Ebook Prescott''s microbiology (9th edition): Part 1
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ninth edition
Prescott's Microbiology
Joanne M. Willey
HOFSTRA UNIVERSITY
Linda M. Sherwood
MONTANA STATE UNIVERSITY
Christopher J. Woolverton
KENT STATE UNIVERSITY
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PRESCOTT'S MICROBIOLOGY, NINTH EDITION
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About the Authors
Joanne M. Willey has been a
Linda M. Sherwood is a member
Christopher J. Woolverton is
professor at Hofstra University on Long
of the Department of Microbiology at
founding professor of Environmental
Island, New York, since 1993, where she
Montana State University. Her interest in
Health Science, College of Public Health at
is Professor of Microbiology; she holds a
microbiology was sparked by the last course
Kent State University (Kent, OH), and is the
joint appointment with the Hofstra
she took to complete a B.S. degree in
Director of the Kent State University (KSU)
University School of Medicine. Dr. Willey
Psychology at Western Illinois University.
Center for Public Health Preparedness,
received her B. A. in Biology from the
She went on to complete an M.S. degree in
overseeing its BSL-3 Training Facility.
University of Pennsylvania, where her
Microbiology at the University of Alabama,
Dr. Woolverton serves on the KSU graduate
interest in microbiology began with
where she studied histidine utilization
faculty of the College of Public Health, the
work on cy anobacterial growth in
by Pseudomonas acidovorans. She
School of Biomedical Sciences, and the
eutrophic streams. She earned her Ph. D.
subsequently earned a Ph.D. in Genetics at
Department of Biological Sciences. He holds
in biological oceanography (specializing
Michigan State University, where she
a joint appointment at Akron Children's
in marine microbiology ) from the
studied sporulation in Saccharomyces
Hospital (Akron, OH). He earned his B.S. in
Massachusetts Institute of Technology
cerevisiae. She briefly left the microbial
Biology from Wilkes College (PA), and his
Woods Hole Oceanographic Institution
world to study the molecular biology of
M.S. and Ph.D. in Medical Microbiology
from West Virginia University, School of
Joint Program in 1987. She then went to
dunce fruit flies at Michigan State
Harvard University, where she spent her
University before moving to Montana
Medicine. He spent two years as a
postdo ctoral fellowship study ing the
State University. Dr. Sherwood has always
postdoctoral fellow at UNC-Chapel-Hill.
filamentous soil bacterium Streptomyces
had a keen interest in teaching, and her
Dr. Woolverton's current research is focused
coelicolor. Dr. Willey continues to
psychology training has helped her to
on real-time detection and identification of
investigate this fascinating microbe
understand current models of cognition
pathogens using a liquid crystal (LC)
biosensor that he patented in 2001. Dr.
and has coauthored a number of
and learning and their implications for
publications that focus on its complex
teaching. Over the years, she has taught
Woolverton has published and lectured
developmental cy cle. She is an active
courses in general microbiology, genetics,
widely on the mechanisms by which LCs act
member of the American Society for
biology, microbial genetics, and microbial
as biosensors and on the LC characteristics
Microbiology (ASM), and served on the
physiology. She has served as the editor for
of microbial proteins. Professor Woolverton
editorial board of the journal Applied
ASM's Focus on Microbiology Education
teaches microbiology, communicable
and Environmental Microbiology for nine
and has participated in and contributed
diseases, immunology, prevention
y e ars and as Chair of the Division of
to numerous ASM Conferences for
and control of disease, and microbial
General Microbiology. Dr. Willey
Undergraduate Educators (ASMCUE).
physiology. He is on the faculty of the
regularly teaches microbiology to
She also has worked with K-12 teachers to
National Institutes of Health National
biology majors as well as medical
develop a kit-based unit to introduce
Biosafety and Biocontainment Training
students. She also teaches courses in cell
microbiology into the elementary school
Program, teaching laboratory safety, risk
biology, marine microbiology, and
curriculum and has coauthored with
assessment, decontamination strategies, and
laboratory techniques in molecular
Barbara Hudson a general microbiology
bioterrorism readiness. An active member
of the American Society for Microbiology,
gene tics. Dr. Willey lives on the north
laboratory manual, Explorations in
shore of L ong Island with her husband
Microbiology: A Discovery Approach,
Woolverton serves on its Board of Education
and two sons. She is an avid runner and
published by Prentice-Hall. Her association
and as the editor-in-chief of its Journal of
enjoy s skiing, hiking, sailing, and
with McGraw-Hill began when she
Microbiology and Biology Education.
reading. She can be reached at
prepared the study guides for the fifth and
Woolverton and his wife, Nancy, have three
joanne.m.willey @hofstra.edu.
sixth editions of Microbiology. Her non
daughters, a son-in-law, and a grandson. He
academic interests focus primarily on her
enjoys time with his family, ultra-light
family. She also enjoys reading, hiking,
hiking and camping, and is an avid cyclist.
gardening, and traveling. She can be
His e-mail address is
reached at
iii
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Laboratory Exercises in
Microbiology, Ninth Edition
John P. Harley has revised this labora
tory manual to accompany the ninth
edition of Prescott's Microbiology. The
class-tested exercises are modular to
allow instructors to easily incorporate
them into their course. This balanced
introduction to each area of microbiol
ogy now also has accompanying Connect content for additional homework
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v
A Modern Approach to Microbiology
Evolution as a Framework
Introduced immediately in chapter 1 and used as an overarching
theme throughout, evolution helps unite microbiological con
cepts and provides a framework upon which students can build
their knowledge.
Aboutthe Authors
Preface
111
24
555
2STheProtists
'----+---
1 TheEvolution ofMicroorganr.;m� andMicrobiology
2 Micro5t:opy
568
TheFungi(Eumycota)
211
Part One Introduction to Microbiology
588
1
22
3 BacteriaiCeiiStructure
42
4
82
ArchaeaiCeiiStructure
5 EukaryoticCeiiStructure
Separate Chapters on Bacteria and Archaea
Actinobacteria: TheHighG + CGram-Positive
Bacteria
iv
Part Six Ecology and Symbiosis
28 BiogeochemicaiCyclingandGiobal
ClimateChange
92
632
29 Methods InMicrobialEcology
6 Viruses and Other Acellular lnfedklus Agents
15415
30 Microorgani5ms inMarine andFre5hwater
In recognition of the importance and prevalence of archaea, the
structure, genetics, and taxonomic and physiologic diversity of
these microbes are now covered in chapters that are separate
from those about bacteria.
Ecosystems
Part Two Microbial Nutrition, Growth, and Control
7 MicrobiaiGrowth
660
31 Microorgani5m5inTerre5triaiEco!i)'stems
32 Microbiallnteractions
133
679
699
8 ControlofMicroorganismsin theEnvironll"lent
9 AntimicrobiaiChemotherapy
189
Part Seven Pathogenicity and Host Response
Part Three Microbial Metabolism
10
lntroductklntoMetaboWsm
33
lnnateHostResistance
34
Adaptivelmmunity
723
753
35 Pathogenicity andlnfection
210
789
11 Catabolism:EnergyReleaseandConservation
12
Anabolism:TheUseofEnergyin8iosynthesis
Part Eight Microbial Diseases, Detection, and Their
266
Control
Part Four Microbial Molecular Biology and Genetics
13 BacteriaiGenomeReplicationandExpression
14 Regulation ofBilcteriaiCellularProcesses
287
andExpression
325
Microbial World
40 HumanDiseasesCaused
18 MicrobialGenomics
structure and function of bacteria and archaea are followed by
the discussion of eukaryotic cells preceding viruses.
854
888
byFungiandProtists
372
Part Nine Applied Microbiology
17 RecombinantDNATechnology
Now covered in chapters 3-6, the separate chapters on the
424
41 MicrobiologyofFood
958
42 Biotechnologyand lndustriaiMicrobiology
43
Part Five The Diversity of the Microbial World
20 TheArchaea
Applied EnvironmentalMicrobiology
Appendix 1
19 MicrobialTaxonomy and the Evolution of Diversity
469
Appendix2
996
AReview of the Chemistry
ofBiologicaiMolecules
A-1
Common Metabolic Pathways
21 TheDeinococci,Mollicutes,andNonproteobacterial
Gram-Negative Bacteria
22 TheProteobacteria
Glossary
489
Credits
509
23 Firmicutes:ThelowG+CGram-PositiveBilcteria
542
Index
G-1
C-1
1-1
Molecular Microbiology and Immunology
Secondary Lymphoid Organs and Tissues
The8plecn is the most highly organized secondary lymphoid
organ.lt is a largeorgan located in the abdominal cavitythat
functions to filter the blood and trap blood-borne particles to
be ass�ssed for foreignness by phagocytes (figure 33.14). Mac
rophages and dendritic cells are present in abundance, and
once trapped by splenic macrophages or dendritic cells, a
pathogen is phagocytosed, killed, and digested. 1he resulting
antigens are presented to lymphocytes, activating a specific im
mune response.
Lymph nodes lk at the junctions oflymphaticvessds, where
macrophages md dendritic cells trap particles that enter the lym
phaticsyslem(figure33.14c).If a parlicle isfollndlobe foreign,il
is then phagocytosed and degraded, and the resulting antigens
arcprcscntcdto lymphocytcs.
Lymphoid tissues are found througholll the body as highly
organi7.ed or loosely associated cellular complexes(figure 33.14).
