Chapter 44 Opening Roadmap.

Chapter 44

Animal Sensory
Systems

授課教師 生命科學系陳瑞芬

僅作為授課教學使用，禁止列印、影印、和公開散佈。
© 2017 Pearson Education, Ltd.

Key Concepts (I) Key Concepts (II)

How do sensory organs convey information to
the brain?
Hearing and equilibrium
Hair cells --- mechanoreceptors
Sensory receptor cells transduce (轉換;轉導) Vision
stimuli to changes in membrane potential
Rods & cones --- photoreceptors
(receptor potential).
(electromagnetic receptors)
Action potentials are sent via specific sensory Taste and smell
neuron to a specific portion of the brain. Chemoreceptors
The signals are processed and integrated in Other sensory systems
specific brain regions.
Thermoreceptors, electroreceptors,
magnetoreceptors, nociceptors, …
3 4
 Sensing and Acting Sensing and Acting
Male moths have large antennae to locate females • Bats use sonar to detect their prey
in total darkness.
receptor cells pheromone • Moths, a common prey for bats
– Can detect the bat’s sonar and attempt to flee

Echolocation
Sensing

Acting

A bat using sonar to locate its prey

5 66

Keeping up with Bats:

Tympanic organ of moths
Dynamic Auditory Tuning in a Moth
Tympanic organ low high
intensity intensity

300 m neuron

at the metathorax Receptor cells

1 mm (or 1 cell)

• Noctuid moths are most sensitive to frequencies at 20–40 kHz.

• Dependent on incident sound intensity, the moth’s ear
mechanically tunes up and anticipates the high frequencies
used by hunting bats.
FROM EGGERS (1919) WITH SOME CHANGES FROM TREAT AND ROEDER (1959).
Current Biology 16, 2418–2423, December 19, 2006. 7 8
http://www.lapshin.iitp.ru/project2/index.php?page=auditory/tympanic
 An insect’s “ear”—on its leg Tympanal Organ

Many insects weta

have a
tympanic
membrane
stretched over
a hollow Tympanic
membrane
chamber.
1 mm Campbell,
Fig. 50.9
Invertebrate phonoreceptor
Journal of Experimental
(Tympanal Organ) weta 沙螽 紐西蘭長鬚無翅大蝗 Biology (2011) 214: 778-785
9 ㄓㄨㄥ 10

Sound sensitivity in most insects depends Reception,

on body hairs that vibrate in response to transduction &
amplification of Integration of Response
sound waves. signal information to signal
• Different hairs respond to different frequencies. Sensory Transmission Transmission
e.g. fine hairs on antennae of male mosquito receptors CNS Effectors
CNS
respond to the hum
produced by the beating wings Bat call
of flying female Sensory Motor
input output Flight muscles
e.g. vibrating body hairs of caterpillars (AP)
Sensory
neuron Ears
detect the buzzing wings of predatory wasp

11 Fig. 44.1 12
 Sensory Information Pathway Sensation 感覺
Stimulus -- an awareness of sensory stimulus
( when action potentials reach the brain)
Reception & transduction of stimulus
Receptor into receptor potential
in sensory receptor Perception 知覺
such as colors, smells, sounds, tastes
Nerve Transmission of action potential -- a meaningful interpretation or conscious
pathway in sensory neuron understanding of sensory data
-- are constructions formed in the brain
Brain Sensation & perception of
region stimulus
in central nervous system
13 14

15 16
 EPSP, IPSP
Graded potential Receptor potential Action potential
voltage-gated Na+ channel
--- variable amplitude & duration
voltage-gated K+ channel
--- passive spread (local potential)
--- all-or-none response
(decremental conduction)
--- reversing polarity
--- no threshold
--- has a threshold & refractory period
--- no refractory period (不反應期)
--- conducted without decrement
--- could be summated

17 18

General Function of Sensory Receptors Coding of stimulus intensity

Reception --- absorb the energy of a stimulus 編碼
• The response of a sensory receptor varies with
Transduction (receptor potential) intensity of stimuli
stimulus energy electrochemical activity
• If the receptor is a neuron, a larger receptor
Amplification --- accessory structure potential results in more frequent action
part of transduction process potentials
Transmission • If the receptor is not a neuron, a larger receptor
action
receptor potential potentials CNS potential causes more neurotransmitters to be
(sensory nerves) released more frequent action potentials
Integration
sensory adaptation in afferent neuron
receptor sensitivity

