Microbiology

1147 Pages·126.819 MB·English

by  Prescott, Lansing M

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Lansing M. Prescott
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ISBN: 0-07-282905-2 h
Description: ©2002 / Hardcover with CDROM
Publication October 2002
Date:
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Overview
Prescott, Harley and Klein's 5th edition provides a balanced, comprehensive introduction to all
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major areas of microbiology. Because of this balance, Microbiology, 5/e is appropriate for students
preparing for careers in medicine, dentistry, nursing, and allied health, as well as research, teaching,
and industry. Biology and chemistry are prerequisites. The Fifth Edition has been updated
extensively to reflect the latest discovaeries in the field.

New to This Edition
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• Every chapter in the book has been updated to reflect the latest discoveries in microbiology, including
information on genomics, biofilms, mechanisms of toxins, classification, and emerging diseases. The most
extensive revision has occurred in the areas of genetics, microbial ecology, and immunology where material
has been updated and reorganized tio allow for easier use.
• New Genomics chapter: Chapter 15. The genetics coverage has been reorganized for clarity and ease of
teaching. The genetics section now ends with a completely new chapter on genomics. New Chapter 28 on
microorganism interactions and microbial ecology!
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• Newly developed art program--much of the art is new or revised! It incorporates color and style consistency
throughout so students will easily identify certain topics.
• New critical thinking questions have been added to provide practice in analyzing data, predicting outcomes,
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and to teach students how to think logically.
• The general organization of the text has been modified to provide a more logical flow of topics and give
greater emphasis to microbial ecology
S

Features
• Prescott's textbook contains briefer chapters than most books, but more of them (42). Students will find the
concise chapters more palatable and less intimidating. Short chapters give the instructor the opportunity to fit
the text more closely to the instructor's syllabus. Topic flexibility is allowed.
• There is an outstanding pedagogical system including outlines, concepts, key terms, cross-referencing,
readings, new critical thinking questions, etc., to help students understand difficult material.
Prescott−Harley−Klein: Front Matter Preface © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
P
REFACE
Books are the carriers of civilization.Without books,history is silent,literature dumb,science
crippled,thought and speculation at a standstill.They are engines of change,windows on the
world,lighthouses erected in a sea of time.
–Barbara Tuchman
Microbiology is an exceptionally broad discipline encom- of microbial interactions such as mutualism,protocooperation,com-
passing specialties as diverse as biochemistry,cell biol- mensalism,and predation that occur in the environment. Parts Nine
ogy,genetics,taxonomy,pathogenic bacteriology,food and Ten are concerned with pathogenicity,resistance,and disease.
and industrial microbiology,and ecology. A microbiologist must The three chapters in Part Nine describe normal microbiota,non-
be acquainted with many biological disciplines and with all major specific host resistance,the major aspects of the immune response,
groups of microorganisms:viruses,bacteria,fungi,algae,and pro- and medical immunology. Part Ten first covers such essential topics
tozoa. The key is balance. Students new to the subject need an in- as microbial pathogenicity, antimicrobial chemotherapy, and epi-
troduction to the whole before concentrating on those parts of demiology. Then chapters 38–40 survey the major human microbial
greatest interest to them. This text provides a balanced introduction diseases. The disease survey is primarily organized taxonomically
to all major areas of microbiology for a variety of students. Because on the chapter level; within each chapter diseases are covered ac-
of this balance,the book is suitable for courses with orientations cording to their mode of transmission. This approach provides flex-
ranging from basic microbiology to medical and applied microbi- ibility and allows the student easy access to information concerning
ology. Students preparing for careers in medicine,dentistry,nurs- any disease of interest. The survey is not a simple catalog of diseases;
ing,and allied health professions will find the text just as useful as diseases are included because of their medical importance and their
those aiming for careers in research,teaching,and industry. Two ability to illuminate the basic principles of disease and resistance.
quarters/semesters each of biology and chemistry are assumed,and Part Eleven concludes the text with an introduction to food and in-
an overview of relevant chemistry is also provided in appendix I. dustrial microbiology. Five appendices aid the student with a review
of some basic chemical concepts and with extra information about
Organization and Approach important topics not completely covered in the text.
This text is designed to be an effective teaching tool. A text
The book is organized flexibly so that chapters and topics may be is only as easy for a student to use as it is easy to read. Readabil-
arranged in almost any order. Each chapter has been made as self- ity has been enhanced by using a relatively simple,direct writing
contained as possible to promote this flexibility. Some topics are style, many section headings, and an organized outline format
essential to microbiology and have been given more extensive within each chapter. The level of difficulty has been carefully set
treatment. with the target audience in mind. During preparation of the fifth
The book is divided into 11 parts. The first 6 parts introduce the edition,every sentence was carefully checked for clarity and re-
foundations of microbiology:the development of microbiology,the vised when necessary. The American Society for Microbiology’s
structure of microorganisms,microbial growth and its control,me- ASM Style Manualconventions for nomenclature and abbrevia-
tabolism,molecular biology and genetics,DNA technology and ge- tions have been followed as consistently as possible.
nomics,and the nature of viruses. Part Seven is a survey of the mi- The many new terms encountered in studying microbiology
crobial world. In the fifth edition, the bacterial survey closely are a major stumbling block for students. This text lessens the
follows the general organization of the forthcoming second edition problem by addressing and reinforcing a student’s vocabulary de-
ofBergey’s Manual of Systematic Bacteriology.Although principal velopment in three ways:(1)no new term is used without being
attention is devoted to bacteria,eucaryotic microorganisms receive clearly defined (often derivations also are given)—a student does
more than usual coverage. Fungi,algae,and protozoa are important not have to be familiar with the terminology of microbiology to
in their own right. The introduction to their biology in chapters use this text; (2)the most important terms are printed in boldface
25–27 is essential to understanding topics as diverse as clinical mi- when first used; and (3)a very extensive,up-to-date,page-refer-
crobiology and microbial ecology. Part Eight focuses on the rela- enced glossary is included at the end of the text.
tionships of microorganisms to other organisms and the environ- Because illustrations are critical to a student’s learning and
ment (microbial ecology). It also introduces aquatic and terrestrial enjoyment of microbiology, all illustrations are full-color, and
microbiology. Chapter 28 presents the general principles underlying many excellent color photographs have been used. Color not only
microbial ecology and environmental microbiology so that the sub- enhances the text’s attractiveness but also increases each figure’s
sequent chapters on aquatic and terrestrial habitats can be used with- teaching effectiveness. Considerable effort has gone into making
out excessive redundancy. The chapter also describes various types the art as attractive and useful as possible. Much of the art in the
xv
Prescott−Harley−Klein: Front Matter Preface © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
xvi Preface
fourth edition has been revised and improved for use in the fifth order to make cross references more precise. The summary
edition. All new line art has been produced under the direct super- now contains boldfaced references to tables and figures that
vision of an art editor and the authors,and designed to illustrate will be useful in reviewing the chapter.
and reinforce specific points in the text. Consequently every illus- 3. New illustrations have been added to almost every chapter.
tration is directly related to the narrative and specifically cited In addition,all figures have been carefully reviewed by our
where appropriate. Great care has been taken to position illustra- art editor,and many have been revised to improve their
tions as close as possible to the places where they are cited. Illus- appearance and usefulness.
trations and captions have been reviewed for accuracy and clarity. 4. All reference sections have been revised and updated.
Besides these broader changes in the text,every chapter has
Themes in the Book
been updated and often substantially revised. Some of the more
important improvements are the following:
At least seven themes run through the text, though a particular
one may be more obvious at some points than are others. These Chapter 1—A box on molecular Koch’s postulates and a new
themes or emphases are the following: section on the future of microbiology have been added.
Chapter 2—Differential interference contrast microscopy and
1. The development of microbiology as a science
confocal microscopy are described.
2. The nature and importance of the techniques used to
Chapter 3—More details on the mechanism of flagellar motion
isolate,culture,observe,and identify microorganisms
are provided.
3. The control of microorganisms and reduction of their
Chapter 5—Phosphate uptake and ABC transporters are
detrimental effects
described.
4. The importance of molecular biology for microbiology
Chapter 6—The chapter has new material on starvation
5. The medical significance of microbiology
proteins,growth limitation by environmental factors,viable
6. The ways in which microorganisms interact with their
but nonculturable procaryotes,and quorum sensing.
environments and the practical consequences of these
Chapter 8—The discussions of metabolic regulation and
interactions
control of enzyme activity have been combined with the
7. The influences that microorganisms and microbiological
introduction to energy and enzymes.
applications have on everyday life
Chapter 9—The metabolic overview has been rewritten to aid
These themes help unify the text and enhance continuity. The stu- in understanding. The sections on electron transport,
dent should get a feeling for what microbiologists do and for how oxidative phosphorylation,and anaerobic respiration have
their activities affect society. been updated and expanded.
