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The Cyprus International Institute for the Environment and Public Health
In collaboration with the Harvard School of Public Health
Constantinos Pitris, MD, PhD
Assistant Professor, University of Cyprus
cpitris@ucy.ac.cy
http://www.eng.ucy.ac.cy/cpitris/courses/CIIPhys/
Lecture 4
The Peripheral Nervous System
(181-240)
Excluded: adaptation of pacinian corpuscle (185), labeled lines
(186), acuity (186-187), phototransduction (200-203), on- off- center
ganglion cells (205)
2
Introduction
• Peripheral Nervous System
• Afferent Division
• Sends information from the PNS to the
CNS
• Efferent Division
• Send information from the CNS to the PNS
• Afferent Division
• Visceral afferents (subconscious input)
• Pressure, O2, temperature, etc.
• Sensory afferents (conscious input)
• Somatic sensation
• Somesthetic sensation from skin
• Proprioception from muscle joints, skin and
inner ear
• Special senses
• Vision, hearing, taste and smell
• Efferent Division
• Autonomic Nervous System
• Cardiac muscle, smooth muscle, most
exocrine glands, some endocrine glands,
adipose tissue
• Somatic Nervous system
• Skeletal muscle
3
Introduction
• Sensation ≠ Perception
• Perception
• Our understanding (conscious
interpretation) of the physical world
• An interpretation of the senses
• Different from what is out there
because
• Our receptors detect limited number
of existing energy forms
• The information does not reach our
brain unaltered. Some features are
accentuated and some are
suppressed
• The brain interprets the information
and often distorts it (“completes the
picture” or “feels in the gaps”) to
extract conclusions.
• Interpretation is affected by cultural,
social and personal experiences
stored in our memory
4
Receptor Physiology
• Receptors
• Detect stimulus (detectable change) from different modalities (energy forms)
• e.g. light, heat, sound, pressure, chemical changes
• Adequate stimulus = the stimulus to which the receptor is most sensitive
• Convert forms of energy into electrical signals (action potentials)
• Process is called transduction
• Types of receptors
• Photoreceptors
• Responsive to visible wavelengths of light
• Mechanoreceptors
• Sensitive to mechanical energy
• Thermoreceptors
• Sensitive to heat and cold
• Osmoreceptors
• Detect changes in concentration of solutes in body fluids and resultant changes in osmotic activity
• Chemoreceptors
• Sensitive to specific chemicals
• Include receptors for smell and taste and receptors that detect O2 and CO2 concentrations in blood
and chemical content of digestive tract
• Nociceptors
• Pain receptors that are sensitive to tissue damage or distortion of tissue
5
Receptor Physiology
• Receptors may be
• Specialized ending of an afferent neuron
Æ receptor potential
• Separate cell closely associated with
peripheral ending of a neuron Æ
generator potential
• Potential generation
• Separate receptor
• Stimulus causes release of chemical
messenger
• Specialized afferent nerve ending
• Stimulus alters receptor’s permeability
which leads to graded receptor potential
• Usually causes nonselective opening of
all small ion channels Æreceptor
(generator) potentials.
• The magnitude of the receptor potential
represents the intensity of the stimulus.
• A receptor potential of sufficient
magnitude can produce an action
potential.
• This action potential is propagated along
an afferent fiber to the CNS.
Chemically gated
channels
Afferent
neuron fiber
Diffusion
chemical
messenger
Receptor
(separate cell)
Afferent
neuron fiber
Voltage-gated channels
Receptor
(separate cell)
6
Receptor Physiology
• May adapt slowly or rapidly to
sustained stimulation
• Types of receptors according to
their speed of adaptation
• Tonic receptors
• Do not adapt at all or adapt slowly
• Muscle stretch receptors, joint
proprioceptors (to continuously
receive information regarding
posture and balance)
• Phasic receptors
• Rapidly adapting receptors
• Tactile receptors in skin (the reason
you don’t “feel” your clothes or
watch)
• Adaptation is not the same as
habituation (synapse changes in
the CNS)
Stimulus
off
Slowly
adapting
Stimulus
on
Time
Receptor
potential
(mV)
Stimulus
strength
Stimulus
off
Rapidly
adapting
Stimulus
on
Time
Receptor
potential
(mV)
Stimulus
strength
Off response
7
Pain
• Pain
• Primarily a protective mechanism meant
to bring a conscious awareness that
tissue damage is occurring or is about to
occur
• Storage of painful experiences in
memory helps us avoid potentially
harmful events in future
• Sensation of pain is accompanied by
motivated behavioral responses and
emotional reactions
• Subjective perception can be influenced
by other past or present experiences
(Are you afraid of your dentist?)
8
Pain
• Three categories of nociceptors
• Mechanical nociceptors
• Respond to mechanical damage such as
cutting, crushing, or pinching
• Thermal nociceptors
• Respond to temperature extremes
• Polymodal nociceptors
• Respond equally to all kinds of damaging
stimuli
• Presence of prostaglandins (released
after tissue injury)
• Lowers nociceptors threshold for activation
• Greatly enhances receptor response to
noxious stimuli
• Aspirin-like drugs inhibit their synthesis Æ
analgesic effect
• Nociceptors do not adapt to sustained or
repetitive stimulation
9
Pain
• Characteristics of pain
Provoked and sustained by release of
bradykinin
Occurs second, persists for longer time,
more unpleasant
Occurs first
Poorly localized
Easily localized
Produces dull, aching, burning
sensation
Produces sharp, prickling sensation
Carried by small, unmyelinated C fibers
Carried by small, myelinated A-delta
fibers
Occurs on stimulation of polymodal
nociceptors
Occurs on stimulation of mechanical
and thermal nociceptors
Slow Pain
Fast Pain
10
Pain
• Two best known pain
neurotransmitters
• Substance P
• Glutamate
• Substance P
• Activates ascending
pathways that
transmit nociceptive
signals to higher
levels for further
processing
Somatosensory
cortex
Thalamus
Hypothalamus
limbic system
Reticular
formation
Noxious
stimulus
Afferent pain fiber
Substance P
Spinal
cord
Brain
stem
Higher
brain
(Location of pain)
(Perception of pain)
( Alertness)
(Behavioral and
emotional responses
to pain)
Nociceptor
Other cortical
areas
(Experience and
deliberation of pain)
11
Pain
• Glutamate
• Major excitatory
neurotransmitter
• Causes hypersensitivity in the
area
• Protection mechanism for
healing and against further
damage
• How does a sunburn feel?
