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posed by a prism, two dark bands make their appearance in the green portion of the spectrum. Stokes repeated and verified the fact, and it soon became in his hands the startingpoint of a new train of research.
He treated a solution of blood-corpuscles with an alkaline reducing agent, and observed that its colour almost instantly changed from scarlet to purple-red, the hue of veinous blood. On examining the spectrum, he now found that the two dark lines had disappeared, and that a single line, intermediate in position between them, had become visible. On shaking the tube with air, the scarlet colour and the two lines at once returned, but, after a few minutes, again disappeared ; and this could be repeated many times. Hence it was evident that the scarlet arterial blood lost its oxygen to the reducing agent, and subsequently recovered it again, when shaken, from the air. The fact is so important that I prefer to give it in Stokes's own words. He says,
“The colouring matter of blood, like indigo, is capable of existing in two states of oxidation, distinguishable by a difference of colour, and a fundamental difference in the action the spectrum. It
may be made to pass from the more to the less oxidized state, by the action of suitable reducing agents, and recovers its oxygen by absorption from the
Hoppe-Seyler had shown that this colouring matter is different from the so-called hæmatin, which is obtainable by artificial means from the blood, and Dr. Sharpey therefore suggested that the true colouring matter should be named cruorine. The name is a good one, and does not, like “hæmato-globulin,” which is adopted by Hoppe-Seyler, involve any hypothesis. In the oxidized—the scarlet state-it is distinguished as scarlet cruorine, and in the reduced state as purple cruorine. It is hardly necessary to point out how intelligible an explanation these facts afford of the oxygencarrying power of the blood-corpuscles. In the lungs the purple cruorine of veinous blood takes up oxygen, and becomes scarlet cruorine; and in the whole of the general circulation, but more particularly in the capillaries, oxidation is effected by means of this oxygen, and the cruorine, to a great extent, passes back to the purple state. Hoppe-Seyler has since found that the blood of a rabbit which has been killed by drowning exhibits the spectrum of purple cruorine. In ordinary states, however, even veinous blood retains enough unreduced cruorine to give the two-line spectrum.
But Stokes has discovered another fact which is of extreme importance in regard to the question of animal oxidation.
Proceedings of the Royal Society," vol. xiii. 357.
found that a solution of the blood-corpuscles from arterial blood-a solution, that is, consisting mainly of scarlet cruorinewhen excluded from the air, slowly reduced itself, and showed, after a time, the purple colour and the one-line spectrum of purple cruorine. On opening the tube and shaking it with air the scarlet colour returned, and with it the two-line spectrum. Hence it is clear that scarlet cruorine is capable of oxidizing a portion either of its own substance or else of the serum, from which it is impossible wholly to free it in the experiment. Whichever it be, it certainly is a part of the blood itself which is oxidized by the cruorine; and this fact is, as we shall presently see, in perfect accord with the theory to which we are led by other considerations.
These curious optical experiments, apart from their physiological interest, have already yielded some practical results of considerable importance. Soon after the publication of Stokes's memoir, Mr. H. C. Sorby contrived an ingenious adaptation of the spectroscope to the microscope, and by its means succeeded not only in repeating all Stokes's experiments, but also in furnishing medical jurisprudence with a new and most valuable means of identifying blood-stains. The spectrum-microscope has since but somewhat improved in construction, and many readers of this journal have no doubt seen it, and the beautiful experiments which its inventor performs with it at some one of the recent ific soirées. It is described in detail in a paper by Mr. Sorby, read before the Royal Society, April 11, 1867. Å scrap of blood-stained fabric, 1-10th of an inch square, containing possibly not more than 1-1000th of a grain of colouring matter, may be experimented upon by its means, and the most certain evidence of the nature of the colour obtained. It has already been found useful in criminal trials.
