The Skull Protects the Eye - Evergreen State College...2015/08/04 · the eye, and the lacrimal...

366 Chapter 10 Sensory Physiology

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~ Vestibular branch of r vestibulocochlear

~ ne~e~lll)

~

Vestibu lar apparatus ~

• FIGURE 10-28 eNS pathways for equilibrium

The Skull Protects the Eye The external anatomy of the eye is shown in Figure 10-29 • .

Like sensory elements of the ears, the eyes are protected by a

bony cavity, the orbit, which is formed by facial bones of the

skull. Accessory structures associated with the eye include six

extrinsic eye muscles, skeletal muscles that attach to the outer

surface of the eyeball and control eye movements. Cranial

nerves III, IV, and VI innervate these muscles.

The upper and lower eyelids close over the anterior surface of

the eye, and the lacrimal apparatus, a system of glands and ducts,

keeps a continuous flow of tea rs washing across the exposed sur

face so that it remains moist and free of debris. Tear secretion is

stimulated by parasympathetic neurons from cranial nerve VII.

Muscles attached to Lacrimal gland external surface of eye secretes tears. control eye movement.

__- - Upper eyelid

~-- Sclera

"'--;~----4~-.-.!-- Pupil

----'-- Iris

------ Lower eyelid

Nasolacrimal duct drains tears into nasal cavity .

• FIGURE 10-29 External anatomy of the eye

The orbit is a bony cavity that protects the eye.

___------~ n-,..~--riJ Somatic

motor neurons controlling eye

movements

The eye itself is a hollow sphere divided into two compart

ments (chambers) separated by a lens (Fig. 1O-30a e ). The lens,

suspended by ligaments called zonuies, is a transparent disk

that focuses light. The anterior chamber in front of the lens is

filled with aqueous humor [hllmidus, moist], a low-protein

plasma-like fluid secreted by the ciliary epithelium supporting the

lens. Behind the lens is a much larger chamber, the vitreous

chamber, filled mostly with the vitreous body [vitnlll'l, glass;

also called the vitreous humor], a clear, gelatinous ma trix that

helps maintain the shape of the eyeball. The outer wall of the

eyeball, the sclera, is composed of connective tissue.

CLINICAL FOCUS

GLAUCOMA

The eye disease glaucoma, characterized by degeneration of the optic nerve, is the leading cause of blindness worldwide. Many people associate glaucoma with increased intraocular (within the eyeball) pressure, but scientists have discovered that increased pressure is only one risk factor for the disease. A significant number of people with glaucoma have normal intraocular pressure, and not everyone with elevated pressure develops glaucoma. Many cases of elevated eye pressure are associated with excess aqueous humor, a fluid that is secreted by the ciliary epithelium near the lens. Normally the fluid drains out through the canal of Schlemm in the anterior chamber of the eye, but if outflow is blocked, the aqueous humor accumulates, causing pressure to build up inside the eye. Treatments to decrease intraocular pressure include drugs that inhibit aqueous humor production and surgery to reopen the canal of Schlemm. Research suggests that the optic nerve degeneration in glaucoma may be due to nitric oxide or apoptosis-inducing factors, and studies in these areas are underway.

The Eye and Vision 367

THE EYE Zonules: Lens bends attach lens to light to focus ciliary muscle it on the retina .

Optic disk (blind spot): ---_, region where optic nerve and blood vessels leave

Canal of the eye Schlemm Aqueous Central retinal artery

humor and vein emerge from ~ center of optic disk. Cornea ~ Optic nerve Pupil changes

amount of light ~ Fovea: region entering the eye. of sharpest vision

Iris Macula: the center of the visual field

Vitreous chamber (b) View of the rear wall of the eye as seen

Retina: layer that through the pupil with an ophthalmoscope contains photoreceptors

Ciliary muscle: contraction alters "--- Sclera is connective tissue. curvature of the lens. FIGURE QUESTION

If the fovea is lateral to the optic disk, (a) Sagittal section of the eye which eye is illustrated in part (b) ?

