Retrospective Theses and Dissertations Iowa State University Capstones, Theses andDissertations

1934

The physiology of the propionic acid bacteriaHarland Goff WoodIowa State College

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Recommended CitationWood, Harland Goff, "The physiology of the propionic acid bacteria " (1934). Retrospective Theses and Dissertations. 13150.https://lib.dr.iastate.edu/rtd/13150

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^a«Hsiteanlam of th® Dlaisimllatlv® produced by th© Pf^jpionle A®id

laettrit In th® F«ra©ntation of alueos®

Sum*!? IVf

Qmsmsmm. u®

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i»R0wefX0s AWi nmiw m tjfiMfW

f3p@pi©»i© »©it lias l®»g l»«®n k«@im m & pwiamet @f fea©t©i^» !

ial mtom %%m eimmXmt aatmr® &f th® e<mp0«n€ | \

was taiowii i% wnt fomS ia t%vmm%i.ng a.i«pjSds« Owimg to tills 1

faet mad to tli© us® ©f Sapai*® ©ttltmr®® tii®i?e Is avteh mntmatm. I

In-tfe# ©ftfly lilat®fy ®f th® prepteiai© aeld f©»®»tstioa«

S"ll.3.a#p CJ.841^ sttiaied th® ©f moist oaieiia

tftftrate md efetiiiii«a * smfestame© lit "faetid©*

®8slgs-aMP®"» ihaagittg fwm tili 'dtssrlption of tti® salts it is

•pTOtoatol® that h& was wayfciiig wttli prepieal© ftoid#

fl.®4S) mft&T & »tm<ly ®f tli® Ited salts dsolawd

til# .Held tQ tm m mlattmr® of toatyf»ie aad aefiti®

a.eidi<«

irigllif CIS47) att«apt®d t® ©leai* mp tlila ©©nfaslon* Hla

d®a©j?ipti@ii ©f tills aeld ana analrsl® ©f tU® silirar salts ©op*

p®sij®ii# t© tHat ®f pi»©pi®jftia atl<a» 1® ©allad »llla«p»s

®a«lisl«i»®* "fttli® t© iiwiltat® its po«itl©ii

a@®ti« %M Imty^le atiAs*

fottli«% C1844) tey th© ©jcidation of siagar^ obtained a

smbatane# h® (sailed ^letaQatonsamr®**#-

l»dt®iitoaeh-@r C1846 J @a fefi»atiRg glfo®!'©!, obtainad a tf

®ttb»tai»® iimilar to §©ttli®b<s '*»®tao«t©n»a'ttP®*#

. memag ««3.a|pti mtA (lii?) ®o!tt®lmd«d fmm « stu€[gr

0f tH® mM mmwwiug %m the Qt ^aleita .

tliat their aslA wa« thm s«a« m

tlllBerts, Mi ile»ri«s»' toiitjnp©*ae®tlqtt«'*»

mm tilt 'first t© tts« tli« tmwm propl©iiiqt»®*«.^

• a©«r (Mm) protest^ ffogicitti# a©l€ fj*©a miarttii*© ©f

oitiPiit® ®urai«d allk*

i3.iS4| i©llt»mlng th© jpy©o©<lur® ©©Msoalf^ msed for

til© pF®pii?ati®tt ®f toatfX'ie fttM ©l>taia#4 for tii® first' tin®,

pr®pl©mi© a©lA fr©m sun®? f®i^iit®ti©ii» I® i?r®par®4 pota®siW|

mMMMt, fearltt®, ©aielia «ad silv#r salti of th® pr®plo»i©

»@i4. fli®tr @har«©t®rl«tl0i Boi»r®ip@iid t© ®«lt8 ©f pi»©pl©i!ii© •

a@|t*

Sl#ii®m l&®li®¥^d that iwi^®r ©ertaia ®ea<3itioiis

htttjrfi© ant ft©«tl@ aeiti m%%&. ©©aMa® t© gif® hatyfo-

»©®tlqtt®*» fhl® ®©M was a««©r4l»#a as hating mil th® pr©33®rtl®a

:©f pr©pl©iil@ ft©l€. Ittier ©©nditt©a# »©t w®ll a»fl»®d. It

d®e©«p©»® lat© feutyri© md ft©@t4© aslda*

fiti p©lat@<a 0ttt that! the ©onfmsioia ©©aoemlng

pr©pl©iii0 a©ld aros® ©hltflr fro»^@ asswiption hf s®'^trttl

w®rk®r» • thfkt oii3.|- ©m® f®wi©atat4©a ©ris®# fi»®M @ai®lwa tartrat®.

1® ®3iplaitts that th®r® ©r® at l®»at thr@® «ltff®r®iit f®wa®atft*

tl@ii8 ©f eal©lm tartrat®. 1®^ «©r@®ir®r, ase®rt*i»@4 that th® .

ii®©©»p©»iti©a ©f »a©li» lmtyr@-a©0t4qm®® mat b@ aaaritoed t©

m imfarltyi ©r »lxtmr®, ©f a©li». After thla mwk th® teran

"rneii#' »@ld an<l-p8#ud©-«a®%ie

meii trm th<& tttmmtmm*

fiti item} l»gan «liat•• t©<lty• is #0»si€@f@d ' ,

%%@ ttmt &i iMportaae# thm efe««iwi ©f th® pfopi©iii<s

A#id 'teaetefifi* 1« wan thm flfst to fttt^iipt • t© eoaa t© m b«tt©r

@f til® #®iiir»ygl0a feifomght uteoat % propionie

a<iit ba0t#Ma« fh® studied w®r® tlioae.of ©aloiti*

rnalat© and #ftl#it« J.®.dtat«» fit« md® a eai»®3^1 st«% of th®

salt a ©f pwipioni® ®ad a@®tl@ aolda^ diatillad: tii#

mlmtll® a@id#,. foiMdd aalta ©f. tti« fFaetioaa -and tli«s .

daflitltalr p«»T®d %f til® 'teoillng poiat md Btlmip oontant tliat

hm ma daaling .with-propiiniie • and aaatie aeita# la ©©nelmdad

that', fi»ct» tliy#,®, »3.®@iiias <sf iialie aoid# two mola.emlaa of

pi»opl©nis aeid ttiiia-©a© ©f atati® aeid mre fomadj,

^liita.® fomi? aol®ott3.es of ©artoa dioxid® a»d ou« moiaenl® of

watar ar«.apa.it off# ffom His wofk ©a ©aaaitsa laetata lia aug- •

i-ptats th® following a^mation fof the pfopioaio aeid famanta- ^

tiofi*

^ %«#g •, V#t V Matie aaid fwpiomi® aoid aaati® «®id

tha liaitad data wMali fits poaaaasad, it ia -

ipaaarkaMa tl»at aa©mi»at® oonaluaioiis aomld liairo feaan dram*

Oonaaimiiig tha oi^faaiisa, ,,fita mw&lf atatad tkat aliOPt

ejliadrisal rods gmm tlia raaults .and that the organia® was a

alandar, loag Mailtesi ©ftan'aa^^sptl'ealla totb fomad togathar'

in Img amrrad #liaias»

tH© ©f fiti, stu% &f tli@ pr-optoni©

aeld Meteria wai ©T#rl®€^«<i mtll th® mf^k of "Ton •

anfi tPla-jrenieH {3li06)« fli©s® two workers Inter-

in th# ©dor and tustes pro«iad®ii by Materia in

olie«s@ ige3.iitt®4 pr^pioiii© «ttid-fr®«S»®iag b»0feitrta froa g1i.0&«@

ttsing a ®iirt@li«®»^ immxm witli an

©amlsion of fro* th® flasks|, in .whlsli volatile

R®ids: w#r# trso®f®ra w@r@ t©- gelatin and

wiitr-agar shak® fh« tw© in^eatlgators aneowtereia

great! diffieiilti©# due to th@ rapid growth #f• B&otari^am. ^aettis -i

mMl istm'ttxicmmu iaetia) • It w«« onlf after persistent 'and

©ar«fml that th»f were afel# to i®%®et an^orgaiilsm but

sligHtlLi' <ilff®#®at froa Battariitti la^tia aeldi*' l<r@n aft®r

tli®y liad «l©t®©t®€ tlie &v0km.tm in tl&i® f#r»Btatl©ii liqmifi, ita

it©l«ti©m ia ,pmr© ©mltrnr® was ©3ctr#m©l|' tiffieu3.t» •f®rsl@tem|i

in mming tr»KSf®rs-li©w®ir@r, l,«d t© t»i® iaolatleu and pirifiea-*-

tien ©f 8©v®ra3.' atraiaa ©f pr#pi©ni® a©id ba©t®ria»

