The Special Senses

Chemical Senses

Chemical senses gustation (taste) and olfaction

(smell)
Their chemoreceptors respond to chemicals in
aqueous solution
Taste to substances dissolved in saliva
Smell to substances dissolved in fluids of the
nasal membranes

Taste Buds

Most of the 10,000 or so taste buds are found on the

tongue
Taste buds are found in papillae of the tongue
mucosa
Papillae come in three types: filiform, fungiform,
and circumvallate
Fungiform and circumvallate papillae contain taste
buds

Taste Buds

Figure 15.1

Anatomy of a Taste Bud

Each gourd-shaped taste bud consists of three major

cell types
Supporting cells insulate the receptor
Basal cells dynamic stem cells
Gustatory cells taste cells

Taste Sensations

There are five basic taste

sensations
Sweet sugars,
saccharin, alcohol, and
some amino acids
Salt metal ions
Sour hydrogen ions
Bitter alkaloids such as
quinine and nicotine
Umami elicited by the
amino acid glutamate

Physiology of Taste

In order to be tasted, a chemical:

Must be dissolved in saliva
Must contact gustatory hairs

Binding of the food chemical:

Depolarizes the taste cell membrane, releasing
neurotransmitter
Initiates a generator potential that elicits an action
potential

Taste Transduction

The stimulus energy of taste is converted into a

nerve impulse by:
Na+ influx in salty tastes
H+ in sour tastes (by directly entering the cell, by
opening cation channels, or by blockade of K+
channels)
Gustducin in sweet and bitter tastes

Gustatory Pathway

Cranial Nerves VII and IX carry impulses from

taste buds to the solitary nucleus of the medulla
These impulses then travel to the thalamus, and from
there fibers branch to the:
Gustatory cortex (taste)
Hypothalamus and limbic system (appreciation of
taste)

Gustatory Pathway

Influence of Other Sensations on Taste

Taste is 80% smell

Thermoreceptors, mechanoreceptors, nociceptors
also influence tastes
Temperature and texture enhance or detract from
taste

Sense of Smell

The organ of smell is the olfactory epithelium,

which covers the superior nasal concha
Olfactory receptor cells are bipolar neurons with
radiating olfactory cilia
Olfactory receptors are surrounded and cushioned by
supporting cells
Basal cells lie at the base of the epithelium

Sense of Smell

Physiology of Smell

Olfactory receptors respond to several different

odor-causing chemicals
When bound to ligand these proteins initiate a
G protein mechanism, which uses cAMP as a second
messenger
cAMP opens Na+ and Ca2+ channels, causing
depolarization of the receptor membrane that then
triggers an action potential

Olfactory receptor cells

Processing of scents in the olfactory bulb

Each of the glomeruli lining
the olfactory bulb receives
synaptic input from only one
type of olfactory receptor
responds to only one discrete
component of an odorant.
The glomeruli sort and file
the various components of an
odoriferous molecule before
relaying the smell signal to
the mitral cells and higher
brain levels for further
processing

Signal transduction in an odorant receptor

Olfactory receptors are G protein
coupled receptors that dissociate on
binding to the odorant.
The -subunit of G proteins
activates adenylate cyclase to
catalyze production of cAMP. cAMP
acts as a second messenger to open
cation channels.
Inward
diffusion of Na+ and Ca2+ produces
depolarization

Olfactory Transduction Process

Odorant
binding
protein

Inactive

Adenylate
cyclase

Odorant
chemical

Na+

Active
Na+ influx
causes
depolarization

ATP

cAMP

Cytoplasm

Depolarization of
olfactory receptor
cell membrane
triggers action
potentials in axon
of receptor

Olfactory pathway

Information is transmitted from the olfactory bulb by axons of

mitral and tufted relay neurons in the lateral olfactory tract.
Mitral cells project to five regions of the olfactory cortex:
anterior olfactory nucleus, olfactory tubercle, piriform cortex,
and parts of the amygdala and entorhinal cortex.
Tufted cells project to anterior olfactory nucleus and olfactory
tubercle; mitral cells in the accessory olfactory bulb project
only to the amygdala.
Conscious discrimination of odor depends on the neocortex
(orbitofrontal and frontal cortices).
Emotive aspects of olfaction derive from limbic projections
(amygdala and hypothalamus).

