Chapter IX: Section IV: From London Lancet, August 8, 1903 (1)

MEANS OF INSURING ADEQUATE MASTICATION

In order to secure the full advantages accruing from the use of the jaws and their appendages, it is, above all, necessary for them to be adequately exercised during the period of development. If this is done, not only will the tendency to dental caries, adenoids, indigestion, and other evils be greatly diminished, but the masticatory instinct will establish itself as a permanent force, so that the individual will tend for the rest of his life to subject even soft foods to thorough mastication. The tongue, the lips, and the jaws of the newly born child find their natural exercise at the mother’s breast, and we should, therefore, do our utmost to get the mother to suckle her child, the bottle affording neither the same kind nor the same amount of exercise. If, unhappily, we fail in this, we must see that the teat of the feeding-bottle is so constructed as to compel the child to earn his meal by, at any rate, some exercise. This kind of exercise promotes the growth of the tongue and thus of the jaws, especially of the mandible. Directly the infant shows a disposition to bite hard things the instinct should be gratified. We may observe a tendency in this direction as early as the third or fourth month, and it becomes more and more pronounced when, the time for the eruption of the teeth approaching, the gums begin to swell up and to get tender, and saliva begins to flow from the mouth; it is now, more than ever, necessary to provide the child with hard substances on which to exercise the jaws and the gums, and a great deal of the trouble of teething is due to the failure to recognise this fact. What, then, are we to employ for this purpose? I am convinced that it is a mistake to rely solely, or even mainly, upon baubles of ivory, coral, and the like useful though these may be in their way; it is far better to give the child something which is not only hard but nutrient and pleasant to the taste, something which will at one and the same time exercise the maxillary apparatus, excite the gustatory organs, and provide a certain amount of nutriment. To this end we may, as the teething time approaches, give a chop or chicken bone, from which most of the meat has been removed; by powdering the bone with white sugar or salt we may increase its attractiveness. From such bones a good deal of nutriment can be extracted, and this of a kind which is most acceptable to the infant stomach, for it must be remembered that the young human is in the main carnivorous. Indeed, since milk is a purely animal diet, all the mammals must be regarded as essentially carnivorous during the period of suckling, while man, as already observed, from the time he emerged from the anthropoid until he learned to cook his food, was throughout life mainly an animal feeder. Therefore we should not hesitate to allow the teething infant animal food in the form suggested. Chicken and chop bones, yielding as they do before the pressure on the gums, are, moreover, just of the right degree of consistence for the purpose in view, while they afford abundant exercise for the tongue; ivory, coral, and the like are, on the other hand, too hard and unyielding, and lack, moreover, the attractiveness belonging to sapidity.

By thus providing the maxillary apparatus with suitable exercise we shall do much to facilitate the eruption of the teeth and to favour the growth of the jaws and their appendages, including the salivary glands, and so to prepare the mouth for the reception of vegetable food. This should, of course, not be given till the teeth appear. The order in which these make their appearance gives some indication as to the order in which vegetable food should be administered to the child. The first teeth to penetrate the gums are the lower incisors which appear from the seventh to the eighth month; then follow the upper incisors from the seventh to the tenth month. These teeth enable the child to _bite_, but not, be it observed, to _masticate_, for which function the molars are necessary. Now the first molars do not appear till the twelfth or fourteenth month; the second molars not till between the fourteenth and the twentieth month; and it seems to me certain that our primitive ancestors could not have obtained starch in any quantity until they reached this age; at the best, pre-cooking man was but scantily supplied with starch, and such slender supply as he had could only be rendered accessible to the digestive juices by vigorous mastication, which broke up the indigestible cellulose framework in which all vegetable starch is contained; hence, until the young human cut his molars, he had little opportunity of securing any starch. These considerations strongly suggest the desirability of giving but small quantities of starch before the twelfth month, and though the facts, that ptyaline appears in the saliva about the time the first incisors are cut, and that pancreatic juice develops its amylolytic ferment at the same time, show that the digestive organs are ready for the reception of some starch at the seventh or eighth month, yet I believe the quantity should be strictly limited. I am ready to admit that the modern child may have, indeed probably has, a greater power of digesting starch than his remote pre-agricultural ancestor; but even so, I am convinced that we should be on our guard not to over-gorge infants with this substance. Only a small quantity should be given before the twelfth month, and it should be gradually increased up to the twentieth month.

I have said that the pre-agricultural infant was unable to secure starch in any quantity by means of his incisors. These teeth enabled him, however, to obtain some soluble nutriment from fruits, and Dr. Sim Wallace has suggested that the early eruption of the lower incisors is for the purpose of enabling the infant to pierce the outer covering of fruits so as to permit him to extract the soluble contents by suction; and, accordingly, when these teeth are cut we may allow the child to bite at such vegetable substances as apples, oranges, and sugar-cane. The latter is a useful article of diet for children, for it provides soluble saccharide in a diluted form, and it is advisable that the child should receive his cane sugar well diluted, for it must be remembered that before the agricultural period man’s supply of pure sugar was limited to wild honey which, consisting as it does almost entirely of mono-saccharide (grape sugar and fruit sugar), is very easily disposed of by the digestive organs. Nowadays, the less digestible cane sugar (which is a di-saccharide) is very largely consumed in the undiluted state, in which it is apt to set up disturbance. When, however, it is obtained by chewing the sugar-cane, it is diluted both by the water in the cane and by the saliva, and I should like to see children obtain most of their cane sugar in this way.

