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BAKTERI ANAEROB

MORFOLOGI, FISIOLOGI,
EPIDEMIOLOGI, DIAGNOSIS,
PEMERIKSAAN
Sy. Miftahul El J.T

Figure 4-1 Catabolism of proteins, polysaccharides, and lipids produces glucose, pyruvate, or
intermediates of the tricarboxylic acid (TCA) cycle and, ultimately, energy in the form of adenosine
triphosphate (ATP) or the reduced form of nicotinamide-adenine dinucleotide (NADH).
Downloaded from: StudentConsult (on 20 October 2009 07:04 AM)
© 2005 Elsevier

Oksigen  berdasarkan keperluan oksigen dibagi dalam 5
kelompok
Group

Environment

O2 Effect

Aerobic

Anaerobic

Obligate
Aerobe

Growth

No growth

Microaerophile

Growth
if
level not too No growth
high

Required but at levels
below 0.2 atm

Obligate
Anaerobe

No growth

Growth

Toxic

Required
(utilized
aerobic respiration)

for

Facultative
Anaerobe
(Facultative
Aerobe)

Growth

Growth

Not required for growth
but
utilized
when
available

Aerotolerant
Anaerobe

Growth

Growth

Not required
utilized

and

not

BAKTERI ANAEROB GRAM POSITIP
BERSPORA
[CLOSTRIDIUM spp]
• Genus ini meliputi kuman berbentuk batang pleimorf, spora lebih
besar dari badan kuman, anaerob obligat
• Gram-positive rods capable of forming endospores was placed in
the

genus

Clostridium.

This

genus

properties:
(1) presence of endospores,
(2) strict anaerobic metabolism,
(3) inability to reduce sulfate to sulfite, and
(4) Gram-positive cell wall structure

was

defined

by

four

oxygen-deprived environment (3)production of numerous histolytic toxins. water. enterotoxins. food poisoning. Most clostridia are harmless saprophytes • The majority of infections seen today are skin and soft-tissue infections. (2)rapid growth in a nutritionally enriched. and sewage and are part of the normal microbial flora in the gastrointestinal tracts of animals and humans. and antibiotic-associated diarrhea and colitis. • The remarkable capacity of clostridia to cause diseases is attributed to their (1)ability to survive adverse environmental conditions through spore formation. and neurotoxins. .• The organisms are ubiquitous in soil.

Species Human Disease C. innocuum C. myonecrosis. sporogenes Gas gangrene Opportunistic infections Gas gangrene Gas gangrene Opportunistic infections Frequenc y pseudomembranous Common Rare Rare Rare Rare Rare .. food poisoning. botulinum C. tetani C. colitis Soft-tissue infections (e. enteritis necroticans. histolyticum C. clostridioforme C. cellulitis. sordellii C. septicum C. novyi C.g. baratii C. septicemia Gas gangrene. suppurative Common myositis. perfringens C. gas gangrene). septicemia Uncommo n Botulism Uncommo n Tetanus Uncommo n Opportunistic infections Uncommo n Botulism Rare Botulism Rare Opportunistic infections Rare C. tertium C. butyricum C. difficile Antibiotic-associated diarrhea.

CLOSTRIDIUM DIFFICILE • Until the mid-1970s the clinical importance of C. difficile is responsible for antibiotic-associated gastrointestinal diseases . self-limited diarrhea to severe. Systematic studies now clearly show. however. This organism was infrequently isolated in fecal cultures and its role in human disease was unknown. difficile was not appreciated. life- threatening pseudomembranous colitis (GAMBAR). that toxin-producing C. ranging from a relatively benign. .

Note the white plaques of fibrin. mucus.Antibiotic-associated colitis: gross section of the lumen of the colon. Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier . and inflammatory cells overlying the normal red intestinal mucosa.

editors: Infectious diseases illustrated. with the characteristic "plaque" (black arrow) overlying the intact intestinal mucosa (white arrow). Farrar WE. 1982.) (From Lambert HP.Antibiotic-associated colitis caused by Clostridium difficile.) Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier . (Hematoxylin and eosin stain. London. A histologic section of colon shows an intense inflammatory response. Gower.

