Читать книгу The Body at Work: A Treatise on the Principles of Physiology - Alex Hill - Страница 8

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From one-fourth to one-third of the whole body is fluid. If the skin be regarded as a water-tight bag, three-fourths or rather less of its contents are solid, one-fourth liquid; and even its apparently solid contents, the tissues, contain much water. Water is an essential constituent of protoplasm. It is also present in cell-juice. The estimate given above does not include the fluid within the cells, but only the fluid with which the cells are bathed. In a general sense this extracellular fluid, excluding blood, is termed lymph. It occupies the spaces of a gauzy “connective tissue,” which connects, or separates—the terms are equally appropriate—muscles, nerves, glands, and other tissues of specialized function. Nowhere, except, in a fashion, in the spleen, does blood come in contact with a cell. The lymph which more or less surrounds them is the bath from which cells receive their food and oxygen, into which they excrete carbonic acid and tissue-waste. The network of lymph-spaces is traversed by capillary bloodvessels with walls composed of flattened connective-tissue cells. Such cells are usually spoken of as elements of an “endothelium.” As the epithelium covers the surface of the body, so endothelium lines its cavities. Endothelial cells are thin scales or tiles with sinuous borders dovetailed one into another. That the tiles which form the walls of capillary vessels are not cemented together in any proper sense is shown by the facility with which white blood-corpuscles, leucocytes, by their amœboid movements, push them asunder when making their way from the blood-stream into the tissue-spaces, or vice versa. They offer no more resistance to a leucocyte than a pair of curtains hanging in front of a door offers to a child. Yet so long as the endothelial cells are alive they keep their edges in such close apposition as to constitute a continuous membrane which shuts off blood from lymph. They are always close enough together to prevent red blood-corpuscles from escaping from the capillary vessels; but their resistance to the passage of the different constituents of plasma varies greatly. The membrane which they compose is more complete and less pervious, or less complete and more pervious, in accordance with the nature of the tissues which surround it, and their varying needs. The blood-passages of the liver may be described as filters. The escape of red blood-corpuscles into lymphatic vessels is prevented, but they offer practically no resistance to the plasma. Plasma—“lymph,” as it is termed as soon as it is outside bloodvessels—passes through the walls of the capillaries of the liver unchanged in constitution. Where they traverse glands (other than the liver), muscles, skin, and various other structures, the walls of capillary vessels, while offering practically no resistance to water and diffusible salts which can pass through membranes, prevent proteid substances from passing from blood to lymph, except in extremely small quantities. In this way an exquisite balance is automatically maintained. Water and salts pass out as they are needed. But they never pass out in excess, because the protein-containing blood-stream tends to keep them in, in virtue of the same attractive force which enables it to suck in the oxidized products thrown into the lymph by the tissues. Whatever a tissue needs it takes from the lymph. Suppose that bone is being formed. Large quantities of lime and phosphates are needed for the calcification of the cartilage in which it is modelled. The cartilage absorbs lime and phosphates from the lymph which bathes it. Lime salts and phosphates immediately begin to diffuse from blood into lymph. The hurrying blood-stream brings up further supplies from the walls of the intestine, products of digested milk and other foods. Lymph contains (although not in the same proportions) everything which blood contains. Many an analogy may be found in the world of economics, although no illustration would be sufficiently complete. From the lymph tissues take the fuel that they need, the oxygen with which to burn it, the foods for their own repair, the raw materials for their arts. Into it they throw their smoke, their drainage, the slag and refuse of their factories. The blood replaces the supplies as they disappear. It absorbs all waste. Lymph occupies streets, market-place, passages, corridors. The blood-stream is a closed system, rolling down the streets and through the market-place, on its never-ceasing circuit from port and mine to open air and open sea. From the alimentary canal it picks up food and fuel; the lungs give it oxygen, and disperse its carbonic acid; the kidneys purge it of non-gaseous waste.

Fig. 3.—A Ductule and Two Acini of a Mucous Gland of the Mouth, with a

Muscle-fibre cut Longitudinally; Capillary Bloodvessels and Connective Tissue.