• FIGURE 10-30

Light enters the anterior surface of the eye through the CONCEPT CHECK

cornea, a transparent disk of tissue that is a continuation of the

sclera. After passing throug h the opening of the pupil, it strikes 24. What functions does the aqueous humor serve' Answers p. 383

the lens, which has two convex surfaces. The cornea and lens

together bend incoming light rays so that they focus on the Light Enters the Eye Through the Pupil retina, the ligh t-sensitive lining of the eye that contains the In the first step of the visual pathway, light from the environphotoreceptors. ment en ters the eye. Before it strikes the reti na, however, the

When viewed through the pupil with an ophthalmoscope light is modified two ways. First, the amount of light that [ophthalmos, eye), the retina is seen to be crisscrossed with small reaches photoreceptors is modulated by cha nges in the size of arteries and veins that radiate out from one spot, the optic disk the pupil. Second, the light is focused by changes in the shape (Fig. lO-30b). The optic disk is the location where neurons of the of the lens. visual pathway form the optic nerve (crania l nerve II) and exit The human eye functions ove r a lOO,OOO-fold range of the eye. Lateral to the optic disk is a sma ll dark spot, the fovea. light intensity. Most of this ability comes from the sensitivity The fovea and a narrow ring of ti ssue surrounding it, the of the photoreceptors, but the pupils assist by regulating the macula, are the regions of the retina with the most acute vision. amount of light that falls on the re tina . In bright sun light, the

Neural pathways for the eyes are illustrated in Figure pupils narrow to about 1.S mm in diameter when a parasympa10-31 • . The optic nerves from the eyes go to the optic chiasm th etic pathway constricts the circular pupillary muscles. In the in the brain, where some of the fib ers cross to the opposite side. dark, the opening of the pupil dilates to 8 mm, a 28-fold inAfter synapsing in the lateral geniculate body (lateral genicucrease in pupil area. Dilation occurs when radi al muscles lying late nilclells) of the thalamus, the vision neurons of the tract ter perpendicular to the circular muscles contract under the influminate in the occipital lobe at the visual cortex. Collateral ence of sympathetic neurons. pathways go from the thalamus to the midbrain, where they Testing pupillary reflexes is a standard part of a neurologsynapse with effer~nt neurons of cranial nerve III that control ical exam ination . Light hitting the retina in one eye activates the the diameter of the pupils. reflex. Signals travel through the optic nerve to the thalamus,

10


(a) Dorsal view (b) Neural pathway for

"'''';;~----'''f-+-+-----1F.l+.- 0 pt i c tract

"--:.....:::I-';H+- Optic chiasm

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~~~

vision, lateral view

Optic Optic Optic Lateral geniculate Visual cortex nerve chiasm tract body (thalamus) (occipital lobe)

(e) Collateral pathways leave the thalamus and go to the midbrain .

,light ,

,,,

,,,

pupillary constriction .

• FIGURE 10-31 Neural pathways for vision and the pupillary reflex.

Collateral pathways synapsing in the midbrain control constr iction of the pupils.

then to the midbrain , where efferent neurons constrict the

pupils in both eyes (Fig. 1O-31c). This response is known as the

consensual reflex and is mediated by parasympathetic fibers

running through cranial nerve III.

CONCEPT CHECK

25. Use the neural pathways in Fig ure 10-31c to answer the following questions.

(a) Why does shining light into one eye cause pupillary constriction in both eyes?

(b) If you shine a light in the left eye and get pupillary constriction in the right eye but not in the left eye, what can you conclude about the afferent path from the left eye to the brain? About the efferent pathways to the pupils?

26. Parasympathetic fibers constrict the pupils, and sympathetic fibers dilate them. This is an example of what kind of control?

Answers p. 383

In addition to regulating the amount of light that hits the

retina, the pupils create what is known as depth of field. A sim

pie example comes from photography. Imagine a picture of a

puppy sitting in the foreground amid a field of wildflowers. If

only the puppy and the flowers immediately around her are in

focus, the picture is said to have a sha llow depth of field. If the

puppy and the wildflowers all the way back to the horizon are

in focus, the picture has full depth of field. Full depth of field

is created by constricting the pupil (or the diaphragm on a

camera) so that only a narrow beam of light enters the eye. In

this way, a greater depth of the image is focused on the retina.

The lens Focuses light on the Retina When light enters the eye, it passes through the cornea and

lens prior to striking the retina. When light rays pass from air

into a medium of different density, such as glass or water, they

bend, or refract. Light entering the eye is refracted twice: first

when it passes through the cornea, and again when it passes

through the lens. About two-thirds of the total refraction

(bending) occurs at the cornea and the remaining one-third

occurs at the lens . Here we consider only the refraction that

RUNNING PROBLEM

The otolaryngologist strongly suspects that Anant has Meniere's disease, with excessive endolymph in the vestibular apparatus and cochlea . Many treatments are available, from simple dietary changes to surgery. For now, the physician suggests that Anant limit his salt intake and take diuretics, drugs that cause the kidneys to remove excess fluid from the body.

