The tonsils are examples of follicular lymphoid structures. They lie one on either side of the entrance to the gullet, between the two folds (the anterior and posterior pillars of the fauces) by which the soft palate is continued to the side of the tongue. Normally the tonsil is not visible, but when inflamed it may project sufficiently to be seen; and its surface may then be covered with mucus and pus. It is liable to become enlarged in childhood, owing to chronic inflammation. A section of the tonsil shows it to consist of clusters of lymph-follicles lying beneath the mucous membrane. The term “follicle” is unfortunate. It conveys no idea of the form or structure of one of these masses of lymph-cells; and it is, besides, applied to things of an entirely different character—for example, the pits of mucous membrane which sink down between the masses of lymphoid tissue in the tonsil. The expression “follicular tonsillitis” does not refer to the lymph-follicles, but to the epithelial pits. It is a condition in which a drop of pus is to be seen in the mouth of each of the pits. A lymph-follicle is a small rounded clump of connective tissue, denser on its periphery than in its centre. Its bloodvessels are disposed chiefly on the periphery. Lymphatic streamlets arise in the centre. Its outer portion is closely packed with dividing lymph-cells and young leucocytes, which as fast as they are formed migrate towards the centre, and eventually escape from the follicle by the lymphatic vessels. The connective tissue which invests and separates the follicles is full of leucocytes. Removal of the tonsils is followed by no ill effects. They are not essential to our well-being. Nevertheless, they have important functions to perform. They are barracks crowded with leucocytes, which guard the pass into the alimentary canal. Their leucocytes incessantly patrol the mucous membrane, capturing germs, removing fragments of injured epithelium, striving to make good the mischief to which this part of the alimentary canal is peculiarly liable. The enlargement of the tonsil which results from frequent sore throat is a response to the demand for an increase in the supply of these little scavengers, in order that they may cope, not only with objectionable things outside the walls, but with the still more pernicious germs which during an attack of sore throat succeed in breaking through the epithelium. It is the invaders which elude the vigilance of the leucocytes that cause fever and other general symptoms. Other notable groups of lymph-follicles are found in the middle portion of the small intestine, where they form oval patches, about three-quarters of an inch long by half an inch broad—Peyer’s patches. The leucocytes which are developed in them search the walls of the intestine for germs. During an attack of enteric fever the patches become inflamed, and one of the greatest risks which the patient runs is the risk of ulceration of a patch and the perforation of the intestinal wall.
The abundant provision for the multiplication of leucocytes shows that the destruction of these cells must occur on an equally large scale. Every day large numbers die. Where this occurs, and how their dead bodies are removed, is not certainly known. Doubtless they are eaten by their fellows, their substance oxidized, and the products—carbonic acid, water, and nitrogenous waste—thrown into the lymph. There is some reason for thinking that a part of the nitrogenous waste is excreted in the form of uric acid ([cf. p. 216]). The daily production, and consequent destruction, of leucocytes shows that their metabolism is a factor which cannot be overlooked when we are making up the body’s accounts.
The fixed tissues receive their nutriment in a digested condition. Leucocytes digest it for themselves. In many cases, although not in all, the cells of fixed tissues last throughout life, so far as their outer form is concerned, although their molecules are oxidized and replaced by new material. It is not improbable, therefore, that there is a difference between the metabolism of the fixed tissues and the metabolism of leucocytes. The whole of a wandering cell, its nucleus included, breaks down and has to be removed. We do not know that this occurs in the case of a fixed cell. On the strength of evidence which points, apparently, to a chemical relationship between nuclear substances and uric acid, it has been inferred that the two chief nitrogenous products which are excreted by the kidney are divisible into the one which in the main represents the oxidation of fixed cells, urea, and the other, uric acid, largely derived from the oxidation of wandering cells.
