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THE WORK OF THE DIGESTIVE GLANDS .
M. ETC. LONDON HON.KC. ST.) KING'S PROFESSOR OF THE INSTITUTES OF MEDICINE. ETC.D. . PARIS. MEMBER IMPERIAL MILITARY ACADEMY OF MEDICINE . LIPPINCOTT COMPANY 1902 [All riyhts cc-scrm/] . ILLUSTRATED LONDON CHARLES GRIFFIN & COMPANY.[SOLE AUTHORISED ENGLISH TRANSLATION] THE WORK OF THE DIGESTIVE GLAFDS ^Lectures bv> PKOFESSOIl J. ST. PETERSBURG . TRINITY COLLEGE. BELFAST EXAMINER IN PHYSIOLOGY. DIRECTOR OK THE PHYSIOLOGICAL DEPARTMENT OK THE INSTITUTE EXPERIMENTAL MEDICINE. ROYAL UNIVERSITY OF IRELAND.. QUEEN S COLLEGE. H. TRANSLATED INTO ENGLISH BY W. (ENCI. AND PROFESSOR IN THE IMPERIAL MILITARY ACADEMY OK MEDICINE. PAWLOW KR . F. B. DUBLIN LATE DUNVILLE PROFESSOR OF PHYSIOLOGY. ETC. LIMITED PHILADELPHIA: J. M. P. AND ROYAL COLLEGES OF PHYSICIANS AND SURGEONS. THOMPSON.Cn. ETC. PETERSBURG FOREIGN ASSOCIATE OF THE ACADEMY OF MEDICINE.S.
TO THE MEMORY OF HIS FRIEND. THE TALENTED PHYSICIAN NIKOLAI PETROWITSCH BOGOJAWLENSKY THE AUTHOR DEDICATES THIS WORK .
to make use of the German . and Salivary Secretion. PHYSIOLOGICAL DEPARTMENT. therefore. J. that in its English form the book will be worthy of its distinguished author. 1902. But the work is of no less value to the PRACTISING PHYSICIAN. A. TRINITY COLLEGE. Succus Entericus. THE TRANSLATOR. . It is to be hoped. text. with more recent notes kindly supplied by the author. therefore. Walther. In carrying this into effect the translator has had the ready permission of Dr.PREFACE TO THE ENGLISH EDITION THE first great importance of the results obtained by Professor Pawlow. DUBLIN. To place the matter. thus together bringing the whole up to date. F. It also contains two new figures. a kindness which he desires to gratefully acknowledge The present edition includes the later work of Pawlow on the Physiology of the Bile. within easy reach of every English speaking medical man it was felt that an English edition was called for. SCHOOL OF PHYSIC. and also of Mr. by others in German and French. September 1. Bergmann. published in collected form in the Russian language in was soon recognised by PHYSIOLOGICAL INVESTIGATORS all the world over. Hence the Russian edition was quickly followed 1897.
But the object of the is experiment. varied. elaborated. experimental investigation which. in these lectures. and of the pancreas. which have been constantly tested.PREFACE TO THE RUSSIAN EDITION IT was not at all my intention. work of my laboratory for nearly ten years experiment which deals with the functions of the gastric glands. has been many times repeated. to treat of every- thing which has been written concerning the work of the digestive I only wished to make known the results of an glands. In the description of the several experiments I always mention the author. spoken of from the point of view of the laboratory. and now appear all to be securely established. I wish to indicate the whole laboratory. lost its fragmentary character and grown into a complete whole. advantage to the reader to see how a uniform guiding principle has developed. and to communicate the same to my hearers. latest views of our laboratory . without giving It is of essential the individual opinions and views of the author. and taken shape in the form of tenable and harmoniously linked experiments. partly by word of mouth and partly by and since every direct demonstration. the material has. it embraces all even the most recent. When I employ the word " we " in the following text. The subject of these lectures represents the . correctly indicates the present position of the subject. frequently corrected. for us at least. In its production my fellow workers have . I am convinced. and extended . its meaning and its position in the whole series. In its main idea the book embodies the the facts.
to all our dear co-workers. PREFACE but it is a joint work. When I look back upon what the laboratory has accomplished in our field of research. with the hope that they have preserved as friendly recollections of us as we have of them. later. ST.x individually taken part principle. but in its totality to the guiding conception which has inspired us all. I should. the result of the It which animates the whole laboratory. owes its existence to the acuity of each individual. therefore. . who are widely scattered over our native land. PETERSBURG. . I know well how much the work of every individual is to be valued. and All the Military Medical Academy. April 1897. in THE AUTHOE. in a shorter form. which were demonstrated before both these the experiments audiences have been included in the work. These lectures were medical first delivered before an audience of men in the Institute for Experimental Medicine. like to take this opportunity of sending heartiest greetings. in the name of the laboratory.
76-92 MECHANICAL STIMULATION OF THE NERVOUS APPARATUS OF THE GASTRIC GLANDS . A GENERAL SURVEY OF THE SUBJECT METHODS . 45-61 IV. THE NORMAL EXCITANTS OF THE NERVOUS MECHANISM OF THE PANCREAS SUMMARY OF MATTERS DEALT WITH. . 62-75 PERIOD OF OCCURRENCE AND IMPORTANCE OF THE PSYCHIC OR APPETITE JUICE IN THE SECRETORY WORK OF THE STOMACH THE INEFFICIENCY OF Pp. Pp. AND OF THE PANCREAS . 1-19 II. AND PROBLEMS FOR FURTHER INVESTIGATION Pp. POTENT EXCITER OF THE GASTRIC SECRETION V. THE CENTRIFUGAL (EFFERENT) NERVES OF THE GASTRIC GLANDS. VI. Pp. . . THE CHEMICAL STIMULI OF THE NERVOUS MECHANISM OF THE GASTRIC GLANDS THE MINIATURE STOMACH A RELIABLE METHOD OF COMPARISON SEAT OF ACTION OF THE CHEMICAL STIMULI HISTORICAL Pp. . 112-130 .CONTENTS LECTURE I. 20-44 III. GENERAL SCHEME OF AN INNERVATION APPARATUSTEE WORK OF THE NERVOUS APPARATUS OF THE SALIVARY GLANDS APPETITE THE FIRST AND MOST . THE AVoRK OF THE GLANDS DURING DIGESTION . Pp. 93-111 VII.
187-189 Pp.xii CONTENTS PHYSIOLOGICAL ACTION AND THE TEACHING OF INSTINCT LKCTURE VIII. . 131-147 LATER RESEARCHES THE COLLABORATION AND MUTUAL BILE INTERDEPENDENCE OF THE DIGESTIVE JUICES AND Succus ENTERICUS . 191-196 INDEX . PRESENT-DAY MEDICAL Pp.. 148-186 Pp. EXPERIENCES OF THE PHYSICIAN IX.. Pp..- MOVEMENTS OF THE STOMACH PATHOLOGY AND EXPERIMENTAL THERA- PEUTICS OF DIGESTION THE METHOD OF EXPERITO MENT INDISPENSABLE REQUIREMENTS LITERATURE . ..
references. I could give a general survey only of the work of many years. therefore. proved to be that which consequence dwells in the stomach and the pancreas. S.. desire. arose to replace the older teaching by bringing into life a fuller and more correct representation. cies In my and to be able to convince present lectures I hope to make good these deficienmy hearers by relating actual experiof the lectures is ments. of myself and my many years. P. * The substance of the taken from work which for in St. A GENERAL SURVEY OF THE SUBJECT : METHODS. ever. With this object I gave. 1X1(4-1)5 (Russian). and was denied the possibility of sustaining my words with documentary Russian clinician. the most important organs mind of the physician. viz. and I believe we have obtained results. howBotkin. an oration* before the Festival Meeting of the Society of Russian Physicians in St. in 1H04. which deserve serious consideration. The work in the alimentary canal.e. Petersburg. The author's operative experience forming a stomach cul-tlr-sac : Importance of surgical methods of a physiological laboratory. and which in A which was dedicated to the memory of the celebrated In the short space of one hour. so far as it concerns the of digestion.THE WORK OF THE DIGESTIVE GLANDS. LECTURE I. has not by any means is represented in text-books. in physiology The surgical department GENTLEMEN. Ti-diixiirtionx Sin: I'lisx. (Icni'i'iil Hut-raj : Introductory The digestive apparatus may be comUnsolved problems in the physiology of pared to a chemical laboratory digestion Methods. A . their ideal requirements Temporary and permanent 1 the latter Difficulties connected with making pancreatic fistula the same combined with oesophagotomy Methods of Gastric fistula. Physicians Peterslury.. attention of for The physiology my whole laboratory of the digestive glands has engaged the fellow workers i. both theoretical of secretion and practical.
? still remain undeare the fluids in why poured out on the raw material in one particular order and not other any Why are the pi'operties of certain reagents often repeated in .THE WORK OF THE DIGESTIVE GLANDS. at another several. where the food. the so-called ferments. each single compartment is provided These are either prepared in neighbouring buiied in the walls of the structure itself. Physiology has learned all this by obtaining either the fluids in question or the pure ferments from the organism. albumin. They are thus brought into a condition in which they are capable of being absorbed into the body fluids and made This factory conuse of for the maintenance of the processes of life. are submitted to an essen- chemical process. But our conception of the digestive process. which are connected. The distant and separate organs. These latter are the so-called secreting glands with their excretory ducts. suffers from many and not unimportant defects. by a system of transmitting tubes. or else in workshops. But even a juice which has only one ferment is a very complex fluid. each of which acts in its own special way. alkalies. since. as in other large chemical factories. on the other. and studying. Many questions many have not even been raised. endowed with definite chemical properties which only act on certain portions of the food. which is essentially reciprocal behaviour towards each deductive. as we say. this latter being ordinarily formed of a complex mixture of different ingredients. the most part has already appeared in print. or even the empirical cided. little factory and. Indeed. own These properties are chiefly determined by the presence of special substances in the reagents. it holds other substances to wit. it may where the raw materials tially the foodstuffs be compared to a chemical factory. sists a series of compartments. is either retained for a time or at once sent on to the of next. in the test-tube. teaching of dietetics. its Each of the workshops furnishes a special particular reagent. their effects upon the constituents of the food as well as their also in solution other. But many unpublished facts which the laboratory is now in possession of will also be referred to. with the main laboratory. it is mainly upon thus acquired that the teaching of the science with regard to knowledge the elaboration of the food or. &c. in addition to the ferment. They thus combine the properties of many individual reagents. acids. A con- siderable gap without doubt exists between such a form of knowledge on the one hand and the physiological reality. attack at one time only a single ingredient of the food. the digestive juices. In view of the chief duty which the digestive canal has to perform in the living organism. fluid. as they are usually termed. indeed. with suitable reagents. according to its properties. The separate fluids. For example. of digestion. is based.
how this digestive fluid is provided. principles. This method has been adopted by many investigators who have studied the progress of the secretion of the various digestive juices. and most carefully adapted to the task in hand. represents one of the most intricate sections Nor is it enough for the physiologist to have a knowledge of the separate elements concerned in the process of digestion. comprehensive knowledge of the processes of digestion may be acquired in one of two ways either by determining in what state of elaboration the raw material is to be found at each separate part of the digestive canal and his pupils this or. include also within the sphere of his observation the actual progress of digestion as a whole. in that one may require a special reagent the activity of which would with that of others on the remaining ingredients ? No one can deny that these questions touch upon the vital facts of the interfere case. beautifully performed. often said. that is to say. we must (t priori admit that for each meal i. therefore. He must. and does this happen with every kind of food hat gains entry to the digestive canal ? Are the reagents subject to variations. by ascertaining the exact quantity of the digestive fluids which is secreted for each individual constituent of the food. If we reflect. work of the digestive canal in every single case is elaborately contrived. the working of the individual agencies.e. stages It is . how and why I do such alterations appeal' ? Do these variations only concern the composition of the fluid as a whole.METHODS. to be wondered with special properties is produced.. This was recognised by many and doubtless would have accomprevious investigators who attempted. and not without truth. A was the method on of Brlicke. plished. and when it is poured into the alimentary canal. apart from some general and empirical each set of materials to be dealt with of therapeutics. for a suitable combination of reagents It is not. at that the subject of dietetics. as well as of Ludwig the other hand. The mechanism of digestion cannot be presented in the abstract manner which is current in the physiological teaching of the present The differences and complexity of the reagents indicate that the time. combination with other properties ? Are all the constituents of a particular fluid simultaneously poured out on the food. in order to fully grasp his subject. that science advances by determined by the results obtained from particular method?. the solution of the problem had it been of an easier description. as well as for the meal as a whole. and if so when. or may the individual constituents alter in different cases and in different directions according to the requirements of the raw material ? How do the reagents vary with augmented or diminished activity of the whole factory ? Is there not a species of contest between the different constituents of the food.
THE WORK OF THE DIGESTIVE GLANDS.
With each advance
in technique \ve reach a higher level from which a open to us, and in which we sec events previously
out of range.
problem consisted, therefore, in the working out of a had to observe how the reagents were discharged upon
the food brought into our digestive factory. To accomplish this in an manner required the fulfilment of many and difficult conditions.
Thus it was necessary to be able to obtain the reagents at all times, otherwise important facts might escape us. They must be collected in
if we were to determine how their composimust be able to estimate their quantities accurately ;
was necessary that the
digestive canal should function
be in perfect
normally, and that the animal under experiment should
but natural that the solution of
these difficult problems
only be gradually achieved by physiology, that not a little trouble should be spent in vain, and that many investigators should
see their efforts fruitless, notwithstanding that several of the most prominent representatives of our science have devoted their attention
to this field.
begin our consideration with the pancreas, as the simpler case.
might appear that here our problem was very light. We have apparently only to seek out the duct through which the secretion of the gland is delivered into the intestine, to fasten a cannula into it, and
thereby afford a free outflow of the fluid towards the exterior, collecting All this, in reality, is very easily done, but it in a graduated cylinder.
our problem is far from being solved for digestion may be very active, yet, as a rule, there is no flow of pancreatic juice from the tube after
the operation ; or if there is one the quantity is very small and obviously sub-normal. In such a case it would be out of the question to observe
the rate of secretion, still more to determine the alterations in the juice dependent upon the nature of the food. On following the matter up, it became evident that the gland is a very sensitive organ, and
suffers such a severe disturbance of its activity
from the unavoidable con-
ditions of the operation (narcotisation, opening of the abdominal cavity), that in the majority of instances not even a trace of normal function
in the literature
under the name of
temporary attempts being made on other
of success naturally led to
was hoped that an improvement might be attained by collecting
the juice some time after the completion of the operation, that
disturbing influence had fully passed away. The fluid was therefore allowed to escape freely from the excretory ducts for a considerable time.
METHODS: 1'ANCREATIC FLSTUL.K.
This was accomplished either by tying a glass tube into the duct and leading it through the abdominal wall (Claude Bernard), or by fastening in a T-shaped piece of twisted lead wire in a similar manner (Lvd-wig's
" These were named " permanent fistulpe. Both modifications proved effective, but only for a short period, generally for three to five After this time days, in exceptional instances for as long as nine days.
the glass tube
out and the fistula closed up
even the lead wire
was unable to prevent this occurrence.
also be regarded as
inhibitory influence of the
was not their operation Lad
passed off after one or two days, another abnormal condition, in many instances set in, viz., an incessant irritation of the gland producing a
whether the dog was fed or not. The question " " then arose, which was the better the " temporary or the " permanent " fistula ? temporary form Evidently neither was faultless. In the
secretion independent of
the conditions were rendered abnormal by the effects of the operation " " form by inflammatory results in the in the so-called permanent
pancreas, which often set in (especially in the older laboratories) within one or two days.
Only one thing remained, namely, to discover a means of access to the gland lumen which would keep the duct open for any length of time that is to say, till the above-mentioned disturbances had completely disSuch a means was first described by me in the year 1*7!', appeared.
and afterwards independently in the year 1880 by Heidenhain.* My method was as follows. It differs slightly from Heidenhain's. From the wall of the duodenum, an oval piece, containing the orifice of
the pancreatic duct,
appreciably narrowed, stitched up,
cut out, the bowel, the lumen of which is not and the isolated piece of intestine
sewn (with the mucous membrane outwards) into the slit in the abdominal wall. The whole heals quickly the opei-ation, which requires no special skill, is only of short duration (about half an hour), and is well borne by the animals. After two weeks they are ready for observation. In the healed-up wound a roundish elevation, 7 to 10
mm. in diameter, is to be seen. This is formed of mucous membrane, and in the more successful cases shows the cleft-like orifice of the duct
If the animal be now supported in a suitable exactly in its middle. the juice may be either directly collected as it drops from the frame, mucous papilla, or if there be a tendency to escape along the abdominal
of a funnel with its wide
fixed in the requisite
Neither of the two disadvantages which beset the investigators
or " permanent
THE WORK OF THE DIGESTIVE GLANDS.
The gland undoubtedly remains in a normal condition, but the difficulties have by no means reached an end. In a very short time the abdominal wall becomes macerated by the escaping juice, and in
of the experimenter
These continuously places even fairly large bleeding patches appear. irritate the animal and prevent the collection of pure juice by means of
What was to be done ? Many things helped e.g., frequent washing of the macerated skin with water and smearing with emollient
funnel tied in position.
however, still better promoted if the dog hours every day in its frame, with the But the best means of all is to allow the
animal constantly to
upon somo porous material, such
except during the hours of the experiment, as a bed of sawdust or sand or old
animals soon discover the best position in which to the escaping juice is at once absorbed by the
In this way the abrasion and maceration of the skin porous material. can most readily be avoided. It is interesting to relate that the hint
which led to the adoption of this last method was given by one of the animals operated upon. I may perhaps take the liberty of giving a fuller account of this In one of the dogs the eroding effects of the juice interesting case.
became evident after ten to fifteen days. The treatment employed At night the dog was tied up in the laboratory, yielded no good results. but one morning, to our great annoyance, we found a heap of mortar beside it, torn from the wall. The animal was then chained elsewhere in the room. Next morning the same thing presented itself to our view, and once more a portion of the wall was damaged. At the same time we noticed that the dog's abdomen was dry and that the appearances of cutaneous
were considerably reduced. It was only now that we grasped the true meaning of the circumstances. prepared the animal a bed of after which the wall ceased to be damaged, and the flow of juice no sand,
longer gave trouble. (Dr. Kuwschinski and I) acknowledged with gratitude that the common sense of the animal had helped us as well as It would be a pity if this fact were lost concerning the psychology itself.
thus overcame another
was not yet attained. Three to four weeks after the operation, the animals, previously well to all appearance, became suddenly ill. Food was almost at oncn This condition refused and a rapidly increasing debility supervened. was accompanied, as a rule, by convulsive symptoms, at times even by violent general cramps, followed, after two or thiee days, by
Obviously here we had a peculiar form of ailment.
of the question, for the animals often died with almost
any form of post-operative
.METHODS: PANCREATIC FJSTUL.K.
peritonitis, hail also to be given up, since neither the conthe animals before death nor the appearances post nwrteiu
afforded ground for such belief. Finally, the possibility of any autointoxication from incomplete and abnormal products of digestion due to the loss of so much pancreatic juice, such as Dr. Agrikoljanski in his
has suggested, was also excluded. Tn the first place, before death, absolutely no symptoms of digestive
we have convinced
vomiting, nor diarrhnea, nor ourselves by means of
special experiments, in which the pancreatic duct was ligatured and There remained divided, that this operation is absolutely harmless.
only one supposition,
creatic juice, lost
that the animals, in the escape of the pansomething essential to the normal processes of life.
Starting with this idea,
we adopted two measures to guard against the had previously known that the nature of the food powerful influence on the composition and quantity of the
pancreatic juice. (Dr. Wassiliew) therefore omitted flesh altogether from the dietary of these dogs, and fed them exclusively on bread and milk. Bearing also in mind the fact that a large quantity of alkali is
the pancreatic juice from the body, we regularly added a certain quantity of sodium bicarbonate to the dietary (Dr. Jablonski). By paying attention to these two rules it is tolerably easy to mainlost in
or even years, in a
for experiment without the necessity of
adopting any other special encountered in the management of
animals naturally vary, but in every four or five dogs one will generally be found which tolerates the operation without any In what way the sodium bicarbonate helps is not yet clear. nursing.
Possibly its administration in the blood, or possibly
makes good an injurious
diminishing the secretion of the juice. of the substance whose loss is so harmful to the organism would
pointed out, by In the latter case the nature
see, then, of
what great importance
have we not here a new pathological condition, capable of being by experimental procedure ? Dr. Jablonski has undertaken
but as yet
in the laboratory the investigation of this matter,
been completed. To return to the subject
the juice is collected by means of a glass or, better, a metallic funnel, so fastened to the abdomen with an elastic band
or thin elastic tube brought round the body that its wide end receives Hooks are fastened to the neck of the orifice of the pancreatic duct.
the funnel, from which a graduated cylinder hangs, the animal being These arrangements are very convenient for fastened in its frame.
We may pass over the old and admittedly inadequate experiments. there entering the intestine. its secretion may not be a considerable quantity still The evolution of a method for obtaining gastric juice and observing was no less difficult and protracted. or. however.8 THE WORK OF THE DIGESTIVE GLANDS. must not. however. A more recent method which has been published by the Italian inves* approximates to a perfectly faultless one. I an apparently easy problem may become ideal one. however. that this loss is rendered less injurious by a "VVe depreciation in the value of that which falls useless to the floor. 18%. or by closing the outer end of the tube. I have intentionally described all these accidents which may arise in connection with the formation of a wished to show how difficult permanent pancreatic fistula. In this way not only would much pancreatic juice be saved to the organism. xvi. When used in the laboratory. when dealing with material of such a peculiar nature. but the possibility of other serious disturbances of the digestive glands from the effects of the fistula would also be excluded. be diverted into the alimentary This experiment possesses. and consider more the making of a carefully the starting-point of the method now in use * Moleschott's Unterswc/tivngen :itr Xttturlclirc dcr Mt'iiscJi-eii >u>d der T'icrc. better to collect the juice from the dogs in the lying posture. One may justly assume that the continued loss of so important a secretion as the pancreatic juice is compensated. which we obtained by this method. outflow from the tube. the observer. He succeeded tigator Fodera in causing a T-shaped metallic cannula to heal into the duct. on the one hand. . could be either collected on the outside. notwithstanding the important defect canal. so that the juice. . Nevertheless. overrate the importance of these somewhat far- Further investigations showed us fetched suppositions. as one must accept it. on the other. but less comfortable for the animal. an we have no guarantee that. for the time being. especially it is if one makes first its position more comfortable by supporting the head. consistent. by an augmented or otherwise altered activity of the remaining digestive glands. Our solution of the problem is evidently by no means an It would be in the highest degree desirable to possess a method which would permit us to collect the juice when desired during the experi- ment and yet allow it to return to the intestine during the intervals. Bil. the animal soon learns to sleep excellently. In such cases it is necessary to employ a suitable vessel pressed more or less firmly to the body-wall beneath the opening of the duct. even under these circumstances. and instructive are the numerous results how clear. for the dog quickly tires and becomes restless.
j Trait6 Analytique de la IHiji-xtion. fected to enable fundamental questions to be In the year 1889. loud in denunciation of the gastric fistula it had justified none of the . In feeding. independently to the French physician. Professor Bassow. Why had not many important observations been made earlier with the help of the gastric fistula ? It required only to be per- by a slight modification. They therefore made an opening through the dog's abdominal wall into the stomach. since only very could be collected from the fistula. the following interesting experithe food drops out again from From the perfectly upper segment of the divided oesophagus. xvi. But. caused both incision. its We divided ends to heal separately into an angle of the skin thereby accomplished the complete anatomical separation mouth and stomach.* and in the year 184:5. and fastened into it a metal tube. since it afforded. little employ an extract made from the and very impure gastric juice It was likewise very difficult to obtain any idea of the rate of flow during digestion. dc Moxrtiu.METHODS: (JASTRIC FISTULA. a permanent opening remained in the abdominal wall which led directly into the stomach. hopes. closed exteriorly by a cork stopper. The tube healed firmly into the opening. when and free access to the cavity of the stomach. or of the properties of the secretion under different conditions. of the cavities of the Witli such animals one can ment. we divided the gullet in the neck. After his recovery.<t'ui (I. solved by its aid. and . Blondlot. and had proved quite valueless.\titnr. and could remain for many years in position without causing the least harm to the animal. as time went all on.f to reproduce artificially in animals a similar condition to that observed by an American physician whose patient suffered from the effects of a gunshot wound. . rfi'x . In the year 1X42. If the make the dog be given flesh to eat : ]l/iU. Dogs so operated upon recover perfectly with careful nursing. gastric fistula. 18-13. This condemnation was naturally exaggerated and was mainly due to the vexatiously slow progress made in our knowledge of the phenomena of secretion in the alimentary canal and especially in the stomach.- In SKI'. desired. and live many years in the best of health. Voices were. and for the purpose of studying the action of the ferment of the gastric juice nearly investigators were obliged to mucous membrane. the idea occurred to our countryman. easy This method raised great hopes at first. T. their food must naturally be brought directly into the stomach. the expectations gave place to disappointment. therefore. we (myself and Madam Schumow-Sirnanow>ki) performed the operation of ocsophagotomy on a dog already possessing a gastric fistula that is to say.
with solutions of commercial pepsin. in no way different from that of is by no means unpleasant . which. I leave the the gastric juice flows under such why to be attributed to the and what importance for the whole question of digestion is phenomenon. One can easily obtain in this way hundreds till matter open conditions. as it appears to me. since it is often considered desirable by the physician. soon commences which continues as long as the animal eats. attention of the pharmaceutist. You can collect on any day or every day from a dog thus operated upon a couple of hundred cubic centimetres of that is to say. an active secretion of gastric juice. it is. animal. indeed in many cases essential. you without any apparent injury to its health a dog almost as one obtains milk from can procure gastric juice from . and with natural gastric juice as obtained from our dogs. has dog by means of a fistula made on the principle been recommended in foreign lands as a Is it to therapeutic agent in various affections of the digestive canal. has not been more used in Russia. juice. indeed. Many experiments in the laboratory upon ourselves bear testimony to its easy The taste of the juice toleration and to the absence of any ill effects. previously washed out with water. although I have frequently drawn the attention of my medical colleagues to it. by Dr. Fremont matter is from the stomach of the of Thiry's intestinal one. and I cannot but express my regret that this substance. however. be the same in our case. To do away with the prejudice one might even procure gastric juice from animals flesh is eaten by mankind. collected by Dr. a cow. showed that the former was incomparably inferior. at all events deserves a trial. Last year pure gastric juice. The wish to try my fortune once more in this whose the cause of this deviation from the description of our procedures. merely remarking for the present that this method has definitely settled the problem of obtaining pure gastric juice. a solution of hydrochloric acid of corresponding strength.10 THE WORK OF THE DIGESTIVE GLANDS. and even fora short time longer. that a product long known tons would have met with greater success had it appeared under a foreign flag ? I now come back to our methods. must also claim the the finest product. to prescribe pepsin and Exact comparative experiments made hydrochloric acid to patients. The dog is converted into an inexhaustible manufactory of This fact. Konowalow. The problem of how to obtain . is The possible objection that the gastric juice procured from a dog can hardly count as a serious obstacle to its employment and distribution as a pharmaceutical preparation. empty stomach. of For pepsin experiments we need no longer prepare an infusion since enormous quantities of the purest pepsin can now be obtained with much greater ease and rapidity from the living mucous membrane. the next lecture of cubic centimetres of gastric juice.
1S7. out of which he formed a pouch which poured its secretion externally.K. In Heidenhain's operation this was obviously not the case. since its glands are microscopic and are embedded in the walls which surround the food receptacle. A tri- angular the stomach. but it does not afl'ord us tlie of observing the secretion of the juice and of studying its pro- perties during digestion. two centimetres from its junction with the pyloric end. llej also isolated a portion of the cardiac end of the stomach. Chigin) have modified Heiden- The first incision. flap is thxis A formed. the apex of which lies in the long axis of second incision is made exactly at the base of this flap. Bd. xviii. When the food way reached the stomach. wise formed by microscopic glands embedded in the intestinal wall and to study it in the act of formation. formed this into a cul-de-sac. l!d.f however. But. in the case of the pancreas the alimentary cavity with (where the gland duct is separate from its food contents) becomes a task of the greatest difficulty in the case of the stomach. in the ordinary In this way the above requirements were fulfilled. for overcoming difficulties of this kind was hit upon In order to procure succus entericus a secretion like- The idea was taken advantage of by Klernensiewicz* in 1875 for the purpose of obtaining the secretion of the pyloric end of the stomach in pure condition.MKTHOPS: fJASTRKJ FISTUI. Obviously to accomplish this there must be the continuance of normal gastric digestion side by side with a quantiThat which was quite simple tative collection of perfectly pure juice.mie. which begins hain's operation in the following way. soon afterwards succeeded in keeping one alive. and divides both the and posterior walls. unfortunately. means 11 pure gastric juice has been already settled. a perfectly clear juice began to flow from the pouch. Pfl iiger's Arrliir. But to draw conclusions with complete the normal work of the organ during digestion it certainty concerning was necessary to retain the nervous connections of the isolated piece intact. xix. which was still in position. To overcome this disadvantage a further improvement of the method was therefore necessary. Sitwngsberichte der Wiener Alcade. he isolated a cylindrical piece of gut. in the fundus of the stomach. is carried in the longitudinal direction for ten to twelve antei'ior centimetres. and sewed its open end into the abdominal wound. . f Heidenhnin :: : J Jf>.~>. Heidenhain. A happy idea by Thiry. his dog lived only three days after the operation. making the transverse incision by which he resected the piece of stomach the branches of the vagus which course lengthwise along the since in wall of the cavity were cut through. With this view we (myself and Dr. and could easily be measured quantitatively.
THE WORK OF THE DIGESTIVE GLANDS.
but only through the mucous membrane, the muscular and peritoneal The margins of the mucous membrane all coats remaining intact. round these incisions are separated for a little way from the submucous on the side of the stomach for a width of one to one and a half tissue
on the side of the
raised edges of
are applied to Out of the piece which belongs to the flap a cupola is formed. Both the stomach and the margins of the flap are then closed by sutures along the edges of the first incision. septum is thus made between
flap for two to two and a half centimucous membrane belonging to the large each other for half their width and sewn
their respective cavities, consisting of
Plexus gastricu. Bnter/or vagi
Plexus gastricus posterior vagi
A.-B. Line of incision.
C. Flap for forming stomach pouch of Pawlow.
one, that of the cupola, being intact, the other stitched along the
was only by forming the mucous membrane of the flap into a cupola that we were able to retain a dog with a permanent and closed No sooner were the two layers of pouch for any length of time. mucous membrane sewn in the middle line than a communication formed after a shorter or longer time between the stomach and the The animal was then useless for our purpose. It is still cul-de-sac. To better to make cupolas out of the mucous membrane on both sides. in a few words, we separated an elongated piece describe the matter from the stomach, formed it into a cylinder, the orifice of which we sewed into the opening in the abdominal wall, and allowed the other end to remain connected with the stomach. The cavity of the pouch the stomach by a septum formed only is separated from that of
METHODS OF FORMINf STOMACH POUCH.
here illustrations of the operation borrowed from the work of Dr. Chigin (Figs. and L>). our addition to the operation of Heidenhain makes it Naturally more difficult, but as will be apparent farther on, we are compensated
as a reward of this increased difficulty
by an intact condition of the
It is clear that nervous relations of the stomach, which was our aim. the fibres of the vagus nerve reach the separated portion of the stomach,
since they course
between the serous and muscular layers
of the flap.
Cavity of stomach. 5'. Pawlow's pouch. A. A. Abdominal wall.
not followed by any serious discomfort, nor does the animal.
We have yet to discuss the question whether the activity of our miniature stomach furnishes a true representation of the secretory work of the large stomach. This is all the more necessary since the food
comes into contact with the walls of the latter during digestion, while O O the former remains empty. A full answer to this question I shall
reserve for a later lecture,
are in possession of
merely state in a few words that, in addition to rigorous inferences drawn from a series of unquestionable facts, there are numerous direct experiments in which the small and large stomachs were compared both as regards conditions of work and
THE WORK OF THE DIGESTIVE GLANDS.
for doubt that we properties of their secretions, and which leave no room with perfect safety employ the small stomach as a means of studying may
In our next lecture even the miniathe function of the normal organ. ture stomach will show itself to be an instructive object worthy of
already been related, Dr. Fremont has recently succeeded in isolating the whole stomach of (since the publication of our method) the dog after the principle of Thiry that is to say, the lower end of
the oesophagus was united to the duodenum and a cannula made to heal This procedure, as I into the stomach, previously closed at both ends.
special experipossesses two or three important defects. First, in ordinary digestion in such dogs we can hardly reckon upon normal conditions of secretion, since the
shall later explain, can, however, only serve for
a general method
mucous membrane can never be
by contact with
food be introduced directly into the stomach, it secondly, mixes with the gastric juice. Finally, as regards the collection of juice
if it appears to us that our combination of the ordinary fistula with cesophagotomy possesses important advantages Our method is incomparably simpler, over Dr. Fremont's procedure. and, under suitable conditions of operation, is not attended by any
for practical purposes,
useless sacrifice of animals
the dogs live for years in the enjoyment of
Can this be said of Dr. Fremont's dogs ? The usual method of obtaining the juice from the miniature
is as follows. small indiarubber or glass tube, freely perThe tube either forated at its deeper end, is led into the pouch. remains in of itself or is fixed in by means of an elastic band brought
round the animal's body.
collected either in
the lying or
standing This method of forming a miniature stomach, so far as I can for the moment imagine, must be regarded as the only one possible which is at
It possesses a few small disadvanthe same time correct in principle. tages, it is true, but these are only matters of detail, such, for instance,
posture of the animal.
as the maceration of the edges of the wound and the loss of some gastric But these defects can easily be counteracted, and, moreover, are juice.
in themselves of trivial importance. They can, I hope, in time be avoided. in the interests of a thorough investigaIndeed, altogether
tion of the whole secretory work of the alimentary canal, a universal of simplification of the technique is to be desired, with a weeding-out
minor defects, so that it may be possible to make several same animal without endangering its life or health.
It is obvious, from the sketch of digestion now given, how important simultaneous and rigidly concordant investigations of the
SUKGICAL METHODS IN PHYSIOLOGY.
several glands, both with regard to periods of activity
and quantitative This can only be achieved, however, relationships of work should be. when the activity of all or many glands is simultaneously observed on one and the same animal.
In bringing the description of methods in this lecture to a close, I it essential to point out the importance to physiology of
It appears to me that the methods of surgery, as surgical technique. contrasted with those of vivisection, must obtain unquestioned recogni-
tion in the series of procedures
the conception and carrying out
which we adopted 1 mean in the perof more or less complicated
operations having for their object either the disconnection of certain the ready observation of deeply seated processes in the organism, the severance of existing relationships between organs, or
establishment of new ones, etc. With these must go hand hand the means of healing the injury inseparable from the operation, and of restoring the animal to its normal condition so far as the nature
rice versa, the
of the procedure permits.
Such a discussion of operative methods appears to me necessary, chiefly because it becomes more evident every day that, in the " acute " ordinary method of the so-called experiment, carried out at
and complicated by free bleeding, many sources of error The crude damage done to the integrity of the organism sets up a number of inhibitory influences which react upon the functions of its different parts. The body as a whole, in which an enormous number of different organs are linked together in the most delicate fashion for the performance of a common and purposive work, cannot in the nature of things remain indifferent to forces calculated to destroy it. It must, in its own interests, restrain some functions while others are allowed free course, and thus, by appropriately economising its energies, rescue that which is possible to save. This circumstance was formerly, and still is, a great hindrance to the
developments of synthetic where it is of value to determine the real course of this physiology, or that phenomenon on the uninjured and normal organism, it continues to be an unavoidable obstacle. Operative discovery, as a means of physiological research, has by no means been played out. On
the contrary, it is only just coming into full activity, as is testified by the achievements of the present day. For example, we need only mention
efforts of analytical physiology, while, in the
the extirpation of the pancreas by Minkowski the transference of the portal blood into the vena cava by Eck and, finally, the amazing operations of Goltz, in which he removed bit by tit the various parts of the central nervous system. Have not many physiological questions been thereby settled, and do not innumerable others arise from the
There can be little doubt that even single operations in the general rooms of the laboratory. performed with the aid of anti. are altogether rare. which consists in the establishment of a communication between the portal of a large surgical vein and the inferior vena cava. number of physical instruments which are yearly invented and introduced for the investigation of physiological phenomena. Take. Petersburg Institute for Experimental Medicine. still impure that the Eck's operation. and vivisecting departments. The physiological institute was and therefore. physical. misfortune attended Dr. and consequently in the surgical sense clean. The physiologists do not regard such problems as essential. microscopic. but of time. It may be objected that I am warmly results already obtained ? Yes. in the operative section of the physiological department of the only St. cleanliness during and immediately after their performance from want department expressly fitted out for the purpose. though carried out by the more experienced hands. for instance. a-septic precautions do not succeed. notwithstanding the endeavours . none set of rooms. the well-known history of the Eck's Fistula. This happy period lasted. well-equipped surgical The general rooms of a laboratory cannot be used for carrying out frequent and complicated operations with safety to the life of the animal afterwards. could not succeed in keeping energy The same for any length of time after the operation. who repeated the operation with It was the assistance of Dr. that any considerable prohowportion of successful cases was attained. and acumen. degenerated into a fruitless waste all This continued for a year.for what is already recognised. the surgical precepts of the age must not be . only for a year. as well as the number of chemical methods and their variations. while provision is made for chemical. then just founded. at that time small. The clearest testimony in proof of the fact that surgical methods have not assumed their legitimate position in physiology is evidenced by the fact that in the buildings for a physiological laboratory of the present day. it is remarkable that many of these operations are introduced by surgeons and not by physiologists. for example. Stolnikow. and only carried out by the few. without the sacrifice of much time and labour moreover. If.10 THE WORK OF THE DIGESTIVE GLANDS. the paucity of the latter stands out in marked contrast to the richness of the former. Again. is made for an efficient. in spite of the employment of every precaution. became so rapidly same. or perhaps because it is almost impossible to maintain a sufficient degree of animal neglected. ever. Eck. but such operations contending. sparing neither trouble nor animals. or perhaps are not in possession of the means necessary for their solution.or are not attempted. notwithstanding his the animals alive In the old laboratories its inventor. be compared with the the number of new physiological operations which permit of the survival of the animal.
SURGICAL METHODS JX PHYSIOU ><. o .- 1J il 3 'C .0 . 31 I 3 Df * -A | f S 0-11 . Y.
space was allotted to I take the liberty of drawing your attention to this. which. and cloths. occupies about four and a half square metres of floor space. in which the animals are kept for the first ten days after the operation. It is only by the arrangethe assistance of this intelligent animal. A passage runs in front of hot air and furnished with electric lights. The long series of operation. till a new physiological department. The of the washed in a bath and dried in a special drying place. to count on . The whole is painted with a white oil colour. In the dog cabinets a water-pipe. when striving to attain surgical results. It consists of a series of rooms along one side.rooms constitutes the best protection against the penetration of dirt into the last and most important room for. so far as I know. in which the preparations for the operation as well as the operaIn the first room (Fig. in the ordinary as well as the surgical meaning of the word. is laboratory devoted physiological institutes. judged by the application of our When I call to mind the the success of Eck's operation. of the operators. The laboratory attendant is not allowed to go beyond the second room of the series. ment of a long series of dirt-catchers. Each cabinet is provided with a large window and ventilating arrangements. In the next it is nar- cotised. although physiology is much indebted to the wisdom of the dog. it would be in vain. for their overalls. was added to the building. it Two buildings. instance of a special section in a physiological Perhaps the example may give my some useful hints for the erection of new colleagues the first to operations. Into this room the narcotised and previously prepared animal is carried (without a table) by the operators themselves. and the donning of The fourth and last is a well-lighted operating-room. oft' from the operating-rooms by massive tightlyThe in floors of with gutters each room.18 DIGESTIVE GLANDS. shut fitting doors. Each is also heated with these dog cubicles. the site of the operation shaved and cleansed with an antiseptic The third is used for the sterilization of the instruments solution. and is more than three and a half metres high. twenty years in different and always upon equally healthy material. a quarter surgical section embraces the half of the upper storey whole laboratory buildings. Separated by a partition-wall from these rooms. the washing of the operators' hands. is a series of cabinets. 2 a) the animal is tion itself are carried out. the department are all made of cement. with constant . runs along the wall. with numerous minute apertures. that one can count on maintaining the operative division at ment have not rendered surgical test results of operations carried out during the last years of work in this departimpure. by means of which the floor can be copiously syringed from the corridor without entering the room. its optimum. in which greater the operative section.
be convinced of their truth.SURGICAL METHODS IN PHYSIOLOGY. It has preserved numerous animals alive and spared our operating staff both time and trouble. attends cleanliness. to reflect on these closing words of my present lecture I beg of you will then. can already be traced by the aid of the present methods. the development of our ingenuity and skill in performing only by chemical work operations en the alimentary canal that the exquisite effected by it will be revealed to us. possibly in even a more striking way success which than the surgeon. the outlines of which I . you am persuaded. . I hope you will pardon this long digression concerning the importI am convinced that it is ance of surgical methods in physiology. 1!) I am convinced of the magnificent repetition of the same operations.
and in the digestive capabilities of differences milk Meaning of these. Having considered the means by which the work of the we may now digestive glands may be more or less perfectly observed. begins to furnish juice within a few minutes after the animal has taken food. * The beginning of gland secretion is connected with the entry of food into the alimentary canal The quantity of juice is proportional to the amount The curve of secretion its importance and exact regularityof food . Thus. Every one of our experiments on dogs gives. is peifectly empty. an unequiThe isolated miniature stomach which. THE WORK OF THE GLANDS DURING DIGESTION. examples Methods <>!' Qualitative changes in the juice during secretion investigating the properties of the juices The gastric juice possesses a constant acidity Meaning of the qualitative variations of the juices . moreover. The juice is only poured into the alimentary canal when the raw material. and.LECTURE II. thanks to the methods now at our command. GENTLEMEN. not however without trouble and difficulty. hardly any physiologists doubt that the activity of the glar. with the same diets The work of the digestive glands under the prowith diets of flesh. the first and gastric fistula and the most elementary facts concerning the activity of the digestive glands were established. The course of secretion and properties of pancreatic juice. in vocal and positive result. in this respect. : It is. . the elaboration of which This is an apparently is demanded. is increased to many times that amount This is a fact which has long been suspected. which is only two to three cubic centimetres per hour. Differences in the rate of secretion gastric juice. fasting animals. it was recognised by all authors that the glands only first began to secrete when the food entered the alimentary canal. By the aid of the older methods (the ordinary earlier forms of pancreatic fistula).ds is strictly dependent upon the taking of food. Similarly with dogs having pancreatic fistula? the quantity of juice. turn to the work itself. after the entry of food. makes its appearance therein. with the requirements of the case. consistent but has only now been fully established. bread and lunged influence of different dietaries.
milk doiilile tin. it was scarcely possible to solve this question.c. regard this result as extremely instructive. We different quantities of the Thus.. How as the food remains in that particular and does segment of the alimentary canal. we are in concluding gastric glands work with great precision. It appears..Ml 42'0 ) c..c. consisting of meat. upon this point. . does the secretion continue so long: .simple fact. food. flesh. give the impression of much . without and precision of the work of the digestive does the work of And now we proceed to other questions Is the requisite quantity of juice poured out once secretion proceed ? and for all on the ingested food or. For a mixed diet.c. IGG'O c. vary regularly with the decreasing quantity and altering properties of the mass ? These questions long ago gave origin to a multitude of investigations from which it appears that the secretion of juice is continuous thixnighit out the whole period of digestion but with a varying rate of progress. the following food taken. Chigin gives the following mean For 100 grms. figures were obtained : . The juice could neither be separated from the food nor its quantity correctly deterAt present we are in command of perfectly accurate data mined.c. by means of the simple fistula alone. to the great accuracy canal. of juice escape. so far as the stomach two was concerned. that there exists an almost exact proportional relationship between the quantity of juice secreted and the amount of isolated stomach. 4(1-0 c. The older methods were quite unable to give an answer to such a question even as the following: Hew does the quantity of juice alter with varying amounts of the same diet ? In other words. values: .ACTIVITY DURING DIGESTION. or do those factors hold a different relationship to each other ? Asa matter of fact. however.c. The problem is easily solved it on the dog with the same simply give food and collect the corresponding quantities of pure juice. SH'2 c. for -MM) for 400 grms. is obvious that it involves a multitude of subtle problems connected with the activity of the glands. i>1 but it. for raw meat. grins. .1(1 I ^TIIIS. :. 2(>-U c. they pour out amount of juice. from these figures that the inasmuch as for varying an exactly proportional quantities of diet administered. Dr. The data in question do not. from these investigations. bread and milk. determined in the first instance by the quality of the justified Hence. I doubt. is the quantity of juice directly proportional to the amount of food taken.c. it points.nO c. meat lin-ad \ With With :i diet of V .aliove i|ii.mtities. of juice were secreted.
As guarantee for what watching the glands under a I have said.MEAL OK 100 GUMS. unknown and under varying conditions quantities.22 THE WORK OF THE DIGESTIVE GLANDS. who did not always endeavour to give the requisite degree of exactness to their researches. partly in the investigators themselves. applied has now become an absolutely constant one. so to speak.S AFTER A . I give here of of two experiments on the gastric glands. Walther : WORK OF THE GASTRIC OLANI. food was often administered in definite composition. The cause is conformity. and likewise two on the pancreas taken from that Dr. almost physical exactness of complex physiological processes imparts a feeling of satisfaction to the experimenter which rewards him for his many hours of perseverance in condition of activity. taken from the work Dr. of inof appetite or the reverse. Thus. . KLESH. a minute degree of exactitude upon the experimental arrangements. Chigin. As a matter of fact. we have bestowed from the first. In our researches. to be sought for partly in the defects of the methods. the course of the secretion when the same conditions nre This. to cjmpare accurately the work of excretion under different conditions.
are not easy of interpretation in all The their separate fe?. nor even in regularly diminishing quantities. the secretion of juice. or gently or suddenly falls. this is almost impossible.tures. Curve of secretion of of flesh. time in work are determined by fixed conditions. line of descent with its fluctuations can more or less satisfactorily be explained on the principle of corresponding variations in the quantity of the of ingesta at any particular part of the digestive canal. indeed. as the case may be. as mentioned in our first lecture. viz. we must admit that this or that rate of secretion is not determined by mere blind chance. The curve is by no means a straight line gradually approximating the It is a special curve which slowly or rapidly ascends. serves for a time a uniform height. only be furnished by physiology when. can How. a definite periodic law. Examples of these will be given later. for example.. pancreatic juice ? A scientific exposition of the curves. Since these curves repeat themselves under the same conditions with stereotyped exactitude.ACTIVITY DURING DIGESTION. that even the variations which occur from time. .) law. or preabscissa. but in Hours all cases follows a necessary 1 Via. in the curve of explained. one which fully and accurately corresponds with the actual facts of the case. The curves. is 23 to to say. The fluid is not poured out secretory follows therefore at the same rate from the beginning to the end of digestion. But the meaning complex line of ascent remains in many cases obscure and inex- plicable. after having attained an initial maximum. can the late appearance of the maximum be which we see during the third hour after a meal. that is. requisite for the due elaboration of the food and therefore beneficial to the organism. ?>. after n men I (Two experiments. for the present. ir:i>trie juice. The work of the glands. however.
