Chapter 16

Sense Organs
 Objectives
– Define receptor and sense organ.

– List the four kinds of information obtained from sensory receptors, and describe how the
nervous system encodes each type.

– Outline three ways of classifying receptors.

– List several types of somatosensory receptors.

– Describe the projection pathways for the general senses.

– Explain the mechanisms of pain and the spinal blocking of pain signals.

16-2
 General Senses

Senses throughout the body:

(touch, pressure, temperature, pain, osmolarity)

16-3
 Introduction

Receptor: a structure specialized to detect a stimulus.

– Some receptors are bare nerve endings.
– Others are true sense organs: nerve tissue surrounded by other tissues that
enhance response to a certain type of stimulus.

16-4
16-5
Unencapsulated Nerve Endings

– Free nerve endings

• For pain, hot and cold
• Skin and mucous membrane

– Tactile discs
• For light touch and texture

– Hair receptors
• Wrap around base hair follicle
• Monitor movement of hair

Encapsulated Nerve Endings

(Connective tissue enhances sensitivity or
selectivity of response)

– Bulbous (Ruffini) corpuscles —Tonic

• Heavy touch, pressure, joint movements, and skin stretching

– Lamellar (pacinian) corpuscles —Phasic

• Deep pressure, stretch, tickle, and
vibration

16-6
General Senses

16-7
Thermoreceptors and Nociceptors
(Temperature and Pain)

16-8
Proprioceptors
(Limb Position)

16-9
 Introduction

What happens when a receptor is activated?

Transduction - Conversion of stimulus energy (light, heat, touch, sound, etc.) into nerve signals.

Sensation- detection of the stimulus via receptors, subjective awareness of a stimulus.

Perception- Conscious interpretation at the cerebral cortex.

16-10
 Classification of Receptors
• By modality
– Thermoreceptors, photoreceptors, nociceptors, chemoreceptors (pH), and
mechanoreceptors (blood pressure)

• By origin of stimuli
– Exteroceptors: detect external stimuli
– Interoceptors: detect internal stimuli
– Proprioceptors: sense body position and limb movement

• By distribution
– General (somesthetic) senses: widely distributed
– Special senses: limited to head
• Vision, hearing, equilibrium, taste, and smell
 Sensory Information
Sensory receptors transmit four kinds of information:
Modality: Type of stimuli (vision, hearing and taste)

Location: (Receptive field) - detects stimuli

• What’s more important to your survival? Sensitivity to fingertips or back???
• Neurons in fingertips have small receptive fields allowing for fine two-point touch discrimination.

Intensity: Brain can distinguish stimulus intensity by:

– Number of receptors stimulated.
– How fast these receptors are firing.

Duration: How long the stimulus lasts. (sensory adaptation)

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 General Properties of Receptors

Sensory adaptation: if stimulus is prolonged, the firing of the neuron gets slower over time, and we
become less aware of the stimulus

– Phasic receptor: generate a burst of action potentials when first stimulated, then quickly adapt and
sharply reduce or stop signaling even though the stimulus continues.
• Smell, hair movement, and cutaneous pressure
• Can you feel your socks? your rings?

– Tonic receptor: adapt slowly, generate nerve signals more steadily

• Proprioceptors—body position, muscle tension, and joint motion
• Nociceptors: Is there a benefit to feeling pain…..chest pain?

16-13
 Somatosensory Projection Pathways
From receptor to final destination in the brain, most somesthetic signals travel by way of
three neurons
• First-order neuron (afferent neuron)
– From body, via spinal nerves
– From head, enter pons and medulla via cranial nerve

• Second-order neuron
– Decussation to opposite side in spinal cord,
medulla, or pons
– End in thalamus, except for proprioception, which ends in cerebellum

• Third-order neuron
– Thalamus to primary somesthetic cortex of cerebrum

16-14
 Pain

• Pain—discomfort caused by tissue injury or noxious stimulation, and typically leading to evasive action

• Injured tissues release chemicals that stimulate pain fibers

– Histamine, prostaglandin, and serotonin also stimulate nociceptors
– Bradykinin: most potent pain stimulus known
– Lost in diabetes mellitus—diabetic neuropathy
• Minor Pressure ulcers go undetected and developed into Major pressure ulcers.