Chapter 11—The chapter now focuses on nucleic acid and
What’s New in the Fifth Edition gene structure,mutations,and DNA repair. New material
on DNA methylation has been added.
Many substantial changes and improvements have been made in Chapter 12—Material on gene expression (transcription and
the fifth edition,including the following: protein synthesis) has been moved here and combined with
an extensive discussion of the regulation of gene
1. The general organization of the text has been modified to
expression. New sections on global regulatory systems and
provide a more logical flow of topics and give greater
two-component phosphorelay systems have been added.
emphasis to microbial ecology. Treatment of nucleic acid and
protein synthesis has been moved to the genetics chapters to Chapter 15—This new chapter provides a brief introduction to
integrate the discussion of gene structure,replication, microbial genomics,including genome sequencing,
expression,and regulation. Recombinant DNA technology bioinformatics,general characteristics of microbial
has been moved to a separate section,which also contains a genomes,and functional genomics.
new chapter on microbial genomics. The three-chapter Chapter 18—Virus taxonomy has been updated and new life
introduction to microbial ecology now follows the survey of cycle diagrams added.
microbial diversity. This places it earlier in the text where Chapter 19—Material on polyphasic taxonomy and the effects
basic principles of microbiology are introduced. Part Nine of horizontal gene transfer on phylogenetic trees has been
now contains a description of nonspecific host resistance as added. The introduction to the second edition of Bergey’s
well as an introduction to the fundamentals of immunology. Manualhas been revised and updated.
Symbiotic associations are discussed in the context of Chapters 20–24—The procaryotic survey chapters have been
microbial ecology. The treatment of microbial pathogenesis further revised to conform to the forthcoming second
has been expanded into a full chapter and placed with other edition of Bergey’s Manual.
medical topics in Part Ten. Chapter 28—This chapter,formerly chapter 40,has been
2. Pedagogical aids have been expanded. A new Critical substantially rewritten and now includes a treatment of
Thinking Questions section with two or more questions symbiosis and microbial interactions (e.g.,mutualism,
follows the Questions for Thought and Review. Section protocooperation,commensalism,predation,amensalism,
numbers have been given to all major chapter sections in competition,etc.). A discussion of microbial movement
Prescott−Harley−Klein: Front Matter Preface © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
Preface xvii
between ecosystems has been added,and the treatment of due to new molecular techniques. A section on developing
biofilms and microbial mats has been expanded. and choosing microorganisms for use in industry has been
Chapter 29—The chapter on microorganisms in aquatic added. Other topics that have been added or substantially
environments has new material on such topics as oxygen revised include the synthesis of products for medical use,
fluxes in water,the microbial loop,Thiomargarita biodegradation of pesticides and other pollutants,the
namibiensis,microorganisms in freshwater ice,and current addition of microorganisms to the environment,and the use
drinking water standards. of microarray technology.
Chapter 30—Microorganisms in cold moist area soils,desert soils,
and geologically heated hyperthermal soils are discussed. The Aids to the Student
effects of nitrogen,phosphorus,and atmospheric gases on
plants and soils are described more extensively. There is a new It is hard to overemphasize the importance of pedagogical aids for
section on the subsurface biosphere. the student. Accuracy is most important,but if a text is not clear,
Chapter 31—This reorganized chapter discusses normal readable,and attractive,up-to-dateness and accuracy are wasted
microbiota and nonspecific resistance. An overview of host because students will not read it. Students must be able to under-
resistance; a discussion of the cells,tissues,and organs of stand the material being presented, effectively use the text as a
the immune system; an introduction to the alternative and learning tool,and enjoy reading the book.
lectin complement pathways; and a summary of cytokine Tobe an effective teaching tool,a text must present the sci-
properties and functions have been included. ence of microbiology in a way that can be clearly taught and eas-
ily learned. Therefore many aids are included to make the task of
Chapter 32—All aspects of specific immunity have been
learning more efficient and enjoyable. Following the preface a
moved to this chapter in order to provide a clearer and more
special section addressed to the student user reviews the princi-
coherent discussion. The chapter contains an overview of
ples of effective learning,including the SQ4R (survey,question,
specific immunity,a discussion of antigens and antibodies,
read,revise,record,and review) study technique. Specific chap-
T-cell and B-cell biology,a discussion of the action of
ter aids are described in the special Visual Preview section.
antibodies,the classical complement pathway,and a section
Besides the chapter aids the text also contains a glossary,an
on acquired immune tolerance. It ends with a summary of
index, and five appendices. The extensive glossary defines the
the role of antibodies and lymphocytes in resistance.
most important terms from each chapter and includes page refer-
Chapter 33—The new chapter on medical immunology contains
ences. Where desirable,phonetic pronunciations also are given.
topics more directly related to the practical aspects of health
Most of the glossary definitions have not been taken directly from
and clinical microbiology:vaccines and immunizations,
the text but have been rewritten to give the student further under-
immune disorders,and in vitro antigen-antibody interactions.
standing of the item. To improve ease of use,the fifth edition has
Previously these were scattered over three chapters. The
a large,detailed index.It has been carefully designed to make text
treatment of vaccines has been greatly expanded.
material more accessible. The appendicesaid the student with ex-
Chapter 34—The treatment of microbial pathogenicity has
tra review of chemical principles and metabolic pathways and
been greatly enlarged and made into a separate chapter.
provide further details about the taxonomy of bacteria and
Several topics have been expanded or added:regulation of
viruses. To aid the student in following the rapidly changing field
bacterial virulence factors and pathogenicity islands,the
of procaryotic taxonomy,appendix III provides the classification
mechanisms of exotoxin action,and microbial mechanisms
of procaryotes according to the first edition of Bergey’s Manual
for escaping host defenses.
of Systematic Bacteriology,and appendix IV gives the classifica-
Chapter 37—In the epidemiology chapter,the treatment of tion used by the upcoming second edition of Bergey’s Manual.
emerging diseases has been expanded. New sections on
bioterrorism and the effect of global travel on health have
Supplementary Materials
been added.
Chapters 38–40—The disease survey chapters have been
Rich supplementary materials are available for students and in-
brought up-to-date,and bacterial diseases are now covered in
structors to assist learning and course management.
one chapter rather than two. New material has been added on
genital herpes,listeriosis,the use of clostridial toxins in For the Student
therapy,and other topics. A new table describing common
sexually transmitted diseases and their treatment is provided. 1. A Student Study Guideby Linda
Chapter 41—New aspects of food microbiology include Sherwood of Montana State University
discussions of modified atmosphere packaging,algal is a valuable resource that provides
toxins,bacteriocins as preservatives,new variant learning objectives,study outlines,
Creutzfeldt-Jakob disease,food poisoning by uncooked learning activities,and self-testing
foods,new techniques in tracing outbreaks of food-related material to help students master course
diseases,and the use of probiotics in the diet. content.
Chapter 42—The chapter on industrial microbiology and 2. TheInteractive E-TEXTavailable on CD-ROM in January
biotechnology has been revised to include current advances 2002 includes all of Microbiology,Fifth Edition,as well as the
Prescott−Harley−Klein: Front Matter Preface © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
xviii Preface
Student Study Guide in an interactive electronic format. The e- multimedia presentations or export images into other
text includes animations and web links to enhance learning. programs. Images may be sorted by a number of criteria.
3. The third edition of Microbes in Motionby Gloria Delisle Features include an Interactive Slide Show and a Slide
and Lewis Tomalty is an interactive CD-ROM that brings Editor.
microbiology to life. A correlation guide on the CD links 4. A set of 50 Projection Slidesprovides clinical examples of
this exciting resource directly to your textbook. This easy to diseases and pathogens to supplement the illustrations in
use tutorial can go from the classroom to the resource the text.
center to students’own personal computers. Microbes in 5. Your McGraw-Hill representative may arrange a Customized
Motionbrings discovery back into the learning and Laboratory Manualcombining your own material with
education process through interactive screens,animations, exercises from Laboratory Exercises in Microbiology,Fifth
video,audio,and hyperlinking questions. The applications Edition,by John P. Harley and Lansing M. Prescott. Contact
of this CD-ROM are only as limited as your good ideas. your McGraw-Hill representative for details about this
4. The second edition of Hyperclinic custom publishing service.
by Lewis Tomalty and Gloria 6. Designed specifically to help you with your individual
Delisle is packed with over 100 course needs,PageOut,PageOut Lite,andMcGraw-Hill
case studies and over 200 Course Solutionswill assist
pathogens supported with audio, you in integrating your
video,and interactive screens. syllabus with the fifth
Students will have fun and gain edition’s state-of-the-art
confidence as they learn valuable media tools. Create your own
concepts and gain practical course-specific web page
experience in clinical supported by McGraw-Hill’s
microbiology. extensive electronic resources,set up a class message
5. The fifth edition of Laboratory board or chat room online,provide online testing
Exercises in Microbiologyby opportunities for your students,and more!
