John Kleyn - The Microbiology of Brewing

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    IBI BIJ K A JA

    Dpatmnt of Bolog, Univit of Pugt Soun Taoa, Wangon anBri coo of Maing an rwg part of Bor

    nrt o Birmga Big England

    HISTORCAL... ... . . . 583 ORGANIZATION O THE BREWING INDUSTRy . 584

    LITERATURE OF BREING . . .... . 584TEACHING AND RESEARCH IN THE INDUSTRY. . .. . .. . . . . 584BACTERIA ENCOUNTERED IN BREWERIES . . 584

    I I BI 585CI BI. . . .. . . . . . . 58I I BI .. .. .. . . . . . . . . . . 58U U . . . .. .. . .. . .. 58 .. . ... 8

    WILD yEASTS......................... 8 OUTLINE O TRADITIONAL BREING PROCESSES .. . .. .. . . 588 OUTLINE O ADVANCED BREING PROCESSES . .... 9

    MICROBIOLOGY O BREWING MATERIALS . 59SEECTON AND PROPAGATON O BREERS' EAST .. 59YEAST MANAGEMENT.. . ..... ........... 59GROTH AND METABOLISM..... 595GROTH AND EMENTATON KINETICS 598YEAST CELL WALL .. ....... .. .... 599BREWING YEAST GENETICS 600MICROBIOLOGICAL CONTROL N BREWING, ERMENTATION

    AND PACKAGING INCLUDING SANTATION ..... 600ASEPTIC PACAGING........ 602

    Man b an ann af bu fnaon a bn undo

    fo only o 30 ya Ya wa pviouy gadd a undabl cumand wh a w nghnd cpon w inad emnaion ouiouy om ya h clinging o adycland qpmn o pnn nld wo Abou 836 uga mnaion wa cibd ialavy o ya and h ungal nau f a wa cognd A hi oonaon ya wa ud only n aaa bu i upad apdly n uop and hn h nd a bcaue mg

    Annu.Rev.Microbiol.1971.25:58

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    KEN &

    o o may ma wma. Pau a h mcoiooical ach dvopd a asoa hoy o mao 8 ad i 6pod o pola y aca ( 83 Pau wo wa ddy as who dvlopd mhods o soai ig yas cs ad o

    a cd c opaga a clo suc o commcascal mao (89 Mod dvopms wig mcoiology la o mai-i yas o acia ad o apd mhod o maio

    ORGANIZATION OF T BREWING INDUSTRY

    h dy wll oad o aioal ad aioal chcaaocao paca o a 1 h Amc ocy o whm h opa wy Covio 3. h u o wia iai ad uaia ad h Ma w Aoco omca cp o 3 aoa cos a oad ulaly

    Collaoav ach ad h dvlom o aalycal mhods au ih wo o ad 3 h s im s ogaaos ho osalish uvsally accpd mhods o aalysis Such dvlopms audoudly couagd y h iaioal shaig o chical owl-d y compa uld ova ws ad y amaamaio owy compai Th s o h y ca y h smadwold podco o o cca X 10 (231).

    LITERATURE OF BREWING

    h a may pcialid oual dvod o w cc adchology clud aut aucaft u wScc Toyo Ccats t Mat w' Aciat fAmerica, Journa ot t Ittt ot Beng Poeeings f merianSoiey ot rewing Che Proeeding ot Europen wery Con-t Wart Labat ouctin. Addoal mcooogical iomaio laig o wi appa i sval iaioal joualdvod oly o micoolog

    TEAClNG AN RESEARCH IN T IDUSTY

    Pacay i uop h wi duy i idvdua cous hashpd o oga cs o achg ad ach w scic uualy v (30. ado w Rach s hav up ca cou y aoal wy assocao ad supplm h wo cad ou y h laoaoi dvdual compai

    ACEA ENONERE IN REWERES

    h m o acal a coued i w mallampov a compis Lctbacs (qu i opmaio

    w ad dcccs (mo commo i oommaio wi Th amgaiv ga a viually cod o Abact(Klebsell) eobe eoon Obeuberum ad Zyoon.Tal how h sa o poducio a whch hy occu (3.

    Annu.R

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    F N

    BREWERY SPOGE RGANM A T SAGE F. lODVCTO HIH V (3)

    Sage Baceria encounered

    Mashng and swee wor Thermophic lacc acd bacera rare

    Cooling of wort o ching wih yeas Acec a.cd baceria rareacic acid bacera rarebmbm rare

    Ferenaon

    V Aer-erenaon saes

    bmbmec acd baceraacc acd baceria

    Acec acd baceraacc acid baceramm rare

    LI R

    y c of acobacus ococcus hv c o boc wh w 12, 59 63 2 oe h of hey we v of a bs asoaus whch hofev pce wh og osh c P aosus hoo fv c wh cocche ce 42 23 . Cohy s g he hoev sce h hoshokosehw 9 cos o hooeee he gcoc hw scve ece ose hexo soee e 50 he eoc o eo ce cc c eho gceo cec c co ox Hoof oc cc c oe s ye o o c 9 whch v ow 2 , c so

    e coo c c ck A w vey of co ocesw ve cy oe 24 oe qe o co ox 95

    oh hoo v heofeve e weg o o c ogeos ses vs 62 heeoegowh e p o coee p o hese es hy Gowh of o h H g 565 hewe he se s y gow H ves eow 5 oe sse oc o h h Co ewes oe o c

    cc c owe h H of he wo hohc ss sch sa

    obacllus lbck o 229 C oc exc o hoohc whch hooy co coaoe cec acd, o o 53 2 h o o,

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    KN & O

    t e ee the nee eet the b uponee ne tt t n ( he te m enfet y etng n the tchng yet o n nontele eqment Cetntn e cle f occltng yet (