16-15
16-16
 Referred pain
• Referred pain—Pain from an organ often mistakenly thought to come from the skin
or other superficial site
– Results from convergence of neural pathways in CNS

– Heart pain felt in shoulder or arm because both send pain input to spinal cord segments T1 to T5
16-17
 Central Nervous System: Pain Modulation
Analgesic (pain-relieving) mechanisms of CNS just beginning to be understood
• Endogenous opioids: internally produced opium-like substances:

– Enkephalins, endorphins, and dynorphins

– Secreted by the CNS, pituitary gland, digestive tract, and other organs

– Act as neuromodulators that can block the transmission of pain signals

and produce feelings of pleasure and euphoria

– Spinal gating –stops pain at the posterior horn of the spinal cord.
• Ex. Rubbing or Massaging an injury is a form of spinal gating.
• Rubbing stimulates spinal interneurons to secrete enkephalins
that inhibit second-order pain neurons

16-18
Special Senses:
Gustation

16-19
 Expected Learning Outcomes
– Explain how taste receptors are stimulated.

– Describe the receptors and projection pathways for taste.

16-20
 Special Senses - Gustation

Gustation (taste)—sensation that results from action of chemicals (tastants) on taste buds
(chemoreceptors).

Lingual papillae, visible to the naked eye

– Vallate (circumvallate): At rear of

tongue in a “V”; contains up to ½ of
taste buds.

– Foliate: taste buds; weakly

developed and degenerate by age 3.

– Fungiform: a few taste buds at tips

and sides of tongue.

16-21
Taste

16-22
 Taste

Taste (gustatory) cells

• Have taste hairs that serve as receptor surface for taste molecules
• Taste pores: pit in which the taste hairs project
• Taste hairs are epithelial cells, not neurons

Supporting cells
• Resemble taste cells without taste hairs

Basal cells
• Stem cells that replace taste cells every 7 to 10 days
16-23
 Taste – Vallate Papilla

Taste buds

Vallate papilla

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von Ebner’s glands
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 Taste (Gustation)
Taste bud Taste Buds Taste pore

Taste cells
Supporting cells
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 Taste
• Five primary sensations: Important for your survival!

– Salty: produced by metal ions (sodium and potassium) – electrolyte balance.

– Sweet: associated with carbohydrates and other foods of high caloric value. Sugars, saccharin, alcohol
and some amino acids – regulate blood sugar.

– Sour: acids such as in citrus fruits, hydrogen ions.

– Bitter: associated with spoiled foods and alkaloids such as nicotine, caffeine, quinine, and morphine.

– Umami: “meaty” taste of the amino acids (aspartic acid & glutamic acid) in chicken or beef
broth.

• Taste is influenced by food texture, aroma (taste is 80% smell) temperature, and appearance.

16-26
 Gustatory Pathway
Three cranial nerves carry impulses from taste buds to the solitary nucleus in medulla oblongata:
– Facial nerve (CNVII) collects sensory information from taste buds over anterior two-thirds of tongue
– Glossopharyngeal nerve (IX) from posterior one-third of tongue
– Vagus nerve (X) from taste buds of palate, pharynx, and epiglottis

16-27
 Projection Pathway of Taste
From there, signals sent to two destinations
– Hypothalamus and amygdala control autonomic reflexes: salivation, gagging, and vomiting.
– Thalamus relays signals to primary gustatory cortex of cerebrum for conscious sense of
taste.
Special Senses:
Olfaction
 Expected Learning Outcomes
– Explain how smell receptors are stimulated.
– Describe the receptors and projection pathways for smell.

16-30
 Olfaction
• Olfaction—sense of smell, action of odorants on
chemoreceptors

– Olfactory mucosa (traps odorants)

• Contains 10 to 20 million olfactory cells, which are
neurons

16-31
 Olfactory Mucosa
• Olfactory cells
– Only neurons in the body directly exposed to the external environment
– Have a lifespan of only 60 days

• Supporting cells

• Basal cells
– Divide and differentiate to replace olfactory cells
Ethmoid Bone and Olfaction

16-33
 Olfaction

Olfactory bulb
Olfactory nerve

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• 1. Chemicals stimulate neurons
Olfaction Pathway
• 2. Axons collect into bundles
• 3. First synapse in olfactory bulb
• 4. Travels olfactory tract →

1. Reach primary olfactory cortex in the inferior surface of the temporal lobe
• signals can reach the cerebral cortex without passing first through the thalamus

2. Secondary destinations: hippocampus, amygdala, hypothalamus, insula, and orbitofrontal cortex;

• Identify odors, integrate with taste, evoke memories, emotions, and visceral reactions.

