Part 19
The X-ray method lends itself less successfully to the study of deglutition in man than in the other animals we have studied. The thickness of the thorax, the distance of the œsophagus from the surface, and the relation to dense tissues render the observation of a swallowed mass difficult, especially when the mass is in rather rapid motion. The few observations which we have to report were made on a seven-year-old girl, placed in the sitting posture. Gelatine capsules containing bismuth were used for solids, and were traced to a point below the heart. The motion was very regular, and apparently due to peristalsis, for the bolus descended without a hitch or irregularity of any kind. Sometimes the capsule became fixed in the upper œsophagus, at about the level of the second rib. Repeated swallows of water would fail to dislodge it. An interesting point was noted here. With each attempt at swallowing, the capsule would rise slightly, as if the œsophagus was pulled up with the rise of the larynx; then the capsule would descend to its former position.
Semi-solids--a mush of bread and milk--could be seen about as far as solids; that is, to just below the heart. The motion of the mushy bolus was the same as with solids, except that the rapidity was perhaps slightly greater.
It should be noted here that with the human subject, as well as with the horse, our results for semi-solids differ from those derived by Meltzer’s method; for according to his statements semi-solids, like liquids, are squirted down the œsophagus, and are not propelled by peristalsis, as has been the case in our observations.
Liquids--bismuth and water--were seen only in the neck and upper thorax. Here there was a decided squirt. With the rise of the larynx the liquid was seen to pass rapidly through the pharynx and well down into the thoracic œsophagus before it was lost to observation. The rate, however, by estimation was less than that of liquids in the horse.
There remains to be considered Meltzer’s latest investigation, in which he endeavoured to ascertain whether liquids remain above the cardia till the arrival of the peristalsis, or ooze down before. An experimental answer was secured by Meltzer by the following method. The abdominal and gastric walls of an anæsthetised dog were incised, and a tube (vaginal speculum) introduced. Through this the entrance of food into the stomach could be observed directly. In repeated experiments no liquid was seen to pass through the cardia before the arrival of the peristaltic wave. An incision through the diaphragm near its anterior origin showed that the swallowed liquid was not squirted as far as a point an inch above the diaphragm. To observe the œsophagus nearer its beginning, the upper three ribs were resected on the left side. Thus the swallowed liquid was seen to shoot along the œsophagus before any peristalsis reached this point. The resection of the fifth rib exposed the œsophagus half-way between the bifurcation of the trachea and the diaphragm. Here a bulging was sometimes observed immediately after the beginning of the act, and the swallowed mass remained there until a peristaltic wave carried it down. If the mass swallowed was small, or was projected with moderate force, it might not even reach as far as the bifurcation. From these experiments Meltzer concluded that in animals, as in man, liquid food is not carried down the œsophagus by peristalsis, but is thrown rapidly into a deep part of the canal. The depth reached depends on the quantity swallowed, the force used, and the tonicity of the lower part of the œsophagus.
The difference between these methods of Meltzer and those employed in our experiments has already been mentioned; and merely his results, which were obtained with liquids alone, need be considered here. According to our observations on the dog, there was no distinct pause at any part of the canal. The movement simply became slower, and continued at this rate until the stomach was reached. Neither was the rate through the diaphragmatic part of the œsophagus slower than through the thoracic. The quick propulsive movement noticed in the dog was observed with solids and semi-solids as well as with liquids, but the liquids descended further down the canal before the movement changed to the slower peristalsis. While this difference was evident to the eye, the total time consumed by liquids in passing from pharynx to stomach was not enough shorter than the time for solids and semi-solids to be determined by our measurements.
SUMMARY.
The phenomena of œsophageal deglutition as determined by our experiments may then be described as follows:—
There is a difference in swallowing according to the animal and the food which is used.
