lateral
geniculate body
7 Inner plexiform layer Ganglion cell
Mller cell
6 Inner nuclear layer
(supporting glial cell)
Bipolar cell
Amacrine cell
5 Outer plexiform layer
Horizontal cell
Rod
Cone
4 Outer nuclear layer
3 Outer limiting membrane

Inner segment
2 Photoreceptor layer
Outer segment
Pigment cells
1 Pigment epithelium
Choroid
Fig. 8.2
Layers of the Retina. Light impinging on the retina would be coming from the top of the figure,
and would pass through all the superficial layers to reach the photoreceptor rods and cones.
tentacle-like processes that extend into the photoreceptor Outer segment Outer segment
layer (layer 2) and surround the outer segments of the Free floating discs
rods and cones. These processes prevent transverse scatter
of light between photoreceptors. In addition, they serve a
mechanical function in maintaining contact between layers Plasma
membrane
1 and 2 so that the pigmented epithelium can (1) provide
nutrients and remove waste from the photoreceptors; (2) Folding of
phagocytose the ends of the outer segments of the rods, membrane
which are continuously shed; and (3) reconvert metabolized
visual pigment into a form that can be reused after it is
transported back to the photoreceptors. Cilium
Cilium
Retinal glial cells, known as Mller cells, play an impor- Inner segment
tant role in maintaining the internal geometry of the retina.
Mller cells are oriented radially, parallel to the light path
through the retina. The outer ends of Mller cells form tight
junctions with the inner segments of the photoreceptors,
and these numerous connections have the appearance of
a continuous layer, the outer limiting membrane (layer 3
of the retina). Synaptic terminals
Rod Cone
IN THE CLINIC Fig. 8.3 Rods and Cones. The drawings at the bottom show
the general features of a rod and a cone. The insets show the outer
The junction between layers 1 and 2 of the retina in adults segments.
represents the surface of contact between the anterior
and posterior walls of the embryonic optic cup during
development and is structurally weak. Retinal detachment
is separation at this surface and can cause loss of vision axons cross the retina helps permit light to pass through
because of displacement of the retina from the focal plane of
the eye. It can also lead to the death of photoreceptor cells, the inner retina with minimal distortion.
which are maintained by the blood supply of the choroid The innermost layer of the retina is the inner limiting
(the photoreceptor layer itself is avascular). Deterioration membrane (layer 10) formed by the end-feet of Mller
of the pigmented epithelium can also result in macular cells.
degeneration, a critical loss of high-acuity central and color
vision that does not affect peripheral vision. Structure of Photoreceptors: Rods and Cones
Each rod or cone photoreceptor cell is composed of a cell
body (in layer 4), an inner and an outer segment that
Inside the external limiting membrane is the outer extend into layer 2, and a set of synaptic terminals that
nuclear layer (layer 4) that contains the cell bodies and synapse in layer 5 onto other retinal cells (Fig. 8.3). The
nuclei of the rods and cones. The outer plexiform layer outer segments of cones are not as long as those of rods, and
(layer 5) contains synapses between the photoreceptors and they contain stacks of disc membranes formed by infoldings
retinal interneurons, including bipolar cells and horizontal of the plasma membrane. The outer segments of rods are
cells, whose cell bodies are found in the inner nuclear layer longer, and they contain stacks of membrane discs that float
(layer 6). This layer also contains the cell bodies of other freely in the outer segment, having completely disconnected
retinal interneurons (the amacrine and interplexiform cells) from the plasma membrane when formed at the base. Both
and the Mller cells. sets of discs are rich in visual pigment molecules, but rods
The inner plexiform layer (layer 7) contains synapses have a greater visual pigment density, which partly accounts
between the retinal neurons of the inner nuclear layer, for their greater sensitivity to light. A single photon can
including the bipolar and amacrine cells, and the ganglion elicit a rod response, whereas several hundred photons may
cells, whose cell bodies lie in the ganglion cell layer (layer be required for a cone response.
8). As previously mentioned, the ganglion cells are the The outer segments of the photoreceptors are connected
output cells of the retina; it is their axons that transmit by a modified cilium to the inner segments, which contain
visual information to the brain. These axons form the optic a number of organelles, including numerous mitochondria.
fiber layer (layer 9), pass along the inner surface of the The inner segments are the sites where the visual pigment is
retina while avoiding the fovea, and enter the optic disc, synthesized before it is incorporated into the membranes of
where they leave the eye as the optic nerve. The portions the outer segment. In rods, the pigment is inserted into new
of the ganglion cell axons that are in the optic fiber layer membranous discs, which are then displaced distally until
remain unmyelinated, but they become myelinated after they are eventually shed at the apex of the outer segment,
they reach the optic disc. The lack of myelin where the where they undergo phagocytosis by cells of the pigmented
Optic disc
16 Cones
Receptor density (104/mm2)

Rods
12
0
80 60 40 20 0 20 40 60
Nasal Fovea Temporal
Fig. 8.4 Graph of a Plot of the Density of Cones and Rods as a Function of Retinal Eccentricity
From the Fovea. Note that cone density peaks at the fovea, rod density peaks at about 20 degrees
eccentricity, and no photoreceptors are found at the optic disc, where the ganglion cell axons leave to
form the optic nerve. (Data from Cornsweet TN. Visual Perception. New York: Academic Press; 1970.)
epithelium. This process determines the rod-like shape of

the outer segments of rods. In cones, the visual pigment is IN THE CLINIC
inserted randomly into the membranous folds of the outer
As mentioned, the axons of retinal ganglion cells cross
segment, and shedding, comparable to that seen in rods, the retina in the optic fiber layer (layer 9) to enter the
does not take place. optic nerve at the optic disc. These axons in the optic
fiber layer pass around the macula and fovea, as do the
Regional Variations in the Retina blood vessels that supply the inner layers of the retina.
The macula lutea is the area of central vision and is charac- The optic disc can be visualized on physical examination
with an ophthalmoscope. The normal optic disc has a
terized by a slight thickening and a pale color. The thickness slight depression in its center. Changes in the appearance
is due to the high concentration of photoreceptors and of the optic disc are important clinically. For example, the
interneurons, which are needed for high-resolution vision. depression may be exaggerated by loss of ganglion cell
It is pale because both optic nerve fibers and blood vessels axons (optic atrophy), or the optic disc may protrude into
are routed around it. the vitreous space because of edema (papilledema) that
results from increased intracranial pressure.
The fovea, which is a depression in the macula lutea,
is the region of the retina with the very highest visual
resolution and, as noted previously, the light from the
fixation point is focused on the fovea. (A major function
of eye movements is to bring objects of interest into view
on the fovea.) The retinal layers in the foveal region are Visual Transduction
unusual because several of them appear to be pushed aside To be detected by the retina, light energy must be absorbed.
into the surrounding macula. Consequently, light can reach This is the job primarily of the rods and cones (a small class
the foveal photoreceptors without having to pass through of ganglion cells are also photosensitive) and is accom-
the inner layers of the retina, and both image distortion plished by visual pigment molecules located in their outer
and light loss are minimized. The fovea has cones with segments. For both rods and cones, the pigment molecule
unusually long and thin outer segments, which allows for consists of a chromophore, 11-cis retinal, bound to an opsin
high packing density. In fact, cone density is maximal in the protein. The visual pigment found in the outer segments of
fovea, providing for high visual resolution, as well as high rods is rhodopsin, or visual purple (so named because it
quality of the image (Fig. 8.4). has a purple appearance when light has been absorbed). It
The optic disc, where ganglion cell axons leave the retina, absorbs light best at a wavelength of 500nm. Three variants
lacks photoreceptors and therefore lacks photosensitivity. of visual pigment, resulting from the binding of different
Thus it is a so-called blind spot in the visual surface of opsins to retinal, are found in cones (in most species, each
the retina (see Figs. 8.4 and 8.9). A person is normally cone expresses one of the three cone pigments). The cone
unaware of the blind spot because the corresponding part pigments absorb best at 419nm (blue), at 533nm (green),
of the visual field can be seen by the contralateral eye and and at 564nm (red). However, the absorption spectrum of
because of the psychological process in which incomplete these visual pigments is broad so that they overlap consider-
visual images tend to be completed perceptually. ably (Fig. 8.5).
CONE ABSORPTION SPECTRA

Violet Rod Blue Green Yellow Orange Red
100
Rod
Blue Red
Normalized absorbance
cone cone
Green
cone
50
419 498 533 564
0
400 500 600
Wavelength (nm)
Fig. 8.5 The Spectral Sensitivity of the Three Types of Cone Pigments and of the Rod Pigment
(Rhodopsin) in the Human Retina. Note that the curves overlap and that the so-called blue and red
cones actually absorb maximally in the violet and yellow ranges, respectively. (Data from Squire LR, etal
[eds]. Fundamental Neuroscience. San Diego, CA: Academic Press; 2002.)
Despite the differences in spectral sensitivity, the trans- Na+ and Ca2+. The resulting current is known as the dark
duction process is similar in rods and cones. The absorption current, and the depolarization it causes leads to the tonic
of a photon by a visual pigment molecule leads to the release of the neurotransmitter glutamate at the photorecep-
isomerization of 11-cis retinal to all-trans retinal, release tors synapses.
of the bond with the opsin, and conversion of retinal to When light is absorbed in a rod (an equivalent sequence
retinol. These changes trigger a second messenger cascade happens in cones), photoisomerization of rhodopsin acti-
that leads to a change in the electrical activity of the rod or vates a G protein called transducin (see Fig. 8.6B). This
cone (discussed later in this section). G protein, in turn, activates cyclic guanosine monophos-
The separation of all-trans retinal from opsin also causes phate phosphodiesterase, which is associated with the
both the loss of its ability to absorb light and bleaching rhodopsin-containing discs, hydrolyzes cGMP to 5-GMP,
(i.e., the visual pigment loses its color). In both rods and and lowers the cGMP concentration in the rod cytoplasm.
cones, regeneration of the visual pigment molecule is a The reduction in cGMP leads to closing of the cGMP-gated
multistep process: the all-trans retinal is transported to the cation channels, hyperpolarization of the rod cell mem-
retinal pigmented cell layer, where it is reduced to retinol, brane, and a reduction in the release of neurotransmitters.
isomerized, and esterified back to 11-cis retinal. It is then Thus cGMP acts as a second messenger to translate the
transported back to the photoreceptor layer, taken up by absorption of a photon by rhodopsin into a change in
outer segments, and recombined with opsin to regenerate membrane potential.
the visual pigment molecule, which can again absorb light. In sum, in all photoreceptors (cones undergo a process
There is evidence that cones also use a second pathway to analogous to that described for rod transduction), capture
regenerate visual pigment. This pathway is much more rapid of light energy leads to (1) hyperpolarization of the
and involves transport of the retinal molecule to and from photoreceptor and (2) a reduction in the release of neu-
the Mller cells (see Fig. 8.2) rather than the pigmented rotransmitters. Because of the very short distance between
epithelial cells. The potential importance of this more rapid the site of transduction and the synapse, the modulation
pathway is discussed in the section Visual Adaptation. of neurotransmitter release is accomplished without the
Ultimately, the transduction process triggered by absorp- generation of an action potential.
tion of photons causes the photoreceptor to hyperpolarize.
To understand this action and its consequences fully, it is Visual Adaptation
necessary to know the baseline state of the photoreceptor in Adaptation refers to the ability of the retina to adjust its
the dark (i.e., before it absorbs a photon). In darkness, pho- sensitivity according to ambient light. This ability allows
toreceptors are slightly depolarized ( 40mV) in relation the retina to operate efficiently over a wide range of lighting
to most neurons because cyclic guanosine monophosphate conditions, and it reflects a switching between the use of the
(cGMP)gated cation channels in their outer segments are cone and rod systems for bright- and low-light conditions,
open (Fig. 8.6A). These channels allow a steady influx of respectively.
Passive
Na+ influx
Membrane
Light
cGMP
GTP
GMP
GC
+ +
+
Na+ Rh T PDE
current Ca++
Disc Na+
Na+ Active Hyperpolarization

Na+ Membrane
K+ efflux
B
Fig. 8.6 A, Drawing of a rod with the flow of current in the dark. With the assistance of the Na ,K
+ +
pump, the rod is kept depolarized. B, Sequence of the second messenger events that follow the
absorption of light through the reduction of cGMP. Because cGMP maintains open Na+ channels in the
dark, the results of light absorption are the closing of the Na+ channels and hyperpolarization of the rod.
cGMP, cyclic guanosine monophosphate; GC, guanylate cyclase; GTP, guanosine triphosphate; PDE,
phosphodiesterase; Rh, rhodopsin; T, transducin.
the visual pigment occurs much more rapidly in cones than

