Sensory Physiology 305

Case Investigation
SUMMARY
Ed was on an international flight, so he was exposed to a
long flight at high altitude (even though the airplane cabin is
pressurized, it still is at a lower than sea-level pressure).
Considering his head cold, his eustachian tube may not
have been able to equalize pressure on both sides of the
tympanic membrane, leading to pain and reduced hearing.
If this is the explanation, the symptoms should resolve with
time and the aid of a decongestant. Ed’s visual problem
suggests that he is experiencing presbyopia, which nor-
mally begins at about Ed’s age. LASIK could not compen-
sate for the inability of the lens to perform accommodation.
If Ed were nearsighted, he might try monovision LASIK
surgery, but because he does not have myopia, he would
still need reading glasses if he had this surgery.

SUMMARY
10.1 Characteristics of Sensory Receptors 264 C. Generator potentials are graded changes (usually depolar-
A. Sensory receptors may be categorized on the basis of izations) in the membrane potential of the dendritic
their structure, the stimulus energy they transduce, endings of sensory neurons.
or the nature of their response. 1. The magnitude of the potential change of the
1. Receptors may be dendritic nerve endings, specialized generator potential is directly proportional to the
neurons, or specialized epithelial cells associated with strength of the stimulus applied to the receptor.
sensory nerve endings. 2. After the generator potential reaches a threshold value,
2. Receptors may be chemoreceptors, photoreceptors, increases in the magnitude of the depolarization result
thermoreceptors, mechanoreceptors, or nociceptors. in increased frequency of action potential production
a. Proprioceptors include receptors in the muscles, in the sensory neuron.
tendons, and joints.
10.2 Cutaneous Sensations 267
b. The senses of sight, hearing, taste, olfaction, and
equilibrium are grouped as special senses. A. Somatesthetic information—from cutaneous receptors
3. Receptors vary in the duration of their firing in and proprioceptors—is carried by third-order neurons to
response to a constant stimulus. the postcentral gyrus of the cerebrum.
a. Tonic receptors continue to fire as long as the 1. Proprioception and pressure sensations ascend on the
stimulus is maintained; they monitor the presence ipsilateral side of the spinal cord, synapse in the
and intensity of a stimulus. medulla and cross to the contralateral side, and then
ascend in the medial lemniscus to the thalamus;
b. Phasic receptors respond to stimulus
neurons in the thalamus, in turn, project to the
changes; they do not respond to a sustained
postcentral gyrus.
stimulus, and this partly accounts for sensory
adaptation. 2. Sensory neurons from other cutaneous receptors
synapse and cross to the contralateral side in the
B. According to the law of specific nerve energies, each spinal cord and ascend in the lateral and ventral
sensory receptor responds with lowest threshold to only spinothalamic tracts to the thalamus; neurons in the
one modality of sensation. thalamus then project to the postcentral gyrus.
1. That stimulus modality is called the adequate
B. The receptive field of a cutaneous sensory neuron is the
stimulus.
area of skin that, when stimulated, produces responses in
2. Stimulation of the sensory nerve from a receptor by the neuron.
any means is interpreted in the brain as the adequate
1. The receptive fields are smaller where the skin has a
stimulus modality of that receptor.
greater density of cutaneous receptors.
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2. The two-point touch threshold test reveals that the bending the stereocilia and electrically stimulating
fingertips and tip of the tongue have a greater density the sensory endings.
of touch receptors, and thus a greater sensory acuity, C. The three semicircular canals are oriented at nearly right
than other areas of the body. angles to each other, like the faces of a cube.
C. Lateral inhibition acts to sharpen a sensation by 1. The hair cells are embedded within a gelatinous
inhibiting the activity of sensory neurons coming membrane called the cupula, which projects into the
from areas of the skin around the area that is most endolymph.
greatly stimulated. 2. Movement along one of the planes of a semicircular
canal causes the endolymph to bend the cupula and
10.3 Taste and Smell 271 stimulate the hair cells.
A. The sense of taste is mediated by taste buds. 3. Stimulation of the hair cells in the vestibular apparatus
1. There are four well-established modalities of taste activates the sensory neurons of the vestibulocochlear
(salty, sour, sweet, and bitter); a fifth, called umami, nerve (VIII), which projects to the cerebellum and to
which is stimulated by glutamate, is now also the vestibular nuclei of the medulla oblongata.
recognized. a. The vestibular nuclei in turn send fibers to the
2. Salty and sour taste are produced by the movement of oculomotor center, which controls eye
sodium and hydrogen ions, respectively, through movements.
membrane channels; sweet and bitter tastes are b. Spinning and then stopping abruptly can thus cause
produced by binding of molecules to protein receptors oscillatory movements of the eyes (nystagmus).
that are coupled to G-proteins.