In fowls the rate is slow and the movement always peristaltic, without regard to consistency. A squirt-movement with liquids is manifestly impossible, as the parts forming the mouth are too hard and rigid. With this diminution of propulsive power in the mouth there is observed a greater reliance on the force of gravity. The head is raised each time after the mouth is filled, and the fluid by its own weight trickles into the œsophagus, through which it is carried by peristalsis.
In the cat the movement is always peristaltic, and slightly faster than in fowls. A bolus takes from nine to twelve seconds in reaching the stomach. Liquids move somewhat more rapidly than semi-solids in the upper œsophagus. In the lower or diaphragmatic part the rate is very much slower than above, and is the same for liquids as for solids.
In the dog the total time for the descent of a bolus is from four to five seconds. The food is always propelled rapidly in the upper œsophagus, and moves more slowly below. This rapid movement is frequently continued further with liquid food. No distinct pause was observed when the movement of the bolus changed from the rapid to the slower rate.
In man and the horse liquids are propelled deep into the œsophagus at a rate of several feet a second by the rapid contraction of the mylohyoid muscles. Solids and semi-solids are slowly carried through the entire œsophagus by peristalsis alone.
THE MOVEMENTS OF THE STOMACH STUDIED BY MEANS OF THE RÖNTGEN RAYS
BY W. B. CANNON, M.D.
_From the Laboratory of Physiology in the Harvard Medical School_
Extracts from _American Journal of Physiology_, 1898
Since the stomach gives no obvious external sign of its workings, investigators of gastric movements have hitherto been obliged to confine their studies to pathological subjects or to animals subjected to serious operative interference. Observations made under these necessarily abnormal conditions have yielded a literature which is full of conflicting statements and uncertain results. The only sure conclusion to be drawn from this material is that when the stomach receives food obscure peristaltic contractions are set going, which in some way churn the food to a liquid chyme and force it into the intestines. How imperfectly this describes the real workings of the stomach will appear from the following account of the actions of the organ studied by a new method. The mixing of a small quantity of subnitrate of bismuth with the food allows not only the contractions of the gastric wall, but also the movements of the gastric contents to be seen with the Röntgen rays in the uninjured animal during normal digestion. An unsuspected nicety of mechanical action and a surprising sensitiveness to nervous conditions have thereby been disclosed.
INTRODUCTORY LITERATURE
The early writings on the subject of gastric movements are characterised by general inferences from physical laws and from the anatomical structure of the stomach. According to Galen the stomach had four functions: to draw the food from the mouth (_facultas attractrix_), to retain the food (_facultas retentrix_) during the process of chemical digestion (_facultas alteratrix_), and, finally, to pass the changed material onward (_facultas expultrix_). In later writings the _facultas attractrix_ failed to appear as one of the functions of the stomach. Fallopius, in the sixteenth century, changed the notion of the _facultas retentrix_ by suggesting that the pylorus alone performed this office, and that the muscles of the gastric wall could help only by remaining quiet. Thus the _facultas alteratrix_ and the _facultas expultrix_ are left as true gastric functions. It is with the latter activity and its effects that this paper is concerned.
The ideas of the early writers concerning the pylorus and cardia are of interest. The cardia, they were agreed, is closed during normal digestion in order to keep the food from re-entering the œsophagus. The pylorus they looked upon as the ruler of the actions of the stomach. Such names as pylorus (keeper of the gate), janitor justus, and rector, which the first investigators gave to the sphincter, indicate their theories of its functions. The passage of chyme into the duodenum, the keeping of undigested food in the stomach, the act of vomiting, were all dependent, they believed, on the “will” of the pylorus.
No substantial advance was made beyond these hypotheses until the beginning of the eighteenth century, when Wepfer and Schwartz applied the experimental method to the study of the gastric movements and laid the foundation of a more accurate knowledge. Wepfer vivisected wolves, dogs, and cats, and observed constrictions following stimulation of the stomach. He remarked a general contraction of the pyloric part in vomiting and noted peristaltic and antiperistaltic movements passing over the organ. About the middle of the stomach he frequently saw a deep constriction. The investigations of Schwartz are more valuable in that his search was for the normal action of the muscular coats. The movements, as he observed them, were generally only slight. They began either at the pylorus and passed to the left, half-way to the cardia, or started at the fundus and went to the pylorus. The contractions and relaxations, following one another, formed larger or smaller depressions and elevations, _i. e._, more or less definite waves.