The consideration of the conditions obtaining for pre-agricultural man not only strongly suggests that the young human of to-day should be given starch in very moderate quantities up to the twelfth month, but it points an even more important lesson--viz., that this substance should be given not, as is the custom, as liquid or pap, but in a form compelling vigorous mastication, for it is certain that early man, from the time he emerged from the ape till he discovered how to cook his vegetable food, obtained practically all his starch in such a form; it cannot too often be repeated that uncooked starch in the natural state, locked up as it is in chambers of indigestible cellulose, has no nutritive value; these chambers need first to be broken up by prolonged and energetic chewing, and in this way much or most of the starch is converted in the mouth into dextrines and maltose, very little passing into the stomach in the crude state to set up disturbance in that organ and later in the bowel. If it is given as liquid or pap it will pass down as starch into the stomach, while if it is administered in a form which obliges the child to chew it properly, not only will the jaws, the teeth, and the gums obtain the exercise which they crave, and without which they cannot develop normally, but the starch will be so thoroughly insalivated that much of it will be converted within the mouth into maltose. How foolish to upset the child’s digestive system by deluging it with liquid starch, and then to endeavour to correct matters by giving the malt extract which the child can and should himself manufacture within the laboratory of his buccal cavity.

Clearly, then, the child should make his first acquaintance with starch, not in the form of a liquid or pappy patent food, but in a solid and somewhat tough form. The best means of achieving this end is occupying my attention, and I hope soon to publish the results of my investigation. Meanwhile, I would point out that hard, well-baked crusts constitute a convenient form in which to administer starch to children. A piece of crust may be put in the oven and re-baked; this not only hardens it but helps to convert the starch into dextrine, which is a stage on the road to maltose. If the crust be then cut into a suitable shape and spread with bacon fat or fresh butter, it constitutes a most agreeable morsel. Later, we may give hard plain biscuits. The same principle should be acted upon during later childhood and youth: we should always give, as far as possible, the starch in a form compelling abundant mastication. Loaves should be shaped so as to give a maximum of crust and a minimum of crumb, and should be baked hard. Such loaves are quite as nutritious as the ordinary ones, and much more digestible, containing as they do an abundance of dextrine and not a little maltose, and compelling efficient mastication, especially if eaten, as they should be, without any fluid. A lady who has the catering for a large number of girls gives the bread in this way, and she tells me that there is keen competition for the most crusty portions.

I do not say that starch in the liquid and pappy form should find no place whatever in man’s dietary at the present day, for this would imply the prohibition of porridge, boiled potatoes, milk puddings, and the like. We cannot put back the hand of time and return to the food of our primitive ancestors, nor is it desirable that we should; but we can, at least, arrange matters so that a large proportion of the starch we consume shall be in a form inviting mastication, such as crusts, stale bread, stale cake, biscuits, and so forth. The less children eat of pastry, or, indeed, of any luxurious foods, the better; if brought up on a healthy dietary and under healthy conditions generally, they will relish their simple fare more than the choicest dishes of the epicure. I do not, I say, object to the child consuming a certain proportion of starch in the liquid or pultaceous form, for if, by bringing him up on a rational dietary, his instinct to masticate be afforded due opportunity to develop he will be likely to subject even soft vegetable food to something like adequate mastication; this will tend to mitigate the evils associated with such food, not only by facilitating the digestion of starch, but by flushing the mouth and promoting the health of the teeth and buccal mucous membrane.

The question how far children should be allowed to crack nuts may here be considered. If the child has been brought up on pappy food, and has in consequence brittle and ill-developed teeth, the cracking of hard nuts will be likely to injure them, and this is _a fortiori_ true if any of the teeth are carious or “filled.” And not only nuts but hard food of any kind, such as ship’s biscuits, may in these circumstances injure the teeth, as many of those who went through the recent South African campaign can testify. But if, on the other hand, the child has from the beginning been fed on coarse, hard foods, so that the teeth have been allowed to grow dense and strong, no harm is likely to ensue from cracking such nuts as filberts and Spanish nuts. If a squirrel or a monkey weighing a few pounds can do so with impunity, surely the young human should be able to also. The cracking should, however, be done by the molars, while such hard nuts as Brazils had best not be tackled at all.

Animal food does not need the same amount of mastication as vegetable food, since it is not digested in the mouth, though some contend that the mixture of proteid with alkaline saliva facilitates its subsequent peptonisation. Cooked animal food is, however, all the better for some mastication, owing to the coagulation of the proteids, and, in order to insure the efficient mastication of meat, fish, and poultry, Dr. Sim Wallace recommends that they should be given in large pieces cut thin. “Flat pieces about one inch square generally _necessitate_ a certain amount of mastication. It is difficult to swallow large flat pieces of meat without mastication, but when finely minced little or no mastication is called forth.” The younger the child the more underdone should the meat be.