    ACA

    hee fclttve neoe come tn of ii i n A at KbUa atht gow n w ly t n b nlwhen the H ove 3 (22 232 he ce we nge f t of metolm whch mt vo n ds t h wot tht m eweet and fty o eemle the mell of cooe ge Col fom me ntoce nto the wot fm wte e fo ng wshg; my e sme the ext femetto tchg

    oly n When wot toe fo contno fementton coos so spoilge

    CC C ACA

    hen at h etcho gell n contt to the ol gell of mole tn of Act ( 3 2 The fome genha tonge owe of oxton the lte eng ccely le to oxeethnol fthe thn cet c (19 Atbt tn m metolegcoe va h hxse mnphsph phwy h rxyic accycle t i ome glcoe oxe the thn hoholte ( 22In stns the M thwy s nomly e t te ccle fl to oete e to k f ctvty f t ehogene noly othe enye (2 Thee theefoe a eqement fo cetnmino cid o the coeponn oxoc n contt A tat tncn ynteze teir entie complement of ntroenou compoun from am-

    mnia n e con gment (5 he mle ntin eqement he mke them lmot qto ome ewee n ece the mt feqent e f t vo n tt (Cetn tn e cle of cng yet cell to e (7 cetc

    c ce my ls oce extno oe n tntl qntten ee ( eqentl the cetc cte gow ge elclen orr o ncree expoure o mopherc oxygn. o s vy tcted n the ence of oygen

    he vlt of ece of cetc c cte o mk tht cl-cton t th level my e of lttle vle (2 A contno ectm oftn m ext wth neghong tn eng only n the lty toe n w enyme

    ObeBmbaerum

    Th nonmotle ece wth ho ft ole cell h the nqe ltf own n cometton wh ctve mlyi east cell (2 t fclttve neoe wth H ot f t 6 whch gow w f-

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    M WN 7

    y v bow 4.5 (208). T gyoy py, e pwy nd TA ye e v en oxdon v e yooe wek 221) Soe n e ee pe o nogeno be nd no d fo gow 4). no ney fo

    n o pn n gow d nd g vy of o ofbon w v (214) T gow f ogn dng ewey enon dpnd on n of y ed nd, onvy e ofeenon y y nned by (215) odo of pnp (204) poy dey de (221), nd o by bo of T foowng foe pon n deveop of ogn n e ey ge ofnon: o of y n dng e of b e p of ewo, nd w f dng feenon (35) f e nm-

    of O tu wn e png y g e n pH of eb w b g n no nd w vd n g nbe n y op (5)

    Zymomon

    n e y gy oe w odk e beng 1-5 o- g (20) Te nob nd n gow ov wdep nge (357). o nd fo omee oe peen nb n oe zd bon o (11) ngy po-vded by EnneDodoo pwy, nd e end pod e no

    nd on dode (155) Mno pod ydogen de nddyde gve gy obonbe odo (20). Te ogn gown bee weend w o nv g nd nd y non qpn ()

    WD ASTS

    d ye e o n pn n wo b o obewey e w, by on do no en, nd ofn po, n pod ny wd y n of Sa v-

    a o clbg nd e ovo, fenon of dxn, nddy of b (244) Ty d o dng fo eye on opoog nd pyoog gond b vey en wokw ve d ben nogng (85). Soog od vebn d ony o dnfy n png y (193) Te nod e bd on ngn onp wn gn Scc- 34 nd in of op y n o S pau obnd fo bb T bobd w e y edn e bewey nd n d fo ng pe of png ye nd Te wd ye y be pen n popoon ow few e peon e of y (19) b nm my be oedond wd y y ng n nb fo go w n opd o on d on ooe

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    5 KN &

    w vo non ps n opo o sns sng el o w ye (194).

    Svey o w yss hve ee e u i i w i phg ye 18, ugh ee 98) Ap fm w

    ye o he eu aaromyc, psnvs m he gee u Candda Dbaromyc, Hannpora Hanl Klokr cRooora Toropi hve ee euee. Spee ccromc o gow on n w ysn s so so o ge, hle he o o ye gee uuly o o ye eele h bw o y n w yss ow ume psn o g nbs o y (163)

    OULNE OF RADIAL BRING PROCESSES

    T bewg pee lue (a mg pe heebyby gn ong , by nsng y o nzyseey eg u ue m ml ju oo p o sson by femenn ; (b bewng o wolue ookng sng wo on k bong n wo oong(c non o wo ( gng o n b ( nsng non; n () bee kn

    Due py o sp pn n en hw ewexp o so o gs ons epe he me he. A p sss ng pue y o by mu e

    elee eue eee o gem ugg 4 jmg sps n gn sog o ns pop my eepg f gn o nssy os onn for mn, ng gn o pp n knng o w ee egnon n oon n povs w s he o nng o s v no wo s p yngw s s sp n gnon n b ng w eely pyso on bwn n osun he no oon hee phe he eeme hehe he ml w

    e p o k 46 I o bov ng e eu eguon o p os n e pon puepopy o R boog pobs sss n- oboog o bwng s.

    s v o bwy n gon o n eeen xon nns hn ng sng. os Ame ee,n on o onn s n sy e hhs by yoyz o sp sgs v yo nzys pee m ,n no p n Gn b he u s gnz y bong n ook w s o o gon .

    oowng gnon ook nns ns mh uw onns he bk o gon x w w e pmyobjeve of m s enz oneon o oy o e

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    poen, pd nd ogn pope o pe weoube foTe eun ex ned wo Te ng epeue n pHe dependen on e pu ng poe Te jo ng poee e nfuon, deoon, nd ed ng w obof e w. nfuon g ue o e, wee deon ng ued fo ge bee nd xed ng fo poduon of b nde bee pe A p eee bewee nfuo eoo ng ng on ng e epue edn wou bng, wee n eoon g p f e wdwn, boed nd euned e u e e epeue f e we . Bon no ne fo nuo gne gy ode e ued w e edy p peponze.ueoe, ow popoon of uned ee dun e ue nee e ofe peke ke