Stellate connective-tissue cells form a labyrinth of intercommunicating lymph-spaces which separate the gland-cells and the muscle-fibre from the walls of the capillary bloodvessels. The capillaries contain circular red blood-corpuscles and nucleated leucocytes. Some of the leucocytes are squeezing their way either out of a capillary into a lymph-space or vice versa. A granular leucocyte is to be seen in a lymph-space at the bottom of the picture.

The facility with which the constituents of blood pass out to the lymph, and the constituents of lymph pass into the blood, depends upon the condition of the walls of the capillary vessels. Water and substances dissolved in water might pass through the wall of a capillary vessel in either of three ways—by filtration, by osmosis, or by secretion. A filter is a porous barrier, which allows water and all substances dissolved in water to traverse it. The solution passes through unchanged in composition. Only solid particles are kept back. The rapidity with which fluid passes through a filter varies as the difference between the pressure on the one side and the pressure on the other. A membrane does not allow of filtration. Water and things dissolved in water pass through it by osmosis. Some things it will not allow to pass; such, for example, as gum, mucin, white of egg. To others it offers resistance in varying degrees. Most of the things that can diffuse through a membrane are capable of crystallization; but the membrane exercises some control over the passage of even crystallizable substances when in solution. If a membranous tube containing water in which proteins, sugar, and various salts are dissolved is hung in a basin of pure water, the proteins remain in the tube; the sugar and the salts pass through its wall into the surrounding water. But they pass at different rates. Those of small molecular weight pass more quickly than those whose molecule is heavy. After a time a condition of equilibrium is established. No more salts pass out of the tube. If now the contents of the tube and the contents of the basin are analysed, it will be found that the tube contains all the proteins, some of the sugar, and some of each of the salts, although not in the proportions in which they were present at the commencement of the experiment. The water in the basin contains some sugar and some of each of the salts, but not in the same proportions in which they are found in the tube. As a matter of fact, the same number of molecules would be present, per unit volume, on each side of the membrane—in the tube and in the basin. In this respect the percentage composition of the two solutions would be the same. But some of the molecules being heavy, others light, the weight of salts which unit volume of the solution in the tube would contain would not be the same as the weight of salts in unit volume of the solution in the basin. A membrane exerts a discriminating action on the substances which pass through it. Secretion is osmosis in disguise. It may be even filtration in disguise. A gland-cell (like an amœba) takes things up and passes them out without regard to their osmotic equivalent. It seems to exercise a choice. It seems to act in disregard of the laws both of filtration and of osmosis. So, at least, it appears to us when we are looking at the result in ignorance of what has happened inside the living cell. The passage from blood to lymph and vice versa through the wall of a capillary vessel is in certain situations or at certain times a mere process of filtration; at others a process of restricted filtration. If the wall is behaving as a perfect membrane, it is a process of diffusion, or osmosis. It seems unnecessary to regard it, in any case, as a process of secretion. The more widely the capillaries are dilated, the less resistance do they offer to exudation. The narrower their calibre, the greater is the restraint which they place on the escape or entrance of fluid. When the skin of the palm of the hand is not sufficiently thick to protect the soft tissues beneath it from the injurious effects of the prolonged pressure of an oar or an axe, the capillary vessels of the under-skin dilate; more lymph transudes; the skin is raised up as a blister. The same thing happens when the capillaries are dilated and paralyzed by scalding water. The fluid of a blister has much the same constitution as blood-plasm, except that it contains less proteid substance. These results might be regarded as purely mechanical—the direct effects of pressure or heat upon the membranous capillary wall. But the “vital” element is more important. The capacity of endothelium to act as a barrier depends upon its nutritive condition—its vital integrity, as it might be termed; which no doubt in the last resort means its chemical relation to the fluids which bathe it. Now and again blebs, like blisters, are formed on the skin—the herpes which appears about the mouth; urticaria, which is more generally distributed; and various other cutaneous disorders. Frequently a connection can be traced between these eruptions and the consumption of a particular food. An attack of urticaria results not uncommonly from eating lobster, mussels, rook-pie, or some few other articles of diet. Various things—bad fish, for example—may produce the same effect; but shell-fish have an especially evil reputation. If extract of lobster or of mussels be injected into the blood of an animal, the amount of lymph which leaves the blood is markedly increased. The extract acts as a poison upon the endothelium of the capillary walls. It increases its permeability in all conditions in which lymph escapes in undue quantity from the blood-stream, or escapes more rapidly than it is absorbed; the nutritive condition of the endothelium is disturbed. Its unusual permeability is due in part, no doubt, to the dilatation of the capillary tube, the stretching of its membranous wall; but it is due also to the diminished vigour of the endothelial cells. They have lost to a certain extent their capacity for holding their edges in perfect apposition.

