Senses:

The ability to generate a sensation of a stimulus

General Senses
Sensations from receptors that are present in already utilized organs
Somatosensory = integument. Touch, vibration, temperature, pain…
Proprioception = skeletal muscle & tendons. Position of your body in space relative to
itself
Special Senses
Sensations from receptors in specialized organs NOT otherwise utilized
Olfaction (smell), Gustation (taste), Vision, Equilibrium (balance), Hearing

Sensory receptor cells

2 types: 1.) Dendrites of specialized neurons 2.) Specialized cells that synapse with sensory
neurons

Depolarization of sensory neuron = a

generator potential
Depolarization of sensory neuron

Module 15.1: Generator potential

Sensory Receptor Cells
Olfactory Sensory receptors
are dendrites of specialized neurons. Dissolved odorants bind to olfactory receptors
Receptors create generator potential
Weak stimulus=Small generator potential & No action potential (AP)
Strong stimulus= Larger generator potential & AP

Sensory Receptor Cells

for taste, vision, equilibrium, hearing are specialized cells with non-excitable membranes
and form synapses with the processes of sensory neurons
Graded potential neurotransmitter release
Neurotransmitter generator potential (neuron)  AP
Action potentials are propagated to CNS
Stimulation graded potential (receptor cell)
No AP

Olfaction = sense of smell

Olfactory Sensory receptors Paired olfactory organs in nasal cavity
are dendrites of specialized neurons. Dissolved odorants bind to olfactory receptors
Receptors create generator potential
Weak stimulus=Small generator potential & No action potential (AP)
 Strong stimulus= Larger generator potential & AP

Olfactory Sensory receptors

are dendrites of specialized neurons. Dissolved odorants bind to olfactory receptors
Receptors create generator potential
Weak stimulus=Small generator potential & No action potential (AP)
Strong stimulus= Larger generator potential & AP

Receptors:
olfactory neuronal receptor cells are distributed along cribriform plate, superior portion of the
perpendicular plate, superior nasal conchae
Each forms knob with up to 20 cilia-shaped dendritesprojecting into mucus
10–20 million receptors in 5-cm2

Olfactory organs have 2 layers:

1.) Olfactory epithelium
Olfactory receptor cells (modified neurons). Each forms knob with up to 20 cilia-shaped
dendrites projecting past epithelial surface into mucus. 10–20 million receptors in a 5-cm2
area. Olfactory receptors are stimulated by odorants
Supporting cells: Basal cells (stem cells). Constantly produce new receptor cells. One of
the few examples of neuronal replacement
2.) Lamina propria
Areolar tissue, blood vessels, nerves
Olfactory glands—secretions absorb water; form thick, pigmented mucus
Odorants are chemicals dissolved in the air that bind membrane receptors (odorant-binding
proteins) on dendrites of olfactory receptor cells. They are generally small organic molecules. As
few as four odorant molecules can activate receptor cell
Olfactory pathway to the cerebrum : Chemicals in air bind odorant-binding receptors stimulate
the sensory neurons in the olfactory organ. Axons leave the olfactory epithelium in bundles
passing through the cribriform plate (ethmoid)
First synapse occurs in olfactory bulb immediately superior to cribriform plate. Axons leaving
bulb travel through olfactory tract to olfactory cortex, hypothalamus, & limbic system .
Distribution to limbic system (emotions) & hypothalamus (ANS) explains why smells produce
emotional & behavioral response

Oddities of the olfactory system are that there are 400 types of receptors for 1 trillion scents
(Nature). Dogs can smell 10k-100k better (PBS). Strong connection to emotions. Comfort food,
smell of a loved one, fresh air, Danger etc. Incorrect smell identifications. Altered sense of smell
when ill.
 Olfactory receptors. The thousands of olfactory receptors, receptors for
the sense of smell, occupy a postage stamp-sized area in the roof of each
nasal cavity.
  Olfactory receptor cells. The olfactory receptor cells are neurons
equipped with olfactory hairs, long cilia that protrude from the nasal
epithelium and are continuously bathed by a layer of mucus secreted by
underlying glands.
 Olfactory filaments. When the olfactory receptors located on the cilia are
stimulated by chemicals dissolved in the mucus, they transmit impulses
along the olfactory filaments, which are bundled axons of olfactory
neurons that collectively make up the olfactory nerve.
 Olfactory nerve. The olfactory nerve conducts the impulses to the
olfactory cortex of the brain.

