The Sense Organs

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Chapter 16
Sense Organs
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Making Sense of the Senses
• Properties and types of receptors
• General senses
• Chemical senses
• Hearing and equilibrium
• Vision
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Chemical Senses- Gustation
• Gustation – sensation that results from action of
tastants on taste buds
• Lingual papillae
– filiform - no taste buds
• important for food texture
– foliate - no taste buds
• weakly developed in humans
– fungiform
• at tips and sides of tongue
– vallate (circumvallate)
• at rear of tongue
• contains 1/2 of all taste buds
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Epiglottis
Lingual tonsil
Palatine tonsil
Vallate
papillae
Foliate
papillae
Fungiform
papillae
(a) Tongue
Figure 16.6a
Taste Bud Structure
• Lemon-shaped groups of 40 – 60 taste
cells, supporting cells, and basal cells
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Vallate
papillae
Filiform
papillae
• Taste cells
– have tuft of apical microvilli (taste
hairs) that project from taste pores
– taste cells are epithelial cells not
neurons
– synapse with and release
neurotransmitters onto sensory
neurons at their base
Taste
buds
(b) Vallate papillae
Figure 16.6b
Foliate
papilla
Taste pore
• Basal cells
– stem cells that replace taste cells
every 7 to 10 days
Taste bud
Figure 16.6c
100 µm
(c) Foliate papillae
c: © Ed Reschke
• Supporting cells
– resemble taste cells without taste
hairs, synaptic vesicles, or sensory
role
Synaptic
vesicles
Sensory
nerve
fibers
Basal
cell
Supporting
cell
Taste
cell
Taste
pore
Taste
hairs
Tongue
epithelium
(d) Taste bud
Figure 16.6d
Projection Pathways for Taste
• Facial nerve collects sensory information from taste buds over
anterior two-thirds of tongue
• Glossopharyngeal nerve from posterior one-third of tongue
• Vagus nerve from taste buds of palate, pharynx and epiglottis
• All fibers reach solitary nucleus in medulla oblongata
• From there, signals sent to two destinations
– hypothalamus and amygdala control autonomic reflexes – salivation,
gagging and vomiting
– thalamus relays signals to postcentral gyrus of cerebrum for conscious
sense of taste
• sent on to orbitofrontal cortex to be integrated with signals from nose and
eyes - form impression of flavor and palatability of food
Chemical Senses- Olfaction
• Olfaction- sense of
smell, involves binding
of odorants to olfactory
hairs
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Olfactory tract
Olfactory bulb
Olfactory nerve
fascicle
Olfactory
mucosa (reflected)
• Olfactory mucosa
– Olfactory cells
– Supporting cells
– Basal stem cells
Figure 16.7a
Olfactory Cells
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• Only neurons in the body
directly exposed to the
external environment
Olfactory bulb
Granule cell
Olfactory tract
– have a lifespan of only 60 days
– basal cells continually divide
and differentiate into new
olfactory cells
Mitral cell
Tufted cell
Glomerulus
Olfactory
nerve fascicle
Cribriform
plate of
ethmoid bone
• Supporting cells
Basal cell
Supporting
cells
Olfactory cell
• Basal cells
Olfactory gland
Olfactory hairs
– divide and differentiate to
replace olfactory cells
Mucus
Odor
molecules
Airflow
(b)
Figure 16.7b
Smell - Physiology
• Humans have poorer sense of smell than most other
mammals
– women more sensitive to odors than men
– highly important to social interaction
• Human Pheromones
– human body odors may affect sexual behavior
Olfactory Projection Pathways
• olfactory cells synapse in olfactory
bulb
– on dendrites of mitral and tufted cells
– dendrites meet in spherical clusters called
glomeruli
• each glomeruli dedicated to single odor, all
fibers leading to one glomerulus have same
receptor type
• tufted and mitral cell axons form
olfactory tracts
– reach primary olfactory cortex in the
inferior surface of the temporal lobe
– secondary destinations –hippocampus,
amygdala, hypothalamus, insula, and
orbitofrontal cortex
– fibers reach back to olfactory bulbs where
granule cells inhibit the mitral and tufted
cells
• reason why odors change under different
conditions
• food smells more appetizing when you are
hungry
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 fluidfilled passages and sensory cells
• fluid is set in motion and how the sensory cells
convert this motion into an informative pattern of
action potentials
Sound: Can You Hear Me?
