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TH I R D ED ITI O N
Microbiology
An Evolving Science
Joan L. Slonczewski
Kenyon College
John W. Foster
University of South Alabama
Appendices 1 and 2 by
Kathy M. Gillen
Kenyon College
0-393-91929-5
W. W. Norton & Company, Inc., 500 Fifth Avenue, New York, N.Y. 10110
wwnorton.com
W. W. Norton & Company Ltd., Castle House, 75/76 Wells Street, London W1T 3QT
123456789
Dedication
We dedicate this Third Edition to the memory of Lynn Margulis
(1938–2011) and Carl Woese (1928–2012), who forever
changed our understanding of evolutionary biology. Margulis
hypothesized that modern-day eukaryotic cells evolved from
symbiotic relationships with bacteria; and Woese used the
evidence of gene sequences to propose the now accepted
paradigm of three, continually evolving, domains of life: Bacteria,
Archaea, and Eukarya. The world will long miss the deep intellect
and larger-than-life personalities of these two scientists. We, the
authors, were profoundly influenced by their thinking and deeply
moved by their passing.
BRIEF CONTENTS
VI
CONTENTS
eTopic Contents!xix
Preface!xxi
About the Authors!xxxiii
PART 1
The Microbial Cell 2
AN INTERVIEW WITH RICHARD LENSKI:
Evolution in the Lab
CHAPTER 1
Microbial Life: Origin and Discovery........................................................................ 5
1.1 From Germ to Genome: What Is a Microbe?!7
1.2 Microbes Shape Human History!10
Special Topic 1.1: How Did Life Originate?!18
1.3 Medical Microbiology!20
1.4 Microbial Ecology!26
1.5 The Microbial Family Tree!29
1.6 Cell Biology and the DNA Revolution!32
CHAPTER 2
Observing the Microbial Cell ....................................................................................41
2.1 Observing Microbes!42
2.2 Optics and Properties of Light!46
2.3 Bright-Field Microscopy!51
2.4 Fluorescence Microscopy!58
2.5 Dark-Field and Phase-Contrast Microscopy!63
2.6 Electron Microscopy and Tomography!66
2.7 Visualizing Molecules!73
Special Topic 2.1: Molecular “Snapshots”: Chemical Imaging!74
VII
VIII ! CONTENTS
CHAPTER 3
Cell Structure and Function .....................................................................................79
3.1 The Bacterial Cell: An Overview!81
3.2 The Cell Membrane and Transport!86
3.3 The Cell Wall and Outer Layers!91
3.4 The Nucleoid, RNA, and Protein Synthesis!101
3.5 Cell Division!104
3.6 Cell Polarity and Aging!107
Special Topic 3.1: Senior Cells Make Drug-Resistant Tuberculosis!108
3.7 Specialized Structures!112
CHAPTER 4
Bacterial Culture, Growth, and Development ................................................. 119
4.1 Microbial Nutrition!120
4.2 Nutrient Uptake!127
4.3 Culturing Bacteria!133
4.4 Counting Bacteria!136
4.5 The Growth Cycle!139
4.6 Biofilms!145
Special Topic 4.1: Sharks and Biofilms Don’t Mix!146
4.7 Cell Differentiation!150
CHAPTER 5
Environmental Influences and Control of Microbial Growth ..................... 157
5.1 Environmental Limits on Growth!158
5.2 Adaptation to Temperature!160
Special Topic 5.1: It’s Raining Bacteria!163
5.3 Adaptation to Pressure!164
5.4 Water Activity and Salt!166
5.5 Adaptation to pH!167
5.6 Oxygen and Other Electron Acceptors!172
5.7 Nutrient Deprivation and Starvation!176
5.8 Physical, Chemical, and Biological Control of Microbes!178
CHAPTER 6
Viruses ........................................................................................................................... 191
6.1 What Is a Virus?!192
6.2 Virus Structure!198
6.3 Viral Genomes and Classification!203
6.4 Bacteriophage Replication!208
6.5 Animal and Plant Virus Replication!213
CONTENTS ! IX
PART 2
Genes and Genomes 234
CHAPTER 7
Genomes and Chromosomes ................................................................................ 237
7.1 DNA: The Genetic Material!238
7.2 Genome Organization!240
7.3 DNA Replication!248
7.4 Plasmids!258
7.5 Eukaryotic Chromosomes!260
7.6 DNA Sequence Analysis!263
Special Topic 7.1: Where Have All the Bees Gone? Metagenomics, Pyrosequencing, and Nature!268
CHAPTER 8
Transcription, Translation, and Bioinformatics .............................................. 275
8.1 RNA Polymerases and Sigma Factors!276
8.2 Transcription of DNA to RNA!280
8.3 Translation of RNA to Protein!286
Special Topic 8.1: Stalking the Lone Ribosome!298
8.4 Protein Modification and Folding!303
8.5 Protein Degradation: Cleaning House!304
8.6 Secretion: Protein Traffic Control!306
8.7 Bioinformatics: Mining the Genomes!312
CHAPTER 9
Gene Transfer, Mutations, and Genome Evolution ....................................... 321
9.1 The Mosaic Nature of Genomes!322
9.2 Gene Transfer!322
Special Topic 9.1: There’s a Bacterial Genome Hidden in My Fruit Fly!330
X ! CONTENTS
9.3 Recombination!338
9.4 Mutations!341
9.5 DNA Repair!348
9.6 Mobile Genetic Elements!355
9.7 Genome Evolution!358
CHAPTER 10
Molecular Regulation ............................................................................................... 365
10.1 Regulating Gene Expression!366
10.2 Paradigm of the Lactose Operon!369
