PERIOD OF OCCURRENCE AND IMPORTANCE OF THE PSYCHIC OR APPETITE JUICE IN THE SECRETORY WORK OF THE STOMACH—THE INEFFICIENCY OF MECHANICAL STIMULATION OF THE NERVOUS APPARATUS OF THE GASTRIC GLANDS
The psychic secretion is the normal commencement, in the majority of cases, of secretory activity on the part of the gastric glands. If the meal be subdivided and administered at intervals, the psychic juice appears each time—Demonstration of “appetite juice” in a dog with an isolated gastric cul-de-sac. The work of the gastric glands if appetite juice be avoided by introducing food through a gastric fistula unperceived by the animal—Digestion of flesh by the stomach with and without sham feeding—Duration of the secretory influence of sham feeding—After the cessation of the psychic effect, how is the secretory work of the stomach maintained?—Experiments to prove the ineffectiveness of mechanical stimulation: excitation of the mucous membrane by means of a glass rod, a feather, a puff of sand, and by rhythmic dilatation of an india-rubber ball—Contact between the food and the stomach-wall may indirectly call the activity of the glands into play by awakening or increasing the desire for food.
Gentlemen,—On the last occasion we made ourselves acquainted with the first normal impulse which, in the natural course of events, calls into activity the innervation apparatus of the gastric glands. This impulse is a mental one, and consists in a passionate longing for food, that which in every-day life, and in the practice of the physician, is called “appetite,” and which everybody, both medical and lay, endeavours carefully to promote. We may now venture to say explicitly, APPETITE IS JUICE, a fact which at once displays the pre-eminent importance of the sensation. Medical science endeavours to assist the debilitated stomach by introducing the active constituent of gastric juice—pepsin—from without, or by prescribing other remedies believed to promote its secretion. It is, however, of interest to follow our experimental investigation still farther. What position is to be assigned to the “psychic” or “appetite-juice”[30] in the course of normal gastric digestion? Is any definite rôle to be attributed to it? What course does gastric digestion take when it is absent? Fortunately to all these important questions satisfactory answers are forthcoming by experiment. We have only to regret that these answers come so late.
Let us recall to memory how the secretion of gastric juice proceeded after feeding with flesh or bread in the case of our dog with the isolated miniature stomach. The following are the quantities and digestive capabilities of the first two hourly portions of juice after the administration of 200 grams of flesh or bread (experiments by Dr. Chigin):
| Hour. | Flesh. | Bread. | ||
|---|---|---|---|---|
| Quantity of juice. |
Digestive power. |
Quantity of juice. |
Digestive of juice. |
|
| c.c. | mm. | c.c. | mm. | |
| 1st | 12.4 c.c. | 5.43 mm. | 13.4 c.c. | 5.37 mm. |
| 2nd | 13.5 | 3.63 | >7.4” | 6.50 |
You see at once that the secretion of the first hour is identical in the two cases both as regards quantity and digestive power, and only in the second is the secretory work differentiated according to the nature of the food. How are we to explain the secretion which takes place at the commencement? Is it not the same which we have already seen in the sham feeding experiments? Is not this first onrush of the stream of secretion the preliminary psychic juice? Unquestionably, gentlemen, this is the case, and we may convince ourselves of the fact in the most diverse ways. Above all, the following is clear: whatever occurs in the so-called sham feeding cannot wholly be absent in the case of normal feeding, since the former is nothing else than the isolated commencement of normal digestion. This justifiable inference is fully confirmed, if the secretion of the first hours after the administration of flesh and bread be compared with that after simple sham feeding. In the case of feeding with flesh and bread, the identically similar and high digestive power of the first hourly portions is striking, and this power coincides with what we have met in sham feeding. Further, if the quantity of juice from the miniature stomach during the first hour be compared with that produced by the non-resected part of the organ,—to do which we must multiply it by ten, since the resected cul-de-sac is approximately one-tenth of the whole organ,—it is here again found that the quantity approximately corresponds to the mean values obtained by sham feeding. Finally, the depression in digestive power or quantity of juice (with flesh, decrease of digestive power; with bread, decline in the quantity of juice), which sets in soon after the taking of food, indicates that the two conditions are connected with the ingestion of food—i. e., with a transitory factor which soon passes away and gives place to other conditions. Our explanation becomes still more convincing when we take into consideration the effects of other foods. If you give the dog, for example, something else to eat which does not interest it to the same degree as flesh or bread, you will find the initial increase in quantity and strength of juice does not appear. Offer the animal milk, for example, which in sham feeding, especially if it does not last long, calls forth, as a rule, no secretion, or at all events only very little, and the rapid flow of the commencement—the already-mentioned initial rise—absolutely fails to appear. You have already seen the figures which deal with this matter; I think it necessary, however, to bring them forward again in order that you may be better able to compare them with the secretion after flesh and bread.