Olfactory pathway

Olfactory Pathway

Olfactory receptor cells synapse with mitral cells

Glomerular mitral cells process odor signals
Mitral cells send impulses to:
The olfactory cortex
The hypothalamus, amygdala, and limbic system

Eye and Associated Structures

70% of all sensory receptors are in the eye

Most of the eye is protected by a cushion of fat and
the bony orbit
Accessory structures include eyebrows, eyelids,
conjunctiva, lacrimal apparatus, and extrinsic eye
muscles

Eyebrows

Coarse hairs that overlie the supraorbital margins

Functions include:
Shading the eye
Preventing perspiration from reaching the eye

Orbicularis muscle depresses the eyebrows

Corrugator muscles move the eyebrows medially

Palpebrae (Eyelids)

Protect the eye anteriorly

Palpebral fissure separates eyelids
Canthi medial and lateral angles (commissures)

Palpebrae (Eyelids)

Lacrimal caruncle contains glands that secrete a

whitish, oily secretion (Sandmans eye sand)
Tarsal plates of connective tissue support the eyelids
internally
Levator palpebrae superioris gives the upper
eyelid mobility

Palpebrae (Eyelids)

Eyelashes
Project from the free margin of each eyelid
Initiate reflex blinking

Lubricating glands associated with the eyelids

Meibomian glands and sebaceous glands
Ciliary glands lie between the hair follicles

Palpebrae (Eyelids)

Conjunctiva

Transparent membrane that:

Lines the eyelids as the palpebral conjunctiva
Covers the whites of the eyes as the ocular
conjunctiva
Lubricates and protects the eye

Lacrimal Apparatus

Consists of the lacrimal gland and associated ducts

Lacrimal glands secrete tears
Tears
Contain mucus, antibodies, and lysozyme
Enter the eye via superolateral excretory ducts
Exit the eye medially via the lacrimal punctum
Drain into the nasolacrimal duct

Lacrimal Apparatus

Extrinsic Eye Muscles

Six straplike extrinsic eye muscles

Enable the eye to follow moving objects
Maintain the shape of the eyeball

Four rectus muscles originate from the annular ring

Two oblique muscles move the eye in the vertical
plane

Extrinsic Eye Muscles

Figure 15.7a, b

Summary of Cranial Nerves and Muscle Actions

Names, actions, and cranial nerve innervation of the

extrinsic eye muscles

Figure 15.7c

Structure of the Eyeball

A slightly irregular hollow sphere with anterior and

posterior poles
The wall is composed of three tunics fibrous,
vascular, and sensory
The internal cavity is filled with fluids called
humors
The lens separates the internal cavity into anterior
and posterior segments

Structure of the Eyeball

Figure 15.8a

Fibrous Tunic

Forms the outermost coat of the eye and is

composed of:
Opaque sclera (posteriorly)
Clear cornea (anteriorly)

The sclera protects the eye and anchors extrinsic

muscles
The cornea lets light enter the eye

Vascular Tunic (Uvea): Choroid Region

Has three regions: choroid, ciliary body, and iris

Choroid region
A dark brown membrane that forms the posterior
portion of the uvea
Supplies blood to all eye tunics

Vascular Tunic: Ciliary Body

A thickened ring of tissue surrounding the lens

Composed of smooth muscle bundles (ciliary
muscles)
Anchors the suspensory ligament that holds the lens
in place

Vascular Tunic: Iris

The colored part of the eye

Pupil central opening of the iris
Regulates the amount of light entering the eye
during:
Close vision and bright light pupils constrict
Distant vision and dim light pupils dilate
Changes in emotional state pupils dilate when
the subject matter is appealing or requires
problem-solving skills

Pupil Dilation and Constriction

Figure 15.9

Controlled amount of light entering the eye

Bright light

Normal light

Dark

The amount of light entering the eye is controlled by the

iris.
Not all the light passing through the cornea reaches the
light sensitive photoreceptors, because of the presence of
the iris, a thin, pigmented smooth muscle that forms a
visible ringlike structure within the aqueous humor

The neural pathways of the pupillary reflexes

Sensory Tunic: Retina

A delicate two-layered membrane

Pigmented layer the outer layer that absorbs light
and prevents its scattering
Neural layer, which contains:
Photoreceptors that transduce light energy
Bipolar cells and ganglion cells
Amacrine and horizontal cells

Retinal layers

Retinal layers

The retinal visual pathway extends from the photoreceptor cells

(rods and cones, whose light-sensitive ends face the choroid away
from the incoming light) to the bipolar cells to the ganglion cells. The
horizontal and amacrine cells act locally for retinal processing
of visual input.

Retina layers

Sensory Tunic: Retina

Figure 15.10a

The Retina: Ganglion Cells and the Optic Disc

Ganglion cell axons:

Run along the inner surface of the retina
Leave the eye as the optic nerve

The optic disc:

Is the site where the optic nerve leaves the eye
Lacks photoreceptors (the blind spot)

Retina seen through the ophthalmoscope

(a)

(b)

(a) A photograph and (b) an illustration of the optic fundus (back

of the eye). Optic nerve fibers leave the eyeball at the optic disc to
form the optic nerve. The arteries, arterioles, and veins in the
superficial layers of the retina near its vitreous surface can be seen
through the ophthalmoscope.