B. The olfactory receptors are neurons that synapse within 10.5 The Ears and Hearing 279
the olfactory bulb of the brain.
A. The outer ear funnels sound waves of a given frequency
1. Odorant molecules bind to membrane protein
(measured in hertz) and intensity (measured in decibels)
receptors. There may be as many as 1,000 different
to the tympanic membrane, causing it to vibrate.
receptor proteins responsible for the ability to detect
as many as 10,000 different odors. B. Vibrations of the tympanic membrane cause movement of
2. Binding of an odorant molecule to its receptor causes the the middle-ear ossicles—malleus, incus, and stapes—
dissociation of large numbers of G-protein subunits. which in turn produces vibrations of the oval window of
The effect is thereby amplified, which may contribute the cochlea.
to the extreme sensitivity of the sense of smell. C. Vibrations of the oval window set up a traveling wave of
perilymph in the scala vestibuli.
10.4 Vestibular Apparatus and Equilibrium 275 1. This wave can pass around the helicotrema to the
scala tympani, or it can reach the scala tympani by
A. The structures for equilibrium and hearing are located in
passing through the scala media (cochlear duct).
the inner ear, within the membranous labyrinth.
2. The scala media is filled with endolymph.
1. The structure involved in equilibrium, known as the
vestibular apparatus, consists of the otolith organs a. The membrane of the cochlear duct that faces
(utricle and saccule) and the semicircular canals. the scala vestibuli is called the vestibular
membrane.
2. The utricle and saccule provide information about
linear acceleration, whereas the semicircular canals b. The membrane that faces the scala tympani is
provide information about angular acceleration. called the basilar membrane.
3. The sensory receptors for equilibrium are hair cells D. The sensory structure of the cochlea is called the spiral
that support numerous stereocilia and one organ or organ of Corti.
kinocilium. 1. The organ of Corti rests on the basilar membrane and
a. When the stereocilia are bent in the direction of the contains sensory hair cells.
kinocilium, the cell membrane becomes a. The stereocilia of the hair cells project upward into
depolarized. an overhanging tectorial membrane.
b. When the stereocilia are bent in the opposite b. The hair cells are innervated by the
direction, the membrane becomes vestibulocochlear nerve (VIII).
hyperpolarized. 2. Sounds of high frequency cause maximum
B. The stereocilia of the hair cells in the utricle and saccule displacement of the basilar membrane closer to
project into the endolymph of the membranous labyrinth its base, near the stapes; sounds of lower
and are embedded in a gelatinous otolithic membrane. frequency produce maximum displacement of the
1. When a person is upright, the stereocilia of the utricle basilar membrane closer to its apex, near the
are oriented vertically; those of the saccule are helicotrema.
oriented horizontally. a. Displacement of the basilar membrane causes the
2. Linear acceleration produces a shearing force hairs to bend against the tectorial membrane and
between the hairs of the otolithic membrane, thus stimulate the production of nerve impulses.
 Sensory Physiology 307

b. Pitch discrimination is thus dependent on the 2. In the dark, more rhodopsin can be produced, and
region of the basilar membrane that vibrates increased rhodopsin in the rods makes the eyes more
maximally to sounds of different frequencies. sensitive to light. The increased concentration of
c. Pitch discrimination is enhanced by lateral rhodopsin in the rods is partly responsible for dark
inhibition. adaptation.
3. The rods provide black-and-white vision under
10.6 The Eyes and Vision 286 conditions of low light intensity. At higher light
A. Light enters the cornea of the eye, passes through the intensity, the rods are bleached out and the cones
pupil (the opening of the iris) and then through the lens, provide color vision.
from which point it is projected to the retina in the back 4. The pigment epithelium has many functions that are
of the eye. required by the photoreceptors in the retina.
