Muscle Interaction and Reciprocal Inhibition Part 2 by Mel Siff
Here continues the information on reciprocal inhibition and muscle
interaction from Basmajian (“Muscles Alive”).
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PART 2
Silent Period and Reciprocal Inhibition
The “silent period” of muscle is the period of cessation of activity which
occurs when a twitch contraction is superimposed on a voluntary effort (e.g.,
electrically or by a tendon tap). Its normal range (in adductor pollicis) is
87 to 151 msec (Higgins & Lieberman, 1968). It also occurs following the
sudden release of a voluntarily innervated muscle provided a certain minimum
rate of shortening is achieved; if shortening is not permitted or is slow, no
silent period results (Struppler, 1975).
Struppler has shown clearly that the silent period is NOT the result of
reciprocal inhibition (autogenic inhibition); rather it is primarily due to
the cessation of facilitatory impulses in the primary afferent fibres from
the muscle spindles. At the end of the silent period, there is a rebound
burst of EMG activity. A sudden stretch of a relaxed muscle does not recruit
a monosynaptic (muscle-spindle) reflex in man (Marsden, Merton & Morton,
1976).
Garland & Angel (1971) showed that during a rapid voluntary movement, the
agonist produces two distinct volleys of EMG activity separated by a relative
silence. When the active limb was unloaded during the movement, the second
burst was significantly reduced. Apparently, the second burst is due to
spinal reflexes. The mechanical properties of the muscle were shown by
Agarwal & Gottlieb (1972) to have a significant influence on the duration of
the silent period that follows the electrically induced H-wave in soleus
muscle. The primary contraction in soleus (M-wave) coincides with reciprocal
inhibition of the tibialis anterior.
Yabe & Tamaki (1976) have shown the voluntary elbow extension is immediately
preceded by a silent period in the unexercised contralateral agonist
(“contralateral agonist silent period”), without any contralateral antagonist
contraction occurring. Obviously this occurrence cannot be a unilateral type
of reciprocal inhibition at the spinal level.
A different kind of silent period has been demonstrated in the muscles of
mastication. Single stimulation of the teeth induces inhibition in actively
contracting temporalis and masseter muscles with a 30 to 40 msec latency. The
same type of inhibition occurs from sudden unload of the muscles (Ahlgren,
1969; Beaudreau et al., 1969; Griffin & Munro, 1969). Munro and I (1971) more
recently demonstrated that the inhibition in all the elevators of the
mandible was almost synchronous; in most subjects there was a synchronous
burst of activity in the chief depressor muscle (anterior belly of digastric)
some 15 to 27 msec after initial tooth contact.
Hannam (1972) has demonstrated that an enhancement of the masseteric reflex
by voluntary contraction of the jaw-closing muscles may be due to autogenic
factors, synergistic factors or both. Stimulation of the muscle spindles and
facilitation caused by the voluntary activity before tooth contact must be
involved.
Effects of Cross Exercise
The hypothesis that there is a transfer of activity to the contralateral limb
during prescr ibed exercise on one side has been frequently postulated, but
now it is being seriously questioned. Probably it is invalid except in very
special circumstances. Gregg, Mastellone & Gersten (1957) of Denver,
Colorado, found that overflow to the unexercised, contralateral muscles did
not occur during simple non-resistive exercises or during isometric
contractions of one biceps brachii. As the exercise stress increased,
however, there was some “overflow” to the opposite triceps and, after even
greater stress, to the biceps. Increasing fatigue played an important role in
the “overflow” but was reversible, for after a rest of two minutes “overflow”
would at first be absent. Moore (1975) found overflow activity to be between
10% and 20% of the maximal intensity of activity in the exercised limb. She
believed that even this small amount of overflow gives sufficient
justification for it to be used in maintaining muscular tone in immobilized
limbs.
Samilson & Morris (1964) confirm the finding that in normal man activity of
one upper limb is not accompanied by activity in the contralateral resting
limb. However, in spastic children, there is such a spread. On the other hand
Podivinsky (1964) of Bratislava, Czechoslovakia finds a slight motor
irradiation occurs from the strong contraction of finger flexors to the
related muscles of the opposite limb (“crossed motor irradiation”).
