Prescott's Microbiology 11th Edition Eleventh 2019

2125 Pages·2022·278.511 MB·English

by  Joanne Willey, Kathleen Sandman, Dorothy Wood

eleventh edition
Prescott’s Microbiology
Joanne M. Willey
HOFSTRA UNIVERSITY
Kathleen M. Sandman
Dorothy H. Wood
DURHAM TECHNICAL COMMUNITY
COLLEGE
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PRESCOTT’S MICROBIOLOGY, ELEVENTH EDITION
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Library of Congress Cataloging-in-Publication Data
Willey, Joanne M., author.
Prescott’s microbiology / Joanne M. Willey, Hofstra University,
Kathleen M. Sandman, Dorothy H. Wood, Durham Technical Community College.
Microbiology
Eleventh edition. | New York, NY : McGraw-Hill Education, [2020] |
Includes bibliographical references and index.
LCCN 2018026319 | ISBN 9781260211887 (student edition)
LCSH: Microbiology—Textbooks.
LCC QR41.2 .P74 2020 | DDC 579—dc23
LC record available at https://lccn.loc.gov/2018026319
The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not
indicate an endorsement by the authors or McGraw-Hill Education, and McGraw-Hill Education does not guarantee the
accuracy of the information presented at these sites.
mheducation.com/highered
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Brief Contents
About the Authors iv 25 Fungi 579
26 Viruses 594
Part One Introduction to Microbiology
Part Six Ecology and Symbiosis
1 The Evolution of Microorganisms and
Microbiology 1 27 Microbial Interactions 619
2 Microscopy 20 28 Biogeochemical Cycling and Global Climate
3 Bacterial Cell Structure 40 Change 632
4 Archaeal Cell Structure 77 29 Methods in Microbial Ecology 646
5 Eukaryotic Cell Structure 87 30 Microorganisms in Marine and Freshwater
Ecosystems 658
6 Viruses and Other Acellular Infectious Agents 106
31 Microorganisms in Terrestrial Ecosystems 675
Part Two Microbial Nutrition, Growth, and Control
Part Seven Pathogenicity and Host Response
7 Bacterial and Archaeal Growth 128
32 Innate Host Resistance 693
8 Control of Microorganisms in the Environment 170
33 Adaptive Immunity 719
9 Antimicrobial Chemotherapy 187
34 The Microbe-Human Ecosystem 752
35 Infection and Pathogenicity 770
Part Three Microbial Metabolism
10 Introduction to Metabolism 209 Part Eight Microbial Diseases, Detection, and
11 Catabolism: Energy Release and Conservation 228 Their Control
12 Anabolism: The Use of Energy in Biosynthesis 265 36 Epidemiology and Public Health Microbiology 786
37 Clinical Microbiology and Immunology 806
38 Human Diseases Caused by Viruses and Prions 825
Part Four Microbial Molecular Biology and Genetics
39 Human Diseases Caused by Bacteria 857
13 Bacterial Genome Replication and Expression 288
40 Human Diseases Caused by Fungi and Protists 900
14 Regulation of Bacterial Cellular Processes 326
15 Eukaryotic and Archaeal Genome Replication and
Part Nine Applied Microbiology
Expression 355
16 Mechanisms of Genetic Variation 375 41 Microbiology of Food 925
17 Microbial DNA Technologies 405 42 Biotechnology and Industrial Microbiology 944
18 Microbial Genomics 425 43 Applied Environmental Microbiology 960
Appendix 1 A Review of the Chemistry of
Biological Molecules A-1
Part Five The Diversity of the Microbial World
Appendix 2 Common Metabolic Pathways A-9
19 Microbial Taxonomy and the Evolution of Diversity 447
Appendix 3 Microorganism Pronunciation Guide A-17
20 Archaea 466
21 Nonproteobacterial Gram-Negative Bacteria 484 Glossary G-1
22 Proteobacteria 504 Index I-1
23 Gram-Positive Bacteria 537
24 Protists 559
iii
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About the Authors
Courtesy of Joanne Willey Courtesy of Adele Anderson Courtesy of Dorothy Wood
Joanne M. Willey Kathleen M. Sandman Dorothy H. Wood
has been a received her has taught
professor at Hofstra University on Long B.A. in Biology from La Salle University microbiology and general biology at
Island, New York, since 1993, where she is and her Ph.D. in Cellular and Developmental Durham Technical Community College in
the Leo A. Guthart Professor of Biology from Harvard University. She was North Carolina since 2004. Dr. Wood
Biomedical Science and Chair of the inspired to a career in science by her older received her B.A. in Biology from Rhode
Department of Science Education at the brother’s experience as an organic chemist Island College where her love of microbes
Donald and Barbara Zucker School of and by the developing technology in began, nurtured by Dr. Charles Owens.