19 20
 (a) Receptor is (b) Receptor regulates
afferent neuron. afferent neuron. frequency
To CNS To CNS action
Action (transmission) Action
potential potential
potential Afferent Afferent
neuron neuron postsynaptic
potential size
Receptor receptor
protein Neurotransmitter potential
Sensory receptor
receptor receptor (graded
potential potential
potential)
Stimulus
(transduction) leads to
(reception) changes in stimulus
neuro-
Stimulus Sensory transmitter
receptor release.
cell Stimulus
Fox, Physiology
Campbell, Fig. 50.3 21 Coding of stimulus intensity 22

(b) Multiple receptors activated How to assess the nature of stimulus:

Sensory receptor Gentle pressure 1. receptor, pathway (brain area) types
Fewer 2. number of afferent axons
receptors intensity
activated 3. frequency of APs
More pressure
More
receptors
activated

weak strong weak strong

Campbell, Fig. 50.4b
23 stimulus stimulus stimulus stimulus 24
 Tonic receptor Phasic receptor Specific sensory neuron specific
張力(感)受器 相位(感)受器 receptor brain area

• The brain distinguishes different

types of stimuli
slow adaptation fast adaptation – Different sensory neurons synapse
with different interneurons in the
brain

25 26

Interneurons Types of sensory receptors:

Sugar
receptor Salt
receptor Mechanoreceptors --- touch, pressure, stretch,
auditory, balance
Brain Chemoreceptors --- internal chemical senses,
taste, smell
Sensory Taste Electromagnetic receptors --- light, magnetic field,
neurons bud
photoreceptors 光受器 electricity
Thermoreceptors --- cold or warm receptors in skin
thermoreceptors in hypothalamus
No sugar No salt
Increasing sweetness Increasing saltiness Nociceptors --- pain receptors
傷害性受器

27 28
 • Exteroceptors 外接受器 Mechanoreceptors 機械性受器

--- detect stimuli originating outside • Mechanoreceptors sense physical

the body deformation
• Interoceptors 內接受器 – Caused by stimuli such as pressure,
touch, stretch, motion, and sound
--- detect stimuli originating inside
the body
e.g. proprioceptors 本體接受器
osmoreceptor
…
30
29

Muscle spindles 肌梭 Extracellular recording of action potential

(mechanoreceptors)
dendrites
of sensory neuron

α-motor
neuron

crayfish, stretch receptor, afferent nerve

Knee-jerk reflex
31 32
 fast adapt. (< 1s) fast adapt.
Baroreceptors Carotic sinus Meissner’s
Merkel’s disc vibration (30-40 Hz)
(Mechanoreceptor) 頸動脈竇 corpuscle
light touch
slow adapt. (Free nerve ending)
vibration Naked dendrites
: Baroreceptors light touch very slow adapt.
(dendrites of pain
sensory nerve) Epidermis temperature
pressure?
Aortic 表皮層
Sensory
fibers arch
主動脈弧 Skin very fast adapt.
Pacinian
Dermis corpuscle
Medulla
真皮層 deep touch
pressure
vibration
(250 - 300 Hz)
Hypodermis
真皮下層
Nerve Connective Hair
33 tissue movement 34
Campbell, Fig. 50.5

Receptive 10 mm
field Two point 22 mm
discrimination
48 mm

42 mm

Perception of
two point of Perception of Sensory
touch one point of neuron
touch 3 mm
2 mm
35 36
receptors sensory nerve Somatosensory Vibrissae
region in S1
cortex

觸鬚
Vibrissae

37 S1: primary somatosensory 38

cortex

Somatosensory
Neuronal activity in cortex
somatosensory Thalamus
cortex (Vibrissae)
觸鬚

Medulla
cross

cross Spinal
cord

pain touch, vibration.

temperature two-point discrimination,
some touch
proprioception
39 40
Vander, Figure 7.19
 Lateral Inhibition in CNS 圓鈍物 Lateral inhibition of afferent pathways
側邊抑制 Skin
輕輕觸壓