;. Mff»»®iitiatioii teai«4 ^i®fl|* mp©ii m©rpli®l0gioai ©Haraete'jw-

iafariitai. th® •traiss, ipt© tw© »p»0t®», ©a® witte t«o strains• •,

Bittt® riiia ' atiii ftr©,pi©alei - a a"fery 'Sfeort- rod ©r

stir®t©li#a ,©0©4«s,j>. •©aail'i'' ©©mfmatd m%%h la©t«rittm • laetit a©idi» •

la®t®yi.^« a©i#i i>r©.®i©niei fe» a abort rod fery similar •

Baetaritia afidi progioniei a tet grewa at hi.A®r t®mp#2*»

•,:atar@i • I ©jftiiMts-leng, irr®galar rods or involution

f&m»0

Baoillus t.oidi ppoplonlel,. a -pliimp irregtiler or

8ll.fi»htly rod^ less snfteroblc than th© other two forma, '

Th®s0 invsstiffatops found all three forajs tO' b® easily

stained, drsw positlv©, non»motll©t gelatin iion-llqti©fying|

faoiiltativ® anaerobes, ha-^lng- th# atbillt|- to ferment lactates

with th© formation of propionic, ao®tlo» and carbonic aeldfl.

fh® oh@mi©al and biological beha'S'lor of the propionic

aold bacteria was sttidl#d and products of fermentation were

d®t«rBiln0d,/partly quantitatively, propionic acid was identi­

fied by tM eharaatarlatio octahsdral crystals of the basic

l«ad salt/ the sll'«f«r, content of th© silver salts of the vola-

tll# acid was d#terain@d, and for th® quantitative datermlna-

tlon of th© volatll® acids the Duclaux analysis was wsed, The

results of thes© analyses substantiated th® ©stabllshed equa­

tion of VltMm

?on Fre«d@niP#|ch and Orla-J@ns@n found that only Bacterliana

acidi proplonici £ followed th® ©qtaation completely sine© th®

ratio of p'ro'pionlc aold to acetic acid dropped below 2 in

lactate fenaantatlon with Bacterima acidl proplonici b and

BftCillua acidi proplonici* fhey asaiima in th© latter two oases

a small amount of th© lactat® is broken down according to th©

following equation#

2.03H#3 3,.-0g«4Og

Van,lfiel (19S8) hai pointed out that this explanation is

not sultabl# sine® with th® three different strains of bacteria

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Bm#iitml fli© spets fS'ev#?! tm to® «l»©st pnm eultupes

©f; trgsntsiis -aliiwliig gr#ftt .ijtellftriti- to wm fremdenfeltgli amfi

a®lt baet®5?lft* ffe®s@ #i»g«ai@®a produsffl

ft aistiact t&6. awA "timmm fi.'ga#»t., fh# amtli@s*a 8tigg6it«d th©

RCidi pgo-p|.€>a|.iti -y«y -guftem f@r- th© •©yganigaas

fomimg teO'Wi pigp^nt itftd acsidl. pi*€n>ioil@i ¥»!* ratomii.

•for >hose fuming -red pin^ent# following th© tiaod

Freu^lteEyeioh tad Oflst'-fisiis©!!, dtt®»inatioiis wer®. mafi®

of th®' pi»©4ttets,« fh© fatlo'of propionie to ,ae.©ti<t

aelis was fm'm&. te to# 0 f©'i» th® f#»i0at6tlott of •ealciim laetat##

fh® of •1 %wmm of fowAo aoi4 w«® alio i?©p©3Pt©a, as in

t.h@--3.m«t ,#i^®tiiILis«tiou th®. ailir©!? salt ihow##- a

'to hteoM . hl.aote# fh#r stat® that the fomlc aeld in,

no ao.tiht,iiot a prolteot of tho-j^lk rogar or laotio aoM f^iwinta'

tioB b«t ftFl»®s fro* th# p?®t®i)a sftterlal# loth org»iileas were

fomd to hti'® tht •'ahllitr to. feiwent «llk smgar, hut the r®tl® ..

of -pmplmiQ iioM'to «e#tle aoia w©e lost.••than two. fhes#

.emthoPS' were mt «hl0 .tm aiir eai# to s&tablish •anmljtlomllj

th® pr®s©iiGe of laotlo' asld im the »lik sugar. f©rffl.eiitatiQii mn&

thm littitel# t© sttpport th® .hypothe-sis that th« Ipetoa®

f@s?s«iitati®» paeaon^ throng th® laotio aeid m «n intermediate ^

•fifo-dm©t« trot#in dooompoaitlon was' fonad to'h®' sligit,. fh®

haettrial pignut wai oart-fttlly sttt€l#a aa€ forafl to hav© th#

l>r©p@rti®s of e&rotln-lilE® OMpoimdt*

laolatJft, pTOpionla aeld baot^Pia

fmm ^Sweitah ^ fha, tlffifittltios ©f'isolation of •

til® ppoplotiie 'aeid hmtmlm were t© « mBA-dd assent i»0ffloir@d

% tli© atdjiistaent of tha n»tpl©iit^ t© a alkalif» ,

^a^itloa. fitii. lirtaiiMting lt.@tle acl4 feii#t©i»ia w&S'

thms' . -ftlag tMa aoMfled pi'0s®€«2'© »h@ ©x»

littl®' .aiffieml.tir' im Isolating and pfmrlf^lag' S0f0pal

strains, fti© mjeii'lty of fcli®;Si® stTOlns were found to ^bilang

BftQtQyjiaa nelAl ^a*pgleiil4l • & amd, b gi*ompi of iron, Fi?®'Ud@ttP®l^

•alld^0l•la-f0ll8#a# • .ja aitltlop, l»©likt«d a itirala-iiftiea®. aimoi'

trm thm® Qt the tw© pyo-rlom®!!*

gTOujis# . ih© ,o®ii#d it Mmt9Ti.vm »0tai ^Ki^^ionioi ©• it i.®

#specially sl4a»ett3?i«#«l toy Its • i» a-lisnglni^ dfop

#i#SNi it lj.as th@ ft|}p«ai?g«0® ef • mn .awf^lutittatd^ eu.ltu:r®«

^QttantltfttlT® .d@t©»t»atloii of th@ wlatila a<sld® -was not #3t* •

temirn'm It was fomndi Ummm, tim% two. »ol#0vl®s of pwtploal©

tola ftlong witli oa© mol#«w.l® of ao@tl© aeid W0» fofwd#

, ^Wolff |1S12) l»olftt®i m fm •mltwpmB of orgsi^sas. kiairiiag

the ability'td .pwda©#'pjmplonto aiit a«®tl® aelds mA eartoon

:ai03ti<l# from l«0tat«ja,.

