PELCZAR Microbiology

Microbiology 900rc & scBvrcEsrNlEhllarlolla! lbr.hiB Mansion,11. Patant ttlttn Microbiology Jn' rcf,ADLJ. PEI/oZA& Un

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Microbiology

900rc & scBvrcEsrNlEhllarlolla! lbr.hiB Mansion,11. Patant ttlttn

Microbiology Jn' rcf,ADLJ. PEI/oZA& Univ€Eity ol M€ryldd

E. C. S. Cf,AII Alsoclqt€ Prol€$or of Micrcbiologv

I{OEI.R, NIIG [email protected] Distins shedPrcIes$rol Mimbioto3v vi4iuia ?olttdhrtc Induut. md sratoUnivdiiv rEi

llltltl tqaz/ll

rIFTH EDIIION

tMl Taia Mccrar4' Ilill Publishing ConlPanv l,inrirccl \[.\v Dlll,til Ne Debi NdYort rnbon [email protected] HanbuB Petu ss lum $o Plulo

Aucktmd sosora cuaremJ' Pbnam Mitan Monuet Madrid Mqi6 svdn€v Totvo rdnro slns.po6

Tat llccraw-Htll Mictobiolog,

Fift Edition

Copyrighl g 1986 197?. 1972, l%5, l95E by Mccraw-Eitl. Inc. AU ights rcs.rvcd. No pan ofrhis publication rnay be reproduc.il or disfibuted in ary fom or by dy m..n!, d srded h a data bai. d rehieval syslcm, withot |[. prior win.r pemission ofih. publishcr. T.ta Mccr.w-Hill

Edition 1993

35ri rcpriot 2007 DZZQCDDYDRDCC Repdnr€d in India by almgddl

wiih Th. M.CEw-Hi[.

Inc., N.w ydt

Salesteritories: India. Pakistan.Nepal.Bangladesh,Sri Lanka and Bhutan Libnry of Cdgress CataloSing-in-Publi6lio! Dar. Pelcar, Mich..l Jos.ph, dae Microbiolory Includes bibliognphies ed indq.s. I- Cha, Eddi. Chin Sm, datc l. Midcbiolory. [. Kricg, [email protected] R. IU. Tille 84-23932 QR 4t.2 P4 l9E6 576 ISBN 0-07-049234-4 ISBN-l31978-M7-46232G6 ISBN-l0: 0{7462320-6 Publish€d by Tai! Mccraw-Hill Publishing Company Linitc4 7 wen Pdcl Nagu, N.w D€lhi 110008, ad pinted at SDR Prilt€rs,west JyothiNagd, t clhi 110094

Contents PFlace P.\RT ON! Chaptd 1 Cbapter 2 chapter 3 chapter 4

PARTTWL} Chapier s Cbapr€. 6 Cbapt€.7 Chapt !I

IN-IRODLICTIONTO MICROEIOLOGY Tho scope of Micrcbiotog! Th6 Hbtort of Micrcliolos, Th6 Chdacterizatiol ClssitetioD, @d [email protected] Exdietiotr oaMicrcor8eistu Th6 MiMpi.

3 1d of MicrcolSanism! 50

NIICROORGANISMSB-{CTERIA The Morpholosy.nd Firo StsduE of Bacteria The CultivatroD of Bacleria Rsprcduction md Crowth [email protected] Cultues ud cultaal chdacteristie

115

r33

PARTTHREE MICiOBIT\L PHISIOLOGY J\^"DGE^'ET]CS chaptd I cb.pter 10 Chapter 11 chiorer 12

Enzlte and Th6ir Regulation Miqobial M6tabolisE: Bturgy Prcducuon Mic.obtdl MotabolieE: Utiliation of Enersy dd Btostnthesis Bacl€rbl csetics

151 777 196 227

PART IIOTIR AIiT I{ORLD OF BACTER]A Chapi€r 13 qhapts. 14 Chapte. 1s Chapte. 16

The Wo.ld ol Betdia Th6 Wodd ol Bdtdia The Wo d ol Bet*ia 'the Wodd of Bactata Compl6x Morpholo8/

I. II: lll: lvi

O.dinary" [email protected]€aativ€ Bacteria "Ordinary Gram"?ositive &ctoria Bacieria with Un$ual Prcpertios Bact€.ia ol Glatrl-Positiae, lileento$

PART FIVE

ALGAE, PROTOZOA.AND VIRIJSES MICROORGA^*ISMS_FUNGI.

Chaptd 17

rutr8i-MoldE

ud Y6ast3

261 245 300

333 365

chapter 19 ch.pter 20 ch.pter 21

!rcrozoa Virus€s ol Bactelia virus€s of Alimds md Pldts

PART SIT

CONTROLOF MICROORGANISIIS

chaltd 22 - chaptd 23 Chapter 24

Contrcl oI Micloorgeisms by Physi.al Aee s Conbol ol Midoo.guisns hy chsnical A8€nls Anlibiotica ud Other CLemothe.ap€utic Ag€nls

TARI'SI]VEN

I:NVIRONMX\'TAI,AND INDUSTRIAL il|ICROBIOLOGY

chdpid 2s Chaprer 26 Chaprer 27 Chsprer 23 Chapier 2s

PARI EIGHT

Mioobiology oi soil Aquatic Misobiology Microbiology of Doh6stic Watd ed Wasresater McrobioloSy of Foods Indusbial Mioobiology

