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SENSORY

RECEPTORS
Dr Kunal MBBS, MD, PGDYS
Associate Prof. Physiology
kunal.kmc@manipal.edu;
8792873242
Specific Learning Objective – Receptor - I
1. What is the definition of receptors and how are they classified –
types of receptors with examples
2. Name the types of touch and pressure receptors found in the skin
and explain their functions
3. Describe the receptors that mediate the sensations of pain and
temperature
4. Explain the electrophysiology of receptor and outline the steps of
transduction from the generation of generator potential to the
transmission of impulse with the example of Pacinian corpuscles

Dr Kunal's Physiology Notes: CNS 02-05-2023 2


Specific Learning Objective – Receptor - II
1. Describe and discuss the properties of receptors
2. Discuss the basic elements of sensory coding in CNS
3. Describe and discuss the coding of sensory information with
respect to stimulus modality, intensity, duration and location
4. Explain the role of lateral inhibition in sensory system

Dr Kunal's Physiology Notes: CNS 02-05-2023 3


Stimulus Sensory Physiology
Sensory Receptors
Transduction
Action Potential

Afferent Nerve
Spinal Cord

Sensory Cortex For


Translation processing and interpretation
5/2/2023 Receptors by Dr. Kunal 4
Sensory Mechanism
• Stimulus: Internal or External: Chemical,
Mechanical, Electrical, Thermal, etc.
• Sensory Receptors: Biological transducer
get depolarized in response to a stimulus
• Transduction: stimulus energy is converted
into electrical energy (Action Potentials (AP)
• Conduction: Transmission of impulses
(APs) to the CNS through Afferent Nerves
• Translation: Integration & Processing of AP
into – Sensation & Perception
• Sensation: decoding the arriving info
• Perception: awareness of sensation
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Receptors
• Biological Transducer - at peripheral end of sensory neuron
• converts different forms of energy into electrical energy (AP)
• Part of Neuron - Free nerve ending or Specialized cell
• Surrounded by connective tissue (capsule)
• Receptor/Generator potential: stimulus changes the membrane
potential of receptor; greater stimulus means larger receptor potential
(Graded Potential)
• The frequency or pattern of action potentials contains information
about the strength, duration, and variation of the stimulus.
• Receptive field: area monitored by a receptor; may be as low as 1mm
to as high as 70 mm
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Sensory
Units and
Receptive
Fields

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Receptors – Points to be discussed

• Evolution of sensation
• Classification of receptor: Based on the following
1. Physiology
2. Structure
3. Stimulus
4. Location
5. Adaptation

5/2/2023 Receptors by Dr. Kunal 8


Evolution of Sensations
1. Epicritic
• Newer, Fine, Discriminative
• Localization is accurate
• Center – cerebral cortex
• Eg: Fine touch, fine gradation of temp 350C to 400C
2. Protopathic
• Older, Crude, non-discriminative
• Localization is inaccurate
• Center – below the Cortex
• Eg: Crude touch, crude temperature sense
• 1st to return, when a nerve fiber regenerates after an injury

Receptors by Dr. Kunal 9


Physiological Classification of Receptors

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Classification
of receptors:
Based on
Structure

Receptors by Dr. Kunal 11


Complex Specialized
Receptors – Receptors
Capsulated
Simple Nerve Ends
Receptor
s – Naked
dendrites
A receptor Cell
synapsing with E.g.
Classification of Afferent Nerve Most
receptors: Based – releases NTs Special
on Structure senses
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Classification of
receptors: Based
on Structure
1. Free nerve endings:
pain
2. Expanded tip endings
Merkel’s disc
3. Hair end organs
4. Encapsulated nerve
endings
Meissner’s corpuscle
Pacinian corpuscle
Krause’s corpuscle
5. Spray nerve endings
5/2/2023 Receptors by Dr. Kunal
-Ruffini’s
13
Stimulus-Based Classification of Receptors
1. Mechanoreceptors – for touch, pressure, vibration, stretch, and itch
2. Thermoreceptors – sensitive to changes in temperature
3. Nociceptors – sensitive to Potentially harmful stimuli such as pain,
extreme heat, and extreme cold
4. Photoreceptors – respond to light energy (e.g., retina)
5. Chemoreceptors – for smell, taste, blood chemistry changes
6. Osmoreceptors – for concentration of solutes, osmolar changes
7. Baroreceptors – detect changes in fluid pressure