10.3 Other Systems of Operon Control!376
10.4 Sigma Factor Regulation!382
10.5 Regulatory RNAs!385
10.6 DNA Rearrangements: Phase Variation by Shifty Pathogens!389
10.7 Integrated Control Circuits!391
10.8 Quorum Sensing: Chemical Conversations!397
10.9 Transcriptomics and Proteomics!401
Special Topic 10.1: Networking with Nanotubes!402
CHAPTER 11
Viral Molecular Biology ...........................................................................................409
11.1 Phage T4: The Classic Molecular Model!410
11.2 Hepatitis C: (+) Strand RNA Virus!417
11.3 Influenza Virus: (–) Strand RNA Virus!424
11.4 Human Immunodeficiency Virus (HIV): Retrovirus!432
11.5 Herpes Simplex Virus: DNA Virus!443
Special Topic 11.1: Cytomegalovirus!448
11.6 Gene Therapy with Viruses!450
CHAPTER 12
Biotechniques and Synthetic Biology ................................................................ 457
12.1 Basic Tools of Biotech: A Research Case Study!458
12.2 Genetic Analyses!458
12.3 Classic Molecular Techniques!463
12.4 Viewing the Interactions and Movements of Proteins!471
12.5 Applied Biotechnology!474
12.6 Synthetic Biology: Biology by Design!477
Special Topic 12.1: Bacteria “Learn” to Keep Time and Signal Danger!478
CONTENTS ! XI
PART 3
Metabolism and Biochemistry 488
CHAPTER 13
Energetics and Catabolism .................................................................................... 491
13.1 Energy and Entropy for Life!494
13.2 Energy in Biochemical Reactions!498
13.3 Energy Carriers and Electron Transfer!501
Special Topic 13.1: Microbial Syntrophy Cleans Up Oil!502
13.4 Catabolism: The Microbial Buffet!510
13.5 Glucose Breakdown and Fermentation!516
13.6 The Tricarboxylic Acid (TCA) Cycle!527
13.7 Aromatic Pollutants!532
CHAPTER 14
Electron Flow in Organotrophy, Lithotrophy, and Phototrophy .............. 539
14.1 Electron Transport Systems!541
14.2 The Proton Motive Force!546
Special Topic 14.1: Testing the Chemiosmotic Theory!548
14.3 The Respiratory ETS and ATP Synthase!551
14.4 Anaerobic Respiration in Organotrophs!559
Special Topic 14.2: Bacterial Electric Power!562
14.5 Lithotrophy and Methanogenesis!563
14.6 Phototrophy!571
CHAPTER 15
Biosynthesis ................................................................................................................585
15.1 Overview of Biosynthesis!586
15.2 CO2 Fixation: The Calvin Cycle!589
15.3 CO2 Fixation: Diverse Pathways!597
15.4 Biosynthesis of Fatty Acids and Polyketides!601
15.5 Nitrogen Fixation!605
Special Topic 15.1: Mining a Bacterial Genome for Peptide Antibiotics!606
15.6 Biosynthesis of Amino Acids and Nitrogenous Bases!612
15.7 Biosynthesis of Tetrapyrroles!618
XII ! CONTENTS
CHAPTER 16
Food and Industrial Microbiology ........................................................................623
16.1 Microbes as Food!625
16.2 Fermented Foods: An Overview!627
16.3 Acid- and Alkali-Fermented Foods!629
16.4 Ethanolic Fermentation: Bread and Wine!637
16.5 Food Spoilage and Preservation!641
16.6 Industrial Microbiology!650
Special Topic 16.1: Companies Take On Tuberculosis!652
Special Topic 16.2: Microbial Enzymes Make Money!654
PART 4
Microbial Diversity and Ecology 662
CHAPTER 17
Origins and Evolution............................................................................................... 665
17.1 Origins of Life!667
17.2 Early Metabolism!676
17.3 Microbial Phylogeny and Gene Transfer!681
Special Topic 17.1: Phylogeny of a Shower Curtain Biofilm!686
17.4 Adaptive Evolution!692
17.5 Microbial Species and Taxonomy!696
Special Topic 17.2: Jump-Starting Evolution of a Hyperthermophilic Enzyme!698
17.6 Symbiosis and the Origin of Mitochondria and Chloroplasts!703
CHAPTER 18
Bacterial Diversity ..................................................................................................... 711
18.1 Bacterial Diversity at a Glance!712
18.2 Cyanobacteria: Oxygenic Phototrophs!718
18.3 Firmicutes and Actinobacteria (Gram-Positive)!723
18.4 Proteobacteria (Gram-Negative)!734
Special Topic 18.1: Carbon Monoxide: Food for Bacteria?!738
18.5 Deep-Branching Gram-Negative Phyla!748
18.6 Spirochetes: Sheathed Spiral Cells with Internalized Flagella!750
18.7 Chlamydiae, Planctomycetes, and Verrucomicrobia: Irregular Cells!752
CONTENTS ! XIII
CHAPTER 19
Archaeal Diversity ..................................................................................................... 757
19.1 Archaeal Traits!758
19.2 Crenarchaeota across the Temperature Range!766
19.3 Thaumarchaeota: Symbionts and Ammonia Oxidizers!773
19.4 Methanogens!775
Special Topic 19.1: Eating Ammonia: Thaumarchaeotes!776
19.5 Haloarchaea!784
Special Topic 19.2: Haloarchaea in the Classroom!786
19.6 Thermophilic and Acidophilic Euryarchaeota!790
19.7 Deeply Branching Divisions!793
CHAPTER 20
Eukaryotic Diversity ................................................................................................. 797
20.1 Phylogeny of Eukaryotes!798
20.2 Fungi!806
Special Topic 20.1: Yeast: A Single-Celled Human Brain?!810
20.3 Algae!818
20.4 Amebas and Slime Molds!824