iroJtki#wl«s '(1183). i8©l«t««l -SOTeral str»iaa of pyoploaio

• sold haQtBTlA fmm mBMim utoleJi found to bo3p-

I*«jsp0«d,.»02*|(h0l®gi0alljf • ttfid i» tlhelr laotste dtooaposition

ftolAi pi»oi?loiiiol In fell# fa went at ion of

laotos® a wii® irai*i«tioB mm fom<l froM tli» tjp@ttkdoim

•msw«ll|- i«s0rlte©a in inveiti-gfttions ©f pfN&pioiii© mM bsoteria#

flj« was ^eia witfei

an ft-dteiafttip# ©f fomi# aeidt' la th® • sam® eas®i m trmo# of

Mgi.#r itQii was • •

•, -fli® flm% lav@atslfa%ioa« of th# fi»opldnie a«14 Meterla

ift %li«' ir«r# out 'lii* sli®w»ii md his 0O*

wmkm9* IS Iftl w0i?lc wm to#gum 1jO'Sa«k ©xplanation .-of .tij®

law •qmlitj' of lasrltsn ^isa follo-wlng tfe# •

of rm amd 3.aet«t» organ?-'

laai were fro«'.iia^f-te'iaa, ehutats.#. m iiipoi».taat «odl*

ftaatlon. of t!»»- @upi-«lia!©ttt ii^dlm of fan fTmuAenml&ti sad O^la*

f©ni#ii was lii%i*odtie#4. In -plmm- of ©slslws last ate, sodium

laetftt® wa» -fliid lir tliti. aoaifloatlom pi of tlia »@Aiw.

was -raised fi»oa -S.g • to, f.*S, ia »<ili, aor# fair®i»at>l@ t©

growth of fsli# •p»pioElo oyganlsai#

ppoplonie tola fe-aottpla Isolalieil-.^toi''^.©iiaaa

ati^ Hia-^ 0o»»oi?iE«i»a miey el@a«ly ••witta. Bft<st#ylii3a:' aoidl

^yo^ploniei a* . •^#1' pog,f«ft#4 th® ability t© fe^a^nt:: gl«eoat,

laotoge, waltos®, nmmm., Imlin# »»<! 'laitfiiiitol. Blffe^eiie®®

»#W: stTUln '8®^ • SSSSSZiS ppopioniol. % mp-

pmF&d sufflsitiit to.'8i*#at# » now 8p#ol®a# fliia apaoies gives

an field i»eaotlon in millc «fid aamaes It to -soigiilatt aftoT a "

tiisi©^ 'It ttlao attft«3(lts gl^osipol'moTO stPO.n.gly then, th©

type. t% w«» nmm Bao.teyitaia toldi pFoploaioi d»

Sliaw sad :Sh#Wikii (3.923) fom«l tlbat th@ typ© prodtieod

oafbofi dio:sii», pFoplonl© a#id mA a«i©tio a©ii- from swii.ein®t©a,

altrogeiiTOi eoapomis ami^ ptAepa to a .aligtit txtent fi*©a •

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»i©ehaftl« ©f tJjf a®i4 fsiwaatatioat AS « nitrogen

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HgO, sr%

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Qaantltati¥@ aaali-ais of the fermentation products of irmrlomt strains in jreast eaEtraofe-soditam lactate inoubation mx 0T

strictly anaerobic oonM t&m at 30 0

Quantities in Milllaols Total

Xaotio vola-Straia aeid tile

f«r* aoid ; ffl@nt#d pro­

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Ratio- P»@* $xiQ* F^ro- pio-ait • li#®ti® darfeom oini©

pionio .acid aeid dioxide aeld acatio pro- ^ro* ' pro» pr®-

duoad dwsad , ^cla©#d duotd

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308^'CHOH*coca 'wnffiig ' unun ' WW* Laotio acid

4

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• ^Qaallttttiw iw#8tlgftti:0m ®f glm®©s# faM^iitatiott

shewed tlie pmrnum^ of pwpimi® • acslAi settle a.©id, ©artooa

4io3rid#t a<i®t|'3y«©tfei|-laai'Ma@3. (tface ) mA twusini© eei€#

^i^antltati^e raaults of -dtot^ln^d hf

lieJ. iiai" fe® hf th# fellowiag tato3.®»

f|I», fan Ii©l il928, p« 13.8)

•^emtltafcl^© ayaal|-sis of th& f©f®#ritation pmdaets. of ¥&rioti« at rains in ©atrstet with glue.©#® »M ©ferbonat®,/

Inouh&tim ttn,<i@r strietlr anae^oMo munitions at 30®0>

—— —"mS'" ————— B03Kt»s# iatlo pi»oi- j|o@ti«s ©artoow 3ti©-

WsmMm f#s^^ til® ?2»o-» piOGi<s aoid dioxide eiaie n@at©i. pioai© aaid ^ pro- pro- itoid

m pm* • s@#ti® pro" dmeed pp©* dglgOg . dm®#d ftOid di*@#d dttOtd

• 1/, , 18S 178 S*7 m#0 4S»0 43,i IQ.O 4 ••.' im %m M isi,f m»f ii,4 M.:'' . 183 , im t«4 %ma §2*4 Sl,4 ' XUQ

Til© followinp5 breakdown @f tfe® Sttfaip is , van Misls

"e'ls"® * -* 0AI®6<^488> • V li®xoa» »onopli©«phate

36«1I''6<«>4«B> — Ws - «3HS02<«'4«E'

fliratt*!.!!# trio«@ pMoa^at® iild®li|'d«

©IP t%lglyoaral

W8<«'A' ' V SVs glf&mlm aldetoyae

©r

-r 41

r ^m m'

<QM

® 01: methyl <?lyoxal

hydrate

,iilpli»-hrdr©3iy- p^opi on» IdeMfd#

Q M # H„0

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pFop,l©iii«f aeii

Glg*SO--0OO« • m p^mvge m&M

i .©a# fi«»- § • ,©001#

pymiflc ftoid hyarst®

1 «g»0OOl # OOg * SB

i# m mtlv&tea ^hfdPi>gm

fk@ pTimipl^ dmimtiom tm& th# »eh#m6 as noted by vaa

Hi#! is that th© ratios «f prooioni® t© ae«tie aeii are lai»g©F

tliais g, varying witii tli# different stains »nd ©ften with tha

saas strain oa dlif&pmt fm&st Mt^s@t »«<ii.|i« fliia a^sa® favor­

ing of pmpl^ula aeid was m0%«d in tli# pyravio aoid feraienta*

ti©it# It it #vi(l#»t that the eiJ»l)on aoatent of the sugar it

f«eov®r®t iB tli« tlirt® f#r«#at»ti©a prodmets pi'opienie. aoid,

aettid »ad tet ®ji ®3i®®8« ©f ^dpogea is present in

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Th® sttcclnle sold df gl»eoi© is pi*#pos#4

Tan li«l' to origltt&ta froa eoiapomiifi»':®f th# fme% escti'get

i^loli, ai*© aetlag m luriwfsn aae^ptofs,#, - It »®®ms p0»slb3.|'

t©, mntrntB ttiat -smesHiiit- A«ld Is foMsd iTtm ©<»^

poim^a of tiJ® by sn aeseptane® ©f Jsi-dPogta wli#a th@

pmmnm ©f redualW© grompg mm cauBing &%

th® smm tim# tli© ©pposlt® mnmmlm n&mXf ioiwitloii &t

.. It h& ©atptettd tliat, a®«fc' of th©

toiy4f0g#a womld la® acc®f%@d % th% m®v group® ^

and thus ©aws# th© yi#l^ ®f to l>« swell,, m thm.