This perhaps is related to the findings of Hellebrandt and her colleagues
regarding indirect learning, i.e., the improvement of strength in one limb by
exercising the opposite limb (Hellebrandt & Waterland, 1962a, b). Its
practical significance in ordinary life is unknown and appears to have been
exaggerated since the days of Scripture et aL (1894). We have shown that at
the finest levels of control in motor unit training the role of
cross-training is not significant (Basmajian & Simard, 1966).
Further, the crossed reflex phenomenon described by Ikai (1956) of Tokyo is
not really the same phenomenon as cross exercise. Ikai showed that the
crossed reflex of limbs in spinal animals can be reproduced under certain
conditions as a brief overflow of monosynaptic reflexes to the opposite limb.
Panin, Lindenauer, Weiss & Ebel (1961) seem to have delivered a serious blow
to the concept of “cross exercise.” In their extensive study they found that
the spread of activity was minimal to insignificant. Insignificant potentials
of low amplitude and frequency appeared in all non-exercised muscles in a
widespread distribution in all four limbs. They appeared most in areas
required for postural stabilization of the subject’s body. Even then the
amount of activity was so slight as not to constitute exercise effect.
Our own studies on quadriceps (p 250) and those of Sills and Olsen (see
above) largely confirm the conclusions of Gregg and his colleagues. We found
in our studies of spastic patients (p 83), however, that an exuberant
overflow occurs to the opposite limb. Walshe (1923) and, more recently, Hopf
et al. (1974) and Soto et al (1974) have written about a similar phenomenon
in hemiplegia. We must conclude that “cross education” is, at best, of
dubious value in normal subjects……
Motor Learning and Control
There is mounting evidence that motor learning and control are not a process
of accretion but depend on patterning of inhibition in motor neurons. Elect
rom yo graphic studies in health and disease indicate that the acquisition of
skills occurs through selective inhibition of unnecessary muscular activity
rather than the activation of additional motor units.
As noted in other sections, almost all resting muscles throughout the bodies
of adults, both human and general mammalian, fall to a level of neuromuscular
silence. This total relaxation occurs unless the muscles are needed to be
tensed for a posture or movement or unless the person suffers from
uncontrolled apprehension or neurotic and neurological disturbances. With
this in mind, MacConaill and I (1969) enunciated the principle that there
should be a minimal expenditure of energy consistent with the ends to-be
achieved. This self-evident principle embraces two laws:
1. The Law of minimal spurt action – no more muscle fibres are brought into
action than are both necessary and sufficient to stabilize or move a bone
against gravity or other resistant forces, and none are used insofar as
gravity can supply the motive force for movement;
2. The law of minimal shunt action – only such muscle fibres are used as are
necessary and sufficient to ensure that the transarticular force directed
toward a joint is equal to the weight of the stabilized or moving part
together with such additional centripetal force as may be required because of
the velocity of that part when it is in motion.
Control of Movement
The neurophysiological literature is encrusted with the barnacle that “the
brain does not order a muscle to contract but orders movement of a joint.”
Recently, Phillips (1975) made a concerted effort to dispel this myth which
has stultified research on the learning of motor behavior. In fact, the best
movements are performed with an economy of muscular movements dependent upon
impulses being sent to only one or two muscles or even a localized area of
one muscle. What the brain has “learned” is patterning of these actions by
means of a progressive inhibition of the inefficient mass responses that were
natural to the child.
Some movements are extremely economical in the well-trained person. For
example, most of us are fairly well-trained in turning our hand over through
pronation and supination of the forearm; in this learned act our nervous
system calls upon only one or two muscles to produce the movements.
Fortunately, the normally plastic human brain quickly adapts to shifts of
function; otherwise tendon-transfer operations would be useless.
Physiologists and even some kinesiologists do not appreciate that each and
every muscle has several (sometime many) component parts which are recruited
in different functions at different times. Many investigations with
intramuscular electrodes in many thousands of muscles lead me to believe that
this local activity is patterned by progressive inhibition of motoneurons
until an acceptable performance is achieved. Our studies of elbow flexion and
thenar muscles, which show the interplay of motor unit functions dedicated to
specific postures and movements, clearly indicate that the positioning of
limbs is predetermined by sets of motor units which are permitted to act for
that position. The same appears to be true for welllearned movements.
