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Describe the neural mechanisms underlying the

stretch reflex. How might this contribute to the


control of posture and movement?
The importance of the stretch reflex has been a subject of some debate and revision.
The stretch reflex was originally identified by Sherrington, who deciphered much of
the relevant anatomy of the reflex and personally instigated a paradigm shift in how
we think about them. Whereas it was previously thought that the stretch reflex was an
intrinsic property of the muscle, Sherrington demonstrated that it was abolished if
either the dorsal or ventral roots were severed, thus elegantly proving a neural
involvement. In 1906 he proposed that reflexes, among them this reflex, served as the
basis for all movement. This stemmed from the observation that a decerebrate cat, that
is to say a cat where only reflexes are seen since all descending control from the brain
is abolished1, can stand upright (though with about as much stability as a stuffed cat),
and can scratch itself, or walk on a treadmill. Graham Brown, on the other hand, who
was working at much the same time, found that cats also retained this ability if their
dorsal roots were severed, abolishing all sensory feedback and thus all reflexes. So
while the stretch reflex is important for the control of posture and movement, it is
important to see it in context as part of a much bigger picture. A more modern view of
the usefulness of the stretch reflex is as controlling the length of a muscle. It can be
seen as working hand in hand with the opposing Golgi tendon organ reflex: the
tension reflex, controlling tension. This is because between them, these two reflexes
control length/tension of a muscle, that is to say its stiffness.

What exactly is a reflex? The traditional view, at least according to Kandel and
Schwartz, was that reflexes are “automatic, stereotyped movements in response to
stimulation of peripheral receptors”. But given that they are variable and subject to
descending control, this cannot be the entire story. For example, clinically, where
stretch reflexes are a mainstay of the neurological examination, if one cannot elicit
stretch reflexes from one’s patient very well, one can ask them to close their eyes and
clench their firsts. This tends to increase the magnitude of the response.
Experimentally it has been found that the reflex action, and even the muscle groups
involved, can be modified depending on the functional set. This is particularly true for
conditioned reflexes. For example, one can train a subject to extend their wrist in
response to an audible tone, to avoid an electric shock, a conditioned flexion
withdrawal reflex. If the subject then turns their hand over, and the audio tone is
repeated, then the subject will normally flex their wrist to remove it from the
electrode and avoid the electric shock. So the purpose of the reflex action remains the
same, to avoid the electric shock, but the muscle groups involved (flexion instead of
extension), have changed completely. What can we conclude? Kandel and Schwartz
state that “reflexes are highly adaptable and control movements in a purposeful
manner”. Perhaps the stretch reflex is in a sense more fundamental than some of the
other reflexes (e.g. conditioned reflexes) and so is less mutable. Nonetheless,
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The cut is made in the midbrain, between the superior and inferior colliculi. These animals have
stereotyped and heightened stretch reflexes (indeed stereotyped and heightened spinal reflexes more
generally). And there is a significant increase in extensor muscle tone, accounting for the stuffed-
animal-like characteristics. In the absence of descending influence from the cortex, descending
influence from the brain stem greatly facilitates the stretch reflex. In normal animals, on the other hand,
the strength of the stretch reflex is much more variable since it is subject to control from the cortex.
examples of how it can be overridden have been demonstrated experimentally, and
will be discussed later. Anatomically, the stretch reflex in common with many other
reflexes requires “interneurons”. Descending motor cortical neurons will typically
synapse on these interneurons, and can thus modulate the stretch reflex. This provides
the anatomical link between movement, posture (both of which from introspection
and experience we “know” can be modulated consciously), and reflexes.

The stretch reflex is the simplest spinal reflex, and consists of the contraction of a
skeletal muscle when it is stretched. There is a brief phasic phase (as with the knee
jerk) and a tonic phase of contraction. When the muscle stretches, this is sensed by the
muscle spindle. The nerve fibres are stretched, and as they stretch, increase their firing
rate. The muscle spindle has two types of endings: bag and chain. Ia and II afferents
supply the bag, ending in spiral endings. These neurons are rapidly adapting, and fire
in response to the change in stretch. The type II neurons supplying the chain
(flowerspray endings) are slowly adapting, signalling the extent of the deformation.
The muscle spindle has a non-contractile centre, where either the bag or chain is
found. At the ends of the muscle spindle there are contractile regions. These are the
gamma efferents. They come in two types: static (innervate both bag and chain fibres)
and dynamic (innervate only the bag fibres). Firing of the dynamic efferents increase
the dynamic sensitivity of the dynamic afferents, while firing of the static efferents
stretches the fibre, so that when sustained contraction does occur, feedback from the
spindles does not cease altogether.