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Sensation – Receptors
1. Fine touch – Meissner corpuscles,
Merkel's Discs, Free Nerve
Endings, Hair End-organ
2. Crude Touch (Pressure) – Ruffini
End organs, Pacinian corpuscles
3. Pain – Free nerve endings of A-δ &
C fibers
4. Temperature – CMR1, VR1, VRL-1
5. Proprioception – Muscle Spindle &
Golgi Tendon Organ

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Location of
Mechanoreceptors
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Location-Based Classification of Receptors
• Exteroceptors (Superficial) – touch, pressure, pain, temperature, itch
E.g. Mechanoreceptors, Thermoreceptors, Nociceptors, etc
• Deep (Proprioceptors) – sensitive to degree of muscular stretch;
Located in muscle, tendon, joints.
E.g. Muscle Spindle, Golgi Tendon organ, free nerve ending
• Interoceptors (visceroceptors) –
E.g. Osmoreceptors, Baroreceptors, Chemoreceptors (taste,
smell, pO2, pCO2, pH),
• Teleceptors (“Wave receivers") – for light (rods & cones), sounds (hair
cells) etc;
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Q1. Why do you stop noticing the
temperature of the room or the
feelings of the clothes on your body?
Q2. You wear a costly perfume, and
enjoy the new smell, but, for how long?

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Adaptation-Based Classification of Receptors
Adaptation: reduction in response due to a constant stimulus
A. Peripheral adaptation: change in receptor activity
1. Phasic (fast-adapting receptors): e.g., Meissener’s, Hair follicles,
Pacinian, Thermoreceptors (usually you stop noticing the temperature of
the room or the feelings of the clothes on your body unless it changes)
2. Tonic (slow-adapting receptor): e.g., Merkel Discs, Ruffini’s End
Organs, Nociceptors, Vestibular,
B. Central adaptation: inhibition of nuclei in the pathway
• E.g., smell (you walk into a room and notice a new smell, but…not for long)
• Adaptation reduces the amount of information reaching the cerebral
cortex and thus to our awareness (only ≈1% of sensory info reach)
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Classification of receptor: Based on Adaptation
1. Tonic: always “on”: slowly adapting receptors
• continue to transmit information for many hours, never get tired
• e.g. muscle spindles and Golgi tendon apparatus continue to send
the status of muscle contraction and load on muscle tendon;
• Macula in the vestibular apparatus, nociceptors, baroreceptors,
chemoreceptors
2. Phasic / Rapidly Adapting / Rate /Movement Receptors
• only active for a short period during stimulus
• Stop transmitting the information very soon, later may respond
again, but only to change in the intensity of stimulus
• e.g. Meissner’s, Hair follicles, Pacinian, thermoreceptors
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Fine Touch Receptors
1. Meissner corpuscles: adapt rapidly – Phasic
• Dendrites (type Aβ)encapsulated in connective tissue
• Respond to fine touch and slow vibrations.
• Non-hairy parts of the skin, specially in the fingertips, lips, and
other areas where one's ability of “tactile localization” is highly
developed
2. Merkel's Discs / Iggo dome / Expanded Tip Receptors:
• adapt slowly – Phasic
• Expanded dendritic endings,
• Respond to fine touch, ability of “tactile localization”

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Iggo Dome Receptor: Group of Merkel's discs
For Light Touch
projects upward
against the underside
of the epithelium,
protruding the skin and
creating an extremely
sensitive dome.
innervated by a single
large myelinated nerve
fiber (type Aβ)

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Fine Touch Receptors
3. Free Nerve Endings: Adapt rapidly
• Found everywhere in the body, in skin and in many other tissues.
• The cornea of the eye contains no other type of receptor but free
nerve endings.
• Even light contact can elicit touch and pressure.
4. Hair End-organ: Adapt rapidly
• Each hair and a nerve fiber entwining its base,
• Detect initial contact of objects with the body and even slight
movement of it on the body surface