Some lymphoid cells are closely associated with specific tissues
such as skin(skin-associated lymphoid tissue,or SALT) and mu
cous membranes (mucosal-associated lymphoid tissue, or
MALT).SALT and MALT arc good examples of highlyorgani7.ed
lymphoid tissues that featuremacrophages surrounded by spe
cific areas of B and T lymphocytes and sometimes dendritic cells.
Loosely associated lymphoid tissue is best represented by the
bronchial-associated lymphoid tissue (BALT), because it lack!!
cellular partitioning. The primary role of these lymphoid tissues
is to efficiently organi7.e leukocytes to increase intc:raction be
tween the innate and the adaptive arms of the immune response.
'lhus, the lymphoid tissues serve as the interface between the in
nate resistance mechanisms and adaptive immunity of a host.
We now discuss these tissues in more detail
Despite the skin's defenses, at times pathogenic microorgan
isms gain access to the tissue under the skin surface. Here they
encounter a spedalized set of cells called the $kin-associated
lymphoid ti!lsue (SALT) (figure 33.15). The major function of
SALT is to confine microbial invaders to the area immediately
underlying the epidermis and to preventthemfrom gaining ac
cess to the bloodstream. One type of SALT cell is the Langc:r
hans cell, a dendritic ccll that phagocytoses microorganisms
that penetrate th� skin. Once the Langerhans cell has int�mal
i7.ed a foreign particle or microorganism, it migrates from the
epidermis to nearby lymph nodes, where it presents antigen to
activate nearby lymphocytes, inducing a specific immnne re
sponse to that antigen. This dendritic cell-lymphocyte interac
tion illustrates another bridge between innate resistance and
adaptive immunity.
The epidermis also contains another type of SALT cell
called the intraepidfrmal lymphocytf (figure 33.15), a spe
cialized T cellhavingpotentcytolyticand immunoregulatory
responses to antigen. These cells are strategically located in the
skin so that they can intercept any antigens that breach the first
line of defense. Most of these specialized SALT cells have limiL.ed
rcceptordiversity andhavelikelyevolvedto recogni7.ecommon
skin pathogen patterns.
39 HumanDiseasesCausedbyBacteria
353
16 MechanismsofGeneticVariation
808
37 Epidemiology andPublic HealthMicrobiology
38 HumanDiseasesCaused byVirusesandPrions
15 Eukaryotic and ArchaealGenome Replication
An Introduction to the Entire
36 ClinicaiMicrobiologyand lmmunology
The ninth edition includes updates on genetics, biotechnology,
genomics, and immunology. The discussion of eukaryotic and
archaeal genetics has been expanded and makes up a separate
chapter to reflect the relatedness of genetic information flow. A
streamlined discussion of immunity with enhanced detail be
tween innate and adaptive linkages helps students grasp the
complexity and specificity of immune responses.
The specialized lymphoid tissue in mllcous membranes is
called mucosal-associated lymphoid tissue (MALT). There
are sneral types of MALT. The system mo$l studied is the gut
associated lymphoid tis�ue (GALT). GALT includes the ton
sils, adenoids, diffllse lymphoid areas along the gut, and
specialized regions in the intestine calledPeyer's patches. Less
well-organized MALT also occurs in the respiratory system and
vi
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A Modern Approach to Microbiology
..
21st-Century Microbiology
Prescott's Microbiology leads the way with updated text devoted
(figurel&ll).Ammoniumrunoffleachei
to global climate change, biofuels, and microbial fuel cells. For
I!to lahsandotream•,frcquentlycausing
eutrophication-an increase in nutrient
level&that stimula�nthegrowthof a lim
more, see chapters 28, 30, 42, and 43.
itednumberoforgani.imll,therebydisturb
ingthe ecoloj;yoftheseaquaticero�m�.
Bycontrasl,microbialniuificationc.anre
lultlntheo!ddationofmunonium tomore
nlt:rattth�.n eanbeinuoobi.li%edby pb.nt!
andmlcrobe$. asorpnlmnneeda$pedfic
rui
: The pr
lhe rnctlft g:Jftnllotue nltro�no.Ddes.
This cycle of nilrific.atioo/denitrification
Metagenomics and the Human Microbiome
fueledbyNH.,+introducedasfertilir.eris
respoosibleforthe highestN00levels in
The updated genomics chapter covers the technical aspects of
650.000)'1lUS.
What are the<:<�n�u..nus of di•
rupting lhccarbon andnitrogencycles!
Globlllclimate change itthemost obvi
ous example. k is important to keep in
metagenomics, and the human microbiome is discussed in the
mind that weaber is oot the same a11 cli
context of microbial interactions in chapters 18 and 32.
mate. WhileNorth America has 1uffered
lomtoftheholl«l tll1llmtnon recordln
the purdecade,a&ingle day<>r week in
Julytlu.t is panio:ubrly hot!8not,by it·
sel£evicknce olgloN\climate ebange.
Globf.l clim•te change is mmuredovtr
decadeswdindudnn1•nypenmete1'1
lucha.&urf�tempenolure on landand
on. and In tb.. atmmpbere and trope·
Flgure28.12 ,..tur•llndHum;m·MidtlnftutnotSonlht"llrO!I"ICJdt.
MICRO INQUIIIV
optMre;rates of precipitation;andfrequency of extreme Wt'ather. Based on
"Mwnorga.rJlmibfntfir(romnirril\caticln1
Laboratory Safety
these analyr.e�o,the average global temperature has incrnK
Reflecting forthcoming recommendations from the American
bustion toCO,(fipre2S.l3).Dtpending on the r;rte ofconlln·
nedincrease in greenhouse gases.the average global surface
Society for Microbiology, chapter 37 provides specific guidance
temperatur e i s predictedtorise betweenl.land6.4"Clr)'2100.
Mo imporunt question is how will microbes re�poOO 10 a
changingworid.ll<>:;auoe for thevast majority ofl'.arilishistory,
m.iCI'O(lrganimu have bun the drtvm of elememzl C)'(llng.
for laboratory best practices to help instructors provide safe con
clwlgesinrniuobialactivitieswillhavea majori.J:rl>acto ntherate
andrnagrtitudeofgreenbouse gas�umul.ationa.ndglobal elimate
ditions during the teaching of laboratory exercises.
eh.a.l!$e.'Ihe rolem.ierobes plty!nba.laneingcvbon andllit.rostn
fluxethunpentdnewavrouesofmearchltiroicroblaleo:ology.
Retrie�. tnfw, ¥rJiy
1. listlhn!e!JtemllouHgom!li.Disa.t!SthtiroriQim
2.DiK11Mthe,_.;bleroleDffureot•in the controiDfCO,.
l. Howdo<�"9"'inthe nitrogencyd"caused bylertilization
lnflu..ncetheurbon�?
4. GIVen !hit "idl mkroblal �roup ha• a n optimum temperalllr e
r;m!JI11or !Jfll'l"th."- migl1t you predict cl1�nges to a soli mlcrolllal
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Flgure28.13 GlobaiAnnuai-MeanSo.ufaceAirTm.per1ture
Change.
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-
Disease
_
26.1
White-Nose Syndrome Is Decimating North American Bat Populations
Special Interest Essays
Bats evoke all kinds of images. Some people immediately
Organized into four themes-Microbial Diversity &
summer evening outdoors on the east coast ofNorth America,
wing. Wings provide a large surface area for colonization,
mosquitoes and the small bats that eat them may come to
and once infected, the thin layer of skin is easily damaged,
leading to adverse physiological changes during hibernation.
white fungal hyphae growing around their muzzles
These in turn result in premature awakening, loss of essential
and connective tissue. Despite the name WNS, the primary
large fruit bats often called flying foxes. If you have spent a
site of infection (and the anatomical site harmed most) is the
Ecology, Techniques & Applications, Historical High
mind. A new scene can now be added to these: bats with
lights, and Disease-these focused and interesting essays
ure).
provide additional insight to relevant topics.
to
�
3.1
and many a budding microbiologist into thinkil
ology cannot be overstated. The Gram stain reaction was for
bacterium has a typical Gram-positive envelope.
many years one of the critical pieces of information used by
gues that by relating cell envelope architecture tot
enies of various bacterial taxa, we may gain insig
in identifying bacteria in clinical settings. The initial studies
evolution of these architectures. He notes that th•
done to differentiate bacteria that stained Gram positive
micutes and Actinobacterta are composed almost
from those that stain Gram negative were done using modd
of monoderm bacteria, whereas almost all othe
organisms such as Bacillus subtilis (Gram positive) and Esch
phyla consist of diderms.
erichia coli (Gram negative). At the time, it was thought that
phylogeny and cell envelope structure. For instanet
However, as the cell walls of more bacteria have been charac
of the genus Mycobacterium (e.g., M. tubercula
terized, it has become apparent that it may be misleading to
to the predominantly monoderm phylum Acti1
refer to bacteria as Gram positive or Gram negative. In other
Mycobacteria have cell walls that consist of pep
words, the long-held models of Gram-positive and Gram
and an outer membrane. The outer membrane is
negative cell walls do not hold true for aU bacteria. Recently
of mycolic adds rather than the phospholipid�
Iain Sutcliffe has proposed that microbiologists stop refer
polysaccharides (LPSs) f�und in the typical Gnu
ring to bacteria as either Gram positive or Gram negative. He
cells' outer membrane.