AT THE CELLULAR LEVEL in rods. This difference is, at least in part, due to the cones
ability to utilize a second pathway for regeneration (see
Rhodopsin contains a chromophore called retinal, which is
the aldehyde of retinol, or vitamin A. Retinol is derived from
previous section). This more rapid regeneration of visual
carotenoids, such as -carotene, the orange pigment found pigment prevents cones from becoming unresponsive in
in carrots. Like other vitamins, retinol cannot be synthesized bright light conditions. In contrast, the slowness of the
by humans; instead, it is derived from food sources. regeneration of rhodopsin molecules means that at light
Individuals with a severe vitamin A deficiency suffer from levels not much above those found in evening hours, essen-
night blindness, a condition in which vision is defective in
low-light situations.
tially all of the rhodopsin molecules are bleached. Thus in
The extraordinary sensitivity of rods, which can signal the bright-light conditions, only the cone system is functioning,
capture of a single photon, is enhanced by an amplification and the retina is said to be light-adapted.
mechanism in which photoactivation of only one rhodopsin When entering a darkened movie theater, a person can
molecule can activate hundreds of transducin molecules. observe evidence of the existing light adaptation (decreased
In addition, each phosphodiesterase molecule hydrolyzes
thousands of cGMP molecules per second. Similar events
light sensitivity in association with the reduced amount
occur in cones, but the membrane hyperpolarization occurs of rhodopsin) in the inability to see the empty seats (or
much more quickly than in rods and requires thousands of much else). The gradual return of the ability to see the seats
photons. while the person remains in the theater reflects the slow
regeneration of rhodopsin and recovery of function of the
rod system, a process known as dark adaptation.
Dark Adaptation
Light Adaptation This process refers to the gradual increase in light sensitivity
As described previously, absorption of a photon causes 11-cis of the retina when in low-light conditions. Rods adapt to
retinal to be converted to all-trans retinal, which then splits darkness slowly as their rhodopsin levels are restored, and,
from the opsin (bleaching). The visual pigments in rods and indeed, it may take more than 30 minutes for the retina to
cones are bleached at a similar rate; however, regeneration of become fully dark-adapted. In contrast, cones adapt rapidly
to darkness, but their adapted threshold is relatively high, present both in the retina and at higher levels of the visual
and so they do not function when the ambient light level pathway and seem to serve to increase the ability to see the
is low. Thus within 10 minutes in a dark room, rod vision contrast between opposing colors.
is more sensitive than cone vision and becomes the main
system for seeing. Retinal Circuitry
In sum, in the dark-adapted state, primarily rod vision A diagram of the basic circuitry of the retina is shown in
is operative, and thus visual acuity is low and colors are not Fig. 8.7. Several features of this circuitry are noteworthy.
distinguished (this is called scotopic vision). However, when Input to the retina is provided by light striking the pho-
light levels are higher (e.g., when the movie is projected) toreceptors. The output is carried by axons of the retinal
and cone function resumes (this is called photopic vision), ganglion cells to the brain. Information is processed within
visual acuity and color vision are restored. There is an the retina by the interneurons. The most direct pathway
intermediate range of light levels at which rod and cones through the retina is from a photoreceptor to a bipolar cell
are both functional (mesopic vision). and then to a ganglion cell (see Fig. 8.7). More indirect
pathways that provide for intraretinal signal processing
Color Vision involve photoreceptors, bipolar cells, amacrine cells, and
The visual pigments in the cone outer segments contain ganglion cells, as well as horizontal cells to provide lateral
different opsins. As a result of these differences, the three interactions between adjacent pathways.
types of cones absorb light best at different wavelengths.
Although the cone pigments have maximum efficiency IN THE CLINIC
closer to violet, green, and yellow wavelengths, they are
referred to as blue, green, and red pigments, respectively Observations on color blindness are consistent with the
(see Fig. 8.5). The differences in the cone absorption spectra trichromacy theory. In color blindness, a genetic defect
underlie humans ability to see colors, as opposed to only (sex-linked recessive), one or more cone mechanisms are
lost. People with normal color vision are trichromats because
shades of gray. they have three cone mechanisms. Individuals who lack one
According to the trichromacy theory, the differences in of the cone mechanisms are called dichromats. When the
absorption efficiency of the cone visual pigments are pre- long-wavelength cone mechanism is absent, the resulting
sumed to account for color vision because a suitable mixture condition is called protanopia; absence of the medium-
of three colors can produce any other color. However, a wavelength system causes deuteranopia; and absence of the
short-wavelength system causes tritanopia. Monochromats
neural mechanism must also exist for the analysis of color lack two or more cone mechanisms.
brightness because the amount of light absorbed by a visual
pigment, as well as the subsequent response of the cell,
depends on both the wavelength and the intensity of the
light (see Fig. 8.5). Two or three of the cone pigments may Contrasts in Rod and Cone Pathway Functions
absorb a particular wavelength of light, but the amount Rod and cone pathways have several important functional
absorbed by each differs according to its efficiency at that differences in their phototransduction mechanisms and their
wavelength. If the intensity of the light is increased (or retinal circuitry. As described previously, rods have more
decreased), all will absorb more (or less), but the ratio visual pigment and a better signal amplification system
of absorption among them will remain constant. Conse- than cones do, and there are many more rods than cones.
quently, there must be a neural mechanism to compare the Thus rods function better in dim light (scotopic vision), and
absorption of light of different wavelengths by the different loss of rod function results in night blindness. In addition,
types of cones for the visual system to distinguish different all rods contain the same visual pigment, so they cannot
colors. At least two different kinds of cones are required signal color differences. Furthermore, many rods converge
for color vision. The presence of three kinds decreases the onto individual bipolar cells and the results are very large
ambiguity in distinguishing colors when all three absorb receptive fields and low spatial resolution. Finally, in bright
light, and it ensures that at least two types of cones will light, most rhodopsin is bleached, so that rods no longer
absorb most wavelengths of visible light. function under photopic conditions.
The opponent process theory is based on observations Cones have a higher threshold to light and thus are not
that certain pairs of colors seem to activate opposing neural activated in dim light after dark adaptation. However, they
processes. Green and red are opposed, as are yellow and operate very well in daylight. They provide high-resolution
blue, as well as black and white. For example, if a gray vision because only a few cones converge onto individual
area is surrounded by a green ring, the gray area appears bipolar cells in cone pathways. Moreover, no convergence
to acquire a reddish color. Furthermore, a greenish red or occurs in the fovea, where the cones make one-to-one
a bluish yellow color does not exist. These observations connections to bipolar cells. As a result of the reduced
are supported by findings that neurons activated by green convergence, cone pathways have very small receptive fields
wavelengths are inhibited by red wavelengths. Similarly, and can resolve stimuli that originate from sources very
neurons excited by blue wavelengths may be inhibited by close to each other. Cones also respond to sequential stimuli
yellow wavelengths. Neurons with these characteristics are with good temporal resolution. Finally, cones have three
Receptor
layer
Outer nuclear
R R layer
Outer
plexiform
Direction of light layer
B H B Inner
nuclear
layer
A A
Inner
plexiform
layer
Ganglion
G G cell
G layer
Optic fiber
Indirect Direct
pathway pathway
Fig. 8.7 Basic Retinal Circuitry. The arrow at the left indicates the direction of light through the retina.
Photoreceptors (R) synapse on the dendrites of bipolar cells (B) and horizontal cells (H) in the outer
plexiform layer. The horizontal cells make reciprocal synaptic connections with photoreceptor cells and
are electrically coupled to other horizontal cells. Bipolar cells reach synapse on the dendrites of ganglion
cells (G) and on the processes of amacrine cells (A) in the inner plexiform layer. Amacrine cells connect
with ganglion cells and other amacrine cells.
different visual pigments and therefore provide for color organization in which the light that strikes the central
vision. Loss of cone function results in functional blindness; region of the receptive field either excites or inhibits the cell,
rod vision is not sufficient for normal visual requirements. whereas the light that strikes a region that surrounds the
central portion has the converse effect. The receptive field
Synaptic Interactions and Receptive with a centrally located excitatory region surrounded by
Field Organization an inhibitory annulus is called an on-center, off-surround
The receptive field of an individual photoreceptor is circular. receptive field (see Fig. 8.8A). Bipolar cells with such a
Light in the receptive field hyperpolarizes the photorecep- receptive field are described as on bipolar cells. The other
tor cell and cause it to release less neurotransmitter. The type of receptive field has an off-center, on-surround
receptive fields of photoreceptors and retinal interneurons arrangement, which characterizes off bipolar cells (see
determine the receptive fields of the retinal ganglion cells Fig. 8.8F).
onto which their activity converges. The characteristics of The center response of a bipolar cell receptive field is
the receptive fields of retinal ganglion cells constitute an due to only the photoreceptors that directly synapse with
important step in visual information processing because all the bipolar cell. Photoreceptor cells respond to light with
the information about visual events that is conveyed to the hyperpolarization and a decrease in glutamate release and
brain is contained in ganglion cell activity. respond to the removal of light with depolarization and
The bipolar cell, which receives input from a photorecep- increased glutamate release. This implies that the difference
tor, can have either of two types of receptive fields, as shown in the center responses of on and off bipolar cells lies
in Fig. 8.8. Both are described as having a center-surround in their response to glutamate. In fact, off-center bipolar
A On-center field F Off-center field

Light
Central spot
of light
Peripheral
spot
B On-center cell responses G Off-center cell responses
C Central illumination H Central illumination
D Annular illumination I Annular illumination
0 0.5 1.0 1.5 0 0.5 1.0 1.5

Sec Sec
E Diffuse illumination J Diffuse illumination
Fig. 8.8 The receptive fields of on-center (A) and off-center (F) bipolar cells and, below them, the
receptive fields of ganglion cells B through E and G through J to which they are connected. Ganglion cell
responses to central spots (upper recording) and peripheral spots (lower recording) are shown in B and
G. Also shown are responses to central (C and H), surround (D and G), and diffuse whole-field (E and
J) illumination in their receptive fields. The ganglion cells and the on-center and off-center bipolar cells
providing input to these ganglion cells have similar receptive fields, but whereas ganglion cells increase
or decrease their spike frequency, bipolar cells depolarize or hyperpolarize, without generating action
potentials. (Redrawn from Squire LR etal [eds]. Fundamental Neuroscience. San Diego, CA: Academic
Press; 2002.)
cells have ionotropic glutamate receptor channels that open channels. Thus on-center bipolar cells are excited by light
in response to glutamate, and thus they are excited by the stimulation of the center of their receptive fields.
removal of light stimuli from the center of their receptive The antagonistic surround response of bipolar cells is due
field. In contrast, on-center bipolar cells have metabotropic to photoreceptors that surround those that synapse directly
glutamate receptors that close their channels in response to on them. These photoreceptors (which also connect directly
glutamate. They are depolarized by light on the center of with their own bipolar cells) synapse with horizontal cells
their receptive field, because the reduced release of glutamate that participate in complex triadic synapses with many
by the photoreceptors results in more open metabotropic photoreceptors and bipolar cells. The pathway through
the horizontal cells results in a response that is opposite from amacrine and bipolar cells. When amacrine cell input
in sign to that produced directly by the photoreceptors dominates, the receptive fields of ganglion cells tend to be
that mediate the center response. The reason for this is diffuse, and they are either excitatory or inhibitory. Most
that horizontal cells are depolarized by glutamate released ganglion cells, however, are dominated by bipolar cell input
from photoreceptors and thus, like off bipolar cells, are and have a center-surround organization, similar to that of
hyperpolarized in the light. Moreover, because they are the bipolar cells that connect to them (see Fig. 8.8).
electrically coupled to each other by gap junctions, they The distances between retinal components are short.
have very large receptive fields. Darkness in the periphery Hence, modulation of transmitter release by changes in
of a bipolar cells receptive field (such as an annulus that transmembrane potential and the resulting postsynaptic
does not affect the photoreceptors to which it is directly potentials are sufficient for most of the activity in retinal
connected) causes neighboring photoreceptors and hori- circuits, and action potentials are not required except for
zontal cells to depolarize. The depolarized horizontal cells ganglion cells and some amacrine cells, which generate
release gamma-aminobutyric acid (GABA) onto central action potentials. It is unclear why amacrine cells have
(and peripheral) photoreceptor terminals, reducing their action potentials, but ganglion cells must generate them to
release of glutamate. Thus when darkness surrounds central transmit information over the relatively long distance from
illumination, there is increased excitation of on-center the retina to the brain.
bipolar cells. There is a complementary effect on off-center
bipolar cells when a bright annulus surrounds a central dark P, M, and W Cells
spot (see Fig. 8.8). Experiments have shown that in primates, retinal ganglion
Bipolar cells may not respond to large or diffuse cells can be subdivided into three general types called
areas of illumination, covering both the receptors that are P cells, M cells, and W cells. P and M cells are fairly
responsible for the center response and those that cause homogeneous groups, whereas W cells are heterogeneous. P
the surround response because of their opposing actions. cells are so named because they project to the parvocellular
Thus bipolar cells may not signal changes in the intensity layers of the LGN of the thalamus, whereas M cells project
of light that strikes a large area of the retina. On the other to the magnocellular layers of the LGN. P and M cells have
hand, a small spot of light moving across the receptive center-surround receptive fields; hence, they are presumably
field may sequentially and dramatically alter the activity of controlled by bipolar cells. W cells have large, diffuse recep-
the bipolar cell as the light crosses the receptive field from tive fields and slowly conducting axons. They are probably
the surround portion to the center and then back again to influenced chiefly through amacrine cell pathways, but less
the surround portion. This demonstrates that bipolar cells is known about them than about M and P cells.
respond best to the local contrast of stimuli and function Several of the physiological differences among these
as contrast detectors. cell types correspond to morphological differences (Table
Amacrine cells receive input from different combina- 8.1). For example, P cells have small receptive fields (which
tions of on-center and off-center bipolar cells. Thus their corresponds to smaller dendritic trees) and more slowly
receptive fields are mixtures of on-center and off-center conducting axons than M cells do. In addition, P cells show
regions. There are many different types of amacrine cells, a linear response in their receptive field; that is, they respond
and they may use at least eight different neurotransmitters. with a sustained, tonic discharge of action potentials in
Accordingly, the contributions of amacrine cells to visual response to maintained light but do not signal shifts in
processing are complex. the pattern of illumination as long as the overall level of
Ganglion cells may receive dominant input from bipolar illumination is constant. Thus a small object entering a P
cells, dominant input from amacrine cells, or mixed input cells central receptive field will change the cells firing, but
TABLE
8.1 Properties of Retinal Ganglion Cells
Properties P Cells M Cells W Cells
Cell body and axon Medium sized Large Small
Dendritic tree Restricted Extensive Extensive
Receptive field
Size Small Medium Large
Organization Center-surround Center-surround Diffuse
Poorly responsive
Adaptation Tonic Phasic
Linearity Linear Nonlinear
Wavelength Sensitive Insensitive Insensitive
Luminance Insensitive Sensitive Sensitive
Fixation point
Monocular Binocular Monocular