When the circulation is sluggish, owing to the inefficiency of the heart, the tissues become œdematous. In other words, lymph accumulates in the tissue-spaces. When the skin of a healthy person is pressed, it returns to its natural position as soon as the pressure is removed. If there is a tendency to dropsy—for ages the term “hydropsia” has been thus familiarly clipped—the finger leaves a pit behind it when pressed upon the skin. It is some little time before the lymph in the connective-tissue sponge readjusts the surface. Excessive escape of lymph from the blood, or its insufficient return into the blood, may also be the result of obstruction to the flow in the great veins. When the veins of the leg are varicose, the weight of the column of blood in the distended vessels impedes its circulation. After standing, the tissues about the ankle become œdematous. The œdema disappears on lying down. A hardening (cirrhosis) of the liver impedes the circulation of the blood which comes to it through the portal vein from the walls of the alimentary canal. The capillaries of the stomach and intestine are distended. Lymph accumulates in the abdominal cavity, producing ascites, another form of dropsy.

It is almost hopeless to attempt to disentangle the various factors which disturb the balance between blood and lymph—excessive outflow from blood, deficient inflow from lymph, stretching of the endothelium of the capillary tubes, imperfect nutrition and consequent imperfect apposition of the endothelial scales, increased permeability of the scales. The exudation which accompanies inflammation would seem to be due to the diminished vitality of the endothelium rather than to a mechanical factor, such as increased blood-pressure in the capillaries, and their consequent distention. Ascites is, apparently, a purely mechanical result of the resistance offered to the passage of blood through the liver; but pleurisy, the accumulation of lymph in the space between the lungs and the chest-wall, cannot be explained in the same way. There is no undue pressure on the vessels in which the blood circulates through the inflamed pleura (the investing membrane of the lungs and lining membrane of the chest), yet the walls of the capillaries fail to maintain a proper balance between blood and lymph.

Hitherto we have spoken of the lymphatic system as a labyrinth of communicating spaces containing stagnant fluid, which is kept in a fitting state by egress and ingress out of and into blood. Such a mental picture is substantially correct. But the system is complicated by the presence of lymphatic vessels. Cells of the connective-tissue sponge-work arrange themselves side by side. They flatten into endothelial scales. The borders of the scales close up. They form lymphatic channels, wider than blood-capillaries, but strictly comparable in every other respect. The lymph capillaries unite into larger vessels. The larger vessels are connected by cross-branches; they form plexuses. Their walls are strengthened with fibrous tissue. Like the veins, they are abundantly provided with valves, which check any tendency to a backward flow on the part of the fluid which they contain. Lymphatic plexuses surround and accompany the larger bloodvessels. They are disposed on the surface of muscles and glandular tissues. They are abundant beneath the skin. Nearly three centuries ago the lymphatic vessels of the mesentery, which collect products of digestion, especially fat, from the walls of the alimentary canal, were recognized owing to the milkiness of their contents after a meal. They were, on this account, termed “lacteals.” Other lymphatic vessels, owing to their transparent walls and colourless contents, are not easily seen; but they are readily injected with mercury or other fluids which render them conspicuous. In the upper part of the thigh, in the armpit, or in the neck, they are about large enough to admit a crow-quill. Those from the lower limbs, from the viscera, and from the walls of the abdomen converge to a receptacle which lies in front of the spinal column. The receptaculum chyli is continued upwards as the thoracic duct, which pours the lymph into the great veins of the left side of the neck and of the left arm just where they join together.