(section 24.1)
tfi
terial cell envelope architectures by focusing on the observa
Members of the genus Deinococcus are anotb
tion that some bacteria have envelopes with a single
ing exception. These bacteria stain Gram positive
derms. Their cell envelopes consist of the plasma ·
positive bacteria-while others have envelopes with two
what appears to be a typical Gram-negative cdl
�
membranes-the plasma membrane and an outer membrane
outer S-layer. Their outer membrane is distinctivt
as seen in typical Gram-negative bacteria. He proposed call
lacks LPS. Deinococd are not unique in this respe'
But why make this change? Sutcliffe begins by pointing
It is now
known that there are several
taxa with c
branes that substitute other molecules for LPS.
out that some bacteria staining Gram positive are actually
diderms and some staining Gram negative are actually
moooderms. By referring to Gram-positive-staining diderms
as Gram-positive bacteria, It is too easy to mislead scientists
WNS was first spotted in 2006 among bats hibernating in
causes mild infection in at least one hibernating bat species.
a cave near Albany, NY. Scientists qukkly became alarmed for
This makes G. destructans an apparent case of pathogen
two reasons. First, it spreads rapidly-it's known to occur in at
pollution-the human introduction of invasive pathogens of
least six bat species and is now found from the mid-Atlantic
wildlife and domestic animal populations that threaten bio
United States, northward into Canada (Ontario, Quebec, and
diversity and ecosystem function.
99%
Soun;e:Sutcliffe,I.C.lfJIO.A phylum level perspectiveonbl�tetUicenMvelope
�tchire c wr•. lren dsU ie robl ol. fB{I0/...64-70.
The capacity of G. destructans to sweep through bat
in any
populations results from a "perfect storm" of host- and
given infected hibernacula (the place where bats hibernate,
pathogen-associated factors. G. destructans is psychrophilic,
which unfortunatdy rhymes with Dracula).
with a growth optimum around trC; it does not grow above
WNS is caused by the ascomycete Geomyces destructans.
20°C. All infected bat species hibernate in cold and humid
It colonizes a bat's wings, muzzle, and ears where it first
environments such as caves and mines. Because their meta
bolic rate is drastically reduced during hibernation, their
body temperature reaches that of their surroundings, be
tween 2 and 7°C. Thus WNS is only seen in hibernating bats
or those that have just emerged from hibernation. When
metabolically active, the bat's body temperature is too Vl-atm
to support pathogen growth.
While it is too late to save the estimated 6 million bats
that have already succumbed to WNS, microbiologists, con
servationists, and government agencies are trying to limit
the continued decline in bat populations. Caves have been
clo.�ed to human traffic, and protocols for decontamination
after visiting hibernacula have been developed to limit the
spread from cave to cave. Although we cannot cure sick
Suborder Corynet
membrane-the plasma membrane as seen in typical Gram
ing the former monoderms and the latter diderms.
humans inadvertently brought it from Europe, where it
There are interesting exceptions to the rela1
all other bacteria would have similar cell wall structures.
suggests that instead we should more precisely describe bac
Where did this pathogen come from and why does it
infect bats? The best hypothesis regarding Its origin Is that
deadly. A population of bats declines from 30 to
The importance of the Gram stain in the history of microbi
fat reserves, and strange behavior.
eliminate the most common bat species In eastern North
New Brunswick), and as far west as Oklahoma. Second, it is
Gram Positive and Gram Negative or Monoderms and Diderms?
bacterial taxonomists to construct taxa, and it is still useful
(box fig
This is the hallmark of white-nose syndrome (WNS),
and if its rate of infection continues unchecked, it is projected
America (Myotis lucifugus) by 2026.
Microbial Diversity& E
erodes the epidermis and then invades the underlying skin
think of vampire bats and are repulsed. Others think of the
bats, it is our responsibility to stop the continued spread of
this pathogen.
Geomyces destructans causes WNS. A little brown bat {Myotis lucifugus)
with the white fungal hyphae(,mow) for which WNS is named.
rylng th$u$1JcausuregioMipopul•tlon
Re•dmorti:Frict,W.F.era/.,2UIU.Aneme
col / 1ps• of• common NarthAm•ric•n Nt 1p1cilr. S�itnca 319:679-682.
Student-Friendly Organization
New! Newsworthy Stories-Each chapter begins with a
real-life story illustrating the relevance of the content cov
ered in the upcoming text.
Viruses and
Other Acellular
Infectious Agents
New! Readiness Check-The introduction to each chapter
cantaloupe
Stille
Mustard, Catsup, and Viruses?
e
�: �� en�:g :'
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States.Hot dogsarJdluochmeats arepopularat outingssuch asbilsebi!ll
!Jilmesandin lunchescarried towarkor schooi.Yeteachyearintfle
includes a skills checklist that defines the prior knowledge
sedan outbreokof listeriosisin20states i n theUnited
ich infected over l:lO and k�led over 20
a student needs to understand the material that follows.
Viruses as agents of good will come as a surprise to many.Typicallywe
thinkofthemasmajorcausesofdisease.However,viruses are>ignilicamfor
otherrea>ens.Theyarevitalmembersofaquatic ecosystems.Ttlerethe
interact with cellulor mkrobesand contribute to the mo\lement
ganic
Urlited Stat,..,apprmimately1,600peopleare sickenedby•bacterium
that can wntominate the meat and. even worse. survive aridgrow when
the"""'a tis properlyrefrigerated.
Thediseasecul,...itisLis teriamonocytogenes,aGram-positive rOO
fourtdinsoil aridmanyotherenvironrnental sites.lt isnot orllycoldtolerant
butsaltandaddtolerantas weii.AithoLJgh itisinthe minorleago.�eswhen
compared to someofthebig hitters offo OObome disease(e.g.,
So/monel/a
fflterico),it isofcoocernfortwo reasons:who itkillsanclhowrTIIlnyitk�ls.
L./'OOil()[ytvgfflesUrgets theyOlllgand old,pregrlilntwomen,and
immunocompromised individuals; about 15%ofthose inf!'Cted die.
ltseffectonpregnantwo"""'n is partkularlyheortbreaking.The
woman usually only suffers mild,flu like symptoms; however.these
innocuous symptomsbelie thefactthatthechildshe carries isin serious
danger. Herpregnaocyoftenendsin miscarriageor stillbirth.Newborns
infected with thebacterium are likely to develop meningitis. Many will die
as a result.Thme whosurviveoftenhave neurologicaldisorders.
Currently,pregnant wamenare coonseledagainst eatingrBldy-to-eat
Bialogica/wntrolafmkroorganisms
tian8.7)
New! Learning O utcomes-Every section in each chapter
Readiness Check:
begins with a list of content-based activities students
flased onwhatyouhilvelearnedpreviousfy,youshoul dbe ableto
II Definetheterm acellular
II Compareand contrast ingeneralterms viruses,viroids,satelites,aOO
prions(sectionU)
should be able to perform after reading.
6.1 Viruses
After reading this section, you should be �ble to
• Define the terms virology,bacterioptloges, and ptloges
• Li>torganism> thatarehoststo viruses
food>unlesstheyhavebeencookedpriorto consumption.f-lowever,
L.monacyrogene� is koown to contaminate manyfoods other than tlot dogs
andthesecan't alwaysbeheated.ln2006th�U.S.FoodandDrug
Administration(FDA)appro•ed a new approachto preventlisteriosis:
spraying •irusesthatattackanddestroythebacteriuman reody-to-eatcold
cutsalldlllncheonmeats.lnother words.the viruseo areafood additive!
The"""'t hodissafet>ecausethe viruses onlyattackL.mooocyrogene<>.not
Sinceapproval.the uoeof virusesto controlthetransmissionof
listeriosisbyotherfoodshasbeenstudied.Unfununately,thosestudiesdid
ootindudefoodssuchas freshfruit.ln2011Lfl"lOIJO!:}'W9enes-contaminated
,--"""'"--'"'"""""-"""'-�"-"'-""-_...�"'"""'-----'--,
Animation I con-Th is sym
bol indicates material pre
sented in the text is also
Micro In quiry- S elect figures
accompanied by an anima
throughout every chapter
tion on the text website at
contain probing questions,
www.mhhe.com/willey9.
adding another assessment
opportunity for the student.
MICRO INQUIRY l'ihydotheemptyw,7'ii'f>rem1ioc;;fc;ch
Cross-Referenced
Notes
In-text references refer stu
dents to other parts of the
book to review.
Retrieve, In fer, Apply
Questions within the nar
rative of each chapter assist
students in mastering sec
tion concepts before mov
ing on to other topics.
viii
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Student-Friendly Organization
Vivid Instructional Art Program-Three
dimensional renditions and bright, attractive
colors enhance learning.
r�>cugllti
l..!LJI d1�m�u," wUdi LUiil.<'i willll�so;,.;x:�t:s _,., iadlii
n::o:nitlon in ''"'linn 333, fiKu.o.in� nn ''""'Pl..'TTI� nt pn liL1TlN md r:..t
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I he ''l'!ionin-ir.:lqx:nd,nt mcchlL\i5mN an: g.· rm - liL\C �r.wd:d.
ret·t'\)'\or l'llf>fd Sl·sttJ:W ..,·J:��".rtill wui�ulu IJ
(f•�;un• 33.11}. A m.u:)xr .,[ .w.�mbr;u"' - bv 1wJ r
More Annotated Figures-All key metabolic
pathways and molecular processes are now anno
tated, so that each step is clearly illustrated and
explained.
Totolml�
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r. - ? r;.-..['ll-:·" -,� • -��; '"''"ffl:->1', vd ,..,... • 1'-."if.';'ll' I' ·. :- -� ·r, "'"*� · �:lll:lllni] -h�- If.'!'-: .. ..,_.�,yr.lll'll' :1;01' ;:c···� .-.- - - r.u:lh Fr.rr.r·•.n rl',R
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Key Concepts-At the end of each
chapter and organized by num
bered headings, this feature dis
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oi >t�lti�li
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tills the content to its essential
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referenced figures and tables.