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Mechanoreceptors
of Skin

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Crude touch / Pressure Receptors
5. Ruffini End organs: (Tonic) adapt slowly,
• dendritic endings with elongated capsules,
• Enlarged and multi-branched, present in deeper tissues
• Continue sending signal about prolonged touch and pressure and
the degree of joint rotation, (the states of deformation)
6. Pacinian corpuscles: (Phasic) adapt rapidly
• Unmyelinated dendritic endings (type Aβ),
• Encapsulated by multiple concentric lamellae of connective tissue
(appearance of an onion peel).
• Beneath the skin and deep in the fascial tissues
• Respond to deep pressure and fast vibration. or other rapid
changes in the mechanical state.

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5/2/2023 Dr Kunal's CNS Lecture notes: Somatic Senses - I 27
Encapsulated Nerve Endings - Proprioceptors

The conscious
component of
proprioception
("body image") is
actually
synthesized from
information coming
not only from
receptors in and
around joints but
also from
cutaneous touch
and pressure
receptors.
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Muscle Spindle &
Golgi Tendon
Organ

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Nociceptors
• Commonly present in: Skin, joint capsules, coverings
of bones, around blood vessel walls
• not present in the brain… then why headache?
• Tonic receptors: no significant peripheral adaptation
as long as the stimulus is present (it keeps hurting),
• Central adaptation may occur (your brain has
capacity to decrease the perception of pain)

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Nociceptors
• Free nerve endings of A-δ fibers • Free nerve endings of C fibers
have smaller receptive fields have large receptive fields
– Diameter: 2–5 µm – diameter <1.2 µm
– Conduction velocity: >12 m/s – Conduction velocity: <2 m/s
– fast pain (cut, prick, etc,); – slow pain “burning, aching”
– easy to localize, – difficult to localize,
– slow adaptation – never adapted

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Too much of any stimulus can
activate pain receptors
• Mechanical Nociceptors: free endings of Aδ fiber (fast)
• Thermal Nociceptors: free endings of Aδ fiber & vanilloid
receptors: (respond to vannilin or capsaicin protins)
– VR1: potentially harmful temperatures above 43 °C.
– VRL-1, activated by severe temperatures around 50°C or and also
by severe cold
• Chemical Nociceptors: free endings of C-fiber (slow)
– Sensitive to Chemicals – a) release by damaged cells; like
bradykinin, histamine, high acidity; b) some environmental irritants
• Polymodal Nociceptors: free endings of C-fiber (slow)
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Thermoreceptors: Warm receptors
– Free nerve endings of C fibers
– in dermis, skeletal muscles, hypothalamus, liver
– respond to temperatures from 30–43°C
– Slow Conduction velocity: <2 m/s
– Phasic: quickly adapting to stable temperature,
respond only to changing temperature

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Thermoreceptors: Cold receptors
– Free nerve endings of Aδ & C fibers
– Both Fast & Slow Conduction
– Cold- and menthol-sensitive receptor 1 (CMR1)
– Firing frequency steadily increases as skin
temperature falls to 24°C and then decreases until
the temperature reaches 10°C
– Respond to temperatures from 35°C – 10°C
– Inactive <10°C and >40°C

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Thermoreceptor Responses

At or Below
5°C, most of the
receptors are
inactive and
thus cold
becomes an
effective local
anaesthetic

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Food For Thoughts

Why do metal objects


feel colder than wooden
objects of the same
temperature?

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Thermoreceptor
Responses
Human Body has 4 to 10 times
as many cold-sensitive spots as
heat-sensitive spots
Cool metal objects feel colder
than wooden objects of the
same temperature because
the metal conducts heat away
from the skin more rapidly,
cooling the subcutaneous
tissues to a greater degree.