    Foown ng e wo eped fo e pen gn nbe ee Wo epo ee o pe w be pe b w een eod o e pun e u ue o ng nd o, wee ge bee pdu, w e u epe vee, ee ue ub o e

    o eo fo wo bog e wo ezon, enze deuo, oguon of unbe d poen ned ub, nd exon obeng ube nd een o fo op Boed wo doe o

    vbe alu nd Clordum poe w, due o e envone,nno popge een (3) n ddo, p en onn ubnex o y pove odped be odng o kebue & e 9, pe of ob n gow n unoppe beeu ew gw e peee f p en. fuey oppg e ny ue w eg 03 b pe bb Uned S o be of bed vue.

    fe bong, pen op e exed by pge of e wo oug ne knwn e op k Te wo nex pupe o oe owo nk foowed by enfugn o eove one fo of poeneoupe ed "o ub w oge n o wo n n, wpooype o wo nk e ofen ued fo eovng o , eeby ovng e need fo enfuge. o onn poen pe e" ub w ogue o n od wo Te e pe e pb-b u eed o e peneu e e ze w ofe o pevu ed w bee n fe y pe nk, op, w ue f wo ng, eeby enbg pepon o od, we o ub pe Te wo nex ood opng epeue b pge oug ooe, wee ue-

    ou need o ep edue e g pe of ye gow oe bewee even ue ed ddon eon upon non of feenoOe ge n e e poedue f dung exe e po

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    59 EYN & O

    n 18 n a oen bewng opean of e abve eeo k bo o n occu n cos ss b ng ae w eiy a e one of feenaon

    e wo ao bee ye e age an e w ae eene wi

    d

    oyet eetey

    Mott ye

    rlbrn pee weea o yea ae rva eieBoo ye oae an ee o e feene boo ne e en eenaion weea ye ein we an f yeay on e w fe As oneene eeo yea ae oee eng uc s f b ws b yeae oee f e feene bo age bee ae e ogo e wo weea ae ee ay one of Bgn e Aean bewee on bo ae an ae e ony ones a bo yea f bo bee ye eeby eiinng e be

    nvove in pogaing an keeping wo ea e epe 188 e poe w boo yea e eignae "b ae veey n nnd "td age poe o non wo bo yea A age ye e o een ae s ay n epe eae enane oyi e ay e ge feenaon e-us C f ae s co o g b 611C An a-n oneene of e ige feenion epeae i a e fenon n o 5 s s o g b 7y Mebe feenaion i a igno e e epaae e w

    yea. arlbrn eboeoie weea r ay negaive 1 n n rva y oe oe ea en arlbrn

    oowing piay eenaon e bee be vo an(geng) an on oee eenng uon e bee ype anabe ie. n o inne e bee s ppey ne n o nn n k u unckg T ow eee s n cbonon an even bioia onanaion oe a beve n anfeng bee o ag-ng nks fwng piay feenon ae: (a bee ian wu u u os ng n b b e ow age epeae ao ep peae a poen-eo i ae opex o o oban oe bn bee a we apeen been ae foion wen e bee e bong Agng s ipve bee o by v ye e nee nges Some bw s gg o en ofaang e bee w bn e og enay eenan.e bewe be e age bee a pay ain an en - b c w cbon ox n nsng nks Pugng w

    C so e ws unsb os ou o bAfe nsng e bee bje a son an oow b

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    MCOBOOY O BW 9

    rfer o og k Afer g, pckge beer re ofen erzed coeo e pezer

    OULIN OF DVNCD RWIN ROSSS

    Dr g o rere copex preco epe o ceveor rro o e edoper o core or d, e e e,preerve e k o c 46 ever rewere, e k redered e re b eer re eg or eepg eepg foowed we g o prodce d erefore ee xer o rered 2 order o redce e reree or -g epe cpc rp reebg cocered wee wor re ed brewere ever core Te rp be derved ro ed cere, b o coo ro ze d be d edegrdo ceved b dr oc eze e ceo ze, cd dro precede ezc degrdo opre bk n su s n c-s; c n nbrewere b vcpcked wder rodced b g op d eectg t rc gd, e t o tt e dee o 3 Aervey, e re y be exrced y orgcsvn eod be ced 9 e coeree e cp er re re oerzed o se (21Sr, e exrc be oerzed bog w de k 7T repro dded er ereo order o ove d

    orpo of e ooe o precped proe e copper d e e ereer (50.

    Ecoed eeo vee re epcg ope oe bece e or s spcn v uc suscepby rborecrob eco, re cooed ore red d ce coeco oeoved crbo doxde 23 cdrococ vee, wc re popr p ncs ns h s sps h hed of fereo o e coe ro wc e rr c e ereoved der g rere e cro doxde prdced ro

    fereo w red dove e beer Lrge oper vee, owed oe brewere or ereo d eer orge e b ded ep or er cpc 2 Te eed wc e n s b hs, e deered o oe dgreee dro of e eer ereer, b cerge re ow cooed o ceve e ero 3

    Cninuus ns cn nc v c s vssscoped ere wc re peror perforce o ge rred ee 1 1 e reced, fro e eergg eer, o h nsncss h v nn u hgh s cncnns e

    e ce dvde o d e of e beer prodced ered Urred coo fereer of ower for per dwee

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    KLEYN &

    n d 9 and ae usd mmea (4 198) . mininuuso cc c sss n us succssu on ! pilsa (83).