Food. are they not also able to extend similar variations to the properties of the secretion ? Judging from a theoretical standpoint. or a varying degree of acidity or position. or a different content of ferment in the fluid. 000 of the digestion of the The total necessary. cal Curve of secretion of pancreatic juice. to the quantity of juice which they produce. experiments performed in we could render no account of these tiue that science has long since answered our question . a juice of varying properties of food can chemiunder the influence of the first portions of respects. both and physical quired. may be rein same mass would be have so altered. with regard seen. the changes both quantitative and qualitative. that it may need for its further digestion a juice of different comThus. alkalinity. without importance. more or less water. cc. throughout the now pass on to a further question. If the glands. 14 10 FK.24 THE WORK OF THE DIGESTIVE GLANDS. as we have are in a position to vary their work so remarkably. are followed.. step by whole alimentary canal. (Two experiments. It is All these separate conditions of juice activity are naturally not But so long as we dealt only with digestive vitro.) food. matters. which the food and admixed secretion undergo. one would expect that in different phases in We IV in IV it. 4. juice. its . milk.step.
it will be found that the greatest interest is attached to variations in the ferment content. viz. Nor were all the details published in merely appeared in Hermann's Encyclopaedic Handbook form. Nor can Heidenpure juice. not with permit but with a mixture of juice and food. Experiments with the ordinary gastric fistula they deal. I must draw your attention for a short period to the methods we employed The proteolytic power of the in our study of the digestive juices. though. This method leaves nothing to be desired so far as convenience of applicaTest experition. although.. strictly speaking. and requires no further watching. in the affirmative.ACTIVITY DURING DIGESTION. fluid egg-white is sucked up into a fine glass tube of 1 to 2 mm. The proteid cylinders in millimetres difference gives the length of the digested and fractions of a millimetre. ments specially carried out (Dr. the length of the pieces of tube. But before I take up the question of our own results. and The tube is coagulated therein at a definite temperature (!>5 C. the food was not of definite composition. the remaining properties of the digestive juices demand an equally careful investigation to arrive at a satisfactory explanation. Ssamojloft) have convinced us that the . unfortunately. The investigations of Heidenhain on the pancreatic secretion in dogs operated upon by his method. it do alter during appears to me. and that of the undigested remains of the proteid columns. then cut into small pieces. Solution of the proteid occurs at the ends of the small glass tubes. which are placed in one or two The whole is kept in cubic centimetres of the fluid to be investigated. in this . has not been appreciated to its full extent. a procedure worked out in iluid was determined by the process of Mett It consists this laboratory and since then constantly retained in use. But the observation. especially as regards gastric juice.. otherwise it would have become an inexhaustible source of persevering inquiry as to why and how these Later I shall adduce instances from our storevariations come to pass. lumen. and clearness with exactness of results is concerned. hain's observations on the isolated fundus be taken as an index of only very hypothetical deductions. since of normal digestive work. after division of its secretory nerves. the thermostat at 37 C. could not be considered sufficient. After the termination of a certain period. must however be recognised as scientifically exact. that the properties of the juice 25 the period of secretion. The available material. are measured oft' with the aid of a millimetre scale and a microscope of low magnifying power. to 38 C. obviously difters greatly from the normal. Naturally.). they condensed . house of observations dealing with these highly interesting qualitative variations of the juice during the separate periods of digestion. since the activity of the isolated stomach.
by 4 and 9.. is the rule which expresses it.. the relationship which lies at the basis of the connection between the length of the digested column of egg-white and the came out with perfect pepsin contents of the fluid under investigation. Dr. These experiments did away upon It may therefore be accepted that the length in mnis. is proportional to the square of the rapidity of digestion to the square of the millimetres) which the juices are capable of An example in figures will period of time. gives us an exact relaIn the researches of tive measure of the digestive power of the fluid. This was the case even when the fluid possessed its highest digesting power. This correspondence in the data furnished by different methods affords I must here express strong assurance of the correctness of the rule. and the genera. for then all observations on the juices of different animals and men would be represented upon a uniform scale. we should have in the second fluid one and a of 8 more ferment.. has not been as widely employed as it in reality deserves. but by the squares of expressed by these numbers i. is without import- . the quantity The following clearness. Moreover. the first ten hours digestion of the proteid columns. viz. of pepsin in the compared fluids i. even within wide limits.e. .We have still to add that the diameter of the tube. which would lead to important deductions concerning variations of ferment in the individual. was published so long ago as regret that the method of Mett. been deduced from comparisons made with half times solutions of pepsin exactly prepared.. with the very natural mistrust that the solution of the proteid may proceed at varying rates at different depths of the tubes. dependent more or less stagnation of the digestive products within the lumen. which the year 1889. carried out upon this question in the laboratory of fessor Tarchanoff. but according to our rule of squares. of course. According to linear measurement.20 THE WORK OF THE DIGESTIVE C! LANDS. The difference is instructive. of egg-white dissolved by the several juices in a certain time. and the other a column mm. column (expressed in digesting in the same make this clear.e. had already the result which been discovered before him. the relative quantity of pepsin in each is not the figures 2 and 3 respectively. Borrisow arrived at independently. of proteid. by Schiitz. so that investigations upon will deny that this is desirable in comparable with each other ? No one my a high degree. ProBorrisow. from polarimetric estimation artificial numerous of the amounts of peptone formed during the digestion of egg-white. If one of the fluids digests a column of 2 mm. This rule has. at least within deal with) is directly proportional (employing the fluids which we have to to the length of the period. the species. the second fluid is two and a quarter times stronger than the first. How easily could it be made a universal means of comparing proteid these ferments would be digesting ferments.
the length of the digested column was measured and exFrom numerous experiments with artificial pressed in millimetres. and in our experiments have been. varies column of digested starch with the square of the length of the measured in millimetres. of Fehling. Arc. ance. and also in that of the number of millimetres of starch-column dissolved. the acidity of the . and was. Glinski and Walther) to estimate the activity of the proteolytic and amylolytic ferments of the pancreas by analogous methods. after it had stood for a certain time (with periodic shaking) at a given temperature. served measure of the amylolytic activity. But naturally our failure will not prevent us from endeavouring to finally to set and we had to solution. and still are. determined in the laboratory by titrimetric estimation. but necessitated a great expenditure of time. 27 and that the white of the hen's egg is of sufficiently constant com- position to be . and recently an attempt was made by the laboratory (I)rs.) the relationship between the quantity of ferment and the length of the column of starch has been and Borrisow is valid here also to its dissolved established. The activity of the amylolytic ferment will therefore be expressed in both ways in tinexperiments given below. The number of milligrammes of sugar produced. remained fruitless. the action. Thin glass tubes were filled with coloured starch paste and then exposed in the thermostat to the action of the ferment for a certain period.. The paste was dissolved from the ends inwards. frequently The activity of the amylolytic ferment of the pancreas was modified. viz. and determine by titration with baryta permanent emulsion formed by fat and pancreatic juice. therefore. action of applies also in its full extent to the tiypsin. The methods of comparing the working of other ferments are less perfect. The number of cubic centimetres of baryta solution which were necessary for the neutralisation of the fat acids. thanks to the colouration. As in peptic digestion. being clearly limit of ferment- visible. full The law i. which is formed from a given of starch paste exposed under certain conditions to the activity quantity for a long time after the method of the ferment. solutions of ferment (pancreatic juice diluted twice. of Schiitz extent the content of ferment in the fluids. thrice.e.Schiitz employed as a and Borrisow The law of test object for the purpose. not wholly satisfactory in a research where numerous as a estimations were required. served as measure of the activity of the fat-splitting ferment. This method furnished good and reliable results.ACTIVITY DURING DIGESTION. up to the present. of the sugar. Unfortunately all attempts to base an evaluation of the fat-splitting ferment upon the same system have. in terms of the milligrammes of sugar formed. Hence a more rapid one was sought for.. usually for half an hour.
or even at still shorter intervals. But this is unavoidable when the properties of the juice have to be followed from hour to hour. and is therefore very tioublesome where numerous estimations have to be made. however. To this it must be added that the results of the method are not always equally reliable.THE WORK OF THE DIGESTIVE GLANDS. Naturally our experiments deal in general only with comparisons of ferment activity. The law of Schiitz and Borrisow was. the method we are compelled to use demands the continuous attention of the experimenter. As matters now stand. and our deductions concerning "quantities" and "total amounts " of ferment must therefore be accepted conditionally.N* o of Column -~ O Mm O . confirmed here also. obtain a method uniform with those applied to the other ferments. In X o c [jj O: O Ln "o -f* o -^ O Proteid .
Experiments of 15th and Hour.i"> (taken 1'nnn the work of I >r.ACTIVITY DURING DIGESTION. L>9 HOURLY VARIATIONS IX IMitKSTIVK 1'OWEK <>F CASTRIC JUK'E AFTER A MEAL OF Ion GRMS. Millimetres of egg-white column <li . OF RAW FLESH. Lolia>sol'l). Itltli May l*'.
the more rapid the latter. This rela- . during given periods of their secretory work. Ferment content milk. that the possesses a conIt is true that stant acidity. which possess a complex chemical activity. fi. that is to say. now one product and now another. and even in our observations. a strong digestive power may recur both with a as well as with a scanty secretion. juice in independently of each other. appears the fact. That which has been said of theferments may also be applied to the quantities of salts present in the juices. therefore. receive the juice directly from the glands. fluctuations are But a all careful investigation leads to the almost indubitable of the data conclusion it is that the juice. . where we deal with absolutely pure juice. always possesses the same degree We do not. as one must accept gastric juice it. All the more interesting.THE WORK OF THE DIGESTIVE GLANDS. In one and the same copious the different ferments may suffer variations running courses juice. as is clearly shown by attributed the apparent numerous observations. diverse relations between the content of water and the richness of the ferments. To this circumstance must be fluctuations of acidity. a HOURS i n in rv if in IV fact which undoubtedly shows that glands such as the pancreas. the more acid the juice. such also to be seen. It is acidity of the juice is closely dependent a rule almost without exception that the upon the rate of secretion . has its acidity reduced.are able to furnish.c. in hourly portions of pancreatic jnico after a c. even method. as FlG. and vice versa. howof acidity. clinical investigations of the secretory activity of the human stomach are almost daily con- cerned with variations of acidity. ever. meal of 600 poured out by the glands. alkaline in our After it is secreted by these it has to flow down over the mucous membrane and inevitably becomes partially neutralised.
prove to exceed the normal.. In the normal stomach a perfectly pure juice may have its acidity reduced to the extent of twenty-five per cent. in nearly all cases. During one five minutes the fistula remained open.ACIDITY OF (J AST LUC JUICE. the greater will During the decline of the secretion we find an absence the low acidity pertaining to a corresponding rate of outflow at the beginning of the experiment. and if the stomach has been washed. by neutralisation with mucus. however. You have seen striking instances of the fact that the juice furnished by may happen to come about. so to speak. the juice acid whether it be rapidly or slowly poured out.) If all this be correct. The greater the quantity of juice the more rapidly will it flow over the The stomach-wall. all the more easily will they arise in a stomach to which food mixed with saliva can Moreover. :!1 tionship is easily to be understood in the light of our explanation. It is. to vary the That it does fluctuate remains a fact. is On equally and strongly the other hand. that the portions of juice obtained in the second way i. and therefore the less will it become neutralised. Obviously this is because the stream of juice has been neutralised by the mucus. possible that the neutralisation of the gastric juice must be looked upon as a purposive and desirable event with a definite aim. possessed a lower acidity than the others. during another it was kept closed. a short time ago we made observations in the laboratory upon a dog suffering from strongly marked hyperacidity of But in no single sample of the juice did the acidity pathological origin. it is quite natural that the first poi'tions of juice secreted. of acidity can occur in this way with pure gastric juice. Dr. acidity observed mate tion. The more be of its acidity. after a delay of five minutes in the And if fluctuations stomach. to that which under these conditions will thus more closely approxiis real and authentic. gain access. for example under the influence of sham feeding. Since is usually covered with a considerable layer mucus. not unfrequently all connection between rate of secretion and degree of acidity can be removed. Ketscher. acidity precisely in this way ! however We discourse once more. Who knows. Ketscher has collected the juice in the following way during the course of the same sham-feeding experiment for periods of five minutes duration each. freely and rapidly the juice flows. perhaps nature has found it serviceable in the interests of the organism.e. In order to test this explana- the wall of the stomach of experiments of various kinds were instituted by Dr. may now take up the threads of our it . in this manner several times in succession. the varying necessity for acid during the course of digestion is supplied by variations in the quantity of juice and not by changes in its acidity. will manifest the lowest acidity. It resulted. or the elaboration of the food. (1'awlow. That is to say. and the juice allowed to escape at one rush.
however. the fact that we are able to consider it is. The facts already communicated indicate that the glands are able to adapt themselves to the separate and successively occurring phases of the elaboration of the food. To this end chemical conditions must be provided in the intestine which help on the action of the pancreatic juice. a brilliant testimony to the essential service rendered by our newer methods. the pancreas and by the digestive glands during the course of the same act of digestion does not remain uniform. viz. The strongest juice is thus poured out when it is most needed. to determine in what way these variations are related to the progress of the act of digestion. Now. This means that the conditions under which gastric digestion has been accomplished must be radically changed. pi-o- We may commence with the stomach. and whether they are of service to it. is furnished chiefly for a particular kind of food. ning of digestion. and since different juices are poured out into the alimentary canal. however. I merely raise these questions now.82 THE WORK OF THE DIGESTIVE GLANDS. Since the food is made up of several con- stituents. with accentuation of certain of its properties. and that a high degree of acidity injuriously influences its activity. where a food material already modified and assorted by the stomach has to be further worked up. We know that trypsin is digested by pepsin. Is this in reality the case ? It is obvious that an answer to this question was impossible by the aid of the older methods. for example. Take. with special perties of the digestive juices. Here the matter concerns a later stage in the work of our factory. Some of the details. this is well timed. since they are injurious to the action of the pancreas.. of itself. disclose an obvious purpose. In connection therewith we may rightly suppose that this adaptability will only appear to its full extent when we compare the variations in the progress of secretion under different dietaries with each other. the first secreted portions of gastric juice they are distinguished from the of this : It is evident that at the beginothers by a stronger digestive power. complete A solution problem must be left to the future. In the case of the pancreatic juice that the alterations in its it is much more difficult to show composition are purposive. Their elucidation will be taken in hand after we have discussed the mechanism of excitation of the glands. the supposition appears natural that each fluid. that every individual kind of food calls forth a particular activity of the digestive glands. when the quantity of food is large and its external structure still coarse. Researches carried out by . but is varied in many It is in the highest degree both interesting and important respects. At present we can convince ourselves by actual experiment of what a priori only appeai-ed probable.
of milk now 7-4 given. is in this respect. and " milk juice by 11 (3-2G ). milk.c. is juice" " 2 1C (8-99-).M 4 received 200 c. bread or meat. 4-2 2-2 The influence of the different foods juice is striking. and also to its quantity. Digestive power 8-0 7-0 7-0 in mm. according to the law of Schiitz and Borrisow. 33 Dr. 3-75 3-30 An additional 400 5 P.M 10 11 1-8 Dog then 12 noon 1 given 200 gruis. 2 represented by 44 (G'G4 )... according to Dr. Chigiu At eight o'clock in the morning the dog was given 200 grms.. The speciality of the work applies not alone to the properties of the juice. Hourly quantity of juice 3-2 4-5 . but more correct. as well as the separate administration of milk.DIGESTIVE POWER OF GASTRIC JUICE.c.. bread to Time.. 8'0 8-8 5-37 3-50 3-75 P. diet of flesh calls forth a juice of 3-99 mm. will deal with these points in order. "flesh juice" by The matter may be illustrated : by the following protocols taken from experiments of Dr. C ."* proteolytic power.. is represented by G-G4 mm. raw meat. in c. and one of milk of :5-2G mm. terms of "juice secreted after the administration of meat and milk" respectively. but to the rate and duration of its secretion. 9-2 8-4 . Chigin.. In other words.. . "bread juice" contains four times as much ferment as "milk-juice.c. four times as concentrated. We greatest digestive power belongs to the juice poured out on Its mean bread. . which for shortness we may name "bread-juice.. * In harmony therewith \ve shall also speak of "meat-juice" and "milk-juice" instead of the longer. of eat.M 2 8-G Dog now 3 P. 2-25 . produce each time characteristic modifications of activity of the gastric glands. 8-9 A.M 6 c. . Chigin on dogs with isolated miniature stomachs have shown that both a mixed diet. upon the digestive power of the In order to exclude the possibility that the order have influenced the result I of administration could append another experiment. digestive power. &c. If we now turn to a comparison of these juices The A with one another we that "bread find." and.
30 received 200 c. and the duration of the secretion is correspondingly protracted.. of milk again given. 1-5 A. * itself with the The acid was estimated titrimetrically. The dog Time. greatest Not alone the with flesh (0-56 per cent... the mean hourly quantity of juice is obtained. each kind of food.) and lowest with bread - (0 4T> per cent. however.). however.M 200 c. The latter is. or its amount of dried substance. and milk the least gastric juice but taking equivalents of nitrogen. which represents the mean different for the different sorts of flesh hourly intensity of gland work.. Quantity of juice Digestive power in nun. or.34 THE WORK OF THE DIGESTIVE GLANDS. 10... reveals itself in the hourly rate of secretion and in the qualitative This time I furnish only one example for variations of the juice.80 . If the quantity of juice produced during a given digestion period be divided by the number of hours in the period.M 3-37 2-0 2. when estimating the food. . but far less. The hourly intensity of gland work is almost equal in the case of milk and flesh diets.30 P. . of bread were now given. in c.30 P. relationship is equally clear whether.. In a similar way the quantity of juice poured out and the duraAnd this tion of its secretion are dependent upon the kind of food...* varied according to the nature of the diet. 8:30-1). and beg you to believe that it repeats same admirable precision which we have already seen. limited to the distinctions given. Nor is this speciality of gland work. one takes into consideration its total quantity. on its Even this is number.c. 7'0 li'O 2-0 145 grms. and is expressed in percentages of HC1. The last. 11. requires the most. of milk.. food.M. exceeds all the others in the time required for its digestion. with bread.c..30 A. 2-0 3'6 . which depends on the variety of It likewise prominently the food.. bread needs the most and flesh the least. Comparing equivalent weights. 1.. 5-4 3-4 .M. . .30 digestive power. . 3'37 5-0 12. lastly. its content of nitrogen (since the gastric juice acts only proteid constituents). ..c< . but likewise the total acidity.
200 GEMS. : (According to mean values obtained by Dr. MILK. 600 C.) . FLESH. 35 QUANTITIES AND PROPERTIES OF GASTRIC JUICE WITH DIFFERENT DIETS 200 GEMS. BREAD.THE JUICES CORRESPOND TO THE DIET. Chigin.C.
I wished merely to bring home to you the . 7. Curves representing the rate of secretion of gastric juice with diets of flesh. COOc. Hours 123456781 2345678910123456 c <u \ \ Flesh. F IG. have burdened the attention and memory of my hearers to an undesirable extent.36 THE WOKK OF THE DIGESTIVE GLANDS. Bread. Hours CC 12345678234 5678923456 I i CD O) Proieid ofColumn -P Mm KO 000 O . 200 grms. '200 grins. Milk. connection. and milk.c. bread. But in cursorily introducing them I did not by any means intend that you were to retain all their complicated details in your minds this would naturally need frequent repetition and a thorough study of the circumstances.
and on GOO c. indeed. since he experimented with different quantities.c. demand is supplied less by an increase in the volume of the juice than by an extraordinary concentration of the fluid poured out. on bread. however. of flesh contain approximately the same amount of nitrogen as (500 c. digestive strength are secreted. different quantities of juice are called forth by the different varieties of food. however. These indicate that. figures I 1(!00 37 430. would be see. and that flesh These nitrogen requires 25 per cent. They may. have obtained MS follows: 100 grms. on proteid in the form of bread. be easily constructed from the data at hand. we obtain the above-mentioned numbers.VARIATIONS DEPENDENT UPON DIET. 1000. 34 c.c. of white bread. strength are secreted. more pepsin than that of milk. comparing the work of we must recognise that it is the gastric glands with different purposive in another sense also. The total quantity of juice hydrochloric acid We secreted on bread is only a little larger than that secreted on milk. juice of 3'1 mm. different kinds of proteid receive. Multiplying. of the digestive strengths (in millimetres) yield 38 for and 10 for milk. and on milk. possibly. These figures afford us the possibility of comparing the ferment concentrations in similar volumes of juice. therefore. The squares bread. that as a matter of fact during the digestion of bread by the stomach an excess of hydrochloric acid is avoided. therefore. This The vegetable proteid of bread requires for its digestion much ferment. on flesh. Since. It . flesh.c. poured out on the corresponding food. These 100 grms. of juice of 4'0 nun. 340. from other properties of the juice in this case. For 250 grms. injurious.c. As shown by Dr. we must take this also into consideration in calculating the quantities of ferment. white bread. 27 c. 1(! for flesh.c. of juice of 6'1G mm. whereas considerable quantities of would remain unused. and 340. on the basis of a proportion between amount of food ingested and the quantity It results that for of juice secreted. 250 grms. One may conclude from this that it is only the ferment of the gastric juice which is here in great requisition. quantities of ferment corresponding to the differences in their digestibility which we already know from physiological chemical experiments. the squares of the numbers representing digestive strengths by the number of c. five times more pepsin is poured out than on the same quantity of proteid in the form of milk. When foodstuffs. bread we find no corresponding data. milk. digestive strength.c. Chigin on ferment units. 42 c. 430. of milk and 250 grms.
which is contained in large quantities in bread. as it a half to five minutes. period occurs when the glands are already filled with gastric wall. Consequently. however. Further. of course. whether the observation be made on the complete always occurs. I mean the long pause of at least five minutes' duration which always intervenes between This interval the taking of food and the beginning of secretion. or on the miniature stomach of a dog normally fed. remains but to in the occurrence of the latent period a definite aim. is period physiological-chemical observation that the digestion of starch. cannot.38 is THE WORK OF THE DIGESTIVE GLANDS. to wit. Nothing. but ? may be termed. it are incapable of responding to a stimulus before the lapse and the secretion spreads out over the would be singular to suppose that the gastric its application. is never less than may often be as long as ten four and minutes. in the digestion of bread. stomach with the aid of sham feeding. or at all events with phenomenon also plays a part which has already been several times mentioned but not yet explained. What have no reason for interpreting its significance occurrence on the basis of extrinsic conditions. This cannot be the the latent juice. is impeded by an excess of acid. since here the adaptation of the secretion In the following table the experiall doubt. a large part of the starch of bread escapes unused from the gastro-intestinal canal. Walther are of the pancreatic gland collected together. requirements is beyond ments of Dr. Perhaps recognise these five to ten minutes are reckoned upon to allow the action of the minutes after But this explanation amylolytic ferment of the saliva to proceed. another present in the stomach during This harmonises well with the Towards aiding the digestion of starch. such time as might be supposed to be necessary for the glands to fill up to their mouths. therefore. From clinical observations we know further that. but the whole little hydrochloric acid of secretion. over a much longer time. in cases of hyperacidity. distributed. upon different diets : which illustrate the work . so long as the of question has not been systematically brought under the scrutiny scientific investigation. relation thereto. since fistular orifice. glands per of five se. All the more gladly do I now proceed to the work of the pancreatic to the nature of the gland. This latent period. or till the juice moistens the inner wall of the stomach and is its We runs in streamlets towards the explanation. while flesh is excellently digested. be regarded as very convincing. so that the mean hourly quantity of juice with bread diet is one and a half times less than after taking milk or flesh.
.PANCREATIC SECRETION AND DIET. 39 Diet.
c. . 3'2. Ki'4. 14'1. and the rate of of food. 38-75. 11-2. both as regards the quantity and property of its juice. me that this fact strengthens our previous supposition. It appears to condition. 36-5. 20-D. 14-G. that in the digestion of bread a large accumulation of hydrochloric acid has to be avoided. 12'7. is specialised The work of the pancreas. milk With 250 grins.c. In any pancreatic case. progress which the secretion takes for the different classes This is represented in the accompanying figures and curves and 10) taken from the work of Dr. 7-6. as yet.c. 9 With GOO c.c. Obviously a rich field of work here. G'<J c. 1-0 c. full of important and. 250 grms. 30'4. 1O7. flesh 8-5. Bread. Walther. 44-6.40 THE WORK OF THE DIGESTIVE GLANDS. 9. unsolved problems.c. JI 111 II HI IV V VI VII VIII 1 II III IV V VI HflHBBfllMH Flesh. 50-2. 100 gnus. The secretion of pancreatic juice in hourly quantities (Figs. Milk. Curves of secretion of pancreatic juice with different diets. 1G'9. O'S c. like that of the gastric glands. FlG. bread With 100 grms. the relationship just discussed between the gastric and secretions I bears interesting testimony to the complexity IV V 1 HOURS. 600 c. and beauty lies of the digestive mechanism.
2'li2 K-it ferment. . ->-s8 3-:. p. 5-0 . 4-5 In view of the foregoing facts and of the tendency of all organised tissues (under the influence of forced work or its opposite) to enter into conditions of a more or less stable nature. 41 VARIATIONS OF FERMENT-CONTENT IN HOl'ULY PORTIONS OF r.-) 1-7 5th (ith .. .C. Wassiljew. Starch ferment. 3-88 2-r>:> i-s BREAD. 10.2 2-2 2-1 I'll 2nd :!rd I . 1-2. . to day.VO 14-3 l'J'7 2nd 3rd I .. FLESH. Proteid ferment.. An investigation of the pancreas with this aim proved fruitful.. and is then brought on to an exclusively flesh diet..V2 3-5 2nd 3rd 4th . . If. 2-SS 2-. .. 2'1 2-:> . the amylolytic power of the juice is found to be continuously on the wane. 3-88 4-12 . ON DIETS OF 100 GRMS. tli .) of milk and one Russian pound .V'. .. .. for example. it is found that the fermentcontent of the juice becomes from day to day more and more adapted to new the requirements of the food. Here is an experiment taken from the work of Dr. MILK RESPECTIVELY. When. 3-12 3-88 . one might imagine that similar eS'ects follow also in the case of the glands.c. while. . . . in feeding animals. a continuous increase of the proteid ferment in the juice The capability of digesting proteid waxes from day on the contrary. .~> .\\ CREATIC JUK'K. V7:> 5-88 4-2. 2-38 2-31 7'0 . 1st .~> 7th . a dog has been fed for weeks on nothing but milk and bread. 2-0 .-> 2-5 . 3. two bottles (1200 c. A fistula dog was given daily for a month is to be observed. -12. 2-75 2-3s 2-C. .) FLESH.l tli ... .... . 4-2.. BREAD. .i . . the kind of food is altered. . and a half.V7 1-1 . . AND GOUC. . 1st . . LH2 <rO 4-7:> 3-1 8th .PANCREATIC JUICE AND DIET. 250 GEMS. Hour. 1st ..VI 2 MILK. which contains more proteid but scarcely any carbohydrate.. . and the diet maintained for a length of time. (Sec Fiji.
and milk respectively. bread.">. the different ferments on the same after the lapse of three clays. KV1S.-}">. . 0'25 For the starch ferment in mgms. <>. The hourly quantities of juice for the six hours after the meal had the following digestive powers: For the prcteid ferment in mm. <. the curves represent the same ferment on different diets if 1<>. 3. of sugar. . we obtained the : - following results. during which time the juice continuously altered in the direction mentioned. 7. On the twenty- third day of the flesh diet. l r>. Ceo. Afterwards the dog was fed daily on a pound and a half I i of flesh.~>.~>. 3.) of white bread. diet. 4. r>. l. while the starch ferment declined. this must be added. but that it had merely . 3'0 . in. !'<>. 0'25. Even HOURS 1 11 ill IV II III IV VVI VHVIIIlim IV FIG. The ferments of pancreatic juice on diets of flesh. 4. in a vertical direction. that in this second oase the starch ferment was allowed to act for twice as long as in the first case. 3'. 0'25. 8' 13. If read horizontally. . likewise for the first six hours after feeding For the proteid ferment.42 THE WORK OF THE DIGESTIVE GLANDS. first (410 grms. Although the result of the foregoing experiment is apparently indubitable. one could see that the pruteid ferment tended to increase. l For the starch ferment.>:>. it is possible the objection may be raised that the quantity To of ferments produced in both cases is the same.
such. or from the effects of a long-continued form of diet maintained perhaps throughout the whole period of life. This shuts out all suspicion that dealing with a spontaneous and inevitable alteration of the glands. The result of the experiment. Mett was exactly 4 mm. Although our laboratory dogs live and are fed under the same conditions. When. so far as concerns the proteid-digesting It is naturally of importance. while. last for proteid experiment carried out twenty-four days was absolutely nil. the thirtieth day of this diet the whole juice secreted in twentyIts power of digesting proteid. was placed on a milk and bread diet. leaves nothing to be desired. ferment. We resolved. neverr theless the pancreatic juice of the different animals often differs very In harmony with the same essentially in the amount of ferment. so far as one could conclude from the experiments of the first six hours after the feeding. to compare the fermentative properties of the juice secreted every hour over the whole twenty-four hours.PANCREATIC JUICE AND DIET. to reverse it several times in one and the same animal. however. The proteid-digesting power of the juice sank continuously. by altering the feeding. in order to give the character of absolute certainty to our results. be further investigated. On four hours was collected. 48 been distributed over different hours of the digestion period. fall. thing. we were Since the food so powerfully affects the nature of the work of the glands. at the showed irregular beginning steadily increased later. as obtains in the case of pedigree dogs ? It appears to me that our tions of experimental material gives some indications in this direction. A third trial was made after the lapse of a further twelve In the fourth and days. which gave a digesting power of 1-25 mm. experiment was carried out by Dr. Ten days later the experiment was repeated when the digestive power of the twenty-four hours' juice had declined to 2'25. therefore. however. This prolonged A dog which had long been fed on flesh. this or that condition of the pancreas had been established in our experiment animals in characteristic form. a change of diet in the case of one clog may very soon manifest itself in altered properties of the juice. we were able. under the influence of a given diet. the . starch ferment it later. Jablonski. the digestive power The however. fluctuations. such as might have arisen from the effects of the operation or other pathological condition. for example. in that of another. with a slight tendency to . and whose pancreatic juice worked very actively on proteid. to investigate the behaviour of the other ferments with equal exactness. The latter circumstance must. is it not possible that a permanent type of pancreatic activity could be produced under the influence of long-standing natural condilife. according to .
up to the present. that the work of the digestive glands so far investigated is unusually complex and elastic. Lobassoft'. in accordance with the burden of work placed upon the digestive canal. remoulding of the pancreas takes place in the slowest manner. such as oatmeal porridge. but at the same time astonishingly exact and purposive. Dealing now with the gastric glands. howfreely every flesh for day on ever. the secretion of gastric juice becomes The dog in gradually less and less (observation of this laboratory). must fact not easy of interpretation. in a dog thus operated upon. After a considerable time they were again tested with the sham feeding. How we the ferment to interpret these results ? Are our methods of estimating content of the gastric juice defective. Towards this problem Dr. pancreatic glands differ in this important respect ? that the pancreas plays the role of a supplementary. so to speak. have never seen any striking or constant differences in the digestive power of the juice obtained from the dogs. In such cases as the latter. in a small number of cases. .and oesophagotomised dogs. or do the gastric and It is. when such division of the vagi. reserve A ever. but. Samojloff" (in experiments not yet published) solving has made observations on three gastro. howalways work at maximum pitch. It is true we have only encountered the latter property v. but the gastric juice showed no are essential divergence from that previously obtained. &c. a certain test meal always yielded a much greater secretion than when the animal had been fed upon other foods. as the first important digestive agency. which possibly indicates that lasting alteratioi^s of the gastric glands may have appear under the influence of prolonged dietetic conditions. notwithstanding. We in our possession a dog with a portion of the fundus of the stomach that is to say. dogs live for a length of time after the operation. the following condition When it had been fed question presented : a considerable period of time. the question of lasting alterations in ferment production must be left unanswered.-ith unquestioned distinctness. of course. an abrupt transition from one regime to a different one can often produce serious illness. while the gastric glands. namely. possible gland.41 THE WORK OF THE DIGESTIVE GLANDS. have up to We the present obtained gastric juice in the laboratory from a very large number of dogs (twenty to thirty) by means of the sham feeding. now diminished. which may be once more repeated All the facts given accord sufficiently with our previous conclusion. too great stress cannot be laid upon this instance. Since. whose duty is now augmented. The dogs were tested beforehand by oft-repeated sham feeding experiments and then placed on different diets. has recently been observed in our laboratory by Dr. involving isolated according to the method of Heidenhain I ought to add that. the gastric glands live and work under altered conditions.
which. Moreover. the late Professor Carl Ludwig. by what means is this made possible ? this apparent instinct of the glands depend and in what does naturally ask ourwhat does it consist ? A probable answer to this question is easily given. N. GENTLEMEN. It is only when such supposition proves itself to be untenable that we must seek for another. however. that the widely renowned physiologist of Leipzig. have taught us an extremely It was made evident that the gastric and pancreatic interesting lesson. .11. It is appropriate that I should mention by way of introduction. 1S. Mrdizln. THE CENTRIFUGAL (EFFERENT) NERVES OF THE GASTRIC GLANDS AND OF THE PANCREAS. We will therefore concern ourselves in the present lecture with a study of the influence which the nervous system exerts upon the activity of the gastric glands and the pancreas. and naturally an explanation of the adaptability of the glands is above all to be sought for in their innervation. We On selves at once. have what we may call a form of instinct. upon the influence of section and excitation of the The gastric glands can be excited by remote influences The sham-feeding experiment Repetition of the same The latter convey secretory fibres to the gastric after division of the vagi glands This is proved by excitation of the vagus nerve in two different forms of experiment The vagus is the secretory nerve of the pancreas Karlier experiments vagi nerves on the gastric glands glands (experiment) It also conveys inhibitory influences to the foregoing These depend upon the activity of inhibitory nerves of secretion. They pour out glands their juice in a manner which exactly corresponds. they secrete precisely that quality of fluid which is most advantageous for the digestion of the meal. to the amount and kind of food partaken of.. On the last occasion we were occupied with wearisome figures and curves.F. both qualitatively and quantitatively. no less than forty-five years agothat the salivary glands possess a special nervewhich immediately * Zt'itxchrijt fiif rat.* was able to prove by a classic experiment. i.LECTUKE III.
Studien des xvii. recognise two kinds of special nerve-fibres which govern the activity of the salivary glands. Physiologists. the production of the watery and inorganic constituents of the secretion. to wit. Heidenhain. Heidenhain. pathological phenomena is nothing but an endless series of the most different and unusual combinations of physiological occurrences which never make their appearance in the normal sti^ange. irttyxtol.40 THE WORK OF THE DIGESTIVE GLANDS." It has also been termed an " anabolic nerve. but by no means isolated. the other leads to an accumulation of the For the former. had fruitlessly endeavoured for decades to arrive at definite results upon this question. iv. They had even come to recognise different morbid conditions of the innervation apparatus. Corresponding to these two subdivisions of the process. The world of course of life. carrying the matter farther. " calls into play brings on the secretion of " " cells. . and which and life institute. resolves itself into two processes . 1878. produced undoubted proof that the secretion of saliva in the glands. and with him the majority of physiologists. The question as to whether the gastric glands have likewise a special secretory innervatiori is now a very old one and has had an interestIn this matter physiology stood for a long time in sharp with practical medicine. and thereby This nerve receives the name of a secretory or " katabolic nerve. Inatltidx zu lirrxluii. at his could scarcely be called into existence by means of the technical resources command. conflict " and looked upon the existence of secretory nerves to the stomach aa undoubted. the specific agency of the secretion. the chemical activity of the gland saliva. on the other hand. Clinical observation will consequently always remain It is therefore only perfectly natural a rich mine of physiological facts. * I'll of the we are fortunately R. instance where the physician gives a more correct verdict concerning physiological processes than the physiologist himself nor is it indeed . the latter he " termed " trophic. that the physiologist should endeavour to maintain a close connection between his science and that of medicine. 18(!8 and iiger's Arckir. the ferment. Bd. The one influences the secretion of water and of the inorganic salts which it holds in solution. and the preparation of a specific organic body. organic body. This is a striking. Heidenhain retained the old name of " secretory nerve. Heidenhain.* the late physiologist of Breslau. Notwithstanding the wide range and perplexing nature literature of the innervation of the gastric glands. had long answered the question in the affirmative. Physicians bringing forward their observations in proof. It is a series of physiological experiments which nature often with such an interlinking of events as could never enter into the mind of the present-day physiologist. ing career.
It furnishes evidence of a nervous influence even where the first direct method remains This consists in the most general evidence of relationship fruitless. cling to many of our physiological methods. or in other way paralyse. deviate from A When the stimulation calls forth each time the same alterations of function. The well-known fact of salivation at the sight of appetising food has in this way always been accepted as a sound proof of nervous influence over the salivary glands. between the organ in question and the nervous system. with clearness and precision. and come nearer the truth. rightly and justly look upon this nerve as governing the Even here. other hand. have three commonly employed methods by means of which the We existence of nervous control over any organ may be determined. we may organ. experiments with a negative result enjoy only a very On the qualified reputation. It is nothing more special than making accurate observations in the clinique and in everyday life. the more completely and perfectly we compare the two conditions. and then submit the organ to accurate observation in order to determine whether its activity has been suspended or increased. or in any other way made to the normal. There is still mode of proof which. unfortunately. Naturally our conclusions regarding the relations between the nerve and the organ will be all the more accurate. and in view of the defects which still. Into this path research also wandered when the innervation of the gastric glands was brought under investigation. certain nerves which are in anatomical connection with the part in question. the alterations of function which appear in an organ on stimulation of this or that nerve may have been brought about in- directly through the intervention of one. one must not disregard the possibility two contingencies. Only a careful and complete physiological (and where necessary anatomical) isolation of the organ can guard us from this source of a third error. however. or several other organs. . perhaps. second and striking proof of the existence of a nei-vous influence over an organ is afforded by the results of excitation of its associated nerve. we can cut through. to grasp the cause of their want of success.NERVOUS CONTROL OF SECRETINC! GLANDS. For this reason. now 47 in a position to pick out. the fundamental truths of the earlier researches. and by many authors are never published. of suffers and when these at once disappear on cessation of the stimulus. First. It may occur that the function of the organ no alteration because the nerve or the organ is placed under abnormal conditions. this is very possible. should have been more correctly given in the first instance. either quantitatively or qualitatively. and from their historical teaching to draw precepts for the performance of an ideal experiment which shall definitely answer our question.
the subdiaphragmatic division of the nerves takes no cognisance whatever of the possibility that the secretory fibres of the vagus may course enter the wall of the digestive tube above the diaphragm and down to the stomach along its deeper layers. We become more and more convinced every day that the animal body is governed on the In this principle of mutual help and defence by the several organs. and is usually followed by death after a few of tenest after two to three days. But one can raise two important objections to the experiments. the survival of the animals cannot be taken as a proof that no deviation in the activity of the from the normal had occurred.48 THE WORK OF THE DIGESTIVE GLANDS. the action of It was consequently hazardous to the gastric glands is also disturbed. as often as the vagi. that. moreover. (This is a good example of the importance of the first of our rules for nerve-division experiments. no matter where or how they stimulated the nerve. First. could claim a result which recalled * a secretory K-nr hi effect. division of the vagi on both sides of the neck is an operation which is accompanied by the severest conse- quences to the animal. it is not to be wondered at. had made no precise and of the secretory activity of the stomach before and detailed comparison gastric glands after vagotomy. were divided in the neck. relative to the experiment of nerve-division. conclude from such an experiment that the vagus bore a direct relationship to the gastric glands. that the vagus stood in intimate relationship to the secretory activity of the stomach.) Secondly. amongst other things. isi!7. case it should have been boi-ne in mind that the sympathetic nerve also sends fibres to the stomach.) attitude towards the experiment seemed all the more justified because Schiff * was able without difficulty to preserve his dogs in good health and nutrition after division of both vagi beneath the diaphragm. the chief anatomical nerves of the of But only very few were convinced organ. The animals increased in weight. These experiments of Schiff have had decided weight with many investigators against the recognition of any secretory innervation whatever. and unfortunately the view is still extant. Schiff. If this operation in the course of such a short time brings the whole functions of the organism to a standstill. Lpqons physiologic de . As is well known. (This is a good illustration of the second rule given Such a cautious above. perhaps even more doubtful. results of The excitation of the vagus nerve proved to be also uncertain. not very Schiff. Hardly any of the authors. Several workers had observed disturbances of the secretory activity the stomach which concerned the quantity or properties of the gastric juice. and the younger dogs grew and thrived as if nothing had happened. The few and la digestion.
one needs only to excite a dog by the sight of food in order to call forth a secretion of gastric juice. Soon after the patient took anything sweet or acid into the mouth. one can always find related by the French authors this or that apparently convincing experiment. have maintained till recent times a rigidly dogmatic attitude against secretory innervation of the organ. under certain circumstances. therefore. amongst the records. however.nd the excitation of nerves have not yielded results when applied to the gastric glands. such as one may well conceive might occur from the contractions of the stomach-wall set up by the nerve excitation. I must here remark. C. constant results. It is interesting to observe how differently the question of secretory innervation of the stomach has been treated by German and French While the German physiologists. demanding precise and physiologists. forty-five minutes after the execution. f Journal de I'Anatomie et de la Physiologic^ 1878. stands the experiment of two French authors. Attempts with the sympathetic turned out to be equally negative. 49 convincing records of positive results received no attention in the loud chorus of confident denial. that this was possibly only an expression of gastric juice out of the glands. Quite alone. Although some authors could not verify this statement. &<_. Schmidt.. or at least not such as have convinced the The third method of procedure was." Progr&s Med. form the point of departure for new researches * Regnard et Loye. Die Verdauungssafte. Bidder u. In the year 1852 Bidder and Schmidt t had observed that. and Richet's observations prove that the nervous system exerts an influence on the secretion of gastric juice. the first two forms of experiment which I have named the division a. and still saw drops of gastric juice forming on the inner surface of the stomach.V t F.THE CENTRIFUGAL GASTRIC NERVES. 1852. Richet was able to perceive Bidder and Schmidt's experiments a secretion of pure gastric juice. Later we shall adduce facts which make the appearance of a true secretory activity but little likely under the experimental conditions of these authors.. D . and this all the more because the experimental ai-rangeuients were similar in all these contradictory investigations. be this direct or indirect. 1SS. ever. or at least may happen upon forms of expression which assume the existence of such an innervation to be probable. howmajority of physiologists. the majority have been able to convince themselves of its truth. consequently.* who stimulated the vagus of a decapitated criminal. strikingly productive. This fact must. "Experiences sur un Supplicie. In recent times the French physiologist Richet J has had the opportunity of making observations on a patient on whom the operation of gastrotomy had been perfoi med for an incurable stricture of the oesophagus.
may proceed the time of making the gastric fistula. I prepared the left vagus free in the neck. passing a loop : of thread round the nerve. the light vagus nerve was In this its recurrent laryngeal and cardiac branches. we have of juice. at further experiments. . stomach has been washed out before the beginning of the lecture. two." It expresses the idea. may continue to feed the dog as long as we the secretion will flow at the same rate for one. . but is continued for a long time out. mouth Its is cut off and has had its oesophagus divided as well. and now. three to four hours gradually dying Without waiting. I give the dog food. already 20 five c. and. not a single drop of fluid escapes from the fistula. the stream steadily becomes greater and greater. we take away the animal's food. so that the from all communication with the cavity of the stomach. It only remained to constant and simple in other words. it has really been fed. the laryngeal and cardiac fibres remained divided below intact. Concerning what constitutes the actual stimulus in this case I experiment is clear. hours. have even had dogs so greedy that they did not tire of eating in this fashion for five or six hours. It possesses an ordinary gastric fistula with metallic cannula. It proves incontrovertibly that the gastric glands are influenced through nerves by " distant effect. secreting during the time a total We We quantity of up to 700 this c. however. or more wish. If now. to facilitate seek out its proper interpretation. before the division. About three hours before the present lecture. but the whole of the food swallowed. After feeding in this way (which for shortness we will henceforth name " sham feeding ")* for five minutes. The animal eats greedily. to for its complete cessation we In the case of this dog. only the pulmonary and abdominal branches on the side in question were thrown out of function.c. the secretion does not cease immediately. minutes after the commencement of secretion." since the phenomenon comes to pass without any immediate contact between the food and the on the subject. It is will deal later.50 THE WORK OF THE DIGESTIVE GLANDS. make its the experiment reproduction and of fact. At present we will carry our experiment a step farther by dividing the vagi nerves. as you see now.c. way. I am now able to demonstrate experiments to which yield absolutely constant and unequivocal results. but not term that The corresponding Russian expression would be better represented by the " make-believe feeding. of the purest gastric juice. perfectly pure gastric juice makes its appearance at the fistula. gastric mucous membrane. We you have here before us a dog operated upon in the manner I have described As a matter in the first lecture. The meaning of obvious that the effect of the feeding is transmitted by nervous channels to the gastric glands. comes out again at the cesophageal opening in the neck. from the dog's point of view.
of carrying The dog which served for the above experiment remained alive for several Later the right vagus was also divided in tJir neck. that the dog. dividing it. and the same result was observed by Dr. Jiirgens with dogs having both vagi severed below the diaphragm. (This is an essential advantage of our procedure over the former method. as right side the laryngeal and cardiac fibres are intact. We may feed the dog as long as we wish. shows no you see. gentlemen. as I have said. possibility of Basing my only necessary to accomplish the division of the vagi nerves under suitable conditions in order to achieve indubitable and invariable results. there can be no question of any general feeling of severe discomfort producing a harmful influence on the secretion of gastric juice. At present the pulmonary and abdominal vagi on both sides are paralysed. We sharp contrast to the previous sham feeding. and always with the same result. The same condition was manifested by another dog which likewise survived a double division of the cervical vagi for * many months. however. but. no feeling of general ill-health whatever. Since the fibres of the cervical vagus which go to the larynx and heart are not completely divided (the abdominal fibres are. in dition. but never again shall we see a secretion of gastric juice in this animal as the result of sham feeding. the dog immediately after the operation quite as lustily as before. fifteen minutes. which it eats with increasing greed for five. :>1 wards. The experiment demon- strated to you will. we do not see a single drop of juice flowing from the stomach. and sever pull gently cm the thread to draw the nerve outwith a sharp snip of the scissors. after division of the left cervical vagus. indeed.* were first carried out by me in conjunction These investigations with Madam Schumow-Simanowski. as fibres in forming the flaps. . while on the I it now The result is.SECRETORY NERVES OF THE STOMACH. Oft-repeated experiments with sham feeding never gave a secretion. and this. totally severed). animal continued in perfect health and enjoyed its life to the full. a like effect was obtained by Professor Ssanozki in a dog with the may be repeated at fundus of the stomach resected after the manner of Heidenhain. There eats is. the usual causes of danger to the animal after complete division of the cervical vagi on the two sides. I take the liberty of asserting that the proof of the matter in question has been raised beyond the You see then. indication whatever of a pathological or otherwise uncomfortable conThere are even no symptoms of cardiac or laryngeal distress. conclusions on these results. again offer the dog food to eat. ten. an is well known. always happens when these conditions are fulfilled. nevertheless the months. that it is all doubt or chance. and repeat our experiment in the next few days as often as we desire. Finally. the division of the vagus operation which involves.
adequate criterion. Pieces of flesh given at intervals. and of a much more watery kind that is to say.52 THE WORK OF THE DIGESTIVE GLANDS out a double vagotomy. the sham feeding experiment can easily be graduated. that the gastric glands are deprived It only shows that certain exciting impulses. glands receive their normal impulses to activity by means of nerve fibres which run in the fibres or Are they special secretory do they only influence the glands indirectly for example. GJ nun. According to the view now through the medium of the vessels ? with regard to secretory phenomena in glands generally. the less specific constituents could it carry away in solution from them. In the act of eating. digestive power. can. or. the prevalent second supposition is but little probable. It is the gastric glands by way of the vagi. however. the best proof of the specific activity of the nerve fibres supplying the If only vasomotor (dilating) fibres for the glands were conglands. offer it a meal but little relishing. give a dog a highly appetising It is food. the more and the richer the juice we receive. less rich in ferment. 2 ) ^ 5^ . but the juice possesses a much lower It follows from this that the stronger we stimulate. perfectly provement as compared with the experiments of Schiff. tained in the vagi nerves./. possible that other forms of stimuli exist. or even in some wholly different way. Pieces of flesli given continuously. on the other hand.. which act on the gastric glands through other nerves. The negative effect of sham feeding after the vagotomy does not of all secreting prove. for instance. when we give pieces of flesh at long intervals. Ketscher : DIGESTIVE POWEK OF THE JUICE. This fact is.) Lastly and this is the most essential characterwe employ in the sham feeding an immediate. and it becomes still less so But what kind could we adduce The stimulating direct proof in effect vagi. In proof of the above I give here a few figures taken from the work of Dr. of We known that a dog eats flesh with much greater greed than bread. other things being equal. an augmented flow of juice from strong The more excitation would mean a lessening of the concentration.. we give it bread we obtain much less juice. istic of our experiments This is an important imand uniform. we obtain not only less juice than if the dog had been fed rapidly. Likewise.] 8 Si 7J . rapidly the fluid passes through the glands. have been removed. the gastric entirely without nerves.. well If . however. of fibres are these? favour of the truth of the first. iiiDi. however. which reach power..
into true secretory and trophic. Petersburg. and. the sciatic nerve for two or three minutes is able to bring gastric From this arose the attempt a standstill. specific and not merely vaso-motor.* and unquestionably inhibit the work of the pancreatic gland for long Dr. it is nevertheless desirable. " " doubtful if the ordinary so-called acute experiment. first. that sensory stimuli frequently conjunction with Professor Afanassjew. that one must divide these fibres. This accounts for the failure of many authors who have previously worked at the question. and later by myself. Fnaug-Diss. reasons. xvi. 1882. . indeed. interpretation of normal conditions We ments many physiological phenomena are misrepresented or. In all 53 instances when the pieces of intervals.THE VAGUS A TRUE SECRETORY NERVE. shown by the fact that the separation of the water and the preparation of the solid constituents obviously take place independently of each have already seen a number of instances in the second lecture where similar hourly quantities of juice were secreted with wholly different amounts of ferment. In our case this doubt was all the more justified because unquestioned proof of the inhibitory effects of sensory and upon the activity of the most important had already been recorded in the literature. of working of a given nerve and the In certain cases great difficulties come the way of the experiment. From the above data fibres. the quantities of juice meat were administered at were much smaller than when the that secretory animal was continuously fed. Bd. can be accepted as a true In such experiin the organism. wholly masked. dependent on the working conother. it follows. * Pfliiger's Archir. We ditions of the glands. Netschajew t has found also that an excitation of periods of time. run in the vagus to the gastric glands.-ml'tfx. Absondemng dm Mniii-it. But while the nerve-section experiment speaks eloquently of the existence of secretory nerves to the in favour stomach. secondly. to take into consideration the excitation only by artificial excitation that we can study accurately all its features the it mode process which controls. as Heidenhain This is did for the salivary glands.* in Ludwig's laboratory. digestion for several hours to to excite the nerves passing to the stomach in such a way that no or accompany the sensory or other reflex impulses could precede experimental stimulation. for many It is method and in in also. performed very at one sitting on a fresh unprepared animal. il'ic f Vber kemmpndc El nfl iixw anf St. in reflex stimuli in general. It was digestive glands shown by Bernstein. In to resort to a carrying out the investigation we have had once more the special is started with the assumption that it arrangement of our own.
narcosis. the right vagus was cut through below the origin of the inferior laryngeal and heart fibres. after a speedy precautions. and were able to see. after the usual interval of five minutes. the left divided in the region of the neck. This occurred. Uschakofi* employed a short chloroform narcosis (10-15 minutes duration). that is to say. In our later experiments. Experiments specially performed with this object on dogs previously gastro. only in half of all the cases. The . By such precautions we invariably succeeded in obtaining a secretion of juice from the empty stomach when the nerve was subsequently excited by slow induction shocks at intervals of one to to two seconds (so-called " rhythmic excitation "). carried out under positive result much oftener. the food passage the region of the neck and also at the pylorus. during which all the above operations were carried out in great haste. Fifteen to twenty minutes after the narcosis the dogs were again lively. After three to four days the stitches were carefully re- moved from the skin and the wound lay free before us. longer or shorter piece of the peripheral end of the latter had been A prepared free. Gastrotomy and cesophagotomy had previously been carried out.and a\sophagotomised had shown that a chloroform narcosis of such short duration was not followed by any serious interference with the glands or their nerves. in his first experiments. divided the spinal cord below the medulla oblongata with the greatest possible rapidity (a few seconds) in order to prevent all reflex effects on the gastric glands from his further operative procedures. placed on a ligature. In dogs prepared as described at one sitting. however. animal was then suspended in a standing position in a frame. when the nerve In this way we avoided appreciable discomfort to the animal before exciting the nerve. during which the we obtained a excitation remained ineffective. and for the time being preserved under the skin. it only appeared after the lapse of a preliminary period. on a dog prepared at the time. ordinary ligatured fistula in The vagi nerves were now sought out and divided an cannula was brought into the stomach. In none of the successful cases was the effect of the stimulus seen immediately.54 THE WORK OF THE DIGESTIVE GLANDS. ate with greed the food set before them. an undoubted and vigorous effect from the stimulation. as we expected. This we (Madam Schumow-Shnanowski and I) achieved with dogs in a manner similar to that which has been now demonstrated prepared to you. This period lasted from fifteen minutes . but naturally with the observance of certain Dr. Uschakoff. we proceeded to excite the nerves. gastric juice of strong digestive power began to flow from the stomach in perfectly normal quantity. and. painlessly opened. but careful tracheotomy. And now that we had the matter under perfect control we attempted " achieve the same thing in the " acute experiment. In his later experiments Dr.
the secretory effect disappeared only gradually. and we must hope that further investigations into the woi'k of the sympathetic nervous system now that we know the relationship between . secretory fibres for the gastric glands. and yet the period of latency is quite as long in this form of experiment with narcosis as without it. notwithstanding the narcosis and the division of the spinal cord. of direct proofs recently established. however. This view would find its simplest expression lating in a hypothesis which assumes the presence of inhibitory nerves acting in antagonism to the secretory. The existence of so lengthened a period during which stimulation no result may be explained. on a dog with Heiclenhain's resected stomach. if the stimulus were reapplied a few minutes later. in a perfectly convincing manner. where we shall find ourselves in possession of a series of appropriate facts and. Atropin. to return however. which restrains secretion. after the nerve began to work. both as regards the commencement of the flow as well as the product formed. 55 an hour or more. that in artificially stimuglands. however. If. however. in the first instance. and this time with rapidity. gives a perfectly normal secretion of juice. indeed. both exciting as well as restraining impulses are transmitted to the glands. exercised any important reflex inhibitory influence on the gastric are forced to conclude. Furthermore. From both forms of experiment. we are fully justified in concluding that the vagus nerve contains It is necessai-y to repeat. Whether this secretion. therefore. Professor Ssanozki has been able to verify the inhibitory effects of atropin. This hypothesis will be more fully discussed in connection with the pancreas. destroys the irritability of the nerves. the stimulus were removed. on the ground that the shock of the operation depressed the excitability of It can. It is hardly conceivable that the operation. is to be of the sympathetic nerves or to some other agency cannot for the present be decided.EFFECTS to Otf DIRECT EXCITATION OF THE VAGtTS. be explained in another and the gastric glands. in a manner similar to what we already know exists in the innervation of the heart. the chronic as well as the acute. and therefore with the vagus fibres divided. that one must not infer that the integrity of the vagus is the only requisite condition for the secretory work of the stomach. the vessels and other organs. therefore. Naturally the work of secretion under these conditions deviates not inconsiderably from the normal. We the vagus. of the nerves gives We soon after the anaesthetic. paralyses secretory nerve mechanisms in a very special manner. The administration of a drug such as atropin. and we ourselves as well. ascribed to the action which occurs after the vagi are severed. have already remarked that sham feeding. very more likely way. have been convinced that the stomach is capable of preparing its specific secretion in the absence of vagus influence. Many previous investigators.
