Part 21

The absence of movement in the fundus would seem to give the food during its stay there little opportunity to become mixed with the gastric juices, and thus to undergo peptic digestion. The truth of this supposition can easily be proved experimentally by feeding a slightly alkaline meal, and later testing the chemical reaction of the contents of various parts of the stomach. A cat which had been without food for fifteen hours was given eighteen grams of mushy bread made slightly alkaline with sodium carbonate. One hour and a half after the cat had finished eating she was killed, and the stomach laid bare by opening the abdomen. A very small hole was then made through the wall in the fundus region, and another similar hole was made into the antrum. By means of a glass pipette food was extracted first from the periphery of the fundus; this food was slightly acid. The cleaned pipette was then introduced two and a half centimetres into the fundus contents, and the food thus extracted gave the original alkaline reaction. Specimens of the liquid contents of the antral and middle regions, taken from various depths, were all strongly acid. A dog killed an hour and three-quarters after eating showed similar differences between the reactions of the food in the fundus and the food in the pyloric portion. So, as a matter of fact, the food does not become acid at a uniform rate in all parts of the stomach, as would be the case if Beaumont’s and Brinton’s theories of mixing currents were true. Moreover, if the facts accorded with their notions, the saliva, which ceases to act in the presence of more than 0.003 per cent free hydrochloric acid, and is destroyed when the percentage of acid proteids is large, would manifestly have its service as a ferment limited to the relatively short time during which the stomach contents, in the process of thorough mixing, were reaching that degree of acidity. There is, however, no movement of food in the fundus, and the alkaline food received from the œsophagus remains alkaline in this region for a considerable period. The nutriment, therefore, if well chewed and thus mixed with saliva, can undergo salivary digestion in the fundus for a considerable period without interference by the acid gastric juice.[32]

From all these observations the conclusion must be that the fundus acts as a reservoir for the food, in which the digestion of sugars and starches may take place; and that the pyloric portion with its simple but marvellous peristaltic mechanism, by a single process, triturates the food, brings it near to the active glands, stirs it thoroughly with their secretions, and expels the products into the intestines.

THE INHIBITION OF STOMACH MOVEMENTS DURING EMOTION

Early in the research a marked unlikeness was noticed in the action of the stomachs of male and female cats. The peristalsis seen with only a few exceptions in female cats failed to appear in most of the males, although both had received exactly the same treatment. Along with this difference was a very striking difference in behaviour when bound to the holder; the females would lie quiet, mewing occasionally, but purring as soon as they were gently stroked. The males, on the contrary, would fly into a violent rage, struggle to be loose from their fastenings, bite at everything near their heads, cry loudly, and resist all attempts to quiet them. On account of this difference only female cats were used for some time, and the significance at first attributed to the action of the males was almost forgotten when the following incident recalled it and suggested that the excitement caused the suspension of the stomach movements. On October 23, 1897, a male cat was fed at 12.00, but was not placed on the holder till ninety minutes later. The waves were passing at the rate of six a minute. The cat fell into a rage and the waves suddenly stopped.

A few days later an observation on a female with kittens explained the absence of gastric movements in the males. While the peristaltic undulations were coursing regularly over the cat’s stomach, she suddenly changed from her peaceful sleepiness, began to breathe quickly, and struggled to get loose. As soon as the change took place, the movements in the stomach entirely disappeared; the pyloric portion relaxed and presented a smooth, rounded outline. I continued observing, and stroked the cat reassuringly. In a moment she became quiet and began to purr. As soon as this happened the movements commenced again in the stomach; first a few constrictions were visible near the end of the antrum, then a few near the sharp bend in the lesser curvature, and finally the waves were running normally from their habitual starting-place. By holding the cat’s mouth closed between the thumb and last three fingers, and covering her nostrils with the index finger, she could be kept from breathing. At the first sign of discomfort the fingers were removed. This experiment was repeated a great many times on different cats, and invariably the evidence of distress was accompanied by a total suspension of the motor activities of the stomach and a relaxation of the antral fibres.

No amount of kneading or compression of the abdomen with the fingers, short of making the cat angry, would cause the waves to stop; so that the cat’s movements, in themselves, were not the source of the inhibition. And since expressions of strong feeling on the part of the animal always accompanied cessation of the constriction-waves, the inhibition was probably due to nervous influence. It has long been common knowledge that violent emotions interfere with the digestive process, but that the gastric motor activities should manifest such extreme sensitiveness to nervous conditions is surprising.

SUMMARY

1. By mixing a harmless powder, subnitrate of bismuth, with the food, the movements of the stomach can be seen by means of the Röntgen rays.