John P. Harley and Lansing M. Prescott has been
prepared to accompany the text. Like the text,the
laboratory manual provides a balanced introduction to Online Resources
laboratory techniques and principles that are important in
each area of microbiology. The class-tested exercises are Through the Prescott 2002 Online Learning Center,everything you
modular and short so that an instructor can easily choose need for effective,interactive teaching and learning is at your fin-
only those exercises that fit his or her course. The fifth gertips. Moreover,this vast McGraw-Hill resource is easily loaded
edition contains recipes for all reagents and media. New into course management systems such as WebCT or Blackboard.
exercises in biotechnology have been added to this Through the Online Learning Center,you will also link to McGraw-
edition. A new appendix provides practice in solving Hill’s new Biocourse.comsite with
dilution problems. a huge dynamic array of resources
6. A set of 305 Microbiology Study Cardsprepared by Kent to supplement your learning experi-
M. Van De Graaff,F. Brent Johnson,Brigham Young ence in microbiology.
University,and Christopher H. Creek features complete Some of the online features
descriptions of terms,clearly labeled drawings,clinical you will find to support your use of
information on diseases,and much more. Microbiology by Prescott, Harley,
and Klein include the following.
For the Instructor
For the Student:
• Additional multiple-choice questions in a self-quizzing
1. A Testing CDis offered free on request to adopters of the
interactive format
text. This cross-platform CD provides a database of over
• Electronic flashcards to review key vocabulary
2,500 objective questions for preparing exams and a grade-
recording program. • Study Outlines
2. A set of 250 full-color acetate Transparenciesis available to • Web Links and Exercises
supplement classroom lectures. These have been enhanced for • Clinical Case Studies
projection and are available to adopters of the fifth edition. • An Interactive Time Line detailing events and
3. The Visual Resource LibraryCD-ROM contains virtually highlighting personalities critical to the development of
all of the art and many of the photos from Microbiology, microbiology
Fifth Edition,as well as the tables that appear in the text. • Study Tips
This presentation software allows you to create your own • StudentTutorial Service
Prescott−Harley−Klein: Front Matter Preface © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
Preface xix
For the Instructor: • Images and tables from the text in a downloadable format
• A complete Instructor’s Manual and Test Item File for classroom presentation.
written by David Mullin of Tulane University. The • Correlation guidesfor use of all resources available with
Instructor’s Manual contains chapter overviews and the text and correlations of text material with the ASM
objectives,correlation guides,and more. The Test Item Guidelines.
File containing over 2,500 questions,and password • Answers to Critical Thinking Questionsin the text.
protected,provides a powerful instructional tool. • Web Linksto active microbiology sites and to other sites
• TheLaboratory Resource Guideprovides answers to all with teaching resources.
exercises in Laboratory Exercises in Microbiology,Fifth • A Course Consultantto answer your specific
Edition,by John P. Harley and Lansing M. Prescott. questions about using McGraw-Hill resources with
your syllabus.
Acknowledgments G.A. O’Donovan, North Texas State James L. Botsford, New Mexico State
University University
The authors wish to thank the reviewers, Pattle P.T. Pun, Wheaton College Alfred E. Brown, Auburn University
who provided detailed criticism and analy- Ralph J. Rascati, Kennesaw State College Mary Burke, Oregon State University
sis. Their suggestions greatly improved Albert D. Robinson, SUNY–Potsdam David P. Clark, Southern Illinois
the final product. Ronald Wayne Roncadori, University of University
Georgia–Athens William H. Coleman, University of
Reviewers for the First and Second Editions Ivan Roth, University of Georgia–Athens Hartford
Thomas Santoro, SUNY–New Paltz Donald C. Cox, Miami University
Richard J. Alperin, Community College of Ann C. Smith, University of Maryland, Phillip Cunningham, Wayne State
Philadelphia College Park University
Susan T. Bagley, Michigan Technological David W. Smith, University of Richard P. Cunningham, SUNY at Albany
University Delaware James Daly, Purchase College,SUNY
Dwight Baker, Yale University Paul Smith, University of South Dakota Frank B. Dazzo, Michigan State
R.A. Bender, University of Michigan James F. Steenbergen, San Diego State University
Hans P. Blaschek, University of Illinois University Valdis A. Dzelzkalns, Case Western
Dennis Bryant, University of Illinois Henry O. Stone,Jr., East Carolina Reserve University
Douglas E. Caldwell, University of University Richard J. Ellis, Bucknell University
Saskatchewan James E. Struble, North Dakota State Merrill Emmett, University of Colorado at
Arnold L. Demain, Massachusetts University Denver
Institute of Technology Kathleen Talaro, Pasadena City College Linda E. Fisher, University of
A.S. Dhaliwal, Loyola University of Thomas M. Terry, The University of Michigan–Dearborn
Chicago Connecticut John Fitzgerald, University of Georgia
Donald P. Durand, Iowa State University Michael J. Timmons, Moraine Valley Harold F. Foerster, Sam Houston State
John Hare, Linfield College Community College University
Robert B. Helling, University of John Tudor, St. Joseph’s University B.G. Foster, Texas A&M University
Michigan–Ann Arbor Robert Twarog, University of North Bernard Frye, University of Texas at
Barbara Bruff Hemmingsen, San Diego Carolina Arlington
State University Blake Whitaker, Bates College Katharine B. Gregg, West Virginia
R.D. Hinsdill, University of Oscar Will, Augustana College Wesleyan College
Wisconsin–Madison Calvin Young, California State Eileen Gregory, Rollins College
John G. Holt, Michigan State University University–Fullerton Van H. Grosse, Columbus
Robert L. Jones, Colorado State College–Georgia
University Reviewers for the Third and Fourth Editions Maria A. Guerrero, Florida International
Martha M. Kory, University of Akron University
Robert I. Krasner, Providence College Laurie A. Achenbach, Southern Illinois Robert Gunsalus, UCLA
Ron W. Leavitt, Brigham Young University University Barbara B. Hemmingsen, San Diego State
David Mardon, Eastern Kentucky Gary Armour, MacMurray College University
University Russell C. Baskett, Germanna Community Joan Henson, Montana State University
Glendon R. Miller, Wichita State College William G. Hixon, St. Ambrose University
University George N. Bennett, Rice University John G. Holt, Michigan State University
Richard L. Myers, Southwest Missouri Prakash H. Bhuta, Eastern Washington Ronald E. Hurlbert, Washington State
State University University University
Prescott−Harley−Klein: Front Matter Preface © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
xx Preface
Robert J. Kearns, University of Dayton Thomas M. Terry, University of Ed Leadbetter, University of Connecticut
Henry Keil, Brunel University Connecticut Donald Lehman, University of Delaware
Tim Knight, Oachita Baptist University Thomas M. Walker, University of Mark Maloney, Spelman College
Robert Krasner, Providence College Central Arkansas Maura Meade-Callahan, Allegheny
Michael J. Lemke, Kent State University Patrick M. Weir, Felician College College
Lynn O. Lewis, Mary Washington College Jill M. Williams, University of Ruslan Medzhitov, Yale University
B.T. Lingappa, College of the Holy Cross Glamorgan School of Medicine
Vicky McKinley, Roosevelt University Heman Witmer, University of Illinois at Al Mikell, University of Mississippi
Billie Jo Mello, Mount Marty College Chicago Craig Moyer, Western Washington
James E. Miller, Delaware Valley College Elizabeth D. Wolfinger, Meredith University
David A. Mullin, Tulane University College Rita Moyes, Texas A&M University
Penelope J. Padgett, Shippensburg Robert Zdor, Andrews University David Mullin, Tulane University
University Richard Myers, Southwest Missouri
Richard A. Patrick, Summit Editorial Reviewers for the Fifth Edition State University
Group Anthony Newsome, Middle Tennessee
Bobbie Pettriess, Wichita State University Stephen Aley, University of Texas at El State University
Thomas Punnett, Temple University Paso Wade Nichols, Illinois State University
Jo Anne Quinlivan, Holy Names College Susan Bagley, Michigan Technological Ronald Porter, Pennsylvania State
K.J. Reddy, SUNY–Binghamton University University
David C. Reff, Middle Georgia College Robert Benoit, Virginia Polytechnic Sabine Rech, San Jose State University
Jackie S. Reynolds, Richland College Institute and State University Anna-Louise Reysenbach, Portland State
Deborah Rochefort, Shepherd College Dennis Bazylinski, Iowa State University University
Allen C. Rogerson, St. Lawrence Richard Bernstein, San Francisco State Thomas Schmidt, Michigan State
University University University
Michael J. San Francisco, Texas Tech Paul Blum, University of Nebraska Linda Sherwood, Montana State
University Matthew Buechner, University of Kansas University
Phillip Scheverman, East Tennessee Mary Burke, Oregon State University Michele Shuster, University of
University James Champine, Southeast Missouri Pittsburgh
Michael Shiaris, University of State University Joan Slonczewski, Kenyon College
Massachusetts at Boston John Clausz, Carroll College Daniel Smith, Seattle University
Carl Sillman, Penn State University James Cooper, University of California Kathleen C. Smith, Emory University
Ann C. Smith, University of Maryland at Santa Barbara James Snyder, University of Louisville
David W. Smith, University of Delaware Daniel DiMaio, Yale University School of Medicine
Garriet W. Smith, University of South Leanne Field, University of Texas William Staddon, Eastern Kentucky
Carolina at Aiken Philip Johnson, Grande Prairie Regional University
John Stolz, Duquesne University College John Stolz, DuQuesne University
Mary L. Taylor, Portland State Duncan Krause, University of Georgia Thomas Terry, University of Connecticut
University Diane Lavett, Georgia Institute of James VandenBosch, Eastern Michigan
Technology University
Publication of a textbook requires effort of many people besides the classification would not have been possible without his assis-
authors. We wish to express special appreciation to the editorial and tance. We also much appreciate Amy Cheng Vollmer’s contribu-
production staffs of McGraw-Hill for their excellent work. In par- tion of critical thinking questions for each chapter. They will sig-
ticular,we would like to thank Deborah Allen,our senior develop- nificantly enrich the student’s learning experience. John Harley
mental editor, for her guidance, patience, prodding, and support. was greatly helped with the section on bioterrorism by James
Our project manager,Vicki Krug,supervised production of this very Snyder. Donald Klein wishes to acknowledge the aid of Jeffrey O.