Another interesting application of the spectroscopic examination of blood was made by Hoppe-Seyler. Claude Bernard discovered, some years ago, that the poisonous action of carbonic oxide gas was due to the circumstance that it had the power of displacing all the loosely-combined oxygen from the corpuscles, and of occupying its place in a somewhat more stable form of combination. All blood, veinons as well as arterial, after treatment with carbonic oxide, acquires a uniform red tint, which it retains with singular persistency, being, in fact, as Bernard expressed it, mineralized by the gas. HoppeSeyler submitted some of the blood so treated to optical examination, and found that it gave a spectrum very similar to, but not identical with, that of scarlet cruorine. But when excluded from the air, instead of reducing itself like scarlet cruorine does, it remained unchanged for an indefinite period of time. Hence the process indicated a delicate and certain test for use in cases of suspected poisoning by carbonic oxide. I myself, in ignorance of Hoppe-Seyler's experiments, mado the same observations. I have by me now a sealed tube, which has for more than a year contained a solution of blood through which carbonic oxide had been passed. The spectrum has not altered in the slightest degree.*
To return from this digression, it is clear that we are now acquainted with the mode in which oxidation is effected in the body, as far as the earlier stages go. Oxygen is absorbed in the lungs, combines with the cruorine, and is afterwards given out again. But at this point we are compelled to pause to consider two more complex and exceedingly important questions. These are, firstly, What is oxidized ? and, secondly, Where is the oxidation effected ? Liebig, as everybody knows, divided the substances oxidized in the body into two great classes, corresponding with the chief constituents of food. These were the non-nitrogenous, or “respiratory” elements, and the nitrogenous, albuminous, or “plastic" elements. The former embraced fat, sugar, starch, etc., and all its members were supposed by him to be oxidized in the blood, and to evolve no force but heat as the result of their combustion. The latter consisted of the organized tissues, and in particular the muscular tissue, the oxidation of which chiefly resulted in the production of mechanical work. It is an obvious corollary from this hypothesis, that the oxidation of a solid tissue must be effected in the tissue itself, outside the walls of the capillaries, and we are therefore compelled to believe in two distinct modes of oxidation. Substances in the blood are in direct contact with the corpuscles, and may therefore be supposed to unite directly with the oxygen of the scarlet cruorine; whereas, for the direct oxidation of a tissue, it is necessary to assume that some of the oxygen leaves the corpuscles, traverses the walls of the blood vessels, and arrives at the comparatively distant fibres in a state of solution, but in an uncombined condition. In its extreme form, Liebig's hypothesis has long been known to be untenable, for it cannot be doubted that nitrogenous substances, as well as non-nitrogenous ones, are oxidized in the blood, and contribute to the animal heat; and it has recently been demonstrated by the conjoined efforts of Traube, Heidenhain, and Donders, and still more distinctly by Fick and Wislicenus, Frankland and Parkes, that the oxidation of nitrogenous substances cannot account for nearly all the work done in the body. Traube, indeed, has started a rival hypothesis, which has been accepted by Fick and Wislicenus in their celebrated memoir ;* namely, that the oxidation of muscle contributes nothing whatever to muscular power, but that the whole of the latter is derived from the oxidation of non-nitrogenous bodies, such as fats and the so-called hydrates of carbon. But as they agree with Liebig in placing the seat of this oxidation in the tissue, there is no great difference, as far as the blood is concerned, between the two views.
* I believe reduced cruorine to be the most delicate, as it certainly is one of the simplest, qualitative tests for oxygen known. If a weak solution of blood is inverted in a test tube over mercury, it reduces itself in a day or two, and a small prism will then show the one-line spectrum. The minutest trace of oxygen will now restore the original spectrum ; a single drop of distilled water will often contain enough. I obtained incidentally in the above experiments a confirmation of the previously known fact, that carbonic oxide is disengaged during the absorption of oxygen by potassic pyrogallate. Air from which the oxygen had been removed by this re-agent, when added to reduced cruorine, caused the latter to give a two-line spectrum, which lasted for weeks.
