Theories about sensing

Aristotle (384-322 BC)

cardiocentric sensing.

Biophysics of sensory Galenus (129-200 AD) raised

doubts about cardiocentric

receptors Cardiocentric sensing

(Medieval reconstruction)
sensing.

Today:
stimulus 
 sensory receptors 
 receptor potential 
 neuron/nerve 
 action potential 
 central nervous system 
 signal processing 
 sensation
fMRI recording during
Sensory homuncle
sensomotoric function

Sensory receptors
Steps of sensing OLFACTION

VISION TASTE TOUCH

HEARING

Case of hearing
Sound recognition
Sound sensation

Intensity Loudness Meissner

Neural
Frequency Physically conductance
Psychologically Pitch body
measured sound
perceived sound
properties
Spectrum effects Tone
Free axon
Duration  Sensory receptor
 Perceived duration

Direction Brain
(ear) (auditory center) Perceived direction Rod Cone

Sensory receptor: Specialized sensory cell, which responds to a given stimulus (e.g., light, sound,
chemicals) and relays the information to the central nervous system.

Cell surface receptor (different meaning!): Proteins which specifically bind hormones,
neurotransmitters and other molecules, and thus iniate specific cellular reactions.
 Five senses? Steps of signal transduction
Most important sensory modalities (First 11: perceived modalities)
Environment STIMULUS
Modality Receptor Organ
(physical-chemical
1 Vision Rods and cones Eye
effects)
2 Hearing Hair cells Ear (organ of Corti) thres
3 Olfaction (smelling) Olfactory neuron mucus membrane
4 Taste Taste receptor cells Taste buds
5 Angular acceleration Hair cells Ear (semicircular canals)
6 Linear acceleration Hair cells Ear (utricle and saccule) Receptor RECEPTOR POTENTIAL
7 Touch, pressure Nerve endings Multiple types
8 Heat Nerve endings Multiple types
9 Pain Nerve endings Multiple types Uthres
10 Cold Free nerve endings ...
11 Joint position and motion Nerve endings Multiple types
Neuron
12 Muscle length Nerve endings Muscle spindle
13 Muscle stress Nerve endings Golgi’s tendon organ
14 Arterial presure Nerve endings Sinus caroticus stretch receptors Uaction ACTION POTENTIAL
15 Central venous pressure Nerve endings Venous, atrial stretch receptors Central
16 Lung stress Nerve endings Pulmonary stretch receptors nervous
17 etc... etc... etc... system

From stimulus to sensation

Sensitivity
Measured signal Step
STIMULUS

thres
speech
ASSOCIATION

recognition

eV - size stimulus is sufficient for

Ugen positive ions
perception
interpretation
hearing hearing and
vision
association
evoking action potential:
RECEPTOR

tial
oten sensory association
p tor p areas areas

Uthres
rece
• sound receptors: thermal motion of
ion channel the molecules of air
SENSORY CENTER

Uact • light receptors: 1-2 photons

tentials Uact
NERVE

o n po
acti
nerve

to activation of sensory
from receptor brain areas
 1. Modality
What is coded by the action potential?

Adequate stimulus
Type of energy for which the receptor is most sensitive (e.g.,
• modality (type) light for the eye).

• intensity (strength)
Action potentials are identical in all nerves. How do we know,
• duration for example, whether an action potential codes for touch and
not cold?
• localization

of the stimulus Principle of specific sensory energies

Sensation is determined by the stimulated cortical region!

2. Intensity 3. Duration, adaptation

Which parameters carry information about stimulus strength? Stimulus

• frequency of action potentials
• number of activated receptor cells Adaptation. During constant stimulus the
Action
potentials

frequency of action potentials gradually

decreases. Receptor
No adaptation
Weber-Fechner potential

psychophysical law Stevens’ law

= sensation strength
n =background intensity Action
  0=absolute threshold Rapidly adapting (phasic) receptors potentials
 = const  lg  = const  
intensity
0  0 n=constant specific for the
E.g., pressure, smell, heat Receptor
Slow adaptation

potential
type of sensation

Action
n<1: compressive function Slowly and partially adapting (tonic) receptors potentials
(hearing, vision)
E.g., cold, pain (dental pain) Receptor Fast adaptation
potential
n>1: expansive function
(pressure, taste)
Weber (1795-1878) Fechner (1801-1887) Stevens (1906-1973)
Seconds
 4. Localization, receptor fields

Branched nerve endings define receptor fields (convergence). Such can be found in
the skin (touch) and in the peripheral retina (rods).

Biophysics of the
eye
Receptor fields with overlap

Optical illusions - intensity Optical illusions – space

 Optical illusions – motion
Optical illusions – direction

Ooptical illusions point out the remarkable and unusual

processing power of the visual system.