complex project with commendable attention to detail. Liz Rudder, Dawson,Frank B. Dazzo,Arnold L. Demain,Frank G. Ethridge,
our art editor,worked hard to revise and improve both old and new Zoila R. Flores-Bustamente,Michael P. Shiaris,Donald B. Tait,
art for this edition. Beatrice Sussman,our copy editor for the second and Jean K. Whelan.
through fourth editions,once again corrected our errors and con- Finally,but most important,we wish to extend appreciation
tributed immensely to the text’s clarity,consistency,and readability. to our families for their patience and encouragement,especially
Each of us wishes to extend our appreciation to people who to our wives,Linda Prescott,Jane Harley,and Sandra Klein. To
assisted us individually in completion of this project. Lansing them,we dedicate this book.
Prescott wants to thank George M. Garrity,the editor-in-chief of Lansing M. Prescott
the second edition of Bergey’s Manual,for his aid in the prepara- John P. Harley
tion of the fifth edition. Revision of the material on procaryotic Donald A. Klein
Prescott−Harley−Klein: Front Matter Visual Preview © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
V P
ISUAL REVIEW
The next few pages show you the tools found Opening Quotesare designed to perk student interest and
throughout the text to help you in your study of provide perspective on chapter contents.
microbiology.
Chapter Prefaceis composed of one or two short paragraphs
that preview the chapter contents and relate it to the rest of the
text. The preface is not a summary,but allows the student to put
the chapter into perspective at the start.
II 5
PART CHAPTER
7.1 Definition of Frequently Used Terms 137
Microbial Microbial Nutrition
stroy microorganisms is no less important:it makes possible the
Nutrition, We all labour against our own cure,for death is the cure of all aseptic techniques used in microbiological research,the preser-
Growth,and diseases. —Sir Thomas Browne vsthcaertii bolaenbd oo irfna f ttoohoriyds ,caahnnaddp h tteohrse ap prietrae alv l(esBnoot eixos sn7e .on1ft) i.dali steoa psee.r sTohnea lt escahfentiyq uine sb doeth-
Control There are several ways to control microbial growth that have
not been included in this chapter,but they should be considered
CMhicarpotbeira l 5Nutrition Sfcootarlmopnhsiy ellasor cwgoehc,ecgnuo sgl dareounwreiunsg tATioltnh oouf gmhhgdi rercmoe rswcoashonteahrysgp, tamtahenneriiscd ss rm uoicnbsoo ,jnrePgatcara totrn otoli psfoI imItfc h samoe r faeni cr ioemrcno obmsonpercengeenecarsfinnfieicie scipdma cr lasow ca.n itnTttirhdchoa ilntls h ae iecnmc hednpas duospaetrtretrsiayrttnr i uaofcdocne--r,. (cfttresohoemrele mlpa epe dipmcr.a a.Ctolu1 rahr2egea 1,epc–nOto3etm12sr , )pi6.an l neC dfdtoehe so arcapdprdi tipceibatrreu etsi4rso e1etn rh ovdeofai n steich fomufoneswi csc( etrssmose beoti ihcfpa relpoo .sguom0rsroge0ow a0toni–tfchi0 s p0maahc)ns.ytd isv aisircuteayr ,lv cpiaovnHnad-l,
CMhicarpotbeira l 6Growth ohdabnuiss mebcaleaossneos sped sas a,uthgbcoaahrgc .a teTesnr h beciomsaiulisas,e,s hsseeunqmtuiaeannl ctwoe seb,lels- aubbcelhien gtaos, mkfioilocl rdao wbspiiadolei la avcgatiervi eiattniyed so fdm imsaeyica rhsoeao.v rTeg huaennridesemfsoisrr eao bri tli eni shc oiebnsi--t 7.1 Definition of Frequently Used Terms
CCohPnahtpyrtoseilc roa 7lf aMndic rCohoergmainciaslm As gbeynts O555...123uTRNRHMtheeuyileqqctdir ruurCinootiiirroogoeeermemmngnamaee,lnnn aoTittnsnyssmd pNf 9oseOu6srx t9Cory7ifaegrnebtnon9,6 C1.oMlcniaanonurn indcvgcsl eercetcor roiqyucofu ecrasaagtcmc natpciotnadairlitrslsstib. e.m aosSsm,hse iyvornedeu qrrpnaauatltsir er otase epatb,n hannoeldeibcddi prsaoa o ioepruudcelnnstsea e,pie1mrn paud0tgherr mitoenet,silytt poseeso ,mhn iafanfr aireeosaere,nn .oy uatzndnssnye egidmednidt eeitsosd, ttacsuFhwpnosreeeodnoic m trfo(ai oT2ffm gli h)tdcarhii n:otnsepotw (ac ih 1mtrbhtyie)hoeasid gtpcniouctrit o onecoad relfnb e m fiiwaosaonntirlacgrn dtuoce ie mholyscrieef ,mhip smfzer iaooemenoto nchatidt ohtocthe,hreage daerlmei ane rncap ddiosgcdy neorhe aotnttsniorhotstod sretrcl.uo gr opo craCsnryftnueih t,mvirbvasoepesmpil etcn taemorsetnof rprftic ohelec3eerscer5gs opti. hasrboi .nan intvairtTssarleim ohnb ddesplsiem usr be agcfiyaceooss tssanrie i cloott. ehhnnidees-, Tctidjaateiinlbreoeslcoglpe rnteo mmpseb or,Tntpeiigiflcndhrcc iatarkoai ehnooncu eleioaftog ss itbmergc meegoaiay nliaantslic in t a npriyiitirsomshgas e tbdm ero puetiu ii aocssstsccelpc oreou fntpuendarnlsocsoadn tilspirrmlicoaeso tuteiol d rlolla. aseay mnbnenSat idsel toioiy.mocc mmns.arb ueo Tpumjenreboshtgctiaierem iatyct c ,wlasa ae bwnipnolste tou rhi eirdnpatewie stsutrqiiht ehlsolruaeia eunkntirsnmir ee oiie os ncenddt nehhnes iilssiv etnyobsise n a,noincptrfi t oy eaishgmnnc r erottatotoroiarfna wrs onileecim lttt lod hucianaome aonotssftnediir itnodr f mkoauuaneinbcstrsile------l.