12
Pain
• Brain has built in analgesic system
• Suppresses transmission in pain pathways as they enter
spinal cord
• Suppress release of Substance P
• Depends on presence of opiate receptors
• Endogenous opiates (morphine like substances) –
endorphins, enkephalins, dynorphin
• Factors which modulate pain
• Exercise (“runner’s high”)
• Stress (survival mechanism)
• Acupuncture
Periagueductal
gray matter
Opiate
receptor Noxious
stimulus
Afferent pain fiber
Substance P
No perception of pain
To thalamus
Transmission
of pain
impulses to
brain blocked
Nociceptor
Reticular
formation
Endogenous opiate
13
Vision
• Eye
• Sensory organ for vision
• Mechanisms that help protect
eyes from injury
• Eyeball is sheltered by bony
socket in which it is positioned
• Eyelids
• Act like shutters to protect eye
from environmental hazards
• Eyelashes
• Trap fine, airborne debris such
as dust before it can fall into
eye
• Tears
• Continuously produced by
lacrimal glands
• Lubricate, cleanse, bactericidal
Canal for tear
drainage
Pupil
Iris
Sclera
Lacrimal Glands
(under eyelid)
14
Vision
• Eye
• Spherical, fluid-filled structure enclosed
by three tissue layers
• Sclera/cornea
• Sclera – tough outer layer of
connective tissue; forms visible white
part of the eye
• Cornea – anterior, transparent outer
layer through which light rays pass into
interior of eye
• Choroid/ciliary body/iris
• Choroid - middle layer underneath
sclera which contains blood vessels
that nourish retina
• Choroid layer is specialized anteriorly
to form ciliary body and iris
• Retina
• Innermost coat under choroid
• Consists of outer pigmented layer and
inner nervous-tissue layer
• Rods and cones
Suspensory ligament
Ciliary body
Conjunctiva
Iris
Pupil
Lens
Cornea
Aqueous humor
Vitreous humor
Blood vessels
Optic disc
Optic nerve
Fovea
Sclera
Retina
Choroid
Extrinsic eye
muscle
15
Vision
• Eye
• Pupil
• Round opening through which light enters the eye
• Iris
• Controls amount of light entering eye
• Contains two sets of smooth muscle networks
• Circular (or constrictor) muscle
• Radial (or dilator) muscle
• Pigment in iris is responsible for eye color
• Unique for each individual
• Basis for latest identification technology
Parasympathetic stimulation
+
Pupillary constriction
Circular
(constrictor)
muscle runs
circularly
Radial
muscle
of iris
Circular
muscle
of iris
Pupil Iris Radial
(dilator)
muscle runs
radially
Pupillary dilation
Sympathetic stimulation
+
16
Vision
• Convex structures of eye
produce convergence of
diverging light rays that reach
eye
• Two structures most
important in eye’s refractive
ability are
• Cornea
• Contributes most extensively to
eye’s total refractive ability
• Refractive ability remains
constant because curvature
never changes
• Lens
• Refractive ability can be
adjusted by changing curvature
as needed for near or far vision
(accommodation)
17
Vision
• Accommodation
• Change in strength and shape of lens
• Accomplished by action of ciliary
muscle and suspensory ligaments
• Age-related reduction in
accommodation ability - presbyopia Suspensory
ligaments
Ciliary muscle
Lens
Pupillary
opening
in front
of lens
Sympathetic
stimulation
Iris
Cornea
Flattened
weak
lens
Relaxed
ciliary
muscle
Taut
suspensory
ligaments
Sympathetic
stimulation
Rounded
strong
lens
Contracted
ciliary
muscle
Slackened
suspensory
ligaments
18
Vision
Far source Near source
No accommodations Accommodations
Normal eye (Emmetropia)
Far source focused on retina without
accommodation
Near source focused on retina with
accommodation
No accommodations No accommodations
Nearsightedness (Myopia)–
Eyeball too long or lens too strong
1. Uncorrected
Far source focused in front of
retina (where retina would be in
eye of normal length)
Focus
1.
Image
out of
focus
Near source focused on retina
with accommodations
Farsightedness (Hyperopia)–
Eyeball too short or lens too weak
1. Uncorrected
Far source focused on retina
with accommodations
1.
Near source focused behind
retina even with accommodations
Accommodations Accommodations
Image
out of
focus
Focus
19
Vision
• Retina
• Several layers of cells
• Receptor containing
portion is actually an
extension of the CNS
• Fovea
• Pinhead-sized depression
in exact center of retina
• Point of most distinct
vision
• Has only cones
Optic nerve
Retina
Direction of light
Direction of retinal visual processing
Front
of
retina
Fibers of
the optic
nerve
Ganglion
cell
Amacrine
cell
Bipolar
cell
Cone Rod
Photoreceptor
cells
Horizontal
cell
Retina
Pigment layer
Choroid layer
Sclera
Back
of
retina
20
Vision
• Photoreceptors
• Rod and cone cells
• Photopigments on the disk
membranes
• Rod Æ one type
• one pigment, high sensitivity
• Cones Æ three different types
• Red, green, blue sensing
pigments, lower sensitivity
• Undergo chemical alterations
when activated by light
• Change the receptor potential
• Induce action potentials
• Unlike other receptors,
photoreceptors hyperpolarize!
Back of retina
Outer
segment
Outer
segment
Inner
segment
Synaptic
terminal
Synaptic
terminal
Inner
segment
Direction
of
light
Cells of
pigment layer
Cone Rod
Discs
Mitochondria
Nuclei
Dendrites
of bipolar
cells
Front
of retina
21
Vision
• Color Vision
• Perception of color
• Depends on the ratio of
stimulation of three different
cones
• Different absorption of cone
pigments
• Coded and transmitter by
different pathways
• Processed in color vision center
of primary visual cortex
• Color blindness
• Defective cone
• Colors become combinations of
two cones
• Most common = red-green color
blindness
Color
perceived
Blue cone Green cone Red cone
Visible
spectrum
Wavelength of light (nm)
22
Vision
• Properties of Rod and Cone Vision
Day vision (lack sensitivity and convergence)
Night vision (from sensitivity and convergence)
Little convergence in retinal pathways
Much convergence in retinal pathways
High acuity
Low acuity
Concentrated in fovea
More numerous in periphery
Low sensitivity to light
High sensitivity to light
Color vision
Vision in shades of gray
3 million per retina
100 million per retina
Cones
Rods
23
Vision
• The sensitivity of the eyes varies through dark and light
adaptation
• Dark adaptation
• Can gradually distinguish objects as you enter a dark area.
• Due to the regeneration of rod photopigments that had been broken down
by previous light exposure.
• Light adaptation
• Can gradually distinguish objects as you enter an area with more light.
• Due to the rapid breakdown of cone photopigments.
• Along with pupillary reflexes increase the range of vision
24
Vision
25
Vision
• Visual field
• Area which can be seen without
moving the head) Æ overlap
between eyes
• Visual Pathway
• Optic nerve
• Optic chiasm
• Thalamus
• Sorts information to appropriate
areas for processing
• Optic radiation
• Primary visual cortex (occipital lobe)
• Higher processing areas
• Information arrives altered at the
primary visual cortex
• Upside down and backward
because of the lens
• The left and right halves of the
brain receive information from
the left and right halves of the
visual field
Left Right
Left
eye
Right
eye
Optic
nerve
Optic
chiasm
Optic tract
Lateral
geniculate
nucleus of
thalamus
Optic radiation
Optic lobe
1
2
3
26
Vision
• >30% of cortex participates in
visual information processing
• “What” and “where” pathways
• Depth Perception
• Visual field of two eyes slightly
different
• Depth perception with one eye
• Other cues (such as size,
location, experience)
27
Hearing
• Hearing
• Neural perception of sound
energy
• Involves two aspects
• Identification of the sounds
(“what”)
• Localization of the sounds
(“where”)
• Sound waves
• Traveling vibrations of air
• Consist of alternate regions of
compression and rarefaction of
air molecules
Region of
compression
Region of
rarefaction
28
Hearing
• Characteristics of sound
• Pitch (tone) of sound
• Depends on frequency of air waves
• Intensity (loudness)
• Depends on amplitude of air waves
• Timbre (quality)
• Determined by overtones
29
Hearing
• Ear
• Consists of three parts
• External ear
• Consists of pinna, external
auditory meatus, and tympanum
• Transmits airborne sound waves
to fluid-filled inner ear
• Amplifies sound energy
• Middle ear
• Transmits airborne sound waves
to fluid-filled inner ear
• Amplifies sound energy
• Inner ear
• Houses two different sensory
systems
• Cochlea (Contains receptors for
conversion of sound waves into
nerve impulses which makes
hearing possible)
• Vestibular apparatus (Necessary for
sense of equilibrium)
Inner
ear
Pinna of
external ear
Tympanic
membrane
(eardrum)
Middle
ear
External
ear
Auditory
ossicles Semicircular
canals
Utricle
and saccule
Oval window
Vestibulocochlear nerve
Cochlea
Round window
Eustachian
tube
To pharynx
External
auditory meatus
(ear canal)
30
Malleus Incus
Stapes
at oval
window
Helicotrema
Cochlea
Scala vestibuli
Scala tympani
Round window
Middle ear
cavity
Tympanic membrane
External
auditory
meatus
Vestibular membrane
Basilar membrane
Tectorial membrane
Organ of Corti
Scala media
Hearing
• Sound Wave Transmission
• Tympanic membrane
• Vibrates when struck by sound
waves
• Middle ear
• Transfers vibrations through
ossicles (malleus, incus, stapes)
to oval window (entrance into fluid-
filled cochlea)
• Amplify the pressure 20x
• Large surface of the tympanic
membrane transferred to the
smaller oval window
• Lever action of the osscicles
• Small muscles change the stifness
of the tympanic membrane
• Protection mechanism
• Slow action (40 msec) Æ protects
only from prolonged sounds
31
Malleus Incus
Stapes
at oval
window
Helicotrema
Cochlea
Scala vestibuli
Scala tympani
Round window
Middle ear
cavity
Tympanic membrane
External
auditory
meatus
Vestibular membrane
Basilar membrane
Tectorial membrane
Organ of Corti
Scala media
Hearing
• Sound Wave Transmission
• Inner ear
• Sound dissipates in cochlea
• Waves in cochlear fluid
(endolymph) set basilar
membrane in motion
• Sound converted to electrical
signals by the Organ of Corti
32
Hearing
• Sound Transduction
• Organ of Corti
• Hair cells with ~ 100 sterocilia
each
• Inner hair cells are tilted as
basilar membrane oscillates
• Mechanically gated channels
open and close Æ graded
potentials
• Communicate via chemical
signals with the nerves which
form the auditory nerve
• Outer hair cells Æ adjust length
with respond to electrical
stimulus (electromotility) Æ
accentuate motion of basilar
membrane and fine tune
response
Vestibular membrane
Tectorial membrane
Scala
media
(cochlear
duct)
Basilar
membrane
Scala tympani
Auditory
nerve
Scala
vestibuli
Outer
hair cells
Hairs
(stereocilia)
Tectorial membrane
Inner hair cells
Nerve fibers
Supporting cell
Basilar membrane
33
Hearing
• Auditory pathway
• Hair cells
• Auditory nerve
• Brain stem
• Signals cross over to opposite site
(unlike visual signals)
• Thalamus
• Sorts and relays signals to cortical
regions
• Cortex
• Cortical Processing
• Primary auditory cortex in the
temporal lobe
• Tonotopically mapped (i.e.