Virulence Factors Associated with Clostridium difficile . induces cytokine production with hypersecretion of fluid. produces hemorrhagic necrosis Induces depolymerization of actin with loss of cellular cytoskeleton Mediates binding to human colonic cells Produces hydrolytic activity Permits organism's survival for months in hospital environment .Virulence Factor Enterotoxin (toxin A) Cytotoxin (toxin B) Adhesin factor Hyaluronidase Spore formation Biologic Activity Produces chemotaxis.

4 × 1. perfringens causes most of the human infections in the United States. perfringens (alpha [α]. tabel). C. perfringens can be associated with simple colonization or can cause lifethreatening disease.Clostridium perfringens • Dahulu dikenal C. welchii. perfringens is a large (0. • This organism is one of the few nonmotile clostridia. is hemolytic.6 to 2. merupakan salah satu penyebab ganggren gas. beta [β]. . Type A C. epsilon [ε]. and is metabolically active. features that make possible its identification in the laboratory.3 to 19. Gram-positive rod (gambar) . keracunan makanan oleh enterotoksin yang termolabil atau enteritis nekrotik • C. and iota [ι] toxins) is used to subdivide isolates into five types (A through E.0 μm). rectangular. The production of one or more major lethal toxins by C. but rapidly spreading growth on laboratory media (resembling the growth of motile organisms) is characteristic (GAMBAR). with spores rarely observed either in vivo or after in vitro cultivation. The organism grows rapidly in tissues and in culture.

Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier .back Gram stain of Clostridium perfringens.

-toxin). back Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier . combined with the characteristic microscopic morphology. Growth of Clostridium perfringens on sheep blood agar.-toxin) and a wider zone of partial hemolysis (caused by the α. perfringens can be made by detection of a zone of complete hemolysis (caused by the Ѳ. spreading colonies and the hemolytic activity of the organism. A presumptive identification of C. Note the flat.GAMBAR 2.

Type A B C D E Lethal Toxins Alph Beta Epsil a on Iota + + + + + + + + + + Distribution of Lethal Toxins in Clostridium perfringens Types A to E back .

The α. This reaction (Nagler's reaction) is characteristic of C.Growth of Clostridum perfringens on egg-yolk agar. This precipitate is not observed when the organism is grown in the presence of antibodies against the toxin (left). producing an opaque precipitate (right). Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier . perfringens.-toxin (lecithinase) hydrolyzes phospholipids in serum and egg yolk.

necrotizing activity Lethal toxin. superantigen .and oxygen-labile hemolysin. necrotizing activity. hemolysin. adenosine diphosphate (ADP) ribosylating Hemolysin Heat. phospholipase C (lecithinase). as seen in myonecrosis Lethal toxin. enterotoxic). hemolysin. necrotizing activity Alters membrane permeability in ileum (cytotoxic. permease Lethal binary toxin. increases vascular permeability.Virulence Factors α-Toxin β-Toxin ε-Toxin ι-Toxin δ-Toxin θ-Toxin κ-Toxin λ-Toxin μ-Toxin ν-Toxin Enterotoxin Biologic Activity Lethal toxin. cytolytic Collagenase. gelatinase. produces necrotizing activity. necrotizing activity Protease Hyaluronidase Deoxyribonuclease.

enterotoksin merangsang enzim adenylate cyclase pada dinding usus →bertambahnya konsentrasi cAMP →hipersekresi air dan Cl dalam usus →menghambat reabsorpsi Na →Diare [selama 1-3 hari] • Diagnosa laboratorium – Spesimen dari luka. agar darah secara an aerob • Pengobatan – Pembersihan luka secara bedah pada jaringan nekrotik – Antibiotik : Penisilin – Hyperbaric oxygen . pus. • Pada keracunan makanan. batang tanpa spora – Biakan: di tanam pada agar tioglikolat.• Pada ganggren gas. terjadi hemolisis intravaskuler. karbohidrat akan dihancurkan dengan pembentukan gas karena ada septikemia. jaringan atau makanan – Mikroskopis: Gram +/P.

Clostridia can be introduced into tissue during surgery or by a traumatic injury. A serosanguineous exudate and subcutaneous gas were present.Clostridial cellulitis. the skin became discolored and bullae and necrosis developed. 1982. Farrar W. editors: Infectious diseases illustrated. This patient suffered a compound fracture of the tibia. London. but there was no evidence of muscle necrosis. Five days after the injury. DownloadedGower. The patient had an uneventful recovery. (From Lambert H.) from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier .

4 × 3. argentinense. sulit bicara. Kerja toksin akan memblokir pembentukan/pelepasan acetyl cholin pada hubungan saraf otot sehingga terjadi kelumpuhan otot. and Clostridium argentinense (type G toxin). the etiologic agent of botulism. kalengan tanpa dimasak terlebih dahulu. . Human disease has only rarely been associated with C.Clostridium botulinum • • • • • • • • Terdapat secara luas di alam. Gejala 18-96 jam makan toksin dengan keluhan penglihatan. Kematian terjadi karena paralisis otot pernafasan atau kelumpuhan jantung [cardiac arrest] Other species of clostridia produce botulinum toxins. butyricum and C. is a heterogeneous group of large (0. butyricum (type E toxin). baratii (type F toxin). diasap. jenis sakarolitik dan tidak proteolitik [Jenis C. fastidious. baratii. botulinum. tetapi menyebabkan keracunan makanan. dan E ] semua jenis kelompok ini menghasilkan sulfida Dosis letal untuk manusia : 1 mikrogram Tidak menyebabkan infeksi luka. akibat toksin yang termakan. D. kadang terdapat pada feses binatang C.0 to 20. B dan F]. Umumnya makanan yang tercemar adalah makanan yang berbumbu. C.2 μm).6 to 1. including C. anaerobic rods Berdasarkan sifaT biokimia ada 2 jenis : Proteolitik [jenis A. hal ini terjadi karena tidak ada koordinasi. and not definitively demonstrated with C. sporeforming. Sulit menelan.

saccharolytic Weakly proteolytic. E. D IV G Phenotyptic Properties Proteolytic. saccharolytic Weakly proteolytic. asaccharolytic Clostridium botulinum Classification and Toxin Production . F C. saccharolytic Nonproteolytic. B.Group I II III Neurotoxin Type A. F B.

5µ. tidak tahan terhadap oksigen. membentuk 3 jenis toksin : – Hemolisin [tetanolisin] • Termolabil. agak bengkok. reflek yang berlebihan dan serangan kejang berulang. ukuran 4.Clostridium tetani • Differential diagnosa – Kuman berbentuk panjang langsing.4 – Tumbuh baik pada media agar darah dengan zona hemolisa alfa kemudian menjadi beta akibat pembuatan hemolisin [tetanolisin] • Toksin. Zat ini akan terikat kuat pada ganglion otak menyebabkan penghambatan neuron spinal pasca sinaps dengan mencegah pelepasan perantara penghambat [inhibitory mediator] → kejang otot menyeluruh. bekerja aktif pada SDM dari sebagian besar binatang[ kelinci. Gram +/P. – Neurotoksin non-spasmogenik yang bekerja di saraf perifer . kuda dll] • Peran belum jelas mungkin bekerja sebagai lekosidin – Neurotoksin [tetanospasmin] • • Tahan terhadap oksigen dan diinaktifkan pada 66 0C selama 5 mnt. terminal [seperti pemukul genderang].8 µ x 0. dapat dibuat toksoid dengan penambahan formaldehid konsentrasi rendah Bekerja seperti striknin dengan menghambat sintesis dan pelepasan asetilkolin sehingga menganggu transmisi neuromuskuler. tidak berkapsul dan motil – Suhu pertumbuhan optimum 370C dan pH 7. tunggal atau berbentuk rantai – Spora bulat.

Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier . Note the terminal spores.Gram stain of Clostridium tetani.

Tetanus neonatorum : akibat pencemaran luka pemotongan tali pusat bayi. akibat Infeksi saluran kelamin oleh alat bantu persalinan dan pembalut 3. Tetanus pasca keguguran dan masa nifas. Splanchnic tetanus: terjadinya kekejangan otot – otot menelan dan pernafasan 4. akan tetap diluka.Clostridium tetani • Patogenesis – Kuman tidak bersifat invasif. angka kematian tinggi 2. dapat menjadi bentuk vegetatif bila kondisi mendukung/anaerob. Cephalic tetanus: luka di daerah kepala akan terjadi kontraksi otot – otot muka unilateral dan bilateral . misalnya adanya: • Jaringan nekrotik • Garam kalsium • Kuman piogenik Beberapa jenis tetanus : 1.

ed 2.Facial spasm and risus sardonicus in a patient with tetanus. Mosby). St Louis. (From Cohen J. Powderly WG: Infectious diseases. Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier . 2004.

) Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier . ed 3. Rowland HAK. Wolfe. (From Emond RT.A child with tetanus and opisthotonos resulting from persistent spasms of the back muscles. 1995. Welsby P: Colour atlas of infectious diseases. London.

Clostridium tetani • Diagnosa laboratorium – Pemeriksaan mikroskopis: sampel diambil dari luka. diwarnai Gram →+/P seperti pemukul genderang – Biakan: potongan jaringan nekrotik dibiakkan pada kaldu daging/agar darah – Percobaan binatang : untuk melihat toksigenitas kuman. kemudian dilihat mulai terjadinya kekejangan • Pencegahan & pengobatan – Pembersihan luka – Imunisasi aktif dengan toksoid – Imunisasi pasif dengan ATS – Pemberian antibiotik :Penicillin . digunakan mencit yang disuntikan 0.2 ml suspensi kuman pada pangkal ekor.

ada jaringan nekrotik] 1 x toksoid 1 x toksoid ATS.PENCEGAHAN TETANUS PADA LUKA Keadaan luka Keadaan kekebalan Kebal Kebal sebagian Tidak kebal Bersih [kecelakaan rumah dlm 6 jam 1 x toksoid 1 x toksoid 3 x toksoid Tercemar [tanah. Antibiotik Terinfeksi 1 x toksoid Antibiotik 1 x toksoid Ats. Antibiotik 3 x toksoid ATS. Antibiotik Catatan : Kebal :Artinya telah menerima dosis lengkap berupa 3 x penyuntikan toksoid Kebal sebagian :artinya telah menerima 2 x suntikan toksoid Tidak kebal : artinya belum menerima toksoid atau keadaan kekebalan tidak diketahui . Antibiotik 3 x toksoid ATS.

makanan yang perlu diperhatikan : Kacang-kacangan.Clostridium botulinum • Diagnosa laboratorium : mendeteksi toksin yang ada didalam serum penderita atau dari sisa makanan. jagung.B dan C] secara IV dan secara simptomatik terutama untuk pernafasan • Pencegahan: Makanan yang diawetkan harus dimasak dahulu secara baik. ikan asap atau ikan segar dalam plastik . • Pengobatan:Pemberian antitoksin polivalen [tipe A. Pendeteksian dilakukan dengan reaksi Netralisasi antigen-antibodi secara agglutinasi SDM yang dilapisis antiserum. Percobaan mencit yang disuntikkan bahan tersangka.

Clostridium botulinum .

Thus the isolation and identification of individual strains are difficult and often time consuming. • These organisms are opportunistic pathogens. • Most of these anaerobes have fastidious nutritional requirements and grow slowly on laboratory media.ANAEROB.spore-forming rods are a heterogeneous group of bacteria that characteristically colonize the skin and mucosal surfaces. . GRAM POSITIP. TIDAK MEMBENTUK SPORA • The anaerobic gram-positive cocci and non. typically responsible for endogenous infections and usually recovered in mixtures of aerobic and anaerobic bacteria.

Schleifer aktinos. curved rod) propionic acid (propionic acid is the primary metabolic product of fermentation) Important Anaerobic Gram-Positive Bacteria . cell (tiny cell) pepto. lactic acid is the primary metabolic product of fermentation) mobilis. aer. cook or digest (the digesting streptococcus) Named after the German microbiologist K. organism originally recovered in milk. milk (milk bacillus. coccus. berry or coccus (anaerobic coccus) Named after the American microbiologist S. cleft.H. good or beneficial (a beneficial rod. mykes. fungus (ray fungus referring to the radial arrangement of filaments in granules) bifidus. uncus. also. capable of movement or being active. air. that is. without. bakterion. Finegold micro. hook (motile. a rod normally present) acto. tiny. monas. ray. small rod (a small clefted or bifurcated rod) eu.Organism Anaerobic Cocci Anaerococcus Finegoldia Micromonas Peptostreptococcus Schleiferella Anaerobic Rods Actinomyces Bifidobacterium Eubacterium Lactobacillus Mobiluncus Propionibacterium propionicum Historical Derivation an.