415 435

510

543 569 593 614 643

MICROORGANIS!'SAND DISEASE

Chapto. 30 Mtcrcbial Flo.a ol ib€ Healthy Hunan ttost Chaltq 31 Host-Mtcrcbe Interactions: The Processot ldeclioD Chaptd 32 Nstural Resisldce and Nonsp€cinc D6feF6 Mecheb4s Chapt6r 33 Bsic dd rhedeti.al As!€cts of tho ImmuD6 R$pors€ Chapt6r 34 Alsalr od Appliotions of the Inmue Respoue Chapt6r 35 Epidomiology of Infectious Diseases Chapt6r 36 Micrcbi6l Agenls oI Disease:Bacreria Chapter 37 lvticmbtal Agenls ol Disease:Vinses Chapter 3S Mi.Nbial Agents oi Dlsease:Pungi dd Prctozoa Glo$4/ lnd€x€s Nmc Ind* OrSuisn IDdex Subject IDd€x

824 450

Preface

''M€3sieuB,c'€st l$ rnimb8s qui auronl le demier mot.' PasIeu.

Fo. rhe major pdt oI the twenlieth ceDtury the physical 3ciences have dominated science and engineering. This situatio! was due to a ldSe degree to the develoDment ol rhe atomic bohb, dnd the achievemenls of the Soviet Union in outer [email protected] Th! succostul launchi.g of the first satellite into slace (Sputntkl in 1ss7 by the Soviet Union accelqaled physicdl science resedh dd ddelopnent progms in lhe Unil6d States by the eovmment, by univerBities, sd by tndustry We became engagedin a race ror loade.ship in scien.s dd tochnology. We de now experieneing d spid shift ol natioMl p.iodties ir reseNh and development- As we spproach th€ iw.nty-fiFt century, we see biology emerSint as one of the top priortties in the n6ld of science,ard amonS the bioloSical sciences micmbiolo8y has sain€d n€w 3rature. Miooorganisms dd their activ ities d.e increa6inSly central to meny of the concerns of society both nattonall, and inlematioMlly. Th€ prcblehs ol the Blobal environnent, th€ recognilion ot the need to recycle natural rcsources, the di&overy of recombinMt DNA md lhe resultin8 hi8h technology ot Benetic en8ircering-ih€e€ and oth€. d€velopmentr hsve plscetl microbiolo8yin the linolighl. MicrcbioloSy is emerginB as tbe k6y biological science. Micrcorganisms !ro' vide th6 models used in nol€cular biolosy fol research. This resedch at the moleculdlevel has prcvided, and continuesto provide, the arswers to numei' ous tundamental queslioN in genelics,helabolism, ad.eU foms ed functions, Midoorganisns also provlde oodel systems lor studyinS the relationships betweenlpecies in mixed population6. ]h€r€ is Sroqng recosnition of rlo potential or [email protected] in msy applted @as. The abtlitv of bic.oorganlsms to decompose materials slch ae herbicides,pesticidqs,dd oils in oil spillsirhe porenrialof midoorgaDismsas iood supplemerrst &s exllottatio! of$idobid activily tD produce energy such as methane 8as lor rusl consumption; dd the polential ol new rherapeulic suhstsnces lroduced by [email protected]'smFlhese and other us* of hidoorSanisms ue becomin8 inra6ingly anractive, Reombinant DNA lechnoloSy, comonly BterEd to as s€netic ensineerinS, b one of the princrial thrusls 6f the emerging hi8h t€chnologies in the biolosical sci€Dces.Recombinad DNA teclhology mak€s ir teasible to consider geneticallj maripulared tensineeredl micrcorg.ris$ lor commerci.l produdion ot new ald valutble prcducls lor a vanely of lDrposes,e-9.,medicirals, tuel. and food, Thi3 ifth edition oI MICROBIOLOGY Ftains narv of the featues tial havd proved succ€sofll in the ffBt fou. edilions, parlicululy the bslarce betwen tunddmeriol or ba6ic nicrcbioloSy and oppljed Fic.obioloel. This approacb basi. empbasizes tb6 iDpo.tmce ol integratirg lew kno$ led8e SaiDedlteud reseaE-hwith applied.eseuih 6rd delelopment plo8raru, A dtrcng @DtinuuD