The thoracic duct provides for the overflow of lymph from the spaces of the body. There is no circulation of lymph. Lymph from the liver and from the intestines is constantly draining into the thoracic duct, and thus returning to the blood-stream by a short direct route, entering it without the necessity for reabsorption through the walls of capillary vessels. By no means all of this fluid has exuded from the blood-stream. Much of it is water which was poured into the stomach as gastric juice, and into the intestines as the secretions of the pancreas and other glands, or imbibed through the mouth and absorbed by the lymphatics of the alimentary canal. The remainder of the water taken up from the alimentary canal enters its bloodvessels. The diluted blood flows to the liver, loaded with digested products which the liver will store. As the blood parts with them the additional water which has served for their transport exudes from the capillaries of the liver into lymphatics, which empty it into the thoracic duct. Large quantities of water are used in washing out digested products. Secreted into the alimentary canal by the digestive glands, it passes out through its wall as the vehicle of digested products. Collected by lymphatic vessels, it is either carried directly into the thoracic duct, or passed from lymph into blood, carried by blood to the liver, again transferred from blood to lymph, and borne by the lymphatic vessels of the liver to the thoracic duct.

Water exuded from blood into lymph may be reabsorbed into the blood near the place where it was poured out, or it may reach the blood via the thoracic duct. It would seem that the former is the natural, the latter the emergency route; the former the course taken when an organ is tranquil, the latter a necessity when the organ is active. If the large lymphatic vessels of a limb are cut, no lymph escapes from them so long as the limb is at rest. When the muscles contract lymph begins to flow. If the limb is flexed and extended by hand, lymph flows. If the muscles are squeezed or massaged, lymph flows. As the flow is set up both by active contraction of the muscles and by passive movements in which the muscles do not take part, it clearly must be due to external pressure on the lymphatic vessels. As they are provided with valves, squeezing them converts them into pumps. The fluid which they contain is bound to go forwards. Additional fluid is squeezed into them from the tissue-spaces. To a large extent, therefore, the outflow of lymph from contracting muscles is to be explained as the result of the pressure which the swelling muscles exert upon the lymphatic vessels within their sheaths. But there is another factor which must not be overlooked, although it cannot readily be estimated. When a muscle is actively contracting its bloodvessels dilate. There is a greater exudation of lymph; and reabsorption by blood is not equal to the exudation. The surplus leaves the limb by the lymphatic vessels. A gland is never at rest. In the intervals between the ejection of its secretion its cells are preparing materials for the next outflow. Lymph is always flowing from a gland; its amount increases as the activity of the gland increases. More lymph leaves the blood when the gland is exceptionally active than when it is relatively quiet. Some of it is not reabsorbed into the blood. A certain proportion of the waste products of the active gland are hurried away by the overflow system in the direction of the thoracic duct.

Lymph is the reservoir of nutriment upon which every cell in the body draws. It is improbable that in health and under normal conditions the activity of any organ is ever restricted for want of sufficient food. As food is removed from lymph, it is instantly replaced by fresh food from the blood. There is some evidence—not very clear—that the removal of waste products offers greater difficulty than the renewal of supplies of food. When the activity of muscles has been excessively prolonged they ache. It has been supposed that their unwillingness to do more work is due, not to the exhaustion of the food which they use up when contracting, but to the inadequacy of the lymph and blood to carry off all refuse. This, at least, is the explanation of fatigue which is usually offered, although it is difficult to understand why the arrangements for removing waste products which have worked to perfection for eight hours should during the ninth hour become rapidly ineffective.

If a frog’s muscle, cut out of the body, has been made to contract until it refuses to work any longer, it again responds to stimulation after a solution of salt has been passed through its bloodvessels. The salt-solution brings no food; the only thing it can do is to wash away waste products. But this experiment upon a tired, isolated muscle does not necessarily throw light upon the nature of fatigue in muscles under normal conditions. The isolated muscle is using up, in contracting, food which it has stored. Cut off from the circulation, it has no means of getting rid of the lactic acid and other products into which food is changed. They may well have accumulated to a poisonous extent long before all the food has been used up. Hardly more cogent is the argument based upon the benefit which a tired man experiences from hot baths, massage, and the like. They take away the feeling of tiredness, but it does not follow that this result is due to the removal of waste products. Quickening the circulation of blood brings about renewal of the lymph. Renewal of lymph means fresh supplies of food as well as removal of waste products. Even human muscles are not perfect as machines. They will not work for an unlimited spell. There comes a time when they must have rest. Something goes wrong in the admirable adjustment which has hitherto provided exactly the right amount of food and exactly the necessary freedom from the products of action. A feeling of fatigue is the signal that the apparatus is not in a condition to work longer; but whether this feeling is due to a dislocation of the balance of supply and loss, or to some deterioration of the apparatus which calls for rest and renovation, it is at present impossible to say. It is not due to the exhaustion of muscle food. A more powerful stimulus, the urgency of fright or some other strong emotion, or an electric current applied directly to the muscle or its nerve, will still induce vigorous contraction. The muscles of a hare that has been coursed until it can run no farther still contain glycogen, muscle food.