segment segment segment
Retina
Fovea Fovea
Optic
chiasm
Left Right
hemisphere hemisphere
Fig. 8.9 Relationships among a visual target (long arrow, top), images on the retinas of the two eyes
(middle), and projections of the ganglion cells carrying visual information about these images (bottom).
The target image is so large that it extends into the monocular segments of the eyes, where one side of
it is seen by only the ipsilateral eye. Note how the axons are sorted in the chiasm so that all information
about the left visual field of both eyes is conveyed to the right side of the brain and all information about
the right visual field is conveyed to the left side.
continued object movement within the field will not be with larger receptive fields that convey information about
signaled. P cells respond differently to different wavelengths illumination and movement. Both exist in on-center and
of light. Because there are blue, green, and red cones, many off-center varieties (Fig. 8.8).
combinations of color properties are possible, but in fact P
cells have been shown to have opposing responses only to The Visual Pathway
red and green or only to blue and yellow (a combination
of red and green). These mechanisms can greatly reduce Retinal ganglion cells transmit information to the brain by
the ambiguity of color detection caused by the overlap in way of the optic nerve, optic chiasm, and optic tract. Fig.
cone color sensitivity and may provide a substrate for the 8.9 shows the relationships among a visual target, the retinal
opponency process observations. images of the target in the two eyes, and the projections of
M cells, on the other hand, respond with phasic bursts retinal ganglion cells to the two hemispheres of the brain.
of action potentials to the redistribution of light, such as The eyes and the optic nerves, chiasm, and tract are viewed
would be caused by the movement of an object within their from above.
large receptive fields. M cells are not sensitive to differences The visual target, an arrow, is in the visual fields of both
in wavelength but are more sensitive to luminance than P eyes (see Fig. 8.9) and, in this case, is so long that it extends
cells are. into the monocular segments of each retina (i.e., one end
Thus the output of the retina consists primarily of of the target can be seen by only one eye and the other end
ganglion cell axons from (1) sustained, linear P cells with by only the other eye). The shaded circle at the center of
small receptive fields that convey information about color, the target represents the fixation point. The image of the
form, and fine details and (2) phasic, nonlinear M cells target on the retinas is reversed by the lens system. The left
half of the visual target is imaged on the nasal retina of the

left eye and the temporal retina of the right eye. Thus the IN THE CLINIC
left visual field is seen by the left nasal retina and the right Interruption of the visual pathway at any level causes a
temporal retina. Similarly, the right half of the visual target defect in the appropriate part of the visual field (see Fig.
is imaged on and seen by the left temporal retina and the 8.9). For example, a tiny lesion in the retina would result
right nasal retina. The lens system also causes an inversion in a blind spot (scotoma) in that eye, whereas a similar
in the vertical axis, with the upper visual field imaged on lesion in the striate cortex would produce corresponding
scotomas in both eyes. Interruption of the optic nerve on one
the lower retina and vice versa. side produces blindness in that eye. Damage to the optic
The axons of retinal ganglion cells may or may not nerve fibers as they cross in the optic chiasm results in loss
cross in the optic chiasm, depending on the location of of vision in both temporal fields of vision; this condition is
the ganglion cell in the retina (see Fig. 8.9). Axons from known as bitemporal hemianopsia and occurs because
the temporal portion of each retina pass through the optic the crossing fibers originate from ganglion cells in the nasal
halves of each retina. A lesion of the entire optic tract,
nerve, the lateral side of the optic chiasm, and the ipsilateral LGN, visual radiation, or visual cortex on one side causes
optic tract and terminate ipsilaterally in the brain. Axons homonymous hemianopsia, which is loss of vision in the
from the nasal portion of each retina pass through the optic entire contralateral visual field. Partial lesions result in partial
nerve, cross to the opposite side in the optic chiasm, and visual field defects. For example, a lesion in the lingual gyrus
then pass through the contralateral optic tract to end in the causes an upper homonymous quadrantanopsia, which
in this case is loss of vision in the contralateral, upper visual
contralateral side of the brain. As a result of this arrange- field.
ment, objects in the left field of vision are represented in the
right side of the brain, and those in the right field of vision
are represented in the left side of the brain.
on the medial surface of the occipital lobe, just below the
Lateral Geniculate Nucleus calcarine sulcus. Axons in the visual radiation that represent
Retinal ganglion cell axons can synapse in several parts the contralateral lower visual field project to the adjacent
of the brain, but the main target for vision is the lateral cuneus gyrus, which lies just above the calcarine sulcus.
geniculate nucleus (LGN) of the thalamus. There is a Together, the portions of these two gyri that line and border
point-to-point projection from the retina to the LGN. The the calcarine sulcus constitute the primary visual cortex (or
LGN thus has a retinotopic map. Cells that represent a Brodmann area 17; Fig. 8.10).
particular retinal location are aligned along projection lines Like the LGN, the striate cortex contains a retinotopic
that can be drawn across the layers of the LGN. map. The representation of the macula occupies the most
The projection from each eye is distributed to three of the posterior and largest part of both gyri, and progressively
layers of the LGN, one of the magnocellular layers (layers more peripheral areas of the retina are projected to more
1 and 2 receive M cell input) and two of the parvocellular anterior parts of these gyri. Overall, there is an orderly
layers (layers 3 to 6 receive P cell input). Color-coded mapping of retinal loci across the surface of the striate
ganglion cells project to groups of cells between the major cortex (see Fig. 8.10).
layers, the intralaminar zones. Thus the properties of LGN The geniculostriate pathway ends chiefly in layer 4 of the
neurons are very similar to those of retinal ganglion cells. striate cortex (Fig. 8.11), whereas the projection from the
For example, LGN neurons can be classified as P or M intralaminar LGN terminates in so-called blobs in layers 2
cells, and they have on-center or off-center receptive fields. and 3. Similarly, axons that represent one eye or the other
The LGN also receives input from the visual areas of the terminate within layer 4C in alternate adjacent patches that
cerebral cortex, the thalamic reticular nucleus, and several define ocular dominance columns. Cortical neurons in
nuclei of the brainstem reticular formation. The activity of such a column respond preferentially to input from one eye.
LGN projection neurons is inhibited by interneurons both Near the border between two ocular dominance columns,
in the LGN and in the thalamic reticular nucleus. These cells neurons respond about equally to input from the two eyes.
use GABA as their inhibitory neurotransmitter. In addition, The receptive fields of neurons in the striate cortex, aside
the activity of LGN neurons is influenced by corticofugal from the monocular cells in layer 4C, are more complex
pathways and by brainstem neurons that transmit signals than those of LGN neurons. Neurons in other layers may be
via monoamine neurotransmitters. These control systems binocular and respond to stimulation of both eyes, although
filter visual information and may be important for selective the input from one eye often dominates (see Chapter 10).
attention. In addition, cortical neurons outside layer 4C often show
orientation selectivity (i.e., they respond best when the
Striate Cortex stimulus, such as a bar or an edge, is oriented and posi-
The LGN projects to the primary visual cortex or striate tioned in a particular way; Fig. 8.12). These simple cells
cortex by way of the visual radiations. The visual radiation appear to be responding as though they received input from
fibers carrying information derived from the lower half of cells whose concentric center-surround receptive fields were
the appropriate hemiretinas (and therefore the contralateral arranged in such a way that their on centers were aligned
upper visual field) project to the lingual gyrus, which lies in a row flanked by antagonistic regions. Complex cortical
Parieto-occipital sulcus
6
4
5 3 2 Cuneus gyrus
1 Calcarine sulcus
Lingual gyrus
Binocular
portion
5
Monocular 3
1
portion Macula
2
4
Left visual field

Fig. 8.10 The left visual field is relayed (via the LGN and visual radiation) to the primary visual cortex
of the right hemisphere, as a point-to-point retinotopic map. The representation of each part of visual
space is proportional to the number of afferent axons with receptive fields in that part of space. As a
result, the area of macular representation (near the occipital pole) is larger than that for the rest of the
binocular and monocular fields. Note that the lower half of the field is represented in the cuneus gyrus
above the calcarine sulcus and the upper half of the field in the lingual gyrus below the sulcus. (Redrawn
from Purves D, etal [eds]. Neuroscience. 3rd ed. Sunderland, MA: Sinauer; 2004.)
LGN
4C 4B
m cell
2+3 5 6
LGN
4C
p cell
LGN
Blobs
intralaminar
(2+3)
p cell
Cortex Cortex SC LGN
Fig. 8.11 Diagram of visual information flow into the visual cortex from the LGN and its projection
to the extrastriate cortex, to the superior colliculus (SC), and back to the LGN. M, magnocellular path;
P, parvocellular path. (Redrawn from Squire LR, etal [eds]. Fundamental Neuroscience. San Diego, CA:
Academic Press; 2002.)
neurons are similar to simple cells in that they are orienta- cortex all tend to have the same orientation selectivity, they
tion specific, but instead of having flanking excitatory and are considered to form an orientation column (Fig. 8.13).
inhibitory zones, they respond best to a particular stimulus As already discussed, color vision may depend on the
orientation anywhere in their receptive field. They may presence in the retina of three different types of cones, as
also display direction selectivity; that is, they may respond well as neurons in the visual pathway that show spectral
when the stimulus is moved in one direction but not when opposition. Retinal ganglion cells, LGN neurons, and some
it is moved in the opposite direction (see Fig. 8.12). The P cells display spectral opponent properties. The spectral
receptive field of a complex cell may be thought of as a opponent neurons in the striate cortex are found in cortical
composite of adjacent simple cells with similar orientation blobs, and these show double-opponency, in which both the
selectivity. Because such neurons in a particular zone of the center and the surround portions respond antagonistically
C D
B
F
Fig. 8.12 Simple and Complex Receptive Fields in the Visual Cortex Can Be Generated From
Multiple Inputs With Concentric Fields. A and B represent on-center and off-center input, respectively,
from the retina. If three on-center cells (A) with adjacent receptive fields converged onto one cortical
neuron (E), that neuron, a simple cell, would respond best to a long bar stimulus at a specific location and
orientation (C). For three off-center inputs (B), the resulting receptive field is shown in D. The convergence
of multiple simple cells onto another cortical neuron (F) would result in a complex cell that responds best
to a bar stimulus with a vertical orientation that can be placed anywhere within its receptive field. (Redrawn
from Squire LR, etal [eds]. Fundamental Neuroscience. San Diego, CA: Academic Press; 2002.)
to two colors. Such a cell, whose center responds to red but areas include several parallel pathways of visual process-
not green (R+G) and whose surround portion responds ing. The P pathway originates with P cells and functions
to green but not red (RG+), is shown in Fig. 8.13A. The in the recognition of form and color. Structures in the
relationships between the ocular dominance and orienta- P pathway include LGN layers 3 to 6, layer 4C of the
tion columns and the cortical color blobs are shown in striate cortex, V4 (Brodmann area 19), and several areas in
Fig. 8.13B. the inferotemporal region (Fig. 8.14). Processing of form
includes recognition of complex visual patterns, such as
Extrastriate Visual Cortex faces. Color information is processed separately from form.
The M pathway originates with M cells and functions in
In animal studies, at least 25 different visual areas have motion detection and control of eye movement. Cortical
been identified in the cerebral cortex, in addition to the structures in the M pathway include layers 4B and 4C
striate cortex (Brodmann area 17, or V1). The extrastriate of the striate cortex and areas MT (medial temporal) and
Stimulus
duration
I
C
I
C
Cortical blobs,
concerned
with color
Orientation
columns
A B
Fig. 8.13 + +
A, The receptive field and responses of a double-opponent (R G /R G ) neuron in a blob of
the striate cortex as it responds to various combinations of red and green bars. The best on response
is to a red bar flanked by two green bars. B, Diagram of the columnar arrangement of the visual cortex.
Ocular dominance columns are indicated by I (for ipsilateral) and C (for contralateral). Orientation columns
are indicated by the smaller columns marked with short bars at varying angles. The cortical blobs contain
neurons like that in A and have spectral opponent-receptive fields.
it depends on convergent input from the two eyes and the