mapped according to tone)
• Higher cortical processing
• Separates coherent and
meaningful patterns
34
Hearing
• Pitch discrimination
• Basilar membrane does not have uniform mechanical properties
• Narrow and stiff to wide and flexible
• Different regions vibrate maximally at different frequencies
• Frequency (or frequencies) are discriminated by the location of hair cells
firing
Wide, flexible end
of basilar membrane
near helicotrema
Narrow, stiff end
of basilar membrane
near oval window
High frequency
Medium frequency
Low frequency
35
Hearing
• Loudness discrimination
• Exquisitely sensitive organ
(motion less than a molecule of
Hydrogen) Æ easily damaged
• Wide range (every 10 dB
means 10-fold increase in
intensity)
• Higher intensity causes larger
basilar membrane movement
• Stronger graded potential of
hair cells
• Faster rate of action potentials
from auditory nerve cells
• Anything > 100 dB can cause
permanent damage
1 quadrillion x
150 dB
Takeoff of jet
plane
1 trillion x
120 dB
Loud rock
concert
1 billion x
90 dB
Food Blender
1 million x
60 dB
Normal
conversation
1000 x
30 dB
Hush of
Library
100 x
20 dB
Ticking of
watch
10 x louder
10 dB
Rustle of
leaves
Comparison to
faintest audible
sound (hearing
threshold)
Loudness
in decibels
(dB)
Sound
36
Hearing
• Localization
• Up-Down localization (elevation)
• External ear (Pinna) shape
changes sound timbre and
intensity slightly according to
elevation
• Left-right localization (azimuth)
• Sound arriving to proximal ear
arrives
• Slightly earlier (~ 0.5 msec)
• Slightly stronger
• Brain uses the electrical activity
changes to these two cues to
localize the direction
37
Hearing
• Deafness
• Conductive
• Sound waves not adequately
conducted through external and
middle portions of ear
• Blockage, rupture of ear drum,
middle ear infection, iddle ear
adhesions
• Hearing aids might help
• Senosineural
• Sound waves conducted but not
translated into electrical signals
• Neural presbycusis, certain
antibiotics, poisoning
• Cochlear implants might help
• Electrical devices stimulating
the auditory nerve directly
38
Equilibrium
• Detection of position
and motion
• Posture and
coordination
• Vestibular apparatus
• Fluid filled tunnels In
the inner ear
• Semicircular canals
• Three circular tunnels
arranged on
perpendicular planes
• Otolith organs (Utricle
and saccule)
• Two bulges arranged
in perpendicular
directions
39
Equilibrium
• Semicircular canals
• Three circular tunnels arranged on perpendicular planes
• Detect rotational acceleration or deceleration in any direction
• Hair cells
• On a ridge in the ampulla
• Have on kinocilium and several sterocilia (mikrovilli)
• Embedded in gelatinous material, the cupula
Semicircular
canals
Perilymph
Endolymph
Ampulla
Oval window
Round window
Cochlea
Vestibular
apparatus
Utricle
Saccule
Vestibular
nerve Auditory
nerve
Hair
cell
Support
cell
Ridge in
ampulla
Vestibular
nerve fibers
Hairs of hair cell;
kinocilium (red)
and stereocilia (blue)
Cupula
40
Direction of
head rotation
Left
horizontal
semicircular
canal
Direction
of fluid
movement in
semicircular
canals
Direction of
bending of
cupula and
hairs of
receptor hairs cells
Right
horizontal
semicircular
canal
Equilibrium
• Semicircular canals
• Signal transduction
• Head is rotated
• Two of the canals are rotated around
their axis in opposite directions
• Endolymph moves opposite to the
direction of motion (inertia)
• Cupula leans in that direction
• Cilia bent and K+ channels open or
close
• The haircells are depolarized or
hyperpolarized
• Neurotransmitter realize from the hair
cells is modified
• Firing of the vestibular nerve is
modified
Kinocilium
Stereocilia
Hair cell
41
Equilibrium
• Vestibular pathway
• Vestibular nuclei in brain stem
• Motor neurons for controlling
eye movement, perceiving
motion and orientation
• E.g. vestibuloocular reflex
• Cerebellum for use in
maintaining balance and
posture,
• The vestibular system detects
acceleration
• Speed is calculated by
integrating circuitry in the brain
stem
Head motion
Eye Movement
Endolymph
Motion
42
Equilibrium
• Otolith Organs
• Detect changes in rate of linear movement in
any direction
• Arranged in perpendicular directions
• Provide information important for determining
head position in relation to gravity
• Hair cells
• As described before
• In addition, calcium carbonate crystals (otoliths)
are embedded in within the gelatinous layer
• Increased inertia
• Sensitivity to gravity
43
Equilibrium
• Otolith Organs
• Signal transduction
• Utricle Æ Forward or backward
motion or tilt (motion due to
gravitational force)
• Saccule Æ vertical motion
• Endolymph and gelatinous
mass with otoliths move in the
opposite direction
• Cilia bent and K+ channels open
or close
• The haircells are depolarized or
hyperpolarized
• Neurotransmitter realize from
the hair cells is modified
• Firing of the vestibular nerve is
modified
Gravitational
force
44
Taste and Smell
• Taste (gustation) and smell
(olfaction)
• Receptors are chemoreceptors
• In association with food intake,
influence flow of digestive juices and
affect appetite
• Stimulation of receptors induces
pleasurable or objectionable
sensations and signals presence of
something to seek or to avoid
• In lower animals also play a role in
finding direction, seeking prey,
avoiding predators and sexual
attraction to a mate
• Less developed and important in
humans
• Really? How much do you spend on
perfumes and colognes
45
Taste (Gustation)
• Chemoreceptors housed in
taste buds
• Present in oral cavity and
throat
• Taste receptors have life span
of about 10 days
• Taste bud consists of
• Taste pore
• Opening through which fluids in
mouth come into contact with
surface of receptor cells
• Taste receptor cells
• Modified epithelial cells with
surface folds called microvilli
• Plasma membrane of microvilli
contain receptor sites that bind
selectively with chemical
molecules
Taste bud
Sensory
nerve fiber
Papilla
Surface
of the tongue
Taste pore
Taste receptor
cell
Supporting cell
Tongue
46
Taste (Gustation)
• Signal transduction
• Tastant (taste-provoking
chemical)
• Binding of tastant with receptor
cell
• Alters cell’s ionic channels to
produce depolarizing receptor
potential
• Receptor potential releases
neurotransmitter
• Initiates action potentials within
terminal endings of afferent nerve
fibers with which receptor cell
synapses
• Signals conveyed via synaptic
stops in brain stem and thalamus
to cortical gustatory area
47
Taste (Gustation)
• Five primary tastes
• Salty
• Stimulated by chemical salts,
especially NaCl
• Sour
• Caused by acids which contain a
free hydrogen ion, H+
• Sweet
• Evoked by configuration of glucose
• Bitter
• Brought about by more chemically
diverse group of tastants
• Examples – alkaloids, toxic plant
derivatives, poisonous substances
• Umami
• Meaty or savory taste (MSG
receptor!)