Medicine at Hofstra/Northwell. Dr. Willey recombinant DNA in the 1970s. Her She earned her Ph.D. in Cell and
received her B.A. in Biology from the graduate work used a transposable element Molecular Pathology from the University
University of Pennsylvania, where her as a mutagen in Bacillus subtilis to study of North Carolina at Chapel Hill, focusing
interest in microbiology began with work gene expression during endospore formation. on pancreatic damage caused by
on cyanobacterial growth in eutrophic She continued in the genetics of Gram- antimicrobial drugs, and investigated
streams. She earned her Ph.D. in positive bacteria with a postdoctoral year alternative therapies based on receptor
biological oceanography (specializing studying Bacillus thuringiensis at the binding by novel compounds. After three
in marine microbiology) from the University of Cambridge in the United years as Assistant Professor at NC Central
Massachusetts Institute of Technology– Kingdom. Another postdoctoral opportunity University, Dr. Wood made the move to
Woods Hole Oceanographic Institution at The Ohio State University provided an the NC Community College System to
Joint Program in 1987. She then went to introduction to the emerging field of focus her attention on her primary interest
Harvard University, where she spent her archaeal molecular biology, where of teaching. Throughout her career she has
postdoctoral fellowship studying the Dr. Sandman discovered archaeal histones developed several courses, including
filamentous soil bacterium Streptomyces and continued research in the structural graduate bacteriology, pathophysiology,
coelicolor. Dr. Willey has coauthored a biology of archaeal chromatin for about and biotechnology. She serves as a visiting
number of publications that focus on its 20 years. She served the National Science scholar at Duke University where she is a
complex developmental cycle. She is an Foundation as a research grant reviewer mentor for the Preparing Future Faculty
active member of the American Society and panelist for the Life in Extreme program. Dr. Wood is a member of the
for Microbiology (ASM), and served on Environments program, and has organized American Society for Microbiology and
the editorial board of the journal Applied conference sessions on archaeal molecular the Association of College and University
and Environmental Microbiology for nine biology and proteins from extremophiles. Biology Educators, as well as several local
years and as Chair of the Division of Dr. Sandman has taught microbiology to organizations that foster pedagogy. She is
General Microbiology. Dr. Willey taught hundreds of students, at both the a digital faculty consultant for McGraw-
microbiology to biology majors for 20 years introductory level and in an advanced Hill and has worked on several textbooks
and now teaches microbiology and molecular microbiology laboratory. in a variety of disciplines, developing and
infectious disease to medical students. She Dr. Sandman has worked as a consultant editing digital content to accompany the
has taught courses in cell biology, marine in a variety of industries, including texts. Outside of the classroom, Dr. Wood
microbiology, and laboratory techniques in industrial microbiology, environmental is a fitness professional, leads health and
molecular genetics. Dr. Willey lives on the geomicrobiology, and technical publishing. wellness seminars, and has been the
north shore of Long Island and has two She lives with her husband in Columbus, treasurer of a nonprofit organization for
grown sons. She is an avid runner and Ohio, and has two grown daughters. She the past 10 years. She enjoys life in
enjoys skiing, hiking, sailing, and reading. enjoys biking, fabric arts, reading, and North Carolina with her husband and two
She can be reached at joanne.m.willey@ genealogy, and can be reached at grown children and can be reached at
hofstra.edu. [email protected]. [email protected].