    MROOOGY OF REWNG MTERIS

    pim ingdiens n b a wa mal ps and as. SmEupan bees a a mal bees weas ms meian bes i pe en ma and pe en adun usu o o

    Poucon f su bng w usull no jo problemsne an b ead i add emv vaus meal ins nd sas(39) I s an lms unvsal pae aden mneae mswa mas w aiu and magnesum suaes eb wing H f s g H s unob f nu of io a ns n bewng pess e.g. a masng saain will n

    ed sm aguain f pen (ub) a biing is nmpeand esuan bes wi a g p a blgal unsab and abl nfn w lai c ce 6 ur e dmginee f naes n femenan s dmnisd (235 ) Oe s f ingani ns g. in a dsussed in e se n gw and fmnon kncs

    so o onc s qun w su b lvf f daing vegan eg alga and ndusal wass su as nls (1 ) sine s nibu s nd ds dul

    mv ()Pdun f suabl mal bgns n fam w a ue vai fbae nea ban vnn gemnan dung mang (6).n mis gwig sesns bal ma b infed wi sos gow ung spng ocss oucng susncs waus gusng in b (3). w fd baea n e usks duingsping as us and an b eimnaed in pa b fun anging f sep qur od gw ung sepng s dud b sng H o s quo w Unoun, s oss nages gw f fmpe baea (2) ne mbilgipbm elae use f gibb ad sped up gemnai dungmang. e as pdued eeb ve ge sluble nen winuages gw f epii abai duing mang ( 189) nain ases, s mig be desiabe paiua unies su as an we us f ds is pd n bwng u ( 11 ) esus , 22 sz onc o ng he o un famn ads in w f avng ssco fenan. psn eef is dpndn b n qu f aw aeas ndmsng pedu (1 84)

    Hos n on o nsc (9), conu o omsng ppes f w and ae mpn fo vng ssco sad pduin n alp bes (51) un nees in fmnig un

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    OBOO OF BE

    ho wo wh h qn aon o a o ho actay colca hs bl (50)

    SELECTON AN PROPAGATON OF BREWERS' YEAST

    s a o sct sans of a fo bwn fo yss ly incoca s. h th acaon of gn ncls ducn nw ans bas' yas has bn ccssl ) th hvebn w nancs of nc hyzaon fo cocal wn ( 1 ) .aton and tansoaon (178) v ls n suggsd fo odcingwng sns w nw rs u h s bn n mrcl xln. Dsbl us n a bwng ys ncd (a th cy duc b f g v n m; (b h aby o fn w dly unil fucs, glucs sus mls and altotos

    v bn s an c th onsy o ow n wo dly unl nnoal bac fnaon l nnn ys s n t ng2.S-SSg y wl n ac naon s sa that thya aa aly o h at h conclon of fnaon lthoh ls ncsay f cnfs a sd fo aaton (3). ltons noaly bas on h suls allscal fnaons (13) . ocontno fntaon n unsd s s ncay o a ys whh s srngly smny hugu h fmnn n d mnn s l a th bas of h ow (4 32 ).

    o ws sola sl nd mnn ys sras b ohna salst laboaos o ov th c h n yswhn a bwy ay b dd fo a snl c o sa solaclls fo ng yas colony o fo val colons (3) Aan sobws hs hv w o o sns my mly n x o n aa naon ssl Pns o stans n a ay howv chan cas of altaton n ms ocd, ndndvdul ans ay b lnad (03) Nvlss ys f svlsrns my d m sussully n sngl cn Cuurs anan t OC on woaa slos a 4C n caohyda a

    sch as 0 pe c ucos wo Wckas al ac mdu(243). bcn s ca ot a la nals (24) ably tlss n -mn nvs Lylzd uus v n n s -nsvly ba h s hh oaly of cll n fez-dyn ndths tans an aans ay scd 25

    Mny ys oaaos a basd on th on wok of ann anKuhle ad oerate eiher semiconiosy o on a bach bass

    w is un n vssl ha s n sili by s n cll xygn s us hug th w nd h ulu f ysfo h aoaoy s noclad Aaon o oynaon ay sld nnuusly u us m mn s usuy inmn 43) In odn a h oaao s cha wh 23 h wor f s ay 1040 an s chd wh 1 f pssd y

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    9 KN & O

    erion i provided or min n ever 0 m minue proi-me 8 exponeni go our fo en ome kg opreed e i vibe in p fermened bee o pei grvi 06Wen e enre onen of e opgor e dirged no of

    fre or ere no g pe 222 i oe rin of evel of beer produe in e propgor re lo n e e reeonge b ee ee re o en e e i ued norm 43oern indrooni fermener m be ued e progor ndrrednk onnuou erener re priur good 6

    ropgion of breer e enbe beer o ee e enreok of e on preeermined bi requen b of e i uedon bo 2 ime before i i dired ere re, oeve breerieiming eir e no been nged o 0 er or more 3

    e nge n e perude breer o dir em ree eiero infeion wi eri or wid e poor eing ner e end of fermenion f boom e or pri o of bii o gro, ermen nproue e expeed qi of beer

    YS MNGEMEN

    n e erge rewery lrge inoulm of ell i ue 515 mil-ion em o or n e ermenion e nmer of e ireeree o ouod. Tereore oneid o oneour o e e rop oe fermenion i ue fo inouion o e nex I e bwy

    o uien ie million bb of beer o more nnu in e UnedSe, ing o e remining e or ue n nim foo uppe-men beoe eonomi eibe ernive e e ued fonfre of e er or for frenon n grn iilere.

    e olleed or repiing i uul mixe wi 23 voe ofie er n ped roug vibring reen o ep reoe biero rub prie 6 In modern reing operion e reenee pe dire no e opper ereb proviing e requiremoun of e o rep iing dori Brewers Diget 2

    One dnger in ing er nge in meboi vi fomermenion o repiion 3 ereb inreing ueibii o uol 1 6 Converel orge under ied er beieve o o uo o mnmum 00 e o be ored for proong period o ime be e f in e ferener under beer 3 One nger of proonge orge i inompee ii o feren upon ree 1) A inmu 4re pero i eieve neer efore reuing e 7 bu preenprie in Brin i op nd boom e n indroon veebeie i beef

    ome uggeion for reuing e oi inlue iron enrimen

    n minenne of ig C o N rio nd e dion of ured d o or 3 22. An morn inde o e uoi iinreed proeoi iv 0

    e onmine i ber poilge beri y eier be reple

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    IOBOO O BW

    w pe le wed w as phosphoc c (45), onm plfe 2) mnn eef 7 eey emnn eneey elemen Ye elemen d wng n eee v ne lly ee eel emenn e ye

    eme lmed ewey 6) eled nfmn n yelemn nd d wn fd n en len n Pgton of e and Microbiologia Contol in Bewing, Fmntation ad Pkagn nlg nn

