coccotdes. · Geographic distribution Peru, Colombia, Ecuador Ubiquitous Ubiquitous Susceptible...
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BARTONELLACEAE' D. PETERS AND R. WIGAND Virusabteilung, Bernhard Nocht-Institut ffir Schiffs- und Tropenkrankheiten, Hamburg, Germany Bartonellaceae (1) form a group of blood para- In electron microscopy, the contrast between sites characterized by their smallness and their B. bacilliformis and the other two species is position close to red blood cells. Bartonella bacil- striking. Cultured B. bacilliformis shows re- liformis (Strong, Tyzzer, Sellards), causing human tracted cytoplasm and cell walls typical of bac- bartonellosis (Oroya fever and verruga peruana) teria (figure 4). Also like bacteria, they are rod- in Peru, Colombia and Ecuador, grows on culture shaped in young cultures and mostly coccoid in media and was early recognized by Noguchi (2) older ones (5). For comparison with animal as a bacterium. Other bartonellae found all over bartonellae, however, B. bacilliformis had to be the world as animal parasites that cause a latent studied as a blood parasite. Also, as such it is infection activated by splenectomy cannot be mostly rod-shaped with cell walls often visible cultured with certainty on artificial media and [figure 5, (4)]. Annular and coccoid particles still withstand definite classification. Weinman which are seen also in light microscopy (3) must (3) in his comprehensive monograph, although be considered degenerate forms, in contrast with listing all bartonellae as bacteria, contrasts the . . S biologic traits of B. bacilliformis with those of the ring forms of E. coccotdes. Such structures animal bartonellae. In his article with Bengtson preferably occur during the final stages of in- in Bergey's Manual of Determinative Bacteriology fection with B. bacilliformis under antibiotic (1) he lists three families within the order of treatmentand in old cultures. Rickettsiales: Rickettsiaceae, Bartonellaceae, H muris shows coccoid particles in electron and Chlamydozoaceae; the family of Bartonel- microscopy. The rod forms of H. muris, as seen laceae comprises four genera: Bartonella, with the in light microscopy, proved to be chains of coccoid only species B. bacillformis; Haemobartonella, or particles in electron optics (figure 7). Similar animal bartonellae; Eperythrozoon; and Gra- findings are valid for Haemobartonella muris hamella. musculi (Schilling) (11), whose light-optical Since light microscopy is not adequate for the aspect is marked by rod forms. No structural minute structural details of these bodies, ranging details or cell walls like those of bacteria could be between the size of rickettsiae and that of large discerned in H. muris [(figures 7, 8 (6)]2. After viruses, electron microscopy was used for B. hemolysis, the identification of H. muris is often bacilliformis of human blood (4) and cultures (5), rendered difficult by the residues of reticulocytes Haemobartonella muris (Mayer) of rat blood (6, present in the anemic blood. Such reticulo- 7, 8), and Eperythrozoon coccoides (Schilling, filamentous particles, however, are translucent Dinger) of mouse blood (9, 10). The results of this and less uniform in size (13), resembling the work and those of other authors on the system- . X atics are summarized in this paper. The main vesicular structures recently described as "endo- features of the three parasites are shown in plasmoi reticulum" (14). table 1. Also in E. coccoides, cell walls and inner struc- Light microscopy shows that all bartonellae ture could not be proved (9). Although ring shapes readily take the Giemsa stain. Rod shapes prevail predominated (figure 9), signet-ring, racket and in B. bacilliformis (figure 1), whereas H. muris comma shapes were also seen. This polymorphism has more coccoid forms (figure 2) and E. coccoides is probably due to injuries of the fragile bodies has more ring-shaped bodies (figure 3). Such during the act of mounting and does not suggest rings probably arise from the process of air- developmental stages (10). drying; wet preparations seen in phase contrast The action of crystallized enzymes on the microscopy suggest that E. coccoides in the circu- previously fixed microorganisms has been studied lating blood has a coccoid or vesicular form. in light and electron microscopy. After trypsin Annular forms rarely occur in H. muris either 2 Eyer and Ruska (12), in a paper on the mor- (8). -phology of typhus rickettsiae, pointed out that 1 We use the term for brevity with the reserva- Haemobartonella muris does not show a cell wall tions as to classification made in the text. like that of Rickettsia prowazeki. 150 on June 29, 2020 by guest http://mmbr.asm.org/ Downloaded from
Transcript of coccotdes. · Geographic distribution Peru, Colombia, Ecuador Ubiquitous Ubiquitous Susceptible...
