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LECTURES 
 
 ON 
 
 THE COMPARATIVE PATHOLOGY 
 OF INFLAMMATION. 
 
LECTURES 
 
 ON THE 
 
 COMPARATIVE PATHOLOGY 
 OF INFLAMMATION 
 
 DELIVERED AT THE PASTEUR INSTITUTE 
 IN 189I 
 
 ELIAS METCHNIKOFF 
 
 Chefde Service a I'lnstitut Pasteur 
 
 TRANSLATED FROM THE FRENCH BY 
 
 F. A. STARLING and E. H. STARLING, M.D. 
 
 WITH 65 FIGURES IN THE TEXT AND 3 COLOURED PLATES 
 
 LONDON 
 KEGAN PAUL, TRENCH, TRUBNER & CO., Ltd. 
 
 PATERNOSTER HOUSE, CHARING CROSS ROAD 
 1893 
 
PREFACE. 
 
 The outline of the natural history of inflammation here 
 offered to the reader, is not intended as an exhaustive 
 work on the pathology of inflammatory processes. My 
 principal object in writing this book being to show the 
 intimate connection that exists between pathology and 
 biology properly so called, I have purposely omitted 
 several points — such, for instance, as the etiology of 
 suppuration, which has recently been so largely investi- 
 gated. 
 
 As the comparative anatomy of former times treated 
 only of man and the higher animals, so medicine has 
 hitherto excluded all the pathological phenomena 
 which occur in the lower animals. And yet the study 
 of these animals, affording as they do infinitely simpler 
 and more primitive conditions than those in man and 
 vertebrata, really furnishes the key to the comprehen- 
 sion of the complex pathological phenomena which 
 are of special interest in medical science. 
 
 If we examine the processes of inflammation from 
 this point of view, we shall be able to form a more 
 complete and definite idea of their real significance. 
 In the following treatise, this theoretical side is indeed 
 the only one considered ; but here again, as in so many 
 
vi PREFACE. 
 
 other cases, the extension of theoretical knowledge must 
 inevitably react upon its practical application. 
 
 The subject of phagocytes is frequently referred to 
 during the course of this study on the biological theory 
 of inflammation. I must however warn the reader 
 that he will not find a full exposition of the phagocyte 
 theory in this work. Many points in connection with 
 this question come under the headings of immunity, 
 regeneration and atrophy — points which I hope to treat 
 separately later on. 
 
 With the exception of some slight modifications, 
 these lectures are published in the form in which they 
 were delivered at the Pasteur Institute, in April and 
 May of 1891. I must, however, mention that in deal- 
 ing with the destruction of the tubercle bacillus in the 
 organism (Lecture X.), I have made use of the reactive 
 phenomena presented by the tuberculous cells of 
 Meriones instead of describing those in Spermophilus, 
 as I did in my course. Rather than repeat here what 
 has already been published in a separate paper, I have 
 preferred to introduce an example which has not yet 
 appeared in print. 
 
 In conclusion, I must express my great indebtedness to 
 Professor H. de Lacaze-Duthiers, who has supplied me 
 with valuable material for the study of inflammation 
 in the lower animals, and also to my colleagues at 
 the Pasteur Institute, Professor Duclaux and Dr. E. 
 Roux, who have spared themselves no trouble in help- 
 ing me in the preparation of this work. 
 
 Elias Metchnikoff. 
 
 Paris, January 15, 1892. 
 
PREFACE TO THE ENGLISH 
 EDITION. 
 
 I WAS much gratified when the proposal was made to 
 me of publishing an English edition of this study on 
 inflammation, the more so as in it I have endeavoured to 
 apply to pathology the principles of evolution which we 
 owe to the genius of English philosophers. I have indeed 
 dared to put forward a new theory of inflammation, only 
 because I felt that I had Darwin's great conception as a 
 solid foundation to build upon, supported as it is by the 
 doctrine of natural selection. The biological theory of 
 inflammation has already endured the test of time. It 
 was first advanced in several papers, the earlier of 
 which were published about ten years ago and have been 
 the subject of manifold attacks and criticisms. The 
 appearance of the French edition of this work, about a 
 year ago, was the signal for a renewal of the objections 
 to the theory, especially by German workers. These 
 criticisms I have endeavoured to meet in an article 
 printed as appendix to this volume, and have shown 
 that all the objections which have been brought for- 
 ward bear only on points of subsidiary importance, 
 without touching the root of the question, and indeed 
 in many cases rest on simple misunderstandings which 
 have arisen through the inadequacy of my treatment. 
 
viii PREFACE. 
 
 In replying to my critics, I have not mentioned some 
 objections which have been raised by English patho- 
 logists. The most important feature in these is the 
 idea that it is principally the eosinophile cells which 
 emigrate through the vessel-wall in inflammation ; that 
 is to say, a variety of leucocytes which never acts as 
 phagocytes. I have already replied to this statement in 
 a special article {Annales de I'lnstitut Pasteur, No. i, 
 1893), and need not repeat all my arguments here. I 
 would only observe that the leucocytes which escape from 
 the vessel in inflammation consist for the greater part of 
 true phagocytes. Special investigations, which my 
 pupils and I have carried out in my laboratory 
 on the eosinophile cells, have shown conclusively that 
 under no circumstances have the eosinophile granules 
 the microbicidal action which has been attributed to 
 them by some pathologists. These granules rather re- 
 present reserve materials, exactly similar to yolk granules 
 and aleurone grains, which are also eosinophile. 
 
 I would conclude by expressing the hope that the 
 history of the evolution of inflammation which is here 
 put forward may be found to withstand the test of any 
 further criticisms that may be directed against it ; and 
 I trust that the book in its present version may be as 
 favourably received as has been the pleasant fate of the 
 original. 
 
 Elias Metchnikoff. 
 
 Paris, April loth, 1893. 
 
CONTENTS. 
 
 PAGE 
 
 Preface ... . . . . v 
 
 Preface to the English Edition . . . . vii 
 
 LECTURE I. 
 
 Infection a struggle between two organisms — Instance of the 
 Sphserophrya — ComparativePathology abranch of Zoology 
 — Elementary principles of this science — Inflammation — 
 General survey of the principal theories in, relation to this 
 phenomenon — Present views on the subject — Necessity of 
 adopting the comparative method in the study of inflamma- 
 tion I 
 
 LECTURE II. 
 
 Are -the unicellular organisms subject to traumatic and in- 
 fectious maladies ? — Merotomy of the Amcebae and Infusoria 
 — Lesions of Vaucheria — Epidemic disease of Amoebae, 
 caused by the Microsphsra — Intracellular digestion in the 
 Protozoa — Digestion of bacteria — Epidemics in the Infu- 
 soria : disease of the nucleus and nucleolus — Division of 
 infected Paramsecia, and the means whereby they rid 
 themselves of the parasite — Acinetas — Chytridia . . 14 
 
 LECTURE in. 
 
 Plasmodium of Myxomycetes — Puncture by a glass tube — 
 Cauterisation — Chemical excitation — Trophotropism — 
 Chemiotaxis — Habituation of the Plasmodium — Negative 
 chemiotaxis — Repulsion of the plasmodium in the presence 
 of bacteria — Digestion of bacteria by the plasmodium — 
 Sensibility of the plasmodium — Fixed plants having no 
 
 a 
 
X CONTENTS. 
 
 PAGE 
 
 true intracellular digestion — Necrosis and regeneration — 
 Waldenburg's experiments — Functions of the cell-wall — 
 De Bary's observations on Peziza sclerotiorum — Tumours 
 ofplants 29 
 
 LECTURE IV. 
 Transition from the unicellular organisms to the Metazoa — 
 Sketch of the phagocytella theory — Protospongia — 
 Sponges : their organisation — their three layers — their 
 nutrition — Intracellular digestion — Ablation of parts of 
 the Sponge— Artificial division — Introduction of pointed 
 bodies — Utilisation of foreign bodies to assist in forming 
 the skeleton — Fate of the organisms which have pene- 
 trated into the interior of the Sponges— Protective function 
 of the ectoderm — Comparison with the Myxomycetes — 
 Comparison with the inflammation of vertebrata . , 42 
 
 LECTURE V. 
 
 Coelenterata, Echinodermata and Vermes — Traumatism and 
 regeneration in Hydra— Accumulation of phagocytes in 
 Acalepha (Scyphomedusffi) — Phagocytes of star-fishes — 
 Inflammation in Bipinnaria — Reactive changes in the 
 perivisceral cells of the Annelida — Phagocytic reaction in 
 the diseases of Nais and Lumbricus — Struggle between 
 the phagocytes of Lumbricus and Rhabditis — Microbic 
 infections of Worms 56 
 
 LECTURE VI, 
 
 Arthropoda, Mollusca, and Tunicata — Their vascular system 
 — Their Phagocytes — Spleen of the Gasteropoda — In- 
 flammatory reaction — Diapedesis in intact Ascidians — 
 Introduction of bacteria into the body of Ascidians and 
 Crustaceans — Infectious disease of sandhopper (Talitrus) 
 — Diseases of Daphnia — Introduction of bacteria into 
 insects — Epidemics among insects 75 
 
 LECTURE VII. 
 
 Vertebrata — Amphioxus — Embryos of Axolotl — Young larvae 
 of Urodela — Comparison with the invertebrata — Tadpoles 
 —Diapedesis— Migratory cells— Fixed cells — Phagocytic 
 properties of leucocytes — Do fixed cells also functionate as , 
 
CONTENTS. XI 
 
 PAGE 
 
 phagocytes ? — Transformation of leucocytes into fixed 
 connective tissue cells — Fate of the leucocytes that do not 
 undergo this transformation — Evolution of inflammation 
 in the organic world 94 
 
 LECTURE VIII. 
 
 Varieties of leucocytes — Origin of these varieties — Mobility — 
 Phagocytic properties — Condition of englobed microbes — 
 Their vitality and virulence — Sensibility of leucocytes — 
 Tactile sensibility — Chemiotaxis — Buchner's investiga- 
 tions — Leucocytosis — Intracellular digestion — Destruction 
 of microbes, especially in immune animals — Action of 
 leucocytes on spores — Multiplication of leucocytes by 
 direct and indirect cell-division — Changes they undergo 
 — Transformation of lobed into single nuclei . . . iii 
 
 LECTURE IX. 
 
 Endothelium of vessel walls — Their development out of the 
 mobile cells of the embryo — Development of the capil- 
 laries — Contractility of the endothelial cells— Star cells — 
 Phagocytosis in the endothelial cells — Fixed connective 
 tissue cells — Clasmatocytes of Ranvier — Ehrlich's cells 
 — Active migration of leucocytes in diapedesis — Experi- 
 ments with quinine (Binz, Disselhorst) — " Itio in partes '' — 
 Dilatation of the vessels — Theory invoking the influence of 
 the surrounding tissue — Influence of the nervous system 
 — Negative chemiotaxis of the leucocytes in cases of severe 
 infection 137 
 
 LECTURE X. 
 
 Chronic ifi/iamma/ions— Tuberculosis as a type of a chronic 
 inflammation — Phagocytic nature of tuberculous cells — 
 Destruction of tubercle bacilli by phagocytes — Power of 
 resistance of j^mt'/i^j to tubercle virus — Leprosy . .157 
 
 LECTURE XI. 
 
 Serous inflammations — Two classes of these inflammations — 
 Bactericidal power of the humours and the serous exsuda- 
 tions — Antitoxic property of serum and the serous in- 
 flammations 172 
 
Xll CONTENTS, 
 
 LECTURE XII. 
 
 PAGE 
 
 Review of other theories of inflammation in light of the ac- 
 quired facts — Nutritional theory of Virchow — Vascular 
 theory of Cohnheim — Experiments of the latter on the 
 tongue of the frog. Introduction of irritative agents into the 
 blood. Argument against Cohnheim's theory furnished by 
 the reaction in vertebrata — Struggle of the organism with 
 external agencies — Use of intracellular digestion — Phago- 
 cytes — Haemitis (recurrent fever, disease of Daphnia) — 
 Tuberculosis. Essential nature of inflammation — Sensi- 
 bility of the phagocytes — Its progressive development — 
 Sensibility of the endothelial cells — Definition of inflamma- 
 tion. Inflammation is not regeneration — Inflammation is 
 not identical with resorption — Objections raised to the 
 biological theory of inflammation — Vitalism — Teleology — 
 Absence of phagocytes in certain infective lesions — Im- 
 perfections in the inflammatory reaction — Surgical inter- 
 ference — Comparative pathology . . i8o 
 
 Description of the Plates 199 
 
 Appendix. — A Consideration of some Criticisms on 
 the BioLOGipAL Theory of Inflammation . .201 
 
 Index . . . .... 213 
 
THE COMPARATIVE PATHOLOGY OF 
 INFLAMMATION. 
 
 LECTURE I. 
 
 Infection a struggle between two organisms — Instance of the Sphae- 
 rophrya — Comparative Pathology a branch of Zoology — Ele- 
 mentary principles of this science — Inflammation — General 
 survey of the principal theories in relation to this phenomenon 
 — Present views on the subject — Necessity of adopting the 
 comparative method in the study of inflammation. 
 
 In deciding to give a few lectures on a subject belong- 
 ing to the domains of pathology, I have resolved to do 
 so solely in my capacity of zoologist. The complexity 
 of the most important pathological processes, as studied 
 according to the universal custom on vertebrates, is so 
 great, even in so low a member as the frog, that it 
 becomes impossible to analyse them or to attain any 
 adequate conception of their real significance. 
 
 It is unnecessary to cite any special proof in support 
 of the doctrine that disease and pathological processes 
 are evolved in the same way as man and the higher 
 animals themselves. In all organisms, starting from the 
 simplest forms of life, we find infectious diseases pro- 
 duced by different classes of parasites. It is therefore 
 only natural to suppose that this parasitism gives rise 
 
 B 
 
INFLAMMATION. 
 
 to a definite series of disturbances in the infected organ- 
 ism, and likewise provokes phenomena of reaction in 
 the latter. 
 
 If we examine the organisation of an animal or a 
 plant, we find that their most characteristic features are 
 their organs of attack and defence. The carapace of 
 the crayfish, the shell of molluscs and the teeth of the 
 vertebrates, as well as many other organs, are so many 
 means of protection to these animals in their perpetual 
 warfare. The mere enumeration of all the organs 
 
 acquired for the 
 purpose of helping 
 them in this strug- 
 gle would involve a 
 complete account 
 of the comparative 
 anatomy of ani- 
 mals. 
 
 Now from active 
 aggression to in- 
 fection, there is but 
 a short step. To take an instance from the lower 
 animals, we have only to consider the biological rela- 
 tions between some kinds of Infusoria. Among these 
 minute animals there is a group of suckers, which put 
 out their feelers in all directions, with the object of 
 attacking other varieties of Infusoria and of absorbing 
 their contents. (Fig. i.) Most of these Acinetae are 
 Infusoria attached to different objects which occur in 
 water, and are entirely predatory in their habits. 
 Observation of their behaviour will explain many points 
 in their organisation, and will also throw light on the 
 mode of resistance offered by the infusoria they attack. 
 
 Fig. I. — Sphizrophrya magna (after Maupas). 
 
LECTURE I. 
 
 Among these Acinetse there are some which may be 
 distinguished by their excessive minuteness, as well as 
 by the fact that, instead of being attached to some 
 object and drawing their prey towards them, they are 
 free and fasten themselves on to other infusoria larger 
 than themselves. These minute 
 AcinetEe pierce the outer covering 
 of the Infusoria they attack, and 
 take up their abode in the proto- 
 plasm of their host where they 
 lead a parasitic existence. (Fig. 2.) 
 
 We see then that organisms 
 which are very nearly related to 
 each other — in fact two members 
 of the same class {Spharophrya 
 magna and Sphcerophrya paramce- 
 ciorum) — may act as voracious 
 aggressors or as parasites with the 
 power of producing a definite in- 
 fection. Since zoological research 
 takes cognisance of the phenomena 
 of attack and defence, it should 
 likewise include the processes of 
 infection and resistance, which are 
 really in such close connection 
 with the former. The phenomena 
 
 of the active struggle among animals, however, being 
 much more prominent, have attracted the attention of 
 naturalists for years, whereas those of infection, which 
 are far less on the surface, have^been but rarely and in- 
 sufficiently studied. 
 
 A branch of zoology should be devoted to the in- 
 vestigation of the various powers of adaptation by 
 
 B 2 
 
 Fig. 2. — Spkarophrya para- 
 fnesczorui7i, 
 
 a. Two parasitic sphsero- 
 
 phryae. 
 h. A parasite adherent to 
 
 the surface of the host. 
 
4 INFLAMMATION 
 
 means of which an animal may penetrate and remain 
 in the body of other animals, as well as to the special 
 study of the reactive phenomena and mechanisms 
 which serve organisms in their resistance to parasitic 
 invasion. A branch of zoology — the comparative 
 pathology of animals — may thus be formed, which 
 would differ from the present comparative pathology in 
 many ways. Whereas the latter, which has been mainly 
 founded on veterinary science, is only concerned with 
 the higher animals and chiefly with mammals, genuine 
 comparative pathology should include the whole animal 
 world, and be treated from the widest biological stand- 
 point. 
 
 The groundwork of such a comparative pathology 
 was laid about five-and-thirty years ago. About this 
 timcj in 1857 and 1858, the theory of natural selection 
 was built up on scientific foundations by Darwin and 
 Wallace, the biological theory of fermentation by 
 Pasteur, and the theory of cellular pathology by 
 Virchow. 
 
 The first of these theories, which now forms the basis 
 of all biological research, proved the genealogical 
 evolution of organized beings, and explained the adap- 
 tation of means to an end observed in them. It de- 
 monstrated that only the characteristics which are 
 advantageous to the organism survive in the struggle for 
 existence, while those that are harmful to the individual 
 are readily eliminated by natural selection. 
 
 The biological doctrine of fermentation, established 
 by Pasteur with the discovery of the lactic ferment in 
 1857 ^"d with that of the butyric ferment in 1861, 
 showed at once in what direction the causes of infection 
 should be sought, so that the forgotten discovery of the 
 
LECTURE I. 5 
 
 anthrax bacillus was readily brought to light again by 
 Davaine, and became the starting point of pathological 
 bacteriology. 
 
 When Virchow demonstrated the important part 
 played by the cells of the organism in pathological 
 processes, a third link was formed in this chain of bio- 
 logical theories, which are indispensable to the founding 
 of a true comparative pathology. 
 
 But although the groundwork of this science was laid 
 more than thirty years ago, we are not yet in a position 
 to treat questions of general pathology from a com- 
 parative standpoint. An inquiry into the pathological 
 doctrines which are at present held concerning the most 
 important morbid processes, will readily prove the truth 
 of this statement. 
 
 As an example we will take inflammation, which is 
 universally admitted to be the most important phe- 
 nomenon in pathology. We will first examine the 
 results obtained by the usual methods of investigation, 
 and then we will proceed to inquire whether it would 
 not be more desirable to treat the question of inflam- 
 mation according to the comparative method. 
 
 As it is impossible to give a complete account of all 
 the theories on the subject of inflammation, we will 
 briefly consider those which are the most generally 
 accepted and taught by pathologists. 
 
 For a considerable time it was principally the ap- 
 pearance of inflammation — the ' rubor ' — which at- 
 tracted the attention of medical men, who were thus 
 frequently induced to regard hyperaemia as the most 
 essential act in inflammation, and even to identify these 
 two conditions. The theories concerning inflammation 
 were thus reduced to an analysis of the hyperaemia, 
 
6 INFLAMMATION. 
 
 which was supposed to be caused either by paralysis of 
 the vasomotor nerves (paralytic theory) or by a spas- 
 modic contraction of the affected arteries, accompanied 
 by an afflux of the blood to the neighbouring parts 
 (spasmodic theory). 
 
 But it was soon seen that hyperaemia of itself is in- 
 capable of producing true inflammation. A temporary 
 hypersemia, or even one of longer duration, may occur 
 without resulting in exsudation, which is an essential 
 condition in cases of typical inflammation. In order to 
 explain the ' tumor,' it was suggested that the tissues 
 at the seat of lesion had an attractive influence on the 
 blood. This view was more definitely formulated in 
 Virchow's conception of an increased nutritive and repro- 
 ductive activity of the cells at the seat of inflammation, 
 which gave rise to the formation of a large quantity of ex- 
 sudation cells at the expense of the cells of the damaged 
 tissues. According to this theory, hyperaemia would be 
 merely a subordinate and entirely secondary phenomenon. 
 
 The definite proof given by Cohnheim, that the cells 
 in inflammatory exsudations arise from the white cor- 
 puscles of the blood, first enabled pathologists to decide 
 accurately upon one of the principal questions in con- 
 nection with inflammation. After having established 
 this undoubted fact, Cohnheim adopted the opinion of 
 Samuel, according to whom the main factor in all in- 
 flammatory states consists in a lesion of the vessels 
 which are attacked by the irritating cause. The 
 inflamed vessels, being more permeable, allow the fluid 
 and corpuscular elements of the blood to flow through 
 them in a purely pas.sive manner. These exsuded 
 products collect at the part where they meet with least 
 resistance, and thus produce the inflammatory tumour. 
 
LECTURE I. 7 
 
 In this theory the tissues at the point of lesion, as 
 well as the hypersemia and the vasomotor phenomena, 
 play a very unimportant part. Although certain ideas 
 in connection with this theory have met with more or 
 less serious objections, it is accepted, especially in 
 Germany, by the majority of contemporary pathologists 
 who have formulated any general conception of inflam- 
 matory processes. Some authorities adopt it in general 
 terms, while laying stress at the same time upon the 
 importance of the phenomena occurring in the damaged 
 tissues and the vasomotor system. As it is not possible 
 to bring these conditions into connection with one 
 another, they content themselves with a simple enume- 
 ration of the changes resulting from inflammation which 
 take place in the tissues and the vascular system. 
 
 Ziegler, ^ who is the author of the best known work on 
 pathological anatomy at the present time, admits in his 
 chapter " On the Definition of Inflammation/' that he is 
 unable to give a clear definition of this condition. In 
 his own words : " The idea of ' inflammation ' includes a 
 whole series of phenomena, which occur partly in the 
 circulation and partly in the tissues, and may be com- 
 bined in various ways. As we are not here dealing with a 
 simple pathological condition, it is impossible to give a 
 short concise definition of inflammation. Even if only 
 certain phenomena, such as those which occur in the 
 circulation, were taken as characteristic of inflammatory 
 processes, a definition of them would certainly fail to 
 convey an adequate conception of inflammation." Ac- 
 cordingly, Ziegler merely gives a summary account of 
 the changes produced by inflammation. 
 
 ' " Lehrbuch der patholog. Anatomie,'' 6th edition, 1889, vol. i. 
 p. 186. 
 
8 INFLAMMATION. 
 
 Recklinghausen > holds that it is " at present impossible 
 to determine the primum movens, the starting point of 
 the changes ; that is to say, the site of the earliest 
 lesion," So he, also, is unable to do more than give 
 a detailed and careful description of inflammatory 
 phenomena. 
 
 The definition of inflammation put forward by Cornil 
 and Ranvier ^ consists simply in an enumeration of the 
 events in this pathological process. They define it as a 
 " series of phenomena, observed in the tissues or organs, 
 and analogous to those artificially produced in the same 
 parts by the action of a physical or chemical irritant." 
 
 In order to simplify this complicated question, it was 
 endeavoured to investigate this pathological condition 
 in parts of the body devoid of blood-vessels, in which 
 inflammation of the tissues only could be observed. 
 Attention was concentrated upon the cartilages, and 
 mesentery, and especially upon the cornea. From the 
 changes observed in the cells of these organs, it was 
 argued that vascular disturbances were not essential 
 for the production of inflammatory phenomena in the 
 tissues. These changes consist in proliferation of the 
 local cells, and their return to " the embryonic con- 
 dition." 
 
 Cohnheim, however, in his experiments on the cornea, 
 demonstrated vascular intervention in experimental 
 keratitis, and proved that immigration of leucocytes com- 
 ing sometimes from the margin of the cornea and 
 sometimes from the conjunctiva, took place into the 
 
 ' "Handbuch d. allgemeinen Pathologic des Kreislaufs," 1883, 
 p. 198. 
 = " Manuel d'histologie pathologique," 2nd edition, vol. i. p. 94. 
 
LECTURE I. 9 
 
 seat of inflammation. These results showed at the same 
 time that the attempts to eliminate vascular influence, 
 even in the most evascular organs of the higher animals, 
 had been quite futile. 
 
 An endeavour was then made to bring the changes 
 produced within the cells themselves into the cycle of 
 inflammatory phenomena, and Virchow's theory of par- 
 enchymatous inflammation was reinstated. The limits 
 of inflammation were thus considerably extended. 
 Brault ' and others have gone even further, and sought 
 to include under this term the acute degenerative phe- 
 nomena of the cells. 
 
 We see, therefore, that the attention of pathologists 
 has of late been mainly concentrated upon the part 
 played by the vascular system and the local tissue- 
 elements respectively. By the discovery of the pheno- 
 mena of karyokinesis, which prepared the way for the 
 solution of many problems dependent on the formation 
 and origin of the cells, a new life has been given to 
 the discussion as to whether the inflammatory cells 
 originate at the expense of the white blood corpuscles, 
 or in consequence of the proliferation of local cells. A 
 discussion on this subject has recently been raised 
 by Grawitz,' an ardent disciple of Virchow's, who 
 maintains that a large proportion of pus-globules are 
 formed from the cells of the connective tissue, and by 
 Weigert,' a faithful pupil of Cohnheim, who upholds 
 the main theory of this pathologist with regard to the 
 origin of the inflammatory cells from the leucocytes 
 
 > " Etude sur I'lnflammation," Paris, 1888, p. 34. 
 
 2 Deutsch. med. Wochensch., 1889, No. 23. 
 
 2 Fortschritte der Medicin, 1889, Nos. 15 and 16. 
 
10 INFLAMMATION. 
 
 that have escaped through the vessel walls. The dis- 
 covery of the karyokinetic phenomena enabled ob- 
 servers to ascertain, beyond a doubt, that very frequent 
 division of the local tissue-cells takes place at the 
 seat of inflammation. But whereas the partisans of 
 Virchow's doctrine regarded this as a proof of the part 
 played by these local elements in the formation of the 
 inflammatory tumour, the adherents of Cohnheim's 
 views interpreted this cell division as a simple pheno- 
 menon of reparation, serving to restore the mischief pro- 
 duced by the primary lesion. As this view became 
 more completely accepted, it gave rise to a distinction 
 of two classes of phenomena in inflammation ; first, in- 
 flammation properly so called, i.e. the lesion of the 
 vessel walls and other disturbances brought about by 
 the irritating cause ; secondly, reparation, consisting in 
 the regeneration of the missing tissues and in the 
 formation of the scar. The most advanced exponent of 
 this classification, Roser,' went so far as to assert that 
 inflammation is a true disease, due to the infection by 
 microbes, and that the reparatory phenomena con- 
 stitute its cure. According to this authority it is even 
 impossible " to give a single definition of inflammation 
 so long as this name is held to include the most hetero- 
 geneous phenomena, such as the disorders due to infec- 
 tion and the processes of recovery." 
 
 But besides this mode of interpreting inflammation, a 
 movement has been for many years travelling in the 
 opposite direction. Instead of separating inflammatory 
 phenomena into two fundamentally distinct classes, they 
 have been regarded as stages of a single process repre- 
 
 ' "Entziindung und Heilung," Leipzig, 1886, pp. 9, 11, etc. 
 
LECTURE I. II 
 
 senting a salutary reaction against some injurious influ- 
 ence. According to this doctrine, not only regeneration 
 and cicatrization but also the primary processes of 
 inflammation, such as emigration and the alteration in 
 the vessel walls, would be considered as reparatory acts 
 serving to counteract the damage brought about by the 
 irritant. This theory, which was clearly stated by L. 
 Sachs more than fifty years ago, has found fresh sup- 
 porters at different times. It was accepted by Buchner ' 
 in general terms and it has recently been developed 
 by Neumann,^ who maintains that true inflammation 
 never occurs except where there has been a primary 
 lesion of the tissues. The definition of inflammation 
 which he endeavours to introduce, is as follows. 
 " Under this name we should include the series of 
 local phenomena which are developed as a result of 
 primary lesions of the tissues (lasio continui ox necrosis), 
 and tend to cure them." ^ 
 
 From this^ review of the present state of our know- 
 ledge on the subject of inflammation, it is apparent that, 
 in spite of all the pains taken by investigators, the 
 methods hitherto adopted are inadequate for the study 
 of phenomena so intricate and variable in their mani- 
 festations. It is not to be wondered that several 
 authors, as, for instance, Thoma," should have proposed 
 to suppress the term ' inflammation •" altogether. 
 
 In spite of many attempts to simplify the experimental 
 conditions and to eliminate certain factors from this 
 
 1 " Prophylactische Therapie der Lungentuberculose," 1882. 
 
 2 " Ueber den Entzundungsbegriflf," Ziegler's Beitragt zurpatho- 
 logischen Anatomie, 1889, vol. v. p. 347. 
 
 3 Loc. cit., p. 363- 
 
 * Berliner klinische Wochewschrift, 1886. 
 
12 INFLAMMATION. 
 
 complex process, we have only succeeded in the case 
 of the elevation of temperature. By studying inflamma- 
 tion in cold-blooded animals incapable of generating 
 heat to any appreciable extent, such as frogs, it is seen 
 that true inflammation can take place in the total ab- 
 sence of heat, one of the four classical factors (dolor, 
 calor, rubor, tumor). In these animals the inflamma- 
 tory nature of the phenomena in question is so evident, 
 that no one has raised any objection to the application of 
 the term ' inflammation ' in cases where the temperature 
 is not raised and where, consequently, the word is no 
 longer applicable in its etymological sense. 
 
 The frog has always been chosen on account of the 
 facilities it presents for experimental study, without 
 realizing that in so doing a method of comparative 
 pathology was adopted. It is along these lines that 
 investigation should proceed, but the comparison 
 should be extended to still lower members of the 
 animal kingdom, in order to eliminate further factors 
 and to study these phenomena in a yet more simple 
 condition. 
 
 As we have already seen, all attempts to obtain 
 inflammation in the higher animals without the inter- 
 vention of the vessels have failed, since it is impossible 
 to exclude the circulatory system, even in the most 
 isolated tissues. In order to obtain definite results, we 
 must direct our attention to the large field presented by 
 the invertebrata, among which there are many animals 
 completely devoid of blood vessels. The comparative 
 method has already rendered good service, not only in 
 the realm of natural sciences, properly so called, but 
 even in the study of the most complicated phenomena. 
 Thus Psychology owes much to the observation of 
 
LECTURE I. 13 
 
 psychical phenomena in the lowest animals, and even 
 in the social sciences, such as ethnology or political 
 economy, investigation has often been extended to 
 the most inferior races. Pathology is almost the only 
 science in which the comparative method has been 
 ignored, although it has to do with phenomena which 
 present complications from every point of view, and it 
 would be particularly profitable to make use of such 
 methods for enlarging the scope of its investigations. 
 
 The question may be thus formulated. Do the 
 factors (traumatic or infective) which evoke the series of 
 phenomena known as inflammation in man and the 
 higher animals, produce any analogous conditions in 
 the lower vertebrata, such as Amphioxus, or in the 
 invertebrata ? Is the existence of a circulatory system 
 essential for the setting up of inflammation, or does 
 this also occur in animals which possess no blood 
 vessels, and in this case how does the nervous system 
 act ? For inflammation to take place, is it necessary 
 that the animal should possess a certain number of 
 differentiated organs or may it consist merely of an 
 agglomeration of non-diff'erentiated cells ? Do we find 
 anything analogous to inflammation in plants? Are 
 there any instances of inflammatory action in uni- 
 cellular organisms .'' In the following lectures we shall 
 discuss these questions one by one, and endeavour to 
 answer them. 
 
LECTURE II. 
 
 Are the unicellular organisms subject to traumatic and infectious 
 maladies ? — Merotomy of the Amoebae and Infusoria — Lesions 
 of Vaucheria — Epidemic disease of Amcebse, caused by the 
 Microsphasra — Intracellular digestion in the Protozoa — Diges- 
 tion of bacteria — Epidemics in the Infusoria : disease of the 
 nucleus and nucleolus — Division of infected Paramaecia, and 
 the means whereby they rid themselves of the parasite — 
 Acinetas — Chytridia. 
 
 We will first inquire whether the unicellular organ- 
 isms, which abound in the media surrounding us, are 
 subject to infectious diseases, and whether they are 
 susceptible to those influences which produce in us a 
 more or less pronounced inflammation. We shall after- 
 wards examine the changes that these influences call 
 forth in these lowly organisms. 
 
 In man and the higher animals a traumatic lesion, 
 even when insignificant, invariably provokes the series 
 of phenomena which characterize inflammation. In 
 unicellular organisms the resulting events are much 
 simpler. If we cut an Amoeba in two, there is not even 
 a wound formed along the line of section, for the edges 
 unite immediately after the passage of the instrument. 
 (Figs. 3, 4.) Two new Amoebae are thus produced ; the 
 one which encloses the nucleus continues to grow and 
 behaves in all respects like a normal individual, while 
 the other, which is without any nucleus, dies at the end 
 
LECTURE II. 
 
 15 
 
 of a longer or shorter period.^ Some other inferior 
 organisms, which contain several nuclei, as for in- 
 stance Actinophrys, can be divided into several 
 pieces, each of which is regenerated in a very short 
 time, provided that it still contain a fragment of 
 nucleus.^ In the Infusoria, which possess a more 
 highly differentiated protoplasm, artificial bisection 
 
 Fig. 3. — An amoeba immediately after 
 bisection. 
 
 rt. The half containing the nucleus «. 
 b. Half without nucleus, 
 v. Contractile vesicle. 
 
 Fig. 4. — The same amoeba five minutes 
 after the operation. 
 
 (After Bruno Hofer.) 
 
 produces a wound which lays bare the inner layer of pro- 
 toplasm. After a short time, however, the edges of the 
 peripheral layer grow over the wound, and secrete a new 
 cuticle, thus securing complete cicatrisation. (Fig 5.) 
 
 ' Bruno Hofer. " Experimentelle Untersuchungen iib. d. Ein- 
 fluss des Kerns auf das Protoplasma." Jenaisclie Zeitschr. 
 f. Naturwiss., 1889, vol. xxiv. p. 109, pi. iv. and v. 
 
 - K. Brandt. " Ueb. Actinosphasrium Eichhornii," 1887, P- 30. 
 
i6 
 
 INFLAMMATION. 
 
 These phenomena are almost exactly the same 
 whether the fragments be provided with a nucleus or 
 not. In those with a nucleus, however, regeneration is 
 complete in a very short time (often in less than twenty- 
 four hours), while the others gradually atrophy and in 
 the end always die. Balbiani,' who has published an 
 important paper on the merotomy of the Infusoria, is 
 indeed of opinion that cicatrisation is never properly 
 completed in the fragments without a nucleus, the 
 latter exercising a decided influence on the secretion 
 
 Fig. 5. — Merotomy of a Sienior. 
 
 a. Anterior fragmemt ; w» middle fragment ; fi, posterior fragment ; 7k^, in^, «z3, 7«S 
 
 stages in the regeneration of the middle fragment. 
 
 (After Balbiani.) 
 
 of the cuticle. In some species, such as Trachelius 
 ovum, the wounds caused by the section are imme- 
 diately covered over by the ectoplasm, and the separate 
 fragments that still possess a nucleus are completely 
 regenerated in less than five hours. 
 
 In the same way unicellular plants may undergo 
 severe injuries without necessarily perishing in conse- 
 quence. Thus Hanstein ^ has observed that, when a 
 
 ^ " Recherches exp^rimentales sur la m^rotomie des infusoires 
 cilies." Recueil zoologique Suisse, vol. v. 1888. 
 
 ' Vide Frank, " Die Krankheiten der Pflanzen," 1880, vol. i. 
 P-97- 
 
LECTURE ir. 17 
 
 part of the unicellular alga Vaucheria is cut or 
 crushed, it is only the damaged part that dies, while 
 the rest of the cell is healed by the secretion of a 
 cuticular layer on the injured surface, and the forma- 
 tion of a sort of sequestrum. 
 
 In these phenomena then, in the lower organisms, we 
 have to do simply with a regeneration that takes place 
 more or less completely and readily. But after trau- 
 matic lesions it is in- 
 fection that most fre- 
 quently provokes in- 
 flammation. Now, in- 
 fectious diseases are 
 very common among 
 the Protozoa and uni- 
 cellular plants. Even 
 the lowest members of 
 these classes are some- 
 times subject to infec- 
 tion. 
 
 Thus in the Amcebae 
 I have observed an Fig. 6.- 
 epidemic caused by a 
 very simple organism, which occurs in the form of 
 a round cell provided with a very delicate wall and 
 a nucleus, and capable of multiplying by division. 
 The large Amoeba with rounded pseudopodia, that 
 feeds upon diatoms, sometimes contains by the side of 
 these brown algae a small number of these round cells 
 (Fig. 6), which I shall allude to under the name of 
 Microsphizra. As the general aspect of the amoeba 
 together with its protoplasmic movements remain 
 normal, no one would suspect this rhizopod to 
 
 c 
 
 -Amoeba infected by the Microsphcera, 
 a. Early stage. 
 
i8 
 
 INFLAMMATION. 
 
 be diseased. Continued observation shows, however, 
 that whereas the enclosed diatoms undergo digestive 
 changes, the Microsphserse divide and multiply without 
 let or hindrance within the protoplasm of the amoeba. 
 This latter ejects the diatoms and becomes gradually 
 less and less active, showing that it is not in a healthy 
 condition. At the same time the protoplasm becomes 
 filled with Microsphserse, and the Amoeba invaded by 
 the parasite finally perishes. (Fig. 7.) 
 
 This case is in- 
 teresting since it 
 shows us that an 
 organism, although 
 composed almost 
 entirely of a proto- 
 plasm which has 
 the power of readi- 
 ly digesting the 
 contents of dia- 
 toms, can never- 
 theless be infected 
 by another organ- 
 ism. The infecting agent, which is to all appearance 
 insignificant, has yet the power of resisting the digestive 
 influence of the AmcEba and of bringing about its death. 
 To explain this fact, we must assume some property in 
 the parasite enabling it to produce in the interior of the 
 amoeba a substance which protects the Microsphaera, 
 and is at the same time toxic for the amoeba. 
 
 Thus the infection develops in spite of the marked 
 power of intracellular digestion possessed by the 
 Amoebae. A closer observation of the group of 
 Protozoa compels us to the conviction that this 
 
 Aronls sc- 
 Fig 7. — A dying Amceba, full of parasitic Microsphaeras, 
 
LECTURE II. 19 
 
 digestive function must play an important part in the 
 mutual relations of these lowly organisms. Many 
 Rhizopoda and Infusoria live in media swarming with 
 other unicellular organisms, including bacteria. The 
 latter, which multiply very rapidly, serve as food to 
 many of the Protozoa. Thus various Amoebae devour 
 bacilli, which undergo certain definite changes in the 
 interior of their protoplasm. Without altering their 
 shape, the bacilli acquire the power of taking up solu- 
 tions of vesuvine, which does not stain these microbes 
 when living in their natural conditions. (Fig. 8.) Since 
 
 Fig. 8. — An Amccba living ia the midst of bacilli of which 
 it has taken up a certain number. 
 
 precisely similar changes are also observed in the interior 
 of Vorticellse and other Infusoria which live on bacteria, 
 it is evident that they are due to a digestive influence 
 exerted by the contents of the Protozoa. This conclu- 
 sion is in harmony with the observation of B. Hofer ' on 
 digestion in Amoebae. This investigator has shown that 
 the more the food is altered in the interior of these Rhizo- 
 pods the more easily does it stain with the aniline colours. 
 We may often see flagellated Monads taking up 
 filaments of Leptotrix several times as long as themselves 
 (Fig. 9), and finally enclosing them in their digestive 
 vacuoles. (Fig. 10.) It is sometimes possible to follow 
 
 ' Jenaische Zeitschrift, vol. xxiv., 1889, p. 109. 
 C 2 
 
20 
 
 INFLAMMATION. 
 
 all the changes undergone by the bacteria within an 
 infusorium, as in the case of the diges- 
 tion by Stentor of the sulphobacterium 
 Thiocystis, observed by Le Dantec' 
 
 It is evident that the digestive func- 
 tion of the protoplasm of the Protozoa 
 must hinder the invasion of these 
 animals by the lower organisms, and 
 it is only in certain special cases that 
 the latter can live as parasites within 
 the Rhizopoda and Infusoria. As I 
 have already mentioned an infective 
 disease of the former class of animals, 
 I will pass to the consideration of an 
 epidemic affecting the ciliated In- 
 fusoria. 
 
 In several species of Infusoria and 
 especially in Paramsecia, attention has 
 long been called to the presence in the 
 nucleus of a number of very fine rods, 
 which J. Miiller, who first discovered 
 them, looked upon as spermatozoids. 
 These bodies were afterwards studied 
 by several observers, of whom I may 
 specially mention Balbiani and 
 Butschli, and were regarded by them 
 as parasitic bacteria. In reality they 
 are organisms quite distinct from 
 the Bacteriaceas, and belong to a 
 special group, consisting of several 
 species. Some of these develop in 
 the nucleus, replacing its contents, 
 ' " Recherches sur la digestion intracellulaire," Lille, 1891, p. 53. 
 
 Fig 9.— A Monad in the 
 process of englobing a 
 filament of Leptotrix. 
 
LECTURE II. 
 
 21 
 
 while another species attacks only the nucleolus. 
 (Fig. II.) The parasite, in its vegetative condition, 
 occurs in the form of elongated fusiform cells or rods 
 which multiply by transverse division or sometimes by 
 budding. Arrived at maturity, the parasites are trans- 
 formed into peculiarly shaped spores resembling in 
 their general appearance either bacilli or spirilla.' 
 
 In spite of the abundance of these microbes in 
 organs so important as the nucleus and nucleolus, the 
 infected Infusoria remain capable of division, when 
 they do not die from exhaustion. In the process of 
 division a certain 
 number of para- 
 sites escape from 
 the nucleus into 
 the surround- 
 ing protoplasm, 
 whence they are 
 expelled just like 
 any other indiges- 
 tible body that has 
 been swallowed 
 by the infusorium. Hafkine has shown that, if placed 
 in exceptionally favourable conditions, the Paramecium 
 may continue to divide and produce successive genera- 
 tions of infected infusoria ; at each division hov/ever, 
 the organism rids itself of a certain number of the 
 parasites, so that finally they may be all expelled and a 
 complete cure result. 
 
 Hafkine has never succeeded in producing infection 
 
 ' Vide the investigation of Hafkine, carried out at my instiga- 
 tion in my laboratory, and published in the Annales de Plnstitut 
 Pasteur. Vol. iv., 1890, p. 148. 
 
 Fig. 10.- 
 
 LeptotrLx enclosed in the digestive vacuole of a 
 Monad. 
 
22 
 
 INFLAMMATION. 
 
 by introducing Paramaecia into capillary tubes contain- 
 ing spores of the parasite ; for the Infusorium, although 
 it swallowed a certain number of the spores, surrounded 
 them with a nutrient vacuole (Fig. 12, 13), and then 
 ejected them as it would any excrementitious matter. 
 In order that a spore should germinate, it must avoid 
 
 the digestive and expulsive 
 action of the protoplasm of 
 the Infusorium, and pene- 
 trate into the nucleus or 
 nucleolus, neither of which 
 has any digestive capacity. 
 
 We see that in this case, 
 as in the disease of the 
 Amoebae, the microbe, in order 
 to infect the Protozoon, has to 
 combat the power possessed 
 by its protoplasm of ejecting 
 or digesting the parasite. 
 The same holds good in all 
 cases where we find the in- 
 vader lodged in the digestive 
 contents of an infusorium. 
 The majority of the in- 
 fectious diseases of these Protozoa are doubtless 
 caused by the parasitism of suctorial Infusoria or 
 Acinetae, which I have already mentioned in the first 
 lecture. In spite of the delicacy of their cuticle, 
 these parasites offer complete resistance to the diges- 
 tive action of the protoplasm of their hosts, although 
 in many cases the latter (as e.g. Stylonychia) are 
 distinguished by their voracity and the ease with 
 which they digest their prey. As already mentioned. 
 
 Fig. Ti. — Paramaecium with its nucle- 
 olus filled with parasites. 
 
 0, niouth ; n. nucleus ; vt, affected 
 nucleolus ; v, c. contractile vesicle. 
 
LECTURE II. 
 
 23 
 
 the young Acinetse fix themselves on to the surface 
 of other Infusoria, and penetrate into the endoplasm 
 of the latter by means of their active movements. 
 Once arrived in the central mass of protoplasm, the 
 parasites grow considerably, divide, and give rise to 
 a number (fifty or more) of young individuals, some 
 of which escape from the body of the infusorium to 
 
 attack another, after a cer- 
 tain period of freedom. 
 
 In order to survive in 
 the interior of the In- 
 fusoria, the Acinetae must 
 exercise some paralysing 
 influence on the digestive 
 function. It is probable 
 that these parasites secrete 
 
 fig. 12.— A Paraiuaecium which has taken Fig, 13.— A vacuole containing 
 
 up some spores of the parasite. spores (very highly magnified) 
 
 a. b. c. d e. f. £. spores surrounded by a 
 
 vacuole ; n. nucleus ; v. c. contractile 
 
 vesicle. 
 
 some toxic substance, since one often sees various Infu- 
 soria fall into a paralysed condition and die in con- 
 sequence of the attacks of free Acinetse.' 
 
 By their growth within the Infusoria the Acinetae 
 give rise to degeneration of the nucleus, which breaks 
 up into small round granules. In many cases, however, 
 
 1 For an account of what is known about the Acinetee, see 
 Biitschli, " Protozoa,'' in Bronn's " Classen u. Ordnungen des 
 Thierreichs," vol. iii., 1889, pp. 1823 and 1842. 
 
34 INFLAMMATION. 
 
 these parasites do not kill their hosts, which may even 
 preserve their power of multiplication. Much more 
 dangerous for the unicellular organisms are the 
 infections produced by the fungi belonging to the group 
 of Chytridiacese, which in most cases attack Infusoria 
 incapable of intracellular digestion, such as those which 
 obtain their nourishm-ent simply by diffusion. They 
 may also attack Infusoria capable of intracellular diges- 
 tion, but in this case the infection occurs while the 
 Infusorium is in an inactive or encysted condition, 
 during which time digestion does not take place. The in- 
 tracellular Chytridium, after penetrating into the interior 
 of the Protozoon, becomes round and immobile, and 
 absorbs the substance of its host, which dies, while the 
 parasite gives rise to zoospores. 
 
 Fig. 14. — Euglena viridis enclosing a Chytridium. 
 
 We will take as an example the Chytridium which so 
 often infects the Euglena viridis, and which was dis- 
 covered by Klebs.' Among these Flagellata, which 
 abound in stagnant water, we may find individuals 
 which are to all appearances in perfectly good health, 
 but enclose a round body^ provided with a nucleus 
 and a very delicate cuticle. (Fig. 14.) This foreign 
 body grows gradually larger, and divides into a number 
 of small cells which become converted into conical 
 
 ' Untersuchungen aus d. botan. Institute in Tubingen, vol. i., 
 1883. See also Hafkine, Annates des sciences naturetles : zoo- 
 logie, 1886, pp. 330, 336, &c. 
 
LECTURE 11. 
 
 25 
 
 zoospores. (Fig. 15.) The zoospores bore their way 
 out of the Euglena and escape into the surrounding 
 water. In the course of this development, the affected 
 individual presents unmis- 
 takable signs of disease. 
 The green chromatophores 
 are rapidly absorbed, and 
 the Euglena becomes highly 
 anaemic. Its contents at 
 the same time undergo ^'^- '^■'■^'tTc^lMilm. '°°"^°'"'" °^ 
 pigmentary degeneration, 
 
 evidenced by the formation of scattered brown granules 
 which become gradually more and more numerous. 
 When the parasite has attained the zoospore stage, the 
 Euglena dies in consequence of the infection. 
 
 The encysted condition, during which the Euglena 
 
 viridis is protected by 
 a capsule, seems to 
 preserve it from the 
 attacks of the Chytri- 
 dium, since this is 
 only found within the 
 mobile Euglens. On 
 the other hand, the 
 cysts of this infu- 
 sorium are often in- 
 vaded hy the Polyp Aa- 
 
 Paridorina with one cell attacked by an 
 Olpidiuni. 
 
 gus Euglencs, which 
 
 represents another genus of the Chytridiace^. 
 
 The colony-forming Flagellata are equally subject to 
 infection by the Chytridiaceas. Pandorina morum (one 
 of the Volvocinese) is often attacked by an Olpidium. 
 The presence of the latter in the body of the flagellated 
 
26 
 
 INFLAMMATION. 
 
 Monad causes a secretion of fluid, which collects to 
 form a vacuole. (Fig. i6.) The parasitic cell, which 
 is small and transparent, increases in size at the expense 
 of its host, and becomes filled with fatty granules 
 interspersed with transparent vacuoles. Soon after- 
 wards it sends out a conical process which pierces the 
 cell-wall of the Pandorina (Figs. 17, 18), and forms an 
 outlet by which the zoospores produced by the seg- 
 mentation of the contents of the parasite escape. At 
 
 Fig. 17.— Another Pandorina with five infected cells. 
 
 other times the parasite, without giving rise to zoo- 
 spores, may secrete a thick external membrane, and be 
 transformed into a cyst. 
 
 As in the case of Euglena, the invaded cell undergoes 
 pigmentary degeneration, and always ends by dying 
 and disintegrating. The neighbouring cells of the 
 colony however remain quite unaffected. They pre- 
 serve complete mobility, show pulsation of their con- 
 tractile vesicles, and divide in a perfectly normal 
 manner. (Fig. 18.) The disease and death of one or 
 even of the greater number of the sixteen members of 
 
LECTURE II. 
 
 27 
 
 the colony has absohitely no effect on the individual 
 cells which have escaped infection by the parasite. 
 
 This brief account of artificial lesions and of the in- 
 fectious diseases of the unicellular organisms, while 
 showing the inadequacy of our present knowledge on 
 the subject, at the same time enables us to appreciate 
 to some extent the general character of these phe- 
 nomena. With regard to those which are called forth 
 by the lesions, the most striking fact is the complete, 
 power of regeneration possessed by these beings. As 
 
 Fig. 18. — Part of an affected Pandorina ; and the zoosporangium 
 of the parasite a. 
 
 we have seen, a detached segment can regain its normal 
 form in a very short time, some hours or even minutes 
 after the section. 
 
 After what has been said in the first chapter, we are 
 justified in assuming generally that the relations be- 
 tween the Protozoa and the micro-organisms which 
 infect them are to be regarded in the light of a struggle 
 between two living species. The parasites are often 
 nothing else than voracious organisms which, in conse- 
 quence of their minute size, do not attack their prey 
 directly, but make their way into the bodies of the 
 Protozoa which serve them for food. This carnivorous 
 
28 INFLAMMATION. 
 
 nature of the parasites is seen not only in tlie Acinetae, 
 but also in the parasitic Flagellata which are allied to 
 the Vampirellae and other voracious organisms. In the 
 cases of infection however, the struggle assumes a more 
 complicated and indirect character. The parasite makes 
 its onslaught by secreting toxic or solvent substances, 
 and defends itself by paralysing the digestive and ex- 
 pulsive activity of its host ; while the latter exercises a 
 deleterious influence on the aggressor by digesting it 
 and turning it out of the body, and defends itself by the 
 secretions with which it surrounds itself. 
 
 Although these phenomena do not come under the 
 heading of the struggle for existence in the strictly 
 Darwinian sense (i.e. competition for the survival of 
 the fittest among individuals of the same species), yet 
 they are all more or less directly connected with the 
 struggle for survival that is always going on between 
 the representatives of the different orders of living 
 beings. In this struggle an important part devolves on 
 the power of intracellular digestion, which is so gene- 
 rally met with in the Rhizopoda and Infusoria and is 
 not entirely wanting even in the Protozoa which obtain 
 their food entirely by osmotic absorption. 
 
LECTURE III. 
 
 Plasmodium of Myxomycetes — Puncture by a glass tube — Cauteri- 
 sation — Chemical excitation — Trophotropism — Chemiotaxis — 
 Habituation of the plasmodium — Negative chemiotaxis — 
 Repulsion of the plasmodium in the presence of bacteria — 
 Digestion of bacteria by the plasmodium — Sensibility of the 
 Plasmodium — Fixed plants having no true intracellular diges- 
 tion — Necrosis and regeneration — Waldenburg's experiments 
 — Functions of the cell-wall — De Bary's observations on 
 Peziza sclerotionim — Tumours of plants. 
 
 We have now to consider the pathological phenomena 
 which occur in multicellular organisms ; and first of all 
 we meet with a group which is very important for 
 several reasons and especially interesting on account of 
 the simple organisation of its members. I refer to the 
 Myxomycetes — a group presenting both animal and 
 vegetable properties and characterised by the fact that 
 it passes through a plasmodium stage. This plasmodium 
 is composed of the largest masses of protoplasm known 
 to occur in nature. 
 
 The Plasmodium, as is well known, represents a colos- 
 sal amoeboid organism formed by the fusion of a large 
 number of zoospores of the Myxomycetes and enclosing 
 numerous nuclei embedded in a common protoplasm. 
 Branching in all directions, the plasmodium is able to 
 move about on the various objects (dead leaves, wood, 
 &c.) on which it lives ; it shows amoeboid movements 
 
30 
 
 INFLAMMATION. 
 
 at the edges of its ectoplasm, while the inner layer of 
 its protoplasm (or endoplasm) is the seat of rapid 
 currents, recalling those of volcanic lava. The Plas- 
 modium readily encloses any solid bodies within its 
 reach, and partially digests them by the help of a peptic 
 ferment and an acid which it secretes around its food.' 
 The residue as well as the indigestible bodies are turned 
 out by the plasmodium, thus forming tracks which 
 mark the places where the protoplasmic processes have 
 been. At a certain time the plasmodium produces 
 
 VJ\0USS9l 
 
 Fig. 19. — A part of the plasmodium enclosing a glass tube. 
 
 sporangia, which are usually in the form of minute 
 fruit, and enclose a number of spores provided with a 
 tough envelope. 
 
 On account of its great extent, which may amount to 
 as much as a foot or more, the plasmodium offers many 
 advantages for the study of protoplasm in general and 
 of pathological phenomena in particular. 
 
 In order to observe the effect of a lesion upon a plas- 
 modium, we may introduce a solid foreign body, such 
 
 ' The peptic ferment was discovered by Krukenberg. Unter- 
 suchungen aus dent physiol. Instit. d. U?iiv. Heidelberg, vol. ii., 
 1878, p. 273. Concerning the acid of the plasmodia, see Annales 
 de rinstitut Pasteur, 1889, p. 25. 
 
LECTURE III. 
 
 31 
 
 as a minute glass tube, into the protoplasm of a Phy- 
 sarum. The puncture by the tube tears a part of the 
 Plasmodium, which diffuses into the surrounding fluid. 
 But the chief mass of protoplasm is in no way affected 
 by the tube, which it moreover englobes after a short time 
 just as if it were a particle of food. (Fig. 19.) After 
 retaining the tube for a longer or shorter time, the 
 Plasmodium ejects it like any other substance that it is 
 unable to utilise as nourishment. 
 We may irritate the 
 
 
 Fig 20.- 
 
 Plasmodium cauterised by nitrate of 
 silver. 
 
 Plasmodium in another 
 way. If we take a 
 specimen (such as the 
 yellow Plasmodium of 
 Physarum) on an object 
 glass and touch its 
 central part with a 
 minute glass rod pre- 
 viously heated in a 
 flame, we shall produce 
 thermal excitation in- 
 stead of a mechanical lesion. Immediately after being 
 touched, the central part of the plasmodium dies and 
 may be clearly distinguished from the living peripheral 
 portions, which remain motionless as if nothing had 
 occurred, and are unaffected by the necrosed portion. A 
 few hours later however the plasmodium wakes from its 
 passive condition and creeps away from the dead part. 
 Chemical irritants operate in a still more powerful 
 way. If we apply a minute fragment of nitrate of 
 silver to the edge of a plasmodium of Physarum lying 
 on a piece of glass, and wash the injured spot directly 
 afterwards with a one per cent, solution of sodium 
 
32 INFLAMMATION. 
 
 chloride, (in order to precipitate any nitrate which may 
 have become dissolved,) we shall see that the edge 
 touched by the nitrate of silver dies and becomes 
 detached from the rest of the plasmodium. (Fig. 20.) 
 The latter reacts immediately by a rapid change in the 
 direction of its movements. Before the operation, the pro- 
 toplasmic currents were tending towards the edge where 
 the nitrate was applied (this spot having been chosen 
 for the experiment on this account) ; directly afterwards 
 
 Fig. 21, — The same Plasmodium 50 minutes after the stage 
 represented in Fig. so. 
 
 they were turned towards the sides of the plasmodium 
 and soon assumed a direction completely opposed to 
 the original one. (Fig 21.) At the end of an hour 
 from the beginning of the experiment, the plasmodium 
 had moved some distance from its first position, leaving 
 the dead residue behind. 
 
 In both of these experiments the plasmodium for- 
 sakes the portions injured by thermal or chemical 
 agencies. The much more rapid removal of the plas- 
 modium in the second case may be attributed to the 
 more powerful influence of the nitrate of silver. 
 
LECTURE III. 
 
 33 
 
 We thus see that irritating agents excite in the Plas- 
 modium either a course of events similar to those which 
 accompany the taking up of any solid nutriment, or else 
 a more or less marked repulsion. In attempting to 
 produce a reaction which should correspond to the con- 
 dition of inflammation in the higher animals, we have 
 brought about the phenomena of attraction or repulsion 
 which occur so frequently in the lives of plasmodia and 
 the inferior animals generally. 
 
 As early as 1 884, Stahl ' discovered that a decoction 
 of dead leaves (on which so many of the Myxomycetes 
 feed) has the property of attracting the plasmodia, 
 whereas solutions of salt, sugar, and numerous other 
 materials, act in a contrary manner, repelling the 
 plasmodia to a greater or less distance. In connecting 
 these phenomena with those of nutrition, Stahl gave 
 them the name o^ positive trophotropism when it was a 
 question of attraction, and of negative trophotropism in 
 cases of repulsion. Pfeffer,^ having found that the female 
 organs of certain cryptogams (ferns, mosses,- and 
 selaginellae) attract the spermatozoids with a different 
 object from that of nutrition, grouped all these various 
 forms of sensibility to chemical agents under the general 
 heading of chemiotaxis (positive or negative), a term 
 which was very soon generally adopted. 
 
 Since, as we have just shown, these phenomena of 
 sensibility play an undoubted part in pathological pro- 
 cesses, it is desirable to consider them in somewhat 
 greater detail. Chemiotactic phenomena are met with 
 not only in the Myxomycetes and the spermatozoids of 
 
 1 Botanische Zeiiung, 1884, Nos. 10-12. 
 
 2 Untersuchungen aus d. botan. Institute in Tubingen, vol. i. p. 
 
 363- 
 
 D 
 
34 INFLAMMATION. 
 
 the above-mentioned cryptogams, but also in Bacteria, 
 Flagellata, Volvocinpse/ and in the zoospores of fungi, 
 such as the Saprolegniaceae.^ Hence it is evident that 
 we have here to do with a phenomenon of general 
 import. 
 
 It cannot be denied that by means of positive chemio- 
 taxis the organisms are directed in their search after 
 nutrient substances, and are enabled to approach the 
 bodies with which they have to establish relations, as 
 in the case of the spermatozoids attracted by an ovum. 
 By means of negative chemiotaxis, on the other hand, 
 they can escape injurious influences. This rule, 
 although true generally, cannot be applied to every 
 particular case. Thus Pfeffer^ has seen spirilla and 
 animalcules (' bodons ') dart into too highly concentrated 
 solutions of sugar and glycerine, to which attracting sub- 
 stances had been added, where they inevitably perished. 
 
 The analogy between these phenomena and the sen- 
 sations of man and the higher animals is obvious. One 
 among many proofs of this analogy consists in the fact 
 that the chemiotaxis of the lower animals is subject to 
 the same law of Weber which has been established for 
 the sense perceptions of man. In order that a bacterium 
 {B. termo) or the spermatozoids of ferns (the organisms 
 on which Pfefifer has made his remarkable observations) 
 may be affected by a change in their surrounding media, 
 it is essential that this change should attain a certain 
 degree. Thus the Bacterium termo placed in a solution 
 of peptone of definite strength, will not move towards a 
 
 ' Untersuchungen aus d. botan. Inst, in Tubingen, vol. ii. i888, 
 p. 582. 
 
 2 Botanische Zeitung, 1890, Nos. 7-11. 
 
 ' Unters. a. d. botan. Inst, in Tubingen, vol. ii. p. 627. 
 
LECTURE III. 35 
 
 more concentrated solution of peptone, until this is five 
 times as strong as the first solution. Having ascertained 
 these proportions, Pfeffer formulated for the chemiotaxis 
 of these unicellular bodies the same law as for the sense 
 perceptions of man, viz. that when the excitation is 
 increased in geometrical proportion, the sensation is in- 
 creased in arithmetical proportion, or, in other words, that 
 the reaction varies as the logarithm of the excitation. 
 
 Now although the two orders of phenomena conform 
 to the same law, there is quantitatively a great difference 
 between them. Man can appreciate a difference of 
 weight equalling one-third, of temperature amounting to 
 one-thirtieth, of light equivalent to one-hundredth, where- 
 as the spermatozoids of ferns are not affected by and do 
 not react to an alteration in the chemical composition, 
 until the quantity of the substance which is acting on 
 them is augmented twenty-nine times. The Bacterium 
 termo is indifferent to an increase of concentration until 
 this amounts to four times the original concentration.' 
 
 In order to test the chemiotactic sensibility of the 
 plas^iodium, I placed several specimens of the Plasmo- 
 dium of Didyntium farinaceum in O'l, O'Oi, 0-05, 0"005, 
 and 0*0005 P^i' cent, solutions of hydrochlorate of 
 quinine. The last two solutions did not prevent the 
 Plasmodium from advancing or even from inserting 
 several processes into them ; whereas the first three 
 solutions exercised a pronounced negative chemiotactic 
 effect. (Plate II., figs. 3-6.) The plasmodium therefore 
 can appreciate the difference between 0*05 and 0005 
 per cent, of hydrochlorate of quinine. 
 
 The Plasmodium, in common with other of the lower 
 organisms^ has the important power of growing gradually 
 
 ' Pfeffer, loc. cit., vol. ii. p. 637. 
 D 2 
 
36 INFLAMMATION. 
 
 accustomed to solutions which in the beginning it 
 avoided. Stahl was the first to notice that the Plasmo- 
 dium of Fuligo, which is at first repelled by a solution 
 of sea-salt in the proportion of 2 per cent, or less, 
 after having been deprived for some time of water 
 finally adapts itself to the changed conditions and dips 
 its processes into the salt-water. Here we have an 
 instance of negative chemiotaxis which, as the result of 
 imperceptible alterations in the protoplasm, is converted 
 into positive chemiotaxis. 
 
 As this fact is of great importance from a general 
 point of view, I was desirous of watching the process 
 myself. For this purpose, I placed a plasmodium of 
 Physarum extended on a glass slip into a vessel con- 
 taining a solution of 0"S per cent, sodium chloride. 
 The Plasmodium immediately showed negative chemio- 
 taxis and moved away from the surface of the liquid. 
 It was then changed to another vessel containing a 0"25 
 per cent, solution of the same salt. The plasmodium 
 was at first repelled, but after the lapse of a few hours 
 it drew near the liquid into which it then immersed the 
 end of its processes. With the view of noting how far 
 this power of adaptation extended, I replaced the 
 Plasmodium in the first vessel with the 0"S per cent, 
 salt solution. It again receded from the fluid ; but at 
 the end of about twelve hours it approached the surface 
 of the water, without however touching it. 
 
 By means of negative chemiotaxis, therefore, the 
 Plasmodium is enabled to avoid injurious influences ; 
 as we have seen, it recedes from bodies which burn it, 
 such as the nitrate of silver, and even from necrosed 
 portions of its own organism, as in the experiment of 
 the application of a heated object. It is probable that 
 
LECTURE III. 37 
 
 the same property may serve to protect the plasmodium 
 against the attack of other organisms, especially of 
 pathogenic microbes. 
 
 Stahl has observed that plasmodia are never attacked 
 by parasites. This fact he seeks to explain by referring 
 it to the facility with which the plasmodia move about 
 from place to place, as well as to their power of 
 expelling foreign substances — a property which is con- 
 nected with the intracellular digestion of solid bodies. 
 Although no direct observations on the expulsion of 
 parasitic organisms by the Plasmodium have so far been 
 made, yet it is extremely probable that this occurs, 
 especially as Pfeffer' has watched the plasmodia of 
 Chondrioderma ejecting living Pandorinae and Diatoms. 
 Moreover the direct observations on the expulsion of 
 parasitic spores by the Paramaecia tend to support 
 Stahl's deduction on this subject. 
 
 The following experiment was made with the object 
 of ascertaining the true significance of the movements 
 of the Plasmodium. I spread out a plasmodium of 
 Physarum on a slide, and placed it midway between 
 two small glass vessels, one of which was filled with a 
 stagnant infusion of dried leaves, full of bacteria, 
 infusoria and other of the lower organisms, the other 
 with the same infusion after filtration through several 
 layers of filter-paper. The two ends of the plasmodium 
 were connected with the liquid in the two vessels by 
 strips of blotting-paper. The plasmodium soon began 
 to approach the filtered liquid, moving along the strip 
 which was soaked in it. Another experiment made in 
 
 > "Ueber Aufnahme und Ausgabe ungeloster Korper." Ab- 
 handlungen d. math. phys. Klasse der k. sacks Gesellsch. d. Wissen- 
 schaften,\o\. xvi. 1890, p. 161. 
 
38 INFLAMMATION. 
 
 the same way, with a few slight modifications, was 
 attended by exactly the same result, showing that the 
 Plasmodium preferred the liquid which was free from 
 microbes. In order to ascertain how far this preference 
 extended, I repeated the experiment, only exchanging 
 the filtered fluid for a fresh infusion of dead leaves in 
 cold water, which was consequently colourless. This 
 time the plasmodium advanced towards the stagnant 
 infusion, in spite of the microbes which it contained. 
 
 The repulsion of the Plasmodium in the presence of 
 the lower organisms is evidently merely relative ; this 
 is in harmony with the fact that the Myxomycetes in 
 their amoeboid condition have the power of englobing 
 microbes. Saville Kent has observed amoeboid 
 zoospores of Physarum tussilaginis which were filled 
 with bacteria. Later on Lister' made some highly 
 interesting researches on the inclusion of bacteria by 
 the zoospores of different Myxomycetes. The bacteria, 
 seized on by the pseudopodia, are dragged into the 
 interior of the amceboid plasma and lodged in the 
 nutritive vacuoles. Here they become less and less 
 distinct, and appear to be almost completely dissolved. 
 A zoospore of Chondrioderma difforme will completely 
 digest a couple of large bacilli in about an hour and a 
 half. 
 
 The powers of digestion and expulsion possessed by 
 the Plasmodium, combined with its property of negative 
 chemiotaxis, are of real service to it in reacting against 
 injurious excitants. 
 
 The Plasmodia, in common with a considerable num- 
 ber of other lowly organisms, are endowed with several 
 forms of sensibility besides that known as chemiotaxis. 
 
 ' Journ. of the Linnaan Soc, Botany, vol. xxv. i8go, p. 435. 
 
LECTURE III. 39 
 
 They avoid sunlight, and are powerfully attracted to the 
 damper places, thus evincing a kind of hydrotropism. 
 Moreover, this positive hydrotropism is converted into 
 negative hydrotropism just before the period of fructifica- 
 tion, when the plasmodium seeks a dryer spot (Stahl). 
 The Plasmodia are also endowed with tactile sensi- 
 bility. 
 
 Fixed multicellular plants, which have no powers of 
 locomotion or of digesting and excreting foreign bodies, 
 react to the various injurious factors with which they 
 may come in contact in a different manner from the 
 Plasmodia, which can either avoid dangerous objects 
 or can eliminate them by digestive or excretory pro- 
 cesses. 
 
 A thorn introduced' into the plasmodium is dealt with 
 like any other foreign body that becomes enclosed in the 
 amcEboid mass. As it cannot be digested, it is simply 
 ejected altogether. If introduced into the tissue of a 
 plant, the same thorn would produce a lesion of the cells, 
 and the injured cells would inevitably perish. The 
 damage is soon repaired with the help of the neighbour- 
 ing cells, which rapidly multiply and form either a mass 
 of cork or a true scar composed of several tissues.^ In 
 both cases there is active cellular neo-formation at the 
 injured spot. Waldenburg,^ who has studied these 
 phenomena from the point of view of their analogies 
 with inflammation in the higher animals, says : " Plants 
 may therefore suffer from inflammation, if by this 
 term we imply only the lesions produced by the irri- 
 tant, as well as the tumour induced by these lesions, 
 
 ' Frank, " Die Krankheiten der Pflanzen." Breslau, 1880, vol. i. 
 
 p. 95, etc. 
 ' Vircha-ufs Archiv, vol. xxvi. 1863, pp. US and 322, Tab. v. 
 
40 INFLAMMATION. 
 
 and leave out of account the blood-vessels and nerves " 
 (p. 344). Inflammation would thus be only an irrita- 
 tion of the tissues (tumefaction, growth) plus a vascular 
 congestion. 
 
 The phenomena of repair in plants have on more 
 than one occasion been quoted in support of the 
 attraction theory of inflammation, and especially in 
 favour of Virchow's theory of a nutritive and formative 
 hyperactivity of the inflamed tissues. But unfortu- 
 nately no account has been taken of the conditions 
 intermediate between plants and the higher animals^ 
 and consequently the most characteristic phenomena of 
 true inflammation have been lost sight of altogether. 
 
 The new cells produced at the seat of injury in 
 plants frequently secrete thicker and tougher cell- 
 walls than usual. The cuticle in fact represents the 
 true protective organ of the plant. The instance 
 quoted in the preceding lecture of a Chytridium which 
 only attacks the Euglena in its mobile and never in its 
 encysted condition, may be taken as a confirmation of 
 this statement. The walls of the vegetable cells are 
 too resistant for many of the microbes, and especially 
 for those which cannot penetrate actively into the 
 tell-contents. This is probably the reason why 
 bacterial infection so rarely takes place in plants. On 
 the other hand, vegetable organisms are very liable to 
 be attacked by fungi, which grow rapidly and in many 
 cases secrete a diastase which dissolves the cellulose 
 cuticle of the plant. The parasitic fungus, once within 
 the cell, absorbs the contents without further hindrance. 
 The cells that are thus invaded die, while the others 
 that remain alive, undergo rapid proliferation which 
 results in the overgrowth of parts, amounting in some 
 
LECTURE III. 41 
 
 cases to a hypertrophy of the entire organism affected, 
 as in the case of Euphorbia Cyparissias when infested 
 by the secidium of Uromyces Pisi. The presence of a 
 parasite, whether it be a fungus or a member of the 
 animal kingdom, often excites the formation of special 
 tumours or galls. 
 
 As in the healing of wounds, infections ' in plants are 
 accompanied with regenerative phenomena, dependent 
 on the rapid proliferation of the cells that are not directly 
 affected. We do not however find processes analo- 
 gous to the essential phenomena of inflammation in the 
 vegetable world. For these we must pass to the con- 
 sideration of the representatives of the animal kingdom. 
 
 ' The most carefully recorded instance is undoubtedly that of 
 the Peziza sclerotiorum on which De Bary {Bofanische Zeitung, 
 1886) made his classical observations. This fungus germinates on 
 the surface of the plant and sends out filaments with which it 
 subsequently penetrates the tissues. (The Peziza sclerotiorum 
 invades a large number of plants.) In order to effect an entry, 
 the parasitic filaments secrete oxalic acid and a ferment which 
 dissolves the cellulose. The parasite feeds on the juice of the cells 
 which have perished under the influence of its secretions, and its 
 mycelium grows into the interstices between the cells, rarely 
 penetrating the latter. De Bary has observed that the Peziza 
 easily finds its way into young plants, but is unable to force an 
 entrance into older plants of the same species. This immunity is 
 probably owing to the fact that the parasite cannot dissolve the 
 cellulose of the old cells. The control experiments showed in fact 
 that the extracted juice of the fungus was able readily to digest the 
 walls of the young cells and was quite without effect on those of 
 the older individuals of the same plant. 
 
 It is evident that the power of resistance possessed by the 
 vegetable cell is specially dependent on the toughness of its 
 cuticle. The parasite in order to infect the plant must first 
 perforate or dissolve this membrane. 
 
LECTURE IV. 
 
 Transition from the unicellular organisms to the Metazoa — Sketch 
 of the phagocytella theory — Protospongia — Sponges : their 
 organisation — their three layers — their nutrition — Intracellular 
 digestion — Ablation of parts of the Sponge — Artificial division 
 — Introduction of pointed bodies — Utilisation of foreign bodies 
 to assist in forming the skeleton — Fate of the organisms which 
 have penetrated into the interior of the Sponges — Protective 
 function of the ectoderm — Comparison with the Myxomycetes 
 — Comparison with the inflammation of vertebrata. 
 
 In passing to the animal kingdom, we have to confess 
 that we are at present ignorant of the way in which the 
 multicellular animals or Metazoa are derived from the 
 Protozoa. The gap between the most highly developed 
 members of the latter class and the lower kinds of 
 Metazoa is too wide and can only be bridged over by 
 hypotheses based on the embryological study of different 
 animals. Putting aside several groups of parasites 
 which have undoubtedly lost much of their primitive 
 character {Dycyemides, Orthonectides) , we find that even 
 the simplest forms of the Metazoa, such as the Sponges, 
 are composed of a multiplicity of organs, arranged in 
 three well-known layers : the ectoderm, the mesoderm 
 and the endoderm. In order to obtain a knowledge of a 
 more simple condition of animal life, we must turn to the 
 embryos of Sponges and of other inferior organisms, such 
 as the Medusae and their allies. Here we may readily 
 
LECTURE IV. 43 
 
 meet with stages in which the animals are composed of 
 two layers, one of which constitutes an enveloping 
 membrane for the larva, the other is formed by the inner 
 cells, grouped in a different manner. These latter cells are 
 sometimes collected into a solid mass, a kind of paren- 
 chyma, composed of amoeboid elements ; or they may 
 be arranged evenly to form an epithelial layer sur- 
 rounding a digestive cavity. The question as to which 
 of these two forms may be considered the more primi- 
 tive has been much discussed. I think that the 
 absence of a digestive cavity, the irregular shape of the 
 cells, together with various other facts afforded by the 
 comparison of the embryogeny of many lower animals 
 (which do not come within the range of this pathological 
 survey), lead to the conclusion that the parenchymatous 
 stage is the more primitive of the two. I have called 
 this stage by the name of Phagocytella^ on account of 
 the power which the cells of the inner layer possess of 
 englobing various solid bodies, and more especially for 
 the reason that the digestive cells of the whole organism 
 are derived from this layer. The latter in the first 
 place produces the endoderm which lines the intestinal 
 canal and its appendages, and secondly the whole or a 
 part of the mesoderm which also includes a large 
 number of digestive cells ox phagocytes. 
 
 The phagocytella stage may readily pass into the 
 Gastrula stage, ^ which has two epithelial layers, one 
 of which represents the wall of the primitive intes- 
 tine that opens by a primordial orifice or blastopore. 
 
 ^ Cf. my account of the Phagocytella theory in my " Embryolo- 
 gische Studien an Medusen," Wien, 1886. 
 
 2 For an account of the Gastrasa theory, see Hseckel " Gastrsea- 
 Theorie." Jena, 1874. 
 
44 
 
 INFLAMMATION. 
 
 This gastrula is so to speak the starting-point for all 
 the Metazoa. 
 
 The structure of the Metazoa when reduced to its 
 simplest condition, that of the phagocytella, is some- 
 what analogous to that of certain colony-forming 
 Protozoa. In these cases the colonies are made up of 
 two kinds of individuals : the flagellated individuals 
 
 Fig. 22. — Protospangia HtEckelii (after Saville Kent). 
 
 forming a sort of outer layer (Fig. 22), and 
 
 the 
 
 amoeboid individuals, situated in the inner mass of the 
 colony. The former would thus correspond to the 
 elements of the ectoderm, which so frequently consists 
 of flagellated cells, and the latter would constitute a 
 kind of inner parenchyma, composed of amoeboid cells, 
 which are at the same time phagocytic in their nature. 
 In these colonies of infusoria, termed Protospongia by 
 
LECTURE IV. 45 
 
 their discoverer Saville Kent,^ the two layers are not 
 yet clearly defined, since the individuals of which they 
 are made up pass readily from the one form into the 
 other. 
 
 By means of these colonies on the one hand and of 
 the organisms resembling the phagocytella on the 
 other, we are almost in a position to bridge over the 
 gap between the Protozoa and the Metazoa. 
 
 I should not have dwelt upon these hypotheses in 
 these lectures on the comparative pathology of inflam- 
 mation, did not their consideration afford an opportunity 
 of studying the general significance of the presence of 
 these amoeboid cells which are able to englobe solid 
 bodies. We have met with them in the various classes 
 of Protozoa, and we find them again even in the most 
 primitive forms of Metazoa. Now the co-operation of 
 amoeboid cells in inflammation as it occurs in the 
 vertebrates is a fact of the greatest importance, which 
 has received almost universal acceptance. 
 
 As low down in the animal scale as the most inferior 
 Metazoa we have to do with these cells. The Sponges 
 are of such undifferentiated organisation that they were 
 long considered to be colonies of Protozoa, consisting, 
 like the Protospongia, of separate flagellated and 
 amoeboid individuals. Later on it was however ascer- 
 tained that they bore a certain relationship to the 
 Polyps and their allies (Ccelenterata). It was then 
 found that they are formed of three characteristic layers. 
 The outer layer or ectoderm covers the whole structure 
 with flat epithelial cells, the contours of which are very 
 clearly defined after the application of a solution of 
 nitrate of silver. The cells themselves are evidently 
 1 " The Manual of Infusoria," 1880-1882. 
 
46 INFLAMMATION. 
 
 contractile — a property which is more readily observed 
 at the free edges of the young cells, where amoeboid 
 prolongations belonging to the ectodermic elements are 
 seen. The contractility of these cells is certainly con- 
 cerned in the remarkable phenomenon of the opening 
 of the numerous pores which are scattered over the 
 whole surface of the Sponge and are bordered by two or 
 more flat cells. These pores open to permit of the 
 passage of a stream of water with the minute particles 
 which it holds in suspension. The liquid first enters 
 a system of efferent canals which are also lined by a 
 pavement epithelium, the origin of which is not yet 
 known. It then passes into canals or into round cavities 
 (the ' ciliated chambers ') which are covered with a cylin- 
 drical epithelial layer, the cells of which possess a single 
 large flagellum. These cells, which present a striking 
 analogy to many flagellated infusoria, form part of the 
 endoderm and represent typical phagocytes, since they 
 attract and englobe a large number of fine granules 
 which are carried along by the stream. 
 
 Besides these flagellated phagocytes of endodermic 
 origin, the Sponges contain a large number of mobile 
 cells, which are typical amoebae and form part of the 
 mesoderm, being situated between the ectoderm and 
 the cylindrical epithelium. Although it has not so far 
 been definitely ascertained how the foreign particles 
 penetrate the mesoderm after they have reached the 
 interior of the Sponge, yet it has been clearly shown 
 that they are largely absorbed by the mesodermic cells 
 themselves. If a coloured substance, such as carmine 
 indigo or sepia, be added to the water in which the 
 Sponges are immersed, it will be found soon afterwards 
 that many of the grains of colouring matter have been 
 
LECTURE IV. 47 
 
 enclosed by the endodermic cells, as well as by the 
 amoeboid phagocytes of the mesoderm. 
 
 In certain Sponges (as for instance in several cal- 
 careous Sponges,) there are very few mesodermic cells, 
 which consequently take but a small part in englobing 
 foreign bodies ; in others again, especially in the 
 silicious kinds, the mesoderm is much better developed, 
 and its more numerous cells can therefore take in a 
 proportionately larger number of these minute particles. 
 There are a few species, such as the Siphonochalina 
 coriacea, whose mesodermic cells alone enclose all 
 foreign bodies, so that the cylindrical cells of the 
 endoderm merely serve to keep up the continuous 
 passage of the fluid through the Sponge. The phago- 
 cytes of both layers have the power of rejecting in- 
 soluble matters, which collect in the larger efferent 
 canals and are expelled from the organism through 
 large apertures of crater-like shape, the walls of which, 
 according to some authors, are furnished with muscular 
 fibres. 
 
 We are however chiefly concerned here with the fact 
 that the mesodermic phagocytes are able to digest the 
 substances as well as to englobe them, and to reject the 
 insoluble residue. Many years ago Lieberkiihn ' ob- 
 served the digestion of Infusoria which had found their 
 way into the mass of amoeboid cells of soft-water 
 Sponges, and pointed out the analogy of this pheno- 
 menon with the digestion of Infusoria by the Rhizopoda 
 or other Protozoa. This has been confirmed by 
 other investigators. Thus I have seen- Oxytricha, 
 Glaucoma, and Actinophrys dissolved in the midst of 
 
 ' Muller's Archiv fur Anat. und Phys., 1857, p. 385. 
 ''■ Zeitschr. f. wissench. Zoologie, vol. xxxii. 1879, p. 371. 
 
48 
 
 INFLAMMATION. 
 
 a mass of the mesodermic phagocytes of a young 
 Spongilla, and afterwards the foreign bodies which had 
 been swallowed by these Protozoa englobed by the 
 same phagocytes. The Euglenae, when sucked into the 
 Spongillse by the current, become likewise surrounded 
 by the mesodermic phagocytes ; but whereas their 
 protoplasm is then digested, the grains of chlorophyll 
 and paramylum remain intact for an indefinite time. 
 
 Fig. 23. — A mesodermic phagocyte of a young Spongilla surrounded by several 
 ectodermic cells. 
 
 The mesodermic cells of young Spongillae, immedi- 
 ately after their escape from the gemmules, can enclose 
 foreign bodies even before the endoderm is developed. 
 The young Sponge at this stage consists only of a layer 
 of flat ectodermic cells and an irregular mass of mesoder- 
 mic cells, a certain number of which soon begin to 
 secrete spicules. If carmine grains be placed in the 
 
LECTURE IV. 49 
 
 water inhabited by the Spongillae, they find their way in 
 without apparently injuring the wall, and are at once 
 englobed by the amoeboid phagocytes of the mesoderm. 
 
 (Fig- 23.) 
 
 The above facts are so invariable in their occurrence 
 and may be so readily observed, that it is very extra- 
 ordinary that de Lendenfeld,' in his monograph on the 
 physiology of Sponges, should seek to throw doubt on 
 the matter. This author asserts that carmine, when 
 added to the water which filters through the Sponges, is 
 very rarely deposited in the amoeboid cells and then 
 only where there is some lesion of the outer layer of 
 cells, and that in a normal Sponge it is only the 
 cylindrical cells of the endoderm which will seize on the 
 carmine. De Lendenfeld lays stress on these con- 
 clusions, in spite of the fact, which he acknowledges, 
 that the fat-globules of milk are readily absorbed by 
 the mesodermic phagocytes. This is a sufficient proof 
 of the part taken by these cells in the intracellular 
 digestion of Sponges. Moreover his memoir con- 
 tains direct indications of the presence of carmine 
 grains in the amoeboid cells of the mesoderm. Thus 
 this author describes that he has seen numerous carmine 
 grains in the phagocytes of Chondrosia reniformis, the 
 species on which he has bestowed the most study. 
 Further, he has even found them in these cells two 
 hours and a half after the introduction of the carmine 
 into the water, at a period when the cylindrical elements 
 of the chambers no longer contained any granules. 
 
 Since these facts remove all doubts that the amoe- 
 
 1 " Experimentelle Untersuchungen iiber die Physiologie der 
 Spongien," Zeitschr. f. wissensch. Zoologie, vol. xlviii. 1889, p. 
 
 406. 
 
 E 
 
so INFLAMMATION. 
 
 boid cells of the mesoderm of the Spongillse have the 
 power of taking in and digesting foreign particles, I 
 have endeavoured to ascertain the conditions which 
 might influence this function. Following up the know- 
 ledge we have acquired concerning the typical in- 
 tracellular digestion in the Protozoa and the Myxomy- 
 cetes, which secrete around the object they have en- 
 globed an amount of acid sufScient to convert the 
 colour of litmus from blue to red, I placed a few grains 
 of blue litmus into the water containing some young 
 
 Fig- 24. — A glass tube surrounded by the mesodermic phagocytes of the Spongilla. 
 
 Spongillae which had developed from gemmules. Like 
 
 most minute particles suspended in the water, these 
 
 grains were soon englobed by the Sponges and were 
 
 found to have been chiefly taken up by the mesodermic 
 
 phagocytes. The litmus however did not change 
 
 colour, even after a prolonged stay in the cells ; hence 
 
 it is evident that the digestion of the Spongillae 
 
 cannot proceed in an acid medium. This fact is in 
 
 complete harmony with Krukenberg's discovery ^ of a 
 
 tryptic ferment in the glycerine extract of several 
 
 varieties of Sponges. 
 
 ' " Grundziige einer vergleichenden Physiologic der Verdauung." 
 Heidelberg, 1882, p. 52. 
 
LECTURE IV. 
 
 51 
 
 If now we introduce a sharp substance, such as a 
 minute glass tube or a spicule of asbestos, into a Sponge, 
 the greater part of it will be situated in the mesoderm, 
 where it will be in immediate proximity to the amoe- 
 boid cells. The latter surround the foreign body either 
 partially or completely, i.e. they react as if the sub- 
 stance were merely a nutritious body of larger size than 
 
 Fig. 25. — Vegetable filament surrounded by the phagocytes of a Spongilla. 
 
 usual (Fig. 24). Sometimes the cells accumulate very 
 sparsely or not at all round the article introduced, 
 showing that too weak a reaction has been excited. 
 On other occasions inert bodies, such as vegetable 
 filaments, will attract a considerable number of phago- 
 cytes which surround them and become partially fused 
 into small plasmodia (Fig. 25). 
 
 In some members of this group the grains of sand 
 and of other hard materials, which they happen to have 
 
 E 2 
 
52 
 
 INFLAMMATION. 
 
 taken up, become surrounded with a mass of spongine, 
 secreted by the mesodermic cells. In such cases the 
 foreign particles are utilised by the Sponge to increase 
 the solidity of its skeleton. 
 
 As I have remarked above, the mesodermic cells can 
 also enclose living organisms which have penetrated 
 into the interior of the Sponge, where they subse- 
 quently undergo digestion by the phagocytes. Organ- 
 
 Fig. 26.— Leptotrix surrounded by the phagocytes of the Spongilla. 
 
 isms of greater resisting powers may escape this fate 
 and may remain for a shorter or longer time within the 
 body cavity of the Sponge without undergoing any 
 alteration whatever. Thus I have seen filaments of 
 Leptotrix living uninjured inside the mesoderm of 
 young Spongillse, and surrounded by a plasmodium 
 formed by the fusion of a number of mesodermic cells 
 (Fig. 26). In certain Sponges {Hircinia echinata and 
 Ceraochalina gibbosa) Keller has found the eggs of 
 Annelida and Crustacea developing undisturbed in the 
 
LECTURE IV. 
 
 S3 
 
 mesoderm, and surrounded by masses of amoeboid 
 cells which formed a regular follicle round them (Fig. 
 27). 
 
 These facts show that any foreign body, which has 
 by some means or other reached the parenchyma of 
 the Sponges, excites the mesodermic phagocytes, which 
 either englobe them, or collect in a mass or even fuse 
 together in large numbers round them. If the foreign 
 bodies are easy to digest, they are soon dissolved ; if 
 however they prove resistant, they remain in the in- 
 
 VJfioirf 
 
 Fig. 27. — Crustacean ova surrounded by the phagocytes of the Ceraosjtoneia, 
 (After Keller.) 
 
 terior ot the Sponge, surrounded by the cells. This 
 phenomenon, which frequently occurs among the 
 Spongiaria, may be regarded as an instance of a kind 
 of commensalism. These soft cellular organisms, 
 being readily penetrated, are very suitable as an abode 
 for many aquatic animals on account of the continual 
 stream of water which passes through them, bringing 
 nourishment to their guests. Hence there are a large 
 variety of the latter, from the Algae {Zoochlorellm and 
 Zooxantellce) which inhabit the interior of the meso- 
 dermic cells, to the Polypi (Stephanoscyphus), Annelida 
 and Crustacea, which take refuge in the canals and 
 
54 INFLAMMATION. 
 
 parenchyma of the Sponges. So far, real parasites, 
 with the infectious diseases they bring in their train, 
 have not been discovered. It may be that the phago- 
 cytes are so effective as to utterly destroy the microbes 
 which enter the Sponges, or it may be that our know- 
 ledge is incomplete on the subject. 
 
 If now we consider the analogy existing between the 
 Sponges on the one hand and the Protozoa and Myxo- 
 mycetes on the other, we find that the digestive and 
 excretory functions take a prominent part in the re- 
 action against the foreign bodies which might injure 
 these organisms. In the case of a sponge or of a Plas- 
 modium, this reaction consists merely in the enclosure 
 of the particles, and their subsequent digestion, if they 
 are digestible, or in their expulsion from the body, if 
 this is not the case. In the Myxomycetes this function 
 is performed by the whole of the protoplasmic contents, 
 whereas in the Sponges it is confined chiefly to the 
 mesoderm, though the endoderm also takes some part. 
 The outer covering or the ectoderm generally is however 
 not unconcerned in the work of protecting the organism 
 against all manner of injurious agents. The flat cells 
 of which the ectodermic layer consists, are contractile 
 and sensitive ; by their contraction the pores are opened 
 and allow the surrounding water to pass into the interior 
 of the sponge, so long as this does not carry any 
 harmful products along with it. It has long been 
 known that in order to follow the mode by which the 
 coloured granules penetrate into the cells to the best 
 advantage, experiments should be made with starving 
 specimens. As soon as the sponge is sufficiently filled 
 with the minute particles which have been brought by 
 the water, the pores refuse to allow the passage of any 
 more by remaining closed. 
 
LECTURE IV. 55 
 
 According to de Lendenfeld, the Sponges do not 
 open their pores to harmful substances, whether these 
 are in solution or consist of granules suspended in the 
 water. Of all the bodies which he made use of, such as 
 carmine, starch, milk, the last was the only one that at 
 once found its way inside the sponge ; in the case of 
 carmine, the pores at first remained shut, although they 
 opened after a short while. The solutions of different 
 toxic substances, as morphine, veratrine, or strychnine, 
 caused contraction of the pores, which did not relax for 
 some little time. 
 
 There is an analogy as well as a difference between 
 the mode of action of the ectodermic cells, which are 
 contractile although not phagocytic, and that of the 
 Plasmodia of the Myxomycetes. The analogy consists 
 in a sensibility towards the chemical composition of the 
 surrounding medium ; the difference is shown in the 
 manner of reaction. The mobile cellular colony — the 
 Plasmodium — is repelled from the object which calls 
 forth the exhibition of its sensibility (negative chemio- 
 taxis, thermotaxis or otherwise) ; the motionless organ- 
 ism — the Sponge — avoids the same object by refusing 
 entrance to it. 
 
 In spite of the limitations of our knowledge, we are 
 in a position to affirm that in their struggle against 
 various injurious bodies, the Sponges make use of all 
 their cellular properties^ more especially of the sensi- 
 bility and contractility possessed by the ectodermic 
 elements, and the power of englobing and digesting 
 common to mesodermic and endodermic cells. These 
 results may serve as a starting-point for the inquiry 
 into the more complex phenomena of reaction in other 
 animals. 
 
LECTURE V. 
 
 Ccelenterata, Echinodermaia and Vermes — Traumatism and re- 
 generation in Hydra — Accumulation of phagocytes in Acalepha 
 (Scyphomedusae) — Phagocytes of star-fishes — Inflammation in 
 Bipinnaria — Reactive changes in the perivisceral cells of the 
 Annelida — Phagocytic reaction in the diseases of Nais and 
 Lumbricus — Struggle between the phagocytes of Lumbricus 
 and Rhabditis — Microbic infections of Worms. 
 
 Although the Ccelenterata are distinguished from the 
 Sponges by their higher organisation, yet there are a 
 number of members of this group which consist of 
 only two layers of cells, the mesoderm being completely 
 absent. Since it is the mesoderm which, as we have 
 seen in the Sponges, plays the principal part in patho- 
 logical processes, it would be interesting to know how 
 those processes are carried out in such animals as Hydra 
 and its congeners, which have only two layers of cells. 
 
 As far back as the last century the phenomena 
 evoked in the fresh-water Polyps by every kind of 
 injury were often the subject of observation. Trembley 
 first pointed out the astonishing power of regeneration 
 possessed by this organism. A Hydra may be cut up 
 into several pieces, pierced with pointed bodies, and in 
 general maltreated to an extraordinary extent without 
 preventing a speedy and complete reintegration. In 
 one experiment of Ischikawa ' the front part of a 
 ' Zeitschr. f. ivissensch, Zoologie, vol. xlix, 1889, p. 433. 
 
LECTURE V. 
 
 57 
 
 Hydra had completely recovered within twenty minutes 
 after the infliction of an injury. Hydrae, cut in two 
 longitudinally, and stretched out on a cork, are able to 
 grow again into complete animals in little more than 
 twenty-four hours. 
 
 In another experiment Ischikawa cut off the head 
 and tentacles of a Hydra, and made a longitudinal slit 
 down the trunk, which he fixed on a piece of cork in 
 such a manner that the endoderm was directed out- 
 wards. In order to injure this layer the hydra was 
 
 Fig, 28. — Regeneration of a Hydra (after Ischikawa). 
 
 taken out of the water in this position, and exposed to 
 the air for five minutes. On then detaching it from the 
 cork and putting it into the water, the Hydra first rolled 
 itself up into a cylinder, the outer surface of which was 
 formed by the endoderm (Fig. 28, i) ; soon, however, it 
 rolled itself in the opposite direction, so that the two 
 layers occupied their normal relative positions. During 
 this inversion, however, the experimenter had inserted 
 a filament of an alga between the edges of the cut, so that 
 they could not fuse together (Fig. 28, 2). The Hydra then 
 changed its position, and finally grew into a closed sac 
 (Fig, 28, 3) which acquired a mouth and tentacles and 
 
$8 INFLAMMATION. 
 
 formed a perfect Hydra (Fig. 28, 4, 5) within six days 
 after the commencement of the experiment. 
 
 Punctures and other artificial lesions heal with extra- 
 ordinary rapidity, without any accumulation of phago- 
 cytes at the injured spot. But although no accumula- 
 tion takes place owing to the absence of a mesoderm, 
 yet we must not imagine that the phagocytic function 
 is completely wanting in these animals. The whole of 
 the endoderm in the Hydra consists of stationary phago- 
 cytes in the form of epithelial cells, which are capable 
 of putting out amoeboid processes from their free sur- 
 face, and ingesting various foreign bodies. 
 
 In the marine colony-forming Hydromedusse, not 
 only the endoderm, but occasionally the ectoderm as 
 well, consists of phagocytes which are of great import- 
 ance on account of their prophylactic action.^ These 
 creatures like the Hydra have the power of regeneration. 
 If the head of one of these Hydrozoa, such as the Podo- 
 coryna, be cut off and the trunk left in contact with 
 the colony, a fresh head will grow, while the detached 
 head becomes fixed and forms a new trunk. 
 
 In all these phenomena we see a regenerative capacity 
 of such rapidity and extent that the danger of infection 
 becomes reduced to a minimum. We may here observe 
 the regenerative side of inflammatory processes, but not 
 the phenomena of inflammation itself or at least not the 
 accumulation of phagocytes at the injured spot. 
 
 This accumulation is not however an uncommon 
 occurrence in the Ccelenterata. Most of these animals, 
 like the Acrospeda, the Ctenophora and the true Polyps, 
 are provided with a tolerably well developed mesoderm, 
 
 ' See my article in Arbeiten des sool. Institutes zu Wien, 
 vol. V. 1883, pp. 143-146. 
 
LECTURE V. 59 
 
 and contain in their intercellular substance a number of 
 amoeboid cells which have all the properties of phago- 
 cytes. 
 
 If we take a large Medusa, known as Rhizostomunt 
 Cuvieri, and introduce some pointed body, as a splinter 
 of wood or even a pin, into its gelatinous bell, the very 
 next day it will be perceptible to the naked eye that a 
 cloudiness has arisen around the foreign body. This, 
 when microscopically examined, will be found to consist 
 of numberless amoeboid cells which have collected round 
 the seat of lesion. The same thing takes place in 
 another of the Acalepha, Aurelia aurita. If the object 
 introduced into the bell of the Medusa has been pre- 
 viously soaked in a colouring matter, such as carmine, 
 the phagocytes which assemble at the injured spot will 
 be found filled with coloured granules. The amoeboid 
 cells which accumulate around the foreign body either 
 remain isolated or unite to form minute plasmodia. 
 
 We therefore see that in these Medusae, which have 
 no kind of vascular system, the mesodermic phagocytes 
 are able to traverse a gelatinous substance, which is 
 sometimes, as in the Rhizostomum, very tough, and to 
 collect for the purpose of englobing the minute bodies 
 or of surrounding the larger foreign bodies which have 
 been introduced. 
 
 The analogy of these events with the reactive pheno- 
 mena in the Sponges is obvious, although there is a 
 considerable difference between the mesoderms in the 
 two cases. Whereas in the Sponges, the mesodermic 
 phagocytes by taking up solid food-particles play an 
 important part in nutrition, in the Medusae and in all 
 the Coelenterata which possess a mesoderm, the func- 
 tion of nutrition is exclusively confined to the endoderm. 
 
6o INFLAMMATION. 
 
 In all the Ccelenterata this consists of a phagocytic 
 epithelium entirely separate from the mesoderm, at any 
 rate in the adult condition. Although deprived of their 
 nutritive function, the mesodermic phagocytes retain 
 their power of approaching foreign bodies, of englobing or 
 surrounding them and of digesting some of them. This 
 power is not only exercised on the foreign bodies which 
 have penetrated into the Ccelenterata as a result of some 
 lesion ; it is equally active in the case of the tissues of 
 these animals themselves. Thus the abortive genera- 
 tive cells, of which there are a large number in Medusae 
 kept under artificial conditions, are duly devoured by 
 the phagocytes, which surround them as with a kind of 
 follicle. It is apparent that these mesodermic cells 
 have not lost their primitive properties of intracellular 
 digestion, and although they have become distinctly 
 separated from the endoderm, the common origin of the 
 two layers may be embryologically demonstrated. 
 
 The development of the mesodermic amoeboid cells 
 at the cost of the endoderm — which is a fact of very 
 frequent occurrence in the animal kingdom — may be 
 readily followed in the various representatives of the 
 Echinodermata, especially in the sea-urchins and the 
 star-fish. We will take a star-fish common in the Gulf 
 of Trieste, the Astropecten pentacanthus, as an example. 
 The segmented ovum becomes converted into an oval 
 body consisting of a layer of ciliated cells enclosing a 
 segmentation cavity. A part of these cells becomes in- 
 vaginated to form the first rudiments of the intestinal 
 canal and its appendages. The larva soon assumes the 
 characteristic gastrula stage, and consists of an ectoderm, 
 or outer layer, and of an endoderm, forming a cul-de-sac 
 with an opening at the lower end. The space between 
 
LECTURE V. 6l 
 
 the two layers represents the general body-cavity and 
 is filled with a homogeneous and semi-fluid substance, 
 containing the amoeboid cells of the mesoderm. These 
 are merely migratory cells which have become budded 
 off from the endodermic invagination.' (Fig. 29.) Al- 
 most immediately upon their arrival in the body-cavity 
 they are able to begin their phagocytic duties. Among 
 the numerous larvae of the Astropecten which float on 
 the sea, some may be found with their delicate ectoderm 
 injured by a sharp body which has pierced the general 
 body-cavity. (Fig. 30.) But as 
 soon as the larva has been thus 
 damaged, the mesodermic cells 
 travel towards the invading object, 
 and surround it completely by 
 fusing into minute plasmodia. 
 (Fig. 31.) The latter, if duly 
 prepared in a o"5 per cent, solu- 
 tion of osmic acid and stained 
 with picrocarmine, are seen to 
 contain a certain number of ^SsmtlTe^krvaofAst'/opiSfn! 
 nuclei, the appearance of which 
 
 shows definitely that no proliferation of cells is taking 
 place. In these larvze, the reaction, which can be 
 followed step by step owing to their transparency, 
 consists merely in an accumulation of mesodermic 
 phagocytes around the foreign body. There can be no 
 question here of the intervention of any vascular, 
 muscular, or nervous systems, since in these larvae such 
 
 ' Cf. my article in the Zeitschrift f. wissensch. Zoologie, vol. xlii. 
 1885. See also the account of the discussion between Selenka and 
 myself, and the more recent paper by Korschelt in the Zoologische 
 Jahrbucher, vol. iv. i88g. 
 
62 
 
 INFLAMMATION. 
 
 systems do not exist. The reaction is effected entirely 
 by the phagocytes themselves, and is accompanied 
 neither by proliferation of cells nor by increased flow 
 of fluid to the part, as shown by the absence of oedema. 
 The non-occurrence of proliferation is readily explic- 
 able by the fact that the foreign body on account of its 
 minuteness has produced but a very slight lesion of the 
 ectoderm. 
 
 In larvas of more highly deve- 
 loped and complex organisation, 
 the reaction takes place in the 
 same manner. I have frequently 
 seen a marine alga, a species of 
 Chcstoceros which is provided 
 with very delicate hairs, settle on 
 and penetrate into specimens of 
 Bipinnaria (larvae of Astropecten). 
 In all these cases, the lesion was 
 followed by an accumulation 
 of mesodermic phagocytes to- 
 with the formation of 
 Plasmodia. 
 In the instances above quoted, the larvae were too 
 minute to admit of artificial experimentation ; it was 
 merely a question of watching the effects of lesions 
 arising under natural conditions. But the reactive 
 phenomena ensuing on artificial injuries ' may be readily 
 observed in the much larger larvae, the Bipimiaria 
 asterigera, which likewise represent a stage in the 
 development of the star-fish. If a delicate glass tube, a 
 rose-thorn, or a spine of a sea-urchin be introduced into 
 
 > See my article in the Arbeiten des zoolog. Institutes zu 
 Wien, 1883, vol. v. p. 141. 
 
 Fig. 30.— Gastrula with a foreign „p\\,pY 
 substance in its body-cavity. gctiici 
 
LECTURE V. 63 
 
 one of these larvae, the amoeboid cells of the mesoderm 
 collect around the foreign body in large masses easily- 
 visible writh the naked eye. All the minute particles 
 adherent to the object introduced, or the granules of 
 carmine or indigo, if the object has been previously 
 immersed in these substances, are eagerly devoured by 
 the mesodermic phagocytes. (Fig. 32.) 
 
 If instead of these sharp solid bodies, a drop of blood 
 be introduced into a Bipinnaria, it will be at once 
 
 Fig. 31. — The oreign body of Fig. 30 surrounded by a Plasmodium of 
 the larva (highly magnified). 
 
 surrounded by the mesodermic cells, which will collect 
 around the masses of blood corpuscles to form true 
 Plasmodia, that is to say, multinuclear protoplasmic 
 masses arising from the complete fusion of the 
 phagocytes (Fig. 33). The changes undergone by the 
 red corpuscles of the blood within the mesodermic cells 
 of the larva correspond exactly with the phenomena of 
 intracellular digestion, and may be observed in the same 
 way with the fat-globules of milk. 
 
 Bacteria if introduced into the Bipinnaria are likewise 
 enclosed by the mesodermic phagocytes. The great 
 
64 INFLAMMATION. 
 
 transparency of the larvae of Echinoderms enables the 
 observer to ascertain the fact that the bacteria are still 
 mobile, and that they have therefore been devoured 
 alive. 
 
 In spite of the differences which distinguish the 
 Sponges, the Ccslenterata possessing a mesoderm, and 
 the Echinoderms from each other, they are all essentially 
 similar in their phenomena of reaction. In the Sponges 
 we have a mesoderm with a plentiful supply of mobile 
 cells which play an important part in the nutrition of 
 ^"m ^"^^k^MT^ a/ these animals. The food, 
 
 ■ ^«^^^&^>" ^^^^"^ entering the body, 
 
 (^J«^^pb^>i>r^ invariably reaches the 
 
 mesoderm, which is in- 
 timately connected with 
 the endoderm. In the 
 Acalepha (Medusae) and 
 the other Coelenterata in 
 
 Fig. 32.— Collection of phagocytes round a which this layCris prCSCnt, 
 splinter, Bi^innaria asterigera. . - . ,. , 
 
 the mesoderm is directly 
 connected with the endoderm only while it is being de- 
 veloped. When its development is complete, the meso- 
 derm becomes definitely separated from the endoderm and 
 has nothing further to do with the function of nutrition, 
 which is relegated entirely to the endodermic phago- 
 cytes. In the larvae of Echinoderms the two layers are 
 equally distinct from each other ; the mesoderm is like- 
 wise excluded from the office of nutrition, while the 
 endoderm which is the sole organ of nutrition, has no 
 power of intracellular digestion . In this animal, digestion 
 is performed by means of ferments secreted by the 
 endodermic cells, and poured into the intestinal cavity. 
 Now although these animals differ thus in organisa- 
 
LECTURE V. 
 
 65 
 
 tion from one another, their mesodermic cells are alike 
 in that they move towards foreign bodies, englobe and, 
 when possible, digest them. In all of them the various 
 lesions produced by such foreign bodies provoke an 
 accumulation of mesodermic phagocytes, with or with- 
 out formation of plasmodia or giant cells. 
 
 Moreover these animals all resemble each other in the 
 fact that the phagocytes of their mesoderm are repre- 
 
 Fig. 33. — Plasmodium formed by the phagocytes of the Bipiniiarla, 
 
 sented by branched connective tissue corpuscles 
 embedded in a semi-fluid or gelatinous intercellular 
 substance. In all the cases that we have considered, 
 there was neither blood nor plasma, blood-corpuscles 
 nor blood-vessels. These structures are not found 
 either in the Sponges or in the Ccelenterata, and are 
 only present in the Echinoderms at a later period of 
 development than that of the specimens which served for 
 these pathological investigations. 
 
 If now we turn our attention to the varied group of 
 the Vermes, we shall at once meet with reactive phe- 
 
 F 
 
66 INFLAMMATION. 
 
 nomena similar in character to those described above. 
 As a representative of the lower orders of worms, we 
 will take the transparent Turbellarium, Mesostomum 
 Ehrenhergi. If we injure any part of its body, the 
 mesodermic phagocytes will, after a certain time has 
 elapsed, assemble round the lesion. The number of 
 granules they contain gives them a great resemblance 
 to the epithelial cells of the intestine which, in the 
 Turbellaria, are likewise true phagocytes. The meso- 
 dermic cells are amoeboid elements embedded in a 
 gelatinous intercellular substance, forming a mucoid 
 connective tissue like the mesoderm of the Sponges, 
 Medusae and Echinoderms. 
 
 In the higher Worms the mesodermic phagocytes are 
 represented by the peritoneal endothelium or by cells 
 suspended in the perivisceral fluid. These two varieties 
 of cells have the same marked phagocytic properties, 
 and this functional analogy may explain the fact that in 
 closely allied species, the perivisceral cells are some- 
 times highly developed while at other times they are 
 completely absent. These mesodermic elements be- 
 sides fulfilling their phagocytic duties, act as respira- 
 tory and excretory organs.^ 
 
 If a splinter be introduced into the perivisceral 
 cavity of an Annelid, such as the Terebella, it will be 
 soon covered with a thick layer of these 'lymphatic' 
 cells, the phagocytic properties of which are shown by 
 the readiness with which they take up the minute 
 grains (of colouring matter or otherwise) attached to 
 the splinter. This is the more interesting, since the 
 
 ^ See Grobben, " Die Pericardialdriise der chaetopoden Anne- 
 liden." Sitzuiigsberichte der k. Akad. d. Wissensch., Wien, vol. 
 xcvii., 1888. 
 
LECTURE V. 
 
 67 
 
 majority of the Annelida are endowed with a highly- 
 developed and completely closed vascular system. The 
 reaction to foreign bodies is however confined to the 
 mesodermic phagocytes, the vessels taking no part 
 whatever in the process, as may be readily seen owing 
 to the bright colour of the blood in them. 
 
 The same phenomena occur in the Annelida which 
 
 
 V^f^omjtl 
 
 Fig. 
 
 3^. — A larva of Gordlus enc>^sted and surrounded by a 
 Plasmodium in a specimen of Nais. 
 
 have a well-developed vascular system, but no peri- 
 visceral leucocytes. If a certain number of Nais 
 proboscidea be examined, some of the individuals will be 
 found to be infected with larvae of Gordius, which after 
 entering the general body cavity, excite phagocytic re- 
 action of the peritoneal cells alone. The latter send out 
 protoplasmic processes and form minute plasmodia 
 around the larvae, which protect themselves by secreting 
 a chitinous covering and enclosing themselves in a sort 
 of cyst. Although these cysts with their surrounding 
 * F 2 
 
68 INFLAMMATION. 
 
 Plasmodia are sometimes found in close proximity to the 
 vessels, the latter do not react in any way to the presence 
 of the parasite. (Fig. 34.) If any exsudation from the 
 vessels took place into the interior of the plasmodium, 
 it would be at once evident, since the blood-plasma is 
 yellow while the perivisceral fluid is perfectly colour- 
 less. 
 
 As it is manifestly impossible to experiment upon the 
 minute Nais proboscidea, the phagocytic properties of 
 the peritoneal cells of this Annelid must be studied 
 in specimens infected by a microsporidium belonging 
 to the microbes of ^ pibrine! These parasites excite a 
 similar reaction on the part of the peritoneal endothe- 
 lium, the minute spores being taken up by the 
 phagocytes which have become separated from the 
 peritoneum. Occasionally these spores may be seen 
 surrounded by vacuoles as in the most typical cases of 
 intracellular digestion. 
 
 The larger Annelida may likewise be employed for 
 these researches. Valuable information may be ob- 
 tained from the study of the common earth-worms, 
 which are frequently invaded by parasites. Among 
 these the most frequent and also the best known are the 
 Gregarinse belonging to the genus Monocystis, which 
 attack the male organs. When once inside the latter,- 
 these mobile Protozoa have to encounter a large num- 
 ber of amoeboid cells, which are among the most active 
 of phagocytes. They are provided with slender and 
 membranous protoplasmic processes (Fig. 35, A), and 
 devour with the greatest eagerness all foreign bodies 
 which come in their way. Even when examined in the 
 aqueous humour of the rabbit or other inert fluid, 
 these cells give evidence of their phagocytic activity by 
 
LECTURE V. 
 
 69 
 
 englobing grains of colouring matter or any other 
 minute bodies that may have been added to the pre- 
 paration. If they meet with a larger object, such as a 
 thread of cotton, they will collect in groups and finally 
 surround it with their protoplasm. (Fig. 35, B.) Now 
 these same cells react to the parasites which have 
 penetrated the earth-worms. 
 While the Gregarinas are in 
 their condition of activity, 
 they repulse the phagocytes 
 by the violence of their move- 
 ments so that the latter are 
 rarely able to fix themselves 
 on the parasite. But as soon 
 as it attains the quiescent 
 state, the phagocytes adhere 
 to its surface, frequently 
 collecting to form a dense 
 mass around it. The Gre- 
 garina evidently objects to 
 this living covering and seeks 
 to defend itself by the secre- 
 tion of a cystic membrane. 
 (Plate II., Fig. I.) Thus 
 protected, it begins to produce spores by dividing 
 into a large number of increasingly smaller oval bodies, 
 and gives rise to the pseudo-navicellae which have 
 been so often described. The mass of surrounding 
 phagocytes continues however to act upon the parasite 
 and frequently succeeds in injuring and even in killing 
 it. The encysted Gregarina goes on defending itself by 
 means of chitinous secretions which assume exaggerated 
 proportions and become fringed with irregular processes. 
 
 f"ig' 35- — -A._ Phagocyte of Lmnhriciis. 
 B. Collection of phagocytes of Liim- 
 hricits round a foreign body. 
 
70 INFLAMMATION. 
 
 presenting a strikingly abnormal appearance. (Plate 
 I., Fig. 2.) Finally the whole interior of the en- 
 cysted Gregarina becomes highly refracting, and the 
 parasite dies, leaving the phagocytes the masters of the 
 day. (Plate I., Fig. 3.) The phagocytes themselves 
 undergo marked alteration around the parasite and, 
 losing their power of movement, are converted into flat 
 closely apposed cells. (Plate II., Fig. i, 2.) Sometimes 
 the capsule thus formed, which has the same structure 
 as connective tissue, remains very thin ; but it usually 
 becomes thickened by the further addition of fresh 
 
 Fig. 36. — A live Rliabditis surrounded by a mass of the phagocytes of Lumbricus. 
 
 layers of cells. Among these may be seen some which 
 are deeply pigmented with a brown colour. During 
 the whole course of the struggle between the parasite 
 and the phagocytes of Lumbricus, the blood-vessels 
 although highly developed remain quite passive, that is 
 to say, there is no visible change in their volume, nor 
 any exsudation of the reddish plasma. 
 
 The spermathecae of Lumbricus as well as its 
 general body cavity, may also be invaded by Nema- 
 toda belonging to the genus Rhabditis. The latter, 
 in spite of their size and mobility and the toughness 
 of their cuticle, have likewise to do battle with the 
 
LECTURE V. 71 
 
 numerous phagocytes of the earth-worm. These cells 
 surround the Nematode forming a thick capsule round 
 it like that which is produced round the Gregarina. 
 Microscopical examination of the Rhabditis in this 
 condition proves directly that the phagocytes have 
 enclosed it while still alive, as it may be seen moving 
 in the midst of the mass of cells. (Fig. 36.) The 
 worm, thus confined in its movements, secretes layers 
 which form, not a true cyst, but a supplementary cuticle 
 which frequently becomes of extraordinary thickness. 
 (Plate I., Fig. 4.) This abundant secretion evidently 
 
 Fig. 37. — A Rhabditis without its pliagocytic covering in order to 
 show the cuticular processes. 
 
 exhausts the parasite, for it loses the granules of fat 
 with which it was at first filled and becomes quite 
 transparent. (Plate I., Fig. 5.) The chitinous layers, 
 as they become thicker and thicker, finally form irre- 
 gular processes, which give a strange abnormal appear- 
 ance to the Rhabditis. (Fig. 37. Plate I., Fig. 4.) In 
 isolating the collections of phagocytes enclosing the 
 worm with its thickened cuticle, it often happens that 
 the Nematode manages to escape, leaving its cuticle in 
 the mass of cells. On the other hand, if the contents 
 of the male organs of Lumbricus be examined, there 
 will frequently be found situated in the midst of 
 phagocytic capsules, highly refracting bodies, which 
 may readily be identified as the shapeless cuticular 
 
72 INFLAMMATION. 
 
 layers and the remains of the Nematode buried in the 
 products of its own secretion. 
 
 Here then we have an example of a struggle be- 
 tween two members belonging to the same group of 
 the animal kingdom. The Nematode worm protects 
 itself by means of cutaneous secretions ; the earth- 
 worm fights by means of an army of mobile cells, en- 
 dowed with phagocytic properties. It is evident that 
 the latter greatly embarrass the parasite by surround- 
 ing it with their solid masses, although we do not yet 
 know the exact nature of the harmful influence exer- 
 cised by the phagocytes on the intruder. They may 
 act by preventing the inflow of nutritive material or 
 of oxygen, or by means of an injurious secretion. 
 These delicate points can only be decided by the 
 most minute research, and must await a perfection of 
 our methods for their final answer. 
 
 At present we must be satisfied with the statement 
 that Lumbricus, like the Annelida generally, reacts 
 to various infective agents by means of the phago- 
 cytes of their perivisceral fluid without any intervention 
 on the part of the blood or of the highly developed 
 blood-vessels. This reaction takes place in the same 
 way against Gregarinse as against Nematoda. In 
 speaking of the latter, I must particularly mention 
 the fact t6at they are devoid of migratory cells. The 
 phagocytic system of the Nematoda is probably re- 
 duced to the muscular phagocytes, which are formed 
 in a very curious manner. These animals protect 
 themselves by the secretion of tough membranous 
 cuticles, resembling in this feature the plants, the cells 
 of which are likewise protected by thick resistant mem- 
 branes. This analogy is borne out by the fact that 
 
LECTURE V. 73 
 
 these animals, like plants, are very frequently attacked 
 by parasitic fungi, which are enabled to penetrate the 
 cuticle by their great power of growth and also by 
 the secretion of ferments which can dissolve the most 
 impervious substances, such as cellulose. 
 
 Among the infectious diseases of the Nematoda, we 
 must name one which is produced by the parasitism 
 of one of the Mucorineae [Mucor helminthophorus, De 
 Bary), which invades the intestine and the genital 
 organs of the Ascaris of the cat [A. mystax)^ as well 
 as the frequent infections of the free Nematoda by 
 several other members of the class of fungi. The 
 most remarkable is certainly the Arthrobothrys oligos- 
 pora Fres., because, according to Zopf," this mould 
 catches the Anguillulides with hooks, and afterwards 
 penetrates their bodies with its filaments. Once inside, 
 the fungus grows freely in the body cavity and causes 
 complete fatty degeneration followed by death of the 
 animal. There is finally nothing left of the Nematode 
 but its cuticle and the chitinous covering of the male 
 genital organs. • 
 
 Besides these epidemics, occasioned by the higher 
 fungi, the Nematoda are liable to invasion by Chytri- 
 diaceae and other inferior organisms, allied to those 
 which infect the nucleus and nucleolus of the Para- 
 maecia.' 
 
 This survey of the pathological phenomena in 
 Ccelenterata, Echinoderms and Worms, has shown 
 
 ^ Zeiischrift f. wissensch. Zool., 1862, vol. ii. p. 135. 
 
 ^ Nova acta Acad. Leopold, vol. xlvii. p. 167; and "Pilze," 1890, 
 p. 240. 
 
 ^ See Biitschli, " Studien iiber die ersten Entwickelungsvorgange 
 der Eizelle." Frankfurt, 1876, p. 360. 
 
74 INFLAMMATION. 
 
 that some of these animals react chiefly by the rapid 
 and active regeneration of the injured parts, while 
 others protect themselves by secretions of chitinous 
 layers. These two methods, however, only hold in 
 certain cases, whereas the usual mode of reaction, to 
 which there are but few exceptions, is by means of 
 amoeboid and mobile cells which accumulate around 
 the injurious body and either surround it entirely or 
 englobe it. The reaction is effected through the sensi- 
 bility of the phagocytic cells themselves, and is in no 
 way influenced by the nervous or vascular system. In 
 all the above-mentioned cases, the phagocytes were 
 mobile connective tissue cells, or cells of the peri- 
 visceral cavity. So far we have not discovered any 
 instance of phagocytic action on the part of the blood- 
 corpuscles. It is true that our observations have been 
 confined to animals which have no formed elements 
 in the blood. Annelida with white corpuscles in 
 their blood do not often occur, and even in cases 
 where these are present, their number is small, and 
 certainly less than the amount found in the perivisceral 
 cavity. 
 
LECTURE VI. 
 
 Arthropoda, Molhisca, and Tunicata — Their vascular system — 
 Their Phagocytes — Spleen of the Gasteropoda — Inflammatory 
 reaction — Diapedesis in intact Ascidians — Introduction of 
 bacteria into the body of Ascidians and Crustaceans — Infec- 
 tious disease of sandhopper (Talitrus) — Diseases of Daphnia 
 — Introduction of bacteria into insects — Epidemics among 
 insects. 
 
 A LARGE number of the invertebrata possess blood- 
 corpuscles in the form of colourless cells which float in 
 the blood-plasma. The circulation of this fluid is 
 effected by the movements of the heart, which is always 
 present. In these animals — Arthropoda, Mollusca, and 
 Tunicata — the vascular cavity is identical with the 
 general body cavity. In the lower representatives of 
 these types (we disregard certain groups without a trace 
 of a vascular system, such as many of the Copepoda, 
 Ostracoda and others) the only vascular organ present 
 is the heart, in the form of a simple sac or tube open at 
 its extremities to expel the blood and provided with 
 lateral apertures for the entry of this liquid. To this 
 central organ are soon added one or several principal 
 arteries which open into a system of lacunae, in which 
 the blood circulates before going back to the heart. 
 In the invertebrata rather higher in the scale, especially 
 in the Mollusca, we find also a venous system, which 
 is sometimes, as in the Cephalopoda, very highly 
 
76 INFLAMMATION. 
 
 developed. But in all cases without exception, even 
 when a large number of vascular ramifications are 
 formed, there is a network of lacuna between the 
 arterial and venous systems. These lacunae are filled 
 with blood and are remains of the general body 
 cavity. 
 
 The blood-corpuscles with a few rare exceptions 
 are represented by colourless cells, possessing one or 
 rarely two nuclei and a protoplasmic body capable of 
 amoeboid movements. In many invertebrates there is 
 only one variety of mobile blood-corpuscles, containing 
 a few sparse granules, whereas in certain others, such 
 as many insects and molluscs, two varieties occur — 
 granular leucocytes, with a large number of coarse 
 granules, and hyaline leucocytes, with few or no 
 granules. The latter kind is the one which will interest 
 us most. 
 
 The leucocytes of Arthropoda, Mollusca, and 
 Tunicata are in most cases amoeboid and phagocytic 
 cells, and differ from the white corpuscles of verte- 
 brates in having a single round or oval nucleus, which 
 is not lobed. In the invertebrates now under considera- 
 tion no multinuclear leucocytes exist, nor do we find 
 in them a vascular system with complete capillary 
 blood-vessels. 
 
 The leucocytes of the three types just mentioned 
 manifest pronounced phagocytic functions. It was in 
 a representative of these invertebrate groups that the 
 discovery was first made (in 1862) that leucocytes 
 possessed the power of taking up foreign bodies into 
 their interior. Hseckel ' showed that, after injecting 
 the mollusc Thethys with indigo, granules of 
 ' " Die Radiolarien," 1862, p. 104. 
 
LECTURE VI. yj 
 
 this colour were to be found within the blood-cor- 
 puscles. Experiments with several other species led 
 him to the conclusion that this was a fact capable of 
 general application — a deduction which has since been 
 confirmed by several observers. It is therefore very- 
 surprising that a recent author, Griesbach, ^ should 
 throw doubts on the occurrence of phagocytosis 
 in the white corpuscles of the acephalous Molluscs. 
 Since he did not succeed in observing any considerable 
 taking up of a powder after injecting it mixed with 
 water, Griesbach concludes that under normal condi- 
 tions phagocytosis does not occur at all in these 
 animals. It is very probable that the unsatisfactory 
 results of this author were due to the fact that he used 
 too much water to dilute the powder, and so caused the 
 leucocytes to swell up. If we proceed more carefully, 
 it is easy to show that in the Molluscs, as in so many 
 other animals, the leucocytes take up greedily any solid 
 bodies with which they happen to be in contact. The 
 transparent Molluscs such as Philliroe, which admit of 
 direct examination under the microscope in the living 
 condition, form very convenient objects for these 
 researches. 
 
 In some gasteropod Molluscs, we find in addition to 
 the white corpuscles, a special variety of phagocytes 
 which form a kind of spleen in these invertebrata. 
 This important fact has been recently discovered by 
 A. Kowalewski," who has shown that solid bodies, 
 injected into the blood of the Pleurobranchsea and 
 several other species (Philina, Gasteropteron, Doris), 
 
 ' Archiv f. mikroskop. Anatomic, vol. xxxvii. p. 86. 
 ^ Menioires de la Societe des Natttralistes de la Noiivelle Russie, 
 vol. XV., 1890 (in Russian). 
 
78 INFLAMMATION. 
 
 accumulate in a certain organ which was first described 
 by de Lacaze-Duthiers under the name of " glande 
 ind^termin6e." The cells of this ' spleen/ as 
 Kowalewski has established, deovur and digest a large 
 number of foreign bodies such as blood corpuscles, yolk 
 granules, and milk corpuscles. 
 
 The phagocytes, so universally present in the inverte- 
 brata which form the subject of the present lecture, 
 react to all sorts of lesions, whether these are artificially 
 or accidentally produced in these animals. We often 
 come across transparent Crustacea, such as Daphnia or 
 Branchipus, with brown spots on their sides, due to 
 
 bites inflicted by other 
 
 individuals. Under- 
 
 r-^'^'U-^fat neath these eschars we 
 
 <Q) -'^'•"^si^a^ generally find a mass of 
 
 5S (X> o"^.^>^^{S^ leucocytes, which re- 
 
 ~1^ main heaped together 
 
 Fig. 38.-I„flamed Cauda appendage of Argulus. ^t the injured SpOt UUtil 
 
 the wound is com- 
 pletely cured.' If we carefully inflict a small injury on 
 one of these animals and obserye it under the micro- 
 scope, we see the leucocytes making towards the 
 affected spot, where they take up their abode. A con- 
 venient object for experiments of this sort is furnished 
 by the caudal appendages of Argulus, in which the leuco- 
 cytes collect directly after the production of the artificial 
 lesion (Fig. 38). 
 
 We may also introduce fairly large foreign bodies, 
 such as wooden splinters, into the larvae of various 
 Coleoptera (cockchafer, Oryctes and others), into 
 
 ' The extremely rapid regeneration of the epidermis in the 
 Arthropoda causes their wounds to heal very quickly. 
 
LECTURE VI. 79 
 
 molluscs such as Thethys or Phylliroe or into AscidiansJ 
 In all these cases a great number of leucocytes may be 
 seen to collect around the foreign body, devouring any 
 little fragments or granules as, for example, carmine, 
 that may have been introduced with it. 
 
 In all cases then a phagocytic reaction is produced 
 by the introduction of a foreign body, the leucocytes 
 either forming a capsule round the intruder, or infil- 
 trating all the surrounding tissues. In this exsudative 
 and inflammatory reaction, which is often accompanied 
 with the formation of giant cells, diapedesis can play 
 no part, for the simple reason that in the Arthropoda 
 and Mollusca there is no closed vascular system 
 properly speaking, the blood cavities being merely part 
 of the general body cavity. 
 
 In the invertebrata only one single example of dia- 
 pedesis occurs ; but this is of so interesting a character 
 that it deserves a detailed description. The Ascidians 
 are covered with a mantle or tunic which is situated 
 outside the epidermis. This tunic, which is composed 
 of cellulose and is often very thick, contains a large 
 number of amoeboid cells with mobile processes. Since 
 it is situated outside the epidermis, it was generally 
 regarded as a cutaneous secretion containing cells of 
 ectodermic origin. Later researches by Kowalewski " 
 have shown that this view was unfounded, and that the 
 cells in the tunic of Ascidians are in reality nothing 
 else than emigrated leucocytes which have wandered 
 through the epidermis. These cells of mesodermic 
 origin are very active phagocytes, and are capable of 
 devouring all sorts of solid bodies, including organs 
 
 ' See \!ii^ Arbeiten des zoolog. Inst.zu lVien,\o\. v., 1883, p. 153. 
 " Loc. cit. 
 
8o INFLAMMATION. 
 
 that are undergoing atrophy. Insertion of splinters 
 into the tunic of Ascidians provokes an accumulation 
 of these phagocytes, so that a sort of infiltration of the 
 tunic is produced. 
 
 We have here an example of diapedesis taking place 
 under normal conditions through the epidermis, quite 
 independently of any inflammation; though this latter 
 process is also carried out by the aid of the same 
 phagocytes^ which collect around the offending particles. 
 
 Lubarsch ' has confirmed the observation that the 
 mobile cells in the tunic of Ascidians congregate around 
 foreign bodies that have been introduced by a puncture. 
 He was not so successful in his experiments on the 
 inoculation of various Ascidians with the bacilli of 
 anthrax. The bacteria which had been introduced 
 into the tunic were only partially taken up by the 
 phagocytes, while those which escaped this fate never- 
 theless showed marked signs of degeneration. As 
 Lubarsch did not investigate the direct influence of the 
 fluid portions of the tunic on the bacteria, we cannot 
 form any definite conclusion from his none too 
 numerous experiments. In considering them we must 
 not lose sight of the facts that they were carried out in 
 the month of March, when the low temperature might 
 have had an injurious influence on the leucocytes, and 
 that the tunic of Ascidians cannot afford a very favour- 
 able soil for the growth of bacteria and the production 
 of their toxines.^ Lubarsch ' has also made some 
 
 1 " Untersuchungen iiber die Ursachen der angeborenen und 
 erworbenen Immunitat." Berlin, iSgr, p. 75. 
 
 ' I must here mention the fact that the tunics of compound 
 Ascidians such as Botryllus, often show phagocytes filled with 
 various bacteria, even when the animals are examined immediately 
 after being taken from the sea. 
 
 ' Loc. cit., p. "JT. 
 
LECTURE VI. 8 1 
 
 similar experiments on " marine Crustacea " without 
 any better result. The experiments are recorded very 
 shortly, so that it is impossible to criticise them. We 
 have a number of facts, however, which prove in 
 the clearest manner the pronounced phagocytic 
 properties possessed by the leucocytes of different 
 Crustacea. By the introduction of a parasitic fungus 
 (allied to Oidium) into the body cavities of sand- 
 hoppers (Talitrus), Hermann and Canu ' have suc- 
 ceeded in producing a disease, which is almost always 
 fatal to these Crustacea. The development of the 
 parasite excites reactive changes on the part of the 
 organism, as shown by a pronounced phagocytosis of 
 the leucocytes. The authors describe these changes as 
 follows : " On the seventh day, the blood, which up to 
 this time has been quite clear, becomes appreciably 
 opalescent, and the disorder becomes more marked on 
 the eighth and ninth days, as the parasites increase 
 in number. This is also the period at which the 
 most active phagocytosis is observed ; if the blood, 
 fixed by osmic acid vapour and stained with picrocar- 
 mine, be examined under a high power, the microbes 
 are seen to be enclosed in the corpuscles, the number 
 in each corpuscle varying from one to twenty. In the 
 protoplasm they are seen to be in various stages of 
 digestion ; they become paler and less highly refracting 
 at the same time that they increase in size, chiefly in 
 consequence of a swelling up of their enveloping mem- 
 brane. Finally the place they occupied in the cell is 
 marked only by a sort of colourless vacuole which 
 preserves for a considerable time the elongated shape 
 of the parasite." Besides the blood-corpuscles, the cells 
 ' Compies rendus de la Societe de Biologie, 1891, p. 646. 
 
 G 
 
82 
 
 INFLAMMATION. 
 
 surrounding the arteries also take part in devouring 
 the parasitic fungi, thus acting as phagocytes, although 
 they are not able to digest their prey. In the end, 
 therefore, the parasite gains the upper hand and brings 
 about the death of the sandhopper. 
 
 Luminous bacteria also, as Giard and Billet have 
 shown, live as parasites on the same species of 
 Amphipoda. 
 
 Fig. 39. — Daphnia infested by Monosporaa. 
 
 A large number of the Crustacea are subject to 
 various infectious diseases which form an interesting 
 subject of study from the pathological point of view, 
 and especially on account of the light they throw on 
 the problem of inflammation. The Daphniae afford 
 especially convenient objects for these researches, in 
 consequence of their transparency and minute size, and 
 the frequency and variety of their diseases. Among 
 the latter we find infectious maladies produced by 
 bacteria, Sporozoa or Saprolegniae. The most interest- 
 ing of these, however, is certainly that provoked by the 
 presence of a fungus multiplying by budding, Motiospora 
 
LECTURE VI. 
 
 83 
 
 bicuspidaia^ This is a sort of yeast, which is found in 
 abundance infesting Daphnia magna in Paris (in the 
 reptile tank of the Jardin des Plantes) and its 
 environs. 
 
 Among the numerous individuals of this Crustacea, 
 we come across specimens distinguished by their milk- 
 white colour. On examining these under the micro- 
 scope, we see that their body-cavity is almost entirely 
 filled with small needle-shaped bodies, either floating 
 freely or adhering to the walls of 
 the heart (Fig. 39). A close in- 
 spection shows at once that we 
 have here very long spores^ en- 
 closed in a capsule (Fig. 40, 5). 
 By the side of these mature 
 spores, elongated cells and oval 
 conidia are seen multiplying by 
 budding, exactly in the same 
 manner as the yeasts. (Fig. 40, 
 1-4.) 
 
 A Daphnia, once invaded by 
 these parasites, always dies, 
 and its body is found filled with ripe spores. Other 
 DaphnisE, feeding on every sort of detritus which they 
 find at the bottom of the water, devour the needle- 
 shaped spores, and thus infect themselves through their 
 alimentary canal. In the intestine the spores lose their 
 capsule, and penetrate the wall so as to lie partly or 
 entirely in the general body-cavity of the Crustacea. 
 Directly the spore, however, appears outside the 
 intestinal wall, it is attacked by leucocytes, which are 
 
 ' Vircho'ufs Archiv., vol. xcvi. p. 177. 
 G 2 
 
 Fig. 40. — Mono'?pora in various 
 stages. 
 
 X. Young conidium. — 2, 3. Bud- 
 ding conidia. — 4. Elongated 
 conidium.— 5. Spore. 
 
84 
 
 INFLAMMATION. 
 
 carried to the spot by the blood-stream. These cells 
 fix themselves on the spore, forming around it a collec- 
 tion of cells, which often fuse together into a Plas- 
 modium. Under this influence the spore undergoes a 
 series of remarkable changes. On being enclosed in 
 the leucocytes the spore first loses its regular contour, 
 becomes sinuous, and finally breaks up into a mass of 
 brownish granules in which it would be impossible to re- 
 cognise the degenerated spores if we had not studied the 
 mode of their formation. (Fig. 41, 1-4.) The proof 
 
 that this degeneration is 
 really due to a phagocy- 
 tic influence is furnished 
 by those cases in which 
 only half a spore is sur- 
 rounded by leucocytes, 
 the other half being 
 embedded in the in- 
 testinal wall, or even 
 projecting outside the 
 skin of the animal (Fig. 41, 5). In these cases it is only 
 the half surrounded by the phagocytes that undergoes 
 the changes just described, while the other half, which 
 is not exposed to the influence of these cells, remains 
 perfectly normal. 
 
 When a large number of spores 
 penetrates the body cavity of the 
 Daphnia at one time, the leuco- 
 cytes congregate round them, in 
 
 a manner resembling an infiltration or a cellular exsuda- 
 tion (Fig. 43, rt). It is in fact a sequence of events 
 exactly similar to that which we have seen produced by 
 a traumatic lesion. 
 
 41. — Spores of Monospora, surrounded 
 by the leucocytes of Daphnia. 
 
 Fig. 42. — Elongated conidium 
 of Monospora surrounded by 
 two leucocytes. 
 
LECTURE VI. 
 
 85 
 
 The phagocytic action of the leucocytes, so evident 
 and easily studied in the transparent Daphniae, destroys 
 the spores of the pathogenic microbe and prevents their 
 development, thus protecting the invaded organism. I 
 have succeeded several times in isolating infected 
 Daphnise and keeping them till they were fully restored 
 
 Fig. 43. — Hind part of a Daphnia. 
 a. Spores of Monospora surrounded by a mass of leuC0C3'tes. 
 
 to health, thanks to the destruction of the spores by 
 their phagocytes. If on the other hand the phago- 
 cytic action is inadequate, owing to the continued in- 
 crease in the number of spores swallowed or for an)' 
 other reason, the latter begin to germinate and give rise 
 to budding conidia. Although the parasite in its 
 vegetative form is also attacked by leucocytes, it 
 obtains the upper hand and the Daphnia inevitably 
 
86 INFLAMMATION. 
 
 succumbs in a short time to its attack. This is owing 
 to the fact that the conidia multiply too rapidly, and 
 also secrete some poison which dissolves the leucocytes. 
 Towards the end of the disease nothing but conidia are 
 to be seen circulating in the body of the Daphnia, all 
 the leucocytes having completely disappeared. 
 
 There can be no doubt that the whole history of this 
 disease of the Daphnise may be summed up as a struggle 
 between two living organisms, the parasitic cells and 
 the phagocytes. In spite of the extraordinary activity 
 of the former, it is the Daphnia in most cases which, 
 under the protection of its phagocytes, gains the upper 
 hand. We can in this way explain the fact that even 
 when an epidemic of Monospora is raging in a tank or 
 aquarium containing Daphniae, the number of these 
 remains as high as ever. While some individuals die 
 every day, stricken with the disease, the rest resist its 
 ravages and multiply, thus filling up the gaps caused by 
 the deaths. 
 
 Very different is the course of the diseases caused 
 by parasites which do not meet with any resistance on 
 the part of the leucocytes. To this class belong the 
 Saprolegnise. The spores of these fungi germinate on 
 the surface of Daphniae or other Crustacea (as e.g. 
 Branchipus) and put forth filaments which penetrate 
 into their body. The filament to effect an entry often 
 makes use of the little fissures or orifices produced by 
 various causes, which may be wounds or little canals 
 pierced by the spores of Monospora. Once inside the 
 body cavity of the Crustacea, the Saprolegnia continues 
 its development in the blood with which this is filled, 
 meeting with no obstacle from any side. The leuco- 
 cytes manifest great indifference towards the developing 
 
LECTURE vr. 87 
 
 mycelium, which in the end dissolves these cells and in- 
 evitably brings about the death of the infested animal. 
 If an epidemic of Saprolegnia; has once broken out in 
 an aquarium, we may be quite certain that it will not 
 stop until it has destroyed the whole of the specimens of 
 Daphniae or Branchipus present.' 
 
 Several of the diseases of Daphniae, such as those 
 caused by the bacteria Pasteuria raniosa ' or Spiro- 
 bacillus CienkowskiP or by the Sporozoa {p^brine and 
 others), meet with but feeble resistance on the part of 
 the phagocytes. As we should expect under the cir- 
 cumstances these diseases, when once established, are 
 never cured, and infallibly end in the death of the 
 animals attacked. 
 
 The feebleness of the phagocytic protection which is 
 so striking in Crustacea, is probably connected with the 
 thickness of their cuticular envelopes, which clothe not^ 
 only the external surface but also the intestine in these 
 animals. The chitinous cuticle is very tough and is 
 quite impermeable to most microbes. Thus we see 
 that the small Crustacea, such as certain Copepoda, 
 which are provided with a very hard covering, can get. 
 on perfectly well without phagocytes, and in fact do 
 not possess any corpuscles at all in their blood. 
 
 The insects, so for as concerns inflammation and 
 resistance to microbic infection, are exactly similar to 
 the Crustacea. Every kind of lesion produces in them 
 an accumulation of leucocytes round the injured spot, as 
 may be easily seen on cauterising the tips of the caudal 
 
 ' Branchipus and Artemia are subject to the disease produced 
 by the Monospora. The pathological phenomena in these cases 
 need further investigation. 
 
 ° Annates de Flnstitut Pasteur, 1888, p. 165. 
 
 * Ibid., 1889, p. 265. 
 
88 INFLAMMATION. 
 
 appendages in the larvae of Ephemeridae or other in- 
 sects. 
 
 Balbiani ' has pubh'shed some very interesting re- 
 searches on the effects of introducing bacteria into the 
 bodies of various insects and Arachnidae, and has found 
 that many saprophytic bacteria are pathogenic and even 
 fatal for a large number of these Arthropoda. But 
 while the insects rich in leucocytes, (such as certain 
 Orthoptera, especially the Gryllidse,) can completely with- 
 stand the introduction of a great number of bacilli, the 
 other kinds that are poor in blood and leucocytes (such 
 as the Lepidoptera, Diptera, and Hymenoptera,) are 
 extremely susceptible to infection by the saprophytic 
 fungi. The power of resistance possessed by the in- 
 sects belonging to the first order " must be ascribed to 
 the action on the bacilli exercised by two varieties 
 of cells. The firs^ of these is represented by the 
 blood-corpuscles which, by means of their pseudo- 
 podia, seize on the bacilli floating in the blood and 
 include them in their protoplasm, where they are quickly 
 destroyed ; the second variety, the elements of the peri- 
 cardial tissue, consists of large cells with multiple nuclei, 
 which surround the heart or dorsal vessel in the form of 
 plates or cords of cells, and are more or less highly 
 developed in different types. Of all the tissues of the 
 body, the pericardial tissue is the only one that has the 
 power of arresting the bacilli carried by the blood, and 
 of taking them into their protoplasm, where they are 
 destroyed as rapidly as in the blood-corpuscles." " 
 
 The insects, although so susceptible to infection by 
 the most widespread and apparently inoffensive of bac- 
 
 ' Comptes renins de l' Academic des Sciences, vol. ciii. p. 952. 
 " Loc. cit., p. 953. 
 
LECTURE VI. 89 
 
 teria, are nevertheless very rarely subject to epidemics 
 of bacterial origin. The cause of this is probably to be 
 sought in the fact that the bacilli lack the means of 
 getting through the thick cuticular wall which covers the 
 skin and lines the intestinal canal and tracheae of insects. 
 Besides the lethargy of silkworms (' flacherie '), discovered 
 by Pasteur,' in which infection takes place through the 
 intestine, there are certain other diseases of insect larvae 
 which are occasioned by bacteria. Thus the larvae of 
 Anisoplia austriaca in the south of Russia are some- 
 times invaded by a bacillus which, in its length and 
 curved shape, recalls the appearance of the anthrax 
 bacillus. At the beginning of the disease the affected 
 larvae are not to be distinguished from normal indi- 
 viduals, and it is only after the complete invasion of the 
 blood that they show signs of illness and shortly after- 
 wards die. 
 
 These diseases, however^ are not nearly so frequent 
 as those caused by the higher fungi or by the Sporozoa, 
 which are much better adapted than the bacteria for 
 penetrating the chitinous coverings of insects. In the 
 strength of their growth the former possess a powerful 
 means whereby to penetrate the cuticular walls, while 
 the Sporozoa have an amoeboid stage during which, by 
 virtue of their mobility, they are able to effect an entry 
 at even the most protected spots. 
 
 An observation of de Bary^ on the conidia of 
 Cordiceps viilitaris that had been englobed by the 
 leucocytes of caterpillars, leads us to conclude that the 
 conidia of fungi which multiply in the insect blood, 
 
 ' " Etudes sur la maladie des vers a soie," 1870, vol. i. 
 
 2 " Vergleichende Morphologic und Biologie der Pilze," 1884, 
 
 P- 399- 
 
90 
 
 INFLAMMATION. 
 
 sometimes meet with a certain amount of resistance on 
 the part of the phagocytes. But in the majority of 
 cases that have been examined to decide this point, the 
 mycehal filaments and conidia develop in the blood 
 without check or hindrance. I can vouch for this 
 especially in the case where Cleonus punctiventris, as 
 larva, chrysalis or perfect beetle, is invaded by the 
 Isaria destructor. The oval green spore germinates on 
 the surface of the body, and gives rise to a small fila- 
 ment. This latter experiences great difficulty in pene- 
 
 Fig. 44. — -A leucocyte of Cleonus, showing two phases of movement. 
 The conidia of Isaria lie by its side, and are not englobed. 
 
 trating the cuticle, which becomes brown round the 
 puncture made by the parasite. But as soon as this 
 obstacle is overcome, the filament gains access to the 
 body cavity where, bathed in the blood, it grows freely. 
 The leucocytes at times approach the filament or de- 
 tached conidia, but do not englobe a single one of these 
 parasites (Fig. 44, 45). The latter therefore speedily 
 invade the whole animal and transform it into the hard 
 mass, so characteristic of the dead bodies of the 
 insects that have perished from one of the varieties of 
 ' muscardine.' These pests, which have but one ob- 
 stacle, the tough cuticular wall, to overcome, are often the 
 
LECTURE VI. 91 
 
 cause of devastating epidemics among insects. We 
 may call to mind the losses that were formerly occa- 
 sioned by the 'muscardine' of the silkworm. The 
 epidemic disease caused by the Isaria destructor affects 
 several species of beetles, especially Cleonus punctiven- 
 tris. It often happens that more than half these insects 
 
 Fig. 45. — Freeconidia oi Isaria in very close promimity to some leucocytes 
 of Cleonus. 
 
 perish from the attacks of the parasite. These insects 
 do a large amount of damage to the beetroot, and the 
 owners of the beet plantations in south-west Russia 
 make their calculations as to the amount of seed that it 
 will be necessary to sow, according to the rate of mor- 
 tality among these insects caused by the " muscardine 
 verte." In fact it is now a rooted conviction that the 
 culture of beetroot in the district mentioned would be 
 
92 INFLAMMATION. 
 
 impossible were it not for the assistance afforded naturally 
 by the Isaria destructor. 
 
 The diseases of insects occasioned by Sporozoa, (a 
 well-known example being the p^brine of silkworms,) 
 have not yet been studied from the standpoint of the 
 comparative pathology of inflammation. We are 
 acquainted with the. microsporidium ' which is the 
 cause of the p^brine, and with its amoeboid condition 
 which enables it to attain access to various cells, such 
 as the young eggs ; but the question whether any con- 
 test takes place between the parasite and the phago- 
 cytes has not yet been investigated. In Daphnia, which 
 is equally subject to the attack of the microsporidia, 
 the resistance offered by the leucocytes is extremely 
 feeble, and is only displayed against the spores. The 
 amoeboid condition of the parasites is developed in the 
 close neighbourhood of the leucocytes, without exciting 
 them to take any part whatever in checking the course 
 of the disease. The microsporidia, which develop so 
 freely in the body cavity of Daphnia, finally invading 
 the entire animal, have absolutely no destructive action 
 on the leucocytes. The latter circulate in the blood, 
 now and then fastening on to the surface of the 
 parasites and dropping off again as if they had to do 
 with some harmless object. 
 
 In reviewing this chapter on the reactive phenomena 
 presented by the invertebrata that are provided with 
 amoeboid and phagocytic blood-corpuscles, we are 
 bound to conclude that in all these animals an agglomera- 
 tion of these cells is produced round any injured spots. 
 This inflammatory reaction takes place as a result 
 
 ' See especially Balbiani, " Le9ons sur les Sporozoaires," 1884, 
 p. 150, et seq. 
 
LECTURE VT. 93 
 
 of any sort of traumatism (cauterisation, introduction 
 of splinters, bites, &c.). It is seen also in the course of 
 certain infectious diseases, as for example tliat caused 
 in Daphnia by the presence of IVionospora. In the 
 case where the phagocytic inflammation occurs to any 
 considerable extent, the cell accumulation consists 
 chiefly of the leucocytes which are brought along by 
 the blood current and are attracted to the injured spot 
 in virtue of their sensibility. The lacunar circulation 
 aids the approach of the leucocytes and renders un- 
 necessary any special arrangements for the passage of 
 these cells, such as we find in the vertebrata. 
 
 As, however, the sensibility (chemiotactic or other- 
 wise) of the leucocytes very frequently remains negative, 
 highly favourable conditions then exist for the inroads 
 of all kinds of parasites. In these cases protection 
 against infection is chiefly provided for by the thick 
 chitinous integument with which the animal is invested, 
 so that we have, in the Arthropoda, a means of defence 
 analogous to that which we have seen to be possessed by 
 Nematoda and plants. In this type of invertebrata, 
 however, members entirely devoid of leucocytes are 
 extremely rare, the large majority of the Arthropoda 
 having a more or less highly organised army of these 
 defenders. 
 
LECTURE VII. 
 
 Vertebrata — Amphioxus — Embryos of AxolotI — Young larvse of 
 Urodela — Comparison with the invertebrata — Tadpoles — 
 Diapedesis — Migratory cells — Fixed cells — Phagocytic pro- 
 perties of leucocytes — Do fixed cells also functionate as 
 phagocytes ? — Transformation of leucocytes into fixed connec- 
 tive tissue cells — Fate of the leucocytes that do not undergo 
 this transformation — Evolution of inflammation in the organic 
 world. 
 
 Thk last survivor of the lower vertebrates, the Amphi- 
 oxus lanceolatus, is curiously distinguished from all its 
 congeners so far as regards its pathology. It possesses 
 no blood corpuscles whatever, and is only furnished 
 with a very small number of amoeboid connective tissue 
 cells. All attempts therefore to provoke inflammatory 
 phenomena in it have given only negative results. The 
 application of nitrate of silver, or an incision does not 
 excite any visible reaction. This is evidently due to 
 the fact that Amphioxus is possessed of a very tough 
 limiting membrane, which serves as an important means 
 of defence to the animal. In this respect it resembles 
 the Nematoda and other animals that are protected by 
 their chitinous cuticles, as well as most plants. 
 
 In order to obtain reactive phenomena analogous to 
 those which take place in most invertebrata possessing 
 a mesoderm, we might turn our attention to the class of 
 
LECTURE VII. 95 
 
 fishes, where we find inflammatory processes similar to 
 those that are known from the study of the higher 
 animals. Since, however, the fishes are ill-adapted for 
 investigation in the living condition, it is better to pass 
 at once to the amphibia. In this class the larval stages 
 serve as classical objects for researches of this nature, 
 the caudal fins of the larvae of Urodela (Tritons and 
 Axolotls) and the tadpoles of Batrachians offering by 
 far the best material for the investigation of inflammation 
 in vertebrata. 
 
 We will first consider the Urodela, which form the 
 lowest group of the amphibia. 
 
 In the embryo of Axolotl ' the rudiment of the fin is 
 completely devoid of blood-vessels and lymphatics. In 
 addition to the epidermis, it is composed of a layer of 
 mesodermic cells which are divided at an early stage 
 into two varieties : fixed cells with processes which 
 ramify like a stag's horn, and amoeboid cells with large 
 mobile processes having few or no branches. Although 
 the fixed cells form a majority of the elements of this 
 connective tissue, the migratory cells occur in fairly 
 large numbers. (Fig. 46.) 
 
 The embryos of Axolotl, if freed from the egg mem- 
 branes at the tenth or twelfth day of development, 
 live readily in the aquarium, and can be used for experi- 
 ments on inflammation. If in one of these embryos 
 (previously curarised), we touch the edge of the fin with 
 a fragment of silver nitrate, and at once wash this off 
 with a stream of saline solution, we obtain a small 
 limited burn ; or we may produce a small lesion of the 
 fin, by introducing a needle charged with powered car- 
 
 ^ I have always used the white variety, since these ar more 
 convenient for researches on inflammation. 
 
96 
 
 INFLAMMATION. 
 
 mine or indigo. By either method we kill a certain 
 number of cells, and lay bare a part of the tissue of the 
 fin, which takes up a certain quantity of water, so that 
 the adjoining cells, especially the stellate cells, become 
 altered and lose to a great extent their highly refract- 
 ing appearance and their vacuoles. A short time after 
 
 Fig. 46. — Connective tissue in the fin of an Axololl embo'O. 
 a. Amffiboid cells. 
 
 the operation we may see a certain number of migratory 
 cells making towards the injured spot, while the epider- 
 mis folds over and covers the wound. The next day 
 some of the amoeboid cells of the connective tissue 
 accumulate round the injured spot and englobe either 
 the coloured granules lying in the wound or the 
 debris of the destroyed cells (Fig. 47). In this collec- 
 tion of cells some are to be found in the process of 
 
LECTURE VII. 
 
 97 
 
 karyokinetic division. The number of mitotic figures 
 however is too small to permit of our ascribing many of 
 the cells assembled at the seat of injury to the division 
 of pre-existing ones. Moreover this hypothesis is un- 
 necessary since direct observation shows clearly that 
 it is the mobile cells which accumulate round the lesion. 
 The stellate connective tissue cells, which can be 
 watched from day to day in the same animal, are entirely 
 
 )t<iusn^ 
 
 yjHuM jc 
 
 Fig. 47. — Inflamed spot in an Axolotl embryo. 
 
 passive in their behaviour. The karyokinetic pheno- 
 mena seen in some of them, are not more pronounced 
 than those occurring in other parts of the fin. The 
 blood-vessels do not take any part in the changes con- 
 sequent on the injury. Although they form large 
 trunks in the tail, they are quite absent in the fin or are 
 present only as small tubes which have no connection 
 with the rest of the circulation. 
 
 Here then in the embryo of a vertebrate animal we 
 have the phenomena of reaction carried out by the 
 mobile cells of the connective tissue alone, without any 
 
 H 
 
98 INFLAMMATION. 
 
 intervention on the part of the blood-vessels or of the 
 white blood-corpuscles. There can be no doubt that 
 this process is analogous with the reaction which, as 
 we have seen, is the result of lesions in the Medusae and 
 the larvae of Echinoderms ; in both cases there is an 
 accumulation of the phagocytes of the connective tissue 
 round the seat of injury. 
 
 Phenomena of the same nature may be also observed 
 in the young larvae of Axolotl, which have blood-vessels 
 in their fins, as well in the larvae of Tritons, which are 
 also provided with capillary blood-vessels. Emigration 
 scarcely or never occurs, owing to the early vessels of 
 these larvae being too minute to allow of the passage 
 through their walls of the large leucocytes, impeded in 
 their movements as they are by the voluminous red 
 corpuscles. Hence arises a remarkable series of pheno- 
 mena. The vessels of the fin in the vicinity of the 
 injured spot remain passive, and do not even undergo 
 any appreciable dilatation, while the migratory cells 
 of the connective tissue travel towards the seat of the 
 lesion. 
 
 As the occurrence of an inflammatory reaction in 
 vertebra without the intervention of vessels is a fact of 
 great importance, I will devote further attention to the 
 same phenomenon in the young larva of Triton tczniatus. 
 The edge of the caudal fin of one of these larvae was 
 touched with a veiy small crystal of nitrate of silver, 
 and washed immediately with salt solution and then 
 with pure water, so that the lesion was confined to a 
 small group of epidermic and connective tissue cells. 
 The branched cells of the connective tissue that 
 were nearest to the injured spot, became less highly 
 refracting and absorbed fluid, so that their protoplasm 
 
LECTURE VII. 
 
 99 
 
 became vacuolated, while at the same time their processes 
 were drawn in and shortened. (Fig. 48.) Two amoeboid 
 cells in the adjoining parts of the connective tissue began 
 to move towards the seat of injury. In a subjacent 
 capillary loop the circulation was completely arrested. 
 Three-quarters of an hour after the application of the 
 
 Fig. 48. — Part of the caudal fia of a Triton embryo, a quarter of an hour 
 
 after tile appiication of nitrate of silver. 
 
 1-5. Fixed connective tissue cells. — a, b. Migratory cells. 
 
 nitrate, the amoeboid cells in the tissues nearest the lesion 
 had increased somewhat in number, and all were making 
 their way towards the cauterised spot. (Fig. 49.) Three 
 hours after the beginning of the experiment the branched 
 cells still retained their relative positions, but had re- 
 gained their normal refractive power, and hardly showed 
 any change in the shape of even their finest ramifica- 
 tions. The migratory cells continued to increase in 
 
 H 2 
 
100 
 
 INFLAMMATION. 
 
 number, but solely at the expense of the neighbouring 
 connective tissue, since in all the vessels the circulation 
 appeared to have resumed its normal course, and no 
 trace of diapedesis was to be seen. Two hours later 
 i.e. five hours after the cauterisation), the eschar formed 
 by the necrosed epidermis came away, disclosing a new 
 
 
 A 
 
 Fig. 49. — The same specimen, three-quarter.'; of an hour after cauterisation. 
 1-5. Fixed cells. — a, &, t. Migratory cells. 
 
 epidermic layer, beneath which lay a mass of migratory 
 cells. (Fig. 50.) Other mobile cells were proceeding 
 in the same direction, whereas the fixed connective 
 tissue cells retained their former positions. The cir- 
 culation was restored in the neighbouring vascular loop 
 and yet, as throughout the experiment, there was abso- 
 lutely no diapedesis. On the following day the injured 
 part was completely restored to a normal condition. 
 
LECTURE VII. lOI 
 
 The Stellate cells (Fig. 51) had resumed their wonted 
 
 Fig. 50. — The same spot five hours after cauterisati 
 1-5. The same fixed cells as in Fig. 47 and 48. 
 
 appearance and again presented their characteristic 
 antler-like processes. The migratory cells, of which a 
 certain number remained accumulated under the re- 
 
102 
 
 INFLAMMATION. 
 
 generated epidermis, were as sparsely scattered in the 
 connective tissue as in a normal animal. 
 
 On seeing how quickly this restitution took place, I 
 again cauterised the same spot, twenty-four hours after 
 the first experiment, with nitrate of silver but more 
 
 A rents sc 
 
 Fig, 51. — The same spot 24 hours after the cauterisation. 
 1-5. The same fixed cells as in Fig. 47-49- 
 
 severely than on the former occasion. Directly after- 
 wards the stationary cells underwent the same changes 
 as before ; they became larger, vacuolated, and less 
 refractive, while their processes were considerably 
 shortened. The circulation in several adjoining capil- 
 lary loops was arrested ; but, in spite of the severity of 
 the lesion, diapedesis only occurred to an insignificant 
 degree. During five hours of observation, I saw only a 
 
LECrURK VXI. 103 
 
 single leucocyte pass through the capillary walls. This 
 fact however did not hinder the accumulation of 
 migratory cells, a certain number of which made their 
 way towards the seat of injury. 
 
 These observations, which have been repeated for 
 several years with uniform results, demonstrate clearly 
 the possibility of an inflammatory reaction in vertebrata, 
 without the intervention of the vascular or nervous 
 system. These phenomena may therefore be classed 
 with the reactive processes in the Annelida. 
 
 Thus a genealogical tree of inflammation can be 
 drawn up, starting with the researches on the reactive 
 phenomena of the invertebrata, and completed by facts 
 observed in the embryonic and early larval stages of the 
 vertebrata. These facts prove that the reactive pheno- 
 mena ensuing on lesions are in their origin essentially 
 the same in the two great branches of the animal 
 kingdom. But, whereas in the invertebrata the pro- 
 cesses have remained stationary, in the vertebrata they 
 have become in the course of development much more 
 complex in character. Even in the older larvae of 
 Triton and Axolotl, which are provided with a larger 
 number of blood-vessels wide enough to allow the 
 passage of leucocytes, the inflammatory reaction takes 
 place in the classical manner that has been so frequently 
 studied during the last twenty-four years. The same 
 lesions still produce first an acceleration, then a slowing 
 of the blood-stream, followed by an accumulation of 
 white corpuscles in the peripheral zone and their emi- 
 gration and movement towards the injured spot. 
 
 The tadpoles of the different Batrachians are well 
 adapted for the investigation of these phenomena ; 
 some, such as those of Bombmator igneus, are distin- 
 
104 INFLAMMATION. 
 
 guished by their large fins, which are so transparent as 
 to admit of direct microscopic examination. If inflam- 
 mation be excited in the living tadpole by the applica- 
 tion of small fragments . of silver nitrate, by a simple 
 puncture or by any other means, all the stages of in- 
 flammation and regeneration can be followed under the 
 microscope for several days or weeks in succession. 
 Hence the great advantage of these researches over 
 those prosecuted on the mesentery or tongue of the frog. 
 If we wish to fix any stage, we have only to adopt 
 Ranvier's method,' and drop the whole animal into 25 
 per cent, alcohol. After it has been for a few hours in 
 this liquid, the epidermis may be detached by means 
 of forceps, the tail cut off and covered with a few drops 
 of an aqueous solution of vesuvine. After some minutes 
 the preparation is sufficiently stained, and may be 
 washed with water and examined just as it is, without 
 being dehydrated and mounted in balsam. 
 
 The tail of tadpoles shows marked anatomical differ- 
 ences compared with the fin of young larvae of Triton 
 or Axolotl. In the former the blood-vessels are much 
 more numerous and more richly branched, while the 
 amoeboid cells normally present in the connective 
 tissue are much less numerous. All the histological 
 elements, the cells of the connective tissue as well as the 
 red and white blood-corpuscles, are smaller. These 
 peculiarities in the tadpole favour the emigration of 
 leucocytes, which is very pronounced after the infliction 
 of the various lesions that I have practised, such as 
 section of the end of the tail, puncture by a foreign 
 body or cauterisation with nitrate of silver. Sometimes, 
 as soon as a quarter of an hour after the infliction of the 
 1 " Traitd technique d'histologie," 2" edition, 1889. 
 
LECTURE VII. 105 
 
 injury, we may see the beginning of inflammation, 
 accompanied by vascular dilatation and a considerable 
 emigration of leucocytes. In cases where the irritation 
 is more lasting, as after the introduction of a splinter, 
 the inflammatory emigration may be observed for 
 several days in succession. The result is an accumula- 
 tion of leucocytes round the seat of lesion, an accumula- 
 tion infinitely more extensive than that which occurs in 
 the young larvae of Triton and Axolotl. A comparison 
 of the reactive phenomena in these Urodela with those 
 in tadpoles of Batrachians demonstrates in the clearest 
 manner possible how much more marked the reaction 
 becomes when the vascular system co-operates. 
 
 It is very probable that the migratory cells of the 
 connective tissue also move towards the injured spot. 
 Their number, however, is so limited that they are quite 
 unimportant in comparison with the mass of emigrated 
 leucocytes. The fixed connective tissue cells undergo 
 the same changes that have been described in the case 
 of the Tritons. Immediately after the application of 
 the irritant, they swell up, become vacuolated and less 
 highly refracting, while their processes become shorter 
 and lose many of their branches. A short time later, 
 however, these cells resume their normal condition, in- 
 cluding their stationary position and other characteris- 
 tics. 
 
 The migratory cells which have collected round the 
 lesion in the young Urodela, as well as the numerous 
 leucocytes which have emigrated from the blood- 
 vessels in the tadpoles and in the older larvae of 
 the same Urodela, at once show marked phagocytic 
 properties. They greedily devour coloured granules 
 which have been either introduced on a foreign body 
 
I06 INFLAMMATION. 
 
 or simply rubbed into the wound ; they also englobe 
 granules of dead pigment-corpuscles or other cellular 
 debris. 
 
 In my first paper on inflammation in amphibia ' I 
 laid stress on the fact that the fixed ramified cells of the 
 connective tissue are also possessed of phagocytic pro- 
 perties. On examining fins which have been inflamed 
 for several days we may readily observe cells containing 
 foreign bodies, such as granules of carmine or broken- 
 down red corpuscles, in their protoplasm, although these 
 cells have antler-like ramifications and must there- 
 fore be regarded as characteristic connective tissue 
 cells. From these facts, which I have confirmed several 
 times, I concluded that these cells were phagocytes, 
 just as much as the migratory cells. But I have since 
 recognised that this deduction was erroneous. In spite 
 of numerous attempts, I have never succeeded in 
 observing the inclusion of foreign bodies by the proto- 
 plasmic processes of the connective tissue cells. My 
 researches on this point, which I have repeated for 
 several years, have convinced me that the fixed cells, 
 once definitely formed, never under any circumstances' 
 englobe granules of carmine or other substances. The 
 granules found enclosed in them have been taken up in 
 a previous stage of their development, when they were 
 still mobile phagocytes. These facts afford therefore 
 certain proof of the conversion of migratory cells into 
 fixed connective tissue cells. Although this conclusion 
 is opposed to the almost unanimous opinion of patho- 
 logists, it is nevertheless correct. 
 
 It does not, however, necessarily follow that all the 
 mobile cells which have collected at the seat of lesion. 
 1 Biologisches Centralblatt, 1883. 
 
LECTURE VII. 
 
 107 
 
 are transformed into fixed branched cells. A great 
 number of phagocytes do not undergo this change ; 
 many of them perish and are englobed by other phago- 
 cytes, as can be seen in every case a few days after the 
 onset of the inflammation (Fig. 52). Several of these 
 wandering cells penetrate into the epidermis, and escape 
 thence to the surface, where they meet a certain death 
 in the surrounding water. Others again pass into the 
 lymphatics (PI. I. Fig. i) and are 
 carried away by the lymph-stream. 
 A certain number of phagocytes 
 remain at the seat of the lesion, in 
 spite of the fact that complete 
 regeneration may have occurred. 
 In the cases where the irritant 
 remains in the tissue, as for in- 
 stance the small glass tube which 
 has been introduced into the con- 
 nective tissue of tadpoles, a num- 
 ber of the emigrated leucocytes 
 remain at the injured spot, some- 
 times surrounding the foreign body 
 for several months. 
 
 This description of inflammation in the tadpole may 
 be taken as a type of the same process in the other 
 Vertebrates, including mammals and man himself. 
 
 Fig. 52. — Phagocytes enclosins: 
 other phagocytes. From the 
 fin of a tadpole of Bombi- 
 7iator. 
 
 We have thus arrived at the final complication of 
 the inflammatory reaction as it occurs in the animal 
 kingdom. Before examining this pathological process 
 in detail, it would be advantageous to cast a look 
 back on the evolution of this important phenomenon. 
 
 Since the chief cause of inflammation — infection — 
 
I08 INFLAMMATION. 
 
 must be considered as a struggle between two organisms, 
 the parasite and its host, and since this struggle brings 
 about adaptations on both sides, we must admit that the 
 organism has elaborated means to defend itself against 
 its aggressors. If the unicellular beings, in which the 
 nature of the struggle is more evident, already possess 
 means of defence, we certainly cannot suppose that the 
 higher animals are deprived of equal powers. 
 
 The plants, which are incapable of motion, defend 
 themselves by the secretion of thick tough membranes, 
 to penetrate which the parasite has to adopt special 
 measures. Thus it must either secrete ferments which 
 dissolve the cellulose, or pierce the membrane by the 
 force of its growth, so that the plant is well defended 
 against a large number of parasites which are not 
 possessed of these powers. 
 
 The penetration of parasites into, or the action of any 
 other injurious agents upon vegetable cells causes their 
 death. But even if these cells are incapable of regenera- 
 tion, others, which remain intact or are even excited 
 by the irritant, multiply and take their place. In the 
 plant world, then, we have lesions and primary ne- 
 crosis ; regeneration also takes place, often to full re- 
 integration, but there is no such thing as inflammation. 
 
 The animal condition of some lower plants, especially 
 the Plasmodium of Myxomycetes, offers an exception 
 to this rule, since here we have a vegetable organism 
 capable of locomotion and cellular digestion. The 
 latter property, which consists in taking up and digest- 
 ing or excreting the deleterious matter, contributes to 
 the protection of the organism. Although it may 
 not constitute inflammation itself, at any rate it repre- 
 sents a primitive condition of this process. 
 
LECTURE VII. 109 
 
 Inflammation makes its appearance only in the 
 animal kingdom and undergoes a slow evolution, which 
 begins in the organisms that have a mesoderm. At 
 first it cannot be distinguished from a simple intra- 
 cellular digestion, effected by amoeboid and phagocytic 
 cells of the mesoderm. Thus in the Sponges the diges- 
 tive and the inflammatory functions are still united ; 
 but as soon as the endoderm becomes definitely 
 separated from the mesoderm, the two functions di- 
 verge. The endoderm now acts exclusively as a diges- 
 tive organ, while the mesoderm alone retains the power 
 of protecting the organism against injurious agents by 
 digesting them when possible. The mesodermic phago- 
 cytes preserve their property of intracellular digestion ; 
 this they effect either by fusing into plasmodia or by 
 collecting to form capsules round the parasites or other 
 foreign bodies. The phagocytic reaction is displayed 
 by all the mesodermic phagocytes. In this process 
 the prominent part is played in some cases by the 
 connective tissue cells, in others by the peritoneal cells 
 or by the corpuscles of the perivisceral fluid or of the 
 blood. In all these cases it is the phagocytes which 
 war against the aggressor by devouring, englobing and 
 digesting it. 
 
 It is apparent that the inflammation of vertebrates, 
 in which the defending phagocytes emigrate from the 
 vessels to proceed against offending bodies, is distin- 
 guished only quantitatively from the analogous phe- 
 nomena in invertebrates and must therefore be also 
 regarded as a reaction of the organism against dele- 
 terious agents. We must conclude that the essential 
 originating factor, the primum movens of inflammation 
 consists in a phagocytic reaction on the part of the 
 
no INFLAMMATION. 
 
 animal organism. All the other phenomena are merely- 
 accessory to this process, and may be regarded as means 
 to facilitate the access of phagocytes to the injured part. 
 
 The morbid phenomena properly speaking, such as 
 the primary lesion or necrosis, as well as the processes 
 of repair, do not form part of the inflammation and 
 must not be confounded with it. 
 
 The discussion as to which stage in the evolution of 
 the phagocytic reaction should have the name of in- 
 flammation applied to it, is really beside the point. 
 We might follow Strieker,' Roser,^ and others, in limiting 
 this term to the phenomena in which the vessels co- 
 operate. In this case the phagocytic reaction of the 
 older larvae of Urodela would fall under the head of 
 inflammation, while the same reaction in younger larvae, 
 where the phagocytes are furnished by the connective 
 tissue, would have no right to this term. If on the 
 other hand we wish to adhere to the etymological 
 meaning of the word, we can only apply the name of 
 ' inflammation ' to those cases where the phagocytic 
 reaction is accompanied by increased local temperature, 
 that is to say, we must reserve it exclusively for the 
 warm-blooded animals. The important points, with 
 which we are concerned, are the recognition of the 
 natural relations between the phenomena and the 
 tracing of their genealogical history. 
 
 This general conclusion, to which we have been led 
 by a comparative survey of the phagocytic reaction, 
 will, I think, facilitate a closer study of inflammation 
 in the vertebrata. 
 
 1 " Allgemeine Pathologie der Infectionskrankheiten." Wien, 
 1886, p. 112. 
 " " Entziindung und Heilung.'" Leipzig, 1886, p. 55. 
 
LECTURE VIII. 
 
 Varieties of leucocytes— Origin of these varieties — Mobility — 
 Phagocytic properties — Condition of englobed microbes — 
 Their vitality and virulence — Sensibility of leucocytes — 
 Tactile sensibility — Chemiotaxis — Buchner's investigations — 
 Leucocytosis — Intracellular digestion — Destruction of mi- 
 crobes, especially in immune animals — Action of leucocytes on 
 spores — Multiplication of leucocytes by direct and indirect 
 cell-division — Changes they undergo — Transformation oflobed 
 into single nuclei. 
 
 For the purpose of simplification, I propose to treat 
 the various parts of the inflammatory reaction in verte- 
 brates separately, beginning with a study of the principal 
 actors in this process — the leucocytes. 
 
 Of these cells, which include the colourless corpus- 
 cles contained in the blood and lymph, several varieties 
 may be distinguished. In the first place, we find small 
 leucocytes with a large round nucleus and a very 
 small amount of protoplasm forming a thin layer round 
 the nucleus. These cells (Fig. 53«) which are often 
 spoken of as lymphocytes because they occur in large 
 quantities in the lymphatic glands, stain readily with a 
 variety of colours, especially with the aniline dyes. The 
 latter stain the nucleus deeply, and the protoplasm only 
 faintly. The lymphocytes gradually grow until they 
 become large-sized leucocytes, provided with a single 
 nucleus, which contains a considerable proportion of 
 
112 
 
 INFLAMMATION. 
 
 nucleoplasm and is easily stained with the aniline 
 colours. The nucleus of these large leucocytes, how- 
 ever, does not stain so deeply, whereas their protoplasm 
 stains as well as or better than that of the lymphocytes. 
 Among these mononuclear leucocytes are some cells with 
 a round or oval nucleus (Fig. ^ib) and others with a 
 kidney- or bean-shaped nucleus. This kind of leucocyte 
 bears a great resemblance to certain fixed connective 
 tissue cells as well as to endothelial cells and cells of 
 the splenic pulp. On this account it is often difficult to 
 distinguish the mononuclear leucocytes from these cells, 
 
 especially when they 
 occur outside the 
 vessels. 
 
 The two next 
 varieties of leuco- 
 cytes on the other 
 hand may easily 
 be recognised, even 
 when surrounded 
 with all sorts of histological elements. The first of 
 these, the eosinophile leucocytes of Ehrlich, are provided 
 with a nucleus, generally lobed and of very variable 
 form. They contain in their protoplasm coarse granules 
 which do not stain with the basic aniline dyes, such as 
 fuchsine, methyl or gentian violet, methylene blue, 
 vesuvine and others, but readily take up the acid 
 aniline colours, especially eosin, which stains them a 
 rose-red. (Plate III. Fig. 2.) 
 
 The fourth variety of leucocytes, the most important 
 quantitatively and even qualitatively, is represented by 
 cells in which the nucleus is either lobed or composed 
 of several portions united by such delicate nuclear fila- 
 
 Flg. 53. — Three forms of leucocytes. 
 
LECTURE VIII. 113 
 
 ments (Fig, 53^) as to give the impression of a multi- 
 nucleated cell. Most of the nuclei are in this form, 
 although they are sometimes really multiple, thus 
 justifying the name of polymiclear leucocytes, which has 
 been given to this kind. The shapes of these com- 
 pound nuclei are very variable. They more frequently 
 resemble a trefoil or a raspberry, but they may occur in 
 the form of a ring. Besides the nucleus these leuco- 
 cytes possess an attraction-sphere composed of fine fila- 
 ments of achromatin and containing a small central 
 body of chromatin. This peculiarity has been lately 
 discovered by Flemming' in the leucocytes of salaman- 
 der larvae. (Plate II. Fig. 7.) 
 
 The polynuclear leucocytes differ from the other 
 colourless corpuscles in their reaction to aniline dyes, 
 which stain the nuclei very deeply, while the protoplasm 
 remains for the most part unaffected. The latter con- 
 tains granules, sometimes, as in man, in great number. 
 These can only be stained by a mixture of the acid with 
 the basic dyes, so that these polynuclear leucocytes are 
 often alluded to as neutrophile leucocytes." 
 
 Although the fact that there are several kinds of 
 leucocytes was shown by Max Schultze'' as long ago as 
 1865, an exact knowledge of the different forms dates 
 only from the discoveries of Ehrlich.'' On examining the 
 relative proportion of the various leucocytes in the blood, 
 
 ' Archiv fiir mikroskop. Analoinie, 1891, vol. xxxvii. p. 249. 
 Plates 13 and 14. 
 
 - In the rabbit and guinea-pig these cells are replaced by 
 amphophile ox pseudoeosinophile leucocytes. 
 
 ■' Archiv f. mikroskop. Anatomie, 1865, vol. i. 
 
 ■• The valuable papers of Ehrlich have been recently collected in 
 a single volume. " Farbenanalytische Untersuchungen zur His- 
 tologie und Klinik des Blutes." Berlin, 1891. 
 
 I 
 
1 14 INFLAMMATION. 
 
 it is found that the neutrophile cells are by far the most 
 numerous, forming three-fourths of the total amount. 
 
 It was at first thought that these varieties depended 
 on a difference in the origin of the leucocytes and that 
 the small cells were formed only in the lymphatic 
 glands, while the others were derived from the bone- 
 marrow. Leucocytes were accordingly divided into 
 two groups, lymphocytes and myelocytes. This classifica- 
 tion had to be abandoned for two reasons. In the first 
 place, small cells can also be formed in other organs, 
 such as the spleen and the bone-marrow ; and, in the 
 second place, the myelocytes do not originate solely in 
 the bone-marrow. Here again our most accurate know- 
 ledge is due to Ehrlich, who has shown that the eosino- 
 phile cells are produced specially by the bone-marrow ; 
 hence an abnormal number of these cells in the blood 
 serves to indicate that this tissue is primarily affected, 
 as in myelogenic leuchaemia. The ordinary polynuclear 
 leucocytes or neutrophile cells develop in the blood itself 
 at the expense of the small cells which have been formed 
 in various organs. This fact has been specially. pointed 
 out by Ouskoff.' 
 
 In considering this question of the origin of leuco- 
 cytes, it must not be forgotten that these cells are to 
 be found in the blood of the inferior fishes, such as 
 Cyclostomata, which possess neither lymphatic glands, 
 bone-marrow nor spleen, and in which the leucocytes 
 are derived from the mesodermic cells of the embryo 
 and perhaps also from the endothelium of the blood- 
 vessels. 
 
 The leucocytes, as Lieberkiihn showed, are mobile 
 cells which, like the amoebae, have the power of putting 
 
 ' " Le sang comme tissu," (Russian), St. Petersburg, 1890. 
 
LECTURE VIII. IIS 
 
 out protoplasmic processes and of moving from place 
 to place. All the leucocytes have this property ; but 
 it is not so well developed in the lymphocytes, which 
 are the youngest of the white blood-corpuscles. Neither 
 these nor the eosinophile cells are able to englobe 
 foreign bodies, and therefore cannot act as phagocytes 
 Hence it is probable that the granules, which are so 
 characteristic of the eosinophile cells and in reptiles 
 and birds are shaped like small rods or crystals, are not 
 taken up by the cells, but are manufactured in the body 
 of the cell itself On the other hand the two remain- 
 ing varieties of leucocytes, the mononuclear and the 
 neutrophile corpuscles are endowed with very pro- 
 nounced phagocytic properties. Even outside the 
 organism these amoeboid cells readily englobe a large 
 number of foreign particles with which they may come 
 in contact, and they may be often seen literally crammed 
 with all sorts of granules. Like the amcebse, they 
 swallow not only inert bodies, such as granules of 
 carmine or other substances that are insoluble in the 
 fluid surrounding the leucocytes, but also a large num- 
 ber of living organisms. Thus the leucocytes of the 
 frog englobe the bacilli which cause a septicaemia in 
 these Batrachians. That the bacilli are in a living 
 condition when swallowed, is shown by the fact that 
 they perform active movements although enclosed in 
 the nutritive vacuoles of the leucocytes.' The same 
 thing is observed if we introduce leucocytes filled with 
 bacteria into a medium which nourishes the bacilli 
 while killing the leucocyte. Thus on introducing the 
 leucocytes of the pigeon filled with anthrax bacilli (to 
 which the pigeon is very refractory) into bouillon, the 
 ' Vide Biologisches Ce?itralblat(, 1883, p. 562. 
 1 2 
 
Il6 INFLAMMATION. 
 
 bacilli grow, pierce the protoplasm of the cells, and 
 form well-developed filaments,' showing definitely that 
 the bacilli were englobed in a living condition. This 
 has also been proved in the case of other bacteria. 
 Thus the Vibrio Metchnikowii that has been taken up 
 by the leucocytes of immune guinea-pigs, will develop, 
 even in an exsudation obtained from the animal itself, if 
 the fluid be put under conditions unfavourable to the 
 life of the animal cells." The conclusion is further con- 
 firmed by the fact that masses of bacilli of tuberculosis, 
 of swine septicaemia (' rouget des pores ') and of mouse 
 septicaemia are englobed by the leucocytes of a large 
 variety of animals whether susceptible or refractory to 
 these diseases. 
 
 But although it may be clearly shown that the leuco- 
 cytes enclose living microbes, we must not conclude 
 that these cells devour all microbes indiscriminately. 
 Thus in many cases we find that the leucocytes of an 
 animal which is very susceptible to a certain kind of 
 bacteria, do not take up these bacteria at all, even 
 though they may be in contact with them. As an 
 example we may quote the behaviour of the leucocytes 
 of mice and guinea-pigs towards the bacilli of anthrax, 
 or that of the leucocytes of pigeons and rabbits towards 
 the bacteria of chicken cholera, or of the leucocytes of 
 guinea-pigs that are susceptible to vibrionic septicaemia 
 when brought in contact with the exciting agent of this 
 disease. 
 
 It is evident then that leucocytes can englobe viru- 
 lent microbes. This fact may be shown in another 
 way. The two kinds of leucocytes which are phago- 
 
 ' Annales de I'Institut Pastetir, 1890, p. 80. 
 ' Ibid., i89i,p. 471. 
 
LECTURE VIII. 117 
 
 cytic in their action, the large mononuclear and the 
 neutrophile leucocytes, vary in their behaviour towards 
 different species of microbes. Thus in man the mono- 
 nuclear leucocytes do not take up either the strepto- 
 coccus of erysipelas or the gonococci, whereas these 
 two microorganisms are readily englobed by the neu- 
 trophile polynuclear leucocytes.' This choice shows 
 that the microbes thus avoided by the mononuclear 
 leucocytes, are by no means inactive bodies, otherwise 
 they would be taken up by this variety of leucocytes, 
 just like any other inert matter. Leprosy bacilli, on the 
 other hand, are never englobed by the neutrophile poly- 
 nuclear leucocytes, but are readily devoured by the 
 mononuclear cells." 
 
 These differences must be due to a varying sensibility 
 to microbes on the part of the two kinds of leucocytes, 
 dependent no doubt in both instances on chemio- 
 taxis. 
 
 Some researches on phagocytosis in amceboid cells ^ 
 suggested that action at a distance must play some 
 part in these phenomena. We owe to Leber,* however, 
 the first clear exposition of the part taken in them by 
 the chemiotactic sensibility of the leucocytes. In his 
 experiments on keratitis produced by a crystalline 
 
 ' Virchow's Archiv, vol. cvii., 1887, p. 227.; 
 
 * Ibid., p. 228, and Sawtchenko, Ziegler's Beitr'dge ziir patholo- 
 (;ischen Anatomie, vol. ix., 1890, p. 252. 
 
 •* " Untersuchungen iiber die intracelluliire Verdauung," in the 
 Arbeiten des zool. Inst.zu Wien, 1883, vol. v. p. 159. 
 
 ■* Fortschritte der Medicin, vol. vi., 1888, p. 460. See also Leber's 
 large work, " Die Entstehung der Entziindung,'' published at 
 Leipzig in May, 1 89 1 . As my manuscript was already written when 
 this monograph appeared, I have not been able to quote from it so 
 often as I should have liked. 
 
Il8 INFLAMMATION. 
 
 substance extracted from cultures of Staphylococcus 
 aureus, he showed that the leucocytes at a distance 
 were attracted towards the point where this substance 
 had been introduced. On putting some small glass 
 tubes filled with this substance, into the anterior cham- 
 ber of the eye, they became filled with a mass of leuco- 
 cytes, although the tubes were so placed that the cells 
 had to move against gravity in order to effect an entry 
 into them. 
 
 This important discovery was the starting-point of a 
 series of researches which proved beyond question the 
 existence in leucocytes of chemiotactic properties 
 absolutely analogous to those of plasmodia and other 
 lower organisms. Lubarsch' showed that the leuco- 
 cytes of the frog are more readily attracted by living 
 bacilli than by the same bacilli if previously killed by 
 heating. Pekelharing '' then pointed out that the leuco- 
 cytes of this animal are much more strongly attracted 
 by the bacilli of anthrax than by a passive substance 
 such as cotton fibres. In this connection many facts 
 of great importance have been brought forward by 
 Massart and Bordet,^ who proved that the leucocytes of 
 the frog are attracted by a number of culture fluids of 
 various microbes, especially of the Staphylococcus 
 pyogenes albus, as well as by the pleuroperitoneal 
 transsudation of frogs that have been poisoned by ox- 
 bile. 
 
 Among the products of the oxidation of albumen, 
 
 ' Fortschritte d. Median, 1888, vol. vi. No. 4, and Centralblaif 
 fur Bacteriologie, vol. vi. Nos. 18-20. 
 
 Semaine medicate, No. 22, 1889, p. 184. 
 
 ■'' " Recherches sur I'irritabilite des leucocytes." Joum. pubt. 
 par la Soc. des Sci. med. et nat. dc Bruxelles, Feb. 3rd, 1890. 
 
LECTURE VIII. 119 
 
 examined by these authors^ leucin was the only one 
 found to exercise an attractive influence on the frog's 
 leucocytes, while others such as creatin, creatinin, 
 allantoin &c., did not give rise to any chemiotaxis. 
 
 Gabritchevsky,' in a research carried out in my 
 laboratory, has shown that the leucocytes of mammals, 
 especially of the rabbit, are much more sensitive to 
 chemical excitation than are those of the frog. He 
 pointed out moreover that whereas leucocytes are 
 strongly attracted by sterilised or living cultures of 
 most pathogenic and saprophytic bacteria as well as 
 . by papayotin, they are repelled by the most virulent 
 bacteria, such as those of chicken cholera, and by lactic 
 acid, ten per cent, solutions of sodium and potassium 
 salts, alcohol, chloroform, glycerine, jequirity, bile and 
 quinine. The leucocytes remain unaffected by many 
 other substances such as water, weak solutions of 
 sodium or potassium salts, peptone, phloridzin, &c. 
 
 Buchner," after confirming these facts on the chemio- 
 tactic properties of leucocytes, has endeavoured to de- 
 termine more precisely the nature of the substances 
 contained in bacterial cultivations which call these 
 properties into plaj^ In conjunction with Lange and 
 Rorner he found that the proteins of various species 
 [bacillus pyocyaneus, bacillus subtilis, the typhoid bacillus. 
 Staphylococcus pyogenes aureus and others) exercised an 
 attractive influence on the leucocytes of the rabbit. He 
 obtained the same effect with solutions of gluten-casein, 
 as well as with some alkali-albumens of animal origin. 
 Buchner considers that these experiments warrant the 
 conclusion that it is only the contents of bacteria, and 
 
 ' Amiales de I'lnstitict Pasteur, 1890, p. 346. 
 
 ^ Berliner klinische U'ochenschrift, 1890, No. 47. 
 
I20 INFLAMMATION. 
 
 not the products of their secretion, which exert a 
 chemiotactic influence on the leucocytes. We must 
 observe, however, that Buchner has not by any means 
 proved his contention. It is evident that the toxic 
 products of the bacteria must play the most important 
 part in evoking the phenomena of chemiotaxis. Now 
 it is precisely these products which adhere most strongly 
 to albuminoid and other bodies, so that we are still far 
 from being able to isolate them and study their effects 
 in a pure condition. On the other hand, the more 
 advanced products of disintegration, such as am.monia 
 and its salts, are not of such great importance in the 
 question of leucocytic sensibility in the animal body as 
 to permit of our drawing conclusions from the fact that 
 they exert no chemiotactic influence. And j'et, even 
 among these substances, we find a body, leucin, which 
 has the property of attracting leucocytes. This fact 
 was observed by Massart and Bordet, and has been 
 confirmed by Buchner himself. 
 
 Physiological chemistry is not yet sufficiently ad' 
 vanced to give a decision in the question raised by 
 Buchner. We have not sufficient evidence therefore to 
 justify us in accepting the dictum laid down by this 
 author, viz. that the chemiotaxis of leucocytes can only 
 be excited by dead or injui'ed bacteria, since only under 
 such conditions are their contents dissolved out into 
 the surrounding fluid. Even if Buchner's hypothesis 
 could be proved, we must remember that in every 
 medium tenanted by bacteria, there are a certain number 
 of dead ones among the living. The former would at- 
 tract the leucocytes, which would thus be brought in 
 contact with the living bacteria, and the final result 
 would be the same as if the living bacteria had them- 
 
LECTURE Virr. 121 
 
 selves exercised the chemiotactic influence. But besides 
 these considerations, there are other facts which tell 
 against this hypothesis, as, for instance, the greater 
 attractive power possessed by living bacteria on leuco- 
 cytes, as shown in the experiments of Lubarsch. 
 Further there are the still more important cases in which 
 the bacteria or other parasites have been englobed by 
 the leucocytes in the living condition. Moreover the 
 evidence brought forward by Buchner himself argues 
 against his idea that the leucocytes are not attracted by 
 active bacteria. In experiments made in conjunction 
 with Romer, Buchner found that injections of proteins 
 of bacteria, especially of the bacillus pyocyaneus, into a 
 vein, produced a pronounced general leucocytosis, the 
 number of white corpuscles in the blood being seven 
 times the normal amount. Now leucocytosis is a con- 
 dition of extremely frequent occurrence in the infectious 
 diseases generally. Although in some, such as typhoid 
 fever in man, the, number of leucocytes has not always 
 been observed to be increased^ yet in the vast majority 
 of the other infectious disorders this has been found to 
 be the case. Thus several observers ' have shown that 
 a pronounced leucocytosis accompanies infection by 
 anthrax in animals which die of this disease (such as 
 guinea-pigs, horses, oxen and others), although masses 
 of actively living bacteria occur at the same time in the 
 blood. 
 
 According to von Limbeck^ and P^e,' leucocytosis 
 
 ' Bollinger, " Milzbrand," 1872, pp. 2, loi. 
 
 ^ Von Limbeck, "Klinisches und Experimentelles iiber die eiit- 
 zundliche Leucocytose," Prag, 1889. 
 
 3 H. Pee, " Untersuchungen iiber Leucocytose," Berlin, 1890, 
 
 p 13- 
 
122 INFLAMMATION. 
 
 is a constant phenomenon in erysipelas in man. It is 
 at its height during the febrile period, while the blood 
 contains a number of living streptococci, and comes 
 to an end after the crisis, when only masses of dead 
 microbes are to be found in the organism. 
 
 This coincidence in time of the leucocytosis and the 
 most active condition of the bacteria has been also 
 shown by von Limbeck in the case of fibrinous pneu- 
 monia in man. The number of the colourless cor- 
 puscles present at any time in the blood corresponds 
 exactly with the temperature ; when the disease ter- 
 minates in crisis, the leucocytes decrease suddenly, 
 while in the cases where the temperature drops slowly, 
 (lysis), the fall in the number of leucocytes is also 
 gradual. 
 
 Working with dogs, von Limbeck pointed out that 
 the injection of Staphylococcus pyogenes aureus into the 
 knee-joint was almost immediately followed by a large 
 increase of the leucocytes in the bjood, before any 
 local symptoms had made their appearance. 
 
 We thus see that the special instance of chemiotaxis 
 known as leucocytosis, occurs at a time when the 
 greatest proliferation of microbes is taking place and 
 that the leucocytosis begins to diminish directly the 
 pathogenic bacteria die. We must remember also that, 
 according to other researches of Buchner, ' these same 
 proteins, which he considers to exist exclusively in the 
 interior of the bacterial cell, are the active agents in 
 producing fever, that is, the symptom which occurs 
 while the pathogenic microbes are actively multiplying, 
 and ceases with their death. 
 
 Like the plasmodium of Myxomycetes and many other 
 ' Berliner klinische WockenscJmft, i8go, No. 30, p. 673. 
 
LECTURE VIIT. 123 
 
 inferior organisms, the leucocytes have other forms of 
 sensibility besides that of chemiotaxis. Their tactile 
 sensibility, which serves them in the inclusion of foreign 
 bodies, is very highly developed. When the leucocytes 
 meet a resistant surface, they react by offering as large 
 a surface of contact as possible (Massart and Bordet). 
 By means of this property, the leucocytes are able to get 
 through the finest pores, and can penetrate elder-pith 
 and even such compact bodies as bone and ivory. The 
 physical changes in the surrounding medium, such as 
 variations in temperature, pressure, or rapidity of move- 
 ment of the liquid, and probably many others, must be 
 appreciated by the leucocytes. It would be very in- 
 teresting to make a systematic study of these physio- 
 tactic properties of the white corpuscles. 
 
 Leucoc3'tes which, by virtue of their chemiotaxis, 
 are attracted at a distance by microbes and other 
 particles, move towards these bodies and on coming in 
 contact with them englobe them owing to their tactile 
 sensibility. The ingested particles are now acted on 
 by the leucocyte. It is an old observation that red 
 corpuscles, when enclosed by leucocytes, partially dis- 
 solve, leaving a pigmented residue. It is also very easy' 
 to follow the changes undergone by pus-corpuscles in 
 the interior of leucocytes, where they gradually lose 
 their staining power and are finally converted into 
 scattered granules which are partially dissolved. 
 These changes are carried out by the protoplasm of the 
 leucocytes, and must be looked upon as an act of intra- 
 cellular digestion. This view is justified by the dis- 
 covery of ferments in leucocytes. Thus Rossbach ' has 
 
 ^ Deutsche medecin. Wochenschrift, 1890, p. 389. 
 
124 INFLAMMATION. 
 
 shown the existence of an amylolytic ferment in leuco- 
 cytes from various organs, especially from the tonsils, 
 and Leber ' has proved that the pus from a hypopion, 
 entirely freed from microbes, will digest coagulated 
 fibrin at a temperature of 25°. The same pus has also 
 the power of liquefying gelatin. This digestive power 
 can be destroyed by heat, showing that in this respect 
 the ferment of the leucocytes resembles all other fer- 
 ments. 
 
 The digestion of proteid substances by the leuco- 
 cytes is well shown by the gradual changes that take 
 place in the muscular fibres which have been englobed 
 by leucocytes in cases of acute muscular atrophy. The 
 presence of peptone in leucocytes, which has been so 
 often proved by Hofmeister, is suflnciently accounted 
 for by this fact of intracellular digestion and need not 
 therefore be referred, as done by this author, to an ab- 
 sorption by these cells of the peptone formed in the 
 alimentary canal. 
 
 Our knowledge of the conditions under which this 
 intracellular digestion takes place is still very incom- 
 plete. In a large number of experiments that I have 
 made on the absorption of granules of blue litmus by 
 leucocytes, I have seen the colour change to red only in 
 a few exceptional cases. ^ It appears then that diges- 
 
 ' " Die Entstehung der Entziindung,'' p. 508. 
 
 ' AniialesdeVInstitut Pasteur, 1889, p. 29. '^^\.'^a&^ {VirchovJ' s 
 Archiv, vol. cxxv., 1891, p. 448) thinks that in the cases where I 
 observed a conversion of the blue into red granules, the change was 
 simply an optical illusion. This he concludes from his own ob- 
 servations on the fate of litmus granules in the interior of leucocytes. 
 My researches, which extend over a series of years, leave me, 
 however, in no doubt as to the reality of the change in colour of the 
 litmus. I must add that the criticism by Netchaeffof my work 
 
LECTURE VIII. 125 
 
 tion is carried out by leucocytes in a neutral or alkaline 
 medium, as in the case of the phagocytes of the 
 Sponges. 
 
 Not only are the disintegrated muscle fibres or blood 
 corpuscles digested by the leucocytes, but in a large 
 number of cases the microbes that have been devoured 
 also undergo the same fate. This digestion of bacteria 
 in the interior of leucocytes can be most readily studied 
 in refractory animals. Nothing is more instructive to 
 ivatch than the changes undergone by the streptococci 
 of erysipelas in the leucocytes of white rats. The bac- 
 teria that have been englobed fuse into irregular masses 
 which do not stain completely, a process closely re- 
 sembling the digestion of the sulphobacteria in the 
 body of Stentor, discussed in the second lecture. 
 Anthrax bacilli are digested in the same way by the 
 leucocytes of a number of immune animals. The frog's 
 white corpuscles, instead of presenting a favourable 
 medium for the development of anthrax bacilli as 
 Koch ' and Petruschky " have asserted, not only prevent 
 their growth and development, but also kill and digest 
 these bacteria. All the phases of this digestion corre- 
 spond exactly with those of the digestion of the sapro- 
 phytic bacilli by amoebse (vide Chap. II,). As in this 
 case, a number of the bacilli in the interior of the 
 leucocytes are stained deeply by an old aqueous solu- 
 tion of vesuvine, which also colours other particles that 
 are being digested by the amoebse, as Bruno Hofer has 
 
 on intracellular digestion shows that this observer has never 
 examined the most classical examples of this process, more espe- 
 cially the digestion in Protozoa. 
 
 ' " Beitrage zur Biologie der Pflanzen," published by Cohn, 1876, 
 vol. ii. p- 300. 
 
 Zcituhrifl fur Hygiene, 1889, vol. vii. p. 75. 
 
126 
 
 INFLAMMATION. 
 
 also shown. This period during which the devoured 
 bacilli are deeply stained, is followed by a stage in 
 which the bacilli are only partially stained, and finally 
 even this property is lost and they remain absolutely 
 uncoloured. At last only the more durable cell-mem- 
 brane of the microbe is to be seen in the leucocyte, and 
 even this disappears at the end of a cer- 
 tain space of time. The bacilli which 
 have been englobed by leucocytes are 
 much more rapidly digested in the case 
 of mammals that are either naturally 
 refractory, as the dog and fowl, or 
 have been rendered artificially immune 
 against anthrax by vaccination, as the 
 rabbit. This fact is shown by the 
 researches of Hess ' as well as my 
 own." 
 
 It is easy to follow the digestion of 
 many other microbes within the 
 leucocytes. Vacuoles are often seen 
 to form around the bacteria that have 
 Fi?. 54.-An aiuhrax bccH swallowed, just as we have 
 
 bacillus, stained by . , . 
 
 vesuvine, in a leucocyte noticed m thc digestion of uutrient 
 
 of the frog. The two 
 
 figures represent two material by the protoplasm of the 
 
 phases of movement of .^ i i 
 
 the same cell. Protozoa and the Myxomycetcs. I 
 
 have been able to observe the changes undergone by 
 the spirilla of recurrent fever in the leucocytes of 
 monkeys,^ as well as those undergone by the vibrio of 
 septicaemia in the leucocytes of immunised guinea-pigs, 
 and those by erysipelas streptococci in the leucocytes 
 
 ' Virchow's Archiv, 1887, vol. ci.\. p. 365. 
 ^ Ibid., 1884, vol. xcvii. p. 502. 
 ■^ Ibid., 1887, vol. cix. p. 176. 
 
LECTURE Vlir. 127 
 
 of man/ &c. We are at present ignorant of the precise 
 manner in which this digestive and destructive action 
 is accomplished, and do not even know whether the 
 substance which kills the microbes is a ferment or not. 
 The fact that the ferments of the higher animals, such 
 as pepsin and trypsin, do not kill bacteria, is no reason 
 
 Fig- 55 — Four leucocytes from the frog, enclosing anthrax bacilli. Some are alive 
 and unstained, others which have been killed have taken up the dye. 
 
 for assuming that there may not be other ferments 
 which are capable of exercising a bactericidal action. 
 
 Some authors have denied to leucocytes the posses- 
 sion of digestive powers. Schafer^ comes to this con- 
 clusion from the observation that neither proteid sub- 
 
 ^ Virchow's Archiv, vol. cvii. p. 209. 
 
 ° British Medical Journal, 1882, No. 1134, p. 573. 
 
128 INFLAMMATION. 
 
 stances nor granules of fat or starch underwent any 
 change when ingested by the leucocytes of Tritons. 
 But as his researches were carried out on leucocytes 
 that had been removed from the body and immersed in 
 normal salt solution, they cannot be taken to refute the 
 facts that have been acquired from a study of the living 
 animal. Netchaeff has recently endeavoured to disprove 
 the intracellular digestion of bacteria by leucocytes, but 
 we have already shown how inadequate are the argu- 
 ments he brings forward. 
 
 It is undeniable then that leucocytes possess digestive 
 powers, and that in particular they are able to digest 
 microbes. But it does not therefore necessarily follow 
 that these cells kill and digest all the microbes they 
 englobe. In certain diseases the leucocytes take in a 
 number of bacteria, such as tubercle bacilli or the 
 bacilli of swine erysipelas or mouse septicaemia, a few 
 of which may be digested while the others resist the 
 digestive action of the leucocytes, multiply in the cells 
 and finally invade the whole organism. 
 
 In several cases where the leucocytes have not the 
 power of killing the microbe, they can nevertheless 
 hinder it from growing and exerting its toxic influence. 
 The best examples of this are afforded by bacterial 
 spores, which are endowed with great powers of resis- 
 tance. Anthrax spores are readily ingested by the 
 leucocytes of many species of animals, and, among 
 others, by those of immune animals such as the frog and 
 fowl. In spite of the fact that the spores germinate 
 and grow easily in the lymph-plasma of the two last- 
 mentioned, they are incapable of doing so in the bodies 
 of the leucocytes themselves, where they cannot exert 
 their injurious effects, although they preserve their 
 
LECTURE VIII. 129 
 
 vitality and their virulence often for a considerable 
 length of time. Once, however, that the vitality of these 
 cells has been extensively lowered, as may be effected 
 in fowls by cooling them or in frogs by heating them 
 arti^cially, the spores germinate within the dead or en- 
 feebled leucocytes, and infect the whole body.' These 
 experiments were repeated several times by Trapez- 
 nikoff" in my laboratory, and show in a most striking 
 manner the important and salutary part played by the 
 leucocytes, in cases where these cells are unable to kill 
 the spore, and where the fluids of the organism are 
 totally inadequate to protect the infected animal. 
 
 The proliferation of leucocytes is at present the sub- 
 ject of much discussion. The polynuclear forms, with 
 fragmented nucleus, can give rise to two new leucocytes 
 by mere division of their protoplasm. This pheno- 
 menon has been described by Ranvier " and observed by 
 several other workers, especially by Arnold.* The fact 
 of the occurrence of simple or direct division of poly- 
 nuclear leucocytes has at length been generally accepted, 
 although for a long time after the discovery of karyo- 
 kinetic or indirect cell division, it was considered doubt- 
 ful, and all cells were supposed to divide in the latter 
 way. The phenomenon of direct cell division once 
 established, people have gone to the opposite extreme 
 and denied to the polynuclear leucocytes the power of 
 dividing in any other manner. All the cases in which 
 karyokinetic division of leucocytes had been observed, 
 
 ^ See Wagner's paper on this subject, Annales de PInstitut 
 Pasteur, 1890, p. 570. 
 
 ^ Ibid., 1891, p. 362. 
 
 3 " Traits d'histologie technique," 2nd edition, 1889, p. 137. 
 
 ■* Archivfur mikroskopische Anatomic, 1888, p. 270. 
 
 K 
 
I30 INFLAMMATION. 
 
 from those of Peremeschko and Koultchitsky ' onwards, 
 were looked upon as open to doubt. Yet it has several 
 times been proved that leucocytes are capable of dividing 
 indirectly. Flemming^ has observed this phenomenon 
 on several occasions, and has recently confirmed it 
 in the case of the leucocytes of the salamander. 
 Spronck,^ too, has seen mitotic division of the leuco- 
 cytes in the blood of the rabbit and has found that at 
 any given time about 2 per looo of these cells are nor- 
 mally undergoing indirect cell-division. I can com- 
 pletely endorse this statement, since I have observed 
 undoubted mitotic division of the rabbit's leucocytes in 
 preparations made by Dr. Muskatbluth in my laboratory 
 at Odessa. I have on several occasions noticed the 
 same phenomenon in the migratory cells of the larvae of 
 Axolotl. In all the above cases it is the large mono- 
 nuclear leucocytes that divide indirectly, while the 
 polynuclear multiply by direct cell-division. All the 
 phases of the karyokinetic division of the leucocytes 
 that have emigrated from the vessels, and the conversion 
 of the new cells into amoeboid cells may be studied in 
 the caudal fin of the living larvae of the white Axolotl. 
 (Fig. 56.) 
 
 In the same way it has been denied that the leuco- 
 cytes possess the power of gradual transformation. The 
 fact that numbers of these cells are found dead in 
 exsudations has given rise to the idea that leucocytes 
 are in most cases destined to perish and are incapable of 
 forming constituent cells of any tissue. The lobed 
 shape of the nucleus in the polynuclear leucocytes has 
 
 ' Centralblatt fur medicmische Wissenschaften. 
 
 ^ Archiv fur mikroskopische Anatomic, vol. xxxvii., 1891, p. 249. 
 
 ^ Nederlandsch Tijdscrift voor Ceneeskunde, March 29th, 1889. 
 
LECTURE VIII. 131 
 
 even been taken as a sign of disintegration. Under 
 the influence of these views, many authors would refuse 
 to the leucocytes the power of conversion into fixed 
 connective tissue cells or even into epithelioid or giant 
 cells. In fact they now hold the exact opposite of the 
 theory formerly maintained by Cohnheim and especially 
 by Ziegler.^ The latter observer at first regarded the 
 granulation-cells as well as the epithelioid and giant 
 cells of tubercle as derived exclusively from mononuclear 
 leucocytes. He has now abandoned this position and 
 
 Fig. 56. — A migratory cell from the fin of a larva of Axolotl in process of division 
 by karyokinesis. 
 
 has adopted the opinion of his former opponents, who 
 believe the leucocyte to be incapable of giving rise 
 to any other form of cell. At the International Con- 
 gress held at Berlin in 1 890, Ziegler/ relying chiefly on 
 researches that had been made by NikiforofF in his 
 laboratory at Freiburg, stated that "the leucocytes do 
 not take any active part in the new formation of tissues." 
 
 ^ " Ueber die pathologische Bindegewebsneubildung," 1875. 
 
 ' Centralblatt fur allgemeine Pathologic, 1890, Nos. 18 and 19, 
 
 P 575- 
 
 K 2 
 
132 INFLAMMATION. 
 
 Marchand ' and Grawitz ^ quoted their own original 
 observations in confirmation of this view. According 
 to them the leucocytes that emigrate during an inflam- 
 mation are not converted into granulation-cells, but 
 perish in the exsudation, and are either absorbed by the 
 lymphatics or devoured by amoeboid cells derived from 
 the fixed connective tissue cells. Ribbert^ holds a 
 similar opinion. 
 
 Nikiforoffs' experiments, on which Ziegler chiefly 
 bases his view, were made on the subcutaneous con- 
 nective tissue of dogs, in which the formation of 
 granulation-tissue was provoked by the introduction 
 of drainage tubes. Although he affirms that this re- 
 generation of tissues is due exclusively to the pro- 
 liferation of the tissue-cells, and that the leucocytes 
 do not take any more active part in this process 
 than the red corpuscles or the fibrin, yet he himself 
 confesses he is unable to bring forward adequate proof 
 that the "cells originating from the blood-vessels are 
 incapable of further development and of conversion into 
 epithelioid cells and fibroblasts." ^ Nikiforoff adds,: 
 " Judging from my observations, I am inclined to the 
 view that in the granulation-tissue the mononuclear leu- 
 cocytes are converted by fragmentation of the nuclei 
 into polynuclear forms and are then taken up by 
 phagocytes." " This sentence is certainly much less 
 positive in tone than his conclusion quoted above and 
 than Ziegler's dictum. If we read Nikiforofif's account 
 of the observations themselves, we shall see that they by 
 
 ' Ceniralblatt fur allge7neine Pathologic, 1890, p. 577. 
 = Ibid., p. 578. 3 Ibid., p. 665. 
 
 * Ziegler's Beitrage zurfathologischen Anatomie, vol. viii., 1890, 
 p. 400. 
 
 ' Ibid., p. 415. '' Ibid., p. 421. 
 
LECTURE VIII. 
 
 133 
 
 no means prove his principal thesis, i.e. the non-par- 
 ticipation of the leucocytes in the formation of granula- 
 tion-tissue. The appearance of granulation-cells at a 
 period when the fixed connective tissue cells are in 
 process of multiplication, does not afford any proof that 
 the leucocytes are not also involved. Another argu- 
 ment brought forward by this author, the occurrence of 
 mitotic division, does not 
 in the least support his 
 conclusion, since we now 
 know that the leucocytes 
 are undoubtedly capable 
 of this mode of division. 
 It is not a matter for 
 surprise that Nikiforoff's 
 results should be want- 
 ing in accuracy and 
 clearness. The material 
 he worked with, the 
 granulations of the sub- 
 cutaneous tissue in dogs, 
 is not well adapted for j 
 the investigation of the 
 processes under discus- 
 sion. In order to obtain reliable results, we must make 
 use of a subject on which we can follow the course of 
 events from day to day. For instance if we take the tail 
 of a batrachian tadpole and injure it in any way and watch 
 it for several days or even weeks in the living condition, 
 as I have done in my researches, it may readily be seen 
 that in these animals the polynuclear cells are converted 
 by a fusion of their nuclei first into mononuclear and 
 then into typical branched connective tissue cells. 
 
 ig. 57. — Transition stages between amceboid 
 and fixed cells. From the tail of a tadpole of 
 Rana asilis. 
 
134 
 
 INFLAMMATION. 
 
 (Fig. 57.) In order to bring out this fact more clearly 
 it is advisable to inflict the wound with an instrument 
 
 XMiduL it. 
 
 Fig. 58. — Inflamed spot of the tail of a larva of Bombinaior igneus 8 days after 
 cauterisation. 
 
 powdered with carmine. The granules of this substance 
 which have been ingested by the emigrated leucocytes, 
 are found in the interior of the connective tissue cells, the 
 
LECTURE VIII. 13s 
 
 nature of which is easily recognised by their charac- 
 teristic antler-like processes. (Fig. 58.) 
 
 The view accepted at the Berlin Congress cannot be 
 maintained. It is not the new theory of Ziegler, but 
 his old way of thinking of 1875-1876 which is correct. 
 Although my personal observations have only been 
 concerned with amphibia, yet they agree perfectly with 
 the facts that have been ascertained in the higher verte- 
 brata by all observers, not excepting Nikiforoff himself. 
 
 In further support of my view I may quote the last 
 communication of Flemming,' who has also seen the 
 lobed or multiple nuclei of the migratory cells become 
 fused to a single rounded nucleus in the gills of sala- 
 mander larvae. 
 
 In the mammals, however, we have no proof that the 
 polynuclear can be transformed into mononuclear 
 leucocytes. Although the latter do certainly become 
 converted into epithelioid and giant cells, the poly- 
 nuclear variety seems unable to be thus transformed. 
 I need however only adduce the fact of the formation 
 of epithelioid and giant cells at the expense of mono- 
 nuclear leucocytes in the interior of the vessels in 
 rabbits which have received an intravenous injection 
 of tubercle bacilli. This phenomenon, which was de- 
 scribed by Yersin,^ has been likewise observed several 
 times by myself.' Tchistowitch '' in a research carried 
 out in my laboratory, has also witnessed all the 
 transition stages between typical mononuclear leuco- 
 
 ' Archiv f. mikroskop. Anatomie. Vol. xxxvii., 1891, p. 277. 
 ^ Annales de rinstitut Pasteur, 1888, p. 257. 
 ' Vtrchow^s Archiv, July, 1888, p. 88. 
 
 ^ Annales de rinstitut Pasteur, July, 1889, p. 347, and PL vi. 
 Figs. 5 and 7. 
 
136 INFLAMMATION. 
 
 cytes and epithelioid and giant cells in the pulmonary 
 alveoli of rabbits. 
 
 In summarising this review of the present state of our 
 knowledge on the subject of leucocytes, we may con- 
 clude that the two varieties of these cells which play the 
 principal part in inflammation— the mononuclear leuco- 
 cytes and the neutrophile cells — are elements endowed 
 with very considerable chemiotactic and physiotactic 
 sensibilities ; that they are capable of amoeboid move- 
 ments, and prone to englobe and digest different foreign 
 bodies, especially a number of living microbes. In the 
 amphibia at least, the polynuclear can be converted into 
 mononuclear leucocytes and can become fixed connec- 
 tive tissue cells. In the vertebrata generally, the mono- 
 nuclear leucocytes may undergo transformation into 
 epithelioid and giant cells. 
 
 All that has been said on the subject of leucocytes 
 refers equally to all migratory cells. 
 
LECTURE IX. 
 
 Endothelium of vessel walls — Their development out of the mobile 
 cells of the embryo — Development of the capillaries — Contrac- 
 tility of the endothelial cells— Star cells — Phagocytosis in the 
 endothelial cells — Fixed connective tissue cells — Clasmatocytes 
 of Ranvier — Ehrlich's cells — Active migration of leucocytes in 
 diapedesis — Experiments with quinine (Binz, Disselhorst) — 
 " Itio in partes " — Dilatation of the vessels — Theory invoking 
 the influence of the surrounding tissue — Influence of the 
 nervous system — Negative chemiotaxis of the leucocytes in 
 cases of severe infection. 
 
 Next to the leucocytes, the vessels and their endothe- 
 lial lining play the most important part in inflam- 
 mation. According to modern embryological research 
 on fishes ' the first endothelial cells of the vessels are 
 formed from the mobile cells on the surface of the 
 yolk sac. It is therefore not a matter for wonder that 
 the endothelial elements should have retained a few- 
 traces of their ancient power of movement. The at- 
 tention of investigators has been especially directed 
 towards the protoplasmic processes of the endothelial 
 cells of the vessels during the new formation of capil- 
 laries. Conical buds of protoplasm make their ap- 
 pearance on the vascular walls. A protuberance show- 
 
 ^ Vide H. Ziegler, " Die Entstehung des Blutes der Wirbelthiere.'' 
 Freiburg i. B., 1889. 
 
138 INFLAMMATION. 
 
 ing very slight signs of movement thus arises and 
 grows gradually longer. These processes often meet 
 each other, and form loops which are at first solid 
 and afterwards become excavated with the forma- 
 tion of a vascular channel. Many observers, such as 
 Strieker, ' Goloubew, ' Klebs, ' and Severini, " have 
 found that the endothelial walls of capillaries are to 
 a pertain extent contractile, — a still further proof of 
 a power of movement inherent in these cells. This 
 property must certainly play an important part in 
 the formation of stomata during inflammation, as 
 Klebs ' thought, and as I imagined and stated in my 
 first papers on inflammation." Arnold based his 
 
 Fig. 59,' — Passage of leucocytes through stigmata. 
 (After Arnold.) 
 
 original theory of stomlta on the supposed pre-existence 
 of these orifices under normal conditions. This is not 
 the case howeverj since they are formed only at the 
 
 ^ Mohschotfs Unterstichtmgm zttr Naiurlekre, vol. x. 
 
 ° Archiv f. niikroskop. Anat., 1868. 
 
 ■' " AUgemeine Pathologic," 1889, vol. ii. p. 384. 
 
 '' " La contrattilitk dei capillari," 1881. 
 
 •'' .See also Leydig, who in his pamphlet " Zelle und Gewebe," 
 1885, p. 17, has expressed the opinion that" in order to permit the 
 passage of the blood corpuscles, the minute pores may often dilate 
 into large orifices." 
 
 ' Biologisches Centralblatt, 1883, p. 564. 
 
LECTURE IX. 139 
 
 time of the inflammatory reaction, as he himself showed ' 
 when he put forward his stigmata theory. (Fig. 59.) 
 Although Arnold does not allude to the contractility 
 of the endothelial cells, he admits that " the spaces be- 
 tween the cells alter according to the conditions of 
 tension and diffusion, so that the respective positions 
 of the endothelial cells may vary very greatly." - He 
 holds that the intercellular spaces are filled with a fluid 
 or viscous material. These pores between the endothelial 
 cells, which open to permit of the passage of the cor- 
 puscles and liquid parts of the blood, and close after 
 their passage, might with greater justice be compared 
 to the pores of the ectoderm in the Sponges, which 
 open and close on the passage of the corpuscles sus- 
 pended in the surrounding water. In both cases we 
 have to do with contractile cells whose movements may 
 be aided by the neighbouring cells (fusiform cells of the 
 Sponges and contractile cells of the endothelium of the 
 vessel wall). 
 
 The contractility of the endothelial cells may be 
 further shown by the fact that in some cases these cells 
 may leave the vessel wall and by means of their amoe- 
 boid movements wander into the lumen of the vessel. 
 This may be observed after injection of tubercle bacilli 
 into the circulation of rabbits. Some time after the 
 injection has been effected, it is found that the vessel 
 wall in some places is composed solely of the adven- 
 titial the endothelial lining having entirely disappeared. 
 This is very frequently found to be the case in the 
 hepatic capillaries,, where, as Eberth and others have 
 shown, the endothelium does not present the typical 
 
 ' Virchow's Archiv, 1875, vol. Ixii. p. 487. 
 ^ Ibid., 1876, vol. xlvi. p. 104. 
 
I40 INFLAMMATION. 
 
 cell boundaries on treatment with nitrate of silver. 
 The endothelial cells of the hepatic capillaries become 
 readily detached from the outer coat, appearing in the 
 form of star-cells ' provided with a number of processes 
 which vary much in appearance. These cells, which 
 were first described by Kupffer,' were originally looked 
 upon as nervous elements on account of their shape 
 and thin elongated processes. It was soon ascertained 
 however that they belong to the endothelial tissue" 
 and that they have the power of ingesting various 
 granules. They are frequently found, especially in 
 frogs, enclosing masses of reddish pigment, and they 
 readily take up granules of pigment, if these be injected 
 into the blood, as Ponfick' and others have shown. 
 
 Star-cells, in common with the other endothelial 
 cells of the vessels, have also the power of englobing 
 various pathogenic and saprophytic microbes which 
 have made their way into the blood. This is the case 
 with the leprosy bacilli, which have been found, some- 
 times in groups, in the endothelial cells of the blood- 
 vessels. They often occur in such large numbers that 
 they completely obscure the nucleus. This discovery 
 was made by Neisser * and Touton, ^ who stated that 
 certain endothelial cells, when filled with bacilli, be- 
 come detached from the vessel-wall and are found free 
 in the lumen, Wyssokowitch * has since found that 
 
 ^ Archivf. mikroskop. Anat., vol. xii. p. 353. 
 
 ^ Asch. " Ueber die Ablagerung von Fett und Pigment in den 
 Sternzellen der Leber.'' Bonn, 1S84. 
 
 ^ Virchow's Archiv, 1869, vol. xlviii. p. i. 
 
 ■* Ibid., 1881, vol. Ixxxiv. 
 
 * Fortschritte der Medicin, 1886, No. 2, p. 48. ( Virchow's Archiv, 
 1886, vol. civ. p. 381.) 
 
 ^ Zeitschr. f. Hygiene, 1886, vol. i. p. i. See also the journal 
 Wratch, No. 44, 1891, p. 991. 
 
LECTURE IX. 
 
 141 
 
 many microbes, when injected into the circulation of 
 rabbits, may be traced in the endothelial cells of the 
 hepatic capillaries. This may be observed some time 
 after the introduction of tubercle bacilli into the vein of 
 the rabbit's ear ; a large proportion of the bacilli are 
 found again in the endothelial cells, especially in the 
 liver. The same thing occurs in natural infections in 
 man. Thus after death 
 from malaria, the en- 
 dothelial cells of the 
 liver are found filled 
 with the characteristic 
 parasite of this disorder.^- 
 Perhaps the most strik- 
 ing instance is furnished 
 by pigeons that have 
 been inoculated with the 
 bacillus of swine septi- 
 caemia. Enormous mas- 
 ses of these microbes are 
 found filling nearly the 
 whole of the endothe- 
 lial cells of the blood- 
 vessels, especially in the 
 liver. (Fig. 60.) As this bacillus, like that of leprosy, is 
 non-mobile, its presence within the protoplasm of the en- 
 dothelial cells can only be regarded as due to an active 
 ingestion on the part of the cells themselves. 
 
 I have laid weight on these details, since the contrac- 
 tility and the phagocytic properties of the endothelial 
 cells are extremely important facts in the question of 
 inflammation. Other varieties of endothelial cells are 
 endowed with the same properties. Thus the lym- 
 
 "ig. 60. — Endothelial cells enclosing the bacilli 
 of swine septicEemia, from a hepatic vein of a 
 pigeon. 
 
142 INFLAMMATION. 
 
 phatic endotlielial cells are still more often the seat 
 oi leprosy bacilli than those of the blood-vessels. 
 Ranvier/ many years ago, showed that in the guinea- 
 pig the endothelial cells of the inflamed omentum were 
 capable of taking up foreign bodies (such as myelin or 
 vermilion) that had been injected into the peritoneal 
 cavity. In a recent study " of the inflammatory changes 
 in the omentum, he has been able to demonstrate the 
 contractility of the cells forming its endothelial cover- 
 ing. Moreover Ribbert,^ in his memoir on the 
 inflammation and regeneration of lymphatic glands, 
 frequently lays stress on the contractility of the endo- 
 thelial cells of these organs. 
 
 The connective tissue cells certainly take some part 
 in inflammation, but this part is evidently much less 
 important or constant than was formerly thought, and 
 is far transcended by the part played by the leucocytes 
 and the endothelial cells of the blood-vessels. Further 
 we must remember that it is extremely difficult to make 
 sure of the co-operation of connective tissue cells in the 
 processes of inflammation. If they be studied in the 
 tails of tadpoles or of Urodelan larvae, the only changes 
 that we see in the fixed connective tissue cells are 
 alterations in their shape and their processes. These 
 changes last only a short time, and must therefore be 
 regarded as the immediate result of the lesion. These 
 facts prove that an absolutely typical inflammatory 
 reaction may occur without the fixed connective tissue 
 cells taking any important part in the process. These 
 cells, however, proliferate in an inflamed organ, and 
 
 ' " Lemons sur le systeme nerveiix,'' 1878, vol. i. p. 304. 
 
 ^ Comptes rendus de I' Acad, des Sciences, April 20, 1891, p. 842.. 
 
 ^ Ziegler's Beitr'dge ziir pathol. Anat., vol. vi. 1889, p. 205. 
 
LECTURE IX. 143 
 
 SO furnish a supply of new cells to replace those 
 killed by the original injury. Hence at a more or less 
 advanced stage of inflammation, karyokinetic figures 
 are to be seen in the connective tissue cells. 
 
 Ranvier' has recently called attention to a variety of 
 connective tissue cells corresponding to the " plasma 
 cells " of Waldeyer, which he designates as clasma- 
 tocytes. (PI. II.. Fig. 8.) These are very large 
 granular cells with processes and are formed from 
 leucocytes or migratory cells, which have increased in 
 size and lost their power of movement. During in- 
 flammation, according to Ranvier, the clasmatocytes 
 regain their former mobility, multiply rapidly and con- 
 tribute to the formation of pus-cells. On this view, the 
 clasmatocytes would form only a temporary constituent 
 element of the connective tissues, and would serve as a 
 source from which leucocytes might be drawn should 
 inflammation take place. These cells are present in 
 large numbers in the peritoneal tissue of tritons, frogs 
 and rabbits, but are not found in the larval stages of 
 amphibians. 
 
 Certain other cells of the connective tissue seem to 
 take some part in the process of inflammation. Thus 
 the basophile cells or " Mastzellen " of Ehrlich (PI. 
 III., Fig. 3) are often numerously represented in the 
 inflammatory products, although we are quite unable 
 to explain their presence under these circumstances. 
 Certain peculiarities in the reaction of these granules 
 to various stains seem to point to a possibility that 
 these cells may act as scavengers and clear away 
 the inflammatory products. Might we not indeed look 
 
 ' Coinptes rendtis de VAcad. des Sciences, April 27, 1891, p. 922. 
 
144 INFLAMMATION. 
 
 upon the granules with which they are filled as the 
 excreta of the other cells ? 
 
 Having considered the principal cellular elements that 
 take part in the inflammatory reaction, we may now 
 inquire how they carry out their functions. I need not 
 insist at length on the fact that the migration of the 
 leucocytes through the vascular wall is due to their own 
 active movements. In spite of all Cohnheim's en- 
 deavours, in spite of the general desire to refer all vital 
 phenomena to mechanical causes, the view that the 
 emigration is effected by the amceboid power of the 
 leucocytes has now found almost unanimous acceptance 
 (Thoma, von Recklinghausen, Lavdovsky and others). 
 This fact must strike anyone who compares the ease 
 with which leucocytes traverse the vascular wall, with 
 the purely passive diapedesis of the red blood-cor- 
 puscles. The latter often remain attached for an 
 indefinite time to the wall or are torn to pieces instead 
 of passing outside the vessels. Even Hering' who 
 originated the theory of the mechanical filtration of 
 leucocytes at the time of inflammation, admits the par- 
 ticipation of the amceboid movements of these cells, 
 although not attributing any great importance to them. 
 According to him the blood must either be circulating 
 or be subjected to a certain pressure for the filtration of 
 leucocytes to take place. Now the complete emigra- 
 tion of leucocytes may readily be observed after entire 
 stoppage of the heart in tadpoles which have, as some- 
 times happens during an experiment, died from the 
 effects of the curare administered to them. (Fig. 6i.) 
 
 ' Sitzungsherichte der k. Akad. der Wissensch. in Wien, 1868, 
 vol. Ivii., Part ii., p. 170. 
 
LECTURE IX. 
 
 I4S 
 
 In the very act of the passage of the leucocytes 
 through the vessel-wall, a certain chemiotactic or 
 physiotactic influence is manifested. If we agree with 
 Massart and Bordet ' in considering this emigration as 
 a mere effect of tactile sensibility, we cannot explain 
 the fact that the leucocytes remain inside the vessels in 
 cases when these are sufficiently dilated but when there 
 may be in the neighbourhood substances which have a 
 repellent action on leucocytes. Some time ago Binz 
 found that no dia- 
 pedesis of white cor- 
 
 puscles took place 
 through the frog's 
 mesentery after this 
 had been moistened 
 with a solution of 
 quinine. As quinine 
 acts poisonously 
 
 upon protoplasm, it ^'^■i'--°S?^^ni%tch'Srd's^^^^^^^ 
 was concluded that 
 
 it paralysed the leucocytes which were consequently 
 unable to pass through the vascular wall. The same 
 experiments were repeated by Disselhorst ^ who con- 
 firmed the fact that the diapedesis of the leucocytes 
 ceased, but was astonished to find that their movements 
 were not paralysed, for when removed from the vessels, 
 the leucocytes showed their wonted amoeboid activity. 
 In order to explain these facts, Disselhorst maintained 
 that the assistance of the vessel-wall was indispensable 
 for the passage of the leucocytes, and that the diapedesis 
 
 ' Journal published by the Soc. roy. d. Sciences medicales et 
 naturelles de Bruxelles, 1890, vol. v. 
 ^ Virchow's Archiv, 1888, vol. cxiii,, p. 108. 
 
146 INFLAMMATION. 
 
 was prevented by the action of the quinine upon the 
 wall. In his arguments, no account is taken of the 
 chemiotactic property of leucocytes, which was not at 
 that time an accepted fact. If this function be ad- 
 mitted, the effect of the quinine may be referred to a 
 negative chemiotaxis of the leucocytes, which, although 
 retaining their mobility, do not move towards the part 
 moistened with this substance. It follows that the 
 leucocytes are able^ while still within the vessels, to 
 detect the presence of the quinine. 
 
 The same explanation may be applied to the cases 
 of infection by highly virulent microbes where the 
 leucocytes, in spite of the dilatation of the vessels, do not 
 cross the vascular wall. Thus in the most rapidly fatal 
 infectious diseases, such as chicken cholera, septicaemia 
 of guinea-pigs and pigeons, and many others, there is 
 scarcely any emigration towards the invaded spot. In 
 these cases no diapedesis takes place although the most 
 favourable conditions exist, i.e. considerable inflamma- 
 tion and hyperaemia accompanied with serous and 
 even haemorrhagic exsudation. This may be explained 
 by a negative chemiotaxis manifested by the intra- 
 vascular leucocytes. This view is moreover borne out 
 by the fact that the microbes of the above-mentioned 
 diseases are not englobed by the leucocytes when 
 brought into contact with them. The occurrence of 
 chemiotactic action within the vessels necessarily follows 
 from the leucocytosis produced by the absorption of 
 various substances formed by the bacteria. 
 
 Massart and Bordet succeeded in stopping diapedesis 
 by depressing the sensibility of the leucocytes with 
 paraldehyde and chloroform, although these drugs did 
 not paralyse their power of movement. 
 
LECTURE IX. 147 
 
 It is highly probable that various other sensibilities, 
 besides those of touch and chemiotaxis, help in bring- 
 ing about the act of emigration. In lesions where the 
 necrosis of the cells is very slight and where conse- 
 quently the attraction due to chemiotaxis can only 
 be insignificant, we may probably regard the physical 
 change in the medium (difference of tensions, &c.) as 
 the chief factor in determining an attraction of the 
 leucocytes. 
 
 The diapedesis of the leucocytes is usually preceded 
 by the peripheral distribution of these cells in the dilated 
 vessel. This phenomenon, to which the name ' itio in 
 partes ' (Samuel) has been given, is generally considered 
 to be the most mechanical event in inflammation. It 
 was first sought to explain it by comparison of the 
 movements of the spherical white corpuscles with the 
 rapid motion of the flattened red blood discs ; but after- 
 wards the explanation suggested by Chklarewsky ' in 
 his account of a research carried out in Helmholtz's 
 laboratory, was adopted. This explanation is based on 
 the fact, discovered by Mach and Bondi, that insoluble 
 corpuscles when suspended in a fluid, increase the 
 density of the mixture. Now as the corpuscles circu- 
 late only in the axial portion of the vessel, while the 
 clear plasma flows along the sides, the density of the 
 latter must be less than that of the central portion. As 
 the leucocytes have a lower specific gravity than the 
 red blood- corpuscles, they are driven from the denser 
 axial current to the peripheral zone. 
 
 Although we must admit that the peripheral distribu- 
 tion of the leucocytes can only be due to a mechanical 
 action, since according to Massart and Bordet the 
 ^ PJViger's Archiv fur Physiologic, vol. 1. 
 L 2 
 
148 INFLAMMATION. 
 
 white corpuscles, after being anaesthetised by chloro- 
 form, still proceed towards the sides of the vessels, yet 
 we are unable to adopt the generally received explana- 
 tion of this phenomenon. It has been known since 
 Cohnheim's time and confirmed by Chklarewsky him- 
 self as well as by other observers that the peripheral 
 distribution, as well as emigration, goes on just as well 
 after the leucocytes have taken up vermilion granules. 
 As these leucocytes now contain a mercurial salt, they 
 must be not only heavier than they were before, but 
 also heavier than the red blood-corpuscles. Neverthe- 
 less the latter remain in the axial current while the 
 weighted leucocytes proceed towards the periphery. 
 
 The attraction of the leucocytes for the vascular wall 
 has been referred by Hering to their adhesiveness. 
 This theory has, however, found few supporters, for the 
 reason that leucocytes are not sticky and do not be- 
 come attached on account of their consistency but 
 solely by means of their amoeboid properties. Anyone 
 may be convinced of the fact that naked protoplasm 
 is not sticky by feeling the large Plasmodium of the 
 Myxomycetes, such as that of the Spumaria alba. 
 
 If it were possible to admit that the sensibility of the 
 leucocytes in Massart and Bordet's experiments was 
 not completely abolished by the action of chloroform, 
 we might attribute their accumulation in the peripheral 
 zone of the dilated vessels to a remnant of sensibility by 
 which they could still appreciate the differences in the 
 external medium, and so would move towards the calmer 
 regions where they are free to extend their protoplasmic 
 processes. 
 
 The vascular wall, by virtue of the power of contrac- 
 tion possessed by its endothelial cells, may no doubt 
 
LECTURE IX. 149 
 
 aid the passage of the leucocytes, although the latter 
 are quite capable of traversing non-contractile mem- 
 branes such as the epithelial tissues of vertebrates 
 (Stohr) or Ascidians. But the contractility of the 
 endothelial cells must have more influence in regu- 
 lating the passage of the red blood-corpuscles and the 
 liquid parts of the blood, especially in those cases where, 
 as in the infectious diseases mentioned above, the leuco- 
 cytes in consequence of negative chemiotaxis remain 
 in the blood, while the plasma and blood-discs exsude 
 through the vascular wall. 
 
 Although the leucocytes may emigrate from the 
 venules even when these are normal or only slightly 
 dilated, (as Recklinghausen first observed in tadpoles 
 under normal conditions,) it is nevertheless undeniable 
 that the dilatation of the vessels accompanied by slow- 
 ing of the blood-stream offers a very favourable condi- 
 tion for the occurrence of diapedesis. This dilatation, 
 which is so constant in the inflammation of vertebrates, 
 is a very complicated phenomenon in which several 
 factors take part. Cohnheim,^ who ascribed it exclu- 
 sively to the action of the vascular wall itself, sought to 
 prove his point by the following experiment. After 
 pinning out the tongue of a frog on a disc of cork, he 
 cut all the tissues connecting it with the body, with the 
 exception of the artery and vein. In this way he con- 
 sidered that he had eliminated any possible nervous 
 influences. He then applied some irritant to the 
 tongue, and found that an inflammatory reaction was 
 produced as usual. Cohnheim would not admit the 
 possibility of local nervous mechanisms in the vascular 
 
 1 " Gesammelte Abhandlungen." 1885, p. 423. 
 
150 INFLAMMATION. 
 
 wall, although it is very probable that such mechanisms 
 do play a part in these phenomena. 
 
 A considerable importance in the production of in- 
 flammatory hypersemia is generally ascribed to the 
 state of tension in the tissues, which is supposed to 
 react on the capillaries. This idea, started by Fluss, 
 was afterwards supported by Landerer,' who undertook 
 a series of experiments to show that the tension in the 
 tissues during inflammation is much lower than under 
 normal conditions. The capillaries surrounded by 
 these inflamed tissues are unable to withstand the 
 pressure of the blood and therefore dilate. The imme- 
 diate effect of this dilatation is an acceleration of the 
 movement of the blood, which rushes in to fill the 
 widened capillaries. But after a certain lapse of time, 
 as the surrounding tissues lose their elasticity more and 
 more, the blood-stream slows, thus producing the con- 
 dition of the circulation typical of inflammation. 
 
 This theory also suffers from the disadvantage that 
 it takes no account of the part played by nervous in- 
 fluence, which has been so much studied of late years. 
 Samuel ^ has in the following experiment strikingly 
 shown the importance of this influence. After cutting 
 the sympathetic nerve in a rabbit on one side, he 
 divided both auricular nerves on the other side of the 
 same animal. Paralysis of the vaso-motors at first pro- 
 duced congestion of the ear on the side of the divided 
 sympathetic and, as an indirect effect, a certain degree 
 of anaemia in the opposite ear. This anaemia increased 
 greatly after section of the sensory nerves, on account 
 
 1 " Ueber die Gewebsspannung," 1884, and "Zur Lehre von der 
 Entziindung." Leipzig, 1885. 
 " Virchow's Archiv, 1890, vol. cxxi., p. 396. 
 
LECTURE IX, 151 
 
 of the impossibility of reflex action by the uninjured 
 sympathetic. If, however, inflammation was produced 
 by treating both ears with hot water (at 54°), totally dif- 
 ferent results were obtained. The ear on the side of 
 the divided sympathetic became very hyperaemic and 
 highly inflamed. In the other ear, no hyperzemia 
 occurred, and inflammation was replaced by a stasis 
 ending in gangrene. This experiment proves the 
 influence of the nervous system on inflammatory re- 
 action and at the same time the beneficial effects of the 
 latter. On the side where' the inflammation is ex- 
 aggerated in consequence of the paralysis of the vaso- 
 motor nerves, the ear is speedily restored to a normal 
 condition, whereas the morbid phenomena are excep- 
 tionally severe on the anaemic side, where, owing to the 
 division of the sensory nerves, the processes of in- 
 flammation are unable to take their usual course. 
 
 In the rabbits in which Samuel divided the sensory 
 nerves (auricularis major and minor) on one side only, 
 the results were much less serious. If the ears of these 
 rabbits were dipped into water at 54°, in adult animals 
 the inflammation followed the normal course, and it 
 was only in young animals which had been treated 
 in this manner that he observed a retardation of the 
 inflammatory reaction and of the subsequent healing 
 process. 
 
 Similar experiments have been made by Roger. ^ 
 But instead of producing inflammation with hot water, 
 he made use of the cultures of the streptococcus of 
 erysipelas. He first divided the sensory nerves of one 
 ear of a rabbit and then inoculated both ears with these 
 bacteria. In the ear of which he had divided the 
 1 Comptes rendus de la Soc. de Biologic, 1890, No. 34, p. 646. 
 
I S 2 INFLAMMATION. 
 
 auricular nerves, the erysipelas lasted a long time, and 
 the healing processes, which were very slow, were 
 accompanied by some mutilation of the organ, whereas 
 in the normal ear of the opposite side the erysipelas ran 
 its usual course. 
 
 The contrary result was obtained by Roger ' on 
 inoculating rabbits with erysipelas, after dividing the 
 sympathetic instead of the sensory nerves on one side. 
 The paralysis of the vaso-motors thus produced gave rise 
 to a condition of hyperaemia, which exercised a favour- 
 able influence on the course of the erysipelas compared 
 with the other ear where the sympathetic was intact. 
 Inflammation began much earlier and the ear healed 
 more rapidly on the side on which the sympathetic had 
 been divided than on the other, although here also a 
 cure finally resulted. 
 
 These experiments confirmed those of several other 
 authorities, as Snellen, K. Danilewsky and others, who 
 had observed an improvement in the course taken by 
 the inflammation after division of the cervical sym- 
 pathetic nerve. Since section of this nerve on one side 
 also affects the ear of the opposite side by rendering it 
 anaemic, Samuel modified his experiment in the follow- 
 ing way. Instead of setting up inflammation in both 
 ears of the same rabbit, he made use of two different 
 rabbits, in one of which he divided the sympathetic on 
 one side, while the other animal was left uninjured. 
 He produced inflammation in the operated rabbit by 
 the application of water at 54° to the ear on the side of 
 the section, and in one ear of the normal rabbit by the 
 same means. He then found that the inflammation 
 
 1 Comptes rendus de la Soc. de Biologic, 1890, No. 16, p. 222. 
 
LECTURE IX. 153 
 
 ran a milder course in the control animal than in the 
 operated rabbit. 
 
 All these experiments prove that the nervous system 
 has a certain influence on inflammation, but at the same 
 time shows that the influence only serves to accelerate 
 or retard its course. While duly acknowledging that 
 it plays this part, we must be careful not to exaggerate 
 its importance, as by so doing we may fail to recognise 
 the true relations of the different factors which co- 
 operate to produce the inflammatory reaction. 
 
 It has been shown by Charrin and Gley ' that reflex 
 dilatation of the vessels can be prevented by injection 
 of 20 c.c. of the soluble products of the bacillus pyocy- 
 aneus into the circulation of a rabbit. After this injec- 
 tion, these authors observed that " the vasodilatation 
 thus produced develops more slowly, besides being less 
 marked and of shorter duration," (p. 734) than under 
 normal circumstances. In a later note ^ they proved 
 that this inhibitory influence on the vascular dilatation 
 is due exclusively to the volatile products contained in 
 the cultures and is not shared by the non-volatile con- 
 stituents, whether soluble or not in alcohol. These 
 facts they refer to a diminution of the excitability of 
 the vasodilator mechanism under the influence of these 
 volatile products. According to them the secretions of 
 the microbes diminish dilatation, hinder diapedesis and 
 in this way favour the course of the infection. They 
 seek to apply these results to the explanation of 
 immunity ; thus vaccination would act by strengthening 
 the part of the nervous system which presides over the 
 dilatation of the blood-vessels and over diapedesis, 
 
 ' Archives de Physiologic, No. 4, i8go, p. 724. 
 ^ Ibid., No. I, 1891, p. 146. 
 
154 INFLAMMATION. 
 
 Charrin and Gamaleia ' have likewise prevented the 
 dilatation of the blood-vessels by the injection of 
 vaccines or of the products of the micro-organism of 
 vibrionian septicsemia, as well as by simple injections 
 of 5-10 per cent, solutions of salt. 
 
 We cannot suppose that in the natural course of an 
 infective disorder there is a sudden entry into the blood 
 of 20 c.c. of bacterial products. In order then to 
 come to a decision on the subject, we must experi- 
 ment on animals which have been inoculated with the 
 microbes in question. If we introduce some bacillus 
 pyocaneus from a culture on agar-agar under the skin 
 of two rabbits, one of which has not been treated, while 
 the other has been vaccinated against the malady pro- 
 duced by this bacillus, the inflammation produced is 
 much more marked in the first rabbit than in the 
 vaccinated animal. In the former the bacterial pro- 
 ducts prevent neither the vascular dilatation nor the 
 increased local temperature, nor the serous exsudation 
 at the point inoculated ; and yet the diapedesis is less 
 than in the vaccinated rabbit, in which on the other 
 hand the vasodilatation and increase of temperature 
 are much less pronounced. The difference is still 
 more striking if we compare the results of injecting 
 the vibrio of septicaemia ( Vibrio Metchnikowit) under 
 the skin of the ear in two guinea-pigs, one of which is 
 susceptible, while the other has been rendered refrac- 
 tory by means of vaccinations. Here we find that the 
 reddening, heat and tumefaction are much more marked 
 in the susceptible than in the immune animal, whereas 
 diapedesis which is well marked in the vaccinated 
 animal scarcely occurs at all in the other. 
 
 1 Centralblatt f. allgemeine Pathologie, Nos. 18, 19, 1890, vol. i., 
 p. 588. 
 
LECTURE IX. 155 
 
 These facts show in the first place that, contrary to 
 the conclusions of Charrin and Gley, the microbes in 
 question do not hinder the vascular dilatation in sus- 
 ceptible animals ; and, in the second place, that dia- 
 pedesis may be almost or entirely absent in spite of 
 considerable dilatation of the blood-vessels. We must 
 conclude therefore that the most important factor in 
 the production of diapedesis is the sensibility of the 
 leucocytes themselves. If this sensibility be positive, 
 the leucocytes emigrate, in spite of the insignificant 
 dilatation of the vessels ; if it be negative, diapedesis is 
 not produced, however dilated the blood-vessels may be. 
 This fact may be easily shown by using for inoculation 
 a microbe exercising considerable positive chemiotaxis 
 on the leucocytes. If two guinea-pigs be inoculated 
 under the skin of the ear, one with a small amount of 
 tubercle bacilli and the other with the vibrio Metch- 
 nikowii, in the first animal the dilatation will be slight 
 and the diapedesis very considerable, while in the latter 
 the dilatation will be marked but will be accompanied 
 by little or no diapedesis. 
 
 The following facts may also serve as indirect argu- 
 ments against Charrin's and Gley's interpretation of 
 their results. In the most acute infections, in which 
 diapedesis is nearly or entirely absent, the presence of 
 leucocytes does not in any way hinder the existence and 
 multiplication of the bacteria, since in consequence of a 
 negative chemiotaxis the leucocytes do not englobe 
 these microbes. Under these circumstances it would be 
 quite superfluous to hinder diapedesis. If then, instead 
 of introducing some of these malignant bacteria (such 
 as the bacillus of chicken cholera in birds and rabbits, 
 or the vibrio of avian septicemia in guinea-pigs and 
 
1 56 INFLAMMATION. 
 
 pigeons) into the subcutaneous tissue, we introduce them 
 directly into the blood, the result is exactly the same. 
 The animals die in a very short time without the 
 occurrence of any phagocytosis ; and it makes no 
 difference to the issue whether, as in this case, the bac- 
 teria are in close proximity to the leucocytes in the 
 blood and spleen, or whether, as after subcutaneous in- 
 oculation, they are at some distance from these cells, 
 which remain in the interior of the blood-vessels. 
 
 We must therefore look upon the sensibility of the 
 leucocytes as the most important factor in inflammatory 
 diseases, not forgetting however the part played in the 
 inflammatory reaction in vertebrates by the sensibility 
 of the endothelial cells, as well as by that of the nervous 
 and other elements.' 
 
 ' See my lecture on " Immunity " in the British Medical 
 Journal of ]2Lnva.ry 31st, 189 1. After I had written this chapter, 
 Massart and Bordet put forward the same ideas in a study of the 
 chemiotaxis of leucocytes in infectious disorders {Annales de 
 r Instittit Pasteur, 1891, p. 417). They concluded from these ex- 
 periments that the absence of diapedesis in certain infectious 
 disorders is due not to a paralysis of the vasomotor centres, but 
 rather to a negative chemiotaxis of the leucocytes. In consequence 
 of this paper, a controversy has arisen between Charrin and Gley 
 on the one side and Massart and Bordet on the other {Comptes 
 rendus de la Soc. de Biologie, 1891, pp. 703-710. See also 
 Bouchard, Comptes rendus de PAcad. des Sc, 1891, pp. 524-529). 
 Without entering into the details of this discussion, we may state 
 generally that it is not a vasomotor paralysis but a negative 
 chemiotaxis which is the real cause of the absence of leucocytes at 
 the seat of the lesion in certain severe bacterial infections. Hertwig 
 (" Physiologische Grundlage der Tuberculinwirkung." Jena, 1891) 
 and Buchner {Munchener medicinische Wochenschrift, 1891) have 
 also pronounced in favour of this view. 
 
LECTURE X. 
 
 Chronic inflammations — Tuberculosis as a type of a chronic in- 
 flammation — Phagocytic nature of tuberculous cells — Destruc- 
 tion of tubercle bacilli by phagocytes — Power of resistance of 
 Meriones to tubercle — Leprosy. 
 
 Having passed in review the principal elements in- 
 volved in inflammation in the vertebrata, it remains to 
 inquire whether the part played by each is invariably 
 the same in all cases of inflammatory reaction. In 
 acute inflammation we find as a general rule vascular 
 dilatation accompanied by an active condition of the 
 endothelium of the vessel-walls and an exsudation with 
 diapedesis, that is to say, three events which concur in 
 producing a considerable afflux of leucocytes towards 
 the injured spot. Is the order of phenomena the same 
 in the chronic inflammations ? We often meet with 
 the idea that whereas in acute inflammation vascular 
 changes accompanied by diapedesis play the most 
 important part, in chronic inflammation the greatest 
 significance must be attributed to local phenomena that 
 take place in the tissues without the co-operation of the 
 cells of the blood or blood-vessels. 
 
 Chronic, like acute inflammations, are due to various 
 causes, which may be chemical, physical, or, as is more 
 usually the case, biological in nature. They may be 
 
158 INFLAMMATION. 
 
 brought about by the slow action of any injurious sub- 
 stance, as lead, phosphorus or alcohol, or by the pro- 
 longed action of heat or other harmful physical means ; 
 but they are more frequently the immediate effect of 
 microbes and their poisonous products. 
 
 We may take as an instance the chronic inflamma- 
 tion which results in the production of tubercles in 
 bacillary tuberculosis or in some other granuloma. We 
 choose this type, not only on account of its great im- 
 portance in pathology, but also because the fixed ele- 
 ments of the tissues have been regarded as the chief 
 factors which contribute to its formation. According to 
 Baumgarten's theory,' which is accepted by most patho- 
 logists, the phagocytic cells in general and the leuco- 
 cytes in particular take little or no part in the formation 
 of the tubercle itself. This he considers to be produced 
 by a proliferation of the local tissue cells, excited by the 
 presence of the tubercle bacillus in their neighbourhood. 
 Thus, according to this view, pulmonary tubercle is de- 
 veloped at the expense of the epithelial cells of the 
 alveoli ; hepatic tubercle at the expense of the 
 liver-cells and the epithelial cells of the bile-ducts ; 
 renal tubercle by a proliferation of the epithelial cells of 
 the urinary tubules, &c. The connective tissue also 
 contributes to the formation of tubercle by means of its 
 fixed cells and endothelial cells. The leucocytes take 
 no part in the formation of the tubercle until a later 
 period and hence must be regarded as of merely 
 secondary importance. According to this theory, 
 tuberculosis would not be a true inflammation, other- 
 wise we must admit that a chronic inflammation may 
 
 ^ " Tuberkel und Tuberkulose." Berlin, 1885. Reprinted from 
 the Zeitschrift f. klin. Medicin. 
 
LECTURE X. 159 
 
 be developed almost entirely independently of the 
 phagocytes or of the leucocytes. 
 
 The formation of tubercle may be accelerated by in- 
 jecting bacilli of avian tuberculosis into the veins of 
 rabbits, which, as is well known, are very susceptible to 
 this disorder. At the end of a few days, microscopic 
 tubercles are developed which may serve as a type of 
 this class of neoplasm. If we examine the liver, we shall 
 see that the epithelioid and giant cells of the tubercles 
 are formed solely at the expense of the phagocytic ele- 
 ments, that is to say, of the large mononuclear leuco- 
 cytes and of Kupffer's star-cells, derived from the 
 vascular endothelium. No hepatic or epithelial cells 
 ever contribute to the production of tubercle. It is true 
 that their nuclei may occasionally be seen about to 
 undergo karyokinetic division ; but this proliferation 
 has no direct connection with the formation of tubercle, 
 and merely serves for the regeneration of the hepatic 
 cells themselves. 
 
 Hepatic tubercle is the result of an accumulation, not 
 a multiplication of the mesodermic phagocytes, which 
 collect in masses and make up the whole of the ori'ginal 
 tubercle. These phagocytes contain bacilli which have 
 been englobed by the amoeboid cells. Several of the 
 epithelioid cells fuse together to form giant cells, while the 
 tubercular elements are reinforced by a large number of 
 lymphocytes and mononuclear leucocytes. Although 
 the former are not yet leucocytes, they soon become 
 converted into epithelioid cells, which represent one 
 variety of leucocytes. 
 
 Hepatic tubercle is therefore not derived from various 
 sources, as Baumgarten maintains, but is produced by 
 the amoeboid and phagocytic cells of the mesoderm. 
 
l6o INFLAMMATION. 
 
 The mode in which pulmonary tubercle is developed 
 confirms this view. This variety of tubercle is formed 
 by the endothelial cells of the blood-vessels together 
 with the leucocytes, and is the result not of the proli- 
 feration but of the accumulation of these cells, which is 
 effected by means of their amoeboid movements. The 
 cells of these tubercles likewise show their phagocytic 
 properties by englobing the tubercle bacilli. In cases 
 where ' dust-cells ' (' Staubzellen ') contribute to the 
 formation of tubercles, we have again to do with phago- 
 cytic elements derived from mononuclear leucocytes.' 
 
 Tubercles of the spleen and of the lymphatic glands 
 are likewise formed by an agglomeration of the larger 
 phagocytes of these organs — phagocytes possessing a 
 single large nucleus. In guinea-pigs and marmots 
 {Spermopkilus) a study of the neoplasms produced by 
 the bacillus of avian or human tuberculosis further con- 
 firms the statement that tubercle is composed of a collec- 
 tion of phagocytes mesodermic in origin, which move 
 towards the spot where the bacilli are situated and englobe 
 them. The phagocytes retain their condition of epithe- 
 lioid cells, or are transformed into giant cells. The 
 latter can also be formed in another way, although in 
 any case the final result is a large mass of protoplasm 
 enclosing a number of nuclei. In some animals, as the 
 marmots, the nuclei proliferate by a process of budding ; 
 more frequently however the multiple nuclei are 
 derived from a number of cells which have fused to 
 form a plasmodium. It is possible that the nuclei may 
 
 ' See Tchistovfitch, Annates de Vlnstitut Pasteur, 1889, p. 337. 
 Afanasieff, in an investigation carried out in my laboratory on the 
 development of pulmonary tubercle in the rabbit after inoculation 
 of tubercle bacilli from man, proved that here also the tubercle is a 
 purely phagocytic product. 
 
LECTURE X. l6l 
 
 also multiply by karyokinesis, but this has never been 
 demonstrated with certainty. 
 
 It is a well established fact that leucocytes take part 
 in the formation of tubercle, but these leucocytes belong 
 to the mononuclear variety. The polynuclear cells 
 englobe the tubercle bacilli readily but perish after a 
 short time, and then with the microbes they contain, are 
 eaten up by various mononuclear phagocytes which 
 may be classed together under the term oi macrophages . 
 These latter cells have a much greater power of resis- 
 tance^ and in some cases are even capable of destroying 
 the tubercle bacilli. Thus after inoculating marmots 
 with avian or human tubercle I have observed a very 
 characteristic degeneration of the bacilli in the interior 
 of the epithelioid and giant cells of these animals, which 
 are but little susceptible to tuberculosis.' The bacilli, 
 evidently under the influence of these cells, swell up 
 and gradually lose their power of retaining colouring 
 reagents. Sometimes the peripheral part but more 
 often the central part of the bacillus is the first to lose 
 its coloration. Later on the bacillus becomes converted 
 into a yellowish sausage-shaped body, enclosing a very 
 delicate canal, and the altered microbes collect to form 
 a conspicuous brownish mass, having the characteristic 
 aspect of a fragment of amber. None of these changes 
 
 ' See my article in Virchow's ArcMv, 1888, vol. cxiii., p. 63. 
 The facts brought forward in this paper concerning tubercu- 
 losis in marmots refer to the bacillus of avian tuberculosis. 
 Later experiments have shown me that this animal is more suscep- 
 tible to the bacilli of human tuberculosis, although, even in this 
 case, a certain number of bacilli are destroyed in the cells and 
 converted into yellow particles. At the Congress in London, 
 Bardach brought forward facts showing the high degree of suscep- 
 tibility possessed by the marmots for the virus of human tubercu- 
 losis, which he made use of in his experiments. 
 
 M 
 
1 62 INFLAMMATION. 
 
 are ever observed in artificial cultivations, even though 
 a number of dead bacilli are present, nor indeed any- 
 where outside the tubercular cells. They must be there- 
 fore regarded as resulting from the phagocytic action 
 of these cells, and recall the degenerative phenomena 
 which have been described in an earlier chapter as taking 
 place in cysts of the Gregarinse and the larvse of the 
 Nematoda, when surrounded by the phagocytes of 
 Lumbricus. In all these cases we have to do with 
 abnormal secretions poured out by the parasites, in order 
 to resist the influence of the phagocytes which enclose 
 or surround them. 
 
 Bacilli have also been ascertained to undergo changes 
 similar to those just described, within the giant cells of 
 rabbits, and very occasionally within those of guinea- 
 pigs. On the other hand I have never been able to 
 detect bacilli which have been destroyed by these means 
 either in the tuberculosis of Bovidse or of man. And 
 yet the resistance offered by the organism in such cases 
 is often of a very marked character. Calcareous de- 
 generation of tubercle has long been recognised as a 
 means by which tuberculosis in human beings may be 
 cured. In order to give a better idea of this reactive 
 phenomenon, I will describe the manner in which the 
 Algerian rat (Meriones) succeeds in resisting the action of 
 the tubercle bacillus. This rodent, although not entirely 
 immune against tuberculosis, receives it in a milder 
 form than many of the allied species. If inoculated 
 under the skin or even in the eye with a culture of the 
 bacilli of human tuberculosis, this animal withstands 
 the effects of the disease for many months. 
 
 If an Algerian rat that has been inoculated six or 
 eight months p^-eviously, be killed, a large number of 
 
LECTURE X. 
 
 163 
 
 tubercles are found in the abdominal organs, the lungs 
 and the lymphatic glands. These tubercles, however, 
 in the majority of- cases, do not present any necrosis or 
 caseation. The tubercular tissue composed of living 
 cells encloses bacilli, the majority of which have under- 
 gone a very remarkable degeneration, which deserves 
 our further consideration. In this animal the main 
 struggle goes on in the spleen, which is studded with 
 small tubercles consisting of non-necrosed epithelioid 
 and giant cells. The 
 former cells enclose a 
 small number of ordi- 
 nary tubercle bacilli, 
 while the giant cells 
 contain characteristic 
 calcareous bodies. (Fig. 
 62.) When examined 
 under the microscope 
 they are seen to be high- 
 ly refracting and in the 
 majority of cases to have 
 the shape of a figure 
 of eight, although some- 
 times they are simply rounded or irregular in form. 
 Under the influence of acids, the lime salt (phosphate of 
 lime) may be dissolved out, leaving a skeleton consisting 
 of a larger or smaller number of fine concentric layers. 
 (Fig. 63.) 
 
 These calcareous bodies have a strong resemblance 
 to the formations described by Schuppel ' as occurring 
 
 Fig. 62.- 
 
 -Tubercle from Weriones enclosing 
 a calcareous body. 
 
 ' " Untersuchungen iiber Lymphdriisen-Tuberkulose.'' Tubin- 
 gen, 1871, p. 104 and PI. I., Fig. 3, 4. 
 
 M 2 
 
164 
 
 INFLAMMATION. 
 
 in scrofulous glands, and observed by several authors ' 
 in many cases of tubercle of the lymphatic glands in 
 man. (Figs. 64, 65.) But whereas the origin of the 
 striated calcareous bodies in man is still quite obscure, 
 it can be demonstrated in Meriones with the utmost 
 facility. The examination of cover-glass specimens or 
 sections doubly stained by the methods of Gram or 
 Ziehl, shows at once that these bodies are the result of 
 a degeneration of the tubercle bacilli in the interior of 
 the giant cells. In the early stages, the bacilli stain in 
 the usual manner, and do not present any noticeable 
 
 
 
 Fig. 63. — The same cal- 
 careous body treated 
 with a dilute acid. 
 
 Fig. 64. — Tubercle contain- 
 ing a calcareous body. 
 From a human scrofulous 
 gland. (After Schiippel.) 
 
 Fig. 65.— Calcareous 
 body fram a mesen- 
 teric gland of man. 
 (After Ziegler.) 
 
 features. But by the side of these, we see other giant 
 cells (Tab. III., Fig. 4), in which the bacilli {a) are 
 covered with a fairly thick layer of an amorphous, 
 colourless substance {6) . This secretion becomes more 
 and more abundant so that the bacilli are finally sur- 
 rounded with a number of concentric layers. In many 
 cases these microbes may be stained with gentian 
 violet or fuchsin ii^ the usual manner, but more fre- 
 quently they lose this power and take up the comple- 
 mentary stain. (Tab. III., Fig. 5.) Sometimes in the 
 centre of a calcareous body, we meet with a bacillus 
 divided into two, half of which still retains the primary 
 
 ' Ziegler, " Lehrbuch der patholog. Anat.," 6th Edition, 1890, 
 vol. ii., p. 98, Fig-. 50. 
 
LECTURE X. l6S 
 
 coloration while the other has already lost it. (Tab. 
 III., Fig. 6.) Thus passing through a series of inter- 
 mediate changeSj the bacillus arrives at a condition 
 when it will no longer take the stain, although its 
 contour is still well-marked. (Tab. III., Fig. 7.) 
 Later on, however, they are not to be distinguished 
 by their coloration from the surrounding substance, 
 (Tab. III., Fig. 8) and finally disappear completely. 
 (Tab. III., Fig. 9.) In the last stage, which is most 
 frequently met with, we see nothing but the stratified 
 calcareous bodies. 
 
 The concentric membranes, which are left after the 
 removal of the phosphate of lime by means of dilute 
 acids, consist of a substance identical with that which 
 constitutes the envelope of the tubercle bacillus. Like 
 the latter, they are dissolved by concentrated acids, 
 they are not affected by alkalies, and do not give a red 
 colour with Millon's reagent. These facts must lead us 
 to the conclusion that the concentric layers are secre- 
 tions of the tubercle bacilli and are exactly analogous 
 to the stratified cuticle which, as we have seen, becomes 
 formed round the Gregarinas and Nematoda when 
 attacked by the phagocytes of Lumbricus (p. 69). 
 Thus in Meriones the bacillus defends itself against the 
 giant cell in which it is lying by the secretion of a series 
 of envelopes. On the other hand it is very probable 
 that the phosphate of lime is laid down by the giant 
 cell itself in its reaction against the bacillus. 
 
 The struggle between these two living bodies — the 
 tubercle bacillus and the giant cell of Meriones — ^is 
 thus carried on by means of secretions. The bacillus 
 defends itself by the secretion of cuticular membranes, 
 and probably also by the production of toxines, while 
 
1 66 INFLAMMATION. 
 
 the giant cell secretes a calcareous deposit by means of 
 which it walls in the bacillus, and usually succeeds in kill- 
 ing it. The giant cell probably also produces digestive 
 fluids which aid it in attacking and digesting the bacillus. 
 
 For the deposition of phosphate of lime to take place 
 in the interior of the giant cell an alkaline medium is 
 required. The alkaline reaction of the contents of the 
 giant cell and of the calcareous body formed from the 
 bacillus rnay be shown by various reagents such as the 
 sulpho-acid of alizarin, which is a very delicate test for 
 the presence of alkalies and gives these structures a 
 pronounced violet colour. Their intense coloration 
 with haematoxylin points to the same conclusion (Tab. 
 III., Fig. 8). 
 
 The exact conditions of this struggle between the 
 microbe and the phagocyte are highly interesting and 
 important and will form the subject of a separate 
 experimental investigation. Here I will only lay stress 
 on the conclusion that the giant cells, so characteristic 
 of tubercle, really represent a special form of phago- 
 cytes, which are particularly energetic in the conflict 
 with the microbes. This is moreover confirmed by the 
 discovery of Soudakewitch,' that the giant cells of lupus 
 are able to digest the elastic fibres of the skin. To 
 attack substances so resistant as tubercle bacilli and 
 elastic fibres requires very active power of digestion. 
 In this respect the giant cells of tubercle may be com- 
 pared to the giant cells known as osteoclasts, which 
 effect the absorption of bone. 
 
 Weigert," whose view on this point is accepted by 
 
 ' Virchoui's Archiv, 1889, vol. cxv., p. 264. 
 
 ' Deutsche med. Wochenschr., \%?il,2iX^i.Fortschritteder Medicin, 
 1888, p. 809, 
 
LECTURE X. \67 
 
 Koch ' and other pathologists, considers that the giant 
 cell of tubercle represents a condition of partial necrosis, 
 by which the division of the protoplasm is prevented. 
 Against this theory, besides the above-mentioned fact 
 of the remarkable power of resistance possessed by 
 these cells, we may cite their property of dividing into 
 smaller cells. It is not surprising that this division 
 should be unaccompanied by karyokinetic phenomena, 
 since here the nuclei are multiple to start with, and all 
 that is still necessary is the division of the protoplasmic 
 body of the cell. 
 
 If we accept the opinion put forward by various 
 authors and lately revived by Chun " that fragmentation 
 of the nucleus, by enlarging its surface, may serve to 
 heighten the activity of the intracellular processes of 
 nutrition, it is easy to understand the teleological reason 
 for the increase in the 'number of nuclei in the cells 
 which have to digest bodies so resistant as those in 
 question. We must not forget that in many inverte- 
 brate animals plasmodia are often formed during the 
 process of intracellular digestion, these fused agglomera- 
 tions of cells being very commonly met with in the 
 phagocytic reaction of Sponges, Medusae, Echinoderms, 
 Molluscs, Daphniae, Sec. 
 
 The cells of tubercle, which serve the organism in its 
 reaction against the microbes, are often destroyed by 
 the action of the bacilli, and then form the necrosed 
 caseous masses so characteristic of this disorder. This 
 degeneration is probably due in some degree to the 
 fact that the tubercles do not contain any blood vessels, 
 so that the activity and vitality of the phagocytes suffer 
 
 1 Detitsch. vied. Woch., 1891, No. 3, p. 102. 
 
 ' Physik.-okon. Gesellschaft zu Konigsberg. April 3rd, 1890. 
 
l68 INFLAMMATION. 
 
 from lack of nourishment. Koch has found that if the 
 inflammatory process in the tubercles be increased by 
 injection of tuberculin, a striking improvement may be 
 produced in the condition of guinea-pigs, and many 
 experimenters have observed a temporary amelioration 
 in man, aiter the same procedure. Probably in this 
 case the cells of the tubercles are better nourished in 
 consequence of the inflammatory hypersemia that is 
 set up, so that they become more active and better 
 fitted to resist the action of the bacilli. 
 
 The type of chronic inflammation that we have 
 chosen has shown us that the essence of the process 
 consists in a reaction of mesodermic phagocytes 
 against the offending agent. Although the macro- 
 phages are more conspicuous than the leucocytic ele- 
 ments, yet we must remember that among the macro- 
 phages of tubercles a large number of mononuclear 
 leucocytes also occur ; and Ehrlich ' has shown that 
 the latter elements are markedly increased in the 
 blood of phthisical patients. 
 
 Another chronic disorder, leprosy, which is analogous 
 in many respects to tuberculosis, must also be regarded 
 as a struggle between the mesodermic phagocytes and 
 the specific bacillus — a struggle which lasts for many 
 years. Although the origin of the macrophages is not 
 yet known with certainty, it is probable that they are 
 derived from the endothelial and connective tissue cells, 
 as well as from the mononuclear leucocytes. These 
 cells unite to form granulomata, and destroy the bacilli 
 that they englobe, thus acting as phagocytes. 
 
 ' Chartte Annalen, 1887, vol. xii., and " Farbenanaiytische 
 Untersuchungen,'' 1891, vol. i., p. 124. 
 
LECTURE X. 169 
 
 The older researches of Neisser ' and Kebner,^ as well 
 as the more recent ones of Soudakewitch ^ and Saw- 
 tchenko * leave no doubt as to the general characters of 
 the cellular pathology of leprosy. Having had several 
 opportunities of studying leprosy of the skin and of the 
 liver (among others in the preparations of Muskatbluth 
 made in my former laboratory at Odessa), I am able to 
 coniirm the fact that in these cases the leprous cells 
 possess all the characteristics of mesodermic phagocytes. 
 In the liver the cells which englobe the leprosy bacilli 
 are chiefly of the endothelial type, with an extra- 
 ordinary number of vacuoles in their protoplasm. 
 From the point of view of the phagocytic theory, this 
 vacuolar ' degeneration ' should rather be regarded 
 as due to an abundant secretion of digestive fluids, 
 analogous to that observed in the Protozoa while intra- 
 cellular digestion is going on. In the interior of 
 leprous cells, especially those of the skin, we often find 
 enormous numbers of dead bacilli, showing the thera- 
 peutic action of these cells. The fact that Boinet and 
 BoreP have not been able to confirm this function of 
 the cells shows either that they had to do with special 
 cases, or what is more probable, that their observations 
 were incomplete. 
 
 Among the tubercular diseases one that has been 
 recently studied by Ebstein and Nicolaier ° has a 
 special interest for us, viz. ; — the tubercular affection 
 
 ' VirclwdJ's Arckiv, 1881, vol. Ixxxiv., p. 520. 
 
 ^ Ibid., 1882, vol. Ixxxviii., p. 299. 
 
 ^ Ziegler's Beitrdge ziir pathol. Anat., vol. ii., Pt. i. 
 
 ■• Ibid., vol. ix. p. 241. 
 
 ' Comptes rendus dela Soc. de Biol., 1890, p. 38. 
 
 " Virchow's Arckiv, 1889, vol. cxviii., p. 432, pi. xiii., xiv. 
 
I/O INFLAMMATION. 
 
 produced by Nematoda in the kidneys of dogs and 
 the lungs of cats. In these cases the larvae become 
 surrounded by veritable tubercles, composed in the 
 dog of epithelioid cells alone, and in the cat of these 
 together with giant cells. In the dog each tubercle 
 contains a living larva enclosed by masses of tubercular 
 cells. Here then we have an undoubted instance of 
 an aggregation of leucocytes around a living animal, 
 recalling forcibly the collection of phagocytes in 
 Lumbricus round the living larvae of Rhabditis, as de- 
 scribed in the fifth lecture. It demonstrates once more 
 the fact that an accumulation of leucocytes may be 
 provoked by the presence of living organisms, and 
 does not require the presence of dead substances, or 
 substances derived from the dead bodies of the para- 
 sites. Unfortunately, however, these cases of zoopara- 
 sitic tuberculosis have not yet been sufficiently studied 
 to give us any accurate knowledge as to the origin or 
 development of the cellular elements composing the 
 tubercles. 
 
 As Brault ' has pointed out, there is a striking 
 analogy between the chronic inflammation brought 
 about by the tubercle bacillus and the processes that 
 give rise to the disease known by the name of hyper- 
 trophic cirrhosis. In both cases we have a reaction of 
 the connective tissues, or rather of the mesodermic cells 
 in general, a reaction which extends over a prolonged 
 period. Even if we regard it as an established fact that 
 the cirrhoses are produced directly by chemical poisons 
 such as lead, alcohol and others, the analogy of the 
 pathological processes in the two instances is by no 
 means destroyed. In the chronic infectious inflamma- 
 ' Archives generates de medecine, 1888, p. 47. 
 
LECTURE X. 171 
 
 tions the phagocytes collect around the parasites, 
 whereas in the cirrhoses they attack the tissue-cells that 
 have been enfeebled under the action of the poisons. 
 The final results then in the two cases may be quite 
 different, since in one the phagocytes destroy the 
 microbes, while in the other they annihilate the cells 
 belonging to the organism itself. In this respect the 
 chronic inflammations are closely allied to the phe- 
 nomena of atrophy, which however are scarcely within 
 the range of these lectures. 
 
LECTURE XI. 
 
 Serous inflammations — Two classes of these inflammations — 
 Bactericidal power of the humours and the serous exsudations 
 — Antitoxic property of serum and the serous inflammations. 
 
 In the chronic, as in the acute inflammations, whether 
 purulent, fibrinous or catarrhal, we find that a pro- 
 nounced phagocytic action forms the key note of the 
 whole process. In the serous inflammations, however, 
 the number of leucocytes present is too small to permit 
 of our ascribing any great significance to their presence. 
 Up to the present our knowledge of these serous in- 
 flammations is very incomplete. So far as we can 
 judge from experiments on animals, they may assume 
 various forms. 
 
 In the first place there are certain inflammations 
 which acquire the character of ' serous ' in consequence 
 of a negative chemiotactic influence on the leucocytes, 
 which do not emigrate or move towards the seat of 
 lesion, and therefore do not take any part in the forma- 
 tion of the exsudation. The walls of the vessels, 
 however, react as usual, thus permitting the passage 
 through their pores of the fluid forming the serous ex- 
 sudation, which is almost entirely devoid of leucocytes. 
 The exsudation often contains numerous pathogenic 
 microbes which multiply rapidly in it without let or 
 
LECTURE XI. 173 
 
 hindrance. Deprived in this way of any phagocytic 
 protection, the animal organism speedily becomes a 
 prey to the microbes. In this class of serous inflam- 
 mations we may include the most acute and fatal 
 diseases, such as the vibrionic septicaemia of birds and 
 guinea-pigs, anthrax in the smaller rodents as mice and 
 guinea-pigs, and some other infectious disorders. In 
 the form of septicaemia just mentioned, the vibrio 
 swarms freely in the serous exsudation with its almost 
 total absence of leucocytes. In the serous exsudation 
 of animals infected with anthrax, the bacilli are only 
 present in small numbers,' although the fluid t:ollects 
 at or in the immediate neighbourhood of the infected 
 spots. 
 
 In a second group of serous inflammations of microbic 
 origin, the exsudation is formed in regions, which may 
 be at some distance from the deposit of microbes, and 
 does not contain any bacteria at all. Roux and Yersin ' 
 have shown that the exsudation produced in the serous 
 pleurisy which often accompanies diphtheria in guinea- 
 pigs does not contain any microbes, which remain 
 localised at the seat of inoculation. In ten cases of 
 serous inflammations accompanying phlegmonous ery- 
 sipelas or other suppurative maladies, the liquid con- 
 tained in the inflammatory oedema was found by 
 Zimmermann - to be absolutely free from microbes. 
 The question at once suggests itself whether this trans- 
 sudation of fluid really represents a phenomenon of 
 reaction on the part of the organism, and if so, what 
 use it is to the infected organism. 
 
 In examining this point we naturally think first of a 
 
 ' Annales derinstitut Pasteur, 1888, p. 635. 
 
 2 Munchener medicinische Wochenschrift, 1889, No. 9, p. 141. 
 
174 INFLAMMATION. 
 
 possible microbicide action of the transsuded liquid 
 which would rid the organism of its aggressors. We 
 find, however, that all kinds of bacteria can live and 
 flourish in the fluid of the inflammatory oedema. 
 Although the serous exsudation of anthrax does not 
 usually contain many microbes, it may nevertheless 
 serve as a nutrient medium for them. Even in animals 
 refractory to anthrax, the spores of this bacillus, if in- 
 troduced into the serous fluid of the cedema produced 
 by a primary inoculation of anthrax,' will germinate 
 and give rise to a new generation of microbes. The 
 small number of microbes found in the exsudation can- 
 not therefore be taken as a proof of any bactericidal 
 action of this fluid. 
 
 In human beings tuberculosis is also frequently 
 accompanied by serous pleurisy in which no bacilli will 
 be found. But their absence is not due to their having 
 been killed by the liquid. If they had been present, 
 they would, as is invariably the case, have attracted the 
 leucocytes. As the microbe of diphtheria remains 
 localised at the seat of inoculation, we cannot doubt 
 that in this disease also there are no bacilli in the 
 serous exsudation from the very beginning of the 
 process. The above-mentioned ten cases of Zimmer- 
 mann were diseases brought about by the streptococcus 
 pyogenes and the staphylococcus, i.e. by two microbes 
 which, as Stern '' has proved, the serous exsudation of 
 human beings is absolutely unable to destroy. And yet 
 Zimmermann was unable to find any bacilli in the serous 
 exsudation which he collected from the above cases. 
 
 ' For a description of experiments of this nature on rats, see 
 Annales de VInstitut Pasteur, l8go, p. 201. 
 2 Zeitschrift f. klin. Mediciit, 1890 , vol. xviii., p. 62. 
 
LECTURE XI. 175 
 
 After this analysis of the facts so far as they are 
 known, we cannot admit that serous inflammation is a 
 means used by the organism for killing the pathogenic 
 organisms. The results obtained with regard to the 
 bactericidal properties of serous fluids generally are in 
 confirmation of this conclusion. In spite of all the 
 endeavours to prove that an active part is played by 
 these fluids in the destruction of microbes and the 
 production of immunity, we must recognise the fact 
 that they are of no importance in this capacity. Further 
 study of the bactericidal powers possessed by the serum 
 has shown that these have no connection either with 
 the phenomena which go on in the organism or with 
 the production of immunity. 
 
 Behring, who was one of the chief originators of the 
 bactericidal theory of the humours, carried out a large 
 number of researches on the subject, and concluded by 
 doubting whether the blood deprived of its cellular 
 elements could really produce immunity.' The results of 
 numerous experiments which he made with Nissen " led 
 him to consider that the bactericidal property of the 
 serum is only met with in a few isolated cases, where 
 natural or acquired immunity exists at the same time. 
 Among these cases, the most marked are the action of 
 the serum of rats upon the anthrax bacillus and that of 
 the serum of vaccinated guinea-pigs upon the Vibrio 
 Metchn ikowii. 
 
 Although the facts discovered by Behring and Nissen 
 are perfectly correct, and we must regard it as esta- 
 blished that the serum of rats has really a bactericidal 
 action on anthrax bacilli, and the serum of vaccinated 
 
 ' Deutsche medicin. Wochenschrift, 1891, No. 19, p. 655. 
 2 Zeitschrift fur Hygiene, 1890, vol viii. p. 424. 
 
176 INFLAMMATION. 
 
 guinea-pigs on the Vibrio Metchnikowii, yet even in 
 these cases these bactericidal powers which are so 
 evident in vitro do not come into play in the body of 
 the living animal. As Hankin,' Roux and I ^ have 
 shown, although rats are not refractory to anthrax, yet 
 their serum outside the body has a bactericidal action 
 on the bacilli of this disease. Again the Vibrio Metch- 
 nikowii is readily destroyed in vitro under the influence 
 of the serum of guinea-pigs which have been rendered 
 refractory by vaccination, although if inoculated under 
 the skin or in the eye of the same animals, these 
 microbes live for some time and finally adapt themselves 
 to their environment, so that they can now live in the 
 serum prepared from these animals.' 
 
 The impossibility of drawing any conclusion as to 
 the events that occur in the living organism, from the 
 results obtained from experiments with the serum out- 
 side the body, was shown in the earlier researches of 
 Lubarsch'' and myself." Recent experiments of 
 Buchner, Ibener, and Roeder,* have also proved that 
 the bactericidal properties of the serum, which are 
 very evident when the serum is allowed to act upon the 
 microbes in test tubes in the usual manner, become 
 much less marked if the same microbes be introduced 
 into the living body enclosed in capsules of filter paper, 
 so that they may be acted upon by the fluids of the 
 body without the possible interference of cells. 
 
 ' Centralblatt fur Bacteriologie, 1891, p. 378. 
 
 - Annales de VInstitut Pasteur, 1891, p. 479. 
 
 ' Ibid., p. 465. 
 
 ■* Centralblatt fur Bacteriologie, 1889. 
 
 = Virchow's Archiv, 1888, vol. cxiv. p. 472. 
 
 ^ Munchener medicin. Wachensckr., 1891, Nos. 32 and 33. 
 
LECTURE XI. 177 
 
 A review of these facts leads us to conclude that the 
 exsudation of a serous fluid in inflammation cannot be 
 regarded as a means which the organism may make use 
 of to destroy the pathogenic microbes, this service being 
 performed essentially by the phagocytes. Since, how- 
 ever, the poisonous chemical products of the bacteria 
 are the most important agencies in producing the general 
 intoxication that we know as the disease, it is possible 
 that the serous exudation may serve to attenuate or 
 modify the action of these products. 
 
 The remarkable discovery of Behring and Kitasato ' 
 that the serum of immunised animals possesses antitoxic 
 properties, seems to tell in favour of this hypothesis. 
 These observers who have been confirmed by Vaillard, 
 Tizzoni, and Cattani, have shown that the serum of 
 rabbits which have been vaccinated against tetanus, can 
 destroy considerable quantities of the toxine of this 
 disease. Behring ^ has obtained analogous, though not so 
 perfect, results for diphtheria ; and the Klemperers ' have 
 also proved that the blood and serum of rabbits which 
 have been inoculated against pneumonia, as well as these 
 fluids in men who have survived the crisis of a pneumonic 
 attack, are endowed with antitoxic properties. 
 
 Is it possible that the serous exsudation, although in- 
 capable of killing the microbes, may serve to destroy 
 the toxines ? This supposition would seem to receive 
 some support from the discovery by Behring of the 
 diphtheritic poison in the pleuritic effusion of rabbits 
 affected with this disorder. But in man, on the other 
 
 ' Deutsche medinische Wochensthrift, 1890, p. 1 1 13. 
 
 "^ Ibid., p. II45- 
 
 3 Berliner klinische W oclienschrift, 1891, Nos. 34 and 35. 
 
 N 
 
178 INFLAMMATION. 
 
 hand, although diphtheria often ends favourably, the 
 production of serous exsudations is never observed. 
 
 If we may regard croupous pneumonia as a disease 
 which terminates by producing an antitoxine (although 
 this cannot be taken as proved, in spite of the statements 
 of the Klemperers), it would furnish us with another in- 
 stance of the destruction of a toxine occurring indepen- 
 dently of any serous inflammation. 
 
 Tetanus is the type of a toxic disease. Yet although 
 vaccination against it gives rise to a rich production of 
 antitoxine, we never find any serous exsudation occur- 
 ring in the disease itself. Vaillard, who has a large 
 experience in the subject of tetanus, informs me that 
 in no case are serous inflammations ever present either 
 in animals affected with tetanus, or in those which are 
 undergoing the process of immunisation by the injec- 
 tion of the serum of vaccinated animals. 
 
 On the other hand tuberculosis, which is so often 
 accompanied by serous exsudations, must be looked 
 upon as one of the diseases in which toxines play the 
 smallest part. This is shown by the fact, among others, 
 that the guinea-pig, which is so susceptible to tuber- 
 culosis, is remarkably insensible to the action of the 
 tuberculin. As another example I may cite the Vibrio 
 Metchnikowii. If this microbe be introduced into 
 guinea-pigs, whether susceptible to or vaccinated 
 against its action, an abundant serous exsudation is 
 formed, and yet no antitoxine is produced in the animals 
 that recover from the disease, as is shown by the fact 
 that these animals, even when vaccinated against infec- 
 tion by this microbe, are still susceptible to the action 
 of the toxine formed by the Vibrio. 
 
 Thus the hypothesis that a serous inflammation is a 
 
LECTURE XI. 179 
 
 salutary reaction of the organism, is not confirmed by 
 the facts so far as we know them. We must keep in 
 mind, however^ that our knowledge on the subject of 
 the production of antitoxines is in its infancy ; and we 
 must wait for much wider and more varied experience 
 before we can come to a conclusion on the subject. 
 
 I may mention one more hypothesis. If we cannot 
 look upon the exsudation of serous fluid as connected 
 with the formation of antitoxines, it is at least possible 
 that it may serve to dilute the bacterial toxincs and so 
 render them less active. 
 
 A study of comparative pathology shows that in the 
 evolution of the inflammatory process the serous in- 
 flammations are much more recent than those which 
 are merely attended with a leucocytic reaction. In 
 none of the invertebrata, in which we have studied the 
 accumulation of phagocytes around foreign bodies, is 
 there any trace of a serous exsudation accompanying 
 this process. This is true not only for the organisms 
 which possess no blood (such as the Sponges, Coelente- 
 rata and Echinoderm larvae) but also for those en- 
 dowed with a circulatory system. Even in the amphi- 
 bians, in which the phagocytic reaction is so pronounced, 
 the inflammation is unaccompanied by any marked 
 transsudation of fluid. Only in a few exceptional cases 
 have I observed a slight accumulation of fluid around 
 inflammatory foci in the tails of tadpoles and of urodelan 
 larvae. 
 
 From whatever point we consider serous inflamma- 
 tion it appears as a phenomenon of minor importance 
 compared with the typical inflammatory process, i.e. an 
 accumulation of phagocytes in the inflamed area. 
 
 N 2 
 
LECTURE XII. 
 
 Review of other theories of infJammation in light of the acquired 
 facts — Nutritional theory of Virchow— Vascular theory of 
 Cohnheim — Experiments of the latter on the tongue of the frog. 
 Introduction of irritative agents into the blood. Argument 
 against Cohnheim's theory furnished by the reaction in verte- 
 brata — Struggle of the organism with external agencies — Use 
 of intracellular digestion — Phagocytes — Hsemitis (recurrent 
 fever, disease of Daphnia) — Tuberculosis. Essential nature of 
 inflammation — Sensibility of the phagocytes — Its progressive 
 development — Sensibility of the endothelial cells — Definition 
 of inflammation. Inflammation is not regeneration — Inflam- 
 mation is not identical with resorption — Objections raised to 
 the biological theory of inflammation — Vitalism — Teleology — 
 Absence of phagocytes in certain infective lesions — Imperfec- 
 tions in the inflammatory reaction — Surgical interference — 
 Comparative pathology. 
 
 Having now reviewed the principal phenomena of 
 the inflammatory reaction in the animal kingdom, we 
 may ask how far the facts thus brought out are in 
 harmony with the various theories mentioned in the 
 first lecture. 
 
 It is unnecessary to insist more than cursorily on the 
 impossibility of explaining these facts by the nutritional 
 theory of Virchow.' This doctrine assumes that a 
 greatly increased flow of nutritive substances takes 
 place towards the inflamed part as well as abnormal 
 
 ' " Cellularpathologie," 4th Edition, 1871, p. 475. 
 
LECTURE XII. l8l 
 
 proliferation of the local cells, that in fact inflammation 
 begins from the moment that nutritional derangement 
 occurs. This derangement consists in " the attraction, 
 that is to say, in the direct absorption and the modifica- 
 tion, according to circumstances, of large quantities of 
 nutrient substances." In this way the cells of the 
 inflamed organ receive an excessive amount of nourish- 
 ment at the expense of the fluid parts of the blood. 
 Considering the matter from this point of view, Virchow 
 regards the most characteristic phenomena of inflamma- 
 tion, not as a salutary means of reaction, but as a 
 process distinguished chiefly by its acute nature and 
 especially by the danger that the organism incurs from 
 it.' 
 
 Samuel and Cohnheim held that inflammation con- 
 sisted essentially in a molecular lesion of the vascular 
 wall, that the latter, modified by some injurious agency, 
 lost its power of retaining the blood corpuscles, which 
 therefore in consequence of the forces acting upon them 
 were pressed out of the vessels, and then wandered to- 
 wards the part of least resistance. According to them, 
 inflammation was in nowise a mode by which the 
 organism reacted to extrinsic influences but was merely 
 a primary lesion of the blood-vessels. One experiment 
 of Cohnheim's well illustrates his way of looking at the 
 subject. If by means of a ligature the circulation of 
 the blood in a frog's tongue be stopped for forty-eight 
 hours, and the ligature be then removed, circulation 
 will again proceed, but accompanied with a condition 
 of inflammation, i.e. with the peripheral distribution of 
 the leucocytes, followed by their diapedesis. Cohnheim 
 explains this fact as a direct result of the lesion to the 
 ' Loc. cit., p. 399. 
 
1 82 INFLAMMATION. 
 
 vessel-wall, consequent upon its previous prolonged 
 state of anaemia. " I consider that the cause of 
 inflammation," he adds, "should undoubtedly be sought 
 in the vessels themselves ; the events outside the vessels 
 appear to be of too subordinate a character to afford an 
 explanation of this phenomenon." ' And yet very 
 important events must take place outside the vessels. 
 The peripheral tissues, deprived of their nutrition and 
 of their protection by the blood, must become the seat 
 of aggression on the part of the numberless microbes 
 which exist in the buccal cavity. The tissues them- 
 selves, or some of them, must at the same time undergo 
 degeneration, which may produce sufficient peripheral 
 excitation to bring about an inflammatory reaction. 
 The instance given by Cohnheim enables us to com- 
 prehend his theory without however proving it. 
 
 In order to produce what we may call a central 
 vascular lesion independently of the lesion in the other 
 organs situated at the periphery, we may adopt a simple 
 expedient, which consists in introducing the irritating 
 object inside the vessels themselves. In the same 
 work of Cohnheim on embolism, he adduces experi- 
 ments which were made with the purpose of producing 
 embolic abscesses. To this end he injected into the 
 arteries of frogs a number of substances (such as 
 globules of mercury, powdered cantharides, particles of 
 putrid meat) which might have been expected to cause 
 considerable inflammation, but had no effect at all. 
 Now these substances if introduced under the skin are 
 followed by a marked reaction. If therefore inflam- 
 mation were only due to a change in the vessel-walls, 
 how is the absence of this change to be explained in 
 ' "Die embolischen Processe," 1873, p. 51. 
 
LECTURE XII. 183 
 
 cases where the irritating agent comes in direct contact 
 with the walls themselves ? 
 
 Many other facts may be quoted which give the same 
 results as the above experiments of Cohnheim. Patho- 
 genic organisms which readily provoke inflammation 
 are frequently found in the blood without causing any 
 exsudation. In recurrent fever the blood is crowded 
 with spirilla which act directly upon the vessel-wall by 
 their spiral movements as well as in all probability 
 by their toxic products. If Cohnheim's theory were 
 correct, this would cause such an alteration in the walls 
 as to bring about the passage of the corpuscles outside 
 the vessels. Nevertheless no inflammation occurs in the 
 organs during the course of this fever, although the 
 whole body of the patient is ' inflamed ' in the highest 
 degree. Many other microbes, such as the anthrax 
 bacillus, cause intense inflammation when 'inoculated 
 subcutaneously, whereas inflammation in the sense used 
 by Cohnheim does not occur at all, when the same 
 organisms are in the blood. This is again the case 
 when the tubercle bacillus is injected into the blood, 
 while if introduced outside the vessels exsudative 
 inflammation invariably results. 
 
 Besides these facts which the theory of the primary 
 lesion of the vascular wall does not take into account 
 the whole comparative pathology of inflammation is 
 an argument against Cohnheim's contention. The 
 reactive phenomena of the invertebrata show that 
 infiltration precedes the vascular events in the evolu- 
 tional history of the process, and that the leucocytes, led 
 by their sensibility and by means of their amoeboid move- 
 ments, themselves proceed towards the injured spot 
 instead of passively filtering through a vessel-wall. 
 
1 84 INFLAMMATION. 
 
 But even if the theories of Virchow and Cohnheim 
 cannot be accepted in the light of our present know- 
 ledge, must we renounce all hope of comprehending 
 imflammatory phenomena, and remain satisfied with a 
 mere description of them as most modern pathologists 
 do? 
 
 The study of inflammation from the point of view of 
 comparative pathology proves first of all that this 
 phenomenon is essentially reactive in its nature. The 
 organism, threatened by some injurious agency, pro- 
 tects itself by the means at its disposal. Since, as we 
 have seen, even the lowest organisms, instead of 
 passively submitting to the attacks of morbid agents, 
 struggle against them, why should not the more highly 
 developed organisms, such as man and mammals, act in 
 the same manner ? We must conclude then that the 
 invaded organism fights against the injurious cause, but 
 in what way ? As the evolution of inflammation shows, 
 it is this phenomenon itself which is both the most 
 general and the most active means of defence among 
 the members of the animal kingdom. 
 
 The essential factor in inflammatory reaction is an 
 endeavour on the part of the protoplasm to digest the 
 harmful object. This digestive action, in which the 
 whole or almost the whole organism of the Protozoa 
 takes part, is undertaken by the entire plasmodic mass 
 of the Myxomycetes, while, from the Sponges upwards, 
 it is confined to the mesoderm. In those cases where 
 the victory remains with the invaded organism, the 
 phagocytic cells of this layer assemble, englobe and 
 destroy the injurious agent. This phagocytic reaction, 
 in the lower scale of animal life, is slow, owing to the 
 progression of these cells towards the injurious body 
 
LECTURE XII. 185 
 
 being dependent solely on their amoeboid movements ; 
 but as soon as a circulatory or vascular system makes 
 its appearance in the course of evolution, it becomes 
 much more rapid. By means of the blood-current the 
 organism can at any given moment send along to the 
 threatened spot a considerable number of leucocytes to 
 avert the evil. When the circulation is partially 
 carried on by a lacunar system there is nothing to 
 intercept the movement of the leucocytes towards the 
 seat of injury. But when these cells are enclosed 
 within the vessels, they are obliged to adapt themselves 
 specially to fulfil their object, which they do by passing 
 through the vascular wall. 
 
 If we accept this conclusion that inflammation in the 
 higher animals is a salutary reaction of the organism 
 and that diapedesis and its accompaniments form part 
 of this reaction, several details of inflammatory pheno- 
 mena will appear clear to us. For instance the lobed 
 and polymorphous shape of the nucleus of the pus- 
 corpuscles has long been remarked. This particular 
 shape is peculiar to the polynuclear leucocytes, which 
 represent the vast majority (75 per cent.) of the 
 total number of white cells. As it was noticed 
 that a quantity of pus- corpuscles died in the exsu- 
 dation, this fact became associated with the curious 
 form of the nucleus ; it was said, and is still maintained, 
 that the polynuclear leucocytes are cells predestined to 
 perish and incapable of any considerable activity. On 
 the contrary these leucocytes are. precisely the most 
 active cells in the organism. The shape of their nucleus 
 may be more adequately explained as a special adapta- 
 tion for passing through the vessel-wall. If the process 
 of diapedesis be watched, the difficulty experienced by 
 
1 86 INFLAMMATION. 
 
 the nucleus in getting through will at once be noticed. 
 Directly this has occurred, the rest of the protoplasm 
 follows rapidly. It is obvious that a nucleus divided 
 into several lobes can pass through the wall more easily 
 than one not so separated. Hence in pus the poly- 
 nuclear leucocytes are more numerous than the mono- 
 nuclear leucocytes, and hence the lobed shape of the 
 nucleus is found only in the leucocytes adapted for 
 diapedesis and does not occur among the inver- 
 tebrata (except in a few Cephalopoda). 
 
 The facts which demonstrate the untenable position 
 taken up by Cohnheim may be readily explained by 
 means of the theory here set forth. If the irritating 
 agent be outside the vessels, it provokes a typical in- 
 flammation, accompanied by diapedesis ; if the same 
 agent be within the vessels no diapedesis takes place 
 but the leucocytes fight against the microbes in the 
 blood itself. For instance in recurrent fever, the 
 spirilla undoubtedly act upon the vascular wall with- 
 out bringing about diapedesis. But the leucocytes in- 
 crease in number ; the leucocytosis is followed by a 
 struggle which is ended by the leucocytes devour- 
 ing the spirilla. We have here a case of inflammation 
 unaccompanied by diapedesis ; the conflict between the 
 phagocytes and the spirilla takes place in the blood 
 itself. Although no diapedesis occurs, there is in re- 
 current fever great elevation of temperature as well as 
 other symptoms which prove it to be an inflammatory 
 disease. It is apparently a case of inflammation in the 
 blood itself, a sort of ' hemitis ' as Piorry considered it 
 might be many years ago. We find the same condi- 
 tions in animals whose vascular system and general 
 body-cavitj' are in communication. Thus as we have 
 seen in the disease of Daphnia, caused by the Mono- 
 
LECTURE XII. 187 
 
 spora, the leucocytes often collect in large numbers 
 around the spores of this parasite — their assemblage 
 taking place in the body-cavity. 
 
 As another instance we may cite tuberculosis. If 
 inoculated subcutaneously, the tubercle bacilli produce 
 inflammation accompanied by considerable diapedesis. 
 But if the same bacilli be injected direct into the blood, 
 no diapedesis occurs, but the phagocytes will gather 
 round the bacilli within the vessels and form intravas- 
 cular tubercles. It cannot be said that in the first case 
 (extravascular inoculation) there is inflammation and 
 that in the second (intravascular inoculation) there is 
 none, especially as the same tubercles are formed in 
 both instances. This is another example of an inflam- 
 mation of the blood itself. 
 
 All these cases of intravascular inflammation without 
 diapedesis, as well as the inflammatory phenomena in 
 the young larvae of Axololotls and Tritons (where it is 
 the migratory cells that collect at the seat of injury), 
 in fact the whole series of reactive phenomena in so 
 many of the invertebrates, prove clearly that the essential 
 and primary element in typical inflammation consists in 
 a reaction of the phagocytes against a harmful agent. If 
 the latter be in the general bddy-cavity, which is filled 
 with blood, the phagocytes will collect here ; if in the 
 interior of the vessels, as in recurrent fever or in 
 intravascular tuberculosis, the phagocytes will assemble 
 in the blood itself; if on the contrary the injurious 
 agent is outside the body-cavity or outside the vessels, 
 the phagocytes will emigrate towards the threatened 
 spot — an emigration without diapedesis in the inverte- 
 brata and young larvae of Urodela, or with diapedesis 
 in the vertebrata. 
 
 Before phagocytic reaction can take place, these cells 
 
1 88 INFLAMMATION. 
 
 must be excited positively. Negative sensibility may 
 also serve as a means of defence in a mobile organism, 
 such as the plasmodium of the Myxomycetes, which 
 retires from the offending cause. In the cases where 
 the latter has penetrated into the organism, negative 
 sensibility on the part of the phagocytes will leave the 
 field of battle to the parasite, so that, as frequently 
 happens, the death of the organism results. Hence as 
 we rise in the scale, we are met by a progressive evolu- 
 tion of positive sensibility in the leucocytes. In 
 Daphnia the observer is struck by the number of 
 diseases in which phagocytosis is entirely or almost 
 entirely absent. By the time we reach the amphibia, 
 positive chemiotaxis is already very marked and, as 
 Gabritchewsky has shown, it is still more highly 
 developed in rabbits. And yet among the rodents, as 
 in the small laboratory animals generally, there occurs 
 a certain number of rapidly fatal diseases (such as 
 chicken cholera, hog cholera, vibrionian septicaemia 
 of birds), in which phagocytosis is often completely 
 absent. In man and the higher mammals, similar 
 diseases are much less frequent. 
 
 But, in addition to the mobile phagocytes adapted by 
 their sensibility to move towards the offending object, 
 there are also fixed phagocytes. A good example of 
 the latter, which are especially developed in the higher 
 vertebrates, is furnished by the endothelial cells of the 
 vessels. Since these cells are contractile and phago- 
 cytic, it is natural to conclude that they must also be 
 possessed of sensibihty. Thus, if we assume a chemio- 
 tactic sensibility of the endothelial cells, we may easily 
 explain the remarkable power of reciprocal attraction 
 possessed by the protoplasmic processes of developing 
 
LECTURE XII. 189 
 
 capillaries, which enables them to meet and form a new 
 vascular loop. We may apply this explanation to 
 account for the fact that in many neoplasms, as in 
 pannus, the vessels penetrate and branch freely in the 
 affected tissue, whereas in the granulomata, such as 
 tubercle, leprosy and actinomycosis, blood-vessels are 
 absent. In the former case there is a positive chemio- 
 tactic influence attracting the vascular loops, in the 
 latter a negative chemictaxis or other form of negative 
 sensibility of the endothelial cells. The co-operation 
 of these cells in the inflammatory process would be also 
 directed by their sensibility, at any rate so far as their 
 active contraction is concerned. 
 
 There is one more form of sensibility that we must 
 mention, namely that of the nervous system, which aids 
 the phagocytic and vascular mechanisms in their re- 
 action against deleterious agents. 
 
 To sum up : Inflammation generally must be regarded 
 as a phagocytic reaction on the part of the organism against 
 irritants. This reaction is carried out by the mobile 
 phagocytes sometimes alone, sometimes with the aid of 
 tJte vascular phagocytes or of the nervous system. 
 
 The theory here indicated might be termed the 
 biological or comparative theory of inflammation, since 
 it is founded on a comparative study of the patho- 
 logical phenomena presented by living cells. 
 
 It is above all necessary to emphasise the fact that 
 the essential phenomena of inflammation represent an 
 actual struggle between the phagocytes and the irritant 
 agent. As we have seen that leucocytes are capable of 
 transformation into connective tissue cells, it might be 
 thought that this was the only object for which the 
 accumulation took place. This idea however is nega- 
 
igo INFLAMMATION. 
 
 tived by the fact that, in the higher vertebrates, the 
 polynuclear leucocytes do not contribute to the forma- 
 tion of granulation-tissue, although they constitute the 
 large majority of the cells taking part in the inflamma- 
 tion. The mononuclear leucocytes are the only cells, 
 besides the endothelial and connective tissue cells, that 
 are actively engaged in the formation of the new 
 connective tissue. These leucocytes are formed 
 directly from the numerous lymphocytes which flock 
 towards the locality where regeneration is going on. 
 
 It has been often thought that the leucocytes which 
 are gathered together in an inflamed area may only 
 serve to effect the absorption of dead cells and microbes, 
 and they have been looked upon as simple ' scavengers ' 
 of the organism. We have already seen that this 
 hypothesis is not justified by facts, and that from the 
 very onset of infection the leucocytes wander towards 
 and englobe the parasites in a living condition. In 
 case these direct proofs may not be thought sufficient, 
 I may mention other considerations which tell against 
 this theory. 
 
 If the main object of the leucocytic reaction in in- 
 flammation were the absorption of solid matters, we 
 should expect to find very pronounced inflammatory 
 phenomena in all cases where a very considerable and 
 rapid absorption is going on. This is however not the 
 case. The metamorphosis of Batrachians is accom- 
 panied by an absorption of the larval organs — tail and 
 gills. This is effected very rapidly (in a few days) by 
 the aid of phagocytes which devour all the tissues no 
 longer required by the animal. And yet in this case 
 there are no signs of inflammation, the phagocytes of 
 the tissues in question being quite sufficient by them- 
 
LECTURE XII. 191 
 
 selves to carry out the work of absorption which is evi- 
 dently a much easier task for them and requires much less 
 activity on their part than does the struggle with parasites. 
 In mammals the real ' scavengers,' that is to say the 
 phagocytes which carry out the work of absorption, are 
 the macrophages, especially the mononuclear leucocytes. 
 The latter cells play an important part chiefly in the 
 chronic inflammations, such as tuberculosis, while in the 
 acute inflammations it is mainly the microphages, or 
 neutrophile polynuclear leucocytes, that are involved- 
 In erysipelas for example the streptococci are englobed 
 only by the polynuclear leucocytes, and are never 
 taken up by the macrophages, which, however, carry 
 out the entire work of absorption, and even englobe the 
 microphages, many of which perish in the struggle with 
 the microbes and have to be themselves absorbed. If 
 absorption were the only function of the inflammatory 
 em.igration, it would be teleologically absurd for this 
 process to end in the formation of pus, i.e. a mass of 
 leucocytes, many of which die and must themselves be 
 absorbed. It is much more natural to assume that the 
 emigration is a reaction on the part of the organism 
 and that in the conflict many of the principal com- 
 batants, the microphages, perish on the field of battle. 
 Absorption afterwards ensues and is carried out by 
 another variety of phagocytes. 
 
 When I first put forward the biological theory of 
 inflammation eight years ago,' I expressed the idea 
 that this reaction is effected by the intermediation of a 
 physiological continuity between " the cells of the 
 connective tissue, those of the endothelial wall and the 
 leucocytes, which form a complete chain and play the 
 ' Biologisches Centralblati, 1883, p. 564. 
 
192 INFLAMMATION. 
 
 principal part in the inflammation of vertebrates." 
 The connective tissue cells which are first attacked, 
 would, I thought, transmit the action to the vascular 
 wall, the cells of which would contract to facilitate the 
 passage of the white corpuscles. At that time only- 
 tactile sensibility of these various cells had been recog- 
 nised, although certain facts, such as the existence of 
 chemiotaxis, was already suspected from observations 
 of the power possessed by the sexual cells of the 
 Hydromedusse to travel considerable distances.' Since 
 then this chemiotaxis has been definitely proved to exist. 
 This biological theory has often been considered too 
 vitalistic in its tendency. I need only quote Frankel's 
 outspoken criticism of my theory from this point of 
 view.* " The phagocyte theory presupposes extra- 
 ordinary powers on the part of the protoplasm of 
 leucocytes, to which are attributed sensations, thoughts 
 and actions, in fact a kind of psychical activity." The 
 sensibility of the phagocytes is not an hypothesis which 
 can be admitted or rejected at will, but an established 
 fact, which cannot be ignored, as it is by Frankel. 
 Whether they possess powers of thought and volition, 
 as this author accuses me of assuming, is quite beside 
 the question, though we are justified in considering that 
 they possess a germ of these qualities and that their 
 sensibility, like that of various vegetable and animal 
 unicellular organisms represents the lowest stage in the 
 long series of phenomena which culminate in the 
 psychical activities of man. 
 
 ' Weismann, " Die Enstehung der Sexualzellen bei Hydromedu- 
 sen." Jena, 1883. 
 
 ' '' Grundriss der Bakterienkunde." 3rd Ed., Berlin, 1890, 
 p. 203. 
 
LECTURE XII. 193 
 
 Asrthe authorities on tlj^se subjects, Herbeft Spencer, 
 Romanes and others have shown, psychical phenomena 
 have nothing specific in them, but have developed as an 
 association of the simple actions which we observe in 
 the lower organisms and the cells of different animals. 
 
 The accusation of vitalism and animism, which is 
 unjustly cast at the phagocyte theory, might really be 
 more appropriately applied to my opponents, who 
 maintain that the psychical acts of the higher animals are 
 fundamentally different in their nature from the more 
 simple phenomena peculiar to the lower organisms. 
 
 It is equally erroneous to attribute a teleological 
 character to the theory that inflammation is a reaction 
 of the organism against injurious agencies. This theory 
 is based on the, law of evolution according to which 
 the properties that are Useful to the organism survive 
 while those which are harmful are eliminated by natural 
 selection. Those of the lower animals which were 
 possessed of mobile cells to englobe and destroy the 
 enenjy, survived, whereas others whose phagocytes did 
 not exercise their function were necessarily destined to 
 perish. In consequence of this natural selection the 
 useful characteristics, including those required for 
 inflammatory reaction, have been established and trans- 
 mitted, and we need not invoke the assistance of a 
 designed adaptation to a predestined end, as we should 
 from the teleological point of view. 
 
 But, it has been urged on several occasions,' if the 
 
 phagocytic reaction has been developed in order to 
 
 protect the organism from danger, how is it that the 
 
 phagocytes refuse to'act just when the orga;nis.m is most 
 
 ' Forinstance by Baumg;arten, Berlin. klm.Woche^sckrift, i8?4, 
 and Burdon Sanderson, Brit. Med. Journ., 1891, p. 1085. , ... 
 
 O 
 
1 94 INFLAMMATION. 
 
 threatened? This objection again arises from an 
 insufficient knowledge of the principles of the theory. 
 It is just because the defence by the phagocytes is 
 developed according to the law of natural selection and 
 is not a designed adaptation to a particular end, that 
 cases naturally occur where the phagocytes do not 
 fulfil their functions, a neglect followed by the most 
 serious danger to or death of the organism. In nature 
 the organism is possessed of many characteristics, which 
 may be either useful or injurious to their owner. The 
 former causes the survival, the latter the death of the 
 possessor. Let us take two organisms : one in which 
 the phagocytes are readily repulsed by the microbe, the 
 other whose phagocytes show a positive sensibility 
 causing considerable phagocytosis. The former will 
 soon fall a prey to the parasite and be eliminated by 
 natural selection, whereas the latter will resist the 
 infection, survive and put forth progeny possessed of 
 the same phagocytic properties. Under these con- 
 ditions the activity of the phagocytes will increase 
 with every successive generation. 
 
 But the curative force of nature, the most important 
 element of which is the inflammatory reaction, is not 
 yet perfectly adapted to its object. The frequency of 
 disease and the instances of premature death are a 
 sufficient proof of this. The phagocytic mechanism 
 has not yet reached its highest stage of development 
 and is still undergoing improvement. In too many 
 cases the phagocytes flee before the enemy or destroy 
 the cells of the body to which they belong (as in the 
 scleroses). It is this imperfection in the curative forces 
 of nature which has necessitated the active intervention 
 of man. 
 
LECTURE XII. 1 95 
 
 The defence of the organism against deleterious 
 agencies, which is at first confined to the phagocytic 
 mechanisms and the somatic system of nerves, by and 
 by spreads to and is undertaken by the psychical 
 nervous apparatus. With the nervous cells which 
 direct the contraction and dilatation of the vessels 
 become associated other cells which control thought 
 and voluntary actions. One function of these psychical 
 cells has been to develop a complete science for the 
 defence of the organism against hostile influences. By 
 its means, methods for assisting the curative inflamma- 
 tion have been devised, as for instance the formation of 
 artificial lesions in order to facilitate the inflammatory 
 reaction. The application of agents which set up in- 
 flammation, such as jequirity, the virus of gonorrhea, 
 tuberculin and cantharidin, is the conscious continua- 
 tion of the defensive measures which have been uncon- 
 sciously evolved by the long series of animals in their 
 struggle for existence. 
 
 But neither the conscious nor the unconscious 
 mechanism is perfect. Medical science should therefore 
 call to its aid all the less complex branches of science, 
 including that of biology, which seeks to investigate 
 organisms and their evolution. Not only inflamma- 
 tion but many other medical problems might be advan- 
 tageously attacked from the standpoint of comparative 
 pathology. In order to trace the first appearance of 
 febrile reaction, the study of fever should begin with in- 
 vestigations on heat production in the lower animals, con- 
 tinuing on to the vertebrates, such as reptiles, &c. As 
 several of these phenomena have originated at an early 
 geological epoch, the conditions in which the first warm- 
 blooded animals lived should also be taken into account. 
 
 O 2 
 
196 INFLAMMATION. 
 
 Comparative pathology could undoubtedly throw 
 light on the important pathological question of tumours- 
 As many of the lower organisms — both animal and 
 vegetable — are subject to the formation of tumours, the 
 paj-t played by parasites in their . etiology could be 
 more ,easily established and the theory of embryonic 
 aberrations refuted. 
 
 The subject of atrophy, intimately connected with 
 that of chronic inflammation, is likewise a suitable one 
 for the application of comparative pathology to its 
 elucidation. Here again the phagocytic phenomena 
 take a prominent part 
 
 If however medical science may learn much from 
 biology, of which it forms but a part, it may at the 
 same time give something in return. General biology 
 may extend its knowledge by including the study of 
 the morbid phenomena of which pathology takes cog- 
 nisance. In biology difficulties frequently present them- 
 selves in the study of the evolutionary processes, be- 
 cause the perfect condition of adaptation in which these 
 phenomena occur in nature, is not a favourable one for 
 analysis. In order to throw light upon the universal 
 law of natural selection, it ,is necessary to study the 
 less stable phenomena, the less perfect mechanisms, in 
 a word, the phenomena in which the working out of 
 this law can be watched day by day. Now it is just 
 the morbid phenomena with the reactions they provoke, 
 the struggle between the organism and its aggressors, 
 which offer the best opportunity for a consecutive study 
 of the course of natural selection. In this struggle 
 every day .some of the combatants, by virtue of natural 
 selection, survive, while others, eliminated by the same 
 means, perish. Now it is the conquering organisms 
 
LECTURE XII. 197 
 
 which survive and the vanquished parasites which be- 
 come eliminated, now the defeated organisms which are 
 eliminated and the triumphant parasites which sur- 
 vive. 
 
 Thus I conclude as I began. General pathology 
 should go hand in hand with zoology or rather with 
 biology, and form one branch of it, that of comparative 
 pathology. This science is only in its infancy, and yet 
 it is already in a position to render good service to 
 medicine. By facilitating the analysis of the reactive 
 phenomena, it indicates the elements which should be 
 especially protected and reinforced in the conflict of 
 the organism against its enemies, and thus contributes 
 to the solution of one of the great problems of 
 humanity. 
 
DESCRIPTION OF THE PLATES. 
 
 PLATE I. 
 
 Fig. I. — Inflamed spot on the caudal fin of a larva oi Bombinator 
 igiieus, 72 hours after cauterisation with nitrate of silver : — a. Pha- 
 gocytes containing pigment and red corpuscles ; b, c. Branched 
 cells enclosing fragments of red corpuscles ; d. A phagocyte inside 
 a lymphatic vessel. 
 
 Fig. 2. — A cyst of the Gregarina of Lumbricus surrounded by a 
 thick and deformed cuticle, and by a follicle formed of connective 
 tissue. 
 
 Fig. 3. — A cyst of the same parasite, with only the remains of 
 its cuticle left. 
 
 Fig. 4. — A Rhabditis in the midst of a mass of phagocytes. 
 c. Thick and deformed cuticle of the parasite. 
 
 Fig. 5. — Another Rhabditis, enclosed in the mass of phagocytes 
 of Lumbricus. c. Cuticle of the parasite, consisting of several 
 concentric layers. 
 
 PLATE n. 
 
 Fig. i. — A cyst of the Gregarina of Lumbricus with its contents 
 divided into cells. The cyst is surrounded by a follicle composed 
 of connective tissue. 
 
 Fig. 2. — A mass of phagocytes of Lumbricus surrounding a 
 cyst of Gregarina which they have destroyed. 
 
200 DESCRIPTION OF THE PLATES. 
 
 Fig. 4. — Example of positive chemiotaxis. A plasmodium of 
 Didymium farinaceum which has dipped its processes into an 
 infusion of leaves. 
 
 Fig. 3. — Example of negative chemiotaxis. The plasmodium of 
 Fig. 4 receding from a o"i per cent, solution of quinine. 
 
 Fig. 6. — Another plasmodium of Didymium repulsed by the 
 hydrochlorate of quinine (o i per cent.) 
 
 Fig. 5.— The same plasmodium 5 hours after the solution of 
 quinine has been replaced by an infusion of leaves. The negative 
 chemiotaxis of Fig. 6 is converted into positive chemiotaxis. 
 
 Fig. 7. — A leucocyte of the salanTander. a. Attraction sphere 
 (after Flemming). 
 
 Fig. 8. — A clasmatocyte from the mesentery of Triton taniatus. 
 
 PLATE III. 
 
 ■ Fig. i. — A cyst of Gregarina surrounded by the phagocytes of 
 Lumbricus. (Magnification : objective D and ocular 4 of Zeiss.) 
 
 Fig. 2. — Eosinophile leucocyte from man (after Gabritchewsky). 
 
 Fig. 3. — One of Ehrlich's cells from a white rat. 
 
 Fig. 4. — A giant cell from the spleen of Meriones, a. En- 
 velope of the bacillus.; b. Bacillus of Koch. (The spleen treated 
 witli Flemming's fluid and stained by Gram's method and eosine. 
 Magnification: objective 1/18, and ocular 3 of Zeiss.) 
 
 Fig. 5. — ^A giant cell froin the spleen of Meriones, containing 
 a calcareous body with a double bacillus. (Stained with hasma- 
 toxylin and Ziehl's fuchsi'n. Same magnification.) 
 
 Fig. 6. — Another giant cell within which the bacillus may be 
 seen surrounded with concentric layers. (Treated and stained as 
 in Fig. 5. Objective 1/18, ocular 2.) 
 
 Fig. 7. — A giant cell with a calcareous body containing only a 
 trace of the bacillus b. , (Fuchsin, hsematoxylin, 1/18x2.) 
 
 Fig. 8. — Another giant cell, in which the bacillus b has becdme 
 converted into a pale rose-coloured body. 
 
 . Fig. g.^A giant cell enclosing a fully ' developed calcareous 
 body. 
 
Missing Page 
 
APPENDIX. 
 
 A Consideration of some Criticisms ' on the 
 BioxoGicAL Theory of Inflammation. 
 
 In the foregoing pages the ontogenetic study of the pheno- 
 mena of inflammation in the animal kingdom has enabled me to- 
 establish a biological theory of this process. I have shown 
 that it really 'represents; an adaptation of the organism to its 
 environment, which has been developed in the course of its 
 never-ceasing struggles against deleterious agencies, " In- 
 flammation generally must' be regarded as a phagocytic reaction 
 on the part of the organism against irritants. This reaction is 
 carried out by the mobile phagocytes, sometimes alone, some- 
 times with the aid of the vascular phagocytes or of the nervous 
 system." (P. 189.) 
 
 In analysing the characteristics of inflammation, pathologists 
 have long recognised that the exsudation forms the principal 
 feature, and that the three other classical symptoms, heat, 
 redness and pain, are only of secondary importance. Now the 
 most important constituents of the exsudation are the leucocytes, 
 which are contained in large quantities, not only in purulent 
 but also in serous and fibrinous exsudations. It is rare to find 
 them completely absent in any inflammatory exsudation. The 
 fact, that in the lower animals the inflammatory reaction consists 
 merely in an aggregation of phagocytic cells similar to leuco- 
 cytes without any accumulation of the body fluids, shows that 
 the existence in higher animals of exsudations free from cells 
 is of secondary importance for our conception of the essential 
 
 ' Podwyssozki. "Memoir in Commemoration of Virchow's 
 Jubilee," Kieff, 1892 (in Russian). Ziegler, " Historisches und 
 Kritisches iiber die Lehre von der Entzundiing," Beitri^e zur 
 ■batholosiischen Anatomie, 1892, vol. xii., p. 152, Weigert, Deutsche 
 niedicinische Wochenschtift, 1893, pp. 17, 37- 
 
202 APPENDIX. 
 
 nature of the process. It is evident however that these 
 phenomena must be taken into account in formulating any 
 scientific theory of inflammation. 
 
 The criticism usually urged against the biological theory of 
 inflammation has been that it totally disregards inflammations 
 in which the exsudation is free from leucocytes. According to 
 Podwyssozki, certain serous inflammations may occur un- 
 accompanied by any phagocytic reaction, and yet we cannot 
 exclude these from the category of true inflammations. He 
 proposes therefore to replace the above definition by the 
 following. " Inflammation is a local reaction, often beneficial, 
 of the living tissues against the irritant substance. This re- 
 action is chiefly produced by a phagocytic activity of the meso- 
 dermic cells. In this reaction however may participate not 
 only changes in the vascular system, but also the chemical 
 action of the blood-plasma and tissue-fluids in liquefying and 
 dissolving the irritant agent." This theory has the advantage 
 of including the purely serous inflammations. 
 
 In the same way Weigert accuses me of ignoring the 
 existence of exsudations free from cells, and suggests that I do 
 not know what to make of the serous inflammations. Ziegler 
 raises the same objection, although he does not formulate it so 
 precisely. These criticisms however rest upon a misconcep- 
 tion which we may remove at once. Although I have devoted 
 a volume to the development of the biological theory of in- 
 flammation, I have had to leave many points insufiSciently 
 discussed, and it is just to these points that the principal 
 objections have been urged. 
 
 The passage of red corpuscles and plasma into the inflamma- 
 tory exsudations must be looked upon as the result of the 
 activity of the endothelial cells of the vessel walls. (P. 149.) 
 The frequent presence of red corpuscles in the exsudations 
 which are most serous and free from leucocytes shows clearly 
 that in these cases there is a direct passage of the constituents 
 of the blood into the exsudation. If the blood discs are able 
 to pass through the vascular wall, it is evident that the blood 
 plasma must undergo the same fate. It is impossible therefore 
 
APPENDIX. 
 
 203 
 
 to ascribe the serous exsudation to a secretory activity on the 
 part of the endothelial cells. These cells on the contrary 
 must permit of the passage of the plasma and corpuscles in 
 consequence of the contractility which is one of their pro- 
 perties. We may compare this phenomenon to those observed 
 in the Sponges, where the ectodermic cells by their contraction 
 open or close the apertures of the canals and so permit or 
 refuse the passage of fluids with the bodies they may hold in 
 suspension. ■ In both cases, the passage of water through the 
 ectodermic wall in the Sponge, and of plasma through the 
 endothelial wall in inflammation, are brought about in con- 
 sequence of the sensibility and contractility of the cells 
 concerned. Although the process cannot be called phagocytic 
 in the strict sense of the term, yet its intimate relationship with 
 phagocytic phenomena is very evident. 
 
 Phagocytosis is a phenomenon of considerable complexity. 
 When it is exhibited by leucocytes, these cells are in the first 
 place affected by various substances which possess an attrac- 
 tion for them. They proceed towards'these substances by means 
 of their amoeboid movements and then englobe them. Intra- 
 cellular digestion may afterwards occur. Here then we have 
 phenomena of sensibility, contraction, ingestion, and produc- 
 tion of digestive fluids. As a matter of fact the sequence is 
 frequently broken at some point. Thus when a guinea-pig is 
 infected with anthrax, the leucocytes, acted upon by the 
 bacterial products, approach the microbes; leucocytosis takes 
 place, but phagocytosis is arrested, and the bacteria are either 
 not englobed at all, or to a very slight extent. In this case 
 there is a phagocytic reaction, which however falls short of 
 the mark. 
 
 In the most complete phagocytic reaction, all the varieties 
 of phagocytes englobe and destroy the irritant bodies. In 
 other instances it is only the mobile phagocytes which fulfil this 
 function. In a third set of examples, the phagocytic reaction is 
 still less complete. The leucocytes remain in the organs and 
 in the blood and do not pass over into the exsudation ; the en- 
 dothelial cells alone react, but do not accomplish all the phases 
 
204 APPENDIX. 
 
 of phagocytosis ; they stop short at a stage of contraction which 
 allows the plasma and the red blood corpuscles to pass through 
 the vessel wall. The most familiar examples of this incomplete 
 phagocytic reaction are met with in certain very acute diseases 
 produced experimentally. It is in these cases that purely serous 
 inflammation has been most studied. Thus in the acute diseases 
 produced in guinea-pigs by injection of vibrio Metchnikowii or 
 in rabbits by the coccobacillus of hog cholera when death ensues 
 after a few hours, the phagocytic reaction is limited to this 
 condition of contraction of the vascular endothelial cells, giving 
 rise to a serous exsudation in the infected regions. Where 
 the disease takes a still more rapid course, as in the most 
 malignant forms of chicken cholera in the rabbit, no exsuda- 
 tion at all occurs. There is no phagocytic reaction whatever, 
 but at the same time there is no inflammation. 
 
 We see then that serous inflammation is included perfectly 
 well by the general conception of the biological theory without 
 modification. In formulating this theory, I have, in order to 
 make it as short as possible, only mentioned ' phagocytic re- 
 action ' in general terms. I have not alluded either to the 
 sensibility or to the contractility of the phagocytes, since these 
 phenomena are already comprised in the one notion of the 
 phagocytic reaction. For the same reason I have not especially 
 referred to the sensibility and contractility of the endothelial cells, 
 which are certainly phagocytic in their nature, although these 
 phenomena play an important part in the production of the 
 serous exsudation. 
 
 Since the comparative pathology of inflammation has 
 established the reactive and beneficial character of this pheno- 
 menon, we should expect to find certain close affinities between 
 it and other processes of the organism. Thus inflammation is 
 Connected by a whole series of intermediate conditions with 
 other phagocytic phenomena, such as the passage of leucocytes 
 through mucous membranes ; and chronic inflammation is inti- 
 mately associated with the atrophy of certain tissues. In fact, 
 from whatevet side we may consider inflammation, we shall 
 always find connecting links with other natural phenomena. 
 
APPENDIX. 205 
 
 Thus from the purely clinical standpointj no hard and fast 
 boundary line can be drawn between inflammation and hyper- 
 aemia, all the intermediate stages between these two conditions 
 being met with at some time or another. 
 
 II. 
 
 The severest critic of the biological theory of inflammation 
 is Professor Ziegler, who considers that it is quite erroneous to 
 attribute a fundamental importance to phagocytosis in inflam- 
 mation. I regret that limits of space prevent my reproducing 
 here all his arguments, and must content myself with quoting 
 his principal objections. He writes, " Metchnikoff affirms 
 quite arbitrarily that the pathological phenomenon which 
 interests him represents the essential part of inflammation. He 
 is moreover inconsequent in his statements, for he sometimes 
 regards the phagocytosis exercised by leucocytes and sometimes 
 the accumulation of mesodermic cells as the essential feature." 
 He continues, " I look upon the phagocytosis which occurs in 
 the course of an inflammation as a purely accidental pheno- 
 menon, which is often brought about for the simple reason 
 that mobile cells happen to be present, together with a material 
 capable of being ingested by them." (Ziegler, loc. cit., p. 200.) 
 
 When we analyse inflammation by means of a study of its 
 ontogeny, we are inevitably forced to the conclusion that 
 phagocytosis is the most primitive phenomenon of the reaction 
 against irritant agents. Inflammation in cold-blooded animals 
 teaches us that increased temperature is not a necessary 
 factor; and the analogous reaction in invertebrates proves 
 that inflammation may occur without any intervention on 
 the part of the blood-vessels. The phenomena thus become 
 more and more simple as we descend the animal scale, till 
 finally the sole phenomenon we have to deal with is phagocy- 
 tosis. Since the leucocytes are derived from the mesoderm, 
 there is no inconsequence in admitting the existence of a 
 leucocytic phagocytosis and a mesodermal phagocytosis. The 
 facts are so clear that there would be no advantage in dwelling 
 on the matter at any greater length. 
 
206 APPENDIX. 
 
 Ziegler continues, " When any bodies, such for instance as 
 bacteria giving rise to attracting substances, are present at any 
 spot in the tissues, the leucocytes move towards these bodies, 
 and in some cases manage to englobe them. When on the 
 contrary the foreign bodies, such as bacteria, exercise a repel- 
 lent or paralysing effect, the cells will move in an opposite direc- 
 tion, or remain where they are. The emigration and pha- 
 gocytosis then are not brought about by the combative 
 ardour of the cells, but by the properties of the foreign body 
 which has been introduced, or of the tissues and tissue-juices 
 modified by the irritant." He concludes, " The idea that 
 the characteristic feature of inflammation is a struggle of 
 phagocytes must therefore be rejected." (loc. at., p. 202.) 
 
 Ziegler forgets that the attraction and repulsion of leucocytes 
 depend not only on the products of the microbes but also on 
 the sensibility of the leucocytes. Substances which repel the 
 leucocytes of susceptible animals attract on the contrary those 
 of animals which are immune by nature or rendered refractory 
 by vaccination. This fact is so general and well established 
 that no one is justified in ignoring it. It is precisely because 
 on the one side we have microbes which defend themselves or 
 attack by means of their toxic products, and on the other 
 phagocytes which approach and englobe the microbes, that I 
 have formulated the idea of a struggle between two living 
 organisms. None of Ziegler's objections can overthrow this 
 interpretation of the facts just mentioned. 
 
 Ziegler proceeds to make use of an argument which has 
 often been brought forward. "In certain cases the phago- 
 cytosis exercised by the leucocytes may help to destroy the 
 foreign bodies. In other cases however the phagocytosis may 
 aid in the generalisation of an infectious disease, e.g. when 
 the bacteria can multiply within the cells, as in leprosy, or 
 when they are carried about by the cells." (P. 202.) It cannot 
 be denied that the phagocytic reaction is far from representing 
 a perfect mechanism, as is evidenced by the frequency of many 
 diseases. But there is also no doubt that the generalisation 
 of bacteria takes place much more rapidly in the cases when 
 
APPENDIX. 207 
 
 they are not englobed by the phagocytes. Thus one often 
 hears that a seat of injury may become infected by tubercle 
 bacilli which have been carried there by leucocytes ; but 
 it must not be forgotten that in tuberculosis the leucocytes 
 serve to localise the disease and prevent the dissemination of 
 the bacilli. The importance of the leucocytes as carriers of 
 infective bacilli has been very much exaggerated. 
 
 Not approving either of the biological theory of inflammation 
 or of the comparative method on which it is based, Ziegler 
 defines " inflammation as a local tissue-degeneration associated 
 with pathological exsudations from the blood-vessels." (loc. 
 cit., p. 173.) This definition is nothing but a statement 
 of a certain number of the phenomena of inflammation, and 
 does not touch the essential factor in this process. The 
 accumulation of migratory cells in Urodela round irritant 
 bodies, which, as I have shown, takes place without any 
 affection of the vessels, as well as all the analogous phenomena 
 in invertebrata, are quite excluded by Ziegler's definition. 
 And yet any scientific definition should take account of the 
 natural affinities which undoubtedly exist between these 
 phenomena and the inflammation accompanied by a vascular 
 reaction. Another still more striking illustration of the 
 insufficiency of this definition is afforded by tubercle. A 
 tubercle formed within the vessels would not come under 
 Ziegler's definition of inflammation, whereas a tubercle which 
 has developed outside the vessels would be a true inflamma- 
 tion. Now it cannot be denied that intravascular and extra- 
 vascular tubercle represent essentially one and the same 
 pathological formation. I must conclude then that Ziegler's 
 definition, which leaves the main question untouched and 
 takes no account of natural affinities, must be rejected. 
 
 III. 
 
 With the exception of the question of serous inflammation, 
 discussed in Section I., Weigert merely makes a few critical 
 remarks with regard to certain subsidiary points in the 
 biological theory of inflammation. My learned critic expresses 
 
208 APPENDIX. 
 
 doubts as to the justice of comparing the destruction of the 
 microbes within the phagocytes to intracellular digestion. He 
 thinks that there is a contradiction in the fact of intracellular 
 digestion taking place in an acid medium in Protozoa while 
 the processes in the phagocytes are carried out in a neutral or 
 alkaline medium. But besides the case in which the phago- 
 cytes show an acid reaction (exsudation-cells of the tail of 
 tadpoles), I may instance the intracellular digestion in multi- 
 cellular animals. In Actinia this takes place in an acid medium, 
 in the Spongillse in a neutral or alkaline medium. And yet these 
 two examples are closely allied to each other. It should be pre- 
 mised as a general rule that intracellular digestion may present 
 great variations, arid may proceed in media of varying reaction. 
 
 Weigert finds another contradiction between my opinion that 
 bacteria are destroyed within the phagocytes and my men- 
 tion of the ferment of leucocytes, discovered by Leber, 
 which acts outside these cells (a ferment which peptonises 
 gelatin). Here a misunderstanding has arisen. I have never 
 asserted that the phagocytic destruction of bacteria is carried 
 out by means of any ferments, still less by means, of those 
 which peptonise gelatin. I have always openly acknowledged 
 that the question as to what substances within the phagocytes 
 harm and destroy the microbes is still quite undecided. 
 They may be ferments, digestive or otherwise, or they may be 
 substances, acid or alkaline, completely different from fer- 
 ments. We shall have to find new and more perfect methods 
 before being able to solve this delicate problem. 
 
 In the question of giant cells, Weigert remains faithful to 
 his old theory, which has been discussed in the " Annates de 
 rinstitut Pasteur," i883j p. 604. Unfortunately he has not 
 chosen to discuss this subject in his criticism. This is the 
 more to be regretted, since the discovery of the special resis- 
 tance offered by the giant cells of the Algerian rat to the 
 tubercle bacillus might give rise to an interesting change of 
 opinion. It is unnecessary to insist in greater detail on the 
 fact that this discovery is an important confirmation of my 
 views as to! the giant cells being a means of phagocytic defence; 
 
APPENDIX. 209 
 
 IV. 
 
 Having 'now replied to the principal objections made to 
 the biological theory of inflammation, there is one point of a 
 different nature left to deal with. In spite of his opposition to 
 the phagocytic theory, Ziegler lays a certain claim to priority 
 in this question. I may take this opportunity of replying to 
 this attack as well. 
 
 To quote Zjegler's own words : " the phenomenon of phagocy- 
 tosis has been known for a long time ; in the sixth decade of 
 our century, experimental researches on the enclosure of coal- 
 dust and of coloured granules by leucocytes were frequently 
 made, as well as on transport by these cells. In 1874 I ob- 
 served that, in granulation-tissue, cells having the characteristics 
 of leucocytes with divided nucleus, together with red corpuscles, 
 were englobed and destroyed by large cells. As the result of 
 my researches I concluded that the material englobed was 
 assimilated and that an act of nutrition consequently took 
 place" (p. 197). Ziegler insists upon the fact that "his 
 researches on intracellular digestion in the mesodermic cells 
 were published eight years before the works of Metchnikoff, 
 and that at the time Metchnikoff's first papers on phagocytosis 
 appeared, this phenomenon was well known to pathologists " 
 {p.199). 
 
 Ziegler is astonished that I gave no historical references in 
 my earlier works, and especially in my treatise on inflamma- 
 tion. But in my first paper ^ where mention was made of 
 phagoc37tes, I quoted " the valuable results of histological and 
 pathological research on the subject of the phenomena - of 
 absorption in vertebrates." I referred the reader to 
 Ziegler's text-book of pathological anatomy itself. I did not 
 dwell particularly on Ziegler's work^ with regard to the inges- 
 tion of leucocytes by granulation-cells, for the reason that 
 
 ' Arbeiten des zool. Inst, zu Wien, 1883, vol. v. p. 157. 
 2 " Experimentelle Untersuch. iiber die Herkunft der Tuberkel- 
 elemente," 1875, p. 68. 
 
 P 
 
2IO APPENDIX. 
 
 this discovery was made four years previously by Bizzozero,^ 
 who suggested that the leucocytes found in pus in the interior of 
 large cells had been devoured by the latter. Later on Ziegler 
 confirmed this conclusion, but in seeking for the analogies of 
 this process, compared it to the conjugation of cells, such as 
 results in the formation of zygospores in Spirogyra or in that 
 of Plasmodia by the fusion of cells, &c. He thus regards the 
 act of ingestion by the leucocytes as a fusion of cells rather 
 than as a preliminary to intracellular digestion. 
 
 I need scarcely say that in none of my publications have I 
 laid claim to the discovery of the ingestion of solid bodies by 
 mesodermic cells, nor have I ignored the large number of 
 researches which have been made on this subject The 
 phagocjrte theory however is still the theme of so much 
 debate that I have not had the opportunity of entering fully 
 into historical details. When it is once firmly established, it 
 will be time enough to determine the exact part taken in its 
 foundation by workers such as Panum, Gaule, Roser, &c., who 
 have much more right than Ziegler to be considered the 
 pioneers of this theory. 
 
 Ziegler is mistaken in thinking that the ingestion of solid 
 bodies by mesodermic cells necessarily involves the conclusion 
 that the animal organism possesses, in its phagocytic cells, 
 (mesodermic and others) a very important means of defence 
 against pathogenic microbes. In order to establish the fact that 
 the phagocytes constitute an important defensive mechanism, 
 it was necessary to prove that the leucocytes englobe rnicrobes 
 in a living and virulent condition and then injure or destroy 
 them in some way or other. In order to show the impor- 
 tance of the phagocytes, it was necessary to prove that their 
 intervention is of usual occurrence. In this task, I have been 
 obliged to demonstrate the inaccuracy of the work directed 
 against the phagocytic theory and carried out in Ziegler's 
 laboratory. Two of the pupils of this author. Palm ^ and 
 
 1 Caz. med. lombarda, iSyi, and Wz'en. med. Jahresber., 1872, 
 p. 160. 
 
 ^ Beiir. z. path. Anatomie, vol. ii. p. 480. 
 
APPENDIX. 211 
 
 Rogowitch, 1 have made some researches on malignant pus- 
 tule in man and 'charbon symptomatique.' Palm came to 
 the conclusion that in anthrax of man " the cells do not play 
 the slightest part in the sense of the phagocytosis of Metchni- 
 koff." Rogowitch expressed the same opinion as regards 
 ' charbon symptomatique ' in several species of animals. And 
 yet it has been definitely shown that both these attacks were 
 based on unreliable data. It is at the present time indubitably 
 proved that the bacilli are englobed in large quantities by 
 the phagocytes both in malignant pustule in nian^ and in 
 'charbon symptomatique.' ^ The attack emanating from 
 Ziegler's laboratory is thus unjustified by facts. 
 
 In this reply to the criticisms on the biological theory of 
 inflammation, I have only considered the more important 
 objections. None of them seem to me to touch either the 
 foundation of the theory or the method upon which it has been 
 built up ; hence I do not think that it can be regarded as having 
 been in any way injured by the opposition raised to it. 
 
 E. Metchnikoff. 
 
 ^ Beitr. z. path. Anatomic, vol. iv. p. 291. 
 
 ^ Karg, Fortschr. d. Med. vol. vi. p. 529, and Lubarsch, " Unters. 
 lib. d. Immunitat," 1891, pp. 1 11- 114. The latter author summarises 
 his chapter in the following words : " The existence of phagocytosis 
 in human anthrax, as well as its parallelism with the course of the 
 disease and the destruction of the bacilli should be considered as 
 established beyond doubt by the facts I have brought forward." 
 It is inconceivable how Roger (Traitd de Medecine de Charcot et 
 Bouchard, vol. i. p. 555) could interpret Lubarsch's article in a 
 directly opposite manner and assert that in the cases quoted by 
 him " there was no connection whatever between the intensity of 
 the phagocytosis and the evolution of the disease." 
 
 ^ Annates de Vlnstitut Pasteur, 1889, p. 194. RufFer, Brit. 
 Med. Journ., May 24, 1890. 
 
 P 2 
 
INDEX. 
 
 Absorption by leucocytes, 190. 
 Acinetse, parasitism of, 2, 22. 
 Actinophrys, 15. 
 Amoeba, merotomy of, 14. 
 Amoeboid cells in Sponges, 45 . 
 Amphioxus, absence of reactive phe- 
 nomena in, 94. 
 Amphophile leucocytes, 113. 
 Anguillulides, 73. 
 Anisoplia austriaca, 89. 
 Annelida, phagocytosis in, 66. 
 
 formation of plasmodia in, 67. 
 Antitoxic function of serous exsudation, 
 
 177. 
 Antitoxines, 178. 
 Arachnidse, 88. 
 Argulus, 78. 
 Arnold, on stomata, 138. 
 
 division of leucocytes, 129. 
 Arthrobotkrys oligospera, 73. 
 Arthropoda, 75 — 93. 
 Ascaris mystax, 73. 
 Ascidians, diapedesis in, 79. 
 Astropecten pentacanthus, 60. 
 Atrophy, 171. 
 Attraction-sphere, 113. 
 Aurelia aurita, 59. 
 Axolotl embryos, inflammation in, 95. 
 
 Bacilli, alteration of, in amoebae, 19. 
 
 englobed in living condition, 115. 
 
 tubercle, degeneration of, l6i. 
 Bacillus pyocyaneus, 121. 
 Bacteria, ingestion by phagocytes of 
 
 Bipinnaria, 64. 
 Bacteria, luminous, 82. 
 Bacterial infection of insects, 89. 
 Bacterium termo, chemiotaxis in, 34. 
 Balbiani, on bacteria in insects, 88. 
 
 on merotomy of Infusoria, 16. 
 
 epidemic in Paramsecia, 20. 
 
 on " pdbrine," 92. 
 
 Bardach, 161. 
 
 Bary, de, on fungi in caterpillars, 89. 
 
 on Peziza scltrotiorum, 41. 
 Basophile cells, 143. 
 Baumgarten, criticism of biological 
 theory, 193. 
 
 formation of tubercle, 158. 
 Bipinnaria asterigera, 62. 
 Behring, on bactericidal action of 
 serum, 175. 
 
 and Kitasato, on antitoxic proper- 
 ties of serum, 177. 
 Billet, 82. 
 Blood-corpuscles of invertebrata, 76. 
 
 white, origin of pus-cells from, 9. 
 Blood-vessels, development of, 137. 
 
 in inflammation, 182. 
 
 behaviour in earthworm, 70. 
 Boinet and Borel, 169. 
 Bollinger, 121. 
 Bombincttor igneus, 103. 
 Bordet, 118, 145. 
 Brault, 170. 
 
 views on inflammation, 9. 
 Bruno Hofer, digestion in amcEbse, 125, 
 Buchner, 156. 
 
 on bactericidal properties of serum, 
 176. 
 
 on chemiotaxis, 119. 
 
 on inflammation, 11. 
 Burdon Sanderson, criticism of bio- 
 logical theory, 193. 
 Biitschli, 73. 
 
 epidemic in Paramsecia, 20. 
 
 on Acinetse, 23. 
 
 Calcareous degeneration of tubercle, 
 
 162. 
 Canu, 81. 
 
 Capillaries, contractility of, 138. 
 Cattani, 177. 
 Cell-division in inflammation, 9, 10. 
 
214 
 
 INDEX. 
 
 Ceraochalina gibbosa, 52. 
 Chsetorceros, parasitic on Bipinnaria,62. 
 Chambers, ciliated, of Sponges, 46. 
 Charrin, 153- 
 Chemiotaxis, 33, 117. 
 
 in developing blood-vessels, 189. 
 
 negative, 146. 
 Chklarewsky, 147. 
 
 Chloroform, action on leucocytes, 148. 
 Chondrioderma, plasmodia of, 37. 
 Chondrosia reniformis, 49. 
 Chronic inflammations, 157-171. 
 Chun, fragmentation of nucleus, 167. 
 Chytridiacese, parasitism of, 24. 
 
 parasitic in Nematoda, 73. 
 Ciliated chambers, 46. 
 Cirrhosis, 170. 
 Clasmatocytes, 143. 
 Cleonus punciiventris, 90. 
 Cockchier, 78. 
 Coelenterata, 56 — 65. 
 Cohnheim, dilatation of vessels in in- 
 flammation, 149. 
 
 views on inflammation, 6, 181. 
 
 experiments on cornea, 8. 
 
 origin of granulation-cells, 131. 
 Coleoptera, phagocytosis in larval, 78. 
 Commensalism in Sponges, 53. 
 Comparative pathology, importance of, 
 
 4, 12, 195. 
 Conidia, parasitic in Daphnia, 83. 
 Connective tissue, formation of, j 32. 
 
 its changes ifl inflammation, 142. 
 Connective tissue cells, behaviour dur- 
 ing inflammation, 97. 
 
 karyokinesis in, 97. 
 
 formation from migratory cells, 106. 
 
 in tubercle, 158. 
 Copepoda, 75. 
 Cordiceps militaris, 89. 
 Cornea, Cohnheim's experiments on 
 
 the, 8. 
 Cornil, on inflammation, 8. 
 Crustacea, phagocytosis in, 78. 
 Cuticle, protective function of, 87. 
 Cyclostomata, leucocytes of, 114. 
 Cyst, formation by Gregarina, 69. 
 
 Danilewsky, 152. 
 Daphnia, fungoid disease of, 82. 
 Delacaze-Duthiers, 78. 
 Diapedesis, 144, 185. 
 
 in Ascidians, 79. 
 
 in vaccinated animals, 1J4. 
 Didymium farinaceum, 35. 
 Digestion, intracellular, in Sponges, 49. 
 Diphtheria, 177. 
 Diptera, 88. 
 
 Diseases, infections, of Infusoria, 
 17—28. 
 in AmcEba, 17. 
 in ciliated Infusoria, 20. 
 Disselhorst, action of quinine, 145. 
 Doris, 77. 
 Dust-cells, 160. 
 
 Earth-worm, diseases of, 68. 
 Ebstein and Nicolaier, 169. 
 Echinodermata, 56 — 65. 
 
 development of, 60. 
 Ehrlich, 112. 
 
 leucocytosis in phthisis, 168. 
 Emigration of leucocytes, 144. 
 Endothelium, development of, 137. 
 
 peritoneal, phagocytic nature of,65. 
 Eosinophile leucocytes, 112. 
 Epidemic in Amoebae, 17. 
 
 in ciliated Infusoria, 20. 
 Epithelioid cells, origin of, 132, 159. 
 Erysipelas, behaviour of leucocytes in, 
 117. 
 
 leucocytosis in, 122. 
 Englena, infection of, 24. 
 Euphorbia Cyparissias, 41. 
 Evolution of inflammation, 109. 
 
 Ferments in leucocytes, 123. 
 
 Fibroblasts, origin of, 132. 
 
 Fishes, 95. 
 
 'Flacherie,'89. 
 
 Flagellated Monads, digestion in, 19. 
 
 infection of, 25. 
 Flemming, on attraction -sphere in 
 leucocytes, 113. 
 
 mitotic division of leucocytes, 130. 
 
 transformation of leucocytes, 135. 
 Fluss, 150. 
 Fraenkel, criticism of biological theory, 
 
 192. 
 Frank, lesions in plants, 39. 
 Fungi, parasitic in Nematoda, 73. 
 
 parasitic onTalitrus, 81. 
 
 Gabritchevsky, sensibility of leuco- 
 cytes, 119. 
 Gamaleia, 154. 
 Gasteropoda, spleen of, 77. 
 Gasteropteron, 77. 
 Gastrsea theory, 43. 
 Gastrula, 43. 
 
 -stage in star-fish, 60. 
 Giant cells, action on tubercle bacilli, 
 162. 
 
 origin of, 135, 159. 
 
 in Arthropoda, 79. 
 Giard, 82. 
 
INDEX. 
 
 215 
 
 ' Glande ind^termin^e,' 78. 
 
 Gley, 153. 
 
 Goloubew, 138. 
 
 Gonococci, behaviour of leucocytes to, 
 
 117. 
 Gordius, parasitic on Nais, 67. 
 Granulation-tissue, origin of, 132. 
 Granuloma, formation of, 158. 
 Grawitz, 9, 132. 
 
 Gregarina, parasite of Earth-worm, 68. 
 Griesbach, on phagocytosis in Molluscs, 
 
 77- 
 Grobben, 66. 
 Gryllid^, 88. 
 
 HiECKEL, Gastrsea theory, 43. 
 
 on ingestion of granules by leuco- 
 cytes, 76. 
 Hafkine, epidemic in Paramsecia, 21. 
 Hankin, 176. 
 
 Hanstein, on lesions of algae, 16. 
 Hemitis, 186. 
 
 Hepatic capillaries, endothelium of, 139. 
 Hering, mechanical theory of inflam- 
 mation, 144. 
 Hermann, 81. 
 Hertwig, 156. 
 Hess, 126. 
 
 Hircinia echinata, 52. 
 Hofer, on digestion in Amoebae, 19. 
 Hofmeister, peptone in leucocytes, 124. 
 Hydra, reaction to injuries, 57. 
 Hydromedusae, 58. 
 Hydrotropism in plasmodia, 39. 
 Hymenoptera, 88. 
 Hyperaemia, J. 
 Hypertrophic cirrhosis, 170. 
 
 Ibener, 176. 
 
 Immunity, 156. 
 
 Infectious diseases, absence in Sponges, 
 
 54- 
 of Infusoria, 2, 17 — 28. 
 Inflammation, author's definition of, 
 189. 
 Comil and Ranvier's definition of, 
 
 8. 
 Neumann's definition of, 11. 
 Ziegler's definition of, 7. 
 theories of, 5, 180. 
 biological theory of, 184. 
 paralytic theory of, 6. 
 spasmodic theory of, 6. 
 Virchow's theory of, 6, 180. 
 Samuel and Cohnheim's views, 6, 
 
 181. 
 evolution of, 109. 
 in plants, 39. 
 in Triton embryo, 98 — 103. 
 
 Inflammation without diapedesis, 186. 
 Inflammations, chronic, 157 — 171. 
 
 serous, 171 — 179. 
 Infusoria, artificial lesions in, 15 — 17. 
 
 biological relations between, 2. 
 
 colonies of, 44. 
 
 infectious diseases of, 17 — 28. 
 Insects, varieties of phagocytes in, 88. 
 Intracellular digestion in Molluscs, 78. 
 Invertebrata, leucocytes of, 76. 
 Isaria destructor, 90, 
 Ischikawa, on Hydra, 57- 
 'Itio in partes^ 147. 
 
 Jequirity, action on leucocytes, 119. 
 
 Karyokinesis, 9, 10. 
 
 in connective tissue cells, 97. 
 Kebner, 169. 
 
 Keratitis, experimental, 8. 
 Kitasato, 177. 
 Klebs, 138. 
 
 on infection of Euglena, 24. 
 Klemperer, on antitoxic properties of 
 
 serum, 177. 
 Xoch, 125. 
 
 giant cells, 167. 
 Korschelt, 61. 
 Koultchitsky, 130. 
 
 Kowalewski, on phagocytes of Asci- 
 dians, 79. 
 
 on spleen of Molluscs, 77. 
 Krukenberg, peptic ferment in Myxo- 
 mycetes, 30. 
 on a tryptic ferment in Sponges, 
 SO. 
 Kupffer, on star-cells, 140. 
 
 Landerer, 150. 
 
 Lange, 117. 
 
 Leber, chemiotaxis of leucocytes, 117. 
 
 digestive ferment in pus, i'24. 
 Le Dantec, digestion in Stentor, 20. 
 Lendenfeld, de, 55. 
 
 physiology of Sponges, 49. 
 Lepidoptera, 88. 
 Leprosy, 168. 
 
 Leprosy bacilli, behaviour of leucocytes 
 to, 117. 
 
 ingestion by endothelial cells, 140. 
 Leptotrix, digestion of, 19. 
 
 in young Spongillae, 52. 
 Lesions, traumatic, in lower animals, 
 
 14. 
 Lethargy of silkworms, 89. 
 Leucin, action on leucocytes, 119. 
 Leuchsemia, 114. 
 Leucocytes, in — 136. 
 
2l6 
 
 INDEX. 
 
 Leucocytes, emigration of, 144. 
 
 in tubercle, 159. 
 
 ofinvertebrata, 76. 
 
 origin of pus-cells from, 9. 
 
 peripheral distribution of, 147. 
 
 psychical activity of, 192. 
 
 varieties of, in. 
 Leucocytosis after injection of pro- 
 teins, 121. 
 Leydig, 138. 
 Lieberkiihn, 114. 
 
 digestion of Infusoria in Sponges, 
 
 47- 
 Limbeck, von, on leucocytosis, 121. 
 Lubarsch, 118, I2i, 176. 
 
 on reactive phenomena in Asci- 
 dians, 80. 
 Luminous bacteria as parasites, 82. 
 Lupus, 166. 
 Lymphocytes, 114. 
 in tubercle, 159. 
 
 Macrophages, 161. 
 as scavengers, 190. 
 
 Malaria, endothelial cells in, 141. 
 
 Marchand, 132. 
 
 Marmot, tubercle of, 160. 
 
 Massart and Bordet, irritability of 
 leucocytes, 118, 145. 
 
 ' Mastzellen,' 143. 
 
 Medusas, inflammation in, 59. 
 
 Meriones, tubercle in, 162. 
 
 Merotomy of Amoeba, 14. 
 
 Mesoderm, formation of, in star-fish, 61. 
 
 Mesostonmm Ehrenbergi, 66. 
 
 Microbicidal action of serum, 174. 
 
 Microsphsera, 17. 
 
 Microsporidium of ' pebrine,' 92. 
 
 Migratory cells, conversion into con- 
 nective tissue cells, 106. 
 
 MoUusca, 75 — 93. 
 
 Monads, digestion in flagellated, 19. 
 infection of, 25. 
 
 Monocystis (Gregarinse) 68. 
 
 Monospora bicuspidata, 83. 
 
 Mucor helminthophorus , 73. 
 
 Miiller, epidemic in Paramsecia, 20. 
 
 Muscardine, 90. 
 
 Muskatbluth, 130, 169. 
 
 Myelocytes, 114. 
 
 Myxomycetes, plasmodium of, 29. 
 
 Nais proboscidea, 67. 
 Neisser, 140, 169. 
 Nemaloda, 70 — 74. 
 
 producing tubercles, 170. 
 Nerves, their influence in inflamma- 
 tion, 150. 
 
 Netchaeff', on litmus-granules in leu- 
 cocytes, 124. 
 
 Neumann, on inflammation, ir. 
 
 Neutrophile leucocytes, 113. 
 
 Nikiforoff, 131. 
 
 Nissen, 175. 
 
 Nucleus, importance for regeneration, 
 IS- 
 
 CEdema, inflammatory, 173. 
 Olpidium, 25. 
 Orthoptera, 88. 
 Oryctes, 78. 
 Ostracoda, 75. 
 Ouskoff, 114. 
 
 Pandorina morum, 25. 
 Paralytic theory of inflammation, 6. 
 Paramsecia, epidemic in, 20. 
 Parasitism in Paramsecia, 20. 
 
 of Infusoria, 2. 
 Parenchymatous inflammation, 9. 
 Pasteur, on diseases of silkworms, 89. 
 Pasteuria ramosa, 87. 
 Pathology, necessity for comparative, 4, 
 
 12, 195. 
 'Pebrine ' infecting Daphnia, 87. 
 
 of silkworms, 92. 
 Pee, on leucocytosis, 121. 
 Pekelharing, 1 18. 
 Peremeschko, 130. 
 
 Pericardial phagocytes of insects, 88. 
 Peritoneal endothelium, phagocytic 
 
 nature of, 66. 
 Perivisceral cells in Worms, 56. 
 Petruschky, 125. 
 Peziza sclerotiorum, 41. 
 Pfeffer, on chemiotaxis, 33. 
 Phagocytella, 43. 
 Phagocytes in HydromedusK, 58. 
 
 their absence in certain Crustacea, 
 87. 
 
 fate of, 107. 
 
 in insects, 88. 
 
 in tubercle, 159. 
 Phagocytic properties of leucocytes, 
 
 114. 
 Phagocytosis in Annelida, 66, et seq. 
 
 in Crustacea, 78. 
 
 in Daphnia, 84. 
 
 in recurrent fever, 186. 
 
 in Sponges, 47, 51. 
 
 in star-fish, 61. 
 Philina, 77. 
 Philliroe, 77. 
 Physarum, 31, 36. 
 Piorry, 186. 
 Plants, inflammation in, 39. 
 
INDEX. 
 
 217 
 
 Plasma-cells, 143. 
 
 Plasmodia, formation of, in Bipinnaria, 
 
 63- 
 
 formation of, in Annelida, 67. 
 
 formation by fusion of cells, 52. 
 
 of Myxomycetes, 29. 
 Pleurobranchsea, 77. 
 Pneumonia, 177. 
 Polyphagus EugUna, 25. 
 Ponfick, 140. 
 Protospongia, 45. 
 Protozoa, digestion in, 18. 
 
 reactive phenomena of, 15—28. 
 Pseudo-eosinophile leucocytes, 113. 
 Pseudo-navicellse, 69. 
 Pus-cells, origin of, 9. • 
 
 Quinine, action on leucocytes, 119, 
 145. 
 
 Ranvier, on inflammation, 8. 
 division of leucocytes, 129. 
 endothelial cells of omentum, 
 142. 
 Rat's serum, bactericidal action of, 176. 
 Reaction of digesting cells of Spongilla. 
 
 50. 
 Recklinghausen, emigration of leuco- 
 cytes, 149. 
 Recurrent fever, 186. 
 Reparation in inflammation, 10. 
 Rhabditis invading Earthvi'orm, 70. 
 Rhizopoda, digestion in, 19. 
 Mhizostomum Cuvieri, 59. 
 Ribbert, 132. 
 
 lymphatic glands, 142. 
 Roeder, 176. 
 Roemer, 119. 
 Roger, 151. 
 Roser, views on inflammation, 10, 
 
 110. 
 Rossbach, 123. 
 Roux, 176. 
 
 and Yersin, on pleurisy in diph- 
 theria, 173. 
 
 Sachs, on inflammation, 11. 
 Samuel, 147, 150. 
 
 theory of inflammation, 6, 181. 
 Sandhoppers, fungoid disease of, 81. 
 Saprolegnise infecting Crustacea, 82, 
 
 86. 
 Saville Kent, inclusion of bacteria by 
 
 Physarum, 38. 
 
 on Protospongia, 45. 
 Sawtchenko, 117, 169. 
 Scavenger function of leucocytes, 190. 
 Schafer, 127. 
 Schueppel, 162. 
 
 Schultze, Max, 113. 
 
 Selenka, 61. 
 
 Sensibility of leucocytes, 117. 
 
 tactile, of leucocytes, 123. 
 Serous exsudation, functions of, 174, 
 
 177. 
 Serous inflammations, 171 — 179. 
 Serum, bactericidal action of, 175. 
 Severini, 138. 
 
 Silkworms, diseases of, 89, 91. 
 Siphonochalina coriacea, 47. 
 Snellen, 152. 
 Soudakewitch, 166, 169. 
 Spasmodic theory of inflammation, 6. 
 Spermatozoids of ferns, chemiotaxis in, 
 
 34- 
 Spermophilus, tubercle of, 160. 
 Sphierophrya magna, 3. 
 Sphserophrya parameeciorum, 3. 
 Spirilla, 186. 
 
 Spirobacillus Cienkowskii, 87. 
 Spleen of Gasteropoda, 77. 
 Sponges, 45—55- 
 
 effects of wounding, 51. 
 
 phagocytosis in, 47, 51. 
 Spongilla, digestion in, 48. 
 Spores, fate of, in leucocytes, 128. 
 Sporozoa infecting Crustacea, 82, 87. 
 
 infecting insects, 92. 
 Spronck, on leucocytes, 130. 
 Spumaria alba, 148. 
 Stahl, on plasmodia of Myxomycetes, 
 
 33- 
 Star-cells, 140. 
 ' Staubzellen,' 160. 
 Stentor, merotomy of, 16. 
 Stephanos typhus, 53. 
 Stern, 174. 
 Stigmata, 138. 
 Stomata, 138. 
 Strieker, no, 138. 
 Stylonychia, 22. 
 Swine septicEemia in pigeons, 141. 
 
 Tadpoles, inflammation in, 104. 
 
 Talitrus (sandhopper), 81. 
 
 Tchistowitch, 135, 160. 
 
 Temperature in inflammation, 12. 
 
 Terebella, 66. 
 
 Tetanus, I77' 
 
 Theories of inflammation, 5, 180. 
 
 Thethys, 76. 
 
 Thiocystis, 20. 
 
 Thoma, on inflammation, 11. 
 
 Tissue-cells, origin of pus-cells from, 9. 
 
 Tizzoni, 177. 
 
 Teuton, 140. 
 
 Trachelius ovum, 16. 
 
 Trapeznikoff, 129. 
 
2l8 
 
 INDEX. 
 
 Traumatic lesions in lower animals, 
 
 14. 
 Triton txniatus, 98. 
 Trophotropism, 33. 
 Tubercle bacilli, degeneration of, 161. 
 Tuberculin, injection of, 168. 
 
 effect on guinea-pigs, 178. 
 Tuberculosis, 157 — 168. 
 
 avian, 161. 
 
 zooparasitic, 170. 
 Tunicata, 75—93. 
 
 Turbellaria, phagocytic reaction in, 
 66. 
 
 Urodela, inflammation in, 95. 
 Uromyces Pisi, 41. 
 
 Vaccination, effect on inflammatory 
 
 reaction, 154. 
 Vacuolar degeneration, 169. 
 Vaillard, 177. 
 Vascular system in inflammation, 8. 
 
 in invertebrata, 75. 
 Vasomotor nerves in inflammation, 
 
 151. 
 Vaucheria, recovery from injury, 17. 
 
 Vermes, 65 — 74. 
 
 Vertebrata, inflammation in, 94—197. 
 
 Vesuvine, reaction of ingested bacilli 
 
 Vibrio Metchnikowii, I54> I7°- 
 Virchow, theory of inflammation, 6, 
 180. 
 
 Waldenburg, inflammation in 
 
 plants, 39. 
 Weber's law applied to lower animals, 
 
 34- 
 Weigert, giant cells, 166. 
 origin of pus-cells, 9. 
 Worms, inflammation in, 65 — 74. 
 Wyssoko witch, 140. 
 
 Yersin, 135. 
 
 ZlEGLER, on inflammation, 7. 
 
 origin of connective tissue, 131. 
 Zimmerman, 173. 
 Zoochlorellsd, 53. 
 Zooparasitic tuberculosis, 170. 
 Zoox.mtellss, 53. 
 Zopf, 73. 
 
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