QontT&vf th« fitld is larg®r In th® glmc^aa fewnsfttatlon.' than

1» til® laette. moid fo»#iitati#»#. •

In 'te-otli IsotlQ ®eld and tli© glmmm f«m@iitatloiia ^

pp©pl0ttt© tli® i^ield ©f «e®tle a©M was InemmmA ,

thTQtA^ i6®^«tiioil.ii • fhs #:^.g®a mt tin® itif fipp##ps to lia^e aoted

as a inyd^^gea

, 'UriAtae®, fci* th« iiit»m0diat# stages of th# gluooa^ tereiJi*

<loisa|,^ ai»® not »miiso3?t fm of

tht glTiQos# 1« ©ff«p©4 t*r f4^%»W1B*S .*©1^1*

fh8 ftrntiitittiQia of, »%»«}! •% fyQB>iQiiiba#teyit»

.iptittlts ,gimilai» t®. .%ii® glmeetf

fli® atttdf ::0f th# .f@»ant®tion of glfmml toy fttn llel

e©nfiw#d the ®aylj woFk'Of that proplonle aeid

is.th® ealF pmAmt of fewieiiitation* .fsti l?l©t fh&t

th« gli-ees-ol is d®h-rilr©g@ftatei.''to tlfs®'?©! aldehyd# which .a«stt

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Sagar disaBT?0a.red lu nilXiMois 6gHgOg

Total aaoTffit of Tola-tile acids tft silllaols

lilsta'blishefi ratio propionicJacetic acid

Total smmmt of pionic acid in ailliaols

pro-'

fst^l eraoijnt ©f scetic aeii in millisol-s

Quantity of acetic acid fresm yeast extract in aillimols Moimt of acetic acid frm Bu$mt ia millimols

Modified ratio prop-•lasie asMiee^stie .seid : twm m§B.T

< m o ® et «t .et

•f •f m MM 0, O p .et » et m » 'H> 3 ,J3* et Ct» o

© © Mct0 »% et- ® 13 1* ® p P* » 13 3 0 et » : IP © » m & **> •y 0- •et Hk' 1 © h* €> 16 1 ^ w • "fufc 1 1 m CbU U m . SS: 1 m p-sa s I* «f * «t e 1- ? » » »i «t M O m aw ct o Sf i

m pf ®. ?9 #«*»•

» c® ® s» m »* s ct *

^ l g m. «

P et ® M o * ® p»ct s- ft* » J* i & 0- ® m © o < «!E| 1 m m m o H- at m : »a i » ® Q: <m m .IS a 4: 9 m m 0. •f 3 €#» m ft •

» * • SO €tp O ' M » .® *»»

® 3 m et © •©. CJh sr g O • |-» a H* m If 0 ® ct ©'•O^ §• 1 tr M» Ct SS m i «t 1 » »*• O .* 3 p- < § ct •O 9 p-

§ *%SS et IX •1 s ® ••i O fit®

• 3 ® » ct ••i O fit® « m 3 8 r M «t. 2 3

» » C» »• 2 4 § GS ct St © «a ® p P § 1 P* f* -w i» m 0> 1 ® St & ® » w* o 01 Ct

^ « 1* *-•• .l-i 3 P 1 < » -» 'S O" 5 1 # a l£ m •» a © S et (9 o o ® Met S c o m a ® O #

3 i er a M et r ct ® *i M » «t r •w ? 9 ® tf • ? o ® • M m ct O. • 1

t m la •

tad 'toifitil® aolto wlileli mmMiBt @M®ay of

if@l4# In muBMemtlon ©f thss© r@swlt@ iram- li®l

that til® aeid, ia #«#s$ of'•that ©bt®lniibl® ffo» th®

mg&T femmt&A wm m&M pm4umd titm th® y-east amt©*

lysat® • m&. li» tppli«d a mitmQtim* I# thus ©b%ftlii#d the vetj,

fftvormhi® i*«ti©s showi la th« table#

fh© qttestion iM»#il®t©l.? pi»#sefitt lts«lf| if'it 1® ii©e©s-

•siify t© intfttdwiegf,« ©©ppeetion h©ip® fox* volatll© «oid i>i»odtio®d

fr©m th® yeast ttutolyimt#, should not a similar eoweotioii b#

ftpi>li«d to *11 f«ra@iitatloiia in whioh y#ast mtrmt was us@df ,.

T®»ist w»t#f (li4 ©f wst#!*) was ut®€ in f«m#nt«tioiis tv®m

whi-eh earbon balan©#® wwe »»€©• A aiailar aorreetioa in

th»s# te«lane#8 wouM l®av# am® sngmw miteeotanted for fey fel-

atil® aeiii* It ir®uld them seem possible thut th® suecinic

Mid foiind f®w»ntfttioa« aay have been fowed directly

froa th© sufiar as propositi by fiitanen#

i. point of iat«?0st in this oonntotion whioh has appai-ently

b®«n owi»*loolr®d fsa Jl#l i# that at th© ©onO:©ijtr«tlon of

y®ast ©aftipaet Immmses -.mt only will th® hyi*»og#n donating

toMpounis of th« yaaat inertai© tout so will also th« hydj*©g@n

aeoepting ©oapot^di., Aooording to Van il®l«s thaory on® shouM

also in aueh a ®»s© expmt an. inoraai® in the yiald of suoeini®

aeift# mfof^tunatoly tha tatawaination® of suooinio aoid wera

oaittad by van Nial in his stmdias involving the nitrogan

matabolisa of thaa« baoteria and this infoiwation is not avail-

abla ffo« hia data#

-fills eQftald«*»8iti@a Idtttoe §©» as to tii© ttooui'iioi-

of faa Mlol»t eonoluiloas, aitioo th®f m^rn tensei upon fowonfca-

fcioBs in liiioh. th® iiiti»oi®» a«teri«l pl&fmA a predominant rol®.-

Ifc is iM®#4 aoufetfttl 'whe-teMr th® hii^i ratios of propioaio, to

a®«ti@ metdB mre •utiFoly th® f«s«3tt of aoRfttiow of hf&rogen

tel- oo»f>omii4s#

Wilion, Wm& C1930) woiPfelmg with II trP®a

of 'propiottio aold baot^riii wMoh sr® not ideiitifiod as. to

S'pwiea, invostigiitod feh# .f#»i@iita.tiott of glwoos®,. Isotoso,

smmMe, i»«l of iiftltod molasio® m& l^drolyzot

slartfe. With par#- ©altttrts ti» diffsrtut sugars w®i»@. fotuad to

oalr partly d0#tr©r»d.# 4a. tb@ -prosono® of x>&ototoaeillm»

oftaoi tli«a@ sw.gitrs wor® f«»©iit©a ^ alsost ©oMplottli-.# with botli.

pur® and ai3»d mltmmB tli» tpsnititj of tho f@«a©»t®d sugar

rtoowro'd m vol&til© aolAs if.®.® lowtr thm that m»

po'rtod C25-TS f#r oontK

i®l«ot«<l typos gro-wn assoeiatiwolf with l^aetofoaoillus

o«yaj@i f©3»m©iitod aoal of amitei oosfi., molaiso®., .laiA hi'drolj^aod

8t«reli» fh® »0l&«s#s and hydrol^'io-d ataroM w#r« quiofely .for*

{90 -por ooat la S €«ya-..)..-ifeil# .noa-wifom roswlta wor®

oMaiao-i with growi# aajtot oo-ni. loJ.ii««o» was foiand'to.'.to® • .

fomomtod without- additiOMl nitrogon ooatslnijii m&torial*

fl» dis-tiXlatian r««ito# frw aootoao^^trntyl •aloohol ..femonta-

tioms- sorfodl a® m nitrogoa aotiroo*

From this w©r1c 1% is ©vident that th« suRsr f@rat©nt»tlons

her#, studied did Eot yl«ld ireXttil© aelds and

dOg 93 in ¥att. liitl*a f«f»«ata%ioiii »ttd %h® stimilatiiif astlon

©f assoeiitt©<l teit.©t#fia wm <pit# evident though I'east

®3CtPaet was 8©»t douht eoaeerning th® ©Imia,

of that thi# stiMlfttioB i® tea.onlj to th® toreakdowa

©f th© iiit3»i©n matfPi&l to t. a©i*® ittlttlii® fom»

fo0t«i, mA |li30) mlng 20 etilfawa ©f

pj?opiowi0 «©id fes©t«i?ia d«t«3P»4»©t their aMlity to f®m©at

araMnts® aiid styles©* fhirt««n #ttltiii?#s wer® fomnd to fement,.