EXAMINATION OF THE MOUTH AND ADJACENT PARTS

If a child be brought up on the lines indicated and under healthy conditions generally, it is tolerably certain that the maxillary apparatus will develop normally, that the teeth will be strong and well opposed, and show little tendency to disease; but, inasmuch as the methods advocated are but seldom put into practice, disorders of the teeth, more especially caries and irregularities, are common, and hence with a view to promote more efficient mastication it is always advisable to examine our patient’s teeth.

Each individual tooth should be inspected in a good light for the presence of caries, and careful note should be taken of the “bite,” a normal bite implying not only a proper opposition of the two rows of teeth but the capacity of the lower ones to move freely across the upper; mere vertical movement of the mandible does not constitute efficient mastication. In this connection it must not be forgotten that an unopposed molar is useless for purposes of mastication, and it is by no means rare to find in a mouth several sound unopposed molars which are for this reason absolutely functionless. Nay, more than this, it may happen that teeth, perfectly sound ones, too, far from helping, may actually interfere with mastication; thus, among the poor, we sometimes find all the teeth gone save the upper canines and the lower incisors, and the teeth and gums being alike unable to come into contact, nothing worthy of the name of mastication is possible; it would be far better to be without any teeth whatever, for the toothless gums would then be permitted to come into contact along their entire extent, under which condition they gradually harden and come to be quite efficient grinding agents.

Next the gums, the alveoli, and the roots of the teeth must be examined, especially for the presence of erosion, tartar, and pyorrhœa alveolaris, this latter condition being evidenced by the welling-up of pus upon pressing the gums against the sides of the teeth.

If our examination of the mouth discloses anything likely to interfere with mastication the aid of the dentist should at once be sought, but every physician should be so far acquainted with disorders of the teeth as to be able to say, in the majority of cases, at all events, when this is necessary. I am convinced that far more illness than is generally supposed is attributable to dental defects, and this even among the more leisured classes. With regard to pyorrhœa alveolaris, it has to be remembered that it not only does harm by causing loosening, lengthening, and shedding of the teeth, and thus interfering with mastication, but also by contaminating the stomach and the blood and thus upsetting the digestion and causing constitutional diseases, such as anæmia and arthritis; and inasmuch as poisonous discharges from the nose, the naso-pharynx, the pharynx, and the tonsils may act in a similar way, these parts also should be inspected in connection with the examination of the teeth. In the dust-laden atmosphere of towns they are very liable to disease, and even when healthy are necessarily dirtied; some go so far as to advise all town dwellers daily to wash out the nasal passages and to gargle the throat; but, whatever may be thought of this, it is certain that under existing dietetic conditions special means are needed in order to keep the mouth and teeth clean. When man fed on raw food this was not necessary, the food itself and the copious flow of saliva, induced by prolonged mastication, effectually cleansing these parts; but, under present conditions, food tends to remain within the mouth, especially between the teeth and in their crevices, and therefore special means are needed to remove it. This is done by “cleaning the teeth” and by rinsing the mouth.

_The tooth-brush._--Probably the ideal method of cleaning the teeth is that adopted by many primitive and not a few semi-civilised peoples--viz., rubbing them with a twig of wood which has been teased out at one end so as to form a sort of brush by means of which the teeth can be burnished and food dislodged from them. The modern tooth-brush requires to be used with great caution, as it is capable of doing much harm, not only by removing the mucoid film, which, according to Dr. Wallace, protects the teeth from corroding agencies,[28] but probably also by injuring the edge of the gum and the neck of the teeth, and thus setting up the condition known as “erosion.” Certain it is that some of the best sets of teeth I have encountered have been wholly unacquainted with the tooth-brush. In any case the brush should be employed with great care; it should be soft, and should always be drawn away from the gums both on the inner and outer aspect of the teeth towards the biting surface, as well as across the latter, never transversely across the outer surfaces, as so frequently is done. The object of these procedures is to dislodge any particles of food that may have collected between the teeth or in their crevices. For this purpose the toothpick may also be employed judiciously. In order to render the enamel of the teeth white it is better to rub each tooth carefully with some soft material, such as chamois leather, rather than to scrub them with a brush. Tooth-powders should not be used as a matter of routine, but only occasionally and for appearance rather than for cleanliness, and should consist of some simple non-irritant material. Antiseptic powders and washes are to be scrupulously avoided, for it is neither desirable nor possible to render the buccal cavity aseptic; myriads of bacteria flourish within it, many of which play a useful part as scavengers. The time of all others for cleaning the teeth is just before going to bed, so that the food shall not be allowed to decompose in the mouth during the night. There will then be no need to use the tooth-brush in the morning.