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mclUhk17.l;
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• l'<:Rna.� mJrno:mu1app�i..:37ifln�. rt.ll!Tcni�u
17.3 Cloning Vectors ilndCreating
Rf'combin.ant DNA
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ix
List of Content Changes
Each chapter has been thoroughly reviewed and many have un
introduces the concept of metabolic flux through the intercon
dergone significant revision. All now feature pedagogical ele
nected biochemical pathways used by cells.
ments, including aReadinessCheck for the chapter and Learning
Chapter 11- The chapter now begins with an introduction to
Outcomes for each section therein.
metabolic diversity and nutritional types.
Chapter 12-Updated coverage ofCOrfixation pathways.
Part I
Chapter 1- Evolution is the driving force of all biological sys
tems; this is made clear by introducing essential concepts of mi
crobial evolution first.
Chapter 3-Coverage of bacterial cellular structure and function.
The chapter now includes a discussion of nutrient uptake in the
section on bacterial plasma membranes.
Chapter 4-G rowing understanding of the distinctive character
istics of archaea has warranted the creation of a new chapter that
focuses on their cell structure and function.Comparisons to bac
teria are made throughout the chapter.
Chapter 5- An introduction to eukaryotic cell structure and
function, with emphasis on eukaryotic microbes. More de
tailed information on protist and fungal cells is presented in
chapters 25 (The Protists) and 26 (The Fungi), which also focus
on the diversity of these microbes. Comparisons between bac
teria, archaea, and eukaryotes are included throughout the
chapter.
Chapter 6- This chapter, entitled Viruses and Other Acellular In
fectious Agents, surveys the essential morphological, physiologi
cal, and genetic elements of viruses as well as viroids, satellites,
and prions. This chapter completes our four-chapter introduction
PartlY
Chapter 13-Now focuses on bacterial genetic information flow
with improved coverage of bacterial promoters, sigma factors,
termination of DNA replication, transcription cycle, and protein
folding and secretion.
Chapter 14-Now focuses on the regulation of bacterial cellular
processes. The coverage of regulation of complex cellular behav
iors has been significantly updated and expanded, including new
material on cyclic dimericGMP.
Chapter 15-A new chapter that considers eukaryal and archaeal
genome replication and expression together. In both cases, the
discussion has been updated and expanded, and reflects the simi
larity of information flow as carried out by members of Archaea
and Eukarya.
Chapter 16-Covers mutation, repair, and recombination in the
context of processes that introduce genetic variation into popula
tions. This is now related to the evolution of antibiotic-resistant
bacteria.
Chapter 17- The use of recombinant DNA approaches to con
struct a synthetic genome is highlighted.
Chapter 18-New principles and applications of genomic tech
of microbial life.
niques, including massively parallel genome sequencing and
single cell genome sequencing, are now reviewed. The growing
Part II
Chapter 7-Reorganized to initially focus on the growth of mi
crobes outside the laboratory (including growth in oligotrophic
environments) and the environmental factors that influence
microbial reproduction. Topics related to laboratory culture of
microbes follow.
importance of metagenomics to environmental microbiology and
its use in exploring the human microbiome are introduced here.
PartY
Chapter 19-Microbial evolution, introduced in chapter 1, is ex
panded with a complete discussion of the endosymbiotic theory,
Chapter 8-Reorganized to reflect emphasis on interruption of nor
mal growth and reproduction functions to control microorganisms.
Chapter 9-Content focuses on the mechanism of action of each
antimicrobial agent and stresses usage to limit drug resistance.
and the concept and definition of a microbial species.
Chapter 20- Expanded coverage of archaeal physiology includes
new figures presenting archaeal-specific anabolic and catabolic
pathways. The evolutionary advantage of each pathway is dis
cussed in the context of archaeal ecology.
Part III
Chapter 21-Now includes mycoplasmas, in keeping with
Chapter 10- This introduction to metabolism includes a new
Bergey's Manual; new figures illustrating the life cycle of Chlamydia
section that outlines the nature of biochemical pathways and
are included.
X
/>
List of Content Changes
Chapter 22-Expanded coverage of proteobacterial physiology
Chapter 34-Reorganized and updated to enhance linkages be
with content on Cl metabolism, including several figures.
tween innate and adaptive immune activities. Discussions inte
Chapter 24- Increased coverage of streptomycetes, with new
grate cell biology, physiology, and genetics concepts to present
graphics illustrating their life cycle and their importance in anti
the immune system as a unified response having various compo
biotic production.
nents. Implications of dysfunctional immune actions are also
Chapter 27-Updated discussion of virus taxonomy and phylog
discussed.
eny, including increased coverage of archaeal viruses and the
Chapter 35- This chapter has been re-titled Pathogenicity and
CRISPR/CAS system.
Infection, reflecting its emphasis on microbial strategies for
survival that can lead to human disease. The essential elements
required for a pathogen to establish infection are introduced
Part VI
and virulence mechanisms highlighted. It follows the immu
Chapter 28-The description of each nutrient cycle is accom
nology chapters to stress that the host-parasite relationship is
panied by a new "student-friendly" figure that distinguishes
dynamic, with adaptations and responses offered by both host
between reductive and oxidative reactions. Expanded cover
and parasite.
age of the interaction between nutrient cycles is also newly
illustrated.
Chapter 29-This chapter continues to emphasize culture-based
Part VIII
techniques as the "gold standard" and reviews some new, innova
Chapter 36-This chapter has been updated to reflect the work
tive approaches. The chapter also discusses a variety of culture
flow and practice of a modern clinical laboratory. Emphasis is on
independent
modern diagnostic testing to identify infectious disease.
techniques
used
to
assess
populations
and
communities.
Chapter 37-Expanded focus on the important role of labora
Chapter 30- Updated and expanded discussion of freshwater
tory safety, especially in the teaching laboratory. Discussion em
microbiology is complemented by discussion of carbon cycling in
phasizes modern epidemiology as an investigative science and
the open ocean and its implications for global climate change.
its role in preventative medicine. Disease prevention strategies
Chapter 31-New and updated coverage of mycorrhizae, with an
are highlighted.
emphasis on host-microbe communication and evolutionary
Chapter 38- Updated and expanded coverage includes viral
similarities to rhizobia.
pathogenesis and common viral infections.
Chapter 32-Microbial relationships are presented along with
Chapter 39-Expanded coverage of bacterial organisms and
human-microbe interactions, helping to convey the concept that
their common methods leading to human disease.
the human body is an ecosystem. New and increased coverage of
Chapter 40-Refocused to reflect disease transmission routes as
the human microbiome.
well as expanded coverage of fungal and protozoal diseases.
Part VII
Chapter 33-Reorganized and updated, this chapter on innate
host resistance provides in-depth coverage of physical and
chemical components of the nonspecific host response fol
Part IX
Chapter 41-Expanded discussion of probiotics in the context of
the human microbiome.
lowed by an overview of cells, tissues, and organs of the im
Chapter 42-This chapter has been reorganized to illustrate the
mune system. This includes a step-by-step discussion of how
importance of industrial microbiology by presenting common
microorganisms and damaged tissues are identified by the host
microbial products-including biofuels-first. This is followed by
using pattern recognition to remove them. Discussions of
an updated discussion of strain development, including in vivo
phagocytosis and inflammation are updated and reflect mo
and in vitro directed evolution.
lecular mechanisms. The groundwork is laid for a full apprecia
Chapter 43-Updated discussion of water purification, wastewater
tion of the connections between the adaptive and innate arms
treatment, and bioremediation. This includes the development
of the immune system.
and use of microbial fuel cells.
xi
Acknowledgments
We would like to thank the Reviewers, who provided constructive reviews of every chapter. Their specialized
knowledge helped us assimilate more reliable sources of information and find more effective ways of expressing
an idea for the student reader.
