TOPICS COVERED

General classification & properties

Muscle spindles
The gamma system
Golgi tendon organs
Articular receptors
Cutaneous receptors

UNIT CONTENT

We have looked at the neuron and should have an idea how impulses
(action potentials) are conducted. However, at this point we have not
discussed just how stimuli are converted into impulses. All sensory
systems are designed to detect forms of energy and the interface
between stimulus and the response of a neuron is normally a structure
called a RECEPTOR. The process by which the various sensory receptors
convert a particular form of energy into another form is known as
transduction.  In other words a receptor converts stimuli into impulses.
Narrated animation about skin receptors
Animation 1

 
Sensory Modalities
 Stimuli can be divided into a range of different types or MODALITIES.
Receptors normally respond to only one type of stimuli (or sensory
modality), and that type of sensory modality is called the adequate
stimulus for a particular type of stimulus. For example, the adequate
stimulus for the eyes photoreceptors (rods and cones) is light and not
touch (mechanical stimuli).

Below is a list of receptor types verses their adequate stimulus:

Adequate Stimulus
Receptor Type
Mechanoreceptors Mechanical displacement

Thermoreceptors Temperature change

Nociceptor Pain
Chemoreceptors Chemicals
Photoreceptors Light
           

With an adequate stimulus, receptors produce a receptor potential when

stimulated.

The Nature of a Receptor Potential

1. All RP's are graded (ie. RP's differ in magnitude).

2. The magnitude (amplitude) of a RP has a sigmoid  relationship with
stimulus strength.
3. The magnitude of the RP is linearly proportional to the firing rate of
the associated afferent neuron.
4. All receptors adapt to stimuli. However, at which they adapt varies
between slow and fast. (What is the significance for this variation?).
In the graphic illustration of the relationship between the receptor
potential and the firing rate along the afferent neuron, as soon as the
excitation threshold of the first node of Ranvier is reached the firing rate
increases linearly as the magnitude of the RP increases.
For example, a touch receptor in the skin (a mechanoreceptor) – impulse
generation
1. The neural part of the Pacinian corpuscle (PC) is  polarized.
2.  The PC is mechanically stimulated.
3. The neural part of the Pacinian corpuscle is depolarized.This forms the
receptor potential.
4. The greater the degree of mechanical stimulation, the larger the
magnitude of the RP.
5. If the excitation threshold at the first node of Ranvier is reached ,
then an AP is generated along the entire length of the afferent nerve
fibre.  

Receptors also work on the same principles as Adequate Stimulation,

spatial and temporal summation as mentioned in unit 2. These are
important concepts with respect to in intensity Coding –

1) means by which we can determine the intensity with which a

particular sensory event occurred

2) property that enables us to distinguish between different sensations

(e.g., hard slap vs. light tap, loud vs. soft sound, strong vs. weak muscle
contraction)

3) the intensity of a stimulus can be conveyed by two mechanisms

Spatial Summation - the stronger the stimulus the larger the

number of different sensory receptors fire

Temporal Summation - the stimulated receptors are fired at a

higher frequency

Sensory Adaptation –enables us to block out irrelevant sensory

information

1) shortly after a sensory receptor registers a stimulus, the firing

rate is reduced or, curtailed

2) different sensory receptors adapt at different rates: touch and

pressure receptors vs. pain receptors and certain proprioceptors
 3) different adaptation rates of receptors determines the nature of
the information provided to the CNS

The receptors important for movement are mostly somatosensory

receptors.  Please refer to the diagram below.

Types: Types:
muscle spindles
Ruffini endings Golgi tendon organs
joint receptors
Pacinian corpusles, Meisner corpusles vestibular apparatus

   
Provide touch and pressure. Distribution is somewhat
disproportionate throughout the body - lips and fingers
contain many more receptors per sq inch of surface
Provide uninterrupted knowledge about the
area
general position of our body in space prior
to and during movement - also provides us
Why? - these areas of the body are involved in very
with information regarding where are body
fine movements and therefore require greater levels of
segments are relative to each other
sensation

 
Receptors that are stimulated by physical deformation are known as
mechanoreceptors

We will limit our discussion to four types of receptors which may be

involved in the position and movement of a limb, and involved in
proprioception. Each receptor provides different information.
 