Near the middle of the last century Haller, after confirming the results obtained by Schwartz and Wepfer, summarised his knowledge of the motor functions of the stomach as follows. In general, contraction alternates with relaxation, so that the stomach is, now here, now there, made narrower by longitudinal or transverse depressions; then in these same places relaxation and bulging occur. So long as both apertures are closed the food is driven hither and thither by the shifting movements. It first takes a definite direction when the cardia or the pylorus opens. If the cardia opens, there is an antiperistalsis followed by regurgitation and vomiting. If, on the contrary, the pylorus relaxes, a contraction, starting at the œsophagus, pushes the contents of the stomach into the duodenum. The pylorus allows the passage of fluids, but if it be stimulated by over distention or by hard pieces of food, it closes tightly.
Such was the knowledge of gastric movements in Haller’s time. A comparison of his descriptions with those in any standard work on physiology published ten or fifteen years ago will show that, despite very many researches, little advance had been made. Examinations of animals and men with gastric fistulas, studies of the stomach through the atrophied abdominal wall, and vivisection have yielded numerous results; but these have not been harmonious, and have led to much controversy. Prominent in this mass of material as a valuable contribution are Beaumont’s careful observations through the gastric fistula of Alexis St. Martin. Beaumont’s work has recently been confirmed by Hofmeister and Schütz, who, with Rossbach, Hirsch, Openchowski, and others, have presented during the last twelve years much new and interesting information. Since, however, it will conduce to clearness to set forth the results of these investigations in connection with my own work, their consideration will be deferred until later.
It will then appear that these later investigations, like the earlier researches, disagree as to the details of the stomach movements. Such differences in results are the proper outcome of the abnormal conditions under which the studies have been conducted. Obviously, in order to see the natural movements of the stomach, the organ should be observed in its natural state, and not after it has been disturbed by removal from the abdomen, or by the adhesions and losses of substance incident to gastric fistulas.
As a means of watching the gastric motor activities under normal circumstances, Dr. H. P. Bowditch, in the autumn of 1896, suggested the use of the Röntgen rays. The present paper is the result of the work thus far completed. The kind assistance and stimulating counsel of Dr. Bowditch throughout the investigation are gratefully acknowledged.
THE ANATOMY OF THE STOMACH AND ITS RELATIONS TO THE SHADOW
It must be constantly borne in mind that the shadows described in this research are cast by the gastric contents, not by the stomach itself. Therefore the movements of the organ are not seen directly, but are indicated by their effect on the contained food. Variations in the length and breadth of the stomach can be inferred from changes in the outline of the shadow, but variations in the front-to-back diameter of the organ must be judged from changes in the intensity of the shadow.
The form of the active stomach soon after food has been taken is shown in outline in Figure 1. Since the several parts of the stomach are to be mentioned frequently, it will be well to recall them here in their relations to the outline. The larger, cardiac part of the organ lies to the left of a line through _w x_. Into it the œsophagus opens through the cardiac sphincter, or cardia, at _c_. The pyloric part, which includes all of the stomach situated at the right of a line _w x_, is closed by the pylorus at _p_. This part has two divisions: the antrum at the right of the line _y z_, and the preantral part of the pyloric portion, or middle region of the stomach, between the lines _w x_ and _y z_. The lesser curvature corresponds approximately to the anterior border of the shadow _c w p_; the greater curvature to the more extensive sweep, _c p_, along the posterior border.
[Illustration: FIGURE 1.]