• Any audible vibration of
molecules
• Pitch (Hertz, Hz) and
Loudness (decibel, db)
– Range 20 – 20000 Hz
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Anatomy of Ear
• Three sections outer, middle, and inner ear
– first two concerned only with transmission of sound to inner ear
– inner ear – vibrations converted to nerve signals
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Ossicles:
Stapes
Incus
Malleus
Helix
Semicircular ducts
Oval window
Vestibular nerve
Cochlear nerve
Vestibule
Auricle
Cochlea
Round window
Tympanic
membrane
Tympanic cavity
Auditory
canal
Tensor tympani
muscle
Auditory tube
Lobule
Outer ear
Middle ear
Inner ear
Figure 16.11
Outer (External) Ear
• Outer ear – a funnel for conducting vibrations to the
tympanic membrane (eardrum)
– auricle (pinna) directs sound down the auditory canal
• shaped and supported by elastic cartilage
– auditory canal – passage leading through the temporal bone to
the tympanic membrane
– external acoustic meatus – slightly s-shaped tube that begins at
the external opening and courses for about 3 cm
• guard hairs protect outer end of canal
• cerumen (earwax) – mixture of secretions of ceruminous and sebaceous
glands and dead skin cells
–
–
–
–
sticky and coats guard hairs
contains lysozyme with low pH that inhibits bacterial growth
water-proofs canal and protects skin
keeps tympanic membrane pliable
Middle Ear
• middle ear - located in the air-filled
tympanic cavity in temporal bone
– tympanic membrane (eardrum) – closes
the inner end of the auditory canal
• innervated by sensory branches of the
vagus and trigeminal nerves
– highly sensitive to pain
– tympanic cavity
• contains auditory ossicles
– auditory (eustachian) tube connects
middle ear cavity to nasopharynx
– auditory ossicles
– stapedius and tensor tympani muscles
attach to stapes and malleus
Anatomy of Inner Ear
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Temporal
bone
Figure 16.12a
Figure 16.12b
Inner (Internal) Ear
• bony labyrinth - passageways in temporal bone
• membranous labyrinth - fleshy tubes lining the bony
labyrinth
– filled with endolymph - similar to intracellular fluid
– floating in perilymph - similar to cerebrospinal fluid
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Endolymphatic
sac
Temporal bone
Dura mater
Semicircular ducts:
Anterior
Figure 16.12c
Posterior
Scala vestibuli
Lateral
Scala tympani
Semicircular canal
Cochlear duct
Ampulla
Vestibule:
Saccule
Utricle
Tympanic
membrane
(c)
Stapes
in oval window
Secondary tympanic membrane
in round window
Anatomy of Cochlea
• cochlea has three fluid-filled chambers separated by
membranes:
– scala vestibuli – superior chamber
• begins at oval window and spirals to apex
– scala tympani – inferior chamber
• begins at apex and ends at round window
– secondary tympanic membrane – membrane covering round window
– scala media (cochlear duct) – triangular middle chamber
• filled with endolymph
• separated from:
– scala vestibuli by vestibular membrane
– scala tympani by thicker basilar membrane
• contains spiral organ - organ of Corti - acoustic organ – converts vibrations
into nerve impulses
Cochlea, Cochlear Duct and Spiral Organ
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Oval window
Figure 16.13
Vestibular
membrane
Cochlear duct
(scala media)
Spiral ganglion
Cochlear nerve
Scala vestibuli
(with perilymph)
(a)
Vestibular
membrane
Tectorial
membrane
Cochlear duct
(with
endolymph)
Hairs
(stereocilia)
Outer
hair cells
Supporting
cells
Basilar
membrane
Scala tympani
(with perilymph)
Tectorial
membrane
Spiral
organ
Inner hair
cell
Basilar
membrane
Fibers of
cochlear nerve
(b)
(c)
Spiral Organ (Organ of Corti)
• spiral organ has epithelium composed of hair cells
and supporting cells
• hair cells have long, stiff microvilli called stereocilia
on apical surface
• gelatinous tectorial membrane rests on top of stereocilia
• spiral organ has four rows of hair cells spiraling along
its length
– inner hair cells – single row of about 3500 cells
• provides for hearing
– outer hair cells – three rows of about 20,000 cells
• adjusts response of cochlea to different frequencies
• increases precision
SEM of Cochlear Hair Cells
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Outer hair cells
Inner hair cells
Figure 16.14
10 µm
Quest/Science Photo Library/Photo Researchers, Inc.