_Rinsing the mouth._--The mouth should be rinsed out as a matter of routine after each meal and on rising, and care should be taken to do this before the early cup of tea, so as not to contaminate the stomach with the buccal secretions which have accumulated during the night. Inasmuch as raw vegetable food has a cleansing effect on the teeth, it is often a wise plan, especially in the case of children, to finish a meal with some kind of fruit, such as an apple or an orange. It hardly seems necessary to insist upon the necessity for keeping all artificial dentures thoroughly clean.

PROFESSOR PAWLOW’S DEMONSTRATIONS OF PSYCHIC INFLUENCE IN DIGESTION

[In presenting a theory of human alimentation involving mental or nervous as well as mechanical and chemical factors which influence it for good, it is not often that an author is able to enlist the assistance of a complete battery of scientific confirmation to fortify his own crude observations taken direct from personal experience in the study of natural requirements.

Professor Pawlow, with his marvellously skilful investigation of the workings of the digestive secretions, and Dr. Cannon of the Harvard Medical School, by aid of persistent and patient X-ray studies, explain how it is that earned appetite and thorough mouth-treatment of food are preliminary necessities of easy digestion, and that disturbance or shock of any sort during the process stop digestive proceedings and endanger health. They show also that when the mouth is used to do _all that it can do_ in the work of digestion all the rest is easily accomplished by the NATURAL AUTOMATIC PROCESSES within the body.

They both show that we have, each of us, a certain responsibility in the matter of right digestion and healthy nutrition, and that all this personal responsibility is located in the head, in the mind, and in the mouth, and that while the alimentation is proceeding it is a sacred duty to do our part _right_, according to the intelligence that these most valuable demonstrations teach.

Professor Pawlow has allowed publication of his lectures in Russian and German, and recently Professor W. H. Thompson of the Physiological Department of Trinity College, Dublin, has made an English translation which is issued by Charles Griffin & Company of London and J. B. Lippincott of Philadelphia.

The author has to express special gratitude to Professor Pawlow, Professor Thompson, Messrs Griffin and Lippincott for permission to reprint herein some entire lectures and extracts that bear especially on the practical understanding of our subject.

Professor Pawlow is one of the Board of Scientific Assessors mentioned in the REPORT of a PLAN for an INTERNATIONAL INQUIRY into the subject of HUMAN NUTRITION.

In one of the lectures, not here reprinted, Professor Pawlow gives merited recognition of the early statements of the French physiologist Blondlot relative to psychic influence on the digestive secretions made some half century ago, but discredited by physiologists since that time, owing to insufficiency of evidence brought forward in support of the statements.

Professor Pawlow’s acknowledgment is so gracefully rendered that it is here given as a model of scientific courtesy.

“I have depicted the work of the gastric glands as we have seen it in our experiments, and as it has developed under our hands. Is the picture a new one? In its details, yes; but not in its fundamental features. However singular it may appear, the sketch of this picture was more than fifty years ago outlined by physiology. May this constitute another reason for our science relinquishing its characteristic shyness of new things and for its conversion to our interpretation of the phenomena under consideration!

“The talented author of the _Traité Analytique de la Digestion_--Blondlot--spoke in plain words of the importance of taking food, and of the specific excitability of the gastric mucous membrane. The facts adduced in the working up of his theory were naturally insufficient, but we must not forget that the first experiments on dogs with artificial gastric fistulæ had only just been performed. It is truly incomprehensible that the researches of Blondlot and his views upon the secretion of gastric juice have experienced during the past fifty years no completion, no additions, but, on the contrary, have passed out of sight, thanks to the faulty experiments and erroneous representations of later authors. Only in the works of a few writers--and those mostly French--has Blondlot’s theory survived. Of other investigators we must give mention to Heidenhain, who has enriched the physiology of absorption in general, but more especially, in connection with the secretory work of the stomach, has discovered many important facts and has given birth to many fruitful ideas. From him proceed the subdivision of the secretory process according to periods and exciting agencies, as well as the suggestion that it would be important to investigate the individual food-stuffs in relation to the work of the stomach. Heidenhain’s results are contained in his well-known article on the secretion of the cardiac glands of the stomach, published in the year 1879 in PFLÜGER’S ARCHIVES. The work of Blondlot and the additions of Heidenhain comprise almost everything of importance which was accomplished by physiology in fifty years concerning the conditions and mechanism of the secretory work of the stomach during digestion. Full of moment, however, for our subject was the obvious error that mechanical stimulation constituted an effective excitant of the gastric glands, and this error was in its turn a result of faulty methods.”—HORACE FLETCHER.]

LECTURE IV

GENERAL SCHEME OF AN INNERVATION MECHANISM--THE WORK OF THE NERVOUS APPARATUS OF THE SALIVARY GLANDS--APPETITE, THE FIRST AND MOST POTENT EXCITER OF THE GASTRIC SECRETION

Constituent parts of a complete innervation mechanism--The special duty of the peripheral terminations of afferent nerves--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 glands; their particular properties--Differences between the exciting agencies of the different salivary glands--Discussion of the sham feeding experiment--Mechanical and chemical stimulation of the cavity of the mouth has no effect on the gastric glands--The experiment of Bidder and Schmidt relative to psychic excitation of the gastric secretion--Conditions for success in this experiment--The passionate longing for food--the appetite--alone brings on the secretory effect in the sham feeding experiment.