Question 5: Why is limiting salt (NaCO intake suggested as a treatment for Meniere's disease? (Hint: What is the relationship between salt, osmolarity, and fluid volume?)

334 · 1339 ~ 3S7 ~ [ 365 e.377 ~\379

occurs as light passes through the lens because the lens is capable of changing its shape to focus light.

When light passes from one medium into another, the

angle of refraction (how much the light rays bend) is influenced by two factors: (1) the difference in density of the two media and (2) the angle at which the light rays meet the sur

face of the medium into which it is passing. For light passing through the lens of the eye, we assume that the density of the lens is the same as the density of the air so that this factor can be ignored. The angle at which ligh t meets the face of the lens

depends on the curvature of the lens surface and the direction of the light beam.

Imagine parallel light ra ys striking the surface of a trans

parent lens. If the lens surface is perpendicular to the rays, the light passes through without bending. If the surface is not perpendicular, however, the light rays bend. Parallel light rays striking a concave lens, such as that shown in Figure 10-32a . ,

are refracted into a wider beam. Parallel rays striking a convex lens bend inward and focus to a point-convex lenses convelge light waves (Fig. 10-32b). You can demonstrate the properties

of a convex lens by using a magnifying glass to focus sunlight onto a piece of paper or other surface.

Concave lens


When parallel light rays pass through a convex lens, the

single point where the rays converge is called the focal point (Fig. 10-32b) . The distance from the center of a lens to its focal pOint is known as the focal length (or focal distance) of the

lens. For any given lens, the focal length is fi xed. For the focal length to change, the shape of the lens must change.

When light from an object passes through the lens of the eye, the focal point and object image must fall precisely on the retina if the object is to be seen in focus. In Figure 10-33a . ,

parallel light rays strike a lens whose surface is relatively flat. For this lens, the focal point falls on the retina. The object is

therefore in focus. For the human eye, any object that is 20 feet or more from the eye creates paralJel light rays.

What happens, though, when an object is closer than 20 feet to the lens? In that case, the light rays from the object are not parallel and strike the lens at an oblique angle that changes

the distance from the lens to the object's image (Fig. 10-33b). The focal point now lies behind the retina, and the object image becomes fuzzy and out of focus .

To keep a near object in focus, the lens must become more

rounded to increase the angle of refraction (Fig. 10-33c). 10 Making a lens more convex shortens its focal length. In this example, rounding the lens causes light rays to converge on the retina instead of behind it, and the object comes into focus.

The process by which the eye adjusts the shape of the lens to keep Objects in focus is known as accommodation, and the closest distance at which it can focus an object is known as the near pOint of accommodation. You can demonstrate chang

ing focus with the accommodation reflex easily by closing one eye and holding yo ur hand up about 8 inches in front of your

open eye, fingers spread apart. Focus your eye on some object in the distance that is vis

ible between your fingers. Notice that when you do so, your fingers remain visible but out of focus. Your lens is flattened for distance vision, so the focal point for near objects falls behind

the retina. Those objects appear out of focus. Now shift your gaze to your fingers and notice that they come into focus. The light rays coming off your fingers have not changed their

• FIGURE 10-32 Refraction ofConvex lens Focal point

light by lenses1

'7

Parallel Parallel i-Focallength-j light rays light rays

The focal length of the lens is the distance from the center of the lens to the focal point.

(a) A concave lens scatters light rays. (b) A convex lens causes light rays to converge.


(a) Parallel light rays pass through a flattened lens and the focal point falls on the retina.

(b) For close objects, the light rays are no longer parallel.

(e) Rounding a lens shortens its focal length .

Light from

distant

source

Light from

distant source \ ,-'

Lens flattened for distant vision

Focal length I" .1

Image distance

Object Object image

Focal length I .. • I

for close vision I .. • I..... • I I .. • I I .. • I Focal length Object Image distance (0) Image distance

distance (P) I" • I now equals focal length Focal length of lens (F)

The lens and its focal length have not changed To keep an object in focu s as it but the object is seen out of focus because the moves closer, the lens becomes light beam is not focused on the retina. more rounded.

FIGURE QUESTION The relationship between the focal length of a lens (F), the distance between an object and the lens (P), and the distance from the lens to the object's image (0) is expressed as 1/ F = 1/ P + 1/0.

(a) If the focal length of a lens does not change but an (b) If an object moves closer to the lens and the image distance 0 must object moves closer to the lens, what happens to the stay the same for the image to fallon the retina, what happens to the image distance O? focal length F of the lens? For this change in F to occur, should the lens

become flatter or more rounded?