iv
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A Modern Approach to Microbiology
Brief Contents
Evolution as a Framework
Introduced immediately in chapter 1 and used as an overarching
theme throughout, evolution helps unite microbiological con- About the Authors iv 25 Fungi 579
26 Viruses 594
cepts and provides a framework upon which students can build
Part One Introduction to Microbiology
their knowledge. 1 The Evolution of Microorganisms and Part Six Ecology and Symbiosis
Microbiology 1 27 Microbial Interactions 619
2 Microscopy 20 28 Biogeochemical Cycling and Global Climate
3 Bacterial Cell Structure 40 Change 632
4 Archaeal Cell Structure 77 29 Methods in Microbial Ecology 646
5 Eukaryotic Cell Structure 87 30 Microorganisms in Marine and Freshwater
An Introduction to the Entire Microbial World 6 Viruses and Other Acellular Infectious Agents 106 Ecosystems 658
31 Microorganisms in Terrestrial Ecosystems 675
Covered in chapters 3–6, separate chapters on the structure and Part Two Microbial Nutrition, Growth, and Control Part Seven Pathogenicity and Host Response
7 Bacterial and Archaeal Growth 128 32 Innate Host Resistance 693
function of bacteria and archaea are followed by the discussion 8 Control of Microorganisms in the Environment 170 33 Adaptive Immunity 719
9 Antimicrobial Chemotherapy 187 34 The Microbe-Human Ecosystem 752
of eukaryotic cells and viruses. 35 Infection and Pathogenicity 770
Part Three Microbial Metabolism
10 Introduction to Metabolism 209 Part Eight Microbial Diseases, Detection, and
11 Catabolism: Energy Release and Conservation 228 Their Control
12 Anabolism: The Use of Energy in Biosynthesis 265 36 Epidemiology and Public Health Microbiology 786
Broad Coverage of Microbial Ecology 37 Clinical Microbiology and Immunology 806
Part Four Microbial Molecular Biology and Genetics 38 Human Diseases Caused by Viruses and Prions 825
39 Human Diseases Caused by Bacteria 857
The importance and multidisciplinary nature of microbial ecology 13 Bacterial Genome Replication and Expression 288 40 Human Diseases Caused by Fungi and Protists 900
14 Regulation of Bacterial Cellular Processes 326
are demonstrated by content that ranges from global climate 15 EEuxpkarerysostioicn a nd3 5A5rchaeal Genome Replication and Part Nine Applied Microbiology
change to the human microbiome. 16 Mechanisms of Genetic Variation 375 41 Microbiology of Food 925
17 Microbial DNA Technologies 405 42 Biotechnology and Industrial Microbiology 944
18 Microbial Genomics 425 43 Applied Environmental Microbiology 960
Appendix 1 A Review of the Chemistry of
Part Five The Diversity of the Microbial World Biological Molecules A-1
19 Microbial Taxonomy and the Evolution of Diversity 447 Appendix 2 Common Metabolic Pathways A-9
20 Archaea 466 Appendix 3 Microorganism Pronunciation Guide A-17
21 Nonproteobacterial Gram-Negative Bacteria 484 Glossary G-1
22 Proteobacteria 504 Index I-1
23 Gram-Positive Bacteria 537
24 Protists 559
34.3 A Functional Core Microbiome Is Required for Human Homeostasis 763 iii
M cell
Microbial
AAAnnnnnntttiiiggggggggggeeeeennnsssssss Antigens Molecular Microbiology and Immunology
DDDDDDDeeeeeeeeeennnnddddrrrrrrriiiiittttiiiiiccccccccccc
ccccccceeeellllllll The eleventh edition includes updates on genetics, biotechnol-
NNaaaiivvvveee TTT ccceellllll IIILLLLLCCC333333 HHHHHaaaaannnnnigtttttiiiih---mmmmmm-aiiiiifficccccrrrrrnoooooibbbbbtyiiiiiaaaa,llll Low-affinity IgA DDDeennnnddrriitttiicc ogy, genomics and metagenomics, immunology, and the hu-
IIIIIggggggggggAAAAAAAAAAAAA cceeellll
TTTHHHH man microbiome. A streamlined discussion of immunity, with
IILLLCCC33
enhanced detail between innate and adaptive linkages, helps
BB cceellll SSttiimmuullaatteedd BB cceeellll IIgggAAAA+++ IIggAAAA+++ BB cceellll students grasp the complexity and specificity of immune re-
IIIIgggggggAAAAAAA++++ BBBB--ccccceeeellllllllll pppppppooooooooooooooooolllll sponses. A new chapter, The Microbe-Human Ecosystem, in-
PPeeyyeerr’’ss ppaattcchhh TT-ccceeeeellllll--dddeeepppeeennndddeeennnttt TTT--ccceeelllll--iiinnnddeeppeendent Diffuse lymphoid tissue
IIIggAAA iiiinnndddduuucccttttiiiiiooonnn IIIIgggAAAA iiiinnndddduuccttiioonn troduces students to the development and impact of the human
Figure 34.7 Production of SIgA Requires ILC3 and Microbial Stimulation. B-cell stimulation can be T-cell dependent (left, Peyer’s patch) or T-cell microbiome.