    A EABO

    ewe w 4) mmnly 814 e en l d f w992 pe en e yde e m yde mnen wot e gucose, uctose mtose, scos, totose, nd g

    e nd pep l ned plye gle nnng men ewe ye e e g p mle n lgemoeces 91. Moe ementbe wts e poduce th ts use e n myly enyme nlned e plly . enge mng empee nee femenly 86) ng e ppo o unte cee e tepetue o mshing nshes wotfemely 13, 1 ) lly, e nenn ngen mel n e w nened y e ml nd e mel ed n wn nd y mn nd w lng ndn 9 w mnly ve 0 mg / ml, nd ngn n

    en lde mmn mle mne, mn d, ne, nd mple ed mle pen ) e m mpn e f ngen e n d Plne, n d, ndn l used 1 3) n, nl, pne d, pydne nd mne e e-en n w nd led y ewe' ye e nen weeen 2 e en e w ld pe lde l-e nd e nn e een w e n C g e Cn Zn. Phospe content s n he nge 1 g/ nde nen n e egn f 4 gle 12) led ygen n-

    en ve fm mgle 4)e gw nd e ewe ye ve eenly een e-vewed 19 ) Ye ell edly e p mnde y ledn 12) nd de ene e ell y mn pemee yem 93) w ndle n me n nve ne. le e l femenle ye e en p.ee l eqene pe f mn d (le 2) ply e-e pen e pemee e eween e v d 13,) e ye le ynee en mn d me el ne , yne, dne, gne, nd ene yed xd w

    e n ned ny exen fm yde melm nd fe nge n e nenn my e e enel mem e ye nd ene e ly e nl e g ntiton om-led weve e mn nd nle

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    6 EYN & O

    B 2 E O OO OF MO FROM ORBY BEE' E

    Group A Group B Group Group D

    Iedae Abobed grada Abobed Oslowlyabsorbed dn erenaon era a aborbe afer

    Arnin Histn Aanne PnAsaran Ioene AonaAsptae Lucn GycnGutamat Mthonn PhenaanneGane Vane rpopann Tyrosn

    n

    Fo Jone ere

    td nto t du pclly wn nging t du trby u tert n brn prbilty (4 3)

    Wn yt ptc nto rtd or ognt wort tr t rtg priod wn t cll ctly tk up tril ro t wort clding

    h olv oygn t not rtn wh th oygn i potnt or thgrt th t bt t rt t o trtd , 23 nunc tochondrl unction 36 h ll o xgbout 1 glitr) nuint or ny gnint robi rpirtionn nd th g of frntbl ugr nur by t rbtrt 21 1 ) tht th tbl nrob h o nrgyydng ptwy i t gyol EbdnMron EM) on btth o onophopt unt ni oprt to litd tntnly for th ynt of nto (102. Pv h h ta, rgos zc carbxt rcto t t

    n rbon o. W t th outtnng tur o t tbol urng br prouton pil or nd ro of b y ro no ohl rton l 3 notl tho tng forvc ac Fr stac str rs ro a tracr rc -

    ong clo opound lool n T ) t tt thu prod fo tyloA nd tnol bot podut o pyu idtbol. rou ftty d lbl wtn t ll ot in tr ynthi cpt tht proponic obutyr nd io id do noturn ty tr Lkg o l oopound o t nt

    hg tt y lo contbut to th ll o t o ncthy prylt 14Estrs otr ta ty strs t s cos c rs ro

    thr coht or ino d toi gng rg oo( 1 111 11) o n o t rquint o t yt

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    MCOOLO O N 597

    S RESOLDS OF OME EER OE /ML

    n watrn lagr n dgassd

    be br

    ehao 36.9 100ano 820Popanol 08 50al 01 002-hyroal 055 002yuanol 41 50yanol 02 50,nyano 0007 47.9 50yl ac 025 93.5 uy acat 00

    soby aca 007 my aca 000 3.44oamy ac 00 0 Dacy 00021 0162 0005

    ro rsls givn n fnc and 202

    be enycay decabyaed e cesndin deyde wc s enedced yied e fse cs (be ) s e ae ise-cne, ecine vaine, and enyaanine f w ess in dcin by

    e yeas f 2 and ey ban isban and eney a ecice f yeas san cndiins f feenain, and w csn eacaec fse ac fan eeby dfyin bee av ad aa andvidn aeia f ese syness

    cein dacey and 2,3enaneine ae na bee cnsens bn ecess ey s e bee by e sy bey, an ey a esecvey e esd f eance f vicina dienes s in e def 020 40 cen s dced f acive aceadeydeydyeyiane ysae and fe aceadeyde n e

    esence f a cabiase Yeas des n idie acein dacey b nsead ends edce diacey s yeas s fen dded eed bee fe eve f vcna dkenes s cive aceadeyde wi eac wiyvc and byic acds ye aceacc and aceydy bycacds esecvey and i is beeved a ese acds wc ay be ecss f vane an secne ise se een f e yeas ces ne bee. y deabyan and dan wn e bee e vicna dkenes ae dced (21 ains f Pedioccc and esiaydecenan f bewes yeass ae sees esnsbe f eves ia kenes .

    Yeas eies sf f e dcin f ens cenyes via-ns ec, and akes anc sf f w cey as enne, andinani s in e f sfa ( 152 Hyden sde is eneaeddin yeas meais and deends in bewey feenans n e

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    8 E & HOH

    S x S I S 2

    chos ldhdes Ox ds min acds

    thn cdhd vc cd nn +Gc G drpvic cid SrnPrpn Ppndhd -Obuc cd -mnuc

    cidsppnBn B dh dn sdhd -Osvric cid Vin lTr. nsm lch svdhd Oiscpc cid cn

    c. m hl c. vldhd Omh v scincd

    n nlpn pn

    Ocic cd spc cid-Og cid Gmi d

    hnhl ch hnpvic cid hnlninTrs dphnl Tsin

    pvc cdTpph Trpphn

    Sminn p d Mi Ofd : Bckw

    yeast san used te tepeate and te wot coposton ) egas unpeasant ove cetan teod level ases ete o leaage osulde ons dung te enyc eduton o sulate o oe lke b to o ee deuhdre on ee ( 133 ecaptans de,an harboxys have been implcaed n te avor o bee ee-teess gowth eat n syntet eda and wot ges se to no sn

    at level of volale orgac ulfu oounds ). ese opons ase fo nonenyc ecton n te bee ) and o t o polage baea ) .