reproduce by fission. and are inhibited by penicillin but not antifungal antibiotics. Actinomyces israelii. • However. slowly developing infections. gram-positive rods. Numerous species have been described. and Actinomyces viscosus are responsible for most human infections. . these organisms are true bacteria in that they lack mitochondria and a nuclear membrane. Actinomyces meyeri. • They grow slowly in culture. They are not acid-fast (in contrast to the morphologically similar Nocardia species). The other species grow best in anaerobic conditions but can grow aerobically. meyeri is a strict anaerobe. Actinomyces naeslundii. Actinomyces odontolyticus.• ACTINOMYCES Actinomyces organisms are facultatively anaerobic or strictly anaerobic. and they tend to produce chronic. • Only A. They typically develop delicate filamentous forms or hyphae (resembling fungi) in clinical specimens or when isolated in culture (GAMBAR.

Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier .Macroscopic colony (left) and Gram stain (right) of Actinomyces.

Actinomycosis is characterized by the development of chronic granulomatous lesions that become suppurative and form abscesses connected by sinus tracts. These colonies.PATHOGENESIS AND IMMUNITY •  Actinomyces organisms colonize the upper respiratory. Macroscopic colonies of organisms resembling grains of sand can frequently be seen in the abscesses and sinus tracts. The organisms have a low virulence potential and cause disease only when the normal mucosal barriers are disrupted by trauma. gastrointestinal. are masses of filamentous organisms bound together by calcium phosphate (GAMBAR). surgery. . • Disease caused by actinomyces is termed actinomycosis (in keeping with the original idea that these organisms were fungi or "mycoses"). These bacteria are not normally present on the skin surface. or infection. and female genital tracts. called sulfur granules because they appear yellow or orange.

Downloaded from: StudentConsult (on 20 October 2009 07:04 AM) © 2005 Elsevier . Delicate filamentous rods (arrow) are seen at the periphery of the crushed granule.Sulfur granule collected from the sinus tract in a patient with actinomycosis.

such as soil or water. the actinomyces that are present in the mouth invade into the diseased tissue and initiate the infectious process. with the disease spread or disease originating from an becoming established in the lungs and then with no evidence of external source.EPIDEMIOLOGY • Actinomycosis is an endogenous infection Patients with thoracic infections generally person-to-person have a history of aspiration. Abdominal infections most commonly occur in patients who have undergone gastrointestinal surgery or have suffered trauma to the bowel. Pelvic infection can be a secondary manifestation of abdominal actinomycosis or may be a primary infection in a woman with an intrauterine device (Central nervous system infections usually represent hematogenous spread from another infected tissue. . spreading to adjoining tissues. In these patients. such as the lungs. Cervicofacial infections are seen in patients who have poor oral hygiene or have undergone an invasive dental procedure or oral trauma. • • Disease is classified according to the organ systems involved.

clindamycin. the antibiotic erythromycin. PREVENTION. . Actinomyces are uniformly susceptible to penicillin (considered choice). extensive tissue destruction. and the tetracyclines have variable activity. or gastrointestinal antibiotic tract is penetrated can lower the risk of these infections. Most species of Maintenance hygiene and of good the use oral of and appropriate are prophylaxis when the mouth resistant to metronidazole. AND CONTROL • • Treatment for actinomycosis involves the combination surgical débridement of of the • The clinical generally response good even is in involved tissues and the prolonged patients who have suffered administration of antibiotics.TREATMENT.

Propionibacterium acnes and Propionibacterium . conjunctiva. external ear. The organisms are anaerobic or aerotolerant. and in the oropharynx and female genital tract. nonmotile. The two most commonly isolated species are propionicus.Propionibacterium • Propionibacteria are small gram-positive rods often arranged in short chains or clumps • They are commonly found on the skin (in contrast with the actinomyces). and capable of fermenting carbohydrates. catalase positive.