The stretch reflex typically involves multiple connections. Ia afferents excite motor
neurons of the same muscle2. This is termed homonymous excitation. In addition, they
excite motor neurons of synergist muscles, which is heteronymous excitation. The
excitation involves glutamate, because transmission is blocked by glutamate
antagonists. The EPSP is partially blocked by NMDA antagonists (ketamines are an
example). Does this imply a role for learning/LTP at a reflex level? So broadly
speaking, excitation of the Ia afferents, as occurs during stretch, leads to contraction
of the muscle that is stretched. Synergists also contract: it is useful to think of a reflex
action, that is to say, movement with a purpose. There is sometimes also a tonic
reflex. If a muscle is stretched and then held at a new length, there is sometimes a
tonic phase of reflex contraction. This is obvious in decerebrate cats, and is
hypothesized (Pierrot-Deseilligny 1990, Hagbarth et al. 1973, Delwaide et al. 1991) to
play a role in some disorders of excess muscle tone in humans (spasticity). Thus, the
stretch reflex is a negative feedback system, in that its action, contraction, serves to
reduce the initial stimulus, that is to say, stretch.

In addition to these exciting observations, there is also an inhibitory aspect to the


stretch reflex, whereby antagonistic muscles are inhibited. There are several different
circuits for this. The simplest is where the Ia afferent excites a Ia inhibitory
interneuron (glycine, for which the snake venom strychnine is an antagonist), which
then inhibits the motor neuron. As mentioned earlier, this interneuron is also acted on
by descending spinal pathways, and in the presence of local anaesthesia these
pathways are sufficient to excite it (Nielsen et al. 1992, 1995). Furthermore, every

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This excitation occurs directly. This was determined by measuring the latency of the response: how
long does it take from excitation of the dorsal roots for their to be an impulse in the ventral roots? The
shortness of the time: less than 1 ms, implied that only one synapse could be involved, and therefore
the excitation was direct: each synapse delays the signal by up to 0.9 ms.
motor neuron has a corresponding pool of Ia interneurons (IaIN’s) that are co-
activated with it. They tend to inhibit antagonistic motor groups. This is the principle
of reciprocal innervation, again established by Sherrington. So the excitation of the
motor neurons by the Ia afferents in the direct reflex pathway could also increase
activity in the Ia interneurons. It must be emphasized that these interneurons are very
important. They play a significant role in voluntary movement, in addition to the
stretch reflex. They receive collaterals of the excitatory motor neurons supplying their
antagonist muscles. If the cortex signals a movement, there is no need to separately
inhibit the antagonists of the muscles involved: the principle of reciprocal innervation
will do this automatically. That said, there is not a stereotyped inhibition of a fixed
degree by IaIN’s on antagonistic motor neurons, but the degree of inhibition will
depend on the context3. It is sometimes advantageous to contract the prime mover and
antagonist simultaneously, since it will increase the stiffness of the joint. For example,
antagonist motor neurons need to contract for activities such as the precision grip. The
antagonist and agonist muscles of the elbow joint will both contract just before
catching a ball4. Descending spinal control of these reflexes is absolutely essential.

Earlier, the gamma efferents were discussed. How do these relate to movement? In
1953 Merton proposed the servo control theory (servo is Latin for “by the slave”).
This stated that the gamma efferents fire to set the length of the muscle spindle to the
desired length of the entire muscle. Assuming the two are different, this will activate a
muscle stretch reflex via the alpha motor neurons, to make the entire muscle contract
to this length. There are a number of reasons why this has to be wrong. First of all, it
does not take the force we are acting against into account. For example, if we are
going to pick up an object, and it turns out to be heavier than expected, we do not just
activate a greater stretch reflex and pick it up, being surprised by the amount of force
we end up using! If everything worked by stretch reflexes, though, this is what we
would expect. Instead, we have to stop, and pick it up again, using the new
information. Even more conclusively, this theory relies on the gamma efferents firing
before the alpha fibres. But detailed measurements show that actually the two activate
synchronously5. This is termed alpha gamma coactivation, and means that as the
muscle shortens (tending to reduce any stretch reflex) the spindle also shortens
(maintaining the sensitivity of the stretch reflex). This is called “servo assistance”. In
addition to this, beta efferents that run from alpha motor neurons to innervate the
spindles are also found: these are part of the so called skeleto-fusimotor system, and
enforce this motor fibre-spindle coactivation.

