Describe the neural mechanisms underlying the stretch reflex.

How
might this contribute to the control of posture and movement?

As a standard clinical diagnostic test, the knee jerk reflex is an example of

the stretch or myotatic reflex in action. This physiological activity comprising an
intricate network of neural fibres is responsible not only for the general
contraction one can witness in muscles but also maintenance of the muscle tone.
In addition it has a significant role in the generation of rhythmic activity such as
swimming in the fish or walking in the human. By cutting the dorsal roots thus
eliminating sensory input from the muscle to the spinal cord,, Sherrington
demonstrated that such an incision led to a loss of muscle tone despite the alpha
motor neurons still being intact. Similarly he discovered that the length of the
muscle i.e. The degree to which it was pulled, was intrinsically related to its
contraction and this was shown through a measured correlation between the firing
of the Ia sensory axons and the length of the muscle of fibre.
The relation between the sensory Ia fibres and the alpha motor neurons
arises from the principle of a monosynaptic arrangement. Only one synapse
separates the primary sensory input and the motor output in this network.
Lengthening of the muscle induces stretching of the muscle spindle fibres in the
equatorial region resulting in opening of mechanosensitive ion channels. Hence a
depolarisation of the Ia sensory nerve is incurred and this relays ultimately to the
alpha motor neurons elevating their action potential firing rate.
Alpha motor neurons are only responsible for the innervation of one type of
the spindle fibres. Those found inside the fibrous capsule of the spindle are known
as intrafusal fibres while those lying outside the spindle are known as extrafusal
fibres and constitute the “bulk” of the muscle. These latter fibres are innervated by
the alpha motor neurons however the intrafusal fibres rely upon a different type of
motor neuron, the gamma motor neuron, for their innervation. Due to the location
of the extrafusal and intrafusal fibres in reference to the muscle, innervation by
both the alpha and gamma motor neurons must coordinate to prevent the spindle
from going “off the air”. Contraction of the extrafusal fibres comes from the alpha
motor neurons however if the spindle went slack then there would be no firing by
the Ia axons and thus no relaying of sensory information. Instead the intrafusal
fibres found at the ends of the muscle spindle are innervated by the gamma motor
neuron and when acitvated, will induce a contraction of the two poles of the
spindle. This pulls the equatorial non-contractile region and maintains the activity
of the Ia axons. Thus the output of the Ia axons is constantly modulated by the
dual inputs of the alpha and gamma motor neurons, with the former decreasing
activity and latter increasing Ia activity. This loop of the circuit comprising
innervation of the intrafusal fibres by gamma motor neurons leading to extrafusal
fibre contraction is known as the gamma efferent loop.
At the junction of the muscle and the tendon, one can find another sensor,
which contributes to our 3d positioning in space, or proprioception – the Golgi
tendon organs. They also monitor the force of contraction and are innervated by
the Ib sensory axons, these being slight smaller in diameter than the Ia fibres.
Contrary to the muscle spindles, which are organised in parallel, the tendon organs
are arranged in series and thus one can observe their responsibility in relaying
information concerning the tension of the muscle as opposed to the length, which
is
accounted for by the spindles. Ib fibres projecting away from the Golgi tendon
organs, travel towards the spinal cord where they synapse onto the ventral horn
interneurons having branched significantly. As in many other neurological aspects
of the human anatomy, there is a concept of lateral inhibition occurring here. A
few of these interneurons in the ventral horn will reconnect with the alpha motor
neurons innervating the same muscle and provide an inhibitory effect. This
principle however is not known as lateral inhibition but plays a role in the reverse
myotatic reflex, which is believed to contribute not only to our fine motor skills
by maintaining muscle tension in an appropriate range but also as a protective
measure against overloading of the muscles. When we lift a heavy object, muscle
tension increases and this results in greater inhibiton of the alpha motor neurons
thus slowing contraction. The opposite happens when we let go of the object. The
importance of the Golgi tendon organs in our locomotion and general
proprioception can be illustrated by investigations made by Burne and Lippold,
who discovered that Parkinson's patients have a loss of inhibition on the Golgi
tendon organs. Using skin electrodes to induce electrical stimulation, they showed
that this deficit was highly likely to play a large role in the rigidity and tremoring,
which is so characteristic of the condition.
Sensors found in the joints also contribute to our proprioception. These are
found particularly in the joint capsules, which are fibrous pieces of tissue
surrounding the joints, and in the ligaments. When we instigate a movement, there
is achange in the angle, speed and direction of the joint and these are sensed by
the nearby mechanosensitive axons. Theses receptors are rapidly adapting
suggesting that a joint experiencing movement provides abundant sensory
information but one devoid of movement would not. This is believed to be
compensated for by the other areas of sensory input e.g, the muscle spindles and
the Golgi tendon organs. This is evident when viewing a patient with a titanium
hip, who is nevertheless able to experience sufficient proprioceptive input.
Opposite to the myotatic reflex, which revolves around a monosynaptic
input, the Ib fibres relaying onto alpha motor neurons from the Golgi tendon
organs are arranged in a polysynaptic manner, their actions being monitored by a
plethora of interweaved spinal interneurons. These interneurons rely upon signals
not only from the primary sensory axons but also descending fibres from the brain
and collaterals of lower motor neuron axons. If we grasp an object in our hand and
perform a flexion at the elbow, we would observe contraction of the biceps just as
much as relaxation by the triceps. If these two antagonistic actions were not
coordinated in such a way, the movement would be impossible. This reciprocal
inhibition arises from inhibitory input by the interneurons. Collaterals of the Ia
nerve fibres synapse onto these inhibitory interneurons in the spinal cord, which
go on to link with the alpha motor neurons supplying the antagonistic muscles.
The example above illustrates spinal interneurons with an inhibitory role but
they do also exist as excitatory forms of input. Such an interneuron concerns the
withdrawal of a body part from a harmful stimulus. When one steps on a pin, the
response is immediate and involves pulling one's leg away but also keeping the
other leg relaxed to maintain standing. Pain axons originating in the sole of the
foot project to the spinal cord and after substantial branching, activate several
interneurons. This leads to stimulation of the alpha motor neurons responsible for
the flexor compartments of the injured limb as well as recruitment of the
inhibitory interneurons essential in reciprocal inhibiton. Additionally, in order to
maintain standing, the crossed-extensor reflex is acitvated. Effectively, this
acitvates the extensor muscles and inhibits the flexors of the other limb. Were this
not to happen, one would fall over after every injury to a single leg.
The crossed-extensor reflex and the concept of reciprocal inhibition both
contribute to the generation of spinal motor programs for walking. In addition, the
process of walking requires some rhythmic input for the motor activity and such a
circuit is known as a central pattern generator, which is thought to originate from
the spinal cord. Sten Grillner demonstrated this notion by dissecting the spinal
cord from the jawless lamprey fish. When he electrically stimulated the axons
descending from the pain, he witnessed an alternating rhythmic activity, which
reflected the procedure during swimming. He also showed that by activating
NMDA receptors in the spinal interneurons, he could induce the rhythmic
locomotor activity evident in walking.
The stretch reflex is an essential component of the guidance governing our
muscular contractions. It is responsible for the maintenance of our muscle tone as
well as our subjective positioning in space i.e. Proprioception. The accuracy of the
stretch reflex also can be used to provide an insight into neurological health. In
upper motor neuron lesions, neurological examination can often reveal a clasp-
knife response, so named because of the similar movement of the limb when
tested. Just by observing how conditions addressing this aspect of the anatomy
can create hugely detrimental effects, illustrates how significant this reflex is in
our survival.