7 MST MT left/right eye inputs are parallel but segregated in the LGN
Visual and in Layer 4 of striate cortex.
cortex The separation of M and P pathways from the retina
V2 V1 through the thalamus and all the cortical regions raises the
issue of how all the parts are combined to account for the
clear, coherent images of events, objects, and persons that
IT V4 humans perceive. It seems unlikely that all the compo-
Laminae
3-6
nents that represent a percept, such as parts of a face and
Dorsal whether that face belongs to a familiar person, are somehow
M-cell pathway LGN converged onto a single neuron that recognizes it. Rather,
P-cell pathway Laminae complex percepts probably arise from the coordinated
1,2
activity of large sets of neurons across multiple regions
Fig. 8.14 Distribution of P and M cell influences on different areas
of the CNS. The process by which a binding of such
of the visual cortex. IT, inferotemporal area; MST, medial superior disparate neuronal elements into a percept is unclear, but
temporal area; MT, medial temporal area; V1, striate cortex; V2 and one working hypothesis is that it may be accomplished
V4, higher order visual areas. by the temporal synchronization of many anatomically
distributed neural events.
MST (medial superior temporal) on the lateral aspect of the

temporal lobe, as well as Brodmann area 7a of the parietal
lobe (see Fig. 8.14). IN THE CLINIC
Both P and M pathways contribute to depth perception Lesions of the extrastriate visual cortex can produce various
or stereopsis, which is dependent on slight differences in deficits. Bilateral lesions of the inferotemporal cortex can
the retinal images formed in the two eyes. Stereopsis is result in cortical color blindness (achromatopsia) or in an
useful only for relatively nearby objects. However, in such inability to recognize faces, even of close members of the
cases, these disparities provide visual cues about depth. family (prosopagnosia). A lesion in area MT or MST can
interfere with motion detection and eye movements.
Interestingly, the anatomy of the visual pathways indicates
that depth perception must be a cortical function because
One cycle = tc
(frequency = 1/tc)
Pressure
Amplitude
Phase
difference
Time
Fig. 8.15 Two pure tones are shown by the solid and dashed lines. Frequency is determined from the
wavelength as indicated. Amplitude is the peak-to-peak change in sound pressure. The two tones have
the same frequency and amplitude but differ in phase.
Other Visual Pathways Audition

The superior colliculus of the midbrain is a layered structure Sound
that is important for certain types of eye movements (see Sound is produced by compression and decompression
Chapter 9). The three most superficial layers are involved waves in air or in other elastic media, such as water. Sound
exclusively in visual processing, whereas the deeper layers frequency is measured in cycles per second, or hertz (Hz).
receive multimodal input from the somatosensory and audi- Each pure tone results from a sinusoidal wave at a particular
tory systems, as well as the visual system, particularly from frequency and is characterized not only by its frequency
cortical areas involved in eye movement. but also, instantaneously, by its amplitude and phase
Another retinal projection is to the pretectum, which (Fig. 8.15). Most naturally occurring sound, however, is
bilaterally activates parasympathetic preganglionic neurons a mixture of pure tones. Noise is unwanted sound and
in the Edinger-Westphal nucleus that cause pupillary con- may have any composition of pure tones. Sound propagates
striction in the pupillary light reflex. The pretectal areas are at about 335m/sec in air. The waves are associated with
also interconnected through the posterior commissure, and certain pressure changes, called sound pressure. The unit of
thus the reflex causes both ipsilateral (direct) and contra- sound pressure is Newtons per meter squared (N/m2), but
lateral (consensual) pupillary constriction when a light is sound pressure is more commonly expressed as the sound
shown in one eye. pressure level (SPL). The unit of SPL is the decibel (dB):
The visual pathways also include connections to nuclei Equation 8.1
that serve functions other than vision. For example, a
retinal projection to the suprachiasmatic nucleus of SPL = 20log P PR
the hypothalamus controls circadian rhythmicity (see where P is sound pressure and PR is a reference pressure
Chapter 37). (0.0002 dyne/cm2, the absolute threshold for human hearing
at 1000Hz). A sound with intensity 10 times greater would
The Auditory and Vestibular Systems be 20dB; one 100 times greater would be 40dB.
The normal young human ear is sensitive to pure
The peripheral parts of the auditory and vestibular systems tones with frequencies that range between about 20 and
share components of the bony and membranous labyrinths, 20,000Hz. The threshold for detection of a pure tone varies
use hair cells as mechanical transducers, and transmit infor- with its frequency (Fig. 8.16). The lowest thresholds for
mation to the CNS through the vestibulocochlear nerve human hearing are, for pure tones, approximately 3000Hz.
(CN VIII). However, the CNS processing and sensory The threshold at these frequencies is approximately 3 to
functions of the auditory and vestibular systems are distinct. 5dB, in comparison with the reference 0dB at 1000Hz.
The function of the auditory system is to transduce sound. In reference to this scale, normal speech has an intensity of
This allows us to recognize environmental cues and to com- about 65dB, and its main frequencies fall in the range of
municate with other organisms. The most complex auditory 300 to 3500Hz. Sounds that exceed 100dB can damage
functions are those involved in language. The function of the peripheral auditory apparatus, and those higher than
the vestibular system is to provide the CNS with informa- 120dB can cause pain and permanent damage. As people
tion related to the position and movements of the head in age, their thresholds at high frequencies rise, thereby
space. Control of eye movement by the vestibular system is reducing their ability to hear such tones, a condition called
discussed in Chapter 9. presbycusis.
140
The Ear
Pain
120 The peripheral auditory apparatus is the ear, which can be
subdivided into the external ear, the middle ear, and the
100 inner ear (Fig. 8.17).
Intensity level (db)
80 External Ear
The external ear includes the pinna and the external auditory
60 meatus (auditory canal). The auditory canal contains glands
that secrete cerumen, a waxy protective substance. The
40
Hearing pinna helps direct sounds into the auditory canal and plays
20
a role in sound localization. The auditory canal transmits
the sound pressure waves to the tympanic membrane. In
0 humans, the auditory canal has a resonant frequency of
about 3500Hz, and this resonance contributes to the low
10 100 1000 10000 perceptual threshold for sounds in that range.
Frequency (cps)
Middle Ear
Fig. 8.16 Sound Threshold Intensities at Different Frequencies.
The external ear is separated from the middle ear by the
The bottom curve indicates the absolute intensity needed to detect tympanic membrane (see Fig. 8.17A). The middle ear
a sound. The dashed curve represents the threshold for functional
contains air. Three ossicles are present and serve to link the
hearing. The top curve indicates levels at which sound is painful and
damaging. tympanic membrane to the oval window of the inner ear.
Semicircular
canals
Malleus Incus Vestibule (saccule and utricle)
Vestibular
nerve
Facial
nerve
External
auditory
meatus Auditory
nerve
Cochlea
Pinna
Tympanic Middle Round