Physiology of Hearing - Middle Ear
• tympanic membrane
– ossicles create a greater force per unit area at the oval window and
overcomes the inertia of the perilymph
– ossicles and their muscles have a protective function
• lessen the transfer of energy to the inner ear
• tympanic reflex
– during loud noise, the tensor tympani pulls the tympanic membrane
inward and tenses it
– stapedius muscle reduces the motion of the stapes
– muffles the transfer of vibration from the tympanic membrane to the
oval window
– middle ear muscles also help to coordinate speech with hearing
• dampens the sound of your own speech
Stimulation of Cochlear Hair Cells
• vibration of ossicles causes vibration of basilar
membrane under hair cells
– as often as 20,000 times per second
– hair cells move with basilar membrane
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Outer ear
Middle ear
Inner ear
Stapes
Oval
window
Incus
Malleus
Basilar
membrane
Sound
wave
Tympanic
membrane
Auditory
tube
Air
Fluid
Secondary
tympanic
membrane
(in round
window)
Figure 16.15
Excitation of Cochlear Hair Cells
• stereocilia of outer hair cells
– bathed in high K+ fluid, the endolymph
• creating electrochemical gradient
• outside of cell is +80 mV and inside about – 40 mV
– tip embedded in tectorial membrane
• stereocilium on inner hair cells
– single transmembrane protein at tip that functions as a mechanically
gated ion channel
• stretchy protein filament (tip link) connects ion channel of one stereocilium to
the sidewall of the next taller stereocilium
• tallest one is bent when basilar membrane rises up towards tectorial
membrane
• pulls on tip links and opens ion channels
• K+ flows in – depolarization causes release of neurotransmitter
• stimulates sensory dendrites and generates action potential in the cochlear
nerve
Potassium Gates
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Unstimulated
Stimulated
Tip link
Mechanically
gated K+
channel
Stereocilia
K+
Surface of
hair cell
K+
K+
gate
closed
K+
gate
open
Figure 16.16
Sensory Coding
• for sounds to carry meaning, we must distinguish between
loudness and pitch
• variations in loudness (amplitude) cause variations in the intensity
of cochlear vibrations
– soft sound produces relatively slight up-and-down motion of the basilar
membrane
– louder sounds make the basilar membrane vibrate more vigorously
• triggers higher frequency of action potentials
• brain interprets this as louder sound
• pitch depends on which part of basilar membrane vibrates
– at basal end, membrane attached, narrow and stiff
• brain interprets signals as high-pitched
– at distal end, 5 times wider and more flexible
• brain interprets signals as low-pitched
Basilar Membrane Frequency Response
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Tympanic membrane (vibrating)
Stapes footplate (vibrating)
Scala vestibuli
Scala tympani
Secondary
tympanic
membrane
Cochlear
duct
Helicotrema
Basilar
membrane
(vibrating)
(a)
Low-frequency sound
(20–800 Hz)
Medium-frequency sound
(1,500–4,000 Hz)
High-frequency sound
(7,000–20,000 Hz)
Figure 16.17
(b)
Proximal end
(attached)
(c)
Distal end
(free)
20,000
5,000
1,000
500
200 Hz
notice high and low frequency ends
Cochlear Tuning
• increases ability of cochlea to receive some sound
frequencies
• outer hair cells shorten (10 to 15%) reducing basilar
membrane’s mobility
– fewer signals from that area allows brain to distinguish
between more and less active areas of cochlea
• pons has inhibitory fibers that synapse near the base
of inner hair cells
– inhibiting some areas and increases contrast between
regions of cochlea
Deafness
• deafness – hearing loss
– conductive deafness - conditions interfere with
transmission of vibrations to inner ear
• damaged tympanic membrane, otitis media, blockage of
auditory canal, and otosclerosis
– otosclerosis - fusion of auditory ossicles that prevents their free
vibration
– sensorineural (nerve) deafness - death of hair cells
or any nervous system elements concerned with
hearing
• factory workers, musicians and construction workers
Auditory Projection Pathway
• sensory fibers begin at the bases of the hair cells
– axons lead away from the cochlea as the cochlear nerve
– joins with the vestibular nerve to form the vestibulocochlear nerve,
Cranial Nerve VIII
• each ear sends nerve fibers to both sides of the pons
– end in cochlear nuclei
– synapse with second-order neurons that ascend to the nearby superior
olivary nucleus
– superior olivary nucleus issues efferent fibers back to the cochlea
• involved with cochlear tuning
• binaural hearing – comparing signals from the right and left