The Retina: Ganglion Cells and the Optic Disc

Figure 15.10b

A.
B.
C.
D.

Fovea centralis
Macula lutea
Optic Nerve
Arteries

The fovea centralis

highest concentration of
cone photoreceptors high
level of cones
Color vision diameter
human fovea centralis is
about 0.04 inches (1 mm)
1% of the total surface
of retina
Contributes to about 50%
of the nerve opticfibers and
about 50% of the brain
volume in the visual cortex

The Retina: Photoreceptors

Rods:
Respond to dim light
Are used for peripheral vision

Cones:
Respond to bright light
Have high-acuity color vision
Are found in the macula lutea
Are concentrated in the fovea centralis

Blood Supply to the Retina

The neural retina receives its blood supply from two

sources
The outer third receives its blood from the choroid
The inner two-thirds is served by the central artery
and vein

Small vessels radiate out from the optic disc and can
be seen with an ophthalmoscope

Inner Chambers and Fluids

The lens separates the internal eye into anterior and

posterior segments
The posterior segment is filled with a clear gel
called vitreous humor that:
Transmits light
Supports the posterior surface of the lens
Holds the neural retina firmly against the
pigmented layer
Contributes to intraocular pressure

Anterior Segment

Composed of two chambers

Anterior between the cornea and the iris
Posterior between the iris and the lens

Aqueous humor
A plasmalike fluid that fills the anterior segment
Drains via the canal of Schlemm

Supports, nourishes, and removes wastes

Anterior Segment

Figure 15.12

Formation and drainage of aqueous humor

Aqueous humor is
formed by a
capillary network in
the ciliary body
Drains into the canal
of Schlemm, and
eventually enters the
blood

Lens

A biconvex, transparent, flexible, avascular structure

that:
Allows precise focusing of light onto the retina
Is composed of epithelium and lens fibers

Lens epithelium anterior cells that differentiate

into lens fibers
Lens fibers cells filled with the transparent protein
crystallin
With age, the lens becomes more compact and dense
and loses its elasticity

Mechanism of accommodation

(a) Suspensory ligaments extend from the ciliary muscle to the

outer edge of the lens

Mechanism of accommodation

(b) When the ciliary muscle is relaxed, the suspensory

ligaments are taut, putting tension on the lens so that it is flat and
weak.
(c) When the ciliary muscle is contracted, the suspensory ligaments
become slack, reducing the tension on the lens, allowing it to assume
a stronger, rounder shape because of its elasticity.

Light

Electromagnetic radiation all energy waves from

short gamma rays to long radio waves
Our eyes respond to a small portion of this spectrum
called the visible spectrum
Different cones in the retina respond to different
wavelengths of the visible spectrum

Light

Figure 15.14

Refraction and Lenses

When light passes from one transparent medium to

another its speed changes and it refracts (bends)
Light passing through a convex lens (as in the eye)
is bent so that the rays converge to a focal point
When a convex lens forms an image, the image is
upside down and reversed right to left

Refraction and Lenses

Figure 15.16

Focusing Light on the Retina

Pathway of light entering the eye: cornea, aqueous

humor, lens, vitreous humor, and the neural layer of
the retina to the photoreceptors
Light is refracted:
At the cornea
Entering the lens
Leaving the lens

The lens curvature and shape allow for fine focusing

of an image

Focusing for Distant Vision

Light from a
distance needs
little adjustment
for proper
focusing
Far point of
vision the
distance beyond
which the lens
does not need to
change shape to
focus (20 ft.)
Figure 15.17a

Focusing for Close Vision

Close vision requires:

Accommodation changing the lens shape by
ciliary muscles to increase refractory power
Constriction the pupillary reflex constricts the
pupils to prevent divergent light rays from entering
the eye
Convergence medial rotation of the eyeballs
toward the object being viewed

Focusing for Close Vision

Figure 15.7b

Problems of Refraction

Emmetropic eye normal eye with light focused

properly
Myopic eye (nearsighted) the focal point is in front
of the retina
Corrected with a concave lens

Hyperopic eye (farsighted) the focal point is

behind the retina
Corrected with a convex lens

Problems of Refraction

Figure 15.18

Lens refrractions

The refractive power of a lens is measured in diopters.

The dioptric power (D) of a lens is the reciprocal of the
focal length measured in meters:

Lens
A converging lens with a focal length of 1 m has a
power of 1 diopter. A lens with a focal length of 10
cm will have a power of 10 D and one with a focal
length of 17 mm (the approximate focal length of the
lens system of the human eye) has a dioptric power
of:

For diverging lenses, the dioptric power is negative,

so that a diverging lens with a focal length of 10 cm
has a dioptric power of-10 D.

Case 1
A patient with hypermetropia has an eye with a focal length
of 59 D but the retina is only 16 mm behind the lens instead
of the usual 17 mm. What power of spectacle lens is
required for correction?
To bring parallel light into focus in 16 mm requires a power
of approximately 62.5D. Thus, in this case, a converging
lens of 62.5 - 58.8 = 3.7 D is needed. (i.e. a converging lens
of 3.7 D).