555...456 RGUPCheerpooqltlFAGawsuakpcri1tcorehth0iieu lvo om0ipFetrfa uaTe TtNcsenrr,tdaatuon sanDtr snsrfslpiidoofeofc r9nurS at8Ntstusii1ool ibft0nnruy1or 11gt00he930en8, 23..ANnrasttihrneetuaoluvqrlltneoomtu rrmcsiuliitpvtsregiiie ooivnchonmrnert rgt opal emeey oaldtnl hsrocep tgsbesalceiea yttvrfuecrn moeosogui resnnodle me csaeriosfa.ib sffe orf lu sbmfcers eoa otipeqhnonnmlur n,a bete.esbhee nmnTre tampe hlabnryle agaaem yjccys o cae,iemrsandma rmn uonribosnfideer tt t acr ohhbc nhnypeereaedoirsnro rsoiotssefgm iaen snsf.ew dcsacctrnooieosdmnrioi tnlamtrfihgmzeaeaecin n tnGidiioansertmnfedee ed scatk h tnwwsieod ibau Htushss etr letnmbohrernieaedlg itwl ze esaurspuit nartliofooslu u snabrys,n up tbderoenva cdf ectauientnsemd iyrtn.iih ucfgToelmahouctette.g ha ThnEbio nutgtdsghiy al etsdpyo uit enisteaxhpgmnieesss .bcta etaMbuenlisdmcloei eostd y abf o f iotbcifodr e e lmiidae netwsigo--, ogaSivscabit rhentjoheriiiesceidlvmt isp ezo srda(ro,s tbcsehiopeeyasno bcasrih[ etcbLaashyt,ap.e tatwAmiennrh di ssci 1tctoaee8hltrr h) iai alelagelirlsr e oe,inln ibutvef,jniieettnacchhbgtteei l o rieccs u edh tltsoeelo ssmapt,tagrrivloeocliyndyaa tulebsf dcirl.se ee W o e csor apho flrfoeflese rnpvmed riss aioa,tnbev vgsrleitei dreol uir mrzifs labreiaotcasnimr,ortor.nae onIa nirnnds-]
Iron Uptake104 Eucaryotic microorganisms also employ
5.7 Culture Media104 endocytosis for nutrient uptake.
Synthetic or Defined 4.Culture media are needed to grow microorganisms
Media104 in the laboratory and to carry out specialized Box 7.1
Complex Media105 procedures like microbial identification,water and
5.8 IsolaTtyiopens o off PMuered iCau1lt0u5res106 fmoiocdro aonraglaynsiissm,asn. dM thaen yis doilfafteiroenn ot fm peadritaic aurlear Safety in the Microbiology Laboratory
The Spread Plate and Streak available for these and other purposes.
Plate106 5.Pure cultures can be obtained through the use of
TChoeGlo Prnooywu Mrth Polr1ap0the8ol1o0g7y and srmepqirceuraiordbe dipa llfa otserp set,hcseite rcsea.arke fpulla stetusd,yo ro pf oaunr ipnldaitveisd aunadl are Pelharabsvooenr anbteeoler insea sfa.e ctIqyt u hsirahesod ub liedne nbt hee eso tfli ammbaoatrjeaodtro tcrhyoa,ntc atenhrdon u misnaa nandlyls mpoeifrc sirnoofnbesico thliooagvnyes sogoaf msdeeios umesaa ssjeuo sri pspe onthtseeino tinina ohl fah llaaizqtiauorindd s oo afrr esao ncl liedina prf.ea Ocrttinicoelu eossf tathheareto mmsooal.sy t A bfrene gqaeuenerenortsa octlea duis sb eyas
died because of such infections. The two most common laboratory- accidents and laboratory operations such as spills,centrifuge accidents,
acquired bacterial diseases are typhoid fever and brucellosis. Most removal of closures from shaken culture tubes,and plunging of contam-
deaths have come from typhoid fever (20 deaths) and Rocky Mountain inated loops into a flame. Accidents with hypodermic syringes and nee-
spotted fever (13 deaths). Infections by fungi (histoplasmosis) and dles,such as self-inoculation and spraying solutions from the needle,
viruses (Venezuelan equine encephalitis and hepatitis B virus from mon- also are common. Hypodermics should be employed only when neces-
keys) are also not uncommon. Hepatitis is the most frequently reported sary and then with care. Pipette accidents involving the mouth are an-
laboratory-acquired viral infection,especially in people working in clin- other major source of infection; pipettes should be filled with the use of
ical laboratories and with blood. In a survey of 426 U.S. hospital work- pipette aids and operated in such a way as to avoid creating aerosols.
ers,40% of those in clinical chemistry and 21% in microbiology had an- People must exercise care and common sense when working with mi-
tibodies to hepatitis B virus,indicating their previous exposure (though croorganisms. Operations that might generate infectious aerosols should
only about 19% of these had disease symptoms). be carried out in a biological safety cabinet. Bench tops and incubators
Efforts have been made to determine the causes of these infections should be disinfected regularly. Autoclaves must be maintained and oper-
in order to enhance the development of better preventive measures. Al- ated properly to ensure adequate sterilization. Laboratory personnel
though often it is not possible to determine the direct cause of infection, should wash their hands thoroughly before and after finishing work.
Chapter Outlinesinclude all major headings in the chapter
Boxed Readingsare found in most chapters and describe
with section and page numbers. This helps the reader
items of interest that are not essential to the primary thrust of
quickly locate topics of interest.
the chapter. Topics include currently exciting research areas,
the practical impact of microbial activities,items of medical
Chapter Conceptsbriefly summarize some of the most significance,historical anecdotes,and descriptions of
important concepts the student should master. extraordinary organisms.
xxi
Prescott−Harley−Klein: Front Matter Visual Preview © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
xxii Visual Preview
Additional Reading 151
Critical Thinking Questions
1.Throughout history,spices have been used as a.Choose a spice and trace its use static agent. How would you determine
preservatives and to cover up the smell/taste of geographically and historically. What is its whether an agent is suitable for use as an
food that is slightly spoiled. The success of common-day use today? antiseptic rather than as a disinfectant?
some spices led to a magical,ritualized use of b.Spices grow and tend to be used 3.Suppose that you are testing the effectiveness
many of them and possession of spices was predominantly in warmer climates. Explain. of disinfectants with the phenol coefficient test
omfetemnb leimrsi otefd t htoe pcorimesmtsu onri toyt.her powerful 2.Danetsimiginc raonb eiaxlp aegriemnte nist atoc tdinegte arms ain cei dwahl eotrher an and obtained the following results. Critical Thinking Questionscontains questions designed
Dilution Bacterial Growth after Treatment to stimulate more analytical and synthetic reasoning.
Disinfectant A Disinfectant B Disinfectant C
1/20 (cid:1) (cid:1) (cid:1)
1/40 (cid:2) (cid:1) (cid:1)
1/80 (cid:2) (cid:1) (cid:2)
11//136200 (cid:2)(cid:2) (cid:2)(cid:1) (cid:2)(cid:2) Questions for Thought and Reviewat the end of the
What disinfectant can you safely say is the most effective? Can you determine its phenol coefficient from these results? chapter contains factual questions and some thought-
provoking questions to aid the student in reviewing,
Additional Reading
General Henderson,D.K. 1995. HIV-1 in the health-care Widmer,A.F.,and Frei,R. 1999. Decontamination, integrating,and applying the material in the chapter.
Barkley,W.E.,and Richardson,J.H. 1994. setting. In Principles and practice of disinfection,and sterilization. In Manual of
Laboratory safety. In Methods for general and infectious diseases,4th ed.,G.L. Mandell, clinical microbiology,7th ed.,P.R. Murray,et al.,
molecular bacteriology,P. Gerhardt,et al., J.E. Bennett,and R. Dolin editors,2632–56. editors,138–64. Washington,D.C.:ASM Press.
