Sensory Systems

CHAPTER
6
Sensory Systems
PowerPoint® Lecture Slides prepared by

Stephen Gehnrich, Salisbury University
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Sensory Receptors
 Range from single cells to complex sense organs

 Types of receptors
 Chemoreceptors, mechanoreceptors, photoreceptors,
electroreceptors, magnetoreceptors, thermoreceptors
 All receptors transduce incoming stimuli into
changes in membrane potential
 Receptor protein detects stimulus
 Opening or closing of ion channel
 Change in membrane potential
 Signal sent to integrating center (central nervous
system)

Sensory Receptors
Figure 6.1
Graded Potentials
Generator potential
 Sensory receptor is also the
primary afferent neuron
 Change in membrane
potential (depolarization)
spreads along membrane
 Triggers AP in axon
Receptor potential
 Sensory receptor is separate
from the afferent neuron
 Change in membrane
potential of sensory
receptor cell triggers
release of neurotransmitter
Classification of Sensory Receptors
Based on stimulus location
 Telereceptors
 Detect distant stimuli
 For example, vision and hearing
 Exteroceptors
 Detect stimuli on the outside of the body
 For example, pressure and temperature
 Interoceptors
 Detect stimuli inside the body
 For example, blood pressure and blood oxygen
 Proprioceptors
 Detect stimuli for the position and orientation of the body
 Receptors in skin, muscles, vestibular apparatus of inner ears

Based on type of stimulus (stimulus modality) the

receptors detect
 Chemoreceptors
 Chemicals
 For example, smell and taste
 Mechanoreceptors
 Pressure and movement
 For example, touch, hearing, balance, blood pressure
 Photoreceptors
 Light
 For example, vision
 Electroreceptors
 Electrical fields
 Magnetoreceptors
 Magnetic fields
 Thermoreceptors
 Temperature

Sensitivity to Multiple Modalities
Adequate stimulus
 Preferred (most sensitive) stimulus modality
Many receptors can be excited by other stimuli, if sufficiently
strong
 For example, pressure on eyelid  perceive light
Polymodal receptors
 Sensitive to more than one stimulus modality
 For example, nociceptors; polymodal receptors for multiple
types of pain; Ampullae of Lorenzini in sharks sense
organs sensitive to touch, electricity, temp.

Stimulus Encoding
 All stimuli are ultimately converted into action

potentials in a primary afferent neuron
 How can organisms differentiate among stimuli or
detect the strength of the signal?
 Sensory receptors and sensory neurons must
encode four types of information
 Stimulus modality
 Stimulus location
 Stimulus intensity
 Stimulus duration

Stimulus Modality and Location
 Receptor location encodes stimulus modality and location
 Integrating center interprets modality and location
 Modality
 Theory of labeled lines
 Brain detects type of stimulus based on the type of
receptor that is stimulated.
 All sensory receptors associated with a single afferent
neuron. Discrete pathway from sensory cell to
integrating center
 Polymodal receptors are exceptions
 Encode modality via temporal patterns of APs
 Location of stimulated receptor
 Theory of labeled lines

Receptive Field and Location of Stimulus
 Receptive field
 Region of the sensory surface that causes a response
when stimulated
 Smaller receptive field allows more precise location of
the stimulus (i.e., greater acuity)
 Improved ability to localize stimuli by
 Using more than one sensory receptor cell and afferent
neurons with overlapping receptive fields ( population
coding)
 Lateral inhibition
 Signals from neurons at the center of the receptive
field inhibit neurons on the periphery
 Enhance contrast
Receptive Field and Location of Stimulus
Figure 6.3
Dynamic Range
 Sensory neurons code stimulus intensity by
changes in action potential frequency
 For example, strong stimuli  high
frequency ( series AP)
 Dynamic range
 Range of stimulus intensities over which a
receptor exhibits an increased response
 Threshold of detection
 Weakest stimulus that produces a
response in a receptor 50% of the
time
Receptor Saturation:
 Below threshold stimulus intensity =
• All ion channels have opened or
no AP
closed
 Saturation • Em for a particular ion
 top of the dynamic range (maximal • Max rate of release of
response) neurotransmitter from receptor cell
 Receptor cell can not increase its • Max frequency of AP in afferent
response neuron
Dynamic Range and Discrimination
 Trade-off between dynamic
range and discrimination
 Large dynamic range
 Large change in stimulus
causes a small change in AP
frequency
 Poor sensory discrimination
 Narrow dynamic range
 Small change in stimulus
causes a large change in AP
frequency
 Good sensory discrimination
within the range
Range Fractionation
 Range fractionation
 Groups of receptors work together to increase
dynamic range without decreasing sensory
discrimination
Figure 6.4c
Encoding Logarithmically
Encode a wide range of stimulus intensities using a single receptor
cell
 Good discrimination at certain intensities (low stimulus
intensities)
 Poor discrimination at other intensities ( high stimulus
intensities)
Figure 6.4d
Tonic and Phasic Receptors
Two classes of receptors encode stimulus
duration:
 Tonic
 Produce APs as long as the
stimulus continues
 Encode duration of stimulus
 Receptor adaptation – AP
frequency decreases if stimulus
intensity is maintained at the
same level
 To focus on novel sensations
 Phasic
 Produce APs at the beginning or
end of the stimulus
 Encode change in stimulus, but
not stimulus duration