2. The stomach consists of two physiologically distinct parts: the pyloric part and the fundus. Over the pyloric part, while food is present, constriction-waves are seen continually coursing towards the pylorus; the fundus is an active reservoir for the food and squeezes out its contents gradually into the pyloric part.

3. The stomach is emptied by the formation, between the fundus and the antrum, of a tube along which constrictions pass. The contents of the fundus are pressed into the tube and the tube and antrum slowly cleared of food by the waves of constriction.

4. The food in the pyloric portion is first pushed forward by the running wave, and then by pressure of the stomach-wall is returned through the ring of constriction; thus the food is thoroughly mixed with gastric juice, and is forced by an oscillating progress to the pylorus.

5. The food in the fundus is not moved by peristalsis, and consequently it is not mixed with the gastric juice; salivary digestion can therefore be carried on in this region for a considerable period without being stopped by the acid gastric juice.

6. The pylorus does not open at the approach of every wave, but only at irregular intervals. The arrival of a hard morsel causes the sphincter to open less frequently than normally, thus materially interfering with the passage of the already liquefied food.

7. Solid food remains in the antrum to be rubbed by the constrictions until triturated, or to be softened by the gastric juice, or later it may be forced into the intestine in the solid state.

8. The constriction-waves have, therefore, three functions: the mixing, trituration, and expulsion of the food.

9. At the beginning of vomiting, the gastric cavity is separated into two parts by a constriction at the entrance to the antrum; the cardiac portion is relaxed, and the spasmodic contractions of the abdominal muscles force the food through the opened cardia into the œsophagus.

10. The stomach movements are inhibited whenever the cat shows signs of anxiety, rage, or distress.

THE MOVEMENTS OF THE INTESTINES STUDIED BY MEANS OF THE RÖNTGEN RAYS[33]

BY W. B. CANNON

_From the Laboratory of Physiology in the Harvard Medical School_

Extracts from _American Journal of Physiology_, 1902

INTRODUCTION

The investigation of intestinal movements has been beset by the same difficulties that characterised the investigation of the gastric mechanism. Pathological subjects or animals subjected to the disturbing

## action of drugs and anæsthetics and of serious operations have been the

only sources of our knowledge. A considerable difference of opinion as to the nature of the normal movements in the intestines has resulted from observations made under these necessary abnormal conditions. The slowly advancing peristaltic wave and the _Pendelbewegung_, or swaying movement, described by Ludwig, have been regarded as true physiological processes. Concerning antiperistalsis and the swiftly running vermicular contraction, observers are not so nearly in agreement. The activity of the large intestine has been described as simply peristalsis of a slower rate than that seen in the small intestine.

The best known of the intestinal movements is the normal peristaltic wave. This wave is slow, having a rate of about two centimetres per minute, is regular, and by most observers is said to move always in one direction. The progress of the contraction, as suggested by Nothnagel’s experiments, and as clearly stated by Mall and by Bayliss and Starling, is dependent upon a local reflex. According to Mall, when an object stimulates the mucosa there occurs above the point of stimulation a constriction which forces the object downward; and as it moves downward new regions immediately above the mass are by this reflex brought into constriction, and thus the wave and its stimulus advance together. “At the same time,” Mall states, “a sucking force, due to active dilatation below the body, may have a tendency to drag it down.” In what manner an active dilatation of the intestinal wall may occur so as to produce a “sucking force” he does not make wholly clear. Bayliss and Starling, in describing normal peristalsis in the intestine, state that the contractions above the bolus increase until there is a strong tonic constriction. This passes the bolus onward, and as it advances the ring of constriction follows it. While the bolus is passing down, the gut above it is traversed by waves running as far as the constricted ring. These observers state the law of intestinal peristalsis thus: “Local stimulation of the gut produces excitation above and inhibition below the excited spot.”

The pendulum movements are characterised by a gentle swaying motion of the coils, and are accompanied by rhythmical contractions of the intestinal wall. They continue with undiminished force after paralysis of the local nervous mechanism by nicotine or cocaine; they have been called, therefore, myogenic or myodromic contractions. Observers have described them variously as shortenings and narrowings of the gut, rhythmically repeated at nearly the same intestinal circumference; as alternating to-and-fro movements of the long axis without changes in the lumen; as local or extensive periodic contractions and relaxations mainly of the circular musculature; and as waves involving both muscular coats of the intestine, and travelling normally from above downward at a rapid rate (2 to 5 cm. per second). They have been seen in the dog, and in the rabbit and cat. In the cat Bayliss and Starling noticed that when the lumen of the gut was distended by a rubber balloon, there appeared rhythmical contractions, which nearly always were most marked at about the middle of the balloon; _i. e._, the region of greatest tension. This form of constriction, which, as my observation shows, is an indication of the manner in which the rhythmical contraction acts in the cat’s intestine, Bayliss and Starling seem to have regarded with slight attention, since it did not accord with the law of peristalsis.