• FIGURE 10-33 Optics of the eye

angle, but your lens has become more rounded, and the light Young people can focus on items as close as 8 cm, but the rays now converge on the retina. accommodation reflex diminishes from the age of 10 on. By age

How can the lens, which is clear and does not have any 40, accommodation is only about half of what it was at age 10,

muscle fibers in it, change shape? The answer lies in the ciliary and by age 60, many people lose the reflex completely because muscle, a ring of smooth muscle that surrounds the lens and the lens has lost flexibility and remains in its flatter shape for is attached to it by the inelastic ligaments ca lled zonules (Fig. distance vision. The loss of accommodation, presbyopia, is the

10-34 e ). If no tension is placed on the lens by the ligaments, reason most people begin to wear reading glasses in their 40s. the lens assumes its natural rounded shape because of the elas Two other common vision problems, near-sighted ness ticity of its capsule. If the ligaments pull on the lens, it flattens (m yopia) and far-sightedness (hyperopia), occur when the

out and assumes the shape reqUired for distance viSion . focal point falls either in front of the retina or behind the retina, Tension on the ligaments is controlled by the Ciliary mus respectively (Fig. 10-35 e ). These conditions are caused by ab

cle. When this circular muscle contracts, the muscle ring gets normally curved or flattened corneas or by eyeba lls that are too smaller, releasing tension on the ligamen ts so that the lens long or too short. Placing a lens with the appropriate curvature rounds. When the Ciliary muscle is relaxed, the ring is more in front of the eye Changes the refraction of light en tering the open and the lens is pulled into a flatter shape. eye and corrects the problem. A third common vision problem,


! E!

~

~ Ciliary muscle

/ /

Ciliary muscle Ciliary relaxed contracted

muscle

Lens CorneaCornea Lens flattened Lens rounded

Ligaments Iris LigamentsLigaments

slackenpulled tight ~

~II

(a) The lens is attached to the ciliary (b) When ciliary muscle is relaxed, the muscle by inelastic ligaments (zonules). ligaments pull on and flatten the lens.

• FIGURE 10-34 Changes in lens shape are controlled by the ciliary muscle.

(e) When ciliary muscle contracts, it releases tension on the ligaments and the lens becomes more rounded .

astigmatism, is usually ca used by a cornea that is not a perfectly

shaped dome, resulting in distorted images.

CONCEPT CHECK

27. If a person's cornea, which helps focus light, is more rounded than normal (has a greater curvature), is this person more likely to be hyperopic or myopic? (Hint: See Figure 10-35.)

AnS\Wls : p. 383

Hyperopia (corrected with a convex lens)

(al Hyperopia, or far-sightedness, occurs when the focal point falls behind the retina.

Myopia (corrected with a concave lens)

(b) Myopia, or near-sighted ness, occurs when the focal point falls in front of the retina.

FIGURE QUESTIONS

Explain (1) what the corrective lenses do to the refraction of light , and (2) why each lens type corrects the visual defect.

• FIGURE 10-35 Common visual defects can be corrected with external lenses.

Phototransduction Occurs at the Retina In the second step of the visual pathway, photoreceptors of the

retina convert light energy into electrical signals. Light energy

is part of the electromagnetic spectrum, which ranges from

high-energy, very-short-wavelength waves such as X-rays and o gamma rays to low-energy, lower-frequency microwaves and

radio waves (Fig. 10-36 e ). However, our brains can perceive

only a small portion of this broad energy spectrum. For hu

mans, visible light is limited to electromagnetic energy with

waves that have a frequency of 4.0-7.5 x 1014 cycles per second

(hertz, Hz) and a wavelength of 400-750 nanometers (nm).

Electromagnetic energy is measured in units called photons. Our unaided eyes see visible light but do not respond to

ultraviolet and infrared light, whose wavelengths border the

ends of our visible light spectrum. On the other hand, the eyes

of some other animals can see these wavelengths. For example,

bees Lise ultraviolet " runways" on flowers to guide them to

pollen and nectar.

RUNNING PROBLEM

Anant's condition does not improve with the low-salt diet and diuretics, and he continues to suffer from disabling attacks of vertigo with vomiting. In severe cases of Meniere's disease, surgery is sometimes performed as a last resort. In one surgical procedure for the disease, the vestibular nerve is severed. This surgery is difficult to perform, as the vestibular nerve lies near many other important nerves, including facial nerves and the auditory nerve. Patients who undergo this procedure are advised that the surgery can result in

deafness if the cochlear nerve is inadvertently severed.