BARTONELLACEAE'
D. PETERS AND R. WIGAND
Virusabteilung, Bernhard Nocht-Institut ffir Schiffs- und Tropenkrankheiten, Hamburg, Germany
Bartonellaceae (1) form a group of blood para- In electron microscopy, the contrast betweensites characterized by their smallness and their B. bacilliformis and the other two species isposition close to red blood cells. Bartonella bacil- striking. Cultured B. bacilliformis shows re-liformis (Strong, Tyzzer, Sellards), causing human tracted cytoplasm and cell walls typical of bac-bartonellosis (Oroya fever and verruga peruana) teria (figure 4). Also like bacteria, they are rod-in Peru, Colombia and Ecuador, grows on culture shaped in young cultures and mostly coccoid inmedia and was early recognized by Noguchi (2) older ones (5). For comparison with animalas a bacterium. Other bartonellae found all over bartonellae, however, B. bacilliformis had to bethe world as animal parasites that cause a latent studied as a blood parasite. Also, as such it isinfection activated by splenectomy cannot be mostly rod-shaped with cell walls often visiblecultured with certainty on artificial media and [figure 5, (4)]. Annular and coccoid particlesstill withstand definite classification. Weinman which are seen also in light microscopy (3) must(3) in his comprehensive monograph, although be considered degenerate forms, in contrast withlisting all bartonellae as bacteria, contrasts the . . Sbiologic traits of B. bacilliformis with those of the ring forms of E. coccotdes. Such structuresanimal bartonellae. In his article with Bengtson preferably occur during the final stages of in-in Bergey's Manual of Determinative Bacteriology fection with B. bacilliformis under antibiotic(1) he lists three families within the order of treatmentand in old cultures.Rickettsiales: Rickettsiaceae, Bartonellaceae, H muris shows coccoid particles in electronand Chlamydozoaceae; the family of Bartonel- microscopy. The rod forms of H. muris, as seenlaceae comprises four genera: Bartonella, with the in light microscopy, proved to be chains of coccoidonly species B. bacillformis; Haemobartonella, or particles in electron optics (figure 7). Similaranimal bartonellae; Eperythrozoon; and Gra- findings are valid for Haemobartonella murishamella. musculi (Schilling) (11), whose light-optical
Since light microscopy is not adequate for the aspect is marked by rod forms. No structuralminute structural details of these bodies, ranging details or cell walls like those of bacteria could bebetween the size of rickettsiae and that of large discerned in H. muris [(figures 7, 8 (6)]2. Afterviruses, electron microscopy was used for B. hemolysis, the identification of H. muris is oftenbacilliformis of human blood (4) and cultures (5), rendered difficult by the residues of reticulocytesHaemobartonella muris (Mayer) of rat blood (6, present in the anemic blood. Such reticulo-7, 8), and Eperythrozoon coccoides (Schilling, filamentous particles, however, are translucentDinger) of mouse blood (9, 10). The results of this and less uniform in size (13), resembling thework and those of other authors on the system- . Xatics are summarized in this paper. The main vesicular structures recently described as "endo-features of the three parasites are shown in plasmoi reticulum" (14).table 1. Also in E. coccoides, cell walls and inner struc-
Light microscopy shows that all bartonellae ture could not be proved (9). Although ring shapesreadily take the Giemsa stain. Rod shapes prevail predominated (figure 9), signet-ring, racket andin B. bacilliformis (figure 1), whereas H. muris comma shapes were also seen. This polymorphismhas more coccoid forms (figure 2) and E. coccoides is probably due to injuries of the fragile bodieshas more ring-shaped bodies (figure 3). Such during the act of mounting and does not suggestrings probably arise from the process of air- developmental stages (10).drying; wet preparations seen in phase contrast The action of crystallized enzymes on themicroscopy suggest that E. coccoides in the circu- previously fixed microorganisms has been studiedlating blood has a coccoid or vesicular form. in light and electron microscopy. After trypsinAnnular forms rarely occur in H. muris either 2 Eyer and Ruska (12), in a paper on the mor-(8).-phology of typhus rickettsiae, pointed out that
1 We use the term for brevity with the reserva- Haemobartonella muris does not show a cell walltions as to classification made in the text. like that of Rickettsia prowazeki.