Eustachian
membrane ear window
tube
cavity
A
Helicotrema
Scala media
Malleus Incus
Basilar membrane
and organ of Corti
Scala tympani
Fig. 8.17 Ear and Cochlear
Scala vestibuli Structure. A, Location of the
right human cochlea in relation
to the vestibular apparatus, the
Tympanic Round middle ear, and the external ear.
membrane window B, Relationships between the outer,
Stapes (sitting in
an oval window) Eustachian tube middle, and inner ear spaces; the
cochlea is depicted unrolled for
B clarity.
Inner Tectorial
hair cell membrane
Outer
hair cells
Stria
vascularis
Scala vestibuli Scala
media
Reissners
membrane
Scala tympani
Basilar
membrane
Spiral
lamina
Spiral Rods
ganglion Nerve fibers of Corti
C
Fig. 8.17, contd C, Drawing of a cross-section through the cochlea. The organ of Corti (see Fig.
8.18A and B) is outlined.
Adjacent to the oval window is the round window, another advantage of the lever system formed by the ossicles. This
membrane-covered opening between the middle ear and impedance matching is sufficient to increase the efficiency
inner ear (see Fig. 8.17A and B). of energy transfer by nearly 30dB in the range of hearing
The ossicles include the malleus, the incus, and the from 300 to 3500Hz.
stapes. The stapes has a footplate that inserts into the oval
window. Behind the oval window is a fluid-filled component
of the inner ear, the vestibule. It is continuous with a tubular
structure known as the scala vestibuli. Inward movement IN THE CLINIC
of the tympanic membrane by a sound pressure wave causes
the chain of ossicles to push the footplate of the stapes into The middle ear also serves other functions. Two muscles
are found in the middle ear: the tensor tympani attached
the oval window (see Fig. 8.17B). This movement of the to the malleus and the stapedius attached to the stapes.
stapes footplate in turn displaces the fluid within the scala When these muscles contract, they damp movements of
vestibuli. The pressure wave that ensues within the fluid is the ossicles and decrease the sensitivity of the acoustic
transmitted through the basilar membrane of the cochlea apparatus. This action can protect the acoustic apparatus
to the scala tympani (described later), and it causes the against damaging sounds that can be anticipated. However,
a sudden explosion can still damage the acoustic apparatus
round window to bulge into the middle ear. because reflex contraction of the middle ear muscles does
The tympanic membrane and the chain of ossicles serve not occur quickly enough. The chamber of the middle
as an impedance-matching device. The ear must detect ear connects to the pharynx through the eustachian tube.
sound waves traveling in air, but the neural transduction Pressure differences between the external ear and middle
mechanism depends on movement in the fluid-filled cochlea, ear can be equalized through this passage. If fluid collects
in the middle ear, as during an infection, the eustachian tube
where acoustic impedance is much higher than that of air. may become blocked. The resulting pressure difference
Therefore, without a special device for impedance match- between the external ear and middle ear can produce
ing, most sound reaching the ear would simply be reflected, painful displacement of the tympanic membrane and, in
as are voices from shore when a person is swimming under extreme cases, cause rupture of the tympanic membrane.
water. Impedance matching in the ear depends on (1) the Unequalized pressure changes as a result of flying or diving
can also cause discomfort.
ratio of the surface area of the large tympanic membrane
to that of the smaller oval window and (2) the mechanical
Inner Ear nerve fibers penetrate the organ of Corti and terminate at
The inner ear includes the bony and membranous laby- the bases of the hair cells (Fig. 8.18; see also Fig. 8.17C).
rinths. The bony labyrinth is a complex but continuous Approximately 90% of the fibers end on inner hair cells, and
series of spaces in the temporal bone of the skull, whereas the remainder end on outer hair cells. Thus approximately
the membranous labyrinth consists of a series of soft tissue 10 afferent fibers supply each inner hair cell, whereas other
spaces and channels lying inside the bony labyrinth. The afferent fibers diverge to supply about five outer hair cells
cochlea and the vestibular apparatus are formed from these each. The inner hair cells clearly provide most of the neural
structures. information about acoustic signals that the CNS processes
The cochlea is a spiral-shaped organ (see Fig. 8.17A for hearing. The sensory function of the outer hair cells is
and B). In humans, the spiral consists of 2 3 4 turns from a less clear.
broad base to a narrow apex, although its internal lumen In addition to afferent fibers, the organ of Corti is sup-
is small at the base and wide at the top. The apex of the plied by efferent fibers, most of which terminate on the
cochlea faces laterally (see Fig. 8.17A). The bony labyrinth outer hair cells. These cochlear efferent fibers originate in
component of the cochlea is subdivided into several cham- the superior olivary nucleus of the brainstem and are often
bers. The vestibule is the space facing the oval window (see called olivocochlear fibers. The length of the outer hair
Fig. 8.17A). Continuous with the vestibule is the scala cells varies; this characteristic suggests that changes in outer
vestibuli, the spiral-shaped chamber that extends to the apex hair cell length may affect the sensitivity, or tuning, of the
of the cochlea, where it meets and merges with the scala inner hair cells. The cochlear efferent fibers may control
tympani at the helicotrema. The scala tympani is another outer hair cell length. Such a mechanism could conceivably
spiral-shaped space that winds back down the cochlea and influence the sensitivity of the cochlea and the way that the
ends at the round window (see Fig. 8.17B). Separating the brain recognizes sound. Other efferent fibers that end on
two, except at the helicotrema, is the scala media enclosed cochlear afferent fibers may be inhibitory, and they may
in the membranous labyrinth. help improve frequency discrimination.
The scala media, or cochlear duct (see Fig. 8.17B and Sound is transduced by the organ of Corti. Sound waves
C), is a membrane-bound spiral tube that extends along the that reach the ear cause the tympanic membrane to oscillate,
cochlea, between the scala vestibuli and scala tympani. One and these oscillations are transmitted to the scala vestibuli
wall of the scala media is formed by the basilar membrane, by the ossicles. This creates a pressure difference between the
another by Reissners membrane, and the third by the stria scala vestibuli and the scala tympani (see Fig. 8.17B) that
vascularis (see Fig. 8.17C). serves to displace the basilar membrane and, with it, the
The spaces within the cochlea are filled with fluid. The organ of Corti (see Fig. 8.18A and B). Because of the shear
fluid in the bony labyrinth, including the scala vestibuli forces set up by the relative displacement of the basilar and
and scala tympani, is perilymph, which closely resembles tectorial membranes, the stereocilia of the hair cells bend.
cerebrospinal fluid. The fluid in the membranous labyrinth, Upward displacement bends the stereocilia toward the
including the scala media, is endolymph, which is very tallest cilium, which depolarizes the hair cells; downward
different from perilymph. Endolymph, generated by the deflection bends the stereocilia in the opposite direction,
stria vascularis, contains high [K+] (about 145mM) and which hyperpolarizes the hair cells.
low [Na+] (about 2mm) and has a high positive potential
(about +80mV) with regard to the perilymph. As a result, IN THE CLINIC
a very large potential gradient (about 140mV) exists across
the membranes of the hair cell cilia that extend into the A common cause of deafness is the destruction of hair cells
endolymph. (These hair cells, which are the sensory recep- by loud sounds. Hair cells can be destroyed, for example,
by exposure to industrial noise or by listening to loud
tors for sound, are discussed in more detail later.)
music. Typically, hair cells in certain parts of the cochlea are
The neural apparatus responsible for transduction of selectively damaged by exposure to high levels of sound at
sound is the organ of Corti (see Fig. 8.17C), which is particular frequencies (as predicted by the place theory), and
located within the cochlear duct. It lies on the basilar mem- thus hearing may be lost over a discrete frequency range.
brane and consists of several components, including three Presbycusis, or the loss of high-frequency hearing with age,
is probably increased by the loss of hair cells as a result of
rows of outer hair cells, a single row of inner hair cells,
long-term noise exposure in urban environments.
a gelatinous tectorial membrane, and a number of types
of supporting cells. The organ of Corti in humans contains
15,000 outer and 3500 inner hair cells. The rods of Corti
help provide a rigid scaffold. Located on the apical surface Sound Transduction
of the hair cells are stereocilia, which can be described as In view of the wide range of frequencies and amplitudes
nonmotile cilia that contact the tectorial membrane. of sound stimuli, it is no surprise that hair cell transduc-
The organ of Corti is innervated by nerve fibers of the tion must provide for a fast response. The fast response
cochlear division of the vestibulocochlear nerve (CN VIII). to deflection of the cilia is based on direct opening of
The 32,000 auditory afferent fibers in humans originate ion channels by so-called tip links that connect the tip of
in sensory ganglion cells in the spiral ganglion. These each stereocilium with the shaft of the next taller one (see
Tectorial K+
Inner membrane Reticular
hair cell lamina
Depolarization
Basilar Nucleus
membrane
Rods Outer
of Corti hair cells
A
Vesicles
Ca++
Transmitter
Afferent
nerve
B C
Fig. 8.18 Detail of the organ of Corti at rest (A) and with upward movement of the basilar membrane
(B). The upward movement causes the stereocilia to bend because of shear forces produced by
relative displacement of the hair cells and the tectorial membrane. C, Diagram of a hair cell with tip link
connections between the hair cell cilia to show how shear forces open mechanoreceptor channels and
depolarize the hair cell.
Fig. 8.18C). With deflection, the tip links are subjected to these cells. This current flow can be recorded extracellularly
a lever action that transiently opens the channels, allows as a cochlear microphonic potential, an oscillatory event
the entry of K+ (because of the high [K+] and high poten- that has the same frequency as the acoustic stimulus. The
tial in endolymph), and depolarizes the hair cell. Several cochlear microphonic potential represents the sum of the
mechanisms have been proposed to account for the equally receptor potentials of a number of hair cells.
important rapid adaptation necessary for a high-frequency Hair cells, like retinal photoreceptors, release an
response. A spring response by the tip links would allow excitatory neurotransmitter (probably glutamate) when
the attachment point of the tip link to be moved along the depolarized. The neurotransmitter produces an excitatory
stereociliums shaft to reset the mechanical leverage of the postsynaptic potential (EPSP) in the cochlear afferent nerve
tip link. In addition, it has been observed that Ca++ can fibers with which the hair cell synapses. In summary, sound
enter and bind to the open channel, change it to require is transduced when oscillatory movements of the basilar
greater opening force, and thereby reduce the statistical membrane cause transient changes in the transmembrane
probability of opening. voltage of the hair cells and, finally, the generation of action
The potential gradient that induces movement of ions potentials in cochlear afferent nerve fibers. The activity of
into hair cells includes both the resting potential of the hair a large number of cochlear afferent fibers in the auditory
cells and the positive potential of the endolymph. As noted nerve can be recorded extracellularly as a compound action
previously, the total gradient across the apical membrane potential.
of hair cells is about 140mV. Therefore, a change in K+ On the basis of differences in width and tension,
conductance in the apical membranes of hair cells results in investigators originally concluded that different parts of
a rapid current flow that produces the receptor potential in the basilar membrane have different resonant frequencies.
Fig. 8.19 Different Frequencies of Sound Result in Different
Amplitudes of Displacement at Different Sites Along the Organ

of Corti. A, Traveling wave produced in the basilar membrane by a
200 Hz a
sound of 200Hz. The curves at a, b, c, and d represent displacement b
of the basilar membrane at different times, and the dashed line is the c
envelope formed by the peaks of the wave at different times. Maximum d
deflection occurs at about 29mm from the oval window. B, Envelopes
of traveling waves produced by several frequencies of sound. Note that
the maximum displacement varies with frequency and is closest to the
stapes when the frequency is highest. (Redrawn from von Bekesy G.
Experiments in Hearing. New York: McGraw-Hill; 1960.)
20 25 30
For example, the basilar membrane is about 100m wide Distance from stapes (mm)
at the base and 500m wide at the apex. It also has higher A
tension at the base. Thus, the investigators predicted that
the base would vibrate at higher frequencies than would
the apex, as do the shorter strings of musical instruments.
Experiments have shown that the basilar membrane moves
as a whole in traveling waves (Fig. 8.19), but displacement
of the basilar membrane is maximal nearer the base of the 25 Hz
cochlea during high-frequency tones and maximal nearer
the apex during low-frequency tones.
In effect, the basilar membrane serves as a frequency ana-
lyzer; it distributes the stimulus along the organ of Corti,
and different hair cells respond differentially to particular 50 Hz
frequencies of sound. This is the basis of the place theory
of hearing. In addition, hair cells located at different places
along the organ of Corti may be tuned to different frequen-
cies because of variations in their stereocilia and biophysical
properties. As a result of these factors, the basilar mem- 100 Hz
brane and organ of Corti have a so-called tonotopic map
Relative amplitude
(Fig. 8.20).
Cochlear Nerve Fibers
Neurotransmitter release by hair cells in the organ of Corti
can evoke action potentials in the primary afferent fibers of 200 Hz
the cochlear nerve. Afferent fibers in the vestibulocochlear
nerve (CN VIII) are bipolar cells with a myelin sheath
around the cell bodies, as well as around the axons. The cell
bodies are in the spiral ganglion, their peripheral processes
synapse at the base of hair cells, and their central processes 400 Hz
synapse in the cochlear nuclei of the brainstem.
Characteristic Frequencies
A cochlear afferent fiber discharges maximally when
stimulated by a particular sound frequency called its char- 800 Hz
acteristic frequency. The characteristic frequency can be
determined from a tuning curve for the fiber (Fig. 8.21).
A tuning curve is a plot of the threshold for activation of 3
the nerve fiber by different sound frequencies. The major
factor that influences the activity of individual afferent
fibers is the location along the basilar membrane of the hair 1600 Hz 0
cells that they innervate. The location of those hair cells is 0 10 20 30
important because for any given sound frequency, there is Distance from stapes (mm)
a site of maximum displacement of the basilar membrane B
as the pressure wave travels along its length (see Fig. 8.19).
Typically, tuning curves are sharp near the characteristic
4,000
800
00
3,0 0 I E
60 20
Threshold intensity (dB)

Apex
5,000 I
2,000
40
0 40
0 20 0
1,00
1,500 60
00
7,0
80
2 4 7 10 15
A Frequency (kHz)
0
00
Threshold intensity (dB)

,0 20
20
40
Base
Fig. 8.20
The tonotopic map of the cochlea. (Redrawn from Stuhl- 60
man O. An Introduction to Biophysics. New York: John Wiley & Sons;
1943.) 80
100
frequency, but they broaden at high sound pressure levels.
Tuning curves can have excitatory and inhibitory areas (see 0.2 0.5 1 2 5 10 20
Fig. 8.21A). The sharpness of the excitatory regions may B Frequency (kHz)
reflect inhibitory processes.
Fig. 8.21 Tuning Curves of Neurons in the Auditory System.
Tuning curves can be considered as receptive field plots. A, Tuning

Encoding
curve with central excitatory frequencies (E) and flanking inhibitory
The different features of an acoustic stimulus are encoded in frequencies (I). B, Tuning curves for cochlear nerve fibers. (A, Redrawn
the discharges of cochlear nerve fibers. Duration is signaled from Arthur RM, etal. J Physiol [Lond] 1971;212:593. B, Redrawn
by the duration of activity; intensity is signaled both by from Katsui Y. In: Rosenblith WA [ed]. Sensory Communication.
the amount of neural activity and by the number of fibers Cambridge, MA: MIT Press; 1961.)
that discharge. For low-frequency sounds, the frequency is
signaled by the tendency of an afferent fiber to discharge
in phase with the stimulus (phase locking; see Fig. 8.22A).
If the tone is much more than 1 kHz, a single fiber cannot
discharge with every cycle, but phase locking can also occur IN THE CLINIC
for sounds with periods shorter than the absolute refrac-
tory period of the afferent fiber. This allows the CNS to An important, although relatively uncommon, condition
that can interrupt the function of cochlear nerve fibers is
detect higher frequency information from the activity of a an acoustic neuroma, a tumor of Schwann cells of the
population of afferent fibers, each of which discharges in vestibulocochlear nerve (CN VIII). As the tumor grows,
phase with the stimulus and which, as a group, signal the irritation of cochlear nerve fibers may cause a ringing sound
frequency of the stimulus (see Fig. 8.22B). This observation in the affected ear (tinnitus). Eventually, conduction in
is the basis of the frequency theory of hearing. cochlear nerve fibers is blocked, and the ear becomes deaf.
The tumor may be operable while still small; therefore, early
For still higher frequencies (>5000Hz), the place theory diagnosis is important. If the tumor is allowed to enlarge
dominates: the CNS interprets sounds that activate affer- substantially, it could interrupt the entire vestibulocochlear
ent fibers supplying hair cells near the base of the cochlea nerve and cause vestibular as well as auditory difficulties. It
as being of high frequency. Thus both the place and the could also impinge on or distort neighboring cranial nerves
frequency theories are necessary to explain the frequency (e.g., V, VII, IX, and X), and it could produce cerebellar signs
by compressing the cerebellar peduncles.
coding of sound (duplex theory) across the entire range
from 20 to 20,000Hz.
(see Fig. 8.21B). By plotting the distribution of the char-

