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Water: Its Physiological Significance.' THE Glasgow Medical Journal. No. I. July, 1926. ORIGINAL ARTICLES. WATER: ITS PHYSIOLOGICAL SIGNIFICANCE.' By E. P. CATHCART, C.B.E., M.D., F.R.S., Professor of Chemical Physiology, University of Glasgow. "Ohne Wasser kein Leben," as Tangl puts it, is literally true, for it can be clearly demonstrated that water is absolutely essential if life, in all forms of living matter, is to continue. Withdrawal of water, indeed, brings about the death of man much more rapidly and more painfully than the absence of food. A mammal may survive thirty, forty, fifty days or even longer, losing almost all the fat in its body and 50 per cent of its protein, if deprived of food, but if water be withheld, death takes place when it has lost little more than 10 per cent of its water content. There is then some reason for the statement of Rubner that the regulation of the water content " of the tissues is, so to speak, anxiously supervised by Nature." Nor is it wonderful that this anxious supervision is necessary when it is remembered what a very large proportion of the * A lecture delivered before the Biological Section of the Royal Philosophical Society of Glasgow, on 12th March, 1926. No. x. Vol. cvi. 2 Prof. E. P. Cathcart? Water: body weight, even of the higher mammals, is due to water. Thus, it has been shown that the average water content of invertebrates ranges from 78 to 88 per cent, although in the case of some (e.g., Rhizostoma Cuvieri) it may be over 95 per cent. In the case of man the average ranges from 58 to 65 per cent, the variation being for the most part dependent on the fat content of the subject. Thus, with 19 per cent fat there was 60 per cent water, and when the fat content was 13 per cent the water rose to 66 per cent. This variation in water content is clearly seen in the data, calculated from the figures of Lawes and Gilbert by Wolff, for animals:? ox. SHEEP. PIG. Per cent. Per cent. Per cent. Fat, 191 301 18-7 35 6 23*3 42*2 Water calculated on Water, 51*5 45 5 57 3 43*4 551 4i .3 / \total live weights. ^ater calcu^ate(J on 70-9 71 *2 76-5 74*3 77 0 76 *8 / \ fat-free tissues. That the variation in water content is conditioned by the fat content is very clearly demonstrated by reference to the of the water uniformity content calculated on fat-free tissue. But, as workers have many shown, the age of the organism an plays important role. As Tangl points out, the alteration of the water content of organised tissue substance during would seem to ontogeny express an alteration which the animal organism also in the course undergoes of phylogeny. He thus correlates the alteration in water content due to age with the observations that the phylogenetically lower standing inver- tebrates, even those which do not live in water, are richer in water than the higher standing vertebrates. The following figures show that and clearly before, at, birth the tissues are much richer in water than they are later:? HENS. CATS. HUMAN. Embryo in egg. Water per cent. Water per cent. Water per cent. 7th . 92*8 day, . Newborn, 80-8 3rd foetal 14th . 94-0 . 87 *3 month, day, 9 days old, . 79*7 . 21st . New-born, 66-68 day, . 80'4 14 days old, 73*8 . Just before A(*ult, . 53.05 hatching, 78*7 83 day8 old, 66*7 is n not possible that the lowered metabolism of old be due in age may large part to a steadily increasing dessication ? Its Physiological Significance. 3 But, naturally, the water is not uniformly distributed throughout the body, nor is it by any means chiefly found in the blood and lymph. Thus, in the human body most elaborate estimations have been made, and the following table gives a very good idea of the average distribution:? PERCENTAGE DISTRIBUTION OF WATER (VOLKMANN). Muscle, . 50*8 Intestine, . 3*2 Fat tissue, . 2*3 Skeleton, . 12*5 Liver,. .2*8 Kidneys, . .0*6 Skin, . .6*6 Brain,. 2*7 Spleen, . 0*4 Blood, . 4*7 Lungs, . 2*4 Rest of body, . 11*0 The muscles, then, are seen to be the great storehouse of water in the body. In the analyses given above the actual amount of water obtained from the muscle amounted to over 20 kilo- grammes, or about one-third of the body weight. It is very manifest that the water of the circulating fluids forms but a small part of the water present in the body. The main part of the water is found, not, it is true, in the free state, but as imbibition water, in the protoplasm of the cell. This, of course, lends point to the statement of Starling that all the chemical changes which are considered under the term metabolism relate to changes in and between substances in solution. This contention may be rendered more general if it be remembered, as Haldane has pointed out, that all biological phenomena can be resolved into metabolic phenomena. One may therefore say that the whole of the processes which make up the life of the living organism, be it animal or vegetable, are ultimately referable to changes which take place in solution. What, then, are the nature and properties of this all-important constituent ? Both in its physical and its chemical properties water holds a unique position even in the physical world. If the physical characters of this familiar, unique, and wonderful fluid be tirst considered owing to its very high specific heat, or heat capacity, it not only plays an all-important rdle in regulating the temperature of the environment, but also within the organism this high specific heat allows of large changes in heat formation with but small alterations of body temperature. If, for instance, the body had a specific heat like the ordinary range of materials it would mean that temperature regulation would become exceedingly difficult, if not impossible. Further, this 4 Prof. E. P. Cathcart? Water: constancy of body temperature plays an important part in the control of the rate of the various chemical reactions which go on in the tissues where a relatively slight rise in temperature might bring about an untoward result, such as the coagulation of protein. The latent heat of evaporation, on the one hand, of water is the highest known, and it plays a very important rdle in the control of body temperature. No other fluid can fix so much heat during evaporation, hence it follows that the loss of a comparatively small amount of fluid in the form of sweat leads to a relatively large loss of heat. On the other hand, the high latent heat of melting is a protection against freezing of the tissues. The thermal conductivity, too, of water is a maximum the among ordinary fluids, and this, of course, plays its part in heat regulation, more particularly in the rapid of heat in the equalisation body, thus doing away with the risk of such as disturbance, overheating at any one point in active muscle. On the chemical side, too, water possesses many unique As a solvent no can properties. fluid compare with water, for, in addition to the fact that water can hold in solution a most wonderful variety of substances, these substances undergo but little chemical One has change. only to consider for a moment the of variety substances which are found in solution in the blood or the plasma, still more wonderful and heterogeneous collection of materials which are present in the urine. Although one infer that might water is chemically inert, it must be borne in mind that in a large number of cases there is good evidence of some kind of association between the molecules of the solvent and the solute. Thus, Bayliss cites the fact that one molecule of saccharose takes six up molecules of water, and that in again, glycerol solution exists, to the extent of 99*96 per cent, in a hydrated form. In watery the solution, again, extent to which ionisation takes place is very a condition high, which renders a " mobile possible most chemistry." And, of all finally, common liquids the surface tension of water is only exceeded that of Surface by mercury. tension plays a most important part in the movement of water in a capillary system like the soil, it also an probably plays equally valuable rdle in the changes which take place Its Physiological Significance. 5 4 within both the living plant and animal, as it is the effective agent in the phenomenon known as adsorption or surface condensation, a phenomenon which leads to the all-important uneven distribution of the solutes in a system. The modern interpretation of the rate of enzyme action, for instance, attributes it, in the main, to adsorption. Familiar as is the fluid-water, equally familiar is the chemical symbol which purports to represent its nature. Perhaps no symbols are so widely recognised as H20. To-day, however, it is generally accepted that only under the most limited of conditions can the fluid we know as water be represented by the simple H20 with a molecular weight of 18. It has been pointed out that if this were uniformly the case, the freezing point should be about ?150? and the boiling point ?100?. Actually the common everyday water must be a polymerised fluid in which a number of molecules are united together.
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