150
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TABLE 1Characteristics of the "Bartonellaceae"
Bartonella bacilliformis Haemobartonella muris Eperythrozooncoccoides
Giemsa-staining Purple Red-purple RedGram-stainingMorphology:Shape Rods Coccoid Annular or coccoidCell walls +Flagella +
Motility + in blood and cultures -
Size 0.4 to 1 AX 0.6 to 2,u 0.3 to 0.5is 0.3 to 0.8,uPosition Eperythrocytic (also in en- Eperythrocytic Eperythrocytic
dothelial cells)Filtrability +Multiplication Binary fission ? ?Growth on culture media +Activation by splenectomy - + +Insect transmission Sandflies Lice LiceGeographic distribution Peru, Colombia, Ecuador Ubiquitous UbiquitousSusceptible animals Monkey Rat, mouse, ham- Rat, mouse, ham-
ster, rabbit ster, rabbitSensitivity to drugs:Arsenic compounds + +Sulfa compounds
in vitro in vivo
Penicillin + +-Streptomycin + + _Chloramphenicol + + _Aureomycin + Not tested + +Terramycin + Not tested + +
treatment, B. bacilliformis shows distinct cell lacked motility in dark-field examinations andwalls [figure 6, (4)]; note that nearly the whole phase-contrast microscopy. Reports of otherplasmatic substance appears to have oozed out, authors (20, 21) agree that active movements areas happens with other gram-negative bacteria not to be seen in these parasites; flagella are(15, 16). With H. muris, trypsin leaves only an absent.ill-defined rest, not to be interpreted as a cell The favorite position of all three microorgan-wall, while E. coccoides is entirely dissolved by isms is on the surface of red blood cells (eperythro-trypsin (17). cytic). Disputes in this respect could be definitelyH. muris and E. coccoides contain ribo- and settled by the pseudo-replica method (6), al-
deoxyribonucleoproteins which are decomposed though the possibility of some parasites enteringby the combined action of nucleases and pepsin the red cell cannot be wholly excluded.(17). An arrangement of nucleoproteins in sepa- B. bacilliformis is found in human bartonellosisrate areas could not be observed. not only as an eperythrocytic parasite, but alsoThe motility of B. baciUiformis seen in the within the endothelial cells. H. muris and E.