independent (right, isolated lymphoid follicle). In both cases, innate lymphoid cells-3 (ILC3s) optimization of dendritic cell activity is needed. ILC3 function requires
interaction with host microbes or their products.
Intestinal Intestinal intrinsic nervous system, that is connected to the CNS via the
lumen barrier CNS vagus nerve. Gut bacteria directly stimulate neurons within the
enteric nervous system, which then communicate with the vagus
Immune system nvaegrvues. nTehrev ev acgauns dniefrfveere innt itautren btreatwnsemenit ss tsiimgnualil sf troo mth en borramina.l Tmhie-
• LPS crobiota and pathogens even in the absence of inflammation, al-
• Oprtohdeur cmtsicrobial Vagus nerve tinhvoeusgthig ahtioown .this fascinating feat is possible remains under
• Microbes signal enteric nervous Finally, as suggested by experiments in which butyrate and
system → vagal stimulation FMT had similar effects on the behavior of GF mice, soluble
microbial products can impact the CNS. To understand this,
Blood-brain barrier we must first consider the CNS as a site that is set apart from the
remainder of the body by the blood-brain barrier (BBB). The
• SCFAs BBB is lined with cells that protect the CNS from normal blood
components as well as toxins and infectious agents. This is
Figure 34.8 The Gut-Brain Axis. Gut microbes influence the central largely due to impermeable connections, called tight junctions,
nervous system using at least three mechanisms. LPS, lipopolysaccharide; between the cells that make up the BBB. Once again, a flurry of
SCFAs, short-chain fatty acids research was triggered by an observation made in GF mice:
(right top) ©GraphicsRF/Shutterstock Their BBBs have the undesirable quality of being more permea-
ble than conventional mice. And once again, it is SCFAs, par-
voifi aippetite, decreased motor activity, loss of sociability, and re- ticularly butyrate, that help maintain the BBB as an impenetrable
duced cognition (“trouble thinking”). barricade. Because SCFAs are most abundant when mice or hu-
Another mechanism that helps mediate the gut-brain axis is mans eat a high-fiber diet, the broader implication is that one’s
a direct pathway from gut to brain. This is possible because the diet can govern the entry of metabolites and other blood-borne
GI tract is lined with a network of nerves, called the enteric or compounds to the CNS.
wil11886_fm_i-xxiv.indd 8 23/10/18 2:13 pm
A Modern Approach to Microbiology
17.5 Cas9 Nuclease Is a Precise Tool for Genome Editing 421
procedures are general enough to be used for any cell into which 21st-Century Microbiology
17.5 Cas9 Nuclease Is a Precise Tool DNA can be introduced and expressed. Responses to viral
for Genome Editing infection (section 14.6)
Like restriction enzymes, Cas9 is an endonuclease that cuts Prescott’s Microbiology leads the way with text devoted to global
After reading this section, you should be able to: both strands of a target DNA. However, there is an important
a. Denisdtoinnguucilseha tshees DanNdA Creacso9g nnuitciloena sfeeatures of restriction dthieffye rreenccoeg nbieztew etheeni rt htea rtgweot styepqeuse nocfe n iunc ldeoasuebsl,e -isnt rtaenrmdesd o Df hNoAw. climate change, biofuels, and microbial fuel cells. For more, see
b. Ea xgpelanionm heow Cas9 nuclease can be directed to cut at a unique site in Rcoensttariccttsi obne tewnezeynm tehse rDeNcoAg nmizoel efcouulre taon dei gamht inboas aec pida isrisd eth crhoauignhs chapters 28, 30, 42, and 43.