16-35
 Olfactory Physiology
• Humans have a poorer sense of smell than most other mammals.
– Humans have only about 350 kinds of olfactory receptors.
– Women more sensitive to odors than men; especially to certain odors at time they are ovulating.

• Pheromones are chemicals (odors) that exerts subtle affects on the behavior and physiology of other people.

• Ovulating women’s vaginal secretions contain pheromones called copulins, that have been shown to raise
men’s testosterone level.
 Special Senses:
Hearing and Equilibrium

16-37
 Expected Learning Outcomes
– Identify the properties of sound waves that account for pitch and loudness.

– Describe the gross and microscopic anatomy of the ear.

– Explain how the ear converts vibrations to nerve signals and discriminates
between sounds of different intensity and pitch.

– Explain how the vestibular apparatus enables the brain to interpret the body’s
position and movements.

– Describe the pathways taken by auditory and vestibular signals to the brain.

16-38
 Special Senses -Hearing and Equilibrium
• Hearing—a response to vibrating air molecules
• Equilibrium—the sense of motion, body orientation, and balance
– Both senses reside in the inner ear, a maze of fluid-filled passages and sensory cells
– Fluid is set in motion and the sensory cells (mechanoreceptors) convert this motion into an informative pattern of
action potentials.

16-39
 Hearing
• Hearing—a response to vibrating air molecules
• A vibrating object pushes on air molecules- a mechanical wave.
• In turn push on other air molecules
• Air molecules hitting eardrum cause it to vibrate

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Ossicles: Semicircular ducts

Stapes Oval window
Helix
Incus
Malleus Vestibular nerve

Cochlear nerve

Vestibule
Auricle Cochlea
Round window
Tympanic
membrane
Auditory Tympanic cavity
canal
Tensor tympani
muscle
Auditory tube

Lobule

Figure 16.11
Outer ear Middle ear Inner ear
16-40
 Pitch & Loudness
• Pitch—our sense of whether a sound is “high” or “low”
– Frequency: cycles per second, or hertz (Hz)
– Human hearing range is 20 to 20,000 Hz, speech is 1,500 to 5,000 Hz, where hearing is most sensitive
– Hearing loss with age is 250 to 2,050 Hz

• Loudness—sound energy, intensity, or amplitude of the vibration

– Expressed in decibels (dB). Prolonged exposure to sounds > 90 dB can cause damage.

16-41
 Anatomy of the Ear

• Ear has three sections: outer, middle, and inner ear

– First two are concerned only with the transmission of sound to the inner ear
– Inner ear: vibrations converted to nerve signals

16-42
 Outer (external) Ear
Outer ear—a funnel for conducting vibrations to the tympanic membrane (eardrum)

Auricle (pinna) directs sound down the auditory canal

• Composed of helix (rim) and lobule (earlobe). Shaped and supported by elastic cartilage

External acoustic meatus (auditory canal)

• Short curved tube lined with skin bearing guard hairs, sebaceous glands and ceruminous glands (earwax).

16-43
 Middle Ear
Tympanic membrane (eardrum) closes the inner end of the auditory canal, separates it from the middle ear.
• Physiology: Vibrates freely in response to sound, transfers sound energy to the bones (ossicles) of the middle ear.

Auditory tube (eustachian tube) connects middle-ear cavity to nasopharynx

• Allows throat infections to spread to the middle ear
Auditory ossicles transmit and amplify sound from the tympanic membrane to the oval window.
– Malleus: attached to inner surface of tympanic membrane
– Incus: articulates in between malleus and stapes
– Stapes: footplate rests on oval window —inner ear begins

16-44
 The Auditory Tube of an
Infant and Adult

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 Tympanic Membrane with Otitis media
Otoscopic Photograph: Lateral

Tympanic
membrane

Wall of
Normal external
tympanic acoustic
membrane meatus

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 Auditory Ossicles (6)

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 Inner Ear
Cochlea—organ of hearing
• Cochlea has three fluid-filled chambers separated by membranes:

– Scala vestibuli: superior chamber Begins at oval window and spirals to apex. Filled with perilymph
– Scala media (cochlear duct): triangular middle chamber
• Filled with endolymph
• Contains spiral organ—Organ of Corti : acoustic organ that converts vibrations into nerve impulses
– Scala tympani: inferior chamber
• Filled with perilymph
• Begins at apex and ends at round window

16-48
 Hearing: Inner Ear
Endolymph (similar to intracellular fluid) and perilymph (similar to cerebrospinal fluid)

16-49
 Potassium Channels of the Cochlear Hair Cells

Excitation of Cochlear Hair Cells- Mechanoreceptors:

Each upward movement pushes hair cells toward the tectorial membrane and causes the stereocilia to bend
which opens a mechanically gated ion channel. K+ flows in the cell is depolarized.

Sensorineural (nerve) deafness: death of cochlear hair cells or any nervous system elements concerned with
hearing
• Factory workers, musicians, construction workers
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Basilar Membrane Frequency Response

16-51
 Cochlea Modiolus

Nerve fibers of cochlea Basilar membrane

Photos © McGraw-Hill Education

Scala tympani Cochlear duct

Scala vestubuli Vestibular

membrane

Spiral ligament of Stria vascularis

Spiral ganglion Spiral organ
cochlea

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 Basilar Vestibular
membrane Cochlea
Photos © McGraw-Hill Education
membrane

Cochlear duct Tectorial

membrane

Nerve fibers Stria vascularis

of cochlea

Scala tympani Scala vestibuli Spiral ganglion Spiral ligament Spiral organ
of cochlea

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 Spiral Organ
Basilar
Tectorial
membrane
membrane

Inner hair Outer

cells phalangeal
cells

Inner Outer
phalangeal hair
cells cells

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 Protection

Bony labyrinth—passageways in temporal bone

16-55
(c)
 Auditory Pathway
Auricle → external auditory meatus (canal) → tympanic membrane → malleus →incus →stapes→ oval window →auditory
hair cells (in spiral organ of Corti in cochlea)→ cochlear nerve → medulla →inferior colliculus (auditory reflex center in
midbrain) → thalamus (sensory integration center) →primary auditory cortex (temporal lobe)

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 Deafness

Conductive deafness: conditions interfere with transmission of vibrations to inner ear

• Damaged tympanic membrane , ossicles, or blockage of auditory canal
– Otitis Media – middle ear infection due to sore throats or infection of the throat that travel up the auditory tube
(eustachian tube)
– Otosclerosis: fusion of auditory ossicles that prevents their free vibration
Sensorineural (nerve) deafness: death of cochlear hair cells or any nervous system elements concerned with hearing
• Factory workers, musicians, construction workers
16-57
Equilibrium

16-58
 Equilibrium
Vestibular apparatus (inner ear)—contains mechanoreceptors for equilibrium in the semicircular canals and vestibule.
– Three semicircular ducts
• Detect only angular or rotational acceleration (dynamic equilibrium) – change in rate of rotation: car turns a corner,
spinning in chair .

– Vestibule: Anterior saccule and posterior utricle

• static equilibrium (head movement)
• dynamic equilibrium (linear acceleration) change in velocity in a straight line (elevator) along vertical or horizontal
planes – Vestibule.

16-59
Special Senses - Equilibrium

16-60
 The Saccule and Utricle

• Macula—a 2 by 3 mm patch of hair cells and supporting cells in the saccule and utricle.
• Each hair cell has 40 to 70 stereocilia embedded in a gelatinous otolithic membrane.

– Macula of saccule - Static equilibrium—when head is tilted, heavy

otolithic membrane sags, bending the stereocilia and stimulating the
hair cells.

– Macula of utricle - Dynamic equilibrium—in car, linear acceleration

detected as otoliths lag behind, bending the stereocilia and
stimulating the hair cells.

16-61
 The Semicircular Ducts

Rotary movements (dynamic angular acceleration) detected by the three semicircular ducts
(spinning in a chair)

– Ampulla (dilated external region):

• crista ampullaris:
– supporting cells
– hair cells with stereocilia
– gelatinous membrane called the cupula

– As head turns, endolymph lags behind, pushes

cupula, stimulates hair cells.

16-62
Vestibular Projection Pathways

Figure 16.22
 Pathologies

• Tinnitus: ringing or clicking sounds in the ears in the absence of auditory stimuli, due to cochlear
nerve degeneration; side effects of aspirin, inflammation of middle or inner ear.