ol rce€arch dd devolopment, f.om tbe basic to the applted, facUiare! rhe dd€lopment ol ben6fits lo! societyOne oI th€ new features of this edition is a D.esentatioD oI the clssification of bacteria in a totally new fomat following the scheme idtuduced i! th€ ff.st volume of the HeDtly published Bersey's Monuol of Sysr€mouc Bdcredolos/. (one or u, No€l R. Krte8, swed as editor ol the ffrst volume.) we have also dpud.d sd rcvis€d the matedal on metabolism, bacte.ial Sonelicsi dd aenetic [email protected] dd EorEdized the sectioD o! Ficrcolg.nism ed di6€ase, Cdelul attedtior has been given to updating of inlolmation i! all [email protected] of the dircipline. Moy !€w summaly rables have ben developed, and new illusbatiols selected. New Eview questiom, dd updated .ef€rcEce., iollow each cha $. Tho subjoct mterial is p.ec€nted iD eiShi pdts, As a new fedture, each pa now opens with m €ssay prcviding added iDsi8ht into the nat€rial that follows. Each chapte. bsSi$ with a chaptd ourline md m inEoduction. Many chaprers now conlain b*d $says hidlishtins inportant dtscoveries dd developnents in Diclobioiory, As in the pdt, the older of dugement of chsples lods its€U to adjatmeDt! ir any sequedce desiEd by the i$truclor A coneiderablo dhoml ot th€ dtwork has been drawn by Di EMin n Le$el (a midobiologitt in ht owu lidtl. W6 have found this to be a distinct assot in terhs of inprcvil8 tbe ped6gogic6l value of iUurbatioN. Thpe valuable supplementdy publications ne available to acoompdy rhis @w edition: m INSTRUCTOR'S MANUAL, a ST!iDEN] S GUIDE, dd L.\BOMTORY EXPERIMENTS IN MICROBIOLOGY Each hs been .evieod to codom with ths subimt natt6r in the SIth edition of MICROBIOTOGY W6 bav€ pr vided extelsiv€ cross-referencing dong these fou. publicdtiotrs, The INSTRUG TOR'S MANUAL includ6 sussssr6d lecturc ald laboratory shedules, chapto. suhmdies, sow€s of audiovisual aids, sounes oI labolatory equipment dd MgeDtr, s w€ll a sdple test qu€stioDs. The STUDEM S GIIDE \as ben d6v6lop6d to ssbt tho studeut ir his or her efforts to comprehend the subjoct halter It provid$ io! e6ch chapter a concise stdtemont ol th€ coDr6Dr (e overvi*), a conprehdsive topicd outltne, dd s le.ies ol sell-study questions of sevdal types, The nitirs oi a tqtbook on a subject d comprehen6ive as nicrcbiolosr requircs couide.able a$ista!.e from a luse nuber ol prclessioial coueagues. ADo.8 th$a, we wbh to ackiowledse th6 followins peFonr who were sen€.ous in th6ir a*i3ldce, particuldly in commenting upon dr.tu ol vaiou chspt6F: Phillip M. Ach6t [email protected] of Flo$da; Romld L. Crawfod, University oi Midqota; toEtta C. Ellias, Fldida State Univ€rsityt Loujs R, Fin6. Kflmas 6tat€ Univ6Fity;[email protected] R. Jew6U,Univ.Fity of Wisconsin-EauClatei Ted R. Iohfton, st. Ol€J Colleser Robed J. laBsen, U versity of Arizona; David Kdkewilz, Rutgds UnivlBity; toseph S, Layne, Mcmphis Siate Untvelsiry; Haideh LiShdoot. [email protected]€m WasbjnstoD Univ€siiy; David Plmdi Rutges UniwB yi &mond J. Seidler O.egon State Universityi Robed Todd, South Dakota Staro U v*ityi AD6 tl. Williu$, Evergreen valley Colles.; ud Fred D. Willios, Iowa Stat€ UDiveFitv, Sp6cial thdk d€ due [email protected] G. Baiaes, Mcciil Unive$ityi john O. Corlise, Univdsilv of M€ryland: A. C, Dombush, Medicsl ResstrLhDivisioD.Amence

C B3t*' CvaMnid Co; lesm6 J Motta, Univ6siiv oI Mdryland; and Robe't r'i"t,"" and stato univoritv, who prcvid€d €xta herp ii*ii[-i'r".!Jr'".

-'S" ,il'*ii,lilll'j;.

rathreD conpsnv atrheMc*aw-lrirBook

w""r".sues sF;lev Editing supeNisori aDd ch'rlos Hss Prc.'".it"-. rlit r i** pr"*mr c-pemiion a'd assist*ce rn the rssk or i".'i.'i s""i-'l.i,i* 'r..; nr""is o aue or"o to Kdrcn Iscquesand il#i"iil;il'';'r'iJuo.t^ ssisrmce in rhe prepdatioo oI nd8stpt t i tt'"i' rc'irir ia,i" ";;;;jii;;t 'uir,l ihi; i"" each ch6pter hd [email protected] the prtm6ry r*poosibiritv of us hs re6d md oitiquod stl the chapteB As .t ."" ir'r'"t.i*."", ."""*tt"v. had lhe b€n6fi1olreviews of eachchapler rNo "*"i""J.-,r"*a. *6octate5 l! the end we tak€ collecrive FlPonsi.'.'r..r*"*t i"i","i bility tor"i the complete content of this ten' Michael t. Pelczar, Jr E. C. s. Ch€n Nool R. Kri6g

