All animals of complex organization, from starfishes and sea-urchins to Man, are inhabited by motile cells. In addition to the bricks which enter into the construction of its fabric, each fixed in its place and definitely united to its neighbours, the animal contains leucocytes which wander through its tissue-spaces or float down the streams of lymph or blood. We are disposed to speak of these wanderers as inhabitants of the body, to distinguish them from the elements which enter into the construction of their habitation. It is difficult to avoid the temptation of describing the body as a habitation. Allegorical as Aristotle’s distinction between body and soul—between the habitation and that which inhabits—may seem, when contrasted with the exact language of modern science, it would save many a periphrasis if we might still use the monosyllable “soul.” The fixed tissues constitute a unity, bound together by nerves. The work done by glands and muscles is done in response to directions conveyed by nerves. It is impossible to say where the control of the nerves ceases—to point out any fixed tissue which is not co-ordinated with other tissues, nor susceptible to the influence of the environment as impressed upon the central nervous system, through the medium of sense-organs. The fixed tissues constitute a habitation for the “soul.” They share in a common life. The wandering cells are as independent of control as the parasites which occasionally find entrance into the body. Each must have a soul of its own in Aristotle’s sense. Like parasites, they carry on all the business of nutrition, respiration, cell division, without reference to the needs of the fixed tissues. They take what they require from the lymph as it leaves the intestines loaded with the products of digestion; they take it from the lymph in the tissue-spaces; they take it from the blood. When nutriment or oxygen runs short, they do not share the privations of the fixed tissues. Only indirectly is their well-being affected by that of the body as a whole; only accidentally is the death of the body the occasion of their death. The same might be said of such parasites as the “blood-worms” of Egypt, or the trypanosomes (the cause of “sleeping sickness”) of Equatorial Africa. Occasionally, in the rare disease lymphocythæmia leucocytes multiply exceedingly, not, apparently, in response to a call for their presence in large numbers, but in defiance of the needs of the economy, and with baneful results. To the indispensable services which wandering cells render, frequent reference will be made. In the present connection, and while we are searching for the principles of construction of the animal body, it would be desirable, if we could do so, to define the status of wandering cells. If they entered the body from without, they would be parasites of commensal type, intruders who share in the food and shelter of the body in return for service. But they do not enter from without. They are cells of the growing body which, detaching themselves from the cells which are forming tissues, assume a wandering life. They are not to be recognized in the embryo until development is considerably advanced. Their origin is far from clear, but histologists believe that, although they are not recognizable as wandering cells in the earliest stages of growth, they, or rather their parent cells, are set apart at a very early date. Probably they are not formed in the embryo proper, but in the “extra-embryonic area,” from which they emigrate into the embryo. In this sense they come in from outside. But, after all, the extra-embryonic area equally with the embryo is a product of the ovum. Looking at the body as a whole, we recognize a common life, a soul in Aristotle’s sense, which inhabits the framework of fixed tissues; and at the same time we see a multitude of independent cells, each an organism in itself, produced, like amœbæ, from similar independent cells by cell division, absorbing the body-fluids, consuming invading germs and fragments of decaying tissues, dying, disintegrating, in their turn absorbed. Wandering cells are autonomous in the largest sense.