BCBlleooncctkkemiaaReS,r,nsndSnoSei- dicd.fccc.tor hSo ioSFoperir.mb et. resD1,yefib,mo,9e sii7fl8de9sgoeo1er72ienrgt5va–. rody M.a–s1S,raet3.0t1it i e41oi3Ccsor.9nr.ti o nolWS9,einsbzd41a taai.fnrt.o,st oho iDvlhDro loeo .iinpi gndsl1e..riyg .g 9neI 4.tPnov8fo,e :he7nJEcAni..,l nttLaicDciodaoeyn.dedncC,leel p.oosmr:hbtpfeAi ieeHcrardm ig:PIlia,iVLerzee reaodsiatcfsiiat.oonnr- MPearkrtiinnNdIdaCsSnstn,iiphee,ssdMrPrwuieJi nia.rnrRl. fcYiisJgeAzn.he. ofca cDisi1.rtltei,,ikli9polo oa4:d6Lllnnntei,C9hni,d svI.i h a nleeiaWPlunn dd.nh:rrdg.eidcie,tC snnod hatGpzchoriilrselsit.anlpahp,lr Le,Llc ol2.see.Rts5i csisvMac.7i ialeCeP9n ann o.–.gon c df1T8sfd e i9t7mhiesnon9.,lofn elf5Ne2m,ete.chdcJ. eatSo. twiesiodtEo.de usYu.r. sisBool iwfrezknaa:ntsiteoetn.t,, 7BSø.r4orhcakSEJDgu,.uciTgn TiLniei,c.edhgeyT nDeCodce.c :le.a1 e oorA1nU 9bpTn9d9ceese8 a2trdrce3gdre.ih a.,feoT o e MemPeolrfdumefi ei ncbPtmmpo clPhieibresc-rr rhyiearmanoestsnua-srbicse.rncih.e oufa iilcaeloltolf gr.,MnayM2t,tr0io1eoa1lsdnt.–t hi: I1senAoo1dn .d.u ,,SssvWae onrli’.ss .4:,
CCCCeeeonnnlltttieeenrtMMUtvpMpihHbDMBrrsmrrsssreiiaaa,u e.n or eoooiaftCfmfnnucCatvhiomorrrlotvsrslt.setotterru.t:hroammee,aa nhr gnDHD-UdbrlDlhlwt.c.ieiso.. iii ises. i.1aoWacnoioWWds,Sssspsrl9aneslereaiiee e.aoeeta oo9l pafne aiohG teegisan2nsnlrikdcsfsaekkentei.o e li .ic oillhbtc yedLsBCv nyyCrafCa oen aey R i BplouoaRRrchnohr onuenamlyntn essueenteivpbtt mmathaoppmreov,tri.lolfr-nr..otiriPitee3occhualrshis33l.tn.8ua-el nsos y -67as.Mt1fri(ss, c d aonoSe((i1i ,9PaaamsnS2fnd r.u938renrs, n 4u 1iepdp8m9emtoNd4dn)yp 9pt 2r8.f :t stie t ueapoh 3h7liGc.wie2hd.n vtnnlet7 r6 123tUiu..he6ugoe4otpogo 7....iWmd2e)in rpbansdn–dO:kr.)t.tdIMAtsmopeai1aeg a8e ai:iooenBiffnbMlafa–to3fnslsm8rlff li a iiattdl.3tInScsosihoibscn.ehco hconnc. poi7M gis–iie B:riwoemooinrsyeoMtl.nnoise1.tboedfCcug n os aisdmnpvi8rfoiantbygtcrmtyedohu ciloSrrlb:yonpohrespy ru.tanbup.airt re l ,pecdmshm,ienseosndnshtt .cece rdt eoo.aodaucotsuranmnfreiol rsneey1onsm e sto 0omte c,lprcornyasle elfza at.er lraMyrntsxlWRSSPeghb .mag cetiueeomerkbaowu ieastlreoprnliaossire ac,lbnoese elilt f7fmpWSedei8LmIAReto,llslnnaenoonily,ddc5ri(mpnaBntl,d.,zisafrr3coaimuuD.i–cbrneyd sEA. iM nti.s,)citlMeOrmn8odcoum .:nheeha.Aoecn.a3ft5r er..rsLxggeied:e1ibi nst Dae8sn.n. To1cR sfecel.u, .9fiat ss9W gloc r n9o.crmotre1hra as18ba;oi-oe–taros7ler v onkr9eoV91odaobH,eurmy4 .sne9nd ea9n9. els:ngiefs 0 U Msi.u on,hi sa 2,5Le rctyBoGt5nssLag.ldltpir., es.iem afoaa.n e lfueoIc rracd6PLpabeglbgnngö ecr,baliitcoator.aiyioet tststhWlyooknMiioroefbr.elrcbe n a reasgwidnerrhoaEi.saccta,artvdyo,ne,er,oteaBnifn7aao.lpDldcelrl,e.u,2.ttlr yla t;ori Pm.3eyiaD9y SoCu gnar.ss3.-l–inysyc .f n aa.Rc:C4oae:ei -H4sfdraeisa5.fncAelsno1 n,nis.tA.Motdcn b– imy11stadMBlNc oiy.io6.ip01n9 uf io uneilsMice6aIr..d..orilcrtsin waitf.roec 1is aefia ANpMaisbgs(ccywPgracLe9s tdfuyyYnarlytineiyru l,mi9uaia el,ilngrndto d,oictibcfGvet5srtbnehtzu ambehJraartlhSide.aodfke.eti.or nimeteneuoecoootriesA:eAoo ittci aatcfac nlasooasrriri.tt..ecti toegigs snirioe1eaRdJto,od enaa,iti r0 . nvne–yudnnn.n,ty2.vosisgceic1iidt.sc:t. tholy0rsmiu Soyro s–utopugdsn1alufof lerm.0 cifomrtnda.araa1g etantct7BBMRRe2eme ne ad nu)d.eouusa ncu.5sasa lsrstaD tykerasim sbhicsren ileonecrnaukDtreeSRars3if itylonon gyoeaen,phn,lcdl,(dtiTo,eufscnso sr2rWaooandSiPuiaui aiosAtitgersr)slrts h.se.lsonsrg tedo:uhel.ihe;ittsh .CiM9.lrc dtaaceasisgdAe n, Di,i Dssi9an entomne by seGdai1lyo.g.s–rcUuiuscbn y .y,,na3scss1ial e..1nkeurf l1 mau,ltitd 39a.necsdaC 1ameahrn9tatat s–cCe7tgdeineamn9eoyrndlo9gsstis43, tmiid ,. lpaliido ir0ehn.tfoi2Wa.nmPn iSnyhonr n.ii.Snlt.RCfff ea onte..co tiBMeeegsu ;W e.slrnehuhM r::Cccba l.L ayibsleif cttADteanocCciaisieemahhrcsooteerrtcvod,clhelreoiunli,etioiwttoDlrciinb,s:ee,vnmirbaY. adir .iAMi gd(ionliltMe.Jo bati yi;l.es.nsosol .lcl, t.iD.a p ee1,nl e cRsaanIeRaao9Hr,e.rcnec dcls1roie9n1Dt v luueehi v6MbAi7s9.Tos,lt.z.)i.i.acC91n o atoEi1 agirihU9c0,untFiwrcmsipd.lv2n .in:nier.i.aay( ao riaoi(s n bdoRih4oatfAApRntont1nnettnesofy bile)shtdpotc)tssee,oorSny.t:s eiiha: thrsm5avrnaDens1cnt oiMrloeee.ianen9 uital 4 yfsteinac.nhacmd.r7cas e7 ld ctn1bbi-P- taDs–p –dcaihieCd9aewerccat67dasdeoemri9 iaot 1cars9gdds.suea9leAysy0o.s,e.srb. s.sy.de P,lat. le eyasra pmenyrfbiht dnomae.o tnrtNDhsit ddtapeAhuscishner hiDcai zeensta ePekdbgnd He i a t osblofn eyserbd toynt oy ahnmtan dhetts hh de phaaeeseaacci sdrssefyit a stalbityt nthesctely teea-,ta CnwraarrtoncioseiAddr, 123...DaoSmstAldhiunenupiceegsds pit dccdacpi piaavrru,euosintansabam.sta sntebeeIob nd dotttstohh h hlegplceiaaseiseea tst smr n aprmceaerento?l iir.mdcaend aHatr tolhiisar coogt erierpninxwloeryaseomor hrcdsdsrtaise igopaac nsahem ni bnipossdtiie ywa nsa Qrtttnmenow hewtqa ueh aeiuwboineceerious iran csdreills sidoatsdc s.g rlia? mibeboBrttuos ao snr.tdwbhyi enseoynifpon ldtlfihegyrsoen amorsdtnei z,T eahough456t... aWaWinWpcnnpoietthphhpndoroieyats cictatg dRrh cereiuea lsnttescln ow u tvc oti pnhoboofihua tenneeetos s?w nsnusttpzia pacteykluha cmeeaonisnbfrne iutposocsslu t da tw?tboci tseh ehtc ehdedurei usesr saCreyeiscnnad ttslgl hvyui nepi lnifs entui phcscrte y ooia cdrcfnplohedoga?rlpayttceeardn 12..IcttyewaIwmThrnnxonohaChheet vaauecimetsenmbaarkrrnetmk leboh r eipo tatoloetoenhalisflrioesdfbeti c tmclsimiytoshymah ao bole rc eioamlypklfmsayo t r eomnTttec c.yaharcn a k teTc reahoirao,exAhmrsthtnsiani ae eediinebtbp m nnd rardcesooskvik apioot lrvrot aoriilte uiaeselrrtano cvbsheircmsnyne yge o aectiorca. dnr uo slflcilrQ a etn n ipaetaR rdeshta lrngeleru l eieib sdehrqmmsdame,e ouerdoicaais tnieetiecaernstennddious lgidlcig i insoyfpc.n eaes s alsnLottui eaeo otntepqiecss fsnlmpu i toaans learsyaugnee r—e2cfk.ee2ws5.