Case 2
A patient with myopia has an eye with a focal length of
59 D, but the retina is 18 mm behind the lens instead of
the usual 17 mm. What power of spectacle lens is
required for correction?
The lens system would need to be 55.5 D to bring
parallel light into focus on the retina. Thus a lens
of 55.5 - 58.8 = -3.3 D would be needed for
correction (i.e. a diverging lens of 3.3 D).

Photoreception:
Functional Anatomy of Photoreceptors

Photoreception process by which the eye detects

light energy
Rods and cones contain visual pigments
(photopigments)
Arranged in a stack of disklike infoldings of the
plasma membrane that change shape as they absorb
light

Photoreception:
Functional Anatomy of Photoreceptors

Figure 15.19

Rods

Functional characteristics
Sensitive to dim light and best suited for night
vision
Absorb all wavelengths of visible light
Perceived input is in gray tones only
Sum of visual input from many rods feeds into a
single ganglion cell
Results in fuzzy and indistinct images

Cones

Functional characteristics
Need bright light for activation (have low
sensitivity)
Have pigments that furnish a vividly colored view
Each cone synapses with a single ganglion cell
Vision is detailed and has high resolution

Cones and Rods

Figure 15.10a

Properties of Rod Vision and Cone Vision

Chemistry of Visual Pigments

Retinene or retinal is a light-absorbing molecule

Combines with opsins to form visual pigments
Similar to and is synthesized from vitamin A
Two isomers: 11-cis and all-trans

Isomerization of retinal initiates electrical impulses

in the optic nerve

Structure of rhodopsin

Position of
retinene1 (R) in the
rod disk membrane

Chemistry of Visual Pigments

Figure 15.20

Ionic Basis of Photoreceptor Potentials

Na+ channels in the outer segments of the rods and cones
are open in the dark current flows from the inner to the
outer segment and synaptic ending of the photoreceptor.
The Na+K+ pump in the inner segment maintains ionic
equilibrium.
Release of synaptic transmitter is steady in the dark.
When light strikes the outer segment initiate close the
Na+ channels hyperpolarizing receptor potential.
The hyperpolarization reduces the release of synaptic
transmitter generates a signal in the bipolar cells
leads to action potentials in ganglion cells The action
potentials are transmitted to the brain.

Effect of light on current flow in visual receptors

In the dark, Na+
channels in the outer
segment are held open
by cGMP.
Light leads to
increased conversion
of cGMP to 5'-GMP,
and some of the
channels close
hyperpolarization of
the synaptic terminal
of the photoreceptor.

Excitation of Rods

The visual pigment of rods is rhodopsin

(opsin + 11-cis retinal)
Light phase
Rhodopsin breaks down into all-trans retinal +
opsin (bleaching of the pigment)

Dark phase
All-trans retinal converts to 11-cis form
11-cis retinal is also formed from vitamin A
11-cis retinal + opsin regenerate rhodopsin

Excitation of rod

In the dark, rhodopsin retinene1 is in the 11-cis

configuration.
Action of light change the shape of the retinene,
converting it to the all-trans isomer alters the
configuration of the opsin activates the associated
heterotrimeric G protein (transducin or Gt1).
G protein exchanges GDP for GTP, and the subunit
separates. This subunit remains active until its
intrinsic GTPase activity hydrolyzes the GTP.
Termination of the activity of transducin is also
accelerated by its binding of -arrestin.

Excitation of Rods

Figure 15.21

Excitation of Cones

Visual pigments in cones are similar to rods

(retinal + opsins)
There are three types of cones: blue, green, and red
Intermediate colors are perceived by activation of
more than one type of cone
Method of excitation is similar to rods

Phototransduction

Light energy splits rhodopsin into all-trans retinal,

releasing activated opsin
The freed opsin activates the G protein transducin
Transducin catalyzes activation of phosphodiesterase
(PDE)
PDE hydrolyzes cGMP to GMP and releases it from
sodium channels
Without bound cGMP, sodium channels close, the
membrane hyperpolarizes, and neurotransmitter
cannot be released

Phototransduction

Figure 15.22

Sequence phototransduction in rods and cones

Light activates retinen1 activates
Gt2
Gt2 in turn activates
phosphodiesterase, catalyzing the
conversion of cGMP to 5'-GMP.
This results in closure of Na+
channels between the extracellular
fluid and the cone cytoplasm:
decrease in intracellular Na+
concentration
hyperpolarization of the cone
synaptic terminals.