• Sharpening (敏銳化) of
sensation Presynaptic 突觸前
Stimulation
• Sensory neurons in inhibition 抑制
the center areas are
stimulated more than
1st 感覺神經元
neighboring fields
• Perceive single touch Sensation

敏銳化

Fox, Fig. 10.6 抑制性 旁邊鄰近區域的

41 1st 感覺神經元 中間神經元  1st 感覺神經元的活性 42

Two-point discrimination
Lateral No lateral
inhibition inhibition

Sharpening (敏銳化)
of sensation

側邊抑制

43 44
 The Mammalian Ear Amplification Incus 砧骨
in middle ear Malleus
錘骨 卵圓窗
Outer Middle Inner Oval window
ear ear ear 聽神經元 22 m
Auditory
neurons Sound waves
(to brain) Sound waves Stapes (in fluid)
鐙骨
(in air) 1 m
Middle Cochlea 耳蝸
Cochlea 耳蝸
ear cavity
Tympanic Eustachian 鼓膜, 耳膜 耳咽管
Ear Eustachian
canal membrane tube 耳咽管 Tympanic membrane (eardrum)
(eardrum) tube
耳道
鼓膜, 耳膜 Force  & surface area  Pressure on oval window

Fig.44.4 46
45 Fig.44.4

1. To protect the cochlea

Attenuation 2. To mask low-freq. sounds in a 正常人耳能聽到的最小音量定為 0分貝
loud noisy environments.
reflex (Acoustic 3. To decrease a person’s hearing
Intensity (amplitude) of sound
reflex) 減弱反射 sensitivity to his/her own speech.
dB = 20 log
Intensity (amplitude) of standard sound
Muscles
contraction
20 dB = 10x
Tensor
osscicles become tympani 40 dB = 100x
more rigid muscle 60 dB = 1000x
鼓膜張肌
conductance
人耳能分辨
50~100 msec (reflex time) 0.1 ~ 0.5 dB difference
Low freq. > high freq.
20 ~ 20,000 Hz (human)
•Increasing the stiffness of ossicles Eustachian Tympanic Stapes Stapedius
•Changing the direction of vibration of tube membrane muscle 0.3 % difference in pitches (freq.)
ossicles so that movement is less 鐙骨肌
effectively coupled to cochlea 47 48
 (perilymph) Cochlea Cochlear (endolymph) Vestibular (perilymph)
前庭管 duct canal
High Na+ Vestibular Auditory nerve Low Na+ High Na+
Low K+ canal (to brain) High K+ Low K+
Bone
耳蝸管 Middle
Cochlear canal
duct Organ of Corti Upper Auditory
(endolymph) Upper canal nerve
canal
柯氏器
Low Na+
High K+
Neurons Tympanic Lower
Lower (to auditory canal canal 柯氏器
Tympanic canal nerve)
(perilymph) Organ of Corti
canal 鼓管
(perilymph) (a) The middle chamber Cross section
of the fluid-filled cochlea through cochlea
Campbell,
contains hair cells
Fig. 44.5a Fig. 50.10
49 50

Bony Labyrinth --- semicircular canals, vestibule, The Organ of Corti 柯氏器
骨性迷路 cochlea (耳蝸) (前庭) 實體纖毛
Tectorial
Membranous Labyrinth --- semicircular ducts, membrane Stereocilia
膜性迷路 utricle & saccule, cochlear duct 覆膜

Hair cells
(receptors)

基底膜
Basilar membrane Axons of sensory neurons

perilymph (b) Hair cells are sandwiched

endolymph between membranes.
Figure 16.20 51 Fig. 44.5b 52
 Stereocilia--- microvili containing actin filaments
Hair cell 實體纖毛 lack of motility
Stereocilia
1 m No kinocilium

Bundled hairs projecting from a hair cell 10–50 m in length

Outer hair cells Inner hair cell
Human Ear
Human
Campbell, Fig. 50.10 53 54

(a) Hair cells have many stereocilia and one kinocilium.