GUSTATION
 Taste buds. The taste buds, or specific receptors for the sense of taste,
are widely scattered in the oral cavity; of the 10, 000 or so taste buds we
have, most are on the tongue.
 Papillae. The dorsal tongue surface is covered with small peg-like
projections, or papillae.
 Circumvallate and fungiform papillae. The taste buds are found on the
sides of the large round circumvallate papillae and on the tops of the
more numerous fungiform papillae.
 Gustatory cells. The specific cells that respond to chemicals dissolved in
the saliva are epithelial cells called gustatory cells.
 Gustatory hairs. Their long microvilli- the gustatory hairs- protrude
through the taste pore, and when they are stimulated, they depolarize
and impulses are transmitted to the brain.
 Facial nerve. The facial nerve (VII) serves the anterior part of the tongue.
 Glossopharyngeal and vagus nerves. The other two cranial nerves- the
glossopharyngeal and vagus- serve the other taste bud-containing areas.
 Basal cells. Taste bud cells are among the most dynamic cells in the body,
and they are replaced every seven to ten days by basal cells found in the
deeper regions of the taste buds.

The tongue has papillae --> Papillae have taste buds --> Taste buds have gustatory receptors -->
Gustatory receptors can detect tastes
Gustation = taste
Receptors: Gustatory receptor cells, Found in taste buds
The receptors for taste and olfaction are classified as chemoreceptors because they
respond to chemicals in solution.

Taste buds: Sensory structures with taste receptors that respond to various chemical stimuli
Superior surface of tongue & pharynx, larynx, epiglottis. Numbers decrease with age. Located in
lingual papillae
Lingual papillae: epithelial projections on tongue surface. Four types of lingual papillae. Differ
in # of taste buds. Differ in location

Lingual papillae =
epithelial projections on tongue surface. Contain taste buds = sensory structures with
taste receptors that respond to various chemical stimuli. Four types of lingual papillae: 1.)
Vallate papillae, 2.) Foliate papillae, 3.) Fungiform papillae & 4.) Filiform papillae
Vallate (vallum, wall) papillae:
Relatively large—each contains up to 100 taste buds. Surrounded by deep epithelial
folds. Form “V” at back of tongue
Foliate papillae
Lateral margins of posterior region of tongue
Fungiform (fungus, mushroom) papillae
Anterior & lateral regions of the tongue. ~5 taste buds each
Filiform (filum, thread) papillae
Provide friction to help tongue move objects around mouth. Anterior two-thirds of
superior surface of tongue. NO taste buds
Four primary taste sensations: Sweet, salty, sour, bitter. No difference in taste bud structure
Taste buds in all portionsof tongue provide all four sensations. They are more prominent in
certain areas
Umami = pleasant, savory taste, characteristic of broths, cheese. Binds receptors for amino
acids, small peptides, nucleotides. Plenty of work on additional tastes (carbs, fats, calcium)
Water receptors
are concentrated in pharynx. Output goes to hypothalamus. Affects water balanceand
regulation of blood volume. Prevent over-ingesting H2O
Taste buds are recessed into epithelium
Gustatory receptor cell (taste receptor cell). Extends slender microvilli (taste hairs) into
surrounding fluids through a taste pore .Typically lives about 10 days before it is replaced .
About 40–100 gustatory cells in each taste bud

Basal cells = stem cells that divide and mature to produce transitional cells that mature into new
gustatory cells
Taste receptor sensitivity: Threshold varies for the four primary sensations.
More sensitive to unpleasant stimuli. 100,000 times more sensitive to bitter; 1000 times more
sensitive to sour (acids) than to sweet or salty
Survival value—acids burn tissues; many toxins are bitter
Sensitivity changes with exposure or lack thereof
AGE Overall sensitivity declineswith age—leads to changing taste sensations with age
Gustation stimulated by dissolved chemicals . Chemicals elicit receptor response by:Diffusion
through plasma membrane leak channels & bind to the receptor proteins of the gustatory receptor
cell.

Different tastes involve different mechanisms. Signals transmitted onto a sensory neuron. ~ 90%
of gustatory receptor cells respond to at least two taste stimuli

Gustatory reception (2 types):

1. ) Salt and sour receptors
Na+ (salt) or H+ (sour) diffuse through Na+ leak channels. Then the membrane/cell
depolarizes. Neurotransmitters are released at synapse with sensory neurons.
Depolarization of sensory neurons leads to generator potential, which can produce action
potentials along gustatory pathway to CNS
2.) Sweet, bitter, and umami receptors:
Binding to these receptors activates G-protein complexes calledgustducins, which are
protein complexes that use second messengers. Activated second messenger causes the
release of the neurotransmitter. Neurotransmitters are released at synapse with sensory
neurons