PARITWO MICROORGANISMSBACTERIA

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N rnt v.,rh lhar bncrerirl ilag . j ,r r roxljts Dhjo.rin-! non . -,r.r,r p\oNfesponsibieiorrioabilit! :,,:: rerl! to syin. Bderi.lacking ttas€tl& and ' r !f\. orl lho abilji! 1(, Drake fl.lella, aro nNrants rllat mat. oDll ,,rnihle lo !qim. inrlicaijtrg that -:Lr a ,rl llJr n)rN. is requira,l ior swilrdDS. plior : r,.\f'\'r : i( \ras n.l clear jir,'r ho!, fiaqelr , i. r.j t)r Lherrr \rtrs thiril tho flas .tl. rrrlddrip ,,. , , li-ir x:1. lnl: .ould causd rh{i flaeelh t! screN rnr frh lLLonrpdn m nNl as a n'lalnrs ad[sirq\' .ar i{ i, ifairii tiec. nf itrtr\. Thp nlsrrrerf oiiisk.s|an€d !i,r.trr,s tr,c li!.oJ b.nl.rldl udsclh !r!'jp(1..t tIrl lli.,"p nlelJ irdqll Lre a '[.8dlar thjt Rnn{] 'n.rtor' .i ,i,.$r{ili to sFi.. I nlorrumrelr. inrlirnluat ta(xe:in1 l-,{.llr \f.trp so lhin 1[.t l1]er.orld n.r bl ob*ih,r r, ,,ri.nli{:i1L! in a.lion on i ln iuJ: l)acldirm i i r!;3. NIi{jhasLSilvernr0n ard :vtel!n! Sinon.t the ( nir..sjil .t Caliiorlja. sdn Dicso, perlomod o\,){in,,irls rhat indi.irFd unequivo.rllr th.r lra.leriat lli qfrh di' r(t dl\'rot4b. T|ev rcalized rlar a bad. al il,L.' !nr ni!hl h. nnrloqoni lo iho shall ot ar .!cdri{: rrrF :,ri,,f il r rr N housnre is h.trDi iir a InLlt, lher i il.ril futr l1,.I n nrq,er. ilrh'r rn.t.ris n r bDli.l '. s lrJsNri [\, lbp shafl anrt hp]rl ,i,. ,I r rr r, I (..rh|lr rriji /}) shl ionorL o"d ln. , :r lr, Fi'! nijl J.r|k Ultrs liris .natog!, S IeF !., ., sjnnn 1i,:iiqt( expelio€lts whd.by rhey irg r.r ri flrspUlm troE htaltnq. The

c.nsftt'Fn.e ot this vroll.l b0 tllrl lhe |octeri.Jl.el waurd rcl.ki insle(d,dnd lhis srle sorn_arhins r!,t couklbe easilyseanwtlh ao ordiniN nricroscole. Silvoma. dd SiDron.hos I nlranl bs.rorium rhal hen a sinsl. slraishl flag0ll\ ! on one sjdo of the .ell TIis nrrtantcoultl nol sNjn be.artserhe{asel!trD \tas Dot hellc.!: oeleihel€ls. il th," darional hlpothesis $'oE,id'cct, !}€ flagellhr slould slillbc lble 10spin. 'l ir.! Frepa&d anliborliesaeojnsl rhe flasellun and lhen adle.l n nirhrre oi tho b..reriaald rh.aDlibodies lt' r nile.Theslidebeoinr ecoar e{i w i theanr r tr i 'J odi es , an(l lhe n88cllaadhercdto Ilie artibodi.s Thus. each _tethered .ell l,ocd e bv its ildgallunrro dre stide.h lhis.:ondilbn rhe narcllaot thesecellscorld nor rorarn: ho\r.r'er each tetlreredcell beratrro sDi'r ljke a pinlr nas difh, !ll 10explain ho( such behsli.r could oc.ur unless. nr a kee sfjnnn'ng cell, rhe tlagetla srrn-prob0ur by meansof a rolary Noror at their b3so.,\rl{litionalorpcrjhenls on !nteLheredcells also srpoorted lhe ideaof fl.Collnr.otarion.Silvcman and siN,n loud th0l sn'all larexb€adscoutd be atrached ln a la.rert.l llaselluD bv hcans oI antibodi66rs!ch btuds, s.hi.h (.re Gsjlv visible \rith. njc.oscope, (1Tc olrsencd lo rotateraDjdlvaborr 6n iD\isibte axis 1lh.sr r ijqhl ll.sclluol! lL ii no\' -r.trarlh a(a|le{l thit brderiat flasola do !)l0li ns rn s bt!.jn!-a Iype oI nolion lh.r nav be llique amoDgliving org.nisb$.

Pltedilg D6re. C6ll eall! of MtrobdcLeriu tubdutdt [email protected] ft€E cqlts *posod ro oxlr€Eslv hlrl Ns 3uos udd lp€cidl coidltloE. Not rb6 ship. h,-hr".d b!, rh€ "holld" 6I [email protected]t! wbrcn t biidtiv6 th6 rtSid lbucruo o{ tb6 6I sdl. x 41,500 (couhry qt B, nibi.J

of

(lryter 5

ITUIIIXE

Ihe Morphology and Fine Str.:reture of Bacteria

Th Sik, Shape, and An !8!b.nr Sizo . Sbape and Anatrsenent

ofBet

rial C€llt

B&terial SrncrN. Stludl$ Enenal ro fte Cell Wdl Flaeells a Molility . Pili llinbriae)

. Cdpstrlsi . Eh€aths Prostheae md Sldlks '

The CeU waU Sttucture and Chenical Conposition Sbu.tures Intemal to the Cell Wall The Ctaoplasnic Menbrane . Protoplosts, Sphercpldt3 t M€mbsro{s Inrrusions &d htracelltrlu M€mbrs€ Syst6ns . The Cyloplasn . Cytoplasmic Inclnsions dd Vadol$ . Nuclea.Material SDoM .nd Cysts