Adaptation

Adaptation to bright light (going from dark to light)

involves:
Dramatic decreases in retinal sensitivity rod
function is lost
Switching from the rod to the cone system visual
acuity is gained

Adaptation to dark is the reverse

Cones stop functioning in low light
Rhodopsin accumulates in the dark and retinal
sensitivity is restored

Visual Pathways

Axons of retinal ganglion cells form the optic nerve

Medial fibers of the optic nerve decussate at the
optic chiasm
Most fibers of the optic tracts continue to the lateral
geniculate body of the thalamus
Other optic tract fibers end in superior colliculi
(initiating visual reflexes) and pretectal nuclei
(involved with pupillary reflexes)
Optic radiations travel from the thalamus to the
visual cortex

Pathways to the Cortex

Visual Pathways

Figure 15.23

Visual Pathways

Some nerve fibers send tracts to the midbrain ending

in the superior colliculi
A small subset of visual fibers contain melanopsin
(circadian pigment) which:
Mediates papillary light reflexes
Sets daily biorhythms

Functions of Visual Projection Areas in the Brain

V1
V2, V3, VP
V3A
V4v
MT/V5
LO
V7
V8

Primary visual
cortex
Continued
processing,
larger visual fields
Motion
Unknown
Motion; control of
movement
Recognition of
large objects
Unknown
Color vision

Transection of visual pathways

A lesion that interrupts one optic nerve causes blindness in
that eye (A).
Lesions affecting the optic chiasm destroy fibers from both
nasal hemiretinas and produce a heteronymous (opposite
sides of the visual fields) hemianopia (B).
A lesion in one optic tract causes blindness in half of the
visual field (C) and is called homonymous (same side of
both visual fields) hemianopia (half-blindness).
Occipital lesions may spare the fibers from the macula (as in
D) because of the separation in the brain of these fibers from
the others subserving vision

Transection of visual pathways

Blindness of the eye (A).

Heteronymous
hemianopia (B)
Homonymous hemianopia
(C)
Occipital lession(D)

Depth Perception

Achieved by both eyes viewing the same image

from slightly different angles
Three-dimensional vision results from cortical
fusion of the slightly different images
If only one eye is used, depth perception is lost and
the observer must rely on learned clues to determine
depth

Depth Perception
Left: two eyes viewing an
arrow lying in the frontal
plane (no stereopsis)
Right: the arrow is inclined
into the third dimensionit
tends to point toward the
observer.
Noncorresponding or
disparate, points on the retinas
can be projected to a single
point, and it is essentially this
fusion of disparate images by
the brain that creates the
impression of depth.

Represents the same object being viewed from

different positions causing the image produced
within each eye to be different

Retinal Processing: Receptive Fields of

Ganglion Cells

On-center fields
Stimulated by light hitting the center of the field
Inhibited by light hitting the periphery of the field

Off-center fields have the opposite effects

These responses are due to receptor types in the
on and off fields

Retinal Processing: Receptive Fields of

Ganglion Cells

Figure 15.24

Thalamic Processing

The lateral geniculate nuclei of the thalamus:

Relay information on movement
Segregate the retinal axons in preparation for depth
perception
Emphasize visual inputs from regions of high cone
density
Sharpen the contrast information received by the
retina

Cortical Processing

Striate cortex processes

Basic dark/bright and contrast information

Prestriate cortices (association areas) processes

Form, color, and movement

Visual information then proceeds anteriorly to the:

Temporal lobe processes identification of objects
Parietal cortex and postcentral gyrus processes
spatial location

Color Vision
Young & Helmholtz Trichromatic theory of color
vision:
There is only one type of rod and this responds strongly
to bluish-green light
Cones are divided into three categories, each of which
has a different sensitivity to light
There are red light receptors, green light receptors and
blue light receptors.
All colors of the visible spectrum can be seen by mixing
the 3 primary colours (red, blue and green)
White objects reflect all colors to eye, black absorbs all
colours so no light to the eye.

Color Vision

Young & Helmholtz Trichromatic theory of color

vision:
There is only one type of rod and this responds strongly
to bluish-green light
Cones are divided into three categories, each of which
has a different sensitivity to light
"red" cones (64%)
"green" cones (32%)
"blue" cones (2%)

Trichromatic Theory

The photocells in our retina, called cones are

responsible for our colour vision. There are 3 types
of cones, each with a different iodopsin (a
photosensitive pigment).
Each type of iodopsin can absorb and respond to a
range of wavelengths:
erythrolabe: max. absorption at 565nm (red)
chlorolabe : max. absorption at 535nm (green)
cyanolabe : max. absorption at 440nm (blue)

Wavelengths of light absorbed by different

cones

Color vision
Color vision is the capacity of an organism or machine to
distinguish objects based on the wavelengths of the light
they reflect, emit, or transmit.
Human's perception of
colors is a subjective
process
Brain responds to the
stimuli that are produced
when incoming light reacts
with the several types of
cone photoreceptors

Color vision
The colour that our brain "sees" comes from the
integration of impulses from the three types of cones.
The resultant color
depends on the
differential
stimulation of
each, i.e. how
many of each kind
are stimulated.