Stimulus: light Stimulus: light

Electromagnetic wave Transverse wave

The eye is sensitive to: wavelength and amplitude (~intensity)

The eye is insensitive to: phase and polarization
 “Receptor-organ”: eye Photoreceptors
pigment
epithelium
light
plasma membrane
receptors discs
rods
optic nerve
fibers
cones membrane

discs
sclera disc
choroid

vitreous humour cytoplasm

retina cones
ciliary body plasma
t
ligh membrane
zonulae ciliares
(suspensory cilia membrane disc
ligaments)

lens mitochondria
rods
ganglion cells
eye axis macula bipolar
cells
anterior fovea nucleus retinal
chamber
optic nerve

synapse
cornea

opsin
iris rod cone
optic disc rhodopsin molecule
posterior
(blind spot)
chamber

Optics of the eye Accomodation and refractive problems

flattened relaxed ciliary
lens muscle bulged contracted
Total refractive power
lens ciliary muscle
of eye: 62 dptr

Optical power entering the eye (P)

2
 d J=intensity (W/m2)
P = J  
 2 d=pupil diameter

relaxed
stretched ligaments
2 ligaments
Pmax  dmax  dmax=8 mm
= = 16
Pmin  dmin  dmin=2 mm
Hypermetropia
Astigmatism LASIK
(Laser Assisted In Situ
Keratomileusis)

air Refractive power of surfaces (D)

cornea n  n' n-n’=refractive index difference Myopia
aqueous D= between refractive media
humour r r=radius of curvature of
refractive surface
lens
Correction with cylindric lens
vitreous
humour
 Spatial resolution of the eye
Generation of visual stimulus
visual angle: the smallest angle at which two object points can be
distinguished.
For a healthy eye: 1’ (angular minute, 1/60 degree)

visual _ acuity =
1'
100% =experimental visual angle
Sensitivity of the human eye


Night vision Day vision

Resolution has wave optics (diffraction) and biological (receptor density) limitations.
Astronomical magnitude

Object Image on receptors Sensation

~2m

Quarter Full moon Twilight

Cloudy Clear moon Sunrise Cloudy Direct sun
night sky night sky sunset day sky exposure

Photoreceptor distribution in retina Properties of receptor cells

blind spot

Rods Rods
Rod Cone
blind spot
Stimulated by very small intensity Smaller sensitivity, but functions at high
Cones Cones (optimally 1 photon!) intensities
fovea optic
nerve
Degree (˚)
Saturates at average intensities No saturation

Found mainly in the peripheral retina In the fovea, mainly central fovea

Many rods per ganglion (convergence);

Small convergence;
Rods greater sensitivity, smaller spatial
greater spatial resolution
Cones resolution

No color sensitivity Sensitivity to colors

Photochemical reaction Color vision

In human: 3 types of receptors. Each senses different

colors - absorbes at different wavelengths (R=64%,
1 rhodopsin absorbs 1 photon G=32%, B=2%).

500 transducin molecules activated

500 phosphodiestherase molecules
activated, and
11-cis-retinal 

Relative absorption
105 cGMP molecules hydrolyzed Light reflection from butterfly
 retina. The different receptors
optical excitation reflect dfferent colors.
250 Na+-channels closed
 rod
Entrance of 106-107 Na+ ions/s inhibited
 cones

cell hyperpolarized (1 mV)


transmitter release reduced (glutamate: Wavelength (nm)

inhibitory neurotransmitter).
all-trans-retinal

X = rR + gG + bB
Additive color mixing

Stimulus: sound

Longitudinal A
mechanical wave
(pressure wave)

Biophysics of 

hearing
Longitudinal wave Transverse wave

y=actual pressure; t=time

Harmonic oscillation: y ( t ) = Asin ( ft +  ) f=frequency; A=amplitude
=phase shift
 Sounds and their spectra Frequency and intensity of sounds
J 
Intensity level: n(dB) = 10 lg  1 
 J2 
Fourier transformation TYMPANIC MEMBRANE
Time domain SUPERSOUND SUPERSOUND Pressure wave therapy
Inverse Fourier transormation Frequency RUPTURE
domain
(Fourier spectrum) PAIN Ultrasound diagnostics,
therapy
pure DISCOMFORT
sound

dri ne
rbi
ll
Tu
Discrete spectra
HEARING
fundamental
harmonics
frequency

Intensity

HYPERSOUND
musical
INFRASOUND MU ULTRASOUND
sound SIC

white
HEARING
noise

Continuous spectra
Cardiac sounds
SILENCE

Frequency
drum beat
Wavelength
(in air)