48
Taste (Gustation)
• Taste Perception
• Influenced by information
derived from other receptors,
especially odor
• Temperature and texture of food
influence taste
• Psychological experiences
associated with past
experiences with food influence
taste
• How cortex accomplishes
perceptual processing of taste
sensation is currently unknown
49
Smell (Olfaction)
• Olfactory receptors in nose
are specialized endings of
renewable afferent neurons
• Olfactory mucosa
• 3cm2 of mucosa in ceiling of
nasal cavity
• Contains three cell types
• Olfactory receptor cell
• Afferent neuron whose
receptor portion is in olfactory
mucosa in nose and afferent
axon traverses into brain
• Axons of olfactory receptor
cells collectively form olfactory
nerve
• Supporting cells
• Secrete mucus
• Basal cells
• Precursors of new olfactory
receptor cells (replaced about
every two months)
Brain
Olfactory bulb
Bone
Soft palate
Olfactory tract
Nasal cavity
Olfactory
bulb
Afferent
nerve fibers
(olfactory
nerve)
Basal
cell
Olfactory
receptor
cell
Supporting
cell
Cilia
Olfactory
mucosa
Mucus
layer 50
Smell (Olfaction)
• Odorants
• Molecules that can be smelled
• Act through second-messenger systems
to open Na+ channels and trigger action
potentials
• To be smelled, substance must be
• Sufficiently volatile that some of its
molecules can enter nose in inspired air
• Sufficiently water soluble that it can
dissolve in mucus coating the olfactory
mucosa
• 5 million olfactory receptors
• 1000 different types
• Afferent signals are sorted according
to scent component by glomeruli
within olfactory bulb
• Two routes to the brain
• Subcortical (limbic system)
• Through the thalamus to the cortex
• The olfactory system adapts quickly
and odorants are rapidly cleared (by
odor-eating enzymes)
Brain
Olfactory bulb
Glomeruli
Mitral cells
To limbic system
and cerebral cortex
Cilia Olfactory receptors
51
Smell (Olfaction)
• Vomeronasal Organ (VNO)
• Common in mammals but until
recently was thought to
nonexistent in humans
• Governs emotional responses
and sociosexual behaviors
• Located about half an inch
inside human nose next to
vomer bone
• Detects pheromones
• Nonvolatile chemical signals
passed subconsciously from
one individual to another
• Role in human behavior has not
been validated
• “Good chemistry” and “love at
first sight”
52
PNS – Efferent Division
• Communication link by which CNS controls
activities of muscles and glands
• Two divisions of PNS
• Autonomic nervous system (ANS)
• Involuntary branch of PNS
• Innervates cardiac muscle, smooth muscle, most exocrine glands,
some endocrine glands, and adipose tissue
• Somatic nervous system
• Subject to voluntary control
• Innervates skeletal muscle
53
Autonomic Nervous System
• Autonomic nerve pathway
• Extends from CNS to an innervated organ
• Two-neuron chain
• Preganglionic fiber (synapses with cell body of second neuron)
• Postganglionic fiber (innervates effector organ)
• Postganglionic fibers end in varicosities
• Two subdivisions
• Sympathetic nervous system
• Parasympathetic nervous system
Central nervous
system
Preganglionic fiber
Preganglionic
neurotransmitter
Preganglionic fiber
Varicosity
Preganglionic
neurotransmitter
Effector organ
Autonomic ganglion 54
Autonomic Nervous System
Postganglionic fibers
release acetylcholine
Most postganglionic
fibers release
noradrenaline
(norepinephrine)
Preganglionic fibers
release acetylcholine
(Ach)
Preganglionic fibers
release
acetylcholine (Ach)
Very short
postganglionic fibers
Long postganglionic
fibers
Preganglionic fibers
are longer
Most preganglionic
fibers are short
Fibers originate from
cranial and sacral
areas of CNS
Fibers originate in
thoracic and lumbar
regions of spinal
cord
Parasympathetic
Nervous System
Sympathetic
Nervous System
Craniosacral
parasym-
pathetic
nerves
Terminal
ganglion
Collateral
ganglion
Adrenal
medulla Blood
E,NE
NE
NE
ACh
ACh
Terminal
ganglion
ACh
Sympathetic ganglion chain
Thoracolumbar
sympathetic
nerves
Spinal
cord
Brain
ACh
ACh
ACh
Effector
organs
Cardiac
muscle
Smooth
muscle
Most
exocrine
glands
and
some
endocrine
glands
55
Autonomic Nervous System
• Most visceral organs innervated
by both sympathetic and
parasympathetic fibers
• In general produce opposite
effects in a particular organ
• Dual innervation of organs by
both branches of ANS allows
precise control over organ’s
activity
• Sympathetic system dominates
in emergency or stressful
(“fight-or-flight”) situations
• Promotes responses that prepare
body for strenuous physical activity
• Parasympathetic system
dominates in quiet, relaxed
(“rest-and-digest”) situations
• Promotes body-maintenance
activities such as digestion
56
Autonomic Nervous System
57
Autonomic Nervous System
• Exceptions to general rule of
dual reciprocal innervation by
the two branches of
autonomic nervous system
• Beyond the scope of this course
• What is the one activity that
requires sympathetic /
parasympathetic
coordination?
58
Spinal cord
Sympathetic
preganglionic
fiber
Adrenal
medulla
Blood
Sympathetic
postganglionic
fiber
Target organs
= Acetylcholine
= Norepinephrine
= Epinephrine
Autonomic Nervous System
• Adrenal medulla is a
modified part of
sympathetic nervous
system
• Modified sympathetic
ganglion that does not give
rise to postganglionic fibers
• Stimulation of preganglionic
fiber prompts secretion of
hormones into blood
• About 20% of hormone
release is norepinephrine
• About 80% of hormone
released is epinephrine
(adrenaline)
• Reinforces the activity of the
sympathetic response
• More long-acting and
sustained
59
Autonomic Nervous System
• Tissues innervated by autonomic
nervous system have one or more of
several different receptor types for
postganglionic chemical messengers
• Cholinergic receptors – bind to ACh
• Nicotinic receptors (bind nicotine)
• Found on postganglionic cell bodies of all
autonomic ganglia
• Opens cation channels Æ Na+ flow is higher
Æ AP
• Muscarinic receptors (bind mushroom
poison)
• Found on effector cell membranes (e.g.
smooth muscle, cardiac muscle, glands)
• Several (five) types
Amanita muscaria
60
Autonomic Nervous System
• Andrenergic receptors – bind to
norepinephrine and epinephrine
• Alpha (α) receptors (α1, α2)
• α1 Æ excitatory
• In most sympathetic target tissues
• E.g. Constriction of skin and GI
arterioles, dilation of pupils, etc.
• α2 Æ inhibitory
• Decreased motility in digestive tract
• Beta (β) receptors (β1, β2)
• β1 Æ excitatory
• Primarily in the heart (increased heart
rate and force of contraction)
• β2 Æ inhibitory
• Dilation of skeletal muscle arterioles and
bronchioles
• β3 Æ ιin adipose tissue
• Lipolysis
61
M3
M3
-constriction
-contraction
α1
-dilation
Iris
Ciliary muscle
Eye
α, β2
β3
β1
-glycogenolysis
-lipolysis
-renin release
Liver
Adipose
Kidney
Metabolism
M
α
-↑ secretion
-↑ secretion
Sweat
-Thermoregulation
-Stress
-↑ watery secretion
α1
-↑viscous secretion (small amounts)
Salivary
Glands
M3
M3
M
-contraction
-relaxation
-erection
β2
α2
α
-relaxation
-contraction
-ejaculation
Genitourinary tract
-bladder wall
-sphincter
-penis
M3
M3
-↑ motility
-relaxes
α2 & β2
α1
-↓ motility
-contraction
G.I. tract
-walls
-sphincters
M3
-constriction
β2
-dilation
Bronchi
α
-contraction
Spleen
M
-dilation
β2
α
α
-dilation
-constriction
-constriction
Blood vessels
-skeletal m.
-skin
-penis and clitoris
Smooth muscle
M2
M2
-↓ rate (atria only)
-↓ contractility (atria only)
β1
β1
-↑ rate
-↑ contractility
Cardiac muscle
Heart
-maintains homeostasis
-conserves energy
-stress response (fight or flight)
-expends energy
General
Homeostasis
Receptor
Action
Receptor
Action
Paraympathetic
Sympathetic
62
Autonomic Nervous System
• Agonists
• Bind to same receptor as
neurotransmitter
• Elicit an effect that mimics that
of neurotransmitter
• E.g.