Gustatory pathway—gustatory receptor cells to cerebral cortex

Gustatory receptor cells respond to stimulation. Information is relayed on cranial nerves
VII, IX, and X. Sensory afferents synapse in medulla oblongata. Postsynaptic neurons
cross over & project to thalamus. Synapse in thalamus, then go to gustatory cortex in
insula for conscious perception. Gustatory receptor cells bind dissolved chemicals.
Generator potential occurs and triggers action potentials. Information is relayed on
cranial nerves facial nerve (VII). Anterior 2/3 of tongue,from the from the tip to the
vallate papillae. Glossopharyngeal nerve (IX). Vallate papillae, posterior 1/3 of tongue.
Vagus nerve (X) Epiglottis. Sensory afferents synapse in the solitary nucleus of medulla
oblongata. Axons of postsynaptic neurons cross over; enter the mediallemniscus ofthe
medulla oblongata. Then Synapse in thalamus; then information is projected to
appropriate portions of gustatory cortex of the insula.
Gustatory reception pathway
Receptors bind chemicals. Generator potential occurs and triggers AP. Information is relayed on
cranial nerves. Sensory afferents synapse medulla oblongata. Axons cross over, synapse in
thalamus; projects to appropriate portions of gustatory cortex of the insula

Taste: Sensation signal is sent to the gustatory cortex. Information from the taste buds is
integrated with other sensory data in association areas to create perception
Texture of food. Taste-related sensations (“peppery” or “burning hot”). Smell of food and sight
of food
Adaptation
Taste receptors adapt slowly
Central adaptation reduces sensitivity quickly
Olfactory stimulation plays important role. We are several thousand times more sensitive to
“tastes” when sense of smell is functioning.

Vision
Eyes are our most complex sense organs. LOTS of processing occurs inthe eye before relayed to
CNS. Resembles detached CNS nuclei
Eye development: Begins as pair of bulges = optic vesicles. Each contains cavity continuous
with neural tube. Form in prosencephalon walls
Lateral bulges indent; form optic cups—connected to diencephalon. The overlying epidermis
formspocket; pinches off to formthe lens. Inner and outer layers form retina. Outer ependymal
cells become photoreceptors. Inner ependymal cells become pigment cells. Nervous tissue of
outer layer forms neurons, ganglion cells, specialized glial cells
Embryonic cells around optic cup form supporting layers of connective tissue. Isolate nervous
tissue . Interior chambers develop; filled with fluid
Eye accessory structures
Eyelashes = hairs that keep matter out of eyes
Eyelids (palpebrae) = continuation of skin; can close to protect eye. Blinking lubricates eye
surface; clears dust/debris
Medial angle of the eye = where eyelids meet medially
Lateral angle of the eye = where eyelids meet laterally
Palpebral fissure = gap between upper/lower eyelids
Iris = colored portion of the eye
Pupil = opening in center of iris; transmits light
Lacrimal caruncle = small, reddish body at medial angle . Produces thick secretions that may
cause gritty deposits after night’s sleep (sleepy dust)
Conjunctiva–mucous membrane and epithelium lining eyelids and covering anterior of eye
Palpebral conjunctiva—inner surface of eyelids
Bulbar conjunctiva—anterior eye surface; continuous with cornea
Fornix—pocket created at junction of conjunctiva
Tarsal glands = modified sebaceous glands. Inner margin of eyelids. Secretions prevent eyelid
sticking
Conjunctivitis, or pinkeye is the inflammation of the conjunctiva. Redness due to dilation of
blood vessels deep to conjunctival epithelium. From pathogenic infection or physical, allergic, or
chemical irritation of conjunctival surface

Lacrimal apparatus
Lacrimal gland. Produces tears which lubricate, nourish, oxygenate, clean cornea . The tear
ducts consist of 10–12 tubes
Deliver tears behind upper lateral eyelid . Forms lacrimal “lake”
Lacrimal canaliculi = connect lacrimal puncta to lacrimal sac
Lacrimal sac—small chamber nestled in lacrimal sulcus of orbit

Internal Visual Anatomy

3 layers (tunics) of the eye:
1.) Outer fibrous layer is the outermost layer of eye, consists of Sclera, corneal limbus, and
cornea.