Pfliiger's ArrJiir. which is the special point in question. sensitiveness of the organ. will yield the selfsame results hands of every investigator. p. but we do Other not conclude therefrom that these organs have no innervation. every observer who has occupied for any length of time with investigating the functions of the himself : pancreas will leave this field with a feeling of dissatisfaction in consequence of the extremely large number of fruitless experiments he is obliged to deduct from the total number of his investigations for not even the greatest care. I need only give here the pancreas. although they have been published for more than seven years." * Bd. The same difficulties which we had to encounter with regard to the innervation of the gastric glands were also met with in the case of the To illustrate these difficulties. such as birds and frogs. and will no longer leave any room for of the doubt in the existence of a secretory innervation to the glands stomach. always clings to the results of such observations. taking 110 precautions against reflex inhibition. following expressive remarks taken from the classic work of Heiden" hain upon the pancreas Indeed.5fi THE WORK OF THE DIGESTIVE GLANDS. x. believe that every single repetition of our experiment. Schmeyer) have obtained more or less positive conand other animals. In the case of many other organs we are able to divide their nerves without stopping their particular forms of activity. but pay no regard to the peculiar alterations of the secretion. continue to disregard the results just given. . will be able to overcome the incomprehensible . Contejean. not only in the Russian but also in the foreign literature of our subject. Only a few results with dogs (Axenfeld. if only fidently We the conditions in the we have given be observed. never undertaken a series of experiments which sacrifice of not set aside even by a numerous I must openly declare that I have dogs and so poor * was so lavish in the in corresponding results. therefore.. Some of the authors speak of the continuance of gastric secretion after severance of the vagi. R90. cannot leave this subject without expressing regret that physiologists have grown accustomed to look upon the gastric glands as being We independent of nervous influences. 1875. A degree of uncertainty. authors adhere rigidly to the traditional formula? of the acute experiment. a function which it does not resume even under the influence of the most favourable secre- tory conditions. in consequence. vagus nerve and the gastric glands will the finally elucidate the problem. nor the ripest experience in the making of pancreatic fistula?. which is repetition of the single experiments. and. which only too often annuls its function for a length of time after completion of the operation.
it was the good fortune of myself ? and my co-workers to be able to solve them all. an undoubted effect upon the gland. the dog remains perfectly still in its frame without Two minutes elapse without any exhibiting the least degree of pain. We with Professor Afanassjew. and I fully recovered . wall where the orifice of the pancreatic duct is situated. I now carefully remove the cutaneous sutures. have shown that sensory stimuli exert an Then Heidenhain. and obtain the same effect. manner cautiously draw forward the ligature with the nerve. however. . Now As you see. On the whole. and the method by which we were first convinced of its activity will be immediately demonstrated to you.NERVES OF THE PANCREAS. upon evei-y dog. Zur Physiologic d. made in the have described in the first lecture. the question of innervation of the pancreas remained very obscure. Why could Heidenhain obtain an effect only in exceptional cases ? By what channels were the in a impulses conducted from the central nervous system to the gland ? To what influences were the inhibitory effects of sensory stimuli to be attributed There was as yet no answer to all these questions. and this is invariably the result. The animal has from the operation and everything is healed. his pupil Landau. and is followed by others in quicker and quicker succession. result from the stimulus this I ask you especially to bear in mind juice flows. more or less satisfactorily. few experiments out of many fruitless ones. It resulted that the vagus is to be looked upon as the secretory nerve For this discovery we have to thank the adoption of a special expei^imental procedure. At the f>7 present moment. and now. with inhibitory influence on the pancreatic gland. and only stops at the end of four to five minutes from the cessation of the stimulus. in the third minute. in Ludwig's laboratory. not a drop of I begin to excite the nerve with an induction current. The dog before you is provided with a permanent pancreatic fistula. obtained. Rresltm. furnished with a ligature. To this must be added that the vagus nerve naturally was stimulated * before. however. I again apply the current. After three minutes I interrupt the excitation. Beginning with the year 1X87. the wide end of which embraces the site on the abdominal funnel. Baitch&peichelabsonderiing. the investigation of the nervous have already relationships of these glands has greatly advanced. Four days ago the cervical vagus was divided on one side the peripheral end of the nerve was laid bare. of the pancreas.* from excitation of the medulla oblongata. and preserved under the skin. but the juice continues to flow spontaneously. stated that Bernstein. 1x7:!. without causing the I beg you to note that from the metallic dog appreciable discomfort. the first drop of juice makes its appearance. and afterwards myself.
On the fourth day after its section. On the other hand. and then. as possible ing only a couple of seconds. In order to comprehend this circumstance it is to necessary excitability of different nerve fibres disappears after section. These specialities are two : The animal cotised. and not even narthe usual custom elsewhere. the glands have neither suffered by the operation nor by the circumstances accompanying the excitation. as is is subjected to no painful sensations. Under such conditions we are able in every experiment. to observe the secretory the vagus on the pancreatic gland. thanks which has elapsed (four days) since the vagus was divided. with the same object. although we may have to excite the nerve perhaps several times at the beginning of the experieffect of ment without once clear. occupybe only carried out on a suitable plan. avoided. may well as in those of earlier authors. The meaning of the foregoing procedure is at By division of the spinal cord the harmful inhibitory effects of the exceedingly exciting the vagus in the thorax. Thus. The reason of our success lies in the nature of our preparations for the experiment. have lost so irritability that the strongest excitation of the nerve much is of their scarcely able to produce even an insignificant and momentary slowing of the heartbeats. in order to prevent it for a long time from reacting to a previously effective stimulation of the vagus. and yet what we are now able publicly to demonstrate was never seen.58 THE WORK OF THE DIGESTIVE GLANDS. In our own experiments. a cannula is tied into the pancreatic duct. with different degrees of rapidity. lengthy operations are prevented. while. It is only necessary to excite its vaso-constrictors for a short period (two to three minutes). for example. : Tracheotomy is order to seek out the vagi below the heart. in this case the remember that the cardio-inhibitory fibres lose their irritability earlier than the secretory fibres for the pancreas. result. or to compress the aorta for tho same length of as itself to time. From these experiments it can . circulatory disturbances are excluded which would otherwise follow the excitation of this nerve. by its influence on the heart's beats is to our notice Further investigations carried out in this manner have brought two conditions under which the secretion of the pancreas be inhibited by nervous influences. In our experiment. But one can ment. therefore. after which artificial respiration is set up. And now we may quietly proceed further. the pancreas has shown be extraordinarily sensitive to circulatory disturbances. The chest is opened in Our procedure performed as quickly and painlessly then the cervical cord is severed from the medulla. all to the time the cardio-inhibitory fibres. after opening the abdominal cavity. is if also obtain a positive result with the acute experi- it as follows .
one is at liberty to make several supposithey may come into play through the vaso-constrictor nerves of the organ. But when groups. one pancreas. to excite the other vagus in like manner. accompanied by the strongest sensory stimulation. seconds to one minute) between the application of the stimulus and the A appearance of the secretory effect. The following are the results of the work of Professor Kudrewezki : If. during the later stages of the excitation. exc'ting influences. pass through the vagus to the to flow only after the stimulus In very many instances the juice had already ceased. it is now only necessaiy. lastly. I will now bring forward some experiments dealing with the influence of the sympathetic nervous system upon the secretion of pancreatic juice. easily be understood 59 why. . because here the inhibitory phenomena are mostly prominently marked. Great importance must also be attached to another circumstance which likewise attracted our attention during the investigation. and consequently by vascular constriction. In the " experiment just now demonstrated. a gentle intermittent of the secretion is observed. even the gland of an animal taken at the height of digestion. the sympathetic nerve excited by means of an induced current. in the ordinary method of operating. after a definite often considerable length of time. the secretion is brought to a All these phenomena have led to the idea that not alone standstill. through genuine inhibitory nerves antagonistic to the secretory. that the question of their existence must be finally decided by a study of the nerves to the stomach and pancreas. often does not yield a single drop of juice. however. began can often observe the following phenomenon (Mett). Before considering this question more fully. Suppose the right vagus be excited for a considerable length of time. It appears to me. or. be in the acute experiment described above. Finally. or through constrictor effects upon the excretory ducts. excitation of the vagus did not call forth the secretion of juice instancertain period of time always elapsed (from fifteen taneously.NEKVKS OF THE PANCREAS. without interrupting the stimulus. In the physiological literature of the last few years one may find references here and there to inhibitory innervation of the glands. when. With tions : respect to the latter. but also inhibitory. It is it the nerves which has been proved beyond doubt for several organs that regulate them may be classed into two opposing why may not the same be rightly maintained for the glands ? quite possible that an antagonism of this nature applies to the genera] principle of innei vation. as also in the acute" experiment. and thereby a steady flow of juice set up. They will fuinish us at the same time with material for the discussion of the above subject. advance but only during the first few seconds. and after its stoppage.
only under special conditions which eliminate the activity of the vasoconstrictor nerves (e. of Francois Frank. know that vaso-constrictor nerves are much less sensitive to mechanical stimuli than many others. therefore. . The possibility of simultaneous action on the part of the motor nerves to the excretory ducts was excluded by ments . influence of the vagus Its inhibitory effect upon the secretion This gives us good reason for believing that the in does not depend upon a contraction of question. with a fresh nerve. however. mechanical excitation. correctly assume (1) that both vaso-constrictor as well as secretory fibres for the pancreatic gland run in the sympathetic nerve (2) that in ordinary electric excitation the vaso-constrictor effects completely mask the secretory and (3) that . or the application of the electric current to a previously divided nerve) can the secretory fibres effects. ing often to complete standstill of the secretion is obtained every time without exception. not. Excitation of the sympathetic. He has. only slows the secretion. Popielski. The question with which we are dealing has recently made an important advance.g. thanks to the work of Dr. but with one which has been divided three or four days before. on the other hand. We may.. employment modes of stimulation mentioned above. the vagus nerve dilates rather than constricts the vessels of the pancreas. the glands Our opinion regarding has suffered. in the first instance. By this a long. The same may be obtained by means of electrical stimulation. During the acute experiment already described a solution of hydrochloric acid is poured into the duodenum. the secretion is completely arrested. If. mechanical stimulation (for example.80 THE WORK OF THE DIGESTIVE GLANDS. full. a different result is to be observed a little : time after the beginning of the excitation a tolerably strong secretion sets in. worked out a plan of experiment in which the inhibitory effect of the vagus upon the pancreas is shown in a constant and striking manner. instead of the electric current. is partially degeThe meaning of these events is easy to understand when nerated. however. of the special no alteration as a result remains to the the vessels.continued and vigorous secretion of pancreatic juice is means If the vagus nerve be now strongly stimulated. and this after the lapse of some time. one mind a few facts from the physiology of the vascular nerves. in consequence. manifest their In the case clearly of the sympathetic nerve we are relationship now the mutual of which exists between able to recognise the vaso- motor and secretory the effect fibres of the the vagus upon of the pancreas. a series of taps with Heidenhain's tetanometer) be employed. and lose their irritability much calls to We earlier after division. a slowcalled forth. A compression of the aorta arrests the secretion only after two to three To this it must be added that. according to the last experiminutes. and which.
vaso-motor nerves constrictor and dilator also pertain to the glands. it is easy to understand their reflex excitation both under normal Nor is the possibility conditions as well as during the operations. From the latter fact we must conclude that in the branches mentioned. a few words concerning the experiment of the two French authors on the stomach of the beheaded criminal. Finally. on artificial stimulation. C. that secretory fibres for the stomach are present not only in the vagus but also in the sympathetic. exist. already noticed nections of the stomach and pancreas You how similar the nervous conhave turned out to be the . have. he called forth by excitation of the medulla. Finally. Moreover. Popielski succeeded also in isolating branches of the vagus which only If such inhibitory nerves inhibited and never called forth a secretion. and that the purely secretory nerves. fore. Why. easily believe that the how extremely I think I may now rest satisfied. he excited at the same time the antagonists of the secretory fibres. Dr. We have and cannot. almost as promptly as the chorda tympani expels saliva. The observations here given furnish an explanation of all the failures and difficulties which the earlier investigators of the innervation of the pancreas had to fight against. by a augmentation careful preparation of the nerves.I administering physostigmin to the animal. after all that has been said and to you. for example. thereauthors were able to set up a true secretory effect from excitation of the vagus forty minutes after the organ had been deprived of blood. In conclusion. did Heidenhain only obtain a positive result in a few experiments on excitation of the medulla oblongata ? To say nothing of the inhibitory effects of the operation . every respect a copy of the other. that the existence of secretory nerves to the stomach and pancreas will appear to you just as real and unquestioned as those for shown the salivary glands in the classic and universally known chorda tympani. cannot innervation of the one is in We doubt. of the secretion made its appearance. a strong contraction of the vessels and an alteration of cardiac activity. it is permissible to fill up gaps in our knowledge of one Consequently of the schemes of innervation by analogy from the other. call into play the activity of the organ without any latent period. the secretory fibres of the pancreas have been anatomically separated from the inhibitory. excluded that reflex inhibition extends also to the secretory centres for the pancreas. in addition to these special nerves. for example. some branches were discovered whose excitation caused a secretion without any latent period. of course.INHIBITORY NERVES OF THE PANCREAS. rather an the activity of smooth muscle. a drug which strongly excites But absolutely no inhibition. seen delicate the digestive glands are. It need hardly be said that. .
two nerves contained two We different classes of fibres. The special The specific qualities of nerve cells Analogy between the innervation mechanism of the salivary glands and that of the deeper-lying glands of digestion The exciting agencies of the nervous mechanism of the salivary Differences between the exciting their particular properties Discussion of the sham feeding agencies of the different salivary glands experiment Mechanical and chemical stimulation of the cavity of the Constituent parts of a complete innervation mechanism duty of the peripheral terminations of afferent nerves glands . in addition. already burdened with many duties. As you have learned in the last lecture. GENERAL SCHEME OF AN INNERVATION MECHANISM THE WORK OF THE NERVOUS APPARATUS OF THE SALIVARY GLANDS APPETITE. The vagus nerve. so far as the pancreas role. and also in part ha\e seen by direct experiment. This is a matter which cannot be doubted. proved itself to be an undoubted exciter of the gastric glands and of the But we must also assign to the sympathetic nerve a similar pancreas. we advanced important experimental evidence to show the existence of special inhibitory fibres to the glands. mouth has no effect on the gastric glands The experiment of Bidder and Schmidt relative for success in to psychic excitation of the gastric secretion Conditions appetite The passionate longing for food the this experiment alone brings on the secretory effect in the sham feeding experi- ment. and is saw good reason for believing that these also highly probable as regards the stomach. the nervous system can influence the work of our glands in the most diverse ways. GENTLEMEN. has. secretory and trophic. is concerned. . farther and have divided Heidenhain's trophic nerves into separate Lastly. a condition which had already been proved to exist by Hoidenhain for the nerves of the As a hypothesis we might even have proceeded a step salivary glands. classes of secretory fibres. THE FIRST AND MOST POTENT EXCITER OF THE GASTRIC SECRETION.LECTURE IV.
We obtained these results by division and artificial excitation of nerves which run to the glands. the effect of the stimulus will be conveyed through the centripetal nerves. their peripheral terminations. which come in from contiguous links of the nervous chain. it may be useful. respond to specific stimuli that is to say. these organs are only called into play by certain definite conditions. impulses. the normal receiving apparatus of the nervous system. as every here without distinction every outagency which has its seat within the applies here to everything'with the single excep^ . the centripetal nerves themselves. The utmost importance is to be attached to the fact that only the peripheral endings of centripetal (afferent) nerves. in contrast to nerve fibres themselves. function as normal sites for the application of external stimuli has still Consequently. it is not borne in mind with sufficient precision by the majority of medical men. and at the same time impart the utmost clearness to our representation. Consequently. The function is of the end organs with which they are . are able to transform definite kinds of external stimuli into nervous . connected therefore of a purposive nature in other words. as through a receiving wire. and. Here it becomes changed into a definite impulse and now comes back along the centrifugal nerves the outgoing wires. and these fibres also (W run in the vagus. in the intact organism these alone constitute impulse. all the more since this scheme is seldom completely followed out or adequately described in physiological text-books. and by what means these nerves are thrown into activity during the normal course of physiological events remains a question. Physiology now accepts it as a settled fact. as well " organism. In order to avoid repetition. the list of whose functions seems the almost interminable. how. the centrifugal (efferent) nerves. The word "external tion of the nervous system itself. Whether the ends of centrifugal (efferent) nerves are likewise able to peripheral to be answered. But when. the nerve cells (a group o? nerve cells connected with each other is " termed a " nerve centre ). when any external agency excites the the receiving stations of centripetal nerves peripheral terminations in this or that organ.AN INNERVATION MECHANISM. to the central station the nerve cells. that nerve fibres serve only as conductors of nervous impulses. and * By the term "external stimulus" I mean ward agency of nature. lastly.to bring before your minds at once the plan of innervation of a given organ. Only the peripheral endings of nerves and the nerve cells themselves have the power of transforming the external stimulus* into a nervous In other words. complete innervation mechanism consists of the peripheral endings A of the centripetal (afferent) nerves.
But. its The same communicated applies to the nerve cells : with specific sensibility. they respond. and other factors which throw this or that end apparatus I always look upon it as a period of scientific into an active condition. we shall only be able to form a true conception of the motive agencies of the whole complicated machine. to Irrespective of obviously they are endowed the excitations which are centripetal nerves. but only to special conditions which are different for the different portions of its length. and cannot therefore have any doubt regarding the specific nature of the end organs of other centripetal nerves. so every organ. at least in the earlier phases of their effects. and consequently presented to the mind To impart to the the physician. This is a sore point in present- day physiology. them from from various pharmacological data. as originators of nervous impulses. or other stimuli arising in the organism. excite or annul the activity of This follows not alone from a number of physiological facts but also Thus we learn that various drugs definite portions of the nervous This specific system. notwithstanding our knowledge of the separate parts of the animal body. We have long known that the peripheral endings of sensory nerves are possessed of a high degree of speciality. excitability of nerve cells. established the specific excitability of the end apparatus of every centripetal nerve. a more correct conception of this matter was my chief object physician in giving these lectures. Just as men and animals in the world are only able to maintain their existence and constantly adapt themselves to changing circumstances by aid of the peripheral endings of their sensory nerves. when we have chemical. indeed.64 THE WORK OF THE DIGESTIVE GLANDS. can only maintain its place in the animal microcosm. chemical. As the work of the digestive canal is now represented in the of majority of text-books. ceivable form of agency. and have discovered all the mechanical. impart the idea of being aware of their purpose. and adapt itself to the activity of innumerable associates. by virtue of the fact that the peripheral end-apparatus of centripetal nerves possesses a specific excitability. mere general excitability that is to say. are admitted to be indistinguishable. does not respond to every con. only or at least mainly to definite forms of mechanical. to furnish you with evidence that the alimentary canal is endowed not with sufficiently convincing. indeed every cell of every organ. of being conscious of their duty. as well as to the general life of the whole. it bears the impress of this period. I hope. inadequacy so long as the effects of the most diverse external agencies upon any normal physiological process. just as much as the same property of peripheral end organs. our next duty is to endeavour to discover the normal exciting . Hence. lies at the bottom of the purposive action of these organs.
Authors naturally expected to see a simple and prompt stimulation-effect sufficed for the salivary glands. In the working of the abdominal glands nervous inhibitory processes play a large part. But the attempts abdominal glands. or. have. as well ae the peripheral endings of the different nerves. advantageous. conclusions drawn from analogy too far. to find out that portion of the nervous mechanism which is for the time being under excitation. abdominal glands behave in some ways different from the salivary glands. It may now be instructive. more correctly. other conditions of experi- ment are necessary than those which held good for the former. and that the work of even the most apparently similar organs should be . for our further conclusions. and this circumstance has decidedly retarded the due development of our knowledge of the nervous control of the abdominal glands. for each phase of the work of secretion. This. but they are almost wholly absent in the case of the salivary This is an additional warning that one must never push the glands. the right to deny the existence of any extrinsic nervous influence upon the abdominal glands. from the same conditions of experiment which and the failure of this gave them. and. tions of nervous inhibition. as they thought. Gii conditions of the centripetal neives belonging to the glands which we had under consideration in our last lecture. The erior is now obvious the . whose in nervation has long ago been investi- gated. have done considerable harm to the usefulness of the analogy and have prevented our arriving at a correct idea of the plan of innervation of the of innervation been accepted analogy and thought of the nervous apparatus of the salivary glands. therefoi e. the mode of activity of the latter. The salivary glands. to glance shoitly at the nervous control of the salivary glands. medical science resorted to a bold as of investigators to apply rigidly to otheis the scheme which holds good for the salivary glands. shall also be We able facts is more fully to comprehend the inner mechanism underlying the which formed the subject of the second lectui-e. which form parts of the nervous apparatus of these glands. of course.SPECIFIC EXCITABILITY OF NERVE ENDINGS. to find out the conditions which excite the centres. but must constantly bear in mind that the life-functions of all organs are extremely complicated. and for their successful investigation. This would include an exact analysis of the stimulating influence which mastication and food exert upon the nervous mechanism of these glands. an ideal programme which we can only follow out as far as the present state of physiology permits. We have already had an example of this nature In the salivary glands we have no clearly marked indicabefore us. and to discover the primary We agency by which this condition is elicited. have generally types of the deeper-lying and when it became necessary to form a conception of digestive glands.
but are specifically excitable. especially since Dr. in the testing process to be useless. On the other hand. for example. dissolve the part of host to every substance and bulky with mucus in order to facilisoluble. everything brought into the mouth may reflexly influence these off' in the effect. if only in brief fashion. thus. as well as numerous experiments upon animals. envelop the hard tate its passage down the narrow oesophagus . a strong acid is to . The saliva is secreted in the compartment of the alimentary canal. their indications erroneously interpreted. food into the mouth. therefore. that the impression is the centripetal nerves of the digestive canal peripheral end-apparatus of The facts were here correctly observed. it nervous centre for the salivary glands.CG THE WORK OF THE DIGESTIVE GLANDS. taken in moisten the dry. drawn between the abdominal and salivary the unjustified analogy glands has to be credited with another important misapprehension. is sufficient to set the salivary glands at once the well-known expression. teach us that a number of substances. The experiences of daily life teach us from the outset. Nor is its duty by any means ended here. The saliva is secreted in the first instance to obviate injurious effects in some way. The great multiplicity of excitants of salivary secretion. glands. has without doubt. To me it appears that submitted to separate and careful observation. for food. the same everyday experience. Hence a psychic event. " to make one's mouth into activity indeed. precisely for this reason I think consideration. are likewise able to call One even acquires the impression that saliva. and it appears to me that it is precisely this which has driven the idea into the background. in a sense play the enters the alimentary canal. some connection with the complicated physiological functions of the saliva. must be accepted as an undoubted excitant of the longing even the thought of . the only difference being a gradual shading the stimulation which the substance dependent upon the strength of forth a secretion of introduced able to exert. which is at the same time Much of what enters the mouth the sorting-room of the organism. And it desirable to bring under the conditions of work of the salivary glands. the eager water. such as starch. and submit certain forms of food material." is based upon this fact. This is the first fluid encountered by everything which It must. or even noxious. that the of activity of the salivary glands begins even before the introduction the sight of food or With an empty stomach. and may prove must either have its deleterious properties neutralised or be comfirst pletely rejected. Glinski has instituted in the laboratory some easily performed experiments which bear upon the matter. to a process of chemical elaboration. when brought into contact with the mucous membrane of the mouth.
If you only think of cleansing how often we are impelled to expectorate. that a feeling of disall in physiology.substances may be simply diluted. that the salivary glands have each particular exciting agencies to which they specially respond. gust prod u s almost as strong a flow of saliva as the sight of a tasty meal. and not a trace taste. can be called forth tioned influence. for it can be supported by facts. Think how often. when enters the mouth. when the removed. the function of the saliva just menprobably the true physiological explanation of the feeling of disgust which many persons experience at the sight of the secretion by mental is itself. To say nothing of the testimony of earlier authors. while other corroding . even after the unpleasant substance has been for a con- more is apparent to the sense of afterwards one has only to recall the circumstances Indeed. the specific excitability of the peripheral endings of the salivary nerves is very comprehensive and widely extended. which. In both ca3e^ the secretion performs the oflice of forerunner: ( it prepares for the washing out of the mouth. Hence I hold that substances which obtain entry to the mouth set a secretion of saliva only because we have here the seat of a definite up physiological sense. as is well known. that is. the saliva plays the rule of injurious substances have to be wholly a washing-out fluid otherwise the . Glinski isolated the orifices of the salivary glands in dogs with portions of the adjoining mucous membrane. (17 a certain degree neutralised. in the second the requisite elaboration of the food. . and caused them to heal into the edges of the skin wounds. this will be clear. and not because the peripheral terminations of the buccal nerves are devoid of specific excitability. with what rapidity the siliva something disagreeable in the first for is poured out.THE SALIVARY GLANDS SPECIFICALLY EXCITABLE. Further. we can demonstrate the following facts from the material collected in our laboratory. In the second place. and capable of being thrown into action by every desired form of stimulus. material. i-i fluid. Such a view finds additional support when we reflect. long to mind in order to bring on anew the secretion of saliva. brought them out of the oral cavity. This is no picture of the imagination. Apparently siderable time removed. to wash out the mouth with s diva after something unpleasant. the psychic excitation of the nerves of salivary secretion also ushers in the act of vomiting. and by mere reduction of concentration have their harm fulness diminished. might in longer or shorter time gain entry into the blood and there develop its noxious influence. This last function of the fluid is hardly taken into account at and yet it evident that the saliva. by clinging to the mucous membrane of the mouth. Dr. In other words. as a must have a wide importance.
collect the saliva. but to our astonishment no collected in We f-aliva tempt the dog with a piece flows. either . whose parotid duct has in a similar manner been diverted outwards. such a comprehensive excitability of the produced. I now offer such an animal and. I respond that Dr. A attached to the funnel. animal the ducts of the submaxillary gland were thus led By means of a Mendeljeff's clip. hang on a new test-tube. Probably ) ou will say there is something wrong. however. But wait a little. One may employ a number substances in this way. to another dog. and obtain at once an abungive is dant secretion. Glinski has had an animal with double parotid and submaxillary fistula?. caused the saliva The results of this experiment permit us to to flow in large quantities. but on individual differences in the dogs. pieces of flesh to eat a piece of flesh. In the first place. and was able to observe on one and the same dog. a like behaviour on the part of the glands to that which Ave have just seen in two different An analogous experiment with bread was also carried out individuals. the plume result that of I obtain of a feather dipped in acid solution. on the other hand. which served to orifice. Glinski. the dog's mouth is opened and a pinch of fine sand thrown in. eager for the savoury meal offered. and now I apply to the buccal mucous Once more a new test-tube is . We now proceed. the innervation apparatus of the parotid manifests a very sharp selective power The mechanical in the choice. and give it a few once more a strong secretion of saliva results. so to speak. innei vation apparatus of the salivary glands that you might readily interpret it as meaning the power of response to all and sundry forms of You stimulation. was attached by a wire. new tube now membrane. by Dr. while dry bread. not on a different specific activity of the glands. first In his outwards. as you see. . and yet the animal of flesh. in the result Should any one happen to think that the variation dependent. Secondly. . as a matter of fact. the several salivary glands are. in respect to the conditions necessary for their activity agencies which excite their nervous mechanisms. draw extremely instructive conclusions. the wide end of a conical the funnel of waterproof material was attached to the skin surrounding To the narrow end a small test-tube. I stop tempting the dog. when a similar eft'ect is always see in this. of an adequate stimulus. to eat again the secretion of saliva Now we now with the method or with the glands of the animal. is most give it some raw flesh is as good as absent only when I come near can I detect one or two drops of saliva running down the sides of the tube. with the a strong flow of saliva. I the dog finely powdered dry flesh. The saliva is the same way. again there is a flow of saliva.C8 THE WORK OF THE DIGESTIVE GLANDS. . The eating of fresh moist bread produced no secretion worth mentioning. very sharply differentiated in the that is to say. the tube fills up at once with saliva.
THE WORK OF THE SALIVARY GLANDS APPETITE. and the intensity of the effect. who is at present carry- ing on the investigation of salivary secretion in our laboratory.y. indeed. when dry is all food (bread or powdered meat) is offered. The parotid meat offers fresh gland. however. as a result merely of the swallowing of food in the so-called sham When one takes into considerafeeding of an oasophagotomised dog. that of which we have just spoken. pointed and yet physiological opinion throughout the length and breadth of the land. tion the absolute independence of this factor. and manifestly important factor. it is true. I am quite convinced that an exact study of the exciting agencies of the three salivary glands will furnish a number of new data bearing upon the question in hand. if at all. At first it and when I previously what does it consist ? drew your attention to powdered flesh thrown into the mouth. has added a very interesting observation to the results of Dr. Glinski already related. and yet it was precisely to the latter that the glands responded. responds with a very active secretion. the exciting effective processes in gastric digestion. <i!> effect of large pieces of flesh is naturally much greater than that of the finely powdered material. in the emphatically that this is not the case. which prepare this juice. e. of the salivary glands an analogous phenomenon to indicate activity \Ve might apply not. We . Dr. but to some other property of the food. Wulfson. previous authors have already out that dry substances cause a specially free secretion of saliva. however. called With the first. events. which makes itself evident in the secretion of a large quantity of agency which brings about such secretion must be recognised as one of the most important and juice of high digestive power. you are already acquainted. similar to the reflex excitation. I assert quite have. has chosen to recognise a universal instead of a specific excitability. which is hardly. therefore. by finely Now. This phenomenon the more surprising since the desire of the animal for eating is much more strongly excited by flesh than by dry bread. have already I refer to the production of gastric juice in the empty stomach. excited when one to the animal. as expressed in text.. which has into play ? a relation thereto. The stimulus is. But in appears sight the fact I expressed the opinion that there is here a simple reflex effect from the cavity of the mouth upon the secretory nerves of the stomach. This other property Our example illustrates how obviously the dryness of the material. that which we may term "purposiveness" comes into play in the working is of our glands and albo how erroneous is the opinion that the mechanical stimulus is all potent.books. and. in the normal course of digestive canal is the gastric juice. not due to the mechanical. The second reagent which is poured out on the raw material in the How. Indeed. is the work of the gastric glands. of the parotid gland. seen.
with the same object.70 THK WORK OF THE DIGESTIVE GLANDS.. acid solution. makes a few chewing movements and The dog on which the acid experiment has just been made serves also for the swallowing of the stones. This play with the stones has now lasted fifteen or twenty minutes (in the laboratory we have often kept it up for hours). from the upper portion of oesophagus. The attendant now places some pebbles iri the front part of the mcmth. in most cases at the chemical least. way with a number local . but always with the same negative result. With can. the acid is. It simply and drop with an audible sound upon the table. the stomach. every conceivable form of stimulus which could possibly come into play in the act of ea.ting. and likewise observe. bitters. same path that the of other food takes in sham We and could experiment in the same saline. decoction. and then swallows them. from the oesophagus. One can easily train a clog to swallow etones which are placed in the anterior part of swallows them down. as if chewing and gnawing them. we may also combine a mechanical stimulus. as you see. In order to prove that the dog is perfectly healthy and normal. although the acid. employ the soluble constituents of flesh in the form of a excitation). the buccal cavity. You see that all the manipulations in this case are carried out with bare hands and without instrumental aid. when the animal rolls them round. We may even. we lay aside the stones and proceed to our old experiment of sham feeding. for example. I will try such an experifistula. the first drop of gastric juice makes its appearance precisely . however. and yet would not obtain the slightest indica- In this dog with a gastric tion of secretory activity in the stomach. and with also a divided oesophagus. As you see. passes along precisely the feeding. mustard. no sign of activity on the part of the gastric glands. passing them back behind the anterior pillars of the fauces and allowing them to fall out It may be added that a again. viz. in spite of continued excitation. may finally give such pieces of sponge. The secretion of saliva begins therefore. or even smooth We stones of considerable size. is swallowed and Hows out again from the upper segment of the oeso- From phagus that is to say. substances: pepper (strong so on. as you see .. a free secretion of saliva. effective. The stones fall out. to the dog to swallow. and yet not a drop of gastric juice is to be seen. well-taught dog puts up with all these procedures without the slightest protest. at once. using the most effective chemical stimulus to the buccal mucous membrane. mixed with the saliva. wipe out the mouth with a sponge soaked in We the solution to be experimented with. and always with the same results but perfect quiescence of the gastric glands. no secretion results. ment.
passing them in front of the dog's nose. of It has. the excitement of the chewing and swallowing produced centres does not imply simultaneous action of the secretory centre of nerves in . as you see. them in another. indeed. the excitement of a keen desire for it. the eager desire for food. thanks to the experiments Bidder and Schmidt. The stomach l. is it is obvious that the excitation <. is sufficient to shall presently cause a flow of gastric juice from the empty stomach. sham feeding In what. neither chemical nor mechanical stimulation of membrane is capable of refiexly exciting the neives Further. likewise having a gastric fistula with divided oesophagus. the buccal mucous of the stomach. in other words.e. and the feeling of satisfaction and contentment derived from its enjoyment. namely. the more certain and intense is the secretory effect. has cnlltd explanation. chop them up. and so on. one can easily observe that the keener and moie eager the desire on the part of the dog for the food. intrinsic to the sham feeding. At one time we even had a dog \\hich the voluntarily took the stones out of one's hand and swallowed them creature had seen our object in previous experiments ai d sagacious doubt that this and function in . and since then not a drop of gastric juice has escaped. of tlie fluid. Here I We bring before you another dog.GASTRIC SECRETION EXCITED BY APPETITE. been known for forty years. therefore. and this alone. chatters its teeth together.as been washed out half an hour ago. that at times. endeavours to get out of its cage and come to the food. does this influence consist which is the gastric glands. Precisely five minutes after first we began to tease the animal in this way the drops of gastric juice appear in the fistula. have occasion to observe the force of this physiological factor. by tens. The secretion grows ever stronger and stronger. Clearly. manifests the liveliest interest in our proceedings. stretches and distends itself.. till it flows in a considerable stream.f theee not the result of a stimulation coincidently that is to say. then. The animal. collected 71 and after a further five niinutrs \ve have more than in I . We begin to get ready a meal of flesh and sausage before the animal as if we meant to feed We take the pieces of flesh from one place. In extreme cases . c the experiment be frequently repeated. swallows saliva. If forth under our eyes a most intense activity of the gastric glands. and so on."> c. consequently there can be no dog both gistric glands and nerves are uninjured normal manner. the offering of food to a hungry dog. After the lapse of a few minutes we can count the number of cubic centimetres of this experiment is o clear as to require no the passionate longing for food. was negative. but which we have not been able to reproduce in our analytical investigation ? There is only one thing to think of. and lay it. The meaning . at the end of five minutes. learned to perform result it of its own accord ! But in tl is case also the.
Thirdly. again. be said that it received general recognition correct. minutes . is the dog had eaten. one may find among the dogs. and then the excitation of the dog with effect of the flesh was begun. 1" c. however. . and this the description... who could never convince themselves of There are authors and in many physio- By way of explanation. the observation of Bidder and Schmidt was perfectly It c innot.72 there the is THE WORK OF THE DIGESTIVE GLANDS. is dependent upon the intensity of the desire for eating. Firstly. was not the case. 10 3 Then followed a sham feeding for six minutes. we logical text-books it is not once mentioned.c. 10 10 10 . positively careless indifferent creatures. have a perfectly uninjured gastric mucous membrane. its reality. 8 10 clear that in this case the tempting instead of being less than the sham feeding. whether with its interest una wakened. 10 1<> 10 10 10 8 8 1!) 1!) . Secondly... the animal must be healthy and vigorous. in the case of many authors who obtained a negative result.s effect and that of an experiment of Professor Ssanozki. After six minutes the secretion commenced and continued as follows : sham Here Duration of the flow. and also upon what a dish that excites its desire or leaves incapable of being perturbed in this way by anything which has not . the success of the experiment. Consequently. and this.. just as men have.c.. must from dependent upon how freely and how long beforehand it is tempted with. on the contrary excelled it. or that it was sufficiently appreciated... i.. even a quantitative relationship between this feeding. A few threads of alkaline mucus had pared just escaped from the stomach. shall now consider how this matter must be dealt with by those who wish to observe the effect. 8 4 4 .. It is known that dogs have very different tahtes. Ib is effective in physiology.. Ib is only under certain conditions that ib it can be seen. 17 minutes ' . Quantity of the juice. 10 10 c. in which the secretory mere tempting of the animal with the sight of food is com with that of sham feeding. as stated above.
If. for instance. a very intense secre- . we have performed many of our experiments on sleeping animals. Fourthly. which even now is apparent. may become in the W e must constantly T fight. one has to reckon with the sense and cunning of the dog. to the keen desire on the part of the animal for food and the satisfaction of enjoying it. therefore. being teased with the food. and turn away offended at what is arrange matters as of If fed in reality. and at the same time see how constant and intense the action of this psychic impression is. this factor. having beforehand convinced ourselves by frequent repetition. so the animals were not going to be disappointed but attention be paid to these conditions the experiment if When one is secretion under different conditions. so to speak. and these investigations have confirmed the opinion at which we had arrived." as we usually term it. impressiongiven them.PSYCHIC EXCITATION OF GASTRIC GLANDS APPETITE. the going out of the room. that is to say. and we should not be far wrong if we said that ascribed in former investigations to the effect of in reality a result of unobserved psychic much which has been influence. every movement. has not eaten for a long time. able and excitable animals are necessary. " excitation of the gastric secretion. taken in hand a number of modifications of We the sham feeding experiment. being done before them. the dog has been prepared by a long fast of two to three days. that sleep exercises no restraining influence on the working of the gastric glands. trouble to arrive at a correct explanation of the mechanism of this have. we are forced to the inevitable conclusion. In view of the importance of the act of eating. we have spared neither time nor factor. eager. was Consequently. a factor which is not lightly to Often the animals perceive at once that they are only be disregarded. and patiently waiting till the food is Hence for success in the experiment. one becomes convinced of occupied for a length of time with the study of the gastric what a dangerous source of error this psychic excitability different experiments. every little triviality may give ri?e to excitation of the gastric glands. agency. in order to verify our own conclusions concerning the effects of this or that condition. such sources of this or that The minutest attention is necessary in order to avoid error. We must. that in our sham feeding experiment the whole secretory effect is due to the psychic stimulus. the appearance of the attendant who ordinarily feeds the animal against If the dog in a word. psychic will be found to be as reliable as the expciiment of sham feeding. keep it ever in view and guai^d against it. When we recall to mind the failure of our attempts to obtain a secretion of gastric juice by any stimulation whatever of the buccal mucous membrane. become annoyed thereat. therefore. 7:5 actually reached their mouths. but which will become still more obvious when the succeeding periods of secretion are investigated.
soup. the excitation of the nerves of the gastric glands depends upon a psychical factor which has here grown into a physiological one. which has not fasted. for naturally the idea does not occur to any of the dogs that all their trouble is in vain. for the sham feeding experiment. in pieces of the same size and at the same rate as before. no matter what may be given it to eat. that is to say. we find dogs which will devour bread with greater In these cases one obtains more and stronger appetite than flesh. Sometimes. With another dog which prefers boiled to raw meat. receives is it eats with greed before one's eyes. The secretion of juice has meanwhile either not begun at all. is just as much a matter of course. is given boiled meat which has been cut into pieces of and the piecfs follow each other at regular intervals The animal eats. develops no particular greed for the meal. in the sham feeding experiment. the more juice will be secreted and the greater the digestive power which it possesses. The raw meat tastes excellently to the dog. from its behaviour.74 THE WORK OF THE DIGESTIVE GLANDS. and refusing altogether a third. it The dog pleasant. tion of gastric juice will always be obtained by the sham feeding experiment. and correspondingly less juice will be produced by sham feeding with bread than with flesh. that is to say. The dog. and this observation is con: a dog the firmed by the fact that after fifteen to twenty minutes it ceases taking flesh. All the conditions which we enumerated above. Broth. you fee that time. and has therefore no reason to feel offended. however. or only after a longer interval than five minutes. under given conditions as any other physiological Regarded . however. corresponding amount and quality of the gastric juice will manifest wide variations. and remains scanty to the end. the food which not only imagines food but actually eats it. has been fed fifteen to twenty hours before. therewith. The majority of dogs prefer flesh to bread. for instance. it eats for hours at a time. bread or coagulated egg-white. either forthwith or next clay. and which are necessary for the successful production of the psychic effect. and. and appears quite as regularly result. milk towards which dogs are usually more indifferent than towards solid food often produce in sham feeding. hold good in combined form. Here is a case in juice in sham feeding with bread than with flesh. Now wait till the secretion has stopped and give the same dog raw flesh. the point of it definite size. the secretion of gastric juice begins precisely after five minutes and is very active. &c. will pick and choose amongst the different foods. by the act of ealing. Consequently. although broth. eating one with great greed. tolerating another. exactly the reverse occurs. either no secretion at all or only very little. but soon. has essentially the same taste as fle-h. The more eagerly the dog eats. It is therefore clear that in sham feeding the psychic effect may readily become an absolute and independent factor. whether boiled or raw flesh.
becomes here at length incorporated into flesh and blood. since they are only excited when the food has already entered the organism or at least has arrived very near it.the food is only found functions. which we have now carefully analysed. to secrete large quantities of the strongest juice. It is acquired not outside the organism in the surrounding world. desire. of is the first and strongest impulse towards the activity This especially applies to the two latter senses. such as judgment. It is by the establishment of the gastric glands. namely. of sight. We are therefore justified in him a large stock of gastric juice wherewith to begin the digestion of the meal. so full of mystery to science. a factor which mightiest embodies in itself a something capable of impelling the empty stomach of the dog in the sham feeding experiment. appetite. 75 the process may be said to be a com- Its complexity arises from this. saying that the appetite is the first and exciter of the secretory nerves of the stomach.RATIONALE UK THE PSYCHIC EFFECT APPETITE. from the purely physiological plicated reflex act. transformed from a subjective sensation into a concrete factor of the physiological laboratory. alone by the exercise of muscular force. means to secure this juice is also absent. of this passionate desire for eating that unerring and untiring nature has linked the seeking and finding of food. easily be predicated. stands in closest connection with an everyday phenomenon of human which life. that the ultimate object is attained by the joint working of many separate organic The material to be digested. will. smell and taste. of hearing. That this factor. A good appetite in eating is equivalent from the outset to a vigorous secretion of the strongest juice . with the commencement of the work of digestion. neous excitation of the different sense organs. where there is no appetite To restore appetite to a man. . but also by the intervention of Hence the simultahigher functions. side. may is so important to life and This agency.
endeavours care- may now venture to say explicitly. and consists in a passionate longing for food. a puff of sand. What position is to be assigned to the . calls into is This impulse activity the innervation apparatus of the gastric glands. On the last occasion we made ourselves acquainted with the first normal impulse which. both medical and lay. a and by rhythmic dilatation of an india-rubber ball Contact between the food and the stomach-wall may indirectly call the : excitation of the activity of the glands into play by awakening or increasing the desire for food.LECTURE V. the psychic juice is The psychic secretion cases. or by prescribing other remedies believed to promote its It is. PERIOD OF OCCURRENCE AND IMPORTANCE OF THE PSYCHIC OR APPETITE JUICE IN THE SECRETORY WORK OF THE STOMACHTHE INEFFICIENCY OF MECHANICAL STIMULATION OF THE NERVOUS APPARATUS OF THE GASTRIC GLANDS. mucous membrane by means of a glass rod. that which in everyday life. of secretory activity appears each time juice be avoided Demonstration of "appetite juice" in a dog with an The work of the gastric glands if appetite by introducing food through a gastric fistula unperceived by the animal Digestion of flesh by the stomach with and without isolated gastric cul-de-sac. in the natural course of events. and in the practice of the physician. by introducing the active constituent of gastric juice pepsin from without. GENTLEMEN. If the meal be subdivided and administered at intervals. however. a mental one. a fact which at once displays the pre-eminent importance of the JUICE. in the majority of on the part of the gastric glands. the normal commencement. Medical science endeavours to assist the debilitated stomach sensation. APPETITE is fully to promote. investigation still We farther." and which everybody. sham feeding Duration of the secretory influence of sham feeding After the cessation of the psychic effect. of interest to follow our experimental secretion. how is the secretory work of the stomach maintained? Experiments to prove the ineffectiveness of mechanical stimulation feather. is called "appetite.