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Transduction: Generator / Receptor Potential

• Transduction: Conversion of one form of energy into other forms


• All sorts of stimuli are converted into Receptor Potential (RP)
e.g. phototransduction, auditory transduction, etc.)
• If stimulus is strong enough, the RP crosses the threshold and
thus causes an AP
• Maximum amplitude of receptor potentials is about 100 mV
(equal to an AP)
• As much the receptor potential rises above threshold level, the
higher frequency of AP is produced in the afferent nerve

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Transduction:
generation of
Generator
Potential or
Receptor
Potential

5/2/2023 39
Transduction: Generator / Receptor Potential
Core of the Pacinian Corpuscle has an
unmyelinated nerve terminal,
surrounded by multiple concentric
layers of membrane
Compressions anywhere around the
capsule elongate, indent, or deform the
central unmyelinated fiber, causing
opening of stretch-sensitive ion
channels (depolarization)
If depolarization reaches threshold due
to Strong stimulus, AP is transmitted in
the sensory nerve
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Transduction: Generator / Receptor Potential
Pressure Stimulus

Compression of Pacinian Corpuscle

Change in the shape of Pacinian Corpuscle

Deformation of central core fiber

Opening or Closing of mechanically Gated Ion channels

Entrance of sodium ions into the core fiber

Development of receptor potential
5/2/2023 Receptors by Dr. Kunal 42
Transduction: Generator / Receptor Potential
Development of local circuit current flow

Spread of local circuit current to 1st node of Ranvier

If crosses Threshold, many VGNa+ channels open

Influx of 1000s of Na+ ions (+ve feedback of Hodgkin Cycle)

Generation of action potential in nerve fiber

Transmission of AP to the spinal cord and Brain

Sensation & Perception – “Your reality vs my reality!”
5/2/2023 Receptors by Dr. Kunal 43
Receptor Potential: Basic Facts

2. “Receptor potential
is generated at the
1. Pressure at the
central core fiber”,
capsule generates
not at the capsule
Receptor potential

3. Putting pressure or 4.“Action potentials


applying anesthetic are generated at the
drugs on 1st node of 1st node of Ranvier”
Ranvier abolishes AP
but does not affect RP
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Food For Thoughts

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Properties of Receptors
1. Law of Differential Sensitivities (specificity) or, Law of
Adequate Stimulus, or, specificity of receptors
2. Specific Nerve Energy (Muller’s Doctrine)
3. Labelled Line Principle
4. Weber-Fechner Law or Power law or Law of Intensity
discrimination
5. Law of projection (sensory homunculus),
6. Adaptation of receptors
7. Lateral inhibition

5/2/2023 Receptors by Dr. Kunal 47


Coding of Sensory Attributes
• Sensory Coding: Converting a receptor stimulus to a
recognizable sensation
• All sensory systems code for 4 elementary attributes of a
stimulus (MILD)
1. Modalities
2. Intensity
3. Location
4. Duration

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1. Modalities:
• Law of Differential Sensitivities (specificity)
• Law of Adequate Stimulus
• Specific Nerve Energy (Muller’s Doctrine)
• Labelled Line Principle Coding of
2. Intensity
• Law of Intensity discrimination
or, Weber-Fechner Law Sensory
or, Power law
3. Location: Attribute
• Law of projection (sensory homunculus),
• Lateral inhibition, the density of receptors
4. Duration:
(MILD)
• Adaptation of Receptors
5/2/2023 Receptors by Dr. Kunal 49
Differential Sensitivities, or
Law of Adequate Stimulus
• Each type of receptor has different sensitiveness to different
types of stimulus
• Each receptor is most responsive to one type of stimulus and
is almost non-responsive to other types of stimuli
• Adequate Stimulus: Form of energy to which a receptor is
most sensitive (have a minimum threshold for action potential)

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Specificity of Receptors
✓ Each receptor is highly specific to respond to one
type of adequate stimulus .
✓ Other type of stimuli may evoke (same) response, if
given in supra-threshold strength, but
✓ Same stimulus applied on different receptors gives
a different sensation
✓ Different stimuli applied on same receptor give the
same sensation

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Transduction by Adequate Stimulus
Stimuli the cause opening of Na+ channels to depolarise the receptors
Mechanoreceptors
– opening of stretch or pressure-sensitive channels
Chemoreceptors
– opening of ligand-gated channels
Thermoreceptors
– Change of temperature opens the temperature-sensitive channels
Auditory Receptors (hair cells)
– vibration due to Sound waves opens stretch-sensitive channels
Photoreceptors (rods & cones)
– Light energy (photons) cause chemical changes to “close” the channel
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Muller’s Doctrine of Specific Nerve Energy
• When a nerve pathway from a particular sense organ is
stimulated, the sensation evoked is that for which the
receptor is specialized, no matter how or where along the
pathway and which type of stimulus is applied. (Muller-1835)
• A pathway is specific for a “modality” (type) of sensation
• Stimulation of Dorsal Column Pathways by a Spinal tumor
pressing over, or by an Electrocautery during neurosurgery
➢ If a dorsal column fibre is stimulated anywhere along its course,
the person always perceives “feather touch” regardless of the
types of stimulus (pressure, electricity, heat, cut, crush, etc.,)