20.5 Alveolates: Ciliates, Dinoflagellates, and Apicomplexans!827
20.6 Trypanosomes and Metamonads!833
Special Topic 20.2: The Trypanosome: A Shape-Shifting Killer!834
CHAPTER 21
Microbial Ecology ...................................................................................................... 839
21.1 Metagenomes—and Beyond!841
21.2 Functional Ecology!849
21.3 Symbiosis!852
21.4 Marine and Aquatic Microbes!857
Special Topic 21.1: Cleaning Up the Deepwater Oil Spill!858
21.5 Soil and Subsurface Microbes!869
21.6 Plant Microbial Communities!876
21.7 Animal Microbial Communities!882
CHAPTER 22
Microbes in Global Elemental Cycles .................................................................889
22.1 Biogeochemical Cycles!890
22.2 The Carbon Cycle and Bioremediation!894
22.3 The Hydrologic Cycle and Wastewater Treatment!896
Special Topic 22.1: Bioremediation of Weapons Waste!902
22.4 The Nitrogen Cycle!903
22.5 Sulfur, Phosphorus, and Metals!908
22.6 Astrobiology!916
XIV ! CONTENTS
PART 5
Medicine and Immunology 922
CHAPTER 23
Human Microbiota and Innate Immunity........................................................... 925
23.1 Human Microbiota: Location and Shifting Composition!926
23.2 Risks and Benefits of Microbiota!934
23.3 Overview of the Immune System!935
Special Topic 23.1: Are NETs a Cause of Lupus?!940
23.4 Barbarians at the Gate: Innate Host Defenses!943
23.5 The Acute Inflammatory Response!946
23.6 How Phagocytes Detect and Kill Microbes!950
23.7 Interferon, Natural Killer Cells, and Toll-like Receptors!953
23.8 Complement’s Role in Innate Immunity!955
23.9 Fever!958
CHAPTER 24
The Adaptive Immune Response ......................................................................... 961
24.1 Overview of Adaptive Immunity!962
24.2 Immunogenicity!965
24.3 Antibody Structure and Diversity!968
24.4 Primary and Secondary Antibody Responses!974
24.5 Genetics of Antibody Production!977
24.6 T Cells Link Antibody and Cellular Immune Systems!982
24.7 Complement as Part of Adaptive Immunity!992
Special Topic 24.1: An Uneasy Peace: Détente at the Microbiota-Intestine Interface!994
24.8 Hypersensitivity and Autoimmunity!995
CHAPTER 25
Microbial Pathogenesis ........................................................................................ 1003
25.1 Host-Pathogen Interactions!1004
25.2 Virulence Factors and Pathogenicity Islands!1010
25.3 Microbial Attachment: First Contact!1013
25.4 Toxins Subvert Host Function!1018
CONTENTS ! XV
CHAPTER 26
Microbial Diseases ................................................................................................. 1051
26.1 Characterizing and Diagnosing Microbial Diseases!1052
26.2 Skin and Soft-Tissue Infections!1054
26.3 Respiratory Tract Infections!1058
26.4 Gastrointestinal Tract Infections!1066
Special Topic 26.1: Sprouts and an Emerging Escherichia coli!1068
26.5 Genitourinary Tract Infections!1075
26.6 Central Nervous System Infections!1083
26.7 Cardiovascular System Infections!1090
26.8 Systemic Infections!1094
26.9 Immunization!1102
CHAPTER 27
Antimicrobial Therapy .......................................................................................... 1107
27.1 The Golden Age of Antibiotic Discovery!1108
27.2 Fundamentals of Antimicrobial Therapy!1111
27.3 Measuring Drug Susceptibility!1112
27.4 Mechanisms of Action!1116
27.5 Challenges of Drug Resistance!1126
27.6 The Future of Drug Discovery!1134
27.7 Antiviral Agents!1135
Special Topic 27.1: Anti-Quorum Sensing Drug Blocks Pathogen “Control and Command”!1136
Special Topic 27.2: Resurrecting the 1918 Pandemic Flu Virus!1139
27.8 Antifungal Agents!1142
CHAPTER 28
Clinical Microbiology and Epidemiology .........................................................1147
28.1 Principles of Clinical Microbiology!1148
28.2 Specimen Collection and Processing!1149
28.3 Conventional Approaches to Pathogen Identification!1152
28.4 Rapid Techniques for Pathogen Identification!1160
28.5 Point-of-Care Rapid Diagnostics!1167
28.6 Biosafety Containment Procedures!1169
28.7 Principles of Epidemiology!1171
Special Topic 28.1: What’s Blowing in the Wind?!1178
28.8 Detecting Emerging Microbial Diseases!1179
XVI ! CONTENTS
APPENDIX 1
Biological Molecules .................................................................................................. A-1
A1.1 Elements, Bonding, and Water!A-2
A1.2 Organic Molecules!A-6
A1.3 Proteins!A-8
A1.4 Carbohydrates!A-11
A1.5 Nucleic Acids!A-13
A1.6 Lipids!A-15
A1.7 Biological Chemistry!A-16
Special Topic A1.1: Calculating the Standard Free Energy Change, ∆G°, of Chemical Reactions!A-19
APPENDIX 2
Introductory Cell Biology: Eukaryotic Cells ................................................... A-23
A2.1 The Cell Membrane!A-24
A2.2 The Nucleus and Mitosis!A-31
A2.3 Problems Faced by Large Cells!A-34
A2.4 The Endomembrane System!A-34
A2.5 The Cytoskeleton!A-38
A2.6 Mitochondria and Chloroplasts!A-40
APPENDIX 3
Laboratory Methods for Microbiology ............................................................A-43
A3.1 Isolating Parts of Cells by Using an Ultracentrifuge!A-44