iii*«.bin©if, «d mlf ©a® ©f th«a© t©' t&wmm% jsrlos®# Of th® 13.

ftraMnos# two ^«ltmr«® w#r© «ip«oiall^

vig®i?oii» in their attaskf ©»# of th#s« two 0talt«r®8 ^feting the

3^1®®# ieimm%»Tn ^•aatit»ti'¥e studies ii®r« aad© of .th® fer*

meiitiition prodwets ,of firatoiiios® and .xylos®! of gltieoa#

for th« pmrpo«« of ««faris@ii.» tn the foment at ion

ttiittg ytatt «xtraot th« ratio ®f propioiiio to ae«tlo moid was

t® ¥ary fTm. t® 8.»0« , fh« irolittil# atidt mfvm.%®^ to ,

from iO-fi.for ®®nt of %h® .fmgur#' fht p©rs®at,ag«, of the sugar

«t@. aoat-'^ol.'ittil.o a.eid8 w«.t thi® that otet«i»«d bj Tan Hlfli •

It oonsisttd ohitflr of siae,oiiil6^««lid Mn& in eortmin ©.sisoa

aaouiittd to as mm% a.» gi.#.® ptr ®«st. of th# sut-gar*

fhe f®»»ntfttioii of th# poato.s.os ,.fti mmpmt^A with glums®

yit-ldod ft lowar saowit^of volstll® aoids'with ft eorresponding

inor#»@« iii nom^f^olatil© acids., fh» riitio of propionio to

tootie aeids d#ors»t®d»

On® eultmr# (2.11 4t4 not show & tfpiml feraeatatloa ©f

gl«.i?ds® &T ia media aot eoatsiiaing ClftS0g» a eomaid*

®f 3.tt0tie asi^ wa® fowwA aad an mmad0ount#d fo#

#«p©iiii4 ft»ain»d ia aft#*' all otltay pi-odtieta bad

hmn hf 4istill»tioii md ©atyaotion#.

: •• i© »«®liaiiism is smgg®st#d toy tii® aiatliopt foi* th® ;lJi»«ak«

ifewa.of th® fii« appears to.'var|r ©©n-

sidtrabll* with .diffeMsat oultujp#®*' fhm gludsa# fementationr.

in ag!r#«@at with %h& py-upesal ©f firfcanen, points t© a 4- and

2» ©aytea tei?#«k<io«ria th® BUg&r* fh® i?atios of pmpioni.^ t© ^

.aQ#tie' aeida atiA asatie aeit to OOg a«®i*aas@ witli. ineraas# in

th# fi»o€tt©ti©m ©f sii«sei»ie aeid a» wo«M fe© aaspaeted if. tto©

mittmn Qmpmm.4,- ii©at«lda%4®, w^m emwertM. to aeeti© aeii#

werkaaw and laadall (ItSl) a e&minl »tm% ©f 32

mltnmB ®f propioaie a©iA ^att®i"ia aooaiit tli# nam® .ffropioni.-

toa©t®yitiai m mlM foir tli« pums# win© si^oias

m&" iat0rib#i,' &m ®f irlii©li g» yaffinoaaeeiaa# is ii©w« fhe

mtikom 'point o«t • ; ditatfantage-a of i®.paf»tion. of th©

ap@ei®« of fyp^i©ni^aet©rium toy 'Miorpliologioal ©liaraetsriatios#

A' aiola©t©}a©«8 lc©r t© til© pi»0,pi©ni® m@M M©t@ria b»s@A upon

^hj-iioioRieal charaet^risties is pr-aaanted#

firtanon -ani Kej«ati»liii {1930) fmrth©x' stApp©t»t th® ©arlj

work ©f fiFtaaan and eritioiz® tli® jp©sult# of van »i®l«' fhey

itat© • that sine© tli© propionie feastaria pFodit©© fyoat papton©

and aspaeialli" fmB% watar aipiifieant aMwints ©f feraanta*

ti®n pi»d«i©ta, it ia ©asr t© m^vlrm at ine©rr®©t ©onclusions

» ^ v . l | a S | 3 i 5 M § g ii © ® ct- « ^ Hf m i a 0 m * $ • • & & m ^ < m m ^ t* m • m & H ^ ® m -m © •«* & # © ; • . 0

g o a t ? » ' i s s § «i ® »* ® » n- g ©• *i.

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letrge moimte of stKSoinio mA furaario aelds#

fymwie mid we® •fmmd.to to® femetited l>jr the living

glS-eegsn-fr®© ^ teaeter*l». and also by the dry eells. In th© lfttt©i»

eas« only in th« pFtteaee of phos^et®. As f®3?m©Bt®«'

tion pwnlxaets prepicitii®* aeatie amd siieeinie aeids, as,, well as

G.Og w«re fome4» itmtliow point out tfe»t wlu@n sm®eini'@

ft«i€t ia. foimed trm &&i<i tfe# ffruvie aeid mat first

eond@nse to. a six ^mrh&n whleh .is tlttn split into 2* mnfi

4- eai*feoa eeap^iBiia,#- fit» m&hmim of th« glwe©s® &r#akioim

is %he sa«® as pjfs^iaw.ilj glv#fi» QotiflMtitloB of th© .ooemi?»

mnm of ti«x©»«»mon6iilia!.spMt# was seeemplisheid hf isolation as-

th® bftriiaa salt»

W9rk»«i and flf33) ©stftljlishtd de.finittly th© syst®®*

atid states of th« p'ropioaie aeid baeteria* By means of a

s@i'Ol©gi0.al elftstifioatiott, they cl^ftrly 0onfli»e€ elasslfi*

eatioas .b««®a .®ii..#tli®? auA t©i6©.ii®trated that th©s.@

M.0t«!»la •©©Estitwt® ft .Hfitmral gi'Otip* two prmimBlf nnd@»

soribei. sp@«>i@s, isolat@<l by litolmer Cl§3g|.,. fro'pioiiib.RGt#ritm

ztft#. aii.4 gyopiomibaeteyiaa' arftbiaosm, *@s^ ©harsete'rig©^*

The int«F®ttiiig f®at mm «8tiibli®li®d that p, ®yBtMjae.s.taai. .i.®

«:fttala@© ii«git%tir« tte« M®®ssitating laodifieatiOR of th©

g©Bfi»ic aiagno«ia# dlffertatiation iudieattd that

oertsia apesits fall into thp®» s«b^a@rlc gpomps. ii.key ie

giftrn fQT th© ®l0f©ii »p©ei«a of th© genu®.*

Fi?osa.g«ot and fetw» {ItaS) report @*p«yl«i0fital Fstult®

i#iioh--tta@y suggest th® n^etasity of a %®xokiiia.seoi»

-ai-

aetiTstor for proplonle aeid. fenaentsfi'bioii <sf gluoos©,. two

«n4#80rlto©d eul.%ui»@s w©s»e-wse<l in this ,stu%| on® dultur©, So.