acteristic frequencies of neurons within a nucleus or in the
auditory cortex, a tonotopic map may be revealed in which
Afferent 1
neurons are ordered according to their best frequencies.
Tonotopic maps have been found in the cochlear nuclei,
Afferent 2 superior olivary complex, inferior colliculus, medial genicu-
late nucleus, and auditory cortex. A given auditory structure
A Low-frequency sounds
may, in fact, contain several tonotopic maps.
Binaural Interactions
Most auditory neurons at levels above the cochlear nuclei
Afferent 1 respond to stimulation of either ear (i.e., they have binau-
ral receptive fields). Binaural receptive fields contribute
Afferent 2 to sound localization. A human can distinguish sounds
originating from sources separated by as little as 1 degree.
Afferent 3 The auditory system uses several clues to judge the origin
of sounds, including differences in the time (or phase) of
Sum
arrival of the sound at the two ears and differences in sound
intensity on the two sides of the head.
B High-frequency sounds
For example, neurons in the medial superior olivary
Fig. 8.22 A, At low frequencies, individual auditory afferent fibers nucleus have medial and lateral dendrites. The synapses on
can respond at each cycle to the signal frequency. B, At higher the medial dendrites are largely excitatory, and they origi-
frequencies, each afferent fiber generates an action potential only at
nate from the contralateral ventral cochlear nucleus. Those
certain cycles, limited by its maximum firing frequency. However, the
overall population of afferent fibers can still signal stimulus frequency on the lateral dendrites are mostly inhibitory and come
by their aggregate firing frequency. from the ipsilateral ventral cochlear nucleus. Differences
in the phase of the sound reaching the two ears affect the
strength and timing of the excitation and inhibition reach-
Central Auditory Pathway ing a particular medial superior olivary neuron. The lateral
Cochlear afferent fibers synapse on neurons of the dorsal superior olivary nucleus processes differences in the sound
and ventral cochlear nuclei. The neurons in these nuclei intensity that reaches the two ears to provide information
have axons that contribute to the central auditory pathways. about the source of the sound. The activity of superior
Some of the axons from the cochlear nuclei cross to the olivary neurons can provide information about sound
contralateral side and ascend in the lateral lemniscus, the localization.
main ascending auditory tract. Others connect with various
ipsilateral or contralateral nuclei, such as the superior Cortical Organization
olivary nuclei, which project through the ipsilateral and Several features of the primary auditory cortex resemble
contralateral lateral lemnisci. Each lateral lemniscus ends those of other primary sensory areas. Not only are sensory
in an inferior colliculus. Neurons of the inferior colliculus mapsin this case, tonotopic mapspresent in the audi-
project to the medial geniculate nucleus of the thalamus, tory cortex but also this cortical region performs feature
which gives rise to the auditory radiation. The auditory extraction. Neurons in the primary auditory cortex form
radiation ends in the primary auditory cortex (Brodmann isofrequency columns (in which the neurons in the column
areas 41 and 42), located on the superior surface of the have the same characteristic frequency), and they also form
temporal lobe. alternating columns, known as summation and suppres-
The input from each ear is bilaterally represented in sion columns. Neurons in summation columns are more
the ascending auditory system pathway at the level of the responsive to binaural than to monaural input. Neurons in
lateral lemniscus and above. Thus the representation of suppression columns are less responsive to binaural than
auditory space is complex, even at the brainstem level. to monaural stimulation, and, accordingly, the response to
Consequently, unilateral deafness may occur with isolated one ear is dominant. Some neurons are selective for the
lesions of the cochlear nuclei or more peripheral structures. direction of frequency change.
Central lesions do not cause unilateral deafness, although Bilateral lesions of the auditory cortex have some effect
they may interfere with overall sensitivity to speech or with on the ability to distinguish the frequency or intensity of
sound localization. different sounds, and they reduce the abilities to local-
ize sound and to understand speech. Unilateral lesions,
Functional Organization of the Central however, have little effect, especially if the nondominant
Auditory System (for language) hemisphere is involved (see Chapter 10).
Receptive Fields and Tonotopic Maps Evidently, frequency discrimination depends on activity at
The responses of neurons in several structures that belong lower levels of the auditory pathway, possibly the inferior
to the auditory system can be described by tuning curves colliculus.
the bony labyrinth. The vestibular apparatus on each side

IN THE CLINIC is composed of three semicircular canals and two otolith
Two simple tests are often used clinically to distinguish
organs (Fig. 8.23; see also Fig. 8.17A). These structures
the most important types of deafness: conduction loss contain endolymph and are surrounded by perilymph. The
and sensorineural loss. Conduction hearing loss occurs semicircular canals are named the horizontal, anterior, and
in disorders of the external ear (e.g., ear canal blocked posterior canals. The otolith organs are the utricle and the
by cerumen) or middle ear (e.g., rupture of the eardrum). saccule (together indicated as Vestibule in Fig. 8.17A).
Sensorineural hearing loss reflects disorders of the inner ear,
the cochlear nerve, or central connections.
Each semicircular canal has a swelling called an ampulla
The Weber test is used to evaluate the magnitude of at the point where it joins the utricle. The saccule connects
conduction hearing loss. In this test, the base of a vibrating with the cochlea, through which endolymph (produced by
tuning fork is placed against the middle of the persons the stria vascularis of the cochlea) can reach the vestibular
forehead, and the person is asked to localize the sound. apparatus.
Normally, the sound is not localized to a particular ear.
However, if the person has conductive hearing loss (e.g., due
The three semicircular canals on one side are matched
to a punctured tympanic membrane, fluid in the middle ear, with corresponding coplanar semicircular canals on the
otosclerosis, or loss of continuity of the ossicular chain), the other side. The horizontal canals on each side of the head
sound is localized to the deaf ear because it is conducted correspond, as do the anterior canal on one side and the
to the cochlea through bone. The sound is also conducted posterior canal on the other side (see Fig. 8.23B). This
to the cochlea of the undamaged ear, but bone-conducted
sound does not activate the organ of Corti as well as does
arrangement allows the sensory epithelia, in corresponding
sound conducted normally through the tympanic membrane pairs of canals on the two sides, to cooperate in sensing
and ossicle chain. One reason why the sound in the Weber acceleration of the head about three nearly orthogonal axes
test is not localized to the normal ear may be that hearing in space. The horizontal canals are not truly horizontal;
in the normal ear is inhibited by the ambient sound level rather, they lie in the horizontal plane if the head is tilted
(auditory masking). Conversely, in people with sensorineural
hearing loss (e.g., due to damage to the organ of Corti, the
down 30 degrees in relation to the horizon.
cochlear nerve, or the cochlear nuclei), the sound is localized The ampulla of each of the semicircular canals contains a
to the normal side. sensory epithelium called a crista ampullaris, or ampullary
In the Rinne test, a vibrating tuning fork is placed against crest (Fig. 8.24). An ampullary crest consists of a ridge,
the bone behind the persons ear, and the person is asked transverse to the long axis of the canal, that is covered by
to indicate when the sound dies out. The tuning fork is then
held near the external auditory meatus of that ear. In people
epithelium containing vestibular hair cells. These hair cells
with normal hearing, the sound is again heard because the are innervated by primary afferent fibers of the vestibular
sound is more effectively transmitted to the cochlea in air nerve, which is a subdivision of the vestibulocochlear nerve
(i.e., air conduction > bone conduction). If the conduction (CN VIII).
mechanism is damaged, the sound is not heard when the Like cochlear hair cells, each vestibular hair cell contains
tuning fork is held near the external auditory meatus. Bone
conduction in this case is better than air conduction. If the
a set of stereocilia on its apical surface. However, unlike
hearing loss is sensorineural, the sound is heard again when cochlear hair cells, vestibular hair cells also contain a large
the tuning fork is placed by the external auditory meatus single kinocilium. The cilia on ampullary hair cells are
because with sensorineural hearing loss, the inner ear and embedded in a gelatinous structure called the cupula. The
cochlear nerve are less able to transmit impulses regardless cupula and the crista occlude the lumen of the ampulla
of whether the sound vibrations reach the cochlea via air or
bone. Thus because air conduction is more effective than
completely. Movement of endolymph, produced by angular
bone conduction, the bone conduction pattern seen with acceleration of the head about an axis perpendicular to the
sensorineural hearing loss is the same as in a normal ear. plane of the canal, deflects the cupula and consequently
bends the cilia on the hair cells. The cupula has the same
specific gravity as endolymph, and thus it is unaffected by
The Vestibular System linear acceleratory forces, such as gravity.
The sensory epithelia of the otolith organs are called the
The vestibular system detects angular and linear accelera- macula utriculi and the macula sacculi (Fig. 8.25). The
tions of the head. Signals from the vestibular system allow utricle is oriented nearly horizontally; the saccule is oriented
the body to make adjustments in posture that maintain vertically. Their hair cells are embedded in the epithelium
balance and trigger head and eye movements to stabilize that overlies each macula. As in the ampullary crests, the ste-
the visual image on the retina. The following description reocilia and kinocilia of the macula project into a gelatinous
of the vestibular system emphasizes the sensory aspects of mass. However, the gelatinous mass in the macula contains
vestibular function, and it introduces the central vestibular numerous otoliths (ear stones) composed of calcium
pathways. The role of the vestibular apparatus in motor carbonate crystals. Together, the gelatinous mass and its
control is discussed in Chapter 9. otoliths are known as an otolithic membrane. The otoliths
increase the specific gravity of the otolithic membrane to
The Vestibular Apparatus about twice that of the endolymph. Hence, the otolithic
Structure of the Vestibular Labyrinth membrane tends to move when subjected to acceleration,
The vestibular apparatus, like the cochlea, consists of a whether linear (such as that produced by gravity) or angular,
component of the membranous labyrinth located within particularly when the center of rotation is outside the head.
Left superior Right superior

semicircular canal plane
canal
Horizontal Temporal
semicircular bone
canal
90
Foramen
magnum
Left posterior
semicircular
canal
Right posterior
canal plane
A B
Fig. 8.23 A, Lateral view of the right semicircular canals of a rhesus monkey that were dissected
after being filled with plastic. Note the ampullae associated with each canal. Scale is in millimeters.
B, Overhead view of the base of the skull showing the orientation of structures of the inner ear. Coplanar
pairs of semicircular canals include the horizontal canals, as well as the anterior and contralateral posterior
canals. (A, Courtesy of Dr. John Simpson, New York University School of Medicine. B, Redrawn from
Haines DE [ed]. Fundamental Neuroscience for Basic and Clinical Applications. 3rd ed. Philadelphia:
Churchill Livingstone; 2006.)
Ampulla Cupula
Crista ampullaris
Cilia
Receptor cells
Endolymph
Head acceleration
Fig. 8.24 Drawing of an Ampullary Crest Inside an Ampulla. The stereocilia and the kinocilium of
each hair cell extend into the cupula, which extends across the entire cross-section of the ampulla. Head
movement (acceleration) to the right would result in endolymph pressure to the left and deflection of the
cupula to the left.
Innervation of Sensory Epithelia of the branches to each of the vestibular epithelia; centrally, it
Vestibular Apparatus accompanies the cochlear and facial nerves as they enter the
The cell bodies of the primary afferent fibers of the vestibular internal auditory meatus of the skull.
nerve are located in the Scarpa ganglion. The neurons are
bipolar, and their cell bodies, as well as axons, are myelin- Vestibular Transduction
ated. Peripherally, the vestibular nerve gives off separate Like cochlear hair cells, vestibular hair cells are functionally
polarized, and the transduction mechanism is presumed to
be similar. When the stereocilia are bent toward the longest
cilium (in this case, the kinocilium), conductance of the
apical membrane increases for cations and, because of the
high K+ concentration of the endolymph, K+ enters, and
the vestibular hair cell is depolarized (Fig. 8.26). Conversely,
when the cilia are bent away from the kinocilium, the hair
cell is hyperpolarized. The hair cell releases an excitatory
neurotransmitter (either glutamate or aspartate) toni-
cally, so that the afferent fiber on which it synapses has a
resting discharge. When the hair cell is depolarized, more
neurotransmitter is released, and the discharge rate of the
Striola afferent fiber increases. Conversely, when the hair cell is
hyperpolarized, less neurotransmitter is released, and the
firing rate of the afferent fiber slows.
Semicircular Canals
Angular accelerations of the head produce minute move-
ment of the endolymph in relation to the head (Fig. 8.27).
This happens because the inertia of the endolymph causes
it to resist the initial acceleration of the membranous
labyrinth. This lag pushes on the cupula, causes the cilia
to bend, and consequently changes the discharge rates
of the vestibular afferent fibers. All the cilia in a given
ampullary crest are oriented in the same direction. In the
Fig. 8.25 Structure of one of the otolith organs, the saccule. Note
the orderly variation in kinocilium orientation, as well as their mirror horizontal canal, the cilia are oriented toward the utricle,
symmetry with regard to the striola. (Redrawn from Lindeman HH. Adv and in the other ampullae, they are oriented away from
Otorhinolaryngol 1973;20:405.) the utricle.
Hyperpolarization
Receptor potential Depolarization
Nerve impulses Resting Increased Decreased

discharge impulse impulse
frequency frequency
Fig. 8.26 Functional Polarization of Vestibular Hair Cells. When the stereocilia are bent toward the
kinocilium, the hair cell is depolarized, and the afferent fiber is excited. When the stereocilia are bent away
from the kinocilium, the hair cell is hyperpolarized, and the afferent discharge slows or stops. (Redrawn
from Kandel ER, Schwartz JH. Principles of Neural Science. New York: Elsevier; 1981.)
Turning motion of head Superior
Ampulla
Axis of Axis of Posterior Anterior
hair cells Utricle hair cells
Striola
Inferior
Fluid motion Fluid motion

in duct in duct
Lateral Striola
Anterior
Horizontal
ducts
Posterior
Medial
Fig. 8.28 Functional Polarization of Hair Cells in the Otolith