 

Awesome Site - read carefully - Required

 
Receptors:
 
Muscle spindles (important for muscle length and velocity and to a lesser
extent, monitoring static length)
Golgi tendon organs (detects muscle force)
Joint afferents (most sensitive to position at extreme joint angles)
Tactile (pressure receptors in muscle and overlying skin, sense flexion or
extension of finger)
 

MUSCLE SPINDLES
·        Detect dynamic and static changes in muscle length

·        considered the proprioceptor of greatest importance

·        serves a dual sensory and motor function

·        higher numbers of spindles per square inch of surface area are

located in muscles subserving fine movements (tongue, hands)

Function of the Muscle Spindle

Muscle spindles are found in all somatic muscles, within the belly of the
muscle and run in parallel with the main muscle fibres.  Whenever
skeletal muscle is stretched the muscle spindles are stimulated. They
detect changes in the length of muscle fibers as well as the rate of
change at which the muscles are lengthening.

1. The Muscle Spindle consists of 2-10 specialized muscle fibers

enclosed in a connective tissue capsule. These muscle fibers are
called intrafusal muscle fibers. Voluntary skeletal muscle fibers are
called the extrafusal muscle fibers.
2. Intrafusal Muscle Fibers:
 ·         Nuclear Bag Fibers:
o        approximately 1-3 fiber per spindle
o        the nuclei are found concentrated in a bag type
central part of the fiber (hence the name 'nuclear
bag' fibers).
o        the ends of these fibers are striated (contain actin
and myosin filaments) and are contractable. The
ends of these fibers are also attached to the
EXTRAFUSAL muscles.
·         Nuclear Chain Fibers:
o        approximately3-7 fibers per spindle
o        the nuclei are spread in a chain-like fashion in the
center of the fiber (hence the name).
o        the ends of these fibers are striated (contain actin
and myosin filaments) and are contractable. The
ends of these fibers are attached to the ends of the 
nuclear bag muscle fibers.

During a muscle stretch the sensory nerve terminals increase their

discharge rate as the sensory ending is stretched. This nerve terminal is
known as the ANNULOSPIRAL ending, so named because it is
composed of a set of rings in a spiral configuration. These terminals
(shown in blue on the diagram below) are wrapped around specialised
muscle fibers that belongs to the muscle spindle (INTRAFUSAL
FIBRES) and are quite separate from the fibers that make up the bulk
of the muscle (EXTRAFUSAL FIBRES).

More about the Nuclear Bag fibers and the GAMMA SYSTEM:

A motor supply to the intrafusal muscle (shown above in red). In this

region on either side of the central area the intrafusal fibres are able to
contract if their motor supply is active. The motor supply comes via
efferent fibres that usually fall into the gamma classification of
diameters. They are often (and better) referred to
as FUSIMOTOR fibres.

Two things can, in principle, cause the annulospiral ending to be

stretched and so increase its discharge.

1. A stretch of the muscle as a whole will stretch the spindles

within it, and thus the sensory endings.

2. Fusimotor activity will cause contraction of the intrafusal

fibres below the fusimotor nerve terminals either side of the
central region. This will result in stretch of the central sensory
region .

Nuclear chain fibre also have annulospiral sensory endings in the central
region (the nuclei are in line). It is a shared branch of the axon that
supplied the central area of the nuclear chain fibre. This sensory nerve is
of group Ia, the fastest found in the body.

Further out we see that there are other sensory endings, more closely
associated with the chain fibres. These fall into the
slower group II division of sensory nerves and are referred to
as SECONDARY endings in contrast to the centrally
located PRIMARY endings.

The two types of intrafusal fibre, (bag and chain) have different
mechanical properties, and respond differently to their largely separate
fusimotor fibres. They also differ in respect to their sensory endings.
Consequently, the information relayed to the CNS by the spindle via
group Ia and group II sensory endings is different.

In simple terms, the Ia afferents respond partly to muscle

length, but respond more powerfully to changes in length
(BLUE). The group II afferents are much better at registering
length alone (RED).

Ia afferents can powerfully excite the ALPHA MOTONEURONES of the

muscle containing the spindle. This is the basis of the classical STRETCH
REFLEX in which extension of the muscle (and thus its spindles) cause a
reflex contraction.

The spindle can therefore register muscle length and velocity.

Furthermore the sensitivity to both length and velocity can be
altered by the CNS via activity in the fusimotor system, the static
gamma system controlling length sensitivity and the dynamic
gamma system controlling velocity sensitivity.