blood (18) and in cultures (5, 19) was found to coccoides, in contrast, are strictly limited to thebe due to unipolar flagella. They regularly showed blood. For this reason, Tyzzer and Weinman (22)up in culture material under both types of mi- have separated animal bartonellae from B. bacil-croscopes (figure 4), whereas in blood material, so liformis and given them the generic name offar, the presence of flagella was not demonstrated. Haemobartonella.Diameter and arrangement of flagella resemble B. baciUiformis and H. muris do not passthose found in bacteria. H. muris and E. coccoides filters retaining bacteria; the opposite is reported
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152 PETERS AND WIGAND [VOL. 19
for E. coccoides (23) as well as for E. parvum and tainty to monkeys only; cultured B. bacilliformisE. suis (24). The filtrability of eperythrozoa may and tissue taken from verruga peruana are morebe explained by variation in size and by structural virulent for monkeys than blood bartonellaeplasticity. H. muris, however, although of similar which merely produce a latent infection. Labora-size, is always attached to erythrocytes or their tory rodents such as mice, rats, and Syrianresidues (8). E. coccoides, however, is also found hamsters (8, 10) are readily infected by bloodfree in the plasma (figure 3). This difference in containing H. muris or E. coccoides.the behavior is a further explanation for the As to chemotherapy, it is known that organicfiltrability of this organism. arsenical substances, such as neoarsphenamine,
B. bacilliformis readily grows on culture media and also arsenic-antimony compounds do not actas well as on chick embryos (25) and tissue in infections with B. bacilliformis (3), but have acultures of the Maitland type (26); H. muris and marked influence on H. muris as well as on E.E. coccoides never multiply with certainty outside coccoides, and may effect complete disappearancethe host blood (7). All authors agree that B. of parasites. All three parasites are refractory tobacilliformis multiplies by binary fission. Nothing sulfa compounds (30, 31, 32, 33). The anti-definite is to be said about H. muris and E. coc- biotics cited in table 1 act bacteriostatically oncoides in this respect. B. bacilliformis in vitro (31). Penicillin (34),Both B. bacilliformis and H. muris cause streptomycin (35), and chloramphenicol (36)
anemia of high degrees, though the pathogenic show a curative effect in B. bacilliformis infec-action and accompanying symptoms differ (3). tions. To date, no reports have been made aboutAnemia due to E. coccoides is an exception (27). the clinical action of aureomycin and terra-Skin eruptions, such as verruga peruana caused mycin. H. muris and E. coccoides are resistantby B. bacilliformis, are never seen in animal to penicillin and streptomycin (33, 37), but notbartonelloses. The spleen seems to exert little, if to aureomycin (7, 33, 38) and terramycin (33, 39).any, influence on the course of B. bacilliformis Chloramphenicol has little, if any, effect. Theinfection in men and monkeys. It plays an obvious action of chemotherapeutic agents is remarkablypart in all animal bartonelloses and eperythro- uniform against H. muris and E. coccoides, inzoonoses, for only after splenectomy does the contrast with their action against B. bacil-animal blood become infested with parasites in liformis.fair number.
B. bacilliformis infection in men and monkeys DISCUSSION
leads to a true immunity subject to differences in As has been outlined, the morphology anddegree and duration. H. muris and E. coccoides biology of Bartonella bacilliformis differs notablyinfections, as known so far, bring about a state of from that of Haemobartonella muris and ofpremunition only. An affinity of B. bacilliformis Eperythrozoon coccoides. B. bacilliformis has char-and H. muris to rickettsiae could not be proved acteristics typical of bacteria, including: size andby serological tests. Complement-fixation re- form, growth on culture media, propagation byactions of bartonella and haemobartonella car- binary fission, flagella, cell walls, and behavior inriers were always negative with rickettsial serological tests. Contrary to the opinion ofantigen, and the Weil-Felix reaction showed only Lwoff (40) no just criteria exist to classify H.occasional positive titers of doubtful specificity muris and E. coccoides among bacteria. Haemo-(7, 28). bartonellae and eperythrozoa can be set apart fromB. bacilliformis is transmitted by sandflies, H. protozoa on account of their small size and lack
muris and E. coccoides by lice. Since lice become of cellular structure. The pleuropneumonia-likeinfected experimentally with B. bacilliformis only organisms (PPLO) resemble haemobartonellaeby the intracelomic injection (29), the human and eperythrozoa in their coccoid and annularlouse most probably will not act as a vector of B. shapes visible with both types of microscopes (41,bacilliformis. The strictly regional occurrence of 42, 43). They differ, however, in that they can beverruga peruana and Oroya fever may be due to grown on culture media.limited habitats of the transmitting sandfly. No A relationship to rickettsiae often supposedsuch restrictions exist for H. muris and E. coc- because of analogies in size and transmission bycoides infections, which are found everywhere. insects may be excluded. In fact, rickettsiae
B. bacilliformis can be transmitted with cer- definitely differ from B. bacilliformis, which is
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flagellated and can be cultivated on artificial 2. NOGUCHI, H. 1927 The etiology of verrugamedia, and also, in structural details, from H. peruana. J. Exptl. Med., 45, 175-189.muris and E. coccoides. Serological relations are 3- WEINMAN, D. 1944 Infectious anemias duelacking and, moreover, multiplication of to bartonella and related cell parasites.bartonellae in the insect vector is questionable. Trans. Am. Phil. Soc., 33, 243-350.