(microtubular) Scanning electron micrograph of several stereocilia
動纖毛 Kinocilium
實體纖毛
Stereocilia
Arrows:
Potassium
channels tip links
joined by
protein “threads”
Nucleus
saccule
Hair cell

Afferent
sensory
neuron

Efferent
sensory 1 m
neuron
Fig. 44.3a Bullfrog 55 56
 A sound wave is a longitudinal wave consisting of
1. Pressure wave a series of compressions (high pressure) and
Pressure wave bends stereocilia. rarefactions (low pressure).
稀薄
endolymph high K+
2. K+channels open more. The speaker C R C R C R C R C R C R
low Na+
More K+ ions flow in. vibrates.

3. Membrane depolarizes.
Depolarization (20 mV)
Synaptic vesicle 4. More calcium flows in. Air pressure amplitude
Calcium channel 5. Synaptic vesicles fuse.
6.  Release of
neurotransmitter
Neurotransmitter
: wave length 
Afferent
released into
synapse neuron Sound wave are traveling
(to brain) Fig. 44.3b 57 http://astronomy.swin.edu.au/~smaddiso/astro/SiS/NPS/sound1.html 58

Sound wave are traveling • These vibrations create pressure waves in the fluid
in the cochlea ( vestibular canal round window)
Point C
Longitudinal wave Axons of
sensory neurons
e.g. sound wave Apex
Oval
window
B Stapes Vestibular
C canal
A

Cochlea

Point B
Tympanic
membrane Basilar
membrane

Base Round
Tympanic
window canal
(a)
http://kiwiphysics.blogspot.com/2014/08/blog-post_96.html Point A
59
Campbell, Fig. 50.12a
 聽覺的閾值 基底膜只需位移 0.3 nm “Hairs” of
maximum: 1.0 m hair cell
More Less
Neurotrans-
NT release NT release
Stereocilia bend outward mitter at
synapse

Sensory Receptor potential

50 50 50

potential (mV)
neuron

potential (mV)

potential (mV)
Membrane

Membrane

Membrane
70 70 70

Action potentials

Signal

Signal
Signal
0 0 0

70 70 70

012 345 67 012 345 67 012 345 67
Time (sec) Time (sec) Time (sec)
Basilar membrane (a) No bending of hairs (b) Bending of hairs (c) Bending of hairs
deflected up in one direction in other direction

Spontaneous Frequency  Frequency 

firing (AP) (AP)

Campbell, Fig. 50.11

61 62

Vibrations in the cochlear fluid

basilar membrane vibrates

hair cells brush against the tectorial membrane

generation of receptor potential (graded )

or  NT release
The firing rate of AP recorded from an
auditory nerve fiber in the squirrel monkey or  action potentials in sensory neurons
Fig.44.2 63 64
How the cochlear distinquishes pitch? 音調 The Basilar Membrane Varies in Stiffness
Different pitches Oval window
stimulate hair cells in different parts of Fig.44.6
the organ of Corti
“place code”
A B C Cochlea
Axons of Point C 3 6,000 Hz Base of cochlea
sensory neurons
(near oval

Relative motion of basilar membrane

Apex
Oval window)
window
B Stapes Vestibular
0 Uncoiled cochlea
C canal 3 1,000 Hz Basilar membrane
A

Cochlea
0
Point B
Tympanic 3 100 Hz 20,000 Hz 2000 Hz 200 Hz 20 Hz
membrane Basilar
membrane

Base Round
Tympanic
Point A canal 0
Narrow part of basilar Wide part of basilar
window 0 10 20 30
Distance from oval window (mm)
membrane is stiff– membrane is flexible–
vibrates in response vibrates in response
(a)
Campbell, Fig. 50.12
(b)
to high frequencies to low frequencies
65 66

Auditory Pathway from the Organ of Corti

Traveling wave in basilar membrane Temporal lobe
Maximal distortion p. 970 聽覺皮層(初級)
Georg Von Békésy auditory cortex
(p. 970 有誤)
Perilymph 1920s-1930s
(中腦)下丘 thalamus 視丘
midbrain
上橄欖核 (inferior colliculi)
耳蝸神經核 下丘
medulla
A high degree
of bilateral
connectivity

67 Figure 16.25 68
 噪音對人體的影響不容忽視 歐盟規定iPod音量不得超過
100分貝
根據研究，長期生活在音量> 70 dB的環境中 如果超過100分貝，每天連
• 聽力受損 續聽2小時以上，只要半年，
• 在神經系統方面，會出現頭昏、頭痛、出冷 聽力就會受損。
汗、兩手發抖等症狀
• 消化系統方面，會食慾不振，甚至腹痛 Live rock band- 100 dB
• 呼吸急促、困難
Conversational speech
50-60 dB
Elevator music- 40 dB
69 70

Sensory Worlds: What Do Other Echolocation 回音定位 ultrasonic sounds

Animals Hear?
Perceive shapes with ears!
• Elephants use infrasounds (sound
frequencies too low for humans to hear) to
communicate (mid-1980s, K. Payne).