All multicellular plants and animals are formed by division of a primitively single cell, the segments remaining in contact. As the scale of life is ascended, the cells which are massed together in the body, whether of a plant or of an animal—we are still unable to find any word other than body for the thing as a whole—tend more and more to differ in appearance. Some are large, others small. Some have cell-walls; others have none. Some remain “protoplasmic”; others are largely composed of “metaplasm.” Better terms are wanted to connote “most living substance” and “less living substance” respectively. It would be easy to coin suitable words, but, alas! the nomenclature of physiology is already hopelessly encumbered, and there is little prospect that a bad word will die when a good one is available in its stead. Differences in structure indicate differences in function. A division of labour has set in. The cell starts with capacities for every function. Its particular situation renders it desirable that it should cultivate one capacity at the expense of the rest. It specializes in a particular direction. If it happens to be placed in the centre of the body on the course of the bloodvessels which bring to the embryo food and oxygen from its mother, it develops a great capacity for taking up food. It accumulates in its substance a vast quantity of nutriment which it cannot consume, holds it, and passes it on into the blood-stream as it is required. Thus the liver is formed. In the embryo it attains to a great size, equal to about one-half the whole body-weight; but whether storing food be its chief function at this stage, or whether the other special functions for which it is responsible are equally important, remains a question for further research. In subsequent life its main work is to store food. After birth, when the child prepares its own food by processes of digestion in its stomach and intestines, the blood-supply of the liver is so modified that the blood from the digestive organs is passed through it. Now and for the rest of life the liver is the storehouse of food, the larder of the body. It is a reservoir from which supplies are distributed as required. A liver-cell retains many primitive characters. It is soft and destitute of envelope. But under the microscope it appears, unless it be taken from a starving animal, unlike any other cell ([Fig. 7]). It is always loaded with masses of glycogen. Sometimes it contains fat globules also. This is perhaps the simplest of all instances of specialization of function. An amœba can take up food. Presumably it always absorbs as much as it can get, the simple law of growth with cell division making it impossible for it ever to get too much. The cells which in the liver are so fortunate as to be placed on the route along which food is carried into the body retain the appetite of an amœba, but lose its capacity for growth and cell division. They return to the blood-stream, when it is deficient in food, the stores which they took up when food was in excess.
The specialization of a gland-cell is opposite in kind to that of a liver-cell. It takes up no more food than it requires, but it has developed a great capacity of producing from the food a substance which would no doubt be needed for its own purposes were it an isolated cell, but which the gland-cell places at the service of the body as a whole. An amœba can digest proteid substances. A cell of the pancreas produces the ferment necessary for the digestion of proteins, and secretes it into the alimentary canal.
To take another instance of specialization. An amœba responds to stimulation by changing its shape. It contracts in one direction, expands in another. A muscle-fibre has developed the capacity of contraction at the expense of all other functions. During the course of its growth it changes from a round cell into one that is elongated. The elongation is in the direction in which it acts with greatest efficiency. Its cell-substance is very highly specialized in order that it may have the maximum capacity of contraction in this direction.
Sensory cells develop to a maximum the capacity of responding to external force; nerve-cells, the capacity of conducting the impulses generated in sensory cells. The body is a republic in which every citizen develops to the highest degree the capacity of doing the thing which his situation makes it desirable for him to do.
The possibility of isolated cell life, and the necessity within certain limits of cell division, have led biologists to dwell too much upon the independence of the separate cells of which the body is composed. Protoplasm organizes itself into cells, but cells are not necessarily anatomically distinct. They may be the partially separate elements of a syncytium, or there may be but the faintest traces of cell separation. The objection to looking upon cells as isolated, self-complete units does not hold good to the same extent when they are viewed from a physiological standpoint. A cell is an administrative area. For purposes of nutrition, respiration, and cell division it is autonomous. It is responsible for its own local affairs. If a part is cut off from it, this part loses its vitality; this, at least, is the conclusion drawn from the atrophy of the axons of nerves when they are cut off from the cells of which they are outgrowths. Apparently we must understand by “the cell,” when speaking of the cutting off of a part, the portion of the cell which retains the nucleus; although we must be careful not to lay too much stress upon the nucleus as the centre of cell life. Red blood-corpuscles, as already pointed out, have no nuclei, and yet they live. Cell growth, estimated by mere increase in size, does not depend upon the nucleus. Many cells of the skin and its appendages increase considerably after the nucleus shows changes which clearly indicate that it is far advanced towards decay. But increase in protoplasm, cell growth in a legitimate sense, and especially cell division, are dependent upon the presence of an active nucleus. While, therefore, histologists no longer formulate the cell theory in the restricted terms in which it was enunciated some years ago, they still regard the cell as the unit of structure and unit of function. The body is built of cells, and whatever is done by the body as a whole is done by its individual cells.
CHAPTER IV
THE FLUIDS OF THE BODY
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.