    RWH AND FEREAIN NEICS

    ewey eentatons ae caateed y te use o a cople edu and a age nocul I bah eentatons te tcng ate s ithe order o 4 g d w/l d e hrve 1 tes saont dependng on te yeast stan used te coposton o te wot ande pocess condtons tat appl Vaatons n tng ate stongl nu

    ence e e o aceve enaos 2 and the aly o eto e atoe ) e a nube o vons nmy u uay eans ta a tue expoeial ow pase s asent an i

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    MCROBOLOGY OF BREWNG 599

    rled by l ler ee i ell . Ae wh eur beue here uey mlle rhydre u myl u he y h e u ue due rmre edm ( ye emue ye-hed ( ye

    e ul le le by ddul ye ppe o e epeen he lopeee ye he verll le iviy (79). evel pH vu lui KZ2+, d H4+ r be r ulzi le anloioe (234) The poane of Z+ leel in wo i ei hey f ye eied (14)

    The rle ye h be eed le ei wh hebee e h wh d ei k he ed e wr 0.5 lved ye eely bee h

    ( e e uri level) led evely e eme (0) . he nge 00 pe en aaio o wo by ye eoe phn ole oygen oneon ely popoion oye op n o feeno e ( 5)

    AS CLL WALL

    The eng ye ell wall ly g lyee 102 m hikopng gl pe en) nnn (4 pe en) poen ( peen ) ld (7 pe en) nogan aeil (3 pe en) hone nly ( pe e Thee rri ie d y d

    ig o e ain ell age, n rwh d (, 56) h ll haoae wh i in ee an on fo invee ai phohae le poee n in he e of S cabg ele (19 1)Glyle pee i n he w o S , wl ye ( 11 ) nyolyze ee ein Ohe eelll enzye n he wll inleglne 14) nnne ( 4) poe lphe ee 3 whh wh he poee e poly epone fo hyolyng he wll( 1 ) n h inene he ng poe n he nal hpe of he (22) n op feeng ye en n e hee y

    lng o eh h f r frm he ell her bewi ye y lu eher r eeilly he bee ur d he reee ivale o(5) The ehm ul rbly vlve he reti lbrde beee ueil hhm dae ell 4 ) an hyogen onng lze he igig ( 95 5) he eionof o r beer liion y eienon r ru an of bew ye hve negive hge eae uel hhe d oyl gopg () n wll heeoe eawih povely hge ning gen h ollagen ( 14) n eain

    geln. he y eenon ye o fo a yeahea ( ilo io of he ell wll fe he ne eponile fohoing he ell gl inee hve bee ide

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    0 YN & HUH

    WNG YT GNT

    een eview y e oo onow owe 7 noe & whone pove ey noion on hee neee bje: ye yoogy poon n genei epeiey Bew-

    ng y gene hogh ovng y h owh b o h oowng (a n o h o pon be neoo ve he ee o ye on ee vo no yeenbe o gene ono 47 Soe o whh e ne o-on 9 25 eenion e n 72 228 oe 78 85228 ooe 228 n en eenon 8 n o yee on n e e 9 ( The bewng he o -oy ye hyb e no wy he e when eve n eenbewee ning heeby h he envonen py oe (0 o

    ebe fo bewng he eveopen o new n fo e n onn- wng n non 4 (c o n e oy-p whh po poo 57 28 Ao hewh i ow o-poe vby 5 Th ppoh fo eng he geno n e ooewh y nion o o hpoon 58 n poe y hve no poe be ewng y 22 h no ng ne o on e eeeo Ohe eho o -ing he genope whh y pove oe ene ine o o-ngove gene onveon n nonnon 7 Thee o hghe feeny hn on n n oee be ne by he oophnllnn noh pon ppoh n on 78 79 hogh o e h no povn o be e whye 94 1 14

    MICROBOLOGICAL CONTROL N BREWN, FERMENTTONND PKN NUN SNN

    In gene ehod o ioioog no hve no hnge gnny ng he p 3 ye eep whee ie by hnge n beepoeng. o hng n bee poeng ne he e o oe

    ope o open vee o ewing, enion ging n pgingenn heewh i he opon o inpe enng ye he poon o nonpeize pkge ee n onno ewing n een on The wo poeng hnge hp ee beoog onnon, b nng h n o n o n o p on pee wohy o onon n hoong oioogqy ono pog e: (a Ue o n wheneve poe e.gn eenng ee wh onie n epbe eve o onnon?T wl vy epenn pon e vel egn n onon f heewey. h opn eene n e ewee wh wooe e

    e i o en n wi no ob hve onby hghe ee ooninion hn oen beeie wh oe eene wh eyoen onde e h ne ee oon ene e i

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    Y

    . br o b psu or rf b o b cosumwihi hor pio of i coi sll ubr of orgsmswh b subct to sptc g c wihst v fw f coi ogss

    o o h o c suv pps o crobologcl coo

    is h 17 41 66 2 12) Th thr o cool s oiog of bws s qul; cto uio coto of cobil cos ; tto of th bwig vio

    Ys qu c b m b poc mum of h foow-i pms u h cou of fmo rmo ( 6) occuo chrcrcs 1 of wo uo (21 ) oub of cs prctg of buig vb cs (122) chh of cls (26) ct o b vo h pc o bc ofoig mcroogsms So of th o ct ovio cu h

    u of uosct s 74) plc of mth blu o ho B( 177) o i t cll vbli h us of Cou cout pc of hoctomt to trmi cl umbrs 1

    Th mot b mtho fo c comt oss s ptgo i owh mum 7 1 1 17 16) Dirc mcoscopc mio s of of o o vu c th como lvl m boo ow o c ths mr (2 .