• Salbutamol
• Activates β2 receptors
• Treatment of asthma
• Phenylephrine
• Stimulates both α1 & α2
receptors
• Vasoconstrictor
• Used as nasal decongestant
• Pilocarpine
• Stimulates muscarinic
receptors
• Useful for both narrow and
wide angle glaucoma
• Side effects include sweating.
63
Autonomic Nervous System
• Antagonists
• Bind with receptor
• Block neurotransmitter’s
response
• E.g.
• Succinylcholine
• Binds to the nicotinic receptor
• Causes prolonged
depolarization marked first by
muscle fasciculations followed
by flaccid paralysis
• Atenolol
• Selective β1 blocker
• Blockage produces
bradycardia and decrease in
blood pressure
64
Autonomic Nervous System
• Regions of CNS Involved in Control of Autonomic
Activities
• Prefrontal association complex
• Influences through its involvement with emotional expression
characteristic of individual’s personality (e.g. blushing)
• Hypothalamus
• Plays important role in integrating autonomic, somatic, and endocrine
responses that automatically accompany various emotional and
behavioral states (e.g. anger or fear)
• Medulla (within brain stem)
• Region directly responsible for autonomic output (cardiovascular,
respiratory, digestive tract)
• Spinal Cord
• Some autonomic reflexes, such as urination, defecation, and erection,
are integrated at spinal cord level but control by higher levels of
consciousness
65
Next Lecture …
Neuromuscular Physiology
(240-249, 253-267,270-286,288-297)
Excluded: muscle length, tension, contraction and velocity,
phosphorylation of myosin

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Lecture 4(0).pdf

  • 1. The Cyprus International Institute for the Environment and Public Health In collaboration with the Harvard School of Public Health Constantinos Pitris, MD, PhD Assistant Professor, University of Cyprus cpitris@ucy.ac.cy http://www.eng.ucy.ac.cy/cpitris/courses/CIIPhys/ Lecture 4 The Peripheral Nervous System (181-240) Excluded: adaptation of pacinian corpuscle (185), labeled lines (186), acuity (186-187), phototransduction (200-203), on- off- center ganglion cells (205) 2 Introduction • Peripheral Nervous System • Afferent Division • Sends information from the PNS to the CNS • Efferent Division • Send information from the CNS to the PNS • Afferent Division • Visceral afferents (subconscious input) • Pressure, O2, temperature, etc. • Sensory afferents (conscious input) • Somatic sensation • Somesthetic sensation from skin • Proprioception from muscle joints, skin and inner ear • Special senses • Vision, hearing, taste and smell • Efferent Division • Autonomic Nervous System • Cardiac muscle, smooth muscle, most exocrine glands, some endocrine glands, adipose tissue • Somatic Nervous system • Skeletal muscle 3 Introduction • Sensation ≠ Perception • Perception • Our understanding (conscious interpretation) of the physical world • An interpretation of the senses • Different from what is out there because • Our receptors detect limited number of existing energy forms • The information does not reach our brain unaltered. Some features are accentuated and some are suppressed • The brain interprets the information and often distorts it (“completes the picture” or “feels in the gaps”) to extract conclusions. • Interpretation is affected by cultural, social and personal experiences stored in our memory 4 Receptor Physiology • Receptors • Detect stimulus (detectable change) from different modalities (energy forms) • e.g. light, heat, sound, pressure, chemical changes • Adequate stimulus = the stimulus to which the receptor is most sensitive • Convert forms of energy into electrical signals (action potentials) • Process is called transduction • Types of receptors • Photoreceptors • Responsive to visible wavelengths of light • Mechanoreceptors • Sensitive to mechanical energy • Thermoreceptors • Sensitive to heat and cold • Osmoreceptors • Detect changes in concentration of solutes in body fluids and resultant changes in osmotic activity • Chemoreceptors • Sensitive to specific chemicals • Include receptors for smell and taste and receptors that detect O2 and CO2 concentrations in blood and chemical content of digestive tract • Nociceptors • Pain receptors that are sensitive to tissue damage or distortion of tissue
  • 2. 5 Receptor Physiology • Receptors may be • Specialized ending of an afferent neuron Æ receptor potential • Separate cell closely associated with peripheral ending of a neuron Æ generator potential • Potential generation • Separate receptor • Stimulus causes release of chemical messenger • Specialized afferent nerve ending • Stimulus alters receptor’s permeability which leads to graded receptor potential • Usually causes nonselective opening of all small ion channels Æreceptor (generator) potentials. • The magnitude of the receptor potential represents the intensity of the stimulus. • A receptor potential of sufficient magnitude can produce an action potential. • This action potential is propagated along an afferent fiber to the CNS. Chemically gated channels Afferent neuron fiber Diffusion chemical messenger Receptor (separate cell) Afferent neuron fiber Voltage-gated channels Receptor (separate cell) 6 Receptor Physiology • May adapt slowly or rapidly to sustained stimulation • Types of receptors according to their speed of adaptation • Tonic receptors • Do not adapt at all or adapt slowly • Muscle stretch receptors, joint proprioceptors (to continuously receive information regarding posture and balance) • Phasic receptors • Rapidly adapting receptors • Tactile receptors in skin (the reason you don’t “feel” your clothes or watch) • Adaptation is not the same as habituation (synapse changes in the CNS) Stimulus off Slowly adapting Stimulus on Time Receptor potential (mV) Stimulus strength Stimulus off Rapidly adapting Stimulus on Time Receptor potential (mV) Stimulus strength Off response 7 Pain • Pain • Primarily a protective mechanism meant to bring a conscious awareness that tissue damage is occurring or is about to occur • Storage of painful experiences in memory helps us avoid potentially harmful events in future • Sensation of pain is accompanied by motivated behavioral responses and emotional reactions • Subjective perception can be influenced by other past or present experiences (Are you afraid of your dentist?) 8 Pain • Three categories of nociceptors • Mechanical nociceptors • Respond to mechanical damage such as cutting, crushing, or pinching • Thermal nociceptors • Respond to temperature extremes • Polymodal nociceptors • Respond equally to all kinds of damaging stimuli • Presence of prostaglandins (released after tissue injury) • Lowers nociceptors threshold for activation • Greatly enhances receptor response to noxious stimuli • Aspirin-like drugs inhibit their synthesis Æ analgesic effect • Nociceptors do not adapt to sustained or repetitive stimulation
  • 3. 9 Pain • Characteristics of pain Provoked and sustained by release of bradykinin Occurs second, persists for longer time, more unpleasant Occurs first Poorly localized Easily localized Produces dull, aching, burning sensation Produces sharp, prickling sensation Carried by small, unmyelinated C fibers Carried by small, myelinated A-delta fibers Occurs on stimulation of polymodal nociceptors Occurs on stimulation of mechanical and thermal nociceptors Slow Pain Fast Pain 10 Pain • Two best known pain neurotransmitters • Substance P • Glutamate • Substance P • Activates ascending pathways that transmit nociceptive signals to higher levels for further processing Somatosensory cortex Thalamus Hypothalamus limbic system Reticular formation Noxious stimulus Afferent pain fiber Substance P Spinal cord Brain stem Higher brain (Location of pain) (Perception of pain) ( Alertness) (Behavioral and emotional responses to pain) Nociceptor Other cortical areas (Experience and deliberation of pain) 11 Pain • Glutamate • Major excitatory neurotransmitter • Causes hypersensitivity in the area • Protection mechanism for healing and against further damage • How does a sunburn feel? 12 Pain • Brain has built in analgesic system • Suppresses transmission in pain pathways as they enter spinal cord • Suppress release of Substance P • Depends on presence of opiate receptors • Endogenous opiates (morphine like substances) – endorphins, enkephalins, dynorphin • Factors which modulate pain • Exercise (“runner’s high”) • Stress (survival mechanism) • Acupuncture Periagueductal gray matter Opiate receptor Noxious stimulus Afferent pain fiber Substance P No perception of pain To thalamus Transmission of pain impulses to brain blocked Nociceptor Reticular formation Endogenous opiate