2.) Middle vascular layer is the Iris, choroid and ciliary body. Vascular layer (uvea) is
blood/lymph vessels and intrinsic (smooth) muscles of eye.
Iris = colored part of eye. Iris consists of blood vessels, pigment cells (melanocytes)
Two layers of smooth muscle—contraction changes diameter of pupil to control amount of light
entering
Ciliary body = thickened region bulging into interior of eye. The Ciliary body consists of ring of
fibers that connect the ciliary body and lens
Choroid = vascular layer underlying sclera; has extensive capillary network supplying oxygen/
nutrients to neural layer
3.) Inner layer, or retina = innermost layer of eye. The outer pigmented layer of the retina
absorbs light.

Eye cavities—separated by lens/ciliary body

Anterior cavity: Cornea to lens; contains aqueous humor fluid, It consists of two chambers: the
anterior chamber—cornea to iris and the posterior chamber—iris to ciliary body & lens.
Posterior cavity : Lens to retina. Vitreous Body filling, gelatinous
Vitreous humor = fluid part of vitreous body
Ciliary body of the anterior cavity:
Ciliary muscle—smooth muscle ring, projects inward
Ciliary processes—folds of epithelium covering ciliary muscles
Ciliary zonule (suspensory ligaments)—holds lens in position so light passes through it
The anterior cavity (continued)
Aqueous humor
Circulates within anterior cavity, passes through pupil. Secreted by epithelial cells of ciliary
processes. Diffuses through vitreous body to retinal surface (vitreous humor). Leaves eye at
scleral venous sinus (canal of Schlemm) = passageway around eye at level of corneoscleral
junction. Flows into veins in sclera. Rate of removal should keep pace with rate of secretion
Functions to transports nutrients and wastes, forms fluid cushion, helps retain eye shape and
tabilizes position of the retina

Iris
Body of iris is highly vascular, pigmented loose connective tissue. Anterior surface—incomplete
layer of fibroblasts/melanocytes . Posterior surface—lined by pigmented epithelium of neural
layer
Oro serrata—jagged edge of neural layer of retina
Eye color—determined by genes that influence density and distribution of melanocytes and
density of pigmented epithelium

Cornea Transparent and clear. Allows light to enter eye

Dense matrix of multiple layers of collagen fibers
Avascular; receives oxygen and nutrients from tears

Lens focuses light onto retina, Manipulated by ciliary body

Posterior to cornea; anchored by ciliary zonule of ciliary body, Concentric layers of cells filled
with transparent crystallins = proteins responsible for clarity and focusing power of lens.
Dense fibrous capsule surrounds cells; blends with ciliary zonule. Primary function of lens is it
changes shape to focus image on photoreceptors.

Retina Light receptors, Absorbs & responds to light. The retina contains photoreceptors,
pigment cells, supporting cells, neurons (continued in detail further down).

Choroid nourishes all eye structures. The chroroid is the middle layer; blood vessels nourish all
eye structures
Sclera Stabilize eye while moving

Sclera (“white of the eye”)—dense fibrous connective tissue with collagen and elastic fibers.
The sclera stabilizes eye shape during movement. Insertion for extrinsic eye muscles

Optic nerve (II)—conveys visual information to brain

Ciliary body—supports lens, controls its shape; tension in ciliary zonule resists tendency of lens
to ball up

LIGHT TURNS TO VISION:

The cornea is the first eye structure hit by light, it is transparent & clear.100% transmission.
Light is bent Gas to solid to liquid transition, the degree of bend is constant.
Lasik-Laser eye surgery altering the shape of the cornea

The pupil is the opening in iris through which light passes. Two pupillary muscles of iris
regulate amount of light entering . The dilator pupillae muscles—extend radially from pupil.
Enlarge pupil; supplied by sympathetic nervous system . The sphincter pupillae muscles—
encircle pupil like a ring. The sphincter pupillae muscles make pupil smaller; supplied by
parasympathetic nervous system
The visual axis is the imaginary line drawn from center of object you are looking at through the
center of the cornea and lens to retina
Focusing is a two-step process:1. Light is refracted (bent) passing from air into cornea
Bending occurs because of the change in density. The amount of refraction at cornea is constant
2. More refraction when light passes from aqueous humor into lens causes light rays to bend
toward focal point—specific point on retina
Focal distance of a lens is the distance between center of lens and its focal point, it's determined
by the distance from an object to the lens and the shape of the lens. The distance from the lens to
the retina cannot change. Focusing is done by changing the shape of the lens = accommodation

Accommodation = change in lens shape to keep focal distance constant and provide clear vision
Ciliary body
Zonules – strings attaching lens to choroid. At rest, choroid pulls on zonules/lens
Muscle – Connected to choroid. Contraction stretches choroid

Accommodation For close vision:

Light enters eye at a greater angle. Ciliary muscle contracts. Reduces tension in ciliary zonule.
Ciliary body moves toward lens. Lens more spherical. Increases refraction. Light from near the
object is focused on retina

Accommodation For distant vision:

Light enters the eye at less of an angle. Ciliary muscle relaxes. Ciliary zonule pulls on lens. Lens
flattens; brings image of distant object into focus on retina

Emmetropia = normal vision. When ciliary muscle is relaxed and lens flattened, distant image is
focused on retinal surface
Myopia = nearsightedness. Focal distance too short. Image focuses in front of retina. Resting
curvature of lens too great. Vision near images is clear, and vision for far images is
[Link]: Concave lens in front of eye, Refracts light rays apart to shift focus forward
onto retina.
Hyperopia = farsightedness. Focal distance is too long, so image focuses beyond retina. Lens is
too flat
Vision near images is blurry, vision for far images is clear. Correction: Convex lens in front of
eye. Refracts light inwards (together).

Surgical corrections:
Photorefractive keratectomy (PRK)
Computer-guided laser shapes cornea, removes 10–20 µm (< 10%) of cornea. This is completed
in less than a minute!

Laser-assisted in-situ keratomileusis (LASIK)

Interior corneal layers reshaped; covered by normal corneal epithelium. About 70% of patients
achieve normal vision, and about 10 million Americans have had it. Immediate and long-term
visual problems can occur with LASIK.

Near point of vision = inner limit of clear vision, determined by lens elasticity, Increases with
age as lens becomes less elastic. In children 7–9 cm (3–4 in.), in young adults 15–20 cm (6–8 in.)
and in ages near 60, about 83 cm (33 in.)

Retina
Formed by photoreceptors, pigment cells, supporting cells, and other neurons
Two main layers 1.) Neural layer which is the superficial layer. The neural layer consists of
photoreceptors, supporting cells and neurons. The neural layer is where the initial
processing/integration of visual information happens. Photoreceptors located closest to
pigmented layer, information travels superficially. 2.) Pigmented layer is the deep layer
consisting of light absorbing cells. The pigmented layer absorbs light that passes through neural
layer. The pigmented layer prevents light from bouncing back, producing visual “echoes”.
Ganglion cells are located in the neural layer . Axons converge at optic disc to form optic nerve ,
this is where te location of the Blind spot is, it is a blind spt because it lacks photoreceptors.
Blood vessels follow optic nerve, supply inner neural layers cells
Photoreceptors are light sensitive cells, they come in two types.
1.) Rods which are for black-and-white vision. Rods are highly sensitive, they allow vision in
very dim light
2. ) Cones are for color vision. Cones give us sharper, clearer images but require more intense
light.

Density of photoreceptors changes along retina

Macula is the region of higher density
Fovea is the region within macula of greatest density. Location of highest visual acuity (sharpest
vision)
Blind spot lacks of photoreceptor due to optic nerve and blood vessels

Retinal layers and cells

Photoreceptors of the retina
Horizontal and amacrine cells Facilitate/inhibit communication between photoreceptors and
ganglion cells.
Horizontal cells—where photoreceptors/bipolar cells synapse
Amacrine cells—where bipolar and ganglion cells synapse
Alter sensitivity of retina; major role in adjusting to dim or bright light
Photoreceptor distribution
Cones: about 6 million per eye. Cone density is highest at fovea [Link] rods there. Cone
density directly correlates with visual acuity (sharpness)
Rods: ~125 million per eye. Maximum density at periphery. Few cones there. Effects are
detection of movement and peripheral vision in dark
Visual pigments transduce light. Derived from rhodopsin (visual purple); pigment in rods
Have opsin (protein; determines wavelength absorbed) and retinal (pigment synthesized from
vitamin A)

Photoreceptor structure
Pigmented epithelium adjacent to photoreceptors. Absorbs photons not absorbed by visual
pigments. Phagocytizes old discs shed from tip of outer segment
Outer segment:
Flattened, membranous discs containing visual pigment
Cones: plasma membrane infoldings; outer segment tapers to blunt point
Rods: discs are separate structures; outer segment forms elongated cylinder

Inner segment
Contains major organelles—responsible for all cell functions other than photoreception
Each photoreceptor synapses with a bipolar cell

