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Clostridial Enteric Diseases of Domestic Animals J CLINICAL MICROBIOLOGY REVIEWS, Apr. 1996, p. 216–234 Vol. 9, No. 2 0893-8512/96/$04.0010 Copyright q 1996, American Society for Microbiology Clostridial Enteric Diseases of Domestic Animals J. GLENN SONGER* Department of Veterinary Science, University of Arizona, Tucson, Arizona 85721 INTRODUCTION .......................................................................................................................................................216 CLOSTRIDIUM PERFRINGENS...............................................................................................................................216 Introduction.............................................................................................................................................................216 Major Toxins ...........................................................................................................................................................216 Disease and Pathogenesis by Toxin Type............................................................................................................217 Type A...................................................................................................................................................................218 Type B...................................................................................................................................................................219 Type C ..................................................................................................................................................................219 Type D ..................................................................................................................................................................220 Type E...................................................................................................................................................................220 Enterotoxigenic C. perfringens ...........................................................................................................................221 Reports of untyped C. perfringens .....................................................................................................................221 Prophylaxis and Therapy.......................................................................................................................................222 Diagnosis..................................................................................................................................................................222 CLOSTRIDIUM SEPTICUM ......................................................................................................................................223 Introduction.............................................................................................................................................................223 Virulence Attributes and Pathogenesis of Enteric Disease...............................................................................223 Prophylaxis and Therapy.......................................................................................................................................224 Diagnosis..................................................................................................................................................................224 CLOSTRIDIUM DIFFICILE ......................................................................................................................................224 CLOSTRIDIUM SPIROFORME.................................................................................................................................225 CLOSTRIDIUM COLINUM........................................................................................................................................226 REFERENCES ............................................................................................................................................................226 INTRODUCTION common in the intestinal tracts of animals (68) and can occa- sionally be found in the environment in areas where disease Members of the genus Clostridium are widely recognized as produced by these organisms is enzootic (292). Because it is a enteric pathogens of humans, domestic animals, and wildlife frequent postmortem invader from the gut, isolation of C. (Tables 1 and 2). Their array of proven and putative virulence perfringens from tissues of dead animals must be viewed with attributes is impressive, and infections take a plethora of forms caution when making a diagnosis (69). in myriad hosts. In spite of the ready availability of inexpensive, usually effective products for immunoprophylaxis, clostridial Major Toxins enteric infections remain a common presentation at veterinary diagnostic laboratories. As many as 17 exotoxins of C. perfringens have been de- This review will be limited to the more common clostridial scribed in the literature (163, 263, 266, 271, 353, 388), but a enteric diseases of domestic animals, although reference will definitive role in pathogenesis has been demonstrated for only be made to human diseases as needed to provide a contextual a few. The species is divided into types on the basis of produc- background. Some infections that might be considered enteric tion of the four major toxins, a, b, ε, and i (Table 3), as (e.g., Clostridium chauvoei infection of gastrointestinal muscu- determined by in vivo protection tests performed by intrader- lature and toxin production by Clostridium botulinum in the mal injection of guinea pigs or intravenous (i.v.) inoculation of gut) are not addressed. Recent reviews (40, 142, 176, 261, 263, mice (47, 263, 387, 388, 434). Toxin (filtered culture superna- 264, 275, 292, 384, 388) may be useful in exploring other tant fluids or eluates from gut contents), both trypsin treated aspects of pathogenesis, diagnosis, and treatment. and untreated, is injected alone or mixed with antiserum. Re- sponses in guinea pigs (dermonecrosis) are noted after 24 and CLOSTRIDIUM PERFRINGENS 48 h and in mice (lethality) through 72 h. Types are determined from the results, with type A defined as strains producing a Introduction toxin, type B as strains producing a, b, and ε toxins, type C as strains producing a and b toxins, type D as strains producing a C. perfringens may be the most widely occurring pathogenic ε bacterium (384) and is certainly the most important cause of and toxins, and type E as strains producing a and i toxins clostridial enteric disease in domestic animals (Table 1). Some (Table 1). Some strains of type A do not produce enough a types of C. perfringens (mainly type A) are consistently recov- toxin to kill mice under typical test conditions, and nonlethal ered both from the intestinal tracts of animals and from the strains are referred to as nontoxigenic by some but as type A by environment, while others (types B, C, D, and E) are less others (176). Based upon production of certain minor toxins, varieties or subtypes have been reported within types A, B, and C (47, 298), although some contend that establishing new types * Phone: (520) 621-2962. Fax: (520) 621-6366. Electronic mail ad- to distinguish minor differences would become unmanageable dress: [email protected]. (397). 216 VOL. 9, 1996 CLOSTRIDIAL ENTERIC DISEASES OF DOMESTIC ANIMALS 217 TABLE 1. Diseases produced by toxigenic types of C. perfringensa TABLE 3. Activities of the major toxins of C. perfringens Toxin type Major toxin(s) Diseases Toxin Activity and effects A a Myonecrosis, food poisoning, a ...........Phospholipase/sphingomyelinase C, hemolytic, lethal, necrotic enteritis in fowl, necrotizing b ...........Induces inflammation, necrosis of intestinal mucosa, lethal enterotoxemia in cattle and ε lambs, necrotizing enterocolitis ............Protease-activated prototoxin; increases intestinal in piglets; possibly equine permeability; central nervous system toxicity; lethal colitis, canine hemorrhagic i.............Ia ADP-ribosylates actin; Ib mediates binding; gastroenteritis dermonecrotic, lethal B a, b, ε Dysentery in newborn lambs, chronic enteritis in older lambs (pine), hemorrhagic enteritis in ε ε neonatal calves and foals, The toxin is produced as the minimally toxic prototoxin hemorrhagic enterotoxemia in (266), which is converted to the .1,000-fold-more-toxic form adult sheep by proteolytic removal of 14 N-terminal amino acids (34). C a, b Enteritis necroticans (pigbel) in Although its precise biological activity has not been identified, humans; necrotic enteritis in ε toxin is necrotizing and lethal (19, 54), and its toxicity is fowl; hemorrhagic or necrotic estimated to be 3.2 3 106 i.v. minimum lethal doses per mg in enterotoxemia in neonatal pigs, mice (363). The ε toxin gene (etx) is thought to reside on a lambs, calves, goats, foals; acute large plasmid, distinct from that bearing cpb, and is found only enterotoxemia (struck) in adult sheep in strains of types B and D (77). The recent cloning and D a, ε Enterotoxemia in lambs (pulpy characterization of etx (181) will yield new information about kidney) and calves, enterocolitis toxin structure and function and provide the basis for second- in neonatal and adult goats, generation vaccines (442). possibly enterotoxemia in adult The i toxin consists of components Ia and Ib (65, 400, 401). cattle Globular skeletal muscle and nonmuscle actin are ADP-ribo- E a, i Enterotoxemia likely in calves and sylated by Ia (380, 427), which
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