Color mixing

Primary additive or substractive

Additive mixing is most intuitive
Add wavelengths:
red+green = yellow
red+blue = magenta
blue+green = cyan
red+green+blue=white
Subtractive mixing is much less intuitive (but much
more common), subtractive mixing happens when
pigments are mix together
Different pigments subtract different wavelengths

Additive primaries

Media that combine

emitted lights to create
the sensation of a range
of colors are using the
additive color system.
Typically, the primary
colors used are red,
green, and blue.
When all pigments in
conus are stimulated
together, it is produced
white color

Subtractive primaries

Media that use reflected

light and colorants to
produce colors are using
the subtractive color
method of color mixing.
Subtraction produce
black color
Colorant is something added to something
else to cause a change in color

Color blindness
Defect of vision affecting the ability to distinguish
colors, caused by a defect in the retina or in other nerve
portions of the eye.
Cause of color blindness:
Genetic
Aging
Eye problems, such as glaucoma, macular degeneration,
cataracts, or diabetic retinopathy
Injury to the eye
Side effects of some medicines, overexposure to lead or
mercury
Lesions of area V8 of the visual cortex

Distribution of cone cells in the fovea of a individual with

normal color vision (left), and a color blind (protanopic)
retina. Note that the center of the fovea holds very few bluesensitive cones.

Genetic of CB

Non CB male: XY
CB male: xY
Non CB female: XX
CB female : xx
CB carrier female: xX

Type of color blindness

Monochromatism: Either no cones available or just one

type of them.
Dichromatism: Only two different cone types, the third
one is missing completely.
Anomalous trichromatism: All three types but with
shifted peaks of sensitivity for one of them. This results
in a smaller color spectrum.

Type of color blindness

Dichromats and anomalous trichromats exist

again in three different types according to the
missing cone or in the latter case of its
malfunctioning.
Tritanopia/Tritanomaly: Missing/malfunctioning Scone (blue).
Deuteranopia/Deuteranomaly:
Missing/malfunctioning M-cone (green).
Protanopia/Protanomaly: Missing/malfunctioning Lcone (red).

Different forms of color vision deficiency

Type

Denomination

Monochromacy

Achromatopsia

Dichromacy

Men

Women

0.00003%

Protanopia

1.01%

0.02%

Deuteranopia

1.27%

0.01%

Tritanopia
Anomalous
Trichromacy

Prevalence

0.0001%

Protanomaly

1.08%

0.03%

Deuteranomaly

4.63%

0.36%

Tritanomaly

0.0002%

Normal

Monochromatism/
Achromatopsia

Normal

Complete green CB
deutranopia

Complete red CB
Protanopia

Complete Blue / yellow

CB =Tritanopia

Color blind diagnoses

Holmgren threads
Pseudo Isochromatic Plate Ishihara and American
Optical HRR
Farnsworth test
Anomaloscope
Medmont C-100 color vision tester

Tes Ishihara
Found by Dr. Shinobu
Ishihara from Tokyo
University in 1917
Most common and easy to
used
Monochromatic dots of
basic color that create
number pattern

Ishihara test from color blind person

American Optical HRR (Hardy-Rand-Rittler)

Pseudoisochromatic Plates
1

Color blindness is not disease

Same condition as hair and skin, just lack
of certain pigment in the eye cone.
Unique characteristic of an individu

The Ear: Hearing and Balance

The three parts of the ear are the inner, outer, and
middle ear
The outer and middle ear are involved with hearing
The inner ear functions in both hearing and
equilibrium
Receptors for hearing and balance:
Respond to separate stimuli
Are activated independently

The Ear: Hearing and Balance

Figure 15.25a

Outer Ear

The auricle (pinna) is composed of:

The helix (rim)
The lobule (earlobe)

External auditory canal

Short, curved tube filled with ceruminous glands

Outer Ear

Tympanic membrane (eardrum)

Thin connective tissue membrane that vibrates in
response to sound
Transfers sound energy to the middle ear ossicles
Boundary between outer and middle ears

Middle Ear (Tympanic Cavity)

A small, air-filled, mucosa-lined cavity

Flanked laterally by the eardrum
Flanked medially by the oval and round windows

Epitympanic recess superior portion of the middle

ear
Pharyngotympanic tube connects the middle ear to
the nasopharynx
Equalizes pressure in the middle ear cavity with the
external air pressure

Middle Ear (Tympanic Cavity)

Figure 15.25b

Ear Ossicles

The tympanic cavity contains three small bones: the

malleus, incus, and stapes
Transmit vibratory motion of the eardrum to the
oval window
Dampened by the tensor tympani and stapedius
muscles

Ear Ossicles

Figure 15.26

Inner Ear
Bony labyrinth
Tortuous channels worming their way through the temporal
bone
Contains the vestibule, the cochlea, and the semicircular
canals
Filled with perilymph

Membranous labyrinth
Series of membranous sacs within the bony labyrinth
Filled with a potassium-rich fluid

Inner Ear

Figure 15.27

The Vestibule

The central egg-shaped cavity of the bony labyrinth

Suspended in its perilymph are two sacs: the saccule
and utricle
The saccule extends into the cochlea
The utricle extends into the semicircular canals
These sacs:
House equilibrium receptors called maculae
Respond to gravity and changes in the position of
the head