ears to identify the direction from which a sound is coming
– function of the superior olivary nucleus
Auditory Projection Pathway
• fibers ascend to the inferior colliculi of the midbrain
– helps to locate the origin of the sound, processes fluctuation in pitch,
and mediate the startle response and rapid head turning in response to
loud noise
• third-order neurons begin in the inferior colliculi and lead to the
thalamus
• fourth-order neurons complete the pathway from thalamus to
primary auditory complex
– involves four neurons instead of three unlike most sensory pathways
• primary auditory cortex lies in the superior margin of the
temporal lobe
– site of conscious perception of sound
• because of extensive decussation of the auditory pathway,
damage to right or left auditory cortex does not cause unilateral
loss of hearing
Equilibrium
• equilibrium – coordination, balance, and orientation in threedimensional space
• vestibular apparatus – constitutes receptors for equilibrium
– three semicircular ducts
• detect only angular acceleration
– two chambers
• anterior saccule and posterior utricle
• responsible for static equilibrium and linear acceleration
• static equilibrium – the perception of the orientation of the
head when the body is stationary
• dynamic equilibrium - perception of motion or acceleration
• linear acceleration - change in velocity in a straight line (elevator)
• angular acceleration - change in rate of rotation (car turns a corner)
Saccule and Utricle
• macula – 2 by 3 mm patch of hair cells and supporting cells in the
saccule and utricle
– macula sacculi – lies vertically on wall of saccule
– macula utriculi – lies horizontally on floor of utricle
• each hair cell has 40 to 70 stereocilia and one true cilium kinocilium embedded in a gelatinous otolithic membrane
– otoliths - calcium carbonate-protein granules that add to the weight and
inertia and enhance the sense of gravity and motion
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Otoliths
Hair cell
Supporting cell
Otolithic
membrane
Vestibular
nerve
Figure 16.19b
(b)
16-32
Macula Utriculi and Macula Sacculi
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Macula utriculi
Macula sacculi
(a)
Figure 16.19
Otoliths
Supporting cell
Hair cell
Vestibular
nerve
Otolithic
membrane
Stereocilia
of hair
cells bend
Otolithic
membrane
sags
(b)
(c)
Gravitational force
•
static equilibrium - when head is tilted, heavy otolithic membrane sags, bending
the stereocilia, and stimulating the hair cells
•
dynamic equilibrium – in car, linear acceleration detected as otoliths lag behind,
bending the stereocilia, and stimulating the hair cells
•
because the macula sacculi is nearly vertical, it responds to vertical acceleration
and deceleration
Crista Ampullaris
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Semicircular ducts:
Anterior
Posterior
Lateral
Crista ampullaris
and cupula
Ampullae
(a)
Cupula
Endolymph
Hair cells
Supporting
cells
Sensory
nerve fibers
Crista
ampullaris
Figure 16.20 a-b
(b)
• consists of hair cells with stereocilia and a kinocilium buried
in a mound of gelatinous membrane called the cupula
(one in each duct)
• orientation causes ducts to be stimulated by rotation in different
16-34
planes
Crista Ampullaris - Head Rotation
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Semicircular ducts:
Anterior
Posterior
Lateral
Crista ampullaris
and cupula
Figure 16.20
Ampullae
(a)
Direction of
head rotation
Cupula
Cupula is
pushed over
and stimulates
hair cells
Endolymph
Hair cells
Supporting
cells
Endolymph lags
behind due
to inertia
Crista
ampullaris
Sensory
nerve fibers
(b)
(c)
• as head turns, endolymph lags behind, pushes cupula,
16-35
stimulates hair cells
Equilibrium Projection Pathways
• information sent to five targets:
– cerebellum – control of head and eye movements, muscle tone,
and posture
– nuclei of oculomotor, trochlear, and abducens nerves (CN III, IV,
and VI) to produce vestibulo-ocular reflex
– reticular formation – thought to adjust blood circulation and
breathing to postural changes
– spinal cord – descend through two vestibulospinal tracts of
spinal cord and innervate extensor (antigravity) muscles
– thalamus - thalamic relay to cerebral cortex for awareness of
position and motor control of head and body
Vision
• Perception of objects in the environment by
means of the light that they reflect or emit
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External Anatomy of Eye
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Pupil
Eyebrow
Upper
eyelid
Eyelashes
Iris
Palpebral
fissure
Sclera
