Propulsion and Mixing of Food in the Alimentary Tract

Voluntary Stage of Swallowing

When the food is ready for swallowing, it is “voluntarily” squeezed or rolled posteriorly into the pharynx by pressure of the tongue upward and backward against the palate, as shown in Figure 64-1 . From here on, swallowing becomes entirely—or almost entirely—automatic and ordinarily cannot be stopped.

Involuntary Pharyngeal Stage of Swallowing

As the bolus of food enters the posterior mouth and pharynx, it stimulates epithelial swallowing receptor areas all around the opening of the pharynx, especially on the tonsillar pillars, and impulses from these areas pass to the brain stem to initiate a series of automatic pharyngeal muscle contractions as follows:

• 1.

  The soft palate is pulled upward to close the posterior nares to prevent reflux of food into the nasal cavities.

• 2.

  The palatopharyngeal folds on each side of the pharynx are pulled medially to approximate each other. In this way, these folds form a sagittal slit through which the food must pass into the posterior pharynx. This slit performs a selective action, allowing food that has been masticated sufficiently to pass with ease. Because this stage of swallowing lasts less than 1 second, any large object is usually impeded too much to pass into the esophagus.

• 3.

  The vocal cords of the larynx are strongly approximated, and the larynx is pulled upward and anteriorly by the neck muscles. These actions, combined with the presence of ligaments that prevent upward movement of the epiglottis, cause the epiglottis to swing backward over the opening of the larynx. All these effects acting together prevent passage of food into the nose and trachea. Most essential is the tight approximation of the vocal cords, but the epiglottis helps to prevent food from ever getting as far as the vocal cords. Destruction of the vocal cords or of the muscles that approximate them can cause strangulation.

• 4.

  The upward movement of the larynx also pulls up and enlarges the opening to the esophagus. At the same time, the upper 3 to 4 centimeters of the esophageal muscular wall, called the upper esophageal sphincter (also called the pharyngoesophageal sphincter ), relaxes. Thus, food moves easily and freely from the posterior pharynx into the upper esophagus. Between swallows, this sphincter remains strongly contracted, thereby preventing air from going into the esophagus during respiration. The upward movement of the larynx also lifts the glottis out of the main stream of food flow, so the food mainly passes on each side of the epiglottis rather than over its surface; this action adds still another protection against entry of food into the trachea.

• 5.

  Once the larynx is raised and the pharyngoesophageal sphincter becomes relaxed, the entire muscular wall of the pharynx contracts, beginning in the superior part of the pharynx, then spreading downward over the middle and inferior pharyngeal areas, which propels the food by peristalsis into the esophagus.

To summarize the mechanics of the pharyngeal stage of swallowing: The trachea is closed, the esophagus is opened, and a fast peristaltic wave initiated by the nervous system of the pharynx forces the bolus of food into the upper esophagus, with the entire process occurring in less than 2 seconds.

Nervous Initiation of the Pharyngeal Stage of Swallowing

The most sensitive tactile areas of the posterior mouth and pharynx for initiating the pharyngeal stage of swallowing lie in a ring around the pharyngeal opening, with greatest sensitivity on the tonsillar pillars. Impulses are transmitted from these areas through the sensory portions of the trigeminal and glossopharyngeal nerves into the medulla oblongata, either into or closely associated with the tractus solitarius, which receives essentially all sensory impulses from the mouth.

The successive stages of the swallowing process are then automatically initiated in orderly sequence by neuronal areas of the reticular substance of the medulla and lower portion of the pons. The sequence of the swallowing reflex is the same from one swallow to the next, and the timing of the entire cycle also remains constant from one swallow to the next. The areas in the medulla and lower pons that control swallowing are collectively called the deglutition or swallowing center.

The motor impulses from the swallowing center to the pharynx and upper esophagus that cause swallowing are transmitted successively by the fifth, ninth, tenth, and twelfth cranial nerves and even a few of the superior cervical nerves.

In summary, the pharyngeal stage of swallowing is principally a reflex act. It is almost always initiated by voluntary movement of food into the back of the mouth, which in turn excites involuntary pharyngeal sensory receptors to elicit the swallowing reflex.