Chemoreception
 Most cells can sense chemical stimuli

 Animals have many types of chemoreceptors
 Olfaction (smell)
 Detection of chemicals in air
 Gustation (taste)
 Detection of chemicals emitted from food
 Olfaction and gustation are distinguished by structural
criteria
 Performed by different sense organs
 Different signal transduction mechanisms
 Processed in different integrating centers

The Olfactory System
Vertebrate olfactory system
 Can distinguish thousands of
odorants
 Located in the roof of the nasal
cavity
 Mucus layer to moisten olfactory
epithelium
 Odorant binding proteins
 Found in mucus
 Allow lipophilic odorants to
dissolve in mucus
 Receptor cells are bipolar neurons
with cilia
 Odorant receptor proteins are
located in the cilia
Odorant Receptors
 Each olfactory neuron expresses only one odorant

receptor protein
 There are 1000s of different receptor proteins
 Each receptor can recognize more than one odorant
 Each odorant can stimulate more than one receptor
 Odorant receptor is linked to G-protein
 Odorant binding causes formation of cAMP
 Opening of ion channels
 Depolarization

Odorant Receptors
Odorant receptors coupled to G-protein
That activates phospholipase C (PLC).
Hydrolyses PIP2 in the plasma
membrane to IP3 and DAG, resulting
to increase Ca2+ i Cl- channels open
Figure 6.7
Pheromones
 Vomeronasal organ
 Detects pheromones
 Chemical signals between
animals
 Structurally and molecularly
distinct from the olfactory
epithelium
 Located in base of nasal cavity in
mammals
 Located in palate in reptiles
 Pheromone Receptor is linked to
G-protein
 Activates phospholipase C
transduction system
 Depolarization
Invertebrate Olfactory Systems
 Located in many parts of the body
 Most near the head
 Primarily on antennae in Arthropods
 Olfactory Sensilla
 Hair-like projections of cuticle
 Sensilla contain odorant receptor
neurons
 Odorant receptor is linked to G-
protein
 Odorant binding causes
formation of cAMP
 Depolarization
 Sensilla involved in detection of
pheromones, gustation, senses of
touch and hearing
The Gustatory System
 Five classes of tastants

 Salty
 Sweet
 Bitter
 Sour
 Umami (savory or meaty)
 Sweet, umami, and salty indicate carbohydrates,
proteins, and ions
 Bitter and sour indicate potentially toxic substances

Taste Buds in Vertebrates
Taste receptors are epithelial cells
that release neurotransmitter
 vertebrate taste receptors are
not neurons
Each taste receptor expresses more
than one kind of taste receptor
protein
Taste receptor cells clustered in
groups
 On tongue, soft palate,
larynx, and esophagus
 On external surface of the
body in some fish

Taste Receptor Transduction Pathways
Figure 6.11a,b
Taste Receptor Transduction Pathways
Figure 6.1c,d
Taste in Invertebrates
 Located on sensilla
 Inside and outside the mouth ( proboscis)
 Along the wing margin
 Ends of the legs
 Receptors
 Bipolar sensory neurons
 G-protein coupled
 Express only a single receptor protein
 Differences between vertebrates and invertebrates
suggest independent evolution

Mechanoreceptors
 Transform mechanical stimuli into
electrical signals
 All organisms (and most cells) sense
and respond to mechanical stimuli
 Two main types of mechanoreceptor
proteins:
 ENaC
 Epithelial sodium channels
 TRP channels
 Transient receptor potential
channels
 Channels are linked to extracellular
matrix
 Mechanical stimuli alter channel
permeability