But are there, indeed, two distinct kinds of oxidation going on in the body, one inside and one outside the walls of the blood-vessels ? Is it probable, or indeed possible, that sufficient oxygen can pass out through the thin walls of the capillaries to account for the enormous force exerted by the body in twenty-four hours ? Mayer thought not, and argued against the notion in his immortal treatise, “ Organic Motion in its connection with Change of Matter," published more than twenty years ago. I believe he was right in this, as in so many other things, and I have elsewhereț drawn attention to his arguments, and endeavoured to add others to them. The question is one of immense theoretical and practical importance, and I will therefore enter into it in some detail.
To begin with, it is necessary to bear in mind another wellknown and most important function of the blood. All the tissues of the body, the muscles among the number, are subject to a ceaseless process of disintegration and destruction. The elementary parts of which a tissue consists, have a definite term of life. They are born, grow, decay, and die, having previously developed new germinal matter from which their successors arise. There is no doubt about this, and it is equally certain that the nutrient matter, the pabulum, from which the new parts are formed and nourished, is derived from the blood, some portion of which must travel through the thin walls of the capillaries, and irrigate the tissue. Extreme uncertainty exists as to the mode in which this exudation takes place. At first sight it would appear to be simply a question of liquid diffusion; but, apart from the colloidal nature of the albuminous bodies of the blood, there are some striking points of difference between the composition of the blood and that of
“On the Origin of Muscular Power," “ Phil. Mag.," June, 1866 (Supplement).
"Die Organische Bewegung in ihrem Zusammenhange mit dem Stoffwechsel." Heilbronn, 1845.
I "Phil. Mag.," May, 1867.
the muscular juice, in respect even of some of the most diffusible substances. Thus common salt, an extremely diffusible compound, is found in large quantity in the blood, but is almost entirely absent in muscular juice, and the blood is invariably and necessarily alkaline; whereas the liquid of the tissue is acid, and may even contain, as Liebig has remarked, enough acid to neutralize the blood. Probably the pressure under which the blood flows, influences in some manner the exudation, but it would be vain to pretend that it explains it.*
The excess of the nutrient fluid, together with the products of the disintegration of the tissues, returns to the blood, a portion perhaps direct to the capillaries, but the great bulk, in all probability, through the lymphatics, which seem to act as overflow-pipes to the tissues. Mayer therefore suggested that the quantity of lymph might be taken as a measure of the quantity of fluid exuded in a given time. Bidder and Schmidt estimate the lymph returned to the blood in twenty-four hours at 22 lbs., but it is safer to assume it to be at least 30 lbs. It would hardly do, however, to take even this quantity as a representation of the average exudation through the capillary walls in twenty-four hours, and I have thought it right to treble it, so as to have a decided over-statement of its probable quantity. We thus get 90 lbs. a day, or about 40 litres. Now if oxygen leaves the blood and passes into the tissues, it is evident that it must pass in solution in this 40 litres of exudate. How much oxygen could possibly be dissolved by this 40 litres ? There is every reason to believe that the exudate does not differ materially from liquor-sanguinis in composition, and we have before seen that liquor-sanguinis is about equal to water in its power of dissolving oxygen. 40 litres of water would dissolve less than two grammes of oxygen; and this quantity of oxygen, whether it were employed in the oxidation of muscle, of fat, or of sugar, could not yield as much as 3000 metre-kilogrammest of force. But it may be urged that, though unlikely, it is still possible that the exuding fluid may be able to carry with it a larger proportion of oxygen than this. Be it so. Let us make the absurd assumption that every hundred volumes of exudate contains more oxygen than the arterial corpuscles themselves do, when saturated
If each hundred volumes of exudate contained forty volumes of oxygen, 40 litres would still only contain about 23 grammes, and this, in uniting with oxidizable
with the gas.
* Some of these arguments were suggested to me by Dr. Marcet, F.R.S., who has studied the bearings of dialysis on pathology with great care and success.
+ A metre-kilogramme is the force required to raise one kilogramme one metre. It is equal to about 74 foot-pounds, and is now almost universally employed as the measure of force.