The valiant leucocytes do their best to cope with all the rubbish, whether living or dead, that needs removal. They flock to any situation in which germs are numerous or tissue has been destroyed. If all goes well they take the foreign matter into their substance—dead tissue is matter foreign to the body—and either digest it in the course of their ordinary progress, or retreat with it, if they cannot digest it, to the nearest lymphatic gland. But in their efforts to reach objectionable matter they are apt to wander too far from the healthy lymph from which they obtain oxygen for their own respiration. Unable to breathe, they die. They lose the power of extruding pseudopodia. Their extensible, prehensile processes are drawn in. Assuming a globular form, they float helplessly in what once was lymph. Their body-proteins are largely changed to fat. As “pus cells,” they are thrown off in the discharge from an ulcer, or accumulate in the cavity of an abscess. A pus cell is a dead and fattily degenerated leucocyte.
The third kind of breeding-place of leucocytes, a lymphatic gland, has a more elaborate structure than the tissues with which we have already dealt. Lymphatic glands are about the size of beans, and of the same shape. They are found in the course of lymphatic vessels in situations where they are not exposed to pressure, such as the back of the knee, the groin, the front of the elbow, the armpit, in the neck above the collar-bone, and on either side of the sterno-mastoid muscle, behind the angle of the jaw. There are a number in the abdomen and in the thorax. Each lymphatic gland is invested by a strong fibrous capsule. Its artery enters, and its vein and efferent lymphatics leave, the concave side (the hilus) of the gland. The lymphatic vessels which bring lymph to it pierce the capsule on its convex side. It is divisible into two parts: (1) The adenoid tissue which surrounds the artery and its branches; (2) the open network of “lymph-ways” which invest this adenoid tissue. Leucocytes divide in the adenoid tissue. The young lymphocytes drop out into the lymph-ways. As a stream of lymph, brought by the afferent vessels, is always flowing into the lymph-ways, and out by the efferent vessel or vessels, the lymphocytes are carried with it towards the thoracic duct. A lymphatic gland is therefore an organ for adding leucocytes to lymph in the course of the lymph-stream. It has, however, another and equally important function. Leucocytes which have picked up germs or other foreign matter pass on with the lymph to a lymphatic gland. After entering its lymph-ways they leave the lymph-stream, squeeze into the adenoid tissue of the gland, and there come to rest with their burden. They remain in the gland until the foreign matter is digested, or, if it be indigestible, until they undergo dissolution, when the particles of soot or pigment are deposited from their débris in a harmless state. When the skin is tattooed, much of the Indian ink and other pigment remains where it was inserted with the needle, but some of it is picked up by leucocytes and carried to the nearest lymphatic gland.
Lymphatic glands are barriers which stop the spread of infection. They are the stations to which our police carry captured germs. The skin of the heel is abraded. Germs from the soil, or elsewhere, which have accumulated in a dirty stocking—owing to the warm moisture enclosed by an impervious boot, the woollen covering of the foot is a peculiarly healthy place for germs—enter the opened lymph-spaces of the subcutaneous tissues. Leucocytes hasten to the spot. They seize the invaders with their pseudopodia, engulf them in their body-substance, enter lymphatic vessels, and are rolled away by the lymph-stream. The instinct which brings them in ever-increasing numbers to the breach in the protecting skin can be explained only in terms of force. From our own conscious action to the causes which determine the movements of a leucocyte, or of an amœba, is so deep a drop that we prefer to recognize in the latter a merely chemical attractive force. “Chemiotaxis” we term the influence which draws leucocytes to the place where food is abundant; although it is also the place, one must admit, where in the interests of the body as a whole they run great risk of asphyxiation. It is appetite which draws a schoolboy to a bun-shop; a sense of duty prompts a fireman to risk his life in a chamber filled with smoke. We have no desire to humanize a leucocyte; but it is difficult to emphasize too strongly its independence. It would be absurd to use terms which imply that a leucocyte has a self-directive power; yet it is equally misleading to describe its migration to the seat of injury, its retreat with ingested germs to a lymphatic gland, its wriggling from the lymph-ways of the gland into the shelter of its adenoid tissue, in terms which imply that the forces which