We have only to regret that these answers come so late. "psychic" or "appetite-juice"'* in the course of normal gastric Is any definite role to be attributed to it? What course digestion? does gastric digestion take when it is absent? Fortunately to all these important questions satisfactory answers are forthcoming by experiment. Chigin) : Hour. . The following are the quantities and digestive recall to Let us memory how the capabilities of the first two hourly portions of juice after the adminis- tration of 200 grms. secretion of gastric juice proceeded after feeding with flesh or bread in the case of our dog with the isolated miniature stomach.APPETITE JUICE. of flesh or bread (experiments by Dr.
long. calls forth. we did not secretory curve. or at all events only very little. 2nd 12-4 have now begun the analytical examination of the variations of our But owing to the importance of the matter. 4'2 c.. which in sham feeding. the ordinary ration of flesh given to our dogs Thus we divided 400 grms. to bring them forward again in order that you may be better able to compare . Quantity of juice. something else to eat w Inch does not interest it to the same degree as flesh or bread. indicates that the two conditions are connected with the inge. Digestive power. If you ing when we take into consideration the effects of other foods. which deal with this matter I think it necessary. 1st Chigin). Hour. or quantity of juice (with flesh. which were administered at intervals to We turned an hour and a half. (Experiments by Privat decent Kotljar and Each time after the dog received its 1 00 grms.c. decline in the quantity of juice) which sets in soon after the taking of food.c.stion i. . for example. The following table shows the figures in question of Dr. of flesh we were able to detect a rise both in the quantity and in the digestive power of the juice. The dog was given 600 c.e. Lobassoff. them with the secretion after flesh and bread. as a rule. 3'57 2-03 mm. of food give the dog. confine ourselves to conclusions which might be drawn from earlier investigations.78 THK WORK OF THE DIGESTIVE GLANDS. with bread. Offer the animal for example. especially if it does not last milk. decrease of digestive power. We new forms of experiment for further proof.) : Half-hour periods. and the rapid flow rise of the commencement the already-mentioned initial You have already seen the figures absolutely fails to appear. however. into four equal parts. you will find the initial increase in quantity and strength of juice does not appear. with a transitory factor which soon passes away and gives Our explanation becomes still more convincplace to other conditions. of milk (experiment by Dr. no secretion.
therefore. minutes from the commencement of the feeding. In the curve which follows. another 3-0 was opened into the main portion of the organ. 123450789 5-0 10 LI 12 sham feeding as they occurred in the case of the dog with divided (esophagus. from both the large and small stomachs. We It endeavoured. The secretion began in both stomachs after the lapse of seven In five 172 grms. these were received back ngain at the orifice of the latter fistula. is connected with the act of taking in food. 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. feeding.). minutes the dog had eaten eighty pieces of flesh (weighing all of which soon afterwards dropped out at the fistula. PreFIG. covered with saliva. Lobassoff. Here is an instance taken from such an experiment performed by Dr. 11. taneously. and proceeded as follows : .APPETITE JUICE. after five minutes the juice began to flow simulcisely as in sham Curve of digestive power constructed from the foregoing table. to imitate the conditions of HALF-HOURLY PERIODS. at the beginning. lated In addition to the iso4-0 fistular orifice leading into the miniature If stomach. only the variations of digestive power are represented. It is clear that the increase. we now fed the dog in the ordinary way with small i-o pieces of flesh. both of digestive power and of juice volume. 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.
Here follows one of these experiments taken from Dr. It has here remained at the same height falling till the cessation of the secretion.sophago- torny on the dog. This experiment proves to us. . secretion from both cavities also The came to an end at the same time. and the intermediate variations of the secretion correspond in both cases. The further course of the secretion was as : follows Hour. without to the lower value which we observed from the beginning of the second hour onwards.is also confirmed later. first. so-called and Secondly.80 THE WORK OF THE DIGESTIVE GLANDS. after normal flesh feeding. The sixth for first drop of juice appeared from both cavities during the minute after commencing the feeding which was kept up half an hour. tion coincides in both. work in perfectly parallel manner with each other. that the main and miniature The stomachs. when we performed an le. the end. This w. beginning. and carried out sham feeding in typical form. the digestive power of the secreis the same which was observed in the sham feeding. Lobassoff's article.
-' 2-C. . 7-5 1-88 2-0 5-0 8th '.c. 81 waking animal. 13. 10th 0-2 I The secretion began twenty-five minutes after introducing now a&k you to compare the following tables : the food. Some kinds of f< od. 1st .c. 7-0 l-ss 5th 6th 7th . but the appearance of the juice is It begins from fifteen to forty-five minutes considerably retarded.snnill and large stomachs. . . is able to excite a secretion. 2nd 3rd 4th 10-0 'J'2 1-63 1-5 . an experiment by Dr. for instance bread and coagulated white of the hen's egg. of flesh were brought into the stomach. . . to 15 is Here and possesses a very low digestive power.c. is under normal circumstances extremely scanty during the first hour (o c. 5-3 3-0 . 5'6 6'6 2-25 . the stage.. .ith . . only the pi-ocess must be performed unnoticed.. the miniature stomach. This holds good both for the . . instead of 12 c. When a glass rod is introduced into the food contained in the organ it remains dry. to 5 c. 3'7 c. Curve of secretion from FIG. . 12.3 n . .. . and the animal's attention must be diverted from thoughts of food. 1:1 Flesh. 2'0 mm.c. .c. do not yield a single drop of juice during the first hour or more afterwards.APPETITE JUICE. and do not in any way resemble the secretion after normal feeding. . Digestive power. The results of this experiment are striking. c. after the feeding. The same from main stomach reduced ten times. . when directly introduced into the stomach.). Lobassoff : 400 grrus. Quantity of juice. . if introduced at this n FIG. . Hour. instead of from six to ten.
. Hour.THE WORK OF THE DIGESTIVE GLANDS.
however add. appetite... Lobassoff in analogous Into the dog's stomach 25 pieces of flesh (100 grms. represents the digestive value of the passage of food through the mouth. 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 and this. it is On not well adapted for class demonstration. Since then an hour and a half have stomach of the elapsed. I give also a series of figures obtained experiments. Each dog received 100 grms. without the assistance of appetite. the free end of which was fastened to the fistular cannula by inserting a cork. the animal being afterwards left alone. is very different in the two cases.. but during the process. The morsels were threaded on a string. pieces of flesh 83 were introduced through the fistula. in the one case and 15 per cent. Without sham feeding. the one hand. The dog was then brought into a separate like number of pieces was introduced into the room and left to itself. it is better before only on dogs accustomed to appear an audience. and may often fail. was digested. Without this longing. and of whose temperament the experimenter is well assured. the flesh remained an hour and a half stomach. that from the nature of this experiment.. with five minutes' sham feeding 15 per . many forms of food-stuff which gain entry to the stomach remain wholly devoid of . was reduced by 30 grms. : was digested. according to our former oasophagus. Once more the flesh remained five hours in the stomach without sham feeding 58 per with sham feeding 85 per cent. 31'6 per cent. in the Again out cent. to carry out this experiment in the lecture theatre. by Dr. it is not at all easy to conceal the introduction of the flesh ings of the less from the dog on the other. means the same thing to the eager desire for focd. the unusual and distracting surroundanimal often cause a short period of sham feeding to have In order to avoid such effect than would otherwise pertain. . or.. In that of the dog without sham feeding.) were The flesh remained two hours in the cavity. G'5 per cent. I must. and consequently the amount of flesh digested. the value of an eager desire for food the value of an . therefore. while the flesh withdrawn from the stomach of the other dog weighs Thi*. A other dog in the same way. a vigorous sham feeding was kept up. cent. in the other. with eight minutes' of the quantity sham feeding. of flesh. the loss of weight amounts to merely G grms. only 70 grms. and now we may draw the pieces of flesh out by mean!- of the thread and weigh them.ANALYSIS OF CURVE OF GASTRIC SECRETION. the balance of undigested food being 42 per cent. brought. in other words experiences. failures. with- sham feeding : 5'G per cent. was digested. that is to say. The loss of weight.
.. 7-2 7-2 C.. ICO 7-4 13-5 ll'O 7-0 . other questions..-.... . is true... why does bread brought unnoticed prehensive manner. . . Here. have especially in the case of animals. . The dog had a gastric fistula and also an opening leading into the esophagus. . . . however... and more powerful. with all the force and reality of a hunger sensation the effective agency.84 THE WORK OF THE DIGESTIVE GLANDS. . 10 10 10 10 10 10 20 20 20 20 20 30 20 .... but the juice poured when we have still more fully recognised the conwhich the secretory work of the gastric glands depends. is not of long duration ? determined many times. . 8-0 <i-8 7-r> . In normal feeding.. . we must consider . <>:> 7-r.. . Naturally the same holds good for the taking of food in the normal way.c. .. .. continue to last ? appetite juice continue to flow after the normal act of eating.. is gastric juice.. 25-5 20-0 13-5 c.. which. How long does the after-effect. ditions tolerably soon (after twenty to forty minutes) provokes this act? This will be explained in the next lecture now. . 8-1 .. 5-5 . 7-2 . the feeling of satisfaction which. it scanty and weak. . .. 8'1 nun. sets in long before the termination of the digestive period .. bear in mind that in sham not satisfied more and prolonged feeding... however.. ... as is well quelling known. After a sham feeding of five minutes the secretion began. is an experiment from the article of Professor Ssanozki which deals with the point. Time in and was continued as follows Digestive power. . ll-o . becomes the eager desire for food more accentuated. not only on our dog with the isolated already We stomach. the echo of the first impulse to the How long does secretory nerves of the stomach. .. feeding is how long the after-effect of sham continued.. excite a secretion. . therefore. while flesh It id only later. .-S .. .. . . . the of the longing. but also on other animals. even after a short period of sham feeding. . ... into the stomach of the dog cause no secretion for hours... and therefore the secretory influence is .. stretches over a length of time. The effect. . Quantity.. out Others. for example..... One must.. .. 11-0 8-5 6-5 ..... : minutes. ll-o r>-5 .. .... however. .. upon that we shall be able to grasp the meaning of these facts in a more comFor instance.
85 filling and distension of the stomach. my opinion. And but it does not happen in the simple. so that we might together bring the matter before the tribunal of fact. then. consequently. the effect of the forced feeding of phthisical and insane How. set forth in many text-books of physiology. When that bread or boiled white of egg. direct physiologists and physicians. either with an unbelieving shake of the head or else with a direct avowal that " it cannot be so. that mechanical stimulation of the stomach wall by food constitutes a reliable and effective means of calling forth the introduce This assertion. To this matter I It is on this ground. which. and the artificial feeding of those with gastric fistula. according to attribute very great importance.(performed " I will on account of stricture of the oesophagus) to be explained ? of my hearers may have " is my answer by a very unexpected pronouncement relative to the asseition. is nothing else than a sad My own statemisconception degenerated into a stubborn prejudice. introduced directly into the stomach.DURATION OF THE SHAM FEEDING EFFECT. even under the most favourable circumstances. therefore. therefore. If once the defenders of the old opinion are inefficiency of left for driven from their position. seek for other exciting agencies of the innervation must. and at the meetings of ment. to the demonstration of which we will now proceed. apparatus of the gastric glands. for the moat part. repeated various medical societies. call forth does not We the secretion instituted by psychic influence first occur to all your minds is naturally the immediate influence which the food exerts upon the walls of the Why and wherefore is maintained? stomach. in the case of certain kinds and quantities of food. must diminish the desire for food. and which consequently has gained hold of the mind of the physician. and. and obliged to admit the mechanical stimulation. patients. which is so categorically secretory work of the glands. is dependent on the factors which we have up to the present investigated." I regret exceedingly that these steadfast unbelievers are not here. This is all the more obvious since a sham of five minutes. What would this is true. feeding a secretion for longer than three to four hours. improbable that the whole secretory process in the stomach. that this dictum is only a picture of the imagination. probably many fashion current in the minds of I said many asked themselves with natural astonishment. there would be nothing further them . bring the secretory effect to an It is. from the mere end. in many published articles. that the whole battle must be fought out between the generally accepted view. and the theory that it is only specific and selected stimuli. thac every agency is capable of exciting the excitable by gasti-ic mucous membrane. may not for hours produce a trace of secretion. lasts from ten to twelve hours. has met.
looks as clear and transparent as water. have assumed a distinct blue tinge. and you see that. and. and you see that they pro- The dog takes the food exactly five duce bright red specks on the blue sheet. After failing with the mechanical stimulation. that. the gastric glands react to it in a perfectly normal fashion. The blue pieces. followed by others faster and faster. alternately. manner. minutes after beginning the feeding. Here is a dog with a gastric fistula on which a I open the it cervical resophagotomy has in addition been performed. than to build up new theories and search out old facts concerning gland work which have hitherto been rigidly kept in the shade. are the streams ! 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. we proceed forthwith to the sham feeding of the same animal. traditional. I will now repeat the experiment of mechanical stimulation of the gastric and classic mucous membrane before you in the well-known.books What can be set aside before your eyes. of juice. the first drops of juice I catch appear from the stomach. juice of we have not been able to find a single spot possessing a of pure gastric noticeable acid reaction. We was washed take the celebrated feather and . fistular orifice have merely been made wet without altering their colour. changing five minutes. fistula as you see. notwithstanding that it appeared so thoroughly reliable and convincing. moreover. Folds of blotting-paper saturated I now ask feather and glass rod. therefore. whose gastric glands from some possible cause. Consequently. You have all seen. offered it with keen appetite. however. caused by the moisture of the alkaline mucous membrane. From the every not even a single drop has escaped. Where. with red and blue tincture of litmus are placed at hand. We cannot. This single objection which we read in text. from one to the other my assistant to continuously move the in all possible directions in the stomach.WORK OF THE DIGESTIVE GLANDS. After thirty minutes' sham feeding we have collected 150 c. has been so little taken into consideration. are unable to react normally. with the most thorough mechanical stimulation of the whole cavity of the stomach. On removal from the stomach each is carefully dried with red and blue blotting paper. then. that Bidder and Schmidt's experiment. We may take it that it is mainly because people were so seized with the belief in the direct and simple mechanical explanation. the drops of moisture. of the excitation of gastric secretion by mental effect. gentlemen. on all the pieces of red blotting-paper I have been able to hand to you. a couple of drops on the blue litmus paper.c. possibly doubt is when the proper stimulus used. which. also a tolerably strong glass rod. that this procedure has now been kept up for half an hour. nothing flows out of the stomach : out clean with water an hour ago. without filtering. .
therefore. by setting up stomach. was placed in no little doubt as to whether his results (O-. howthat this lecture certain very obvious rules are have not observed. is proved by this. contractions of the not surprising that the glass tube. and his assistant at the time solutions. venture to think will quit the field. This apparently simple ever. 87 From this it irrefutably follows. that the mucous membrane of the stomach. obtained a secretion from mechanical stimulation pure and simple. is perfectly indifferent to mechanical excitation. stomach was emptied by removing the stopper from the fistular cannula. when determining the acidity of the juice first obtained from the resected stomach. but it was not washed out till an acid reaction was no longer given. It is only necessary to call to mind that Heidenhain. experiment from now onwards and give place to the one I have just shown you. was the means of expressing small quantities of acid fluid from the fistula-tube. it is necessary that the stimulus.) That matters are as I state. really we may adduce the fact that none of them made mention of the To overlook constant and precise period of five minutes' latency. stomach should be clean. It is very necessary that the gastric glands be not already in activity at the . completely emptied and imperfectly washed-out organ. and that nothing shall gain entry to it from It is true the Such conditions were not formerly fulfilled. Of no less importance is the second condition when we wish to perform the experiment of mechanical stimulation in the correct way. First. saliva from the cavity of the mouth could gain entry. (The relationship between mechanical stimulation and the motor functions of the stomach is not to be confounded with what we are here speaking of. however. viz. experiment of mechanical stimulation can. namely. and folds of consequently preformed gastric juice was left behind between the the mucous membrane. that only one explanation is to be found for the negative result in the first half of our experiment. probably on These. furnishing a healthy gastric juice. this was not possible if a genuine excitation of the glands had been obtained.INDIFFERENCE TO MECHANICAL STIMULATION. without.J to ()(!) were correct. physiologists account of a preconceived belief in the effectiveness of the mechanical when These rules are two. and that the facts correspond to the explanation. At the same time. that nobody till now has obtained genuinely pure gastric juice of an acidity to 0-5 or O'G per cent. which quickly became acidified in the in- It is. (Gscheidlen) was set to verify the correctness of his standard amounting The " acidity of the purest" juice known at that time was scarcely O3 As a further proof that none of the older observers ever per cent.. And yet this mechanical stimulus is demonI strated as an exciting agency in the physiological lecture-theatre. only be successfully performed followed. so far as secretory activity goes.
lesophagotomy have been performed. which I have already dwelt upon. And yet we are here dealing with a strong and widely diffused stimulus. both of our conditions have been fulfilled on the dog before you. could excite the glands to secretion. or that the hands of the attendant who has prepared the food should smell of it. Look for a moment at the performance of the bellows outside the stomach. sand. bring their glands to rest. with regard to mechanical stimulation. As you have just seen. that in order to obtain results. will be made answerable for the excitation of the gastric glands. diameter) at its rounded end. with which an india- rubber pump can be connected and a blast By rhythmic compression of the india-rubber . possible objection I will now show you two a similar dog is used. It is only necessary that some food be near the dog. Nobody has as yet said. On the contrary. Into the fistula I bring a thick glass tube containing a number of small openThe other end ings (2 to 3 mm. sand falls out again between the side of the cannula and the glass tube. out clean and is at present in a state of complete rest. and the glass tube. Again. and the result of the experiment stands in irreconcilable contradiction to those of the laboratory and lecture experi- ment of former times. and also that during the experiment no impulse comes into play. which of itself. apart from mechanical excitaNor have we any proof that tion. one on But in order to meet this new modifications. but in no case is it able to turn blue litmus red. which both gastrotomy and The stomach has been washed of the tube is connected with a glass ball containing tolerably coarse Leading into the ball is a second tube.88 THE WORK OF THE DIGESTIVE GLANDS. the mechanical agency must simultaneously come into contact with numerous points of the inner surface of the stomach. observers formerly waited for hours before commencing the experiment. or at least it is necessary to wait for The experimenter must strain his whole attention to preserve such an experiment free from objection. and this play is kept up for ten to The fifteen minutes nevertheless. that is to say. 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. or that some other similar circumstance should come into play. beginning of the experiment. quite undeservedly. and convinced themselves that the And some sible to dogs are so easily excited in this way that it is almost impos- hours. From every opening of the tube . The importance of the experiment. and it is either dry or scarcely moistened. we see no trace of gastric juice. of sand ball I inject blown through. gastric glands had ceased working. sand with considerable force into the stomach. justifies me in making still further demands upon your attention in order to show you two modifications of it.
when the operation has over a year folds of mucous membrane are often lasted a long time formed in the neighbourhood of its inner orifice. The surface of And here also subsequent sham feeding shows that the dog is in a suitable condition I must add that in making this observation the for the experiment. and yet not a drop flows through the tube. and if any of our luethods for the study of gastric secretion are reliable. fistula. that is to say. If one dispassionately regards this question. our experiment is ended. Moreover. it is a daily occurrence to find in the stomach of the dog. During the experiment a glass or india-rubber tube is brought sufficiently far into the cul-de-sac to catch the juice. and yet their presence in no way hinders the arrest of the secretion. Such an occurrence would have been specially obvious in our dog with the isolated stomach. of the uselessness of mechanical stimulation. when that the grains of sand strike with considerable force. and yet the manipulation of itself never calls forth a passed Further. How could this be the case if the mechanical stimulus were effective. you feel quite distinctly. Into its empty and resting stomach an india-rubber ball is introduced. In the case of dogs with an ordinary gastric special reason. And now. so long as true secretory conditions are absent. in easy and unquestionable fashion.INDIFFERENCE TO MECHANICAL STIMULATION. must have been fed within dog must ten to twelve hours before. The same holds good continuously in contact with for the dog with the tube has tolerably often to be taken out and set light. we may convince ourselves by sham feeding. thick rolls of hair. one must be convinced step by step in the laboratory. which occurs when digestion has ceased. not a drop of gastric juice ever and failing some escapes from the stomach other than during the digestive period. *!) ejected. secretion. which completely close In these cases a long thick perforated metal tube has to be the tube. Yet another experiment on a similar dog. that the innervation of the dog's stomach is perfectly normal. afterwards being allowed to collapse. since it was bedded with sawdust in order to guard Very often against maceration of the wound by juice trickling out. numbering considerably more than ten a strong stream of sand is If you hold your hand against it. the ball taken out of the organ is everywhere alkaline. otherwise a psychic excitation of the secretion can readily be induced. During this time not a single drop of juice has appeared from the stomach. in deeply. . nor does its inner surface ever become acid. In the ordinary gastric fistula in dogs. is kept up for ten to fifteen minutes. since the inner rim of the fistula-tube ? is the gastric mucous membrane resected stomach. This is distended with air by means of a syringe till it is as large as a child's head and main- The procedure tained in this condition for a time. not be too hungry.
to set the neuro-secretory apparatus of the stomach into activity. We may therefore take it that through the mouth and throat produces less enjoyment and excites. certainly acted as mechanical stimuli. In other words. it of general hunger. gastric glands. the other hand. very weak. even treat of in articles which secretion. and especially of satisfaction from the enjoyment food. that by direct mechanical stimulation. but also by impulses awakened by the passage of the food along the deeper portions of the oesophagus and by its entry into the stomach. the dog had licked the wound from weight. is referred solely to the stomach. especially when the latter happened to be empty. There are. a denied its influence. and give it a subordinate position in the series but as yet I know of no other physiologist who has wholly agencies. They obtain their sensations of of general well-being. can in the end distinctly perceive sensations and even call them up to consciousness. is its entry into the stomach devoid of the secretory process? It can hardly be is connection with doubted that. food which merely passes is by other sensations and impressions. connected with the matter of food with the gastric we will take into consideration a question we have just discussed. which in other people are normally masked caused not only by stimulation of the mouth and throat. . however. and function in some text-books. fcow they were able to follow a special tit-bit. The feeling On a mouthful of a favourite wine. not to be very mechanical stimulation. which to its nose. never series of facts ought to suffice to carry the to me that this adherent sawdust. as a rule. in the This sensation to say. obviously it then swallowed. Since the contact mucous membrane has no direct influence on all the secretion. under normal conditions. we found enormous half a pound as much as quantities of sawdust in the stomach. and the majority of people become accustomed to pay no heed to normal course of digestion. which yet It caused a secretion. all of us have met with men who could describe or exactly and with gusto. without taking cognisance of the factors contributing to them. yea. To conclude this lecture. its surface has a is. And appears long one is able supposition to its grave. the stomach is the seat of certain definite sensations. as exciters of the specially gastric few physiologists who hold in relation to gastric secretion. the whole way through the oesophagus down to the stomach. Naturally the gourmand. And yet the feather and the glass tube continue the even tenor of their ways to this moment. of exciting effective. who directs his attention continuously to the act of eating. that certain degree of tactile sensibility. and who has not held it least possible to obtain at some juice by it. the satisfaction derived from eating. together with that sticking these particles of sawdust of themselves. it is true.THE WORK OF THE DIGESTIVE GLANDS.
as they express it. that which is called " "appetite and not the latent need of the organism for nourishment. 1H therefore. In this way one can also conceive how the appetite becomes lost in cases of long-continued obstruction of the The patients forget their stomachs. together with a normal healthy For this reason it is easy to feeling in all parts of the digestive tract. owing to some keenly It is interesting occupation. . Perfectly well. a very complex sensation. the initial impulse towards in the awakening an appetite may originate in the stomach and not buccal cavity. to the production of an appetite to a How often does it happen that the passionate craving for food. the eager craving after food. the . does not lead immediately. may suddenly bring back the appetite. and dislike the idea of eating because the food. of entry of food. not the least desii-e for food is felt ? known first sets to everybody. appears to into a strange empty sack. this complete indifference Fearing that I should collapse. indeed it has become a proverb. and often not merely the need of the organism for food material is necessary for its excitement. I had lost all desire for food. people with gastric anaesthesia suffered of from this loss of appetite. no desire for food. but also a condition of thorough well-being. that real appetite in with eating. we naturally being meant the passionate and conscious longing for food.iNGESTION OF FOOD INCREASES THE APPETITE. recovered. ordinary hour for a meal has struck. indeed. moment perceived the onset of a strong appetite. As from a further illustration. and yet. or in other words. When we spoke above of the desire for food the excitant of the secretory nerves of the stomach. the creation of a necessity for food. and who have no feeling of appetite. instances direct introduction of food into the organ. I my own but transient be permitted to give an instance After an illness with which a personal experience. nor in all cases. If this be true. is. even when they Cases are known to neuropathologists where are no longer present. I felt it and the literally at that quite distinctly pass along the oesophagus into the stomach. Such patients are no longer conscious fall having stomachs. than the food which passes the whole way into the stomach. I resolved on the second or third day to endeavour to create an appetite by swallowing a mouthful of wine. I only differed from others in that I could with ease abstain from all food. remember the sensations. although otherwise fully high may in There was something curious towards eating. fever was associated. after an operation. understand how patients who have diseased sensations in these organs. The appetite. known that withholding food from the organism. and in such alimentary tube. whether consciously or unconsciously. a less feeling of appetite. is capable of It is This observation teaches that the tactile sensation of the stomach at moment awakening or increasing the appetite.
the its mechanical stimulus came to be considered this point it effective. lead to a reconciliation between assertion concerning the inefliciency of the mechanical stimulus anil belief. tlie distension by the food mass. but indirectly. the juice. Viewed from my may. be submitted to experimental proof. good example which enables us to differentiate is furnished by our dogs with sham feeding. It is therefore quite possible that in the case of some dogs. I hope that the foregoing will in no way lead to a confusion of ideas in your minds. . necessity for food exists in such cases. which has not yet been transformed into a concrete passionate desire.92 THE WORK OF THE DIGESTIVE GLANDS. to a certain degree. may give the impulse which appetite. in the old experiment. after it has firttt awakened and enlivened the idea of food in call into the generally prevailing excitation will at times I further also admit that mechanical the dog's consciousness. and thereby called forth the passionate desire. For such. not however directly by means of a simple physiological leflex. bears more or less of a hypothetical character. however. of course. only begins to flow as soon as this need has taken between these two factors The the form of a passionate longing. it is only necessary to com pa re the influence which sham feeding exercises in an oesophagotomised dog with that in one having a simple gastric fistula. which previous simple explanation of the facts. even before the experiment . but will assist you to an exact and concrete analysis of the This representation. the touching of the gastric mucous membrane with any object at hand. and when the appetite is awakened the juice This is possibly a third reason why. excites flows. its mechanical excitation. could. and at a certain stage of hunger. A lack of nutrition. play the work of the gastric glands.
The investigations of Blondlot and of Heidenhain on the secretory work of the stomach. and of hydrochloric acid on the gastric glands. In the last lecture its we had is . are unable to call forth a direct secretion of gastric juice. secretion (2) that the mechanical properties of the food in themselves GENTLEMEN. when a second fistula was opened in the main stomach. and solutions of Liebig's Extract exciting agencies Neither starch nor fat is able to call forth a gastric secretion Chemical excitants are also produced in the tive as excitants reliable are augmenting peptic digestion of proteids Starch influences the quality of the juice by its content of pepsin. since it conand is is tains fewer sources of error less troublesome for the observer. The introduction of the sound unpleasant to the animal. notwithstanding importance. THE CHEMICAL STIMULI OF THE NERVOUS METHE CHANISM OF THE GASTRIC GLANDS MINIATURE STOMACH A RELIABLE METHOD OF COMPARISON SEAT OF ACTION OF THE CHEMICAL STIMULIHISTORICAL. : Water as an excitant of the gastric glands The effects of watery solutions of the ash of flesh. both from a quantitative and from a qualitative point of view The miniature cul-dc-sac furnishes a true picture of the work of the large stomach. meat juice. settled (1) that psychic excinot the only source of gastric tation. The fluid substances to be tested were at first passed into the stomach by means of a sound. In order to answer the question as to what circumstances within the cavity of the stomach may act as stimuli to secretion. and may . of sodium chloride. we must turn to the chemical properties of the food. Solutions of egg-white are ineffec- Meat broth. they were introduced directly through it. The chemical excitants of the nervous mechanism of the gastric glands have their seat of action at the surface of the mucous membrane.LECTURE VI. of soda. Later. Our experiments on this point have for the :uust part been performed on dogs with isolated miniature stomachs. Fat inhibits the work of the gastric glands. Obviously the latter method is incomparably the better.
of water be injected into the large stomach of a dog with the isolated pouch. owing to bility For instance. in the case of our to of two stomachs. spite of drops is. The constancy of the result. for instance. tion. as soon as water was introduced into the main cavity.04 influence THE WORK OF THE DIGESTIVE GLANDS. in may in certain ways affect the work Further. as water was poured into the stomach. both earlier and later experiments at hand which remove every doubt concerning the stimulating influence of water. or in some In passing the t-ound.c. We have. very often that is to say. From a that it long series of experiments has. It may be employed in this way. I have myself always obtained a secretion from the introduction of water in dogs which I succeeded in keeping healthy for several months after the vagi nerves were divided in the neck. and these the glands. however. It is only a prolonged and widely spread contact of the water with the gastric mucous mem- brane. All this by the fistula leading into the large stomach. howtion of gastric juice as the result of sham feeding. the secretory process either by causing pain. and especially the regularity of the quantity of juice secreted. clearly indicated that no accidental condition. from its wide occurrence. up. I wish further to emphasise the fnct that section of . if only a weak one. without waking it. in dogs whose the division of the vagus nerve fibres.c. Dr.c. Heidenhain had cul-de-sac. only 100 c. dog with the water into the larger we always obtained a secretion of gastric juice (Dr. a few drops of the liquid injected not infrequently fell on the mucous membrane of the mouth. Jiirgens never saw a secrevagi As soon. Moreover. the most important constituent Has water an exciting effect upon the gastric glands? of the food. may have awakened the idea of food in the dog's mind. we introduced 400 cavity. The same phenomenon was likewise observed In such a case the possiat a later period by Professor Ssanozki. though not a large one. ever. of cannot be denied. Chit/in). avoided substances. we have 500 c. but also more consistent of course. isolated long ago shown that a secretion began from the gastric by his method. of a j sychic excitation of the secretion is excluded. Before passing. if instead of 500 c. such as a mental effect. The substances may even be introduced when the animal is asleep. Hence water must be accepted as a chemical excitant of gastric secreThus.. came into play. were cut below the diaphragm. arrived at the conviction When. Finally. to 1 50 e. in about half the cases not the least trace of secretion is to be seen. which. which gives a constant and positive result. vomiting movements may be set it other way. an undoubted secretion occurred. not only fluids.c. during the withdrawal of the sound. every precaution. was natural to commence the investigation with water the simplest and.
for example. we have a keen desire for eating. itself. lies in this that in cases where. their watery solutions had precisely the same effect as water To sodium bicarbonate an inhibitory influence must be ascribed. I believe. tion caused For. AN EXCITANT OF GA8TRIC SECRETION. and the instinct for it thirst is even more pressing and persistent than the desire for solid food. a little mucus escaped. when brought in quantities of 50 c. upon them. If a dry meal be eaten without appetite. Thus. take the place of the sympa- of the vagi in have. into the large stomach. which are constituents of the food or are employed in the practice of medicine. were able At most to expel even a single drop of juice from the small cavity. We But all probability different physiological duties to perform. chloride and bicarbonate of sodium. remains without finding a use. why The chief reason. does water act as an exciter ? The fluid needs no digestive juice. with an empty stomach. before us the transmitting the psychic impulses. thirst will compel one to drink water afterwards. Water is very widely distributed in nature. strength). testing the effect of must be kept in view when we are any other substance upon the gastric glands. explanation. when. and this fluid suffices to ensure the beginning and continuation is of no and cannot weigh as a serious objection against our consequence. After water. seen. a number of different inorganic substances.secreted juice when water alone is drunk. but are unable to do so for some reason or by water is not of itself other. cised the least influence with the exception of bicarbonate of sodium. nor could the secretory fibres whose existence is almost certain. were also tested. that secretory fibres which course in different nerves interesting fact. even the free flow of psychic juice may at times be secreted when there is no use for it for example.WATER. We must always compare the results produced by a watery solution of the influence of water The stimulating given substance with the effects of a like quantity of water alone. of the secretory at times. the constituents of meat ash. the secre- important and secondly. therefore. work of the glands. That the . 95 the vagi nerves. It appeared that not one of these substances. Chigin). But this does not make us doubt the great physiological import- ance of the psychic juice. have in : no psychic juice is present. does not in the least prevent the stimulating effect of water thetic. which suppresses all psychic influence upon the gastric glands.c. as we have already . Hence the presence of sodium bicarbonate 1 . exeron the secretory work of the stomach that is to say. and hydrochloric acid were repeatedly brought under investigation (Dr. Not one of the soda solutions (varying from 0*05 to 1 per cent. till we were fully satisfied of the certainty and accuracy of the results. in the fiivt place. the impulse to the secretory work of the stomach may be given by means of water.
of serious consideration. obtained when we injected a peptone product manufactured by the firm of Chapoteau. on further investigation. bodies. that if the gastric were specially adapted to act on the proteids. gave constant effects.. although it calls forth fact. is able to influence the output of nitrogen in the urine. would not reply. Nencki's laboratory. Naturally. and kindly communicated to us. it appeared of special interest to study the viz. when investigating the way in which egg-white. so-called food-stuffs the carbohydrates. return to them effects of the Next. led . however. later.96 THE WORK OF THE DIGESTIVE GLANDS. that the experiment with egg-white was repeated of its reliability could exist. We will. we obtained no greater secretion than a This seemed to us so peculiar. and only to the smallest extent of peptone. obtained by the action of strong and pure gastric juice on raw fibrin. however." positive result from chemical excitation of the gastric A mucous was. in which a preparation obtained repeated. How astonished were we then to find. meat broth. the peptone solutions acted wholly negative results that Dr. similar volume of water had caused. is Petersburg was employed. that the peptone of Chapoteau contained as much as merely as water acts. these substances juice It was would also prove to be chemical stimuli of the mucous membrane of the stomach. which it sets up. of pure peptone. It would be difficult to find either who. however. either pure or diluted with an equal volume of water. to be erroneous effects. if asked what stomach by means altogether a wholly unexpected a physiologist or a physician to egg-albumen when introduced into the happens " of the sound. that when fluid egg- white was introduced into the stomach of our dog. meat juice. On the other hand. Chigin to infer that peptone must be the chemical stimulant of the neuro-glandular apparatus of the stomach which we sought. These facts are worthy on account of their clinical interest as well as on physiological grounds. 50 per cent. A comparison of these chemical results with the physiological myself and Dr. till no doubt whatever The result was also corroborated in the laboratory by Professor Rjasanzeff at a later period. no digestive work. proved. The experiments with this membrane preparation always gave a marked secretory effect every time they were Other experiments. Dzierzgowski analysed both preparations for a special purpose in Professor V. it is is This digested by the secretion of gastric juice. both prevents the stimulating properties of the water. and solutions of meat extract proved to be constant and active exciters of . the fats. and the proteid to be expected. from Stoll and Schmidt in St. This belief. furnished to say. from a priori reasons. howfor neither pure peptone nor the products ever. after introduction into the alimentary canal. while that of Stoll and Schmidt consisted almost entirely of albumose.
sh. such as starch and fat. exact nature of &c. but rather than water alone. kreatinin. that absolute alcohol. for which purpose the funnel and india-rubber tube for pouring in the secreted in the second hour 2'6 c. in addition to water.c. Starch. from the experiments digested with of Dr. remaining food-stuffs. In many cases. Milk and solu- this undiscovered chemical stimulus of gastric secretion. effects. The first drop of juice appeared thirteen minutes after the introduction of the fluid. What the effective agency in each of these cases is. in the extractive materials of tion of gelatine were. chemical excitant. or finally. The inactivity of starch as a chemical stimulus was made the basis of the following interesting investigation u . Up to the present we only know. whether and mixed in less. From this it seemed natural to conclude that the actual substances which produced the result mentioned must also be contained in the peptone of Chapoteau. especially with solutions of meat extract. of juice with a digestive power of 4'25 mm. however. 97 the secretory process in the stomach. the active Liebig's Extract is bodies for the most part when remain behind in the residue. fluid naturally had to be connected beforehand with the fistula.. One must bear in mind. and possibly the secretion remains undecided. Chigin's hands not to have any exciting boiled or unboiled.MEAT EXTRACT A CHEMICAL EXCITANT. after the water has started a secretion. Thus. is however. In the course of the first hour 5'3 . also found to be direct chemical stimuli to gastric secretion. in possessing a greater degree of stability. on the track of We plete analysis of the constituents of venture to hope that a more commeat extract will finally bring us at present only found one other fle. of water in which grms. we have viz. . Lobassoff}. as in the case of meat. remains wholly obscure whether some exciting constituent is actually present in the substance itself. The experiments carried out with the above products. The reality and significance of the fact. The same applies to grape-sugar and cane-sugar.c. of Liebig's Extract were dissolved.c. these experiments were carried out on sleeping animals. proved Dr. whether the material comes into existence under the influence of some other alteiation. such as kreatin. in no way thereby diminished. were injected through the fistula into the large stomach of a dog. however.. The individual extractives. or whether the effective agent must first be formed during the course of digestion. had no greater effect. were found to be ineffective. 1 I give one as an example : 150 c. Lobassoff. that eggalbumen contrasts with the constituents of milk and with gelatine. excited by water it is not sufficient to alter it to such an extent as to in render The capable of exciting the gastric glands. were with a digestive power of 4'0 mm. different proportions with water. are already to be numbered by tens (Dr. c. The the chemical excitant remains as yet unexplained.
In all these cases the fat was directly introduced into the stomach without swallowing.. .. . judging from (Dr.98 THE WORK OF THE DIGESTIVE GLANDS. therefore. 2-2 2-8 1-8 1-2 6-25 5-88 G-25 nth C.. 0-7 .. water and juice. . and. solution of Liebig's Extract of Meat. They were tested not alone on a dog with isolated miniature stomach. we obtained. J-G-5 8th 0-2 Total . ... . . . . when we made a mixture juice would be secreted. . 5-0 fvO mm. as was anticipated.. Digestive power. it the stomach. in a absorbable form. . certain as yet unknown constituents of flesh belong. Quantity of juice. 0-6 7th .jelly. and finally on dogs which had survived the operation of section neck performed several months previously. if the ingredients of the meat extract could be somewhat longer retained in the stomach. 1st figures from one of the experiments : 2nd 3rd 4th . Lobaasoff). of the vagi in the Hence..c. meat extract acted through the blood. from the mucous membrane. the majority of food-stuffs showed an absence of stimulating effect upon the secretion of gastric To the minority which yielded a positive result. . Here are the Hour. If was to be expected that. if the would be much more effective less than when mixed with starch.. is a stimulus of only moderate strength. .. . 12-3 This experiment is also interesting because it materially supports the assumption which w e have tacitly made. The reason of this may possibly lie in the fact that the solution quits A so. . . and animal Particular care was also devoted to the investigation of vegetable fats. twice as much juice as would be yielded by the same quantity of meat extract in simple watery solution. is it It or even on the glands themselves.. . a larger quantity of As a matter of fact..th . and divided the cooled jelly into pieces which were afterwards inserted into the stomach. the quantity of juice secreted. . whose surface it should specifically excite. in solution at once apparent that... of meat extract with starch solution. ... but also on animals with gastric and oesophageal fistula?. and not that they are absorbed into the and then act directly on the peripheral neuro-glandular apparatus. too quickly. The result was invariably negative. 2-8 c. . when separately investigated. . that all the substances hitherto employed excite the nervous machinery by a reflex stimulus r blood. . . .
then see what the combined influence of the composite food may be. When analogous experiments were undertaken with bread and boiled egg-albumen. and then pushed the flesh into the stomach finished in the with a suitable rod. and this may naturally lead to a psychic excitation of the secretion. and then saw that all its introduction into the stomach occasioned either no secretion at or It was only necessary. however. arrangement which was used for bringing the flesh into the stomach (Dr. To prevent this mishap. Lobassoff. and it drops off to sleep again immediately. was always obtained. and is easily to be explained. it is true. Liebig's Extract to the sodden flesh in order to restore to it the activity proper to raw meat. either on the one hand that these unexpected results were due to the unfavourable physico-chemical condition of the material (locking-up of or.FLESH MEAT A CHEMICAL EXCITANT. It is justifiable to suppose. who undertook substances dissolved in the meat juice. from fifteen At this point I must not overlook an to thirty minutes afterwards. the dog guesses what is happening. If the what has happened. Thus he boiled the flesh thoroughly for several days. for the dog should wake up. to add some only a very weak one. stuffs into the We shall When considerable quantities of finely divided raw flesh are intro- duced unnoticed into the main stomachs of our dogs the secretion of juice begins (as already stated in the fifth lecture) at the earliest. and learn in how far the former may be explained by the latter. but the procedure always awakens it. When meat is introduced piece by piece into the fistula. introduced glass it gently into the fistula-tube. From to others. further. Sometimes. the investigation of this question. of the already mentioned simple effects. under these conditions. Dr. But flesh always sets up a secretion even This was to be expected after what has been said in the beginning of the present lecture. however. by introducing the food-stuffs into the stomach in a manner which wholly excluded psychic influence. made up as it is. that is to say. the animal maybe asleep. and. avoiding at the same time the act of eating. as has been said. Lobassoff found that watery infusions . They remained (as long as the observation was kept up) without exciting the least trace of secretion. the water). on the other hand. Obviously the secretion is chiefly to be attributed to chemical Dr. mind. and the feeding has then to be waking condition. it can no longer guess whole thing is finished. that direct chemical excitants are The negative effects of fluid albumen must be borne in really absent. Lobassojf). in 99 the experiments with simple substances we may now pass on which we introduced different combinations of ordinary food- stomach. a two or three hours negative result. adopted several modifications of the experiment in order to test the validity of these conclusions. we filled a wide tube with the meat-pulp from the mincing-machine.
I beg you to the experiments with starch jelly. a chemical agency comes into existence early in digestion. however. The source of the remaining portions remains as yet undiscovered. nlays In support of this If from experimental data. Lobassoff). a much stronger and more constant secretory effect is a like quantity of water or fluid eggproduced than is yielded by The formation of this product cannot. which is tolerably clear. which count from the end of the first three or four hours. correspondence. in the one case eaten. as a matter of fact. and them directly into the main stomach of a dog with an isolated miniature cavity. a secretion lasting two to mentioning. in compare the other directly brought into the stomach. Only the initial secretory period. that during the elaboration of bread and egg-albumen by the psychic juice. egg albumen is eaten. some other cause. calls forth if no secretion worth eaten by the animal. as already stated. Probably it is a digestive product identical or similar to the substance which the role of exciter in the case of flesh. hour after the eating of the egg-albumen possesses no specially strong while the psychic juice. namely. while. With Their relabread and boiled-egg albumen the case is otherwise. some other excitant. no longer obtains when bread or boileddiscloses its at once. That. Are we then to rest content with these results Do they furnish a complete explanation of the progress of secretion in normal digestion ? In the case of flesh diet the condition of matters Obviously. make the importance starch paste. As I told you previously. must that the juice of the second or third exist. . follows from this. is explained . which sets the neuro-secretory mechanism of the stomach into activity. To of these questions more manifest. Not more than one-half or even one-third of the juice secreted in these cases can be ascribed to the psychic effect. The most natural conclusion is. effect has its origin in the psychic effect. bread had no stronger stimulating influence on the gastric glands ? than equal quantities of water. is of the digestive power. A The juice poured out in the second meaning " " case exactly corresponds to the secretion. most active kind. careful consideration of this experiment three hours is set up. the stomach of injects explanation we are able to bring forward one obtains the fluid digestive products a dog which has eaten egg-albumen. No ! The secretion here is partly set up by psychic and partly by chemical substances peculiar to flesh. when directly introduced. tions are by no means clear. the energy of psychic This which we have already learned in the sham feeding experiment. since only a portion of the secreted juice be accounted for by conditions at present revealed to us. albumen (Dr. the further periods.100 of THE WORK OF THE DIGESTIVE GLANDS. are dependent upon an iinknown can process.
the remainder.EXCITANTS PRODUCED DURING DIGESTION. In the case of flesh this juice affords important assistance to the pre-existing excitant. two is KM however. here plays the role of the igniting material which sets the stove ablaze. delivery of these lectures I had to repeat my experiment on the influence of vagus division on the secretory effect produced by sham feeding. Further. as I had anticipated from my own previous experience and also from the publications of other authors. and in this way to test For the of the secretory processes is correct. which lasts to three hours. is in dogs with divided vagi. only formed during digestion. as stones would without the least appearance of digestion. or still better. is always followed by a considerable augmentation of the secretion (Dr. The psychic juice the digestion of these foods spontaneously proceeds. meat broth or meat extract. in the light of our new discoveries. Chigin. however. Part of the chemical body is exciting substance already exists in the is meat . of verifying all these results in actual practice. I have been afforded an opportunity play the part of igniting material. the psychic wholly and for ever done away with. just lie. or introduced unobserved. this statement . and thereby determines a rapid digestion. Chiyin). In such cases the appetite juice is the sole initiator of the secretory process and the necessary condition for its continuation. may be made to . It is obvious that when bread or egg-albumen is eaten without appetite water. supported by the following experiment. I resolved. When a secretion is already going on in the stomach. because after the psychic secretion. This investigation brings under our notice a very special and exceedingly important property of the psychic or appetite juice. caused either by psychic effect or continued from the last period of digestion. " and for this reason it has been called " Igniting-juice by Dr. especially Ludwig and Krehl. an indispensable condition to their digestion. to try and aid the animal's digestion. the unobserved introduction of fluid egg-albumen. Probably it is also. The result was that a greatly disordered condition of digestion was set whether our analysis up in the dog. I . be very active. comes to an end the hourly quantity of juice poured out upon bread or egg-albumen is very small. this juice is With other foods for example. shortening the stay of the raw product with bread If in the digestive canal. they lie there for a long time. If started by its assistance. on this account. that the psychic juice with every food possesses a uniform and tolerably strong digestive power. How is this phenomenon to be interpreted if one is not to suppose that at the commencement of egg-white digestion a formed which stimulates the mucous membrane ? The same explanation which holds good for bread and egg-albumen may be extended and applied to the proteids of flesh. into the stomach bread or egg-albumen be eaten without appetite. secretion of gastric juice Now.
the variations of digestive power in the juice secreted during the later hours after the eating of food. finally to the amalgamation of starch difference The first supposition was easily set aside. Not say. of almost exclusively confined ourselves to the quantity of juice secreted that upon the different foods. Consequently. a juice of like digestive power that secreted on ordinary bread. . In dealing with the secretory processes. which was satisfactorily digested. as a to matter of fact. then introduced 200- meat broth and waited till it became strongly acid. It due to the physical properties of the food. must be produced by the dissimilar chemical influence of the different foods. or bread again moistened. Before each feeding we washed out the dog's stomach. or with the proteid of the bread. We mixed flesh and pure starch paste in the same proportions in which the proteid and starch of bread are found. then tested (Dr. Chigin\. therefore endeavoured to supply its place by some other mean?. till the gastric glands were till then did \ve introduce solid food. we have up to the present :>(>() c. otherwise began to decompose. without the relationships in the digestive powers The third hypothesis was of the juices being altered (Dr. Lobassoff).c. or to the special might be nature of the proteid substances of bread and flesh respectively. is this alteration produced all In our investigation we started from the fact that a much stronger On what does this than upon flesh. How the psychic juice for As we have already repeatedly stated. gave this artificial bread to our dogs to eat. But from the second lecture. Meat may be desiccated. and obtained. juice is poured out on bread depend ? There were a whole series of possibilities. we know the quality of the juice with the different kinds of foods also varies.102 THE WORK OF THE DIGESTIVE GLANDS. Hour. ? kinds of food possesses a uniform digestive power. By this means the food. that is to thrown into vigorous activity.
but it to light which is important towards a knowledge of secretory of the mixture of starch and flesh revealed processes. from the second lecture. and hence in themselves constitute a valuable enrichment of our knowledge in this field. but a totally different curve if of gastric secretion was observed to that obtained the same food. fat also when tested by itself has no influence on the gastric does not necessarily follow that it would remain ineffective if introduced as the forerunner of. be. cubic centimetres of Provence oil. which. enable us to settle possible this question positively . another one. namely. the As in the case of similar effect of fat. usual five had previously been given (Dr. as you already no secretion. Half an hour to an hour later. just as the study in the effect produced by the former upon the proan interesting point fact came perties of the juice. of confirmation of our third theory did away with the But although the same specially testing the second. might its effects in various ways. other In the further investigation of the fat influence. the decline after the second hour must be ascribed to the less potent influence of so. when it again becomes more effective and since the psychic juice. the mechanism of their production required still further investiThe combination of starch with the proteids of flesh. that after a meal of flesh the produce may We juice from the second hour onwards continuously declines in digestive power till near the end of secretion. If this be that the starch paste may in some way restrain or prevent the depressing action of the meat constituents. gation. were poured into the main stomach of a dog know. of flesh. know already. namely. Instead of the to ten minutes. a new food materials. that a material which of itself is unable to excite a gastric secretion can alter the work of the glands in a decided and special we are able to raise Whatever the answer may manner when combined with the active With the question just discussed. starch. stands in natural connection. necessity results were repeatedly observed from the above experiment. but nevertheless it it argues considerable pro- gress that and take it experimentally in hand. which is present during the earlier . always possesses a fairly strong digestive power. or in conjunction with. however. is how starch able to the secretion of gastric juice. the dog was One hundred given its usual ration of 400 grms. influence constituents of as to flesh. The material in possession of the laboratory does not yet. Chigiti). hours after the taking of food. without oil. glands.CHEMICAL EXCITANTS OF GASTRIC SECRETION Tliis brilliant 10. produces by means of the sound. we have here before us a new fact. But another explanation is also possible. it is the chemical excitant of the flesh. It is conceivable that the paste strongly stimulate the trophic nervous mechanism without affecting the true secretory. we had to wait half an hour to an hour before .