The swift vermicular wave may pass the whole length of the intestine in about a minute. It is often seen just after death, as well as in pathological states such as intestinal anæmia or hyperæmia, and when the bowel contains gases and organic acids from decomposing food. Starling is inclined to regard this type of intestinal activity as an exaggeration of the rhythmic type; Mall, on the other hand, places it in a class by itself, and declares that its service is to rid the intestine rapidly of irritating substances. Nothnagel, who designates this form of movement with the term _Rollbewegung_, thinks it is transitional between a physiological and a pathological activity.

The existence of antiperistalsis has been so much questioned that it will be considered in a special section of this paper, where my observations may be conveniently introduced.

The common understanding of the manner in which food passes through the intestinal canal is that the chyme ejected from the stomach is pressed downward by a peristalsis, which passes slowly over a part or all of the small intestine. The peristaltic waves of the colon are supposed to constitute an independent group, similar to those of the small intestine, but weaker and slower. Movements of the food other than the uninterrupted advance have been mentioned by some observers. Starling states that the effect of the pendulum movement is to mix the contents of the intestine and bring them into intimate contact with the mucous membrane. Grützner writes that he has been brought “by strange and peculiar observations” to believe that the fluid contents of the small intestine move irregularly forward, then forward and back, then perhaps remain quiet for some time, only to pass backward for a long distance, and finally to move forward steadily to the end. In this manner the food is mixed and brought into contact with the absorbing walls. The to-and-fro shiftings of the food Grützner ascribed to advancing and retrograde contractions of the intestinal musculature, and he argued that even circular constrictions must force the liquid contents away in both directions. To support his contention, Grützner introduced mercury into the intestine and observed it with the Röntgen rays. After noting a backward and forward movement of the mercury he dismissed the method, saying, “Many a flash must come from the Röntgen tube before the _normal_ movement of the intestinal contents is made entirely clear by this method.”

The following account is a summary of many repeated observations on different animals, and is a contribution to a clear understanding of the normal movements of the intestines and their contents.

THE MOVEMENTS OF THE SMALL INTESTINE

When the food has been distributed through the intestine so as to present the appearance shown in Figure 1, a noticeable feature in most or all of the loops is the total absence of movement. If the animal remains quiet, however, only a few moments elapse before peculiar motions appear in one or another of the loops, or perhaps in several, and last for some time. These motions consist in a sudden division of one of the long, narrow masses of food into many little segments of nearly equal size; then these segments are again suddenly divided and the neighbouring halves unite to make new segments, and so on, in a manner to be more fully described. I have called this process the rhythmic segmentation of the intestinal contents. Further observation reveals peristalsis here and there, and under certain circumstances the typical swaying movements may be seen. All these phenomena are now to be considered in detail.

[Illustration: FIGURE 1.--Appearance of food in the intestines 5¾ hours after eating. This and other radiographs reduced two-thirds.]

[Illustration: FIGURE 2.--Diagram representing the process of rhythmic segmentation. Lines 1, 2, 3, 4 indicate the sequence of appearances in the loop. The dotted lines mark the regions of division. The arrows show the relation of the particles to the segments they subsequently form.]

_Rhythmic segmentation of the intestinal contents._--This is by far the most common and the most interesting mechanical process to be seen in the small intestine. The nature of the process may best be understood by referring to the diagram in Figure 2. A string-like mass of food is seen lying quietly in one of the intestinal loops (line 1, Fig. 2). Suddenly an undefined activity appears in the mass, and a moment later constrictions at regular intervals along its length cut it into little ovoid pieces. The solid string is thus quickly transformed, by a simultaneous sectioning, into a series of uniform segments. A moment later each of these segments is divided into two particles, and immediately after the division neighbouring particles (as _a_ and _b_, line 2, Fig. 2) rush together, often with the rapidity of flying shuttles, and merge to form new segments (as _c_, line 3, Fig. 2). The next moment these new segments are divided, and neighbouring