2-gj W68 t :stareh fe»ef5it#F« the netl^ato? wa® t®«oiistPat;@fi in,

pi^oteln-fiNi# ey^wmtB of' of p®as» fh® teasal in@diw

eoRslst«d of m eom »asli» It was feund that in

©,. mm meiitm pdtato #3ctjp®®t was iadigfeuslbl#

for th@ growth of u-tt'ittJJ*# 11 aad »®e#l#2*ftt®A (Imt was not es­

sential for) the gTOwth of -CHltwre 12, I» no ©»i© did th#

potato o? mxtv-mt support th® gx-owth ©f th# propionie aoiA,

baoterls in th« •ahs'iraie© of o©m-itt®«h» fh®8# ©xtyaets w@re not,

neeessary for th# titillE,«tioE of laetli® fh« restAlts &r&

interpreted sa shorn in th« seheaes#

Hultmr©. IS .tttltUF© 11 StaMllK®€ gluoos®

!• I »alto«t 4eti¥Rtor

•i i I»aMl® gl»0®s« -« l<&@ti© aoit

I i j i St&Mlli®d fol«tti« aeid

'''.Meti| s6ia^32tlw|o!L

•f olatil#. atid

A "hexekinaa#* it .tuggaittd as n«@»iaa.i»|- for th© activation, ©f

•aiS0O'li'^#d or ttaMliied glueogs' whll# labile gltjoose, proposed

to oTigiaat® froa. hfdpolraoi starish in f^Tmnted withotit •aeti«»

•Vfttion •(eultiare !§)• fhe f«et' that th© extracts er© not.in-

dispensibls to tha wtiliaation of laetio aoia indloatod they

ar« not nemamfj In. gltieos® ferraontation. .heoause ©f th#ir

nmt.ritionftl eonstitmeats. fh# ©Jttr&ots when «»h@d show#a ®om®

' iil

fswratol# laflmea©©* fh® aetl-^atlon was pvlm&Tilf <3ta© to th©

c5i»gani« 0oaatitm®nts ©f tlie ©xtrasta, hmmeTm

fwo addltloaal int«i«0«tlag feots. wero @st«to3lish.©a with

sultiii?# 12# In nash-oaleiiBa laetate- aodltaa there was no

i#9r©as# In th# quantity ©f ls,stic noli tmt on the eontrarf ait

Xmm'Bm was rioted, th# aeld feeing tovme^A hf the feraitntstioa

<jf 'BtsLmUrn Xn mm maBh or earn mmah pim laetls aeia &p

gXutSfO§0 mn in&mme in th« 2»<i<Sttelng po-w&r of th@ mBdlim was

astahllsh^d di»iag th# fermentatloa, aM a«id h^'drolysla l®d

to^''fti3Pth«r liies»«'ast« this ir«.a i«t0i*pr©ted m Axe %&• th@ iomrn*

tiou' of glme®«© aM aaXtos# tmm »t»r®h thyou^i th® eaa^olyti®

actloii ©f e«ltti3?« • 12#

It is ©f int«r®»t in thit- mmimtton that miPkM&n ant

l#nAa3.1 C1930| motsd that v«getahl® entm&t-M stlaulat® gp©wth

of tha pTOfleai© aeid bstttpia#

?®tt mnd Wfnmm |3.i33^) iiiif®stigst«<S th® phosphatss©

af&t&m ©f giloitytaitiitt . a##t-6teTityl.i®ti3i and gropiaalto&eteyitim

.1@iii«nii using so#l«ra phosphate, aoditiBi h©xos©diphos«

phate aM py^epho^iphttt® as suhftretes# In asking th@

t®st 0«1 haotdflft'wsa mmd p®r 10 e©# of «edl«m and

th@ iaoftast 1» ime^gaai© ph©sph«t® was dtt«miB0d» the

phOiphatas® of the €Fl©d h«et#i*ia wms fotiM to he quit© atabl«#

la @1% months ia, a deiiQefttof at Tom t®fflp®j*stiir@ about om»

hftlf of th© aetlvitr -ilsa^peitred. Various p3P0|(®rtt©s and p®*

l«tioa»hif» wm0 ©sts.hl4#h0d for th® mzymmu* fh© followinf^

w«i?# d®te»ih#d with fyopioaihaateyjiia jmrnniit, fh® pi

optimum for th# hydrolysis of cc-o? /r-glyesrol^ospHat© was

for sodium hexosedlphdsphafc© 6«0, for pypophoipihat© 7,0,'

Of tfe® Ions l*iir®etlgatt»a mapiealua was fch© only Ion

stlaalat«a «ll ©f th© thi-©® phmphB%0 systeas, fh©

optlwttM oenoentratiea, 0«0Q3M,. wm th© sea# for th© thP®e

sy«t®ait 2iBe • aetlfftt©d th© p3r^ophespli.at«®# ot:li®r

ion® •©ithe.f liad no e f t r n t of i*#tara©4 tli# hfdroljrits. fh«

a|>®«d of r«fe©tioii wm fowi.4 |)yopo:rti0n.sl to the ©oncentretion

©f • the pho$]^.et,its# toluem toad n© mttm% m th® rwotioii.

fSi©. siiftiifioanea' of -..til#®® . e-rS' €i»©ttei®d la i^ela* /

tien. t© th# quastion 6f .s.pselfieiti-^ ©f aetl*?ati@35t

mnA th© .©-©tiwi-ale n&^mm. of tn# l©n,, attd also with

r©gaM to tli« gpeeifioiti" ©f mmfmm*

. f®tt atii Wftm^ Cl.t34j f&tm& tliat th# li'biiWtioii'of' phes'-

phat© froa h»j:m«phmph%tm and .glyeereipliosphat® with dry

pr&paffitlon of the propl©iii0:« was &ee«l#fat©i by ans^net© and

fti».foiitfe© Thajr s«gipst tii« aoetlereting ©ff©ct-' la .to® to

gtlsiulatloa 'Of the a|^it©» ytsponslbl# fov th@ depyafift*'