A3.2 Agarose Gel Electrophoresis!A-45
A3.3 Protein Identification on 2D Gels with Mass Spectrometry!A-47
A3.4 RNA and DNA Identification by Northern and Southern Blots!A-49
A3.5 Sanger Method of DNA Sequencing!A-51
A3.6 Gene Fusions Identify Regulatory Mutants!A-52
A3.7 Primer Extension Identifies Transcriptional Start Sites!A-52
A3.8 DNA Microarray!A-54
A3.9 Multiplex PCR!A-55
A3.10 Fluorescence In Situ Hybridization (FISH) and CARD-FISH!A-55
A3.11 Immunoprecipitation Techniques!A-57
APPENDIX 4
Taxonomy ................................................................................................................... A-59
A4.1 Viruses!A-60
A4.2 Bacteria!A-62
A4.3 Archaea!A-66
A4.4 Eukarya!A-68
1.1 An Interview with Rita Colwell: The Global Impact 10.5 The Phage Lambda Lysis/Lysogeny “Decision”
of Microbiology Is to Kill or Not to Kill
1.2 Discovering the Genetic Code 11.1 Poliovirus: (+) Strand RNA
1.3 Clifford W. Houston: From Aquatic Pathogens 11.2 The Filamentous Phage M13: Vaccines
to Outer Space—An Interview and Nanowires
2.1 Confocal Microscopy 11.3 Genetic Resistance to HIV
3.1 Isolation and Analysis of the Ribosome 12.1 Mapping the E. coli Interactome
3.2 How Antibiotics Cross the Outer Membrane 12.2 GFP Proteins Track Cell Movements in Biofilms
3.3 Outer Membrane Proteins: Isolation for Vaccine 12.3 DNA Vaccines
3.4 Experiments That Reveal the Bacterial Cytoskeleton 12.4 Gene Therapy and Gene Delivery Systems
4.1 Transport by Group Translocation: 12.5 Directed Evolution through Phage Display
The Phosphotransferase System Technology
4.2 Eukaryotes Transport Nutrients by Endocytosis 12.6 DNA Shuffling Enables In Vitro Evolution
4.3 Biofilms, Antibiotics, Garlic, and Disease 12.7 Site-Directed Mutagenesis Helps Us Probe Protein
5.1 The Arrhenius Equation Function
5.2 Some Alkaliphilic Enzymes Produce Useful Drug 13.1 Observing Energy Carriers in Living Cells
Delivery Systems 13.2 Swiss Cheese: A Product of Bacterial Catabolism
5.3 Signaling Virulence 13.3 Genomic Analysis of Metabolism
5.4 Oligotrophs 13.4 Pyruvate Dehydrogenase Connects Sugar
6.1 How Did Viruses Originate? Catabolism to the TCA Cycle
6.2 West Nile Virus, an Emerging Pathogen 13.5 Genetic Analysis of Aromatic Catabolism
7.1 Genes and Proteins Involved in DNA Replication 14.1 Caroline Harwood: A Career in Bacterial
Photosynthesis and Biodegradation: An Interview
7.2 Trapping a Sliding Clamp
14.2 Measuring Dy and DpH in Microbes by the Uptake
7.3 Replication Mechanisms of Bacteriophages
of Molecules
7.4 Plasmid Partitioning and Addiction
14.3 Environmental Regulation of the ETS
7.5 Equilibrium Density Gradient Centrifugation
14.4 ATP Synthesis at High pH
8.1 Building the Ribosome Machine
15.1 The Discovery of 14C
8.2 Discovering the mRNA Ribosome-Binding Site
15.2 Metagenomic Screening for Polyketide Drugs
8.3 Ubiquitination: A Ticket to the Proteasome
15.3 Antibiotic Factories: Modular Biosynthesis
8.4 What Is the Minimal Genome? of Vancomycin
9.1 F Pili and Biofilm Formation 15.4 Riboswitch Regulation
9.2 Mapping Bacterial Chromosome Gene Position 16.1 From Barley and Hops to Beer
by Conjugation
16.2 Caterpillar Viruses Produce Commercial Products
9.3 Deinococcus Uses RecA to Repair Fragmented
17.1 The RNA World: Clues for Modern Medicine
Chromosomes
17.2 Horizontal Gene Transfer in E. coli O157:H7
9.4 Mutation Rate
17.3 Leaf-cutter Ants with Partner Fungi and Bacteria
9.5 The Transposase for a Bacterial Transposon
Resembles the Integrase for HIV-1 18.1 Karl Stetter: Adventures in Microbial Diversity Lead
to Products in Industry
9.6 Integrons and Gene Capture
20.1 Oomycetes: Lethal Parasites That Resemble Fungi
10.1 CRP Interactions with RNA Polymerase
and CRP-Dependent Promoters 20.2 A Ciliate Model for Human Aging
10.2 Glucose Transport Alters cAMP Levels 21.1 Mapping Bermuda Phytoplankton
10.3 Slipped-Strand Mispairing 21.2 Cold-Seep Ecosystems
10.4 Toxin-Antitoxin Modules: Mechanisms 22.1 Wetlands: Disappearing Microbial Ecosystems
for Self-preservation or Altruism? 22.2 Metal Contamination and Bioremediation
XVII
XVIII ! ETOPIC CONTENTS
I
n the first two editions of Microbiology: An Evolving Science, we worked to write
the defining core text of our generation—the book that would inspire under-
graduate science majors to embrace the microbial world. Our emphasis on genet-
ics and ecology, the use of case histories in the medical section, and the balanced
depiction of women and minority scientists, including young researchers, drew—and
continues to draw—enthusiastic responses from our more than one hundred adopt-
ers. Our focus on evolution, and our modern organization reflecting changes in the
field, proved so successful that other textbooks have adjusted their chapter sequence
to parallel An Evolving Science. In the Third Edition, we maintain this chapter orga-
nization to facilitate year-to-year course transitions for instructors. In addition, we
incorporate exciting new research advances to ensure that An Evolving Science is the
most current and engaging microbiology textbook available.
