FUNCTIONAL NEUROANATOMY AND

PHYSIOLOGY OF THE MASTICATORY SYSTEM

69
Function of the Sensory Receptors

The dynamic balance of the head and neck muscles previously described is
possible through feedback provided by the various sensory receptors. When a muscle is
passively stretched, the spindles inform the CNS of this activity. Both the Golgi tendon
organs and the muscle spindles monitor active muscle contraction. Movement of the
joints and tendons stimulates the pacinian corpuscles. All of the sensory receptors are
continuously providing input to the CNS. The brainstem and thalamus are in charge of
constantly monitoring and regulating body activities. Information concerning nomal
brought into the
body homeostasis is dealt with at this level, and the cortex is not even
to
regulatory process. If, however, incoming information has significant consequence
for conscious evaluation
the person, the thalamus passes the information to the cotex
influence on function
and decision. Therefore the thalamus and brainstem have powerful
of the individual.

Reflex Action69

stimulus that passes as an

A reflex action is the response resulting from a
root or its cranial equivalent,
impulse along an afferent neuron to a posterior nerve
leading back to the skeletal mus
where it is then transmitted to an efferent neuron
higher centers, the response is
cle. Although the information is sent to the
or brainstem influence. A
independent of will and occurs normally with no cortex
monosynaptic reflex occurs
reflex action may be monosynaptic or polysynaptic. A
the efferent fiber in the CNS. A
when the afferent fiber directly stimulates
neuron stimulates one or more
polysynaptic reflex is present when the afferent
efferent nerve fibers.
interneurons in the CNS, which in turn stimulate the

system: (1)
Two general reflex actions are important in the masticatory
masticatory
myotactic reflex and (2) nociceptive reflex. These are not unique to the
as well.
muscles but are found in other skeletal muscles

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yotactic reflex. 69

The myotactic or stretch reflex is the only monosynaptic jaw reflex. When
skeletal muscle is quickly stretched, this protective reflex is elicited and brings
bout a contraction of the stretched muscle.

The myotactic reflex can be demonstrated by observing the masseter while

sudden downward force is applied to the chin. This force can be applied with a
small rubber hammer. While the muscle spindles within the masseter suddenly
stretch, afferent nerve activity is generated from the spindles. These afferent
impulses pass into the brainstem to the trigeminal motor nucleus by way of the
trigeminal mesencephalic nucleus, where the primary afferent cell bodies are
located. These same afferent fibers synapse with the alpha efferent motor neurons
leading directly back to the extrafusal fibers of the masseter.

Stimulation of the alpha efferent fibers by the la afferent fibers causes the
muscle to contract. Clinically, this reflex can be demonstrated by relaxing the jaw
muscles, allowing the teeth to separate slightly. A sudden downward tap on the
chin will cause the javw to be reflexly elevated. The masseter contracts, resulting in
tooth contact.
eof te spindie
pat
es. Thie leses

es the sens Trigeminal

mesencephalic nucleus
Spindle afferent
fiber
Trigeminal main
sensory nucleus
Iy-Efferent
fiber
Muscle spindle
Trigeminal
motor nucleus

a-Efferent fiber

Jaw-closing muscle

72
 Faefin

Ihe yotattic reficx sces ithut scif resse fnon ie cortex srd
is very impoMan in determining the resting ositin of the jaw. If there were
eomplete reiaxation of ai the msc kes that sappot the jaw, the forces of gravity
wuid wct to lower he jaw and separate the articiar sarfaces of the TMJ. To
prevent this disiocatiTm, the elevtOR Cies (and other mscies) are maintained itn
a mild state of contraction called mascie tonas. This pruperty of the cievator mus
cles coutrteracts the eftect of gravity om the manditie ard maintains the articuiar
sarfaces f the jovnt in comstar COniact. The myotactic retlex is a principai
deerminast of muscle tos in the cicVor muscies. While gravity pulis down on
the mandibie, tthe eievatos mles ae passively stretchei, which aso creates
stretching of the mscle spies. This infornmation is reflexly passed from the
afierent eurns cenginstig in spinds te the alpa motor neurons that tead
hack to the extrafusad bers of the eevôor auscies, Thus pasRIve stretehing causes
reative contraction that neleves tôe strenh ot the ie spindie. Muscie tonus
can also be infsensed By aferex inpu mcthe? sesory receptors, suxh as
those from the skin o te crai a s

The myotatis reflex dresuting msc le tcOmus can alse be influenced by

the highcr centers vis the fusiotor system The curtex and brainstem çan bring
about increased ganma eflerent activisy o the intrafusal fibers of the spindle.
While this activity inceascs, the inirafkaal tibers contract, cusing a partial
stretching of the nuclear hag asd tucicar chain areas of the spindies. This lessens
the atnouni of stretch nedod in te overall muscle before the spindie afterent
activity is elicited. Thetefore the higher cesters can use the fusimotor system to
altcr the sensitivity of the muscle spindies to stretch. Increased gamma efferent
activity increases the sensitivity of the myotactic retlex, whereas decreased garmma
efierent activity decreases the sensitivity of this reflex.

