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Plasmids and the Virulence of Proteus Mirabilis a Thesis Submitted to the University of London for the Degree of Doctor of Philo

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Plasmids and the Virulence of Proteus Mirabilis a Thesis Submitted to the University of London for the Degree of Doctor of Philo PLASMIDS AND THE VIRULENCE OF PROTEUS MIRABILIS A THESIS SUBMITTED TO THE UNIVERSITY OF LONDON FOR THE DEGREE OF DOCTOR OF PHILOSOPHY BY ALI AKBAR MOHAMMADI DEPARTMENT OF BIOLOGY UNIVERSITY COLLEGE LONDON LONDON WC1E 1 ProQuest Number: 10608873 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a com plete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 10608873 Published by ProQuest LLC(2017). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C ode Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 Acknowledgment My sincere thanks are due to my supervisors Professor R. J. Rowbury and Doctor D. G. Smith, for their valuable advice, help and patience throughout this work. I am also grateful to Razi Institute and the Ministry of Culture and Higher Education for their financial support. Thanks are also due to Dr. Gruneberg and Karen Jones of Microbiology Department UCH for providing me with the urinary strains of P. mirabilis. I am thankful to Margaret Goodson for her advice and John Mackey for his Photographic help. 2 ABSTRACT: The effects of large plasmids on different virulence characteristics of Proteus mirabilis strains mostly from clinical origin were studied. Moreover the inhibitory effect of urea and its derivatives on the swarming property of the strains was investigated. A. All strains were screened for plasmid detection, antibiotic resistance and swarming ability. B. Three multiresistant plasmid—carrying strains (PM5, P49 and P991) were cured and two transconjugants (G9pPM5 and G9pP49) were obtained by conjugation between two p+ donors (PM5 and P49) and one p- recipient <G9). C. By comparing the virulence properties of cured and transconjugant strains with their parental isolates it was f ound t hat: 1. Plasmids confer resistance to P. mirabilis strains against one or more antibiotics. 2. The presence of most plasmids reduces the swarming ability of the strains. 3. Plasmids affect the motility and flagellation of P. mirabilis strains. 4. Plasmids enhance the adherence property of their host strains to inert surfaces and uroepithelial cells as well as aut oagglut i nat i on. 5. Plasmids increase the hydrophobicity of P. mirabilis strains. 6. The presence of plasmids reduced the growth rate of the strains. This effect was more apparent in iron-restricted medium. 7. Plasmids reduced the growth rate of their host strains in the presence of detergent <SDS). 8. The presence of plasmids reduced the survival of P. mirabilis strains in human and rabbit serum. 9. Plasmids decreased the survival of the strains in aquatic systems. 10. Plasmids reduced the production of urease and increasedsome others such as haemolysin and protease. D. Urea and some of its relatives inhibited the swarming property of P. mirabilis strains and this effect was concentration dependent. 4 Index. Content. Page No. Abstract. 2 Acknowledgment. 4 Index. 5 List of Tables. 15 List of Figures. 20 I. Introduct ion:. 21 I. 1. Identification. 25 I. 2. Typing of P. mirabilis. 26 I. 3. Swarming. 26 I. 3. a. Types of swarming. 28 I. 3. b. Mechanism of swarming. 29 I. 3. c. Swarming inhibitors. 30 I. 4. Urea. 32 1. 4. a. Properties of urea. 33 I. 4. b. Urea as osmotic agent. 33 I. 4. c. Urea as a nitrogen source. 34 I. 4. d. Urea as an antibacterial agent. 35 1.5. Plasmids. 37 I. 5. a. Conjugation. 37 I. 5. b. Mechanism of conjugation. 38 I. 5. c. Sex factor mediated transfer. 39 I. 5. d. Transduction. 39 I. 5. e. Transformation. 40 I. 5. f. Curing of plasmid DNA. 41 I. 5. g. Replication of plasmid DNA: 41 1. 5. h. Mechanism of replication. 42 1. Crains* model 43 2. Gilbert and Dressier model. 43 3. Rolling circle model. 43 1.6. Cell envelope structure in Gram—negative bacteria. 44 I. 6. a. Cytoplasmic membrane. 44 1.6. b. Cell wall. 47 I. 6. b. 1. Peptidoglycan layer. 47 I. 6. b. 2. Peri pi asm! c region. 48 I. 6. b. 3. Outer membrane. 50 I. 6. b. 3. a. Lipopolysaccharide. 51 I. 6. b. 3. b. Lipoproteins. 54 I. 6. b. 3. c. Major proteins. 55 I. 6. b. 3. d. OmpA. 55 I. 6. b. 3. e. Porin proteins. 56 I. 6. b. 3. f. Minor proteins. 56 I. 6. b. 3. g. Phospholipids. 