= a a Fee os =, Fe: ‘ i oo cn =< ? 7 ¥ . . , ‘ i . a “hs ' ' . ‘ - . j i ~~ - ~ ‘ a : ‘i : i. > - , 5 a o ' ’ , ae. , po sh ON THE ORIGIN OF THE FIBRIN FERMENT! THE ‘fibrin ferment’ which makes its appearance in shed blood is generally, I believe, supposed to arise from the cellular elements of blood, either from ordinary white corpuscles or from some special kind of corpuscles, the cells so concerned discharging the ferment into the blood or setting it free by their actual disintegration. Without wishing to deny that this may be one source of fibrin ferment, I am able, I think, to bring forward evidence that ferment may make its appear- ance in blood-plasma perfectly free from cellular, and indeed from all formed elements, in which case it must arise from some constituents of the plasma itself, and not from cells of any kind. It will be most convenient, perhaps, if I state the facts which I have to bring forward in connection with two series of experiments. | I. A measured quantity of blood was received directly from the carotid of a dog into a vessel containing an equal bulk of a 10 per cent. solution of common salt, great care being taken that the complete admixture of the blood and salt solution was effected as rapidly as possible. By the help of the centrifugal machine plasma was separated from this ‘ salted blood,’ and this plasma was again subjected to the action of the machine until all traces of formed elements were removed. As is well known, a portion of such a plasma diluted with five times its bulk of water coagulates rapidly, whereas the undiluted plasma remains hquid for an almost indefinite time. According to commonly received opinions, such a ‘salted plasma’ contains all the fibrin factors, including the ferment, 1 [From the Proc. Roy. Soc. 1884, p. 417. | — ae 120 ON THE ORIGIN OF THE FIBRIN FERMENT the latter having already passed out of the cells into the plasma ; and the reason given for the absence of coagulation in such a salted plasma and its occurrence upon dilution is, that the presence of the salts presents a hindrance to the action of the fibrin ferment, and that this obstructive influence of the salt is removed by the dilution of the mass. No one, however, as far as I know, has taken the trouble to ascertain whether fibrin ferment is present in such salted plasma. And, as a matter of fact, it is not; whereas it does make its appearance as soon as dilution with water has taken place, as the following experiment shows :— A portion of the undiluted salted plasma was treated with absolute alcohol in large excess, and the precipitate, after being allowed to remain under the alcohol for three or four weeks, was dried at a low temperature and extracted with water ; that is to say, the plasma was treated in the way usually adopted for obtaining a solution of ferment fairly free from proteids, &c. A portion of the diluted plasma, or rather of the serum resulting from the coagulation of the diluted plasma, was treated in an exactly similar manner. The aqueous extract of the diluted plasma brought about coagulation in specimens of magnesium sulphate plasma (such as is usually employed for testing the presence of fibrin ferment) in from ten to fifteen minutes. The aqueous extract of the undiluted plasma brought about no coagulation in specimens of the same magnesium sulphate plasma even after the lapse of elghteen hours. The conditions of each experiment were made as exactly alike as possible; and the conclusion seemed inevitable that ferment is present in the diluted and coagulated plasma, but absent from the undiluted plasma. This conclusion is, moreover, supported by the following experiments :—T'o a portion of the undiluted plasma above mentioned a small quantity of fibrin ferment was added, in the form of the dried precipitate thrown down by alcohol—z.e. a mixture of coagulated proteids and ferment. Coagulation took place. I have no record of the exact time elapsing between the \ ‘ \ ON THE ORIGIN OF THE FIBRIN FERMENT 121 addition of ferment and the appearance of the clot, but it was certainly not longer than three or four hours. II. Of the so-called peptone-plasma (7.e. plasma of the blood of a dog after the injection of peptone into the veins, such blood, as is well known, coagulating with great difficulty), freed from all cellular elements by the centrifugal machine, two portions were taken. To the one (A) a quantity of een was added, the lecithin being rubbed up with the plasma so as to be diffused through it; the other (B) was left untouched. Through both a stream of carbonic acid was passed, with the result that while A clotted in about ten minutes, B after the lapse of half an hour showed no disposition whatever to coagu- late. Both portions were then treated with excess of alcohol for the extraction of fibrin ferment in the usual way. The aqueous extract of A proved to be exceedingly rich in ferment, producing coagulation in magnesium sulphate plasma in about ten minutes. ‘The similarly prepared aqueous solution of B produced no coagulation at all. Now I have elsewhere,' in discussing the action of lecithin in promoting coagulation, shown that the coagulation which is brought about by the addition of lecithin is not due to the lecithin, or to any of its products of decomposition acting after the manner of a ferment, or to its carrying a fibrin ferment with it. In this case, therefore, as in the previous case of ‘salted’ plasma, the ferment appears to be absent before coagulation, but to be present after coagulation. I may here call attention to an observation made by Rauschenbach.” This observer found that the addition of yeast to plasma, prevented from coagulating by exposure to cold, brought about coagulation, and at the same time gave rise to the appearance of a large quantity of fibrin ferment. Never- theless, he completely failed to extract any fibrin ferment from the yeast itself. Now yeast is very rich in lecithin, and it seems highly probable that the coagulation caused by yeast was due 1 Journ. of Physiol. vol. iv. 1883, p. 226. [Collected Papers, p. 100. ] 2 ¢Blutplasma und Protoplasma,’ Inaug. Diss., Dorpat. 122 ON THE ORIGIN OF THE FIBRIN FERMENT to the lecithin contained in it, and hence the appearance of the fibrin ferment after the addition of yeast, and consequent coagulation, is quite parallel to the result of the experiment with lecithin and peptone-plasma recorded above. In both cases the ferment appears to have arisen out of the plasma itself. It is possible to obtain a coagulation in peptone-plasma without the addition of lecithin. or this purpose large dilution is necessary, followed by the passage of a stream of carbonic acid gas, But in such a case, however, coagulation is not only long in making its appearance, but the fibrin is formed, so to speak, in successive crops. Thus a feeble coagulation first appears, and if the clot so formed be removed, a succeeding coagulation is observed some time later, to be followed in turn by a third, and so on. When lecithin, on the other hand, is added, without previous dilution, the clotting is speedy and complete. If the serum thus resulting from the coagulation of peptone- plasma brought about by large dilution and treatment with carbonic acid be examined for fibrin ferment in the usual way, it will be found to contain ferment, though much less than could be obtained from a corresponding quantity of the same plasma coagulated rapidly by the addition of lecithin. The relative amount of ferment appearing under different circum- stances is illustrated by the following experiment :— Of three equal portions of the same peptone-plasma, one portion was simply treated with a stream of carbonic acid gas, without any dilution, and did not coagulate; a second was treated with a stream of the same gas after large dilution, and coagulated slowly; to a third lecithin was added, and a stream of carbonic acid passed through it, with the result of producing a rapid and complete coagulation. All three portions were treated in the same way for the extraction of the fibrin ferment, and the activity of the three aqueous extracts then prepared was tested under exactly the same conditions, with the help of magnesium sulphate plasma. The first produced no coagulation after the lapse of twenty hours. a 7 rd 4 J ~*~ ON THE ORIGIN OF THE FIBRIN FERMENT E28 The second produced coagulation in four hours. The third produced coagulation in five minutes. The amount of ferment seems to be in proportion to the energy of coagulation ; and the presence of ferment after simple dilution and the action of carbonic acid gas shows that the ferment appearing after coagulation by the help of lecithin does not come from the lecithin itself. Thus there is a remarkable coincidence between the occur- rence of coagulation itself and the appearance of the fibrin ferment, and that in plasma freed most carefully from all cellular elements. I believe, therefore, that I am justified in concluding that though fibrin ferment does not pre-exist in normal plasma, it may make its appearance in that plasma in the absence of all cellular elements, and must therefore come from some constituent or constituents of the plasma itself. I am still engaged in investigations directed to find out what that constituent is, or what those constituents are. 124 ON A NEW CONSTITUENT OF THE BLOOD ON A NEW CONSTITUENT OF THE BLOOD AND ITS PHYSIOLOGICAL IMPORT’ In a paper on the Origin of the Fibrin Ferment, published in ‘Proc. Roy. Soc.,’ vol. 36, 1884, I showed that there exists, dis- solved in the plasma, a body which can give rise to fibrin ferment. I have proceeded with my investigations, and have succeeded in making some additions to our knowledge of this subject, which I here describe. As my researches are not complete, | confine myself to as brief an account as possible. The subject is best studied in the blood of peptonised dogs ; but, as I showed in the above quoted paper, similar results are obtained from normal salt-plasma, so that the results are not peculiar to peptone-blood. The body the presence of which gives rise to fibrin ferment can be isolated from peptone-plasma in the following very simple manner :—The plasma, having been completely freed from all corpuscular elements by means of the centrifuge, is cooled down to about 0°. The plasma, which was _ previously perfectly clear, becomes rapidly turbid, and after standing for some time in the cool, a very decided flocculent precipitate forms. I have already described this observation in a short note, ‘ Ueber einen neuen Stoff des Blut-Plasmas,’ in Du Bois-Reymond’s ‘ Archiv fiir Physiologie,’ but it is necessary for me to allude to it here. Now it is this body which gives rise to the fibrin ferment. So long as the former is present in considerable quantity, the latter clots readily on passing through it a stream of carbonic acid, or on dilution, and at the same time a very considerable quantity of fibrin ferment makes its appearance. 1 [From the Proc. Roy. Soc. 1885, p.70. (An account of this new constituent of the blood appeared also in Du Bois’ Archiv, 1884.—Ed.). ] 7 \ ~ AND ITS PHYSIOLOGICAL IMPORT 125 By prolonged cooling the greater part of this substance can be removed, and with its gradual removal the plasma clots less and less readily with CO,, and less and less ferment is formed, till finally it becomes practically incoagulable—z.e. forms only a faint trace of fibrin after several days. If some of the substance be again added to the plasma, it regains its power of clotting with CO,. (The substance must be added before it has stood very long : see under.) It must be understood that the plasma, previous to the pas- sage of the CO,, is quite free from fibrin ferment, so that there can be no question of the ferment being mechanically removed by the precipitate. | Moreover, that it is really the body removable by cold which gives rise to the fibrin ferment, and not any second body which is mechanically carried down with the former, is shown by the fact that the diffusion of a large quantity of inert finely divided precipitate through the plasma, and its subsequent removal by the centrifuge, does not in any way do away with the power of the plasma to clot. It is, therefore, justifiable to assumethat when peptone-plasma clots readily and completely with CO,, it must contain this new body in some quantity, and that when it will not clot, or only very imperfectly, after repeated treatment with CO, or dilution, this new body must be present in very small quantity. Now | have found that the behaviour of peptone-plasma with CO, varies very considerably with the diet on which the animal is fed, and whether the animal is fasting or has been recently fed. In some cases it clots readily, in others practically not at all. Out of eight dogs fed on very lean meat only one gave a plasma which clotted at all fully, and in this case the clotting went on for two days. rom all the others the plasma, in spite of repeated treatment with CO,, only gave rise, after two or three days, to a scarcely perceptible fibrin membrane. The animals were killed about eighteen hours after the last meal. Of six dogs fed on fat and meat for several days, all gave a 126 ON A NEW CONSTITUENT OF THE BLOOD plasma which clotted rapidly and fulfy in from twenty minutes to one hour after the CO, treatment. The animals were killed about eighteen hours after being fed. Of two dogs fed on bread and meat, both gave a readily coagulable plasma. One day’s feeding on fat does not produce any effect; that is, the blood of a dog thus fed behaves like that of a dog fed on a lean meat. A dog fed for some days on fat and meat was for five days previous to being killed put on fat alone; as a consequence it practically starved, as it ate scarcely anything. The blood from this dog clotted very incompletely. Simple starvation for three days did away with the influence of fat in another case. These results only hold good for dogs in health. Ina dog with a suppurating wound, kindly placed at my disposal by Mr. Horsley, the plasma, in spite of a lean meat diet, clotted with very great rapidity, and contained an enormous quantity of the new body. All the other dogs were healthy, but were badly nourished when they came into my hands. It is necessary for these experiments that the peptonisation should be complete. For the better understanding of these results I must return to a further consideration of this new constituent of the plasma. The turbidity which appears on first. cooling the plasma, if examined microscopically, is found to consist of a great number of minute pale transparent bodies of a rounded shape, much resembling small organised bodies; such, for instance, as the stroma of the red corpuscles, except that they are of very various size, but generally much smaller than red corpuscles. They have a great tendency to run together into granular masses. At first the precipitate is soluble on re-warming the plasma slightly, but it soon undergoes change, and loses the power of redissolving by heat. If the substance be collected by means of the centrifuge, it forms a disc or thin membrane at the bottom of the tube, much reminding one of fibrin, but closer examination we X i a \ ig | | 2 AND ITS PHYSIOLOGICAL IMPORT 1b4"( shows that it presents marked differences from the latter, and that, in truth, it much more closely resembles the peculiar viscid body obtained by destroying leucocytes with dilute alkalies, &c. On longer standing, however, it becomes in most cases still further changed, and is then undistinguishable from ordinary fibrin, swelling in dilute HCl like the latter. For further details as to the properties of this substance I refer to my paper quoted above. We have already seen that this substance gives rise to fibrin ferment, but it does more than this in inducing coagulation. Peptone-plasma is not coagulable with fibrin ferment. If we take some plasma rich in this new substance, and by means of CO, induce coagulation, we obtain, on removing the clot, a serum which has the power of inducing exceedingly rapid coagulation in a new portion of plasma, and this when the serum has regained its alkalinity. This serum contains ferment ; but, inasmuch as ferment is not sufficient to induce coagulation, it must also contain some other substance. Now leucocytes have exactly the same power. ‘They give rise to ferment, but they also give rise to the other substance necessary for coagulation. We see, therefore, that we have dissolved in the plasma a body exerting the same influence on the induction of coagulation as the leucocytes. I think this is the strongest chemical proof that can be brought that the leucocytes break down to make, at any rate, a part of the proteid constituents of the plasma, and have shown above the influence which diet, &c., has on the extent of this process, a fact of obvious interest for the question of assimilation. There is, however, another important. conclusion to be drawn from these observations, viz., that one must admit, in addition to the ordinary fermentative fibrin formation, that fibrin may be deposited from blood by simple physical means, without any fer- ment process ; for this new substance becomes, as I have stated, true fibrin, and yet the plasma does not contain ferment. Pos- sibly this mode of fibrin formation is of importance in the formation of a thrombus. es ON A NEW CONSTITUENT OF THE BLOOD 128 re 4 The peculiar microscopical characters should also be noted, as possibly affording an explanation of the observations made by Osler, Bizzozero, Hayem, and others. I refer, of course, to the granules, Blutplittchen, heematoblasts, described by these authors. A oe As I am actively engaged on this subject, and as I hope before long to produce a complete account of my researches on » * i i “ 7 ; _ - ‘ Ray = . . bent be \ ri fi * 2 Pic. wry CLASS IV THE CROONIAN LECTURE ON THE COAGULATION OF THE BLOOD ; > 7 Bd vo or tia. b= : a F ; | E rf 141 THE CROONIAN LECTURE! ON THE COAGULATION OF THE BLOOD I. The Relation of the Formed Klements of the Blood to Coaqulation Ir is generally accepted, at the present time, that the inter- vention of formed elements is necessary for coagulation, that the blood-plasma is lacking in certain. of the constituents necessary for coagulation, and that these factors are supplied either by the white blood-corpuscles or by certain special corpuscles, to which the name of ‘ Blutplittchen’ is generally given. My obser- vations have led me to an opposite conclusion. I find that the plasma contains, in solution, everything necessary for coagula- tion. And although [ will not go so far as to affirm that the formed elements cannot exert any influence, yet, as I hope to show, it is very doubtful whether they play any part at all, and it is certain that they are not necessary. The proof of this is as follows. By injecting peptone into the blood of a dog, the blood drawn off directly after the injection is prevented from clotting. For the present we may leave out of the question how it is that peptone produces this effect. The result is always constant, and we can obtain from the blood drawn off, by subjecting it to the action of a centrifugal machine, a perfectly clear plasma free from all formed elements. This plasma, if left to itself, will re- main fluid for a very long period—in fact, until putrefactive changes set in—but it will clot on subjection to very simple in- fluences, influences which cannot be looked upon ag being in themselves fibrin factors, but which must be regarded as simply neutralising the restraining power exerted by the peptone injec- tion on coagulation. ‘Thus the plasma will clot when a stream 1 Delivered April 8, 1886. 142 THE CROONIAN LECTURE of carbonic acid gas is passed through it, or when diluted with water, or more slowly when diluted with 4 per cent. solution of common salt; further, it will clot imperfectly if filtered through a clay cell. Peptone-plasma! is entirely free from fibrin ferment, but in the process of coagulation fibrin ferment makes its appear- ance. By receiving blood direct from the blood-vessels into solutions of certain neutral salts the blood can also be prevented from clotting; and, in the same way as in the case of peptone- plasma, a plasma can be obtained entirely free from corpuscles. The plasma will be found to have different characters according to the solution which has been employed to prevent coagulation. Thus, if we receive into a 10 per cent. solution of common salt an equal volume of blood, we shall obtain a plasma’ which clots readily on dilution with four or five times its bulk of water. It will be convenient for me to allude to this plasma as ‘ sodium chloride plasma.’ This plasma, before dilution, is quite free from fibrin ferment; after clotting has taken place ferment is found to be present. If, instead of using a 10 per cent. solution of sodium chloride, we use a solution of sulphate of magnesia, we shall get a plasma which does not clot or form ferment on dilu- tion. In order to make this plasma clot, ferment must be added. This variety will be referred to as‘ magnesium sulphate plasma.’ It is obtained most conveniently, in the case of the dog, by taking 4 vol. of saturated solution of sulphate of magnesia to 1 vol. of blood. Another plasma can be got from the blood of the horse by cooling it to 0° and allowing the corpuscles to settle; by filtering it, still at a low temperature, through several folds of thick filter-paper the plasma becomes almost entirely free from corpuscular elements, and has but very little power of clotting by itself. Apparently then, these two last kinds of plasma afford no support to the statement made concerning the spontaneous coagulability of the plasma. But the result is explained by the following fact :— The body which has the power of initiating coagulation is readily separable from the plasma by cooling or by a certain strength of magnesium sulphate. This body is a new substance 1 Peptone-plasma—the plasma from peptonised blood. ON THE COAGULATION OF THE BLOOD 143 I have discovered ; it forms such an important element in the question of coagulation that without a knowledge of its existence the most experienced observer could hardly fail to fall into error. It has been mentioned that peptone-plasma clots when a stream of carbonic acid gas is passed through it, and that at the same time fibrin ferment is formed. If the plasma be placed in ice it rapidly becomes turbid, and in course of time a considerable precipitate forms. This is the new substance. If the cooling is prolonged until all this substance disappears, the plasma will be found to have lost its power of © clotting with CO, and of giving rise to fibrin ferment. If only part of the new substance is removed, the plasma coagulates more slowly and less ferment is formed; and by again adding some of this body to the plasma which has been deprived of it, the power of spontaneous coagulability is restored to the plasma. If peptone-plasma is mixed with an equal volume of 10 per cent. NaCl solution, it retains its power of clotting on dilution, and cooling does not give rise to a precipitate. In fact, peptone- plasma, treated in this way, behaves just as normal NaCl plasma does; it retains its power of clotting on dilution for an almost unlimited period. If, however, about + of its bulk of a saturated solution of magnesium sulphate is put to peptone- plasma, it is at first capable of spontaneous coagulation; but on standing—for a short time at ordinary temperature, and more quickly at a low temperature—there arises in the plasma a pre- cipitate, upon removal of which the plasma will not clot on dilution ; it completely resembles the normal MgSO, plasma. It may now be understood how it is that this form of plasma i.e. normal MeSO,—presents an exception to the general state- ment brought forward. The ‘ cooled’ plasma must be considered in somewhat greater detail, for the doctrine of the participation of the white cor- puscles in coagulation has been founded in great measure upon experiments made with this variety of plasma. The researches of Alexander Schmidt are well known, but it may be advisable to quote some of the more essential observations bearing on this subject. ‘The blood of the horse is used, and is prevented from 144 THE CROONIAN LECTURE clotting by collection in vessels surrounded with ice. The blood is thus maintained at a low temperature until the cor- puscles sink, the red falling much more readily than the white. After a time, in spite of the cold, coagulation sets in, beginning first and most extensively in the lower layers of the colourless. plasma, above the sediment of red corpuscles, where there are most white corpuscles; it 1s scanty and later in the upper layers of the plasma, where there are but few corpuscles. Now it must be remembered that this new substance separates from the plasma when cooled. Schmidt did not know of the existence of this substance, nor could he, from his method of experimenta- tion, obtain any definite evidence of such a substance. * But it will be obvious that the greater coagulability of the lower layers of the plasma may be due, not to the greater number of white corpuscles present, but to this new substance being precipitated out by the cold and sinking to the lower layers of the plasma. Indeed, Schmidt himself mentions that in these lower layers of plasma not only are great numbers of white corpuscles to be. seen, but also a large quantity of granular matter. He discusses’ this granular matter, and comes to the conclusion that it is the débris of white corpuscles which have broken up in spite of the cold. But he has no conclusive evidence to offer in support of this view. I think that my experiments leave no doubt as to what this granular precipitate is-—.e. this new body separable by cold from the plasma; and inasmuch as we have seen, from the account of my observations above given, that this substance is of great importance in initiating coagulation, these experi- ments of Schmidt’s lose their force as conclusive proof of the - part taken by the white blood-corpuscles in coagulation. The same objection can be urged against the experiments of Schmidt. on the cooled plasma filtered through thick filter-paper. This plasma clots very slowly and forms but little ferment. But these results cannot be attributed to the absence of corpuscles, since, by the process of cooling, the new body has been removed as well as the corpuscles. So far, then, the participation of the white corpuscles is an open question, but with lymph-corpuscles the case is different. : c 5 ON THE COAGULATION OF THE BLOOD 145 In a paper read before the Royal Society in 1881,! I showed that isolated lymph-corpuscles had the power of inducing co- agulation in a marked degree. This was the first definite proof that any form of corpuscle could contribute to the process of coagulation. My experiments were made with peptone-plasma, and as they have been repeated with the same result by other observers on different kinds of plasma, there is no doubt as to their correctness. If lymph-corpuscles are added to peptone- et plasma, or to cooled plasma, clotting rapidly takes place. This might be regarded as strong evidence in favour of the white corpuscles of the blood also being agents in the process of coagu- lation, and at the time these experiments were made I did so regard it. But it is only valid if there is adequate proof of the identity of white blood-corpuscles and lymph-corpuscles. This supposition is generally made; but some observations have led me to doubt whether it is a true one, and have also thrown _ grave doubts on the capability of the white blood-corpuscles to initiate coagulation. ‘The experiments are as follows :—A dog having been injected with peptone, a small quantity of its blood was drawn off, which remained fluid, but clotted immediately on the addition of a small quantity of lymph-corpuscles. certainly not under 50 per cent.; and yet all these substances. form clear solutions on treatment with aqueous dilute alkali. Lecithin itself is not in the least dissolved by weak alkali. Further, it is not at all clear that lecithin is easily ‘carried down’ from its solutions mechanically except when it occurs associated with proteid, and proteid is the ‘ carrying-down agent, for there is not the slightest proportionality between the bulk of the precipitate and the amount of lecithin separated, but quite the contrary. The digestion precipitate from tissue- fibrinogen, though very slight in bulk, is enormously rich in lecithin. The same bulk precipitate from the original fibrinogen solution contains certainly one hundred times less. It is clear, therefore, that, unless we suppose the lecithin to be chemically united with proteid, there are great difficulties and very much vague assumption. A-fibrinogen, the pseudo- crystalline substance, contains lecithin, certainly not under 5 per cent. (vide p. 283). The same general reasoning which renders it probable that A-fibrinogen is a chemical individual would be strongly in favour of the lecithin of tissue-fibrinogen being in chemical union. VIII. On the Processes termed ‘ Coaqulation ’ 1. Coagulation studied on extravascular plasmata.—In deal- ing with this, nothing but perfectly pure, absolutely haemoglobin- free plasmata are used. ‘The most scrupulous precautions are adopted as to complete removal of form-elements. In dealing with salt-plasma the greatest care must be taken to ensure that the admixture should take place with extreme rapidity—an absolute necessity, the non-recognition of which completely frustrates all attempts at exact work. Dog’s blood chiefly used. For salt blood large animals with rapid flow from carotid (dogs) ; horses also from carotid. I shall confine myself chiefly to two aspects of the chemical side of coagulation processes—t.e. 1, fibrin formation; 2, fibrin- ferment formation. These two aspects by no means exhaust the question. There 286 SECOND REPORT TO THE GROCERS’ COMPANY is a third subject, in particular, to which I have already alluded— viz. the formation of new altered fibrinogens simultaneously with the fibrin. I shall, however, but briefly allude to this division, as my knowledge of it from a chemical standpoint is not yet sufficient. 2. Coagulation of A-fibrinogen in peptone-plasma.—As it separates from the plasma, A-fibrinogen is certainly not fibrin. Its solubilities are quite different. Nevertheless it speedily undergoes changes in the direction of conversion into fibrin, Simple cooling, therefore, of peptone-plasma suffices to produce a certain amount of jibrinous coagulation. As I have already stated, A-fibrinogen on simple separation undergoes ‘alteration,’ which I regard as analogous to the ‘ altera- tion’ of all the other fibrinogens (which ‘ alterations’ are shown, as we shall shortly see, in a very high degree by the fibrinogen of plasma). But the case of A-fibrinogen is special; its separa- tion takes place in a fluid which is not indifferent, for it can be shown distinctly that A-fibrinogen possesses in the most marked degree the power of influencing the coagulation of the plasma in toto. It is impossible to collect A-fibrinogen except in an altered condition. Its appearance and physical characters are exactly like fibrin, its solubilities are in the main similar, its distin- suishing features are its solubility in dilute HCl and its becoming opaque with acetic acid, together with its behaviour on digestion. Now one obtains every gradation between typical A-fibrin- ogen and typical ‘ fibrin,’ that is to say, the degree of ‘ alteration ’ shown by A-fibrinogen varies in different cases. In all cases it separates as a substance quite different from fibrin; the altera- tion may never go further than what I have described previously (vide section, ‘ A-fibrinogen,’ p. 274), but there is every gradation between this alteration and the formation of true fibrin. An important property of A-fibrinogen is its power of in- ducing coagulation in peptone-plasma when a current of CO, is passed through it. As I have pointed out in Section I., a com- mencing precipitation is helpful to this interaction. Now ON THE PROCESSES TERMED ‘COAGULATION’ 287 A-fibrinogen can be collected and re-added to plasma, and again confer on the latter the power of coagulation with CO,. But this is not invariably the case; if the change of the fibrin- ogen have gone too far in a ‘fibrinous’ direction, the addition is then without influence. A-fibrinogen, then, separates out and remains for a certain time a substance quite distinct from fibrin. But it frequently passes, without any re-solution or any further treatment, into fibrin. I do not in the least mean to assert that in these changes A-fibrinogen is converted bodily into fibrin. Chemical interaction probably takes place between the substance and the body of the plasma. But if two portions of the same plasma are taken and cooled, and one portion examined quickly and the other portion after an interval, the precipitate is, in the former case, A-fibrinogen ; in the latter case, either fibrin, un- distinguishable from the ordinary fibrin obtained from the co- agulation of plasma, or something intermediate between the two. There are two important factors which favour the more or less fibrinous modification of A-fibrinogen. These two factors are under certain conditions helpful to one another. Factor I.—I have alluded to what I call strong and weak peptone-plasma. Strong peptone-plasma is much less readily coagulable with CO, than is weak. From previous statements it will be remembered that the coagulation of peptone-plasma with CO, is apparently due to the interaction of A-fibrinogen with the rest of the plasma. Now the fibrinous alteration of A-fibrinogen on simple cooling invariably takes place much more rapidly, and is more complete in weak than in strong peptone-plasma. The position is, therefore, as follows (vide Section I. p. 244) :— 1. With CO, genuine undoubted coagulation of plasma. 2. Partial separation of A-fibrinogen necessary condition of CO, action. 3. Interaction with CO, much more rapid in ‘ weak’ peptone- plasma. 4, Simple separation in weak peptone-plasma always leads to greater alteration of A-fibrinogen in the direction of fibrin. 288 SECOND REPORT TO THE GROCERS’ COMPANY I consider, on these grounds, that the fibrinous alteration of A-fibrinogen on cooling is due partly to an interaction between the A-fibrinogen and the rest of the plasma. : Factor II—The second factor is very remarkable. It depends on the greater or less quantity of separated A-fibrin- ogen distributed in a given quantity of plasma. It is ‘ Massen- wirkung.’ If you take two equal quantities of peptone-plasma mode- rately peptonised, place one (1) in a flat large dish and allow to cool slowly, the other (2) in a test-tube and allow to stand upright (both being otherwise under the same conditions), a widely different distribution of A-fibrinogen takes place in the two cases. In the one case (1) it sinks over a wide area, and is in intimate contact with a large part of plasma; in the other (2) it sinks to the bottom of the tube, and is in intimate connec- tion with a small part of plasma. Supposing we assume, as I have pointed out is probable, that the precipitated A-fibrinogen acts as a new fibrinogen added to the plasma, we shall in the case (1) have a greater quantity of A-fibrinogen acting on a given quantity of plasma. than in case (2). Now if reference be made to statements in the section on Quantitative Determinations (p. 293), dealing with the influence. ~ of tissue-fibrinogen on plasma, it will be seen that a certain quantity of tissue-fibrinogen must be added to produce an obvious result. A weighable quantity can be added without. producing any coagulation. And in the case of A-fibrinogen, when it is allowed to collect and interact with only a limited quantity of plasma, as in the test-tube, the ‘change towards fibrin’ is infinitely greater and more rapid than when it ig collected in flat dishes and the interaction spreads over a large area. In dealing with the fibrinogens—.e. the substances which in animal fluids are more immediately concerned in coagulation— it has been pointed out that they are by no means simple substances, that in particular they contain a very large per- centage of lecithin. It has always been assumed that in « ON THE PROCESSES TERMED ‘COAGULATION’ 289 dealing with the material substratum of fibrin we had only to consider alterations and modifications of proteid. So far as I can judge, the view which many chemists hold on the question of coagulation is much as follows: Proteids are substances which easily pass from the state of apparent solution to the coagulated state. In the coagulation of the blood a part of the proteid, probably under a ferment influence, passes into the solid state at a lower temperature than usual. ‘There is, therefore, nothing so especially remarkable about this. It is part of the general chemical question as to what is the nature of the change in the passage of proteid from the soluble to the insoluble modifi- cation. But this notion is utterly wrong. In the coagulation of the blood the formation of fibrin is but one thing—a striking phenomenon—but only a very small part of the whole process. Although it is paradoxical, coagulation processes occur without any coagulation. ‘The formation of fibrin is the possible out- come of a fibrinogen interaction, but only one part of a most complex chemical interaction. There are two processes in coagulation, I have already stated, which can be considered in something approaching chemical detail —the formation of fibrin and the formation of fibrin ferment. I pick out these two for the present, because the one is a very obvious, easily recognised process (fibrin formation); but it is essential to consider it with the formation of fibrin ferment, not because the fibrin ferment formation is a constant con- comitant of fibrin formation, as this is not the case, but for the reason that in coagulable fluids, especially if there is any tendency to spontaneous coagulation, there are many influences, the chemical bearing of which is quite obscure, which may accelerate the formation of fibrin. Since the formation of fibrin ferment essentially depends on chemical conditions, it is of the greatest value as a control in coagulation experiments. (For special precautions needful in testing for fibrin ferment, vide Section I. p. 222.) Now in the coagulation of shed blood all admit that the formation of fibrin and the formation of fibrin ferment occur. It is certain that neither of these bodies is present in blood when U 290 SECOND REPORT TO THE GROCERS’ COMPANY it leaves the vessels, and yet it is certain that fibrin forms and that the serum contains fibrin ferment. Hence it has long been known that, chemically considered, the coagulation of the blood involves more than the question of proteid coagulation. But, although it has long been known, it was not till my work on the coagulation of corpuscle-free plasmata that it came fully to the front. The fibrin ferment formation was wrapped up in a morphological expression. It was stated to be formed by the ‘ breaking up’ of white blood-corpuscles. However true this may be, it is obvious that, chemically, we get no further, and the notion of an enzyme action necessarily drew for a time one important chemical process out of the chemist’s vision. Hence the misunderstandings in coagulation. I have shown that fibrin ferment formation goes on in plasma—.e. goes on in a fluid.’ ‘The chemical conditions of its investigation are therefore now possible, and its origin is not necessarily connected with form-elements. It is, in fact, no longer a morphological expression. To illustrate this I take the following experiment :— Peptone-plasma is quite free from fibrin ferment. A solu- tion of tissue-fibrinogen is added, clotting ensues rapidly; the serum from this coagulation contains abundant fibrin ferment. Now we can investigate more nearly the chemical conditions under which these processes occur. ‘Tissue-fibrinogen is not a simple substance, it undoubtedly contains lecithin. (There could, I presume, be no dispute that the stromata of the red corpuscles contain lecithin.) And if, instead of adding tissue-fibrinogen, we add lecithin? prepared from tissue-fibrinogen, clotting also 1 «On the Origin of the Fibrin Ferment,’ Proc. Roy. Soc. 1884. [Collected Papers, p. 119.] * Lecithin.—The lecithin used for these clotting experiments I prepared with the greatest care. The solution and treatment with alcohol and ether was made over and over again. I have also treated (1) brain lecithin, and (2) lymph-gland lecithin prepared with platinum chloride as follows :—1, Solution absolute alcohol. 2, Precipitation PtCl,. 38, Washing with absolute alcohol. 4, Solution in chloroform, clear filtrate obtained. 5, Re-precipitation—alcohol, and treatment of mixture with H,S, 8-9 hours required. A little solid Na,CO, was added to neutralise any acid formed. 6, Evaporation, re-solution, treatment with freshly precipitated hydrate of silver. 7, Removal of ON THE PROCESSES TERMED ‘ COAGULATION’ 291 ensues ; fibrin ferment formation also takes place. This is a fact which is quite beyond any doubt ;! and it is of fundamental importance with regard to all our ideas concerning the chemistry of proteid matter. ; The extremely frequent association of lecithin with proteid has long been known. If now, in a process like coagulation of the blood, it can be shown definitely that lecithin can play an important chemical réle, it is absurd to suppose that the presence of lecithin with proteid means nothing. At the present time it is customary to divide up proteids by their solubilities, their being precipitated by this or that, or coagu- lated by this or that. It is known that we are not dealing with a single proteid, but for convenience we ignore that there is another substance present, and we say the substance is a parti- cular modification of proteid! We treat an organ or a fluid with this and that reagent. We see a coagulation occurs at such and such a temperature. Proteid, varieties of proteid ! Just so we might call chlorine, hydrochloric acid, chloride of sodium, varieties of chlorine; sodium, soda, carbonate of soda, varieties of sodium ; and the oxygen, the carbonic acid, &c. mere extractives! The procedure is as follows: A complex mixture is examined, its properties detailed. The mixture is then carefully boiled for hours with alcohol, &c. &c.; elementary traces of silver by H,S. 8, Re-solution and re-evaporation some six or eight times. The lecithins so prepared give all the reactions of lecithin, and I have most carefully tested their influence on coagulation. They possess the power in the highest degree. It must, of course, be understood that the term ‘lecithin ’iscertainly a genericone. Itis perfectly clear, from the only modern research (Hundeshagen), that, in the first place, there are polymeric substances closely allied to lecithin ; and, secondly, that the chemical structure of lecithin may be a very different one in different individuals. Thus both in hens’ eges and fish eggs there occurs, apparently dissolved in fat and not connected with proteid, a very easily extractable lecithin which has no influence on coagulation, although its physical characters are very similar indeed to the lecithin obtained from proteid. It is easy to make out that there are also very decided chemical differences—e.g. the platinum salt of the egg-lecithin is very easily soluble, that of proteid-lecithin hardly at all. But this is a line which, while doubtless very important to investigate, is not in the scope of my inquiry. 1 [Here, and in several other parts of this Report, the author refers forwards to sections which he intended to write. | 292 SECOND REPORT TO THE GROCERS’ COMPANY analysis made; conclusion, substance having the composition of the elementary analysis possesses the properties of the complex mixture. I have chosen in this section to take ferment formation as. the additional field of observation, because nothing can more clearly bring out the chemical bearing of lecithin in coagulation. Now, there is not the slightest doubt (I haye given many instances of such observations) that lecithin is most effectual in causing the appearance of fibrin ferment. It is quite easy to obtain specimens of peptone-plasma in which the CO, coagulation and the lecithin CO, coagulation are not very strikingly different until we come to estimate the ferment, and then we find that in the coagulation in presence of lecithin the ferment formation is increased a hundredfold. There is no other proximate principle which I have found to possess this power in any way. Great production of ferment is evoked by a variety of complex Substances (e.g. all the ‘fibrinogens’), but they all contain lecithin in abundance, and from consideration of this fact I have never had the slightest doubt of the chemical influence of lecithin. It must be understood that although lecithin causes clotting and the appearance of fibrin ferment, the clotting is not caused by the fibrin ferment. They are generally associated phenomena, but that there are not two processes—e.g. 1, formation of ferment influenced by lecithin; 2, formation of fibrin influenced by ferment—is seen from the following :— 1. Lecithin produces clotting under conditions in which the strongest fibrin ferment is ineffectual.’ 2. Under special experimental conditions lecithin produces coagulation, but no ferment. (Details and analysis of process to follow.) Not only does lecithin unquestionably enormously favour the formation of fibrin ferment, but there is probably never any fer- ment formed unless either free lecithin, or lecithin in a particular form (loosely combined with proteid) be present. This last statement is at present only an extreme probability. It is not a proven fact, because we are so utterly ignorant of the possible 1 Vide ‘ Uebersicht ti. d. Gerinnung.’ [Collected Papers, p. 186. | ~ QUANTITATIVE RELATIONSHIPS 293 relationships of proteid and lecithin, and it is so very difficult to get at the problem in a manner satisfactory to chemical criti- cism. I will merely shortly illustrate the point. A-fibrinogen, under given conditions which have already been mentioned, produces coagulation and ferment formation. After precipitation, it may again be added to plasma and still do this. But it does not do this if it has undergone too much ‘alteration’ (see page 274). Now A-fibrinogen, both before and after this alteration, contains lecithin, and it is impossible to say whether more or less; but coincident with its ‘alteration’ and loss of coagu- lation-producing power, it undergoes remarkable changes in its chemical behaviour. The alteration consists in the fact that it has become fibrinous instead of merely fibrinogenic; this is an obvious alteration. But the chemical point is that its behaviour on digestion is profoundly modified. AsI have pointed out, the fibrinogens have a very characteristic behaviour on artificial digestion. They readily yield a precipitate which is extraor- dinarily rich in lecithin. But after they have undergone the fibrinous modification they do not do this, and consequently I am inclined to think that in the latter case—i.e. in clotting— the lecithin-proteid relationship must have been altered. The exact meaning to be attached to the experiment is open to question, and requires very full treatment, which I am not at present prepared to give. This is the only illustration I will give at present. It is a very wide and important question, which I am and have for a long time been dealing with. Although I have confined myself to the occurrence of clotting and the formation of fibrin ferment, it must be thoroughly under- stood that these are but a limited part of the chemical aspect of coagulation. IX. Quantitative Relationships iv Fibrinogen Interactions From the general account given in my Croonian Lecture, it was clear that the process described under the head of Fibrinogen 294 SECOND REPORT TO THE GROCERS’ COMPANY Interaction was quite different from any fermentative process. This will be the more apparent from a consideration of the following quantitative determinations. The quantitative estima- tions adduced are purposely limited in number, because, as will be clear from subsequent statements, the greatest caution is necessary in drawing conclusions from such estimations. The previous quantitative work seems to have had in the main but one object, viz. to determine whether or not there was any ‘fibrinoplastic’ influence to be admitted in coagulation. Schmidt, in fact, relied mainly for the support of his theory on certain quantitative determinations. Certain transudation fluids yielded unmistakably larger quantities of fibrin when, in addition to ferment, paraglobulin was added. ‘This observation has been adduced to prove that paraglobulin takes a material part in coagulation, and it is really the only point on which Schmidt relied. But, as Hammarsten has pointed out, it is not necessarily a proof that paraglobulin is at all concerned in coagulation. It — is easy to adduce an instance parallel to those brought forward by Hammarsten from my own experience. Ferment added to peptone-plasma causes little or no coagu- lation, but with the addition of a stream of CO, there is infinitely more coagulation. It does not, however, follow that the CO, is a factor in the material building up of fibrin. The same objections can be raised, and have been raised, to Schmidt’s results. Now, although Iam quite certain that Schmidt afforded no convincing proof of a ‘fibrinoplastic action’ whatever, it appears to me that Hammarsten was wrong in his endeavour to show that there was no ‘fibrinoplastic action’ in Schmidt’s sense. I think Schmidt undoubtedly observed one special in- stance of a very important process which Hammarsten entirely overlooked, and which I have termed the ‘ fibrinogen interaction.’ The attempt to prove the participation of ‘ fibrinoplastin’ by quantitative determination of the amount of fibrin formed from any particular coagulable fluid only narrowed extremely the full consideration of the process of coagulation. It could only be effectual in the case where the quantity of fibrin formed exceeded the total fibrinogen of the coagulable fluid. This condition has, QUANTITATIVE RELATIONSHIPS 295 however, never been shown to occur, and hence Hammarsten’s objection that the ‘ fibrinoplastic action’ was merely an indirect influence must always hold good. Existence of the ‘ fibrinogen interaction’ could not possibly be adduced entirely from quantitative estimations of the fibrin formed ; for, as I have pointed out, a very pronounced fibrin- ogen interaction may take place without any fibrin at all being formed, but the very opposite condition is set up—.e. in intra- vascular injection of small quantities of fibrinogen the negative phase is evoked. For the material participation of tissue-fibrinogens (serum- fibrinogen, &c.) in the process of coagulation, therefore, it is obvious that only secondary importance can be attached to quantitative fibrin determinations, although they are not without interest and importance. The following observations on this subject refer to the inter- action of peptone-plasma and a solution of fibrinogen from the thymus gland. I would again recur to the point that this inter- action is not necessarily identical with processes occurring in nature. On adding a solution of tissue-fibrinogen to peptone-plasma coagulation occurs and the tissue-fibrinogen disappears—.e. the resulting serum does not contain tissue-fibrinogen. ‘This can be very clearly demonstrated when using the fibrinogen from thymus, because, as I have pointed out, this substance is extremely insoluble in acetic acid, whereas the fibrinogens of plasma and of serum are easily soluble in excess. But the power of a given quantity of plasma to effect this change is strictly limited, as will be obvious from the following :— Exp. A.—By tests the quantity of tissue-fibrinogen which could be added to a specimen of plasma and certainly disappear was deter- mined, when it was found that 20 c.c. of a solution of tissue- fibrinogen (from thymus) could be added to 30 c.c. of plasma and disappear. If more were added the absence of tissue-fibrinogen from the serum was doubtful. The acetic acid test was used. The 20 c.c. of tissue-fibrinogen solution contained ‘120 grm. tissue- fibrinogen. 296 SECOND REPORT TO THE GROCERS’: COMPANY Exp. B.—25 c.c. plasma totally clotted (i.e. the serum gave no further trace of clotting on adding tissue-fibrinogen) with 10 c.e. tissue-fibrinogen ; clotting occurred in five minutes. Weight of fibrin, 089. The dry weight of tissue-fibrinogen was ‘056, The plasma contained °548 per cent. fibrinogen. (For method of deter- mination see Note A at the end of this Report.) In this experiment the weight of the fibrin is greater than that of the tissue-fibrinogen added. Some of the fibrin must, therefore, have come from the plasma. The total fibrinogens concerned in the reaction were : Of plasma . ; ‘ ‘ ‘ . , o | *hSF Of tissue-fibrinogen ; ‘ : ; . 056 "293 Of fibrin’ ay S., due id es a ee The amount of fibrin is thus much less than that of the fibrinogen of the plasma alone. Exp. C.—20 ¢.c. plasma treated with CO,, gave slow clotting, Fibrin formed, 0°71. 20 c.c. plasma with tissue-fibrinogen in excess. Fibrin formed, 0°82. 20 c.c. of the plasma contained 1°28 fibrinogen. In this experiment there is a slight increase of fibrin in the tissue-fibrinogen coagulation. But determinations of this nature zive us no information as to the tissue-fibrinogen being materi- ally concerned in coagulation, and the amount of CO, coagula- tion varies greatly with external circumstances. It may be nothing or very extensive. Experiments of this kind are analogous to the fibrinoplastic determinations of Schmidt. They really only obscure the coagulation question. Proportionality between the amount of tissue-fibrinogen added and the fibrin formed. (The proportionality is marked, as explained in Section I., only when using strongly peptonised plasma.) Exp. D.—(1) 20 cc. plasma; 5 ¢.c. solution tissue-fibrinogen. Fibrin, ‘025 grm. (2) 20 c.c. plasma, 10 ¢.c. solution tissue-fibrinogen. Fibrin, ‘061 grm. Clotting in (1) did not occur for twenty minutes and was very incomplete. In (2) it took place in five minutes, This difference in time in the action of large and small quantities of the fibrinogen is always marked. The serum of experiment (1) QUANTITATIVE RELATIONSHIPS 297 remained quite clear for hours; clotted, however, through and through on addition of more tissue-fibrinogen. In Experiment B the amount of fibrin formed is greater than that of the tissue-fibrinogen added. It may be less, for small but weighable quantities of tissue-fibrinogen may be added to peptone- plasma without causing any coagulation at all. In Experiment B 10 cc. tissue-fibrinogen (‘056 dry weight) acting on 25 c.c. plasma produced total coagulation yielding 0-89 fibrin. With the same plasma, 2 c.c. tissue-fibrinogen (‘01 grm. solid) acting on 15 ¢.c. plasma produced none. It will be obvious, from the numbers quoted above, that esti- mations of the amount of fibrin formed cannot give us precise information as to the chemical process which has taken place, and for this reason. The fibrin formed is always less than the fibrinogen present; in other words, a part of the fibrinogen does not separate as fibrin, but remains in solution. Now quanti- tative estimations cannot give us full information unless we can exactly characterise and estimate the total quantity of the pro- ducts of fibrinogen interaction—i.e. not only the fibrin but the soluble substance or substances—~and this we cannot at present do. The soluble products of a fibrinogen interaction not only vary greatly in quantity but they vary in quality, and it is not possible to give any positive method of determination at the present time. The direction in which my researches have gone will be later pointed out. The study of the process of coagula- tion in peptone-plasma, and of the intravascular effect of fibrin- ogen injections, forces this question of the soluble products of “coagulation processes’ very much to the front. Probably it forms a part of what I have called the negative phase of the fibrinogen interaction, but not the whole. In dealing, then, with the question of whether a particular proteid enters materially into the formation of fibrin, it must be clearly understood that the weight of fibrin formed does not give us information. To take Experiment C. With CO,, 20 c.c. of the plasma gives ‘071 fibrin. The total fibrinogen in 20 ¢.c. of the plasma is ‘128; a great part, therefore, of the fibrinogen of the plasma is not separated as fibrin. 298: SECOND REPORT TO THE GROCERS’ COMPANY The same quantity of the plasma is treated with tissue- fibrinogen, and yields :08 fibrin—.e. a slight increase ;' but could the soluble products of the interaction be estimated, they would probably be greater too, and until this can be accurately done the quantitative estimation of fibrin is of little avail. Evidence obtained in other ways, to which I shall subsequently refer, is of far greater force than quantitative determinations. NOTE A FIBRINOGEN DETERMINATION In order to avoid side issues, and to make it clearly understood how little value is really to be found in quantitative estimation, I have not discussed the method of fibrinogen determination. This is naturally a matter of great importance, but at present we are not able to do much more than guess at the facts. The methods which heer been adopted by previous investigators ‘ have been those of taking the fibrin formed as a measure oo the amount of fibrinogen present, or of taking the amount of coagulum formed on heating to 56° as the measure of the fibrinogen. Now it is possible that under certain conditions the amount so obtained might give results mainly corresponding to the amount of fibrinogen. But it has also been shown that under certain conditions—z.e. when ferment acts on certain solutions of fibrinogen—a part of the fibrinogen always remains in solution, and also that when solutions. of fibrinogen are boiled a part remains in solution, and that the ratio of soluble to insoluble parts is variable (Hammarsten). Clearly, then, neither of these methods can pretend to accuracy, and they are in most cases instances of working entirely in the dark. The method I have used is open to objection, but it is at the present time the only one available. It depends on the following facts. Paraglobulin is soluble, though not easily, in saturated solu- tion of NaCl. Fibrinogen is not thus soluble. I therefore precipitate the plasma by saturation with NaCl. The precipitate is washed with saturated NaCl solution until all proteid reaction is gone from the wash-fluid ; this ought to remove the paraglobulin. The precipi- tate is then dried at 100° C. and washed with water till freed of all salt. The results are probably almost certainly too low. 1 The slight increase, ‘07-08, is not an indication of ‘ fibrinoplastic action, because the CO, coagulation is most variable. FIBRINOGEN DETERMINATION 299 I give an illustration of the importance of this question of fibrinogen determination. As is well known, Hammarsten states that the serum contains. more paraglobulin than the plasma, and this result has very gene- rally been adopted as true. I quote an illustration of the sort of evidence on which it rests. His method is this : he takes the cooled filtered plasma and the serum of the same animal. In both he determines the total globulins by the MgSO, method. In the plasma he determines the fibrinogen and subtracts this from the total globulins, the residue representing the paraglobulin. It follows from this method that if the serum really contains more paraglobulin than the plasma, it must come from something outside the plasma. The source is supposed to be the white cor- puscles—this being a pure hypothesis. The following is an instance :— Globulins in plasma . ‘ ° : : . » 4°87 Globulins in serum : , ‘ ° : : « 4:48 Fibrinogen of plasma (a) determined by heating . 0°32 Be “ (>) determined by fibrin . . 0°62 Paraglobulin of plasma incase (@) . ‘ ‘ . 4°55 (i.e. more than serum.) Paraglobulin of plasma in case (db) . : : . 4:25 (z.e. less than serum.) Now, although the percentage error of determination in the two. cases is not very great, it is obviously of great importance in dealing with such small quantities to know whether 0°32 is the amount, or the double of that, 0°62 ; because in this particular instance it is just this difference which tells for or against the assertion of Hammarsten. There is also to be considered in regard to this result that neither number, by Hammarsten’s own showing in subsequent papers, represents the real amount of fibrinogen, but only an un- known fraction, and also that the filtered cold plasma has already lost a considerable and unknown quantity of its fibrinogen (vide Section I.). All these numbers of Hammarsten are, therefore, purely fancy numbers, and it is impossible to deduce any result whatever from them. Now Hammarsten is an observer of whom I have the highest opinion, and I introduce this criticism merely with the object of showing that the attempt to solve the coagulation question by such chemical methods is at present likely to lead to false con- clusions, and by giving such false conclusions an appearance of scientific accuracy to tend to perpetuate error 300 SECOND REPORT TO THE GROCERS’ COMPANY It cannot be too clearly understood that we must have the chemical problems of coagulation sharply marked out; we must know for certain what our results mean. To continually draw conclusions which in reality rest on a pure assumption is a futile proceeding, because the assumption may turn out to be wrong. NOTE B ON THE REACTION OF FIBRINOGENS TO GASTRIC DIGESTION It is this reaction with an artificial digestive fluid which is so d characteristic of the fibrinogens, and which renders them so closely allied to the casein of milk. I think these bodies have long been known and spoken of as nucleo-albumins. It is an unfortunate expression. It seems to have arisen in the case of Plosz, working with the fibrinogen of the liver. On artificial digestion this yielded an albuminous precipitate, soluble in alkali, and very rich in phosphorus. Plosz concluded this was nuclein, without investigating the condition of the phosphorus. Now there appear to be proteid substances containing large quantities of phosphorus not in the form of lecithin, at any rate not extractable by alcohol. The term nuclein was certainly not meant to include proteids, which were mixtures or possible compounds of proteid and lecithin. And these digestion precipitates of the fibrinogens contain very little, if any, phosphorus after complete extraction with alcohol ; and, owing to the great difficulty of com- pletely removing the lecithin, it is difficult to say whether the phos- phorus which remains behind in small quantities is due to incomplete extraction or not. I am at present occupied with determining, as accurately as possible, the quantitative relationships of the Jecithin to the proteid in the precipitate. The following numbers are approximations ; they certainly underestimate the quantity of lecithin :— A. Fibrinogen from Thymus. On digestion with artificial gastric juice gave the usual precipitate. Weight of dry washed precipitate . ; . 1°68 gram. Weight of lecithin extracted from precipitate 24 ,, Percentage amount of lecithin in precipitate . 14:2 % B. Fibrinogen from Plasma. Dry weight of digestion precipitate : . 1:12 gram. Weight of lecithin extracted . ; : oD Percentage amount of lecithin , é + 28°5 % EiiebEN ex IO PATE meee N DEX «TO PART. T NOTE ON THE COAGULATION OF THE BLOOD} In a paper read before the Royal Society, April 26, 1888, Dr. Halliburton offers some criticism of my views respecting the coagu- lation of the blood. In this note I shall briefly summarise and traverse the objections Dr. Halliburton raises to my theory and experiments. I. Dr. Halliburton suggests that the substance I call ‘ A-fibrin- ogen’—which I obtained by cooling peptone-plasma—is not a normal constituent of the blood-plasma, but that it is a precipitate of a hemi-albumose, supposed by him to be present in the peptone which is injected into an animal for the purpose of obtaining peptone- plasma. Ido not use Witte’s peptone, as Dr. Halliburton appears to have done, on account of its recognised impurity, but that obtained from Dr. Grubler’s well-known laboratory in Leipzig. This peptone is prepared according to Henniger’s method. Ya 806 APPENDIX have endeavoured to give a short exposition of the work that I had already communicated. In 1883 I discovered the close connection existing between lecithin and the formation of fibrin, and in the same year I pub- lished three papers on this subject.!_ I also considered the relation of lecithin to the formation of fibrin ferment in the ‘ Proceedings of the Royal Society’ for 1884. As the papers of Samson-Himmelstjerna and Nauck appeared two or three years later, it was obviously impossible for me to quote them. Should Dr. Krier still maintain that these facts have been firmly established by these authors, I can only say that I am very pleased to have my discovery thus confirmed. With regard to the other ‘retrogressive proteid metamorphoses’ and their rela- tion to fibrin formation, I will gladly leave the priority of the discovery to these gentlemen, since I am of opinion that they are of no importance at all in the coagulation of the blood. Dr. Kriger has somewhat misconceived my position with re- ference to the co-operation of cellular elements. I certainly do not deny that leucocytes from lymph-glands and other tissue-cells have eminently the power of inducing coagulation in extravascular plasma ; in fact, I was the first to discover this.” But they lose this power so soon as they are introduced into the circulating blood. In my paper in Ludwig’s ‘ Festschrift’ I have shown that if leucocytes from lymph-glands be injected into the circulation they do not produce any intravascular clotting, and, moreover, the blood when drawn off will not coagulate ; they must, therefore, have lost their power of causing clotting during their sojourn in the living blood. It follows that the effects observed on adding cells to plasma cannot be taken to prove the participation of white blood-corpuscles in coagulation. The further objections of Dr, Kriger will be considered in a detailed account of my work on coagulation. THE COAGULATION QUESTION? In a paper read before the Royal Society in April, 1888, and recently in this journal, Dr. Halliburton has published his belief 1 «Zur Gerinnung der Blutes, Du Bois’ Archiv, 1883; ‘ Further Observa- — *, tions on the Coagulation of the Blood,’ Journ. of Physiol.; ‘On the Coagula- tion of the Blood,’ Zbid. [Collected Papers, pp. 106, 100, 112.] 2 Du Bois’ Archiv, and Proc. Roy. Soc. 1881. [Collected Papers, p. 89. ] 3 [From the Journ. of Physiol. 1889. (This paper was preceded by the follow- ~ THE COAGULATION QUESTION 307 that the fibrin ferment is a cell globulin, and also a somewhat extensive criticism of my work on coagulation. In the following discussion of the chief points which Dr, Halli- burton has raised against my work and views I will first take the last point handled by Dr. Halliburton in his ‘criticism.’ An important feature in my work on coagulation is the produc- tion of intravascular clotting by the substances I call ‘tissue- fibrinogens.’ As an explanation of this intravascular clotting, Dr. Halliburton said, in his Royal Society paper, there was fair reason to suppose _ the solutions of these substances were of a very slimy nature, so that they would, when injected, naturally block up the small vessels and thus cause coagulation ; just as an iron wire will cause coagula- tion when introduced into the sac of an aneurism. From the very first papers I had published on this subject (1886) it was clearly impossible that this suggestion could be correct, and in the light of what I had subsequently published, previous to Dr. Halliburton’s first paper, it was manifestly out of the question. With the greatest ease Dr. Halliburton could, by repeating my experiments, have seen whether his assertions were true or not. This he has now done, and he withdraws his observations unre- servedly. In his paper before the Royal Society Dr. Halliburton stated also that I had spoken of a substance, B-fibrinogen, in my abstract of the Croonian Lecture, and that this substance had disappeared from the paper which was published in Ludwig’s ‘ Festschrift,’ or had become identical with Hammarsten’s fibrinogen. Now since a large part of my paper in Ludwig’s ‘ Festschrift’ was devoted to describing B-fibrinogen, and the differences between it and Ham- marsten’s fibrinogen, Dr. Halliburton’s assertions were quite un- intelligible to me. In my researches on coagulation I lay great stress on a substance which separates from peptone-blood-plasma on cooling ; I call it ‘A-fibrinogen.’ I have taken the greatest and most conscientious trouble to investigate this substance, and I regard it, and I am sure with truth, as of the greatest importance in coagulation processes. Now Dr. Halliburton plainly indicated in his first paper that this ing Editorial Notice: The terribly sad event, the untimely death of the author on June 6, by which Physiology has been robbed of a bright and enthusiastic inquirer, has laid on me the duty of carrying out by myself the final revise of this paper. It has already been revised by the author; and I have en- deavoured to make as few changes as possible.—M. FOSTER.) | x 2 >» ce hl ae a pe ee ee ae ee ‘ : - a : . - , re) — Pe / 808 APPENDIX precipitate was merely an albumose mixed with the peptone injected, — and was in no way a constituent of the blood. The principal point in his criticism was, and is still, that solutions of Witte’s and Grub- _ ler’s peptone yield on cooling a precipitate of rounded granules. I pointed out that the peptone I use (Grubler’s) gives no precipitate on cooling, and it disappears entirely from the blood. I was, of course, aware at the time when I first found the ‘cold body’ (A-fibrinogen) that peptone (using the word as a generic term) had been obtained in rounded granules (Schmidt-Miilheim), and also — that other proteids—i.e. vegetable albumen and paraglobulin—could ~ be obtained in rounded granules ; for I had worked for a long time at Leipzig at crystallised proteids, the first step towards crystallisa- tion always being the production of rounded granules. The mere fact that A-fibrinogen can be obtained in rounded granules is, of course, no more entirely characteristic of it than would the state- ment that a body was crystallisable suffice to distinguish it. But from the very first there could be no doubt that this substance was not peptone. I had seen it for a long time before recognising what it meant. As I saw it, its appearance was precisely similar to a fibrinous clot, and for a long time I thought it was a partial coagu- lation of the plasma due to the presence of leucocytes. Iwasstruck by the fact, however, that this clot only appeared when I kept my __ plasma overnight in ice, and that it was absent from my plasma when kept in the warm room. Its characters, both chemical and physical, and its influence on coagulation, I have described in detail in my papers ‘ Ueber einen neuen Stoff,’ &c. ;! ‘On a New Constitu- ent of the Plasma’ ;? in Ludwig’s GWastdenrice and ‘ Beitrage zur Frage der pentane If Dr. Halliburton had suggested that — A-fibrinogen was merely fibrin it would have been intelligible but — not correct. But, although it is difficult from Dr. Halliburton’s statements * that cooling peptonised serum and other solutions of peptone give a precipitate exactly similar to that of A-fibrinogen (Wooldridge), and that, as Neumeister has shown, the albumose or peptone injected ‘ exists as such or in a loosely combined condition — in the blood for a time,’ to conceive that he meant or means any- thing else than that my A-fibrinogen is but such precipitated — albumose, we are now to understand (p. 280, the same on which the above statements are made) he means nothing of the sort. All he — means is that albumose alters the normal proteids of the plasma. i 1 Du Bois’ Archiv, 1884, * Proc. Roy. Soc. 1885. [Collected Papers, p. 124) 3 Du Bois’ Archiv, 1888. [Collected Papers, p. 253.] * Proc. Roy. Soc. vol. xliv. p. 266; and Journ. of Physiol. p: 280. THE COAGULATION QUESTION 809 This is quite a different view, and is, of course, a necessary conclusion from my own experiments and publications. In fact, Dr. Halli- burton tries to discredit my observations on this subject by— I. Hinting that it is nothing but albumose precipitated by cold ; indeed stating, page 278, $ 2, ‘that the occurrence of the so-called A-fibrinogen in peptone-plasma is produced by the peptone, is supported by the fact that if one takes a solution of Witte’s peptone and cools it to 0° C. a precipitate is produced consisting of rounded granules having very much the appearance of blood tablets.’ II. Asserting that, nevertheless, it is not albumose, is not the actual precipitate from the peptone injected. The flagrant contradiction of these two ways of looking at the subject is only consummated by Dr. Halliburton’s suddenly dropping the subject he raises—viz. what is the nature of the body I call A- fibrinogen ? But he proceeds to add remarks the justification for which rests on his ignorance of my researches on this subject. For instance, Dr. Halliburton complains that in my short answer to him in the ‘ Proceedings of the Royal Society ’ I do not give a list of the properties of A-fibrinogen. Am I compelled to rewrite all my descriptions and facts because Dr. Halliburton will not take the trouble to read them? On page 278 of the Journal paper Dr. Halliburton says the following in reference to the occurrence of A- fibrinogen in salted plasma :—‘ The statement that sodium chloride plasma contains A-fibrinogen is thus a matter of pure inference ; and the reasoning adopted by Wooldridge is a very obvious case of reasoning in a circle as follows: this form of salted plasma coagu- lates spontaneously, therefore it must contain fibrinogen A, and because this plasma contains fibrinogen A therefore it coagulates spontaneously.’ What I have really said on this subject is as follows : Peptone- plasma free from ferment and corpuscles— 1. Yields on cooling a substance which can be collected and examined, A-fibrinogen. 2. Clots spontaneously—z.e. by means not ! fibrin factors. 3. When it clots spontaneously yields fibrin ferment. 4, Cooled, and with A-fibrinogen thus removed, loses the power of spontaneous coagulation and of forming fibrin ferment. 5. Treated with sodium chloride solution so that the mixture contains 4 to 5 per cent. of the salt, yields no precipitate on cooling and retains the power of spontaneous coagulation. 6. If treated with MgSO,, as I have described, a precipitate 1 ? Independently of.—Ep. 810 : APPENDIX “9 is formed in the plasma closely resembling the cold precipitate, 7 A-fibrinogen. | 7. After the removal of this precipitate the plasma no longer coagulates spontaneously and forms no fibrin ferment. To judge of the action of salts on plasma we must first have it. — Normal blood of the dog (i.e. blood the moment it leaves the vessels), _ treated severally with 10 per cent. NaCl solution and MgSO, solution, __ yields plasmata exactly corresponding, as regards spontaneous coagu- lability and formation of fibrin ferment, to those obtained by the action of salts on peptone-plasma. Consequently I consider I am justified in the deduction that A-fibrinogen is present in salted plasma. In his own paper Dr. Halliburton says: ‘Magnesium sulphate — plasma is generally the slowest to coagulate when simply diluted with water ; this is, no dowbt, because some of the fibrinogen of the plasma is precipitated by the use of this salt.’ It is not clear why _ Dr. Halliburton says ‘no doubt.’ It is precisely the statement I _ have made in consequence of my study of the action of magnesium sulphate on peptone-plasma. Nor is it clear why, with his views, Dr. Halliburton thinks the removal of this fibrinogen should delay the occurrence of clotting. Has he any evidence to show that a slight diminution in the quantity of fibrinogen makes the rest clot slower with ferment? Why should it ? Now there is this difference between the two kinds of salted plasmainthedog. Thesodium chloride plasma clots in from three to ten minutes on simple dilution. The magnesium sulphate plasma — does not clot for forty-eight hours. Such a striking difference isonly __ explicable by the assumption that the fibrinogen removed by the MgSO, has a special power of initiating coagulation. What I have said is that NaCl plasma and uncooled peptone- plasma contain a special fibrinogen (A-fibrinogen) which has the — power of initiating coagulation. Dr. Halliburton first tries to throw ridicule on my results, and then for his own purposes practically admits they are true. He states in his paper in the Journal that I infer that so- lutions of tissue-fibrinogen cause clotting in extravascular plasma, — from its action on intravascular blood. But this again is quite inaccurate. In my paper in Ludwig’s ‘ Festschrift,’ and in a paper published _ in March, 1888, in Du Bois’ ‘ Archiv,’ I state and describe fully the action of tissue-fibrinogen on extravascular plasma. I quote from Ludwig’s ‘ Festschrift’ (which, Dr. Halliburton fre- THE COAGULATION QUESTION 311 quently refers to). ‘Diese Stoffe (Gewebes-Fibrinogen) bewirken nicht nur Gerinnung im extravascularen Plasma, wie Pepton-Plasma, sondern sie bringen auch beim Einspritzen u.s.w. Die genannten Stoffe sind, ahnlich wie Lymphkorperchen und Lecithin, ganzlich ohne Wirkung auf stark verdiinntes Bittersalzplasma—bringen sie dagegen extravasculares Plasma zur Gerinnung, so entsteht gleichzeitig viel Ferment.’ I think the illustrations I have given above indicate fairly the nature of Dr. Halliburton’s criticism. Having sufficiently dealt with what I must characterise as a travesty of my researches, I now pass to a consideration of the actual observations of Dr. Halliburton. Dr. Halliburton, to his own satisfaction, conclusively overturns all my work by the following highly remarkable dictum : ‘ Now Dr. Wooldridge admits too much ; he admits that the fibrin ferment causes coagulation,’ and ‘The formation of fibrin either is or is not a ferment action, it cannot be both.’ . Sugar is formed from starch by ferment action ; does Dr. Halliburton think it can be formed in no other way? In sucha state of mind does Dr. Halliburton approach the coagulation question. Truly, I admit that fibrin ferment sometimes causes the coagulation of fibrinogen ; to state that fibrin cannot be formed in any other way is to beg the question at issue, and consequently to assume most’ unwarrantably perfect knowledge of the whole process of coagulation. The destruction my observations undergo in Dr. Halliburton’s hands, when he treats them experimentally, is quite on a par with that treated logically. He completely ‘upsets one of the very foundations on which’ my theory is constructed. He describes six experiments in which he prepares, according to Schmidt’s method, solutions of fibrin ferment from lymphatic glands. He consequently ‘feels justified in concluding that the cells of lymph-glands yield as one of their disintegration products a substance which has the same properties as fibrin ferment.’ ‘This is a fact which has been observed by others ; it is, however, denied by Wooldridge.’ I have, of course, never denied in the least that leucocytes might as the result of disintegration yield fibrin ferment ; quite the contrary. The real statement I have made is reproduced in the very same sentence by Dr. Halliburton himself, and among the ‘others’ whom he quotes as supporting his view, and diametrically opposed to me, is Rauschenbach. 312 APPENDIX — Dr. Halliburton’s own work on leucocytes i is in many respects an — incomplete repetition of Rauschenbach’s. Rauschenbach, a pupil of Alexander Schmidt, repeated anda extended my original observations, using cooled plasma. He is the os originator of the statement that the leucocytes from lymph-glands _ contain no fibrin ferment and that they have no action on certain fibrinogen solutions. He is the originator of the statement that there are ‘mother substances’ of the fibrin ferment. From my own papers ~ eight years ago and from Rauschenbach’s onwards there has beena __ complete revolution in the coagulation question. I quote from Rauschenbach.! ‘When the cells are treated with distilled water and immediately filtered, ten parts of the filtrate with one part of salt-plasma (the test fluid) gave signs of a weak fermentative activity (clotting in four or five hours)’ (p. 27). ‘ ‘The slimy mass of leucocytes which had been produced by the action of salts, treated with alcohol yielded oy traces of fibrin ferment’ (p. 25). ‘ After the cells had been acted on by distilled water for many hours an active extract was obtained ; heating to 50° quickened the formation of this active extract’ (p. 27). ‘Whether the traces of fibrin ferment which even the freshest collection of cells contains, and which pass into the watery extractof the alcohol coagulum and into the rapidly filtered watery extract of the cells themselves, are of vital origin or are formed during the pressing out and filtration of the mass, I must leave for the present undecided’ (p. 28). ‘IT must expressly emphasise here that fresh pressed-out cell fluid, in a quite undiluted condition, exerted no perceptible action whatever on my fibrinogen fluids nearly free from paraglobulin. The same is | true of the watery extract obtained immediately after pressing out the cells’ (p. 30). My observations in regard to the absence of ferment from fresh isolated leucocytes are quite in accord with those of Rauschenbach. As Dr. Halliburton quotes him, one would gather that this observer’s statements were quite in harmony with those of Dr. Halliburton and opposed to mine ; whereas the exact contrary is the case. Now these observations of Rauschenbach clearly point to the fact that fresh isolated leucocytes contain none or dubious traces of fibrin ferment ; that by artificially breaking them up fibrin ferment i may be eel Dr. Halliburton, in dealing with me, says that — » 1 «Protoplasma und Blutplasma,’ Dorpat, 1882. THE COAGULATION QUESTION Sis all my complicated and in his eyes apparently ridiculous observa- tions fall to the ground because the leucocytes contain such a powerful fibrin ferment. He says, speaking of peptone-plasma : ‘Leucocytes contain a very powerful ferment, therefore they cause clotting in peptone-plasma, whereas serum and Schmidt’s ferment are not so powerful.’ There is not a particle of evidence in the whole of Dr. Halliburton’s paper in support of this statement—the one experimental point he endeavours to make against me. From the six experiments I have alluded to it is impossible to judge whether there is any fibrin ferment present at all. Since the only approximately reliable experiment, with strong magnesium sulphate plasma, is spoilt and rendered dubious by the very slight dilution of the plasma, and as no weight or measures or comparisons with serum are given, there is not even an attempt to prove his statement. Lymph-cells and leucocytes yield very active ‘preparations of fibrin ferment,’ says Dr. Halliburton ; although he in no way proves it, it may be true. But if it should prove to be true that leucocytes, without any other organic addition, can be made to yield active preparations of fibrin ferment, this would not in the least alter or injure what I have said on coagulation. I have said they do not contain it; I have not said they might not, by various destructive processes, be made to yield it. On the contrary, I have always said that when they cause clotting in peptone- plasma they also cause the appearance of fibrin ferment. Whether any other form of destroying the cells may cause the formation of fibrin ferment I cannot personally say. Dr. Halliburton seems to think that a leucocyte in a lymphatic gland alive and well is the same thing as a leucocyte treated with various salt solutions, dialysed, treated with alcohol, distilled water, and soon. It is, of course, easy to say, as Dr. Halliburton does, that leucocytes cause the clotting of the plasma because they contain a very active ferment. His experiments prove nothing of the sort. My own observations are quite opposed to this statement. We can only judge of the activity of a ferment solution by the rapidity with which it causes the clotting of a test fluid. Now solutions of fibrin ferment which cause clotting in highly diluted strong magnesium sulphate plasma in two or three minutes cause, when added to peptone-plasma in the same proportions, no clotting whatever or only traces after hours. Yet leucocytes added to the same peptone-plasma cause clotting in a few minutes. If ee APPENDIX these experiments mean anything, they mean that the leucocytes act quite differently to strong fibrin ferment. And it was perfectly apparent from my first experiments, published in 1881, that the inter- action which occurs between the leucocytes and plasma was of a totally different nature to a fermentative process. Now concerning Dr. Halliburton’s observations I would say the following :— They leave the chemical aspect of coagulation quite untouched ; his theory is therefore naturally simpler than mine. For to say that coagulation is due to a ferment is merely to envelop the whole question in the hopeless mist in which it has been for the last twenty _ years. Dr. Halliburton says the white corpuscles liberate fibrin ferment when blood is shed, and he says his theory explains why the blood coagulates outside the vessels. This is, of course, the old theory of Alexander Schmidt ; a theory which has long been shown by its author and his pupils to be hopelessly untenable. It was shown to be impossible in my first papers in 1881. Dr. Halliburton’s repetitions of Schmidt’s experiments and his deductions therefrom are as follows. By destroying lymph-cells in various ways he obtains solutions which he says contain the fibrin ferment. Therefore, he says, when the white corpuscles leave the blood-vessels they break up and yield this ferment. Finally, he says, leucocytes contain a very active ferment, a much more active ferment than serum. Now if, for the sake of argument, we accept these observations as correct, it is clear that there is a very large gap between Dr. Halliburton’s deductions and his observations, because we know with certainty that white blood-corpuscles when they leave the vessels contain no fibrin ferment. Leucocytes and white blood-cor- puscles are then, according to this, totally different, if Dr. Halli- burton’s statements are correct and he means what he says. Under these circumstances, therefore, the ferment theory as propounded and maintained by Schmidt falls to the ground in Dr. Halliburton’s hands. He himself does not appear to grasp the fact that Schmidt was quite aware that the leucocytes in the blood contained no fibrin ferment, and that he accordingly put forward the hypothesis that a specific form of protoplasmic change occurred which led to the development of ferment from the disintegrated corpuscles. Now, unfortunately, the views of Schmidt and Hammarsten, edited by Dr. Halliburton, are supported by him by an experimental method which is open to fatal objections, THE, COAGULATION QUESTION 315 He uses, as a test for his supposed fibrin-ferment solutions, a fluid which he says clots rapidly with substances which are not fibrin ferment. It is, therefore, extremely difficult for him to know when he is dealing with the ferment and the non-ferment. He says that because his solutions cause clotting in his test plasma it cannot be concluded that they contain ferment, because other substances, e.g. muscle, cause rapid clotting. (In his first paper in the Royal Society he omits this.) It is, therefore, not at all clear how he knows he is dealing with fibrin ferment at all. His identification of the ferment would appear to depend on the temperature at which his solutions lose their coagulating activity, and even this is, according to his own statement, a somewhat precarious method of identification. The fibrin ferment is, he asserts, ‘ cell-globulin,’ a substance con- taining no phosphorus. ‘It is a true ferment, an unstable substance, the product of a living cell, and that induces changes in the sub- stances with which it comes in contact by a catalytic action—+.e. without itself apparently participating in the chemical changes its presence induces.’ Dr. Halliburton, it may be seen, is very precise on the subject. But, as a matter of fact, all this rests purely on his upse dre«it. Long before he took up the subject of coagulation it had been abundantly shown that all kinds of different substances when added to plasma cause coagulation. These substances were not fibrin fer- ment. And therefore it became of the utmost necessity to adopt the greatest precautions in testing for this substance. The two fluids which afford reasonable certainty that we are dealing with fibrin ferment are (1) the test fluid of Alexander Schmidt—7z.e. highly diluted strong magnesium sulphate plasma, and (2) the fibrinogen of Hammarsten—7.e. the fibrinogen prepared from the strong MgSO, plasma of the horse by repeated precipitation and re-solution. They afford a reasonable certainty of the existence of ferment, because, so far as is known, they have little or no tendency to clot with ‘ other substances.’ Dr. Halliburton appears very seldom to use these test fluids. As I have pointed out before, the serous fluids which he employs are very unreliable ; they vary in different cases, although as a general rule they resemble Hammarsten’s fibrinogen in being easily affected by ferment and not by ‘ other substances.’ It must be remembered that the original and continued belief in the fibrin ferment depends almost entirely on our being able to separate it from proteids. Its recognition by Schmidt depended on this. The whole of Hammarsten’s work depended on this. If it — cannot be separated from proteid, then the proof that it produces 316 APPENDIX ‘changes without itself participating in the chemical changes becomes an exceedingly difficult one, and has never yet been pre- sented to science. Dr. Halliburton says ‘that his ferment does not participate,’ but I cannot see that he has afforded any material proof whatever. In- deed, he is not clear that his proteid solutions, which he calls ‘ fer- ments,’ do not influence the amount of fibrin formed, and he affords no direct proof whatever that they are not materially altered. He does not give convincing evidence that the various substances he speaks of as ‘fibrin ferment’ are one and the same body. It is, of course, extremely hazardous to assert that one proteid is identical with another. But first there is one point Dr. Halliburton seems to have omitted to profit by. His cell-globulin from lymph-glands is ‘quite free from phosphorus.’ ‘This is the first proteid, if this is true, ever found free from phosphorus (Bunge, Ritthausen). It isa pity he has not applied this method of identification to his other cell-globulins. Then Dr. Halliburton makes a very remarkable statement about the fibrin ferment, which is entirely opposed to all previous experi- ence. He says that even } or | per cent. solutions of sodium chloride hinder very considerably the action of the ferment. I hope Dr. Halliburton has made direct observations on this point, since it is another of the sweeping and unsupported statements he has made concerning my work. ‘The action of sodium chloride has been very fully worked out by the Dorpat school. The presence of salt solu- tion up to 3 per cent. causes not a hindrance but a decidedly increased rapidity in fermentative coagulation. Against this well-known fact he brings no experimental evidence. The sole original point which Dr. Halliburton has made is that the fibrin ferment is a proteid. All the observations he gives on clotting have been made before. We have known for years that leucocytes from lymph-glands cause clotting, that extracts containing proteid from leucocytes cause clot- ting, that a proteid from serum causes clotting, and it is still for Dr. Halliburton to prove that these substances are the fibrin fer- ment, or that they act as an enzyme at all, and even then we shall not have got much further in the knowledge of the intimate nature of coagulation. How extremely little value can be placed on the effect of heating as a test for an enzyme can be seen from an earlier observation I have long ago recorded. Tissue-fibrinogen causes intravascular clot- ting ; after the solutions have been boiled they lose this power. Yet they are just as active on extravascular plasma as on peptone-plasma. THE COAGULATION QUESTION S17 In conclusion, then, Dr. Halliburton’s sole experimental point against me is that leucocytes cause clotting in peptone-plasma because they contain a ferment so very much more active than the ferment of Schmidt or than serum. He does not, however, adduce one single observation which proves that leucocytes contain or even yield any ferment whatever, and there is no attempt to prove that their alleged ferment is so very active. The observer he quotes in support of his views flatly contradicts him. Surely an observer in Dr. Halliburton’s position, who brings such sweeping criticism to bear on my work, is bound to show that his one observation ‘which completely upsets one of the very foundations upon which my theory rests’ is a true and correct one. But at present Dr. Halliburton has only made the assertion, he has not attempted to prove it. Part II PAPA OwOGlOAT: PAPERS 321) NOTE ON PROTECTION IN ANTHRAX? HITHERTO in the few cases in which protection against zymotic disease has been found possible, it has been effected by the communication to the animal of a modified form of the disease against which protection is sought. I have succeeded in protecting rabbits from anthrax by an altogether different process, and although this is scarcely, at present, of practical utility, it may perhaps be found to be of some interest as regards the general nature of protection in this and other diseases depending on micro-organisms. I use as a culture fluid for the anthrax bacillus a solution of a proteid body which is obtained from the testis and from the thymus gland. I have described this substance to the Society on a previous occasion,” so that I need not repeat the descrip- tion of the process used in its preparation. The proteid substance is dissolved in dilute alkali and the solution sterilised by repeated boiling. It is then inoculated with anthrax and maintained at 37° C. for two or three days. The growth is generally not very abundant, and at the end of the period mentioned is removed from the culture fluid by filtration. A small quantity of the filtered culture fluid is in- jected into the circulation of a rabbit, and it is then found that the animal will not take anthrax. A subcutaneous inoculation of extremely virulent anthrax blood made at the time of the injection of the protecting fluid, and two subsequent inoculations at intervals of five and ten 1 [From the Proc. Roy. Soc. vol. xlii. 1887. ] ? «Intravascular Clotting,’ Proc. Roy. Soc. 1886. [Collected Papers p. 135.] Y 822 PATHOLOGICAL PAPERS days, remain entirely without effect. The animals used as a control invariably die. Four rabbits have been protected in this way. If the anthrax grown in the fluid be inoculated it either kills or it has no effect. It does not protect in the slightest degree. The injection of the culture fluid in which no anthrax has grown is without effect. The animals die as usual when inocu- lated. The injection of the fluid itself causes no ill symptoms whether anthrax has grown in it or not. If other albuminous fluids-—e.g. blood serum—be used as a culture medium and the filtered culture fluid be injected, it exerts no protection. It may be fairly concluded that the growth of the anthrax bacillus in the special culture fluids used in these experiments gives rise to a substance which, when injected into the organism, protects against an immediate and subse- quent attacks of anthrax. It would obviously be of very great advantage if some such method as this could be used for the zymotic diseases affecting man for which no protective inoculation in the ordinary sense appears possible. I am indebted to the Medical Officer to the Local Govern- ment Board for permission to publish this short account of these experiments, the full description of which will appear in his report. I must also express my thanks to Dr. Klein, F.R.S., for kindly supplying me with many anthrax cultivations. Note added The following experiments give additional weight to the previously described results :— In the one case the anthrax grew with very great rapidity in the culture fluid, and the clear filtrate contained but a very small quantity of proteid matter. Forty cubic centimétres of this fluid were injected into a rabbit, and the rabbit immediately inoculated in the ear with virulent anthrax blood; in two days there was very marked cedema at the seat of inoculation, which increased to an enormous extent during the next few days, and NOTE ON PROTECTION IN ANTHRAX 323 then gradually subsided. The rabbit is now perfectly well, twenty-four days after the inoculation. In the second case the growth of anthrax had been very slight; 20 c.c. of the filtered fluid was injected, and the animal immediately inoculated in the leg with virulent anthrax blood. In three days there was marked cedema at the seat of inocula- tion. This spread up the leg to the back, so that there was enormous cedema occupying nearly the whole posterior part of the animal ; this persisted for ten days, and then gradually sub- sided. The animal is quite well, twenty-eight days after inoculation. These cases are of interest, since they are obviously instances of partial protection. The animals are still affected by anthrax, but it is only as a severe local affection, and does not kill them. Tid 324 PATHOLOGICAL PAPERS PRELIMINARY REPORT ON THE MODE OF ACTION OF PATHOGENIC ORGANISMS! THERE can be scarcely any doubt that the ultimate action of pathogenic organisms in producing disease is a chemical one; and, although suggestions and theories have not been wanting as to the nature of this action, the subject is still involved in very great obscurity. Dr. Klein has placed at my disposal, for the purposes of research, a bacillus which produces a rapidly fatal form of septicemia in guinea-pigs and rabbits. Dr. Klein has himself studied and described the morphological and physiological characters of this organism, so that it will only be necessary for me to give an account of the observations I have been able to make as to the mode in which this organism produces its fatal effects. It must be added that, owing to the short time I have been engaged on this subject, these observations are still very incomplete. It has recently been shown? that, under the influence of certain non-pathogenic organisms, poisonous substances of the nature of alkaloids are produced in organic media; and it has been assumed that such poisons may arise in the animal body as the result of the activity of pathogenic organisms, thus account- ing for the toxic influences of the latter. But satisfactory proof that pathogenic organisms are capable of producing such poisons does not at present exist; and as this is a very important sub- ject, attention was particularly directed to this point in the case of the septiczemic bacillus now under consideration. For an inquiry of this nature there are objections to the use 1 [From the ‘ Report of Medical Officer,’ Local Government Board, 1887, | 2 Brieger, ‘ Ueber Ptomaine.’ MODE OF ACTION OF PATHOGENIC ORGANISMS $25 of the more complex media more generally employed for purposes of culture, such as peptone-broth, or peptone and sugar; and in the case of this particular organism it was found more suitable to make use of a vegetable albumen. Vegetable albumens are obtainable commercially from various sources, and the one used in these experiments has been obtained from the Brazil nut. The raw product is extracted with dilute alkali, filtered, and the clear solution precipitated by neutralisation with acetic acid. The solid matter is collected by means of the centrifugal machine and washed. It is again dissolved in dilute alkali, and this solution forms the cultivating medium for the septic bacillus under consideration. With regard to this solution, it is to be observed that it may be injected in large quantity into the circulation of the rabbit without producing any obvious dis- turbance. Before inoculating the bacillus into the solution of vegetable albumen, the solution is thoroughly sterilised by prolonged boil- ing on several successive days. Provided sufficient alkali be pre- sent, the only change the albumen solution undergoes on boiling is that 1t becomes very slightly opalescent. It will be necessary hereafter to ascertain the exact amount of proteid and of alkali which the solution should contain; here it may be said that it is of great importance to avoid any excess of alkali, as this exerts an unfavourable influence on the growth of the bacillus. When the solution of vegetable albumen has been fully steri- lised, it is inoculated from a pure culture of the bacillus on peptonised gelatine broth, and the flask containing the albumen solution is kept in the incubator at a temperature of 36°-37° C. After a short period, varying from two to four days, a volumin- ous precipitate occurs in the fluid. The time which elapses be- tween inoculation of the solution and the appearance of the precipitate depends on conditions not yet understood, among them the degree of alkalinity of the fluid probably being the most important. When the precipitate has appeared, a small quantity of the contents of the flask is drawn off by means of a sterilised pipette, 326 PATHOLOGICAL PAPERS and from this gelatine tubes are inoculated, so that a control as to the purity of the culture may be obtained. ‘The general characters of cultivations on gelatine are given in Dr. Klein’s paper. . Microscopically examined, the precipitate is found to contain large numbers of bacilli, but at the same time it is found on care- ful examination that these bacilli do not make up the bulk of the precipitate. That consists principally of a very finely granular substance, which chemical examination shows to be chiefly proteid matter. Indeed, when the precipitate has fully developed, the supernatant fluid is found to contain only a very small quantity of proteid. The culture fluid is still markedly alkaline when the precipi- tate is fully developed. Hence the precipitate cannot be referred to the production of an acid body by the bacillus. The growth, then, of the bacillus has caused the precipitation of the bulk of the albuminous matter contained in the original solution. This matter can hardly be regarded as representing the débris of or- ganisms ; for, in the first place, the rapidity with which the pre- cipitate is formed renders this almost impossible ; and secondly, the bulk of the growth and of the débris of growth which is formed in other fluid media (for instance, peptone-broth) is very much smaller than the precipitate under consideration. The precipitate caused by the operation of the bacillus can be readily collected and washed by means of the centrifuge. When so collected it is found to be much less readily soluble in dilute alkali than was a precipitate formed by acetic acid from the original solution of vegetable albumen. The bacillary precipitate having been treated with dilute alkali is filtered through a fine linen cloth, and in this way a fluid is obtained, turbid, but free from any visible solid particles. This fluid possesses very marked toxic properties, for if injected into the circulation of a rabbit it will cause death in less than a minute. To produce this result, however, a considerable quantity of the fluid must be injected, or, more exactly stated, the fluid in- jected must contain a considerable quantity of the precipitate in MODE OF ACTION OF PATHOGENIC ORGANISMS 827 suspension. I estimate the quantity necessary to kill a rabbit as 0°5 grm., but it must be understood that this is only an ap- proximation. A less quantity produces no very obvious immedi- ate effects; butas the animal is under the influence of an anes- thetic when the injection takes place, milder symptoms may be masked. ‘The animal is killed before the anzesthesia is over. The fluid which was separated from the precipitate, that is to say, the original culture fluid after the removal of the precipitate, does not exert any immediate toxic effect when injected in large quantity into the blood.! No doubt the precipitate caused by the growth of the bacilli contains, after being taken up by the dilute alkali, a great num- ber of the bacilli in suspension. But then, in like manner, there are a large number of bacilli to be found in that solution of the proteid which had shortly before been inoculated with them, and in which the precipitate has not yet made its appearance; yet the injection of this solution is not followed by any immediate toxic effect ; so that the rapid toxic effect cannot be referred to the bacilli themselves. The actual cause of the rapid death, above referred to, appears to be respiratory paralysis, the heart continuing to beat for some time after the respiration has stopped. In one or two instances I have observed intravascular clotting, but this is not constant. In four cases, however, a marked acceleration of the clotting of the blood after it left the vessels was noticed. These changes in the blood are of interest because it has been shown ? that a very poisonous substance can be obtained from many animal tissues. ‘The substance is a complex proteid body, and it presents some analogies in its mode of action with the precipitate produced by our septiceemic bacilli. As has been stated, the latter consists chiefly of proteid matter ; whether itis this which has the toxic power, or whether this power is due to some other substance contained in the precipitate, must be left an open question till further experiments can be made. 1 «Immediate,’ because the animal is killed before the expiration of the aneesthesia. 2 Proc. Roy. Soc. Feb. 4, 1886. [Collected Papers, p. 135.] 328 : PATHOLOGICAL PAPERS One conclusion may be safely drawn from these observations, viz., that the growth of this bacillus in vegetable albumen solu- tion does: not result in the production of any soluble poisons, since only the precipitate has any toxic action—the fluid is inert. If it can be definitely shown that growth of bacilli in solutions of innocuous proteids can rapidly convert these into noxious bodies, a great advance in our knowledge of the action of patho- genic organisms may be possible, and perhaps these observations may be taken as an indication in this direction. They are certainly not favourable to the ptomaine theory. In addition to the observations already described, one other point of interest was brought out. It was found that the organisms could be cultivated in the following fluid :— Kreatin, 1 grm. Glucose, 0°5 grm. Potassium phosphate, 0°5 grm. Lime and magnesia, traces. 4 per cent. solution of common salt, 100 c.c. The growth produced was always of very limited extent, but the toxic influence of the organisms was preserved through many successive generations. But although the organism still retains its activity when inoculated, it is very much less virulent than when grown in peptone-broth or other albuminous fluid. Inoculations from a growth of peptone-broth cause death in guinea-pigs in from two to four days; whereas the average time elapsing between inoculation and death in the case of the erowth in the kreatin solution was seven days. 329 RESEARCHES ON CHEMICAL PROTECTION! It is well known that in many zymotic diseases the blood is profoundly affected, and that in these diseases we meet with symptoms of general intoxication, associated with localised pathological processes, and with the remarkable phenomenon of protection against a subsequent attack of the disease. In the course of my investigations on the coagulation of the blood I have met with results so obviously related to the processes alluded to above as to inspire me with the hope of approaching more nearly the ‘Chemismus’ of zymotic disease. I must express my thanks to my revered teacher, Professor C. Ludwig, for having allowed me to carry out many experi- ments, which are recorded in this paper, in his laboratory. I. The Action of Tissue-fibrinogen * on the Blood Hyery watery extract of any fresh tissue may be considered as a solution of tissue-fibrinogen. To purify the solution it is precipitated with acid. The precipitate 1s washed, and dissolved in very dilute alkali. The injection of this purified slightly alkaline solution into the vein of a rabbit produces total throm- bosis of the whole vascular system. In the dog, on the other hand, thrombosis is confined to special regions, viz., the portal venous system, as I have already described in these ‘ Archives,’ 1888, page 174.3 The dogs recover, in many cases, from ' [Translated by the author from Du Bois’ Archiv, 1888, p. 527.] * Tissue-fibrinogen, vide ‘Intravascular Clotting,’ Proc. Roy. Soc. 1886; ‘ Ueber intravasculire Gerinnung,’ Du Bois’ Archiv, 1886 ; ‘ Beitrige zur Frage der Gerinnung,’ ibid. 1888; ‘On Hem. Infarct. of Liver,’ Brit. Med. Journ. ‘Nov. 1887. [Collected Papers, pp. 135, 253, 346.] 3 [Collected Papers, p. 253. ] 330 PATHOLOGICAL PAPERS this lesion. But if, some hours later, a second injection of this solution be made, it has practically no effect, even if it be in much larger quantity. The first injection produces not only the constantly occurring local thrombosis, but the blood is so altered that it is rendered indifferent towards a second injection. This condition may last, according to the amount. of the first injection and other circumstances, for several days. Thrombosis and diminished coagulability of the blood are quantitatively closely associated. If small quantities are in- jected, only insignificant thrombosis results, and the shed blood clots slowly. If large quantities are used, there is ex- tensive coagulation, and the shed blood clots with the greatest difficulty. ‘To illustrate the above I describe the two following experiments :— 1. Forty c.c. of a solution of very freshly prepared tissue- fibrinogen were injected into the jugular vein of a dog (weight 54 kilos.). After an interval of from one to two minutes there was a very slight convulsion and the respiration stopped com- pletely; after three minutes the respiration returned and con- tinued for five minutes, then ceased permanently ; no recovery of consciousness. Sec. caday. was made at once. The portal vein and all its branches leading to the liver were completely thrombosed. Also most of the mesenteric veins, the splenic vein in particular, very strongly thrombosed. The lymphatic vessels. leading from the liver were full of blood. Blood was effused beneath the peritoneal covering of the gall-bladder. The liver, which was otherwise pale, showed extensive and distinct heemor- rhagic infarctions. In the right heart was a fibrous shrunken coagulum, the animal being in full digestion.!. Elsewhere there were no coagula. The blood from the heart was collected, and not being spon- taneously coagulable it was centrifugalised ; the plasma of this. blood coagulated on further addition of tissue-fibrinogen, and also on the addition of leucocytes from lymph-glands. Left to itself, coagulation began in twenty-four hours. 2. Dog. Weight, 7 kilos. during digestion. At 11.40 A.M. ! Vide these ‘ Archives,’ 1888. [Collected Papers, p. 253.] RESEARCHES ON CHEMICAL PROTECTION 831 50 c.c. of the same solution were injected. Temperature of the: dog before the injection 388° C. Immediately after the injection convulsions, cessation of respiration ; pulse in the cruralis not to. be felt. The animal recovered quickly and was let loose. After re-. covery from the (chloroform) narcosis, it was observed to have marked weakness of the hind legs for a considerable period. At 3.30 P.M. the animal was again narcotised (the temperature of the dog was 40°1° C.), and had 50 c.c. of a solution of the same tissue-fibrinogen, but of double strength, injected into the other jugular vein. A scarcely perceptible struggle occurred; the breathing went on quietly. After five minutes the carotid is opened, the blood streams out under high pressure, and without difficulty 280 c.c. are collected. (On the other hand, as I have previously recorded, the result of a first injection is to cause so ereat a fall in blood-pressure that it is difficult to get a single drop of blood from the carotid.) The animal was killed by chloroform. In the vena porta is a shrunken thrombus 2 to 3 cm. long; the margins are already distinctly decolourised. The clot is continued upwards into several branches of the portal vein in the liver. It is prolonged downwards into a very thin white centrally-placed fibrin thread not thicker than ordinary sewing-cotton. Such threads are also. to be found in the mesenteric veins also distinctly decolourised. The splenic vein contains a large shrunken and distinctly de- colourised thrombus. After very careful examination I found two mesenteric veins with fresh coagula not decolourised. The right heart contains a very small quantity of fibrous completely decolourised coagulum. The pulmonary artery, left heart, venz cave, subclavian veins, iliac veins, contain completely fluid blood. The liver shows unimportant hemorrhagic spots; the kid- neys are microscopically unaltered; the urine is coloured with methemoglobin, due, I am inclined to think, to the rapid decolourisation of the clots. The other internal organs present no abnormality. The blood was centrifugalised ; the plasma was slightly fatty,, hye PATHOLOGICAL PAPERS so that in large quantity it appeared whitish, but in small test- tubes it was quite clear. This plasma has the following properties. It did not clot on addition of tissue-fibrinogen during six hours—time of observa- tion. With leucocytes from lymph-glands no coagulation in six hours. With a very active solution of fibrin ferment no coagu- lation in five hours. (To 10 c.c. plasma 5 c.c. ferment solution added. Twoc.c. of the same ferment solution produced coagu- lation in 25 c.c. dilute MgSO, plasma [ferment test] in fifteen minutes. ) With lecithin a rapid trace of coagulation occurred, but did not increase. That the plasma contains none of the injected tissue-fibrinogen is evident from the following: The solution of tissue-fibrinogen, even after it has been diluted twenty-five times, gives a permanent and distinct precipitate after addition of acetic acid in large quantity (4 c.c. acetic acid 35 per cent. to 2 c.c. fibrinogen solution). The plasma treated with acetic acid in the same proportion remains perfectly clear. The plasma remained entirely fluid for three days. Nevertheless, the plasma contains much fibrinogen. With dilute sulphuric acid (0-4 per cent.) added till a marked acid re- action is obtained, it gives a voluminous proteid precipitate. In - another portion of the plasma the fibrinogen is precipitated in the usual way with NaCl. ‘The precipitate is washed with satu- rated NaCl solution till disappearance of proteid reaction, the salt removed, and the precipitate dried and weighed. I found 0-93 per cent. fibrinogen. If it be remembered that the great mass of the blood has remained fluid, and that only in the portal and splenic veins extensive thrombi have formed, the large amount of fibrinogen still left in the blood is in no way surprising. The point of interest is that, although the solution is added to the whole circulating blood, it only produces pathological changes in certain defimte regions, and that the first wyection so changes the Sibrinogen of the blood that vt remains indifferent to a second wyec- tion, both when in the vessels of the living animal and when shed. RESEARCHES ON CHEMICAL PROTECTION 333: II. Use of Boiled Tissue-fibrinogen as Culture Fluid. Haperi- ments with Anthrax In order to study the changes which solutions ot tissue- fibrinogen undergo owing to the action of pathogenic organisms, it appeared to be of importance to use a bacillus which acts. quickly and certainly. I have, therefore, confined my obser- yations chiefly to the bacillus anthracis.’ The solution is always sterilised by boiling before being inoculated; and as the result of boiling, the fluid undergoes changes in its chemical and physiological characters which I will first describe. The boiled solution has lost its power of producing coagu- lation when injected into the circulating blood, but it retains the power of producing coagulation in extravascular plasma,? e.g. peptone-plasma. As the result of boiling, a greater or less quantity of the tissue-fibrinogen is coagulated. The coagulum cannot, however, be regarded as ordinary coagulated albumen ; for the coagulated (by boiling) fibrinogen possesses in a high degree the power of inducing coagulation (fibrin) in peptone- plasma. With regard to the quantity and properties of the coagulum, different solutions are in no way identical. We can conveniently describe three varieties. Hither the greater part of the fluid coagulates, or the fluid becomes merely opalescent, or it is not appreciably changed. The different behaviour depends certainly in part, but not exclusively, on the amount of alkali present in the solution. By means of a certain degree of alkalinity, solutions can always be obtained which are not changed by boiling. But the amount of alkali requisite to produce this result cannot at present be generally stated, because tissue-fibrinogen is a very changeable body, and because the extracts which are obtained from different specimens of thymus are not precisely identical. For convenience I will style such solutions, which undergo no change on boiling, ‘strong alkaline solutions.’ It must be 1 Vide ‘Report of Medical Officer,’ Local Government Board, 1887, and forthcoming Report. [Collected Papers, pp. 324, 340.] 2 Thid. . OO4 PATHOLOGICAL PAPERS understood that this term is used relatively—z.e. sometimes such solutions are really very alkaline, sometimes only weakly alkaline. The different behaviour of different boiled solutions of fibrinogen is a marked feature in their behaviour as culture fluids. In the ‘strong alkaline solutions’ the bacilli grow with very great rapidity and in great abundance, and they kill, after many days’ culture, with great rapidity when inoculated. But the fluid itself is not poisonous. It can be easily separated by ordinary filter-paper from the bacilli, which form a coagulum- like mass of long fibres. The filtrate can be injected into the circulation of arabbit without producing any evil effect whatever. The animal acquires no immunity as the result of the injection. It dies in the usual period if it be inoculated subcutaneously with anthrax blood. ‘Weak alkaline solutions ’ behave quite differently. In these the anthrax bacilli sometimes will not grow at all, or they grow to a certain extent and lose their virulence, or they rapidly exhaust the proteid of the fluid and are still very poisonous. With such ‘weak alkaline’ culture fluids I have been repeatedly able to render rabbits immune as regards anthrax; but the bacilli must not be injected into the blood together with the fluid. They can be separated from the fluid by filtration, as has just been described. This procedure, however, with scanty growth, is uncertain, and it is better to kill the bacilli by boiling. But now it is found that the fluid—which, as described, has already been boiled previous to inoculation—has again become sensitive to boiling, and displays a greater or less tendency to form an additional coagulum. ‘This point will be referred to subse- quently. If the solution, now freed from bacilli, be injected into the vein of the animal, the latter as a rule acquires immu- nity towards anthrax—i.e. it can, immediately or later, be subcutaneously inoculated with the most virulent anthrax blood without any ill effects. I have protected a large number of animals in this way, so that there is no doubt about the observation being correct. RESEARCHES ON CHEMICAL PROTECTION 335 The protective action lasts very long. In one case it was in full power after an interval of fifteen months. The best way to proceed to obtain protection is as follows: The watery extract of thymus, or the very weak alkaline solu- tion of the acetic acid precipitate, is boiled, diluted, and filtered through linen. The fluid is inoculated with anthrax, and the culture incubated for two or three days. Now the fluid is boiled to kill the bacilli. Should the fluid display a tendency to form extensive coagula on boiling, alkali must be added. After boiling it is filtered through linen, and it is now ready to inject. With such a fluid I have been able in one series to protect eight out of nine rabbits used. In the case of the ninth, the proteid coagulated so firmly on boiling that it would not filter ; consequently the fluid transfused was almost proteid free. Cultivations in which the tissue-fibrinogen so completely coagulates on boiling that clear filtrates are obtained giving no proteid precipitate with acetic acid are useless. The injection of such solutions has no influence on the course of inoculated anthrax. I record here two observations which have been made for special purposes. ‘They will serve as illustrations for describing in detail the procedure. 1. November 10.—The watery, very feebly alkaline extract of a thymus yields on boiling an extensive coagulation. The filtrate is opalescent, and contains abundant proteid easily pre- cipitated with acetic acid. The fluid is sterilised and inoculated with anthrax. After three days in incubator, the anthrax settles as a coagulum-like mass at the bottom of the vessel. The fluid above is removed, filtered, and boiled. Two rabbits are inoculated with this fluid. Rabbit I., of 4 lbs. weight, has 30 c.c. of the solution injected into the jugular vein. Thereupon it is inoculated with blood from the heart of a guinea-pig dead of anthrax. Rabbit IT., of 24 lbs. weight, has 25 c.c. of the solution, in several portions, injected subcutaneously. Then it is inoculated with the same blood as Rabbit I. | A guinea-pig is simultaneously inoculated with the same blood. November 12.—Guinea-pig found dead. 836 PATHOLOGICAL PAPERS November 13.—Rabbit II. found dead. November 16.—Rabbit I. is quite well, and is again in- oculated with anthrax. Six months later still alive. From this it must be concluded that the subcutaneous injec- tion of the fibrinogen does not protect. 2. April 28.—The watery extract of a thymus gives a dis- tinct coagulum on boiling. After addition of water it is filtered through linen. ‘The strongly opalescent filtrate, after repeated boiling, is inoculated with anthrax. After two days in in- cubator, the culture—in which the bacilli have grown abun- dantly—is, without boiling, filtered through paper. The fluid is at first not clear, the pores of the filter soon become stopped, and the filtrate becomes clear; consequently the filter is fre- quently changed in order to prevent the filtrate becoming too poor in proteid. The collected filtrates are strongly opalescent and contain anthrax bacilli. Without previous boiling, 40 c.c. are injected into the jugular vein of a rabbit. Simultaneously it is inoculated in the ear with anthrax blood. May 1.—The animal is dead. No cedema of the ear, but cedema of skin of abdomen. Blood full of bacilli. Thirty c.c. of the solution are boiled for several minutes and filtered through linen. The filtrate is injected into the blood of a second rabbit. Immediately afterwards the animal is in-~ oculated with anthrax blood. With the exception of a very slight redness at the point of inoculation, there is no result. ‘ - — H . a ’ - : cm ed im 3 i ce F ra ‘ +e Th Cr ~ 3 “ é = - i t v bs . LU te Z ‘ 4 c _ ¥ % 3 . - ae she = % . - : os ‘ 4 LP 4 i‘ 2 A 7 + 7: er « - ee a _ ; ve oak 3 ho . ” y ‘ Sg 4 i. - ' 2 “ys . woe. = - ~ nig ‘ be ah * ie ite . - ¥ ’ 4 r 4 e - ~~ Sy 4 - . ae as noe 2 = 5 & aba Gh eo, toe. Shh AY } ) he ed, e. ’ - i P oe } & i ' ‘ ~ . \ é f . , ’ . » ON AUTO-INFECTION IN- CARDIAC DISEASE B00 fibrinogen be injected into the circulation through the jugular vein, and the femoral be then ligatured, the effect produced is - most pronounced, and is as follows: either the most extensive and rapidly developing simple cedema of the leg occurs or an enormous hemorrhage ‘ per diapedesin’ takes place throughout the tissues of the limb; or the two are combined—there is hemorrhage and cedema. The injection of fibrinogen,’ then, in addition to the obvious effects of clotting or delay in clotting, produces a totally dis- turbed relationship between the blood and the vascular wall, since, after the injection, a slight mechanical disturbance to the circulation causes a greatly increased exudation of the fluid of the blood, or this associated with a free passage of the red cor- puscles. ‘The tendency the injection has to cause hemorrhage I have already pointed out in a previous publication ;? the fact that it produces a simple but severe and sudden cedema is new. Now, to produce this altered state of the blood, leading to oedema, the same conditions of admixture of blood and fibrinogen are necessary—1.e. the admixture must be rapid. I will illus- trate this by an experiment. Exp. 8.—Used the NaCl fluid of thymus free from cells. Dog I.— Weight, 17 lbs. 12 c.c. of solution rapidly injected into the jugular. Right femoral vein tied close to Poupart liga- ment. Dog killed the next day. The portal system thrombosed. The whole right leg extremely cedematous. Large hemorrhages over upper part of leg and lower part of abdomen. Dog IIl.—17 lbs. 12 c.c. of solution injected, but ten times diluted, and injection lasting five minutes. Femoral vein tied close to ligament. Dog killed next day. No trace whatever of clotting anywhere. Leg absolutely free from the slightest trace of oedema or hemorrhage. So far as my observations go, the tendency to cedema is the first symptom of fibrinogen intoxication—1v.e. it is more easily produced than any other. 1 The fibrinogen used to produce this effect may be lymph-fibrinogen, tissue-fibrinogen, or certain varieties of blood-fibrinogen. 2 «On Hemorrhagic: Infarction of the Liver,’ Zrans. Path. Soc, 1888. [Collected Papers, p. 346. ] A A 354 PATHOLOGICAL PAPERS One of the most important features in these observations lies in their relationship to many important diseases. I have pointed out the conditions which must prevail to produce a fibrin- ogen intoxication. It is improbable that diseased conditions are often set up by a sudden large flow of lymph into the blood, but it is certain that the other condition, the slowing of the circulation in the neighbourhood of the thoracic duct, is a common incident ; particularly I may mention valvular disease of the heart and obstruction to the circulation through the lungs, as conditions which necessarily produce this result. It is a dogma of medicine that cardiac dropsy, as a symptom of cardiac failure, is due to the mechanical obstruction of the circulation. My observations lead me to the conclusion that the danger in cardiac disease is fibrinogen intoxication ; and that the symptoms of cardiac disease—e.g. dropsy, formation of intravascular clots, hemorrhagic infarction, fever, &c.—are largely dependent on this condition. PRINTED BY SPOTTISWOODE AND CO., NEW-STREET SQUARE LONDON CHEMISTRY, MEDICINE, &c. 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