    Rc s hv s k pov i foulo of fo tcto of cobil cot u lg to b ust

    g of hir ct utitol qus (21 ) A c mpl skgw's u (169) for cto of b rc ois 1.os souc ot us b lctobc 2 scobc c fo h-bo o obc c 3 co o ihii bw s lowpH v o hbi cssv bc

    wo o msurs u b m bws o os s como r c w of hv co o Y M) poc rplcm of st wh fh cuu (s co oY Popgio) h cutu o po o h coc ih cs uch uc of fo hh

    b h ult of gc chg th populo.O of h st s to mcob cotmio of bwrs

    pc ioi. 1969 nn w 11967) whch rc s hs hp volutoz bw qu coo. A ct h ucc o uch opo o hvig o fo k boil o fo io pc ci i h poc of cicuig c soluos houh procss qupmt rov h so mlb chmc cto wo tps of crcuts us cos crcut o

    h chgs pp is op ccuit fo brwhou vslsfos h lk Auoo s posbl spg of clgsoluo pt g vv stg s lt b rgg for th

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    KLEYN

    puc inan a g pu an anng un wcic canng an aniing agn avaia prp ic pnin in par n nau maria an ( m mr cmmny u anng agn nu vau han 76uarnay ammnum mpun 9 an manu cmpun ua rmah 7 an ramn

    anain wy nvinmn i aciv in pa wn u uravi ig r riiain war u inupmn an n urav g uc may ncrpra icy n cy war in. n rr acv 1 p cn riain ii impran a war rn parc uc a rn piwc can acia rm g uc. (yn J 93 navain ( b Uav gh may may u ar ran (2 Hwvr i n ug n w u w pi

    cpin apc n rm ran () Dumcan vaiu car an packan aa ry inin m rw wic inurn w p mn muy a n pn 3aag a ug a umcan ym a p ruc num crganim prn in ai ( d) Can rm cniquuc a u piiv a ru in an ap ng rm Mn a an pa ra pgam ) Iprain ngc n guing mn u n mnan ca

    ASPC PACKAGNG

    akagng a ng p y a gay apacag pruc y man an cnvnna unn pauiain Curn cmmrcia pacic incu a paurian ran ran an cmca aiiv Or pp m incuning aan an iguncy ica ( 3 A mar avanag a prucng wi an imprv a k avrCnvry, r ar rw w wi n ap prc nc unnpauian i iv mak a piiv cnriuin avr r cn vanag n can apc pacagng h a paagg a pa vna un pauza-in (9 A r avana ra a avng pac rug iminan cnvnina unn paurir wc ur cnia rr F viw n p pkgng nu n 7

    On prm p y an mman ran an m n y cmica av nciy avig an nay r pacagng aa cup w wry wrkr ain in micga nu ary a pakan. na um aa apic pakagng ncu a prucn a r wi

    rmna ugar prn Anc wi ni pucin m camnan mcrranm ir anuan can h hgh ya ran up h g rwin

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    OOO O BREWING

    n eenon es b otion o ee wit low oxygen onent ygen ses ot ioi gow n oxe niy nso eeoe e ee o ni Meos oose fo eingte ontent of ee ine te se o nitogen in ios sges o teotling oss (99) c oon of ee w low ne of og-niss e o gow n kge ee. A o eqieent eein is see-ton o ewes yest stn w is ne to ow in wettentee oi ontinnts e esent in ig nes eoe ton, one w no e e to elne e w on 12) Hig nes lso ose oe wi e ies sne te tty e-enent n t on e onenttion o ontnnt ioogniss esen n te ee

    One of te o onnns fo n sol kge ee s Saccha-rmyc datatcu , 124, 219), yest le o lze ns et n

    ee w el ewes yess, S crlsbrns n S. crv nnot ege (0 . Gensn ( eene tt s ew s o st- s o net kge e o ny sie ste & eniste eine tt seon o onnnt o kge ee, Lct o inet wt few s one e e o ote of ee Btof e oe ogniss gow wel in kge ee in tt tey n tete ogn onnons Oe ogniss fon oionly insoe ge ee ine ees o e gens Pc, O-bctr n Brttnc

    Mjo oes o s steon ne efets in ee o en t to neen eng o ee n e eqeent fo et llnge tte qen is lso nesy wen sng ene ton tis oe to soe exent y ei ities ny 6) n os Glno 1 ) oe ets n eeits of te st two entineoesses. e tes eny se n ee inle ety yoyenoe 6) n oy gll o oon, teee, e e itons eey ong oony o new ne ites. oe oe esetes ete t not in seo e e yoone ) yogen eoie (29) n sls of ylen ne te 121)

    e ens of on of ey yoxyenote is togt tonole eson o te yosi ene ( 13 wees A sogt fnon y nng el ons s as M+ essent yetgwt ( nteestingy eng te tty o eyyoxyen-ote s enne y te ition o + M+ ( 13) , wees ese os wi DTA ( .

    1 . As R W 1970 Unpubshedlt2 Andran A A Si J B

    J.

    23-36 At, R. G 6 [ 776

    G Hampton, N N

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    4 Y &

    R Roberts, R. H. 9 In Brew Lndn 7:26077

    5 R G Wdward D. 9 uaran 4:1-3

    6. yaaa 96 r. Brew n! Conr (Nie) 276-7

    7. Bah McKn W. E. 19 Can Microbl : 309-

    8 Bae A. 169 rocesBiohem. 4:234

    9 B G hg J. Dwg H 199 Natwisenhafen 28

    10 Begander E., Bahmann K 197Nahrn 7487

    Bexn Dae E . 970. Gen Mrobil 60421-23

    2. Bhandai R R Wakr 1953. Gen. Mcrob. 8:33032

    3. Be . Hn R MaW C 1963 I. nst Brew69239

    s L R 96 Prc. S. Brew hem. 64-6

    . Bho L R 1970. n. Bw.767-1

    Bkemann J. B 9 Proc A ..Maer Be . -

    7. rge C 1968. Btechnque 3:239

    ay B 98. Ins Brew44-7

    9 nne M W nd H 1966.Tech. Quar Maer Brew Asoc A 319399

    0 Bnn M 1970 Tch QuMast Bew A Am. 49

    Bdn E Y 969 ab. Eup Digt fm nt Bew p 969, 44

    rw M Hg J. S 1965. Brew. onv. Proc. ngr.kh 3-37

    2. n hn B. J. 97 0 .Gen. Micrbiol. 427987

    C M Wg R rocess Biohe 3:2-22 2

    5. Bn G D Ran C 95. IGen. Mo 1

    6. Bn M 90 nst rew7-

    27 ruch C. W Hmn . neR M. Benn M W 94 IIns. Br 70226

    rmsed D D. Gene P. 19 Proc. A.M Am. S. Br.Che 12-

    gav N

    ben

    o A. .