The dog was given 600 c.c. of milk (experiment by Dr. Chigin).
| Hour. | Quantity of juice. c.c. |
Digestive power. mm. |
| 1st | 4.2 | 3.57 |
| 2nd | 12.4 | 2.63 |
We have now begun the analytical examination of the variations of our secretory curve. But owing to the importance of the matter we did not confine ourselves to conclusions which might be drawn from earlier investigations. We turned to new forms of experiment for further proof.
Thus we divided the ordinary ration of flesh given to our dogs—400 grams—into four equal parts, which were administered at intervals of an hour and a half. (Experiments by Privat docent Kotljar and Dr. Lobassoff.) Each time after the dog received its 100 grams of flesh we were able to detect a rise both in the quantity and in the digestive power of the juice. The following table shows the figures in question:
| Half-hour periods. |
Quantity of juice. |
Digestive power. |
Remarks. |
|---|---|---|---|
| c.c. | mm. | grms. flesh given. | |
| 1st | 3.1 | 5.13 | 100 |
| 2nd | 5.0 | 4.63 | |
| 3rd | 4.7 | 4.50 | |
| 4th | 5.4 | 4.88 | 100 |
| 5th | 5.5 | 3.38 | |
| 6th | 4.7 | 2.75 | |
| 7th | 6.0 | 3.75 | 100 |
| 8th | 5.4 | 2.50 | |
| 9th | 5.9 | 2.50 | |
| 10th | 5.4 | 3.88 | 100 |
| 11th | 5.3 | 3.0 | |
| 12th | 4.2 | 2.5 |
In the curve which follows, only the variations of digestive power are represented.
It is clear that the increase, both of digestive power and of juice volume, is connected with the act of taking in food.
It appeared of interest definitely to determine the volume and properties of the secretion called forth by the act of eating in the dog with the isolated stomach. We endeavoured, therefore, at the beginning, to imitate the conditions of sham feeding as they occurred in the case of the dog with divided œsophagus. In addition to the fistular orifice leading into the isolated miniature stomach, another was opened into the main portion of the organ. If we now fed the dog in the ordinary way with small pieces of flesh, these were received back again at the orifice of the latter fistula, covered with saliva. Precisely as in sham feeding, after five minutes the juice began to flow simultaneously, from both the large and small stomachs. The secretion ran a corresponding course in the two cavities and ceased at the same length of time in both after the administration of food was stopped. Here is an instance taken from such an experiment performed by Dr. Lobassoff.
In five minutes the dog had eaten eighty pieces of flesh (weighing 172 grams), all of which soon afterwards dropped out at the fistula. The secretion began in both stomachs after the lapse of seven minutes from the commencement of the feeding, and proceeded as follows:
| Hour. | Miniature stomach. | Main stomach. | |||
|---|---|---|---|---|---|
| Quantity of juice. |
Digestive power. |
Quantity of juice. |
Digestive power. | ||
| c.c. | mm. | c.c. | mm. | ||
| 1 | 7.7 | 83.2 | 5.35 mm. | ||
| In consequence of a mixture with bile (10-15 c.c.) the digestive power was greatly reduced. | |||||
| 2 | 4.5 | 6.25 mm | 58.1 | ||
| 2½ | 0.6 | 8.5 | |||
The secretion from both cavities also came to an end at the same time.
This experiment proves to us, first, that the main and miniature stomachs work in perfectly parallel manner with each other. The beginning, the end, and the intermediate variations of the secretion correspond in both cases. Secondly, the digestive power of the secretion coincides in both, and is the same which was observed in the so-called sham feeding. It has here remained at the same height till the cessation of the secretion, without falling to the lower value which we observed from the beginning of the second hour onwards, after normal flesh feeding.
This was also confirmed later, when we performed an œsophagotomy on the dog, and carried out sham feeding in typical form. Here follows one of these experiments taken from Dr. Lobassoff’s article.