Glycogen is stored in the liver. Fat, if it is assimilated in excess of the needs of the body, accumulates in the connective tissues. Proteins, if in excess, are either destroyed by oxidation, or partly destroyed and partly converted into fat. Increasing the amount and richness of the food does not, if nutrition is already at its best, improve the quality of the blood. The surplus of food is either stored or burnt. The composition of lymph is unaffected. Its quality is not improved by taking more food than enough. A perfect balance is maintained. Every cell is able, when conditions are normal, to obtain as much nutriment as it needs. It cannot get more. It cannot lay by food and shirk work. If it did it would grow. Reaching its optimum size, it would divide. Additional tissue would be formed. But when it does more work it needs more food; and it is a matter of common experience that the system is so adjusted that food is supplied to the tissues, not reluctantly, but with a slight tendency towards generosity. Working harder than usual, they find the lymph by which they are bathed somewhat richer in the materials that they need than the necessities of the case demand. They are able not merely to obtain all they want, but a little more. Activity favours growth.

Many attempts have been made to show that if a part of the body has more than its share of food it grows to an excessive size. John Hunter grafted a cock’s spur into its comb. It grew to monstrous dimensions. Such a result favours the view, but it is not quite conclusive. Undoubtedly the comb was richly supplied with blood, but it does not follow that the cells of the spur were able in their new situation to take advantage of this supply. Besides, the spur when projecting from the head was not subject to the accidents to which it was exposed whilst on the leg. Its size was not kept down by friction. Nor was it as hard and compact as it would have been in its normal situation. It is scarcely possible to devise any experiment that would be satisfactory now that the relations between blood and lymph and lymph and tissues are understood. In certain pathological conditions, however, hypertrophy is the result of the hyperæmia of chronic inflammation; and there is little doubt that, if we could arrange for a certain group of cells to receive lymph richer in food and freer from waste products than the perfect adjustment of supply to needs normally allows, the cells would grow.

Under perfectly healthy normal conditions growth can be induced only by use. Nature supplies the fuel which is used during activity, and a balance of food available for the construction of additional machinery. The muscle which is called upon to do work develops a greater capacity for work.

When nutrition is not at its best, the growth of muscle may be favoured by external pressure which squeezes lymph out of its tissue-spaces, and therefore leads to increased exudation from the blood. It is not improbable that in badly nourished tissues the circulation of blood is somewhat torpid and the lymph stagnant. A feeble circulation usually results in some œdema. The muscles, or rather the connective tissue which envelops and penetrates them, feels doughy, instead of being, as it should be, firm and elastic. Under these conditions massage is undoubtedly of service. Squeezing the muscles displaces lymph, and, if the pressure is properly directed, drives it along the lymphatic vessels. Fresh lymph exudes from the capillary bloodvessels, and the muscle-fibres, surrounded with a more abundant supply of nutriment, benefit, as, in a vigorous person, they benefit from use.

Lymph is an exudate from blood. Its composition therefore depends upon that of blood-plasma, but it tends to differ from it owing to the influence of two causes. In the first place, the walls of the capillary bloodvessels restrict exudation. Red blood-corpuscles cannot pass through them. Proteins which are non-diffusible are, according to the circumstances of the tissues, held back to a greater or to a less extent. The pseudo-capillaries of the liver let them pass, as has already been said. The capillaries of the limbs restrict their passage to such proportions as, it may be supposed, are absolutely necessary for the nutrition of the tissues. In the second place, tissues remove food from lymph and add to it waste products. Hence the lymph issuing from a limb, after full contact with the tissues, contains less of the former and more of the latter—less sugar, for example, and rather more oxidized nitrogenous substances, lecithin and other things termed collectively “extractives,” because they can be extracted from dried blood or lymph by ether. The reaction of lymph is alkaline. After a time it coagulates, but coagulation is slower, and the clot less firm than in the case of blood.