The wall of the cat’s stomach consists of three coats, but as this paper deals only with the functions of the muscular coat, that alone will be described. The gastric muscular fibres are disposed in three sets: an outer longitudinal layer, a middle circular layer, and a set of inner oblique fibres. The longitudinal fibres continue those of the œsophagus, and, radiating over the cardiac end, become more marked along the curvatures than on the front and back surfaces. Over the antrum they lie in a thick, uniform layer. The circular fibres form a complete investment, and are arranged in rings at right angles to the curved axis of the stomach. Towards the pyloric end they become denser and stronger, and at the pylorus form a thick bundle, the pyloric sphincter. Separating the antrum from the rest of the stomach, at _y z_, is a special thickening of the circular fibres, called by the early writers the “transverse band,” and described by Hofmeister and Schütz as the “sphincter antri pylorici.” The oblique fibres start from the left of the cardiac orifice and pass as two strong bands along the anterior part of the dorsal and ventral surfaces, giving off fine fasciculi to the circular musculature; towards the antrum they gradually disappear.
The musculature of the stomach consists of smooth muscle fibres, the chief physiological characteristics of which are slowness of contraction, rhythmic alternation of contraction and relaxation, and a very great tonicity, or power of prolonged contraction. The action of these muscles in the process of gastric digestion is now to be considered.
THE NORMAL MOVEMENTS OF THE STOMACH
Since the time of Haller the chief contributors to the knowledge of the mechanics of the stomach have been Beaumont, Hofmeister and Schütz, and Rossbach.
Beaumont’s famous investigations on Alexis St. Martin are recorded in almost all general works on physiology. Through a gastric fistula he introduced a thermometer-tube and observed how it was affected by the motions of the stomach. His conclusions are as follows: “The circular or transverse muscles contract progressively from left to right. When the impulse arrives at the _transverse band_, this is excited to a more forcible contraction, and closing upon the alimentary matter and fluids contained in the pyloric end, prevents their regurgitation. The muscles of the pyloric end, now contracting upon the contents detained there, separate and expel some portion of the chyme.... After the contractile impulse is carried to the pyloric extremity, the circular band and all the transverse muscles become relaxed, and a contraction commences in a reversed direction, from right to left, and carries the contents again to the splenic extremity to undergo similar revolutions.”
In close accord with Beaumont’s description of the activities of the human stomach are the records of the investigations on the stomach of dogs by Hofmeister and Schütz. They removed the stomach from the body and placed it in a moist chamber, kept at body-heat and covered with glass. Under such conditions the organ remained active for from sixty to ninety minutes. A typical movement is described by these observers as composed of two phases. In the first phase a constriction of the circular fibres, deeper on the greater curvature, starts a few centimetres from the cardia and passes towards the pylorus. As the constriction proceeds it increases in strength until a maximum is reached about two centimetres in front of the antrum. This annular contraction, called by Hofmeister and Schütz the “preantral constriction,” closes the first phase. Immediately thereafter the strong sphincter antri pylorici, or transverse band, contracts. Now, while the preantral constriction is relaxing, the sphincter antri pylorici tightens still more, and the antrum is shut off from the rest of the stomach. As soon as this has occurred a general contraction of the muscles of the antrum follows. Relaxation begins at the sphincter antri pylorici and progresses slowly towards the pylorus; it is sometimes accompanied by an antiperistaltic movement.
Although Rossbach also used dogs, his results vary considerably from those of Hofmeister and Schütz. This discrepancy is possibly accounted for by a difference in method, for Rossbach left the stomach in the body. The dogs were treated with morphia and curare, and the abdomen was then widely opened, so that the movements could be clearly seen. When the stomach was full Rossbach saw deep constrictions begin near the middle and pass in waves to the pylorus. At first these movements were weak; later, however, they became more vigorous. The fundus remained in tonic contraction about its contents and took no part in the peristalsis.
Before attempting to explain the difference in the records of these observers I shall give an account of what may be seen in a cat by use of bismuth subnitrate and the Röntgen rays.