    nn 94 r. Ves.uh. Ild. Ins PivvarPro. 43 A hem.

    958 52:19020. ge A H. 964.

    ultiatin an Utilization Ldo: Leoad H 00

    B 97 Brw.Garian 1 7

    32 . B I w d. 09-9

    Cagna-La C. 188. Ann ehii hyue 68:06

    34 9 u Pr ong ar 4

    a A 19 nst r 7:250-5

    6 Chan C ae W 168. Biohem 4

    3 7 Clk D F 1967 roces Biohem22

    3. me A A D 95 Bew. G

    4:6

    8

    69 mrie D 96 J tBrw 733351

    40 ksy K E Ribw 1962 Ien Mcbil 27: 3-42

    4. ran H . 965. eh Qa.Matr Brw Aoc Am 2:21

    Cost H R. 9 Gen. Mirobil 3 7: 3 1

    4. r N . ar G Brw Cv o. CngCoenhae 49-6

    zck, H T Van Eg E L99 Be D 2-4. Dean B. 97. Inst. Bew

    36346 e e . 9 A exboo f

    wn V r K BtWgh. nn: hpmn andHa Ld 87 pp

    47. e R H 966 Gn Mcrb 44496

    4 d 9 J Ge rbl24:350

    4 Desle A L., Phaf H ,

    roc. A o. Brew he 0-18

    50 Dn 67 J. n B 7348893

    Dx J K H 969. In. Br 54

    52 Dw B ech H OceH I Th F M 96Monch Bu 9-

    53 Dnan L K., Se H 9I. Gen 0297308

    ddy A A 198. It Bw 143

    Ed A . in . D. 95Prc R. Soc. S B. ndn1819

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    MCOBOLOY OF BEWN

    56 ms . . 98 Buer wiss Be:063

    57. e C. 19 Monatscr Bau9-9

    8 Emei . 966. Z Naturfoh168167

    9 Ee 1969. nasshr ra28

    60. Eme C 0 Brew Dg 4660

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    69 Gilliland, R B 9 J Inst Brew.888

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    MCOOOY OF W 6

    5 MMurough ose A. . 197Bioche 105:189-203

    17 Mddebr, 19 Walle-sin L" Cun 1172

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    338 Mthel, W M 970. B uadan 993)18

    . me, , Ian, K., Tnke, 1 Appl Mcb (Tkyo)11926

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    1 Mch, G 11 Bissschaf147

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    170 Naayama, O M., Fly, W. 1968 Pro A c w. Chm19802

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    184 Pend! A. 198 B 1084182 8 Pend, A. 9 auwnhft

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    7491300188 Pchl . 18 h. Q. M

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    n h as ed A Rose, J J Hriso 14724ndn Ne Yok Academ.90 pp.

    9 hds, M Cld T . 197 nt Bw 78

    3 hard, M 18 n. w.74;4333

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    0 Seg, G M., Lws M. J Wokw. 17 r. Am. c Bw.Chm 14

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    205. Shimwell J L 195. n. Bw78

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    Annu.Rev.Microbiol.1971.25:583-608

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    08 KY & HOU

    208 Shimwe, . . 1964 Brew 99 :759-6

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    u 5512 omk P 1958. S Chm Indg No 37-0

    22 tvn R. 1967 h R 6:191

    1 Sn J. 1966 Bw7369

    2 Srandov F B Bcklann J B1955 Poc A S. Bw Chem342

    25 Sandov, B Bocmann J. B Proc. Am So Bew Ch.94-97

    2 Stndov F B Zoo H. Brescia J A ocman, J. B1965 P,o A So. Bw h194

    7 Suomaan H. 1968 Aspts of KM H. reb 17 Oxfod:Baw

    28 ua H, Rkinen, P1968 N (Lod) 07929

    2 9 Takahsh, T 96 Bl. Bw Si1914

    20 Tkang S Hada 196E Bw. nv (Mdd) 47-88

    22 Toa M 1970 A stdy o Obsubu ptu mn bu PhD h Ung g

    222 To, 1964 Bw. ! 50599-605

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    nha :119, 198484

    23 ited ao Satc YearBoo 1967

    2 an J 1967 Ksn 1

    1133 Vermen V 196 d lafan d Aoc Roy Acn Eeed 'Inu Su d mnaon Ga. Belgm vls16 pp

    234 . Vur K Ko B H 1970 ! w. 763666

    2 V, ,W. 196 M nahr Ba 20 1 6

    23 Wacbaur K 96 Montssch

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    24 W D B aac L

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    Bioh J 984 Wiceham, L J 1951 u

    U. S Dpt Ag. N A E 9 n Bw 5

    65245. Williams, P T. C B Rainbow, C

    1964 I. Gen. Micrbiol. 35:237-

    4726 Wmon D H 1959 App

    Bt. :39-0

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    rsg hs 479949 ood B A Ranow 19

    hm J 780-950 W W A 1961 Fmntaon f

    aohdat and Ratd oound, - Th Bcrad C Gna, R Y SaneNe Y cac 7 pp

    5 ant J 196 . ns. Bw 68

    5 Yshid T 1968 Rp R Krn 11786

    Annu.R

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    .Downloadedfromwww.annualreviews.org

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