Golgi tendon organ (GTO)

The GTO detects:

a) the rate of increase of tension on a muscle during active

contraction

b) the absolute amount of tension on a muscle during a

isometric contraction Prevents damage during excessive force
generation
GTOs are relatively simple sensory structures which consist of
elaborated nerve ending in a capsule which lies within the tendon at the
musculotendinous junctions of somatic muscle. Each skeletal muscle
contains a large number - approximately 1 GTO for every 10 muscle
fibers.  These ending produce generator potentials and action potential
discharges with frequency proportional to the force exerted on the
capsule. The tendon organ acts as a "force transducer", and because it is
in series with the tendon is only activated in a sustained fashion when
the muscle is active. These sensory structures appear to be quite
localized, that is they will monitor force within a subdivision of tendon
which represents the physical connection of a motor unit to the global
tendinous structure of the muscle. Thus the CNS has information about
the force output of individual motor units and can make feedback
adjustments appropriately in the efferent side.

Figure: muscle spindle structure compared to GTO structure.

GTO’s are innervated with a 1b afferent fiber. This afferent information

may also contribute to preventing the muscle from developing too much
tension through its inhibitory connections with alpha motor neurons. 
See unit 7.

Joint/ Articular receptors

The joint capsules and ligaments of all synovial joints in the skeletal
system are well endowed with proprioceptors.  Collectively, joint
receptors provide us with information about: a) static position of a joint
in space (i.e. joint angles) b) endpoint positions of joints during active
movement
Joint Receptors
Receptor Type
Location
Response
Function
Associated Afferents
 
Type I
Ruffini-like
ligaments & joint capsule
Mechanoreceptor;
Slow adapting;
Active at end ranges (passive)
change of direction, amplitude, pressure, velocity to cerebellum
II
 
Type II
Pacinian-like
joint capsule & fat pads
Mechanoreceptor;
Rapid adapting;
Active at end ranges-dynamic
boost muscle at beginning of movement
II
 
Type III
GTO-like
ligaments
Mechanoreceptor;
Slow adapting;
Active under tension
protective reflex
Ib
 
Type IV
Free nerve-like
ligaments, joint capsule, fat, periosteum
Nocioceptor;
Slow adapting;
flexion reflex -->prevent further movement
A delta & C
Tactile (cutaneous) receptors

Cutaneous sensations (review skin structures)

Cutaneous receptors:  Glabrous (smooth, hairless skin) and hairy skin

both contain a wide variety of receptors for the purpose of detecting
mechanical, thermal, or painful stimuli applied to the body surface. 
Three types of receptors are common to glabrous and hairy skin:
Pacinian corpuscles, Merkel's discs, and free nerve endings.   The other
receptors found in glabrous skin are Meissner's corpuscles.  The most
important receptors in hairy skin are the hair follicle endings and Ruffini
terminals.

1. Merkel Disk:  Slowly adapting mechanoreceptors structured to

respond to maintained deformation of the skin surface.  Typically, an
afferent fiber branches to form a cluster of Merkel's discs, situated at
the base of a thickened region of epidermis.  Each nerve terminal
branch ends in a disc enclosed by a specialised accessory cell called a
Merkel cell.  Movement of the epidermis relative to the dermis will
exert a shearing force on the Merkel cell.  The Merkel cell plays a role
in the sensing of both touch and pressure.

2. Meissner Corpuscle: Terminal branches of a myelinated axon

intertwines in a basketlike array of Schwann cells.  Found just
underneath the epidermis in glabrous skin (nipples, fingertips, soles
of feet), these receptors are quickly adapting mechanoreceptors
providing discriminative touch.

3. Ruffini Corpuscles:  Spray-like dendritic endings in the dermis

hairy skin which are involved in the sensation of steady, continuous
pressure applied to the skin. 
 

4.  Pacinian Corpuscles:  Pacinian corpuscles are pressure

receptors. They are located in the skin and also in various internal
organs. Each is connected to a sensory neuron.

5. Free Nerve Endings:  Made up of branching nerve axons, which

are entirely or partially surrounded by Schwann cells.  The
axon/Schwann cell complex is further surrounded by a basement
membrane.  Free nerve endings originate from fine myelinated or
unmyelinated fibers that branch extensively in the dermis and may
penetrate into the epidermis.  These endings respond to strong
mechanical and thermal stimuli, and they are particularly activated by
painful stimuli.

Type Adaptation
  rate
(receptive fields)
Merkel Type I Slow
 Ruffini Type II Slow

Meissner Type I Fast

Pacinian Type II Fast
Corpuscle
 

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