• Meniere’s syndrome: disorder that affects the cochlea an semicircular canals. Causes vertigo,
nausea, and vomiting.

16-64
Special Senses:
Vision

16-65
 Expected Learning Outcomes
– Describe the anatomy of the eye and its accessory structures.
– Discuss the structure of the retina and its receptor cells.
– Explain how the optical system of the eye creates an image on the retina.
– Discuss how the retina converts this image to nerve signals.
– Describe the mechanism of color vision.
– Trace the visual projection pathways in the brain

16-66
 Accessory Structures of the Eye
• Eyebrows provide facial expression
– Protect eyes from glare and perspiration

• Eyelids (palpebrae)
– Block foreign objects, help with sleep,
blink to moisten
– Meet at corners (lateral and medial commissures)

• Eyelashes help keep debris from eye

• Conjunctiva—a transparent mucous membrane that lines

eyelids and covers anterior surface of eyeball, except
cornea. Richly innervated and vascular.
– Conjunctivitis – inflammation of the conjunctiva;
results in irritated, red eyes
– Pinkeye – highly contagious bacterial or viral
infection of the conjunctiva

16-67
 Accessory Structures of the Eye
.

Trochlea

Optic nerve Trochlea

Muscles: Superior oblique

Muscles: tendon
Superior oblique Superior oblique
Superior rectus
Medial rectus Muscles:
Medial rectus
Superior rectus
Lateral rectus Inferior rectus
Lateral rectus

Inferior oblique Levator palpebrae

superioris (cut)
Inferior rectus
Figure
16.24a

(a) Lateral view

(b) Superior view

• Six extrinsic muscles attached to exterior surface of eyeball.

– Attached to bony orbit,
– Enable the eye to follow moving objects.
– Maintain the shape of the eyeball
– Strabismus – cross-eyed due to eye rotating medially or laterally; aka Lazy Eye
16-68
 Extrinsic Muscles
• Innervated by cranial nerves
• CN IV innervates superior oblique
• CN VI innervates lateral rectus
• CN III innervates other four extrinsic muscles

Figure 16.25c
 Accessory Structures of the Eye: Lacrimal
Apparatus

• Lacrimal secretions (tears), a dilute saline solution containing mucus, are produced by the lacrimal gland, flow across
eyeball to help wash away foreign particles, deliver O2 and nutrients, and prevent infection with a bactericidal lysozyme and
antibody A.

• Tears flow through lacrimal punctum (opening on edge of each eyelid) via the lacrimal canal to the lacrimal sac, then
into the nasolacrimal duct emptying into nasal cavity 16-70
 Anatomy of the Eye
Sclera
. Ora serrata
Choroid
Ciliary body
Retina

Macula lutea Suspensory

ligament
Fovea centralis

Optic disc Iris

(blind spot)
Cornea
Optic nerve Pupil

Lens
Central artery
and vein
of retina Anterior
chamber
Posterior
chamber
Hyaloid canal

Vitreous body

• Three principal components of the eyeball

– Three layers (tunics) that form the wall of the eyeball
• Optical component admits and focuses light
• Neural component retina and optic nerve
16-71
 The Tunics

• Tunica fibrosa—outer fibrous layer. Dense avascular

connective tissue
– Sclera: dense, collagenous white of the eye, protects
and shapes eye
– Cornea: transparent area of sclera that admits light into
eye

• Tunica vasculosa (uvea)—middle vascular layer

– Choroid: highly vascular, deeply pigmented layer behind
retina, blocks stray light rays.
– Ciliary body: extension of choroid that forms a muscular
ring around lens
• Supports lens and iris
• Secretes aqueous humor
– Iris: colored diaphragm controlling size of pupil, its
central opening

• Tunica interna (sensory) - retina and beginning of optic

nerve

16-72
 The Optical Components
• Cornea: transparent anterior cover.

• Aqueous humor: Serous fluid secreted by ciliary body

into posterior chamber: posterior to cornea, anterior to
lens.

• Lens: Changes shape to help focus light (primary

function).