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ulnr \fri.rtidrs,,1 lh,,sr lLrr. l,rsjL . . . ,. c.niir sDdrics.l l)a'ihrir lIo (rlls dn ,r, q"l ot $hrch ari pairs. chsrFrs., hJir\. tri.h.ir.s. r r,i r..a!n!z.lh.sc I'n1l.n: ol sl)iDr rrrl .lNr rr,rrr"r. a.tenslic ol a laxononri. group, e.!., n lefus ;oJnf r,,r r.r,rs.Do\5o!(rrl p"nddp,'. sh:,h a. n.dp .,rrblp l. .,. . n,.; ',' plecr-o m: ro,cnn\..\jl. r '.r"sc,r,,rrhi oer, morpholosical.hdacleristics of bAct€rinj cFlts The bddcri.l r:erl posy'ss{is | (loraiLql I i. liq,r.,, ,, lhis inl.flral srf(ctur! \'as nrad. pojsjl,l0 lj\ if,t,,,tq ,j ,,,1rj,f.rri n.n cr.s..!.r.rhniquesrn.l,rfinsrrunrcnrsin.st(Ln!.tt,!1,.i ,,.rl,j r,.r,.tr thin sc.tjon. 'fhe lornrs mi.robi6l .vl',logl r!1t lja( lrrral r:nrlDor t,jr. tr i r* commonDl ace itr mi{robiolojri.al lilorarirl 'rh. r.riors s[u.trrs .l ! ba{.t.rirt {:.i] rliit, : ,, .r I ri!.rtri nnr or \ r I lh'iir phlsic.l l.afur.s blrl also in th.ir(hrN,t1J r..i.r,,.rr(L i,, 1r)rL, ], lunctions. T[rs Ltioloeisls t(xli\ se'][ l,i rr,)url j,r ., t]dr(rL. in(t -. ,,,

functional p.opedies of the hcte.ial cell. This area or r€sea.ch studied by biolosists is sometimes relened to €s biochenicat rvrolos!.

Tfif, SIZE. SHAPf,JATI) A-RRANG[MENI OF BACTIRIAI,CEI,LS Siz?

Shapeand

hctgria ale very small, most being app.oximately 0.5 to 1.0 pm in djamelei An impoltant consequence of the small siz of mic.oo.ganisms is rhal the snrn.,c arpn\t,lomr raln' of bacteda is exceedinsly high cohpared io the same mtio for iaryer o|8anisms ofsimild shape lTable 5-1J.A relstively ldee surface throush which nutdenls €an ent6r (or waste pDducts leavel compared io a smsll voiume of cell subsiance to be nourirhed accounis lor the unusually hieh rate of gowth and metabolism of bacte.is. Moreover, becaose of rhe hish sudace a.ea voiume ratio, the mass ol cell substance to be nourish€d is very clo6e to the surface: theEfore. no circulatorv m6chanism is needed ro disribute rhe nulrients thal are taken in, and there b thousht to be liitle o! no cytoplasmic movement withtu a bactorial cell. D*pite lhese advantaSes,a hish su.face arca/ volume .6tio limits the size of bacteria to microscolic dimensions. The shape of a bacteium is govemed by its risid cell wal| howeve!, exacrly what atlribute oI thk rigid mate.ial dete.mines that a cell will have a pa.ticular shape is ro( yet undeNtood. Typical ba€terial cells are spherical (jo,:ci:sinsular, cocc!s);Et.6ight lod, (b'cilli: singular.bacillus); or rods that are helicallv curved (spirilr.: B:trEular,spiiltum ) as illusuated in FiE. 5-1. AlthouSh most b.cte.jal species have cells thal a.e of a fairly constant and cha.acterisri€ shape, some h8ve cell6 that a.e pleomord,'.. i.e., ihat can exh'bit a variety of shapes (Fis.

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risft 5-lO Drawing of Lhemaior s;ctures which occu! within lhe bactelial csll wall. Certainsloc_ commo; to all bacL€rislc€lb fEF win F. lessel, iliustrotor'l

73 SIRU-CTURIS LXf[RijAL TO THt Ctl,L l-lAl.t Fldgella aDd yolility

Fitw !.11. DEwing3 ol v&tou megielt! of bactsrial Aa86Ua.(A) Mono, ticbo$; a silglo pold !6gelluD. (Bl Lophotricho{r; 6 clust€! ol lold O.g6Ua, (C) Anphibichos; It s6na, eithe! 3ingl. o! clustss, at both c6U poles. (D) Perudchouer s'mund€d by lardsl nas6Ua.{EMin F. !a$el, jl,

wcrooRc,{vlsMs_B{cr|ru BaLtedal nd8.tta {sinSulai fla8e[umj sre hatrlit€. helicatappendases thar prcI'u de throu8h the ( eil wa | | and are respongbte tor s wiDmi n8 morility. ThHy d rc mDch lhrnner than rhs flagella or crlia or eucsryotFs.beingb.ol to 0.02 !m in diameler, dd 6re also much simpler ir sljucture. T_heirtocation on the 'hev on the cel varies depending bacte.ial speciesand nay be Dotar (at oqe or both ends of the bacierium) or hbfal lalon8 the edes of rh;L;ieriumt. Some ananSenents of bacterial flagelts ee shown jn pis 5-11. A naqellum is composed of three parrs (Fi8. 5_12):a lusal bodv aslo;iated wjth !f,e cytollasDic rmhbree and cell wall. a short hook, abJ a'h6licat filanetrr whici is nsud,,y several rimes as lons as rhe celt. sore cram_n€8ari;;G"ii,ii" r,a"" , sunoundins rhe flasellu& this shearh is conrinuous with th6 outer hemb.ane " "r,"a of the cran-nesarive cell wall. The chemicel compos ion of rh6 basd ;oqy E unknown, but rhe hook and fftament m composed ot protein subun s {mDron€rsl dran8ed in a ielical fashion. The pdein of ihe filam€nt is known as flac. in. Unlile a hair, a llageilum srows at its rip rarhe. rhan at rhe base. FiaSeliin monomers synthesized wiihin the celt are betieved ro pass alonq rhe h;Iow cenrer ot thF flase um and de added ro ine disrdl end oi the ffldmnr