Also in this Third Edition, we maintain our signature balance between cutting-
edge ecology and medicine, while adding new research topics and emerging
microbial-human partnerships. The book opens with a new Part 1 Interview with
Richard Lenski, in which he presents his personal perspective on the groundbreak-
ing bacterial evolution experiment. Experimental evolution now fills a new section
in Chapter 17, Origins and Evolution. Other chapters that underwent major revision
include Chapter 3, Cell Structure and Function, with a tightened opener and a new
section on cell aging; and Chapter 21, Microbial Ecology, which opens with a new
section on metagenomics and the culturing of “unculturables.”
In many chapters, we relate topics to current events, to keep students interested
in and informed on the role of microbiology in the world today. One example is
synthetic biology, the construction of microbes with genetic circuits engineered for
commercial use (Chapter 12, Biotechnology and Synthetic Biology). Another exam-
ple is the use of of viral replication cycles to develop lentiviral treatments for cancer
and inherited disorders, including the first possible “cure” for pediatric leukemia
(presented in Chapter 11, Viral Molecular Biology).
Our Third Edition continues as a community project, drawing on our experi-
ence as researchers and educators as well as the input of hundreds of colleagues
to create a microbiology text for the twenty-first century. We present the story of
molecular microbiology and microbial ecology from its classical history of Koch, Pas-
teur, and Winogradsky, to twenty-first-century researchers Rita Colwell and Bonnie
Bassler. The Third Edition includes many contributions recommended by colleagues
from around the world, at institutions such as Washington University, University
of California–Davis, University of Wisconsin–Madison, Cornell University, Florida
State University, University of Toronto, University of Edinburgh, University of Ant-
werp, Seoul National University, Chinese University of Hong Kong, and many more.
We are grateful to you all.
XIX
XX ! PREFACE
While we have expanded and developed new topics, we also recognized the need
to keep the length and “core” of the book to a size reasonable enough for the under-
graduate student. In order to contain length while adding new material, we continue
transferring certain topics online as “eTopics.” The eTopics are called out in the text,
hyperlinked to the ebook, and their key terms are fully indexed in the printed book.
Therefore, returning adopters can be confident of keeping access to all of the mate-
rial they taught from the Second Edition, but now they also have new topics on Myco-
bacterium tuberculosis cell aging and drug resistance (Chapter 3) and on bacteria that
convert phage genes into toxin secretion systems (Chapter 25), and many more.
Major Features
Our book targets the science major in biology, microbiology, or biochemistry. Sev-
eral important features make our book the best text available for undergraduates
today:
! New research on contemporary themes such as evolution, genomics, meta-
genomics, molecular genetics, and biotechnology enrich students’ understanding
of foundational topics and highlight the current state of the field. Every chapter
presents numerous current research examples within the up-to-date framework of
molecular biology. Examples of current research include measuring the movement
of a single translating ribosome; transplanting a whole genome; determining the
“pangenome,” the overall set of genes available to a species; and the spectroscopic
measurement of carbon flux from microbial communities.
! A comprehensively updated art program with engaging figures that are also
dynamic learning tools. A fresh, contemporary new design and an updated art
program presents content in an engaging, visually dynamic manner. New in-
figure Thought Questions encourage students to interpret and analyze visuals of
important concepts. Figures that pair with a process animation online include a
QR code in the text that students can scan using their smartphones to immedi-
ately view online.
! Core concepts are presented in a student friendly way that motivates learning.
Ample Thought Questions throughout every chapter challenge students to think
critically about core concepts, the way a scientist would. In addition, scientists
pursuing research today are presented alongside the traditional icons. For exam-
ple, Chapter 1 introduces historical figures such as Koch and Pasteur alongside
genome sequencer Claire Fraser-Liggett and young microbial ecologist Kazem
Kashefi growing a hyperthermophile in an autoclave, and undergraduate students
conducting transcriptomics in E. coli. Medical microbiology is presented using the
physician-scientist’s approach to microbial diseases. Case histories present how a
physician-scientist approaches the interplay between the human immune response
and microbial diseases.
! An innovative media package provides powerful tools for instructors and stu-
dents. A new Micrograph Database for instructors includes hundreds of micro-
graphs from the book and beyond tagged by easy-to-browse categories as well as
by chapter. For students, a new ebook integrates powerful new self-study ques-
tions, process animations, quiz questions, weblinks, eTopics, and more to encour-
age the use of multimedia to enhance their learning of core concepts.
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first appears above ground, it has a sweet taste like milk, and will
not bear carriage. As it grows older, it gets strong, stones being piled
round to protect it from the sun. The root of the plant is not used as
medicine. There are no date trees in Cabool, though they are to be
found both east and west of it—at Candahar and Peshawur. There
the people are ignorant of the art of extracting an intoxicating juice
from them, as in India. Peshawur is celebrated for its pears; Ghuzni
for its plums, which are sold in India under the name of the plum of
Bokhara; Candahar for its figs, and Cabool for its mulberries; but
almost every description, particularly stone fruits, thrive in Cabool.
Fruit is more plentiful than bread, and is considered one of the
necessaries of human life. There are no less than fourteen different
ways of preserving the apricot of Cabool: it is dried with and without
the stone; the kernel is sometimes left, or an almond is substituted
in its stead; it is also formed into cakes, and folded up like paper. It
is the most delicious of the dried fruits.
