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Craniofacial Muscles

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13 Tongue Biomechanics and Motor Control

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enough sizes to fi t through the pharynx and esophagus. All of the tongue muscles

are active in manipulating the food into the correct position for the teeth. The SG

and the PG lifts the posterior tongue to close off the pharynx while the intrinsic

muscles manipulate the bolus. If the food moves too quickly to the pharynx or a

tongue depressor depresses the tongue and stimulates the oral pharynx, the gag

re fl ex is stimulated. The vagus nerve senses the stimulation and makes a monosynaptic

connection with the hypoglossal nerve.

The pharyngeal phase propels the bolus to the esophagus while coordinating

inhibition of respiration, closing the palatopharyngeal isthmus, and constricting the

larynx. The genioglossus and the anterior digastric may be active initially, followed

by the mylohyoid, stylohyoid, and geniohyoid, working to lift the hyoid anteriorly

(Ono et al. 2009 ) . Finally, the posterior tongue is depressed to form the anterior wall

of the pharynx as the pharyngeal muscles take over in the effort to propel the bolus

toward the esophagus.

It is also important to consider the role of the mandible during eating and speech.

Obviously, several of the muscles attach to the mandible and so need the stability of

the bone on which to move the tongue. The temporomandibular joint facilitates the

synchronicity of anteroposterior movement of the mandible and tongue but limits

horizontal movement. The hyoid, on the other hand, moves with the tongue to produce

large amplitude horizontal movements.

There has been a lot of exploration of a masticatory CPG focused on the medial

bulbar reticular formation, with most of the studies using in vitro isolated mammalian

brainstem preparations (Katakura et al. 1995 ; Nakamura et al. 1999 ) . It became

apparent that sucking rhythm generators coupled with cranial nerves V, VII, and XII

were initially necessary to coordinate suckling during late gestation and infancy

(see Barlow 2009 for review). At birth, the system has to develop an interaction

between the swallow and a protection of the airway. Once again, sensory experience

is important during this critical period to develop a brainstem pattern for swallowing

pro fi ciency. As the infant develops, the tongue begins to discern the size of

the food bolus and how to move it to the back of the tongue for swallowing. The

more mature patterns of mastication involve the rhythmic opening and closing of

the jaws but rhythmical bursts also occur in hypoglossal motoneurons (Dellow and

Lund 1971 ) .

13.5.4 Speech

Manipulation of the tongue during speech is highly variable depending on the language,

acoustic feedback, and learned pattern of speech. Therefore the movements

of the muscles are extremely complex and only somewhat follow a pattern. Models

of the tongue and vocal tract have been enhanced by computer power. Speci fi cally,

the models have tried to relate muscle recruitment and tongue shape (Perkell and

Zandipour 2002 ) . More recently 3D biomechanical models of the tongue and oral

cavity have provided a view of the activity of parts of muscles rather than whole

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