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Bacillus Cereus Spores and Cereulide in Food-Borne Illness

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Bacillus Cereus Spores and Cereulide in Food-Borne Illness Recent Publications in this Series: SHAHEEN RANAD 13/2009 Liliya Euro Electron and Proton Transfer in NADH:Ubiquinone Oxidoreductase (Complex I) from Escherichia coli 14/2009 Dario Greco Gene Expression: From Microarrays to Functional Genomics 15/2009 Elena Gorbikova Oxygen Reduction and Proton Translocation By Cytochrome c Oxidase Bacillus cereus 16/2009 Mari J. Lehtonen Rhizoctonia Solani as a Potato Pathogen - Variation of Isolates in Finland and Host Response 17/2009 Kristiina Kaste Transcription Factors ∆FosB and CREB in Drug Addiction: Studies in Models of Alcohol Preference and Chronic Nicotine Exposure Illness in Food-Borne and Cereulide Spores 18/2009 Mari K. 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Leo Faculty of Agriculture and Forestry Structural and Functional Studies on Trimeric Autotransporters 33/2009 Päivi Uutela University of Helsinki Liquid Chromatography-Tandem Mass Spectrometry in Studies of Neurotransmitters and Their Metabolites in the Brain Helsinki 2009 ISSN 1795-7079 ISBN 978-952-10-5846-2 Dissertationes bioscientiarum molecularium Universitatis Helsingiensis in Viikki 34/2009 34/2009 Bacillus cereus spores and cereulide in food-borne illness Ranad Shaheen Department of Applied Chemistry and Microbiology Division of Microbiology Faculty of Agriculture and Forestry University of Helsinki Academic Dissertation in Microbiology To be presented, with the permission of the Faculty of Agriculture and Forestry of the University of Helsinki, for public criticism in Auditorium 2402, at Viikki Biocenter 3, Viikinkaari 1, on December 18th, 2009 at 12 o’clock noon. Helsinki 2009 Supervisor: Prof. Dr. Mirja S. Salkinoja-Salonen Division of Microbiology Department of Applied Chemistry and Microbiology University of Helsinki, Finland Cosupervisor: Dr. Maria A. Andersson Division of Microbiology Department of Applied Chemistry and Microbiology University of Helsinki, Finland Reviewers: Doc. Dr. Maria Saarela VTT Biotechnology Espoo, Finland. Prof. Dr. Edward R.B. Moore Department of Clinical Bacteriology University of Göteborg Göteborg, Sweden Opponent: Dr. Carl E. Cerniglia, Director National Center for Toxicological Research, Food and Drug Administration (FDA) Jefferson Arkansas, USA ISBN 978-952-10-5846-2 (paperback) ISBN 978-952-10-5857-8 (PDF) ISSN 1795-7079 Yliopistopaino Helsinki, Finland 2009 The Cover image: Field emission scanning electron micrograph (FESEM) taken using the instrument Hitachi S-4800 (Tokyo, Japan) of spores of the Bacillus cereus strain UM 98 isolated from milk of dairy silo tank. For the microscopy, the spores were fi xed with phosphate-buffered glutaraldehyde, dehydrated in an ethanol series and dried in hexamethyldisilazane. In the fi gure, the spores adhered to steel surface. The image was taken by Ranad Shaheen. To my beloved Parents and Family Table of Contents LIST OF FIGURES LIST OF TABLES LIST OF ORIGINAL PUBLICATIONS THE AUTHOR’S CONTRIBUTION ABBREVIATIONS DEFINITIONS ABSTRACT 1. REVIEW OF THE LITERATURE ..........................................................................1 1.1. Taxonomic view on B. cereus .............................................................................1 1.1.1. B. cereus group .........................................................................................1 1.1.2. The species B. cereus (sensu stricto) ........................................................1 1.2. B. cereus spore ....................................................................................................2 1.2.1. Ultrastructure of the Bacillus spore ..........................................................2 1.2.2. Properties of the Bacillus spores and those of B. cereus spores ..............3 1.2.3. Sporulation and germination of Bacillus spores ......................................5 1.3. Diversity of B. cereus .........................................................................................6 1.4. Interactions of B. cereus with living and nonliving environment .......................7 1.4.1. Adhesion of B. cereus to nonliving and living surfaces ............................7 1.4.2. Antagonistic property of B. cereus towards microorganisms ...................8 1.4.3. B. cereus as a probiotic .............................................................................8 1.5. Human virulence ...............................................................................................10 1.5.1. B. cereus as a human pathogen ...............................................................10 1.5.2. Cereulide, the emetic toxin of B. cereus .................................................11 1.5.2.1. Structure and properties of cereulide .........................................11 1.5.2.2. Toxic mechanisms and detection of cereulide ............................13 1.5.2.3. Toxicity of cereulide in whole animals ......................................14 2. AIMS OF THE STUDY ..........................................................................................20 3. MATERIALS AND METHODS ............................................................................21 4. RESULTS AND DISCUSSION ..............................................................................22 4.1. Strategies for isolating and identifying cereulide producers among strains and isolates of B. cereus ............................................................22 4.2. Cereulide productivity of B. cereus isolates of geographically and temporally diverse origins ................................................................................23 4.3. Occurrence of cereulide producing B. cereus in some environmental niches ..24 4.4. Potential for cereulide production in selected infant food formulas .................25 4.5. Strategies of B. cereus spores for persistence in the dairy process environment ......................................................................33 4.6. Are the cereulide producing B. cereus successful in competition with the non producing B. cereus in food? .......................................................34 5. SUMMARY AND CONCLUSIONS ......................................................................36 6. ACKNOWLEDGEMENTS ....................................................................................38 7. REFERENCES ........................................................................................................39 LIST OF FIGURES Figure 1. Transmission electron micrograph of a sporulating culture of the emetic B. cereus strain F4810/72. The culture was grown for 10 days on tryptic soy agar at 22 °C .................................................................................2 Figure 2. Structure of cereulide. ..................................................................................11 Figure 3. Gene cluster for the biosynthesis of cereulide in B. cereus AH187 (F4810/72) carrying the plasmid pCER270. ................................................19 Figure 4. A fl ow chart for distinguishing the cereulide producing B. cereus isolates from the nonproducers in foods or in environmental samples. ....................22 Figure 5. Intraspecies antagonistic activity of B. cereus. For testing the sensitivity, the target strain of B. cereus was plated as a lawn and the potential antagonists were spotted on the lawn. In this fi gure the lawn of B. cereus ML60 was spotted with antagonistic strains F4810/72, B208 and ML30 and the nonantagonistic strain RIVMBC0067. The clear zone around the spotted F4810/72, B208 and ML30 show these were antagonistic towards ML60. .35 LIST OF TABLES Table 1. Surface characteristics reported for B. cereus spores. The studied strains were the type strain ATCC 14579T, one dairy isolate, 3 isolates from patients with diarrhoeal food poisoning and 2 strains isolated from the environment (Tauveron
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