Touch and Pressure
Three classes of receptors

 Baroreceptors
 Interoceptors detect pressure changes
 Tactile receptors
 Exteroceptors detect touch, pressure, and vibration
 Proprioceptors
 Monitor the position of the body

Vertebrate Tactile Receptors
 Widely dispersed in skin
 Receptor structure
 Free nerves endings
 Nerve endings enclosed in
accessory structures
 (e.g., Merkel’s disks are tonic
receptors , Pacinian
corpuscles are phasic
receptors)
 Meissner’s corpuscles-
discriminative touch, localize
tactile sensation

Vertebrate Proprioceptors
 Monitor the position of the body

 Three major groups
 Muscle spindles
 Located in skeletal muscles
 Monitor muscle length
 Golgi tendon organs
 Located in tendons
 Monitor tendon tension
 Joint capsule receptors
 Located in capsules that enclose joints
 Monitor pressure, tension, and movement

Insect Tactile Receptors
Two common types of sensilla
 Trichoid
 Hairlike projection of cuticle
 Bipolar sensory neuron
 TRP channel
 Detects air movements
 Campaniform
 Dome-shaped bulge of
cuticle
 Bipolar sensory neuron
 Detects deformation of
cuticle as insects moves to
make coordinated movement

Insect Proprioceptors
 Scolopidia
 Bipolar neuron and
complex accessory cell
(scolopale)
 Can be isolated or
grouped into chordotonal
organs
 Most function in
proprioception
 Can be modified into
tympanal organs for
sound detection

Equilibrium and Hearing
 Utilize mechanoreceptors
 Equilibrium (“balance”)
 Detect position of the body relative to gravity
 Hearing
 Detect and interpret sound waves
 Vertebrates
 Ear is responsible for equilibrium and hearing
 Invertebrates
 Organs for equilibrium are different from organs of
hearing

Statocysts
 Organ of equilibrium in
invertebrates
 Hollow, fluid filled
cavities lined with
mechanosensory
neurons
 Statocysts contain
statoliths
 Dense particles of
calcium carbonate
 Movement of
statoliths stimulate
mechanoreceptors

Insect Hearing
 Strong vibrations sensed by trichoid sensilla (TRP
channels)
 Weak vibrations and sounds are detected by
chordotonal organs
 Clusters of scolopidia
 Located on leg
 Mechanosensitive ion channels
 Tympanal organs
 Most sensitive ears; located on insect body (abdomen,
legs, thorax, wing base)
 Thin layer of cuticle (tympanum) overlays
chordotonal organ
Vertebrate Hair cells
 Mechanoreceptor for hearing and balance
 Modified epithelial cells (not neurons)
 Cilia on apical surface
 Kinocilium (a true cilium); absent
in ears of adult mammals
 Stereocilia (microvilli)
 Tips of stereocilia are
connected by proteins (tip
links)
 Mechanosensitive ion channels in
stereocilia
 Movement of stereocilia 
change in permeability
 Change in membrane potential
 Change in release of neurotransmitter
from hair cell

Signal Transduction in Hair Cells

Fish and Amphibian Hair Cells
 Hair cells detect body position

and movement
 Neuromast
 Hair cells and cupula
 Stereocilia embedded in
gelatinous cap
 Detect movement of water
 Lateral line system
 Array of neuromasts within
pits or tubes running along
the side of the body

Vertebrate Ears
 Function in both equilibrium and hearing
 Outer ear
 Not in all vertebrates
 Pinna
 Auditory canal
 Middle ear
 Not in all vertebrates
 Interconnected bones in air-filled
cavity
 Inner ear
 Present in all vertebrates
 Series of fluid-filled membranous
sacs and canals
 Contains mechanoreceptors (hair
cells)
 Bony labyrinth: Cochlea ( hearing);
vestibule and semicircular canals (
balance)
Inner Ear: Vestibular Apparatus and Cochlea
 Vestibular apparatus detects
movements
 Three semi-circular canals with
enlarged region at one end
(ampulla)
 Two sacklike swellings (utricle
and saccule)
 Lagena
 Extension of saccule
 Extended in birds and mammals
into a cochlear duct or cochlea for
hearing
 Hair cells present in vestibular
apparatus and lagena (cochlea)

