Microbiology, 2021

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592 14 • Antimicrobial Drugs Mechanisms for Drug Resistance There are several common mechanisms for drug resistance, which are summarized in Figure 14.18. These mechanisms include enzymatic modification of the drug, modification of the antimicrobial target, and prevention of drug penetration or accumulation. Figure 14.18 There are multiple strategies that microbes use to develop resistance to antimicrobial drugs. (Not shown: target overproduction, target mimicry, and enzymatic bypass). (credit: modification of work by Gerard D Wright) Drug Modification or Inactivation Resistance genes may code for enzymes that chemically modify an antimicrobial, thereby inactivating it, or destroy an antimicrobial through hydrolysis. Resistance to many types of antimicrobials occurs through this mechanism. For example, aminoglycoside resistance can occur through enzymatic transfer of chemical groups to the drug molecule, impairing the binding of the drug to its bacterial target. For β-lactams, bacterial resistance can involve the enzymatic hydrolysis of the β-lactam bond within the β-lactam ring of the drug molecule. Once the β-lactam bond is broken, the drug loses its antibacterial activity. This mechanism of resistance is mediated by β-lactamases, which are the most common mechanism of β-lactam resistance. Inactivation of rifampin commonly occurs through glycosylation, phosphorylation, or adenosine diphosphate (ADP) ribosylation, and resistance to macrolides and lincosamides can also occur due to enzymatic inactivation of the drug or modification. Prevention of Cellular Uptake or Efflux Microbes may develop resistance mechanisms that involve inhibiting the accumulation of an antimicrobial drug, which then prevents the drug from reaching its cellular target. This strategy is common among gramnegative pathogens and can involve changes in outer membrane lipid composition, porin channel selectivity, and/or porin channel concentrations. For example, a common mechanism of carbapenem resistance among Pseudomonas aeruginosa is to decrease the amount of its OprD porin, which is the primary portal of entry for carbapenems through the outer membrane of this pathogen. Additionally, many gram-positive and gram- Access for free at openstax.org.
14.5 • Drug Resistance 593 negative pathogenic bacteria produce efflux pumps that actively transport an antimicrobial drug out of the cell and prevent the accumulation of drug to a level that would be antibacterial. For example, resistance to β- lactams, tetracyclines, and fluoroquinolones commonly occurs through active efflux out of the cell, and it is rather common for a single efflux pump to have the ability to translocate multiple types of antimicrobials. Target Modification Because antimicrobial drugs have very specific targets, structural changes to those targets can prevent drug binding, rendering the drug ineffective. Through spontaneous mutations in the genes encoding antibacterial drug targets, bacteria have an evolutionary advantage that allows them to develop resistance to drugs. This mechanism of resistance development is quite common. Genetic changes impacting the active site of penicillin-binding proteins (PBPs) can inhibit the binding of β-lactam drugs and provide resistance to multiple drugs within this class. This mechanism is very common among strains of Streptococcus pneumoniae, which alter their own PBPs through genetic mechanisms. In contrast, strains of Staphylococcus aureus develop resistance to methicillin (MRSA) through the acquisition of a new low-affinity PBP, rather than structurally alter their existing PBPs. Not only does this new low-affinity PBP provide resistance to methicillin but it provides resistance to virtually all β-lactam drugs, with the exception of the newer fifth-generation cephalosporins designed specifically to kill MRSA. Other examples of this resistance strategy include alterations in • ribosome subunits, providing resistance to macrolides, tetracyclines, and aminoglycosides; • lipopolysaccharide (LPS) structure, providing resistance to polymyxins; • RNA polymerase, providing resistance to rifampin; • DNA gyrase, providing resistance to fluoroquinolones; • metabolic enzymes, providing resistance to sulfa drugs, sulfones, and trimethoprim; and • peptidoglycan subunit peptide chains, providing resistance to glycopeptides. Target Overproduction or Enzymatic Bypass When an antimicrobial drug functions as an antimetabolite, targeting a specific enzyme to inhibit its activity, there are additional ways that microbial resistance may occur. First, the microbe may overproduce the target enzyme such that there is a sufficient amount of antimicrobial-free enzyme to carry out the proper enzymatic reaction. Second, the bacterial cell may develop a bypass that circumvents the need for the functional target enzyme. Both of these strategies have been found as mechanisms of sulfonamide resistance. Vancomycin resistance among S. aureus has been shown to involve the decreased cross-linkage of peptide chains in the bacterial cell wall, which provides an increase in targets for vancomycin to bind to in the outer cell wall. Increased binding of vancomycin in the outer cell wall provides a blockage that prevents free drug molecules from penetrating to where they can block new cell wall synthesis. Target Mimicry A recently discovered mechanism of resistance called target mimicry involves the production of proteins that prevent drugs from binding to their bacterial cellular targets. For example, fluoroquinolone resistance by Mycobacterium tuberculosis can involve the production of a protein that resembles DNA. This protein is called MfpA (Mycobacterium fluoroquinolone resistance protein A). The mimicry of DNA by MfpA results in DNA gyrase binding to MfpA, preventing the binding of fluoroquinolones to DNA gyrase. CHECK YOUR UNDERSTANDING • List several mechanisms for drug resistance.