Reviewers
Mark McBride, University ofWisconsin-Milwaukee
Tamarah Adair, Baylor University
Vance McCracken, Southern Illinois University Edwardsville
Richard Adler, University of Michigan-Dearborn
Donald Mcgarey, Kennesaw State University
Fernando Agudelo-Silva, College of Marin
Robert McLean, Texas State University
Shivanthi Anandan, Drexel University
Tamara Mcnealy, Clemson University
Penny Antley, University of Louisiana at Lafayette
Rita Moyes, Texas A&M University
Suzanne Barth, The University of Texas at Austin
Karen Nakaoka,Weber State University
Larry Barton, University of New Mexico
Comer Patterson, Texas A&M University, College Station
Nancy Boury, Iowa State University
Ed Perry, Faulkner State Community College
Ginger Brininstool, Louisiana State University-Baton Rouge
Thomas Pistole, University of New Hampshire
Linda Bruslind, Oregon State University
Ronald Porter, Penn State University-University Park
Alison Buchan, University of Tennessee
Jackie Reynolds, Richland College
Jim Buritt, University ofWisconsin-Stout
Margaret Richey, Centre College
Martha Smith Caldas, Kansas State University
Veronica Riha, Madonna University
Joseph Caruso, Florida Atlantic University-Boca Raton
Timberley Roane, University of Colorado Denver
Andrei Chistoserdov, University of Louisiana at Lafayette
Jerry Sanders, University of Michigan-Flint
Carlton Cooper, University of Delaware
Pratibha Saxena, The University of Texas at Austin
Susan Deines, Colorado State University
Mark Schneegurt, Wichita State University
John Dennehy, Queens College
Sasha A. Showsh, University ofWisconsin-Eau Claire
James Dickson, Iowa State University
Khalifah Sidik, University of Illinois College of Medicine at Rockford
Ronald Dubreuil, University of Illinois at Chicago
Deborah Siegele, Texas A&M University
Paul Dunlap, University of Michigan-Ann Arbor
Jack Steiert, Missouri State University
Mary Farone, Middle Tennessee State University
Raji Subramanian, NOVA Community College Annandale
Babu Fathepure, Oklahoma State University-Stillwater
Karen Sullivan, Louisiana State University-Baton Rouge
Kathy Feldman, University of Connecticut Storrs
Cristina Takacs-Vesbach, University of New Mexico
Bernard Fry e, University of Texas Arlington
Monica Tischler, Benedictine University
Sandra Gibbons, University of Illinois at Chicago
Virginia Young, Mercer University
Elizabeth Good, University of Illinois at Urbana-Champaign
Jianmin Zhong, Humboldt State University
Melanie Griffin, Kennesaw State University
Janet Haynes, Long Island University, Brooklyn
The authors wish to extend their gratitude to our editors, Kathy
Michael Ibba, The Ohio State University
Lowenberg, Kathleen Timp, Angela FitzPatrick, Sandy Wille,
David Jenkins, Ihe University of Alabama Birmingham
and Lynn Breithaupt. We would also like to thank our photo
Dennis Kitz, Southern Illinois University Edwardsville
editor, Mary Reeg, and the tremendous talent and patience
James Koukl, Ihe University of Texas at Tyler
displayed by the artists. We are also very grateful to the many
Shashi Kumar, Saint Mary Mercy Hospital
reviewers who provided helpful criticism and analysis. Finally,
Jeffrey Leblond, Middle Tennessee State University
we thank our spouses and children who provided support and
Richard Long, University of South Carolina
tolerated our absences (mental, if not physical) while we
Jean Lu, Kennesaw State University
completed this demanding project.
xii
/>
Contents
About the Authors
Preface
5.3
5.4
iii
iv
Part One Introduction to Microbiology
0
G
0
The Evolution of Microorganisms
and Microbiology
1.1
1.2
Members of the Microbial World
1.3
Microbiology and Its Origins
1.4
Microbiology Today
Microbial Evolution
1
1
4
11
17
Microscopy
22
2.1
2.2
2.3
2.4
2.5
Lenses and the Bending of Light
22
Light Microscopes
23
31
34
39
Preparation and Staining of Specimens
Electron Microscopy
Scanning Probe Microscopy
Bacterial Cell Structure
3.1
3.2
3.3
3.4
The "Prokaryote" Controversy
A Typical Bacterial Cell
Bacterial Plasma Membranes
Bacterial Cell Walls
3.5
3.6
3.7
3.8
3.9
0
(s
Cell Envelope Layers Outside the Cell Wall
Bacterial Cytoplasm
External Structures
Bacterial Motility and Chemotaxis
Bacterial Endospores
Archaeal Cell Structure
4.1
4.2
4.3
4.4
4.5
A Typical Archaeal Cell
Archaeal Cell Envelopes
Archaeal Cytoplasm
External Structures
Comparison of Bacteria and Archaea
Eukaryotic Cell Structure
5.1
5.2
A Typical Eukaryotic Cell
Eukaryotic Cell Envelopes
96
Organelles of the Secretory
and Endocytic Pathways
97
5.5
Organelles Involved in Genetic Control
of the Cell
5.6
5.7
Organelles Involved in Energy Conservation
External Structures
101
103
104
Microbial Diversity & Ecology 5.1
5.8
There Was an Old Woman Who Swallowed a Fly
106
Comparison of Bacterial, Archaeal,
and Eukaryotic Cells
108
Viruses and Other Acellular Infectious Agents
6.1
6.2
Viruses
Virion Structure
112
112
113
Microbial Diversity & Ecology 6.1
Host-Independent Growth of an Archaeal Virus
6.3
6.4
6.5
6.6
6.7
Viral Multiplication
Types of Viral Infections
Cultivation and Enumeration of Viruses
114
119
124
127
129
130
42
42
43
47
53
Part Two Microbial Nutrition, Growth, and Control
54
0
Microbial Diversity & Ecology 3.1
Gram Positive and Gram Negative or
Monoderm s and Diderms?
Cytoplasm of Eukaryotes
61
62
69
72
76
82
82
84
87
88
90
92
92
94
Viroids and Satellites
Prions
Microbial Growth
7.1
7.2
Reproductive Strategies
Bacterial Cell Cycle
133
133
134
Microbial Diversity & Ecology 7.1
Cytokinesis Without FtsZ
7.3
7.4
7.5
7.6
7.7
7.8
(s
Influences of Environmental Factors
on Growth
Microbial Growth in Natural Environments
Laboratory Culture of Cellular Microbes
Growth Curve: When One Becomes
Two and Two Become Four ...
Measurement of Microbial Population Size
Continuous Culture of Microorganisms
Control of Microorganisms in the Environment
8.1
8.2
8.3
Principles of Microbial Control
The Pattern of Microbial Death
Mechanical Removal Methods
137
141
149
154
160
164
168
172
172
174
175
xiii
8.4
8.5
8.6
c;
Physical Control Methods
Chemical Control Agents
Evaluation of Antimicrobial
Agent Effectiveness
177
180
11.7 Anaerobic Respiration
11.8 Fermentation
11.9 Catabolism of Organic Molecules Other
184
186
11.10 Chemolithotrophy
Antimicrobial Chemotherapy
189
Acid Mine Drainage
9.1
9.2
The Development of Chemotherapy
189
General Characteristics
of Antimicrobial Drugs
190
8.7
Biological Control of Microorganisms
Than Glucose
247
248
251
253
Microbial Diversity & Ecology 11 .1
9.3
9.4
9.5
9.6
9.7
9.8
Determining the Level
of Antimicrobial Activity
Antiprotozoan Drugs
193
195
201
203
205
Factors Influencing Antimicrobial
Drug Effectiveness
206
Antibacterial Drugs
Antifungal Drugs
Antiviral Drugs
11.11 Phototrophy
255
256
G 2 Anabolism: The Use of Energy
in Biosynthesis
266
12.1
12.2
12.3
12.4
12.5
12.6
266
268
269
272
274
Principles Governing Biosynthesis
Precursor Metabolites
C0 2 Fixation
Synthesis of Carbohydrates
Synthesis of Amino Acids
Synthesis of Purines, Pyrimidines,
and Nucleotides
12.7 Lipid Synthesis
281
283
Part Three Microbial Metabolism
~0
Introduction to Metabolism
210
10.1 Metabolism: Important Principles
and Concepts
10.2 ATP: The Major Energy Currency of Cells
10.3 Redox Reactions: Reactions of Central
Importance in Metabolism
Part Four Microbial Molecular Biology and Genetics
211
213
~3
215
10.4 Electron Transport Chains: Sets
10.5 Biochemical Pathways
10.6 Enzymes and Ribozymes
10.7 Regulation of Metabolism
216
219
220
224
Catabolism: Energy Release and Conservation
230
of Sequential Redox Reactions
~
11.1 Metabolic Diversity
and Nutritional Types
11.2
11.3
11.4
11.5
11 .6
Chemoorganotrophic Fueling Processes
Aerobic Respiration
From Glucose to Pyruvate
Tricarboxylic Acid Cycle
Electron Transport and Oxidative
Phosphorylation
230
232
235
235
239
239
~4
Bacterial Genome Replication
and Expression
287
13.1
13.2
13.3
13.4
13.5
13.6
13.7
13.8
Protein Maturation and Secretion
288
288
293
301
304
309
311
319
Regulation of Bacterial Cellular Processes
325
14.1 Levels of Regulation
14.2 Regulation ofTranscription Initiation
14.3 Regulation ofTranscription
326
326
DNA as Genetic Material
Nucleic Acid and Protein Structure
DNA Replication in Bacteria
Bacterial Gene Structure
Transcription in Bacteria
The Genetic Code
Translation in Bacteria
Elongation
14.4 Regulation of Translation
14.5 Regulating Complex Cellular Processes
xiv
/>
333
336
338
Contents
~5
Eukaryotic and Archaeal Genome Replication
and Expression
18.5
18.6
18.7
18.8
353
15.1 Why Consider Eukaryotic and Archaeal
Genetics Together?