This last point is interesting, since it shows that the nervous system has already
evolved a system for co-activation of muscle fibre and spindle (which serves to keep
the spindles sensitive). So why is there this capability for the separate excitation of the

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These interneurons receive descending input from most of the spinal tracts, in particular the
corticospinal tract. Another important source of the variability in the strength of their activation is the
Renshaw cells, which produce recurrent inhibition of the motor neurons that stimulate them (negative
feedback), and also influence Ia interneurons and synergist motor neurons, thus providing a site for the
control of all the processes being discussed.
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Although when taught how to catch, for example a cricket ball, one is told to consciously relax the
joints to slow the ball down, and diminish the force acting on the hands. Another example of the
conscious modification of these reflexes?
5
In the 1960’s Vallbo and Hagbarth used the technique of microneurography, which they invented, to
demonstrate that as the muscles of the finger slowly contracted, output from the Ia afferents actually
increased. The only way to explain this was alpha gamma coactivation.
spindles, via the gamma efferents? No-one yet has a definitive answer, but what
follows are some potential uses (from Zigmond and Bloom). The strength of gamma
activation seems to depend on the task. In slow movements, feedback can be useful to
counteract fatigue or to alter ongoing movements (a sort of servo-control). Gamma
activation in these circumstances is high (Hulliger et al. 1989, Schieber and Thach,
1985, Hagbarth, 1993), although Kandel and Schwartz claim that gamma activation
tends to decrease in these circumstances. When walking on a balance beam, or doing
some other difficult task, it is useful in general to suppress stretch reflexes, and
gamma activation is suppressed (Zigmond and Bloom. Kandel and Schwartz quote
Prochazka et al. 1988, and again seem to say the opposite!) Presumably, this gamma
activation provides one way of altering the stretch reflex, at least in part. Certainly,
the stretch reflex and its role in movement are affected by conditioning. In Royal
Danish ballerinas, calf reflexes seem weaker than in the less well trained population:
ballerinas presumably need more control over these muscles, and cannot be perturbed
by excessive stretch reflexes. (Nielsen et al. 1993)

In addition to the interneurons, there are other sites at which the stretch reflex can be
modulated. Two examples are the alpha motor neurons themselves, and the
presynaptic terminals of the Ia afferents. Descending pathways terminate at both these
sites. For example, tonic excitation of the alpha motor neurons causes their resting
potential to depolarise, so they need less stimulation to fire. Input presynaptically can
increase or decrease the probability of transmitter release. This control can be tonic
(depending on the functional set), or dynamic, allowing the stretch reflex to be
modulated throughout the movement. A good example of the use of the latter is
during walking, when the strength of the presynaptic input is continually being
modulated. This sets the sensitivity of the stretch reflexes to change during each step.

This is all very nice and interesting, but how does it relate to voluntary movement?
This control of the amplitude and nature of the stretch reflex allows the body to use
the stretch reflex to support voluntary movement. Key in this is to move away from
thinking of the stretch reflex as merely a reflex, and more as a system for feedback
and control. When we activate a stretch reflex, there is the initial phasic contraction,
whose physiology has been discussed above. This is termed the M1 response. Later,
there is a second contraction, the M2 response. This still occurs too rapidly for it to be
the result of voluntary activity6, and it transpires that this reflex passes through the
cortex: the so called long loop reflex7. This is the mechanism by which the stretch
reflex can provide feedback during voluntary movement. They are particularly
important for control of fine, distal muscles and movements, whereas the medial
musculature, typically concerned with posture, makes do with the subcortical control.

Another potential use for the stretch reflex in voluntary movement was proposed by
Ragnar Granit in 1963, and draws on the concept of servo assistance, as well as
negative feedback, discussed earlier. He suggested that the motor cortex output
merely sets the desired position of the limbs, and signals this via both the alpha and
gamma efferents. The stretch reflexes serve to attain this position. For example, if the

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Although the SMA provides a conscious gating mechanism for this reflex.
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This was shown by patients who have Klippel-Feil syndrome. In this, the neurons from the motor
cortex bifurcate, and innervate motor neurons on both sides of the body. In these people, part of the
stretch reflex serves to activate the contralateral muscles as well as the ipsilateral ones one would
expect, so the motor cortex must be involved in the reflex.
contraction of the muscle is excessive, and it overshoots, there is insufficient stretch
acting on the stretch receptors, they stop firing, and this can be taken as a signal to
relax the muscle somewhat. Conversely, if the contraction brought about by the alpha
efferents is insufficient, the stretch reflex fires to increase its contraction. In theory,
this would help in overcoming unexpected loads. When picking up a cup of coffee, if
there were no such stretch reflexes we would have no way of knowing (at least not
sufficiently quickly) that the force of contraction was insufficient to bring the cup to a
horizontal position, due to the weight of the mug. So it would spill. Servo assistance
provides a way of getting around this thorny issue. But there are limits, as the suitcase
example given above demonstrates.