direct it are known, and their mode of action understood. The success which attends the inroads of germs is due to their amazing capacity for multiplication when they reach lymph or blood. It is useless to attempt to form an idea of the rapidity with which they divide, since we have no data upon which to base calculations. If the leucocytes fail to deal with the first few that enter, germs soon swarm within the lymph-vessels. This leads to an inflammation of the walls of the vessels, which may then be seen as red lines beneath the skin. These red lines lead upwards towards the nearest lymphatic gland. The glands in the space behind the knee are not usually affected when the focus of infection is in the foot. The red lines can be traced up the inner side of the knee and the front and inner side of the thigh to the groin. The glands in this situation swell until they can be easily felt. If the mischief is in the hand, the gland at the elbow may be affected, but most of the lymphatics pass by it on their course to the glands in the armpit. If a sore throat is the source of infection, the glands beneath the angle of the jaw enlarge. Thus various glands block the further progress of infection. In doing this their resources may be strained to the uttermost; they may enlarge, become tender, grow soft, fill with pus, break down and discharge the pus without the aid of a surgeon’s knife, although as soon as pus is recognizable within them it is wise to let it out. If germs pass through these first stations into the lymph-vessels beyond them, abscesses are formed in other situations. A condition of “blood-poisoning,” so called, is set up.
The readiness with which leucocytes sacrifice themselves in their efforts to remove germs and decaying tissue is a matter of almost every-day experience. The fatty matter produced in the sebaceous glands of the skin normally overflows on to the surface. It serves to render the skin supple and impervious to water. Germs get into one of the sebaceous glands of the face or of the eyelid. The contents of the gland begin to decompose. Leucocytes enter it for the purpose of removing the putrescent substance. They lose their vitality and turn into pus corpuscles. The pimple or the stye bursts, and pus and fatty matter are discharged together.
That the conversion of leucocytes into pus cells is due to want of oxygen has been shown by the following experiment: A minute piece of phosphorus is placed beneath the skin. Leucocytes gather round the spot with a view to removing the tissue which the phosphorus has destroyed. But phosphorus has so strong an affinity for oxygen that it exhausts the supply in the area of tissue which surrounds it. The leucocytes die before reaching the tissue immediately adjacent to the piece of phosphorus. Their dead bodies form round it a raised ring of pus cells. We can explain this readiness of leucocytes to sacrifice themselves in their efforts to reach foreign matter which needs to be removed, only by saying that the attraction of the food is greater than the repulsion of lymph destitute of oxygen. An amœba placed in comparable circumstances gives up the quest of food, however strongly chemiotaxic, and retreats towards water which contains oxygen sufficient to provide for its respiratory needs.
Blood.—A portion of the body fluid is enclosed within vessels and kept in circulation by the heart. The heart pumps blood into the aorta. This trunk gives off large arteries, which in turn divide until the finest capillary vessels are reached. The capillary tubes reunite to form veins, which, with the exception of those which collect food from the digestive organs, convey the blood right back to the heart. The veins which drain the stomach and intestines (the organs in which food is prepared for absorption) and the spleen (the organ in which worn-out red blood-corpuscles are in a sort digested) break up in the liver into a second set of small vessels. The pseudo-capillary vessels of the liver reunite to form the hepatic veins, which add the blood that has passed through that organ to the rest of the blood which is passing up the inferior vena cava to the heart. A second capillary circulation is found in the kidney also.
The heart is four-chambered ([Fig. 10]). Its left ventricle drives the blood round the systemic or greater circulation, the blood returning to the right auricle. The right ventricle drives the blood through the lesser or pulmonary circulation, from which it returns to the left auricle. The walls of all bloodvessels, except capillary tubes, are sufficiently thick to prevent the escape of any of the constituents of blood. To support the pressure of the blood which they contain, the arteries and the larger veins need walls of considerable thickness. The walls of the capillaries allow an interchange between blood and lymph in the manner already described ([cf. p. 39]).