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Microbiology SENIOR CONTRIBUTING AU
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OPENSTAX OpenStax provides free, pe
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Contents Preface 1 CHAPTER 1 An Inv
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9.2 Oxygen Requirements for Microbi
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18.1 Overview of Specific Adaptive
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Appendix C Metabolic Pathways 1155
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Preface 1 PREFACE Welcome to Microb
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Preface 3 that often confer critica
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Preface 5 further. Our features inc
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Preface 7 and science, and pursues
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CHAPTER 1 An Invisible World Figure
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1.1 • What Our Ancestors Knew 11
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1.2 • A Systematic Approach 17 Th
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1.2 • A Systematic Approach 19 Fi
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1.2 • A Systematic Approach 21 Ta
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1.3 • Types of Microorganisms 23
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1 • Summary 31 SUMMARY 1.1 What O
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1 • Review Questions 33 17. The p
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CHAPTER 2 How We See the Invisible
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2.1 • The Properties of Light 37
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2.1 • The Properties of Light 39
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2.2 • Peering Into the Invisible
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2.3 • Instruments of Microscopy 4
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2.4 • Staining Microscopic Specim
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2 • Review Questions 71 REVIEW QU
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CHAPTER 3 The Cell Figure 3.1 Micro
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3.1 • Spontaneous Generation 75 F
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3.2 • Foundations of Modern Cell
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3.3 • Unique Characteristics of P
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3.4 • Unique Characteristics of E
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3 • Summary 125 SUMMARY 3.1 Spont
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3 • Review Questions 127 3. Which
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3 • Review Questions 129 Critical
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CHAPTER 4 Prokaryotic Diversity Fig
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4.1 • Prokaryote Habitats, Relati
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4.2 • Proteobacteria 139 for or s
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4.2 • Proteobacteria 141 Class Al
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4.2 • Proteobacteria 143 Class Be
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4.2 • Proteobacteria 145 Figure 4
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4.2 • Proteobacteria 147 Class Ga
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4.3 • Nonproteobacteria Gram-Nega
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4.4 • Gram-Positive Bacteria 157
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4.6 • Archaea 167 The class Therm
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4.6 • Archaea 169 thus is a livin
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4 • Summary 171 SUMMARY 4.1 Proka
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4 • Review Questions 173 REVIEW Q
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4 • Review Questions 175 39. Expl
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CHAPTER 5 The Eukaryotes of Microbi
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5.1 • Unicellular Eukaryotic Para
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Figure 5.7 5.1 • Unicellular Euka
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5.2 • Parasitic Helminths 193 hel
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5.2 • Parasitic Helminths 195 Fig
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5.2 • Parasitic Helminths 197 Fig
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5.2 • Parasitic Helminths 199 MIC
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5.3 • Fungi 201 Figure 5.25 Multi
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5.3 • Fungi 203 Figure 5.27 zygos
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5.3 • Fungi 205 diploid stages (F
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5.3 • Fungi 207 Figure 5.32 prese
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5.3 • Fungi 209 MICRO CONNECTIONS
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5.4 • Algae 211 and other photosy
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5.5 • Lichens 213 Figure 5.37 Chl
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5.5 • Lichens 215 marina. (b) Thi
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5 • Review Questions 217 REVIEW Q
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CHAPTER 6 Acellular Pathogens Figur
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6.1 • Viruses 221 assembly of vir
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6.1 • Viruses 223 Viral Structure
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6.1 • Viruses 225 Figure 6.5 (a)
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6.1 • Viruses 227 LINK TO LEARNIN
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6.2 • The Viral Life Cycle 229 on
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6.2 • The Viral Life Cycle 231 Fi
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6.2 • The Viral Life Cycle 237 by
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6.2 • The Viral Life Cycle 239 Ca
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6.3 • Isolation, Culture, and Ide
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6.4 • Viroids, Virusoids, and Pri
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6.4 • Viroids, Virusoids, and Pri
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6 • Summary 253 SUMMARY 6.1 Virus
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6 • Review Questions 255 18. A/an
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CHAPTER 7 Microbial Biochemistry Fi
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7.1 • Organic Molecules 259 Figur
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7.1 • Organic Molecules 261 Figur
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7.2 • Carbohydrates 263 and the m
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7.2 • Carbohydrates 265 Figure 7.