Among the public buildings in Cabool, the Bala
Bala Hissar, or
palace prison of the
Hissar, or citadel, claims the first importance; but
princes. not from its strength. Cabool is enclosed to the
south and west by high rocky hills; and at the
eastern extremity of these the Bala Hissar is situated, which
commands the city. It stands on a neck of land, and may have an
elevation of about 150 feet from the meadows of the surrounding
country. There is another fort under it, also called the Bala Hissar,
which is occupied by the governor and his guards. The citadel is
uninhabited by the present chief; but his brother built a palace in it
called the “Koollah i Firingee,” or the Europeans’ Hat, which is the
highest building. Dost Mahommed Khan captured the Bala Hissar, by
blowing up one of its towers: it is a poor, irregular, and dilapidated
fortification, and could never withstand an escalade. The upper fort
is small, but that below contains about five thousand people. The
King’s palace stands in it. The Bala Hissar was built by different
princes of the house of Timour, from Baber downwards. Aurungzebe
prepared extensive vaults under it, to deposit his treasure; and
which may yet be seen. While it formed the palace of the kings of
Cabool, it was also the prison of the younger branches of the royal
family, in which they were confined for life. They tell a story, that,
when set free from their prison, after murdering their keeper, they
looked with astonishment at seeing water flow—so close had been
the confinement in their walled abode. It is difficult to say, whether
these unfortunate men were not happier than in their present state,
which is that of abject poverty. Many of the sons of Timour Shah
came in absolute hunger to solicit alms from us. I advised them to
make a petition to the chief for some permanent relief, but they said
that they had no mercy to expect from the Barukzye family, now in
power, who thirsted after their blood.
Near the Bala Hissar, and separated from it and
Persians of Cabool.
every part of the city, the Persians, or
Kuzzilbashes, as they are called, reside. They are
Toorks, and principally of the tribe of Juwansheer, who were fixed in
this country by Nadir Shah. Under the kings of Cabool they served as
body-guards, and were a powerful engine of the state. They yet
retain their language, and are attached to the present chief, whose
mother is of their tribe. I had an opportunity of seeing these people
to advantage; being invited to a party given by our conductor from
Peshawur, the jolly Naib Mahommed Shureef. I met the whole of the
principal men, and their chief, Sheereen Khan. The entertainment
was more Persian than Afghan. Among them, I could discover a new
people, and new mode of thinking; for they have retained some of
the wit that marks their countrymen. As the evening was drawing to
a close, the chief called on a person to display his powers, not in a
tale, but in depicting the peculiarities of the neighbouring nations.
He began with the Afghans; and, after an amusing enough
exordium, which excepted the Dooranees or chiefs, (who, he said,
were not like other Afghans,) he described the entry of some twenty
or thirty nations into paradise. When the turn of the Afghans came,
he went on blasphemously to relate, that their horrid language was
unintelligible, and that, as the prophet had pronounced it to be the
dialect of hell, there was no place in heaven for those who spoke it.
The fellow had humour, and brought in some Afghan phrases, much
to the amusement of the company. He then attacked the Uzbeks for
their peculiar way of making tea, and their uncouth manners. He
now levelled his batteries against the whining, cheating and deceitful
Cashmeerian; and these people must be belied indeed, if they be
not masters in vice.[16] All parties, however, admit their talents and
ingenuity, which is a considerable counterbalance. The natives of
Herat, and their peculiar dialect, exercised the powers of this
loquacious Meerza: he imitated the roguery of their custom-house;
and allowed himself, as the officer on duty, to be bribed out of his
due, by accepting some wine, which he pretended was not for
himself.
The difference between Eastern manners, and
Difference between
Asiatic and
those of Europe, is nowhere more discernible than
European manners. in their manner of saying good things. An
European enjoys an anecdote; but he would be
very much surprised to be called on in a company to tell one for its
amusement. In the East, there are professional anecdote makers; in
the West, we are content with a bon-mot as it flows in the course of
conversation. Both may be traced to the government: for, in the
East, though there is much familiarity, there is little social
intercourse; and, in Europe, good manners teach us to consider
every one at the same board on an equality.
During our stay, the “Eed” occurred, which is the
Eed, or festival.
festival kept in commemoration of Abraham’s
intention to sacrifice his son Isaac. It was
observed with every demonstration of respect: the shops were shut;
and the chief proceeded to prayer at an appointed place, with a
great concourse of persons. In the afternoon, every one was to be
seen flocking to the gardens; nor could I resist the impulse, and
followed the crowd. In Cabool, you no sooner leave the bazar, than
you find yourself on the banks of the river, which are beautifully
shaded by trees of mulberry, willow, and poplar. Almost all the roads
round the city lead by the verge of aqueducts or running water. They
are crossed by bridges; and the large river has three or four of these
edifices; but they cannot boast of architectural beauty. The finest
gardens of Cabool lie north of the city; and they, again, are far
surpassed by those beyond, in the district of Istalif, under the first
snow-clad mountains, towards Hindoo Koosh. Their site is to be seen
from Cabool. I was conducted to the tomb of
Tomb of Timour
Shah.
Timour Shah, which stands outside the city, and is
a brick building of an octagon shape, rising to the
height of 50 feet. The interior of it is about 40 feet square, and the
architecture resembles that of Delhi. The building is unfinished. A
lamp was formerly lighted on this sepulchre; but the sense of this
king’s favours, like that of many others, has faded. Timour Shah
made Cabool his capital, and here is his tomb. His father is interred
at Candahar, which is the native country of the Dooranees.