Color vision: Rods all contain same opsin; responds to wavelengths of light (normal = 400-700
nm)
Three types of cones : Blue cones (16% of cones), Green cones (10%), Red cones (74%)
Three cone types have different opsins; sensitive to different wavelength ranges, with overlap
If all three types are stimulated equally, we see white
Color blindness—nonfunctional cones. Inability to distinguish certain colors
1+ types of cones are nonfunctional. Absent or do not make necessary visual pigment. Most
common type is red–green colorblindness, Cannot distinguish between red and green. Often
inherited. 10 percent of males, 0.67 percent of females and 1 in 300,000—no pigment

Photoreception process:
1. )Resting state (in the dark)Chemically gated sodium ion channels of outer segment stay open
if cGMP present. Inner segment continuously pumps sodium ions out of cytosol. This movement
of ions = dark current. Resting membrane potential = –40 mV. Photoreceptor continually
releases neurotransmitters to bipolar cells
2.) Retinal molecule in rhodopsin changes shape (activation) from a bent 11-cis form to more
linear 11-trans form.
3.) Opsin activates transducin, a G protein bound to disc membrane. Transducin activates
phosphodiesterase (PDE). 4.) Phosphodiesterase breaks down cGMP, inactivating gated sodium
channels. Sodium entry decreases
Active state: Decreased sodium entry reduces dark current. Membrane potential drops to –70
mV (hyperpolarized). Reduces neurotransmitter release.
Rhodopsin cannot respond to another photon until original shape of retinal is regained
Three step process
1.) Bleaching
Entire rhodopsin molecule first broken into retinal and opsin
2.) Retinal converted back to cis shape—requires ATP
3.) Opsin and retinal are reassembled as rhodopsin

Light reception to perception

Visual pathways: Photoreceptors to ganglion cells. About 1 million ganglion cell axons converge
at optic disc. Continue to diencephalon as the optic nerve (II). Two optic nerves (right & left)
reach diencephalon at the optic chiasm. Some wires cross and some wires continue. Fibers
continuealong optic tracts. Go to lateral geniculate nucleus. Projection fibers connectto Visual
cortex. Diencephalon. Brainstem
Optic radiation is the bundle of projection fibers linking each lateral geniculate body with
visual cortex,in occipital cortex on same side. Collaterals from fibers synapsing in lateral
geniculate bodies go to subconscious processing centers in diencephalon and brainstem
Pupillary reflexes : Triggered by collaterals going to superior colliculiin thalamus
Depth perception
The ability to judge depth or distance by interpreting 3-D relationships
Each eye collects visual information: Color, Brightness and Position
Position (per eye) is only direction
Info sent to visual cortex and integrated to provide a general image (shapes, colors), changes in
the image (movement) and differences between overlapping right and left visual field (depth)
Visual field
The spectrum of your vision. Can be altered by damage to ganglion cell axons or projection
fibers. Each location alters visual field in specific ways. Described loss can dictate where
damage is located
Image formation
Your eyes see upside-down and reversed.. Images are not single point, but rather consist of large
numbers of individual points each focused on retina. Image is inverted and reversed; brain
compensates—learned from experience
Visual sensation is upside down and backwards
The lens causes all images to flip
Up is down, down is up
Right is left, left is right
Your brain “knows” this is wrong based on other input
Corrects it prior to perception of vision

Auditory
Equilibrium and hearing involve the internal ear
Olfactory receptors—specialized sensory neurons
Gustatory receptors—communicate with sensory neurons
Photoreceptors—respond to light
Both route information directly to the CNS. All located in epithelia exposed to external
environment

Internal ear sensory receptors

All other special senses have receptors exposed to the external environment
Equilibrium and hearing receptors
Isolated and protected from external environment
Located in internal ear
Information integrated and organized locally before CNS
Hair cells = sensory receptors in internal ear
Specialized non-motile processes cover free surface
Stereocilia = resemble microvilli; 80–100 per hair cell
Kinocilium = single large cilium

External ear collects/directs sound waves toward middle ear

Auricle elastic cartilage
External acoustic meatus is the passageway in temporal bone
Ceruminous glands secrete waxy cerumen (earwax); keeps foreign objects out; slows growth
of microorganisms. Hairs trap debris.