Vestibular Apparatus
 Mechanoreceptors of the inner ear
 Macula
 Present in utricle ( horizontal, side to side
tilting of head; righting reflexes) and
saccule (vertical head/body;
forward/backward tilting )
 Mineralized otoliths suspended in a
gelatinous matrix
 Stereocilia of hair cells embedded in matrix
 >100,000 hair cells
 Detect linear acceleration and tilting of
head
 Cristae
 Present in ampullae of semicircular canals
 Gelatinous matrix (cupula) lacks otoliths
 Stereocilia of hair cells embedded in matrix
 Detect angular acceleration (turning) of
head
Maculae Detect Linear Acceleration
and Tilting

Cristae Detect Angular Acceleration
Figure 6.24
Sound Detection by Inner Ear
Fish
 Sound waves cause otoliths to move
 Displacement of cilia on hair cells
 Some fish use the swim bladder to amplify sounds
Figure 6.25
Sound Detection by Inner Ear
 Terrestrial Vertebrates
 Hearing involves the inner,
middle, and outer ear
 Sound transfers poorly
between air and the fluid-
filled inner ear
 Amplification of sound waves
 Pinna acts as a funnel to
collect more sound
 Middle ear bones increase
the amplitude of vibrations
from the tympanic
membrane to the oval
window

Mammalian Inner Ear
 Specialized for sound detection
 Perilymph
 Fills vestibular and tympanic
ducts
 Similar to extracellular fluids
(high Na+ and low K+)
 Endolymph
 Fills cochlear duct
 Different from extracellular
fluid (high K+ and low Na+)
 Organ of Corti
 Hair cells on basilar membrane
 Inner and outer rows of
hair cells
 Stereocilia embedded in
tectorial membrane in cochlear
duct (filled with endolymph)

Sound Transduction
 Sound waves vibrate tympanic
membrane
 Middle ear bones transmit vibration
to oval window
 Oval window vibrates
 Pressure waves in perilymph of
vestibular duct
 Waves in endolymph Basilar
membrane vibrates
 Stereocilia on the inner hair cells
bend (detect sounds)
 Hair cells depolarize
 Hair cells release neurotransmitter
(glutamate)
 Glutamate excites sensory neuron
Round window serves as a pressure valve
Encoding Sound Frequency
 Frequency ( Pitch) Detection
 Basilar membrane is stiff and
narrow at the proximal end (
close to oval and round
windows) and flexible and
wide at distal end
 High frequency sound vibrates
stiff end
 Low frequency sound vibrates
flexible end
 Specific regions of auditory
cortex brain respond to
specific frequencies
 Place coding

Encoding Sound Amplitude
and Amplification
 Amplitude Detection
 Loud sounds cause larger movement of basilar membrane than quiet
sounds
  depolarization of inner hair cells
  AP frequency
 Outer hair cells amplify quiet sounds
 Change shape of prestin in response to sound waves and
depolarization
 Do not release neurotransmitter
 Change in shape increases movement of basilar membrane
 Increased stimulus to inner hair cells
 Feedback loop ( loud noise) : protective mechanism for inner hair cells
damaged by loud sounds: efferent neuron ( releases acetylcholine ) which
synapses with outer hair cells reduces response of outer hair cells
Detecting Sound Location
 Brain uses time lags and differences in sound

intensity to detect location of sound
 Sound in right ear first
 Sound located to the right
 Sound louder in right ear
 Sound located to the right
 Rotation of head helps localize sound

Photoreception
 Ability to detect visible light

 A small proportion of the electromagnetic spectrum
from ultraviolet to near infrared
 Ability to detect this range of wavelengths supports
idea that animals evolved in water
 Visible light travels well in water; other wavelengths do
not

Electromagnetic Spectrum
Figure 6.27a,b
Photoreceptors
 Range from single light-sensitive cells to
complex, image-forming eyes
 Two major types of photoreceptor cells:
 Ciliary photoreceptors
 Have a single, highly folded
cilium
 Folds form disks that contain
photopigments
 Rhabdomeric photoreceptors
 Apical surface covered with
multiple outfoldings called
microvillar projections
 Microvillar projections contain
photopigments
 Photopigments
 Molecules that absorb energy from
photons
Vertebrate Photoreceptors
 Vertebrates have ciliary

photoreceptors
 Rods
 Cones
 Both have inner and outer
segments
 Inner and outer segments
connected by a cilium
 Outer segment contains
photopigments
 Inner segment forms synapses
with other cells
Characteristics of Rods and Cones
Table 6.1
Diversity in Rod and Cone Shape
 Diverse shapes of rods

and cones among
vertebrates
 Shape does not
determine properties
of photoreceptor
 Properties of
photoreceptor depend
on its photopigment
Figure 6.30
Photopigment Complex
Photopigments have two covalently bonded parts
( rhodopsin, iodopsin, porphyropsin,
melanopsin, VA opsin)
 Chromophore
 Derivative of vitamin A
 For example, retinal
 Contains carbon-carbon double bonds
 Absorption of light converts bond
from cis to trans
 Opsin
 G-protein-coupled receptor protein
 Opsin structure determines
photopigment characteristics
 For example, wavelength of light
absorbed