57 I. 6. b. 3. h. Properties of the outer membrane: 57 1. Resistance to enzymes. 58 2. Resistance to hydrophobic molecules. 58 3. Resistance to polycatlonic agents. 59 1. 7. Antibiotic resistances. 61 6 I. 7. a. Antibioics. 61 I. 7. b. Antibiotic resistance in bacteria. 62 I. 7. c. Mechanisms of antibiotic resistance. 64 I. 7. c. 1. Chromosomal resistance. 64 I. 7. c. 2. Plasmids and antibiotic resistance. 65 1.7. d. Resistance to antibacterials acting on the cell wall. 69 I. 7. d. 1. Chromosomal resistance. 69 I. 7. d. 2. R-plasmid encoded resistance. 70 1. 7. e. Resistance to ant ibacterials acting on the cytoplasmic membrane. 71 I. 7. e. 1. Chromosomal resistance. 72 I. 7. e. 2. Mutational resistance. 72 1.7.f. Resistance to agents acting on nucleic acids. 73 I. 7. f. 1. Chromosomal resistance. 74 1.7.g. Resistance to antibacterials acting on folic acid metabolism. 75 I. 7. g. 1. Chromosomal resistance. 77 I. 7. g. 2. Plasmid resistance. 78 1.7. h. Resistance to antibacterial agents interfering in protein synthesis: 81 1.7. h. 1. Chloramphenicol 81 1.7. h. 1. a. Chromosomal resistance. 81 I. 7. h. 1. b. Plasmid resistance. 82 I. 7. h. 2. Tetracyclines. 83 7 t I. 7. h. 2. a. Chromosomal resistance. 84 I. 7. h. 2. b. Plasmid resistance. 85 1. 7. i. Resistance to aminoglycosides. 86 1. 7. i. 1. Chromosomal resistance. 87 I. 7. i. 2. Plasmid resistance. 88 I. 7. j. Resistance to nitrofurans. 89 I. 7. j. 1. Chromosomal resistance. 92 I. 7. j. 2. Plasmid resistance. 92 1.8. Bactrial attachment. 94 1.8. a. Pili. 95 I. 8. b. Types of pili. 95 1.8.c. Bacterial adherence as a virulence factor in urinary tract infections. 97 I. 8. d. Adherence of P. mirabilis strains to surfaces. 99 I. 8. e. The effects of plasmids on bacterial adherence. 102 I. 9. Bacterial hydrophobicity. 104 1.9. a. Plasmids and bacterial hydrophobicity. 106 1.10. Factors influencing bacterial growth. 106 I. 10. a. The significance of iron in bacterial growth. 109 I. 10. b. Growth and nutritional requirements of P. mirabilis. 110 1.10.c. Effects of plasmids on the growth and nutrition of bacteria. 112 1.10.c. The effect of anionic detergents on bacterial growth. 114 8 I. 11. Plasmids and the motility of P. mirabilis. 115 I. 11. a. Proteus flagella. 116 I. 12. Survival of P. mirabilis in different envi ronment s. 117 1. 12. a. Survival in serum. 117 I. 12. a. 1. Plasmids and serum resistance. 120 I. 12. b. Survival and growth of bacteria in sewage. 122 1.12.c. Survival of bacteria in effluent. 124 I. 12. d. Survival of bacteria in river water. 127 1.12.e. Effect of plasmids on the survival of bacteria in aquatic systems. 128 I. 13. Enzymes in P. mirabilis. 130 I. 13. a. Urease. 130 1. 13. a. 1. Plasmids and urease production. 133 I. 13. b. Haemolysin. 134 1. 13. b. 1. Plasmids and haemolysin activity of bacteria. 136 I. 13. c. Protease. 137 I.14. The aims of this work 140 II. Materials and Methods: 141 II. 1. Bacterial strains. 142 II. 2. Media. 142 II. 3. Methods. 143 II. 3. a. Identification of Proteus strains. 143 II. 3. b. Plasmid detection. 144 9 II. 3. c. Sizing of plasmid DNA. 145 II. 3. d. Antibiotic resistance tests. 146 II. 3. e. Conjugation. 146 11.3.f. Curing the plasmid-bearing strains of P. mirabilis. 150 II. 3. g. Swarming test. 151 II. 3. h. Attachment to glass beads. 151 II. 3. i. Attachment to glass cover slips. 152 11.3.j. Adherence to epithelial cells. 152 II. 3. k. Hydrophobic!ty. 153 II. 3. 1. Measurement of the growth. 154 II. 3. 1. 1. Growth in nutrient broth. 154 II. 3. 1. 2. Growth rate in iron depleted nutrient broth. 154 II. 3. 1. 3. Growth in the presence of detergents. 155 II. m. Measurement of viable count. 155 II. 3. n. Measurment of optical density. 155 II. 3. o. Clump formation test. 156 II. 3. p. Motility test. 156 11.3. q. Effects of detegent on clump formation and motility. 156 II. 3. r. Inhibition of swarming. 156 11.3. s. The inhibitory effect of urea and its analogues on swarming. 157 11.3.t. Checking efffects of urea on flagellation. 158 II. 3. u. Checking urea effect on long cell formation. 159 10 II. 3. v. Haemolysin assay. 160 II. 3. w. Protease assay. 160 II. 3. x. Lecithlnase assay. 161 II. 3. y. Amylase secretion assay. 161 II. 3. z. DNase production assay. 161 11.3. a* . Qualitative test for production of urease. 162 II. 3. b* . Comparative assay for urease secretion on solid medium. 162 11.3.c*. Production of urease in liquid medium. 162 II.
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