Middle ear (tympanic cavity) Air-filled chamber from tympanic membrane to auditory ossicles
Connects to pharynx by auditory tube
Tympanic membrane (eardrum) = thin, semitransparent sheet that separates external ear and
middle ear
Auditory ossicles = three tiny bones; connect tympanic membrane and inner ear. Auditory
ossicles are the malleus, incas and stapes.
Auditory tube (pharyngotympanic or eustachian tube). Connects middle ear to nasopharynx
Allows pressure equalization across tympanic membrane. Can allow microorganisms into middle
ear, causing infection (otitis media)—can impair hearing, may invade internal ear
Malleus (hammer)— auditory ossicles that attaches to tympanic membrane
Incus (anvil)— auditory ossicles that attaches malleus to stapes
Stapes (stirrup)— auditory ossicles that attached to oval window, opening to internal ear
Middle ear Muscles: Tensor tympani muscle connects malleus to temporal bone . Contraction
stiffens tympanic membrane, reduces vibration. Innervated by mandibular branch of trigeminal
nerve (V)
Stapedius muscle
Connects stapes to posterior wall of middle ear. Reduces stapes movement at oval
window

Amplification and protection

Sound waves vibrate tympanic membrane; convert sound into mechanical movement
Auditory ossicles conduct vibrations to internal ear. Focuses sound on oval window and
amplifies it. Contractions of tensor tympani and stapedius muscles protect tympanic membrane
and ossicles from violent movement under very noisy conditions
Internal ear contains sensory organs for hearing and equilibrium. Cochlea (hearing).
Semicircular Canals (equilibrium)
Internal ear receives amplified sound waves from middle ear. Superficial contours established by
layer of dense bone = bony labyrinth
Sound propagation
The auricle catches sound waves
Directed into the auditory canal
Sound waves hit the tympanic membrane
Tympanic membrane vibrates
Sound is converted into mechanical movement
Auditory ossicles conduct vibrations to internal ear
Focuses sound on oval window and amplifies it
Bony labyrinth
Shell of dense bone surrounding membranous labyrinth
Filled with perilymph = liquid similar to CSF; between bony labyrinth and membranous
labyrinth
Three parts: Semicircular canals, Vestibule and Cochlea
Membranous Labyrinth
Collection of fluid-filled tubes/chambers
Houses receptors for hearing and equilibrium
Contains fluid called endolymph
Internal ear
 Membranous Labyrinth
 • Vestibular Complex
1. Semicircular ducts (within semicircular canals)
o Receptors stimulated by rotation of head
2. Utricle and saccule (within the vestibule)
o Provide sensations of gravity and linear acceleration
• Hearing
1. Cochlear duct (within cochlea)
o Sandwiched between pair of perilymph-filled chambers
o Resembles snail shell
o Receptors stimulated by sound

Vestibular Complex
Semicircular ducts (within semicircular canals)
Receptors stimulated by rotation of head
Utricle and saccule (within the vestibule)
Provide sensations of gravity and linear acceleration
Hearing
Cochlear duct (within cochlea)
Sandwiched between pair of perilymph-filled chambers
Resembles snail shell
Receptors stimulated by sound

Vestibular System Sense of Equilibrium

Semicircular ducts
Membranes found within semicircular canals (bone). Three ducts (anterior, posterior, lateral) .
Continuous with utricle. )Filled with endolymph (Vestibular System Sense of Equilibrium).
Ampulla = enlarged part of duct that houses receptors
Ampullary crest = region in wall of ampulla with receptors
Ampullary cupula = gelatinous structure extending through ampulla with kinocilia and
stereocilia of hair cells embedded in it
Detecting an equilibrium change in rotational movement
Head rotates in the plane of a duct, Endolymph moves, Hair cell stereocilia are pushed to one
side, distorting receptor processes.
One direction causes stimulation
Opposite direction causes inhibition
Ampulla rebounds when endolymph stops moving
Each duct detects a different plane of movement
Sagittal = Anterior
Frontal = Posterior
Transverse = Lateral
Utricle and saccule
Provide equilibrium sensations, lets you know if the body stationary or moving
Connected by slender passageway continuous with endolymphatic duct that ends in
endolymphatic sac
Sac projects into subarachnoid space
Endolymphatic duct continuously secretes endolymph; returns to general circulation at
endolymphatic sac
Utricle and saccule
Detects linear movement and position
Utricle is horizontal and detects horizontal movement
Saccule is vertical and detects vertical movement
Contain hair cells clustered in maculae
Hair cell processes embedded in gelatinous otolithic membrane
Surface has densely packed calcium carbonate crystals(otoliths, or “ear stones”)
As otoliths shift, a signal is generatedin the maculae detectinglinear movement
Utricle and saccule Detecting changes. Normal upright position. Otoliths in utricle sit on top. Tilt
head back
Otoliths in utricle shift. Movement distorts hair cells. Macular receptors stimulated
Signal is sent.