GENTLEMEN,--As you have learned in the last lecture, and also in part have seen by direct experiment, the nervous system can influence the work of our glands in the most diverse ways. The vagus nerve, already burdened with many duties, has, in addition, proved itself to be an undoubted exciter of the gastric glands and of the pancreas. But we must also assign to the sympathetic nerve a similar _rôle_. This is a matter which cannot be doubted, so far as the pancreas is concerned, and is highly probable as regards the stomach. We also saw good reason for believing that these two nerves contained two different classes of fibres, secretory and trophic, a condition which had already been proved to exist by Heidenhain for the nerves of the salivary glands. As a hypothesis we might even have proceeded a step farther and have divided Heidenhain’s trophic nerves into separate classes of secretory fibres. Lastly, we advanced important experimental evidence to show the existence of special inhibitory fibres to the glands, and these fibres also run in the vagus, the list of whose functions seems almost interminable.

We obtained these results by division and artificial excitation of the nerves which run to the glands. But when, how, and by what means these nerves are thrown into activity during the normal course of physiological events remains a question.

In order to avoid repetition, and at the same time impart the utmost clearness to our representation, it may be useful to bring before your minds at once the plan of innervation of a given organ, all the more since this scheme is seldom completely followed out or adequately described in physiological text-books. Consequently, it is not borne in mind with sufficient precision by the majority of medical men.

A complete innervation mechanism consists of the peripheral endings of the centripetal (afferent) nerves, the centripetal nerves themselves, the nerve cells (a group of nerve cells connected with each other is termed a “nerve centre”), the centrifugal (efferent) nerves, and, lastly, their peripheral terminations. Physiology now accepts it as a settled fact, that nerve fibres serve only as _conductors_ of nervous impulses, which come in from contiguous links of the nervous chain. Only the peripheral endings of nerves and the nerve cells themselves have the power of transforming the external stimulus[29] into a nervous impulse. In other words, in the intact organism these alone constitute the normal receiving apparatus of the nervous system. Whether the peripheral ends of centrifugal (efferent) nerves are likewise able to function as normal sites for the application of external stimuli has still to be answered. Consequently, when any external agency excites the peripheral terminations--the receiving stations--of centripetal nerves in this or that organ, the effect of the stimulus will be conveyed through the centripetal nerves, as through a receiving wire, to the central station--the nerve cells. Here it becomes changed into a definite impulse and now comes back along the centrifugal nerves--the outgoing wires.

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, respond to _specific_ stimuli; that is to say, are able to transform definite kinds of external stimuli into nervous impulses. The function of the end organs with which they are connected is therefore of a purposive nature; in other words, these organs are only called into play by certain definite conditions, and impart the idea of being aware of their purpose, of being conscious of their duty. We have long known that the peripheral endings of sensory nerves are possessed of a high degree of speciality, and cannot therefore have any doubt regarding the specific nature of the end organs of other centripetal nerves. This is a sore point in present-day physiology. But, notwithstanding our knowledge of the separate parts of the animal body, we shall only be able to form a true conception of the motive agencies of the whole complicated machine, when we have established the specific excitability of the end apparatus of every centripetal nerve, and have discovered all the mechanical, chemical, and other factors which throw this or that end apparatus into an active condition. I always look upon it as a period of scientific inadequacy so long as the effects of the most diverse external agencies upon any normal physiological process are admitted to be indistinguishable. As the work of the digestive canal is now represented in the majority of text-books, and consequently presented to the mind of the physician, it bears the impress of this period. To impart to the physician a more correct conception of this matter was my chief object in giving these lectures. I hope, indeed, to furnish you with evidence sufficiently convincing, that the alimentary canal is endowed not with mere general excitability; that is to say, does not respond to every conceivable form of agency, but only to special conditions which are different for the different portions of its length. 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, so every organ, indeed every cell of every organ, can only maintain its place in the animal microcosm, and adapt itself to the activity of innumerable associates, as well as to the general life of the whole, by virtue of the fact that the peripheral end apparatus of its centripetal nerves possesses a specific excitability.

The same applies to the nerve cells: obviously they are endowed with specific sensibility. Irrespective of the excitations which are communicated to them from centripetal nerves, they respond, as originators of nervous impulses, only or at least mainly to definite forms of mechanical, chemical, or other stimuli arising in the organism. This follows not alone from a number of physiological facts but also from various pharmacological data. Thus we learn that various drugs excite or annul the activity of definite portions of the nervous system, at least in the earlier phases of their effects. This specific excitability of nerve cells, just as much as the same property of peripheral end organs, lies at the bottom of the purposive action of these organs.