  • 4. 13 Vision • Eye • Sensory organ for vision • Mechanisms that help protect eyes from injury • Eyeball is sheltered by bony socket in which it is positioned • Eyelids • Act like shutters to protect eye from environmental hazards • Eyelashes • Trap fine, airborne debris such as dust before it can fall into eye • Tears • Continuously produced by lacrimal glands • Lubricate, cleanse, bactericidal Canal for tear drainage Pupil Iris Sclera Lacrimal Glands (under eyelid) 14 Vision • Eye • Spherical, fluid-filled structure enclosed by three tissue layers • Sclera/cornea • Sclera – tough outer layer of connective tissue; forms visible white part of the eye • Cornea – anterior, transparent outer layer through which light rays pass into interior of eye • Choroid/ciliary body/iris • Choroid - middle layer underneath sclera which contains blood vessels that nourish retina • Choroid layer is specialized anteriorly to form ciliary body and iris • Retina • Innermost coat under choroid • Consists of outer pigmented layer and inner nervous-tissue layer • Rods and cones Suspensory ligament Ciliary body Conjunctiva Iris Pupil Lens Cornea Aqueous humor Vitreous humor Blood vessels Optic disc Optic nerve Fovea Sclera Retina Choroid Extrinsic eye muscle 15 Vision • Eye • Pupil • Round opening through which light enters the eye • Iris • Controls amount of light entering eye • Contains two sets of smooth muscle networks • Circular (or constrictor) muscle • Radial (or dilator) muscle • Pigment in iris is responsible for eye color • Unique for each individual • Basis for latest identification technology Parasympathetic stimulation + Pupillary constriction Circular (constrictor) muscle runs circularly Radial muscle of iris Circular muscle of iris Pupil Iris Radial (dilator) muscle runs radially Pupillary dilation Sympathetic stimulation + 16 Vision • Convex structures of eye produce convergence of diverging light rays that reach eye • Two structures most important in eye’s refractive ability are • Cornea • Contributes most extensively to eye’s total refractive ability • Refractive ability remains constant because curvature never changes • Lens • Refractive ability can be adjusted by changing curvature as needed for near or far vision (accommodation)
  • 5. 17 Vision • Accommodation • Change in strength and shape of lens • Accomplished by action of ciliary muscle and suspensory ligaments • Age-related reduction in accommodation ability - presbyopia Suspensory ligaments Ciliary muscle Lens Pupillary opening in front of lens Sympathetic stimulation Iris Cornea Flattened weak lens Relaxed ciliary muscle Taut suspensory ligaments Sympathetic stimulation Rounded strong lens Contracted ciliary muscle Slackened suspensory ligaments 18 Vision Far source Near source No accommodations Accommodations Normal eye (Emmetropia) Far source focused on retina without accommodation Near source focused on retina with accommodation No accommodations No accommodations Nearsightedness (Myopia)– Eyeball too long or lens too strong 1. Uncorrected Far source focused in front of retina (where retina would be in eye of normal length) Focus 1. Image out of focus Near source focused on retina with accommodations Farsightedness (Hyperopia)– Eyeball too short or lens too weak 1. Uncorrected Far source focused on retina with accommodations 1. Near source focused behind retina even with accommodations Accommodations Accommodations Image out of focus Focus 19 Vision • Retina • Several layers of cells • Receptor containing portion is actually an extension of the CNS • Fovea • Pinhead-sized depression in exact center of retina • Point of most distinct vision • Has only cones Optic nerve Retina Direction of light Direction of retinal visual processing Front of retina Fibers of the optic nerve Ganglion cell Amacrine cell Bipolar cell Cone Rod Photoreceptor cells Horizontal cell Retina Pigment layer Choroid layer Sclera Back of retina 20 Vision • Photoreceptors • Rod and cone cells • Photopigments on the disk membranes • Rod Æ one type • one pigment, high sensitivity • Cones Æ three different types • Red, green, blue sensing pigments, lower sensitivity • Undergo chemical alterations when activated by light • Change the receptor potential • Induce action potentials • Unlike other receptors, photoreceptors hyperpolarize! Back of retina Outer segment Outer segment Inner segment Synaptic terminal Synaptic terminal Inner segment Direction of light Cells of pigment layer Cone Rod Discs Mitochondria Nuclei Dendrites of bipolar cells Front of retina
  • 6. 21 Vision • Color Vision • Perception of color • Depends on the ratio of stimulation of three different cones • Different absorption of cone pigments • Coded and transmitter by different pathways • Processed in color vision center of primary visual cortex • Color blindness • Defective cone • Colors become combinations of two cones • Most common = red-green color blindness Color perceived Blue cone Green cone Red cone Visible spectrum Wavelength of light (nm) 22 Vision • Properties of Rod and Cone Vision Day vision (lack sensitivity and convergence) Night vision (from sensitivity and convergence) Little convergence in retinal pathways Much convergence in retinal pathways High acuity Low acuity Concentrated in fovea More numerous in periphery Low sensitivity to light High sensitivity to light Color vision Vision in shades of gray 3 million per retina 100 million per retina Cones Rods 23 Vision • The sensitivity of the eyes varies through dark and light adaptation • Dark adaptation • Can gradually distinguish objects as you enter a dark area. • Due to the regeneration of rod photopigments that had been broken down by previous light exposure. • Light adaptation • Can gradually distinguish objects as you enter an area with more light. • Due to the rapid breakdown of cone photopigments. • Along with pupillary reflexes increase the range of vision 24 Vision
  • 7. 25 Vision • Visual field • Area which can be seen without moving the head) Æ overlap between eyes • Visual Pathway • Optic nerve • Optic chiasm • Thalamus • Sorts information to appropriate areas for processing • Optic radiation • Primary visual cortex (occipital lobe) • Higher processing areas • Information arrives altered at the primary visual cortex • Upside down and backward because of the lens • The left and right halves of the brain receive information from the left and right halves of the visual field Left Right Left eye Right eye Optic nerve Optic chiasm Optic tract Lateral geniculate nucleus of thalamus Optic radiation Optic lobe 1 2 3 26 Vision • >30% of cortex participates in visual information processing • “What” and “where” pathways • Depth Perception • Visual field of two eyes slightly different • Depth perception with one eye • Other cues (such as size, location, experience) 27 Hearing • Hearing • Neural perception of sound energy • Involves two aspects • Identification of the sounds (“what”) • Localization of the sounds (“where”) • Sound waves • Traveling vibrations of air • Consist of alternate regions of compression and rarefaction of air molecules Region of compression Region of rarefaction 28 Hearing • Characteristics of sound • Pitch (tone) of sound • Depends on frequency of air waves • Intensity (loudness) • Depends on amplitude of air waves • Timbre (quality) • Determined by overtones
  • 8. 29 Hearing • Ear • Consists of three parts • External ear • Consists of pinna, external auditory meatus, and tympanum • Transmits airborne sound waves to fluid-filled inner ear • Amplifies sound energy • Middle ear • Transmits airborne sound waves to fluid-filled inner ear • Amplifies sound energy • Inner ear • Houses two different sensory systems • Cochlea (Contains receptors for conversion of sound waves into nerve impulses which makes hearing possible) • Vestibular apparatus (Necessary for sense of equilibrium) Inner ear Pinna of external ear Tympanic membrane (eardrum) Middle ear External ear Auditory ossicles Semicircular canals Utricle and saccule Oval window Vestibulocochlear nerve Cochlea Round window Eustachian tube To pharynx External auditory meatus (ear canal) 30 Malleus Incus Stapes at oval window Helicotrema Cochlea Scala vestibuli Scala tympani Round window Middle ear cavity Tympanic membrane External auditory meatus Vestibular membrane Basilar membrane Tectorial membrane Organ of Corti Scala media Hearing • Sound Wave Transmission • Tympanic membrane • Vibrates when struck by sound waves • Middle ear • Transfers vibrations through ossicles (malleus, incus, stapes) to oval window (entrance into fluid- filled cochlea) • Amplify the pressure 20x • Large surface of the tympanic membrane transferred to the smaller oval window • Lever action of the osscicles • Small muscles change the stifness of the tympanic membrane • Protection mechanism • Slow action (40 msec) Æ protects only from prolonged sounds 31 Malleus Incus Stapes at oval window Helicotrema Cochlea Scala vestibuli Scala tympani Round window Middle ear cavity Tympanic membrane External auditory meatus Vestibular membrane Basilar membrane Tectorial membrane Organ of Corti Scala media Hearing • Sound Wave Transmission • Inner ear • Sound dissipates in cochlea • Waves in cochlear fluid (endolymph) set basilar membrane in motion • Sound converted to electrical signals by the Organ of Corti 32 Hearing • Sound Transduction • Organ of Corti • Hair cells with ~ 100 sterocilia each • Inner hair cells are tilted as basilar membrane oscillates • Mechanically gated channels open and close Æ graded potentials • Communicate via chemical signals with the nerves which form the auditory nerve • Outer hair cells Æ adjust length with respond to electrical stimulus (electromotility) Æ accentuate motion of basilar membrane and fine tune response Vestibular membrane Tectorial membrane Scala media (cochlear duct) Basilar membrane Scala tympani Auditory nerve Scala vestibuli Outer hair cells Hairs (stereocilia) Tectorial membrane Inner hair cells Nerve fibers Supporting cell Basilar membrane