These then are some of the uses of the stretch reflex in voluntary movement.
Intuitively, it would seem that it is also useful in this straightforward manner in the
retention of posture. For example, when we stand up, if for example a calf extensor
muscle is stretched, then it would make sense for a stretch reflex to cause it to
contract, thus regaining posture. But things are not so simple. In 1976 Nashner
performed an experiment to demonstrate this. Volunteers were asked to stand on a tilt
table. The table would move backwards. Because of their inertia, the volunteer’s body
swayed forwards, and the calf extensor muscle was stretched. The stretch reflex then
is altogether appropriate in bringing one’s centre of gravity above one’s feet. But
then, the table tilted backwards, again stretching the calf extensor muscles. Here, a
stretch reflex is totally inappropriate, since it would throw you backwards still further.
Nevertheless, on the first tilt after the backwards movements, the stretch reflex
operated (as measured by the EMG for the muscle) and the volunteers tended to fall
off the tilt table. So the stretch reflex is not always useful for maintenance of posture.
Furthermore, on subsequent repetitions of the table tilting backwards, the stretch
reflex was suppressed, and subjects remained on the tilt table.

Thus it seems that the traditional idea of the control of the posture: antagonistic
vestibular and cervical reflexes, acting on the extensor muscles via the spine and on
the neck (to decrease the vestibular stimulation at source) to retain posture regardless
of head position in relation to the body, are more important. Although of course
control of posture is extremely complicated, involving multiple interacting factors,
and lying well outside the scope of this essay, one interesting question is to what
extent the cervical reflexes can be regarded as modified stretch reflexes. Very briefly,
input from both vestibular organs and peripheral proprioceptors combine to generate
reflexes which aid in the maintenance of posture. The vestibular organs sense
displacement of the head, and so head movement in a particular direction will excite
vestibular afferents, which will in turn excite central neurons. These will excite motor
neurons in the body or neck, and will stimulate movement of the head in the direction
opposite to that which caused the original stimulus. Thus there is a negative feedback
system at work. The vestibular collic reflex acts on the neck. It would be unfair to
regard this as a bona fide stretch reflex: after all the original stimulation is taking
place at a different site from the response, unlike the stretch reflex8. The vestibulo
spinal reflex affects the rest of the body musculature, and as such is clearly not a
stretch reflex.

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In addition to this role of the vestibular reflexes in stabilising head and body, they are most sensitive
to displacement at 2-3 Hz, which coincidentally is the natural resonance frequency of the head and
neck. So they damp this natural resonance!
The cervical reflexes, however, have sensory input from the muscle spindles of the
neck muscles. These spindles also participate in neck muscle stretch reflexes, which
clearly have some role in the maintenance of posture since they directly determine
where the vestibular systems are and how they are moving! The cervical spinal reflex
tends to oppose the vestibulo spinal reflex: for example moving the neck downward in
a quadruped tends to cause flexion of the front limbs via the cervico spinal reflex,
while the same movement acting on the vestibular system would cause their
extension. The end result of all this is to make the orientation of the neck muscles
irrelevant to posture, and only the orientation of the body important. But the
orientation of the body has to be sensed, at least in part, by the vestibular system, and
for that we need the proprioceptive information about the position of the neck to turn
the head centred information from the vestibular system into body centred
information useful for posture. The cervico collic reflex is thus very important in
stabilizing the head in relation to the body, by opposing neck muscle stretch. And how
does this relate to the humble stretch reflex? The short answer is that we don’t know.
There are certainly homonymous monosynaptic stretch reflexes, but similarity in
response between the cervicocollic and vestibulocollic reflexes implies that there are
also central mechanisms involved in the cervicollic reflex. So it is not as simple as a
straightforward stretch reflex.

Thus the stretch reflex is one of the oldest and most distinguished research areas in
neurophysiology, bearing the stamp of Sherrington. In recent times, it seems to have
been rediscovered and re-evaluated, and is “rearing its head” in many aspects of the
control of movement and posture.