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Sensory Systems. 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 - PowerPoint PPT Presentation
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Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
PowerPointPowerPoint®® Lecture Slides prepared by Lecture Slides prepared by Stephen Gehnrich, Salisbury UniversityStephen Gehnrich, Salisbury University
6C H A P T E R
Sensory SystemsSensory Systems
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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
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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
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Vertebrate Tactile Receptors
Widely dispersed in skin Receptor structure
Free nerves endings Nerve endings enclosed
in accessory structures (e.g., Pacinian corpuscle)
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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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Insect Tactile Receptors
Two common types of sensilla
Trichoid Hairlike projection of
cuticle Bipolar sensory neuron TRP channel
Campaniform Dome-shaped bulge of
cuticle Bipolar sensory neuron
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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
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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
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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
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Insect Hearing
Strong vibrations sensed by trichoid sensilla Weak vibrations and sounds are detected by
chordotonal organs Clusters of scolopidia Located on leg Mechanosensitive ion channels
Tympanal organs Thin layer of cuticle (tympanum) overlays chordotonal
organ
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Vertebrate Hair cells Mechanoreceptor for hearing and
balance Modified epithelial cells (not neurons) Cilia on apical surface
Kinocilium (a true cilium) 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
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Signal Transduction in Hair Cells
Figure 6.18
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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
Fish Neuromast
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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) Mammalian Ear
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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)
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Vestibular Apparatus (1)
Mechanoreceptors of the inner ear
Macula Present in utricle and
saccule 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
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Vestibular Apparatus (2)
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
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Maculae Detect Linear Acceleration and Tilting
Figure 6.23
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.24
Cristae Detect Angular Acceleration
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.25
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
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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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.26a
Mammalian Middle and Inner Ear
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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)
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.26a,b
Mammalian Inner Ear
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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 Basilar membrane vibrates Stereocilia on the inner hair cells bend Hair cells depolarize Hair cells release neurotransmitter (glutamate) Glutamate excites sensory neuron
Round window serves as a pressure valve
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Encoding Sound Frequency
Frequency Detection Basilar membrane is stiff and narrow at the proximal
end and flexible and wide at distal end High frequency sound vibrates stiff end Low frequency sound vibrates flexible end
Specific regions of brain respond to specific frequencies
Place coding
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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 in response to sound
Do not release neurotransmitter Change in shape increases movement of basilar
membrane Increased stimulus to inner hair cells
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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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.27a,b
Electromagnetic Spectrum
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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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.28
Phylogeny of Photoreceptors
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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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsTable 6.1
Characteristics of Rods and Cones
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.30
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
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Photopigments
Photopigments have two covalently bonded parts 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
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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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.32
Phototransduction
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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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.33a
Types of Eyes
Flat sheet eyes Some sense of light direction and intensity Often in larval forms or as accessory eyes in adults
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.33b
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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.33c
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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.33d
Types of Eyes
Convex Eye (annelids, molluscs, arthropods) Photoreceptors radiate outward
Convex retina
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Compound Eyes of Arthropods Composed of ommatidia
(photoreceptor) Each ommatidium has its
own lens Images formed in two ways
Apposition compound eyes Ommatidia operate
independently Each one detects
only part of the image
Afferent neurons interconnect to form an image
Superposition compound eyes
Ommatidia work together to form image
Resolving power is increased by reducing size and increasing the number of ommatidia
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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
Tapetum Layer in the choroid of nocturnal animals that reflects
light
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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 Vitreous humor
Gelatinous mass in the posterior chamber
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Image Formation
Refraction – bending of light rays Cornea and lens focus 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
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Image Accommodation
Accommodation Light rays must converge 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
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Image Accommodation
Distant objects Light rays are parallel when entering the lens Ciliary muscles contract Lens is pulled and becomes thinner
Little refraction of light by lens
Close objects Light rays are not parallel when entering the lens Ciliary muscles relax Lens becomes thicker
More refraction of light by lens
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.36
Image Accommodation
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The Vertebrate Retina Arranged into several layers
Rods and cones are 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”)
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The Fovea
Region in center of retina Overlying bipolar and ganglion cells are pushed to the
side Contains only cones
Color vision
Provides the sharpest images
Image is focused on the fovea
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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
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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 cells connects to a single ganglion cell Ganglion cell has small receptive field High resolution
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.38a,b
Convergence in the Vertebrate Retina
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.39
Signal Processing in the Retina
Complex “on” and “off” regions of the receptive fields of ganglion cells improve their ability to detect contrasts between light and dark
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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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.40
Lateral Inhibition in the Retina
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The Brain Processes the Visual Signal Action potentials from
retina travel to brain Optic nerves optic
chiasm optic tract lateral geniculate nucleus visual cortex
Binocular vision Eyes have overlapping
visual fields Binocular zone
Combine and compare information from each eye to form a three-dimensional image
Depth perception
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Color Vision
Detecting different wavelengths of visible light Requires photopigments with different light
sensitivities Most mammals: see two (dichromatic) colors Humans: see three (trichromatic) colors Birds, reptiles and fish: see three, four
(tetrachromatic), or five (pentachromatic) colors
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.42
Color Vision
Retina and brain compare output from each type of receptor and infer the color
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Thermoreception
Central thermoreceptors Located in the hypothalamus and monitor internal
temperature
Peripheral thermoreceptors Monitor environmental temperature
Warm-sensitive Cold-sensitive Thermal nociceptors – detect painfully hot stimuli
ThermoTRPs Thermoreceptor proteins TRP ion channel
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsFigure 6.43
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)
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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