Question 6: Why would severing the vestibular nerve alleviate Meniere's disease?

369 I» 319


380 nm

450 nm

500 nm

Visible 550 nm light

I

600 nm

650 nm

700 nm

>2> c (j) 750 nm

UJ

Gamma rays

X-rays

UV

Infrared

Microwaves

Radio waves

f-

10·Snm

10·3 nm

1 nm

103 nm

106 nm

109 nm (1 m)

103 m

e FIGURE 10-36 The electromagnetic spectrum

Pho tot ransduction is the process by which animals con

vert li ght energy into e lectrical signals. In humans, phototrans

duction takes place when light hits the retina, the sensory

organ of the eye. The retina develops from the same embryoniC

tissue as the brain, and (as in the cortex of the brain) neurons

in the retina are organized into layers (Fig. 10-37d e ). There are

five types of neurons in the retinal layers: photoreceptors, bipo

lar cells, ganglion cells, amacrine cells, and horizontal cells.

Backing the photosensitive portion of the human retina is

a dark pigment epithelium layer. Its function is to absorb any

light rays th at escape the p hotoreceptors, preventing distract

ing light from reflecting inside the eye and distorting the visual

image. The black color of these epi thel ial cells comes from

granules of the pigment melanin.

Pho toreceptors are the neurons that convert light energy

into electrica l signals. There are two main types of photorecep

to rs, rods and cones, as well as a recently discovered photore

ceptor that is a modified ga nglion cell (see Emerging Concepts

Box: Melanopsin) . You might expect photoreceptors to be on

the surface of the retina fa cing the vitreous chamber, where

light will strike them first , but the retina l layers are actually in

reverse order. The photoreceptors are the bottom layer, with

their photosensitive tips aga inst the pigment epithelium . Most

light entering the eye must pass through severa l relatively trans

parent layers of neurons before striki ng the photoreceptors.

One exception to th is organizational pattern occurs in a

small region of the retina know n as the fovea [pit]. In this area,

photoreceptors receive light directly because the intervening

neurons are pushed off to the side (Fig. 10-37c). The fovea is

also free of blood vessels that would block light reception. As

noted earlier, the fovea and the macula immediately surround

ing it are the areas of most acute vision, and they form the cen

ter of the visual field.

EMERG I NG CONCEPTS

MELANOPSIN

Circadian rhythms in mammals are cued by light entering the eyes. For many years, scientists believed that rods and cones of the retina were the primary photoreceptors linked to the suprachiasmatic nucleus (SCN), the brain center for circadian rhythms. However, in 1999 researchers found that transgenic mice lacking both rods and cones still had the ability to respond to changing light cues, suggesting that an additional photoreceptor must exist in the retina. Now scientists believe they have

found it: a subset of retinal ganglion cells that contain an opsin-like pigment called melanopsin . Axons from these ganglion cells project to the SCN, and it appears that these newly identified photoreceptors join rods and cones as the light-sensing cells of the mammalian retina.

When you look at an object, the lens focuses the object

image o n the fovea. For example, in Figure 10-38 e, the eye is

focused on the green-yellow border of the color bar. Light from

that section of the visual fi eld falls on the fovea and is in sharp

fo cus. Notice also tha t the image falling on the retina is upside

down. Subsequent visual processing by the brain reverses the

image again so th at we perceive it in the correct orientation.

Sensory in for matio n about light passes from the photore

ceptors to bipo lar neurons, then to a layer of ganglion cells.

The axons of ganglion cells form the optic nerve, which leaves

the eye at the optic disk . Beca use the optic disk has no photore

ceptors, images projected onto this region cannot be seen, cre

ating what is called the eye's blind spot.

CONCEPT CHECK

28. Animals that see well in very low light, such as cats and owls, lack a pigment epithelium and instead have a layer ca lled the tapetum lucidum behind the retina . What property might this layer have that would enhance vision in low light?

29. How is the difference in visual acuity between the fovea and the edge of the visual field similar to the difference in touch discrimina tio n between the fingertips and the skin of the arm?

30 . Macular degeneration is t he lead ing cause of blindness in Americans over the age of 55 . Impaired function of the macula causes vision loss in which part of the vi sua l field?

Answers: p. 383

Photoreceptors Transduce Light into Electrical Signals There are two types of photorecep tors in the eye: rods and

cones. Rods function well in low light and are used in night

pOint

(a) Dorsal view of a section

(b) Axons from the retina exit via the optic nerve.