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7.3 • Lipids 267 Figure 7.11 Star
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7.3 • Lipids 269 Figure 7.13 This
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7.3 • Lipids 271 Figure 7.15 Five
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7.4 • Proteins 273 Figure 7.17 Am
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7.4 • Proteins 275 Figure 7.19 Re
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7.4 • Proteins 277 Figure 7.23 Pr
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7.5 • Using Biochemistry to Ident
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7.5 • Using Biochemistry to Ident
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7 • Review Questions 283 as hormo
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7 • Review Questions 285 19. A tr
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CHAPTER 8 Microbial Metabolism Figu
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8.1 • Energy, Matter, and Enzymes
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8.2 • Catabolism of Carbohydrates
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8.3 • Cellular Respiration 301 8.
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8.3 • Cellular Respiration 303 Fi
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8.4 • Fermentation 305 If respira
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8.4 • Fermentation 307 Common Fer
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8.5 • Catabolism of Lipids and Pr
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8.6 • Photosynthesis 311 independ
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8.6 • Photosynthesis 313 Oxygenic
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8.7 • Biogeochemical Cycles 315 L
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8.7 • Biogeochemical Cycles 317 N
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8.7 • Biogeochemical Cycles 319 F
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8.7 • Biogeochemical Cycles 321 F
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8 • Summary 323 oxygen as the fin
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8 • Review Questions 325 4. To wh
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8 • Review Questions 327 Matching
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CHAPTER 9 Microbial Growth Figure 9
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9.1 • How Microbes Grow 331 and a
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9.1 • How Microbes Grow 333 The G
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9.1 • How Microbes Grow 335 Susta
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9.1 • How Microbes Grow 337 chang
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9.1 • How Microbes Grow 339 Figur
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9.1 • How Microbes Grow 341 trans
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9.1 • How Microbes Grow 343 micro
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9.2 • Oxygen Requirements for Mic
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9.3 • The Effects of pH on Microb
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9.4 • Temperature and Microbial G
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9.4 • Temperature and Microbial G
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9.5 • Other Environmental Conditi
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9.6 • Media Used for Bacterial Gr
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9 • Summary 361 SUMMARY 9.1 How M
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9 • Review Questions 363 7. Filam
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9 • Review Questions 365 Matching
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9 • Review Questions 367 Critical
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CHAPTER 10 Biochemistry of the Geno
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10.1 • Using Microbiology to Disc
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10.2 • Structure and Function of
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10 • Summary 401 SUMMARY 10.1 Usi
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10 • Review Questions 403 3. Whic
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10 • Review Questions 405 Short A
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CHAPTER 11 Mechanisms of Microbial
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11.1 • The Functions of Genetic M
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11.2 • DNA Replication 411 Figure
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11.2 • DNA Replication 413 Figure
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11.2 • DNA Replication 415 strand
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11.2 • DNA Replication 417 telome
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11.3 • RNA Transcription 419 Figu
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11.3 • RNA Transcription 421 the
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11.4 • Protein Synthesis (Transla
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11.5 • Mutations 429 Effects of M
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11.5 • Mutations 431 In recent ye
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11.5 • Mutations 433 Figure 11.20
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11.5 • Mutations 435 formation of
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11.6 • How Asexual Prokaryotes Ac
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11.7 • Gene Regulation: Operon Th
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11 • Summary 457 SUMMARY 11.1 The
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11 • Summary 459 undamaged DNA st
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11 • Review Questions 461 11. Whi
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11 • Review Questions 463 46. ___
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71. The following figure is from Mo
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CHAPTER 12 Modern Applications of M
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12.1 • Microbes and the Tools of
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12.2 • Visualizing and Characteri
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12.3 • Whole Genome Methods and P
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12.3 • Whole Genome Methods and P
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12.4 • Gene Therapy 499 Figure 12
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12.4 • Gene Therapy 501 overall w
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12 • Summary 503 SUMMARY 12.1 Mic
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12 • Review Questions 505 11. The
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CHAPTER 13 Control of Microbial Gro
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13.1 • Controlling Microbial Grow
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13.2 • Using Physical Methods to