Hence, our next duty is to endeavour to discover the normal exciting conditions of the centripetal nerves belonging to the glands which we had under consideration in our last lecture, or, more correctly, to find out the conditions which excite the centres, as well as the peripheral endings of the different nerves, which form parts of the nervous apparatus of these glands. We have, therefore, for each phase of the work of secretion, to find out that portion of the nervous mechanism which is for the time being under excitation, and to discover the primary agency by which this condition is elicited. This would include an exact analysis of the stimulating influence which mastication and food exert upon the nervous mechanism of these glands. We shall also be able more fully to comprehend the inner mechanism underlying the facts which formed the subject of the second lecture. This, of course, is an ideal programme which we can only follow out as far as the present state of physiology permits. It may now be instructive, and, for our further conclusions, advantageous, to glance shortly at the nervous control of the salivary glands.

The salivary glands, whose innervation has long ago been investigated, have generally been accepted as types of the deeper-lying digestive glands, and when it became necessary to form a conception of the mode of activity of the latter, medical science resorted to a bold analogy and thought of the nervous apparatus of the salivary glands. But the attempts of investigators to apply rigidly to others the scheme of innervation which holds good for the salivary glands, 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 abdominal glands. We have already had an example of this nature before us. In the salivary glands we have no clearly marked indications of nervous inhibition, and this circumstance has decidedly retarded the due development of our knowledge of the nervous control of the abdominal glands. Authors naturally expected to see a simple and prompt stimulation-effect from the same conditions of experiment which sufficed for the salivary glands, and the failure of this gave them, as they thought, the right to deny the existence of any extrinsic nervous influence upon the abdominal glands. The error is now obvious; the abdominal glands behave in some ways different from the salivary glands, and for their successful investigation, other conditions of experiment are necessary than those which held good for the former. In the working of the abdominal glands nervous inhibitory processes play a large part, but they are almost wholly absent in the case of the salivary glands. This is an additional warning that one must never push the conclusions drawn from analogy too far, but must constantly bear in mind that the life-functions of all organs are extremely complicated, and that the work of even the most apparently similar organs should be submitted to separate and careful observation. To me it appears that the unjustified analogy drawn between the abdominal and salivary glands has to be credited with another important misapprehension. And precisely for this reason I think it desirable to bring under consideration, if only in brief fashion, the conditions of work of the salivary glands, especially since Dr. Glinski has instituted in the laboratory some easily performed experiments which bear upon the matter.

The experiences of daily life teach us from the outset, that the activity of the salivary glands begins even before the introduction of food into the mouth. With an empty stomach, the sight of food or even the thought of it is sufficient to set the salivary glands at once into activity; indeed, the well-known expression, “to make one’s mouth water,” is based upon this fact. Hence a psychic event, the eager longing for food, must be accepted as an undoubted excitant of the nervous centre for the salivary glands. On the other hand, the same every-day experience, as well as numerous experiments upon animals, teach us that a number of substances, when brought into contact with the mucous membrane of the mouth, are likewise able to call forth a secretion of saliva. One even acquires the impression that everything brought into the mouth may reflexly influence these glands, the only difference being a gradual shading off in the effect, dependent upon the strength of the stimulation which the substance introduced is able to exert, and it appears to me that it is precisely this impression which has driven the idea into the background, that the peripheral end apparatus of the centripetal nerves of the digestive canal are specifically excitable. The facts were here correctly observed, but their indications erroneously interpreted.

The great multiplicity of excitants of salivary secretion, has without doubt, some connection with the complicated physiological functions of the saliva. This is the first fluid encountered by everything which enters the alimentary canal. It must, therefore, in a sense play the part of host to every substance taken in--moisten the dry, dissolve the soluble, envelop the hard and bulky with mucus in order to facilitate its passage down the narrow œsophagus; and submit certain forms of food material, such as starch, to a process of chemical elaboration. Nor is its duty by any means ended here. The saliva is secreted in the first compartment of the alimentary canal, which is at the same time the sorting-room of the organism. Much of what enters the mouth may prove in the testing process to be useless, or even noxious, and must either have its deleterious properties neutralised or be completely rejected. The saliva is secreted in the first instance to obviate injurious effects in some way; thus, for example, a strong acid is to a certain degree neutralised, while other corroding substances may be simply diluted, and by mere reduction of concentration have their harmfulness diminished.

In the second place, when the injurious substances have to be wholly removed, the saliva plays the _rôle_ of a washing-out fluid; otherwise the material, by clinging to the mucous membrane of the mouth, 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 all in physiology, and yet it is evident that the saliva, as a cleansing fluid, must have a wide importance. If you only think of how often we are impelled to expectorate, that is, to wash out the mouth with saliva after something unpleasant, this will be clear. Such a view finds additional support when we reflect that a feeling of disgust produces almost as strong a flow of saliva as the sight of a tasty meal. In both cases the secretion performs the office of forerunner: in the first it prepares for the washing out of the mouth, in the second for the requisite elaboration of the food. Think how often, when something disagreeable enters the mouth, with what rapidity the saliva is poured out, even after the unpleasant substance has been for a considerable time removed, and not a trace more is apparent to the sense of taste. Indeed, long afterwards one has only to recall the circumstances to mind in order to bring on anew the secretion of saliva. Apparently the psychic excitation of the nerves of salivary secretion also ushers in the act of vomiting, which, as is well known, can be called forth by mental influence. Further, the function of the saliva just mentioned is probably the true physiological explanation of the feeling of disgust which many persons experience at the sight of the secretion itself.