Cochlear SystemSense of Hearing

Cochlear duct (scala media)Filled with endolymph . Between 2 separate one-way perilymph
chambers
Scala vestibuli (vestibular duct) – moving away from vestibule
Scala tympani (tympanic duct) – moving towards vestibule
Encased by bony labyrinth except at oval/round windows
Interconnect at tip of cochlear, forming single long chamber from oval window to round
window
Vestibular membrane
Separates cochlear duct/scala vestibule
Basilar membrane
Separates cochlear ductfrom scala tympani
Hair cells for hearing located in cochlear duct in the spiral organ (organ of Corti) on basilar
membrane

Cross-sectional anatomy of cochlea:

Scala vestibuli and tympani filled with perilymph. Cochlear duct filled with endolymph.
Contains spiral organ (receptors for hearing). Spiral ganglion—cell bodies of sensory neurons
monitoring adjacent hair cells in spiral organ
Axons from spiral ganglion in cochlear nerve of vestibulocochlear nerve (VIII)

Spiral organ : Stereocilia are in contact with overlying tectorial membrane. Sound waves create
pressure waves in perilymph. Pressure waves vibrate inferior membrane up & down. Stereocilia
pressed into tectorial membrane- distortion. Distortion triggers nerve impulse. Sensory neurons
relay signal through spiral ganglion to cochlear branch of vestibulocochlear nerve (VIII)

Physiology of Hearing
Hearing is the perception of sound
Sound are waves of pressure
In air, pressure wave causes alternating areas of compressed/separated molecules
Wavelength of sound = distance between adjacent wave crests (peaks) or adjacent troughs
Frequency = number of waves (cycles) passing fixed point in given time
Measured as cycles per second (cps); units = hertz (Hz)
Wavelength and frequency inversely related
Pitch
is our perception of frequency. Location of vibration interpreted as pitch
High frequency (short wavelength) is a high pitch sound.
Low frequency (long wavelength) is a low pitch sound
Volume
Intensity is the amount of energy in sound waves. Amplitude of soundwave reflects amount of
energy
Greater energy equates to larger amplitude which equates to louder sound. Measured in decibels
(dB)
Sound waves and vibration. Energy of sound waves is physical pressure
Sound waves strike tympanic membrane. At particular frequency and amplitude, object will
vibrate at same frequency, which equals resonance
Tympanic membrane resonates generating movement of stapes at oval window
Stapes pushes on the oval window
Inward movement causes distortion of basilar membrane toward the round window
Opposite action when stapes moves outward
Cochlear duct inferior membrane regions resonate at different frequencies

Events involved in hearing

For hearing:
Flexibility of basilar membrane varies along its length. Different sound frequencies affect
different parts of the membrane. Location of vibration interpreted as pitch
Number of stimulated hair cells interpreted as volume
Nerve signals for hearing are carried on the cochlear nerve, which is part of the vestibulocochlear
nerve

Vestibulocochlear nerve function

Hearing
Most auditory info is projected to the auditory cortex on opposite side
Hemispheric lateralization
Some reaches auditory cortex on its same side
Aids in localizing sounds (left/right)
Reduces functional impact of damage to one cochlea or ascending pathway

disorders
Olfaction disorders
Head injury—damage to olfactory nerve (I)
Age-related changes:
Olfactory receptors are regularly replaced by stem cells, but number declines with age.
Remaining receptors become less sensitive

Gustation disorders
Problems with olfactory receptors
Decreased smell dulls taste. Damaged taste buds
Mouth infection, inflammation
Damaged CN VII, IX, or X
Trauma or compression
Natural age-related changes
Vision disorders
Senile cataract—lens loses transparency
Natural consequence of aging
Progresses—person needs more light to read; acuity may decline to blindness
Can be surgically corrected
Presbyopia
Age-related farsightednessdue to loss of lens elasticity
Less accommodation possible for close vision

Equilibrium disorders
Vertigo
False perception of spinning From conditions altering function of: Internal ear receptor complex
Vestibular nerve (of vestibulocochlear nerve VIII). Sensory nuclei and CNS pathways. Can be
due to vision problems or drug use (including alcohol)
Vertigo
Stimulated by anything that sets endolymph in motion
Motion sickness is most common cause
Symptoms—headache, sweating, flushing of face, nausea, vomiting

Hearing disorders
Partial hearing deficit affects ~37.5 million in United States
Two types: conductive and sensorineural
Conductive hearing loss—problem conducting sound waves
Causes include impacted earwax, infection, perforated tympanic membrane
Sensorineural hearing loss—damage to cochlea or nerve pathways from internal ear to brain
Causes include exposure to loud noise, head trauma, and aging
Age changes
Tympanic membrane loses flexibility
Articulations between auditory ossicles stiffen
Round window may start to ossify