When the flow at length commenced In the space of two to three hours. Either the fat in two ways. . as previously compelled to adhere to the second hypothesis of flesh always begins with shown.. .104 THE WORK OF THE DIGESTIVE GLANDS.c. which is inhibited precisely this secretion. instead of the it was very scanty.c. a (experiments of Dr. above everything. by reflexly stimuits nerve endings the inhibitory nerves of the glands. from the oil had been for 1st same ration of flesh. ... with a flow of central origin. . as may be clearly seen from the experiment quoted. curve of secretion was obtained when the fat was introduced into the stomach immediately after the meal of flesh. ..... ... the beginning of the secretion. . and only later make its appearance. . and A similar influence did the inhibitory energy at the usual time after feeding. the secretory activity after a meal the psychic juice that is to say.c. How is upon our experimental and remembers that the secretion was collected from the arrangements. 17'8 c.. of flesh Hour. we are those nerves. of one of these experiments. . f>-2r> mm. . and it .. Fat depresses inhibits the normal energy ? of the secretory process.c. given after 75 one and a half hours in the stomach : c. is by the . usual lOc.. per hour. : Normal secretion after a meal of 400 grins.. .c. of olive 2nd 3rd 4th . 5-3 4-5 3-8 . 4-3 c. quantities appeared. or the inhibitory centres of lating After a careful examination of all the facts. to 5 c. by and preventing the covering over the mucous membrane of the stomach this effect to be interpreted When one reflects or else indirectly.. I give here an lowering of its digestive power was also observed. that here the secretion began with normal was very much later we collected only when the normal 3 c.c. and also by way of contrast I give example when the normal flow after a meal of flesh without fat. 1st 2nd 3rd 4th .. .. Lobassoff).c. 1'75 T75 A that new and very is. excitation of for. Quantity of juice. ..... besides the diminution of the juice. fat.. 13-8 12-0 8-5 .. The only difference consisted in this. . Digestive power. striking fact is here before us.. to 15 it c. gastric cul-de-sac. it might be explained hinders the secretion directly in a mechanical way for example... 4-5 . the same results were obtained In these instances the dog was given fat mixed with flesh to eat. Finally. . 3-75 . 3-38 Secretion.. 4-25 3-0 mm. .
It is of great interest to note that a prolonged able to overcome the inhibitory effect of the fat. When the influence of such a powerful excitant sham feeding as the psychic impulse can be diminished by for the stimuli.c. milk with an . 4-75 mm. During the course of an hour 1*8 c. Occasionally we allowed the oil to flow out of the stomach immediately before the sham feeding.c. to 100 c. feeding we took the which the secretion commenced. without exception.ve solved this problem by the We administration of cream to our dogs that is to say. of oil were juice were accurately determined. which we fixed into the fistula. One hundred c. of juice.. and after a quarter to half an poured hour.c. During the course of two hours nothing was secreted by the cul-de-sac. which we have just discussed. After thirty minutes there was another sham feeding for six minutes. "j Digestive power. The action of fat. Lobassqff") to vai'y the experiment in On a gastro.INHIBITORY INFLUENCE OF FAT.and oesophngotouiised dog a sham conceivable way. . stomach layer of ? Whether the covering of the mucous membrane with a fat contributes in any way to the diminution of the secretion cannot be answered with certainty from the material to hand. Hour. In view of the great importance 105 of the inhibitory influence of fat. both with regard to duration of the feeding and to the quantity of food. . the sham feeding was repeated precisely as before. 2nd 3rd . the quantity. In every case. or in some cases still later. The sham feeding in this case lasted six minutes. may possibly explain the slow progress of the secretion after the taking of milk. is were collected. present also it began much later. and also the low digestive power of the juice.. into the stomach of the same dog. 1st .c. juice. the quantity was much less and the juice much weaker.c. which act how much more must this hold good directly on the mucous membrane of the fat.. and when it was secretion was observed. greatest pains (Dr. a marked diminution of the psychic Often there was none. The experiment on the dog with isolated stomach and divided oesophagus was particularly convincing. Quantity of . with a digestive power of 4v> mm. 1-0 0-5 . . .tube. The secretion of juice in these cases was observed by means of a wide glass tube closed at its outer The specifically heavier juice end. every The time at of short duration (one minute) was carried out. 4-0 c.. thus collected at the bottom of the tube and was at once visible. olive oil were then poured into the stomach.. Once more a sham feeding for six minutes was made. Is the fat of the milk not to be credited with this ? hoped to ha. and the properties of the Then 50 c.
This.should be The following still weaker. as a matter of fact. If the fat is to be credited at all with the increased low digestive power of " milk juice.106 THE WORK OF THE DIGESTIVE GLANDS. . table gives a comparison of the secretion with milk and with cream (Dr. is the case. amount of fat. Lobassoff)." that secreted upon cream .
Such an assertion is incontrovertible when all the data two preceding lectures are taken into consideration-. of this and the The secretion begins with the psychic excitation of the secretory nerve centres. the miniature stomach remains constantly empty of food. the same effect when the substances in question are introduced into the . the chemical materials which excite the secretion are absorbed from the centres digestive canal. The condition of matters has here radically changed when contrasted with former times. from the small stomach after the psychic secretion course. and indeed a complex one. we must (to preserve uniformity in our conception) admit a probability. what occurs when the psychic influence has come to an end ? At all events. . for instance. If. we may state with all positiveness that this apparently weighty consideration is. conceive the that One may. and carried by the blood either to the secretory which they excite or even to the glands themselves. to which a work must be assigned. and its absence in the other. would involve of the an enormous difference in the working conditions After a careful investigation into the facts of two stomachs. after all. as happens when the food is placed unobserved into the stomach of the dog. then. attempts to disBut such a mechanism has now been discovered. This supposition is If it be correct. that the nervous system acts in a similar way during the remaining phases of the secretory process. 107 It must occur to every one who becomes acquainted for the first time with our investigations upon the secretion of gastric juice. . success cover a nervous mechanism was not for attained gastric in the glands. that while the main stomach during digestion is filled in the ordinary way. These facts we shall take as the basis of our argument.SMALL STOMACH A MIRROR OF THE LARGE ONE. of no moment. How are they to be explained ? How can local events in the large stomach produce an effect. the beginning of the secretory process is identical in the two stomachs what is the condition. the case. we still see a flow its has run . after the secretion has set in. and this excitation naturally spreads in identical fashion to all points of the mucous membrane and its glands. on the small ? The functional connection between the two stomachs can only be effected by one or other or both of two systems of the body for instance. or even when it has not at any time been present such. One might suppose that the contact of food in the one case. which we assume has its causation in a local excitatory effect produced by the food in other words. where notwithstanding the endeavours of several investigators. whether in the large or small cavity. we ought to obtain capable of being easily tested. circulatory or the nervous. But with this proved. When juice flows at the beginning of the meal from the gastric cul-de-sac. its activity at the time must be accepted as identical with that of the large stomach.
of stimulation It is. which the vagus fibres are severed. out the rectum and investigating the washwater both physiologically and chemically. . even after a full meal. by reflex stimuli from the larger cavity. Furthermore. As a matter of food itself fact. and not a mere local one a definite area of mucous this area but over . since the in constant motion. The dog operated upon by our method. is excited through nervous channels that is to say. growing less and less from hour to hour. after all. a gradual decline of the secretion is noticeable. under the conditions of gastric digestion. In such animals the variations are occasioned by differences in the water contents of the food eaten. only three to five hours. In the beginning. therefore. The results on this point speak most decidedly against the hypoexperiments thesis. with one made after the method of Heidenhain. and is rapidly moved from the point to another. that is membrane occasions a the whole mucous membrane to say. even in the later phases of secretion. administered to his dogs. it may be taken that. the dogs thus operated upon do not manifest the characteristic alterations in the work of the glands dependent on differences in the food. These conclusions receive important confirmation when we compare the activity of a gastric cul-de-sac. Many investigators have injected meat broth and solutions of circulation in other of Liebig's Extract per rectum into animals. We also further conclude that secretion in the large stomach likewise depends on reflex events. per rectum. they work very energetically after a full meal their secretion is copious : and If. isolated in according to our method. the reflex must be a diffuse. is only a piece of the large). however. if the nervous connections of the cul-de-sac effects of excitation of have been preserve! uninjured. quite natural that the the inner surface of the large stomach should be constantly and accurately transmitted to the small one (which. which we depicted in the second lecture. But if these exist. and a month or six weeks after the operation. it lasts for lasts for several hours (Heidenhain. the animals live long. still manifests an exactly similar course of secretion in the two cavities under the same conditions. Ssanozki).108 THE WORK OF THE DIGESTIVE GLANDS. He Dr. Lobassoff has also investigated this question with great care. ways than by means of the stomach. is a localised response would have little meaning. the excitation of secretion not alone at of the stomach. cids-de-sac But the formed by Heidenhain's method in the course of time have their secretory capabilities very essentially altered. but have never seen an indication of gland activity. Hence we are driven by a process of exclusion to the imperative deduction that our miniature stomach. he convinced himself that By washing the excitatory material of the extract had disappeared. much larger doses of meat if extract than would suffice to induce a secretion injected into the stomach. now three and a half years ago.
having a made according to our first own method. a dog with an isolated small stomach. only in definite situations. the figures were given. for instance. however. have severally been repeated and confirmed on a number of cesophagotomised dogs with ordinary gastric gastric cul-de-sac fistula?. there This happens. it is simultaneously proved that the exciting substances first. of reflex origin. the stomach and intestine were separated from each other near the pylorus well as from the stomach. fistula and also a duodenal . upon the mucous membrane of the alimentary canal. show that such an effect arises both from the small intestine as In a dog which had an ordinary gastric one (provided with a metallic cannula). on the is no reflex effect on the gastric glands from the rectum . that the whole work of secrenervous and. Water produces a secretion both in the large and in the small orgar. has been proved by direct observation. The same applies to Liebig's Extract. with a second dog. other hand.MINIATURE STOMACH A MIRROR OF THE LARGE. with the possible exception of the psychic effect. Egg-albumen and starch. 109 But these results do not constitute the only reasons which convince us of the reliability of our method. the most imporanimal. It is not difficult to see that our second question concerning the seat of action of the exciting substances is also answered at the same time as the When it is shown. The absence of the sham effect in dogs with Heidenhain's isolated stomach is in perfeeding fect harmony with the fact that in animals with intact stomach but having the vagi divided in the neck. in which As you may remember. and divided oesophagus. recent experiments of my own. the juices from both We stomachs were in every way the same. and. tion is of act on the peripheral end-apparatus of the centripetal nerves. the solutions of which act more strongly on both stomachs than water. were reproduced in a tant of the facts observed on the veritably stereotyped manner. In short. to recall the facts bearing upon the matter and to arrange them in In the fifth lecture we described a sham feeding experiment on order. Fat produces a secretion in neither. and at the same time evidence was obtained show- A have here only ing that the above deductions are fully justified. indeed manifests rather an inhibitory influence. whether in fluid or solid form. in consequence. the sham feeding is also ineffective. which were obtained on the dog with isolated stomach. are ineffective on both cavities. complete parallelism between the work of the large and that of the small stomach. I think it is also essential to mention here that many of the facts. Recently also. we know of no single instance where the secretory process takes a different course in the two stomachs. which are not yet published. The above similarity in the working of the two stomachs is also to be seen when those excitants which act immediately upon the mucous membrane are employed. As we have seen.
so far as I see. It will only gradually by giving way before the discovery of other neuro-glandular will attract ever phenomena. in order to excite the gastric glands. In the immense majority with a strong central (automatic ? ) excitation of the secretory and trophic fibres of the glands. however. . fluid is directed from the lumen of organ. but this In plants the nervous system has not analogy is scarcely justified. was able to penetrate into the thick mucous membrane. In support yet been differentiated into a separate tissue or factor its functions are shared by all the cells in our case. one necessity admitted.110 THE WORK OF THE DIGESTIVE GLANDS. How little Now. Sooner vation of cases gastric digestion begins or later after the taking of food the influence of the reflex excitant comes into play. the analogy of insect-eating plants has been brought forward. however. the gastric glands . into the mouths of the glands. the supposition becomes physiological this idea sounds all the more improbable since according to the latest investigations the of the inner surface of the stomach are very limited. it must not be forgotten that even a long-sustained absorption might not of necessity lead to a penetration of the substances into the Nor can one even admit that the substances gain entry peptic glands. absorbing capabilities ! Moreover. disprove this die outj improbable hypothesis by direct experiment. Unfortunately. matter once more to you as a working hypothesis of the inner- Now apparatus of the gastric glands. while the automatic psychic effect gradually dies out. If meat have been eaten. In this dog a gastric secretion could be set up when the exciting substance was introduced either into the stomach or into the intestine. But a remote possibility may still exist that the food (apart from its reflex effect) could act also in a more immediate and direct manner upon the glands. the reverse effects happen. of by a septum formed mucous membrane in a way similar to that described in our operation for making the miniature stomach. I must present the stomach. when nothing was known concerning gastric innervation. which formerly. We must confess that this hypothesis. the secretory centre will still be strongly excited in a reflex manner from the stomach and intestine. appears now quite inconceivable. however. depends on nervous processes. because during secietion the stream of of the gland towards the cavity of the the view for which we are contending. . When After the cessabread. whilst at the same time the trophic centre receives only weak impulses from the peripheral terminations of the nerves in question. had to admit that the food. one cannot. was of To accept it. have placed at their disposal the services of a very intricate nervous mechanism. which certainly to themselves. more and more attention that I have shown you that the secretory activity of the as a whole. is eaten.
and as it has developed under our hands. and this error was in its turn a result of faulty methods. we has Blondlot's theory survived. Only in the works of a few writers and these mostly tion. reflex inhibitory impulses proceed to the centres which affect the activity both of the secretory and of the trophic nerves. are strongly influenced. thanks to the faulty experiments and erroneous representations of later authors. on the other hand. however. published in the year 1879 in Pfliiger's Arc/tires. have passed out of sight. Is the In its details. French must give mention Of other investigators. work of Blondlot of and the additions of Heidenhain comprise almost importance which was accomplished by physiology in everything of the secretory fifty years concerning the conditions and mechanism work of the stomach during digestion. who has enriched the physiology of absorption in general. . the secretory fibres are now only weakly excited through the end apparatus. stomach. but more especially in connection with the secretory work of the stomach. but not in its fundamental picture a new one ? may appear. the sketch of this picture outlined by physiology. that the researches of Blondlot and his views upon the secretion hensible. I have depicted the work of the gastric glands as we have seen it in our experiments. The of the specific excitability of the gastric mucous membrane. proceed the subdivision of the secretory as well as the suggesto investigate the individual food-stuffs would be important in relation to the work of the Heidenhain's results are con- tained in his well-known article on the secretion of the cardiac glands The of the stomach. the trophic.MECHANISM OF GASTRIC SECRETION SUMMARY. However singular it was more than shyness of new things and for its conversion to our interpretation of the phenomena under consideration The talented author of the Traits Analytique dela Digestion Blond! lot spoke in plain words of the importance of the act of taking food. yes. but. to Heidenhain. From him process according to periods tion that it and exciting agencies. of gastric juice. of his theory were naturally insufficient. for our subject was the obvious error that mechanical stimulation constituted an effective excitant of the gastric glands. Full of moment. May this constififty years ago tute another reason for our science relinquishing its characteristic features. on the contrary. In cases where fat has been added to the food. Ill tion of the psychic stimulus. has discovered important facts and given birth to many fruitful ideas. have experienced during the last fifty years no comple- but we must not forget that the first gastric fistulse had only just been. no additions. facts adduced in the working up and experiments on dogs with artificial It is truly incompreperformed.
and which has consequently to adapt itself to peculiar chemical conditions. and consequently there can be tigation which arise no question of a direct penetration of the food into the gland ducts. GENTLEMEN. moreover.LECTURE VII. The intestinal canal is completely counted by separated from the lumen of the gland. in our lecture to-day.1 properties of this excitant duodenal mucous membrane Probable significance of the relationship established by the acid between the gastric and pancreatic secretionsStarch does not augment the secretion of the pancreas. AND PROBLEMS FOR FURTHER INVESTIGATION. considerably disthe following advantage. Fat is a reliable exciter of pancreatic and also increases the amount of fat-splitting ferment in the juice Sleep does not hinder pancreatic secretion Although psychic exmay be accepted as existing. however. excite the pancreas chiefly because of the acid reaction of the gastric juice The acid reflexly excites the pancreas by acting on the is . Water is an independent exciter of the pancreas Solucitation of the pancreas tions of neutral and alkaline salts of the alkalies inhibit pancreatic to be investigated in connection with the work of the digestive glands The definite establishment of a fundamental specific excitability of the digestive canal constitutes the secretion Grouping of problems still result of all the experiments related Summary of the results from a general point of view Outlook and programme of future investigations. THE NORMAL EXCITANTS OF THE NERVOUS MECHANISM OF THE PANCREAS: SUMMARY OF MATTERS DEALT WITH. content of amylolytic ferment. a supplementary which works on food already affected by a previous agency. and by what means the nervous mechanism of the pancreas comes to be excited during digestion we must from the beginning exand unexpected facts. yet it plays only an unimportant rule. pect to encounter both complicated relationships The secretion of the pancreas it is is richer in ferments than that of the fluid stomach and. In turning. but does increase its secretion. . how. Acidity powerful excitant of the pancreatic gland The specific The stomach contents on passing into the duodenum. to the questions as to when. The difficulties of inves- from these sources are.
energetic.c.c. tensity of the juice-secreting influence of acids. acts here as the exciting agency. chloric acid I introduce of by means of the sound and makes no protest whatever against the procedure. of for solutions particular of neutral on the other hand. the animal was fed hours ago.c. I bring into the stomach 500 c. and the flow faster. just as did the disIn the search covery of the p>ychic excitation of the gastric glands. tested. by means of the sound. and we may now proceed to the important initial experiment of our investigations iipon this subject. solution of hydrointo the stomach.. the secretion at present to three drops in the minute. and pancreatic juice no longer flowed. taken from the work of Dr. perfectly Two to three minutes after the injection you notice that the drops fall We can already count twenty-five in the minute.. but. 0-4 c. two is As you : see.IT. hand. I give here an example who had the investigation in The dog was fed twenty-two hours beforehand. and you see not only that the secretion does not increase. pancreatic stimuli we (Dr. .C'. grows stronger and stronger. Dolinski. In order to meet the objection that water still half per cent. water the alkali metals It resulted that a species of carbon dioxide gas. the carbon dioxide distinctly more These results directed our attention to the effects of acids.ACIDS EXCITE A FLOW OF PANCREATIC JUICE. '. This powerful influence of acids upon the pancreas is one of the most securely established facts in the whole physiology of the gland. of lime-water. grows weaker and weaker. indeed almost stops. creatic flow that by their The acids are such strong stimulants to panmeans one can excite the activity of the gland is more effectively than by any other. the for We Becker) effects . on the one hand. Two hundred and fifty c. 113 'of begin with a form of stimulus which aroused the attention the laboratory in a very striking manner. 9-5 9-5 . salts of and alkaline saturated with excitants of pancreatic juice than water. hardly worth mentioning. into the stomach. reasons. The dog remains or the influence of fluids generally.c. antagonism was found to exist between these substances in their The saline solutions proved to be weaker effects upon the pancreas. of hydrochloric acid of the strength of the gastric juice were poured. and even fifteen this quite easily accounted for 1">0 c. The quantity of juice secreted every five minutes is shown below : (i'O C. so much this the case. the effect of acids has become a crucial test of the normal To illustrate the incondition of the alimentary canal in this respect. 2-i . . The dog which I bring before you possesses a is permanent pancreatic fistula. that in the laboratory. on the contrary.
c... 0-2 13-5 15-0 10-0 13-0 15-0 Secretion stopped .c.. 7-0 .e. an exceptional fact.c. 7-0 8-0 1-0 0-2 0-8 "'I ..c. the effects of increasing strengths of hydrochloric acid (from O'Oo to 0-5 per cent.c.. therefore.c. indeed. In the 2nd hour. In the 2nd hour.. c. In the 1st hour..THE WORK OF THE DIGESTIVE GLANDS.. therefore. citric. 0-0 - . 3-D c. . and simultaneously we remembered that the contents of the stomach normally possess an acid reaction. at once occurred to vis that we had discovered in acids the specific excitant of the pancreatic gland. It. in addition to the hydrochloric. Thereupon 250 c.c. No noticed. 82 . water were poured in no secretion followed for thirty minutes. l. tions had to be proved and established experimentally. (VO (1-2 2-0 0-5 . results : Into the stomach 250 of HC1 of the following strengths were poured : - 5 per cent.) were tried and with the following c. In the first place. The constancy and intensity of the acid influence stands out as a very remarkable. Then another 250 c.c. .VO 10-5 y-0 7'5 io-r. In the 1st hour. . 13-0 c. 74 - c. particular difference in the exciting effect of various acids was The acids investigated were phosphoric."> c. appeared to us that this acid reaction probably serves as a connecting link between two neighbouring compartments of the But all these interesting and pregnant supposialimentary canal. lactic and acetic.c. . The idea. of the same hydrochloric acid solution were introduced and the quantities of juice recorded every ten minutes : : 1*5 c. 65'5 c.
We The solutions caused distinct burning on the tongue but not the least in- dication of a stimulating effect on the pancreas. must be interpreted in a different way to that which the author indicates. The experiments of Dr. an excitant appeared all fectly indifferent to the the more forcible since the gastric glands remained persame acids. while weak solutions of acids invariably caused a flow of the juice at once. As a logical sequence to the above. or through a fistula. to be excitants of the pancreas only when If neutral or alkaline. with the same substances. caused by the large doses of the substances in question. confirmed us in the belief that they behave not merely as general and indiscriminate This conclusion exciters. When at the height of digestion. our hypotheses became fully established when we succeeded in destroying the exciting effect of the gastric contents by neutralisation of the mixture. but as specific stimulants of the pancreas. in some cases less. or lime-water. 115 It was conceivable that in the employment of 0'5 per cent. which alone possesses specific It appears to me that these irritability) were in consequence excited. with a even free secretion of pancreatic juice. Finally. have. The proportionate nature of the effects. pepper and mustard.HOW ACIDS ACT IN PROMOTING PANCKEAT1C FLOW. however. even more con- We have often compared the effects (Prof. the sensitiveness of the pancreas is about as great as that of the organs of taste. for a fluid which just tastes acid acts distinctly as of the gland. secreto-motor effect was no greater than that of water. With the former we could perceive no trace of exciting effect. acid we had not yet reached the highest degree of gland activity. solutions of soda. and the great sensitiveness to the influence of acids. which were performed on rabbits. Decoctions of red pepper and mixtures of oil of mustard and water were used as strong as they could be employed without producing vomiting. their they possessed a strong acid reaction. and which gave contrary results. one introduces into the stomach of an animal by means of the sound. when introduced into the stomach. so far as one can judge from experiments not systematically carried out. Solutions of different sugars. Obviously there must have been an injury to the mucous membrane. or pancreatic juice. of such stimulating substances as pepper and mustard with those of acids. and the centripetal neives themselves (not their terminal apparatus. we convinced oxirselves that To begin pure gastric juice was just as powerful an excitant of the gland in question as an acid solution of equal strength. came the further conclusion that the gastric contents must have an exciting effect on the gland by virtue of the acid reaction which they possess. with. It naturally was not difficult to test this supposition. SchirokicJi) vincing proofs. of peptone. one always . data suffice to give a positive answer to the question as to whether acids are specific exciters of the pancreas or not. after a few minutes. Gottlieb. On the other hand. and of ovalbumen proved.
however. Ixix. a diminution which often : observes a diminution of the normal secretion lasts for here one experiment as an example The secretion was recorded every five minutes. Hence the) conclude that its action is not a reflex one through nervous channels. 1902. 5-0 6-8 6-0 5'7 . p. as are introduced from without. p. 352. also and Starling have recently shown (Proceedings Royal Society.* as The question how these substances could act in the blood its alkalinity. I give 5-6 c. but a substance produced by its action upon the mucous membrane of the duodenum and of the upper jejunum. very simple. we must decidedly accept the Let us attentively consider the facts to of the acid effect.116 THE WORK OF THE DIGESTIVE GLANDS. London. 682) that the exciting agency is not the acid itself. also after the injection of atropin. which they term "seeretin" is effective when injected into the blood stream. therefore.c. the saliva which moistens everything is able to act as an exciter of gastric secretion by virtue of its dry.c. and this can be the only alteration which * Bayliss vol. and thus the mutual content of water. after extirpation of the solar plexus and destruction of all nervous fila- ments passing to the isolated loop of intestine. 1902. pancreatic juice is expelled. its point of action either locally by exciting the peripheral end-apparatus of the centripetal nerves in the mucous membrane. upon each other is is clearly We are. We see in the above of the segment an instructive instance of how the work of one alimentary canal is connected with and dependent upon that of the previous one. CntralMatt f. 6-6 7-2 7-4 1-4 c. the facts already communicated. The acid works are two possibilities. In the stomach itself it is in this way ensured. that the psychic secretion which is the forerunner of digestion is continued by the influence of the saliva. is first hypothesis. and so on. together with the results of analyse some new experiments. influence which the digestive glands exert manifested. The acid of the gastric juice acts in its turn as an excitant of the pancreatic gland.c.c. were here poured into the stomach. by reducing If. namely. therefore. Bd. the alkalinity of the blood soon as acids must thereby be reduced. Physiologic. 5'6 c. or else it is absorbed into the blood and stimuIf we lates either the secretory centre or the gland cells directly. a long time. Thus. The substance is not destroyed by boiling or neutralisation of the acid extract of 1 . pancreatic gland. xv. 1-0 1-0 1-1 7-2 6-8 1-5 1-6 Of the dog's own pancreatic juice 70 c. This substance. 2'2 . justified of tbe in saying that the acid is a specific exciter ? There Where.
Popielski. which in its turn is derived from the blood. however. This fact was first indicated by Dr. are therefore unable to admit an acid effect through the But we are in possession of direct experiments which speak in the same direction. In normal digestion.THE ACID EFFECT UPON PANCREATIC SECRETION. From these theoretical considerations alone we medium of the blood. the alkalinity of increased. Here is one of his experiments on a dog with a per- manent pancreatic fistula: Time. calls forth the secretion of pancreatic juice. happens in the the blood is 117 fluid. according as the pancreas is excited to action by acids introduced from without or by that prepared in the stomach. Gottlieb. The food of the stomach. The increased alkalinity of the blood during digestion is also in harmony with the well-known fact that the alkaline content of the urine is greater during the same period. When acid solutions are injected In the same into the rectum the pancreas remains in perfect rest. by virtue of its hydro- chloric acid. without entering the intestine. . but the question has since been more fully investigated by Dr. acids do not act on the pancreas if they continue in the stomach way. consequently diametrically opposite changes would occur in the blood.
is less. was In addition we have had a dog (Dr. This fact is undisputed. and thus to provide a suitable medium for the activity of its ferments. is prepared from the sodium chloride of the blood. when acid was poured into the pyloric portion. PopielsM) whose stomach divided into two parts near the pyloric region. but only when the acid passed on into the duodenum. mean chemical composition of the fluid. to forced conclusions. for pancreatic juice protects itself against the destrucwhich a neutral or alkaline medium is very fruitful idea of Thus the Biiicke that the bile arrests the action of pepsin in the for intestinal digestion duodenum and provides favourable conditions may also be extended to the pancreatic juice. into each of which a fistula-tube was brought. and this must. a secretion of pancreatic juice This appeared. but why does the acid and not something else serve as the ? Naturally. and thereby in turn lead to a marked diminution of the alkalinity of the blood. in order to An preserve the it. of physiology. according to the present teaching acid possible. Consequently the reaction of this fluid would suffer great variations in both directions during digestion . On the other hand.118 THE WORK OF THE DIGESTIVE GLANDS. falls in that in the case of the pancreas. we can only bring forward some hypotheses. as we have already said. this is a factor the constancy of which is of great importance to the chemical processes which transpire in the . tive action of the pepsin. interesting question remains of how we are to account for the fact that the acid has an effect. We may mention here once more. would again be absorbed. but as we know. excess of alkali thus arises in the blood. If now acid were introduced into the larger segment the pancreas remained at rest. and with any pronounced degree it Hence may be conceived that the pancreatic juice comes in to neutralise the acid which causes it to flow. however. is in harmony with the fact that the flow of pancreatic juice rises and undulatory fashion. the pancreatic ferments act best in an alkaline medium. after fulfilling its function in the alimentary canal. At the same time the unsuitable. we cannot as yet claim to have solved this question scientifically. connecting link When of acidity weakly acid their effect it soon becomes nil. At juice is secreted with the most concentrated strength of hydrochloi'ic This hydrochloric acid. who interprets the action of acids upon the pancreas as anything else but reflex. The substance. obviously from its connection with the entry One must be very partial of the acid food mixture into the intestine. the same time another important significance of this relationship For a peculiar and as yet not quite obvious reason. be removed from The hydrochloric acid. forms The a connecting link between gastric and intestinal digestion. As is known. the necessity is completely removed of of the discussing a direct penetration of food materials into the lumen gland . the gastric lies near.
an acid-exciting substance evokes the production of alkali. and never merely for the neutralisation of acids. Determinations of the amount of ash. with the object amongst others. the basic element. is This shows that the relation- The juice is only one of relative importance. Walther. is that is acid gastric acidity and in to say. or. as well as titration both of the . the sodium. always produced for digestive purposes. the ferments of the juice adapted themselves to the kind of food ingested at the time being. ash and of the unaltered pancreatic juice. organism. apart from the content of ferment. was found to be increased. and with milk the fat-splitting ferment. we should in consequence expect to meet with variations in the alkalinity of the juice. that juice hours after partaking of this food approaches very nearly in inorganic to constituents the " acid juice. juice causes secretion of pancreatic fluid l>y its direct proportion to the sodium chloride it. Recently this explanation has received support from the experiments of . while the acid constituent of The taken up by the peptic glands and then passed on into the cavity of the stomach. know [t if the acid always plays a part in the production of the appears.THE ACID EFFECT UPON PANCREATIC SECRETION. In the first lecture we saw that analogous to one previously described. which is poured out on acids is never wholly deprived of its ferment-properties. serves for the preparation of pancreatic fluid. have incontestablv shown that a connection exists between the nature of the secretory excitant and the amount of inorganic substance in the pancreatic fluid. The juice excited by acid solutions shows a very unimportant amount of organic substances with a maximal content of inorganic in fact. And thus the two constituents of the sodium chloride meet again in the alimentary canal and reproduce the salt. for " " flesh secreted by the pancreas during the first instance. The juice. Thus the rapidity of the secretion has no decided influence. Likewise here. the latter is two or three times as great as that of the former. of neutralising itself. while the organic constituents which are not required are extraordinarily reduced. to juice. With bread the amylolytic ferment. If the acid excites a ilow of pancreatic juice. " the " acid juice retains its characteristic properties even when the This occurrence is perfectly hourly quantity of secretion varies. however. in other words. the quantity of . ship just mentioned By a fuller study of the alkalinity we shall probably soon be in a position to indicate that part of the general flow of pancreatic fluid which is evoked by the acid. It shows also a very high degree of alkalinity both in the ash and in the juice itself. and determined by the acidity of the exciting fluid and this is indeed the case." This harmonises beautifully with . 1 1 These difficulties are all removed when we consider the relationships of the digestive juices just discussed.Dr.
that the starch happen here. Science has not attempted. greater. we were unable to prove an exciting effect on the part of starch. namely the acid. Solutions of starch of varying concentration expelled the juice no more energetically than water alone. or different therefrom. to reproduce a synthesis of real digestion that is to say. the acid of which appears to be at the same time the chief exciter of pancreatic flow. far as our experiments go. the bread juice a lesser degree of fat-splitting action. and of the probable importance of the acid The experiments dealing with the influence of fat upon the pancreatic gland yielded much more positive results and were much simpler. So for. This does not. Finally. It may. Consequently. The further investigation. has proved itself to be the strongest excitant of the neuro-secretory apparatus of the pancreas. to combine the often conflicting interests of the different foodstuffs among themselves with organism as a whole.120 THE WORK OF THE DIGESTIVE GLANDS. the digestion may be dependent on some other condition continued production of lactic acid from the carbohydrate constituents of the food. preclude the possibility that there may also be other exciters. requires that the minutiae of the requisite conditions for the starch possible still for instance. Some experiments When he fed a dog with bread. after a meal of flesh. those of the digestive canal and of the Here I beg to remind you of the relationships of digestion. Possibly this is the explanation of the chemico-physiological fact in question. augments the quantity exercises only a trophic influence effect may of the ferment without increasing the of Dr. the meaning and importance of which have as yet been but little cleared up. however. since the field of action is wider in the case of the ferments of the pancreas. Hence the natural question arises whether starch and fat may not also be exciters of the pancreas. that is to say. the mere comparison of known facts made it very probable that fat is an independent exciter of the pancreas. as we know. however. either identical with those for the gastiic glands. fat to gastric effect. ticularly interesting that in the same specimens of juice the fat-splitting ferment behaved in just the opposite way the flesh juice revealed a Walther furnish a basis of fact for these hypotheses. and with the same rate of flow. as in the case of gastric juice. Indeed. for it is quite question. and could not up to the present venture. total quantity of juice. the possibility is not excluded that the progress of starch for example. for the time being have escaped our observation. the gland has special relations to these substances. Fat restrains the secretion . the fact that in the early hours after a meal of flesh a very vigorous secretion of gastric juice takes place. a non-nutrient substance. the pancreatic fluid possessed a much stronger amylolytic action than the juice obtained at a corresponding It is parperiod.
When the acid secretion from the stomach ceased. The tube at the end of the At arm of the T-piece remained open to allow the gastric contents to escape freely. the beginning of the experiment the connecting tube was between the funnel and the T-piece with a Mohr's clamp. of oil.c. and consequently we could not fancy that. to 10 c. attained a rate of 7 c. A perfectly healthy dog. in fifteen minutes. effect or. the secretion of pancreatic juice after the administration of fat takes place even when every trace of acid contents is absent from the stomach. Dolinsky poured fluid oil info the stomach flow of of dogs. The gastric was closed by a cork through which a glass tube was led. and constantly observed a more or less considerable Bearing in mind the strongly inhibitory effects of fat on gastric secretion. but three to five minutes after the pouring in of the oil the flow was quite distinctly increased. was secreted. to 115 c. Often at the beginning of the experiment the stomach secreted a clear acid fluid the psychic gastric juice. At the same time only a very small quantity of alkaline mucus collected from the stomach in the lower division of the india-rubber tube . consequently. Dr. having two fistula? one a gastric. the experimenter then cautiously closed the outflow tube and opened the clamp on the oil So long as the stomach cavity remained unclosed. A confirmed sceptic might. or at most O5 c. a pancreatic secretion after fat is indirectly caused by the acid of the gastric juice. and after fifteen to thirty minutes funnel. in fifteen minutes.c. Beneath the orifice of the pancreatic duct a metallic funnel and graduated cylinder were fastened. pancreatic juice. so clamped that the oil could not leave the funnel. Fifteen to thirty minutes after the introduction of the oil. The animal gave up the hope of receiving anything to eat. as we previously indicated ourselves. More rarely it transverse contained only a self in little a separate alkaline mucus. however.c. and which in turn was connected with an india-rubber tube and funnel. The funnel was hung up at a suitable height. The india-rubber tube was also provided with a T-piece. of 121 gastric juice. the clamp on the lower . fistula the transverse arm of which was connected with a second india-rubber tube. reply that possibly an acid fluid had accumulated in the stomach before our experi- ment.c. and finally slept. and contained 110 c. that a strong psychic might have overcome the inhibitory influence of the fat. But here is an experiment instituted by Dr.EFFECT OF FAT ON PANCKEATIC SECRETION. The experimenter secluded himroom with the dog and quietly waited.c. the constancy of this result afforded good assurance that we had here to deal with a direct influence of fat upon the pancreas. Some- times the experiment was varied in the following way.c. Damaskin which complies with the strictest requirements. either no pancreatic juice. to I'O c. the other a pancreatic fistula was last fed twenty hours before the experiment. under ordinary conditions.
however. The Dr.c. For the most part it contained only 15 c. mucous membrane of the duodenum. in the course of an hour or still longer. Thus we have often observed a strong and lasting secretion of pancreatic juice. means When it is borne in mind that fat of fat scarcely requires a special disis a chemically indifferent But substance. immediate idea that an acid reaction is rapidly set up in the intestine by anti-peristaltic action. If the milk. when the milk is synthetically reconstructed the fat ferment in the pancreatic juice increases to the previous amount. and 3 c. reacted alkaline. be deprived of its fat by filtration. the fat may well excite the peripheral end-apparatus of the nerves which are specially adapted to react to every possible influence. cleavage of the fat. such as chemical. with the described above. either early or late. india-rubber tube was opened and the contents of the stomach emptied out. and the like. cluding that fat is investigations of that fat excites also first affords us adequate grounds for conan independent exciter of the pancreatic gland. even when the stomach had been perfectly emptied of fat.c. some bile or bile-stained fluid This latter fluid contained suspended escaped from the stomach. fat In our case the contents consist of mixed with the fluids bile and pancreatic juice which act upon . suggests the idea that the duodenum forms a cavity. We might from the experiments related. of oil. But the mechanism of And pancreatic excitation by cussion. that the stimulus acts on the suppose. fat When. c. which is characteristic of "milk juice. together quantities of oil flowed out. to 5 with small alkaline mucus. Walther go still farther. during the whole of this time These facts exclude the pancreatic juice freely flowed from the fistula.122 THE WORK OF THE DIGESTIVE GLANDS.c. without any other alteration in the progress or rate two hours commonly of the secretion.c. mechanical. an effect through the blood can scarcely be thought of. showed no trace of acid reaction. however. after a meal of milk a juice is furnished which is unrich in fat-splitting ferment. but In the increase of the fat-splitting ferment. the milk filtrate is again mixed with to say. fat. of with this mucus. as regards the seat of influence of the fat. flows for one to two hours from the stomach. possibly. bounded somewhat and that the contents of this cavity are continuously driven from one end to the other. like the stomach. This fact. and then acts as an exciter of the pancreas. to 20 c. the juice presents a very low fatsplitting power. The intestinal contents. I now wish oil to introduce some i-emarks relative to the experiments fluid." that is now. and show undoubtedly The experiment leads to an not only a free secretion of pancreatic fluid. while later only mucus mixed In other cases. An emulsion-like as has been stated. and was obviously driven into the stomach by Nevertheless.
In the interests. and allowed its body to hang in the In this way the skin was straps of the boot. We have intentionally deferred this question near the end of the . The investigator who made it was. we should be inclined to admit the activity of the above-named exciters. Thus a distinct but purely extraneous and accidental connection came to exist between the sleeping of the dog and the arrest of the pancreatic flow. This fact is differently stretched. is have shown that this was an error. As I have also possible. however. sleep does not exercise the least influence on the work of the gastric glands. the feeling of appetite and the presence of water are necessary to ensure a beginning of gastric secretion. of a greater degree of independence for the pancreas. 123 a fact Tt Under after their influence it is emulsified and split." When the animal slept. although the secretion is essentially dependent (through the acid) on gastric digestion. and fluid it is that is one or two hours the emulsified becomes acid. When But why? the animal slept the secretion abruptly declined Instead of the connection which was it was found that a purely extraneous circumstance explained the relationship between the sleep of the animal and the arrest of the flow. If retically. apparently.PSYCHIC EFFECT AND PANCREATIC SECRETION. the answer might be either a positive or a negative one. in position. but we of pathological conditions where the hydrochloric acid was absent from the gastric juice for months. The dog in this case was retained in a frame by a special assumed. it. Will they not also have an influence on the pancreas ? I refer to the psychic effect. Theotogether with that of water and also of extractive substances. it naturally assumed an easy. in secretory previously stated. even when it is in full progress. the distinct statement went forth from the laboratory some creatic years ago that sleep almost completely arrests the secretion of panFurther observations fluid. But facts till must decide the case. the case of an empty stomach. which we named " boots. fell to nil. We shall now pass on to the other exciting agencies which proved to be effective in the case of the secretion of gastric juice. and perfectly right. With regard to the pancreas. and the origin of the mistake not without interest. In affections of the stomach the pancreas might remain without its chief exciter. passive position. arrangement of leather supports many parts being displaced from its ordinary and at the same time the duct of the pancreas which passes through it was kinked and pressed upon. further evidence of the necessity for constant watchfulness over the minutest details in the performance of physiological experiments. that the duodenal The second remark cavity may expand at the expense of the former. the same might also apply to the flow of pancreatic fluid. therefore. concerns the sleeping of the animal during the experiment. and yet digestion as a whole pro- know ceeded tolerably well.
Now. the gastric glands. is not easy to the acidity of the gastric juice. indeed. In the experiapplication of the exciting agency animal by offering it food. part of the juice obtained entry into the was another way by which the matter might be decided. begins to flow two to three minutes after the juice. probably. which was at that time perfectly justified. and also operative procedure. remain in perfect rest. an acid. They are not also capable of acting independently and directly. as the case might be. e/en with an open gastric divergent conclusions. and were able to observe the onset or augmentation. This appears to me to point generally begins to a direct psychic influence through the secretory nerves of the pancreas. of this experiment is ambiguous. gastric juice. however. was that the nerves of the pancreas can be excited by psychic influence. which one may often notice when the flow of pancreatic juice is After a rumbling sound in the abdomen animal. however.124 THE WORK OF THE DIGESTIVE GLANDS. have been decisive if no juice had flowed. viz. of pancreatic The result. very simple.. however. One is at liberty to conceive that a passing desire for food has thrown the centres for the motor nerves of the intes- . It would flow. observed in a such as has fasting a more or less active secretion often sets in. the stomach. as we have seen. made both gastric and pancreatic fistula?. with the gastric fistula open. apart from And this. and is never less than four and a half minutes. But there fistula. we may draw It may be that. decide. We then submitted the gastric juice. The latent period of the gastric secretion in dogs has a sharply marked lower The pancreatic limit. but this with obviously now requires further examination. duodenum. Are we not dealing a stimulating effect of the gastric juice collected in the stomach under the effect of the psychic impulse ? It was necessary to repeat the experiment in a manner which eliminated the intervention of the In the beginning we placed our hopes on a complicated We performed cesophagotomy on a dog. Dr. dog to a sham feeding. pancreas through It is necessary to determine whether they actual question in hand. otherwise with those agencies which are at the same time excitants of established relations. His conclusion. its lecture because solution is intimately connected with previously The experiments which concern the excitants but it is quite special to the pancreas are. the determination of the latent period of sham feeding for the pancreas. And with this. on the contrary. for example. will naturally also be indirect excitants of the But this does not solve the the acid of the latter. ment of teasing the of long been established for the secretory mechanism a phenomenon is connected. the pancreatic flow also after two to three minutes. Kuwschinski showed long ago that tempting a hungry dog with food at times called forth a very lively secretion of pancreatic juice.
that water exciter of the nervous occurred in the latter. moreover. acids little and fat. continues for a time after the emptying of the stomach. . and then empties increased. however. as we know. it is said that in severe hunger the stomach yearns. those The experiments extractives of flesh ? which we have found in the on this point were arranged in the same way as those with pure water. the strong craving for to be included. and has In any instance. When water is poured But how it ? into the stomach Is it because it a secretion of pancreatic juice. or may only have The conclusion is clear and is an independent and direct mechanism of the pancreas. We must. free from objection. the question of psychic excitation of the pancreas requires further investigation. the extractive subspecific stimuli is . How do other chemical excitants of the gastric glands behave. while the impulse is not sufficient to excite the more inert gastric glands. for a universally recognised fact. we are in a position to say that there to the gastric glands and to the pancreas. into action. and gave precisely the same When solutions of meat extract were poured in. in addition. although no secretion whatever may have appeared after the lapse of ten minutes. It is. Damasking as in the case of When 150 c. Amongst their food and water. independently excites the gland. the glands of which are resting.c. one sees after two or three minutes that the secretion of pancreatic juice either begins or becomes distinctly If one waits for a minute or two longer. both organs have for the gas-trie glands. stances of meat and for the pancreas. even become a byword when. or rather a neutral or possibly alkaline fluid. some water. dog. to be borne in Similar considerations to those in the case of psychic excitation have mind with regard to the relationship which exists between Avater and pancreatic is secretion. or because the result has led beforehand to acidification of the gastric contents? In the experiments to decide this question the same method was adopted (Dr. of water are poured unnoticed into the stomach of a oil. which. and was in no case began greater. dwell a longer on the inhibitory phenomena .CHEMICAL EXCITANTS OF PANCREATIC SECRETION. But. however. is Occasionally the secretion of pancreatic juice generally still found. the stomach. intestine. the secretion results. If we now sum up these are some excitants common these facts. and also for pancreatic secretion. after the same length of time as with water. namely. namely. Finally comes the question. 125 tine. own probably the psychic effect. have a longer latent period. The psychic excitation of intestinal case. possible that the centre in agreement with the fact that the for the nerves of the pancreas gland belongs to the abdominal set of digestive organs should be more or less closely associated with the centre for the motor nerves of the movements is.
of given in water.. seen. here some figures taken from the article of Dr.c. the experiment more exactly.c. some substances diminish the secretion of pancreatic juice very striking.c. which comes down to And here an unusually low point.V4 r>3-. water poured into the stomach. water poured into the stomach. The dog was given 1200 c. Becker. The secretion was recorded every hour. which became evident in certain cases during the activity of the panAs we have already related. Becker's article. 250 c. c I ask you to recall to mind what I said to you in the first mcerning the efl'ects of a continued addition of sodium bicarbonate to the food. In every case a marked diminution of the secretion was to be an example from Dr. 46-4 is-i 21-4 14-5 . whether an influence is also produced through the blood. 56-3 21-5 1-V7 12-0 . 22-4 18-7 14-4 13-1) . of meat. of Essentucky water.c.c. 250 c. in question The secretory influence of the solutions was compared with that of water.120 THE WORK OF THE DIGESTIVE GLANDS. in which was particularly observed how long such influence lasted. 7-3 o-o 1-0 . How we are to interpret the mechanism of inhibition remains still obscure. That the . and cei'tainly deserves our consideration."> .c. Total 204-7 Total 152-1 Total 200-6 lecture. 32'2 c. The same food without the Essentucky water. and in every case the I give flow of pancreatic juice was considerably less with the former. 9-9 6-2 18-0 c. Such an addition for a length of time markedly depresses the secretory activity of the pancreas. 5"6 c.c. milk and 2 Ib. It is at present difficult to decide whether we have only a local Hence the fact that is effect or on the peripheral terminations of the reflex-transmitting nerves. Once more I give Two hours before the feeding the dog was given 400 c. 2 grins. An hour later the animal received its ordinary meal and the resulting secretion was compared with the normal. it The inhibitory efl'ect was also investigated in another way. The solution to be investigated was poured into the stomach of a dog by means of the tube.c. 250 c. creatic gland. of NaHCO 3 4-2 c.c. 4G'(5 c. solutions of alkalies and of alkaline salts of the alkali metals not only do not excite a flow of I will describe pancreatic juice but even exert an inhibitory action. 42-3 c. 62-1 4.c. The fluid was collected and recorded every half-hour.c.