Medial
commissure
Figure 16.22
Lateral
commissure
Lower
eyelid
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Conjunctiva
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Frontal bone
Levator palpebrae
superioris muscle
Orbicularis
oculi muscle
Superior rectus
muscle
Figure 16.23a
Tarsal plate
Tarsal glands
Cornea
Conjunctiva
Lateral rectus
muscle
Inferior rectus
muscle
(a)
• transparent mucous membrane that lines eyelids and covers anterior
surface of eyeball, except cornea
• richly innervated and vascular (heals quickly)
– secretes a thin mucous film that prevents the eyeball from drying
Lacrimal Apparatus
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Lacrimal
gland
Figure 16.23b
Ducts
Lacrimal
sac
Lacrimal
punctum
Lacrimal
canal
Nasolacrimal
duct
Inferior
meatus
of nasal
cavity
Nostril
(b)
• tears flow across eyeball help to wash away foreign particles, deliver O2 and
nutrients, and prevent infection with a bactericidal lysozyme
• tears flow through lacrimal punctum (opening on edge of each eyelid) to
the lacrimal sac, then into the nasolacrimal duct emptying into nasal cavity
Extrinsic Eye Muscles
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Trochlear nerve (IV)
Abducens nerve (VI)
Levator palpebrae
superioris muscle
Superior
oblique
muscle
Superior rectus
muscle
Medial rectus
muscle
Lateral
rectus
muscle
Oculomotor nerve (III)
Inferior rectus
muscle
Figure 16.24c
Inferior oblique
muscle
(c) Frontal view
• superior and inferior oblique mm. turn the “twelve o’clock pole”
of each eye toward or away from the nose
Anatomy of the Eyeball
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Sclera
Ora serrata
Choroid
Ciliary body
Retina
Macula lutea
Suspensory
ligament
Fovea centralis
Optic disc
(blind spot)
Iris
Cornea
Optic nerve
Pupil
Lens
Central artery
and vein
of retina
Anterior
chamber
Posterior
chamber
Hyaloid canal
Figure 16.25
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 – the retina and optic nerve
Anatomy of the Eyeball
Retina
• Conversion of light energy into action
potential
• Structure
– Pigment epithelium
– Neural components
• Rods and cones
• Bipolar cells
• Ganglion cells
Figure 16.34b
Retina
Visual Pigments
• rods contain visual pigment - rhodopsin (visual purple)
– two major parts of molecule
• opsin - protein portion embedded in disc membrane of rod’s outer
segment
• retinal (retinene) - a vitamin A derivative
– has absorption peak at wavelength of 500 nm
• can not distinguish one color from another
• cones contain photopsin (iodopsin)
– retinal moiety same as in rods
– opsin moiety contain different amino acid sequences that
determine wavelengths of light absorbed
– 3 kinds of cones, identical in appearance, but absorb different
wavelengths of light to produce color vision
Scotopic System (Night Vision)
• rods sensitive – react even in dim light
– extensive neuronal convergence
– 600 rods converge on 1 bipolar cell
– many bipolar converge on each ganglion cell
– results in high degree of spatial summation
• one ganglion cells receives information from 1 mm2 of
retina producing only a coarse image
• edges of retina have widely-spaced rod cells, act
as motion detectors
– low resolution system only
16-47
– cannot resolve finely detailed
images
Color Vision
Photopic System (Day Vision)
• fovea contains only 4000 tiny cone cells (no
rods)
– no neuronal convergence
– each foveal cone cell has “private line to brain”
• high-resolution color vision
– little spatial summation so less sensitivity to dim
light
16-48
Color Vision
• primates have well developed
color vision
– nocturnal vertebrates
have only rods
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S cones
420 nm
Rods M cones L cones
500 nm 531 nm 558 nm
100
80
• three types of cones are named
for absorption peaks of their
photopsins
– short-wavelength (S) cones peak
sensitivity at 420 nm
– medium-wavelength (M) cones peak
at 531 nm
– long-wavelength (L) cones peak at
558 nm
• color perception based on
mixture of nerve signals
representing cones of different
absorption peaks
16-49
60
40
20
400
500
600
700
Wavelength (nm)
Wavelength
(nm)
400
450
500
550
625
675
Percentage of maximum
cone response
Perceived hue
(S:M:L)
50 : 0 : 0
72 : 30 : 0
20 : 82 : 60
0 : 85 : 97
0 : 3 : 35
0: 0: 5
Violet
Blue
Blue-green
Green
Orange
Red
Figure 16.40
Projection Pathway for Vision
• First order neuron- bipolar cells
• Second order neuron- ganglion cells
– Axons form optic nerve
– Hemidecussation
– End in the… (guess where)
• Third order neuron- primary visual cortex
– Occipital lobe
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