4. WIGAND, R., PETERS, D., AND URTEAGA, 0.
Troansmissio bycteri dvinuset s too ormmcriterion 1953 Neue Untersuchungen uber Bartonellaprotozoa, bacteria and viruses to form a criterion bacilliformis. 4. Mitteilung: Elektronenop-for taxonomy. tische Darstellung aus dem Blut. Z.The electron-optical findings of de Robertis Tropenmed. u. Parasitol., 4, 539-548.
and Epstein (44) in Anaplasma margrnale sug- 5. PETERS, D., AND WIGAND, R. 1952 Neuegesting a likeness to H. muris are not sufficient Untersuchungen uber Bartonella bacilli-to link haemobartonellae to the anaplasma group formis. 1. Mitteilung: Morphologie der Kul-as proposed in older publications (45). Relations turform. Z. Tropenmed. u. Parasitol., 3,to grahamellae, which can be cultivated on 313-326.artificial media, are likewise questionable. 6. NAUCK, E. G., PETERS, D., AND WIGAND, R.
It is tempting to compare H. muris and E. 1950 Elektronenoptische Untersuchung der* Bartonelia muris Mayer. Z. Naturforsch.,
coccoides with viruses on account of their small 5b, 259-264.size, which allows E. coccoides to pass filters that . WIGAND, R., AND PETERS, D. 1950 Neuereretain bacteria, and because of their inability to Untersuchungen uber Bartonella murisgrow on culture media. Yet viruses of the psit- Mayer. 1. Mitteilung. Z. Tropenmed. u.tacosis (46) and pox group (47, 48), approaching Parasitol., 2, 206-220.haemobartonellae in size, are structurally differ- 8. WIGAND, R., AND PETERS, D. 1952 Neuereent. They possess cell walls that are missing in Untersuchungen uber Bartonella murisH. muris and E. coccoides, and, moreover, nothing Mayer. 2. Mitteilung. Z. Tropenmed. u.is known about an intracellular multiplication, Parasitol., 3, 437-452.
9. PETERS, D., AND WIGAND, R. 1951 Zur
At present it seems advisable to reserve a Morphologie und Klassifzierung vonAtpresenit seems advisable to reservea Eperythrozoon coccoides. Z. Naturforsch.,special place within the system of microorganisms 6b, 326-333.for haemobartonellae and eperythrozoa and to 10. WIGAND, R., AND PETERS, D. 1952 Blut-set animal bartonellae even farther apart flrom parasiten der weissen Maus. I. Studien tiberB. bacilliformis than has been customary up to Eperythrozoon coccoides Schilling, Dinger.now. Although B. bacilliformis must keep its Z. Tropenmed. u. Parasitol., 3, 461-472.place among bacteria, H. muris and E. coccoides 11. WIGAND, R., AND PETERS, D. 1952 Blut-should be excluded. It is to be presumed also that parasiten der weissen Maus. II. Zur Kennt-other species of haemobartonellae and epery- nis der Bartonella muris musculi Schilling.throzoa resemble H. muris and E. coccoides as
Z. Tropenmed. u. Parasitol., 4, 1-10.
dehrozoainthisresemblew.min .o ea 12. EYER, H., AND RUSKA, H. 1944 tVber dendSicred mthischarateview.cs of H. and E.