In the second lecture we learned of the great complexity. And that the idea obtained no general recognition. because it was not founded on any definite basis. ? How variations of gland activity come about ? We have previously spoken of the foods and subdivided them into their separate component materials. very much. completed the part of these lectures which and pancreatic secretions. deals with gastric I have now. of the work of the gastric It is glands and of the pancreas. but we have not by a long way . but I am far from believing that the subject is thereby in reality exhausted. local effect in 127 any case is not confined to these substances is apparent from the fact that the inhibitory influence is not confined to them. that in recent articles is and text-books only a stimulating the food as a whole mentioned. have now many more open questions before us than we had a short time ago. although the idea of a special stimulating effect caused by acids and by the acid chyme was long ago expressed. but which we now valuable. must be borne in mind. The questions are so many that we must group them together for discussion. In particular we must answer the following ments questions not any other : Why is at any given moment a certain quantity ? secreted other ? In what way of juice and certain properties and not any do its quantitative and qualitative variations Why has it advance the digestion of the food are they of service to the as a healthy condition of the digestive canal and of the organism such as. However. We believe we are justified in characterising them as new. belongs Oae gathers the impression that certain substances which are easily soluble in water diminish its ordinary properties acting as a local stimulant. now for every step of this complicated process.EFFECTS OF FOOD ON PANCREATIC SECRETION. Damaskin). has still to be achieved before we are able to congratulate ourselves on That which is gained is already very a final conquest of our territory. gentlemen. it may serve as a sign-post to guide future research. necessary to seek an explanation and in doing bo the require- of the individual food constituents. and the conditions necessary for the welfare of the digestive canal. and prevent it from Such are the facts which this laboratory . how do nil these whole? To these questions others are linked. but also to other bodies readily soluble in water (Dr. because they testify to the existence of a wide field of inquiry which we have studied from a general point of view. and all these questions mean progress in our investigations. it is a long way from the mere expression of an opinion to a clear and precise statement of facts. because We wish to submit to exhaustive research.has collected concerning the normal excitants of the pancreatic gland. but at the same time great constancy and accuracy. and for the organism as u whole. Much. is shown by effect of this.
one could decide whether its secretion was caused by acids or not. but also by impulses from other organs as well. A and in what way gland complete answer to the two groups of questions. On the other hand. It is probable that the same questions likewise apply to those digestive fluids. of course. I hope. will only be obtained when an exact investigation of the contents of the digestive canal is joined hand in hand with observations upon secretory activity when. for any given period of digestion. Although much more remains to be done. such as the bile and the succus entericus. have shown quite as exact and intimate a connection between this event and the nature of different sorts of food as we have already learned for the gastric and pancreatic juice. they deal with the chain of phenomena in the central nervous system. interwoven in the closest manner.128 brought THE WORK OF THE DIGESTIVE GLANDS. their mode of working. have at once and for ever done away with the crude and fruitless idea that the alimentary canal is universally . jected to. is we know to be found. separate point of the curve of secretion which we observe after a com- In order to solve this problem we must plex meal. determined not only by peripheral impulses from the digestive canal. must successively synthetically build up the food step by step. and for every point of the digestive . must be explained. on the one hand. and the combined results of the local specific excitants. for instance. because their physiology. Bruno. moreover. all their real constituents under consideration. combine the individual constituents with each other. from the point of view of these Recent experiments of lectures. their seat of action. Our results. Dr. A correspondence in the results obtained and the synthetic The systematic investigation by both methods the analytic furnishes the best assurance of their reliability. the effects of the elementary food constituents. has not been sufficiently worked out. and. of the elements of the food must undoubtedly lead to the discovery of many unexpected relationships between the food-stuffs on the one hand and the digestive glands on the other. In the case of a complex food we shall then be in a position to draw conclusions from the properties of the juice as to what is the effective Thus from the degree of alkalinity of the pancreatic juice excitant. The questions in these two groups are. which have not as yet found place within the limits of our subject. We must naturally have a knowledge of all of these and determine their importIt is from the effects of the elementary constituents that each ance. precisely where a certain constituent of the food and what alterations it is at that moment being subThe latter group of questions concern. that is to say. canal. carried out in the laboratory upon the entry of bile into the intestine. why activity varies. we have reason to be satisfied with what has been accomplished. must submit the properties of the juice at each phase to an exact analysis.
when 1 look back upon their results. thanks to a suitable interlinking of the several processes. in nature. see now the outlines of a skilled mechanism which. convert a considerable part of the reaction. as we may say in contrast to cell-physiology. which are initial rate of is modified to harmonise with the altered now in such a condition that they are capable of acting directly upon the end organs of the neuro-secretory In the interest of all the constituents a certain equilibrium apparatus. however. before the final solution of the problem is arrived at. and. when once started on the right way. chemical or thermal agency. The working out of these presumably will furnish. our department of life. the instinct is at the same time the first and . in organ physiology. one may reasonably hope that I . and not as normal definite guiding factors in the Instead of a crude indefinite scheme. Even in the Far West a lively interest is taken in our researches. without distinction. I expressed the opinion that in ten years we should have as good a knowledge of the chemical work of the digestive canal as we have now of the physical apparatus of the eye. the one ingredient is promoted. that is. a theme of engrossing but ever-insatiable In interest. Since then two years have fled. are not dealing with assistance of We questions concerning the nature of life or the physics and chemistry of the cell. by every mechanical. the other to a certain degree restrained a species of contest for the ingredient needed is fought out between the several components of the food. as with everything and in An and instinctive craving for food essential advantage to the digestive process is derived from the for. to the beginning of these lectures. food into a soluble half-fluid condition. and thus allow the chemical constituents of the food-mixture to take effect. According to the prevailing view of matters. The : secretory work which began with the ingestion of food is thus propagated farther and farther along the alimentary canal. the pronounced action of those agencies could serve merely as favouring or restraining influences. in addition to the impulse to seek out partake of food. 129 excitable. In consequence of this.VALUE OF THE INSTINCTIVE CRAVING FOR FOOD. in regard both to the quantity and strength of the digestive fluids as a whole . and with the numerous European colleagues together with our workers the investigations. is established. The fluids of varying strongest exciter of several digestive glands. In my which I referred in address before the Association of Russian Medical Men. will rapidly here. lead to a complete accomplishment of the task. we secretory work of digestion. for a long series of generations. the gland activity constituents of the food. secreted under its influence. I see no reason to retract my words. proves itself to be adapted with the utmost delicacy the most suitable manner to the work which it has to perform. independent oF the speciality which attaches to each single phase of digestion.
within certain . What is the portals does it perceive peripheral end -apparatus of a centripetal nerve ? How this or that form of excitation ? What are the nervous phenomena by cell lead to which reactions and molecular changes in the secretory the formation of this or that ferment. and investigated the rules and laws of their working in the apparatus as a whole. the digestive canal) and the bearing of the whole towards objects of external nature (in this case to the food) will receive a complete explanation. We . or to the preparation of this or that reagent? We have hitherto taken these properties and elementary functions as granted.130 there are THE WORK OF THE DIGESTIVE GLANDS. have learned how this or is be set into activity that limits we have been able to understand it. At the very of organ-physiology we find such questions as. that apparatus may to say. many territories where the reciprocal relations of the individual component parts of a system (for example.
we would.LECTURE VIII. Nor should we be encounter insuperable Thanks likely to to the advances of bacteri- ology. the same methods should be applied from the same standpoint to the experimental investigation of the pathology To bring our knowledge by the physician in disorders of the digestive to full fruition. in a sense. that the methods described in these lectures should be employed in experimental investigations into the pathology and therapeutics of the digestive canal on the The fact that the beginning of the secretory work in the stomach depends upon a psychic effect harmonises with the experiences of everyday life. have to deal with external ailments. difficulties. namely. PHYSIOLOGICAL ACTION AND THE TEACHING OF INSTINCT: EXPERIENCES OF THE PHYSICIAN. that food should be eaten with attention and relish To restore the appetite has from all ages been the endeavour The indifference of the present-day physician towards of the physician lines laid clown Probable causes of this Curative remedies based upon a restoraappetite The therapeutic effects of bitters depend upon the excitation of appetite The usages of the mid-clay meal are in agreement with tion of appetite physiological requirements customs and empirical regulations food Dietetics of fat Physiological reasons for certain instinctive Importance of an acid reaction of the and its is therapeutic application The peculiar position of milk among food-stuffs of the curative effects based on physiological reasons Explanation of sodium bicarbonate and sodium chloride The work of the digestive glands Participation of the inhibitory nerves of secretion in the production of pathocauses of individual differences in the logical effects. since our present methods enable us to obtain access to any desired part of the inner surface of the digestive canal. Moreover. It would be desirable. and so secure for it and therapeutics of the alimentary canal. many of the pathological processes can now be experimentally produced in the laboratory. and with the regulations prescribed apparatus. GENTLEMEN. In such pathological animals the functional diseases of the apparatus could . the most useful application. To-day we shall endeavour to bring the previously communicated results of our laboratory investigations into reconciliation with the customs observed in the ingestion of food. in the interests of medicine.
. on the dog whose vagi nerves were divided in the neck. study of therapeutic questions in this But this does not exclude_the prob- . The investigator obtained through the fistula a mixture of substances from which it was difficult. because clinical study of the same subject (notwithstanding the zeal devoted to it during the last ten years and the results derived therefrom). At the beginning of the ailment. It can hardly be doubted that scientific. while local excitants almost completely failed. which was previously practised on animals. and thus easily thrown into activity by central impulses. One may conceive that the deeper layers of the mucous membrane with the gastric glands were still healthy. in addition to an Experimental Physiology and Pathology. On such animals therapeutic remedies could also be tested. is more uncomfortable than the ordinary form of gastric We fistula of the stomach.132 THE WORK OF THE DIGESTIVE GLANDS. indeed before it became fully established. the whole process of healing and the final result experimentally observed. Nor is this difficult to understand. whilst the surface of the membrane with the end-apparatus of the centripetal nerves was already distinctly damaged. or even at times impossible. the properties of the fluids. I have already described one of such pathological therapeutic experiments. has to contend with serious difficulties. the alterations of secretory activity. and yet the physiology even with the aid of the latter. Other similar cases I can also call to mind. while the conditions of secretory activity during every phase of the healing process could be investigated. but of strong digestive power. Our dog with the two at one time from a slight and transitory gastric was then very interesting to observe that the pathological process (which we were usually able to wholly guard against) spread from the large to the small stomach. still furnished juice in appropriate quantity). the pathological conditions of the digestive organs and their treatment. It continuous slimy secretion of very slight acidity. which I may call impressions rather than precise observations. gress for Hence the exact region still scientific belongs to the future. I mention these. the chief clinical instrument. It manifested itself here in an almost stomachs suffered catarrh. and the conditions under which they appear could be examined. namely. to decide anything. the psychic stimulation was remarkably effective (that is to say. there has also been built up an Experimental Therapeutics. because I wish to point out what a fruitful field awaits the investigator who wishes to study. with the aid of our present methods. proof that this is possible is furnished can only take its A by the recent vigorous strides made by the science of bacteriology. that is to say ideal. must not forget that the sound or stomach-tube. proper position as a science when. be studied in a precise and detailed manner that is to say. medicine. Such an investigation is all the more desirable. made no material pro- many long years.
because it will It is thus that be better adapted to the special needs of the case. however. in relation here more than anywhere else. medicine. everything is directed to take c. Certain preparations are carried out (in England a change of raiment is usually effected.THE IMPORTANCE OF APPETITE. ability 133 that the influence the newer acquirements of physiology may fruitfully But physiology naturally can work of the physician. induces a mild narcosis. Probably this also explains the use of alcoholic beverages at meals. to the foregoing facts. Naturally this highly developed hygiene of eating is only found in the intelligent and well-to-do classes. that food should be eaten with interest and enjoyment. because here also the food is served in greater quantity . based upon exact knowledge. the treatment of the diseased organ will be more effective. discussions and it is also plain why heated serious readings are held to be unsuitable during meal times. If it be at all admitted that We instinct is the outcome of an everyday experience. since the make no pretence knowledge at its disposal is incomplete. made out a brilliant case when Perhaps the old and empirical requirement. for alcohol. much more restricted than that of As a recompense for this. is the most imperatively emphasised and strengthened of all. may now return to our subject. to guide the field of medicine. the mode of action of which is not clear. being daily enriched by new physiological facts. and often a blessing is asked upon the meal by the oldest of the family). From this point of view even in the lighter phases of its action. applied physiology. physiological knowledge is often able to explain the causation of an illTo employ a ness and the meaning of empirical curative methods. musical and other guests are invited to while away the time at meals in a word. In the case of the well-to-do. because here the mental activity is more strained and the various questions of life more burning . or. a special room for meals is set apart . first. which has led to the unconscious adoption of the most favourable conditions for life. chosen. it The expresis particularly so with regard to the phenomena of digestion. which contributes towards distraction from the pressing burden of the daily work. is quite a different remedy In the latter case.ires of daily life. a company of relatives. instinct has often brought before the tribunal of physiology. from the and to concentrate away the thoughts them on the repast. namely. will at length grow into what it ideally must become. in other words. In every land the act of eating is connected with certain customs designed to A suitable time of day is distract from the business of daily life. and secondly. and is the broad world of clinical reality. the art of repairing the damaged machinery of the human body. human sion that physiology merely confirms the precepts of instinct is justified It appears to me also that. acquaintances or comrades assemble. thing from knowing precisely what we are doing.
amount of muscular activity and the constant lack of more than sufficient nourishment ensure a strong and lively desire for food in a normal manner.134 than is THE WORK OF THE DIGESTIVE GLANDS. I believe that in this they are not only animated by an endeavour to free their patients from trouble- some symptoms. as we say. Thus. a dog which has not fasted for more than some hours will not eat everything with equal pleasure which dogs usually eat. soon becomes worse than useless. Hence the presence of a certain kind of spice is a general requirement. not necessary. which are foretastes of the actual repast. without reason that it is often said cient enjoyment. are obviously designed to awaken the curiosity and interest. the physician has effectively employed measures to restore specially it. with perceptible enjoyment. required for the wants of the organism. and the instinct strives against it. is only right up to a certain point. eating to order or from conviction. The same conditions explain why the preparation of food is so choice in the case of the upper classes and so simple in that of the lower. for some degree of appetising taste is desired by everybody. that to give to excite them. although naturally individual tastes differ. In the case of the where mental activity is less highly developed. however. that to the same extent to which the patient wishes back his appetite. It was here only necessary is. an impulse to the organs of their activity taste. Every one knows that a normal. It may be said. and to augment the desire for food. but also of itself will favour the restitution of by the conviction that the return of appetite normal digestive conditions. appetising. This short discussion as to how different people behave with regard to the act of eating is of itself testimony that care should ever be taken to keep alive the attention and interest for food and to promote enjoyment of the repast that is to say. useful food is a food eaten with appetite. Medical men of all times and of every land have held it to be a pressing duty. One of the most frequent requests addressed to the physician is to restore the appetite." This dictum. How often do we see that a person who begins his customary meal with indifference. Further. Every other form of eating. after overcoming the fundamental illnesses of their patients. afterwards enjoys it with obvious pleasure when his taste has been awakened by something piquant or. Hence we have not a few remedies which are named " gastric . in order that might be later maintained by less powerful excitants. without recourse to any special regulations or customs. the greater poorer classes. that " Hunger is the best sauce. that care should be taken of the appetite. but will seek out the food which it relishes best. therefore. all the accessories of the meal. of course. The quelling of hunger in his case affords of itself suffiIt is not. even by animals. For a person who feels hungry such extra inducements are. to pay special attention to the restoration of the appetite.
has been shown to be Each of the contending factors has at length been purely imaginary. Naturally we do not doubt that a movement in this direction indicates a great advance. however. but even here. it is remarkable how is and whose action to little attention in a is paid to appetite as a is symptom or to its special therapy. as with every undertaking of mankind. one may meet statements in which the physician is recommended to adopt no special means for counteracting so unimportant a subjective symptom as a bad appetite After what I have said and demonstrated to you in these lectures. now that our knowledge is more complete. they are valued only in so far as in the laboratory or not into medical science. To a certain degree we can understand and this contributes to an explanation of matters how medical science of our time has come to regard so lightly the loss of appetite as a special object for treatment. and then merely in short parenthetic phrases. We often do not know all the essential conditions for the production of the phenomenon in question. and that which corresponds to the real conditions. Now. tonics. appetite. in disorders of digestion. reads current text-books on disorders of digestion. assigned to the mechanical stimulus. We must not consider an event to be a mere picture of the imagination because it is not realisable under given experimental conditions. As stated above. with the result that they can be verified by laboratory experiments. Unfortunately medical from this. mention in physiology. Great importance was. one ! can only designate such views as gross misconceptions. the efficiency of which. On the other hand. to promote secredifferent remedies. the correct treatment of the If one appetite. the experimental method has penetrated more and more many pathological factors and are judged of according to whether they hold good therapeutic agents that is to say.THE IMPORTANCE OF APPETITE. of digestion assistance was sought for in the but nothing was there met with which had a relation to laboratory. so far as quantity and quality of the juice are concerned. nor are we yet able to grasp the connection between all the separate Thus in the clinical treatfunctions of life as fully as may be desired. things do not proceed without mistakes and exaggerations. the psychic gastric juice obtained only cursory ." 135 science has latterly deviated promote appetite. it When the is precisely here that symptomatic treatment is essential. on the other hand. If anywhere. necessary. and consequently this factor was overlooked in medical prac- ment and pathology tice. this object can most certainly and tory activity by completely be achieved by endeavouring to restore the appetite. physician finds it We have already seen that no other excitant of gastric secretion. can compare with the passionate craving for food. . and this not even by all authors and when it was noticed it was related more as a curiosity. Only few of them its importance indicated.
and if clinical medicine maintains her worthy desire of following out the experimental investigation of her problems. This was an exact experimental reproduction of the customary treatment of a weak stomach. But notwithstanding the itself. From this point of view the meaning of removing a patient. which should take up the stimulus of the chemical excitant. the subject of chronic weakness of is the stomach. a mentally customaiy surroundings. in the commonest disorders of the stomach. since. or even wholly absent. is regulated according to physiological needs in all these cases the physician seeks to awaken appetite. many therapeutic measures are based on the promotion And in this. In the first case. only the surface It may. layers of the mucous membrane are affected. is not able to fulfil its duty. from his Take. or eats and carries on his work at the same time. and particularly the eating.136 assigned THE WORK OF THE DIGESTIVE GLANDS. which ordinarily lasts for a long time. particularly in the case of people who live . is obvious in these cases that the indication is to promote digestion by exciting a flow of appetite juice. its proper place. and relies enjoined to eat sparingly. the truth of empiricism makes itself irresistibly felt. ovei'strained individual. for instance. in addition to preventing the overfilling of a weak stomach. ask you here to call to mind one of our experiments in which food was given in small portions to a dog. she must in actual practice accord to appetite its old claim for con- sideration and treatment. consequently. as yet unaffected layers of the mucous membrane. may evoke the secretory is A impulse in the central nervous system and send it unhindered to the glands which lie in the deeper. indifference of physicians to appetite in of it. where the food is prescribed in small portions. for the most part disturbed. An instance of this. I have already related at the beginning of this lecture. strong psychic excitation. When the patient is from eating at all he is (for instance. or again. during convalescence) removed from his ordinary surroundings and sent to an establishment where the whole life. and thus led to a secretion of much stronger juice than if the whole ration had been eaten at once. or a responsible official how of ten does it happen that he cannot for a moment distract his thoughts from his daily work. when upon it as a factor in the cure. taken from the pathological material of the It laboratory. and the period of chemical secretion. This often happens. which is so rich I in quantity and so strong in digestive power. also plain. the oft-recurrence of appetite juice. a keen feeling of appetite. and not to rely upon that excited by chemical stimuli. i?s of great importance. happen that the sensory surface of the stomach. or when he is restrained till the physician expressly permits. And such a regulation of diet is all the more necessary. : He eats without noticing it.
The conclusion had already been empirically arrived at. or eats. that inclination of the taste ultimately determines the relation of . There is no " igniting juice. degrading to indulge excessively and exclusively in culinary enjoyments. however. but. The fundamental cause of membrane of disease. or even of whole communities. digestive canal than is necessary. As so often is the case. of With the establishment mental effect upon the secretion of juice the influence of condiments enters upon a new phase. It is. but also to have a care that in wider circles of by placing him the community a due conception of the importance of eating should be disseminated. and to substitute for a time.THE EFFECTS OF BITTERS. to take him completely away. to free him from his pursue. inattention to the act of eating prepares the way for digestive disturbances in the near future. but that should also be tasty. at most. to interrupt the interminable train of thought. into a state of decomposition the alimentary canal and brings it into a condition of medicinal treatment can help such a patient while he remains surrounded by his old conditions. in the interest of the public If this or weal. 137 The systematic in the midst of the incessant turmoil of great cities. The secretory appetite juice. for want of sufficient which irritates the mucous digestive juices. further. often do the people who have charge of the commissariat pay attention solely to the nutritive value of the food. as his only object in life. For this reason. It precisely in the so-called intelligent classes of Russians that a proper conception of life generally is often found wanting. who has often to express an opinion upon the suitability of the dietaries of different persons. This is attained by sending the patient to in a hydropathic establishment. No There is only one course to in progress. have made a species of cult of the act of eating. or place a higher value on every- thing else than taste ? We must. direct attention especially to the feeding of children. methodical nations. But how secretion . no the food remains much longer in the activity comes slowly into play . or passes. should constantly bear in rniud the question of psychic that is to say. the care of his health. very little. It is the duty to regulate not only the life of individual patients of the physician according to such rules. and where an absois More lutely unphysiological indifference towards eating often exists. of course. and his illness still continues a regard for what he travel." or. the physician. a lofty contempt for eating is also reprehensible. occupation. This is particularly so with the Russian physician. with all their consequences. namely. whether with or without enjoyment. like the English. the best course here also lies between the two extremes. Now. on the other hand. that it was not alone sufficient for the food to be it composed exclusively of nutrient substances. he should inquire after and learn how the food has been eaten. we know why this is so.
grown-up individuals towards food. . for instance. A and also to himself when in health. of the therapeutic influence of the so-called bitters. In either case there is no longer indifference. moreover. They are. drawn. They not only arouse no desire for eating. The whole question of the therapeutic importance of the bitters. which by contrast awaken the idea ones. this object is most quickly attained by exciting sharp. therefore.138 THE WORK OF THE DIGESTIVE GLANDS. that the conditions of the experiment possibly had not corresponded with the actual state of affairs. as how do bitters affect the appetite ? It is and later physicians that bitters increase the appetite. they became greatly discredited. or at best a perverse one. which are agreeable to other people. dislike that the gustatory apparatus should receive a necessary. and this pleasant . in since the appetite. not by any means strange that this had not previously been observed in the laboratory. When tested in the laboratory. it was forgotten directly excites secretory activity. . The substances were either introduced directly into the stomachs of normal dogs or else injected into the circulation. however. so that many were quite ready to discard their use altogether. other conditions in after life. such. it would seem undesirable to habituate children solely to a nicety and uniformity of gustatory senSuch might affect their capabilities of adapting themselves to sations. real secretory stimu- everything lants. now appear tasteless. however. a blunted taste. is the strongest of all stimuli to the digestive glands. Obviously. a long period of high repute these substances have been almost expelled from the list of pharmaceutic remedies. and was bitter taste common to them all. a matter with which the commencing phase of digestion is closely linked. however. were grouped together mainly on account of a certain nerves. therefore. in the eyes of the juice. But their action is chiefly it bound up with their effect upon the gustatory not. as has many times been repeated in these It is. the simple conclusion was produce a flow of gastric clinician. lectures. strong stimulus in order to restore a normal sensation. they were unable to justify their old and valued reputation when directly introduced into the stomach. a certain degree of gustatory The ordinary foods. but may even cause a feeling of It is there is no sense of taste. As experience teaches. is the universal opinion of the earlier and if this be so consequence. indifference. said. that a weak digestion could only be assisted by a remedy which In this. acquires a different significance when we link it with another question. without some reason that this large of remedies. unof pleasant gustatory impressions. many of them were unable to Consequently. The question appears to me. consisting of substances of the most varied chemical group composition. it After bears the closest connection with that of appetite. person who suffers from digestive disturbance has.
both of which are designed to awakea the appetite. and in many text-books of medicine. awakens an appetite. such as condiments. The customs of the chief meal of the day also corresecretory spond with our physiological results. whether applies to other substances. Thus it is represented that some medicinal remedy exactly reversed. of different kinds . and so This explanation of the appetising effects of bitters proceeding on. the laboratory therefore. which The same probably indirectly excites a physiological secretory activity. consisting mostly of meat broth (bouillon. the question of the therapeutic effect of bitters is settled in the affirmative the moment we acknowledge that these substances awaken appetite. the repast proper begins. different soups. it has been confirmed by many clinicians that after the administration of bitters effect of these some such special sensations do arise in the stomach. of assurance that the investigation has been correctly carried out that It to say. that it dealt exactly with the point under consideration. in addition. is is interesting to observe that the connection between appetite and gastric juice is by many physicians. by its presence in the Here we have to deal with a false stomach. is 139 may be awakened. As has been already stated in the fifth lecture. our explanation corresponds to the actuality or not. and has not hitherto been attempted in the lo is not sufficient to hand over clinical observations to the laboratory. The question is a difficult one. with something hot. calls forth a secretion of gastric juice.THE RATIONALE OF THE MENU. as experimental proofs. perhaps also with a liqueur of brandy (especially customary in Russia). lishing the fact an experimental investigation of bitters consists in estabthat they have an effect upon the appetite. and this. After this comes the really nourishing food meat and so on). the enjoyment of blithesome health more intense after illness. there is some reason for believing that certain impulses from the cavity of the stomach are likewise necessary for the It is possible that bitters not only act directly on excitation of appetite. One must have. but in the production of a certain psychic effect. a sound louder after silence. and that because it was not recognised that effect could by any possibility be a powerful excitant of nerves. The problem. from the mouth does not exclude the possibility of some such similar influence coming also from the stomach. and. . The remedies consists. in the majority of cases. but that they also act on the mucous membrane of the stomach in such a way that sensations are generated which contribute to the passionate craving for food. In any case. and the foundation upon which an appetite for this or that kind of food here a general physiological law is illustrated. After this or that hors d'ceuvre. the gustatory nerves in the mouth. a psychic explanation of a true fact. As a matter of fact. The light appears brighter after darkness. not merely in the generation of a simple reflex. therefore.
when a person has no appetite. and. from the standpoint of physiology. chemical excitant is used for awakening the early secretion. in two ways to secure a free secretion of gastric juice to act on the first in the excitement of the appetite juice by the chief food rational. . would be very useful. It contains a considerable quantity of lactic acid. is further to be borne in mind that the quantity of the digestive juices in general stands in close connection with the content of water in the organism. a strong chemical excitant for example. or. If one arranged the ordinary fluid foods in descending order. . and everybody knows that sweets are of The usual termination the * Jtioas is a favourite Russian drink. according to the influence of the chemical excitants. in the case of poorer people. holds good for healthy people. milk thirdly. therefore. or meat extract to people who have no appetite. different kinds mixed with flour. for instance. whatever. stews made with vegetables. is quite Meat broth. from the physioThe chief meal is generally to be understood. strong broth. particularly if they do not tives of flesh. water. . he has no psychic juice or only very little consequently. and therefore rich in carbohydrate material. and consequently with the poorer classes a less expensive. since the food can only be In any case the addition of meat extract introduced in a fluid form. some acetic acid. such as meat juice and the like secondly. of foods. of meat. The same applies also to forced feeding. &c. repast is also. hors d'oeuvre. or only a weak one. For example. A good meat broth can only be afforded by well-to-do people. This has been shown by the experiments of Dr. logical standpoint. easy ended with something sweet. of the insane. in every way desirable to prescribe meat juice. Walther for the If this sequence pancreatic juice and by my own for the gastric juice. : . the preparations of the flesh. while in Germany. as we have already seen. where the price of consisting of water meat is high. This sequence of foods. the meal must in every case be begun with strictly . with a solution of the extracOtherwise solid foods. kwas * serves in this way with the Russian population. visions for the digestion of the chief meal. and other products of fermentation. It is. indeed. is an important An attempt is therefore made chemical excitant of gastric secretion. with malt and yeast. It of soups are used. It is true that the method of introduction in this case necessarily secures the presence of a chemical excitant. it must be even more adhered to in pathological conditions. served in various ways. bread or meal. prepared from water. the following would be the series first. Thus. would remain long in the stomach without any digestion consist therefore. also a less effective. and secondly in the promotion of the flow by the action of It is in this way that human instinct has made prothe meat broth.140 THE WORK OF THE DIGESTIVE GLANDS. bread.
intentionally discard digestion in it the stomach. After thus dealing in general with the usual arrangement of our meals. since it brings the gland a digestive organ at once so powerful and so difficult of access under the control of the physician. demands a greater activity upon the part of the pancreatic gland. FAT. 141 The meaning of this is easy to guess. Above all comes the acid reaction of the food. are prescribed in digestive disturbances. acid fruits and green vegetables are used and they are either of themselves acid. and thus supplement the weak action of the stomach. in certain affections of directly brought about by the acid. especially of hydrochloric and phosphoric acids. and thus transfer to the bowel by prescribing substances . It appears to me that a knowledge of the special relations of acids to the pancreas ought to be very useful in medicine. but is at the same time It is even conceivable the strongest excitant of the pancreatic gland. number of acid substances. that in certain cases the whole digestion may depend upon the stimulafor ting properties of acids. we make use of acids. It is easy.THE EFFECTS OF pleasant. must also. The enormous quantity may either is assist digestion of starch which he consumes. in the stomach where too little gastric or bring about vicarious digestion by the pancreas present. many kinds of wine have a somewhat acid In Russia. consequent on the pressing need for food. Further. as food. associated with loss of appetite. acid solutions. which figures in a number of sauces and such like. The repast. the explanation being that they excite an increased activity of the pancreatic gland. We could. great quantities. It is apparent that use quite a acidity enjoys a special preference in the human taste. it is only the gustatory nerves which should be agreeably excited. and this is Further. notwithstanding the stilling of hunger. the stomach. or made so in the preIn medicine this instinct is likewise often made use of. juice where it is wholly absent. lactic acid in Finally. be terminated with an agreeable sensation. we may now speak of some special points. endeavours to prepare hydrochloric acid. Thus. one of the commonest We seasoning substances is vinegar. ktvas. and paration. since the pancreatic juice exerts a ferment In this way. is consumed in taste. to understand why the Russian peasant enjoys his kwas with bread. At the same time the digestive canal must not be burdened with work at this stage. begun with pleasure. both from instinct as well as medical direction. acids action upon all the constituents of the food. for example. therefore. and is consequently always present. Moreover. either as bread or porridge. for instance. nature itself constantly the stomach in addition to the The former arises from the food introduced. especially in the acid form. These facts are all physiologically comprehensible when we know that an acid reaction is not only necessary an efficient action of the peptic ferment.
which do not excite the gastric glands. There is no struggle in this and therefore no one of them In harmony also with daily experience. especially when and but little acid. cases. fatty food. and in the case of Aveak stomachs heavy. Everybody knows that fatty foods are that is. and is given in cases of weak digestion as well as in a whole series The of severe illnesses. in comparison with nitrogenous equivalents of other foods. consequently.142 THE WORK OF THE DIGESTIVE GLANDS. as might a priori be inferred from its wide employment. or fat as emulsion. are now in a position to understand this they are usually avoided. as may be seen from the fact that large quantities of lard are consumed in certain districts of Russia with impunity. cases of weakness of the stomach. but little gastric juice while the fat which excites the pancreatic gland itself ensures a rich production of ferment both for and also for the Fat alone does not count by any means starch and proteid of bread. since the inhibitory influence of the fat in this case does not prevent the digestion of any other food-stuff. As we already know. by lessening the acidity of the gastric juice we could reduce the activity of the pancreas. in suffers. In pathological however. for itself. in heart and kidney affections. which we have so strikingly seen in our experiments. Amongst and this is all the articles of human food. a combination of fat and proteid-holding foods is particularly difficult to physiologically. extreme importance of this substance. No upon less instructive is fat. unanimously recognised. or even in some general disturbances of the digestive apparatus. difficult of digestion. milk takes a special position. &c. milk which secure it an exceptional position. both in daily experience and in the By everybody milk is considered a light food. the weakest . This also is comprehensible. dietetics We of fat in large quantities in the chyme interest the further secretion of gastric juice. restrains in its The existence own and can only be borne by those who have good stomachs and The combination of bread and butter is less difficult. and thus impedes the digestion of proteid substances . a comparison of the results of our experiments with the demands of instinct and also with the precepts of and therapeutics. the physician. for example. we can now well understand. game. totally excludes fatty food and recommends meat of a fat-free kind. case between the several food constituents. for example. where an excessive activity of the gastric glands is And here manifested. a food prepared by nature There are three properties of itself. and these are matters which might be made use of in various special diseases. as a heavy food. calculated per unit. Bread requires digest. medicine has empirically brought to its aid the restraining action of fat. On the other hand. practice of medicine. keen appetites. is prescribed. and is con- ducive to the assimilation of the fat itself.
than in the case of milk (Experiments of Prof. It gets excites not only a really The secret of the relation of milk to the secretion of the digestive juices can. and which is probably only an expression of the first. which the organism pays for the nitrogen of milk. If the hourly rate of absorption and which milk and bread are respectively used up be taken into consideration. We of importance for the inhibition and the alkalinity on the other for the restraint Thus the gastric glands and the pancreas. when it is introduced unobserved into the stomach of an animal it causes a secretion both in the stomach and also one from the pancreas. Ejasanzew) consequently. however. at present be submitted to no are at liberty. milk possesses a further important property. notwithstandof the pancreas. unfortunately. consequently the secretory activity requisite for its In addition. be accepted that milk effective. appears to be an independent chemical excitant of the digestive and in this action it is remarkable that we perceive no essential in the effect difference when is the milk is the stomach from that which occurs to lap. it results that the increase during the first seven to ten hours after the milk (compared with the excretion beforehand) amounts only to from 12 per cent. must be expressions of the functional activity of the digestive canal itself. while after bread. and that of this activity in the case of bread is three or four times greater . Thus. secretion. of nitrogen. 143 gastric juice and the smallest quantity of pancreatic fluid are poured out on milk. consists in the When one administers to an animal equivalent quantities following. therefore. to 15 per cent. consequently it canal. a matter which is in every way is suppose that the fat on the one hand of the gastric glands. in the case of milk a much larger fraction of its nitrogen is free to be used up by the organism at large (irrespective of the organs of digesIn other words. are maintained by milk at a certain ing but not too high degree of activity. and afterwards estimates the hourly output of nitrogen in the urine. it is by no into the stomach. assimilation is much less than with any other food-stuff. in the one case as milk. the third characteristic which is observed to belong to milk. the presence of excitants. to further analysis or investigation. when it it is brought unnoticed into given to the animal Although means a matter of flesh a better chemical indifference how must. Finally.A NEW TEST OF THE NUTRITIVE VALUE OF FOODS. excitant. of the nitrogen taken in. and also that the appetite is unable to stimulate this secretion into a more active or abundant flow. desirable in consideration of the easy digestibility of its constituents. in the other as bread. 50 per cent. the price tion) than in that of any other kind of food. in the form of work . it has to be admitted that these augmentations it amounts extent to the to urinary nitrogen which appear soon after feeding. but at the same time a very economic.
for instance. those food-stuffs are less useful whose nutritive value little . much ! less than that for other admirably. to be proved. of view those materials must be taken as less nourishing and le?s point digestible. that in the digestion of a given food the alimentary canal has been given a certain work to perform if it be in . how much nutrient material was contained in its digestibility You will the food. for example. Fibrin. a good deal of other non-nitroThe digestibility and nutritive value of foods must genous material. boiled and roast meat. The energy used up in gland metabolism must be deducted from that of The remainder will then indicate the value of the food food taken in. to the organism. although milk contains. both in regard to the quantity and quality of the juices poured out on a given amount of nutrient material. is on the part foods.144 THE WORK OF THE DIGESTIVE GLANDS. more than covers the cost of their digestion. cannot alone be trusted to solve the question in a satisfactory manner. that is to say. The older must frankly make room for the new or else be displaced by them. Suppose. therefore. &c. consequently it is of great practical importance to compare from this aspect the same foods. in which food is normally partaken of are quite from those in the test-tube. obviously be decided by an estimation of the real work which they entail upon the digestive apparatus. hard and softdifferently prepared boiled eggs. is clear from the observation of Dr. health. where we have to deal with only one juice. boiled and unboiled milk. Kor- experiments settle the question of digestibility. regarded by all as the most easily digested proteid. for experiments different tinction. when compared with a nitrogen equivalent of milk. In your experiment you do not the alimentary canal to extract ean artificial digestion know how all great an effort it has cost the nourishment from the food. make a disfood constituents. of its digestive apparatus. with complete abstraction of everything nutrient. will give the amount available for use by all From this the other organs exclusive of the digestive apparatus. which is in addition to nitrogenous substances. as well as what is absorbed into the body fluids. but the question of remains as obscure as before. that thus learn to say. as a matter of fact. . Walther in our laboratory. the is work will be accomplished in the best possible manner. and not with the interaction of different juices and different That one must here. which are in large part used up to make good the expenditure entailed by their digestion on the part of the alimentary canal that is to say. a much stronger excitant of the pancreas. Experiments upon the utilisation of food-stuffs. the food prepared by nature disitself when compared with all others tinguishes The facts just related bring forward a new aspect from which the relative nutritive values of different foods criteria How may be judged. in which what remains undigested is determined.
however. the gaps in the experiment were happily made good by the clinician. received daily during the course of several weeks an addition of soda to its food. Consequently. that these salts promote a flow of gastric juice. on supposed physiological datn. the effect is due to the fact of are succagogues. I may relate the following observation had survived the performance. yet it often it turns to the explanation of facts. to : and of the alkaline salts of sodium.EXPLANATION OF THE EFFECTS OF ALKALIES. pharmacological. recovery. a^pancreatic fistula. medical science has fallen into error. and has returned to normal conditions. and not to a direct influence of the alkalies. they . That sodium salts (the chloride and bicarbonate) are useful in disorders of the do they act. concerns the therapeutic use of the neutral and alkaline salts of sodium. The assistance afforded by the however. one after the other. however. for any experimental foundation to support We this doctrine. It appears to me that here. both on the succagogue influence on the part of these salts. they began with methods either false in themselves. This means. This latter. When The experiments brought forward cannot be regarded Blondlot sprinkled sodium bicarbonate upon flesh. 145 A discussion of some further medical questions The first may here be taken up. In clinical. It frequently tries to explain complicated healing processes in the And this is true in the simplest way. as conclusive. that it has recovered from a disordered state. for the experiment appeared to be confirmatory of clinical experience. which is exactly the opposite of that generally accepted. must be specially proved. however. look in vain. When we know that an effect takes place it does not by any means imply that occurs. We effect. and although medicine How we know is the mechanism by which it broad enough and comprehensive empiricism in practice. sometimes begins to secrete a greater quantity of juice. ever. The animal K . which affords an example of prevalent medical reasoning the alkalies work favourably in digestive disturbances therefore. and physiological text-books it is may stated now. however? digestive apparatus there can be no doubt. or far removed from normal conditions. and an cosophagotomy. Naturally the stomach. chloride solutions directly or Braun and Griitzner introduced sodium into the blood. were unable to convince ourselves of any Indeed. under the influence of alkalies. howexample. In this case. that which is ordinarily given. alkalies for the organism might be capable of another explanation In this case. stomach and pancreas they proved in our hands to have an inhibitory In addition to the experiments which I previously brought : forward concerning the relation of alkalies to gastric and pancreatic A dog which fortunately juice. as in some other cases. as ever. I venture to offer a reason for the effects of sodium chloride. of a gastric fistula. . free enough to make use of thinks in narrow grooves when present case.
when an affection is once set up by this or that cause. and thus to a restoration of the normal state. that is. Without doubt the digitalis aids by breaking through this vicious this leads (from known the heart. is generally prescribed. suppose that. In this consists. weakly acid gastric juice. With regard to the salivary glands. which means that a certain amount . of lowered activity of several digestive glands. of recovery. contrary to the ordinary were hardly at all insalivated. the digestive glands. therefore. with the use of the salts in question. pancre- and salivary glands. from the upper end of the oesophagus. the circumstance was naturally submitted to closer I believe that the inhibitory influence of the alkalies on investigation. The due nourishment. physiological causes) to an increase in the number of beats. further. a greatly viz. When. may in this way contribute to the removal of the pathological condition. a it remedy effectively restrains the excessive work of a diseased organ. proceeds best during rest. in many cases the affection begins with a hypersecretion. the quickening leads to weakening of the organ. atic. during which the latent work of restoration is accomplished. An uncompensated heart beats rapidly. and thereby gives new With our explanation of the action of the alkalies power to harmon- a strict diet the further circumstance that. therefore. that is an abnormal excitability of the secretory apparatus which makes flow. which was here proved experimentally.14 G THE WORK OF THE DIGESTIVE GLANDS. The weak action of the heart lowers blood pressure the lowering of A . One might draw a parallel between the action of these substances in digestive disturbances and that of digitalis in compensatory disturbances of the heart. the gastric. the relatively small effect of At otherwise very active juice-exciting procedure at once struck the same time we observed that the pieces of flesh which rule. and thereby only aggravates its condition. for continuous activity has undoubtedly a harmful influence on the glands.. characterised by an incessant or very protracted secretion of slimy. such a condition sets in after operations It is. In the normal course of events. in my opinion. after a period of active work follows a pause. simultaneously existed. it may later maintain itself independently. of vicious cycle is set up. When the first carried out. sham feeding experiment was this us. fell enjoyed good health and had an excellent appetite. the healing effects of the alkalies. Its time of rest. itself evident in a superfluous and to useless of The same must be conceived : happen in disorders the pancreatic gland performed for at least. Further. In this dog. and the restoration of organs after activity. justifiable to physiological purposes. ises cycle in that it greatly slows the pulse. restitution of the organ is shortened. may furnish a basis for the following representation of their mode of action in Catarrhal affections of the stomach are producing healing effects.
the erroneous belief that alkalies promote a flow of juice obviously lies in this. such. it appears to me. and is well borne and useful. Chigin). Every food determines a certain amount of digestive work. especially from a sparse to a rich diet. instituted if a change be suddenly made from one dietetic regime to and another. Finally. clinical investigations of a restraining effect. which from some : new demands made upon them. Consequently. In one and patients react to the same diet in wholly That which is agreeable to one. contribute to a clearing up of the situation. definite fixed types of glands are set up which can only slowly and with In consequence. that people omitted to compare the effects of the saline solutions with those of like quantities of water mention being now more frequently made The cause of (Dr. illness. different the same different ways. with the stomach-tube. may be rank poison to another. and when a given dietary is long continued. The second point which we may consider chief difficulty of the physician who wishes is the The following. for instance. it may be of some use cause or other has been excessively and abnormally stimulated. These disturbances are expressions of the temporary insufficiency of the digestive glands to meet the to relate the following here There are often cases of sudden and unaccountable digestive disturbances From the standpoint of modern physiology they might be explained by an activity of the secreto-inhibitory nervous system. a new phase has also set in.THE GOLDEN RULE IN of rest is DIETETICS. to regulate the diet of patients when they suffer from digestive disturbances consists in the fact that idiosyncrasy plays a very important role. account. does all this indicate ? Till now physiology had no experimental answer to the question. digestive disturbances are often difficulty be altered. the golden rule in dietetics is to give no directions with regard to food till one has made What inquiries concerning the inclinations and habits of the patient. after a period when the alkalies were looked upon as succagogues. as happens after the long Russian fasts. In any this system is now a factor of which the physician has to take due case. But our facts. 147 It is interesting that in secured for the digestive glands. .
L. 0. N. G. and A. L.wccvs en- but only of watery constituents Pancreatic juice a specific The favouring influence of avccvs entcrlcus upon the trypsin of pancreatic juice depends upon the transformation of the zymogen The amount of trypsin in the form of zymogen depends on the into trypsin nature of the diet The spleen and the pancreas The propulsion of food excitant of the kinase along the digestive canal its functional adaptation thereto Experiments on the movements by which the stomach is emptied of its contents The .LECTUEE LATER RESEARCHES THE COLLABORATION AND MUTUAL RELATIONSHIPS OP THE DIGESTIVE JUICES BILE AND SUCCUS ENTERICUS MOVEMENTS OF THE STOMACH PATHOLOGY AND EX- PERIMENTAL THERAPEUTICS OF DIGESTION THE METHOD OF EXPERIMENT IS THE ONLY ONE ADEQUATE TO PRESENT-DAY MEDICAL REQUIREMENTS. N. A. A. Nagorski. C. D. H. It was also dedicated to the memory of S. Wulfson. the following collaborateurs J. C. 0. Lobassow. A. Walt her. 8. Bruno. Sawriew. Kladnizki. Sokolow. Krewer. The lecture was afterwards published as a separate pamphlet under the title. Schepowalnikow. P. Botkin in the names of Professor Pawlow and of J. Mechanical excitation its effects a secretion of . G. P. : N. Wolkowitsch. E. A. R. Popielski. Shirokich.} An indispensable Requirement of Medical Research. P. the is ENTEROKINASE iericiis. W. passage of * chyme into the intestine is regulated by impulses generated in This lecture was delivered at the anniversary meeting of the Society Russkm Medical Men five years later than those preceding. Soborow. W." . S. of salivary secretion Later researches on the digestive glands -The physiology and psychology New method for the separate study of gastric secretion The multiplicity of functions attributed to the bile is an indication of our lack of knowledge of the physiology of this secretion Method of studying the biliary secretion The entry of bile into the intestine is dependent on bile is to the presence and the nature of the food The true digestive function of the augment the activity of the ferments of pancreatic juice The svccus enterieiis has also the same function A ferment of ferments. P>. Hanicke. Glinski. J. " The Experimental Method ( Translator. A. N. Serdjukow. P. A.
but in addition we have the bile and the snccus entericus. and our later results have even a GENTLEMEN. I think it desirable to bring them all under consideration. nearly all cases been It may well be understood that even at this stage. the saliva. closer connection with the practice of medicine than the former. since this was already fully known. Further. In the case We direct our whole attention to an investigation of the factors which determined the normal activity of these glands. mainly with the more important of the digestive organs. still further at the physiology of these glands. but now these subjects have been made matters of direct research.SALIVA. of the gastric glands and the pancreas we were able to the question of the main physiological import of their secredisregard were therefore free to tions. and the pancreas. 149 of the duodenum Keflex closure of the pylorus by and importance The pathology of the stomach as studied by our experimental methods The protection of the mucous membrane of the stomach Physiological function of the epithelium of the gastric mucous membrane Experimental production of gastric asthenia and of a condition the reverse of this The point of attack of the round ulcer of the stomach Substitution of the small stomach for the large in our physiological and pathological experiments Nervous inhibition in aii'ections of the stomach Experimental therapeutics of the stomach based on our investigations Survival of vagotomised dogs Treatment acids. namely. The result? of the several investigations have in . At that time I could only theoretically discuss the pathology and therapeutics of digestion. we have also made obser- movements of the food along the digestive canal. However. Since the delivery of the preceding lectures. it was necessary to establish at the outset their functional importance. With regard to the glands of the stomach and also the pancreas the investigations of the past five years have confirmed our fundamental deductions while they have also added a greater completeness to them. the gastric During the last five years we have worked glands. vations on the included that of the remaining digestive secretions. carried on without interruption. The beautiful adaptation to the requirements of the food undergoing . purety physiological questions have taken the first place in our researches. and that each single piece of work will find its proper place in the whole scheme. the facts concerning which I am about to speak will be already known to many of you. its significance membrane of gastric hyper-secretion by alkalies. But in the case of the latter fluids. now five our investigations into the physiology of digestion have been years ago. BILE the mucous AND SUCCUS ENTEMCUS. communicated to this society consequently. but our field has My former lectures dealt naturally grown ever wider and wider. It is only in this way that the general bearing of the investigations can be made clear.