Phototransduction
 Steps in photoreception
 Chromophore absorbs energy from photon
 Chromophore changes shape
 Double bond isomerizes from cis to trans
 Activated chromophore dissociates from opsin
 “Bleaching”
 Opsin activates G-protein
 Formation of second messenger
 Ion channels open or close
 Change in membrane potential depends on light intensity
 Bright light: decrease cGMP ---- close Na+ channels----hyperpolarization-
 Dim light: slight decrease cGMP---- few close Na+ channels ---
depolarization
 Dark: high cGMP--- open Na+ channels--- depolarization

Phototransduction
Figure 6.32
The Eye
 Eyespots
 Cells or regions of a cell that contain photosensitive
pigment
 For example, protist Euglena
 Eyes are complex organs
 Detect direction of light
 Light-dark contrast
 Some can form an image

Types of Eyes
Flat sheet eyes

 Some sense of light direction and intensity
 Often in larval forms or as accessory eyes in adults
Figure 6.33a
Types of Eyes
 Cup-shaped eyes (e.g., Nautilus)

 Retinal sheet is folded to form a narrow aperture
 Discrimination of light direction and intensity
 Light-dark contrast
 Image formation
 Poor resolution
Figure 6.33b
Types of Eyes
 Vesicular Eyes (present in most vertebrates)

 Lens in the aperture improves clarity and intensity
 Lens refracts light and focuses it onto a single point on
the retina
 Image formation
 Good resolution
Figure 6.33c
Types of Eyes
 Convex Eye (annelids, molluscs, arthropods)

 Photoreceptors radiate outward
 Convex retina
Figure 6.33d
Compound Eyes of Arthropods
Composed of ommatidia (photoreceptor)
 Each ommatidium has its own lens
Images formed in two ways
 Apposition compound eyes (diurnal insects)
 Ommatidia operate independently
 Each one detects only part of the
image;
 Afferent neurons interconnect to form
an integrated image
 Superposition compound eyes ( nocturnal
insects)
 Ommatidia work together to form
bright superimposed image on retina
 Resolving power is increased by reducing size
and increasing the number of ommatidia

Structure of The Vertebrate Eye
 Sclera
 “White” of the eye
 Cornea
 Transparent layer on anterior
 Retina
 Layer of photoreceptor cells
 Choroid
 Pigmented layer behind
retina; contains blood
vessels; absorbs light in
diurnal animals
 Tapetum
 Layer in the choroid of
nocturnal animals that
reflects light
Structure of the Vertebrate Eye
 Iris
 Two layers of pigmented smooth
muscle
 Pupil
 Opening in iris allows light into
eye
 Lens
 Focuses image on retina
 Ciliary body
 Muscles that change lens shape
 Aqueous humor
 Fluid in the anterior chamber;
secreted by ciliary body
 Vitreous humor
 Gelatinous mass in the posterior
chamber
Image Formation
 Refraction – bending of light
rays
 Cornea and lens (both
convex lenses) focus
converging light on the
retina
 In terrestrial vertebrates,
most of the refraction
occurs between air and
cornea
 Lens does fine focusing
 Lens changes shape to focus
on near or far objects
 Accommodation

Image Accommodation
 Accommodation
 Light rays must converge
and focal point must fall on
the retina to produce a clear
image
 Focal point
 Point at which light waves
converge
 Focal distance
 Distance from a lens to its
focal point
 The eye is set for distance vision
(over 20 ft away)
 The lens must change shape to
focus for closer objects