Feinbau der Fleckfieberrickettsie. Z. Hyg.Since the characteristics of H. muris and E. Infektionskrankh., 125, 483-492.
coccoides reported in table 1 correspond in nearly 13. PETERS, D., AND WIGAND, R. 1950 Elektro-every respect, it must be questioned whether two nenmikroskopische Untersuchung der Struk-generic names are justified by trifling differences tur hamolysierter Reticulocyten. Klin.in morphology. Wochschr., 28, 649-653.
14. PALADE, G. E., AND PORTER, K. R. 1954ACKNOWLEDGMENTS Studies on the endoplasmic reticulum. I. Its
Research work was aided by funds of Deutsche identification in cells in situ. J. Exptl.Forschungsgemeinschaft. We are indebted to Dr. Med., 100, 641-56.W. Cordes, who kindly translated this paper. 15. PETERS, D., AND WIGAND, R. 1953 En-
zymatisch-elektronenoptische Analyse derREFERENCES Nucleinsaureverteilung, dargestellt an Es-
1. BREED, R. S., MURRAY, E. G. D., AND cherichia coli als Modell. Z. Natur-HITCHENS, A. P. 1948 Bergey's manual of forsch., 8b, 180-192.determinative bacteriology, Suppl. I, pp. 16. WIGAND, R., AND PETERS, D. 1954 Licht-1083-1123, 6th ed. The Williams & Wilkins und elektronenoptische UntersuchungenCompany, Baltimore, Md. uber den Abbau gramnegativer Kokken mit
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Nucleasen und Proteasen. Z. Naturforsch., 31. WIGAND, R. 1952 Neue Untersuchungen9b, 586-596. uber Bartonella bacilliformis. 2. Mitteilung:
17. WIGAND, R., AND PETERS, D. 1954 Abbau- Verhalten gegenuber Sulfonamiden undversuche an Haemobartonella muris und Antibiotica in vitro. Z. Tropenmed. u.Eperythrozoon coccoides. Z. Tropenmed. u. Parasitol., 3, 453-460.Parasitol., 5, 482-492. 32. EMERY, F. E. 1940 Treatment of Bartonella
18. STRONG, R., TYZZER, E. E., BRUES, C. T., muris infections with sulfanilamide. Proc.SELLARDS, A. W., AND GASTIABURU, J. C. Soc. Exptl. Biol. Med., 44, 56-57.1915 Report of first expedition to South 33. THURSTON, J. P. 1953 The chemotherapy ofAmerica, 1913. Harvard University Press, Eperythrozoon coccoides (Schilling, 1928).Cambridge, Mass. Parasitology, 43, 170-174.
19. NOGUCHI, H., AND BATTISTINI, T. S. 1926 34. MERINO, C. 1945 Penicillin therapy inCultivation of Bartonella bacilliformis. human bartonellosis (Carrion's disease).J. Exptl. Med., 43, 851-864. J. Lab. Clin. Med., 30, 1021-1026.
20. MAYER, M., BORCHARDT, W., AND KIKUTH, W. 35. ALDANA, G. L., GASTELUMENDI, R., AND1927 Die durch Milzexstirpation ausl6sbare DIEGUEZ, J. 1948 La estreptomicina en lainfektiose Rattenanamie. Arch. Schiffs- u. enfermedad de Carrion. Arch. peruanosTropen-Hyg. (Beihefte), 31, 295-317. patol. y clin. Lima, 2, 323-338.