Increase Decrease Organs. A, The saccule. B, The utricle. The striola in each case
in firing in firing
is indicated by the dotted line. (Redrawn from Spoendlin HH. In:
Fig. 8.27 Effect of Leftward Head Movement on the Activity of Wolfson RJ [ed]. The Vestibular System and Its Diseases. Philadelphia:
Vestibular Afferent Fibers Supplying Hair Cells in the Horizontal University of Pennsylvania Press; 1966.)
Semicircular Canals. The perspective in the figure is from above
the head looking down. In this case, head rotation (acceleration)
to the left causes endolymph pressure to the right and the result is
increased output from the left canal and decreased output from the IN THE CLINIC
right canal. Small arrows in the ampulla indicate the functional polarity
of the hair cells. The large curved arrow at the top indicates movement Irritation of the vestibular labyrinth, as in Menieres disease,
of the head; smaller curved arrows indicate relative movement of the can result in rhythmic conjugate deviations of the eyes,
endolymph. followed by quick return saccades. This condition is known
as nystagmus (see Chapter 9). These eye movements are
The way in which angular acceleration of the head affects accompanied by a sense of vertigo and often nausea.
the discharge of vestibular afferent fibers is exemplified by The brain interprets a difference in the input from the two
sides of the vestibular system as head motion. Irritation (or
the activity that originates from the horizontal canals. Fig.
destruction) of one labyrinth produces an asymmetry of
8.27 shows the horizontal canals and utricle, as seen from input that results in abnormal eye movement and associated
above as the head is rotated (accelerated) to the left. As psychological effects.
acceleration to the left begins, the inertia of the endolymph
in the horizontal canals increases pressure toward the right
side. This causes the cilia to bend on hair cells of the ampulla
of the left horizontal canal toward the utricle and bends the direction. Instead, they are oriented in relation to a ridge,
cilia of the right canal away from the utricle. These actions called the striola, along the otolith organ (see Fig. 8.25).
increase the firing rate in the afferent fibers on the left and In the utricle, the hair cells on either side of the striola are
decrease the firing rate of the afferent fibers on the right. polarized toward the striola, whereas in the saccule they are
Once the head is moving at a constant velocity of rotation polarized away from the striola. Because the striola in each
(i.e., no acceleration), there would be no force on either otolith organ is curved, there are hair cells with all orienta-
cupula, and therefore the hair cells of both canals would tions in the plane of the organ (Fig. 8.28). In any particular
be firing as they do at rest. However, when the indicated orientation of the head, the cilia of the hair cells are bent to
rotation is stopped, the inertia of the endolymph creates varying extents according to their orientation in relation to
a force on both cupulas, but in the direction opposite to the gravitational vector. This results in a particular pattern
that caused by the original acceleration. This results in an of input from the otolith organs to the CNS. When the
increase in the discharge rate of afferent fibers on the right head is tilted to a new position, the orientation of the
side and a decrease in the discharge rate on the left. This otolithic membranes in relation to the gravitational vector
postrotatory effect is of functional and clinical significance. changes, and so the cilia of the hair cells are bent in a new
way. This change in the bending of the cilia of the hair
Otolith Organs cells changes the pattern of input from the otolith organs
Unlike the hair cells in the ampullary crests, not all the to the CNS and creates the sensation of movement, as well
hair cells in the otolith organs are oriented in the same as possibly triggering various reflexes. Similarly, a linear
acceleration caused by other forces, such as might occur in

a fall or the angular acceleration when a car turns around Taste Receptors
a curve (angular accelerations have linear centripetal and The sensation of taste depends on the activation of che-
instantaneous tangential components), also affects output moreceptors located in taste buds. A taste bud consists of
from the otolith organs. a group of 50 to 150 receptor cells, as well as supporting
cells and basal cells (Fig. 8.29A). The chemoreceptor cells
Central Vestibular Pathways synapse at their bases with primary afferent nerve fibers,
The vestibular afferent fibers project to the brainstem and their apices have microvilli that extend toward a taste
through the vestibular nerve. As previously mentioned, the pore. Chemoreceptor cells live only about 10 days. They
cell bodies of these afferent fibers are located in the Scarpa are continuously replaced by new chemoreceptor cells that
ganglion. These primary afferent fibers terminate in the differentiate from basal cells located near the base of the
vestibular nuclei (see Fig. 4.6D to E), which are located in taste bud.
the rostral medulla and caudal pons and in specific regions Chemoreceptor molecules, each specialized for one type
of the cerebellum, most prominently in the nodulus. of taste stimulus, sit on the microvilli of chemoreceptor cells
The vestibular nuclei give rise to various projections, and detect molecules that diffuse into the taste pore from
including projections through the medial longitudinal the overlying mucus of the tongue, part of which originates
fasciculus (see Fig. 4.6C to E) to the oculomotor nuclei. from glands adjacent to the taste buds. Some stimuli can
Therefore, it is not surprising that the vestibular nuclei pass directly into the cell to depolarize it (Na+ for salty and
exert powerful control over eye movements (the vestibule- H+ for sour) or open cation channels to generate a receptor
ocular reflex). Other projections give rise to the lateral potential (also salty and sour), whereas others (sucrose,
and medial vestibulospinal tracts, which, respectively, quinine, and glutamate for sweet, bitter, and umami)
provide for the activation of trunk and neck muscles and activate a second messenger that can either open cation
thereby contribute to equilibrium and to head movements channels or directly activate intracellular Ca++ stores (see
(vestibulocolic reflex). There are vestibular pathways to Fig. 8.29B). In each case, depolarization of the receptor
the cerebellum, the reticular formation, and the contra- results in the release of an excitatory neurotransmitter and,
lateral vestibular complex, as well as to the thalamus. The consequently, action potentials in the primary afferent nerve
pathway to the thalamus, via a projection to the cerebral fiber that are transmitted to the CNS.
cortex, mediates conscious sensation of vestibular activity. Coding of taste, however, is not based entirely on the
Vestibular reflexes and clinical tests of vestibular function selectivity of the chemoreceptors for the different primary
are described in Chapter 9. qualities because each cell responds to a range of stimuli,
although most intensely to one. Because most natural tastes
The Chemical Senses have chemicals that effect responses from a number of che-
moreceptors, recognition of taste quality appears to depend
The senses of gustation (taste) and olfaction (smell) help on the patterned input from a population of chemorecep-
detect chemical stimuli that are present either in food tors, each responding differentially to the components of
and drink or in the air. In the evolution of humans, these the stimulus. The intensity of the stimulus is reflected in
chemical senses apparently did not have the survival value the total amount of activity evoked.
of some of the other senses, but they contribute consider-
ably to quality of life and food selection, and they are Distribution and Innervation of Taste Buds
important stimulants of digestion. In other animals, the Taste buds are located on different types of taste papillae
chemical senses have greater survival value, and their activa- found on the tongue, palate, pharynx, and larynx. Types
tion evokes a number of social behaviors, including mating, of taste papillae include fungiform and foliate papillae
territoriality, and feeding. on the anterior and lateral aspects, respectively, of the
tongue and circumvallate papillae on the base of the
Taste tongue (see Fig. 8.29C). The circumvallate papillae may
contain several hundred taste buds. The tongue in humans
The stimuli that we commonly know as tastes are actually may have several thousand taste buds. The sensitivity of
mixtures of five elementary taste qualities: salty, sweet, sour, different regions of the tongue for different taste qualities
bitter, and umami.b Taste stimuli that are particularly effec- varies slightly because taste buds responding to each type of
tive in eliciting these sensations are, respectively, sodium taste are widely distributed. The taste buds are innervated
chloride, sucrose, hydrochloric acid, quinine, and mono- by three cranial nerves. The chorda tympani branch of
sodium glutamate. Umami has been described as having a the facial nerve (CN VII) innervates taste buds on the
proteinaceous, meaty character. anterior two thirds of the tongue, and the glossopharyn-
geal nerve (CN IX) innervates taste buds on the posterior
third of the tongue (see Fig. 8.29C). The vagus nerve (CN
b
The existence of a sixth, taste, fat (free fatty acids), is currently being X) innervates a few taste buds in the larynx and upper
debated. esophagus.
Central Taste Pathways in the face area of the S1 cortex and the other in the
The cell bodies of taste fibers in cranial nerves VII, IX, insula. An unusual feature of the central gustatory pathway
and X are located in the geniculate, petrosal, and nodose is that it is predominantly an uncrossed pathway (unlike the
ganglia, respectively. The central processes of the afferent central somatosensory pathways, which are predominantly
fibers enter the medulla, join the solitary tract, and synapse crossed).
in the nucleus of the solitary tract (see Fig. 4.6D to E). In
some animals, including several rodent species, the second- Olfaction
order taste neurons of the solitary nucleus project rostrally
to the ipsilateral parabrachial nucleus. The parabrachial The sense of smell is much better developed in some animals
nucleus then projects to the small-celled (parvocellular) part (macrosmatic animals) than in humans. The ability of
of the ventroposterior medial (VPMpc) nucleus of the dogs to track other animals on the basis of odor is legendary,
thalamus. In monkeys, the solitary nucleus projects directly as is the use of pheromones by insects to attract mates.
to the VPMpc nucleus. The VPMpc nucleus is connected However, olfaction contributes to humans emotional life,
to two different gustatory areas of the cerebral cortex: one and odors can effectively conjure up memories. It also helps
Ligand-gated 2nd Messenger
Taste stimuli Taste stimuli
Receptor Transmembrane
ion channel receptor
Direct
Conductance Na+
change
2nd
messenger
2nd messenger
Taste pore
+
Depolarization ++
+
Generator
Na+
potential
Calcium Release of
stores Ca++ stores
R K+
S R R S
Ca++ Ca++ Ca++ Ca++
influx influx
Transmitter Transmitter
release release
Afferent nerve
B
A B
Fig. 8.29 A, A taste bud is shown with the taste pore at the top and its innervation below. B, basal
cell; R, ciliated taste receptor cells; S, supporting cells. B, Taste receptor cell showing second messenger,
ligand-gated, and direct depolarization resulting in depolarization of the cell.
nonmotile cilia on the apical surface of these cells contain

Glossopharyngeal Chorda tympani Taste buds
nerve nerve
chemoreceptors that detect odorant chemicals dissolved in
the overlying mucus layer. From its opposite side, the cell
gives off an unmyelinated axon that joins other olfactory
nerve filaments and penetrates the base of the skull through
openings in the cribriform plate of the ethmoid bone.
These olfactory nerves synapse in the olfactory bulb, a
Circumvallate portion of the cerebral hemisphere of the brain located at
the base of the cranial cavity, just below the frontal lobe
Taste buds
(Fig. 8.31).
Humans have about 10 million olfactory chemorecep-
tors. Like taste cells, olfactory chemoreceptors have a short
life span (about 60 days), and they are also continuously
replaced. However, olfactory receptor cells are true neurons
Serous gland
and, as such, are the only neurons that are continuously
regenerated throughout life.
Foliate
The olfactory mucosa is exposed to odorant molecules
by ventilatory air currents or from the oral cavity during
Taste buds feeding. Sniffing increases the influx of odorants. The
odorants are temporarily bound in mucus to an olfactory
binding protein that is secreted by a gland in the nasal
cavity.
Olfactory coding resembles taste coding in that most
natural odors are complex and consist of many molecules
Fungiform that excite a wide variety of olfactory chemoreceptors.
C Coding for a particular perceived odor depends on the
Fig. 8.29, contd C, Distribution of the tastebuds on the tongue responses of many olfactory chemoreceptors, and the
and their innervation. (Redrawn from Squire LR, etal [eds]. Fundamen- strength of the odorant is represented by the overall amount
tal Neuroscience. San Diego, CA: Academic Press; 2002.) of afferent neural activity.
Central Pathways
people avoid consuming spoiled food and detect dangerous The initial synapse of the olfactory pathway is located in
situations. For example, an unpleasant odorant is added the olfactory bulb, which is a specialized portion of the
to odorless, colorless natural gas so that people can easily cerebral cortex and located on the underside of the frontal
detect a leak. lobe. It contains mitral cells, interneurons (granule cells;
Odor has more primary qualities than taste does.c As periglomerular cells), and distinct synaptic clusters (glom-
many as 1000 different odor receptors are coded in the eruli; see Fig. 8.31) in which theses interact with olfactory
human genome, and although only approximately 350 afferent fibers. As the olfactory afferent fibers reach the
types are functional, they represent the largest population of olfactory bulb from the olfactory mucosa, they branch as
G proteincoupled receptors in the genome. The olfactory they approach an olfactory glomerulus to synapse on the
mucosa also contains somatosensory receptors of the tri- dendrites of mitral cells. Each glomerulus is the target of
geminal nerve. When performing clinical tests of olfaction, thousands of olfactory afferent fibers, but all the afferent
clinicians must avoid activating these somatosensory recep- fibers to a single glomerulus convey input from the one
tors with thermal or noxious stimuli, such as the ammonia type of olfactory receptor. This is all the more remark-
used in smelling salts. able because olfactory receptor cells are being regenerated
continuously and new axons must therefore navigate their
Olfactory Receptors way to a correct glomerulus.
The olfactory chemoreceptor cells are located in the olfac- The granule and periglomerular cells are inhibitory inter-
tory mucosa, a specialized part of the nasopharynx. Olfac- neurons. They form dendrodendritic reciprocal synapses
tory chemoreceptors are bipolar nerve cells (Fig. 8.30). The with the dendrites of mitral cells. Activity in a mitral cell
depolarizes these inhibitory cells, and they in turn inhibit the
original and adjacent glomeruli. Because each glomerulus is
c
specialized by being the target of afferent fibers for a unique
The conscious perception of flavor, particularly of foods, is the result of combination of odor qualities, this appears to be a way of
both olfactory and gustatory input based on directly inhaled odor, taste
from the food as it is macerated in the mouth, and retronasal odor from enhancing stimulus contrast, much the way horizontal cells
the volatile molecules that are released by maceration and pass up into do in the retina. In addition, it provides a mechanism for
the nasal cavity from the pharynx. adaptation to continuous stimulation.
Axons
Basement
membrane
Olfactory
mucosa
Supporting
cell
Receptor cells
Fig. 8.30 Olfactory chemoreceptors and supporting cells. (Redrawn from de Lorenzo AJD. In: Zotter-
man Y [ed]. Olfaction and Taste. Elmsford, NY: Pergamon; 1963.)
Granule cell
Periglomerular cell
Mitral cell
Olfactory glomerulus
Cribriform plate
Olfactory nerve fibers