tion- ©f^'tlie oipfenie part &f %h% #st®r, rather than to th® stlia*

lalsttoB. of the •phoaphfttm®©*

p©tt Slid Wpm# (a.i33|,) i4eEtifi#d M&thylgXfQXBX .©btainet

as. th® 2, 4- •idittltre-'phtEyliayarozon® from feraentations by

an®# propioml© aold b&eterla with msgnsslwis h®35os®di{5hosph&t@

as a gubstrete# Slialltr stt@apts't© identify Interiaediat®

produete from 3«gly©erol phQsph&te wer© not d«flnlt» ©Ithomgh

pr«ll»iiiary eft<i©R®« was obt<iin©d for th# foraatlon of

methylglfoxal and of glyoerol aldehyde ^or aihydrojtysoeton©)» -

Sufficient material was not svallabl# •howe¥©r to «nshle ©on-

olusiv# identifloation*

wood and W«rk«an (studied th© raetatoolism of

fro.oiottibAetegitim arsMitostm.. fh®y, (1934^) -from repeated

analysis diiriii!^ th@ oourse of «luoos» femantstlons eonstTOctad

earbon b®l«no0s and demonstrstod for th© first tlin© in pro-

plonlo aold fermentation th© formstlon and doatmiotlon of a

non*r®dmolng ccaapound." fh© Intorraodlato broakdown of pyruvic

aold OS proposed by van WI®1 r©o®iv©d substantial stapport,

throu!^ their (1934|5) isolation of this aold using sulphite aa,

& fistativ®. in glueoa® f®ra#ntfttions, ivideno© rolatl'u^© to tho

Intormodist® aeohanlsa of,th® foraatlon of propionic aold was

also obtalnsd by the isolation of propionaldohyd© (1934^) frosi

glycerol fermentetiona uwlng a aimllar teohnlo* Those.result#

together with thoi# obtslntd with other apeoles of proplonl®

aold baoterle. will b® dlsou®s®d in detail in the oxperlTOntal

sootlon of this thoals*

Suamary of titorftturo Oltod

Claaalfloatlon and nomenolatur© of the propionic aold

bmoterla app@ar to bo in satisfaotory oonditlon and thor© is

general Rgr#®m«nt among th® different invoatlgators# Phy8lolo,gf

of thoso b&Otorlft roquiros mora study., particularly with regard

to th# moohanlsia of glmcos® disslailatlon# fh® schemos which

hav® boon proposod do not satisfaotorily fulfill th® ro-qulro- ' •

-45-^ •

mesits of ail tli© data wMeh hmB b#an i«©oo2»dtci in th©

turn, ' im tfele cermeetioa partlaulayif the aod® of fomatlss

of succlttio eald reqmlrsa f^irthar %m»s%ig&tlon, TI10 .tery

thopotigtt work of.f&a ii«l oa^fch® ©na-haM points to tii© faot

that proplottto acid, saetic aoid, and O'Og ai?® the only ppodiaet®,

of fementation of , glM0oa0* fli© eonvinolng work of flrtanen,

and others, ©b th& otU&r hand indieategi tliet aiicelnie aoid is

a f©mentation product of gltioos©. In comeotion with van

li©l*s results it ftppenrs mther iinjtastifi©<j to ass^ra© that

mQ&inlQ R0id is th# oal|- px»o&iot: reooTaiPea in tht analysis

whieh^eomes from th© nitJ'ogenoiis ociftitituents of the •medirat#

:fhl0 is ©spesifillr so tin©# s©ir»fsl .workers report oth«p pro-

duetss from ftast wafctr anfl p®ptott#« firtanen*® sohem® oontaias

m-mflMnt fl«w-ia that th® two-earbon eoapound, aoeteldehy^'

(ei» its, • eo.ftvtrsion pnofltiet) hm not |»eii fowsid &» & profeot of

f«»eftt»,tlo»# If til® ©.e®tald@ii|'a© i6"©xidi2©d t.€>'•aettie. aeii

.«ts • s#,©its probmbl® n' aorrt.^poniiEg redmotlon produet amst

.fouwfi* fhis ro^duotiom pt»o<lu<st would probably "b© prooionie

eoid,# , Asgualrifi such a QomBmlon th^ ratios propionle aeld to

©.eetio 'aeid amdi GO.g te aeetic aeld wouli tee smaller than those

of f#m©ntmtion§ not, involving aiach a eonve-rsion* Van Il«l ®imI

firtanen report th# ratio' GOg to aettic aoid as always.--equal

to !• Foot®, 'Wm^- f&t&mou report som© fementations fora*

ing 9-«eelni<5 mt& in whioh. this ratio was. less than, 1. fheir

results ar« pr®fi&iit®d as aTeragea of stveral fenaentations end

Binm th® (pantiti#® o,f @»oh prodmst i^ary oonsidtrehly for

^44**

faraantatioas it is- imposalM^ to r«aeh deflnlt©

aottci:lii»l0iis fi?oa th$ia» ?ir-taii©ii lias aot siippllsdl any' data in

til© %& smp^QT% Ms atat«a#nt to the quantltf

of SOg,'in faot h.% i3tat®a -(1923) tliat the Q©2 ^©teminatioii

waa not qTiantltatl^©# With ip«gai»3 to the yatlo of proploni©

aold .to aeetie- aeld a littraturs indlastes - It

smf fe© ©qiisl t©|, lass tta.,aa, or gftattf tha.a S» It .lits

pointed Qiit that the .©xplsnatlon of llel, mreposliag

that ratios gPtttsr t!iiiit t*o are the i-egttlt of hydrogen, dona­

tion bi- til© y#ast mmpoimdBf is .not adeqttttttly s«.-ppoi*t©d. "by

.his @xp#.rim«.iits# fhe i»stio« less tH-aa two saggtst a 4-» mA S-

• ciiyboii eldtmg# of the gliaeoss aoleeml® wltli the fernetloni of .

©a®, ml#awle of stise-iBis told, one ,»iol©o«l« of tcetle'aoid and ,

tw©.' fttoss of -lif#Pog#ii wlilc?h. %n a ye-dtietiOB pfobablj' to

'fom. proploBle a-sld# It Is •appfi.rent tl»«t ftis»tli®i' Iwestlga-

tion will M: mc^XmA to estsblish tii« sel-tttion ©f this pi?otol«a»

0©iis'ld-®iPtbla ittforaatiott has b-®«-n o-fetsia®^ oone^ra-ing

pcissibl® liitewiediat® ppodusts sinet th® piibllo.8.tloii of ir.»ii

Ilel*3. aist©l'tati«9a+ f!i® following have i3-«®m detected, and

Ideiitlfi^ds-

1,. Monohoxoaeohospltat® i« Lactic acid 3» pyruvic acid 4« pyoi>lon8ia#li|'d«

5,, Methylojlyoxtl

fhese r@gttlts need to b© lnt®s»pp@%@a Itt a ®ingl#, achea©

of tevrnrntrntlmm

I sttxdi* attenptlag to copfalat# the faots es-batolishod

coa3@raing the gtliwilatini aetion of ssa©oiatlve bftCtePlal

growthj gyowth aeUiirato?s, %t3colclaase® aM tho mltrogtia i-®-

qtil?«a®iit§ sf'IH® pr05il0ttia« is siigg0st#d hf t>i0 llte'ralui?#

I'aifiew* "Bils phase ©f the pm'bl^m tiaa not bten attidltd sx-

t@ssi7®ly hf It .is hoped, that aysoh an In-festigit*

t.lofi msf hB made .In tb© .fiittip© ^

*4§»

II

Slimp%«T Qm

Methodg

Section 1* Cons ions,

fhm pvlmmr^ pwppos® of tbif iiiv#®tigfttion has to

ftt«niife imfo»»ti©i» m»mvmtng th,® «©eiiani»m ©f gluoos® di»-

slailation and to d®FiT# # s^ea® t© r«pr«s«nt this ateohanisa.

f® a®e«pl.lili tmeh a pitti»p©s^ it is #«s®iitial to hav® not onlf ,

aeturat® Imowltd®® ®f tli« ^aatiti#a of prudmeta @f far-

»entati©tt femt it' is als© naeaasai^ t© lia¥® infemation e©no©ra«»

ing til# intamadiat© eoapotiuds wtiioli ar® iwolvad in tk@ staps

laading t©. tHa®® final ©osfersion produeta* In obtaining stieh

infemation fixation »#tlj©ds ha¥# feeen aaployad# Pui'th®!' avi-

danaa-li«.a-TO'twltad .by lnt@»lttent anais-sis of single' f«i*Hi®nta-»

%%mM thma daaionstrating an inoraas® and subsaqaant daofaasa

ill @ai*tftin pyadmats^-* fariation ©f th# eanditions of fairoanta-'

tion has lad t# an aaotarettlation easpoMnds whie^; nnus^lly do

not @aaw;j» aa and-pyoi&ots* fha «,p#eias ®f tha ganus show a

^irariation in fsataltlii® thus giving inforaation ishioh it help­

ful in fowttilating a lehaaa#

^sstiaing ©na has a knowlad^ of th© intawiadiata ooapounda.