THE RETINA ~~~~J::L~;:;z . -_

.L-ii::=g;;I.UI

Fovea

Rod

Cone

Optic nerve

Fixation Light Lens Retina Optic nerve - ---t-- Sclera

The choroid layer contains blood vessels.

Pigment epithelium of retina absorbs

of the left eye. excess light.

Bipolar neuron

Horizontal cell

Amacrine cell

Ganglion cell

(c) Light strikes the photoreceptors in the fovea directly because overlying neurons are pushed aside.

Light

Rod

Ganglion

Neurons where signals cell Cone (color vision) from rods and cones { Bipolar

are integrated cell Rod (monochromatic vision)

(d) Retinal photoreceptors are organized into layers. (e) Convergence in the retina

FIGURE QUESTION

How many rods converge on the ganglion cell in (e)?

• FIGURE 10-37 Drawing of photoreceptors in the fovea adapted from E. R.

Kandel et al., Principles of Neural Science, 3rd edition (McGraw Hill: New York, 2000) .

373


PIGMENT EPITHELIUM

Old disks at tip are phagocytized by pigment epithelial cells.

Melanin granules

OUTER SEGMENT

Visual pigments in membrane disks

Connecting stalks

INNER SEGMENT Mitochondria

Location of major organelles and metabolic operations such as photopigment synthesis and ATP production

Cone Rods ~"'\I'-'Irl

SYNAPTIC TERMINAL Synapses with bipolar cells

LIGHT

e FIGURE 10-39 Photoreceptors: rods and cones. The dark pigment epithe

lium absorbs extra light and prevents that light from reflecting back and distorting

vision . Light transduction takes place in the outer segment of the photoreceptor.

Changes in photoreceptor membrane potential alter neurotransmitter release onto

bipolar cells.

Fovea

The projected image is upside down on the retina . Visual processing in the brain reverses the image.

e FIGURE 10-38 Images project upside down onto the

retina.

vision, when objects are seen in black and white rather than in color. They outnumber cones by a 20:1 ratio, except in the fovea, which contains only cones.

Cones are responsible for high-acuity vision and color vision during the daytime, when light levels are higher. Acuity

means keenness and is derived from the Latin acuere, meaning "to sharpen ." The fovea, which is the region of sharpest vision, has a very high density of cones.

The two types of photoreceptors have the same basic structure (Fig. 10-39 e ) : (1) an outer segment whose tip touches the pigment epithelium of the retina , (2) an inner segment that contains the cell nucleus and organelles for ATP and protein synthesis, and (3) a basal segment with a synaptic terminal that releases glutamate onto bipolar cells.


In the outer segment, the cell membrane has deep folds that form disk-like layers, like candy mints stacked in a wrapper. Toward the tip of the outer segments in rods, these layers actually separate from the cell membrane and form free -floating membrane disks. In the cones, the disks stay attached.

Light-sensitive visual pigments are bound to the disk membranes in outer segments of photoreceptors. These visual pigments are transducers that convert light energy into a

change in membrane potential. Rods have one type of visual pigment, rhodopsin. Cones have three different pigments that are closely related to rhodopsin.

The visual pigments of cores are excited by different wavelengths of light allowing us to see in color. White light is a combination of colors, as revealed when you separate white light by

passing it through a prism. The eye contains cones for red, green, and blue light. Each cone type is stimulated by a range of light wavelengths but is most sensitive to a particular wavelength (Fig. 10-40 e ). Red, green, and blue are the three pri

mary colors that make the colors of visible light, just as red, blue, and yellow are the three primary colors that make different colors of paint.

The color of any object we are looking at depends on the wavelengths of light reflected by the object. Green leaves reflect green light, and bananas reflect yellow light. White objects reflect most wavelengths. Black objects absorb most wavelengths, which is one reason they heat up in sunlight while white objects stay cool.

Our brain recognizes the color of an object by interpreting the combination of signals coming to it from the three differ

ent color cones. The details of color vision are still not fully understood, and there is some controversy about how color is processed in the cerebral cortex. Color-blindness is a condition in which a person inherits a defect in one or more of the three types of cones and has difficulty distinguishing certain colors. Probably the best-known form of color-blindness is red-green, in which people have trouble telling red and green apart.