The Pharyngeal Stage of Swallowing Momentarily Interrupts Respiration

The entire pharyngeal stage of swallowing usually occurs in less than 6 seconds, there by interrupting respiration for only a fraction of a usual respiratory cycle. The swallowing center specifically inhibits the respiratory center of the medulla during this time, halting respiration at any point in its cycle to allow swallowing to proceed. Yet, even while a person is talking, swallowing interrupts respiration for such a short time that it is hardly noticeable.

The Esophageal Stage of Swallowing Involves Two Types of Peristalsis

The esophagus functions primarily to conduct food rapidly from the pharynx to the stomach, and its movements are organized specifically for this function.

The esophagus normally exhibits two types of peristaltic movements: primary peristalsis and secondary peristalsis. Primary peristalsis is simply continuation of the peristaltic wave that begins in the pharynx and spreads into the esophagus during the pharyngeal stage of swallowing. This wave passes all the way from the pharynx to the stomach in about 8 to 10 seconds. Food swallowed by a person who is in the upright position is usually transmitted to the lower end of the esophagus even more rapidly than the peristaltic wave itself, in about 5 to 8 seconds, because of the additional effect of gravity pulling the food downward.

If the primary peristaltic wave fails to move all the food that has entered the esophagus into the stomach, secondary peristaltic waves result from distention of the esophagus by the retained food; these waves continue until all the food has emptied into the stomach. The secondary peristaltic waves are initiated partly by intrinsic neural circuits in the myenteric nervous system and partly by reflexes that begin in the pharynx and are then transmitted upward through vagal afferent fibers to the medulla and back again to the esophagus through glossopharyngeal and vagal efferent nerve fibers.

The musculature of the pharyngeal wall and upper third of the esophagus is striated muscle. Therefore, the peristaltic waves in these regions are controlled by skeletal nerve impulses from the glossopharyngeal and vagus nerves. In the lower two-thirds of the esophagus, the musculature is smooth muscle, but this portion of the esophagus is also strongly controlled by the vagus nerves that act through connections with the esophageal myenteric nervous system. When the vagus nerves to the esophagus are cut, the myenteric nerve plexus of the esophagus becomes excitable enough after several days to cause strong secondary peristaltic waves even without support from the vagal reflexes. Therefore, even after paralysis of the brain stem swallowing reflex, food fed by tube or in some other way into the esophagus still passes readily into the stomach.

Receptive Relaxation of the Stomach

When the esophageal peristaltic wave approaches the stomach, a wave of relaxation, transmitted through myenteric inhibitory neurons, precedes the peristalsis. Furthermore, the entire stomach and, to a lesser extent, even the duodenum become relaxed as this wave reaches the lower end of the esophagus and thus are prepared ahead of time to receive the food propelled into the esophagus during the swallowing act.

Function of the Lower Esophageal Sphincter (Gastroesophageal Sphincter)

At the lower end of the esophagus, extending upward about 3 centimeters above its juncture with the stomach, the esophageal circular muscle functions as a broad lower esophageal sphincter, also called the gastroesophageal sphincter. This sphincter normally remains tonically constricted with an intraluminal pressure at this point in the esophagus of about 30 mm Hg, in contrast to the midportion of the esophagus, which normally remains relaxed. When a peristaltic swallowing wave passes down the esophagus, “receptive relaxation” of the lower esophageal sphincter occurs ahead of the peristaltic wave, which allows easy propulsion of the swallowed food into the stomach. Rarely, the sphincter does not relax satisfactorily, resulting in a condition called achalasia. This condition is discussed in Chapter 67 .

The stomach secretions are highly acidic and contain many proteolytic enzymes. The esophageal mucosa, except in the lower one-eighth of the esophagus, is not capable of resisting the digestive action of gastric secretions for long. Fortunately, the tonic constriction of the lower esophageal sphincter helps prevent significant reflux of stomach contents into the esophagus except under abnormal conditions.

Prevention of Esophageal Reflux by Valvelike Closure of the Distal End of the Esophagus

Another factor that helps prevent reflux is a valvelike mechanism of a short portion of the esophagus that extends slightly into the stomach. Increased intra-abdominal pressure caves the esophagus inward at this point. Thus, this valvelike closure of the lower esophagus helps to prevent high intra-abdominal pressure from forcing stomach contents backward into the esophagus. Otherwise, whenever we walk, cough, or breathe hard, stomach acid might be expelled into the esophagus.