1. Light rays are bent, or refracted, by the cornea a. The pigment epithelium phagocytoses the shed
and lens. outer segments of the rods and cones, absorbs stray
2. Because of refraction, the image on the retina is light, and has many other important functions.
upside down and right to left. b. The photoreceptors cannot convert all-trans retinal
3. The right half of the visual field is projected to the left back into 11-cis retinal; this is done by the pigment
half of the retina in each eye, and vice versa. epithelium.
B. Accommodation is the ability to maintain a focus on the c. The 11-cis retinal is then moved back into the
retina as the distance between the object and the eyes is photoreceptors, where it can associate with opsin to
changed. regenerate the photopigment; this is known as the
1. Accommodation is achieved by changes in the shape visual cycle of retinal.
and refractive power of the lens. B. In the dark, a constant movement of Na+ into the rods
2. When the muscles of the ciliary body are relaxed, the produces what is known as a “dark current.”
suspensory ligament is tight, and the lens is pulled to 1. When light causes the dissociation of rhodopsin, the
its least convex form. Na+ channels become blocked and the rods become
a. This gives the lens a low refractive power for hyperpolarized in comparison to their membrane
distance vision. potential in the dark.
b. As an object is brought closer than 20 feet from 2. When the rods are hyperpolarized, they release less
the eyes, the ciliary body contracts, the neurotransmitter at their synapses with bipolar cells.
suspensory ligament becomes less tight, and 3. Neurotransmitters from rods cause depolarization of
the lens becomes more convex and more powerful. bipolar cells in some cases and hyperpolarization of
C. Visual acuity refers to the sharpness of the image. It bipolar cells in other cases; thus, when the rods are in
depends in part on the ability of the lens to bring the light and release less neurotransmitter, these effects
image to a focus on the retina. are inverted.
1. People with myopia have an eyeball that is too long, C. According to the trichromatic theory of color vision, there
so that the image is brought to a focus in front of the are three systems of cones, each of which responds to one
retina; this is corrected by a concave lens. of three colors: red, blue, or green.
2. People with hyperopia have an eyeball that is too 1. Each type of cone contains retinene attached to a
short, so that the image is brought to a focus behind different type of protein.
the retina; this is corrected by a convex lens. 2. The names for the cones signify the region of the
3. Astigmatism is the condition in which asymmetry spectrum in which the cones absorb light maximally.
of the cornea and/or lens causes uneven D. The fovea centralis contains only cones; more peripheral
refraction of light around 360 degrees of a circle, parts of the retina contain both cones and rods.
resulting in an image that is not sharply focused on 1. Each cone in the fovea synapses with one bipolar cell,
the retina. which in turn synapses with one ganglion cell.
a. The ganglion cell that receives input from the fovea
10.7 Retina 293
thus has a visual field limited to that part of the
A. The retina contains rods and cones—photoreceptor retina that activated its cone.
neurons that synapse with bipolar cells. b. As a result of this 1:1 ratio of cones to bipolar cells,
1. When light strikes the rods, it causes the visual acuity is high in the fovea but sensitivity to
photodissociation of rhodopsin into retinene and low light levels is lower than in other regions of the
opsin. retina.
a. This bleaching reaction occurs maximally with a 2. In regions of the retina where rods predominate, large
light wavelength of 500 nm. numbers of rods provide input to each ganglion cell
b. Photodissociation is caused by the conversion of (there is a great convergence). As a result, visual
the 11-cis form of retinene to the all-trans form acuity is impaired, but sensitivity to low light levels is
that cannot bind to opsin. improved.
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E. The right half of the visual field is projected to the left 1. The receptive field of a ganglion cell is roughly
half of the retina of each eye. circular, with an “on” or “off” center and an
1. The left half of the left retina sends fibers to the left antagonistic surround.
lateral geniculate body of the thalamus. a. A spot of light in the center of an “on” receptive
2. The left half of the right retina also sends fibers to the field stimulates the ganglion cell, whereas a spot of
left lateral geniculate body. This is because these light in its surround inhibits the ganglion cell.
fibers decussate in the optic chiasma. b. The opposite is true for ganglion cells with “off”
3. The left lateral geniculate body thus receives input receptive cells.
from the left half of the retina of both eyes, c. Wide illumination that stimulates both the center and
corresponding to the right half of the visual field; the the surround of a receptive field affects a ganglion
right lateral geniculate receives information about the cell to a lesser degree than a pinpoint of light that
left half of the visual field. illuminates only the center or the surround.
a. Neurons in the lateral geniculate bodies send fibers 2. The antagonistic center and surround of the receptive
to the striate cortex of the occipital lobes. field of ganglion cells provide lateral inhibition, which
b. The geniculostriate system is involved in enhances contours and provides better visual acuity.
providing meaning to the images that form on B. Each lateral geniculate body receives input from both
the retina. eyes relating to the same part of the visual field.