As the composition of lymph depends upon the source from which, and the conditions under which, it has been obtained, it is unnecessary to state the results of a chemical analysis. It suffices to say that lymph contains all the substances which are present in the plasma of blood, but not necessarily in the same total amount or in the same relative proportions. Speaking generally, leucocytes are present in about the same numbers as in blood—6,000 to 8,000 to the cubic centimetre; but leucocytes are everywhere present: in blood, in the lymph, in lymph-vessels, in the tissue-spaces. As they are not passively floating bodies like red blood-corpuscles, but active migratory organisms, they tend to accumulate in one situation and withdraw from another, in accordance with the opportunities which the different localities afford. They desert effused lymph, blisters, ascitic fluid, and the like. They are not found in the lymph in the pericardium. There are fewer in the lymph coming from the intestines after a meal than in the same lymph during the intervals between meals. Their departure from effused lymph might easily be explained. It is not so easy to account for their comparative absence from the lymph in the lacteals when it is heavily charged with fat and other products of digestion. Such leucocytes as are present at this time are loaded with fat granules which they have stolen from the chyle, as the lymph in the lacteals is usually termed. One would need to be very intimate with a leucocyte before one ventured to give reasons for all its movements. Lymph contains the same proteid substances as blood, and in the same relative proportions, but usually in smaller quantity.

Incidental reference has been made to the great lymph-spaces—peritoneal, pleural, and pericardial. The brain and spinal cord are separated from their outer membranes by a lymph-space. There are also spaces within the brain—the ventricles—and a central canal in the spinal cord. The aqueous and vitreous humours of the eye are also lymph-spaces, although the latter contains some remnants of tissue. The joint cavities are lymph-spaces. So also are the bursæ which surround tendons or separate them from bones. It is not, however, justifiable to include all these cavities in a single category, either from the point of view of their purpose, their mode of formation, or the nature of their contents. The peritoneal, pleural, and pericardial spaces are parts of the great primitive body-cavity, or cœlom. The two first are potential rather than actual. Normally they contain just sufficient fluid to moisten the apposed surfaces of the endothelium which lines their walls and covers the organs which they contain. There is no fluid in them which can be collected and labelled “peritoneal” or “pleural” fluid. The purpose of the spaces is to allow of movement without friction—in the one case of the intestines, in the other of the lungs. It is possible to take a spoonful or so of fluid out of the space which surrounds the heart. It has the usual composition of lymph. It contains proteins, but is not spontaneously coagulable. Leucocytes are absent, a fact which probably accounts for its not clotting. The fluid inside the cerebro-spinal system is extremely dilute. Its principal salt—its principal constituent, indeed—is sodic chloride. It contains hardly a trace of proteins, and these in a modified condition—proteoses. It also contains pyro-catechin, a benzoic alcohol. This substance has long been recognized as a constituent of cerebro-spinal fluid, owing to the fact that, like sugar, it reduces copper salts when heated with them in an alkaline solution. It appears to be one of the products of proteid decomposition. Although exuded as lymph from the bloodvessels of the chorioid plexuses, the composition of cerebro-spinal fluid has been profoundly changed by the activity—it might almost be called the digestive activity—of the epithelium which lines the cerebro-spinal canal. There is a theory that the ancestors of all vertebrate animals were organized on a very different plan from that of their distant descendants. Our cerebro-spinal canal was their stomach and intestine. It would appear that the lining epithelium of these organs, although disused for millions of years, cannot resist the temptation to digest the lymph which they contain! The fluid in joints contains mucin (the essential constituent of mucus), or a substance resembling mucin. In this case the joint-membrane has added something to lymph without removing or destroying any of its other constituents.