The first drop of juice appeared from both cavities during the sixth minute after commencing the feeding, which was kept up for half an hour. The further course of the secretion was as follows:
| Hour. | Miniature stomach. | Main stomach. | ||
|---|---|---|---|---|
| Quantity of juice. |
Digestive power. |
Quantity of juice. |
Digestive power. |
|
| c.c. | mm. | c.c. | mm. | |
| 1st | 7.6 | 5.88 | 68.25 | 5.5 |
| 2nd | 4.7 | 5.75 | 41.5 | 5.5 |
| 3rd | 1.1 | 5.5 | 14.0 | 5.38 |
| —— | —— | ——— | —— | |
| 13.5 (total) | 5.75 (mean) | 123.75 (total) | 5.5 (mean) | |
The secretion came to an end in both stomachs at the same time.
The above is represented in curves in Figs. 2 and 3, the scale on which that for the main stomach is drawn being ten times less than that for the small. As you see, the progress of secretion is identical in both.
The existence of a fistula leading into the large stomach affords us also the possibility of performing an experiment upon our dog which is exactly the converse of the sham feeding experiment, and which constitutes a real experimentum crucis. While in sham feeding, we had only, so to speak, the beginning of digestion before us, we are now able in our cross experiment to start at the continuation of this beginning. For this purpose it is only necessary to bring the food into the stomach through the fistula, without attracting the dog’s attention. Since in this experiment it is above all necessary not to excite the dog’s appetite, it is best to carry out the procedure on the sleeping animal. I may add at once, however, that the same result can be obtained on the waking animal, only the process must be performed unnoticed, and the animal’s attention must be diverted from thoughts of food.
The results of this experiment are striking, and do not in any way resemble the secretion after normal feeding. Some kinds of food, for instance bread and coagulated white of the hen’s egg, when directly introduced into the stomach, do not yield a single drop of juice during the first hour or more afterwards. This holds good both for the small and large stomachs. When a glass rod is introduced into the food contained in the organ it remains dry. Flesh, if introduced at this stage, is able to excite a secretion, but the appearance of the juice is considerably retarded. It begins from fifteen to forty-five minutes after the feeding, instead of from six to ten, is under normal circumstances extremely scanty during the first hour (3 c.c. to 5 c.c. instead of 12 c.c. to 15 c.c.), and possesses a very low digestive power.
Here is an experiment by Dr. Lobassoff:
| 400 grms. of flesh were brought into the stomach. | ||
|---|---|---|
| Hour. | Quantity of juice. c.c. |
Digestive power. mm. |
| 1st | 3.7 | 2.0 |
| 2nd | 10.6 | 1.63 |
| 3rd | 9.2 | 1.5 |
| 4th | 7.0 | 1.88 |
| 5th | 5.6 | 2.25 |
| 6th | 6.6 | 2.63 |
| 7th | 7.5 | 1.88 |
| 8th | 5.3 | 2.0 |
| 9th | 3.0 | 5.0 |
| 10th | 0.2 | — |
The secretion began twenty-five minutes after introducing the food. I now ask you to compare the following tables:
| Hour. | Fed with 200 grms. of flesh (Chigin). |
Flesh (150 grms.) brought into stomach (Lobassoff). |
Sham feeding (Lobassoff). |
Total quantity of juice in two experiments |
||||
|---|---|---|---|---|---|---|---|---|
| Quantity of juice. |
Digestive power. |
Quantity of juice. |
Digestive power. |
Quantity of juice. |
Digestive power. |
|||
| c.c. | mm. | cc. | mm. | cc. | mm. | cc. | ||
| 1st | 12.4 | 5.43 | 5.0 | 2.5 | 7.7 | 6.4 | 12.7 | |
| 2nd | 13.5 | 3.63 | 7.8 | 2.75 | 4.5 | 5.3 | 12.3 | |
| 3rd | 7.5 | 3.5 | 6.4 | 3.75 | 0.6 | 5.75 | 7.0 | |
| 4th | 4.2 | 3.12 | 5.0 | 3.75 | — | — | 5.0 | |
The progress of juice secretion in the above is also represented in the following curves:
As you see, the curve which represents the results of the direct introduction of flesh ascends much more slowly and does not attain anything like the height of that caused by normal feeding with the same food. But if the quantities obtained by direct introduction of the flesh be added to those of sham feeding, the resulting curve is almost identical with the normal.