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Th. Molpf,ol.3y .d I|!. Srtu run or B..t.rt. tarse motile bodies such as boats and fish make use of rhe ineriia ot water to! their propulsion. when pushed a8ainst with, lor exanple, an oar, a propeller blsde, or ins, the water lemporarily acts as a solid, thereby enablinE rhe boal o. fish io Beneratea forward propukiveforce. However,the small sizeofbacle.ja p.ohibils their use of the inertia of water lo Said propulsive force, becausethe drag forces drte to lheliscosilt, olwater become thousands ol rimes Ereaterthan a n r to r.es lhdr can be CpnF,aledhom r ne ia. Ihe d i i fi c ul r v {oul J bp s i m i l dr lo whal we would en.ounier if we atempred to row a boat on a tske fflled wilh lhick molasses.However. bactelia can swim manv lihes their own lelpth Def F tw 5.12, The m€chsnism s" co n d undp, analoSouscondr tion\: :i ailachmentof flas€llato a Bacteria propel themselves by rotalins thei! heiical flasella. The p.inciple :b-negative bacl€lial ceil involved .an be illusbatod by jmasioins the penehatio! ot a prece of cork by a :i€!domonos oe.ucinosdl. corksc.ew It one tries to ram ihe corksoew directly through the cor&, Breat : Prior lo electron,nicroIorce will probably be needed. On the other hand if one merely lotates lhe i.!e ex6m'narion,the cells cor!6cre!v, the cork can be easjly penet.ated. In lhe case of bacie.ia, the co.k is 'de.e pa.tially lysed and analosous lo the viscous medium and the corkscrew to the heljcal fldsellum. 1l ::en negatively stained to is apparent lrom lhis analo8y that a muiant bacte.ium havins slraishr rather the pojnl of flaselld than helical flagella would be unable to swim. The nature oi rhe rotarv motor -ie i::achn€nt (bual bodyj rh d l 5 p i n, edcnLor kslFw+haped fldeellumis.r ill Dor unJ er s r ood bur r he r ns s :tF visible (X80,000apfound in lhe basal body (Fig. 5,12) are probably involved. rt is known ihai rhe i ). (B) lsolated flarella flasellsr moto. is d.iven by the proloninotive force, i.e., the torce de.ived fron :::\fing basal.bodyat one lhe elecl.ical polentiai and the hvdlo8en-ion sladient across lhe cvtoplasmjc i:d. lC) Model of basal memb.ane {see Chap. 10). Moreover, recent studies sussest thal lhe conceltsr:dt iliushating its struction oi CCMP (guanosine 3',s'-cyclic phospho.ic acidl w hin the cell govems :r:e and attachhenr to a the di.ection in which the .otation occu.s. ::d-negalive b6cte.ium. Bacteria having polar flasella swim in a back-andJo.lh tashion: thev rcveree ::e flasells ot c.am posi, ::e bacl€riahave only two thei. djreclion ol swimmin€ by reversing lhe direction of nasetlar roiation. :d dngs. (Coudesy ol T. Bacteria haviDs lateral flagella swim jn a more complicated manner. Their :i:e. University ot Tokyo.) flagellaopemte in synchrony to lorin a bundle thar ertends behind rhe.eil t!.is.

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Fi8w 5.13, Dbgrm ofthe conffgurationand aF rangehenl oi pe.itrjchous flagells du.ine swimming and tumblins. The snalt uows indi.aie the direc, tjon of propa8ationoI heljcal waves along the flaseu6. (AJ Duing swinnirs the flaSellaare in the forn of left-handed helices and Dtate counterciociwis, in synchhD) ,o lo m a bundle. The srse driow indicalo6 lhe directjon of swinnins. (B) Durins tumbliD€ the flagella revese then rctstio!, po.tjons oi the flag€lle acqune a short wavelenSih and ri8hr handed configuration,and the bundle flies apait. The cell carnot swim unde! th€second ions and instead exhibits a chaotic motion, as symbolized by the luse c.ossedd.ows. [Cou.tesyofn. M. MdcNob ond M, K Ohston,.l Mol Biol7t2.1,19?7,)

5-13).Honever. $hen the fla8ellar nolom reverse.conJormalioDal .hanaes occur d i o n S th e lldgelld.' hF bundl" ni"..pdr r . thp ...1 tum b,F s $i tdti F i Idtty . "nd lhe flagellar motom.esume their nomal direcrion. the flascllar bundle aeain l o rF r. a n o,h",pl bFsin" lo,uim bur r oq i1 d d. er ;nr a,.r i on. i t,, sequence oi evenls occurs .epealedlr, so that lhe morility becomes a serjes of periods ol swimming (rurs) punctuatcd bv pe.iods of tumblin8 (twiddles), with a chanse ln direction afle! each tunble.

SwimmingMotjlity Withoutflagella

Ce.tain helical bacteria {spi.ochetes) ex}ibit swimmins motility, parricula.ly in highly viscous media, yet they lack external flagella. However, they have flagellalike structures located within the cell, just benearh the oute. cell envetope (seeFis.13'1). These are calledlefirtasn,i. tas.|a; they have also been re.ned axialnbrils orcndonas€lla. They are resporsiblefo. the motilityof spirccheres, but how lhey a.complish this is not yei clear. Other helical bacreia called sp,r.ptasmas are able to swih in viscous media, yei iack any apparent orSanelles lor notility, even periplasnic flagella. The mechanism for rheirlmotility is completely unknown.