I moved about every where during the day, and
Alchymy and
minerals.
had the pleasure of many sociable evenings with
our host the Nawab, whom I found, like many of
his countrymen, in search of the philosopher’s stone. Such an
opportunity as our arrival seemed to promise him a rich harvest. I
soon undeceived him, and laughed at the crucibles and recipes,
which he produced. I explained to him, that chemistry had
succeeded alchymy, as astronomy had followed astrology; but as I
had to detail the exact nature of these sciences, my asseverations of
being no alchymist had little effect. He therefore applied himself to
the doctor, from whom he requested recipes for the manufacture of
calomel and quinine plasters and liniments; which it was no easy
matter to furnish. He could not credit that the arts of giving and
manufacturing medicines were distinct; and set us down as very
ignorant or very obstinate. He would not receive the prepared
medicines, as they would be of no use to him after we had left. We
found this feeling generally prevalent; and woe be to the doctor in
these parts who gives medicines which he cannot make. We kept the
Nawab in good humour, though we would not believe that he could
convert iron into silver. We heard from him the position of many
metallic veins in the country. He produced among other curiosities
some asbestos, here called cotton-stone (sung i poomba), found
near Julalabad. The good man declared that he must have some of
our knowledge in return for what he told so freely.
Freemasonry.
I informed him that I belonged to a sect called
Freemasons, and gave some account of the craft,
into which he requested to be admitted without delay. But, as the
number of brethren must be equal to that of the Pleiades, we put it
off to a convenient opportunity. He confidently believed that he had
at last got scent of magic in its purest dye; and had it been in my
power, I would have willingly initiated him. He made me promise to
send some flower-seeds of our country, which he wished to see in
Cabool; and I faithfully forwarded them. I cut the plates out of Mr.
Elphinstone’s History of Cabool, and presented them to the Nawab at
a large party; and not only is the costume exact, but in some of the
figures, to their great delight, they discovered likenesses. Pictures
are forbidden among the Soonee Mahommedans; but in the present
instance they proved very acceptable. Among the Nawab’s friends
we met a man 114 years old, who had served with Nadir Shah. He
had been upwards of eighty years in Cabool, and seen the Dooranee
dynasty founded and pass away. This venerable person walked up
stairs to our rooms.
From the crowd of people we constantly met at
Afghans. Jewish
origin.
the house of our host, I was resolved on gathering
some information on the much disputed point of
the Afghans being Jews. They brought me all their histories, but I
had no time to examine them, and wished for oral information. The
Afghans call themselves, “Bin i Israeel,” or children of Israel; but
consider the term of “Yahoodee,” or Jew, to be one of reproach.
They say that Nebuchadnezzar, after the
Their traditions.
overthrow of the temple of Jerusalem,
transplanted them to the town of Ghore, near
Bameean; and that they are called Afghans, from their chief
Afghana, who was a son of the uncle of Asof (the vizier of Solomon),
who was the son of Berkia. The genealogy of this person is traced
from a collateral branch, on account of the obscurity of his own
parent, which is by no means uncommon in the East. They say that
they lived as Jews, till Khaleed (called by the title of Caliph)
summoned them, in the first century of Mahommedanism, to assist
in the wars with the Infidels. For their services on that occasion,
Kyse, their leader, got the title of Abdoolrusheed, which means the
Son of the mighty. He was also told to consider himself the “butan”
(an Arabic word), or mast of his tribe, on which its prosperity would
hinge, and by which the vessel of their state was to be governed.
Since that time, the Afghans are sometimes called Putan, by which
name they are familiarly known in India. I never before heard this
explanation of the term. After the campaign with Khaleed, the
Afghans returned to their native country, and were governed by a
king of the line of Kyanee, or Cyrus, till the eleventh century, when
they were subdued by Mahmood of Ghuzni. A race of kings sprung
from Ghore, subverted the house of Ghuzni, and conquered India.
As is well known, this dynasty was divided, at the death of its
founder, into the divisions east and west of the Indus; a state of
things which lasted till the posterity of Timourlane reduced both to a
new yoke.
Having precisely stated the traditions and history
Opinions regarding
these traditions.
of the Afghans, I can see no good reason for
discrediting them, though there be some
anachronisms, and the dates do not exactly correspond with those
of the Old Testament. In the histories of Greece and Rome we find
similar corruptions, as well as in the later works of the Arab and
Mahommedan writers. The Afghans look like Jews; they say they are
descended from Jews; and the younger brother marries the widow
of the elder, according to the law of Moses. The Afghans entertain
strong prejudices against the Jewish nation; which would at least
show that they had no desire to claim, without a just cause, a
descent from them. Since some of the tribes of Israel came to the
East, why should we not admit that the Afghans are their
descendants, converted to Mahommedanism? I am aware that I am
differing from a high authority[17]; but I trust that I have made it
appear on reasonable grounds.
As the chief desired, I passed another evening
Party at the chief’s.
with him; and the doctor, being convalescent,
accompanied me; Mr. Wolff had proceeded on his
journey to India. Dost Mahommed Khan pleased us as much as
ever; he kept us till long past midnight, and gave us a full insight
into the political affairs of his country, and the unfortunate
differences that exist between him and his brothers. He expressed
hopes of being able to restore the monarchy,
Politics.
evinced a cordial hatred towards Runjeet Sing,
and seemed anxious to know if the British
Government would accept his services as an auxiliary to root him
out; but I replied, that he was our friend. He then promised me the
command of his army, if I would remain with him; an offer which he
afterwards repeated. “Twelve thousand horse and twenty guns shall
be at your disposal.” When he found that I could not accept the
honour, he requested me to send some friend to be his
generalissimo. On this occasion, we had some
The Kaffirs a
singular people.
highly interesting conversation regarding the
Kaffirs, who live in the hills north of Peshawur and
Cabool, and are supposed to descend from Alexander. The chief, on
the former occasion, had produced a young Kaffir boy, one of his
slaves, about ten years old, who had been captured for two years.