15.2 DNA Replication
15.3 Transcription
354
354
Proteomics
437
Systems Biology
440
440
443
Comparative Genomics
Metagenomics
358
Part Five The Diversity of the Microbial World
363
367
(, 9 Microbial Taxonomy and the Evolution
15.4 Translation and Protein Maturation and
Localization
15.5 Regulation of Cellular Processes
~6
Mechanisms of Genetic Variation
372
16.1 Mutations
16.2 Detection and Isolation of Mutants
16.3 DNA Repair
372
378
16.4
16.5
16.6
16.7
16.8
16.9
~7
Bacterial Transformation
380
383
385
387
393
Transduction
396
Evolution in Action: The Development of
Antibiotic Resistance in Bacteria
398
Creating Additional Genetic Variability
Transposable Elements
Bacterial Conjugation
Recombinant DNA Technology
17.1
Key Developments in Recombinant
DNA Technology
404
405
Techniques & Applications 17.1
Streptavidin-Biotin Binding and Biotechnology
17.2 Polymerase Chain Reaction
17.3 Cloning Vectors and Creating
Recombinant DNA
410
411
412
Techniques & Applications 17.2
How to Build a Microorganism
416
17.4 Construction of Genomic Libraries
17.5 Introducing Recombinant DNA
417
into Host Cells
418
17.6 Expressing Foreign Genes
in Host Cells
c;s
419
Microbial Genomics
424
18.1
18.2
18.3
18.4
424
429
431
433
Determining DNA Sequences
Genome Sequencing
Bioinformatics
Functional Genomics
of Diversity
447
19.1 Introduction to Microbial Taxonomy
19.2 Taxonomic Ranks
19.3 Exploring Microbial Taxonomy and Phylogeny
448
449
450
19.4 Phylogenetic Trees
19.5 Evolutionary Processes and the Concept
456
of a Microbial Species
19.6 Bergey's Manual of Systematic Bacteriology
459
464
Microbial Diversity & Ecology 19.1
"Official" Nomenclature Lists- A Letter from Bergey's
c;o TheArchaea
~1
465
469
20.1 Overview of the Archaea
20.2 Phylum Crenarchaeota
20.3 Phylum Euryarchaeota
470
476
480
The Deinococci, Mollicutes, and
Nonproteobacterial Gram-Negative Bacteria
489
21.1
Aquificae and Thermotogae
21.2
21.3
21.4
21.5
21.6
21.7
21.8
Deinococcus-Thermus
Class Mollicutes (Phylum Tenericutes)
Photosynthetic Bacteria
Phylum Planctomycetes
Phylum Chlamydiae
Phylum Spirochaetes
Phylum Bacteroidetes
21.9 Phylum Verrucomicrobia
(i2 The Proteobacteria
22.1 Class Alphaproteobacteria
22.2 Class Betaproteobacteria
22.3 Class Gammaproteobacteria
490
490
491
494
501
501
504
506
507
509
510
518
522
Microbial Diversity & Ecology 22.1
Bacterial Bioluminescence
22.4 Class Deltaproteobacteria
22.5 Class Epsilonproteobacteria
530
533
538
XV
G 3 Firmicutes: The Low G + C
Gram-Positive Bacteria
542
23.1
543
Class Clostridia
23.2 Class Bacilli
~4
547
Actinobacteria: The High G + C
Gram-Positive Bacteria
555
24.1
Order Actinomycetales
557
24.2 Order Bifidobacteriales
566
65
0
Biogeochemical Cycling and Global
Climate Change
632
28.1
633
Biogeochemical Cycling
28.2 Global Climate Change
The Protists
568
25.1
569
Overview of Protists
Part Six Ecology and Symbiosis
G9
642
Methods in Microbial Ecology
646
29.1
647
Culturing Techniques
29.2 Assessing Microbial Diversity
651
29.3 Assessing Microbial Community Activity
655
(io
Microorganisms in Marine and Freshwater
Ecosystems
660
25.2 Supergroup Excavata
571
25.3 Supergroup Amoebozoa
573
30.1
662
Water as a Microbial Habitat
661
25.4 Supergroup Rhizaria
574
30.2 Microorganisms in Marine Ecosystems
25.5 Supergroup Chromalveolata
577
30.3 Microorganisms in Freshwater Ecosystems
672
25.6 Supergroup Archaeplastida
584
Microorganisms in Terrestrial Ecosystems
679
31.1
680
c;6 The Fungi (Eumycota)
~
Soils as a Microbial Habitat
588
31.2 Microorganisms in the Soil Environment
683
590
31.3 Microbe-Plant Interactions
684
26.2 Chytridiomycota
593
31.4 The Subsurface Biosphere
696
26.3 Zygomycota
593
26.4 Glomeromycota
594
Microbial Interactions
699
32.1
Microbial Interactions
700
Microbial Diversity & Ecology 32.1
Wolbachia pipienris: The World's Most
Infectious Microbe?
701
26.1
Overview of Fungal Biology
26.5 Ascomycota
595
26.6 Basidiomycota
598
0
Disease 26.1
White-Nose Syndrome Is Decimating
North American Bat Populations
26.7 Microsporidia
c;7 Viruses
27.1
Virus Taxonomy and Phylogeny
27.2 Double-Stranded DNA Viruses
713
Microbial Diversity & Ecology 32.2
601
Do Bacteria Make People Fat?
32.3 Normal Microbiota of the Human Body
604
714
715
604
606
Microbial Diversity & Ecology 27.1
What Is a Virus?
32.2 Human-Microbe Interactions
599
617
Part Seven Pathogenicity and Host Response
~3
Innate Host Resistance
723
724
27.3 Single-Stranded DNA Viruses
617
33.1
27.4 RNA Viruses: Unity Amidst Diversity
619
27.5 Double-Stranded RNA Viruses
620
33.2 Physical and Mechanical Barrier
Defenses of Innate Resistance
725
27.6 Plus-Strand RNA Viruses
622
33.3 Chemical Mediators in Innate Resistance
728
27.7 Minus-Strand RNA Viruses
624
33.4 Cells, Tissues, and Organs
of the Immune System
735
27.8 Retroviruses
626
33.5 Phagocytosis
743
27.9
628
33.6 Inflammation
748
Reverse Transcribing DNA Viruses
Innate Resistance Overview
xvi
/>
Contents
G4
Adaptive Immunity
753
34.1
34.2
34.3
34.4
753
755
756
757
760
764
767
Overview of Adaptive Immunity
Antigens
Types of Adaptive Immunity
Recognition of Foreignness
34.5 T-Cell Biology
34.6 B-Cell Biology
34.7 Antibodies
37.3 Measuring Infectious Disease Frequency
37.4 Patterns of Infectious Disease
in a Population
776
34.8 Action of Antibodies
34.9 Acquired Immune Tolerance
34.10 Immune Disorders
777
35.3 Exposure and Transmission
Diseases and Pathogens
37.6 Health-Care-Associated Infections
37.7 Prevention and Control of Epidemics
The First Immunizations
37.8 Bioterrorism Preparedness
778
779
839
841
843
789
790
793
802
846
848
Historical Highlights 37.6
1346- The First Record ed Biological
Warfare Attack
~8
Historical Highlights 35.1
The First Indications of Person-to -Person
Spread of an Infectious Disease
837
37.5 Emerging and Reemerging Infectious
Historical Highlights 37.5
Monoclonal Antibody Therapy
35.1 Pathogenicity and Infectious Disease
35.2 Viru lence
836
Historical Highlights 37.4
"Typhoid Mary"
Techniques & Applications 34.1
G 5 Pathogenicity and Infection
835
803
849
Human Diseases Caused by Viruses
and Prions
854
38.1
38.2
38.3
38.4
855
865
865
878
Airborne Diseases
Arthropod-Borne Diseases
Direct Contact Diseases
Food-Borne and Waterborne Diseases
Historical Highlights 38.1
A Brief History of Polio
Part Eight Microbial Diseases, Detection,
38.5 Zoonotic Diseases
and Their Control
~6
Clinical Microbiology and Immunology
Laboratory
from Specimens
36.4 Clinical Immunology
f 7 Epidemiology and Public Health Microbiology
37.1
Epidemiology
Viral Hemorrhagic Fevers: A Microbial
History Lesson
808
809
812
820
830
830
37.2 Epidemiolog ical Methods
~9
Human Diseases Caused by Bacteria
888
39.1 Airborne Diseases
39.2 Arthropod-Borne Diseases
39.3 Direct Contact Diseases
888
898
901
Disease 39.1
A Brief History of Syphilis
Biofilms
831
909
39.4 Food-Borne and Waterborne Diseases
910
915
Techniques & Applications 39.3
832
832
Historical Highlights 37.3
SARS: Evolution of a Virus
885
Disease 39.2
Historical Highlights 37.2
John Snow, the First Epidemiologist
882
38.6 Prion Diseases
808
Historical Highlights 37.1
The Birth of Public Health in the
United States
881
Disease 38.2
36.1 Overview of the Clinical Microbiology
36.2 Biosafet y
36.3 Identification of Microorganisms
881
833
Clostridial Toxins as Therapeutic Agents:
Benefi ts of Nature's Most Toxic Proteins
39.5 Zoonotic Diseases
39.6 Opportunistic Diseases
9 19
924
926
xvii
~0
Human Diseases Caused by Fungi and Protists
932
40.1
Pathogenic Fungi and Protists
~2
Biotechnology and Industrial Microbiology
979
980
932
42.1
40.2 Airborne Diseases
934
42.2 Biofuel Production
982
40.3 Arthropod-Borne Diseases
937
42.3 Growing Microbes in Industrial Settings
983
938
42.4 Microorganisms Used in
Industrial Microbiology
985
944
42.5 Agricultural Biotechnology
990
42.6 Microbes as Products
992
Applied Environmental Microbiology
996
43.1
996
Disease 40.1
A Brief History of Malaria
40.4 Direct Contact Diseases
40.5 Food-Borne and
Waterborne Diseases
948
40.6 Opportunistic Diseases
952
~3
Major Products of Industrial Microbiology
Water Purification and Sanitary Analysis
Techniques & Applications 43.1
Part Nine Applied Microbiology
~1
Waterborne Diseases, Water Supplies,
and Slow Sand Filtration
999
Microbiology of Food
958
43.2 Wastewater Treatment
1001
41.1
959
43.3 Microbial Fuel Cells
1008
41.2 Controlling Food Spoilage
961
43.4 Biodegradation and Bioremediation
1009
41.3 Food-Borne Disease Outbreaks
964
41.4 Detection of Food-Borne Pathogens
967
41.5 Microbiology of Fermented Foods
969
Microbial Growth and Food Spoilage
Techniques & Applications 41.1
Chocolate: The Sweet Side of Fermentation
41.6 Probiotics
Appendix 1
A Review of the Chemistry
of Biological Molecules A-1
Appendix 2
Common Metabolic Pathways
970
976
Glossary
Credits
Index
G-1
C-1
1-1
xviii
/>
A-9
Brief Contents
About the Authors
Preface
24 Actinobacteria: The High G + C Gram-Positive
Bacteria 555
iii
iv
25 The Protists
568
26 The Fungi (Eumycota)
Part One Introduction to Microbiology
27 Viruses
588
604
1 The Evolution of Microorganisms and Microbiology
2 Microscopy
22
3 Bacterial Cell Structure
42
4 Archaeal Cell Structure
82
5 Eukaryotic Cell Structure
Part Six Ecology and Symbiosis
28 Biogeochemical Cycling and Global
Climate Change 632
92
6 Viruses and Other Acellular Infectious Agents
29 Methods in Microbial Ecology
112
646
30 Microorganisms in Marine and Freshwater
Ecosystems 660
Part Two Microbial Nutrition, Growth, and Control
7 Microbial Growth
31
Microorganisms in Terrestrial Ecosystems
32 Microbiallnteractions
133
8 Control of Microorganisms in the Environment
9 Antimicrobial Chemotherapy
172
189
Part Seven Pathogenicity and Host Response