• Bats use ultrasonic sounds (sound

frequencies too high for humans to hear) to
echolocate (navigate by sound).

• Hair cells in the lateral line system allow

fishes and amphibians to sense pressure
changes in water.

71 ~Fig. 44.7 72
 Echolocation
buzz
(回音定位)

Ultrasound
20-120 kHz

口 200 dyne/cm2
喉 或 (close to the mouth)
鼻
Jet plane takes off & passes
only 100 m overhead

73 74

Frog ear Lateral line → nearby (water movement & vibration)

Hearing system → from a greater distance (vibration)
骨膜 Otolith utricle
organs saccule
(inner ears) lagena
(hair cells inside)

耳柱骨
Weberian ossicles

Air-filled
Swim bladder
(resonating chamber) Carp 鯉魚
Herring 鯡魚
75
** an amplifier of sound (助聽器?) 76
 側線系統 Nearby (water movement & vibration)  Salmon have a lateral line system (側線系統), seen
here as the dark line along the sides of the fish
 This enables the salmon to sense the direction
and velocity of water currents (水流的方向和速
度) and thus distinguish which direction is
upstream (上游)
Lateral line  Unfortunately for
the salmon, it
cannot perceive
a bear's paw
descending
from above
Migrating salmon
Fig. 44.8a 77 78

(b) Water enters the canals through pores and

bends kinocilia & stereocilia on hair cells,
activating sensory neurons.
動纖毛

實體纖毛

Fig. 44.8b
79 Fig. 44.8c 80
 Fig. 44.9
81 82

鯰魚

孔雀魚

Fig. 47.9
Fig. 44.9 83 84
 Vestibular system (前庭系統) balance
半規管 Semicircular
canals
Dynamic Vestibular
nerve 前庭神經
equilibrium

前庭 Vestibule
Static 橢圓囊
equilibrium Utricle
Saccule
球囊
Fig. 44.9
85 Campbell, Fig. 50.13 86

Static equilibrium Static equilibrium

橢圓囊
utricle horizontal acceleration linear
static head position acceleration
耳石 saccule vertical acceleration
球囊
Gelatinous
cap

gravity tilt

87 88
 壺腹 Left ear Right ear
ampulla 頂蓋 Rotation
Flow of fluid Cupula speed
Flow of fluid

Firing
rate
Hairs
Hair
cell Firing rate
of sensory
neurons Leftward rotary
Nerve fibers
Cupula from acceleration of
horizontal the head
Direction of body movement semicircular
canal

Campbell, Figure 50.13 89 90

Head
Vestibular-ocular
reflex (VOR) Lateral Medial
recti Lateral
rectus rectus
Keep your eye muscle Left Right muscle
pointed in a particular
direction even while
you are dancing like + + + +
(Midbrain)
a fool. + +
前庭器 Oculomotor
nucleus (III)
eye
movements (Pons)
Abducens
可作為軀體平衡及眼 nucleus (VI)
球運動之依據 — +
腦幹 counter Vestibular
中腦 head nucleus
脊髓 movement +
動眼中樞
+ Excitatory, inactive
Horizontal
+ Excitatory, active semicircular
91 — Inhibitory, active canals 92
 Nystagmus slow component (VOR)
眼球震顫 fast component (saccade)
自發性眼球震顫

93 94

What causes motion sickness? Statocyst 平衡胞

(Mechanoreceptors)
• Motion sickness is a result of the brain Pressure
receiving signals from equilibrium receptor
cells
receptors in the inner ear that conflict with
visual signals from the eyes
Cilia
gravity

Statolith
平衡石
Sensory
nerve fibers
(axons)

95
Campbell, Fig. 50.8 96
www.solarnavigator.net/sea_sickness.htm www.kathycrowe.com/.../motionsickness.html