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• A pathway or “line” is specific for a “modality” of sensation
• Stimulation of Dorsal Column Pathways by a Spinal tumor
pressing over, or by an Electrocautery during neurosurgery
• If a dorsal column fibre is stimulated anywhere along its course,
the person always perceives “feather touch” regardless of the
types of stimulus (pressure, electricity, heat, cut, crush, etc.,)

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Receptor 1A Receptor 2
Receptor 1B

➢ This specificity of nerve fibers


for transmitting only one
modality of sensation from
specific part of the body is
called the labelled line principle Cortex
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• Sensation is felt at the point of origin of sensory tract, no
matter where the stimulus is being applied, along the course
• AP from each receptor, via its afferent pathway, reach to a
specific area of brain meant for that receptor only
• Sensory Homonculus, is the part of sensory cortex (area
no. 1, 3 & 5) where whole body is systematically represented
• Systematic representation of whole body is maintained all
over the nervous system

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• Sensation is felt at actual location of receptor, even if the
pathway is stimulated anywhere from receptor to cortex.
• No matter where a particular sensory pathway is
stimulated along its course from the receptor to the
cortex, the conscious sensation felt is always
referred to the receptor site
• Eg: Phantom Limb

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Phantom Limb
➢ A person complains of severe pain and burning sensation in left
arm, which had been amputated 7 years ago!!
➢ Explained on the basis of “Law of Projection”
➢ Irritation of nerve fibers of the amputated limb at the
a) Amputee stump b) any where along the course c) in spinal
cord d) in thalamus
➢ Irritation of designated “cortical area” of the amputated limb
by overgrowth or overstimulation of surrounding cortical areas

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Amount of sensation felt is Proportional to intensity of stimulus
1. Increasing frequency of action potentials
2. Recruiting more & more receptors, because
Week stimuli activate only the receptors with lower threshold,
Stronger stimuli also activate receptors with higher threshold

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 Stimulus Intensity →
 Magnitude of Generator Potential (GP) →
 Number of Action Potential reaching to CNS per second →
 Magnitude of sensation felt
• “Frequency” of AP is directly proportional to Stimulus “Intensity”

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• Weber’s law: Frequency of Action Potential α Intensity of stimulus
f AP α ΔI = Kw. I
(Kw = Weber Fraction; ΔI = Intensity difference)
• Fechner’s law:
Magnitude of sensation felt α log of intensity of stimulus
Ψ = k logS
(Ψ = sensation felt, k = constant & S = physical intensity of the stimulus)
• Fechner’s law provides an explanation for Weber’s law
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The “intensity of sensation” varies with the logarithm of the
“intensity of stimulus” and a constant”
S = K log I + C
(K& C-constant; S-intensity of sensation; I-Intensity of stimulus
Now modified into,
R = KAS
K & A are constant for any specific sensory modality
R-intensity of sensation; S-Intensity of stimulus

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a. Intensity
Coding by
Recruitment

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b. Intensity
Coding by
Recruitment

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Intensity Coding by frequency of APs

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Hand has four types of mechanoreceptors; their combined
activation produces the sensation of “Touch” with an object.

Sensory systems encode four attributes of stimuli: modality,


location (receptive field), intensity, and duration (timing)

• Stimulus intensity: signalled by rates of firing


• Stimulus duration: signalled by time course of firing.

The spike trains: action potentials elicited by pressure from


a small probe at the center of each receptive field.
5/2/2023 Receptors by Dr. Kunal 66
• Location of a stimulus is encoded by spatial distribution of
the population of receptors activated.
• Deeper receptors have “Larger” receptive fields and thus,
have less precise Localization. (MMRP)
• Most precise localization: Meissner's corpuscles &
Merkel discs
• have the smallest receptive fields and
• are most sensitive to small probe pressure and touch.