  • 9. 33 Hearing • Auditory pathway • Hair cells • Auditory nerve • Brain stem • Signals cross over to opposite site (unlike visual signals) • Thalamus • Sorts and relays signals to cortical regions • Cortex • Cortical Processing • Primary auditory cortex in the temporal lobe • Tonotopically mapped (i.e. mapped according to tone) • Higher cortical processing • Separates coherent and meaningful patterns 34 Hearing • Pitch discrimination • Basilar membrane does not have uniform mechanical properties • Narrow and stiff to wide and flexible • Different regions vibrate maximally at different frequencies • Frequency (or frequencies) are discriminated by the location of hair cells firing Wide, flexible end of basilar membrane near helicotrema Narrow, stiff end of basilar membrane near oval window High frequency Medium frequency Low frequency 35 Hearing • Loudness discrimination • Exquisitely sensitive organ (motion less than a molecule of Hydrogen) Æ easily damaged • Wide range (every 10 dB means 10-fold increase in intensity) • Higher intensity causes larger basilar membrane movement • Stronger graded potential of hair cells • Faster rate of action potentials from auditory nerve cells • Anything > 100 dB can cause permanent damage 1 quadrillion x 150 dB Takeoff of jet plane 1 trillion x 120 dB Loud rock concert 1 billion x 90 dB Food Blender 1 million x 60 dB Normal conversation 1000 x 30 dB Hush of Library 100 x 20 dB Ticking of watch 10 x louder 10 dB Rustle of leaves Comparison to faintest audible sound (hearing threshold) Loudness in decibels (dB) Sound 36 Hearing • Localization • Up-Down localization (elevation) • External ear (Pinna) shape changes sound timbre and intensity slightly according to elevation • Left-right localization (azimuth) • Sound arriving to proximal ear arrives • Slightly earlier (~ 0.5 msec) • Slightly stronger • Brain uses the electrical activity changes to these two cues to localize the direction
  • 10. 37 Hearing • Deafness • Conductive • Sound waves not adequately conducted through external and middle portions of ear • Blockage, rupture of ear drum, middle ear infection, iddle ear adhesions • Hearing aids might help • Senosineural • Sound waves conducted but not translated into electrical signals • Neural presbycusis, certain antibiotics, poisoning • Cochlear implants might help • Electrical devices stimulating the auditory nerve directly 38 Equilibrium • Detection of position and motion • Posture and coordination • Vestibular apparatus • Fluid filled tunnels In the inner ear • Semicircular canals • Three circular tunnels arranged on perpendicular planes • Otolith organs (Utricle and saccule) • Two bulges arranged in perpendicular directions 39 Equilibrium • Semicircular canals • Three circular tunnels arranged on perpendicular planes • Detect rotational acceleration or deceleration in any direction • Hair cells • On a ridge in the ampulla • Have on kinocilium and several sterocilia (mikrovilli) • Embedded in gelatinous material, the cupula Semicircular canals Perilymph Endolymph Ampulla Oval window Round window Cochlea Vestibular apparatus Utricle Saccule Vestibular nerve Auditory nerve Hair cell Support cell Ridge in ampulla Vestibular nerve fibers Hairs of hair cell; kinocilium (red) and stereocilia (blue) Cupula 40 Direction of head rotation Left horizontal semicircular canal Direction of fluid movement in semicircular canals Direction of bending of cupula and hairs of receptor hairs cells Right horizontal semicircular canal Equilibrium • Semicircular canals • Signal transduction • Head is rotated • Two of the canals are rotated around their axis in opposite directions • Endolymph moves opposite to the direction of motion (inertia) • Cupula leans in that direction • Cilia bent and K+ channels open or close • The haircells are depolarized or hyperpolarized • Neurotransmitter realize from the hair cells is modified • Firing of the vestibular nerve is modified Kinocilium Stereocilia Hair cell
  • 11. 41 Equilibrium • Vestibular pathway • Vestibular nuclei in brain stem • Motor neurons for controlling eye movement, perceiving motion and orientation • E.g. vestibuloocular reflex • Cerebellum for use in maintaining balance and posture, • The vestibular system detects acceleration • Speed is calculated by integrating circuitry in the brain stem Head motion Eye Movement Endolymph Motion 42 Equilibrium • Otolith Organs • Detect changes in rate of linear movement in any direction • Arranged in perpendicular directions • Provide information important for determining head position in relation to gravity • Hair cells • As described before • In addition, calcium carbonate crystals (otoliths) are embedded in within the gelatinous layer • Increased inertia • Sensitivity to gravity 43 Equilibrium • Otolith Organs • Signal transduction • Utricle Æ Forward or backward motion or tilt (motion due to gravitational force) • Saccule Æ vertical motion • Endolymph and gelatinous mass with otoliths move in the opposite direction • Cilia bent and K+ channels open or close • The haircells are depolarized or hyperpolarized • Neurotransmitter realize from the hair cells is modified • Firing of the vestibular nerve is modified Gravitational force 44 Taste and Smell • Taste (gustation) and smell (olfaction) • Receptors are chemoreceptors • In association with food intake, influence flow of digestive juices and affect appetite • Stimulation of receptors induces pleasurable or objectionable sensations and signals presence of something to seek or to avoid • In lower animals also play a role in finding direction, seeking prey, avoiding predators and sexual attraction to a mate • Less developed and important in humans • Really? How much do you spend on perfumes and colognes
  • 12. 45 Taste (Gustation) • Chemoreceptors housed in taste buds • Present in oral cavity and throat • Taste receptors have life span of about 10 days • Taste bud consists of • Taste pore • Opening through which fluids in mouth come into contact with surface of receptor cells • Taste receptor cells • Modified epithelial cells with surface folds called microvilli • Plasma membrane of microvilli contain receptor sites that bind selectively with chemical molecules Taste bud Sensory nerve fiber Papilla Surface of the tongue Taste pore Taste receptor cell Supporting cell Tongue 46 Taste (Gustation) • Signal transduction • Tastant (taste-provoking chemical) • Binding of tastant with receptor cell • Alters cell’s ionic channels to produce depolarizing receptor potential • Receptor potential releases neurotransmitter • Initiates action potentials within terminal endings of afferent nerve fibers with which receptor cell synapses • Signals conveyed via synaptic stops in brain stem and thalamus to cortical gustatory area 47 Taste (Gustation) • Five primary tastes • Salty • Stimulated by chemical salts, especially NaCl • Sour • Caused by acids which contain a free hydrogen ion, H+ • Sweet • Evoked by configuration of glucose • Bitter • Brought about by more chemically diverse group of tastants • Examples – alkaloids, toxic plant derivatives, poisonous substances • Umami • Meaty or savory taste (MSG receptor!) 48 Taste (Gustation) • Taste Perception • Influenced by information derived from other receptors, especially odor • Temperature and texture of food influence taste • Psychological experiences associated with past experiences with food influence taste • How cortex accomplishes perceptual processing of taste sensation is currently unknown