4. Instead of synapsing in the geniculate bodies, some 1. The neurons receiving input from each eye are
fibers from the ganglion cells of the retina synapse in arranged in layers within the lateral geniculate.
the superior colliculus of the midbrain, which controls 2. The receptive fields of neurons in the lateral geniculate
eye movement. are circular, with an antagonistic center and surround—
a. Because this brain region is also called the optic much like the receptive field of ganglion cells.
tectum, this pathway is called the tectal system. C. Cortical neurons involved in vision may be simple,
b. The tectal system enables the eyes to move and complex, or hypercomplex.
track an object; it is also responsible for the 1. Simple neurons receive input from neurons in the
pupillary reflex and the changes in lens shape that lateral geniculate; complex neurons receive input from
are needed for accommodation. simple cells; and hypercomplex neurons receive input
from complex cells.
10.8 Neural Processing of Visual Information 302 2. Simple neurons are best stimulated by a slit or bar of
A. The area of the retina that provides input to a ganglion light that is located in a precise part of the visual field
cell is called the receptive field of the ganglion cell. and that has a precise orientation.

REVIEW ACTIVITIES
Test Your Knowledge
Match the vestibular organ on the left with its correct
5. Tonic receptors
component on the right.
a. are fast-adapting.
1. Utricle and saccule a. Cupula
b. do not fire continuously to a sustained stimulus.
2. Semicircular canals b. Ciliary body
c. produce action potentials at a greater frequency as the
3. Cochlea c. Basilar
generator potential is increased.
membrane
d. are described by all of these.
d. Otolithic
6. Cutaneous receptive fields are smallest in
membrane
a. the fingertips. b. the back.
4. The dissociation of rhodopsin in the rods in response to
c. the thighs. d. the arms.
light causes
7. The process of lateral inhibition
a. the Na+ channels to become blocked.
a. increases the sensitivity of receptors.
b. the rods to secrete less neurotransmitter.
b. promotes sensory adaptation.
c. the bipolar cells to become either stimulated or
inhibited. c. increases sensory acuity.
d. all of these. d. prevents adjacent receptors from being stimulated.
 Sensory Physiology 309

8. The receptors for taste are 16. Which of these sensory modalities is transmitted directly
a. naked sensory nerve endings. to the cerebral cortex without being relayed through the
thalamus?
b. encapsulated sensory nerve endings.
a. Taste
c. specialized epithelial cells.
b. Sight
9. Which of these statements about the utricle and saccule
are true? c. Smell
a. They are otolith organs. d. Hearing
b. They are located in the middle ear. e. Touch
c. They provide a sense of linear acceleration. 17. Stimulation of membrane protein receptors by binding to
specific molecules is not responsible for
d. Both a and c are true.
a. the sense of smell.
e. Both b and c are true.
b. sweet taste sensations.
10. Because fibers of the optic nerve that originate in the nasal
c. sour taste sensations.
halves of each retina cross over at the optic chiasma, each
lateral geniculate receives input from d. bitter taste sensations.
a. both the right and left sides of the visual field of both 18. Epithelial cells release transmitter chemicals that excite
eyes. sensory neurons in all of these senses except
b. the ipsilateral visual field of both eyes. a. taste.
c. the contralateral visual field of both eyes. b. smell.
d. the ipsilateral field of one eye and the contralateral field c. equilibrium.
of the other eye. d. hearing.
11. When a person with normal vision views an object from a
distance of at least 20 feet,
Test Your Understanding
a. the ciliary muscles are relaxed.
19. Explain what is meant by lateral inhibition and give
b. the suspensory ligament is tight.
examples of its effects in three sensory systems.
c. the lens is in its most flat, least convex shape.
20. Describe the nature of the generator potential and explain
d. all of these apply. its relationship to stimulus intensity and to frequency of
12. Glasses with concave lenses help correct action potential production.
a. presbyopia. 21. Describe the phantom limb phenomenon and give a
b. myopia. possible explanation for its occurrence.
c. hyperopia. 22. Explain the relationship between smell and taste. How are
these senses similar? How do they differ?
d. astigmatism.