c. Dhoiamgoralomg ohuosw r ea cnoemwb gineanteio mnay be inserted into a chromosome by icno nthseis etinnzgy omf ea (pfiogluyrpee 1p7ti.d2)e. aCnads 9a, ghuowideev RerN, iAs a ( griRboNnAuc).l eRoepcroogteniin-
tion of target DNA for cleavage occurs by hybridization of about
CuobfatitloIenhaanrfatsdaictheet ecleetty s esdthe,rnb9e r s nioao yr tspgborcfrin aae eeyopoJlCcfl nye drlu eeFaso nogerst dsecnhmredfu9 neoenei as idfgserf olhne Cs u l maeor y,Ztnt a dlor hgtehDcsat eiee 9tatdaroeip ni nsjgioncluaoan geenogcdClmgm,n ime nw nhRosigneadcopameetIn sru orSnDattoe egn orPonsm fNahe dmR drapontoiAi sicfetCmtEdo c osimclRormrs y iae avwonIu CmnpbdeSs sh RereavtPeabei apcibcdnIRedvtotSha us o, imdc .PClen.t oiti wlRtA eaerCcl ee esr-owuodsa9icC sanisaCrt tt,a 9leefsh iayhoscn s eon 9glraedgm ufd ,eare. mg s tpnrvorCaethceoeeeedrnehafnmclrcieo ose fmhthgrm9iepidaaeeer o e nenronedersd ac iuude tase,a, igpci w tnndtnwbliusidecgidin,ulthd a a igotdtttesne r nhhheeirllgiynyeeeeess-. vftittknf(2qnshhaeru0iaieowrgeerzre geam e ybn C elgDstIofa c ht Rniaam tNseagoi ssteN c umutA9eoisf anrsqAe b g cien abwuciftek unloheer1ein .codsrten n4 ow l b S gcue.dbgmt2eeeeehaceaes7n eqelsessn,ienee.)ut eso o,rt tk e s Imhuiittaa.encnniheclnyd ce me tead ece iC (lot so soegfomRt n nimshR wgi.ttIigne rucSNeiTe calrnrCPntleAshoeth Rtorasebc , et 1ms ui aa (9aeal7tinCboex l n. dantc 1rtRclheerc uuy3eeeIsidtsc )tleSmt .iusllrrgtieP inisieseaccnoR pi ntsotqgpsie heomopu hlc emobneeapip cfe,c gsre liueciooeece,fn astm uvaioe auczelrdcnenliyscnyttne edemys a r t i tvaieoneptacvheraf lroey arectagn tah s h ehpfgcDmal teeareest2Nri ril ,m0grdg lriAe neeR-sefadbv)vc r ,Nr eoaaoe sibsrsdugemlAyyyoe---- MTmvehirecetrte oaibmbrgaepetsoe nirsnto ai sman lctlihe cer senoa vfa dimrneodden t tmathghereonenut osHg mhauonicmudst iatinhnn e e u xMtnpedilxcoetrr.r iosAntbag nn isodeywmimn gecb hitoahnpett ser roo,l feT ihnoef-
Microbe-Human Ecosystem, explores the human microbiome.