21. DINGER, J. E. 1929 Neues uber das 36. PAYNE, E. H., AND URTEAGA, 0. 1951Eperythrozoon coccoides. Zentr. Bakteriol. Carrion's disease treated with chloro-Parasitenk. Abt. I, Orig., 113, 503-509. mycetin. Antibiotics and Chemotherapy,
22. TYZZER, E. E., AND WEINMAN, D. 1939 1, 92-99.Haemobartonella, n.g. (Bartonella olim pro 37. HAVLIK, 0. 1950 The influence of penicillinparte). H. microti, n. sp., of the field vole, and streptomycin on experimental barto-Microtus pennsylvanicus. Am. J. Hyg., 30, nellosis of rats. Bull. State Inst. Marine141-157. and Trop. Med. Gdansk, Poland, 3, 57-58.
23. NIVEN, J. S. F., GLEDHILL, A. W., DICK, G. 38. MAYER, M. 1949 La aureomicina en laW. A., AND ANDREWES, C. A. 1952 Further anemia de las ratas causada por la Barto-light on mouse hepatitis. Lancet, 263, 1061. nella muris Mayer (nota preliminar).
24. SPLITTER, E. J. 1952 Eperythrozoonosis in Arch. venez. Patol. trop. 1, 329-335.swine-filtration studies. Am. J. Vet. Re- 39. STANTON, M. L., LASKOWSKI, L., AND PINKER-search, 13, 290-297. TON, H. 1950 Chemoprophylactic ef-
25. PINKERTON, H., AND WEINMAN, D. 1937 fectiveness of aureomycin and terramycin inCarrion's disease. I. Behavior of the etio- murine bartonellosis. Proc. Soc. Exptl.logical agent within cells growing or sur- Biol. Med., 74, 705-707.viving in vitro. Proc. Soc. Exptl. Biol. 40. LWOFF, M. 1953-54 Abstracts dealing withMed., 37, 587-590. references 5, 8, 33. Bull. Inst. Pasteur, 51,
26. JIMINEZ, J. F., AND BUDDINGH, G. J. 1940 525-526; 52, 444-445.Carrion's disease. II. Behavior of Bartonella 41. DIENES, L. 1945 Morphology and nature ofbacilliformis in the developing chick embryo. the pleuro-pneumonia group of organisms.Proc. Soc. Exptl. Biol. Med., 45, 546-551. J. Bacteriol., 50, 441-458.
27. THURSTON, J. P. 1954 Anemia in mice 42. KLIENEBERGER-NOBEL, E., AND CUCKOW, F.caused by Eperythrozoon coccoides (Schilling, W. 1955 A study of organisms of the1928). Parasitology, 44, 81-98. pleuropneumonia group by electron mi-
28. REESE, J. D., MORRISON, M. E., AND FOWLER, croscopy. J. Gen. Microbiol., 12, 95-99.E. M. 1950 Complement fixation and 43. MORTON, H. E., LECCE, J. G., OSKAY, J. J.,Weil-Felix reaction in rabbits inoculated AND COY, N. H. 1954 Electron microscopewith Bartonella bacilliformis. J. Immunol., studies of pleuropneumonialike organisms65, 355-358. solated om man andahike J.gan29. WIGAND, R., AND WEYER, F. 1953 Neue teriol.a 68f 697-703.Untersuchungen uber Bartonella bacillifor- tEr 68,6 .,mis. 3. Mitteilung: t7bertragungsversuche 44. DE ROBERTIS, E., AND EPSTEIN, B. 1951auf Rhesusaffen und auf Kleiderlause. Z. Electron microscope study of anaplasmosisTropenmed. u. Parasitol., 4, 243-254. in bovine red blood cells. Proc. Soc.
30. JARAMILLO, R. 1939 Contribucion al estudio Exptl. Biol. Med., 77, 254-258.de la bartonelosis en Colombia. Rev. Hig., 45. NEITZ, W. O., ALEXANDER, R. A., AND DUBogota, 20, 13-70. TOIT, P. J. 1934 Eperythrozoon ovis (sp.