His complexion, hair, and features were quite European; his eyes
were of a bluish colour. We made him repeat various words of his
language, some of which were Indian. The Kaffirs live in a most
barbarous state, eating bears and monkeys. There is a tribe of them
called “Neemchu Moossulman,” or half Mahommedans, who occupy
the frontier villages between them and the Afghans, and transact the
little trade that exists among them. It is curious to find a people so
entirely distinct from the other inhabitants, and unfortunately every
thing that regards them rests in obscurity. I have hereafter stated
the particulars which I collected regarding the Kaffirs, whom I take
to be the aborigines of Afghanistan, and in no wise connected with
the reputed descendants of Alexander the Great, as has been stated
by some authors.
We had passed nearly three weeks in Cabool;
Preparations for our
journey.
which appeared as a few days. It was now
necessary to prepare for our journey, which
seemed no easy matter. No caravan was yet ready; and it was even
doubtful if the roads were passable, as snow had fallen during the
month. It occurred to me that our best plan would be to hire a
Cafila-bashee, or one of the conductors of the great caravans, as
one of our own servants; and we might thus proceed at once,
without the delay attendant upon a caravan, and, I hoped, with
equal safety. The Nawab did not altogether relish the plan, nor our
precipitate departure. He would have willingly kept us for months.
We, however, entertained one Hyat, a sturdy but hale old man, who
had grown grey in crossing the Hindoo Koosh. When the Nawab
found our determination to depart, he urged his relative, the Ameen
ool Moolk, a nobleman of the late Shah Mahmood, who carries on
commercial transactions with Bokhara and Russia, to despatch one
of his trusty persons with us. It was therefore determined that a
brother of his Nazir, or steward, named Doulut, a respectable
Afghan, also styled the Nazir, should proceed with us. He had
business in Bokhara, and was even going on to Russia: our
movements expedited his departure. Every thing looked well, and we
were furnished by the Nawab’s kindness with letters to the Afghans
in Bokhara. The most influential of these was Budr-oo-deen. His
agent in Cabool, who brought me the letters, was resolved on being
rewarded for doing so by an enjoyment of our society. His name was
Khodadad, and he was a Moollah. He stopped and dined with us; but
declared, that whatever might be our wisdom as a nation, we had
no correct ideas of good living. He did not like our English fare,
which was cooked with water, he said, and only fit for an invalid.
Khodadad was a very intelligent man, who had travelled in India and
Tartary, and was well read in Asiatic lore. He had also studied Euclid,
whom his countryman, he said, nicknamed “Uql doozd,” or wisdom-
stealer, from the confusion which he had produced in men’s heads.
He was not fond of mathematics, and wished to know our motive for
studying them: he had not heard that it improved the reasoning
faculties; and only considered the persons versed in Euclid, as
deeper read than others. The chief also prepared his letters; but
there is little communication between the Afghans and Uzbeks, and
we found them of no service; that for the King of Bokhara was lost
or stolen. One of Dost Mahommed Khan’s court, however, the
governor of Bameean, Hajee Kauker, furnished us with letters, which
were of real use, as will afterwards appear. This man, though serving
under the chief of Cabool, is more friendly to his brother of
Peshawur, by whom we were introduced to him. I held my
intercourse with him secret, and he tendered the services of fifty
horsemen, which it was prudent to decline.
Before our departure from Cabool, I made the
Shikarpooree
merchants.
acquaintance of many of the Hindoo or
Shikarpooree merchants. The whole trade of
Central Asia is in the hands of these people, who have houses of
agency from Astracan and Meshid to Calcutta. They are a plodding
race, who take no share in any other matters than their own, and
secure protection from the Government by lending it money. They
have a peculiar cast of countenance, with a very high nose: they
dress very dirtily. Few of them are permitted to wear turbans. They
never bring their families from their country, which is Upper Sinde,
and are constantly passing to and from it; which keeps up a national
spirit among them. In Cabool, there are eight great houses of
agency belonging to these people, who are quite separate from the
other Hindoo inhabitants. Of them, there are about three hundred
families. I met one of these Shikarpooree merchants on the Island of
Kisham, in the Gulf of Persia; and were Hindoos tolerated in that
country, I feel satisfied that they would spread all over Persia, and
even Turkey.
With such an extensive agency distributed in the
Money
arrangements.
parts of Asia which we were now about to
traverse, it was not, as may be supposed, a very
difficult task to adjust our money matters, and arrange for our
receiving a supply of that necessary article, even at the distance we
should shortly find ourselves from India. Our expenses were small,
and golden ducats were carefully sewed up in our belts and turbans,
and sometimes even transferred to our slippers; though, as we had
to leave them at the door of every house, I did not always approve
of such stowage. I had a letter of credit in my possession for the
sum of five thousand rupees, payable from the public treasuries of
Lodiana or Delhi; and the Cabool merchants did not hesitate to
accept it. They expressed their readiness either to discharge it on
the spot with gold, or give bills on Russia at St. Macaire (Nijnei
Novgorod), Astracan, or at Bokhara, which I had no reason to
question: I took orders on the latter city. The merchants enjoined
the strictest secrecy; and their anxiety was not surpassed by that of
our own to appear poor; for the possession of so much gold would
have ill tallied with the coarse and tattered garments which we now
wore. But what a gratifying proof have we here of
Great proofs of the
civilisation by
the high character of our nation, to find the bills of
commerce. those who almost appeared as beggars cashed,
without hesitation, in a foreign and far distant
capital. Above all, how much is our wonder excited to find the
ramifications of commerce extending uninterruptedly over such vast
and remote regions, differing as they do from each other in
language, religion, manners, and laws.
CHAP. VI.
JOURNEY OVER THE HINDOO KOOSH, OR SNOWY
MOUNTAINS.