Image Accommodation
Distant vision
 Light rays are parallel when entering
the lens
 Ciliary muscles relax
 Increase tension on suspensory
ligaments
 Lens is pulled and becomes thinner
 Little refraction of light by lens
Near Vision
 Light rays are not parallel when
entering the lens
 Ciliary muscles contract
 Decrease tension on suspensory
ligaments
 Lens becomes thicker
 More refraction of light by lens
The Vertebrate Retina
 Arranged into several layers
 Rods and cones are in the
retina and their outer
segments face backwards
 Other cells are in front of
rods and cones
 Bipolar cells, ganglion
cells, horizontal cells,
amacrine cells
 Axons of ganglion cells join
together to form the optic
nerve
 Optic nerve exits the retina
at the optic disk (“blind
spot”)
The Fovea
 Region in center of retina
 Overlying bipolar and
ganglion cells are pushed
to the side
 Contains only cones
 Allows light to strike the
photoreceptors without
passing through several
layers of neurons
 Provides the sharpest
images
 Image is focused on the
fovea
Cephalopod Eye and Retina
 Photoreceptors are on
the surface of the retina
 Project forward
 Supporting cells are
located between
photoreceptor cells
 No other layers of
cells associated
with photoreceptors
 Axons of
photoreceptors form
optic nerve

Signal Processing in the Retina
 Rods and cones form different images
 Rods
 Convergence
 Many rods synapse with a single bipolar cell
 Many bipolar cells synapse with a single ganglion cell
 Ganglion cells has large receptive field
 Poor resolution (fuzzy image)
 Cones
 Each cone synapses with a single bipolar cell
 Each bipolar cell connects to a single ganglion cell
 Ganglion cell has small receptive field
 High resolution
Convergence in the Vertebrate Retina
Figure 6.38a,b
Signal Processing in the Retina
“On” and “off” regions of the receptive field
of ganglion cells improve contrast of light
and dark
“Center-surround” organization of receptive
field
 “On-center” ganglion cells
 Stimulated by light in center of
receptive field
 Inhibited by light in periphery of
receptive field
 “Off-center” ganglion cells
 Stimulated by dark in center of
receptive field
 Inhibited by dark in periphery of
receptive field
Photoreceptors in center and periphery
inhibit each other by lateral inhibition
Lateral Inhibition in the Retina
Figure 6.40
The Brain Processes the Visual Signal
 Action potentials from retina ( temporal

and nasal parts of the retina) travel to
brain
 Optic nerves  optic chiasm 
optic tract  lateral geniculate
nucleus (thalamus)  visual cortex(
occipital lobe)
 Binocular vision
 Eyes have overlapping visual fields
 Binocular zone
 Combine and compare information
from each eye to form a three-
dimensional image
 Depth perception
Color Vision
 Detecting different
wavelengths of visible light
 Requires photopigments
with different light
sensitivities
 Most mammals: see two
(dichromatic) colors (
green , blue)
 Humans: see three
(trichromatic) colors
 Birds, reptiles and fish:
see three, four
(tetrachromatic), or five
(pentachromatic) colors

Color Vision
Figure 6.42
Thermoreception
 Central thermoreceptors
 Located in the hypothalamus and monitor internal
temperature
 Peripheral thermoreceptors
 Monitor environmental temperature
 Warm-sensitive ( above 300C; capsaicin)
 Cold-sensitive ( menthol)
 Thermal nociceptors – detect painfully hot stimuli ( starting
at 450C )
 ThermoTRPs
 Thermoreceptor proteins
 TRP ion channel
Specialized Thermoreception
 Specialized organs for detecting heat radiating

objects at a distance
 Pit organs
 Pit found between the eye and the nostril of pit vipers
 Can detect 0.003°C changes (humans can detect only
0.5°C changes)
Figure 6.43
Magnetoreception
 Ability to detect magnetic fields

 For example, migratory birds, homing salmon
 Neurons in the olfactory epithelium of rainbow trout
contain particles that resemble magnetite
 Responds to magnetic field

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CHAPTER

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 Range from single cells to complex sense organs

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Based on type of stimulus (stimulus modality) the

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 All stimuli are ultimately converted into action

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 Most cells can sense chemical stimuli

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 Each olfactory neuron expresses only one odorant

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 Five classes of tastants

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Three classes of receptors

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 Monitor the position of the body

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 Hair cells detect body position

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 Brain uses time lags and differences in sound

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 Ability to detect visible light

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 Vertebrates have ciliary

 Diverse shapes of rods

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Flat sheet eyes

 Cup-shaped eyes (e.g., Nautilus)

 Vesicular Eyes (present in most vertebrates)

 Convex Eye (annelids, molluscs, arthropods)

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 Action potentials from retina ( temporal

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 Specialized organs for detecting heat radiating

 Ability to detect magnetic fields

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