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13.3 • Using Chemicals to Control
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15.4 • Virulence Factors of Eukar
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15.4 • Virulence Factors of Eukar
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15 • Review Questions 647 protect
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15 • Review Questions 649 Critica
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CHAPTER 16 Disease and Epidemiology
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16.1 • The Language of Epidemiolo
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16.1 • The Language of Epidemiolo
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16.2 • Tracking Infectious Diseas
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16.3 • Modes of Disease Transmiss
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16.4 • Global Public Health 673 C
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16.4 • Global Public Health 675 S
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16 • Summary 677 SUMMARY 16.1 The
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16 • Review Questions 679 Matchin
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16 • Review Questions 681 20. Wha
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CHAPTER 17 Innate Nonspecific Host
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17.1 • Physical Defenses 685 Over
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17.1 • Physical Defenses 687 know
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17.1 • Physical Defenses 689 Figu
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17.2 • Chemical Defenses 691 Phys
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17.2 • Chemical Defenses 693 sali
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17.2 • Chemical Defenses 695 prod
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17.2 • Chemical Defenses 697 Figu
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17.2 • Chemical Defenses 699 Clin
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17.3 • Cellular Defenses 701 Hema
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17.3 • Cellular Defenses 703 stra
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17.3 • Cellular Defenses 705 Clin
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17.3 • Cellular Defenses 707 Figu
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17.4 • Pathogen Recognition and P
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17.5 • Inflammation and Fever 713
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17 • Summary 719 SUMMARY 17.1 Phy
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17 • Review Questions 721 8. Hist
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17 • Review Questions 723 Short A
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CHAPTER 18 Adaptive Specific Host D
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18.1 • Overview of Specific Adapt
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18.2 • Major Histocompatibility C
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18.3 • T Lymphocytes and Cellular
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18.4 • B Lymphocytes and Humoral
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18.4 • B Lymphocytes and Humoral
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18.5 • Vaccines 751 pathogen—bu
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18.5 • Vaccines 753 Eye on Ethics
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18.5 • Vaccines 755 for Disease C
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18.5 • Vaccines 757 Classes of Va
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18 • Summary 759 SUMMARY 18.1 Ove
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18 • Review Questions 761 5. MHC
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18 • Review Questions 763 20. Mat
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CHAPTER 19 Diseases of the Immune S
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19.1 • Hypersensitivities 767 Fig
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19.1 • Hypersensitivities 769 bin
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19.1 • Hypersensitivities 771 mec
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19.1 • Hypersensitivities 775 CHE
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19.1 • Hypersensitivities 777 Cli
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19.1 • Hypersensitivities 779 MIC
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19.2 • Autoimmune Disorders 783 t
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19.2 • Autoimmune Disorders 785 F
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19.2 • Autoimmune Disorders 787 M
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19.2 • Autoimmune Disorders 789 F
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19.3 • Organ Transplantation and
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19.4 • Immunodeficiency 793 and m
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19.4 • Immunodeficiency 795 CHECK
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19.5 • Cancer Immunobiology and I
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19 • Summary 799 SUMMARY 19.1 Hyp
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19 • Review Questions 801 13. All
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CHAPTER 20 Laboratory Analysis of t
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20.1 • Polyclonal and Monoclonal
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20.2 • Detecting Antigen-Antibody
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20.3 • Agglutination Assays 823 a
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20.3 • Agglutination Assays 825 F
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20.3 • Agglutination Assays 827 e
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20.3 • Agglutination Assays 829 s
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20.4 • EIAs and ELISAs 831 Mechan
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20.4 • EIAs and ELISAs 833 Figure
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20.4 • EIAs and ELISAs 837 • Wh
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20.5 • Fluorescent Antibody Techn
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20 • Review Questions 849 20.4 EI
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20 • Review Questions 851 15. Sup
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CHAPTER 21 Skin and Eye Infections
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21.1 • Anatomy and Normal Microbi
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21.3 • Viral Infections of the Sk
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21.4 • Mycoses of the Skin 881 se
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21.4 • Mycoses of the Skin 883 fr
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21.4 • Mycoses of the Skin 885 Fi