The receptive fields of different mechanoreceptors


differ in their size and response to touch.

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Stimulus Intensity
Coding
More the receptor
potential rises above the
threshold level, the
greater becomes the
action potential frequency

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The 
frequency of
APs from
sensory
receptors in
proportion to
the  in
receptor
potential.

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Extreme range of response,
from very weak to very intense
• With progressively stronger
mechanical compression (
stimulus strength), the receptor
potential  rapidly at first but then
less rapidly at high stimulus
strength.
• Applicable to almost all sensory
receptors, allows the receptor to be
sensitive to very weak sensory
experience and yet not reach a
maximum firing rate until the
sensory experience is extreme.
5/2/2023 Receptors by Dr. Kunal 70
Food For Thoughts
Why are you having Problems reading this?
How do Blind People Read?
Which specialty of the nervous system benefits them?
Braille symbols are separated by what distance?
5/2/2023 Receptors by Dr. Kunal 71
Two Point Discrimination
• Tactile Acuity: ability to feel two closest points as separate points
• Two-point discrimination threshold: minimum distance between
the two points that can be felt as separate points
• Blind individuals benefit from the higher tactile acuity of fingertips
which facilitates the ability to read Braille; the dots forming Braille
symbols are separated by 2.5 mm.
• Two-point discrimination is used to test the integrity of – dorsal
column (medial lemniscus) system, parietal lobe of cerebral cortex

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Acuity is improved
by Lateral Inhibition
Lateral Inhibition:
• Afferent neurons of stimulated receptors
have inhibitory connections with the
surrounding afferent neurons
• Inhibitory influence of stimulated afferent
neurons is seen at each level (primary,
secondary & tertiary)

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Action potentials
Activity in second-
order neurons

Frequency of
Second-
order X2 Y2 Z2
neurons Inhibitory Inhibitory
interneurons interneuron
s
X2 Y2 Z2
2nd-order neurons Acuity is
improved
Activity in
by

Action potentials
afferent neurons

Frequency of
Afferent
neurons Lateral
(first- X1 Y1 Z1
order X1 Y1 Z1
Inhibition
neurons) Afferent neurons

Location of stimulus

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Acuity is
improved
by
Lateral
Inhibition

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Acuity:
Two Point
Discrimination

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The cortical Sensations

1.Tactile Localization
2.Two-Point Discrimination
3.Stereognosis

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Stereognosis
• Recognizing an object without looking, just by touching it. Perception of
the form and nature
• Astereognosis: Loss of stereognosis = Tactile Agnosia
✓ an early sign of damage to parietal lobe (posterior to postcentral
gyrus) and may occur without any detectable defect in touch and
pressure sensation
✓ Also occur when dorsal columns tracts are damaged

Agnosia: failure to identify an object


after seeing it (Visual), by its sounds or words
(Auditory), just by knowing the color (Color), or by
the location or position (Position)

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Mechanisms of Receptor Adaptation
• Viscoelasticity of Pacinian Corpuscle:
• distorting force is transmitted by the viscous component of the
corpuscle to the central nerve fiber
• within a few milliseconds, fluid within corpuscle gets redistributed
and receptor potential can no longer be elicited.
• Accommodation: due to time-bound "inactivation" of Na+ channels in
nerve fiber membrane (Inactivation of all or most Na+ channels are
Time-bound)
• Rapidly Adapting Receptors (Rate, or Movement, or Phasic) are
stimulated only when there is a change in stimulus strength
• Rods and cones adapt by changing the concentrations of their light-
sensitive chemicals
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Slow Versus Fast Adaptation

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Adaptation of Receptors

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Sustained Light Touch

Free
Nerve
Endings Slowly adapting

Rapidly adapting

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Encapsulated Nerve Endings
Spiraling nerve ending surrounded by Schwann cells

Rapidly
adapting

Slow
adapting

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Dr Kunal's Physiology Notes: CNS 02-05-2023
5/2/2023 Receptors by Dr. Kunal 87
Principle Sensory Modalities
Types of Skin Receptors (Exteroceptors)

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