• Acne is unrelated to the effectiveness of skin cleansing because the lesion develops within the sebaceous follicles.g...g. acne is managed primarily through the topical application of benzoyl peroxide and antibiotics. • Antibiotics such as erythromycin and clindamycin have proved effective for treatment. artificial heart valves or joints) or intravascular lines (e. but this finding usually represents contamination with bacteria on the skin at the phlebotomy site. For this reason. catheters. . cerebrospinal fluid shunts). acnes is responsible for two types of infections: (1) acne (as the name implies) in teenagers and young adults and (2) opportunistic infections in patients with prosthetic devices (e. • Propionibacteria are also commonly isolated in blood cultures.•  P.

Despite their appearance in Gram- • The organisms are fastidious. stained specimens they are classified as gram-positive rods slowly enriched media supplemented ends.Mobiluncus •  Members of the genus Mobiluncus are obligate anaerobic. gram-variable • or growing gram-negative. and in (3) Are susceptible to vancomycin. and ampicillin but resistant to low numbers but are abundant colistin. mulieris. curved rods with tapered because they even on with rabbit or horse serum. . in humans. have been identified (2) lack endotoxin. clindamycin. curtisii and Mobiluncus Mobiluncus (1) have a gram-positive cell wall. colonize the genital tract in erythromycin. • Two species. women The with organisms bacterial vaginosis (vaginitis).

g. and tract. intestines. and (3) opportunistic septicemia in an immunocompromised patient. after childbirth or a gynecologic procedure).Lactobacillus •  Lactobacillus species are facultatively anaerobic or strictly anaerobic rods. The organisms are most commonly isolated in urine specimens and blood cultures. genitourinary of the mouth. Invasion into blood occurs in one of the following three settings: (1) transient bacteremia from a genitourinary source (e.. • They are found as part of the normal flora stomach. . (2) endocarditis. Because lactobacilli are the most common organism in the urethra • The reason lactobacilli rarely cause infections of the urinary tract is their inability to grow in urine.

Lactobacillus • Treatment of endocarditis and opportunistic infections is difficult because lactobacilli are resistant to vancomycin (an antibiotic commonly active against grampositive bacteria) and are inhibited but not killed by other antibiotics. . A combination of penicillin with an aminoglycoside is required for bactericidal activity.

bacteria can be and vagina. large intestine. isolated in These clinical specimens but have a very low virulence potential and usually represent clinically insignificant contaminants.Bifidobacterium and Eubacterium •  Bifidobacterium and Eubacterium species are commonly found in the oropharynx. .

the endogenous anaerobic pathogen. prototypical . Porphyromonas. and Prevotella and the cocci in the genus Veillonella • Among these pathogens. Fusobacterium. gastrointestinal.Anaerobic Gram-Negative Bacteria •  The most important gram-negative anaerobes that colonize the human upper respiratory. and genitourinary tracts are the rods in the genera Bacteroides. the most important is Bacteroides fragilis.

Virulence Negative Rods Virulence Factor Adhesins Capsule Factors in Anaerobic Gram- Bacteria Bacteroides fragilis.. IgM. Prevotella melaninogenica B.Table . fragilis. P. gingivalis Fusobacterium nucleatum Fimbriae Hemagglutinin Lectin Resistant to Oxygen Toxicity Superoxide dismutase Many species Catalase Many species Antiphagocytic Capsule B. IgG Porphyromonas spp. melaninogenica Immunoglobulin (Ig)A. fragilis. Porphyromonas gingivalis P. proteases Prevotella spp. Lipopolysaccharide Fusobacterium sp Succinic acid Many species .

New York. 2005. ed 6. . Porphyromonas. 1994. and Fusobacterium species.Virulence Factor Tissue Destruction Phospholipase C Hemolysins Proteases Collagenase Fibrinolysin Neuraminidase Heparinase Chondroitin sulfatase Glucuronidases N-Acetylglucosaminidase Volatile fatty acids Toxin Enterotoxigenic toxin   Bacteria Fusobacterium necrophorum Many species Many species Many species Many species Many species Many species Many species Many species Many species Many species B. and Lorber B: Bacteroides. Churchill Livingstone. In Mandell. Prevotella. Douglas and Bennett's principles and practice of infectious diseases. fragilis Modified from Duerden B: Clin Infect Dis 18(suppl 4):S253-S259.