23. Explain how the vestibular apparatus provides information
13. Parasympathetic nerves that stimulate constriction of the about changes in the position of our body in space.
iris (in the pupillary reflex) are activated by neurons in
24. In a step-by-step manner, explain how vibrations of the
a. the lateral geniculate. oval window lead to the production of nerve impulses.
b. the superior colliculus. 25. Using the ideas of the place theory of pitch and the
c. the inferior colliculus. tonotopic organization of the auditory cortex, explain how
d. the striate cortex. we perceive different pitches of sounds.
14. A bar of light in a specific part of the retina, with a 26. Describe the sequence of changes that occur during
particular length and orientation, is the most effective accommodation. Why is it more of a strain on the eyes to
stimulus for look at a small nearby object than at large objects far away?
a. ganglion cells. 27. Describe the effects of light on the photoreceptors and
b. lateral geniculate cells. explain how these effects influence the bipolar cells.
c. simple cortical cells. 28. Explain why images that fall on the fovea centralis are
seen more clearly than images that fall on the periphery of
d. complex cortical cells. the retina. Why are the “corners of the eyes” more
15. The ability of the lens to increase its curvature and sensitive to light than the fovea?
maintain a focus at close distances is called 29. Explain why rods provide only black-and-white vision.
a. convergence. Include a discussion of different types of color blindness in
b. accommodation. your answer.
c. astigmatism. 30. Explain why green objects can be seen better at night than
objects of other colors. What effect does red light in a
d. amblyopia.
darkroom have on a dark-adapted eye?
310 Chapter 10

31. Describe the receptive fields of ganglion cells and explain and 19). Explain what functions could be disturbed by a
how the nature of these fields helps to improve visual damaged pigment epithelium, and relate this to the disease
acuity. of macular degeneration.
32. How many genes code for the sense of color vision? How
many for taste? How many for smell? What does this Test Your Quantitative Ability
information say about the level of integration required by Refer to figure 10.26 (p. 287) and use the figure reproduced
the brain for the perception of these senses? below to answer the following questions.
33. Discuss the different functions of the pigment epithelium
of the retina, and describe the visual cycle of retinal.
34. What makes the lens of the eye clear? What happens when S (blue) cone M (green) cone L (red) cone
100
cataracts form?

Relative sensitivity (%)

Test Your Analytical Ability
35. You are firing your laser cannon from your position on
the bridge of your starship. You see the hostile enemy
starship explode, but you hear no accompanying sound.
50
Can you explain this? How do receptors for sight and
hearing differ?
36. People with conduction deafness often speak quietly. By
contrast, people with sensorineural deafness tend to speak
louder than normal. Explain these differences.
37. Opioid drugs reduce the sensation of dull, persistent pain
0
but have little effect on the initial sharp pain of a noxious 400 500 600 700
stimulus (e.g., a pinprick). What do these different Wavelength (nanometers)
effects imply? What conclusion can be drawn from the
fact that aspirin (a drug that inhibits the formation of
43. At which wavelength of light are the blue and green cones
prostaglandins) functions as a pain reliever?
equally stimulated? What color is this light?
38. Compare the role of G-proteins in the senses of taste and
44. Which cones are most sensitive to light at 600 nm? Which
sight. What is the advantage of having G-proteins mediate
are the least sensitive?
the effect of a stimulus on a receptor cell?
45. Which wavelength of light stimulates two cone systems to
39. Discuss the role that inertia plays in the physiology of the
the greatest degree? What color is this light?
vestibular apparatus. Why is there no sensation of
movement in an airplane once it has achieved cruising 46. Which wavelength of light stimulates all three of the cone
speed? systems to the greatest degree? What color is this light?
40. Explain why menthol feels cold on the tongue and chili
sauce feels hot. What does this reveal about our
perceptions of reality?
41. In order to see a dim star at night, it is better not to look
directly at it. If you see it from the “corner of your eye”
and then turn to look at it, it may disappear. Explain how
this occurs. Also, you can stare directly at an object in Visit this book’s website at www.mhhe.com/Fox12
daylight and continue seeing it, despite the bleaching of for chapter quizzes, interactive learning exercises, and
photoreceptors. Explain how this is possible.
other study tools.
42. The pigment epithelial cells may undergo apoptosis in
response to the oxidative stress of free radicals (chapters 3