In like manner the digestive power of the secretion in the foregoing experiments can be dealt with, and with the same result. It is a good instance of how a secretion curve can be synthetically constructed from its constituent factors.
Finally, I am able to demonstrate to you the following instructive experiment. In the presence of some of my listeners, whom I had invited to attend an hour before the lecture, I carried out the following procedures on two dogs, both of which had ordinary gastric fistulæ and were, besides, œsophagotomised. Into the stomach of one, while its attention was distracted by patting and speaking kindly to it in order to avoid arousing any thoughts of feeding, a definite number of pieces of flesh were introduced through the fistula. The morsels were threaded on a string, the free end of which was fastened to the fistular cannula by inserting a cork. The dog was then brought into a separate room and left to itself. A like number of pieces was introduced into the stomach of the other dog in the same way, but during the process a vigorous sham feeding was kept up, the animal being afterwards left alone. Each dog received 100 grams of flesh. Since then an hour and a half have elapsed, and now we may draw the pieces of flesh out by means of the thread and weigh them. The loss of weight, and consequently the amount of flesh digested, is very different in the two cases. In that of the dog without sham feeding the loss of weight amounts to merely 6 grams, while the flesh withdrawn from the stomach of the other dog weighs only 70 grams, that is to say, was reduced by 30 grams. This, therefore, represents the digestive value of the passage of food through the mouth, the value of an eager desire for food, the value of an appetite.
I give also a series of figures obtained by Dr. Lobassoff in analogous experiments. Into the dog’s stomach 25 pieces of flesh (100 grams) were brought. The flesh remained two hours in the cavity. Without sham feeding 6.5 per cent, with eight minutes’ sham feeding 31.6 per cent, of the quantity was digested.
Again: the flesh remained an hour and a half in the stomach; without sham feeding 5.6 per cent, with five minutes’ sham feeding 15 per cent, was digested.
Once more: the flesh remained five hours in the stomach; without sham feeding 58 per cent, with sham feeding 85 per cent, was digested, the balance of undigested food being 42 per cent in the one case and 15 per cent in the other.
I must, however, add that from the nature of this experiment it is not well adapted for class demonstration, and may often fail. On the one hand, it is not at all easy to conceal the introduction of the flesh from the dog; on the other, the unusual and distracting surroundings of the animal often causes a short period of sham feeding to have less effect than would otherwise pertain. In order to avoid such failures it is better before an audience to carry out this experiment only on dogs accustomed to appear in the lecture theatre, and of whose temperament the experimenter is well assured.
I hope you have now been convinced of the great importance which is to be attached to the passage of food through the mouth and œsophagus, or, in other words—and this, according to our former experiences, means the same thing—to the eager desire for food. Without this longing, without the assistance of appetite, many forms of food-stuffs which gain entry to the stomach remain wholly devoid of gastric juice. Others, it is true, excite a secretion, but the juice poured out is scanty and weak.
It is only later, when we have still more fully recognised the conditions upon which the secretory work of the gastric glands depends, that we shall be able to grasp the meaning of these facts in a more comprehensive manner. For instance, why does bread brought unnoticed into the stomach of the dog cause no secretion for hours, while flesh tolerably soon (after twenty to forty minutes) provokes this act? This will be explained in the next lecture; now, however, we must consider other questions.
How long does the after-effect, the echo of the first impulse to the secretory nerves of the stomach, continue to last? How long does appetite juice continue to flow after the normal act of eating, which, especially in the case of animals, is not of long duration? We have already determined many times, not only on our dog with the isolated stomach, but also on other animals, how long the after-effect of sham feeding is continued.
Here, for example, is an experiment from the article of Professor Ssanozki which deals with the point. The dog had a gastric fistula and also an opening leading into the œsophagus. After a sham feeding of five minutes the secretion began, and was continued as follows:
| Time in minutes. |
Quantity. | Digestive power. |
|---|---|---|
| c.c. | mm. | |
| 10 | 25.5 | 8.1 |
| 10 | 20.0 | 8.0 |
| 10 | 13.5 | 6.8 |
| 10 | 11.0 | 7.5 |
| 10 | 8.5 | 8.1 |
| 10 | 6.5 | 9.0 |
| 20 | 13.5 | 7.4 |
| 20 | 11.0 | 7.2 |
| 20 | 7.0 | 7.2 |
| 20 | 11.5 | 6.8 |
| 20 | 11.0 | 6.5 |
| 30 | 6.5 | 7.6 |
| 20 | 5.5 | 7.2 |
The effect, therefore, even after a short period of sham feeding, stretches over a length of time. Naturally the same holds good for the taking of food in the normal way. One must, however, bear in mind that in sham feeding, with all the force and reality of a hunger sensation not satisfied, the eager desire for food, the effective agency, becomes more and more accentuated, and therefore the secretory influence is prolonged and more powerful. In normal feeding, however, the quelling of the longing, the feeling of satisfaction which, as is well known, sets in long before the termination of the digestive period from the mere filling and distension of the stomach, must diminish the desire for food, and, consequently, bring the secretory effect to an end.
It is, therefore, improbable that the whole secretory process in the stomach, which, in the case of certain kinds and quantities of food, lasts from ten to twelve hours, is dependent on the factors which we have up to the present investigated. This is all the more obvious since a sham feeding of five minutes, even under the most favourable circumstances, does not call forth a secretion for longer than three to four hours. We must, therefore, seek for other exciting agencies of the innervation apparatus of the gastric glands.
Why and wherefore is the secretion instituted by psychic influence maintained? What would first occur to all your minds is naturally the immediate influence which the food exerts upon the walls of the stomach. And this is true, but it does not happen in the simple, direct fashion current in the minds of many physiologists and physicians. When I said that bread or boiled white of egg, introduced directly into the stomach, may not for hours produce a trace of secretion, probably many of my hearers may have asked themselves with natural astonishment, “How, then, is the effect of the forced feeding of phthisical and insane patients, and the artificial feeding of those with gastric fistulæ (performed on account of stricture of the œsophagus) to be explained?” I will introduce my answer by a very unexpected pronouncement relative to the assertion that mechanical stimulation of the stomach wall by food constitutes a reliable and effective means of calling forth the secretory work of the glands. This assertion, which is so categorically set forth in many text-books of physiology, and which consequently has gained hold of the mind of the physician, is nothing else than a sad misconception degenerated into a stubborn prejudice. My own statement, repeated in many published articles, and at the meetings of various medical societies, that this dictum is only a picture of the imagination, has met, for the most part, either with an unbelieving shake of the head or else with a direct avowal that “it cannot be so.” I regret exceedingly that these steadfast unbelievers are not here, so that we might together bring the matter before the tribunal of fact, to the demonstration of which we will now proceed. To this matter I attribute very great importance. It is on this ground, according to my opinion, that the whole battle must be fought out between the generally accepted view that every agency is capable of exciting the gastric mucous membrane and the theory that it is only excitable by specific and selected stimuli. If once the defenders of the old opinion are driven from their position and obliged to admit the inefficiency of mechanical stimulation, there would be nothing further left for them than to build up new theories and search out old facts concerning gland work which have hitherto been rigidly kept in the shade. We may take it that it is mainly because people were so seized with the belief in the direct and simple mechanical explanation that Bidder and Schmidt’s experiment of the excitation of gastric secretion by mental effect has been so little taken into consideration, notwithstanding that it appeared so thoroughly reliable and convincing.
I will now repeat the experiment of mechanical stimulation of the gastric mucous membrane before you in the well-known, traditional, and classic manner. Here is a dog with a gastric fistula on which a cervical œsophagotomy has in addition been performed. I open the fistula; as you see, nothing flows out of the stomach; it was washed out clean with water an hour ago. We take the celebrated feather and also a tolerably strong glass rod. Folds of blotting-paper saturated with red and blue tincture of litmus are placed at hand. I now ask my assistant to continuously move the feather and glass rod, alternately, in all possible directions in the stomach, changing from one to the other every five minutes. On removal from the stomach each is carefully dried with red and blue blotting-paper. You have all seen, gentlemen, that this procedure has now been kept up for half an hour. From the fistular orifice not even a single drop has escaped, and, moreover, the drops of moisture on all the pieces of red blotting-paper I have been able to hand to you have assumed a distinct blue tinge, caused by the moisture of the alkaline mucous membrane. The blue pieces, however, have merely been made wet without altering their colour. Consequently, with the most thorough mechanical stimulation of the whole cavity of the stomach, we have not been able to find a single spot possessing a noticeable acid reaction. Where, then, are the streams of pure gastric juice of which we read in text-books! What objection can be raised against the conclusiveness of this experiment? In my opinion only one: that we are dealing with a dog out of health, whose gastric glands from some possible cause are unable to react normally. This single objection can be set aside before your eyes. After failing with the mechanical stimulation, we proceed forthwith to the sham feeding of the same animal. The dog takes the food offered it with keen appetite, and you see that, exactly five minutes after beginning the feeding, the first drops of juice appear from the stomach, followed by others faster and faster. I catch a couple of drops on the blue litmus paper, and you see that they produce bright red specks on the blue sheet. After thirty minutes’ sham feeding we have collected 150 c.c. of juice, which, without filtering, looks as clear and transparent as water.
We cannot, therefore, possibly doubt that, when the proper stimulus is used, the gastric glands react to it in a perfectly normal fashion, furnishing a healthy gastric juice. From this it irrefutably follows that only one explanation is to be found for the negative result in the first half of our experiment, viz., that the mucous membrane of the stomach, so far as secretory activity goes, is perfectly indifferent to mechanical excitation. And yet this mechanical stimulus is demonstrated as an exciting agency in the physiological lecture theatre. I venture to think that this lecture experiment from now onwards will quit the field, and give place to the one I have just shown you. This apparently simple experiment of mechanical stimulation can, however, only be successfully performed when certain very obvious rules are followed. These, however, physiologists have not observed, probably on account of a preconceived belief in the effectiveness of the mechanical stimulus. These rules are two. First, it is necessary that the stomach should be clean, and that nothing shall gain entry to it from without. Such conditions were not formerly fulfilled. It is true the stomach was emptied by removing the stopper from the fistular cannula, but it was not washed out till an acid reaction was no longer given, and consequently preformed gastric juice was left behind between the folds of the mucous membrane. At the same time saliva from the cavity of the mouth could gain entry, which quickly became acidified in the incompletely emptied and imperfectly washed-out organ. It is, therefore, not surprising that the glass tube, by setting up contractions of the stomach, was the means of expressing small quantities of acid fluid from the fistula-tube. (The relationship between mechanical stimulation and the motor functions of the stomach is not to be confounded with what we are here speaking of.) That matters are as I state, and that the facts correspond to the explanation is proved by this; namely, that nobody till now has obtained genuinely pure gastric juice of an acidity amounting to 0.5 or 0.6 per cent. It is only necessary to call to mind that Heidenhain, when determining the acidity of the juice first obtained from the resected stomach, was placed in no little doubt as to whether his results (0.5 to 0.6) were correct, and his assistant at the time (Gscheidlen) was set to verify the correctness of his standard solutions. The acidity of the “purest” juice known at that time was scarcely 0.3 per cent. As a further proof that none of the older observers ever really obtained a secretion from mechanical stimulation pure and simple, we may adduce the fact that none of them made mention of the constant and precise period of five minutes’ latency. To overlook this was not possible if a genuine excitation of the glands had been obtained.
Of no less importance is the second condition when we wish to perform the experiment of mechanical stimulation in the correct way. It is very necessary that the gastric glands be not already in activity at the beginning of the experiment, and also that during the experiment no impulse comes into play, which of itself, apart from mechanical excitation, could excite the glands to secretion. Nor have we any proof that observers formerly waited for hours before commencing the experiment and convinced themselves that the gastric glands had ceased working. On the contrary, we have not the slightest evidence to indicate that the authors had attempted to guard against accidental psychical stimulation of the glands—a matter which we have seen is of considerable difficulty. And some dogs are so easily excited in this way that it is almost impossible to bring their glands to rest, or at least it is necessary to wait for hours. The experimenter must strain his whole attention to preserve such an experiment free from objection. It is only necessary that some food be near the dog, or that the hands of the attendant who has prepared the food should smell of it, or that some other similar circumstance should come into play, and the glass tube, quite undeservedly, will be made answerable for the excitation of the gastric glands. As you have just seen, both of our conditions have been fulfilled on the dog before you, and the result of the experiment stands in irreconcilable contradiction to those of the laboratory and lecture experiment of former times.
The importance of the experiment, which I have already dwelt upon, justifies me in making still further demands upon your attention in order to show you two modifications of it. Nobody has as yet said, with regard to mechanical stimulation, that in order to obtain results the mechanical agency must simultaneously come into contact with numerous points of the inner surface of the stomach. But in order to meet this possible objection I will now show you two new modifications. Again a similar dog is used, that is to say, one on which both gastrotomy and œsophagotomy have been performed. The stomach has been washed out clean and is at present in a state of complete rest. Into the fistula I bring a thick glass tube containing a number of small openings (2 to 3 mm. diameter) at its rounded end. The other end of the tube is connected with a glass ball containing tolerably coarse sand. Leading into the ball is a second tube, with which an india-rubber pump can be connected and a blast of sand blown through. By rhythmic compression of the india-rubber ball I inject sand with considerable force into the stomach, and this play is kept up for ten to fifteen minutes; nevertheless, we see no trace of gastric juice. The sand falls out again between the side of the cannula and the glass tube, and it is either dry or scarcely moistened, but in no case is it able to turn blue litmus red. And yet we are here dealing with a strong and widely diffused stimulus. Look for a moment at the performance of the bellows outside the stomach. From every opening of the tube—numbering considerably more than ten—a strong stream of sand is ejected. If you hold your hand against it, you feel quite distinctly that the grains of sand strike with considerable force. And now, when our experiment is ended, we may convince ourselves by sham feeding, in easy and unquestionable fashion, that the innervation of the dog’s stomach is perfectly normal.
Yet another experiment on a similar dog. Into its empty and resting stomach an india-rubber ball is introduced. This is distended with air by means of a syringe till it is as large as a child’s head and maintained in this condition for a time, afterwards being allowed to collapse. The procedure is kept up for ten to fifteen minutes. During this time not a single drop of juice has appeared from the stomach. The surface of the ball taken out of the organ is everywhere alkaline. And here also subsequent sham feeding shows that the dog is in a suitable condition for the experiment. I must add that in making this observation the dog must not be too hungry, that is to say, must have been fed within ten to twelve hours before, otherwise a psychic excitation of the secretion can readily be induced.
If one dispassionately regards this question, and if any of our methods for the study of gastric secretion are reliable, one must be convinced step by step in the laboratory of the uselessness of mechanical stimulation. In the case of dogs with an ordinary gastric fistula, and failing some special reason, not a drop of gastric juice ever escapes from the stomach other than during the digestive period. How could this be the case if the mechanical stimulus were effective, since the inner rim of the fistula-tube is continuously in contact with the gastric mucous membrane? The same holds good for the dog with resected stomach. During the experiment a glass or india-rubber tube is brought sufficiently far into the cul-de-sac to catch the juice, and yet not a drop flows through the tube, nor does its inner surface ever become acid, so long as true secretory conditions are absent. Moreover, the tube has tolerably often to be taken out and set right.
In the ordinary gastric fistula in dogs, when the operation has lasted a long time—over a year—folds of mucous membrane are often formed in the neighbourhood of its inner orifice which completely close the tube. In these cases a long, thick, perforated metal tube has to be passed in deeply, and yet the manipulation of itself never calls forth a secretion. Further, it is a daily occurrence to find in the stomach of the dog thick rolls of hair, and yet their presence in no way hinders the arrest of the secretion, which occurs when digestion has ceased. Such an occurrence would have been specially obvious in our dog with the isolated stomach, since it was bedded with sawdust in order to guard against maceration of the wound by juice trickling out. Very often we found enormous quantities of sawdust in the stomach, as much as half a pound weight; obviously the dog had licked the wound from adherent sawdust, which it then swallowed, together with that sticking to its nose. And yet these particles of sawdust of themselves, which certainly acted as mechanical stimuli, never caused a secretion. It appears to me that this long series of facts ought to suffice to carry the supposition to its grave that by direct mechanical stimulation one is able to set the neuro-secretory apparatus of the stomach into activity.
And yet the feather and the glass tube continue the even tenor of their ways to this moment and function in some text-books, yea, even in articles which specially treat of gastric secretion as exciters of the gastric glands. There are, it is true, a few physiologists who hold mechanical stimulation, in relation to gastric secretion, not to be very effective, and give it a subordinate position in the series of exciting agencies, but as yet I know of no other physiologist who has wholly denied its influence, and who has not held it possible to obtain at least some juice by it.
To conclude this lecture, we will take into consideration a question connected with the matter we have just discussed. Since the contact of food with the gastric mucous membrane has no direct influence on the secretion, is its entry into the stomach devoid of all connection with the secretory process?
It can hardly be doubted that, under normal conditions, the stomach is the seat of certain definite sensations, that is to say, its surface has a certain degree of tactile sensibility. This sensation is, as a rule, very weak, and the majority of people become accustomed to pay no heed to it in the normal course of digestion. They obtain their sensations of general well-being, and especially of satisfaction from the enjoyment of food, without taking cognisance of the factors contributing to them. The feeling of general hunger, however, is referred solely to the stomach.
On the other hand, all of us have met with men who could describe exactly, and with gusto, how they were able to follow a special tit-bit, or a mouthful of a favourite wine, the whole way through the œsophagus down to the stomach, especially when the latter happened to be empty. Naturally the gourmand, who directs his attention continuously to the act of eating, can in the end distinctly perceive sensations, and even call them up to the consciousness, which in other people are normally masked by other sensations and impressions. We may therefore take it that the satisfaction derived from eating is caused not only by stimulation of the mouth and throat, but also by impulses awakened by the passage of the food along the deeper portions of the œsophagus and by its entry into the stomach. In other words, food which merely passes through the mouth and throat produces less enjoyment and excites, therefore, a less feeling of appetite than the food which passes the whole way into the stomach. The appetite, the eager craving after food, is, indeed, a very complex sensation, and often not merely the need of the organism for food material is necessary for its excitement, but also a condition of thorough well-being, together with a normal healthy feeling in all parts of the digestive tract. For this reason it is easy to understand how patients who have diseased sensations in these organs, and who have no feeling of appetite, no desire for food, remember the sensations, whether consciously or unconsciously, even when they are no longer present. Cases are known to neuro-pathologists where people with gastric anæsthesia suffered from this loss of appetite. Such patients are no longer conscious of having stomachs, and dislike the idea of eating because the food, as they express it, appears to fall into a strange empty sack. In this way one can also conceive how the appetite becomes lost in cases of long-continued obstruction of the alimentary tube. The patients forget their stomachs, and in such instances direct introduction of food into the organ, after an operation, may suddenly bring back the appetite.
As a further illustration, I may be permitted to give an instance from my own personal experience. After an illness with which a transient but high fever was associated, although otherwise fully recovered, I had lost all desire for food. There was something curious in this complete indifference towards eating. Perfectly well, I only differed from others in that I could with ease abstain from all food. Fearing that I should collapse, I resolved on the second or third day to endeavour to create an appetite by swallowing a mouthful of wine. I felt it quite distinctly pass along the œsophagus into the stomach, and literally at that moment perceived the onset of a strong appetite. This observation teaches that the tactile sensation of the stomach at the moment of entry of food is capable of awakening or increasing the appetite. It is known that withholding food from the organism, or in other words the creation of a necessity for food, does not lead immediately, nor in all cases, to the production of an appetite, to a passionate craving for food. How often does it happen that the ordinary hour for a meal has struck, and yet, owing to some keenly interesting occupation, not the least desire for food is felt? It is known to everybody, indeed it has become a proverb, that real appetite first sets in with eating. If this be true, the initial impulse towards awakening an appetite may originate in the stomach and not in the buccal cavity. When we spoke above of the desire for food being the excitant of the secretory nerves of the stomach, we naturally meant the passionate and conscious longing for food, that which is called “appetite,” and not the latent need of the organism for nourishment, the lack of nutrition, which has not yet been transformed into a concrete passionate desire. A good example which enables us to differentiate between these two factors is furnished by our dogs with sham feeding. The necessity for food exists in such cases even before the experiment; the juice, however, only begins to flow as soon as this need has taken the form of a passionate longing. It is therefore quite possible that in the case of some dogs, and at a certain stage of hunger, the touching of the gastric mucous membrane with any object at hand, its mechanical excitation, its distension by the food mass, may give the impulse which excites the appetite, and when the appetite is awakened the juice flows. This is possibly a third reason why, in the old experiment, the mechanical stimulus came to be considered effective. Viewed from this point it may, to a certain degree, lead to a reconciliation between my assertion concerning the inefficiency of the mechanical stimulus and the generally prevailing belief. I further also admit that mechanical excitation will at times call into play the work of the gastric glands, not however directly by means of a simple physiological reflex, but indirectly, after it has first awakened and enlivened the idea of food in the dog’s consciousness, and thereby called forth the passionate desire. I hope that the foregoing will in no way lead to a confusion of ideas in your minds, but will assist you to an exact and concrete analysis of the previous simple explanation of the facts. This representation, which bears more or less of a hypothetical character, could, of course, be submitted to experimental proof. For such it is only necessary to compare the influence which sham feeding exercises in an œsophagotomised dog with that in one having a simple gastric fistula.