When a muscie contracts, the muscle spindles are shortened, this causes the
affenent activity output of these spindles to shut down. If the electrical potential of
the afferent nerve activity is monitored, a silent period (no electrical activity) will
be noted during this contraction stage. Gamma efferent activity can influence the
kength of the silent period. High gamma efferent activity causes contraction of the

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 Functional Neuroanatomy and Physiology of the Masticatory system

intrafusal fibers, which lessens the time the spindle is shut down during a muscle
contraction. Decreased gamma efferent activity lengthens this silent period.
69
Nociceptive reflex.

The nociceptive or flexor reflex is a polysynaptic reflex to noxious stimuli

and therefore is considered to be protective. Examples are present in the large
Jimbs (e.g., withdrawal of a hand when it touches a hot object). In the masticatory
system this reflex becomes active when a hard object is suddenly encountered
during mastication. When the tooth is forced down on the hard object and the
periodontal structures are overloaded, a sudden, noxious stimulus is generated. The
primary afferent nerve fibers carry the information to the trigeminal spinal tract
nucleus where they synapse with interneurons. These interneurons travel to the
trigeminal motor nucleus. The motor response taken during this reflex is more
complicated than the myotactic reflex in that the activity of several muscle groups
the
must be coordinated to carry out the desired motor response. Not only must
but
elevator muscles be inhibited to prevent further jaw closure on the hard object,
away from
also the jaw opening muscles must be activated to bring the teeth
potential damage.

Efferent (motor) neuron

Trigeminal
motor nucleus

Interneurons

Afferent
Trigeminal (sensory)
spinal tract neuron
nucleus

receptors reaches the

As the afferent information from the sensory
interneurons, two distinct actions occur:

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 Functinnal Neuroanatomy and Physiology of the Masticutory system

control of mandibular movement to be

achieved. For the skeletal relationship of the
skull, mandible, and ncck to bc
maintained, each of the antagonistic muscle groups
must remain in a constant state of
mild tonus. This will overcome the
skeletal
imbalances of gravity and keep the head in what is termed the
postural
previously discussed, muscle tonus plays an important role in the position. As
position, as well as in resistance to any passive mandibular rest
Muscles that are in full contraction activate most of
displacement of the mandible.
the muscle fibers that can
compromise blood flow, resulting in fatigue and pain. By contrast,
muscle tonus
requires contraction of a minimal number of muscle fibers
and the contracting
fibers are constantly being rotated. This type of activity
allows proper blood flow
and does not producce fatigue.

Regulation of Muscle Activity 69

To create a precise mandibular movement, the CNS must receive

input
from the various sensory receptors through the aferent fibers. The brainstem and
cortex must assimilate and organize this input and elicit appropriate motor activities
through the efferent nerve fibers. These motor activities involve the contraction of
some muscle groups and the inhibition of others. It is generally thought that the
gamma cfferent system is permanently activated, though it docs not necessarily set up
movement. The gamma discharge keeps the alpha motor neurons reflexly prepared to
receive impulses arising from the cortex or directly from the afferent impulses of the
spindles. Most mandibular movements are probably controlled by a link between the
gamma efferents, the spindle aflerents, and the alpha motor neurons. This combined
output produces the required contraction or inhibition of the muscles and allows the
neuromuscular system to keep a check on itsclf.

Various conditions of the masticatory system greatly influence mandibular

movement and function. The sensory receptors in the periodontal ligaments,
periosteum, TMJs, tongue, and other soft tissues of the mouth continuously
feedback information, which is processed and used to direct muscle activity. Nox
ious stimuli are reflexly avoided so that movement and function can occur with
minimal injury to the tissues and structures of the masticatory system.

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 Functional Neuroanatomy and Physiology of the Masticatory ystem

Influence from the Higher Centers

As previously mentioned, the brainstem and cortex function together to

assess and evaluate incoming impulses. Although the cortex is the main determiner
of action, the brainstem is in charge of maintaining homeostasis and controlling
normally subconscious body functions. Within the brainstemn is a pool of neurons
that controls rbythmic muscle activities, such as breathing, walking, and chewing.
This pool of neurons is collectively known as the central patern generator
(CPG).10-13. The CPG is responsible for the precise timing of activity between
antagonistic muscles so that specific functions can be carried out. During the
process of chewing, for example, the CPG initiates cotraction of the suprahyoid
and infrahyoid muscles at the precise time the elevator muscles are told to relax.
This allows the mouth toopen and accept food. Next the CPG initiates contraction
of the elevator muscles while relaxing the suprahyoid and infrahyoid muscles,
producing closure of the mouth onto the food. This process is repeated until the
particle size of the food is small enough to be swallowed easily. For the CPG to be
most efficient, it must receive constant sensory input from the masticatory
structures. Therefore the tongue, lips, teeth, and periodontal ligaments are
constantly feeding back infomation that allows the CPG to determine the most
appropriate and efficient chewing stroke. Once an efficient chewing pattern that
minimizes damage to any structure is found, it is learned and repeated. This
learned pattern is called a muscle engram.

Therefore chewing can be thought of as an extremely complex reflex

sensory
activity that is primarily controlled by the CPG with input from numerous
activity that
receptors. Like many other reflex activities, chewing is a subconscious
walking are other
can be brought to conscious control at any time. Breathing and
but can also be
CPG reflex activities that generally occur at subconscious levels
brought under voluntary control at will.

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