4.Macromolecular components often undergo nitrate reduction catalyzed by the enzymes 17.Triacylglycerols are made from fatty acids and
self-assembly to form the final molecule or nitrate reductase and nitrite reductase. glycerol phosphate. Phosphatidic acid is an
complex. 12.Nitrogen fixation is catalyzed by the important intermediate in this pathway. Additional Reading
5.Pthheo Ctoaslyvninth ceyticcl eC aOn2d fmixaayti obne idsi vciadreride din otou tt hbryee nniittrrooggeenna isse r ecdoumcpelde xto. Aamtmmoospnhiae,rwich micohl eisc uthlaern 18.Pcahno sbpeh soylinptihdess ilzikeed pfrhoomsp phhaotisdpyhleatthidainco alcaimd ibnye General Yoon,K.-S.; Hanson,T.E.; Gibson,J.L.; and Current topics in bioenergetics,vol. 16,
67..GGapgfnpp1drnrlhh0uuollueduuaa.rcmc4ci sslccnruoe)eeoo e.gortns, nlshsT :saeaetoeeithohtn,odremhegrsdfgdeee. re; e i uet dsnncgnh uciiacAeaeptcgrosol Th barairisesorPnopces gsttxosi, oespef yraa rrsnhatlmonnehetaamdi dertteioiea s do mnts tnwsdyiin oaoaoyne otnnpnfnrte hginh Ntosvpheloa nauAhoesssetciarseeiDs ovis, Cm zaeosePteOer hsfaH(ed .edfg2 isB e.wlgr u eaaudicdrctrioehutree secuerctsiitaeolydn 1134..AfaiaAo(iwMnnifnnfxinhmctfaadgea aoinpfurltip1reirynmlrpoole0 oeieoanrem.t r1 on rd a1osra6t aiceu0ttita)pichiaep..ctd.l1ecdep e r ts7 blrciei io)oniteea.ootntsncio ss ctktby i.r ae oeianTmoennlthp shsziaye nyetmr nhotmegitepp clhaey lh eccsapoiyti cbaixcdccetoaysl h t eTpll(awia rfcCilteinya egAcp zy ubucaes yrarct shCelbycaoswlrcOner al1sa ce2ni.0yesc. sh1a4lso 1290..PPfapttafazhroomocnreeaerorrndppnmmio cetnlts1isieisioepdpdsn0 sd ioo iosga.ntd rg2 hgc b it Clt9inelscyyhdy) vDa ceNc..ctroa reParcAlanlin-velen dM l isrsmslniy apyb-wgnecNena tymapbthcAllhtotelgbioe.Gtsdl rhspEyiaa-si rcnpxst Ueoie eieonisrcD.nsont oc itCPla nula,(, rgrpfctdowsih eoge pseiphmrusenntii-p rvic plddateiahliinedstedsi k dcxovu1sirgen0ne asli.gty2t er ascea8nandn GGLDCeaaaohlwrtdrtnseewicpSBN1mPtsnteh,h9cu,gieelaoIiy9bRelwte.l,csa3rlkn.Wh,iiD b.Y,ostHAeoiPh.Gl.omf,ol.i.Rirr,in.acgkiisL a.sng1n ym: P.tn2,c.9d;rS u di0C28 y2NppSb ,0d6dGolrule20e . iime lterncds.Bshid d ag mosMoeae.t.he n rifdcBuBiroa l,ctb.-anmnoe eDVrinNirssoolc,edm.i.tebcaaroC,LwhlitolnIaai. .e.on m,;glYMm W ntMa.pe,osin.thCs.r, aa dk1ty1Isarb :s9ns9CyloiiS9.c,9o:fol.9a2i.2lBx:sou.d.m,S gln MaMeyt,dcad i2eakr.c.rd nwMrsN od.eeeb.dlwil.al 1BD0reia.ll4n,,mmeAANTWDdamiice.itbc.aPTttr.JriaoRod.to2hbh rage.1b3-o;,meoei 96inlBnFos i7(iA-.asl3cop7comsR )l.hsehPgi: .n.6Bi syr ome,2I8ei,ifnmroJs0–e2, .sul08tdE3o.aTi10 s4lglne...la,;9ic od Acat–Syc.ai,nc5luuloudvl 8tonnsol o.Zftpi tle tSur.ehror oaod1esrfpng:in,, ah IeSgJDa inn.,toc nri LLf ofeuCdige.crx Odo t1aguNe:9t2arir9eoibn3tsner,.i.r cogS,gcie.n 1DMPe0ooty.er1lares,03i5NbM,,nJ6i4RJio.i9t(N .crc13.–Wroeh9PF)Jo9gu:e.9b.2;0oaer;m 0i 9o.CnoFrt .i3sS atlhcimGp–.sseaaaoeh34nlrllau r-eo09 sngaDtrut4:ta,e3r sip.icmuKno3e krcoeg5nia.atnt;of–us,i ea c:s6rJC na eA.6mp; d.. a eecae MnrDalntsldadidpe cb eW aferVomuncolint,biianisccvDitml oteePli.lro. r., Ra neARVns.:n d.es1A nv. 19c.u9y9.8 c5R8l.ie.nvg.
and algae synthesize glycogen and starch from 15.Purines and pyrimidines are nitrogenous bases is selectively degraded by autolysins so new York:Worth Publishers. organization,and synthesis. J. Bacteriol. Turnover of cell walls in microorganisms.
8.Paodrhgeoansnopishcio nprehu ods siippsh hooabsttepa.hinaeted gflruocmo sien.organic or fproiuburoninsdee i5snk- pDehlNeotsAopn,h RiastN es Ayann,tdah neidnsi ioztietahdlel ybr empgroionldenucincuegles ws. iTthhe material can be added. MMaanthdeBLewliososntc,adhCmoe.n,mK:Ji.Bs.;, tlMraayncc kdoQw fv ubeaialnlllc eHStneco,riilKeadnle. t;,gi afKrinoc.dw PE Dtu.h ba1,wl93ic9dea6 set,.idoI..n 1s9.82. Dilwo1l9re7(tg1h5u2,(m2)M:15e.) 1,:n69aon7–dd32u 73Gl–.el 4we4no.nr,kA? .TRre.n 1d9s8 B4i.o Hchoewm d. oSecsi .a HarolMtmdh,ieiFcc cr.rooaMoubrs.i goe1als9. n o9Rifs0e m.mv .Tso5o. r2 Mps(hh4ioac)prg:5oee5b na4ieo –cslie6.s l7R lo:.efAv.n inquiry into
Biochemistry,2d ed. Redwood City,Calif.: Glenn,A.R.,and Dilworth,M.J. 1985. Ammonia 54(4):381–431.
Benjamin/Cummings. movements in rhizobia. Microbiol. Sci. Höltje,J.-V. 1998. Growth of the stress-bearing and
Key Terms Moat,A.G.,and Foster,J.W. 1995. Microbial 2(6):161–67. shape-maintaining murein sacculus of
physiology,3d ed. New York:John Wiley and Howard,J.B.,and Rees,D.C. 1994. Nitrogenase: Escherichia coli. Microbiol. Mol. Biol. Rev.
aaandcyaelpn licenarerortiiec2r 1rpe7raoctteiionn (sAC2P1)6220 gggllluuutttaaammmaaintteee dssyyenhntythhdaerstoaegseen2a1s21e11211 ppphhuoorissnppehhaota2idd1ei6cn oasciinde 52′-2p0hosphosulfate 210 NeidhSMmaoro.dn l1tes,9c.Fu9.l0aC. r.P ;a hIpnypsgrirooalahocaghmy. ,oSJfu .tnLhde.e; bralaancndtd eS,rciMahlaa ecscse.hl:lt:eAr, KnowoAAlefn snmn,uuiRcc. l.rRe o2oeb0tvii0.od 0Bleo.i- goNdycei,hptr2eeodmng de.eed6nn.3 ,ct: yv2moc3oll5e.l –.e3 6cI,nu4J l..Ea Lrn ecsdwyecirltobcpehre.gd,ia Höltje6EJ,.2n JL(c.1-eyV)dc:.le1 o2r8pb01ee0–dr02gi.a0, C 3eod.efil tlmo wirc-airlnol-sbc,ihobilaeocfgt,ey7r,5i2a9dl–. e7Ind1.., Svaonl. 1,
assimilatory nitrate reduction 211 glyoxylate cycle 216 pyrimidine 216 Sinauer Associates. editor-in-chief,379–91. San Diego:Academic Diego:Academic Press.