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21.5 • Protozoan and Helminthic I
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21.5 • Protozoan and Helminthic I
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21 • Summary 891 SUMMARY 21.1 Ana
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CHAPTER 22 Respiratory System Infec
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22.4 • Respiratory Mycoses 931 ha
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22.4 • Respiratory Mycoses 933 Fi
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22.4 • Respiratory Mycoses 935 re
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Figure 22.29 22.4 • Respiratory M
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22 • Review Questions 939 200 vir
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CHAPTER 23 Urogenital System Infect
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23.5 • Fungal Infections of the R
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23 • Summary 975 SUMMARY 23.1 Ana
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CHAPTER 24 Digestive System Infecti
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24.2 • Microbial Diseases of the
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24.4 • Viral Infections of the Ga
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24.6 • Helminthic Infections of t
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24 • Summary 1033 SUMMARY 24.1 An
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CHAPTER 25 Circulatory and Lymphati
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25.1 • Anatomy of the Circulatory
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25.1 • Anatomy of the Circulatory
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25.3 • Viral Infections of the Ci
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Figure 25.26 25.3 • Viral Infecti
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25.4 • Parasitic Infections of th
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25 • Review Questions 1089 • To
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25 • Review Questions 1091 Critic
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CHAPTER 26 Nervous System Infection
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26.1 • Anatomy of the Nervous Sys
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26.1 • Anatomy of the Nervous Sys
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26.2 • Bacterial Diseases of the
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26.3 • Acellular Diseases of the
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Figure 26.19 26.3 • Acellular Dis
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26.4 • Fungal and Parasitic Disea
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26 • Review Questions 1133 are fa
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26 • Review Questions 1135 Matchi
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26 • Review Questions 1137 59. Th
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A • Fundamentals of Physics and C
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B • Mathematical Basics 1149 APPE
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B • Mathematical Basics 1151 Mult
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B • Mathematical Basics 1153 The
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C • Metabolic Pathways 1155 APPEN
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C • Metabolic Pathways 1157 Entne
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C • Metabolic Pathways 1159 Figur
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C • Metabolic Pathways 1161 Elect
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APPENDIX D Taxonomy of Clinically R
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D • Taxonomy of Clinically Releva
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E • Glossary 1177 APPENDIX E Glos
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E • Glossary 1179 destruction by
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E • Glossary 1181 pollutants from
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E • Glossary 1183 chromosome disc
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E • Glossary 1185 cysts are forme
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E • Glossary 1187 occurring ectop
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E • Glossary 1189 molecules of DN
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E • Glossary 1191 glycoprotein co
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E • Glossary 1195 discontinuously
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E • Glossary 1197 infection cause
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E • Glossary 1199 target cells by
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E • Glossary 1201 intact ribosome
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E • Glossary 1205 complexes forme
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E • Glossary 1207 antibody is fir
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E • Glossary 1209 the number of c
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E • Glossary 1211 the reciprocal
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E • Glossary 1213 vector animal (
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1215 ANSWER KEY Chapter 1 1. D 2. D
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1217 B 18. A, C, B 19. peristalsis
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Index 1219 INDEX Symbols +ssRNA 237
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Index 1221 antigen-presenting cells
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Index 1223 Bordetella 142 Bordetell
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Index 1225 chromosomes 372, 394 chr
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Index 1227 de Duve 113 dead zone 35
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Index 1229 Enteroinvasive E. coli (
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Index 1231 Gardnerella vaginalis 47
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Index 1233 shunt 298 Hfr cell 443 H
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Index 1235 iron lungs 1118 iron-sul
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Index 1237 maturation 230 mature na
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Index 1239 Health 501 National Noti
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Index 1241 patterns (PAMPs) 710 pat
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Index 1243 primary lymphoid tissue
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Index 1245 596, 1048, 1051, 1051 ri
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Index 1247 sterilant 511, 541 steri
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Index 1249 toxigenicity 633 toxin 6
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Index 1251 water activity 522 water
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Microbiology, 2021

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