## particles unite to make a third series, and so on. At the time of the

second segmentation (line 3, Fig. 2) the end particles are left small. Observation shows that these small pieces are not redivided. The end piece at A simply varies in size with each division; at one moment it is left small, at the next moment it is full size from the addition of a part of the nearest segment, and a moment later is the small bit left after another division. The end piece at B (probably the rear of the mass) shoots away when the end mass is divided, and is swept back at each reunion to make the large end mass again, only to be shot away and swept onward with each recurrence of the constrictions. Thus the process of repeated segmentation continues, with the little

## particles flitting towards each other and the larger segments shifting

to and fro, commonly for more than half an hour without cessation. From the beginning to the end of a period of segmentation the food is seen to have changed its position in the abdomen to only a slight extent; whether this change is a passing of the food along the loop, or a movement of the loop itself, it is impossible to tell from the shadows on the screen. The change of position, however, is much less conspicuous than the lively division and redivision which the mass suffers so many times from the busy, shifting constrictions.

From this typical form of rhythmic segmentation there are several variations. Sometimes, and especially when the mass of food is thick, the constrictions do not make complete divisions and are so far apart that the intermediate portions are relatively large. Moreover, the constrictions do not take place in the middle of each portion, but near one end; thus each portion is constricted, not into halves, but into thirds. If a little pointer is placed at the middle of a segment, when the segments are completely divided into halves, in a few seconds the pointer will be in the middle of the clear space between two segments; but in a few seconds more the first phase will return and the pointer will again be indicating a segment,--two operations intervene between similar phases. When, however, the portions are constricted into thirds, the indicator shows it, since three operations intervene between similar phases. The manner of these changes is made clearer by reference to the diagram in Figure 3. That each portion is constricted into three pieces is proved also by watching the gradual reduction of the portion at the left end of line 1 through lines 2 and 3, and also in the gradual formation of a full-sized portion at the right end of lines 2, 3, and 4. When food undergoing this process is watched, it appears to be affected by a series of constrictions, each of which starts at one end of the mass and marches through to the other end, leaving its impress at short intervals along the length. The progression of the dotted lines from right to left in _a_, _b_, _c_, and _d_, etc., Fig. 3, gives a notion of these advancing constrictions.

Another variation of the segmentation is shown in Figure 4. In this type there are evidently divisions and subdivisions, _i.e._, one more operation between the appearance and the reappearance of the same phase than is present in the simple division of the small segments in a long string of food (Fig. 2). This form of segmentation is fairly typical for the constrictions seen in food advancing through the intestine. Sometimes the divisions occur in the middle of a long string of food and leave the ends wholly unaffected.

[Illustration: FIGURE 3.--Diagram showing the relations of the portions when they are constricted into three pieces. The dotted lines indicate regions of constriction; the arrows indicate the relationship of the pieces to the portions they subsequently form.]

A remarkable feature in the segmentation of the food is the rapidity with which the changes take place. The simplest way of estimating the rate of division is to count, not the number of times the partition of the food recurs in the same place, but the number of different sets of segments observed in a given period. Thus in Figure 4 the appearances of lines 1, 2, 3, 4, etc., would be counted, and not merely lines 1, 4, etc. Repeated observations on different animals have shown that the most common rate of division in long, thin chains of food varies between twenty-eight and thirty times in a minute; _i. e._, there is a change from one set of segments to another set every two seconds, and a return of the same phase every four seconds. In some cases the rate is as low as twenty-three times per minute. The larger masses seem to be associated with a slower segmentation; the operations indicated in Figure 3, for example, occurred from eighteen to twenty-one times in a minute, so that the same phase reappeared only once in eight or nine seconds. The segmentation frequently continues for more than half an hour; in one instance it was seen to persist with only three short periods of inactivity for two hours and twenty-two minutes. At the rate of thirty segmentations per minute it is clear that a slender string of food may commonly undergo division into small particles more than a thousand times while scarcely changing its position in the intestine.

I have seen once, in a cat only lightly etherised, the exterior of an intestine which was dividing the food as above described. An hour and a half after a meal of salmon the anæsthetic was given, the abdomen opened, and the flaps raised so as to form walls. Warm salt solution was then poured into the abdominal cavity, and the floating coils left covered with the transparent omentum. The gastric peristaltic waves were running regularly; on the intestine there were visible at various places during the period of observation regions of constriction which had the appearance shown in Figure 3, except that the rings were relatively nearer together. New rings of constriction took place on the same side of all the bulging parts at the margin of the constricted portion (_cf._ dotted lines, Fig. 3). As new rings occurred the old relaxed, but apparently with tardiness, for the contents gurgled as if forced through the narrowed lumen. The constrictions recurred irregularly and at much longer intervals than in the normal animal. The contracted rings were pale and bloodless.