1. _Movements of the pyloric part._--Within five minutes after a cat has finished a meal of bread, there is visible near the duodenal end of the antrum a slight annular contraction which moves peristaltically to the pylorus; this is followed by several waves recurring at regular intervals. Two or three minutes after the first movement is seen very slight constrictions appear near the middle of the stomach, and pressing deeper into the greater curvature, course slowly towards the pyloric end. As new regions enter into constriction, the fibres just previously contracted become relaxed, so that there is a true moving wave, with a trough between two crests. When a wave swings round the bend in the pyloric part the indentation made by it deepens; and as digestion goes on the antrum elongates and the constrictions running over it grow stronger, but until the stomach is nearly empty they do not entirely divide the cavity. After the antrum has lengthened, a wave takes about thirty-six seconds to move from the middle of the stomach to the pylorus. At all periods of digestion the waves recur at intervals of almost exactly ten seconds. So regular is this rhythm that many times I have been able to determine within two or three seconds when a minute had elapsed simply by counting six similar phases of the undulations as they passed a given point. It results from this rhythm that when one wave is just beginning several others are already running in order before it. Between the rings of constriction the stomach is bulged out, as shown in the various outlines in Figures 2, 3, 4, and 5. The number of waves during a single period of digestion is larger than might possibly at first be supposed. In a cat that finished eating fifteen grams of bread at 10.52 A.M., the waves were running regularly at 11.00 o’clock. The stomach was not free from food until 6.12 P.M. During that time the cat was fastened to the holder at intervals of half an hour, and the waves were always observed following one another in slow and monotonous succession. At the rate of three hundred and sixty an hour, approximately two thousand six hundred waves passed over the antrum during that single digestive period.
From the above review it will be manifest that my observations of the movements of the pyloric part agree closely with those of Rossbach, but differ considerably from the harmonious results of the work of Beaumont, and Hofmeister and Schütz. Beaumont’s methods, however, may be justly criticised on the ground that the thermometer-tube which he held in the stomach was wholly unlike food and very liable to bring about unwonted contractions in so sensitive an organ as the stomach. Further, the movements observed by Hofmeister and Schütz, as Ewald has pointed out, may easily have resulted from the abnormal stimulus due to lack of blood--a potent cause of peristalsis. And it will be shown later that the accounts given by these investigators describe very well the actions of the stomach when stimulated by an unusual irritant. In this connection it may be added that since the publication of the preliminary notice of my work, Roux and Balthazard, using the Röntgen rays, have published the results of observations on the stomachs of the dog and man similar to those thus far described in this paper.
The fact that my observations and those of Roux and Balthazard were conducted under normal conditions, and that the conditions of Rossbach’s experiments were more nearly normal than those of the other observers mentioned, warrants the conclusion that the pyloric part has a more important function than that of merely expelling the contents of the stomach into the intestines. After summarising the description given by Hofmeister and Schütz, Ewald, for _a priori_ reasons, declares: “I cannot accept this view. The plain fact that the pyloric portion secretes a strongly digesting fluid containing pepsin and hydrochloric acid proves it to be an important part for the peptonising function of the stomach.” The account of the remarkable manner in which the pyloric portion performs this function must be deferred until the movements of other parts of the stomach have been considered.
2. _Movements of the pyloric sphincter._--Rossbach mars his otherwise careful work by declaring that the pylorus is tightly closed during the whole digestive period of from four to eight hours, and that then the sphincter relaxes and the peristaltic waves empty the stomach. That this is not the normal action of the sphincter has been shown by several observers. Hirsch watched dogs with duodenal fistulas and saw food come from the stomach at intervals of one-fourth of a minute to several minutes. Roux and Balthazard maintain that in dogs food enters the duodenum at the completion of each wave of constriction. Observations on the cat, however, do not support their view, but agree rather with the statement of Hirsch.