aabusastcoitmolypislraientnosoryl 2su22l32fa1te reduction 210 gmmuaoacnnroionmmeeorlse2c1u72l0e5205 rsthiebylufm-laoisnsseee-m1,b52l-1yb7isp2h0o7sphate carboxylase 208 WVoheitt,eNpD,rDe.o,w.c aa1 nYr9dyo9o r5Vtk.eo :TseJ.htoN,ehJ epn.wh GWy Y.s oii1olre9lkyo9:g 5aOy.n xaBdfn ioSodrocd bhn Uieso.mnchiivseetmrrsyiis,tty2r ydP oreefds.s. KuykePaEnnrneddcsa yEslc.ll,lkoLap.neD,dGi.a;. DoHfa. d m2si0oc0nr0o,.bR Ni.oiBltor.go; ygH,ea2nsd hf ieexdma.t,,iovFon.l.M .I 3n.;, KNoanchn,iYWnAogo.ralkL,d:Nr. iC1n.;h9g Wa9hp5,im.Ce Bna.tanLjec .&st e1, r9HFi9.aa2Bll .lg. .;Er oMnwvuetlhldo aeprne,d Eg fr.o;o rwamntdh. Ninew
Calvin cycle 207 nitrate reductase 212 transaminases 221 Zubay,G. 1998. Biochemistry,4th ed. Dubuque, J. Lederberg,editor-in-chief,392–406. San Escherichia coli—Spatial and temporal
cdcCyaiOsrtsob2isomfixinxiyelaastotioom2rny1e 7ssu2l1f2a60te7 reduction 210 nnnniiiutttcrrrioolteggeo eersnneia ddfseiuexcatta2is2o1e1n73221122 tttuurrrairaannccsoiyplvlegeprl2yti1cde72ar0toi5oln222023 1Sc0h.l2egIoewl,oTaHf:h .WCeG COP.,Bha2/nModt cBoGsorywanwiet-nHh,ieBllt..,ice dFitoixrsa. t1i9o8n9 . Lens,DEJDP.n .iiL,eecaggeynoodcd::leo rAAPpbocceeaaldr,gddiLa,ee emm.odHfiii cctm.o PP2irc0rr-eeir0noss0-ssbc... ihSoilueolfgf,uy4r,9 2c5yd–c e5lde0..5 ,I.vn Soal.n 4, ofeUudrnngiticavotneriiosrz,snai1,ttyi8So 5.Pn –M.r2 eI2osns2h .P.a nrNo,ekCwa.r YDyoootrwikc:, saCtnraudmc Jtb.urAried. g aCenodles,
ffaattttyy aacciidd sy2n1th8etase 218 npuhcolsepohtaidtaese212710 uridine diphosphate glucose (UDPG) 209 ATeuctoht rPoupbhliicsh bearcs.teria.Madison,Wis.:Science Ludene,nPe.rgWy. f1o9r9 b1i.o Elongeircgael tnicistr oogf eann dfi xsoautirocne.s I onf
gluconeogenesis 209
Chapter Summariesare a series of brief numbered statements Additional Readingsare provided for further study. Most are
designed to serve more as a study guide than as a complete, reviews,monographs,and Scientific Americanarticles rather
detailed summary of the chapter. Useful tables and figures are than original research papers. Publications cited in these reviews
cited in the summary. introduce sufficiently interested students to the research
literature. References through early 2001 have been included.
Key Termsis a list of all boldfaced terms and is provided at The reference sections are organized into topical groups that
the end of the chapter to emphasize the most significant facts correspond to the major sections in each chapter. This
and concepts. Each term is page-referenced to the page on arrangement provides ease of access for students interested in
which the term is first introduced in the chapter. particular topics.
Prescott−Harley−Klein: Front Matter Visual Preview © The McGraw−Hill
Microbiology, Fifth Edition Companies, 2002
Visual Preview xxiii
11.3 DNA Replication 235
nucleosome.Thus DNA gently isolated from chromatin looks like Patterns of DNA Synthesis
Rmeavjoier wse Qctuioensst.i oTnhseasep pqeuaers itnio snms ahlell bpo txhees s atut dtheen te mnda sotfe rm thoest aoptutihhn sarseegeoit r rms1fmsioh.n1e ilaHg.tds9po ,iiobessntf)h itgos o.eb fWna oe letnifahnh ddHDeke s mne1vN. r iTfae Asropihieol bpgedislne iieisontsa to cnrgr( seh,s m rtertvecooeoaah m rarcf ieiosheog sfsecsuo sDooirc nmemaiN a l4mpeetAe.snl2ae gwsb0xxete)i ihem.ctt hwnhfu r rteiomhonemem n,e l tau1tihhntc4eiekan t cer obyshr e toorrraoetvudigmeccsir t oacu o1tnerri0elsn ln0s s tt u o ab(dc fkaauilesgeirdes--- WApTfnbreoahopaaswemrteryis e ll o p dihon1na yne9 iiaprne 5uino n3pdawtg.h n a—ChAeolisortclihAimnzhcegek orwd t tshph atiteu tehnhyeb adt x lwtT si asstu,ochhegt Gepeplg ydaa tew wr rsotaehtoitnnetee hetds ia tr (rlCm fh adis ognoet(rwudsnfaicsrt gnhr eDoudi pfrslN1 tae tit1t Ahohe1.enr1r 1o , m0d.ou7a)ofig. ) gu s.DhF hbe WcNrtlc eeooAbh emhne e ed nsnpl rtu ilrpexetcuhpam lcuepleitneseocuenwratr itetdniaae neupridndys--.
sweictthi othne’s cfhaacptutearl. material and major concepts before continuing 123...DWeWWaDuneNhhcdhsaAaa acartttr n y iaiadbtsorrio eetepoee ainrshsngr .uiai astcslt nlolmoeeimnzilec?eeea dsa n d ca ietinndot das tshi ?nlae Hyut hc cothelhwe rasoo ttdmsr tuoohocm eDst uoetNwrsme?Ao e oD sasf net otrsdhfac enpRr dirDNbose NcAa atrA hredye idcof fwooteemuarsb y ipaln lein en sdhm twreuelhicnxittcu.ahrrey? crsCasoetolnrrresimdaiguocn iltekRentid ,d’uo..se e ec fphasSR alE cyryaiehcy.prns a eoocetct htotaloiehioelrnnsiecsn.tkshii a ssieF pis dnco iaoc niottrncoto pegcregsirru xueeaneradbt csshpm,st e.aearaprrqert eelbp uen tyle,sthin acooewltamn Dthreiyeeoe NewonnparA hlr btmisa hces teaotg hr tdriaacieninosi fser dfn cneap uarztrfe nlyooaandmvr rts e k aieDdin ,snn g,N teWlthpwAewer aolp otyccpso rhoaflieaonrrnpoycrt l,memoiact tnheoeaaddess-t
which the DNA helix is unwound and individual strands are repli-
Numbered Headingsidentify each major topic and are used for 11.3 DNA Replication cthaeteyd h. aTvwe oc oreppielidc athtieo nw hfoorlkes r mepolviceo onu,ttwhaatr dp ofrrotimon t ohfe tohrei ggienn uonmtiel
easy reference throughout the text and the accompanying The replication of DNA is an extraordinarily important and com- tchaatito cno nfotariknss mano voeri gairno uanndd tihs er ecpilricclaet,ead astsr ua cutnuirte. Wshhapened t hleik ree pthlie-
plex process,one upon which all life depends. We shall first dis- Greek letter theta ((cid:3)) is formed (figure 11.11). Finally,since the
laboratory manual. cuss the overall pattern of DNA synthesis and then examine the bacterial chromosome is a single replicon,the forks meet on the
mechanism of DNA replication in greater depth. other side and two separate chromosomes are released.
5′ 3′
G C Parental helix Origin
A T
G
C
T A
T A
G C
AT Replication forks
TAA
C G Replication fork
C G
A T A T
G C G C
G G
A T A T
A T A T
T A T A
CCG CCG Replicas
TA TA
A T A T
3′ G C 5′ 3′ G C 5′
Parental New New Parental
Figure 11.10 Semiconservative DNA Replication.The replication
fork of DNA showing the synthesis of two progeny strands. Newly Figure 11.11 Bidirectional Replication.The replication of a circular
synthesized strands are in maroon. Each copy contains one new and bacterial genome. Two replication forks move around the DNA forming
one old strand. This process is called semiconservative replication. theta-shaped intermediates. Newly replicated DNA double helix is in red.
3.5 The Procaryotic Cell Wall 59
Porin
Osi-dsep cehcaifiincs Blipraoupnro’stein Multimedia-Supported Illustrationsappear throughout the
Lipopolysaccharide text. To facilitate finding corresponding full-color video,
animations,or interactive screens from the third edition of
Outer Microbes in Motion,a correlation guide is provided on the CD-
membrane
ROM,on the Student Online Learning Center,and in the
Pspearicpela asnmdic Student Study Guide.
peptidoglycan
Phospholipid Peptidoglycan Pmleamsmbarane Microbes in Motion,Third Edition,CD-ROM is organized into
Integral protein 18 topical “books,”the books are divided into “chapters,”and
Figure 3.23 The Gram-Negative Envelope. the chapters have numbered “pages.”For each multimedia-
supported illustration,the correlation guide directs the reader to
the book,chapter,and page on the CD-ROM where
corresponding material can be found.
Figure 3.23 Bacterial Structure and Function Book Cell
Wall Chapter
Peptidoglycan Topic pp.2–3