Hence I hold that substances which obtain entry to the mouth set up a secretion of saliva only because we have here the seat of a definite physiological sense, and not because the peripheral terminations of the buccal nerves are devoid of specific excitability, and capable of being thrown into action by every desired form of stimulus. In other words, the specific excitability of the peripheral endings of the salivary nerves is very comprehensive and widely extended. This is no picture of the imagination, for it can be supported by facts. To say nothing of the testimony of earlier authors, that the salivary glands have each particular exciting agencies to which they specially respond, we can demonstrate the following facts from the material collected in our laboratory.

Dr. Glinski isolated the orifices of the salivary glands in dogs with portions of the adjoining mucous membrane, brought them out of the oral cavity, and caused them to heal into the edges of the skin wounds. In his first animal the ducts of the submaxillary gland were thus led outwards. By means of a Mendeljeff’s clip, the wide end of a conical funnel of waterproof material was attached to the skin surrounding the orifice. To the narrow end a small test-tube, which served to collect the saliva, was attached by a wire. I now offer such an animal a piece of flesh, and, as you see, the tube fills up at once with saliva. I stop tempting the dog, hang on a new test-tube, and give it a few pieces of flesh to eat; once more a strong secretion of saliva results. A new tube is now attached to the funnel, the dog’s mouth is opened, and a pinch of fine sand thrown in; again there is a flow of saliva. Once more a new test-tube; and now I apply to the buccal mucous membrane, the plume of a feather dipped in acid solution, with the result that I obtain a strong flow of saliva. One may employ a number of substances in this way, when a similar effect is always produced. You see, in this, such a comprehensive excitability of the innervation apparatus of the salivary glands that you might readily interpret it as meaning the power of response to all and sundry forms of stimulation. We now proceed, however, to another dog, whose parotid duct has in a similar manner been diverted outwards. The saliva is collected in the same way. We tempt the dog with a piece of flesh, but to our astonishment no saliva flows, and yet the animal is most eager for the savoury meal offered. Now we give it some raw flesh to eat; again the secretion of saliva 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. Probably you will say there is something wrong, either with the method or with the glands of the animal. But wait a little. I now give the dog finely powdered dry flesh, and obtain at once an abundant secretion. Should any one happen to think that the variation in the result is dependent, not on a different specific activity of the glands, but on individual differences in the dogs, I respond that Dr. Glinski has had an animal with double parotid and submaxillary fistulæ, and was able to observe on one and the same dog, a like behaviour on the part of the glands to that which we have just seen in two different individuals. An analogous experiment with bread was also carried out by Dr. Glinski. The eating of fresh moist bread produced no secretion worth mentioning, while dry bread, on the other hand, caused the saliva to flow in large quantities. The results of this experiment permit us to draw extremely instructive conclusions. In the first place, the several salivary glands are, as a matter of fact, very sharply differentiated in the conditions necessary for their activity--that is to say, in respect to the agencies which excite their nervous mechanisms. Secondly, the innervation apparatus of the parotid manifests a very sharp selective power in the choice, so to speak, of an adequate stimulus. The mechanical effect of large pieces of flesh is naturally much greater than that of the finely powdered material, and yet it was precisely to the latter that the glands responded. The stimulus is, therefore, not due to the mechanical, but to some other property of the food. This other property is obviously the dryness of the material. Our example illustrates how that which we may term “purposiveness” comes into play in the working of our glands and also how erroneous is the opinion that the mechanical stimulus is all potent. Indeed, previous authors have already pointed out that dry substances cause a specially free secretion of saliva, and yet physiological opinion throughout the length and breadth of the land, as expressed in text-books, has chosen to recognise a _universal_ instead of a _specific_ excitability. Dr. Wulfson, who is at present carrying on the investigation of salivary secretion in our laboratory, has added a very interesting observation to the results of Dr. Glinski already related. The parotid gland, which is hardly, if at all, excited when one offers fresh meat to the animal, responds with a very active secretion, when dry food (bread or powdered meat) is offered. This phenomenon is all the more surprising since the desire of the animal for eating is much more strongly excited by flesh than by dry bread. 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.

The second reagent which is poured out on the raw material in the digestive canal is the gastric juice. How, in the normal course of events, is the work of the gastric glands, which prepare this juice, called into play? With the first, and manifestly important factor, which has a relation thereto, you are already acquainted, and, indeed, have already seen. I refer to the production of gastric juice in the empty stomach, as a result merely of the swallowing of food in the so-called sham feeding of an œsophagotomised dog. When one takes into consideration the absolute independence of this factor, and the intensity of the effect, which makes itself evident in the secretion of a large quantity of juice of high digestive power, the exciting agency which brings about such secretion must be recognised as one of the most important and effective processes in gastric digestion. But in what does it consist? At first sight it appears--and when I previously drew your attention to 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, similar to the reflex excitation, _e.g._, of the parotid gland, by finely powdered flesh thrown into the mouth. Now, however, I assert quite emphatically that this is not the case. We have, it is true, in the activity of the salivary glands an analogous phenomenon to indicate--not, however, that of which we have just spoken. We might apply every conceivable form of stimulus which could possibly come into play in the act of eating, and yet would not obtain the slightest indication of secretory activity in the stomach. In this dog with a gastric fistula, and with also a divided œsophagus, I will try such an experiment, using the most effective chemical stimulus to the buccal mucous membrane, viz., acid solution.