Further. has been even experiments. we have that both Russian and foreign representatives of clinical medicine have made use of the results of our physiological investigations and with good effect. I now come to a new subject in our researches.150 THE WORK OF THE DIGESTIVE GLANDS. We have determined the relationship of each individual Pojnelski. Walther. 0. and this encourages us to extend our work. that is to bears a definite relationship to the requirements which must say. Everything does not mean advance we often meet with a standstill. which purport to sum up the achievements of our science. such as the act of mastication. mixed with the proteid. Schiff. endeavoured to solve the interesting and natural questions of how much and what kind of saliva is poured into the mouth mider different conditions. thermal. an essential characteristic of the more clearly expressed in our later digestion. determined by the fact that starch Lobassow. element of the food to the receptive part of the nervous apparatus of the glands. ifec. that every form of stimulus applied to the cavity of the mouth. excites the . and others. L.) . the entry of foods of different tastes and different It resulted and the same is to be expected degrees of dryness. And yet we read in modern from every is organ in the body that the work of of itself to text-books of physiology. after the appearance of the classical work of Mitscherlich. been able to explain nearly (Dr. (Experiments of Dr.) Further. Thus. is all the quantitative and qualitative changes in the secretions of the digestive glands which occur during the whole It is also very gratifying to me to be able to state period of activity. a number of talented investigators. in which we see that what is true of science generally is likewise true of every individual scientific problem. the fat-splitting ferment predominates. with a diet containing a large proportion of fat. and so on. all our hypotheses and discoveries have gained wide recognition in foreign lands. example of this is . sometimes even take a retrograde movement. be fulfilled by the saliva produced.shown in the investigation of the salivary glands. which we saw then to be work of these glands. the salivary glands capable adapting given conditions. be it mechanical. we have gained a better knowledge of the mechanism of this adaptation we have discovered new factors in the nervous machinery of these glands nerves which restrain their activity. About the middle of the last century. with is its strong digestive power. especially in connection with the pancreatic secretion.) Similarly. A striking . J. such as Claude Bernard. The ferment activity of the pancreatic juice harmonises in the most beautiful manner with the nature of the food to be digested. or otherwise. chemical. And in this way we have convinced ourselves by direct experiment that the special nature of the gastric juice poured out on bread. (Experiments of Dr. B.
be poured. and we could not The experiments readily give our assent to the contrary representation. and that they respond in haphazard fashion to every form of stimulus. The secretions of the sub-maxillary and sub-lingual glands were jointly collected. bitter and caustic substances saliva. or washed out of the buccal cavity. Wulfson. mind the accuracy of a sensation so finely differentiated as that presided over by the nerves of taste ? In the case of the deeper-lying glands. diluted. This explanation is fully confirmed will likewise by the absolutely constant and striking fact that a thin watery saliva. or at most only one or two drops. Again. without distinction. Thus if a handful of small quartz pebbles be thrown from a certain height into the dog's mouth. D. therefore. Glinski and afterwards continued S. because the sand cannot otherwise be got rid of than by a free stream of fluid. salts. the dog will gnaw them. if ice-cold water be poured or snow thrown in. a kernel of truth always of the earlier authors The utter inadequacy of any hypotheses to the contrary must. Upon all substances which the clog rejects for example. How could it come about that the work Where of the salivary glands should be robbed of its adaptation ? when we bear in is the reflex that could determine this. move them backwards and forwards. to be repeated. while upon eatable substances a slimy mucin-holding These experiments were begun by Dr. as to the why and the One is led to suppose that the work not one word. Obviously it is not required in these cases. without exception. special factor in the case. so that the mechanical stimulus is fairly strong. L. because. containing mere traces of mucin. be clear. This could not be ascribed to the occasional discrepancies in the observations of these different authors. as a rule. to the skin around the orifice. But cast in some sand. namely. is poured out by the mucous salivary be glands upon every substance. H. But. acids.* We easily convinced ourselves that it was by no means the case that every form of mechanical and thermal stimulus. that of the parotid separately. because these require to be neutralised. concerning salivary secretion had. During the experiments a small funnel (with a cylinder for collecting and measuring the saliva) was attached * by Dr. and yet no saliva flows.THE SECRETION OF SALIVA. is said ! of the salivary glands is of no import. into the mouth. sometimes swallow a few of them. we had previously learned many facts pointing indubitably to adaptation. in spite of such. 151 in the case activity of the salivary glands in one and the same way. and the saliva flows in quantities. however. Obviously the fruitful ideas had passed into complete oblivion. which requires to fluid is removed. no saliva will be seen. Only of a f ew authors has the -state of dry ness of the food received mention as a wherefore of all this. . those of the stomach and the pancreas. especially remained. They were performed on dogs in which the normal orifices of the salivary ducts were displaced outwards. excited a flow of saliva.
we find all the elements of what we usually " attribute to " mental activity namely. a watery saliva escaped from the mucous glands if food. when we pretended to throw pebbles into the dog's mouth. finally. for instance. choice. flesh much less saliva appeared than in the previous case. when we merely drew the animal's attention to the substances in question. when we offered this or that kind of food. proteids.152 THE WORK OF THE DIGESTIVE GLANDS. Further. For example. must now . For these. be proved by special experiments designed to exclude the play of In the psychology of the salivary glands. therefore. And if the food were dry for example. which I. as such. as it psychological factors. the physiology and psychology of the salivary glands have come to be associated together . we un" observations upon the " adaptation of this ferment. concerns some antitoxic precautionary measure. or. even more than this. can be unhesitatingly accepted where formerly only pure physiological relationships were thought of whereas the physiological influence. quantity of saliva flowed out. on the other hand. as the introduction of the stimulating substances into the buccal cavity. the psychology has in many cases displaced the physiology. proceed to another aspect of the question in the highest degree interesting. the psychological influence. If we pretended to throw in sand. a moist food was presented for example. however eagerly the dog may have desired the food. a secretion either immediately appeared. such. reappeared in exactly the same manner under the influence of psychological conditions that is to say. Thus. dispassionate . dry bread a large juice . in accordance with the properties of the substance which we had pi'eviously seen to regulate the quantity and nature of the when physiologically excited to flow. in a quite unexpected way. This peculiarity is is still Perhaps it Since ptyalin fortunately could make no almost absent from the saliva of the dog. even when it excited no special interest on the part of the dog. or. In the course of our it appeared that all the phenomena of adaptation which we experiments is saw in the salivary glands under physiological conditions. or to pour in something disagreeable. sensation. in some cases. a striking proof that the first of the digestive glands adopts itself to the physical conditions of the food. or to cast in sand. or it it did not appear. has displayed itself to us. Of special interest is the peculiar relationship between the parotid secretion and acids. the quantity of saliva secreted is The drier this is. the more closely related to the clryness of the food. bolus and facilitates its descent secreted which lubricates the food through the oesophagus. a saliva is always secreted which is particularly rich in saliva flows without an explanation. When. Thus. a slimy saliva. This latter effect is particularly obvious in the case of the parotid gland.
to renders every beneficial influence which acts upon. towards a synthetic study of the whole indivisible life. It is. even here. if we could analyse step by step the adaptation in the latter organ. It is quite clear that the activity of even such apparently insignificant organs as the salivary glands penetrates unconsciously into our everyday psychical conditions through sensations. In order to relative definitely settle some new juice. the physical and psychical being. for example. to the secretion of gastric we found as well as old problems it necessary to perform still more complicated operations upon our dogs than we had heretofore done. which seizes hold of one's whole mental existence. and finally the cavity of the large stomach was shut off from the duodenum by a septum formed in the region of the pylorus To permit of the dog being fed daily in of mucous membrane only.AUTO-REGULATION OF GASTRIC SECRETION. by means of such unconscious impressions that the usual physiological processes of our bodies are guided. We see no reason why the same should not apply to the other organs of the body. should like to go more fully into two conclusions to be drawn from them the one of a more practical. the ordinary way. desires. Hence. (Inoperated upon are exceedingly convenient to experiment upon. reacts upon the body and On the other hand. it is clear that the adaptation of the salivary glands is a phenomenon of the same order as that. A. Viewed from this aspect. Further. and thoughts which in their turn exert an influence on the work of the glands themselves. Thus a way is open to us. of the pancreas. Further. the other of a more theoretical nature. the gastric and duodenal fistulas were connected Dogs thus externally by means of glass arid indiarubber tubes.) . and 153 judgment with respect to the substance introduced These latter facts are of such importance that I into the buccal cavity. P. indeed. we have here a clear physiological scheme for the study of the development of psychological phenomena. Thus the following operations were successively In the first place an ordinary carried out on one and the same animal. Sokolow. a cheerful disposition tends to develop and strengthen the body by increasing its susceptibility to every invigorating emotion of life. With such animals it can easily vestigations of Dr. and then an isolated cul-de-sac of the stomach were gastric fistula formed. it an easy prey to every form of disease. the rationale is at once apparent which underlies the prevalent conviction that a deep and lasting sorrow. a duodenal fistula (provided with metallic cannula) was made. consideration. and if it be open to us to regard the process in the salivary glands as the more primitive form.
supposed uses. and of the of the different exciting substances. selective excitability of the mucous membrane of the stomach ? have only brought forward a few of the results which we obtained by our new method. the less is the real The reason of this is quite plain. prevents the further secretion of gastric juice when it has accumulated It is in any considerable quantity within the cavity of the organ. and would merely remark. the inhibitory effect of fat originates chiefly of the duodenal mucous membrane and not from that a With the same dogs we were able. and so But when we come to the bile we have to read whole series of on. I effects cutting out of the influence of the movements of the food. So it is learn the functions of the gastric juice. In every text-book one can straightway number of insignificant that we do not know its it. of the pancreatic juice. that the experiments on such dogs were characterised by an astonishing degree It might seem that the of accuracy and constancy in the results. gentlemen. that the greater the number of remedies recommended for a disease. of the promotion of peristaltic . moreover of the greatest interest that other acids. for example. without new form of auto-regulation on the part difficulty. be applied to our knowledge of the organs of the body. and perhaps will not even then learn Mention will be made of the emulsification of fat. which It appears that the acid of the Butyric acid. on the phoric. When a functions are assigned to any organ.154 THE WORK OF THE DIGESTIVE GLANDS. from these parts of the digestive canal.. of the moistening of the intestinal wall. phos- stomach. we may The same criterion possess a really good remedy we want no other. and conclusively be decided that the second or chemical period of gastric secretion is determined by a reflex proceeding mainly from the inner surface of the stomach. a full secretory effect is only obtained from the In the latter the secretion is very small stomach in the former case. strongly connected with the fact that hydrochloric acid strongly inhibits butyric What could be more striking as an instance of the acid fermentation. it means real function or have not properly appreciated with the bile. excites gastric secretion. You all know. in addition. do not exert this inhibitory action. the other hand. I turn now to the bile. would make it otherwise. When efficacy of any one of them. sentences about its its chief function. scanty. a matter which is probably contrary. On from the surface of the stomach. &c. If chemical excitants (solutions of meat extract and so on) be alternately introduced into the large stomach and into the duodenum. to demonstrate concerns the secretion of hydrochloric acid. but this was not so.
otherwise it would not be poured into the intestine at such a remarkable situation. b. Further. it is conducted from thence In all cases. applied. .). It because of this that a special reservoir is provided for it in the shape of the gall-bladder. since. the bile continuously flowed from the gall-bladder whether the dog had been fasting or fed. And this is easy to understand. Consequently. was fastened with sutures. of the intestinal 155 action. cannot doubt that the bile is necessary for digestion. See Fig. ' (Experiments of The operation is not an easy one. that it plays an important rule in this process. namely. a receptacle where the bile (continuously bile The formed by the liver) is temporarily stored up till required for use. the loop was stitched into the abdominal wound. 18 (a. the academic teacher will have little or nothing on. In the first instance the orifice of the duet. Are all clear and incontestable to demonstrate concerning the bile. has been collected from the most widely different animals. After some preliminary trials we adopted the following procedure. of the precipitation of the proteid bodies. however ? In how far is the one essential and the other not ? To such questions you will find no satisfactory answer. with a piece of the surrounding mucous membrane. although the bile in the first instance enters the duct. This done. a fluid the And yet we appearance and composition of which are so special. But how are we to set ? functions of the bile are One about determining what the digestive of the most direct ways consists in examining how much and what kind of bile is poured into the digestive It is remarkable that this method has never been correctly canal. was cut out of the intestine and transplanted upon its serous coat. but always by means of a fistula leading into the gall-bladder. different conclusions. although numbers of physiologists have worked at the subject. since the formation of bile in the liver and its into the gall-bladder. where it of intestine at the point in question it healed.FUNCTIONS OF THE BILE. and so these correct. in order to determine what the function of the bile during digestion enters the alimentary canal and not how is. both during digestion as well as in fasting. It was on these lines that we proceeded. after deviating the natural orifice of the bile duct (with a piece of the surrounding mucous membrane) towards the exterior.* study of the results showed that the entry of bile into the intestine is regulated by the same laws A that govern the flow of the other digestive juices. The experiments have led the authors to very employment during digestion are naturally two is different things. of the excitement villi. of the disinfection of the intestinal contents. Experiments performed on dogs with an artificial opening into the common bile duct differ little from the foregoing. r. one must observe how accumulates in the it it gall- bladder. where the acid peptic gives place to the alkaline pancreatic digestion.
But eaten. nor boiled starch paste.} drop of bile entered the intestine. Bruno and Dr. N. a flap of skin long as the digestion lasted. 18. but with definite fluctuations in quantity and quality. One cannot has quite as important a part to play in the chemical elaboration of the food as any of the other digestive Hence we proceeded further in our investigations. raw directly into the stomach. or brought them It resulted that neither water. G. Kladnizki. and gave the juices. dependent upon the nature of the food. the flow began at a certain definite time afterwards. Showing the different stages in transplanting the (c) orifice of the bile is duct outwards. on the other hand. Fat. G. resist the idea that the bile ingredients of the food separately to the dog to eat. as well as the . In represented in situ over the bowel. N. In the fasting animal not a Dr. The fluid continued to flow as when the dog had FlG.156 THE WORK OF THE DIGESTIVE GLANDS. whether in solid pieces or as a thin egg-albumen fluid. acids. caused a flow of bile. This time is different for the different kinds of food.
carried out. it Different investigators have long ago shown that In addition. The bile. Further. Williams and laboratory Martin. Further. open. however. which have the effect of causing it to flow into the intestine. we have discovered the main Numerous experiments on dogs. in systematically examining the variations ferment and the relationships which they bear to digestive activity. however. But. but are this action is only a weak one. begun originally by Professor Nencki in his Berne and carried on by Heidenhain. fact that in print. less . which were only increased about twofold. resembles other up digestive juices in that it possesses its own particular combination of excitants. extractives of meat. have shown us that when a definite quantity of pancreatic juice is which varies for the different ferments. there are very old experiments which point to an inhibitory action of the bile upon. do not enjoy great popularity amongst physiologists. and only with those of the pancreatic bile We shall see that in the favouring action of upon the ferments feature of its systematically of bile. ferments. digestive importance. all of them. therefore. the ferment of the gastric juice. therefore. It was ments dealt. It has long been known that the ferments of the stomach and pancreas manifest different degrees of digestive activity. The lack of esteem from which they suffer is shown by the But in what does the work of the bile consist this question. and consequently in the main with zymogen and not with the ready-made ferment. to doubt whether they would hold good for the actual Bachford alone carried out experiments with conditions of digestion. Dr.FUNCTIONS OF THE BILE. added to pan- creatic juice. the action of which was increased two to three on the other two. but n^t with juice of the rabbit. with extracts of the pancreas. ? In order to answer which now took a definite shape. ferment. has a slight amylolytic action. The bile. Rachford. investigations. have shown that the bile has a favouring action on the fer- The majority of these experiments of the pancreatic juice. according to the chemical properties of the medium in which they work. set 157 and the products of the digestion of egg-albumen. many The bile possesses text-books of physiology they are consigned to small only a weak direct chemical action upon the food constituents. during the past year. indeed. it produces a constant and decided accentuation of the The effect was most pronounced on the fatactivity of its enzymes. on the other hand. There remains the possibility of a chemical effect upon the other juices with digestive which the bile mixes in the intestinal canal. we adhered to certain facts which. a free discharge of the fluid. splitting fold. Gegaloff discovered in this laboratory that the bile of carnivora also its contains a proteolytic ferment. it must be admitted. We of now engaged. it was observed that this augmenting action showed adaptive variations dependent upon the nature of the food.
cases. c 1? after bread. secretion (upper series) and the entry of bile into the intestine (lower series) . we may with certainty conclude that the chief duty of the bile is to . 6. The experiments dealing with the inhibitory action of bile on pepsin We have further deterhave also been repeated and confirmed by us. a^ after the ingestion of milk . In comparing the curves. or. after flesh. their general form only is to be taken The scale of the ordinates was different in the different into account. 19. 6j. mined the extent of this influence.158 proved THE WORK OF THE DIGESTIVE GLANDS. their mixture beforehand in a common excre- tory duct. I am also able forward another striking proof of the close relationship between the bile and the pancreatic juice. is not without a purpose. intestine under the striking. Their similarity is most Is it not obvious that the two fluids have a reciprocal chemical relationship towards each other. same conditions of diet. and have come to the conclusion that it must have a definite When we link all these facts together physiological significance. and in consequence must act hand-in-hand ? Their discharge at the same place into the intestines of many animals. using for our purpose pure gastric juice. a. I pray to examine the following curves in which the hourly rate of the you secretion of pancreatic juice is compared with the entry of bile into the a Curves representing in each case the hourly rate of pancreatic Fig. as is often the case. itself to juice. a fluid which be a constant and powerful auxiliary of the pancreatic is of such importance for digestion and in itself to bring already so complicated. and c.
In the case of the latter. was. the one contains pure pancreatic juice.SUCCUS ENTERICUS. the importance of the succus for which they entericus as a digestive fluid a fluid long known to physiologists. particular the fat-splitting ferment. This research was started with the same question which proved to be so fruitful in the case of the bile. the increase often reaches to an astonishing degree. He who has once convinced himself of this by experiment will never doubt for a moment that this accentuating influence is the most important function of the succus entericus. not powerfully aided than either of the other two. the other a mixture of pancreatic and intestinal juice. Judging from extreme opinions. P. its existence was even doubtful. arrests the action of the pepsin. I think it necessary to demonstrate to you the fact itself. three pieces of fibrin. Now while in the second. But quite recently it has been the good fortune of our laboratory (experiments of Dr. or at least to it. since closer ? showed very strikingly that in the favouring influence of investigation upon pancreatic juice. and all fluid is an inverting action on sugar. Pieces of fibrin of equal size are placed in the two vessels. but had discovered no precise use. that regarding the physiology of the succus entericus. in the first vessel the solution of fibrin has . therefore. In view of the novelty and importance of our facts. Only in one particular It was denied are investigators agreed namely. and more especially the proteolytic. the fat -splitting ferment was much more It was. Still more indefinite and unsatisfactory than the teaching concerning the digestive functions of the bile. N. It was only considered to have a weak solvent efl'ect on starch. one after the other. have been fully digested before your eyes. Our anticipations have been fully confirmed by sesses the striking capability of The succus entericus undoubtedly posaugmenting the activity of the pancreatic ferments. Upon a screen the shadows of two vessels are thrown. to the pancreatic juice bile Does not the succus entericus also function as a second adjuvant This seemed all the more likely. in a similar manner to the way in which the bile influenced the fat-splitting. discovery. I. all digestive influence may. to elevate to a high position. Sckepowalnikow) at one stroke.VJ It facilitate the transition from gastric to intestinal digestion. that the digestive action of the supposed to be very slight. and favours the ferments of the latter . be authoritatively said that there is not a single point connected with the physiology of the succus entericus which has not at some time been contested. which is injurious to the ferments in of the pancreatic juice. unlikely that the succus entericus would be found to actuate one or other of the remaining ferments. until recent times. indeed. almost of insignificant importance.
on the contrary. a secretion of succus entericus would be unnecessary at a place where the food could only arrive some minutes or even some hours later. the two fundamental features of digestive activity. Sawilsch) that a tube was introduced into the fistula. it was found (experiments of Dr. a wide field was open to us in which we could study the most subtle problems of adaptation on the part of these combined fluids to the substances undergoing digestion. Our preliminary experiments therefore afforded that in the study of the chemical action of the three hoping ground united fluids. therefore. adaptivity and response to a specific stimulus. for entericus. stood out more prominently in connection with the succus entericus than with any fluid. discovered a ferment. The secretion of the succus entericus seems also to follow laws of its it own. taught us to steadfastly believe in this specific character. and the succus entericus other when afterwards collected in separate portions. not for this or that constituent of the food.160 THE WORK OF THE DIGESTIVE GLANDS. only just begun. is most increased by intestine. us. When the secretion produced in an isolated loop of intestine under the stimulus of the cannula is studied for several consecutive hours. For the latter. activity in very small quantities. a specific secretion mainly of water and not of kinase. namely. destruction by boiling. had. Indeed. stimulus had to be sought. and . in so far that is a purely local secretion. and this stimulus was ultimately discovered in the ferment constituents of the pancreatic juice. of everything learned in connection with the other secretions. From cur previous experience the fact appeared quite extraordinary. the bile and the succus the duodenal secretion. A flow follows only in that segment of the alimentary canal which is immediately stimulated. I propose to give it the name of Enterokinase to distinguish other possible ferments of a similar kind. especially in some cases. moves very slowly forwards along the digestive canal. both by the secretion of the duodenum and by that of other portions of the small The proteid ferment. This fact has obviously a rational significance. namely. and convinced us that in this case we were dealing in point of fact so on with a ferment. Wherein lay the Why ? review of all the facts. was a puzzling question which long troubled specific the mechanical stimulus should here prove to be efficient. The application of the usual tests for ferment action namely. but a FERMENT OF OTHER FERMENTS. the pancreatic juice. the amount of kinase in the secretion became steadily less and less obviously the tube excited a . and the belief has not nature of the stimulus A that we had deceived us. Thus. Since the food mass. it We from I may renmrk that the activity of the fat-splitting and amylolytic ferments of the pancreatic juice is promoted equally well.
Wassiljew and Jablonsky likewise saw a very weakly acting juice in normal animals. But. 161 when at length the juice contains little or no kinase. have fully confirmed the pancreatic juice. only dissolved fibrin after four to six hours in A the thermostat. pancreatic juice has not this effect. and had not even attacked coagulated egg-white after ten hours. hypothesis. It was therefore con- physiological cluded. that the proteolytic ferment in these cases. learned of analogous phenomena in the case of the salivary already We One may further assume that the severe diarrhoea which glands. or to all of them. was secreted in the form of a zymogen. and excites a secretion of water. essential fact appears that their assistance is variable. It was therefore conceivable that the operation might have caused some injury to the gland and have brought about an abnormal condition in it. the secretion of the fluid part is more sharply separated from that of the ferment than anywhere else. The experiments of Dr. What is and inconstant the meaning of this assistance. if now a few cubic centimetres of pancreatic juice be poured in and left for half an Boiled hour. acts as a crude indigestible foreign body. obtained from a temporary fistula in an "acute " experiment. being now greater and now different less. often acts very weakly or not at all upon proteids. we may is take it that every cannula which is introduced into the fistula. SchepowalinkoAv that the kinase worked all the more energetically tho weaker in general was the pancreatic juice. I must further add that this peculiar physiological reaction to the ferments of the pancreatic juice (whether to one only. secreted immediately after the formation of a fistula. remains an open question) From the facts here communicated one extraordinarily sensitive. Lintwarew. and why is it so ? It was long ago observed that pancreatic juice. and the coagulated egg-white in L . who started with this idea. have merely with the object of being washed along the intestine. must therefore conclude that in the case of the succus entericus. as has already been related in the second lecture. occurs in certain acute forms of enteritis results as a consequence of some powerful impulse to of the intestinal irritation of the this water-secreting glands. a fluid will afterwards be secreted having much kinase. for some reason or other. With regard to the watery constituents. The flow is and cleansing function here excited by the extreme mechanical or chemically injurious contents of the bowel.PANCREATIC JUICE AND SUOCUS ENTERICUS. Hence the bile and the succus entericus have revealed themselves to be adjuvants of the And in their action the pancreatic juice. under the influence of certain diets. there was obviously a purposive It was remarked in the experiments of relationship. In this latter. But on the addition of some succus entericus the fibrin was dissolved in three to seven minutes.
has its ferments. Hanicke. the juice was an extremely active one. while one which has been actuated by succus entericus. for some reason or other it often slightly diminished the proteolytic action of the fluid. between albumen on the one hand. But whether. which might possibly be injured by the other in its ripe and very active form. that such a protection of the ferments is chiefly required in significant the gland-lumen and along the passage to the digestive canal. preserves the activity of its ferments even after A several hours in the thermostat. With dogs. Our in- power of the juice and in their became still greater. however. A. milk. on the contrary. proved also by the experiments of E. and so on) the gland furnished a juice with a very weak initial effect upon proteids. The relationships of the amylolytic ferment were. The same result was obtained with the juice which first flowed from a permanent fistula. and indeed to investigate the fact that an antagonism was often observed necessary. It This latter was was further shown that the promoting effect of bile upon the fatsplitting ferment likewise depended on definite conditions of diet. the zyrnogen condition of the fei'ment almost wholly disappeared after a short time. A. and the succus entericus had no augmenting effect. because possible dependence upon in this way we hoped to be able to discover the meaning of the remarkterest in the normal variations of digestive diet. which were fed chiefly on starch and fat (bread. the action of which was rendered very energetic by an addition of Obviously the zymogen stage of the ferment predominated here. the appearance of the proteolytic ferment under different forms depends upon definite conditions. That is Mett's tubes in from three and a half to six minutes. where the proteicl ferment is made active by the kinase. mainly containing zymogen. The experiments of E. or contained trypsiu to begin with. juice. and the actuating influences of the succus entericus and the bile on the other. confirmed this.162 THE WORK OF THE DIGESTIVE GLANDS. the proteicl fei'ment is excreted in a latent form. indeed. Thus. the augmenting action of spleen extract upon infusions of the pancreas first observed . therefore. The juice contained then only trypsin. It is especially the amylolytic and fat-splittirig. Hanicke have. the presence of proteid food and of bile. the newly discovered kinase soon found useful employment. however. in able relationships in question. The idea at once occurred to us that. however. under similar circumstances. Additional experiments are also desirable. with the object of protecting the amylolytic and fat-splitting ferments. In the bowel. succus entericus. less obvious in this respect. in the case of starchy and fatty foods. namely. to say. we found that. As a matter of fact. new conditions arise which protect the fat-splitting and starch ferments . dogs fed exclusively on flesh. Indeed. Consequently. very easily destroyed.
The discovery of this and the possibility which it affords of a real synthetic construction of digestive processes. the food is to be found at any given amount. To constantly remember that and quickly parts of the field organisms work together sheds a bright light over the special under review. especially in his latest contributions. would probably be very applicable. the artificial own work. moment. My must. But what was the known physiology of the motor functions of stand the variations in the curves i:5 the digestive canal able to do for us in this connection is ? How extensive this section of physiology ? in its investigation How methods have been employed many nerves have been stimulated and how ? How many . 1GB by Schiff. Herzen. in what and in what condition. esteemed colleague. Naturally these tive juices new results concerning the interaction of the digesall our earlier experiments upon the the pancreatic ferments under different compel us to repeat production of quantitative conditions. and by myself in the digestion of fibrin. Popielsky with the aid of proteid in tubes. be convinced of this when I say that the pancreatic juice of a splenectomised animal digested a quantity of fibrin in thirty to forty minutes at thermostat temperature. who firmly believes the contrary. I at the same time venture to indicate that a method if of procedure similar to that which underlies our employed in other of physiology. the freshly collected pancreatic it .PKOPULSION OF FOOD ALONG THE CANAL. that we can at once distinguish the accidental from the essential. the question of the In our analysis of the curves of secretion of the different digestive PROPULSION OF THE FOOD ALONG THE ALIMENTARY CANAL became more and more pressing. It is only when we are able to bring into view the whole train of normal occurrences in this or that portion of the organism. by Herzen. as has already been shown by Dr. from the natural. juices. Might relationship. I think. and departments lead to fruitful results. juice from animals previously deprived of their spleens (dogs and cats) contains large quantities of proteolytic ferment in the form of trypsin. is without doubt exaggerated. Hence the chemical agencies of digestion form an alliance of a complicated nature in which the individual members are linked together mutually to relieve and support each other. then more exactly investigated by Herzen. I would like to designate as the most important general result of the work of our laboratory. and still more has any significance. and does not correspond with the facts of the case for. all detect bygone errors. find out new facts. In order fully to underit necessary to know where. which filled a test-tube fifteen centimetres high and 1'5 centimetres in diameter. I am unable to recently confirmed by Pachon I wish only to remark that the importance which is ascribed to say.
no acid be injected into the duodenum. because upon the food. who simultaneously discovered that the passage of food from the stomach into the intestine is quantitatively regulated from the upper segment of the latter. and have already yielded interesting In the first place. Mering). a reflex act is set up which temporarily occludes the . and a solution of soda be poured into the duodenum. take place. If. in German such a way that the expelling movements of the stomach are temporarily inhibited by a reflex from the duodenum. This cannot be accounted for by a reflex mechanically called into play. same lines. we have observed that the is not prevented. the one is moved quickly and the other slowly. and from what combination of separate elementary conditions is this propagation of the food effected. Serdjukow} that the duodenal mucous membrane. But as to why the one kind of food is held back while the other is moved forwards. either small quantities of acid solutions.164 THE WORK OF THE DIGESTIVE GLANDS. it was discovered (experiments of Dr. apart altogether from the fulness of the tube. of acid solutions out of the stomach is remarkably slower in passage the case of dogs with a pancreatic fistula than in those without one. as a matter of a mixture of is fact. for if all the other conditions remain unaltered. When one continuously injects. been of the actual progress. It offered to us the analytical consideration of a long series of facts without any inner connection. and with the purposiveness of a delicate and ingenious mechanism of ? The synthesis have the movements. varying combinations. which closes the pylorus each time after a portion of the stomach contents has passed into the intestine. A. a solution of soda which had previously been introduced into the stomach may be kept there for an unlimited time. and the mechanism till recently as little brought under investigation as had the synthesis of the secretory work of the digestive The credit of having begun this synthesis belongs to two apparatus. however. which different substances. the solution of soda generally leaves the stomach very quickly. the escape of the solution from the stomach On the other hand. Each time that the intestine receives a portion of the acid contents of the stomach. there was no reply and yet is not the complex food mixture somehow separated into its components during the onward movement? All these processes must. investigators (Hirsch and v. quantitatively and Why. or of pure gastric juice. Our investigations have been carried on along the results. regulates the passage of food from the stomach into the intestine by its behaviour towards the acid reaction of the gastric contents. numerous the forms of stimuli employed ? And yet this great mass of work could not in the smallest degree answer our question. or why . and in qualitatively. through a duodenal fistula. the various juices are poured out in different parts of the alimentary canal. both therefore. 8.
The completely arrested. it by thin king exclusively of a mechanical reflex discharged from preted investigators the duodenum and acting upon the pyloric orifice of the stomach. egg-albumen. or is actually happening. /SWm'o&ic/i). if the dog for instance. pyloric orifice of the organ. however. and the acid contents of the stomach passed without control into the duodenum. now. solution of sodium bicarbonate. the stomach happened to passed on be at rest to begin with. neutral or (e. the evacuating .g. Although. When. be But as matters stand this cannot happen. the fluids. the following phenomenon was observed by In fasting animals. causes in turn 165 and at the same time inhibits the propulsive movements The acid food mass allowed through by the pylorus an increased secretion of pancreatic juice. ^ per cent. of the stomach can be repressed in a purely psychic manner. clear and direct references are to the fact that acid and alkaline fluids pass from the stomach into the duodenum at different rates. it remained several minutes in spontaneously occurring movements These without moving from the spot. or milk) be poured through the fluids are all physiological saline. under certain circumstances. probably as the result of a psychic impulse. fluid fistula into the stomach. of a further portion of acid contents from the stomach is perescape This regulatory action prevents disorder in the progress of digestion. P. when an ingestion of food is immediately to take place. the ferments of the pancreas are afforded the opportunity of unfolding their activities in the most unrestrained manner. neither this author nor the other made who have worked at the subject (v. If it were otherwise. on mixing with it. without the animal perceiving it. they misinterthe food contents into the intestine. alkaline solutions If. Hirsch. Consequently. or still better.. while the insufficient reduction of would hinder the activity of the pancreatic ferments. the very rapidly that is. which dealt with the movements cf the stomach itself. In other experiments (Dr. movements for emptying the stomach are from us. together with the entry of those powerful excitants of the pancreatic juice. in the course of a few minutes largely into the intestine. introduced with the same precaution. and at the same time ensures regularity in the transition from mitted. if with an lesophagotomised animal a sham feeding be carried out. and is thus It is only when this has been achieved that the gradually neutralised. the acid gastric digestion to the alkaline intestinal. the bile. namely. acidity digestion injurious effect of pepsin upon intestinal digestion is arrested. the bile and the succus entericus. The of the food might thus.THE EMPTYING OF THE STOMACH. Mering and Marbaix) appreciated the nature of the acid reflex which regulates the passage of On the contrary. be greatly roused by the sight of food. while owing to the neutralisation of the food mass. 0. in the paper of Dr. might arrest or greatly weaken the action of the pepsin. time to time discharged.
in the study of stomach movements. be seen. we have encountered the same earlier investigation? namely. namely. of movements the stomach are arrested. The apparatus of defence is immediately called into play. "VVe are far from having the matter fully under control. like the others. At first such accidents annoyed us considerably. a general struggle begins on the part of the organism against these agencies. but it soon became evident that our discontent arose from an obvious misconception of the nature of the facts. in contrast to that intro- duced unobserved through the fistula. The struggle in question leads either to a repulse of the .166 THE WORK OF THE DIGESTIVE GLANDS. The animals employed in these researches. have given you a short summary of our later I see quite well how much yet remains to physiological experiments. more correctly said. and this is a *urther confirmation of the relationship already mentioned. gentlemen. although evacuating movements may have been going on immediately before. S Lintwarew have shown that fat also. that experiinfluences. or to the attack of pathogenic micro-organisms to a degree beyond the usual extent of these influences. occasionally became ill. and indeed at the very threshold characteristics of the subject. and which for the most part served for observation during many months or years. But physiology learns of them only through illness at other times their work is not to . Thus milk lapped up by the dog. The spontaneous movements also cease when acid fluids are poured into the stomach through the fistula. when poured into the duodenum. that. but I In the foregoing the next advances are clearly indicated. should a pathological condition of the digestive apparatus not to us ? What is a pathological condition ? Is it not the effect appeal Why produced upon the organism by the encountering of an unusual condition. This consists of parts of the body which. and we are justified in hoping that in the future our field of work will be even more accessible to investigation than in the past. which we saw in our ments of Dr. and share in the maintenance of the general equilibrium of the whole living organism. does not pass at once into the duodenum. live within it. be done. clearly with the object of allowing the food to remain in the cavity and be worked up in a suitable manner. or. an unusually intensified ordinary condition ? Suppose one received a mechanical shock or became exposed to the effects of great cold or heat. and sometimes the affected organ was that actually under investigation. You see. Consequently they are worthy objects for physiological investigation. the purposive nature of the phenomena and the intervention of psychical I have one further remark to acid. regulates the emptying of the stomach in even a more striking manner.
a method is quite commonly used in physiology for the investigation of the functions of a given organ. it is most competent and to logically is apply them to the study of vital phenomena. namely. established in this new condition of body gradually provided. may lead to the discovery primaiy seat in the of action of the process these relationships which. organism Is not this from beginning to end true physiology. which brings as a result the injury or destruction of this or that part of the body. . or it leads to a conquest by the enemy. as well as the by which continuity of function has been impaired. we have again a method of recognising followed up. it appears to me. upon to decide precisely the physiologist who is the value of such investigation methods. when the work of defence ceases. has again proved to be of inestimable assist- ance in pathological investigations. but is invaluable in the analysis of the pathological condition. its which function naturally ceases. When pathogenic agencies (such as great heat or cold. ifec. if of the cause. a method put into operation by nature with a delicacy which is quite unattainable by our crude technical measures? If the destruction be limited to a single organ. But if an organ be destroyed. even the most minute. The experimental method indeed soon proved triumphant in our new field.) were caused to act on the surface of the miniature stomach. other supplementary organs come into . 1G7 enemy. are calculated to excite the interest of the whole clinical world. in which we seek to penetrate into the relationships between important parts of the body ? Only an incurable scholastic could say that it is not part of our work. Have we not therefore in this. It not only permits all the details of the diseased condition of the glands to be laid bare.PATHOLOGY OF GASTRIC SECRETION. the injury be not limited to a single organ. already known to the members of this society. compensation for the loss of its function is is equilibrium play. These at present apply only to the pathology of the gastric glands. A We learn way to recognise new and finer relations between the organs. in his special field of research. Nay more. Every drop of the altered secretion of the mucous membrane could be collected. various strong chemical reagents. but spreads still wider owing to the functional relationships between organs. could be seen. however. therefore. Every detail of the pathological The diseased state condition. and discover functions which were previously hidden. the deviations of the activity of the gastric glands from the normal could be observed in an ideal manner. The method of isolating a gastric cul-de-sac. If. the EXPERIMENTAL PATHOLOGY OF DIGESTION. I am already able to communicate some data which. Although only two workers in the laboratory have given special attention to it. The Physiologist here.
only mucus instead of secreted by the irritated surface. sublimate solution. a 10 per cent. that is. of their cells. which immediately appeared upon the application of the irritating substance. solution of nitrate of silver. operates on the walls of the small cavity. without the intervention of complications of any kind. a 0'2 per cent. the pathogenic This might reflexly influence. : were introduced for a few minutes into the small stomach. an acute mucous Is it. /Saiurieiv. The digestive apparatus or from disturbances of general latter are excluded. It acts upon no food. J. or a strong emulsion of oil of mustard. and yet I juice ask again. either from consecutive trouble in neighbouring parts of the nutrition. as well as those of the glandular layer. Has not the true physiological function of the surface epithelium . which on the day of the experiment had completely suppressed the normal secretion of gastric juice. intensity of the The contrast between the disappeared without leaving a trace behind. to made of the short interval when the unusual stimulus. indeed. the large stomach we are able on the one hand to is acted upon by noxious agencies. phenomenon and its short duration is really striking. however. such as absolute alcohol. was obtained during the whole period of secretion. An exception is. catarrh. Our results were as follows When powerfully acting substances.368 THE WORK OF THE DIGESTIVE GLANDS. could be observed day by day or hour by hour from the very beginning to the end. C. may. the more or less copious secretion of mucus. the diseases of the reflex transmitting part of separately investigate the surface of the stomach. be But in these experiafiect the remaining parts of the digestive canal. have . they produced a more or less considerable. after the lapse of an hour or two.) One might think this was a serious pathological condition. (Experiments of Dr. to one's utter astonishment. since it is always empty. while everything pursued its usual course in other portions of the digestive canal. see the reflex effects in the small cavity as well as to observe the disturbance provoked by a general disIn this way it is possible to order of digestive activity on the other. however. had wholly exhausted itself or an enormous flow of mucus. however. and no stimulus takes origin in it which reacts either on the large stomach or on the intestine. resist the idea that in the cases described no morbid conhad as yet been established. more than one hundred times the normal amount of mucus was At times. but rather that the pathogenic influences had been successfully battled with and conquered before our One cannot dition eyes. is it a morbid state? Often. since the miniature stomach takes no part in the general work of digestion. a condition of disease ? In extreme cases. on the next day. ments we are almost wholly concerned with a study of the pathological conditions of the peptic glands themselves. in many cases an enormous secretion of mucus. When.
and that it gives a general indication of the adaptive mechanisms of the enabled to encounter pathogenic influences. at the same time. that extraordinary stimuli which come in as pathogenic agencies. (150 grms. leaving those of the surface quiescent. furnished a rich have already cells. The surface epithelium thus wards off the danger which threatens the more important elements of the mucous membrane beneath. may We brane. a large quantity of mucous fluid is combipoured out which dilutes the noxious substance. it appears to me. we have already seen in the case of cells of which when brought into the stomach excites only the the peptic glands. according to the nature of the causal con- dition.} of the hourly quantities of juice. A similar differentiation of stimulating effect flesh. C. or forms chemical nations with it. attention to the following. of those forms of apparatus which are adapted to overcome I believe that this applies to all diseases.) of flesh : . In this way we succeeded in establishing different pathological conditions of the contribution to peptic glands. secreted by the isolated miniature asthenia. with all its power of or that adaptation.PATHOLOGY OF GASTRIC SECRETION. an unusually weighty fact We have here namely. which in its also details may vary greatly. the disease producing effects. that stomach. are at the same time specific excitants of the protective mechanisms of the organism namely. and drives it away at the same time from the stomach wall. Sawriew. (Experiments is. leaving the other unaffected. Naturally the effects of the substances named. By your solution of nitrate of silver we have been able to produce a condition of a considerable store of remarkable observations. in striking contrast extreme activity of the surface epithelium. Indeed. but I the peptic glands. That this explanation is correct is also shown by the fact that the to the peptic glands remain absolutely at rest. is dependent upon the specific excitability of this animal body. before us. the intricate progress of normal life. notwithstanding the energetic protection of the epithelium. The chemical substances mentioned stimulate therefore the one kind of epithelium only. before and during the pathological condition experimentally The animal was fed each time on the same quantity provoked. the physiological characteristics of the gland We wish only to call the application of a 10 per cent. but which for the most part assumes a phasic character. of the 169 stomach been here revealed a function of which we could form no adequate conception in the normal course of aft'airs ? By virtue of its wonderful power of secretion. J. when of a certain extend to the deeper layers of the mucous memstrength. then see an altered form of activity in the peptic glands. by which it is apparatus. of irritable debility of In the following two columns are given Dr. and.
5-3 4-3 4-4 )! 3-5 o 2-5 1-2 2-8 4-4 o-o )) Total 24-7 Total 15-6 You see. . . 6'5 c. Obviously such a state is not merely a special result of the nitrate of silver.. So far as I know.c.c. In the gland-cell has become much more inert.t the same time they were extremely easily fatigued.. 1st . 8-4 c. . . and at length the secretion stopped prematurely. The gland-cells had become more excitable than before. this interesting fact has been first established by experiment in my laboratory. . but n. clinical observation has to contend with much greater difficulties than is the case with experimental investigation in the laboratory. therefore. . Kasanski).170 THE WORK OF THE DIGESTIVE GLANDS. Pathological secretion. 8-4 3-5 1-9 1-3 . . 2nd 3rd 4th 5th . . tinued in the third hour.. .. 11-6 c. 6'2 c. 11-6 10-8 5-6 3-6 . therefore. pate that a knowledge of this condition We may confidently anticiwill influence both the methods of clinical investigation. state : Normal secretion. where the normal secretion is compared with that during the pathological Hour. Total 26-7 Total 37'8 it The condition was produced by the application of intense cold.. set in.c. a typical form of depressed cell activity. It is a striking confirmation of the fact that. The significance of this condition of the cell is at once clear. after much less than the normal amount of juice had been . . and represents. but must also appear under other conditions. . an This conexceptionally steep decline to abnormally low values. that exactly the opposite condition of the glands can be set up. Normal secretion. Recently it has also been demonstrated (experiments of Dr. as well as the therapeutics of such diseases. hour markedly exceeded the normal produced.. notwithstanding the infinity of cases where it might previously have been observed in the clinic. This is seen in the following table. but in the second hour. more difficult to set in . .. Pathological secretion. in the analysis of morbid phenomena. . . that in the diseased state the secretion assumed The quantity for the first quite an unusual and special character..c.
Further. it shows observed. the flow finally much greater interest. after perforation of the stomach-wall. Wolkowitsch. than normal. This was especially marked after feeding with bread. as was first pointed out by Dr. as and one of much weaker psychic and centrally excited flow. How is ? this deviation from the ordinary progress to of secretion to be explained Since the centrally excited secretion of the first that the glands. chemically do" with the round ulcer the secretion of the first hour departed in no L o free in the second it remained at its previous instead of dropping to about one-half. (Observations and Experiments of produced a secondary peritonitis. gave rise from time to gastric cul-de-sac.c. The following also. hour is normal. Here.} During the development of the ulcer. in all . revealed that two different periods of secreinstances a further analysis tion were here sharply separated from each other. of We the conditions have. 171 motion. and the corresponding nerve centres are all in normal condition. it must be taken to prove that the augmented has originated either excitability of the secretory apparatus at this stage. therefore. 26-6 15'S . an unstable and an inert. The normal secretion after bread is characterised by a copious flow during the first hour. exceeding the normal by three to four times. It Dr. the secretion was considerably greater way from the normal. a period of to. Of than this hypersecretion was the fact that a sharply prohowever. and finally. but once set under normal conditions for the same going. then a great fall in the second As in previous a rule. is in the centripetal nerves or in their nerve endings. it does more work than amount typical of stimulation. and more tardy in its action when it begins. 26-2 c. about half the former value. appeared. in all probability. 13-0 13-0 . 26-2 c. happened that in one of our dogs a ROUND ULCER formed in the which steadily increased in size. : Pathological secretion. a con- tinuous and increasing hypersecretion was observed. and we know that this secre- tion a reflex one. nounced deviation from the ordinary hourly rate of flow. namely. N~. P. P.ROUND ULCER OF THE STOMACH. then. Chigiri. the centrifugal nerves. A. time to violent bleeding. but are the figures in question Normal secretion. In the succeeding hours height. In our excited secretion. when an increased secretion in the second hour is The following appears me to be the most correct view. brought about by reflex action. two living substances when thrown out of equilibrium namely.C.
the size of the small stomach. of the inhibitory nervous system. which is thrown into activity by the more than ordinary degree of stimulation. us that these opposite conditions represented different phases of one and the same affection. The thought suggests itself that. which finally produced as much secretion as the large did normally. C. the alcohol is also able to set up an abundant to protect the deeply lying cells secretion in the large. especially when. free secretion of gastric juice begins from the small cavity. have shown with great regularity that the first reaction of the peptic glands to a powerful and unusual influence consists in a marked depression of their activity. an extremely alcohol. J. which here represents a healthy part of the large stomach manifested a striking compensatory activity. in the disorders of the large stomach. (Experiments of Dr.172 THE WORK OF THE DIGESTIVE GLANDS. the activity of the peptic glands is at once inhibited by means still of a special reflex. which we several times produced. The only exception to this. whose object is further against harmful influence. more especially in the This happens not only in the large cavity but also in the first hours. has not hitherto been discovered or defined in the clinical investigation of the disease. at lea>t for some kinds of food. On one occasion. regarded as the primary and which the secondary ? in which diseased conditions of the large or small stomach were experi- mentally provoked. Which of them. the walls of which latter at no time come into direct contact with the injurious substance. was about one-tenth of the large. so far as I know. for instance. however. . lasting for several hours or even This depression is of a reflex nature. for several days. (Experiments of Dr. It is due to the influence days. the secretion which is subsequently produced by an ordinary meal is less than normal. C. Sawriew. is observed after the action of strong When alcohol is poured into the large stomach. is to be Our experiments. . As a general rule. /Soborow.) Further. probability. C. or a solution of nitrate of silver. When one. J. In the accidental illnesses of our experiment-animals we have frequently observed an augmented or 'a diminished activity of the digestive occurred to It has often glands when contrasted with the normal. Conversely. we had completely arrested the work of the large stomach. judging by the amount of secretion under normal circumstances.) As soon as a diminution of secretion below the normal appeared in the large stomach an increase was seen in the small. an enormous activity gradually developed in the small cavity. we have often distinctly observed that the isolated miniature organ. into the large stomach (experiments of Dr. is the special point of attack of the disorder a matter which. pours ice-cold water. Soborow). by the application of very hot water. by acting on the latter. J. as soon as the stomach encounters an unaccustomed stimulus. small.
in the form. allowed the contents to escape. Thus. and compared the secretions from both cavities. the whole problem under compare it with the normal. again. and firmly convinced that further endeavours along such lines will lead to still more important results. continues hour for some time. Any of the food which does not of itself drop out at the fistula may be removed by washing. Or. however. when was augmented. in certain pathological conditions of the large stomach. be ascribed to the facts of experimental pathology It appears to me that. towards the end of the digestion we often found. In the case of fasting animals. of mince-meat. a secretion. at once. The astonishing capabilities of the organs for vicarious activity drove us irresistibly to an analysis of the purely physiological problem namely. A comparison of relationships of the the quantities of juice. certain length of time. with the fistula tube open. 173 in the above case. by investigations such as these. in point of fact. also. and of our having mastered. can be furnished. But I should like to indicate a certain importance which is. Occasionally. may be determined in different ways. then. 1 think. in the third or fourth of normal digestion. the secretory two stomachs. under normal or pathological conditions of either. if the fistula tube of the large stomach be opener]. as experimentars. gave. the mechanism of and have grasped causation. the physiological defensive arrangements more sharply defined from the purely pathoitself is subdivided into phases. am alimentary canal.NECESSITY FOR EXPERIMENTAL THERAPEUTICS. we opened the fistula of the large stomach. in such cases. the its secretory activity was increased tenthe work of the glands in the large stomach activity of those in the small organ became diminished. The relationship between the secretions of the large arid small stomachs. on opening the fistula. the small organ inversely reflects the activity of the large. which. while the pathological state I accurately localised. we. of the mechanism by which these compensatory events are brought about. flesh chopped up in small pieces. or. may be given to eat for a measured. that the large stomach was free from food while the secretion from both the large and small still continued. then washed out the interior. as accurate and complete as that which possess of its we now I Are admirably beautiful work under normal conditions. we naturally wish to come back and It is Qnly by so doing that the final proof of our physiological knowledge being complete. Conversely. and that we shall in the end arrive at a knowledge of the processes of disease in the logical. When we its see a deviation from the normal. Hence fold. to just communicated. . the conditions of disease are better differentiated. to rest satisfied with this? think not. and then the juice from both stomachs collected. which can be may be excited in both stomachs by offering the animal food. still better. Or lastly.
explain The following example may my meaning. united in reality the whole of medicine. and in every way self-competent. from a new side. and therapeutics. however. since it presents the phenomena of life. misunderstanding. As the first example of our therapeutics. disregarding for the moment its practical aims. the youngest and most branch of the series. pathology. unfettered by physiology. from the sudden cutting-out of its most important secretory and motor nerve. I should wish. I think. Hence. is the only one which has developed to vigorous the natural and full extent of its own inhei-ent capacity. Physiology should be able to do much the same. is by no means new. cherish a well-grounded hope of expanding this method of research in the future to vestigations in physiology an extent commensurate with the results and experimental pathology.174 THE WORK OF THE DIGESTIVE GLANDS. will be able to give valuable indications to the clinician. to which I now But we may. proceed. Bacteriology. almost every trace of digestive action on the part of the stomach disappears during the earlier The ingested food soon undergoes decomposition. . born of the laboratory. It entry into the of is our innatural that upon our to be guided first new field we should grow allow ourselves by the experience of clinical therapeutics. What I have said now about experimental therapeutics. In these animals. No one can say that he fully comprehends the physiology of an organ till he is able to restore its disordered function to a normal state. The subject itself affords a new and fruitful method for the study of living events. It is only an expression of the prevailing opinion of medical science. the experimental therapeutics. by the experimental method. the traditional settings and mouldings which constituted. for the older investigators. and often reveals to us gaps in our physiological knowledge. supported by scientific knowledge. but I am con- vinced that our new therapeutics will soon to be an indepen- dent source of experimental physiological and pathological knowledge. which it is our duty to investigate. lines of separation between the different fields of work. Then. are for the present not very important. an EXPERIMENTAL THERAPEUTICS spontaneously arises. Our own investigations in experimental therapeutics. is due to modern bacteriology. experimental therapeutics to avoid a is essentially a test of physiology. Thus the necessity for investigation. and this in periods. This science is at one and the same time It proceeds from beginning to end along experimental lines. Undoubtedly. the great honour of having. and previously about pathology. may I bring before you the treatment and care of the dogs in which the vagi nerves were divided in the neck. A mechanic only lays aside the study of a machine when he is able to take the parts asunder and put them back again in their original places.