The Vestibule

Figure 15.27

The Semicircular Canals

Three canals that each define two-thirds of a circle

and lie in the three planes of space
Membranous semicircular ducts line each canal and
communicate with the utricle
The ampulla is the swollen end of each canal and it
houses equilibrium receptors in a region called the
crista ampullaris
These receptors respond to angular movements of
the head

The Semicircular Canals

Figure 15.27

The Cochlea

A spiral, conical, bony chamber that:

Extends from the anterior vestibule
Coils around a bony pillar called the modiolus
Contains the cochlear duct, which ends at the
cochlear apex
Contains the organ of Corti (hearing receptor)

The Cochlea

The cochlea is divided into three chambers:

Scala vestibuli
Scala media
Scala tympani

The Cochlea

The scala tympani terminates at the round window

The scalas tympani and vestibuli:
Are filled with perilymph
Are continuous with each other via the helicotrema

The scala media is filled with endolymph

Fluid movement in cochlea

Fluid movement in cochlea

Fluid movement within the perilymph set up by
vibration of the oval window follows two pathways:
Through the scala vestibuli, around the heliocotrema,
and through the scala tympani, causing the round
window to vibrate. This pathway just dissipates sound
energy
A shortcut from the scala vestibuli through the basilar
membrane to the scala tympani. This pathway triggers
activation of the receptors for sound by bending the
hairs of hair cells as the organ of Corti on top of the
vibrating basilar membrane is displaced in relation to
the overlying tectorial membrane

Basilar membrane (partly uncoiled)

Different regions of the basilar membrane vibrate maximally

at different frequencies.

Basilar membrane (completely uncoiled)

The narrow, stiff end of the basilar membrane nearest the oval
window vibrates best with high-frequency pitches.
The wide, flexible end of the basilar membrane near the
helicotrema vibrates best with low-frequency pitches

The Cochlea

The floor of the cochlear duct is composed of:

The bony spiral lamina
The basilar membrane, which supports the organ of
Corti

The cochlear branch of nerve VIII runs from the

organ of Corti to the brain

The Cochlea

Figure 15.28

Sound and Mechanisms of Hearing

Sound vibrations beat against the eardrum

The eardrum pushes against the ossicles, which
presses fluid in the inner ear against the oval and
round windows
This movement sets up shearing forces that pull on
hair cells
Moving hair cells stimulates the cochlear nerve
that sends impulses to the brain

Properties of Sound

Sound is:
A pressure disturbance (alternating areas of high
and low pressure) originating from a vibrating
object
Composed of areas of rarefaction and compression
Represented by a sine wave in wavelength,
frequency, and amplitude

Properties of Sound

Frequency the number of waves that pass a given

point in a given time
Pitch perception of different frequencies (we hear
from 2020,000 Hz)

Properties of Sound

Amplitude intensity of a sound measured in

decibels (dB)
Loudness subjective interpretation of sound
intensity

Figure 15.29

Transmission of Sound to the Inner Ear

The route of sound to the inner ear follows this

pathway:
Outer ear pinna, auditory canal, eardrum
Middle ear malleus, incus, and stapes to the oval
window
Inner ear scalas vestibuli and tympani to the
cochlear duct
Stimulation of the organ of Corti
Generation of impulses in the cochlear nerve

Transmission of Sound to the Inner Ear

Figure 15.31

Resonance of the Basilar Membrane

Sound waves of low frequency (inaudible):

Travel around the helicotrema
Do not excite hair cells

Audible sound waves:

Penetrate through the cochlear duct
Vibrate the basilar membrane
Excite specific hair cells according to frequency of
the sound

Resonance of the Basilar Membrane

Figure 15.32

The Organ of Corti

Is composed of supporting cells and outer and inner

hair cells
Afferent fibers of the cochlear nerve attach to the
base of hair cells
The stereocilia (hairs):
Protrude into the endolymph
Touch the tectorial membrane

Excitation of Hair Cells in the Organ of Corti

Bending cilia:
Opens mechanically gated ion channels
Causes a graded potential and the release of a
neurotransmitter (probably glutamate)

The neurotransmitter causes cochlear fibers to

transmit impulses to the brain, where sound is
perceived

Excitation of Hair Cells in the Organ of Corti

Figure 15.28c

Deflection of the basilar membrane

The stereocilia (hairs) from the hair cells of the basilar

membrane contact the overlying tectorial membrane. These
hairs are bent when the basilar membrane is deflected in
relation to the stationary tectorial membrane.
This bending of the inner hair cells hairs opens mechanically
gated channels, leading to ion movements that result in a
receptor potential.