The secretion of saliva begins at once, as you see; the acid is, therefore, effective. From the stomach, however, in spite of continued excitation, no secretion results, although the acid, mixed with the saliva, is swallowed and flows out again from the upper segment of the œsophagus--that is to say, passes along precisely the same path that the food takes in sham feeding.

We could experiment in the same way with a number of other substances: saline, bitters, pepper (strong local excitation), mustard, and so on, and always with the same results; a free secretion of saliva, but perfect quiescence of the gastric glands. We may even, with the same object, employ the soluble constituents of flesh in the form of a decoction, and likewise observe, in most cases at least, no sign of activity on the part of the gastric glands.

With the chemical we may also combine a mechanical stimulus. We can, for example, wipe out the mouth with a sponge soaked in the solution to be experimented with, but always with the same negative result. We may finally give such pieces of sponge, or even smooth stones of considerable size, to the dog to swallow, passing them back behind the anterior pillars of the fauces and allowing them to fall out again, from the upper portion of œsophagus. It may be added that a well-taught dog puts up with all these procedures without the slightest protest. You see that all the manipulations in this case are carried out with bare hands and without instrumental aid. One can easily train a dog to swallow stones which are placed in the anterior part of the buccal cavity. It simply makes a few chewing movements and swallows them down. The dog on which the acid experiment has just been made serves also for the swallowing of the stones. The attendant now places some pebbles in the front part of the mouth, when the animal rolls them round, as if chewing and gnawing them, and then swallows them. The stones fall out, as you see, from the œsophagus, and drop with an audible sound upon the table. This play with the stones has now lasted fifteen or twenty minutes (in the laboratory we have often kept it up for hours), and yet not a drop of gastric juice is to be seen.

In order to prove that the dog is perfectly healthy and normal, we lay aside the stones and proceed to our old experiment of sham feeding. As you see, the first drop of gastric juice makes its appearance precisely at the end of five minutes, and after a further five minutes we have collected more than 15 c.c. of the fluid; consequently there can be no doubt that in this dog both gastric glands and nerves are uninjured and function in normal manner. At one time we even had a dog which voluntarily took the stones out of one’s hand and swallowed them; the sagacious creature had seen our object in previous experiments and learned to perform it of its own accord! But in this case also the result was negative.

Clearly, therefore, neither chemical nor mechanical stimulation of the buccal mucous membrane is capable of reflexly exciting the nerves of the stomach. Further, it is obvious that the excitation of these nerves in sham feeding is not the result of a stimulation coincidently produced; that is to say, the excitement of the chewing and swallowing centres does not imply simultaneous action of the secretory centre of the gastric glands. In what, then, does this influence consist which is intrinsic to the sham feeding, but which we have not been able to reproduce in our analytical investigation? There is only one thing to think of, namely, the eager desire for food, and the feeling of satisfaction and contentment derived from its enjoyment.

It has, indeed, been known for forty years, thanks to the experiments of Bidder and Schmidt, that at times, the offering of food to a hungry dog, in other words, the excitement of a keen desire for it, is sufficient to cause a flow of gastric juice from the empty stomach. We shall presently have occasion to observe the force of this physiological factor. Here I bring before you another dog, likewise having a gastric fistula with divided œsophagus. The stomach has been washed out half an hour ago, and since then not a drop of gastric juice has escaped. We begin to get ready a meal of flesh and sausage before the animal as if we meant to feed it. We take the pieces of flesh from one place, chop them up, and lay them in another, passing them in front of the dog’s nose, and so on. The animal, as you see, manifests the liveliest interest in our proceedings, stretches and distends itself, endeavours to get out of its cage and come to the food, chatters its teeth together, swallows saliva, and so on. Precisely five minutes after we began to tease the animal in this way the first drops of gastric juice appear in the fistula The secretion grows ever stronger and stronger, till it flows in a considerable stream. After the lapse of a few minutes we can count the number of cubic centimetres by tens. The meaning of this experiment is so clear as to require no explanation; the passionate longing for food, and this alone, has called forth under our eyes a most intense activity of the gastric glands. If the experiment be frequently repeated, one can easily observe that the keener and more eager the desire on the part of the dog for the food, the more certain and intense is the secretory effect. In extreme cases there is even a quantitative relationship between this effect and that of the sham feeding.

Here is an experiment of Professor Ssanozki, in which the secretory effect of the mere tempting of the animal with the sight of food is compared with that of sham feeding. A few threads of alkaline mucus had just escaped from the stomach, and then the excitation of the dog with flesh was begun. After six minutes the secretion commenced and continued as follows: