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Originally appearing in Volume V27, Page 347 of the 1911 Encyclopedia Britannica.
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TRYPANOSOMES, or HAEMOFLAGELLATES, minute Protozoan parasites, characterized by the possession of one or two flagella and an undulating membrane, and specially adapted for life in the blood of a vertebrate?. Of late years considerable progress has taken place in our knowledge of these organisms, research upon them having been stimulated by the realization of their extreme importance in medical parasitology. Not only has the number of known forms been greatly multiplied, but the study of the biology and life-history of the parasites has been attended in some cases with remarkable and unexpected results. Historical.—The first observation of a trypanosome is usually ascribed to Valentin (55), who in 1841 announced his discovery of certain amoeboid parasites in the blood of a trout. In the two or three years following several other observers recorded the occurrence of similar haematozoa in various fishes. The generic name of Trypanosortia was conferred by Gruby in 1843 upon the well-known parasite of frogs. E. Ray Lankester (18) subsequently described this same form (under the name of Undulina ranarum) A (From Lankester.) is shown. and was the first to indicate the presence of a nucleus in the cell-body. To Mitrophanow (1883–1884) and Danilewsky (1885–1889) we owe the first serious attempts to study the comparative anatomy of these haematozoa. Trypanosomes were first met with in cases of disease by Griffith Evans, who in 188o found them in the blood of horses suffering from surra in India. In 1894 (Sir) David Bruce discovered the celebrated South African parasite (T. brucei) in cattle and horses laid low with nagana or the tsetse-fly disease; and this worker subsequently demonstrated, in a brilliant manner, the essential part played by the tsetse-fly in transmitting the parasites. The credit for first recognizing a trypanosome in human blood, and describing it as such, must undoubtedly be assigned to G. Nepveu (1898) . Trypanosomes were next seen in human blood 1 Trypanophis, although lacking (so far as is known) a haemal habitat, is included here, since it is undoubtedly closely related to Try anoplasma. 2 The illustrations in this article are from H. M. Woodcock's " Trypanosomes," in the Quarterly Journal of Microscopical Science. B in Senegambia in Igor, in a European suffering from intermittent fever. Forde discovered the parasites, but was uncertain of their nature; he shewed them to E. Dutton, who (ii) gave this form the name of Trypanosoma gambiense. A year later A. Castellani (6) found the organisms (most probably the same species) in the cerebro-spinal fluid of patients suffering from sleeping-sickness in Uganda; and it has since been conclusively proved by Sir David Bruce and D. Nabarro (4) that they are the true cause of that dreadful malady. More important,' from the standpoint of protozoology, than these interesting medical discoveries have been the investigations by A. Laveran and F. Mesnil (20-24), L. Leger (30-35), S. Prowazek (47),F. Schaudinn (50) and others, upon numerous tolerated (i.e. non-pathogenic) forms; these researches supply, indeed, practically all the material facts on which to base an account of the Haemoflagellates at the present day. Trypanosomes are harboured by members of all the chief classes of vertebrates with the exception of cyclostomes. By occurrence. far the greater number of hosts are furnished by fishes, birds and mammals. Among batrachians the parasites have been found, up till now, only in frogs; and among reptiles their occurrence has only been observed in one or two solitary instances (T. damoniae, fig. 3 J). Data with regard to the frequency with which individual species occur, in any kind of host, are as yet somewhat scanty; in one or two cases the parasites are fairly common, T. lewisi, for example, being met with in a considerable percentage of sewer-rats throughout the world. In considering the occurrence of Trypanosomes in mammals, careful distinction must be drawn between natural or true hosts, which are tolerant of the parasites, and casual ones, which are unaccustomed and unadapted to them. A Trypanosome usually produces markedly harmful effects upon gaining an entry into animals which have never been, by their distribution, liable to its invasion previously. Such a state of affairs is produced by the march of civilization into the " hinter-lands " of the various colonies, when man, together with the numerous domesticated animals which accompany him, is brought into proximity to big game, &c., and, what is equally important, into the zone of the particular blood-sucking insects which prey upon the same. Very many of the common domestic mammals can be successfully infected (either thus accidentally or else on purpose) with different " pathogenic " Trypanosomes, to which they succumb more or less readily, but they cannot be regarded as the natural hosts of those Trypanosomes. In dealing with disease-causing forms, the more narrowly the original source of the parasite concerned is defined, the closer do we get to the true vertebrate host or hosts. In the case of the naganaparasite, various Antilopidae (e.g. the gnu, bushbuck and koodoo) can certainly lay a strong claim to the honour. The capybara, again, is most probably the native host of T. equinum of mal de caderas of horses in South America. Similarly with regard to the many other pathogenic Trypanosomes now known, there is undoubtedly, in each case, some indigenous wild animal tolerant of that particular form, which serves as a " latent source of supply " to strange mammals. The transmission of the parasites from one vertebrate individual to another is effected, in the great majority of cases,l Trans- by a blood-sucking invertebrate, and by this means mission; alone. The " carrier " of a Trypanosome of warm-Anernation blooded vertebrates is, in all instances so far deafHosts. scribed, an insect, generally a member of the Diptera; in the case of parasites of cold-blooded vertebrates the same role is usually played by an ichthyobdellid leech (piscine forms), but possibly, now and again, by an Ixodes (amphibian or reptilian forms). Until lately it remained quite uncertain, however, whether the invertebrate merely conveys the Trypanosomes or whether 1 Trypanosoma equiperdum, the cause of dourine in horses and asses, is apparently only conveyed by the act of coitus. This direct mode of transmission is most likely a secondary is a true alternate . host, one i.e. in which definite stages of the parasite's life-cycle are undergone. Schaudinn (50), who investigated certain avian Trypanosomes, considered the latter view to be correct, and believed, that the carrier—in this in-stance a gnat— is indeed the definitive host, i.e. the one in which sexual conjugation occurs. Many other workers have since studied the subject and, so far as the parasites of fishes are concerned, there can be little doubt, thanks to the researches of E. Brumpt (5a), L. Leger (32, 33) and others, that leeches are true alternate hosts for these forms, in which certain phases of the life-cycle are normally undergone. We cannot write quite so confidently with regard to the relation of the various pathogenic Trypanosomes to Tsetse-flies (Glossinae). In the first place experiment has shown that biting-flies, other in all probability than the true, natural hosts, may at times transmit the parasites—as it were—accidentally, if, after feeding on an infected animal, they are allowed to bite a fresh one within a limited time. One very helpful factor in determining which is the principal carrier of any form is the coincidence of the zone of a particular insect with that of any disease. By this means it has been ascertained with practical certainty that, among the family of Tsetse-flies (Glossinae) for instance, at least four species are the natural carriers of different Trypanosomes. Of these perhaps the best-known is G. palpalis, of Equatorial Africa, whose bite transmits the human parasite (T. gambiense). Nevertheless, the fact, commented upon by several observers, that even here an infected fly is only infectious for a comparatively short period suggests that this species of fly, at any rate, is not the true alternate host in which the life-cycle of that particular Trypanosome is completed. However, indications furnished by Koch (16a) point in this connexion to G. fusca. Lastly, before leaving this interesting and important subject, F. Stuhlmann's work (54a) on developmental phases of T. brucii, the nagana parasite in G. fusca and G. tachinoides, does render it probable that the pathogenic forms also have true invertebrate hosts. Schaudinn had fully described the relations of certain avian Trypanosomes to their invertebrate host, Culex pipiens (females). The distribution of the parasites in the gnat is closely Habitat: connected with the process of digestion. The Try- Effects on panosomes ultimately overrun practically all parts float. of the body, sometimes not even the ova escaping. Thus true hereditary infection of a succeeding generation of gnats may be brought about. The life of the parasites while in the insect is characterized by an alternation of active periods, during which multiplication goes on, with resting-periods, when the Trypanosomes become attached to the epithelial cells of the host. According to S. Prowazek (47), the behaviour of T. lewisi in a louse (Haematopinus) is, in its main features, similar. On gaining an entry into the blood of a vertebrate the organisms pass rapidly into the general circulation, and are thus carried all over. Considering them first in a tolerant host, the trend of observation is to show that they are never abundant, but on the contrary usually somewhat scarce. One reason for this scarcity is to be sought in connexion with the fact that multiplicative stages are very rarely met with, at any rate in the general circulation. The parasites are frequently more numerous in the spleen, bone-marrow, kidneys, &c., than else-where, and it has been found that multiplication goes on rather more actively in the capillaries of these organs. The Trypanosomes, in the active phase, are of course always free in the blood plasma (interglobular). In the majority of cases it is very uncertain whether they actually come into relation with the blood corpuscles or not. Schaudinn has stated, however, that Trypanomorpha becomes, in certain phases, attached to a red blood-corpuscle (ectoglobular), and, in others, penetrates inside one and eventually destroys it (endoglobular); while his other avian parasite, Trypanosoma ziemanni, apparently draws up into itself the white corpuscle (leucocyte) to which it becomes attached. In addition, there are two or three obervations to hand which shew that piscine, amphibian and mammalian Trypanosomes may also become attached. Probably most forms possess a resting, attached phase at some period or other, in the invertebrate, if not in the vertebrate host. Considering now the Trypanosomes in an unaccustomed, mammalian host, they may either remain infrequent or rare (sometimes, indeed, being unnoticed until shortly before death), or, on the other hand, they may soon become numerous and go on increasing (fig. 2). In the latter case the disease is acute and rapidly fatal; in the former it is more chronic and lasts much longer, often several months. The main features of trypanosomosis, or illness caused by a Trypanosome, show a general agreement, whichever variety is considered; one symptom may be, of course, more marked than another in any particular case. Death is due either to weakness and emaciation (in chronic cases), or to blocking of the cerebral capillaries by the parasites (where these are abundant), or to disorganization of the nervous system (paraplegic and sleeping-sickness cases). In post-mortem examination, the most obvious pathological lesion is hypertrophy of the spleen, which may be very pronounced; the lymphatic glands in the neck, inguinal region, &c., are also often greatly swollen. These are undoubtedly the organs which react most strongly to the parasites, and their enlarged condition is to a great extent due to their enhanced activity in elaborating blood-corpuscles and leucocytes to cope with the enemy. Ingestion and dissolution of the Trypanosomes by phagocytes has frequently been observed; and it is probable also that the haematopoietic organs secrete some substance which exerts a harmful action on the parasites, and causes them to undergo involution and assume weird-looking " amoeboid " and " plasmodia) " forms. A peculiar feature in the behaviour of the parasites, which is most probably caused by unfavourable biological conditions Ageomera-in the host, is that known as agglomeration. The don• process is readily brought about artificially by the addition of sera or chemical solutions to blood containing the parasites. Agglomeration consists in the grouping or union together of several Trypanosomes around a common centre; this leads to the formation of rosette-like clusters, or even of Iarge masses composed of several rosettes. The end by which the parasites join is typically, in the case of Trypanosoma, the non-flagellate (anterior) end. If a favourable change in the surrounding medium sets in, the Trypanosomes are able to undergo the reverse process, namely disagglomeration; the parasites liberate themselves and the rosette is dissolved. Trypanosomes vary greatly with regard to size; even in one and the same species this variation is often noticeable, especially under morphology. different conditions of life. The common Trypanosoma rotatorium of frogs (fig. 4, A and B) is, taking it all in all, one of the largest forms so far described. Its length (inclusive of the flagellum) varies from 4o-6o µ, while its greatest width (including the undulating-membrane) is from 8-3o µ; in the very wide individuals breadth is gained more or less at the expense of length. Conversely, T. gambiense, the human parasite (fig. 3 C), is one of the smallest forms known, its average size being about 2I-23.e by 14--2 µ. There is equally great diversity in respect of form. Typically, the body is elongated and spindle-shaped ; it is usually more or less curved or falciform (fig. 3, A-D), and tends to be slightly compressed laterally. It may be, however, anything from extremely slender or vermiform (fig. 3, H) to squat and stumpy (fig. 3, G, 4, A). Moreover, apart from the fact that a full-grown adult, ready to divide, is in many cases much plumper than a young adult (cf. T. lewisi, fig. 6, A and B), there can be no doubt that considerable polymorphism also sometimes occurs (e.g. T. rotatorium). In many cases, at any rate, this indicates a difference in sexuality; and it is particularly necessary to bear this factor in mind when consideringthe avian Trypanosomes, where, perhaps, the extremes of form are to be met with. 'That one and the same species may appear entirely different in different phases of the life-history is manifest on comparing, for instance, the chief " forms " of Trypanosoma somes, to illustrate the chief morphological characters. A, Try panosoma lewisi, after Bradf. and Plimmer. B, T. brucei, after Lay. and Mesnil. C, T. gambiense (blood, T-fever), after Bruce and Nabarro. D, T. equinum, after Lay. and Mesnil. E, Trypanomorpha (Trypanosoma) noctuae, after Schaud. F, Trypanosoma avium, after Lay. and Mesnil. G, Hanna's Trypanosome from Indian pigeons. J , T. ziemanni, after Schaud. T. damonia, after Lay. and Mesnil. c.g, Chromatoid grains; v, vacuole; 1.s, fold or striation. ziemanni described by Schaudinn. The asexual or indifferent type (fig. 3, H) is extremely thread-like, greatly resembling, in fact, a Spirochaete; on the other hand, both male and female individuals have the form of a very wide spindle. In Trypanoplasma and Trypanophis there are two flagella, inserted into the body very close to the anterior end (fig. 4, F ar_d G). One flagellum is entirely free and directed forwards; the other at once turns backwards and is attached to the convex or dorsal side of the body for the greater part of its length. In all other Trypanosomes there is only one flagellum, which is invariably attached to the body in the same manner as the posterior one of biflagellate forms. This flagellum, however, is most probably not to be considered homologous in all cases. (See Woodcock, loc. cit.) In Trypanomorpha (fig. 3, E), which is to be derived from a Herpetomonadine type, the single, anterior flagellum of the ancestral parasite has been drawn backwards along one side of the body and now originates in the posterior half. Hence in this genus the end bearing the free part of the flagellum is the anterior one. The genus Trypanosoma, in which are included at present the great majority of Trypanosomes, is rather to be regarded as derived from a Heteromastigine ancestor, such as Trypanoplasma, by the loss of the anterior flagellum. Hence in this type the single flagellum represents the posteriorly-directed one of Trypanoplasma, and the end at which it becomes free is the hinder end. The point of origin of the flagellum in Trypanosoma is usually near the anterior end, but may vary considerably (cf. figs.) ; and its free portion may be very short or lacking. Along the dorsal side runs the characteristic fin-like expansion of the body, the undulating-membrane, which is the organella principally concerned in locomotion. This always begins at the place where the attached flagellum emerges from the body; and its free edge is really constituted by the latter, which forms a flagellar border. The membrane is usually more or less sinuous in outline, and is sometimes thrown into broad folds (fig. 3, F and J). Distally it thins away concurrently with the body. (After Doflein.) a, Parasites. b, Blood-corpuscles. The body appears to be in all cases naked. A differentiation of the peripheral cytoplasm in the form of an ectoplasmic layer has been described in one or two instances, and it seems Minute probable that in most Trypanosomes there is such a structure. layer, although only poorly developed, as a rule, around the body generally. On the other hand, the undulating-membrane is largely if not entirely an ectoplasmic development. This is usually much clearer and more hyaline than the general cytoplasm. In many forms deep-staining grains or granules, of a chromatoid nature and of varying size, are to be seen in the cytoplasm. In C, T. inopinatum, after Serg. D, T. karyozeukton, after Dutt. and Todd. E, T. netspruitense, after Lay. and Mesnil. F,G, Trypano plasma borreli (living and stained), after Leger. H, T. cyprini, after Plehn. J, Try panosoma soleae, after Lay. and Mesnil. K, T. granulosum, after Lay. and Mesnil. L, T. remaki, var. magna, after Lay. and Mesnil. h, Clear zone or halo around kineto- a.fl, Anterior flagellum; nucleus. p fl, Posterior flagellum; th,Chain of chromatic rodlets run- l.s, Longitudinal striations fling from trophonucleus to nemes); kinetonucleus. v, Cytoplasmic vacuole. most cases these granules are, if not confined to, chiefly distributed in the posterior (flagellate) half of the body (figs. 3, B, D and E, 4, E and G). In certain Trypanosomes a well-defined, usually oval vacuole is often, though not constantly, to be observed, situated at a varying distance from the anterior end (figs. 3 and C, G, 4, F). There is no reason to doubt that this vacuole is a normal cell-constituent, for it has been described in parasites in quite normal surroundings and conditions. A Trypanosome always possesses two distinct nuclear bodies, one the trophonucleus, regulating the trophic life of the cell, the other, the kinetonucleus, directing its locomotor activities. The recent investigations of Schaudinn and Prowazek (II. c) have shown that, in some forms at any rate, the finer structure and detailed development of the nuclear apparatus is extremely complex. Notonly is there an intimate correspondence in this respect between the two principal organellae, but the flagellar apparatus itself is really of nuclear origin and remains closely connected with the kinetonucleus (cf. fig. 7). In most cases, however, little beyond the position and general appearance of the nuclei has been so far made known. The trophonucleus is usually situated somewhere about the middle of the body. The kinetonucleus is typically near the anterior end; but in a few instances it lies more centrally (e.g. T. inopinatum, T. rotaiorium, fig. 4, A–C) ; in Trypanomorpha it is in the posterior half of the body (fig. 3, E). In certain forms the occurrence of prominent myonemes or muscle-fibrillae has been described, and, more-over, a nuclear origin assigned to them also. In Trypanomorpha they are confined to the undulating-membrane (fig. 3, E), but in other cases—Trypanosoma ziemanni, T. lewisi, T. brucei, and T. soleae—they are arranged laterally, half running down each side of the body (fig. 4, J). In Trypanoplasma borreli there is only a single myoneme on either side. All Trypanosomes are capable of binary longitudinal fission, and this appears to be the chief method of multiplication. The division of the nuclear appa- ratus is the first to take place (fig. 5, A). The Muttiplicakinetonucleus more often leads the way, but tion. sometimes either kinetonucleus or trophonucleus may do so indifferently. The duplication of the flagellum begins at its proximal end, that which is in relation with the kinetonucleus. Until recently the process has been considered as an actual longitudinal splitting of the flagellum, following upon the separation of the two daughter-kinetonuclei. Both Schaudinn (in the case of Trypanomorpha) and Prowazek (in the case of Trypanosoma lewisi and T. brucei), have found, however, that the new flagellum is developed quite independently and laid down alongside the old one. It is at present somewhat uncertain, therefore, in what cases actual splitting occurs. The same applies equally to the formation of the undulating-membrane. If the flagellar border splits, the membrane doubtless divides also; but where the flagellum is a new formation the membrane will be too. The division of the cytoplasm in most forms is equal or sub-equal, and two approximately equal daughter-Trypanosomes result (fig. 5, C). In some instances (e.g. T. equinum, T. equiperdum) the longitudinal fission is apparently multiple, three or even four descendants being produced simultaneously. A B (After Lay. and Mesnil.) 4 T. lewisi differs from most Trypanosomes in that the cytoplasm divides in a very unequal manner (fig. 6). The process is more comparable to budding, since the larger or parent-individual may produce, successively, more than one " daughter "; moreover, the daughter-individuals may subdivide before separating, the whole family remaining attached by the non-flagellate (anterior) end (fig. 6, F). In this type of division it may be noted that the kinetonucleus comes to lie alongside the trophonucleus, or even passes to the other side of it (i.e. nearer the flagellar end). Easily derivable from this method is the other one characteristic of T. lewisi, viz. segmentation. The chief difference is that in the latter no parent-individual is distinguishable, a rosette of many equal daughter-parasites being formed. The small Trypanosomes resulting from either of these modes of division differ from typical adults by their stumpy, pyriform shape, the position of the kinetonucleus near the flagellar end of the body, and the absence, during the first part of their youth, of an undulating-membrane. At this period they have, in fact, what may be termed a " pseudo-Herpetomonadine " aspect. These young individuals can themselves multiply by equal binary fission, giving C (myo- rise to little fusiform parasites; with growth, these gradually assume the adult appearance. Comprehensive researches (1905, seq.) have made it evident that Trypanosomes have a much more varied and complex develop- Develop- ment and life-history than was previously supposed. meat and This has now been found to be the case in widely- Lile-cycle. differing parasites, occurring in widely-different hosts. The following examples have been investigated: Try panosoma lewisi (also, but much less completely, T. brucei),' among mammalian forms, described by Prowazek (47) ; T. ziemanni and Trypanomorpha noctuae, among avian parasites, described by Schaudinn (5o) ; Trypanosoma inopinatum, among batrachian forms, described by A. Billet (la and 2), T. barbatulae and Trypano- plasma varium, described by Leger (32 and 33), and T. borreli, by (A-E, after Lay. and Mesnil; F, after Wasiel and Senn.) G. Keysselitz (16), from fishes; also several other piscine Trypanosomes have their development phases in leeches worked on by Brumpt (5a). In addition, a Trypanosome whose vertebrate host is yet unknown (T. grayi) has been studied in detail by Minchin (41a). It is impracticable here to consider fully all the various develop-mental phases and modifications of the life-cycle described as occur-ring in the above parasites. In view, however, of the great interest excited by Schaudinn's work on avian parasites, as well as on account of the far-reaching importance of his conclusions to the study of the Haematozoa, a brief summary of his celebrated research is necessary. According to Schaudinn's account, he was dealing with two separate Trypanosome parasites of the Little Owl (Athene noctua), viz. Trypanomorpha (Trypanosoma) noctuae and Trypanosoma (Spirochaete) ziemanni. The latter organism, in certain phases, very closely resembles a Spirochaete. In the blood of the owl resting, intracellular phases of both parasites alternate with active trypaniform ones; and, when in the former condition, Schaudinn considers that the parasites are identical with what have been formerly regarded as distinct Haemosporidia, Halteridium and a Leucocytozoon respectively. In other words, he considers that these two Haemosporidian forms are really only phases in the life-history of particular Trypanosomes. To this life-cycle belongs the formation of sexual individuals and their conjugation on arrival in the gnat (Culex) ; the process is described as agreeing in the main, in both cases, with what has already been made known by Mac-Callum for another species of Halteridium. The male gametes, it may be noted, are said to possess the essential characters of a Trypanosome. The motile copula or ookinete formed in the gnat gives i T. brucei has also been studied in a Tsetse-fly (G. fusca) by Stuhimann (54a).rise to one of three types of Trypanosome individual: indifferent, male or female. The development of an indifferent ookinete into an indifferent Trypanosome is shown in fig. 7, from which it will be seen that the cytological details are very complex. The indifferent parasites exhibit an alternation of resting, attached phases with active periods, during which they multiply actively and become very abundant in the insect. The male forms, which are very small and the homologues of the microgametes developed in the blood, appear to die off soon. The female Trypanosomes, on the other hand, grow to a large size, laying up a store of reserve nutriment. They are very sluggish and do not divide. They are the most resistant to unfavourable conditions of environment, and are able, by a process of parthenogenesis, to give rise to ordinary, indifferent forms again, which can repopulate the gnat. So far as regards the remarkable connexion between Trypanosomes and Haemosporidia indicated by Schaudinn, this has met with a great deal of criticism on the part of Novy and McNeal among others, and it must be admitted that up to 1909 no definite corroboration can be said to have been brought forward. Again, the spirochaetiform Trypanosoma (T. ziemanni) described may have been really a true Spirochaete, i.e. a Bacterium. In short, it is quite possible Schaudinn did not sufficiently distinguish between the life-cycles of four distinct parasites of the Little Owl: a Trypanosome, a Spirochaete, a Halteridium and a Leucocytozoon; though, on the other hand, this is by no means proved. However this may be, the research of subsequent workers—e.g. Brumpt (5a), Leger (32, 33), Keysselitz (16), Prowazek (47), Minchin (41b) and others—has undoubtedly shown that much of Schaudinn's scheme of the life-history of a Trypanosome is well-founded. It is certain, for instance, that the three types of form which he discovered, viz. indifferent, male or female, can be recognized in many cases, often in the vertebrate, but always more sharply differentiated in the invertebrate. Moreover, it is very probable that conjugation occurs soon after the arrival of the parasites in their specific invertebrate host; and this act may perhaps give rise to an aflagellar copula, which is gregariniform and comparable to an ookinete. Different investigators, it may be noted, have described various (After Schaudinn.) A–D shows the formation of the two nuclear elements (trophonucleus and kinetonucleus) from the definitive nucleus (synkaryon) of the ookinete. E–H shows the formation of the myonemes and the flagellar border (flagellum) of the undulating membrane, by means of a greatly elongated nuclear-spindle. Lehr, Trophonuclear chromo- some. m, k.chr, Kinetonuclear do. f.b, c, Centrosomic granule. a.s, First axial spindle. a,s2, a.s', Second and third do. c.3, t, Trophonucleus. k, Kinetonucleus. k.c, Kinetonuclear centrosome. G. Trophonuclear centrosome. Myonemes. Flagellar border of undulating-membrane (3rd axial spindle). Its (proximal centrosome ts distal one vanishing as such). complicated nuclear changes and divisions undergone by Trypanosomes; these are considered, in many cases, to represent some kind of parthenogenesis. A very interesting modification of the life-cycle of a Trypanosome which must be mentioned has been made known by Minchin, in his account of T. grayi, in a tsetse-fly (G. palpalis). Unfortunately the vertebrate host of this form is not yet known. Certain individuals of a particular character form definite rounded cysts in the rectum of the fly; in this condition, the only sign of Trypanosome structure is afforded by the two nuclei, which remain separate. These cysts are doubtless for dispersal by way of the anus, and the vertebrate host is in all likelihood infected by the mouth and alimentary canal. This reveals a quite novel mode by which infection with a Trypanosome may be brought about; so far, however, T. grayi remains the only known example. As remarked in the section on morphology, the Trypanosomes Ciasstt'tca- as a whole are preferably regarded as including don. two entirely distinct groups, Monadina and Hetero- mastigia. SUB-ORDER MONADINA Family: Trypanomorphidae, Woodcock.—Haemoflagellates de-rived from a uniflagellate, Herpetomonadine form, in which the point of insertion of the single (anterior) flagellum into the body has travelled backwards from the anterior end for a greater or less distance, the flagellum itself having become, concurrently, attached to the body for a portion of its length by means of an undulating membrane. Genus Trypanomorpha, Woodcock, 1906.—With the characters of the family. The only species yet known is the type species, T. noctuae (Celli and San Felice). [Syn. Trypanosoma n. (C. & S.F.), Schaud.=Halteridium n. (C. & S.F.)]. See figs. 3, E, 7. Vertebrate host, Athene noctua, Little Owl; invertebrate host, Culex pipiens. There are, in addition, other forms, which are probably to be placed in this family, but which are not yet sufficiently well known for their systematic position to be settled. It is, for instance, quite likely that certain Herpetomonadine parasites described by Leger (29, 34) from various blood-sucking insects are really only stages in the life of a Haemoflagellate. Some of these are placed by Leger in a newly discovered genus, Crithidia. SUB-ORDER HETEROMASTIGINA Family: Trypanosomatidae, Doflein.—Flagellates, in the great majority of instances haemal parasites, derived from a biflagellate, Bodo-like type, in which the posteriorly-directed (trailing) flagellum is always present and attached to the body by an undulating membrane, of which it constitutes the thickened edge. The other, the anterior flagellum, may or may not persist. Genus Trypanoplasma, Lay. and Mesnil, 1902.—The anterior flagellum is present. Both flagella are inserted close together, near the anterior end of the body. Two sub-groups may be distinguished. In one, exemplified by T. borreli (fig. 4, F and G) from the rudd and minnow, the anterior flagellum is well-developed, and the free parts of both are of about equal length. In the other, exemplified by T. cyprini (fig. 4, H) from carp, the anterior flagellum is much shorter than the free part of the posterior one, and evidently tending to disappear. Known invertebrate hosts for different species are Hemiclepsis and Piscicola, leeches. Genus Trypanophis, Keysselitz, 1904.—The body resembles that of Trypanoplasma in general appearance, but the locomotor apparatus does not appear to be so well-developed, especially in T. grobbeni. The anterior flagellum is longer than the free part of the posterior one. The species included are not, so far as is known, haemal parasites. T. grobbeni occurs in the coelenteric cavity of various Siphonophora. An interesting form, " Trypanoplasma" intestinalis, which resembles both the above genera, occurs in the alimentary canal of Box boops. Probably this is not a haemal parasite, and lacks an alternate host. Genus Trypanosoma, Gruby, 1843.—(Principal synonyms: Undulina, Lank., 1871; Herpetomonas, Kent, 188o, only in part ; Paramoecioides, Grassi, 1881; Haematomonas, Mitrophan, 1883.) There is no anterior flagellum. The point of insertion of the attached (posterior) flagellum into the body, and, consequently, the commencement of the undulating membrane may be almost any-where in the anterior half of the body, but is usually near the extremity. Among the more important and better-known forms are the following: Parasitic in mammals: T. lewisi (Kent), the well-known natural Trypanosome of rats (figs. 3, A, 6, A) ; T. brucii, Plim. and Bradf., the cause of nagana among cattle, horses, &c., in South Africa (fig. 3, B) ; T. evansi, Steel, the cause of surra to horses in Indo-Burmah; T. equiperdum, Dofl., the cause of dourine in horses in Algeria and other regions of the Mediterranean littoral; T. equinum, Voges, causing mal de caderas or " hip-paraplegia " in South America (fig. 3, D) ; T. theileri, Lay., a very large form, the cause ofgalziekte or bile-sickness to cattle in the Transvaal; and T. gambiense, Dutton (syn. T. ugandense, Castellani, T. castellanii, Kruse), the cause of human trypanosomosis in central Africa, which becomes sleeping-sickness when the organisms penetrate into the cerebro-spinal fluid (fig. 3, C). Parasitic in birds: T. avium (Danil., Lay. emend.), probably the form to which Danilewsky's original investigations related, parasitic in owls and (according to Novy and McNeal) also in other birds (fig. 3, F) ; T. johnstoni, Dutt. and Todd, a very spirochaetiform type, from little birds (Estrelda) in Senegambia; and Hanna's peculiar wide species from Indian birds, with a remarkably tapering anterior end (fig. 3, G). Lastly, there is T. ziemanni, Lay., [syn. Spirochaete z. (Lay.), Schaud, " Haemamoeba" z., Lay., the " Leucocytozoon " of Danil.], from various owls, and Culex pipiens, whose life-history has been described by Schaudinn (fig. 3, H). (As above mentioned, this form may not be a true Trypanosome.) Only one reptilian form is well known, T. damoniae, Lay. and Mesn., from a tortoise, Damonia reevesii (fig. 3, J). Parasitic in batrachia: T. rotatorium, Mayer (syn. Amoeba r., Mayer, July 1843, T. sanguinis, Gruby, November 1843, Undulina ranarum, Lank., 1871), the best-known parasite of frogs, which exhibits remarkable polymorphism (fig. 4, A and B) ; T. mega and T. karyozeukton, Dutt. and Todd, even larger than T. r. (fig. 4, D), with peculiar cytological differentiation, may be only sub-species; T. inopinatum, Sergent, and T. nelspruitense, Lay., also from frogs (fig. 4, C). Parasitic in fishes: T. remaki, Lay. and Mesnil, from pike, a relatively small form (fig. 4, L); T. barbatulae, Leger, from loach ; T. granulosum, Lay. and Mesnil, a very long vermiform parasite, from eels (fig. 4, K) ; T. soleae, Lay. and Mesnil, from soles, with a relatively small flagellum (fig. 4, J); and T. scyllii and T. rajae, from those Elasmobranchs, both very large forms, described by Lay. and Mesnil. Undoubtedly closely allied to the Haemoflagellates, although no actual trypaniform phase has yet been observed, are the important parasites usually known as the " Leish- The man-Donovan " bodies, without some consideration Leishmanof which an account of the Haemoflagellates would Donovan-hardly be complete. These bodies are constantly Wright found in certain tropical fevers (e.g. dum-dum fever, Bodies. kala-azar) particularly prevalent throughout Indo-Burma, of which they are generally held to be the cause. They were discovered by W. Leishman in 1900, but before his first account of them (36) was published they were also seen quite independently by C. Donovan. Moreover, organisms very similar to these (morphologically, indeed, the two sorts appear scarcely distinguishable) are found in various sores or ulcers (e.g. Delhi boil, Oriental sore, " bouton d'Alep ") to which people in different parts of the East are liable. These were first described by J. H. Wright (58). The chief distinction between the parasites in the two cases is in their habitat. In the one case they are entirely restricted to the neighbourhood of the boil or ulcer, whereas in the other there is a general infection of the body, the organisms spreading to all parts and being met with in the spleen, liver, bone-marrow, &c., and (rarely) in the peripheral circulation. The parasites are either free or intracellular. In the latter case they invade cells of a leucocytic or phagocytic character as a rule; Leishman's form is particularly abundant in large macrophageal cells originating from the vascular endothelium of the spleen (fig. 8, I. M). The parasites themselves are very minute and usually ovoid or pyriform in shape (fig. 8, I. a), the latter being, perhaps, the most typical. The splenic type is somewhat smaller than Wright's parasite; the former, when pear-shaped, is from 31 to 4 A in length by 11 to 2 µ in width, the latter being about 4 µ by 3 A (fig. 8, III.). The body is probably not limited by any distinct membrane. The cytoplasm is finely granular and fairly uniform in character. The most interesting point about the morphology is the fact that two chromatic bodies, of very unequal size, are almost invariably to be recognized. The larger nuclear body, which corresponds to the trophonucleus of a Trypanosome, is usually round or oval ; the smaller one, representing a kinetonucleus, has the form either of a little rod or of a round grain, and is generally separate from the larger nucleus. The parasites multiply in two ways—(a) by binary fission, and (b) by multiple division or segmentation. The principal stages in the first method are well known (fig. 8, I. b) ; they offer strong resemblance to the process in Piro plasma. Multiple division has not yet been so satisfactorily made out. It appears to con-form more or less to the radial or rosette type of multiplication, enlarged rounded parasites, with a varying number of nuclei (up to about eight) uniformly arranged near the periphery, having been often noticed (fig. 8, I. c and IV. b). The details of the process are somewhat differently described, however, by different observers. Laveran and Mesnil (27) gave the name Piro plasma donovani to Leishman's form; and there is no doubt that the parasites are closely allied to that type of organism. This does not, however, preclude in any way the supposition that they—equally with certain other Haemosporidia—represent, nevertheless, only a phase of a complete life-cycle; and this supposition has in fact been definitely proved to be true by the work of Rogers (48). Rogers cultivated the parasites obtained from cases of kala-azar in artificial media, and found that what were unmistakably flagellate a 0 (D. 'T> 4 I. I. Piroplasma (Leishmania) donovani, Lay. and Mesnil. a, Typical pear-shaped or oval forms; b, various stages in longitudinal division ; c, nuclear division preparatory to multiple fission; d, endoglobular forms, in red blood-corpuscles (p= pigment grains) ; e, bacillary form of the parasite in a corpuscle; M, large macrophageal cell with many parasites (after Donovan). II. Uninuclear leucocyte (L) containing several parasites (after Lay. and Mesnil). a, Single individuals; b, dividing forms (from Mesnil, mostly after Wright). IV. P. donovani in cultures of different ages. a, Ordinary forms of varying sizes; b c, stages in multiple division; d, binary fission; e, f, g, flagellate forms (after Rogers). stages developed in the cultures at different intervals (fig. 8, IV. e, f, g). These forms were elongated and spindle-like; and to one end of the body, near which the smaller nuclear element was situated, a well-developed flagellum was attached. Since then many other workers have obtained similar stages [see Leishman and Statham (38), Christophers (7)] ; but however slender and Trypanosome-like the flagelliform parasites may appear, up till now no indications of an undulating membrane have been seen, and the kinetonuclear element is never far from the insertion of the flagellum. Nevertheless, the general appearance and structure of these motile forms so greatly resemble that of a Herpetomonad, or of the " pseudo-Herpetomonadine " forms of a Trypanosome which are obtained in cultures, that it cannot be doubted that the " Leishman-Donovan-Wright " bodies are closely connected with the Haemoflagellates. That being so, it is quite possible that, in normal conditions and circumstances, these parasites also possess, at some period of the life-cycle, a trypaniform phase. Nothing definite is yet known with regard to the transmission of the parasites by an alternate invertebrate host, although there is presumptive evidence in favour of this supposition? A word or two must be said in conclusion with reference to Supposed the supposed connexion of the Spirochaetae with the Connexion Trypanosomes. In Schaudinn's great memoir he otthespiro- regarded Trypanosoma ziemanni as possessing, in chaetae with certain phases, the actual characteristics of a Trypano- Spirochaete as then known; and, further, he was comes. inclined to think that other Spirochaetae (e.g. S. obermeieri of relapsing fever) were also only phases in the i R. Ross (49), regarding the parasites as a quite different kind of Sporozoan, termed them Leishmania; and Wright named his variety from tropical ulcers Helcosoma tropicum. s Patton (Sci. Mem. India, No. 27, 1907) has brought forward evidence to show that the bed-bug (Cimex macrocephalus) is the invertebrate of a particular Haemoflagellate. As a result of his more recent investigations on S. plicatilis (the type-species of Ehrenberg) and other forms (51), he finds, however, that this is not the case, but that the organisms exemplified by S. plicatilis are to be widely separated from the Trypanosomes, and placed rather with the Bacteria. In addition, it is most probable that, at any rate, certain other spirilliform parasites, e.g. S. balbianii, S. refringens, agree fundamentally in structure with the type-species. On the other hand, evidence has lately been brought forward to show that certain parasites which greatly resemble a Spirochaete are really related to the Trypanosomes. This is the case with the celebrated organism first described by Schaudinn and E. Hoffmann (52) from essential syphilitic lesions, and now known as Treponema (Spirochaete) pallida, Schaud. F. Krzysztalowicz and M. Siedlecki have published an important account (17) of this parasite, which they consider possesses a true trypaniform phase, and for which they have proposed the name Trypanosoma leis. This view requires, however, corroboration. Nevertheless the resemblance between the biology of this organism in relation to syphilis (as regards mode of infection, habitat, &c.) and that of Trypanosoma equiperdu'm, the cause of dourine or " horse-syphilis," may not be without significance. `QD O ova % donovani, Lay. et Mesn.) parasite d'une fievre de 1'Inde, " C. r. ca. sci. (1903), 137, p. 957, figs, ; (28) idem, " Sur la nature bacterienne du pretendu trypanosome des huitres, T. balbianii, " C. r. soc. biol. (1901), 53, p. 883; (there are numerous other papers by these tuthors in the C. r. ac. sci. and the C. r. soc. biol. from 1900 inwards) ; (29) L. Leger, " Sur un flagelle parasite de l'Anopheles maculipennis," C. r. soc. biol. (1902), 54, p. 354, figs. ; (30) idem, " Sur la morphologie du trypanoplasma des vairons," C. r. ac. sci. (1904), 138, p. 824; (31) idem, " Sur la structure et les affinites des trypanoplasmes " (1904), t. C. p. 856, figs. ; (32) idem, " Sur les hemoflagelles du Cobitis barbatula, L. I. Trypanosoma barbatulae, n. sp.," C. r. soc. biol. (1904), 57, p. 344; (33) idem, " Trypanoplasma varium, n. sp., parasite du sang de Cobitis barbatula, L." (1904), t. C. p. 345; (34) idem, " Sur les affinites de l'Herpetomonas subulata et la phylogenie des trypanosomes " (1904), t. C. p. 615; (35) idem, " Sur la presence d'un trypanoplasma intestinal chez les poissons," op. cit. (1905), 58, p. 511; (36) W. Leishman, " On the possibility of the occurrence of trypanosomiasis in India," Brit. Med. Journ, (1903), i. 1252, figs.; (37) idem, " Note on the nature of the parasitic bodies found in tropical splenomegaly," op. cit. (1904), i. 303; (38) Leishman and Statham, " The development of the Leishman body in cultivation," Journ. Army Med. Corps (1905), 3, 14 pp., I pl.; (39) J. Lignieres, " Contribution a 1'etude de la trypanosomose des equi es sud-americains connue sous le nom de Mal de Caderas," Rec. med. vet. (8) (1903), 10, p. 51, 2 pls.; (40) A. F. Mayer, " Spicilegium observationum anatomicarum de organo electrico in raiis anelectrias et de haematozois," (Bonn, 1873), 18, pp., pls.; (41) F. Mesnil, F. Nicolle and P. Remlinger, " Sur le protozoaire du bouton d'Alep," C. r. soc. biol. (1904), 57, p. 167; (41a) E. A. Minchin, " On the occurrence of encystation in Trypanosoma grays," &c., Proc. Roy. Soc. (1907), 79 B, p. 35.; (41b) idem (with Gray and Tulloch), Glossina palpalis in relation to Trypanosoma gambiense," &c., op. cit. (1906), 78 B, p. 242, 3 pls.; (42) Mitrophanow, " Beitrage zur Kenntniss der Hamatozen," Biol. Centbl. (1883), 3. p. 35, figs.; (43) G. Nepveu, " Sur un trypanosome clans le sang de 1'homme," C. r. soc. biol. (1898), 50, p. 1172; (44) F. G. Novy and W. J. McNeal," OntheTrypanosomes of Birds," Journ. Inf. Dis. (1905), 2, p. 256, pls.; (45) W. S. Perrin, " The life-history of Trypanosoma balbianii;" Proc. Roy. Soc. (1905), 76 B, p. 367, figs., also in Arch. Protistenk. (1906), 7 pls.; (46) M. Plehn, " Trypanoplasma cyprini, n. sp.," Arch. Protistenk. (1903), 2, p. 175, I pl. ; (47) S. Prowazek, " Studien fiber Saugethiertrypanosomen," Arb. kais. Gesundheitsamte (1905), 22, 44 pp., pls. ; (48) L. Rogers, " On the development of flagellated organisms (Trypanosomes) from the spleen Protozoic parasites of cachexial fevers and Kala-azar," Quart. Journ. Mic. Sci. (1904), 48, p. 367, 1 pl. ; 149) R. Ross, " Notes on the bodies recently described by Leishman and Donovan, " Brit. Med. Journ. (1903), 1. 1261, 1401, figs. ; (50) F. Schaudinn, " Generations- and Wirthswechsel bei Trypanosoma and Spirochaete," Arb. kais. Gesundheitsamte (1904), 20, p. 387, figs.; (51) idem, " Zur Kenntniss der Spirochaete pallida," Deutsch. med. Wochenschr. (1905), No. 42, p. 1665; (52) Schaudinn and E. Hoffmann, " Vorlaufiger Bericht fiber das Vorkommen von Spirochaeten in syphilitischen Krankheitsproducten," Arb. kais. Gesundheitsamte (1905), 22, p. 527; (53) E. and E.'Sergent, " Sur un trypanosome nouveau parasite de la grenouille verte," C. r. soc. biol. (1904), 56, p. 123, fig. ; (54) idem, " Hemamibes des oiseaux et moustiques ' Generations alternantes,' de Schaudinn,'" op. cit. (1905), 58, p. 57; (54a) F. Stuhlmann, " Beitrage zur Kenntniss der Tsetsefliege,"&c., Arb. kais. Gesundheitsamte (1907), 26, p. 83, pis; (55) Valentin, " Ober ein Entozoon im Blute von Salmo f ario, " Muller's Arch., 41, p• 435; (56) O. Voges, " Mal de Caderas," Zeitschr. Hyg. (1902), 39, p. 323, 1 pl. ; (57) Wasielewsky and G. Senn, " Beitrage zur Kenntniss der Flagellaten des Rattenblutes," op. Cit. (1900), 33, p. 444, pls. (58); J. H. Wright, "Protozoa in a case of tropical ulcer (Delhi sore), " Journ. Med. Research, Boston (1903), 10, p. 472, pls. (H. M. Wo.)
WILLIAM TRYON (1729–1788)

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