CONCEPT CHECK

31. Why is our vision in the dark in black and white rather than in color' Answers: p. 384

Phototransduction The process of phototransduction is Similar for rhodopsin (in rods) and the th ree color pigments (in cones). RhodopSin is composed of two molecules: opsin, a protein embedded in the membrane of the rod disks, and retinal, a vi tamin A derivative that is the light-absorbing portion of the pigment (see Fig. 10-39). In the absence of light, retinal binds

snugly into a binding site on the opsin (Fig. 10-41a e ). When activated by as little as one photon of light, retinal changes shape to a new configuration. The activated retinal no longer binds to opsin and is released from the pigment in the process known as bleaching (Fig. 10-41b).

Blue Green Red cones Rods cones cones

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Violet Blue Green Yellow Oranae Red

400 450 500 550 600 650 700

Wavelength (nm)

GRAPH QUESTIONS

• Which pigment absorbs light over the broadest spectrum of wavelengths?

• Over the narrowest? • Which pigment absorbs the most

light at 500 nm?

e FIGURE 10-40 Light absorption by visual pigments. There are three types of cone pigment, each with a characteristic

light absorption spectrum.

How does rhodopsin bleaching lead to action potentials traveling through the optical pathway? To understand the pathway, we must look at other properties of the rods. As you learned in Chapters 5 and 8, electrical signals in cells occur as a result of ion movement between the intracellular and extracellular compartments. Rods contain three main types of cation channels: cyclic nucleotide-gated channels (CNG channels) that allow Na+ and Caz+ to enter the rod, K+ chan

nels that allow K+ to leak out of the rod, and voltage-gated Caz+ channels in the synaptic terminal that help regulate exo

cytosis of neurotransmitter. When a rod is in darkness and rhodopsin is not active,

cyclic GMP (cGMP) levels in the rod are high, and both CNG and K+ channels are open. Sodium and Caz+ ion influx is greater than K+ efflux, so the rod stays depolarized to an average membrane potential of -40 mV (instead of the more usual-70 mY).

At this slightly depolarized membrane potential, the voltagegated Caz+ channels are open and there is tonic (continuous) re

lease of the neurotransmitter glutamate from the synaptic portion of the rod onto the adjacent bipolar cell (Fig. 10-41a).

When light activates rhodopsin, a second-messenger cascade is initiated through the G protein transducin (Fig. 1041b). (Transducin is closely related to gustducin, the G protein

~

10


(a) In darkness, rhodopsin is inactive, cGMP is high, and CNG and K· channels are open.

Pigment epithelium cell

Transducin -!-!----=Ui~~~-;;:::;;---~ (G protein)

Inactive ---"'------:-;-~o;{

rhodopsin (opsin and retinal)

levels high

~=::::!- Ca2+eNG channel Na+

open

Membrane potential in dark = - 40mV

Tonic release of neurotransmitter

onto bipolar neurons

FIGURE QUESTION

(b) Light bleaches rhodopsin . Opsin decreases cGMP, closes CNG channels, and hyperpolarizes the cell.

Membrane hyperpolarizes

to -70 mV

Neurotransmitter release decreases in proportion

to amount of light.

(e) In the recovery phase, retinal recombines with opsin.

One rod contains about 10,000 CNG channels open in the dark. One photon of light activates 1 rhodopsin. Each rhodopsin activates 800 transducin. Each transducin cascade removes 6 cGMP. A decrease of 24 cGMP closes one CNG channel. How many photons are needed to close all the CNG channels in one rod?

• FIGURE 10-41 Photo transduction in rods uses the visual pigment rhodopsin.

found in bitter taste receptors.) The transducin second-messenger cascade decreases the concentration of cGMP, which closes the CNG channels. As a result, cation influx slows or stops.

With decreased cation influx and continued K+ efflux, the

inside of the rod hyperpolarizes, and glutamate release onto the bipolar neurons decreases. Bright light closes all CNG channels and stops all neurotransmitter release. Dimmer light causes a re

sponse that is graded in proportion to the light intensity. After activation, retinal diffuses out of the rod and is trans

ported into the pigment epithelium. There it reverts to its inactive form before moving back into the rod and being reunited

with opsin (Fig. 1O-41c). The recovery of rhodopsin from bleach

ing can take some time and is a major factor in the slow adaptation of the eyes when moving from bright light into the dark.

CONCEPT CHECK

32. Draw a map or diagram to explain phototransduction. Start with bleaching and end with release of neurotransmitter.

Answers: p. 384

Signal Processing Begins in the Retina We now move from the cellular mechanism of light transduction to the processing of Ught signals by the retina and brain, the third

and final step in our vision pathway. Signal processing in the

--

On-center/ Off-center/ off-surround on-surround

field field

Bright light Q onto center 0 Bright light onto surround

Brighflignt onto surround ~ Diffuse light ~o on both center and surround

Excites Inhibits ganglion cell ganglion cell

Inhibits Excites ganglion cell ganglion cell

Weak response Weak response from from

ganglion cell ganglion cell

The retina uses contrast rather than absolute light intensity for better detection of weak stimuli.