“Receptor-organ”: ear Physical schematics of the ear

Inner ear The pressure wave
Outer ear Middle ear Ossicular Lever system of ossicles
of the fluid generates
results transmits waves with
muscles inceased forces surface waves on the
basal membrane
Semicircular
canals (vestibular
organ) Outer ear Middle ear Inner ear
The basal membrane is
Auricula Oval Axis of rotation narrow and taught at the
window of lever system beginning of the cochlea
The basal membrane is
wide and slack at the
end of the cochlea

Apex
Cochlea extended (35 mm)
Auditory
nerve

External auditory canal: Auditory nerve

External resonator with quarter
Apex wavelength.
auditory canal cochleae Hair cells on the vibrating
basal membrane are bent
Auditory ossicles and excite auditory nerve
Eardrum (malleus, incus, stapes) The smaller surface
endings
Cochlea area of the oval window,
compared with that of While the oval window
Round the eardrum, results in is pressed inside, the Pathway of
window pressure increase. round window is atmospheric pressure
Eustachian pressed outside equilibration
tube
 Middle ear: mechanical amplifier
Outer ear: sound collector

Auricula Auditory ossicles (malleus,

incus, stapes)
Sound is steered into the external auditory
canal. They amplify eardrum
resonance and transmit it to
the oval window. Auditory
External auditory canal ossicles
Oval
Eardrum
Conducts pressure waves towards the Amplification:
window

eardrum. More efficient in certain freqency due to area ratio: 17 

due to lever action: 1,3 
range (2000-5000 Hz). water

air

Total amplification: 22 drum

(pressure increase)
Eardrum
Brought into resonance by sound waves. drum

Inner ear: sensor Ultrastructure of the inner ear

Vestibular organ: semicircular canals Inner hair cells. Appr. 500

Cochlea: 2.5-pitch, 35-mm-long fluid-filled channel.

It is halved in length partly by an osseous, partly by a membranaeous
wall, the basal membrane.
Sensory organ of sound.

Semicircular
Stapes
canals

Oval window
Cochlea

Eardrum
Basal membrane

Round window

Outer hair cells. Appr. 12-20000

 Békésy: propagating surface waves on basal membrane Function of the organ of Corti

Due to the bending of the basal membrane, hair cells become tilted and depolarized.

Oval window
“Surf” wave

Apex
Oval window Velocity of wave propagation
Frequency
Velocity of “surf” wave is smaller
than that of sound (1440 m/s)

2. Tectorial membrane
becomes lifted

x
Basa

Ape
l mem
brane
Inner hair cell
Pivot point

Location of propagating wave at Outer hair 3. Shear

subseqent times cells force

Relative deflection
Apex
Oval window

Surface curve of
propagating wave Distance from oval window (mm)
György Békésy
Nobel-prize 1961 Basal membrane at rest Pivot point 1. Basal membrane
becomes distorted

The frequency-dependence of the location of propagating wave maxima

provide a rough frequency-discrimination.

Inner hair cells:

Mechanoelectric transducers Outer hair cells: amplifiers
endolymph mechanical
effect, stereocilia
deform Passive mechanism
Loose Tight tip
tip link force (basal membrane)
link
stereocilia Active mechanism

Oscillation amplitude
cation gates open (tip link
effect), K+ ions enter (outer hair cells)
support cells

(log)

Base

Apex
hair cell becomes
depolarized
hair cell Basal membrane
nucl.
Ca2+ channels open,
Ca2+ ions enter
Distance from oval membrane (mm)
synapse Outer hair cells
neurotransmitter is
released into synaptic cleft

afferent Regenerative amplifier:

efferent
afferent neuron positive feedback
depolarizes, frequency of
action potentials increases (large amplification in narrow frequency range)
neurons
 Psychoacoustics: loudness (Fletscher-Munson)
Coding of acoustic information ethalon sound
loudness Hson (son)
loudness level Hphon (phon)
pain threshold

Intensity level
Location theory Volleyball theory

Pressure

Intensity
Disco
120 phon
stimulus oval wido surface wave
w
base basal membrane
apex
Frequency sensing coded
loc
locally. Street
noise
80 phon
oscillatio
n
amplitu
de
Basis: Loud
speaking
1. Weak frequency-dependence speak
ing ra
60 phon
nge
of the amplitude maxima of t (tim
e)
hair
propagating surface waves. cells

2. Active amplification. action Whisper

poten 30 phon
tials
3. Frequency senitivity of
thr
afferent neurons (threshold simu
ltane es
ho
o
pote us action ld
of
stimulus depends on freqency). ntials he
arin
Simultaneous appearance reference intensity, J0 g

of action potentials in the

auditory nerve may
encode large frequencies. frequency f (Hz)

Phon and son scales

n)
(so
’law
v ens
Ste
law
Weber-Fechner ale (phon)
cte d (lo ga rith mic) loudness sc
Constru
loudness level
loudness

a
at
ald
e nt
rim
pe
ex

Intensity

Intensity level