gRNA Cas9 nuclease
DNA:RNA hybrid L876a CbHAPoTERr 39a | Htuomarn yDis eSaseas Cfauesedt byy Bacteria
Genomic DNA Reflecting recommendations from the Centers for Disease Con-
Cas9 nuclease cuts we know about the progression and
both DNA strands. trol and Prevention, along with the Amerpirocgnaosnis ofS syophciliis eist ybas edf oon r
shameful studies performed in the
Serologically positive United States during the twentieth
Tbhy en odno-uhbolme-ostloragnodu sb reenadk jios inreinpga.ired Donor DNA Thohem odloougboleus-s rteracnodm bbrienaakti oisn rwepitahi rdeodn boyr DNA. % MtdCases reactiveou1r0987ir0000ycir nobgbe tishocPthria elmnoapc rryetgreaySearec,uccp ottncihoidnahciryneasgpRseere ut ccporoteutniododrfenassr ycl3ehnat7eb lElsaotpaptaerglnyrer taoitnvosirdtyreLlsatau tatesgeenect xsteoprrecsTsceti arisgtipeaferyirsco. vgiudcctttaiohilegeoirdeninny nt vi, u Dc ae((rasaVnRyilnane DPag(drhDRen Rcifaio)isLm,lstse eeo)dimm sari a ysesounee,aff cnda s 3poossu 9oyherara. ypesr1rps shen)t aiisi.hdcylele adi.as pa l rp Tliciareswbhos xttm odahlioa umriobtncoeoirt usninrrgotaaeasnh---,s -
60 of antibody made in response to host
cells damaged by T. pallidum. These
Figure 17.13 Genome Editing with Cas9 Nuclease. 0 10–90 3–8 weeks 4–12 weeks acareu scea tlhleedy dnoo nntroet pdoenteemct aslp itreosctsh ebtee--
MICRO INQUIRY How could you assemble the donor DNA molecule for homologous recombination? Exposuredays 66 mweoenkthss– 2af tyeera rs 1a0ft eyre ars sbpye cpiafiicr ianngt iba ondoy.n Dtreiapgonnoesmisa ils masasdaye
interval primary primary (i.e., VDRL or RPR) with a trepone-
between primary and chancre chancre mal specific test, such as the T. pal-
secondary stages lidum particle agglutination (TPPA)
Special Interest Essays Figure 39.18 The Course of Untreated Syphilis. test, which detects antibodies made
Organized into four themes—Microbial Diversity & Ecology,
DISEASE
Techniques & Applications, Historical Highlights, and Disease—
these focused and interesting essays provide additional insight 39.1 Syphilis and the Tuskegee Study
into relevant topics. A research investigation named “Tuskegee Study of Un- The “Tuskegee Study,” as it was known, started with a
27.2 Mutualism and Cooperation Are Two-Way Interactions 621 treated Syphilis in the Negro Male” would be unthinkable racist objective—to develop syphilis treatment for black peo-
today. But it was the reality in 1932 Macon County, Alabama, ple. The enrollees were provided free medical checkups,
when the federal Public Health Service began the study on meals, and burial assistance, but were not told the study had
600 black men (399 with syphilis, 201 without the disease). anything to do with syphilis. Rather, they were informed that
MICROBIAL DIVERSITY & ECOLOGY The tale of this study and its participants is a stain on the his- they were being treated for “bad blood” (box figure). Except
tory of U.S. public health, for which President Bill Clinton they weren’t being treated. Even after penicillin was shown to
27.1 Wolbachia pipientis: The World’s Most Infectious Microbe? formally apologized in 1997. be a highly effective cure for syphilis in 1947, treatment was
Mpipoisetn ptieso, pbluet hthavise rnicekveetrt shiae ainrdfe cotfs tmheo rbea ocrtegrainuimsm Ws othlbaanc ahniya belonHgo twheev geer,n uthse A medoessq (ubitoox vfeigcutorres) ,o afr eth neoset nvaitruursaels h, owshtsi ctho wonit fhohre 4ld0. yTehaer sp. roject was supposed to last 6 months; it went
other microbe. It is known to infect a broad range of crusta- Wolbachia. Nonetheless, infection can be established by Finally in July 1972, a newspaper story broke the news
ceans, spiders, mosquitoes, millipedes, and nematodes, and transinfection, the process of transferring the microbes from that men were unknowingly enrolled in this highly unethical
mwsimhayapt liydn,of eetchst i Wsm .eo npredip otihseyanmnti sb2 i oomwnitel l iiiosts n a ei xnmtsreaacsottr esdrpi neaact iremys aswuncoipcrluedslwas?tii dnQeg.u Tiittoes asdcnersoe,t ehanenerd di nsfesoevrc ettrh asolp sseetc atibhelsae.t Mihnasavenecy tt hWsetor amlbinoassc thh siaaevv eies rboele aeetnfef see chsttasav boel nibs heAeeedn­ sptaundtys haandd bae egno vmerisnlmede nant dp aanpeplr ocpornifaitrem meedd tihcaatl tsrteuadtym penart thicaid-