• FIGURE 10-42 Ganglion cell receptive fields

retina is an excellent example of convergence [ ~ r . 282], in

which multiple neurons synapse onto a single postsynaptic cell (Fig. 1O-37e). Depending on location in the retina, as many as

15 to 45 photoreceptors may converge on one bipolar neuron. Multiple bipolar neurons in turn innervate a single ganglion

cell, so that the information from hundreds of millions of retinal photoreceptors is condensed down to a mere 1 million axons leaving the eye in each optic nerve. Convergence is min

imal in the fovea, where some photoreceptors have a 1: 1 relationship with their bipolar neurons, and greatest at the outer edges of the retina.

Signal processing in the retina is modulated by input from two additional sets of cells that we will not discuss (Fig.

10-37d). Horizontal cells synapse with photoreceptors and bipolar cells. Amacrine cells modulate information flowing between bipolar celis and ganglion cells.

Bipolar Cells After glutamate is released from photoreceptors onto bipolar neurons, signal processing begins. There are two types of bipolar celis, light-on and light-of( Light-on bipolar cells are inhibited by glutamate release in the dark and are acti

vated (released from inhibition) in the light. Light-off bipolar cells are excited by glutamate release in the dark and inhibited in the light. Whether glutamate is excitatory or inhibitory de

pends on the type of glutamate receptor on the bipolar neuron. In this fashion, one stimulus (light) creates two responses with a single neurotransmitter. Bipolar cells then form either excita

tory or inhibitory synapses with ganglion cells, the next neurons in the pathway.


Ganglion Cells We know more about ganglion cells because they lie on the surface of the retina, where their axons are the most accessible to researchers. Extensive studies have been

done in which researchers stimulated the retina with carefully I I I

placed light and evalua ted the response of the ganglion cells. Each ganglion cell receives information from a particular

area of the retina. These areas, known as visual fields, are

similar to receptive fields in the somatic sensory system I I I r

[ ~ p. 336]. The visual field of a ganglion cell near the fovea is quite small. Only a few photoreceptors are associated with each

I ganglion cell, and so visual acuity is greatest in these areas. At

the edge of the retina, multiple photoreceptors converging onto a single ganglion cell results in vision that is not as sharp.

I

I An analogy for this arrangement is to think of pixels on

i your computer screen. Assume that two screens have the I same number of "photoreceptors," as indicated by a maximal

screen resolution of 1280 x 1024 pixels. If screen A has one photoreceptor becoming one "ganglion cell" pixel, the actual screen resolution is 1280 x 1024, and the image is very clear.

If eight photoreceptors on screen B converge into one ganglion cell pixel , then the actual screen resolution falls to 160 x 128, resulting in a very blurry and perhaps indistinguish

able image. Visual fields of ganglion cells are roughly circular (unlike

the irregular sha pe of somatic sensory receptive fields) and they are divided into sections: a round center and its doughnutshaped surround (Fig. 10-42 e ). This organization allows each

ganglion cell to use contrast between the center and its surround to interpret visual information. Strong contrast between the center and surround elicits a strong excitatory response (a

series of action potentials) or a strong inhibitory response (no action potentials) from the ganglion cell. Weak contrast between center and surround gets an intermediate response.

There are two types of ganglion cell visual fields . In an oncenter/off-surround field, the associated ganglion cell responds most strongly when light is brightest in the center of the field (Fig. 10-42). If ligh t is brightest in the off-surround region of the

field, the ganglion cell is inhibited and stops firing action potentials. The reverse happens with off-centel/on-surroul1d fields.

What happens if light is uniform across a visual field? In that case, the ganglion cell responds weakly. Thus, the retina

uses contrast rather than absolute light intensity to recognize objects in the environment. One advantage of using contrast is that it allows better detection of weak stimuli.

Scientists have now identified multiple types of ganglion cells in the primate retina. The two predominant types, which

account for 80% of retinal ganglion cells, are M cells and P cells. Large magnocellular ganglion cells, or M cells, are more sensitive to information about movement. Smaller parvoce/llliar ganglion cells, or P cells, are more sensitive to signals that pertain to form and fine detail , such as the texture of objects in the

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