The role of stereocilia in sound transduction

The role of stereocilia in sound transduction

The role of tip links in the responses of hair cells

When a stereocilium is pushed toward a taller stereocilium, the tip

link is stretched and opens an ion channel in its taller neighbor.
The channel next is moved down the taller stereocilium by a
molecular motor, so the tension on the tip link is released.
When hairs return to their resting position, the motor moves back
up the stereocilium.

Ionic composition (mmol/L) of perilymph

Auditory and vestibular pathways

Pathway for sound

transduction

Auditory Pathway to the Brain

Impulses from the cochlea pass via the spiral

ganglion to the cochlear nuclei
From there, impulses are sent to the:
Superior olivary nucleus
Inferior colliculus (auditory reflex center)

From there, impulses pass to the auditory cortex

Auditory pathways decussate so that both cortices
receive input from both ears

Simplified Auditory Pathways

Figure 15.34

Auditory Processing

Pitch is perceived by:

The primary auditory cortex
Cochlear nuclei

Loudness is perceived by:

Varying thresholds of cochlear cells
The number of cells stimulated

Localization is perceived by superior olivary nuclei

that determine sound

Deafness
Conduction deafness something hampers sound
conduction to the fluids of the inner ear (e.g., impacted
earwax, perforated eardrum, osteosclerosis of the ossicles)
Sensorineural deafness results from damage to the neural
structures at any point from the cochlear hair cells to the
auditory cortical cells
Tinnitus ringing or clicking sound in the ears in the
absence of auditory stimuli
Menieres syndrome labyrinth disorder that affects the
cochlea and the semicircular canals, causing vertigo, nausea,
and vomiting

Mechanisms of Equilibrium and Orientation

Vestibular apparatus equilibrium receptors in the

semicircular canals and vestibule
Maintains our orientation and balance in space
Vestibular receptors monitor static equilibrium
Semicircular canal receptors monitor dynamic
equilibrium

Anatomy of Maculae

Maculae are the sensory receptors for static

equilibrium
Contain supporting cells and hair cells
Each hair cell has stereocilia and kinocilium
embedded in the otolithic membrane

Otolithic membrane jellylike mass studded with

tiny CaCO3 stones called otoliths
Utricular hairs respond to horizontal movement
Saccular hairs respond to vertical movement

Anatomy of Maculae

Figure 15.35

Effect of Gravity on Utricular Receptor Cells

Otolithic movement in the direction of the kinocilia:

Depolarizes vestibular nerve fibers
Increases the number of action potentials generated

Movement in the opposite direction:

Hyperpolarizes vestibular nerve fibers
Reduces the rate of impulse propagation

From this information, the brain is informed of the

changing position of the head

Effect of Gravity on Utricular Receptor Cells

Figure 15.36

Crista Ampullaris and Dynamic Equilibrium

The crista ampullaris (or crista):

Is the receptor for dynamic equilibrium
Is located in the ampulla of each semicircular canal
Responds to angular movements

Each crista has support cells and hair cells that

extend into a gel-like mass called the cupula
Dendrites of vestibular nerve fibers encircle the base
of the hair cells

Crista Ampullaris and Dynamic Equilibrium

Figure 15.37b

Activating Crista Ampullaris Receptors

Cristae respond to changes in velocity of rotatory

movements of the head
Directional bending of hair cells in the cristae
causes:
Depolarizations, and rapid impulses reach the brain
at a faster rate
Hyperpolarizations, and fewer impulses reach the
brain

The result is that the brain is informed of rotational

movements of the head

Rotary Head Movement

Figure 15.37d

Role of the otolith organs

otoliths

The hairs (kinocilium and

stereocilia) of the receptor
stereocilia
hair cells protrude into an
kinocilium
overlying gelatinous sheet,
whose movement displaces
the hairs changes in hair
cell potential.
The otoliths (ear stones):
crystals of calcium carbonate
suspended within the
gelatinous layer, making it
heavier giving more
inertia than the surrounding
fluid

Role of the otolith organs tilt the head

Tilt the head in any direction other than vertical (other than
straight up and down) the hairs are bent in the direction of
the tilt because of the gravitational force exerted on the topheavy gelatinous layer produces depolarizing or
hyperpolarizing receptor potentials depending on the tilt of the
head.
The CNS thus receives different patterns of neural activity
depending on head position with respect to gravity.

Role of the otolith organs - walking

As walk forward is started , the top-heavy
otolith membrane at first lags behind the
endolymph and hair cells because of its
greater inertia. The hairs are thus bent to the
rear, in the opposite direction of the forward
movement of the head.
When walking pace is maintain, the
gelatinous layer soon catches up and moves at
the same rate as your head so that the hairs
are no longer bent.
When walking is stopped, the otolith sheet
continues to move forward briefly as the
head slows and stops, bending the hairs
toward the front.

Input and output of the vestibular nuclei

Balance and Orientation Pathways

There are three modes

of input for balance
and orientation
Vestibular receptors
Visual receptors
Somatic receptors

These receptors allow

our body to respond
reflexively
Figure 15.38