hosts’ reproductive biology. with Wolbachia from Drosophila. Cytoplasmic interference been withheld. At this time, it was also revealed that the men
W. pipientis inhabits the cytoplasm of its host’s cells and is extensive in these strains of Wolbachia. To establish a were never given the opportunity to quit the study. Three
ietshg egt rsfa eonrfts ifilenizrfraeetcidtoe nfdr aofnemmd avolnieaesb .i glTietony es ourafr tviinoivfnee ,ct toWe doth lebega gncseh xwiat h mtihleuro sdtu egecnhrs eutahrsee- tisantansbet lcteth ,pa ota psth ueilta itnmiofunec sott ifo cAnoe mndoepste itomeu tpwsoidisteeh taha edn rdlaa bmionartteaigtco rbrayut,er idti einnst oiom nwp toihlrde- montThhse l aftoelrl,o twhein pga nyeela rs,h ua t cdlaoswsn-a tchtieo snt uldawy.suit was filed on
ing the likelihood that uninfected eggs survive. The mecha- populations. behalf of the study participants. In 1974, a $10 million settle-
nwcoiasmsmpp sab tyaib nwidl hitmiyc,oh ws tqhhuiicisth oi esm sa,e caWcno.s m pthpipalitis eehnmetidbs rdyceaopunesince dsd sec voyentol otphplaems hemnoitsc tw . iiInln-l mZioksaqI anun itado epdsre onamgreui seei fnvfgeir cudtisevevese .la oEtp xbmploeecrnkitmi, neWgn ottshlb eian tc rAhaniuass-mtirnaifsleisacio thenad vo efA becdooneths- mpreonvti dwe alsif eretiamche emd.e dTihcae l Ub.eSn.e fgiotsv earnndm beunrti aall ssoe rpvriocems isfoerd atlol
be abnormal if only the male is infected. For instance, when firmed the ability of these insects to persist and spread in lo- enrollees. In 1975 the wives and children of the men partici-
infected sperm of the wasp Nasonia vitripennis fertilizes un- cal mosquito populations, validating this approach to insect pants were added. The last study participant died in 2004 and
isnpfeercmte pdr eemggastu, rcehlyr otmryo tsoo maleigsn fwroimth tthhee eWgg. ’sp ciphireonmtioss–olamdeesn. vector control. Source: CDC there are currently 12 offspring receiving benefits.
These eggs then divide as if never fertilized. However, chro-
mosomes behave normally when an infected female mates
with an uninfected male. This yields a normal sex distribu-
tion, and all progeny are infected with the rickettsia.
In other infected insects, W. pipientis may simply kill all the
male offspring and induce parthenogenesis in infected females;
that is, the females simply clone themselves. This limits genetic
diversity but allows 100% transmission of rickettsias to the next
generation. In still other hosts, the microbe modifies male hor-
mones so that the males become feminized and produce eggs.
Another effect of Wolbachia infection is interference with
viral replication. Viruses normally spread by insects may not
be transmissible if the insect is infected with Wolbachia. This
has led to the notion that this infection could be used as a
means of biological control. Aedes mosquitoes transmit numer-
ous viruses, like Zika, dengue, chikungunya, and West Nile. Female Aedes aegypti mosquito. Infection with Wolbachia may render
In humans, these viral infections have no cure and no treatment these insects incapable of spreading arboviruses to humans.
beyond supportive care, so insect control is the best prevention. Source: CDC/James Gathany
For most microbial ecologists, the nonobligatory aspect between Mutualism defines the relationship in which some reciprocal
host and symbiont differentiates cooperation from mutualism benefit accrues to both partners. This is an obligatory relationship
n(Lataetilyn, mitu tuisu so, fbteonr rodwifefdic ourlt retoci pdroisctainl)g u(fiisghu reo b2li7g.1a)t.o Uryn fforrotum- Iinn wmhaincyh tchaes ems,u tthuea liinsdt iavniddu tahle ohrogsatn airsem dse wpeinlld nenott osnu revaicvhe owthheern. ix
nonobligatory because that which is obligatory in one habitat separated. Several examples of mutualism are presented next.
mmoayst nuoste fbuel dinis tainnocttihoenr b(ee.tgw.,e ethne cloaobpoerraatotiroyn) . aNndo nmetuhteulaelsiss,m t hies Microorganism-Insect Mutualisms
the observation that cooperating organisms can grow indepen- Mutualistic associations are common between microbes and in-
dently, although they may not function as well. sects partly because insects often consume plant sap or animal
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