• Vitreous body (humor): Jelly like substance that fills

space between lens and retina, keeps retina flat
against sclera, helps maintain shape of eye.
 The Optical Components
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Cornea

Anterior chamber
Scleral Iris
venous sinus
Ciliary body: Posterior
Ciliary chamber
process
Lens
Ciliary Vitreous
muscle body

Figure 16.26

• Aqueous humor is released by ciliary body into posterior chamber,

passes through pupil into anterior chamber, then reabsorbed into
scleral venous sinus 16-74
 Refraction
Emmetropia—state in which the eye is relaxed and focused on an object more than 6 m (20 ft) away. Normal vision (20/20)
• Light rays coming from that object are essentially parallel
• Rays focused on retina without effort
Myopia (nearsightedness) – focal point is in front of the retina, e.g. longer than normal eyeball
Hyperopia (farsightedness) – focal point is behind the retina., e.g. in a shorter than normal eyeball
Presbyopia – age related disorder where lens becomes inelastic and cannot accommodate (focus on near objects). Occurs normally around
45 years old, and is fixed by magnifying reading glasses.
Astigmatism – caused by unequal curvatures in different parts of the cornea or lens.

16-75
 The Neural Components
• Optic nerve (optic disc)

• Retina
– Attached to the rest of the eye only at optic disc (posterior) and at ora serrata (anterior)
• Optic Disc (blind spot) site where the optic nerve leaves the eye and there are not photoreceptor cells.

• Blind spot: use test illustration above

• Close right eye, stare at X and red dot disappears
• Visual filling: brain fills in green bar across blind spot area

– Pressed against rear of eyeball by vitreous humor

– Macula lutea—cells on the visual axis of eye (3 mm)
– Fovea centralis: center of macula; finely detailed images due to packed photoreceptor cells. Concentration of cones.
– Detached retina causes blurry areas in field of vision and leads to blindness

16-76
The Fundus (Rear) of the Eye
 Retina
Ophthalmoscopic Photograph - Anterior

Fovea centralis Optic disc

Macula lutea Retinal vessels

Photos © McGraw-Hill Education

 Photoreceptors

Cone cells (color, photopic, or day vision) Bright light, high acuity, central
– Three types of cones:
• Long-wave lengths cones (red) respond to low frequency (vibration)
Wavelength = frequency (vibrations)
• Medium-wave lengths cones (green)
• Short-wave lengths cones (blue) respond to high frequency (vibration)

Rod cells (night, or scotopic, vision or monochromatic vision) Dim light, peripheral vision
– Provide indistinct, fuzzy, non-color peripheral vision
– Perceived input is in gray tones only – black/white

16-79
 Visual Pigments

Rhodopsin Hyperpolarizes the photoreceptor (OFF SWITCH)

16-80
Visual Pigments

16-81
 Visual Pigments

Rods = rhodopsin (visual pigment):

– Opsin—protein portion embedded in disc membrane of rod’s outer segment
– Retinal (retinene)—a vitamin A derivative
– Night Blindness – when rod function is impaired due to Vitamin A deficiency.

Rhodopsin production increases at night!

Cones = photopsin (iodopsin)

– Opsin contains different amino acid sequences than rods that determine wavelengths of light absorbed
– Retinal same as in rods
– Color Blindness – congenital lack of one or more of the cones
16-82
Sensory Transduction in the Retina

16-83
 The Visual Projection Pathway

Cornea → aqueous humor of anterior segment → lens → vitreous humor of posterior segment → retina
(photoreceptors → bipolar cells → ganglion cells) → optic nerve → optic chiasm → optic tract → thalamus
(sensory integration center) → primary visual cortex (occipital lobe) 16-84
 The Near Response

Near response—adjustment to close-range vision requires

three processes

– Convergence of eyes
• Eyes orient their visual axis toward object

– Constriction of pupil
• Blocks peripheral light rays and reduces spherical
aberration (blurry edges)

– Accommodation of lens
• Ciliary muscle contracts, lens takes convex shape
• Light refracted more strongly and focused onto retina

16-85
 Cataracts and Glaucoma

• Cataract
– Induced by diabetes, smoking, drugs,
ultraviolet radiation, and certain viruses
– Lens fibers darken with age, fluid-filled
bubbles and clefts filled with debris appear
between the fibers —clouding of lens
– Replace natural lens with plastic one

• Glaucoma—elevated pressure within the eye

due to obstruction of scleral venous sinus
and improper drainage of aqueous humor
– Death of retinal cells due to compression
of blood vessels and lack of oxygen

16-86