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Cryptosporidiosis in Animals and Humans SAUL TZIPORI Attwood Veterinary Research Laboratory, Westmeadows, Victoria 3047, Australia

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Cryptosporidiosis in Animals and Humans SAUL TZIPORI Attwood Veterinary Research Laboratory, Westmeadows, Victoria 3047, Australia MICROBIOLOGICAL REVIEWS, Mar. 1983, p. 84-96 Vol. 47, No. 1 0146-0749/83/010084-13$02.00/0 Copyright © 1983, American Society for Microbiology Cryptosporidiosis in Animals and Humans SAUL TZIPORI Attwood Veterinary Research Laboratory, Westmeadows, Victoria 3047, Australia INTRODUCTION................................. 84 CHARACTERISTICS OF THE ORGANISM ................................. 84 Classification ................................. 84 Life Cycle ................................. 84 Species Specificity ................................. 87 Diagnosis .................................. 88 Studies on Oocysts .................................. 88 CHARACTERISTICS OF THE DISEASE................................. 89 The Disease in Calves................................. 89 The Disease in Lambs ................................. 89 The Disease in Goats ................................. 89 The Disease in Humans ................................. 89 The Infection in Birds ................................ 90 The Infection in Other Species ................................ 90 EXPERIMENTAL CRYPTOSPORIDIOSIS ............. ................... 90 Studies with Calf Isolates ................................ 90 Studies with Human Isolates................................ 91 TREATMENT................................ 92 CONCLUSIONS ................................ 92 SUMMARY ................................ 94 LITERATURE CITED ................................ 94 INTRODUCTION CHARACTERISTICS OF THE ORGANISM Classification Cryptosporidium is a protozoan parasite which completes its life cycle on intestinal and Cryptosporidium is a genus in the family respiratory surface epitheliums of mammals, Cryptosporidiidae, suborder Eimeriina, order birds, and reptiles (28). The infection, until Eucoccidiida, subclass Coccidia, class Sporo- recently, was thought to be uncommon and the zoa, phylum Apicomplexa (30). At present the organism was thought to be opportunistic and, suborder Eimeriina contains 13 families with like other coccidia, highly host specific. over 1,500 named species. The great majority of Cryptosporidium was first recognized in the these species, however, belong to the genera gastric glands ofthe laboratory mouse by Tyzzer Eimeria and, to a lesser extent, Isospora. Both in 1907 (58). Up to 1975, some 15 reports de- are intracellular parasites which primarily infect scribing the infection in eight species of animals the intestinal tract of vertebrates. Toxoplasma, a were published; only 5 of them associated cryp- tissue cyst-forming coccidia, is another impor- tosporidiosis with some illness, 3 of which were tant member of the suborder Eimeriina. in calves (36, 42, 52). Since 1975, over 60 Life scientific publications have appeared in the liter- Cycle ature, the great majority being since 1980. Dur- The life cycle of Cryptosporidium has been ing this period our concept of cryptosporidiosis elucidated for a number of representative host was transformed from that of a rare and largely species and generally follows that of other enter- asymptomatic infection to an important cause of ic coccidia (10, 22, 77). However, there remain enterocolitis and diarrhea in several species, areas of discrepancy among authors regarding including humans. the presence (77) or absence (7, 22, 59) of a In this communication an attempt was made second-generation schizont. It has been suggest- to gather the available information on the orga- ed that the second-generation schizont may sim- nism, to examine the nature of the infection it ply be sporocysts containing four sporozoites produces, and to demonstrate that Cryptospo- (44). However, sporocysts have not been identi- ridium infection can, under certain circum- fied in structures considered to be oocysts (7), stances, cause serious disease in some species of and the genus is not considered to have sporo- animals. cysts by some (29). The morphological charac- 84 VOL. 47, 1983 CRYPTOSPORIDIOSIS IN ANIMALS AND HUMANS 85 FIG. 1. Electron micrograph of a macrogamete with characteristic dense polysaccharide granules and central nucleus (N). Note: (i) the feeder organelle (F) at the attachment zone; (ii) the parasitophore (P) between the inner parasitic plasma membrane and the outer membrane which, like the microvilli of enterocytes, is coated with glycocalyx. Whether the outer membrane is derived from the host cell, therefore making the organism intracellular (10, 22, 77), or is of parasitic origin and therefore extracellular (44, 59) is a point of dispute (x 30,000). teristics and the nature of the parasite-host rela- ever the nature of the host-parasite