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nov.) infection in sheep. Onderstepoort J. 47. GREEN, R. H., ANDERSON, T. F., AND SMADEL,Vet. Sci. Animal Ind., 3, 263-269. J. E. 1942 Morphological structure of the
46. KUROTCHIKN, T. J., LIBBY, R. L., GAGNON, E., virus of vaccinia. J. Exptl. Med., 75, 651-AND COX, H. R. 1947 Size and morphology 656.of the elementary bodies of the psittacose- 48. PETERS, D., AND STOECKENIUs, W. 1954lymphogranuloma-inguinale group of Structural analogies of pox viruses andviruses. J. Immunol., 55, 283-287. bacteria. Nature, 174, 224.
PLATES I-IV
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PLATE I
_ S li~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
Figure 1. Bartonella bacilliformis, human blood, light microscopy, Giemsa stain. [Original pub-lished in Z. Tropenmed. u. Parasitol., 4, 539-548, 1953 (4)1
Figure 2. Haemobartonella muris, Syrian hamster blood, Giemsa stain. [Original published in Z.Tropenmed. u. Parasitol., 3, 437-452, 1952 (8)1
.TrpnFigure 3. Eperythrozoon coccoides, mouse blood, Giemsa stain. [Original published in Z rpnmed. u. Parasitol., 3, 461-472, 1952 (10)1
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PLATE II
Figure 4. Bartonella bacilliformis from 7 days' culture, electron microscope, Pd-shadowed. Rod-likeparasites, retracted cytoplasm, cell walls and flagella. [Original published in Z. Tropenmed. u. Par-asitol., 3, 313-326, 1952 (5)1Figure 5. Bartonella bacilliformis, pseudo-replica from human blood. Thin blood smears are covered by
collodion films which are taken off by diluted hydrofluoric acid and transferred to electronoptical grids.The erythrocytes have left their impression on the film as less opaque "negative" areas, while theparasites stick to the film and appear "positive." Rod with cell wall and "annular" form on erythrocytereplica. [Original published in Z. Tropenmed. u. Parasitol., 4, 539-548, 1953 (4)]
Figure 6. Bartonella bacilliformis, pseudo-replica, Chabaud fixation, trypsin-treated. Empty cellwalls, marginal folds. [Original published in Z. Tropenmed. u. Parasitol., 4, 539-548, 1953 (4)]
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PLATE III
_B_(;::_
l.
i isS
-,,z,, 7,:-: f .. 4s,. t
- r
A.:nL i,, , 'l
__fif fff fft':'--'' '
Sr ;:
1:;7n
::.::
.... ff f; _ ..s:}i. 7: . . g6 f 00 0; 0 A : 00 f *.s *.r .00002|y ,fF t000000 f7. s
Figure 7. Haemobartonella muris, attached to red cell ghost after osmotic hemolysis, Pd-shadowedCoccoid particles without structure, partly in chains. [Original published in Z. Tropenmed u Par
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PLATE IV
________,____s_
__
_s_____S____s_:Xf
_f;:g t"?3!9 !Et" h''' ,: -
| !_I E §, Ei!|- | | g E | W_ W | | ! | |_ - I RE lli,2sw__ | l | | | |_ * l * l l E |
__
- | l * | l | |_ * l | l | I_ | I * I | l_ | l * l |_ | l * l |- - | - | - |
_ | I * I l |_ | l * l l |
l - ll | ll| - - | - | | - |
_ | l * l l || l * l l |
- - l | l l | l-
* |11 11 11 1-* r11 11 111rr | | | ! | E gX - X 0 ! ! ! y y !
Figure 8. Haemobartonella muris, pseudo-replica. Parasites on erythrocyte replica. [Original publishedin Z. Tropenmed. u. Parasitol., 3, 437452,1952 (8)]
Figure 9. Eperythrozoon coccoides, pseudo-replica. Annular parasites on erythrocyte replica. [Originalpublished in Z. Naturforsch., 6b, 32S333, 1951 (9)]
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