GlidingIiotilily

Somebacteria, e.9.,Cr'tophoso species,are motile only when they are itr contaci wilh a solid surface. As they glide they exhibil a sinuous, flexing norion_ Thjs kind of novenent is com!'aratively slow, only a few pm per second. The mechantsm ol glidirs motility is unlnown; no or8anelles responsible tor notilitv have been observed.

BacterialChemoraxis

Many, perhaps host, motile ba.teda are capable of di.e.ied swimminq toward o r d w a y fr om var ious.hFmjcal compoundH phen om enon c atted bar r er i al chen,oiaxis. Swimingioward a chemical is termedposirive chemotdisi swimmins away is oqarive chemorais. Allhough chemicals may act as attractanls or Epellenls, the stimulus is in fact not the chemi€al sell but rather a chanse i n ,h e , o n cenlidr iohol lhe Lhpm ! al wr $ r ime, i."., a r" door al 6adi enr . Suan srddrpnls drF sFnled by mFans of prcrein. h"morcLeploh which-de toLatpd on lhe cyloplasmic membrane and are speciffc lor va.ious att.actants and.eDellents. By mFdns oi ils [email protected] bar(erium conribuallv co^p*i its i.mediate environmeni with ihe enllronment it had experienced a few momenrs

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ii;i"i-:'ll [:ffi'::Ii]::ii:fi'"11i:i,:l: ff:1il..':i;,x';;"i:l',i:,:ii:"xT l;::'l: 1;*l*l,ijj:::'l':li 1li.li""iiil;iii",i'"',i"i:;:l;l#':il no r hangF:n Sl eoqF c onr m r r dr i oi

cn se -cD p (r hL( nem or e' ppl

;il:T; ; r""llfl r;,;:i]:;: ;'iT;T:fi:i; -:i;l:n$ ::l'""t",,"."11,1;l[: :,""17 iti:"J;'""Jff :$:l'J'l;' ;:,#lilT*:fi il""':::,rillt"':r' :*iT''lH"; ;*; l;jl::l;:;,i",1;:l',;l:l:m:"r:ff i:"'.""'"i; "lil*:";:r;!:,[: :l;r;"'";ii:;]:;l,l:;",.""r"' ;:1,]: i. eu.o_ ;",r, _:::;:.1Jff,f11,liilT iitj"r...,i;" "*

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di changes iumbLes, ::::?T,""X'illi:li;#:lii,i,l"i'"i,111*lihil ";'i.oon dircction is achieved {ln a filaLlv the "right" i.:";;t"r; i#,i",

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lrolhic bacte.ia exhiblt positive fhotol'rus low

. theeanhs mssneli'f i" 'r.lil''i,'pi",'"tp."." ro d c'r"inor n"enerire 'i''1..1"* rurruerrhis-rq'r' hrr'eo "J;;;; 'r;

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Fisre 5-14, Negalivelt r::red cell of AqudsPnif i- ndgneloloc!icum show -. , .article chain t?C) oI : ::ir: electron-densemas r:::! lnclusions (nagneto_ rLithin the ceu The :::i!l :!: :.presents 1 pm {cour D '. -.i D. L. Eolkwill, : : : : 3 o o n d R P B l o ke _ _ : ': j B d c l e . i o l14 1 :1 3 9 9

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Pili (Firnbriae)

piti {srn8ulu, Dituli are hollob, nonhelic€t. filuFntous sppebdaSesrhat ar€ rnrmer, short6r, .bd thao flag€lls tFi8. 5-1s). Thsy do not tuoclion in motililv. since they de foDnd on oonmorile as we as motile spocies. There are, howeve., several funcrions associated wlth difier€nl rypes ;f piij. One type, known as lhe F piius {or s6x pilus), serves aE the po.t of onr.t of Senetcrateiral during bdltFrial matinS (see Chap. l2l. Some pili play s malor rcle in human inJecrjonby sllowrng pathoSetricbacreria to auach to epitheliat cells liDilg the respiratory, intestinal, or genitourinary bacts. This att;chment plev€nts ihe bacteria from beins wsshed away by the flow ot mucous o! body fluids and pefmits the infection to be established,

Caprules

Sooe bacterial cells @ surounded by a viscous substance fomiDg a cov6.ing layer or envelope around the cell wall. r lhis laye. can be visuali;ed hy lishi miooscopy using special staining methods, it is termed a .a,rsute. f the laye. is too thin to.be seen by liBht nicroscopy t is telm€d a m- ;;;;sule. it it is so abunder that ney cells ale enbedded ir a conmor ddii,"iii;';;ierial is

'r11"4.,ri'."..