  • 13. 49 Smell (Olfaction) • Olfactory receptors in nose are specialized endings of renewable afferent neurons • Olfactory mucosa • 3cm2 of mucosa in ceiling of nasal cavity • Contains three cell types • Olfactory receptor cell • Afferent neuron whose receptor portion is in olfactory mucosa in nose and afferent axon traverses into brain • Axons of olfactory receptor cells collectively form olfactory nerve • Supporting cells • Secrete mucus • Basal cells • Precursors of new olfactory receptor cells (replaced about every two months) Brain Olfactory bulb Bone Soft palate Olfactory tract Nasal cavity Olfactory bulb Afferent nerve fibers (olfactory nerve) Basal cell Olfactory receptor cell Supporting cell Cilia Olfactory mucosa Mucus layer 50 Smell (Olfaction) • Odorants • Molecules that can be smelled • Act through second-messenger systems to open Na+ channels and trigger action potentials • To be smelled, substance must be • Sufficiently volatile that some of its molecules can enter nose in inspired air • Sufficiently water soluble that it can dissolve in mucus coating the olfactory mucosa • 5 million olfactory receptors • 1000 different types • Afferent signals are sorted according to scent component by glomeruli within olfactory bulb • Two routes to the brain • Subcortical (limbic system) • Through the thalamus to the cortex • The olfactory system adapts quickly and odorants are rapidly cleared (by odor-eating enzymes) Brain Olfactory bulb Glomeruli Mitral cells To limbic system and cerebral cortex Cilia Olfactory receptors 51 Smell (Olfaction) • Vomeronasal Organ (VNO) • Common in mammals but until recently was thought to nonexistent in humans • Governs emotional responses and sociosexual behaviors • Located about half an inch inside human nose next to vomer bone • Detects pheromones • Nonvolatile chemical signals passed subconsciously from one individual to another • Role in human behavior has not been validated • “Good chemistry” and “love at first sight” 52 PNS – Efferent Division • Communication link by which CNS controls activities of muscles and glands • Two divisions of PNS • Autonomic nervous system (ANS) • Involuntary branch of PNS • Innervates cardiac muscle, smooth muscle, most exocrine glands, some endocrine glands, and adipose tissue • Somatic nervous system • Subject to voluntary control • Innervates skeletal muscle
  • 14. 53 Autonomic Nervous System • Autonomic nerve pathway • Extends from CNS to an innervated organ • Two-neuron chain • Preganglionic fiber (synapses with cell body of second neuron) • Postganglionic fiber (innervates effector organ) • Postganglionic fibers end in varicosities • Two subdivisions • Sympathetic nervous system • Parasympathetic nervous system Central nervous system Preganglionic fiber Preganglionic neurotransmitter Preganglionic fiber Varicosity Preganglionic neurotransmitter Effector organ Autonomic ganglion 54 Autonomic Nervous System Postganglionic fibers release acetylcholine Most postganglionic fibers release noradrenaline (norepinephrine) Preganglionic fibers release acetylcholine (Ach) Preganglionic fibers release acetylcholine (Ach) Very short postganglionic fibers Long postganglionic fibers Preganglionic fibers are longer Most preganglionic fibers are short Fibers originate from cranial and sacral areas of CNS Fibers originate in thoracic and lumbar regions of spinal cord Parasympathetic Nervous System Sympathetic Nervous System Craniosacral parasym- pathetic nerves Terminal ganglion Collateral ganglion Adrenal medulla Blood E,NE NE NE ACh ACh Terminal ganglion ACh Sympathetic ganglion chain Thoracolumbar sympathetic nerves Spinal cord Brain ACh ACh ACh Effector organs Cardiac muscle Smooth muscle Most exocrine glands and some endocrine glands 55 Autonomic Nervous System • Most visceral organs innervated by both sympathetic and parasympathetic fibers • In general produce opposite effects in a particular organ • Dual innervation of organs by both branches of ANS allows precise control over organ’s activity • Sympathetic system dominates in emergency or stressful (“fight-or-flight”) situations • Promotes responses that prepare body for strenuous physical activity • Parasympathetic system dominates in quiet, relaxed (“rest-and-digest”) situations • Promotes body-maintenance activities such as digestion 56 Autonomic Nervous System
  • 15. 57 Autonomic Nervous System • Exceptions to general rule of dual reciprocal innervation by the two branches of autonomic nervous system • Beyond the scope of this course • What is the one activity that requires sympathetic / parasympathetic coordination? 58 Spinal cord Sympathetic preganglionic fiber Adrenal medulla Blood Sympathetic postganglionic fiber Target organs = Acetylcholine = Norepinephrine = Epinephrine Autonomic Nervous System • Adrenal medulla is a modified part of sympathetic nervous system • Modified sympathetic ganglion that does not give rise to postganglionic fibers • Stimulation of preganglionic fiber prompts secretion of hormones into blood • About 20% of hormone release is norepinephrine • About 80% of hormone released is epinephrine (adrenaline) • Reinforces the activity of the sympathetic response • More long-acting and sustained 59 Autonomic Nervous System • Tissues innervated by autonomic nervous system have one or more of several different receptor types for postganglionic chemical messengers • Cholinergic receptors – bind to ACh • Nicotinic receptors (bind nicotine) • Found on postganglionic cell bodies of all autonomic ganglia • Opens cation channels Æ Na+ flow is higher Æ AP • Muscarinic receptors (bind mushroom poison) • Found on effector cell membranes (e.g. smooth muscle, cardiac muscle, glands) • Several (five) types Amanita muscaria 60 Autonomic Nervous System • Andrenergic receptors – bind to norepinephrine and epinephrine • Alpha (α) receptors (α1, α2) • α1 Æ excitatory • In most sympathetic target tissues • E.g. Constriction of skin and GI arterioles, dilation of pupils, etc. • α2 Æ inhibitory • Decreased motility in digestive tract • Beta (β) receptors (β1, β2) • β1 Æ excitatory • Primarily in the heart (increased heart rate and force of contraction) • β2 Æ inhibitory • Dilation of skeletal muscle arterioles and bronchioles • β3 Æ ιin adipose tissue • Lipolysis
  • 16. 61 M3 M3 -constriction -contraction α1 -dilation Iris Ciliary muscle Eye α, β2 β3 β1 -glycogenolysis -lipolysis -renin release Liver Adipose Kidney Metabolism M α -↑ secretion -↑ secretion Sweat -Thermoregulation -Stress -↑ watery secretion α1 -↑viscous secretion (small amounts) Salivary Glands M3 M3 M -contraction -relaxation -erection β2 α2 α -relaxation -contraction -ejaculation Genitourinary tract -bladder wall -sphincter -penis M3 M3 -↑ motility -relaxes α2 & β2 α1 -↓ motility -contraction G.I. tract -walls -sphincters M3 -constriction β2 -dilation Bronchi α -contraction Spleen M -dilation β2 α α -dilation -constriction -constriction Blood vessels -skeletal m. -skin -penis and clitoris Smooth muscle M2 M2 -↓ rate (atria only) -↓ contractility (atria only) β1 β1 -↑ rate -↑ contractility Cardiac muscle Heart -maintains homeostasis -conserves energy -stress response (fight or flight) -expends energy General Homeostasis Receptor Action Receptor Action Paraympathetic Sympathetic 62 Autonomic Nervous System • Agonists • Bind to same receptor as neurotransmitter • Elicit an effect that mimics that of neurotransmitter • E.g. • Salbutamol • Activates β2 receptors • Treatment of asthma • Phenylephrine • Stimulates both α1 & α2 receptors • Vasoconstrictor • Used as nasal decongestant • Pilocarpine • Stimulates muscarinic receptors • Useful for both narrow and wide angle glaucoma • Side effects include sweating. 63 Autonomic Nervous System • Antagonists • Bind with receptor • Block neurotransmitter’s response • E.g. • Succinylcholine • Binds to the nicotinic receptor • Causes prolonged depolarization marked first by muscle fasciculations followed by flaccid paralysis • Atenolol • Selective β1 blocker • Blockage produces bradycardia and decrease in blood pressure 64 Autonomic Nervous System • Regions of CNS Involved in Control of Autonomic Activities • Prefrontal association complex • Influences through its involvement with emotional expression characteristic of individual’s personality (e.g. blushing) • Hypothalamus • Plays important role in integrating autonomic, somatic, and endocrine responses that automatically accompany various emotional and behavioral states (e.g. anger or fear) • Medulla (within brain stem) • Region directly responsible for autonomic output (cardiovascular, respiratory, digestive tract) • Spinal Cord • Some autonomic reflexes, such as urination, defecation, and erection, are integrated at spinal cord level but control by higher levels of consciousness
  • 17. 65 Next Lecture … Neuromuscular Physiology (240-249, 253-267,270-286,288-297) Excluded: muscle length, tension, contraction and velocity, phosphorylation of myosin