33 Innate Host Resistance
34 Adaptive Immunity
Part Three Microbial Metabolism
10 Introduction to Metabolism
11 Catabolism: Energy Release and Conservation
230
12 Anabolism: The Use of Energy in Biosynthesis
266
13 Bacterial Genome Replication and Expression
14 Regulation of Bacterial Cellular Processes
287
325
789
36 Clinical Microbiology and Immunology
808
37 Epidemiology and Public Health Microbiology
830
38 Human Diseases Caused by Viruses and Prions
854
39 Human Diseases Caused by Bacteria
15 Eukaryotic and Archaeal Genome Replication
and Expression 353
18 Microbial Genomics
753
Part Eight Microbial Diseases, Detection, and Their
Control
Part Four Microbial Molecular Biology and Genetics
17 Recombinant DNA Technology
723
35 Pathogenicity and Infection
210
16 Mechanisms of Genetic Variation
679
699
888
40 Human Diseases Caused by Fungi and Protists
372
Part Nine Applied Microbiology
404
424
41
Microbiology of Food
958
42 Biotechnology and Industrial Microbiology
43 Applied Environmental Microbiology
996
Part Five The Diversity of the Microbial World
19 Microbial Taxonomy and the Evolution of Diversity
20 The Archaea
932
447
469
Appendix 1
Appendix 2
21 The Deinococci, Mollicutes, and Nonproteobacterial
Gram-Negative Bacteria 489
Glossary
22 The Proteobacteria
Credits
509
23 Firmicutes: The Low G + C Gram-Positive Bacteria
542
Index
G-1
C-1
1-1
A Review of the Chemistry
of Biological Molecules A-1
Common Metabolic Pathways
A-9
979
/>
1
The Evolution
of Microorganisms
and Microbiology
Artist's rendition of the six planets orbiting a star called Kepler-11.
The drawing is based on observations made of the system by the
Kepler spacecraft on August 26,2010. Some are Earth-sized and
may be habitable by life.
Over 2,000 Potential Planets Discovered
thus a major contributor to the functioning of the biosphere. In addition to
these familiar types of metabolism, other microbes are able to use inorganic
molecules as sources of energy in both oxic (oxygen available) and anoxic
I
n February 2012, the National Aeronautics and Space Administration
(NASA) reported that over 2,000 potential planets had been discovered
by the 2009 Kepler mission. Using a telescope in space, the light
emanating from stars as far as 3,000 light-years away had been
monitored every half-hour. The Kepler telescope identified planets as
they circulated their star and caused a brief decrease in emitted light; just
as an object is detected as a blip by radar, a blip of "darkness" indicates a
planet.
Unless you are a science fiction fan, you might wonder why NASA is
interested in finding planets. By finding other planets, scientists can
gather evidence to support or refute current models of planet formation.
(no oxygen) conditions. It is these microbes that are of particular interest to
NASA scientists, as it is thought that the organisms on other planets may
have similar unusual metabolisms.
Our goal in this chapter is to introduce you to this amazing group of
organisms and to outline the history of their evolution and discovery.
Microbiology is a biological science, and as such, much of what you will learn
in this text is similar to what you have learned in high school and college
biology classes that focus on large organisms. But microbes have unique
properties, so microbiology has unique approaches to understanding them.
These too will be introduced. But before you delve into this chapter, check to
see if you have the background needed to get the most from it.
These models predict a process that is chaotic and violent. Planets are
thought to begin as dust particles circling around newly formed stars. As
Readiness Check:
these particles collide, they grow in size, forming larger chunks. Eventually
Based on what you have learned previously, you should be able to:
a series of such collisions results in planet-sized bodies. Astrobiologists are
tl List the features of eukaryotic cells that distinguish them from other
interested in identifying characteristics of a planet that may allow it to
support life. Using Earth as a model, they hypothesize that life-supporting
cell types
tl List the attributes that scientists use to determine if an object is alive
planets will share many features with Earth. But how will life be recog
nized? Again, scientists look to life on Earth to answer this question, and
increasingly they are turning to microbiologists for help.
Earth formed 4.5 billion years ago. Within the next billion years, the
1.1 Members of the Microbial World
first cellular life forms-microbes-appeared. Since that time, microorgan
After reading this section, you should be able to:
isms have evolved and diversified to occupy virtually every habitat on Earth:
• Differentiate the biological entities studied by microbiologists
from oceanic geothermal vents to the coldest Arctic ice. The diversity of
cellular microorganisms is best exemplified by their metabolic capabilities.
Some carry out respiration, just as animals do. Others perform photosynthe
sis, rivaling plants in the amount of carbon dioxide they capture, forming
organic matter and releasing oxygen into the atmosphere. Indeed,
Prochlorococcus, a cyanobacterium (formerly called a blue-green alga), is
thought to be the most abundant photosynthetic organism on Earth and
from those studied by other biologists
• Explain Carl Woese's contributions in establishing the three
domain system for classifying cellular life
• Provide an example of the importance to humans of each of the
major types of microbes
• Determine the type of microbe (e.g., bacterium, fungus, etc.) when
given a description of a newly discovered microbe
2
CHAPTER 1
I
The Evolution of Microorganisms and Microbiology
Organisms and
biological entities
studied by
microbiologists
I
can be
includes
e.g.
e.g.
•
Yeasts
Molds
Figure
1.1
•
Algae
Protozoa
Slime molds
includes
e.g.
•
Escherichia
coli
e.g.
composed of
composed of
composed of
ckJ
•
Methanogens
composed of
~
Concept Map Showing the Types of Biological Entities Studied by Microbiologists.
M 1 C RO IN Q u 1 RY How would you alter this concept map so that it also distinguishes the cellular organisms from each other?
Microorganisms are defined as those organisms and acellular
into compartments ("rooms") by membranes ("walls"). The
biological entities too small to be seen clearly by the unaided
most obvious characteristic of these cells is that they lack the
eye
(figure 1.1). They are generally
1 millimeter or less in diam
membrane-delimited nucleus obser ved in
eukaryotic cells
eter. Although small size is an important characteristic of mi
(Greek eu, true, and karyon, nut or kernel). Eukaryotic cells
crobes, it alone is not sufficient to define them. Some cellular
not only have a nucleus but also many other membrane-bound
microbes, such as bread molds and filamentous photosynthetic
organelles that separate some cellular materials and processes
microbes, are actually visible without microscopes. These mac
from others.
roscopic microbes are often colonial, consisting of small aggre
These observations eventually led to the development of a
gations of cells. Some macroscopic microorganisms are
classification scheme that divided organisms into five kingdoms:
multicellular. They are distinguished from other multicellular
Monera, Protista, Fungi, Animalia, and Plantae. Microorganisms
life forms such as plants and animals by their lack of highly dif
(except for viruses and other acellular infectious agents, which
ferentiated tissues. Most unicellular microbes are microscopic.
have their own classification system) were placed in the first three
However, there are interesting exceptions, as we describe in
kingdoms. In this scheme, all organisms with prokaryotic cell
chapter 3. In summary, cellular microbes are usually smaller
structure were placed in Monera. The five-kingdom system was an
than 1 millimeter in diameter, often unicellular and, if multi
important development in microbial taxonomy, but it is no longer
cellular, lack differentiated tissues.
accepted by microbiologists. This is because not all "prokaryotes"
The diversity of microorganisms has always presented a
are the same and therefore should not be grouped together in a
challenge to microbial taxonomists. The early descriptions of
single kingdom. Furthermore, it is currently argued that the term
cellular microbes as either plants or animals were too simple.
prokaryote is not meaningful and should be abandoned. As we
For instance, some microbes are motile like animals but also
describe next, this discovery required several advances in the
have cell walls and are photosynthetic like plants. Such mi
tools used to study microbes.
crobes cannot be placed easily into either kingdom. An im
(section 3.1)
�I The ''prokaryote" controversy
portant breakthrough in microbial taxonomy arose from
Great progress has been made in three areas that profoundly
studies of their cellular architecture, when it was discovered
affect microbial classification. First, much has been learned
that cells exhibited one of two possible "floor plans." Cells that
about the detailed structure of microbial cells from the use of
prokaryotic cells (Greek pro, before, and
electron microscopy. Second, microbiologists have determined
came to be called
karyon, nut or kernel; organisms with a primordial nucleus)
the biochemical and physiological characteristics of many dif
have an open floor plan. That is, their contents are not divided
ferent microorganisms. Third, the sequences of nucleic acids and
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1.1 Members of the Microbial World
3
proteins from a wide variety of organisms have been compared.