Other illustrations might be given showing how the plasma of blood is altered in composition while it is passing out of, or after it has passed out of, capillary bloodvessels. Perhaps it would be more logical to start on the outer side of the walls of the capillaries; since blood may, very properly, be regarded as a tissue, dependent, like all other tissues, upon diffusion from lymph for the nutrient materials that it needs. In the wall of the alimentary canal it receives supplies via the lymph. It drops them in the liver, its garde-manger, to pick them up again as they are wanted. The torrent of lymph which the thoracic duct discharges into the veins of the neck conveys the fat which could not traverse the walls of the capillary bloodvessels, and much of the reserve of food which the blood had deposited in the liver. Only about one-quarter of the fluid of the body (one-thirteenth of the body-weight) is included within the blood-system; but this enclosed fluid, owing to the fact that it is kept in circulation by the heart, replenishes and purifies the much larger quantity which does not circulate. The unenclosed lymph has in particular situations a chemical composition which varies widely from that of the blood. Imagine a marsh through which a river flows—the vast plains of water-plants on the Nile above Fashoda, for example. There is a constant interchange between the flowing water of the river and the stagnant water of the marsh. In any given part of the marsh the quality of the water will depend upon what it has been able to take from, and what it has given back to, the river; upon what the water-plants have taken from it, and what they have added to it. Boats which cannot penetrate the walls of reed keep to the open channel of the Nile. Fish swim, now in the river, now in the narrow passages and open pools of the marsh. So it is, in a way, with the fluid in the spaces and cavities of the lymphatic system and in the bloodvessels which traverse them, and with its migratory inhabitants. In our extravagant analogy read leucocytes for fish. Fish have two reasons for wandering from river to marsh. Amongst the water-weeds they hunt for food; they seek quiet places in which to breed. In this matter the analogy holds good. A leucocyte may be overtaken with cell division anywhere—in the blood-stream or in a lymph-vessel. But cell division very rarely occurs except in certain favoured spots. The breeding-places chosen by leucocytes are sheltered situations in connective tissue where the blood-supply is abundant, and the eligibility of such a spot is much increased by its being near to a field where their services are likely to be called for. The nests of connective tissue made by the leucocytes are of three kinds, termed respectively diffuse adenoid tissue, lymph-follicles, and lymphatic glands. The connective tissue beneath the mucous membrane of the whole of the respiratory tract—trachea, bronchi, and bronchioles—is diffuse adenoid tissue. It presents no special structure, but its spaces are packed with leucocytes in various stages of cell division, and young leucocytes, or lymphocytes, as they are usually named. Some of the lymphocytes make their way into the blood or into the lymph. Others, acquiring their full dimensions, scour the epithelium which lines the respiratory tract for germs and other foreign bodies which are drawn into the tract with inspired air. They may be seen pushing aside the cells of the lower strata of the epithelium, on their way to the surface, or returning to the subepithelial connective tissue with germs, or particles of soot, or débris of epithelial cells which they have taken into their substance (Fig. 4, B).

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).

Blood fresh from the lungs, whether still in the pulmonary veins or in the systemic arteries, is scarlet in colour. Venous blood is darker and purple-red, the depth of its tint varying with the extent to which it has parted with its oxygen. It looks less opaque than arterial blood. With this exception, the physical properties and chemical composition of blood are remarkably constant in all parts of the body. Arterial blood contains more oxygen, venous blood more carbonic acid. Other chemical differences can be recognized, but they are relatively very small. The constancy in the constitution of blood is its most notable character. Bleeding, unless excessive, does not greatly affect it. The number of corpuscles is of course diminished, but even these are replaced with great rapidity. The plasma, after bleeding, soon recovers its proteins and salts. A similar readjustment occurs if normal saline solution (water containing 0·9 per cent. sodic chloride), or even a strong solution of salt, is injected into the blood. Within certain limits it is very difficult to disturb the balance of its constituents. It gets rid of substances added in excess, or replaces substances removed, with remarkable facility. If sugar (glucose) be injected into a vein, it escapes through the capillary walls into the lymph. After a short interval the lymph contains more sugar than the blood. If an excess of protein, whether of a kind foreign to the blood or its own serum-albumin, be injected, it is removed by the kidneys. The blood has various sources from which it can draw out reserves of anything that is lacking, and various ways of getting rid of anything that is in excess. It draws upon the lymph in the tissue-spaces for water. It discharges salts into the lymph. It also takes salts from the lymph. It draws upon the liver for sugar, and probably for proteins also. In a starving animal the blood still contains sugar long after fresh supplies have ceased to reach it from the intestines. The lungs remove its carbonic acid. The kidneys free it from everything which cannot be otherwise removed. It is essential to the well-being of the organism as a whole that a uniform standard of composition should be maintained by the blood.