!v i'8nr mrcroscopy,capsules spped to b€ amorphous Selatinous deos suF roundine a cell (Fig. 5-16AJrhowever, special techniques desitned to preseive delicate stNciures for observattoo by elecbon mic.oscopy have revealed rhat capsuiesconsist of a mesh or network ol ffne srrands (Fis. 5-1681, In baDy ined.es capsuld matF,iatis nor hiShly waler-sotubleand rherefore does nol readily d ffus6 awal fiom rhe (ells that Droduce ir. Io other iNLaDces [email protected] 5.15, Fjnbriated bscteria. (A) Sliseuo tene.ii dividtls bqcilli with nun6} ous frdbnae sunourdin8 lhe ceus (x20,000).(BJ60lmonelld typhii dividing bscilli with nunerous finbrtse md a few flasella (lhe very lorg appendages)(X12,500), lcourtesy ot /. P. Dusuid ond /. F. Wlhnson ond Tle socjety of GeEtul Mi.robi1961,)

Figw 5'16, Bacterialcap' :ules as seenby liehl ni:.scopy (A) 4d electton lrroscopy (B). (A) India' jr preparatioD oi a capsua:edbactenlm isolated :oD a paper-mill operation. lle panicles oI cdbon in ae ink cannotpenetale the rouDd tle cells).Cou.r4y r' P. M. Boric&,Wdllo.e :reeze-etchplep6.aliotroi Cran-posiliverod-shaped lcreria isolaredLon acid rire water,showinSa fi:.illd polyn€. nclworl su! iunding tha celk. The iEeze-IracluF prcc4s hs dso !€vealedvdious interral and surlace st uclures oI ae cells.{court€syotP. n. tugon, C. B. MocMillon, od n. M. Ptiste., /. Bocr€-

Sh€aths

and dissolvesin the mediuh, sometines ih€ material is hishly ,l.aqatically inoeasilg'vate.+oluble the viscosity ol the brolh in Nhich tbe or8anismsare Capsulesc6n seNea number of functions, depetrdinso! lhebacteial species t1) They may plovide prolection asainst temporary drying by bindin8 $aler nol€cules. (2) They nay block altachmentof bacteriophages.(31They may be antiphagocylicii,e,, lhey inhibit lhe enSullmentof pathogenicbacleria by white blood cells and thus conhibute lo invasive o! irleclive ability {virulencej. {4) They may piomote attachnent ofbacteria lo suriacesrfor exam!16, Streplococcus mrtons, a bacteriumassociatedwith produ.ing dental cades,firmly adheres to the shooth su.laces ol leeth becaus€of its sedelion of a wale.-itsoluble capsularglucd. (s) lf capsulesa.e conposed of cdnpdunds having an electri cal chege, such as suga!-umnic acids, they may pronot€ lhe stability of bac_ terial suspension by preventin8 lhe cetls flon agg.eg6tin8and oetllin8 out, becansecells bearins sinilarly charsed surfacestend to repel one another Most bacterialcapsulesa.e composedol polysa.cha.ides.Capsulescomposed are usually ol a sinele kind oi susa. are termed honopotlsaccha.id€s, synlhesized oltside lhe cell hon disacchalidesby exocellular enzymes.The (a mutdns is an oi from suclose by S. synthesis of slucan lolyn€. slucosel exampic.Other capsules8re composedofsevelal kinds oi susarsand are temed hcr.rnDolvsac.haridesrthesea.e us!ally synthesizedJrom sugarprecursorsthal are aciivired lenereizeJ)within the cell, attached to d lipid carjer mol€cule. bansportedados6 the cytoplasnic memblane,and lolymeized outside the cell The capslle ol Xlebsielld pneuhonide is an examlle A few capsulesa.e polypeplides. Ior example, the capsule of the anth.ax orSanism,B. dnlhmcis, is composed entirely of a polyme. ol Slutamjc acid Mo.eover, this peptide is an unusu6l one becauselhe glutamic acid is the Fle D oplical isoner rdther lhan the lsual L isoner conmonlv found in nature Some speciesof ba.lelja, pdticularly those from freshrlater and marine eDvi rcDmenis,form chains o! t chomeslhat are enclosedb,r a hollow tube called a This sbucture is most readilv visualized when sone of the cells have

ri8ure 5.17, Sheatnedbacteria. SheadraDdcells o{ SpnoerotilB noronsstained with njSrosin-Dinensiols oi iDdividual cells are 1 Em sheaths may reach a length oi severalmillinele8. [cou'1$y ot I. l. stoles,I Bocte.ial671279,1951) mjg.aled i.otu lt (FiB. 5,12). Shearhsnay sonerimes b*ome imp!6snate.l with Ienrc or nanganese hyd.oxides, which srenpthen rhem.

Sialkr

THf, CIII WAT,I

t|'i \rh 'r.'d s nculdr .r ,,iar h,,.,l r e "em,r i8id extensl onsof r hp c el l w ai t and cyroprasnrL mFnbrdne dnd hdve a didmerer th"r is alw")s tessrhan rh6t of the cell. They are chdacterislic ofanuobe. ofaercbic bacteda trom heshwate. an.l mdioe mrironmenls- Some bacterial senem such as Coutoboder have a single prosihe.ai others such as Sr€Uoand Ancoloni.rcbJum hsve several (FiB. 5-1-s). Prosrhecaeinoease rh€ sultace 6ea of rhe celts fo! nuiljent abso4lion, which 6 a d va n raseolri1 dilu,e envir oim ents.Sonp pr osr heL" r pba.l er i ; m ay l or n a n e $ L e l l tbud) ar the end of s pr osr hsLdr other shav" "1 adhc s i v es ubs l an.eat rne end ot a prooihp-d thor dids in r!hm"nr " "urta.ps. Althoush the telm srrtk is sometimes used'o inierchangeabty with the ierns p'osltrp.o u hvpno. ir i! perhdp! beller lo rcsllct irs us; to (erlain nonhvinp uDoon]ll