The comparison of ribosomal RNA (rRNA), begun by Carl
Woese in the 1970s, was instrumental in demonstrating that
there are two very different groups of organisms with prokary
otic cell architecture:
Bacteria and Archaea. Later studies based
Protista is not a cohesive
on rRNA comparisons showed that
taxonomic unit (i.e., taxon) and that it should be divided into
three or more kingdoms. These studies and others have led many
taxonomists to reject the five-kingdom system in favor of one
that divides cellular organisms into three domains:
(sometimes referred to as true bacteria or eubacteria),
Bacteria
Archaea
(sometimes called archaeobacteria or archaebacteria), and
Eukar ya (all eukaryotic organisms) (figure 1.2). We use this
system throughout the text. A brief description of the three
domains and of the microorganisms placed in them follows.
�I Nucleic acids (appendix I); Proteins (appendix I)
Members of domain Bacteria are usually single-celled or
ganisms.1 Most have cell walls that contain the structural mol
ecule peptidoglycan. Although most bacteria exhibit typical
prokaryotic cell structure (i.e., they lack a membrane-bound
nucleus), a few members of the unusual phylum Planctomycetes
1--1
rRNA sequence change
inconsistency is another argument made for abandoning the
� Unresolved branching order
have their genetic material surrounded by a membrane. This
term "prokaryote." Bacteria are abundant in soil, water, and
air, including sites that have extreme temperatures, pH, or sa
linity. Bacteria are also major inhabitants of our skin, mouth,
and intestines. Indeed, more microbial cells are found in and
on the human body than there are human cells. These microbes
begin to colonize humans shortly after birth. As the microbes
Figure
1.2 Universal Phylogenetic Tree.
These evolutionary
relationships are based on rRNA sequence comparisons. To save space,
many lineages have not been identified.
establish themselves, they contribute to the development of the
MICRO 1 N Q u 1 RY How many of the taxa listed in the figure include
body's immune system. Those microbes that inhabit the large
microbes?
intestine help the body digest food and produce vitamins. In
these and other ways, microbes help maintain the health and
well-being of their human hosts.
�I Phylum Planctomycetes
(section 21.5)
Unfortunately, some bacteria cause disease, and some of
Members of domain Archaea are distinguished from bacte
ria by many features, most notably their distinctive rRNA
these diseases have had a huge impact on human history. In 1347
sequences, lack of peptidoglycan in their cell walls, and unique
the plague (Black Death), an arthropod-borne disease, struck
membrane lipids. Some have unusual metabolic characteristics,
Europe with brutal force, killing one-third of the population
such as the methanogens, which generate methane (natural) gas.
(about 25 million people) within four years. Over the next
Many archaea are found in extreme environments, including
80 years, the disease struck repeatedly, eventually wiping out
those with high temperatures (thermophiles) and high concen
75% of the European population. The plague's effect was so
trations of salt (extreme halophiles). Although some archaea are
great that some historians believe it changed European culture
members of a community of microbes involved in gum disease
and prepared the way for the Renaissance. Because of such
in humans, their role in causing disease has not been clearly
plagues, it is easy for people to think that all bacteria are patho
established.
gens, but in fact, relatively few are. Most play beneficial roles,
from global impact to maintaining human health. They break
Domain
Eukarya includes microorganisms classified as
protists or fungi. Animals and plants are also placed in this
down dead plant and animal material and, in doing so, cycle
domain.
elements in the biosphere. Furthermore, they are used exten
most bacteria and archaea. They have traditionally been di
Protists are generally unicellular but larger than
sively in industry to make bread, cheese, antibiotics, vitamins,
vided into protozoa and algae. Despite their use, none of these
enzymes, and other products.
terms has taxonomic value as protists, algae, and protozoa do
1
In this text, the term bacteria (s., bacterium) is used to refer to those microbes belonging to domain Bacteria, and the term archaea (s., archaean) is used to refer to those that belong to domain Archaea.
In some publications, the term bacteria is used to refer to all cells having prokaryotic cell structure. That is not the case in this text.
4
CHAPTER 1
I
The Evolution of Microorganisms and Microbiology
not form cohesive taxa. However, for convenience, we use
them here.
The major types of protists are algae, protozoa, slime molds,
and water molds.
Algae are photosynthetic. They, together with
cyanobacteria, produce about 75% of the planet's oxygen and are
Retrieve, Infer, Apply
1.
How did the methods used to classify microbes change, particularly
in the last half of the twentieth century? What was the result of
these technological advances?
the foundation of aquatic food chains. Protozoa are unicellular,
2. Identify one characteristic for each of these types of microbes that
animal-like protists that are usually motile. Many free-living
distinguishes it from the other types: bacteria, archaea, protists,
protozoa function as the principal hunters and grazers of the
fungi, viruses, viroids, satellites, and prions.
microbial world. They obtain nutrients by ingesting organic
matter and other microbes. They can be found in many different
environments, and some are normal inhabitants of the intestinal
tracts of animals, where they aid in digestion of complex materi
als such as cellulose. A few cause disease in humans and other
animals.
Slime molds
are protists that behave like protozoa in
one stage of their life cycle but like fungi in another. In the pro
tozoan phase, they hunt for and engulf food particles, consum
ing decaying vegetation and other microbes.
Water molds
are
protists that grow on the surface of freshwater and moist soil.
They feed on decaying vegetation such as logs and mulch. Some
water molds have produced devastating plant infections, includ
ing the Great Potato Famine of 1846-1847 in Ireland ...I
protists (chapter 25)
Fungi are a diverse
The
group of microorganisms that range
from unicellular forms (yeasts) to molds and mushrooms. Molds
and mushrooms are multicellular fungi that form thin, thread
like structures called hyphae. They absorb nutrients from their
environment, including the organic molecules they use as
sources of carbon and energy. Because of their metabolic capa
bilities, many fungi play beneficial roles, including making
bread rise, producing antibiotics, and decomposing dead organ
isms. Some fungi associate with plant roots to form mycorrhi
zae. Mycorrhizal fungi transfer nutrients to the roots, improving
growth of the plants, especially in poor soils. Other fungi
cause plant diseases (e.g., rusts, powdery mildews, and smuts)
and diseases in humans and other animals. ..I
(chapter 26)
The Fungi
The microbial world also includes numerous acellular infec
tious agents.
Viruses
are acellular entities that must invade a
1.2 Microbial Evolution
After reading this section, you should be able to:
• Propose a time line of the origin and history of microbial life and
integrate supporting evidence into it
• Design a set of experiments that could be used to place a newly
discovered cellular microbe on a phylogenetic tree based on small
subunit (SSU) rRNA sequences
• Compare and contrast the definitions of plant and animal species,
microbial species, and microbial strains
A review of figure 1.2 reminds us that in terms of the number of
taxa, microbes are the dominant organisms on Earth. How has
microbial life been able to radiate to such an astonishing level of
diversity? To answer this question, we must consider microbial
evolution. The field of microbial evolution, like any other scien
tific endeavor, is based on the formulation of hypotheses, the
gathering and analysis of data, and the reformation of hypotheses
based on newly acquired evidence. That is to say, the study of
microbial evolution is based on the scientific method
.mhhe.com/willey9).
(see
www
To be sure, it is sometimes more difficult to
amass evidence when considering events that occurred millions,
and often billions, of years ago, but the advent of molecular meth
ods has offered scientists a living record of life's ancient history.
This section describes the outcome of this scientific research.
Evidence for the Origin of Life
host cell to multiply. The simplest viruses are composed only of
Dating meteorites through the use of radioisotopes places our
proteins and a nucleic acid, and can be extremely small (the
planet at an estimated 4.5 to 4.6 billion years old. However, con
smallest is 10,000 times smaller than a typical bacterium). How
ditions on Earth for the first 100 million years or so were far too
ever, their small size belies their power: they cause many animal
harsh to sustain any type of life. Eventually bombardment by
and plant diseases and have caused epidemics that have shaped
meteorites decreased, water appeared on the planet in liquid
human history. Viral diseases include smallpox, rabies, influ
form, and gases were released by geological activity to form
enza, AIDS, the common cold, and some cancers. Viruses also
Earth's atmosphere. These conditions were amenable to the ori
play important roles in aquatic environments, and their role in
gin of the first life forms. But how did this occur, and what did
shaping aquatic microbial communities is currently being ex
these life forms look like?
Viroids
satellites
are infectious agents composed
Clearly, in order to find evidence of life and to develop
only of ribonucleic acid (RNA). Viroids cause numerous plant
hypotheses about its origin and subsequent evolution, scien
diseases, whereas satellites cause plant diseases and some im
tists must be able to define life. Although even very young
plored.
and
prions, infec
children can examine an object and correctly determine
tious agents composed only of protein, are responsible for
whether it is living or not, defining life succinctly has proven
causing a variety of spongiform encephalopathies such as scra
elusive for scientists. Thus most definitions of life consist of a
Viruses and other acellular in
set of attributes. The attributes of particular importance to
portant animal diseases such as hepatitis. Finally,
pie and "mad cow disease." .. I
fectious agents (chapter 6)
paleobiologists are an orderly structure, the ability to obtain
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