I believe that a mosaic of spinal motoneurons is dedicated to the learned
response of a specific posture or movement of a joint through space. The
ultimately superior performance of a skilled movement depends on the
reproducibility of the ideal, an economically spare mosaic of motoneuronal
activity (Basmajian, 1977).
With different objects in mind, Payton et al. (1976) put it slightly
differently: they found no statistically significant difference between
pre-learning and post-learning of a simple task in regard in the EMG
activity, movement time and range of movement. They concluded that all the
prime movers that are going to contribute to the final learned act take part
even before the skill is learned; thus as motor learning takes place, there
is a marked reduction of activity only in the auxiliary muscles while the
prime movers neither gain nor lose (Payton et al., 1976).
When I first described the precision possible in controlling single
motorneutons, I believed (as did many others) that this type of control was
the building block of motor performance. Given visual and auditory cues
through electronic amplification and feedback, subjects could be quickly
trained to consciously activate single motoneurons with great precision. But
conscious activation of single motoneurons in the single-motor-unit training
paradigm depends on the same principles as the learning of any other novel
task, that is, progressive (and sometimes rapid) inhibition of the
motoneuronal activity that adds no useful function in producing a desired
motor response (Smith, Basmajian & Vanderstoep, 1974).
Training, whether it is the unconscious process of the child learning simple
social motor responses or the preparation for a specific skilled act (such as
those of a musician or athlete), is a progressive inhibition of many muscles
that flood into play when one first attempts to produce the required
response. The athlete’s continued drill to perfect a skilled movement
exhibits a large element of progressively more successful repression of
undesired contractions. Among others, O’Connell (1958) has demonstrated this
convincingly. A group of physical education majors required to perform “head
stands” while being studied electromyographically could be graded as to their
actual experience by the amount of overflow of undesired activity in muscles
that were only casually related to the exercise.
The young animal has enormous amounts of overactivity and reactive
contractions in muscles that are serving no directed purpose in producing the
desired movement or posture. Among others, Janda & Stara (1965) demonstrated
in children a high incidence of mass responses in a predictable pattern even
in muscles that are far removed from those which produce a required movement.
As children mature, this overactivity disappears and is absent in normal
adults. It reappears in adults under psychological stress, but people can be
trained to inhibit it to varying degrees. In patients with diseases and
injuries of the central nervous system, the normal inhibition pattern is
lacking; then mass responses from local interoceptive and exteroceptive
bombardments of the motoneurons result in an exaggerated mass response
described as spasticity.
The Moscow investigators led by Yusevich (see, for example, Okhnyanskaya et
al., 1974) attribute normal motor hyperactivity in infants and children to
synkinesis or synergies of suprasegmental origin, pointing out the fact that
they normally disappear by the time a person is adult.
The patterning of the inhibition would seem to come in part from obscure
processes in diffuse centers of the cerebral cortex; since inhibition is a
central feature, one must consider the possibility that brainstem centres and
perhaps the cerebellum are critically important in the imprinting of the
learning. It is too simplistic to consider a schema where an impulse is
started at a tiny area of the cerebral cortex and is thence passed directly
along a facilitatory path to a desired set of motoneurons. The motor learning
process probably employs a neuronal network with the “main” pathway for motor
activation being almost a small part of the whole.
Proprioceptive Effects
Gellhorn (1960) has described electromyographic studies which disclose the
effects of central proprioceptive influences on movements elicited by the
electrical stimulation of the motor cortex. Movements so produced are
strongly reinforced by proprioceptive impulses which also determine, by and
large, the type of movement that results. He showed, for example, that the
contraction of triceps and flexor carpi muscles when stimulated through the
cerebral cortex is greater if the elbow is at 45 degs than if it is at 110
degs or 160 degs. Furthermore, a cortical stimulus that is below threshold
when a muscle is slack may become effective when the muscle is put on the
stretch…………
END OF SERIES
Mel Siff