P. are suitable. J. and this result is solely to be ascribed to the fact that the causes of the disturbances which set in after the operation. fully confirmed. In a sway in the clinic. It is one differing greatly from the current belief. and the stomach be freed. Katschkowski. . more than ordinarily * But naturally the mechanism of the alkali influence has still to be physiologically analysed. before the feeding. we have a instance of a rational therapy. E. The author had already made several communications dealing with these investigations. founded upon laboratory knowstriking ledge. by systematic washing out. Thus. the indications for a rational employment of alkalies have been still further accentuated.per cent. But if each time. has at length been answered in the affirmative by physio- restored. Soborow) the hypersecretion proved to be readily amenable to the influence of alkalies. It is to be observed that. and its digestion regulated as above indicated (the cavities of the mouth and stomach having been severed by means of an oesopha- gotomy so as to prevent the gastric contents from passing along the gullet into the lungs. C. In this. solution of sodium bicarbonate) by way of treatment. arising from spontaneous illness. it is true. Indeed. from the remnants of the previous meal. and a tolerably good condition I should like here to transgress the narrower bounds of my and once more expressly state. life and an excellent condition of health. In all the cases observed by us (Prof. with regard to the effect of these substances.* that the question of the subject. in the state of increased excitability which alkalies. survival of dogs after vagotomy. with the experimental establishment of asthenia of the gastric glands. in the laboratory.) may return once more to digestion. the normal psychic excitant of the gastric glands (which is now absent) be replaced by a chemical stimulant. Pawlow and Dr. if a gastric fistula be made in the dog. a double vagotomy in the neck ceases to have a fatal effect. P. have been sub- mitted to an exhaustive physiological analysis. or set up intentionally.ASTHENIA OF THE GASTRIC GLANDS. should vomiting occur). P. the difficulties are soon overcome. directed against a severe and fatal lesion of the organism. and the greatly exalted excitability of the glands was fully and permanently set aside. the operation is consistent with long Prof. It diminished markedly. which I explained five years ago to this esteemed assembly. J. J. with their inhibitory effects. which has been for so long a matter of uncertainty. (Experiments of We series of dogs with well-marked hypersecretion. producpd also. its 175 turn makes matters worse. logy. which still holds Pawlow and Dr. The result was that we had the satisfaction of seeing the mode of action of alkalies. In the irritable debility of the cell that is soon leads to exhaustion to say. we have used alkalies (a |.
which the organism Our instead of holding back will. J. This unexpected result has only been made possible by the These are. Let us now turn to MEDICINE. One of these favouring cir- cumstances this we water into the system. Hence we are able to assist a weakly which only abstracts the water with difficulty. processes are exceedingly complex It cannot be denied that biological when contrasted with other pheno- mena of nature. that mankind from the co-operation of two conditions. experiments have confirmed this hypothesis. P. that the secretory cells attract In certain circumstances the blood opposes If sufficient a considerable resistance to this abstraction. I might say all mankind the present achievements of medicine seem remarkable. largely in this. accomplished. Pawloio). Her task seemed endlessly large and hopeShe has lessly complex. discovered in the introduction of large quantities of based (Experiments of Dr. acting cell. but they are not as yet concluded. by intentionally diluting the blood with an excess of water. indeed. at least to a. fortunately arrived at many correct solutions of the problems which confronted her. that in this search after health nume. lie in effectively combating the different forms of hypo- We have further endeavoured to come to the aid of an enfeebled condition of activity of the gastric glands by supplying favourable conditions for the preparation of the juice. water be not present. glands is readily amenable to treatment. this condition of the gastric The chief difficulty will. But if indeed. however. it cannot be rous individuals . These experiments are now being carefully pushed forward. but yet it is. took part. the cells could not withdraw an adequate quantity for the preparation of the juice. We have already taken of food act up the question of how the individual kinds upon the condition of hypersecretion (Dr.) We (experiments of Prof. Judging from the material to hand. showing that the quantity of juice was strikingly dependent upon the amount of water in the organism. earliest times constantly and passionately strove after the maintenance of life and health and. considerable degree. and. endeavour to expel. long before they had become matters of scientific investigation. notwithstanding the countless numbers of other possible ones. Soborow). secondly. it was ordained by the inevitable dispensations of life that medicine should hold sway over biological phenomena. secretion. Nevertheless. accomplished that which was expected of her. It is clear that the formation of juice upon earlier facts by the glands consists water from the blood. at least in part. and that the difficulty of establishing a causal relation- ship which would give us control over them is very great. And medicine has. on the contrary.176 THE WORK OF THE DIGESTIVE GLANDS. Saiurietv.
such as is at least read of by the medical student. observato practice the implement. 177 doubted that they are only very small in comparison with what she must ultimately accomplish. she is limited in the majority of cases to one implement of natural investigation.THE VALUE OF EXPERIMENTS. M . If an educated man. It appears to me that the most remarkable advance of modern medicine consists in this. and within relatively narrow limits. The true character of this connection can only be guessed at by the observing investigator he has to choose between a number of possible hypotheses. but also into that of theraThere is no reason why this influence should become less. while the other limits than the inventiveness of the human brain. Observation. by artificial but simplified combinations. This revolution has. for. experiment. nature of the conditions. to come into play. namely. he must certainly be astounded at the sovereign power with which modern physiology holds sway over the complicated animal organism. in another only an indirect and accidental connection. from nature what it will. grapples with the problem. carefully conducted course of demonstrations in animal physiology. for instance. peutics. and between which there is in one case an essential. And his astonishment will further increase when he learns that this power has been acquired not in thousands or hundreds of years. It permits now one condition. It is only by experiment. The power of biological research is experiment wrests immense. In other words. the solution of which is sought for. It has created in the short space of seventy to eighty years almost the whole of what must be described as the very comprehensive subject of the organ physiology of the animal body. that the possibility is opened up of extending experimental investigation into all its important branches. however. the development of which knows no other tion . And now we see the method of experiment extend its influence not only into the subject of pathology. she will never attain complete success. These future advances. what the real connection of the observation collects what is offered to it phenomena by nature is. : on the contrary. in different from only. that the crowning work of medicine can be achieved. which only suffices for the investigaThe more complicated these become tion of simpler phenomena. . should attend an ordinary. will not be attained by merely calling in the results of increased knowledge branches of natural science. and learns in this way. to assist in her diagnostic So long as medicine devotes herself and therapeutic measures. encounters a number of individual phenomena in the animal organism which proceed together. however. Experiment. and what is more complex than life the more necessary is experiment. Observation is a method. dare only be employed by her with great precaution. not otherwise closely acquainted with biological science. but in a few decades. now another.
and then only. It is the full etiology of disease is known that the medicine of . laboratory experiment alone that is capable of the whole problem of disease. and of exactly differentiating the reparative from the compensatory events. collected an of material concerning the finer processes of disease. and where the secondary which It is by such knowledge alone that approit has called forth. the protective phenomena from the lesion itself. It is unravelling in the organism. and begins to work in the organism before But the recognition of the patient becomes an object of medical care. in recent times. and effective assistance can be rendered to the diseased organism. indeed. Etiology.178 THE WORK OF THE DIGESTIVE GLANDS. it is is well known. since in it we always with a causal factor which is intentionally set to work. but on the contrary always bring help. the origin of disease is one of the most essential problems of medicine. conditions. Pathological anatomy too crude an instrument for this. priate real cause of a disease only by experiment that we can ultimately discover the and estimate its importance. as Again. still what it one can only fight the etiological factor. of a thorough knowledge of all the processes of a disease was first by the method of experiment. will our interference be followed by no evil results. pathology again. of proper knowledge On the other hand. It is only by such experience that the interweaving of these latter can be detected. is the weakest branch of medicine. and clinical observation provides without experiment is powerless in face of the complexity of the attained phenomena. with a is to be aimed at. yet the possibility of a complete analysis of the whole course of a disease from its inception to its curethat is. is more important. and render only harmless before when it has penetrated into the organism. For in the first place. It is begin precisely here that the power of clinical medicine is least. when the cause is known. for the for was brought into the laboratory. While clinical medicine has clearly distinguished the different types of diseases. still continue to do so. One is now able to in- vestigate almost every pathological phenomenon in the laboratory. that one can it only under these conditions. pathogenic microbes that the experimenter had the whole field of pathological physiology opened up to him. that it can be proved where the primary injury lies. and. the cause usually steals in. yet the working up of the material was greatly restricted for want of knowledge of so important a factor It was only after the discovery of in disease as micro-organisms. As a matter of fact. most part been accomplished by bacteriology. and has given an almost perfect morphology of pathological and while macroscopic and microscopic anatomy in associ- ation enormous amount with clinical investigation have. Then. and this is forestall the cause of disease. Although some time before the development of the latter subject.
the other hand. One can readily conceive himself in the the laboratory. to an experimental analysis of the phenomena of disease. it is usually appears.EXPERIMENTAL THERAPEUTICS. taken its proper place as an experimental science. of the obvious justice and importance of these consideraone cannot pass on without expressing a regret that pathology has not yet. however. and not to conclusions and abstractions drawn its materials. It was for this reason that therapeutics several years ago called in the method of experiment to its aid. if The pharmacologists have. the endeavours of the clinician to grasp the mode of action of his remedies are easily to be understood. in his measures against this or that disease. and when thorough exploration the difficult position of physician who. the place of honour must be assigned to experimental pathology. and there their analysed. often does not know what in a given case. and pursues its special theoretical aims. or in his use of this or that remedy against a given symptom. The connection. At first. bit by original goal. or at least not everywhere. as pathological physiology. of academic instruction. bit deviated little. how much room for all sorts of chance occurrences Hence. In the ordinary In view tions. HYGIENE in its widest sense. No when the field of pathological investigation is becoming more and more its dependent upon promises to be so fruitful and so engrossing. at in the healing action of a given substance. the remedy in reality effects in the organism. Therapeutic measures were given over to laboratory investigation. our clay 179 nan become the medicine of the future that is to say. It investigates the action of chemical agencies on the animal body. and from this experimental pharmacology sprang up. it appears to me that in the training to which we at present give the name of general pathology. namely. however. programme appendix pathology. and now interest themselves but from their all. and in one laboratory. which often only amount to another grouping of very important scientific advantage is likely to accrue from a purely theoretical treatment of general pathology at a time from special pathology. or how it aids How insecure and indefinite must his interposition ! often be. at least in a way that will afford justice to On each. either as an lost in the subject of general of pathological anatomy. by a natural process of development. more especially if academic teaching be added to the duties. Pharmacology has thus. between the mateiials of pharmacology and the aims of practical medicine has thereby been . grown to be a section of physiology. Against this in itself no objection can be raised. or But the methods of pathological anatomy and of experi- mental pathology are too different to be combined under one representative. effects on the healthy organism were to be chemical medicaments were experimented with.
they obtain no proper recognition. general. has not been set about. by experiments on healthy animals. pharmacology should be completed by an experimental therapeutics. and. at the same time. the indications and centra-indications to its therapeutic applications are then related without any connection with the previously It is in consequence of this that many discussed physiological action. are subjected in the laboratory to a bold and constantly controllable therapeutics. yet in many cases it has grown to be very lax and purely nominal. lay in the difficulty of procuring the necessary diseased animals in the laboratory. that we are to witness an of the interest of investigators as soon as other pathological processes. thanks to the advances of experimental pathology. likewise itself. it is only when pharmacology is blended with experimental therapeutics. in the right way. overcome. For in spite of the fact that this connection formed an essential factor in the original working plan of pharmacology. simply because the animals were healthy. make clear the real importance and mode of action of a therapeutic The necessity of studying the effect of such remedies on remedy. both the experimenter and the practical physician. we may be con- vinced that the experimenter may reckon on not a few triumphs the moment he sets aside the exclusive point of view of the would-be . the regrettable possibility will be remedies being undeservedly thrown aside solely because On their pharmacological analysis. At present. and even still finds expression in the name of the science. as above indicated. This difficulty is now. damaged. and corresponding requests have already been uttered. unfettered by extraneous considerations. it will. deepen our knowledge of the reaction <f the organism to chemical agencies. should also find a place. but also with the little of diseased It will then study not alone the action in animal body. Nay more. in a large measure.180 THE WORK OF THE DIGESTIVE GLANDS. that merited oblivion. or perhaps could not be set about. not without reason. of the different remedies. of many much therapeutic delusion will pass into long the other hand. but also their healing influence on It will then in its own interests expand and the diseased organism. avoided. in the comprehensive programme of medical teaching. In the teaching plan of experimental therapeutics the experimental investigation of other remedial measures than the mere administration of chemically active substances. Then it will deal not alone with the healthy. But an essential barrier to the fulfilment of these requests. Thus in the ordinary textbook. our knowledge also of the organism In the interests of the practical physician. not merely the bacteriological. Indeed. after the author has dealt with the physiological action of this or that remedy. One may enormous awakening hope. diseased animals has long been recognised. physicians are so mutual interests In the satisfied with modern pharmacology.
Thus without any special good to the science itself. biological. Many have hoped to bring pharmacology and medicine more together by recommending and bringing about the establishment of clinical departments in pharmacological institutes. either the experimenter will be lost in the clinician. Further.EXPERIMENTAL MEDICINE. in its whole extent a conscious and purposively acting have an example in proof of this in modern surgery. form an of devotees. Aided by the plasticity of the organism. Every physician who has the mind. 181 and takes upon himself the initiative of therapeutic treatment. it can now treat its subject from the purely On what are mechanical standpoint. and in stimulating you to real activity ? If I have succeeded at all. And now for our conclusions It : is only when medicine is able to stand the crucial test of experiment that it can become what it should be. can take part in general scientific medical work. special clinical pharmacology would have no superiority in a systematic and competent handling of therapeutic remedies. but also by actively supporting experimenters in their efforts. We its brilliant results founded ? Simply on its perfect knowledge of how to achieve its aims. and ultimately become an important and ceaseless worker in this field. because in the interests of every way the and also of medical science. the sick person in it can be made just as little the subject of experiment as elsewhere. the talent. since it is only within the . and more prospect of a practical result. namely. not only by personally taking part in it to the fullest extent possible. since this competence is sought after by every clinical teacher. and secured by asepsis and antisepsis against its arch enemy the micro-organism. and the energy. With unfeigned interest I now stop to ask myself in how far I have succeeded in convincing you of the extreme importance for practical medicine of the method of experiment. A lasting and stable combination of these separate activities is scarcely to be achieved in practice. But it appears to me that laboratories for the study of experimental therapeutics would have had more scientific justification. and suitable means are necessary. suitable conditions. it is your duty to forward in interests of biological experiment. the following important difference between the representatives of clinical and of experimental medicine The scientific representatives of practical medicine are drawn from the : whole mass of practising physicians. guided by a knowledge of the anatomical structure and physiological importance of the several parts of the body. expert. than special pharmacological clinics. inappreciably small number Experimenters. on the other hand. Do not forget. suitable men. gentlemen. It matters not what the clinic is called. medical art. or the clinician in the experimenter.
for in them the sick. therapeutists. of Consequently. generally. in the laboratory. whether it be to clear the mode of action of a feel the necessity at the present time out the problems in the laboratory which they encounter in working the hospital. erection of institutes physiology. one to pathological. places for benevolent activity. as well as in specialisation in the laboratory affords them. Gentlemen. after all. Work . afterwards. and one to therapeutic required by science. Because it is your duty. is what must engage our attention. whether they spring from These institutes are. on the one hand. And now to turn away from the fostering care Every human being will welcome the founding and of all kinds devoted to the care of the sick. Nevertheless. I maintain that . persons who. considered from a broad standpoint. Hence. and will ever remain so. with the clinician. no misuse of my great powers of adaptation. great credit. private or public initiative. since many better It is known that clinicians. experiment. in the struggle for existence. even more than the physiologists.182 THE WORK OF THE DIGESTIVE GLANDS. leaves to him. to encourage the beginners of laboratory work. work I . am making . demands a full surrender requires that the worker shall devote his whole energies to it. chances in life. In the curriculum of medical science there should logy everywhere be three chairs given to experimental physioone to normal. Think of one who mentally grasps the unfathomable depths of his problem. The clinicians. whether they be of a pathological or therapeutic nature. have encountered greater or less injury persons sacrificed to the general conditions of life are taken care of. life narrow range of the laboratory that they can be recruited. and . in by far the greater number of instances the initiative to investigations of an experimental pathological or experimental theraThis is to their peutic nature at present proceeds from clinicians. that is to say. or to test a proposed surgical procedure. this kind of work has always to take a second place it fills the leisure hours which his first duty. however. to make therapeutic measure. and surgeons in many cases turn to the fruitful method of analyse a pathological process. these institutes are fields of for people who in life are truly called upon to bear an excessive burden to solve problems which are as yet insoluble. of this. the care of the sick. Such endeavours naturally are to be hailed with pleasure. both in the scientific institute. On the other hand. they are in methods and conception nothing else than branches of physiology) should be given the most favourable conditions and the most independent position possible. life as it words. our special departments of experimental pathology and experimental therapeutics (for. I have in mind life with all its concerns the majority of man- kind and this.
for instance. therefore. in his heart nurses a bitter feeling of impotence. Thus. or that so great a number is not necessary. vie with each other and equipment. which is in striking con- ment has to the extraordinary growth of the pi-oblems biological experito solve. gentlemen. for in case of the least doubt or suspicion the laboratory can repeat and control its earlier observations. animals alive for months or years. For. in Germany the by scientific laboratories. Unfortunately. a general lacerability of the blood vessels. In many other scientific institutes a great want of accommodation makes itself felt. of the Institute for Experimental Medicine. in addition to a series of special rooms for the several experiments. within your power and it will not be too much. I have at present. How many and how profound pathological processes have. the same cannot by any means be said. I am convinced from various occasional observations during the last few years. in connection with a ! disturbance of the functions of the liver. Hardly any one would be so bold as to say that these animals have not been employed to good purpose. for investigations in experimental pathology and therapeutics. of our laboratories. at another time an ascending paralysis of the central nervous system or. come into existence before my eyes I have seen. in the laboratory of the Institute for Experimental Medicine. I had by no means the intention at first to set up conditions of disease kept my I operated solely for physiological purposes. an enormous ascites develop .REQUIREMENTS OF SCIENCE. and so on. a large number of experiment animals favours the starting and solution of new problems. where That a the events to be observed stretch out over months or years ? fruitful field is open in the prolonged observation of experiment animals. and . On the other hand. again. 183 Give him everything tributes which Moreover. . which in the splendour of their design owes its existence to the lofty ideas and enlightened benevolence of Prince Alexander of Oldenburg. with the one wellknown exception. the great cultured nations. a number of sufficiently large and suitably trast furnished compartments are now unavoidably necessary for the different animals under experiment. . our beautiful hospital buildings are only we pay to human misery and infirmity. especially the physiological. should that in which mankind beholds its dignity and pride be likewise deserving of palaces. It is in fact the number of animals which has given security to our results. and which must be so kept that their state of health leaves nothing to be desired. up to thirty dogs on which the physiology of digestion has been or is being studied. But these animals have been necessary solely for the How many more will be required study of physiological problems. where the power of man and his Such palaces have been built strength of mind may be enlarged. under these circumstances. All the more.
in an out-ofthe-way village. and the regrettable but unavoidable outward presentment is seized hold of. the more certainly will patients be cured. gentlemen I appeal to the medical men in my audience to assist us here. You move about every day amongst and come into contact with its highest and lowest it by the most intimate relationships. with possibly serious consequences. as you suffering from goitre. And yet does not tell nature and "religion of us that animals are provided for the service ? mankind. and the less frequently will they have to submit to trials between this and that remedy. to be unnecessarily . society.184 THE WORK OF THE DIGESTIVE GLANDS. who pass their whole time in the laboratory. how- tutes. You are linked with Therefore. this form of experiment has often met with the most bitter hostility. there fore. for example. cannot influence the opinion of the public with regard to experiments and experimenters. Take. to come back to it. it is not confined to out-of-the-way villages. it lies with you to spread the conviction amongst the public. But the needs and wishes of experimenters are mostly repulsed. ever. You must make it understood that the greater the perfection attained by means of experiments upon animals. the experience choice of therapeutic remedies. unfortunately. Make it manifest. when. was gained at the expense of a This can be judged of by instances. They can neither find. With us. to the public that modern medicine has passed the stage of the gruesome experiment upon man himself. would not have occurred. There resulted. the patient attempts' at healing on the part of succumbs to the revolting torture of uncalled-for some quack. and of the greatest conceivable importance to it. If more had been extirpation of the the early unhappy results of its removal from patients thyroid body. and have no regular intercourse with the outside world. which even now"are by no means uncommon. and. or uselessly sacrificed not. the following instance. an incurable condition of cretinism. requires therefore. It is your duty. of course. Should you wish to speak in defence of our science. for example. Investigators themselves. Experiments on animals are often depicted in the most malicious way as animal torture. known by experiment upon animals concerning know. grades. nor reckon upon sympathy with their objects. instiwhich will cost hundreds of thousands of roubles. Private individuals and authorities give willingly to the building of new hospitals or clinics. and the keenest sorrow's of mankind. which serves the life and health of mankind. Biological experiment. You take an active part in the greatest joys. The underlying lofty idea is obscured. It is well known But that medicine in its has drawn largely upon application. that experiments upon animals are unavoidably necessary for the advancement of medicine. acquired by their popular this experience great sacrifice of mankind. your words will be listened to.
How is it possible therefore. whether it be devoted to scientific purposes or to teaching. physiological thought. have set forth the work of our laboratory. The income of the only physiological laboratory in Russia namely. of the physiological laboratory of such a colossal medical institute as the But it only approaches Military Medical Academy of St. 185 But when large and specially fitted up apartments are required for experiments. during the course of ten years. and my own views with regard to the relationship of experiment to medicine.THE VALUE OF EXPERIMENT. these are naturally performed not only in the interests of science but also for the purposes of instruction. experimental research and experi- mental teaching must take comfort in tho Once more let it be repeated : The final hope of better things. recording drums. a mere fact of his visit often had an effect. Petersburg. I have had the honour of being associated. triumph of medicine can With this conviction I only be achieved by laboratory experiment. P. the parsiactivity. the physiological laboratory of the Military Medical Academy. that of the Institute for exceeds by three-and-a-half times.. Botkin. so far as it concerned his laboratory. In this respect we are far behind our Western neighbours. for example. has How can a department with so small a staff only one assistant. under present requirements. venture to predict that in any given country. for an experimental chair to develop a tolerably extensive scientific and teaching Further. into astonishment by his rare gift of recognising a and of finding the best remedy for How frequently have I heard the confession from his clinical pupils. and in the laboratories costly apparatus. who was thrown disease. such practical exercises are carried out on a large scale. etc. with the work of the depeirted clinician. the name of the clinician its To-day I whose memory we celebrate. word from him. that the same prescription which for the patient a really magic power. the future. the income Experimental Medicine. upon an income of a thousand roubles. Ten other years have now fled since the death of S. for example in England. With us. it was the clinician yet his memory lives with us all. under the segis of a great name. and in any given medical institute. For example. teach the students a course of practical physiology ? And yet a direct with the materials of physiology. His personality possessed it. mony is just as great in the personnel of the laboratory. its fundamental idea. and If anybody. the average for the corresponding institutes of the German Universities. and a schooling in acquaintance is of the utmost importance for the physician of Elsewhere. the progress of medicine will go hand in hand with the care and attention paid to experimental departments. are provided in large numbers for the use of the students. or even the . Had I a right to do this ? I should not have done so had I not been convinced of that right.
THE WORK OF THE DIGESTIVE GLANDS.
had worked wonders
of the master,
had remained without
One might, perhaps, in theirs, in apparently identical cases. suppose that the celebrated clinician would have been satisfied both
inwardly and outwardly with such results.
standing, unalloyed by these triumphs, always sought in the laboratory, by the aid of experiment upon animals, the key to the great puzzle ;
eyes he has
man and how is he directed many of his
to be helped ? pupils to the
this great appreciation of the
opinion, does no less
experiment on the part of a honour to the name of S. P.
Botkin than his
to all Russia.
this I close
what has here
been communicated will no doubt be welcome to the practical physician. He will often find in our physiological facts an explanation of pathological
be led to the
phenomena, and by a knowledge of the true state of affairs will employment of effective remedial measures. Physicians
would, however, secure to themselves further advantage, if they imparted to the physiologists in what way the explanations, in their and also if they further pointed out opinion, may need adjustment new phenomena in the subject of digestion which have cropped up in
the broad world of clinical observation, but which have not yet entered the field of cognisance of the physiologist. My belief extends to this
only by an active interchange of opinion, between the physiocommon goal of physiological science logist and the physician, that the and of medical art will be most quickly and securely reached.
INDEX OF THE PUBLICATIONS OF THE AUTHOR AND HIS CO-WORKERS REFERRED TO IN THESE LECTURES.
N. M. BECKER.
1'influence des solutions de bicarbonate
de soude, de
sel marin, d'acide carbonique, et
de quelques eaux alcalines, sur
sue pancreatique." Archives des Sciences Biologiques, ii. 433. The same in Russian (Inaug. Dispert. St. Petersburg, 1893). "The Bile as an Important Agency in Digestion." (Inaug. 2. G. G. BRUNO. Dissert. St. Petersburg, 1898 Russian.) The same in French, Archives des
T. vii. 1 and 2, 1899. CHIGIN (French Khigine). " The Secretory Work of the Stomach of The same in French, Archives des (Inaug. Diss. St. Petersb. 1894.)
"The Influence of Fat on the Secretion
of Pancreatic Juice."
Trans. Soc. of Russian Physicians, St. Petersburg, 1896
d. Sc. Biolog.,
stimulant de la secretion pancreatique." in Russian. (Inaug. Dissert. St. Petersb.
The Secretory Work
Pancreatic Duct, and the
Dissert. St. Petersb., 1900.)
7. D. L. GLINSKI. municated by Prof. J.
Stomach after Ligature of the Ferment of the Bile." (Inaug.
Experiments on the Work of the Salivary Glands (comTrans. Soc. Russ. Physicians St. Petersb., Pawlow).
On the Physiological Causes of the Conservation and Destruction of the Ferments of Pancreatic Juice." Trans. Soc. Russ. Physicians,
9. J. JABLONSKI. () "The Specific Form of Disease which Affects Dogs, " " The Influence of a Diet of Milk (b) Permanently deprived of Pancreatic Juice and Bread upon the Activity of the Pancreas." (Inaug. Dissert. St. Petersb., 1894 Russian.) The second part also in French in the Archiv. d. /Sci. Biolog., iv.
secretion stomacale, chez les chiens qui ont subi la Archiv. d. Sci. des nerfs pneumogastriques."
i. 323. The same in Russian. (Inaug. Dissert. St. Petersb., The chemical part of this work was carried out in the chemical department Institute for Experimental Medicine under the late Prof. M. v. Nencki.
THE WOEK OF THE DIGESTIVE GLANDS.
KASANSKI. "Two Typical Pathological Conditions of the Peptic Trans. Soc. R-uss. Physicians St. Petersb., 1900-1901. 12. P. E. KATSCHKOWSKI. "The Survival of Dogs after Double and Simul(Inaug. Diss. St. Petersb., 1899
taneous Division of the Vagi in the Neck."
tion of Gastric Juice."
"Reflex Excitation of the Buccal Cavity and the Secre(Inaug. Diss. St. Petersb., 1890 Russian.)
14. N. N. KLADNIZKI. "Unpublished Observations on the Escape of Bile from the Natural Orifice of the Common Bile Duct."
"Comparisons of Preparations
with Normal Gastric Juice."
(Inaug. Dissert. St. Petersb., 1893 Russian.) analysis of the Secretory Work of the Pancreas.
Inaug. Dissert. St. Petersb. (Russian.)
"Material zur Physiologic der Bauchspeicheldruse."
Anat. u. Physiolog., 1894.
(Inaug. Dissert. St.
the Influence of some Food Stuffs and MediJuice." (Inaug. Dissert. St. Petersb.
cines on the Secretion of Pancreatic
of the Contents of the
Trans. Soc. Russ. Physicians St. Petersb., 1900-1901. J. LINTAREW. "On the Diverse States of the Ferments of Pancreatic
Juice under different Physiological Conditions.'' Kid. 21. J. O. LOBASSOW. "Secretion gastrique chez le chien."
Biolog., v. 425.
The same in Russian. (Inaug. Dissert. St. Petersb., 1896.) "Zur Innervation der Bauchspeicheldruse." Archiv. f. Anat. 22. S. METT. The same in Russian. (Inaug. Dissert. St. Petersb., 1889.) Physiolog., 1894. " 23. J. P. PAWLOW. The Methods of Making a Pancreatic Fistula." Trans.
"Die Innervation der Bauchspeicheldruse."
in the Russian Weekly Clinical Gazette, 1888. " 25. J. P. PAWLOW. Surgical Methods for Observing the Secretory Work of the Stomach." Trans. Soc. Russ. Physicians St. Petersb., 1894 (Russian). " On 26. J. P. PAWLOW. the Death of Animals after Division of the Vagi
u. Physiol., 1893.
Ibid. 1895 (Russian).
27. J. P.
tion der Magendriisen
and MDME E. beim Hunde."
u. Physiol., 1895.
1890 (Russian). A summary of the chief results of this investigation was published in Wratsch and also in the Centralblatt fur Physiologic, 1889.
Gastric Secretion in Dogs."
29. J. P.
Pathological-therapeutic Experiment upon the Trans. Soc. Russ. Physicians St. Petersb., May 1897
Case of Experimental Ascites observed in a Dog in
Survival of Dogs after Division of
30. J. P.
Ibid. 1896 (Russian).
Nerves in the neck."
31. J. P.
Ibid. 1897 (Russian).
Stomach during Fasting."
Botkin's Hosp. Gazette, 1897 (Russian).
Reflexes from the
in the Laboratory on Pathological Trans. Soc. Russ. Physicians St. Petersb.,
33. J. P.
for the Experimental Study of the Gastric
die Secretionshemmenden Nerven der BauchPhysiol.,
digestion et 1'excretion de 1'azote
Arch. d. Sci. Biolog.,
"The Excito-secretory Agencies of the New Ferment of Trans. Soc. Russ. Physicians St. Petcrsb., 1900-1901. "Contributions to the Physiology and Pathology of the 37. J. G. SAWRIBW.
Gastric Glands in the Dog." (Inaug. Dissert. St. Petersb., 1900.) "Results obtained by the New Method for the Experi38. A. SOKOLOW. mental Study of the Gastric Secretion." Trans. Soc. fiuss. Physicians St. Petersb.,
Influence of Different Acids on the Secretion of
la peps-ine," par le
SSANOZKI. The same in Russian.
"Determination du pouvoir fermentative des liquides precede de M. Mett. Arch. d. Sci. Biol. ii. 699. "Sur les stimulants de la secretion du sue gastrique." Ibid.
(Inang. Dissert. St. Petersb., 1892.) "The Physiology of Succus
(Inaug. Dissert. St. Petersb. 1899.)
Sur 1'inefficacite des irritants locaux, comme stimu43. J. SCHIROKICH. lants de la secretion pancreatique." Arch. d. Sci. Biol., iii. 449. " 44. P. O. SCHIROKICH. Unpublished Experiments on the Entry of Chyme
into the Intestine."
45. A. S. SERDJUKOW. "One of the Conditions Essential for the Passage of the Food from the Stomach into the Intestine." (Inaug. Diss. St. Petersb. 1899
46. J. C.
in Pathological Conditions of the Digestive Canal."
(Inaug. Dissert. St. Petersb.,
d. So. Biol., iv. 429.
Le nerf vague comme nerf secreteur de 1'estomac." The same in Russian. (Inaug. Dissert. St. Petersb.
" Le travail secretaire du pancreas." Arch. d. Sci. The same in Russian. (Inaug. Dissert. St. Petersb., 1897.) 49. W. WASSILJEW. "De 1'influence des diverses especes d'aliments sur le fonctionnement de la glande pancreatique." Arch. d. Sci. Biol., ii. 219. The same in Russian. (Inaug. Dissert. St. Petersb., 1893.) 50. A. N. WOLKOWITSCH. "Physiology and Pathology of the Gastric
Biol., viii. 1899.
(Inaug. Dissert. St. Petersb., 1898 Russian.) WULFSON." The Work of the Salivary Glands."
St. Petersb., 1898.)
disturbing in- Atropin. of deter- stomach without the animal's knowledge. 141 ct scq. effective when introduced . 152 of bile. . 115. exciter during diges- creas. 21 relation between diet of. ingastric juice. P. ineffective as an exciter of 28. 100. 126 influence of. 101 . 138 influence of "Acute" experiments. 118. for digestion. 70. " . ignorance as to function faulty earlier tion. 156. becomes an tion. 161. of. into the Amylolytic power. on flow of pancreatic juice. as a remedy for effect of. 139. . 157 influence of. when . use . 26 Botkin. 133 etseq. of gastric juice. effect of. Nerve apparatus of. 1. 146. 148. of bile. 178 Bile. mixed with meat and milk. tion of effect of. 175 Alkalinity as a factor in digestion. . 140 . influence of. 39 et scq. 117. in increasing the the sugar paste milk diet. 115 . in physiological investigations. 168 Alkaline compounds as stimuli of the pancreas. non157 . in 29. bitters on. 68. effect of. 142 . 118 . 156. 186 Bread. 15 BACTERIOLOGY mental in relation to experi- Afferent or centripetal nerves. 157 of gastric Butyric acid as an exciter secretion. means mining. 99 effect of. in inducing hypersecretion of mucus in stomach. 113. 100 but . digestion. . 157 and on its rate exciters of. .. complex character of 90. 26. on the parotid secretion. 157.INDEX ACID. digestion of. and with starch Mett's tubes. Anabolic nerve. 154. by estimation of formed. influence of bread and meat on. as exciters of the bile. . from white of egg in determining proteolytic power. 120 . 185. on sub-maxillary gland. amylolytic action of the pancreatic fluid. Gastric Juice.. action of. 156 Acidity of gastric juice. Salivary Glands. of bile. 70 on the pancreas. 113. ineffective as an exciter of gastric juice. 155 . 25. of flow. tribute to. use of. 174. 111 Borrisow on Mett's method of estimating digestive power. 118 et scq. and digestive power et seq. 154 . 162 138 Blondlot's theory of digestion. 118 . Food " under Sham the the sensation. 158 . fluence of the succus entericus on the therapeutic influence of. 91 importance of . on the ferments of the pancreatic juice. . 120 advantages of combining butter or fat with. 96 Alcohol. 131 ct seq. secretion of gastric juice. 157 in. 156. . inhibitory effects of on gland secretions. . and exciters of the flow the work of the. 116. 30 of pancreatic juice. 34 et scq. on the panineffective in causing flow . 55 fluence of. pathology. on pepsin. 99. 156 digestive products of. 116. physiology and therapeutics. 46 Appetite (see feeding). in exciting secre102 . effect of. at meals. See Albumen. 29 flow of gastric juice. new method. 27 after a . in exciting the Albumose. it is inhibitory effect of. methods of investiga- 155. 118 et seq. S. 133. 68 gastric hypersecretion. 87 of digestive fluids as factor in digestion. on . 145 .. 158 Bitters. Desire for as an excitant of the parotid secreineffective in causing flow tion.
on salivary secretion,
on the pancreas, 113
Centrifugal nerves. See Nerve apparatus
Centripetal nerves. See Nerve apparatus of Chigin, Dr., method collecting
gastric juice, 11-13
151, 152 Duodenum, reflex effect on flow of gastric juice from, 110; effect of acids
entering the, on the flow of pancreatic
citer of gastric juice, 95
effect of alkalinity the, in regulating the
state of during digestion and of fat in
Cold ineffective in promoting flow of effect on peptic glands, saliva, 1 51
the stomach, 164
or centrifugal nerves.
on the flow and
Enterokinase, the ferment of the succus flow of, excited by the entericus, 160
power of gastric
Psychic effect) Diarrhcea, explanation of, in enteritis, 161
pancreatic juice, 160, 161 Essen tucky water, inhibitory effect on flow of pancreatic juice, 126
Experimental pathology, 167 need for chair of, 182
Experimental therapeutics, 174
rleed for chair of, 182
defects in our physiology of, 2, 3 Briicke's teaching as regards, 3
and Ludwig's methods of investigating problems of, 3 the method adopted in this inquiry, 4 gastric,
Experiments essential for the solution of problems in medicine, 177 in and in therapathology, 177 et scq.
peutics, 177 ct scq. demand for institutes specially devoted to biological,
effect of exciting the sciatic nerve on,
upon an inner
Blonvation apparatus, 110 et seq. Heidendlot's researches on, 111
FAT, not an
exciter of the flow of
important additions to the knowledge of, 111 experimental pathology of, 167 ct scq.
gastric juice, 97; effect associated with flesh, 103
a probable exciter of the pancreas,
effect of, in increas-
Digestive fluids, specific activity of, 2 defective character of the deductive
ing the activity of Ihe fat-splitting ferment, 122 use of, in food, 142 ; duodenum, the seat of inhibitory
Blondlot's methods of collecting, by
effect of, 154
relationship of to their required
of bile, 156, 157 and on the ing of the stomach, 166
functions, 118, 119; inter-dependent relationship of the, 163
Digestive glands, activity of, dependent on the presence of food, 20 adapta;
power, 27, 28 power of, after a milk diet, 29, 30, 122 diges-
bility of, to special foods, 32 et seq.
Digestive system compared to a chemical factory, 2
on the heart, 146 Diseases as aids to experimental pathology, 166, 167, 173 method of experiment essential for ascertaining the
Digitalis, action of,
bable seat of influence of foregoing, action of bile on, 157 122, 123 Ferment, special nerve fibres for inducing secretion of, in saliva, 46
causes and origin
by the succus
Fibrin, influence of the succus entericus
studying pancreatic methods of forming gastric, devised by Bassow and by Blondlot, 9 dogs with several kinds of, 153 Flesh, influence of, in exciting the
excitation of special food, 33 ct scq. flow of, by "sham feeding," 50 by
the desire for food, 49 ct scq., 70 ct 76 et scq., 86, 133 ct scq. by scq. the act of swallowing, 69 ct scq. duration of psychic influence on the
secretion of gastric juice, 21, 23, 33, 74 effect of, when it is mixed with
wall to excite flow
inefficiency of stomachof
bread and milk, 21
the digestive power of gastric juice, ct seq., 77 ct scq. effect of, on submaxillary gland, 68 on parotid gland,
degree of acidity of, 87 water in exciting flow
of, 94 ct seq.
influence of chloride of sodium
effect of direct introduction of,
of hydrochloric acid, 95 inhibitory effect of sodium bicarbonate, 95, 96
stomach on quantity and
of gastric juice, 78 ct scq., 102 effect of mixing starch with, 102, 103
non-exciting effect of peptone, 96 the exciting effect being due to some;
to Chapoteau'ssan:) le
of mixing oil with, 103 et scq. preparations of, at meals, 136
and certain extracts of meat, 96 et seq.
Fodera, method of forming pancreatic fistulas devised by, 8
starch, fat, sugars, do not chemically excite flow of, 97, 98 nor do bread
of, in exciting the secretion of digestive fluids, 20 quantity of gastric juice regulated by the
quantity of, 21; gastric juice adapted to the kind of, 33 ct seq., 119 in;
evidence for the formation of chemical excitant during importance of digestion, 100 et scq. that secreted under the influence of effect of cream on the appetite, 101
or albumen, 99, 100
ability of, to excite the gastric glands by direct contact, 110 necessity of
flow and digestive excited by reflex
investigating the relation between the action of the digestive glands and each constituent of the, 128
cites flow of
pancreatic juice, 115, effect of saliva on the flow of
Fremont, Dr., isolation of dog's stomach
inhibitory effect of hydrochloric acid, 154 ample supply of water needed for formation of, 176
Fruits, use of, at meals, 114
Gastric cul-de-sac, Heidenhain's method Pawlow's method of, of forming, 11
catarrh, experimental obser-
Gelatine, flow of gastric juice excited by solutions of, 97
vations on, 132, 146
by Klemensiewicz, modification Heidenhain, 11
of Thiry used 11 method of
disease, digitalis as a
Heidenhain's method as adopted by Dr. Chigin and Dr. Pawlow, 11-14 Dr. Fremont's method, 14
Heidenhain, collection of gastric juice important additions to the by, 11
Gastric juice, ready method of obtaindefects in Heidening pure, 10
of digestion by, 111
hain's method of forming a gastric the amount of food pouch, 108 taken regulates the quantity of, 2123 qualitative changes of, during 24-26 variadigestion, hourly tions of digestive power of, after a meal of flesh, 28, 291 constant
Hydrochloric acid as a factor in digesas a stimution, 37, 38, 118,123, 141
lator of the pancreas,60, 113 ct scq ineffective as an exciter of gastric juice,
effect of varying the strength
on the flow of pancreatic
as a factor in dietetics,
flow of gastric use of, as a
Igniting juice, 101 Institute for Experimental Medicine, St. Petersburg, work of the laboratory
of the, 1 surgical department of, 16 et seq., 183 ; animals used by 183,
food, 140, 142
diet, effect of
Mucus, hypersecretion of, in stomach under the influence of irritants, 168 a normal physiological defensive
(duodenum), gastric glands influenced by reflex action from the,
function, 168 ct seq. Mustard, without action as a stimulant on the pancreas, 115 effect of oil of,
collection of gastric
inducing hypersecretion of mucus in stomach, 168
juice by, 11 Kreatin, ineffective as
an exciter of the
flow of gastric juice, 97 Kreatinin, not an exciter of the flow of gastric juice, 97
secretory, for salivary glands, methods for determining ;
the function of nerves, 47
Nerve apparatus, complete form
Kwas, a Eussian drink,
a possible factor in
centrifugal nerves, as parts of, 63 centripetal nerves, as parts of, 63 Nerve-cell as part of a complete nerve
digestion, 120, 140, 141 Latent period in the secretion of gastric
apparatus, 63 64
specific sensibility of,
on the response of 38, 124 the pancreas to stimuli, 59, 124 Lime-water, effect of, on the flow of pancreatic juice, 113-115
Nerve impulse, place of origin
Literature on which these lectures were
paths of conduction of, 63 Nitrate of silver, effect of, in inducing hypersecretion of mucus in stomach, 168 disease of peptic glands set up artificially by, 169, 170 Nutritive value of food, new criterion of, 144
nected with, 133 et seq. Meat-broth as an excitant of the gastric glands, 96, 139, 140
CESOPHAGOTOMY practised for separating the mouth from direct connection
Oil, effect of,
as exciters of flow of
of bile flow,
Mechanical stimulus of stomach-wall
with the stomach, 9 on the digestive process when associated with flesh, 103 et seq. on the flow of pancreatic juice, 121
capable of producing a flow of gastric juice, 86 et seq., 135
Medicine, past and future, 176 et seq. progress dependent on experiment, 177 et seq. Mett's method of determining the
PANCREAS, method of studying rate of its secretion by temporary fistulas, 4 " " fistula 5 the by permanent method devised and adopted by the
proteolytic 25, 26
of digestive fluids,
Milk, influence of, in exciting the secretion of gastric juice, when mixed with
Fodera's method, 8 ; inauthors, 5 fluence of milk on its activity, 22, 24 ; methods of determining the amylo;
lytic activity of secretion from, 27
and bread, 21
relation 22, 122, 143 to digestive power of pancreatic juice, 29, 30, 143 and to that
digestive power of secretion from, after a milk diet, 29, 30 ; action of
secretion of, on different foods, 38 adaptability of to required seq.
of gastric juice, 33
effect of sen-
sory stimuli on, 53 57; the vagus as a secretory nerve of the, 57 ct seq. the stimuli of the, 113 differing
Peristaltic action of the digestive canal.
effects of acids
stimulating substances as pepper, &c., 115 the gastric juice as an exciter of the, 116
Pharmacology, 179 it scq. Phosphoric acid as an aid to digestion,
the real exciting agent of secretion of, 116, note ; probable influence of fat on the secretions of the, 120 ct seq. correction of error as to the effect of can a desire for food sleep on, 123 excite the ? 124, 125 water as an ex;
Prosecretin, 117, note Proteids, flow of gastric juice not excited by, 96
acids as exciters
Proteolytic power, Mett's method of determining, 25, 26 of the pancreatic
Pancreatic juice, effect of introducing it into the stomach on pancreatic
flow, 115, 116
of bile, 157 of pancreatic juice, influence of the succus entericus on, 159
effect of bile
effect, influence of desire for
ferments of the, 157
rate of flow
and on the as an exciter of
See under Gastric juice,
the secretion of kinase, 160, 161 influence of the succus entericus on the digestive power of, 161, 162 the zymogen condition of ferments of,
vary glands, Ptyalin, 152
SALIVA flow of, nervous mechanism of 46 mixed nature of, 46 func;
Parotid saliva, effect on flow of flesh, 68 of bread, 68, 69 of acids, 152 Pawlow, the method of forming pancreatic fistulse devised by, 5-8
properties of several glands, 68 ;
application of acids, &c., to buccal cavity without influence on, 70 ; as
stomach-pouch, mode of forming, 11-13 question as to how
and swallowing, 70
the gastric secretion, 116
correctly it represents the effects of influences on the main stomach, 107
bility of, to special foods, 150, 151
action of stimuli on flow of, 151
use in the study of the pathology
of digestion, 167, 168
in disturbed states of the
supply of, Salivary glands, nerve nervous apparatus of, 65 ; 45 et seg.
large stomach, 172, 173
Pepper has no stimulating action on
the pancreas, 115 Pepsin,
excitation of, by the desire for food, 66 by noxious influences, 66 ct scq.
ready method of obtaining pure, 10 comparison between the natural and commercial forms of, 10 digestive power of, 26 ; as a remedy
Sand, effect of, on submaxillary gland, 68, 151, 152
relation to gastric
for loss of appetite, 76
bile on, 118, 157, 158 Peptic glands, disease of, set up arti-
digestion, 53 Secretin as an excitant of the pancreas, 116, note
by means of nitrate of silver, by cold, 170, 171 Peptone ineffective as an exciter of
Secretory nerve, 46, 53 Sham feeding, 50 curve of gastric secretion with, 82 gastric digestion
gastric juice, 96, 97 creatic juice, 115
and of pan;
with and without, S3
Sleep, correction of error as to the flow of the pancreatic juice being stopped
their Peripheral endings of nerves, 63 functions as receiving stations, 63, 64
Succagogues. . 166 . use of. in Mett's tubes. Stomach-pouch Sublimate (see Stomach. 102. 94 ct seq. 30 . 46 . and to that of pan96. use of. 113. 134. 120 ineffective in causing flow of bile. for determining the amylolytic power of pancreatic fluid. experimental value of. 139 Starch. supposed action of sodium salts as. 11 46. 159 influence of. Trypsin. VAGI. connection of. 156 . Surgery. on the proteolytic ferment of the . 90 et seq. et seq. . 108 et seq. 168 Submaxillary gland. 168 defensive purpose of such secretions. 98 . Sodium salts. round. importance of. 126.145. 156 THIRY. with the pancreas. Printed by BALLANTYNE. 159 flow of water not of. 109 on the large. 141 . pathological conditions associated with. . London &* Edinburgh . Stomach. burg. 141 effect of irritants in inducing hypersecretion of mucus in. 146 creatic juice. 16 as exemplified by the . for animals with pancreatic fistulas.196 INDEX. 109 et seq. 172 but they do receive reflex impulses from the duodenum. digestion of. HANSON &* Co. in determining amyloaction of. for preparation of gastric juice. 27 pancreas. with . flesh and. 103 exciter of pancreatic juice. 97. 57 Vegetables. 145. use of. 164 et seq. 68 as an of starch. except when eaten by the animal. at meals. use of. in physiological study. 141 Sympathetic nerve. 145 Spices. induced by mechanical stimulus. 100. . at meals. special nerve for inducing the secretion of. digestive fluids. not influenced by stimulation of the rectum. 115 . 107 . as a clue to the effects of influences Trophic nerves. special set of rooms in the Institute of Experimental Medicine. 102. 39 the flow of gastric juice. as also by fat in the duodenum. regulated by alkalinity being set up in the duodenum through the increased flow of pancreatic juice. method of forming a gastric pouch. 11 functions of the. 103 action of. properties of saliva from. ignorance as to the obtaining. presence of. ULCER . . 176 Wine. . Thiry's method of use of. ducing hypersecretion of mucus in stomach. 171. 140 ineffective in causing flow of bile. defects in Heidenhain's 107 et seq. 150 . therapeutic use of. 27. use of. 16-19 Sweets. 29. 125 . digestive power of. . on the pancreas. . at meals. discharge of acid contents of. the probable seat and cause of the diseased state. 140. 160 Sugar. inhibitory to the secretion of gastric juice. St. . 26 . 171 ct seq. 102 effect of mixture with flesh. . use of. . 169 . . from salivary glands. 163 . Peters- not an exciter of for digestion of. . 126 Sodium bicarbonate. entericus devised by. need of ample supply of. 137. connection process. in in- as an exciter of the flow of gastric effect of a mixture juice. WATER. 48 59 et seq. 168. use of. glands of. with the digestive with the pancreas. use of. after a milk diet. 38 power of the pancreatic secretion . on the lytic power. 53 the functional connection between the two stomachs. Soda inhibitory of free flow of pancreatic juice. effect of. 7 . 95. at meals. 141 Vinegar. 146 . method of obtaining succus tactile sensations of. in stomach. in the pancreatic juice. 115. miniature. 48 of. at meals. the stomach. . 15. 145 Succus entericus. miniature) solution. at meals.
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