THE MOVEMENTS OF THE FOOD IN THE ŒSOPHAGUS
By W. B. Cannon and A. Moser
From the Laboratory of Physiology in the Harvard Medical School
Extracts from American Journal of Physiology, 1898
The movements of deglutition, in common with many other physiological processes, were explained by the older physiologists on anatomical grounds. Thus, Magendie divided the act into three parts, corresponding to the anatomical regions of the mouth, pharynx, and œsophagus. The muscles of each of these divisions were considered the active agents in propelling the food onward. The function of moving the mass to the pharynx was variously ascribed to the tongue itself, to the mylohyoid muscles, and to gravity. For the second part, the movement through the pharynx, there was more unanimity of opinion, since the constrictors, especially the middle and lower, were evidently concerned.
Direct observations on the movement of swallowed masses in the œsophagus were first made by Mosso. The œsophagus of a dog was laid bare, and a transverse incision made through it, or a piece of it excised. A small wooden ball was placed in the canal below the excised part, and the animal was then stimulated to swallow. One or two seconds after the contraction of the pharyngeal muscles a peristaltic wave began to traverse the œsophagus. This wave did not stop at the point of excision, but in due time reappeared below, and carried the ball to the stomach. Thus the act was shown to be controlled by the central nervous system. Peristalsis was so plainly the motive power that the action was never doubted. Yet this belief was soon to be questioned.
In 1880 Falk and Kronecker studied the movements in the mouth and pharynx, and advanced the theory that deglutition was accomplished by the rapid contraction of the muscles of the mouth. During the act of swallowing the air-tight buccal cavity shows a manometric pressure of twenty centimetres of water. The same pressure was demonstrated to be present also in the œsophagus, but not in the stomach. This pressure was considered sufficient to force food through the œsophagus before the peristaltic wave traversed it. Another argument for rapid descent was found in the fact that cold water can be felt in the epigastric region almost immediately after being swallowed. Further, when strong acids pass through the gullet, they corrode but small parts of it, and not the entire mucous membrane, as would be the case were the acid carried to the stomach by peristalsis.
Over a year and a half ago it was suggested by Prof. H. P. Bowditch that if some substance opaque to the Röntgen rays were swallowed, it could be seen in its passage to the stomach, and the nature of its movement thus determined. Anæsthesia could be dispensed with,—a desirable condition, since observers had found that it interfered greatly with the deglutition reflex. It would be unnecessary to open either the abdominal or the pleural cavity. The reflex stimulus of food, moreover, would be better than electrical stimulation of the superior laryngeal nerve. In short, the animal would swallow normal food under practically normal conditions. At Dr. Bowditch’s suggestion and with his valuable assistance—which we gratefully acknowledge—we made the following series of experiments.
To render the swallowed mass opaque, subnitrate of bismuth was used. The salt is tasteless, practically inert, and can be fed in large quantities without harm. In order that observations could be made by more than one person, all experiments were conducted in a dark room. On the side of the animal opposite the Crookes tube was placed an open fluorescent screen, on which the different tissues of the animal were outlined with varying degrees of light and shade. Among these shadows the swallowed mass appeared as a darker object, and thus its motion could be studied.
For the first experiments the goose was selected. The head and neck were held stationary by a tall pasteboard collar, which allowed free movement of the head without constriction of the neck. The fluorescent screen was placed against this collar at a uniform distance of thirty centimetres from the tube. When a bolus of corn-meal mush mixed with bismuth was placed in the pharynx, it descended slowly and regularly, and occupied about twelve seconds in passing over a distance of fifteen centimetres. The screen was marked at intervals of two centimetres with cross lines, by means of which the relative rate in different parts of the œsophagus could be studied. A vibrator marking tenths of a second was interrupted whenever the bolus crossed a line. An average of over one hundred such observations showed that the rate became slightly slower as the bolus proceeded.
In order to test liquids, molasses was mixed with bismuth to such a consistency as to drop easily from a glass rod. When this was fed with a pipette, it passed slowly and regularly down the œsophagus, clearly by peristalsis. The rate was about the same as for solid food. In both these experiments the addition of water would sometimes cause irregularities in the descent. Microscopic sections from four different parts of the œsophagus of the goose showed no histological difference.
In the experiments on the cat, the animal was placed on its back and left side on a holder. The extremities were secured by straps. The head was held between two upright rods, connected above by a thong; this allowed free movement of the head, without resistance to the passage of food. Shreds of meat dipped in bismuth were ordinarily masticated and swallowed without difficulty. For soft solids, bread and milk were used, so fluid as to be easily drawn up into a pipette. The insolubility of the bismuth salt rendered the study of liquids more difficult. Strong solutions of potassic iodide and other salts, and suspension of bismuth, in acacia and molasses were tried; but a simple mixture of milk and bismuth, shaken in a test tube and immediately drawn up into a pipette, was found most practicable.
Inasmuch as the movement of these different foods varied in different parts of the œsophagus, it will be convenient to divide the latter into three sections. The first or cervical portion extends from the pharynx to the thorax; the second or thoracic, from here to the lower half of the heart; and the third comprises the rest of the canal. The relative length of these three parts is about in the ratio of 9:8:6.