association, tionship have been studied in some detail in a the physical location of the organism outside the number of representative host species, and the cell boundaries (Fig. 1) distinguishes it from question of whether the parasite ought to be other coccidia which develop and multiply with- regarded as intracellular (7, 10, 22, 77) or extra- in the cytoplasm of the cell. Although some cellular (44, 59, 60) remains unresolved. What- differences in the various morphological studies 86 TZIPORI MICROBIOL. REV. emerge, they presumably reflect differences in surface; oocysts without sporocysts with 4 na- methodology and interpretation rather than spe- ked sporozoites; microgometes without flagella; cies variation. There has been some doubt as to one genus and about 11 named species" (29). the existence of an oocyst (10, 77), but it is now Cryptosporidium has largely been observed in believed that spherical bodies measuring 3 to 4 association with the epithelium of the small or ,um (Fig. 2) and containing four naked sporozo- large intestine or both of clinically healthy mice ites are the exogenous infective stage (7, 22, 44). (17, 59, 60), rabbits (21, 50, 61), chickens (61), It has been demonstrated in mice recently that geese (47), guinea pigs (24, 76, 77), and cats (22); these sporozoites sporulate endogenously and or of clinically ill calves (2, 7, 36, 42, 43, 46, 52, therefore are infective when discharged in the 55, 70), lambs (4, 63), humans (2, 27, 35, 41, 49, feces (49). Figure 3 provides a diagrammatic 54, 57, 66, 78, 79), goats (33, 71), turkeys (20, 48, presentation of an interpretation of the life cycle 53), deer (65), monkeys (13, 26), and immuno- of Cryptosporidium which was adapted from logically deficient foals (56). Iseki (22). The generic character of Cryptospori- Cryptosporidium has also been observed in- dium is described as "Homogeneous: develop- fecting the trachea (20, 32, 48), cloaca (14), mentjust under surface membrane of host cell or bursa of Fabricius (15), and conjunctival sacs within its brush border and not in the cell (32) of birds, the stomach of mice (58) and proper. Oocysts and meronts with a knoblike snakes (11, 34), and the bile ducts of a monkey attachment organelle at some point on their (26) and immunodeficient foals (56). FIG. 2. Electron micrograph showing an oocyst (left) and a mature macrogamete, attached to the surface epithelium of lamb ileum experimentally infected with Cryptosporidium (x50,000). VOL. 47, 1983 CRYPTOSPORIDIOSIS IN ANIMALS AND HUMANS 87 OUTSIDE HOST FIG. 3. Diagrammatic representation of the life cycle of Cryptosporidium. (1 to 4) Asexual cycle of the endogenous stage: 1, sporozoite or merozoite invading a microvillus of a small intestinal epithelial cell; 2, a fully grown trophozoite, 3, a developing schizont with eight nuclei; 4, a mature schizont with eight merozoites. (5 and 6) Sexual cycle: 5, microgametocyte with many nuclei; 6, macrogametocyte. (7) A mature oocyst containing four sporozoites without sporocyst. (8) Oocyst discharged in the feces. a, Merozoite released from mature schizont; b, sporozoite released from mature oocyst. Adapted from Isaki (22). Species Specificity serially passaged three to four times in mice (unpublished data). The human isolates in our Based on early transmission experiments experience also appear to infect baby mice less Cryptosporidium was thought to be not only readily than do isolates from other species (Tzi- host (22, 58, 76) but site specific (60). More pori, Bhathal, et al., submitted); on the other recent experiments, using specific-pathogen- hand, human isolates studied by Reese and co- free (SPF) newborn animals, indicate that Cryp- workers (49) were said to infect adult mice as tosporidium from some species of animals can well as newborns. Tyzzer was able to transmit infect a wide variety of other animals, with or infection between adult mice (60), whereas a without causing illness (64). Thus, isolates from bovine isolate only infected mice younger than calves, humans, deer, goats, and lambs readily 21 days (51). These differences could be attribut- infect other species such as lambs, calves, and ed to either host or parasite factors or both. The piglets, causing diarrhea (37, 62, 64, 67, 72, 75; exact nature and indeed the existence of such S. Tzipori, P. S. Bhathal, M. Smith, and C. strain differences will become clear when more Halpin, submitted for publication), and mice, information is available, including the develop- rats, guinea pigs, chickens, and foals without ment of more sensitive serological tests and the causing illness (49, 51, 64; Tzipori, Bhathal, et availability of techniques for accurately estimat- al., submitted; unpublished data). However, ing the infectious dose in experimental animals. some differences between isolates have been Passage of Cryptosporidium experimentally noted which could indicate "strain" variations. through animals
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