Basement
membrane
Olfactory Olfactory
receptor mucosa
Supporting cell
Fig. 8.31 Drawing of a Sagittal Section Through an Olfactory Bulb, Showing Terminations of the
Olfactory Chemoreceptor Cells in the Olfactory Glomeruli and the Intrinsic Neurons of the Olfactory
Bulb. The axons of the mitral cells are shown exiting in the olfactory tract to the right. (Modified from
House EL, Pansky B. A Functional Approach to Neuroanatomy. 2nd ed. New York: McGraw-Hill; 1967.)
The axons of mitral cells leave the olfactory bulb and

enter the olfactory tracts. From here, the olfactory connec- IN THE CLINIC
tions become highly complex. Within the olfactory tracts Olfaction is not generally examined in a routine neurological
is a nucleus, called the anterior olfactory nucleus, that examination. However, smell can be tested by having the
receives input from the olfactory bulb and projects to the patient inhale and identify an odorant. One nostril should be
contralateral olfactory bulb through the anterior commis- examined at a time while the other nostril is occluded. Strong
sure. As each olfactory tract approaches the base of the odorants, such as ammonia, should be avoided because
they also activate trigeminal nerve fibers. Smell sensation can
brain, it splits into the lateral and medial olfactory striae. be lost (anosmia) after a basal skull fracture or after damage
Axons of the lateral olfactory stria synapse in the primary to one or both olfactory bulbs or tracts by a tumor (such as
olfactory cortex, which includes the prepiriform cortex an olfactory groove meningioma). Concussions can cause
(and, in many animals, the piriform lobe). The medial anosmia because the sudden movement of the brain inside
olfactory stria includes projections to the amygdala, as the skull can shear the small unmyelinated olfactory nerve
fibers. An aura of a disagreeable odor, often the smell of
well as to the basal forebrain. These structures are por- burning rubber, occurs during uncinate seizures, which are
tions of, or directly connected to, the limbic system (see epileptic seizures that originate in the medial temporal lobe.
Chapter 10 ).
Of note is that the olfactory pathway is the only sensory
system that does not have an obligatory synaptic relay in
the thalamus before signals reach the cortex. However,
olfactory information does reach the mediodorsal nucleus narrow range of gustatory receptors with the wide reper-
of the thalamus, and it is then transmitted to the prefrontal toire of olfactory receptors. In addition, via its intimate
and orbitofrontal cortex. The functional roles of olfaction, connections with limbic and, by extension, hypothalamic
in addition to the conscious perception of odor, include structures, it provides input to subconscious mechanisms
providing much of the subtleties of taste by enhancing the related to emotions, memory, and sexual behavior.
Key Points
1. Light enters the eye through the cornea and lens but ganglion cells respond by generating action
and is focused on the retina, which lines the back of potentials.
the eye. The cornea is the most powerful refractive 5. The axons of ganglion cells in the temporal retina
surface, but the lens has a variable power that allows project to the brain ipsilaterally; those in the nasal
images of near objects to be focused on the retina. retina cross in the optic chiasm. Because the lens
The iris regulates depth of field and the amount of inverts the image that falls on the retina, each side of
illumination that enters the eye. the visual field is projected to the contralateral side
2. The outer segments of the photoreceptor cells of the brain for both eyes. In the lateral geniculate
transduce light. Photoreceptors synapse on retinal nucleus (LGN) of the thalamus, the input from each
bipolar cells, which in turn synapse on other eye terminates in separate layers, and the M ganglion
interneurons and on ganglion cells. The ganglion cells cells (sensitive to movement) and P ganglion cells
project to the brain through the optic nerve. The (sensitive to detail and color) project to separate
optic disc, where the optic nerve leaves the retina, layers as well.
contains no photoreceptors and is therefore a blind 6. The LGN projects to primary visual (striate) cortex
spot. The portion of the retina with the highest via the visual radiation and terminates largely in
degree of spatial resolution is the fovea and the layer 4, where there is an orderly retinotopic map.
surrounding macula. Within the map, information from each eye maps to
3. Rod photoreceptors have high sensitivity, do not alternating adjacent points to create ocular dominance
discriminate among colors, and function best under columns that extend vertically in the cortex. Striate
low light levels. Cone photoreceptors have lower cortical neurons outside of layer 4 respond best to bar
sensitivity but higher spatial resolution. Color vision or edge stimuli oriented in a particular way. Cells that
relies on the three types of cones that have different prefer a particular stimulus orientation are grouped
spectral sensitivities. in orientation columns.
4. Bipolar cells and many ganglion cells have concentric 7. The extrastriate visual areas have different functions.
receptive fields with an on-center/off-surround or Some in the inferotemporal cortex are influenced
off-center/on-surround organization. Horizontal chiefly by P cells, and they function in form
cells mediate this center-surround antagonism. detection, color vision, and face discrimination. M
Photoreceptor, bipolar, and horizontal cells respond cells influence regions of the middle temporal and
to stimulation by modulating their membrane parietal cortex, which function in motion detection
potential and their release of neurotransmitters, and the control of eye movements.
8. A pure tone is characterized in terms of its cross-section of the endolymph-filled canal. Angular
amplitude, frequency, and phase. Natural sounds head acceleration displaces the endolymph and the
are combinations of pure tones. Sound pressure is cupula, bending the cilia. If the stereocilia bend
measured in decibels (dB), in relation to a reference toward the kinocilium, the hair cell is depolarized,
level. which causes an increase in the firing rate in the
9. The pinna and auditory canal convey airborne afferent fiber.
sound waves to the tympanic membrane. The three 16. In the otolith organs, the cilia project into an
small bones (ossicles) of the middle ear transmit the otolithic membrane. Acceleration of the head, as
vibrations of the tympanic membrane to the oval with linear movement, or change in position in
window of the fluid-filled inner ear. Hearing is most relation to gravity displaces the otolithic membrane
sensitive at about 3000Hz because of the dimensions (because of the mass of the otoliths) and changes the
of the auditory canal and the mechanics of the firing patterns of the hair cells, depending on their
ossicles. orientation.
10. The cochlea of the inner ear has three main 17. Central vestibular pathways include afferent
compartments: the scala vestibuli, the scala tympani, connections to the vestibular nuclei and the
and the intervening scala media (cochlear duct). The cerebellum. Activation of the vestibular afferent
cochlear duct is bounded on one side by the basilar fibers is detected by the brain as head acceleration
membrane, on which lies the organ of Corti, the or position change and is relayed via the vestibular
sound transduction mechanism. nuclei to pathways that mediate compensatory eye
11. When the basilar membrane oscillates in response to movements, neck movements, and adjustments to
pressure waves introduced into the scala vestibuli at posture.
the oval window, the stereocilia of the hair cells of the 18. Taste buds contain chemoreceptor cells arranged
organ of Corti are subjected to shear forces, which around a taste pore. Taste buds are located on
open mechanoreceptor K+ channels. This results in several kinds of papillae on the tongue and in the
a membrane conductance change that modulates pharynx and larynx. Five types of taste-receptor cells
the release of neurotransmitters on to cochlear nerve detect the five elementary qualities of taste: salty,
fibers. sweet, sour, bitter, and umami. Complex flavors are
12. High-frequency sounds best activate the hair cells signaled by the patterned activity of multiple classes
near the base of the cochlea, and low-frequency of taste receptor and by central correlation with
sounds activate cells near the apex. Such a tonotopic accompanying olfactory input.
organization is also present in central auditory 19. Afferent taste fibers synapse in the nucleus of
structures, including the cochlear nuclei, superior the solitary tract. The thalamic relay is via a part
olivary complex, inferior colliculus, medial geniculate of the ventroposterior medial nucleus to the
nucleus, and primary auditory cortex. taste-receiving areas located in the S1 cortex and
13. Auditory processing at many sites in the central the insula.
auditory pathway contributes to sound localization, 20. Odors are detected by olfactory chemoreceptor
frequency and intensity analysis, and speech cells, which are continuously regenerated in the
recognition. olfactory mucosa. These cells are true neurons that
14. The vestibular apparatus is part of the inner ear. are endowed with a wide array of G proteincoupled
It includes three semicircular canals (horizontal, receptors that enable the detection of hundreds of
anterior, and posterior) and two otolith organs odor molecules.
(utricle and saccule) on each side. These transduce, 21. Individual olfactory axons project to olfactory
respectively, angular and linear accelerations of the glomeruli, specific for each stimulus type, in the
head. The three semicircular canals are mutually olfactory bulb. They synapse on the dendrites of
orthogonal, so they can resolve angular acceleration mitral cells, which have reciprocal synapse with
of the head about any axis of rotation. inhibitory interneurons. This synaptic organization
15. In each semicircular canal, there are sensory hair cells in the glomerulus underlies stimulus adaptation and
whose cilia extend into a cupula, which blocks the contrast enhancement.
Additional Reading
Squire L, Berg D. Fundamental Neuroscience. 4th ed. New York:
Academic Press; 2012.

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8

The Special Senses

Visual axis Optic axis

Iris Aqueous humor

occurs; that is, until the ciliary muscle contracts, causing

Retinal layers Components

7 Inner plexiform layer Ganglion cell

3 Outer limiting membrane

Receptor density (104/mm2)

epithelium. This process determines the rod-like shape of

CONE ABSORPTION SPECTRA

419 498 533 564

Na+ Active Hyperpolarization

the visual pigment occurs much more rapidly in cones than

A On-center field F Off-center field

C Central illumination H Central illumination

D Annular illumination I Annular illumination

0 0.5 1.0 1.5 0 0.5 1.0 1.5

Monocular Binocular Monocular

half of the visual target is imaged on the nasal retina of the

Left visual field

it depends on convergent input from the two eyes and the

MST (medial superior temporal) on the lateral aspect of the

Other Visual Pathways Audition

Tympanic Middle Round

Fig. 8.19 Different Frequencies of Sound Result in Different

Amplitudes of Displacement at Different Sites Along the Organ

Threshold intensity (dB)

Threshold intensity (dB)

Tuning curves can be considered as receptive field plots. A, Tuning

(see Fig. 8.21B). By plotting the distribution of the char-

the bony labyrinth. The vestibular apparatus on each side

Left superior Right superior

Nerve impulses Resting Increased Decreased

Turning motion of head Superior

Fluid motion Fluid motion

Fig. 8.28 Functional Polarization of Hair Cells in the Otolith

acceleration caused by other forces, such as might occur in

Ligand-gated 2nd Messenger

Taste stimuli Taste stimuli

nonmotile cilia on the apical surface of these cells contain

man Y [ed]. Olfaction and Taste. Elmsford, NY: Pergamon; 1963.)

Olfactory nerve fibers

The axons of mitral cells leave the olfactory bulb and

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