Wm K'lwologlit stttdfing M®shanls»i throu:^ s 4««

t«wilfiatloii ©f the ©lid prodmets i« yet ©onfrouted with th®

pr©hl#a 0f Judging th« #f his anftlys®®# fh© balaaa®

0f th@ ©f th® iiibstrat® f®f®©iit®d and that'ia th© d®-

t®raiis@d pi»0d«©t« giir@s on® lapertant iJh®®k ©a th® d#t®rmina-

ti©iiQ« fh® lia"«r«stl,g®tor of moMmhtfn f®»®nt®tioa also ha®

airallslil® m i®«0iid eh®slr In the foi* ©f m oxidation and r@-

diiotlott halaii®® (jr0h»»®n» f«t®r«©ia ®iQd fred ©oMpotadiB

®«eh as gitae®#® liiish mntsdn and ©ay^s iu th© »fim®

ratio m wst®r, giv® h|- this halaao® th® • simt iffioimt af r®du©®d

m ©xidizad |»rodti®ti pr©¥idlag e®ap®miids other th®» thos® ®ris*

lag from th® d«®o®p6sitioii #f th® ar® not eetlng as

hfdr0g®n' ®0o«ptors ®r ddttatora# Th«a® -oheek® ®n the aocuracy

©f analysis: hair® b®©a m®®d soiittstently threti^out thi# in-

wstlgmtion as s has!# of Judging th® reliability of data to •

b® ms@d' in d«t«»lni»g th® f®r®®ntati6n s0h®«®«

S®etioa S# Kethods of m&lf9l9»

§mmm pr®i».»tg8* Qaalitativ© ^anftiys®® ®f pr®li«lnary^'

f«ra@iit®tiopa «®iifl»ed th® obsewation ®f f0ra«r inirestigators

that th® gas prodattd'by propioaie' aeid baet@ria ©onsiets ex*

@ltjsit®ly of carb©!! dloxid®,# th® qwaiitltatl'?® d®temination

®f th® S^lHi®rat®d d^riiig f®mentation, a® well as that o^f

th® r®tldmal G©g ®f th® ffl»di\iaij| was ao©©i^Il®h®d by absorbing

th®' gas in l©w®n potaah buiba ®®nt«lnlng 4S par ®®nt 101, or

in 'U-tubss -©ontalning soda-lla© (SO p«r ®«nt taOE and 50 p®r

;*4%»

0®nt ill trains we^® ari»»iig«a *ltli s«lt«lsle C?a0lg

.dt^inf tjr-tmb«s«. fw© lowtn Imltes, six tmh %ub@a fill®4

witli ioda-lim# w@m ii8@d, wltit. ®«eti loaf i>h#ii9lp!ithiilei»

was. %@ tli« slteall ®f th® B©w@ii temltos.aa^ r©plft©«ii®iits

m&m ffl&i© tli® iRdieator feetam# y©A#

; la ^o»0 f®iwi»talioii# in whiiaM ItXQig. was us®€ ai a

tomff#f'^,.l;lj0\. QOg @f til® a®<ii«a te#f©y© and »ft®r f©m®iitafcioii

was i#temi»#d In &Mqm% pwtioat of th® a@diaii» •lU# 00^

wa®. llfe®i?ftt#d toy asliif^img tTtt© to ©eng© feA papef with. .

HgSO^ and Tmrnovisl wa®. aeeomplishai. fef boiling and

aerating tli® ll{|tiid under « refliix eoaa#a»®r» f#meritationa

In whlBh OaOOg «« tt.ea a. . »mrf« reqalMd « .odlfled teolml*

foy d@%@wifi«ti©a ©f th# fesidual OOg feseausa of th® diffieulty^

Qf «»#pif###iitatit» samplts* X«i tbas#-f©»®ntatio.as

til® OaO'ig was ir«i#i®i 'Oiit s®#tirat#ly^ ott a gi'atimatPi© balanea,

pla®®A in, tha f®HM,ntatioa flask with dlatillad water aiid

atariliaed at 20 lto«, praaattpa for om*h&lf liour. i.ft©r addi­

tion of it«rilii®d eonstitwaats ®f tlia madiuia and

til#/ iuootiliia, t«ii oe# of the supamataiit liquid w®'r© ramo^ad

witli a itarila pipatta for sugar anali-ils, oar# Isting taken

aot to raaov# m a ras-wlt of stlrriag# Tha original GO^

of th® meditja was oalsulatad acpi'ralant to tha waited quantity

of 0a0©g« At th® oonoluaion.of the fanasntation 25 oe* of th®

supairaatant liquid war# raaovad. for stagar dat«rBiinatlo,ns and

for msa iii f'ariiying th# piiritj ©f tba owlt^ra* fha fawanta*-^

tion flask' was %hmn attaoliad to a ra^wr oondanaar ^i.Qh lad

%© & A. irolmi© 0t ii IgSO^ sXiglitly in

mMB of %h&t mmMw&Tf te r#»©t with %li# eaCOg wa® «M®(1 slowlf

tti3?oii#i a, dropping A k1©w mrmn% of CQg-ff®® air wa«

fass«d throu^ ttie li^td# Aft#r »pprejriaat©3.f ©n® hour a lour

h«at WHS applied to th® fl».sk and the mn%mt& w®re bro-ugit to •

a, boil in ttppfoxlaatolr ©n® ftourt fh# litfttiiiR and a«ration

fftr# ®oiitinti«d f©r an additional hour «M tii© liberitted. 0Og

wa« d®t®»im»d grtt¥i»@tri®all|r# fh© residwal 00^ of th® aedltm

plui th© OOg oolleoted aurlng fe™«ntatlon «lnu. th. OOg .ddea

fti 0*€©g th® 00^ fr©dtt#©d'toy th® f©OT@ntfttloii» D@t@r» •'

minations miing kmmi qiitntities of sterilised GaQOg proved

that'th® m#th©d'gav# aeouriit® 'reamlts.

MmW»l. yoltttil# piroduetfr* fh.® f#ra®iit#d llqwor was,

«@ldlfi®d with, %§©4, aad ft V:Olm® ©f dlstlllat© ©qtii^alent to

on@»h®lf thf ©f tha ©rigin#! liquid mm ®oll®0t®d msing,

».a i@®d eo.tid«ii-i<©r# Aeid .di,stillatl©n was fownd n®e©ssary in

erder t© ,a-fold, the deeoaposltlon lAieh QttQn results-• when

sugars or other ©©njstitTiients .-of a hflet®rlol©e:ie»l mediiAa are

h@at„«d in &n alkalin® soliitian* fh# distillate was natatrali2«i,

t© litTOS with laOI and r#dlstlll@d t© oua-half volme# fh®

raiidmas @f th® distillations war#- eomhlnad for iis® in the

Tolatil® a©id datawitiatloii »hd th« distillate was tested for ,

v©l^tila ale©h©ls, aldah^das ahd ketenas. fha ©nidation mathod

'•pf Stahly* Oabmrh a»d W^rkmm {lfS4) wa« uiad for th® quanti-

tativa d«%ar»iuati©h of th® aleohols. Qualitatlir® t©,8ta for

. aldahj'des arid ,k®t0fi#a w#r« mad© with 8, 4»diiiitroph©ii5'lh5-dra2in®,f

s&divm. attyopimssid® an'd by the l0d#f0m peaetlon* flies©

t®0t8 pmveA that volatll# alsohols, Bld&hf&ma aiwi l£:@%oit©s

w©i»® ttstaally n©t |>rd»©iit in eseteatfatioms whitli eould to® A©-

ftud 'Vftim dfft®®!;#!® they oesmrf^d ejaly in ti»a0@Sj. fli«

d®t&Miii®lioa of Haittpal volntll© pyeduetis was 0»itt®d ^®n

prt^iotis f©fre#nt«ti6ii® mttJf slMilar #Qaditi®ii« had given

ii@gliilM® Qma»titi®s of t}i®s© p^odnets# fh@ diatillat®# were-

t#8tj®d for a@®tjr3.-a@t.hyl*0®FM^©l by fell® 0*l®a^a Metfeed' (193l):«.

fli#y w©i»® 0m.m%Qm.%lf po»itiv«# toaly#®® of -f^wtiited a®di&

toy ••a.® «®%liod" ©f li»©elcii«]aa »mi (li33) showe-d

wis« prostnfc oaly' a^trace of ae«tyl»mtthyl-©a^Miiol and that i

th#i?© was m t>, i*bttfcylsa& gly®©l«

In'®®rt«in #3ip«i»la#at8 in wlii«h • »uifite was us«d as &

fisrative, pf^pionnldeliyd# oeou»©<i. as « fermentation prodticfc.

It wm €9'teimtm& by distilling into lalsO-^ solution, titrating

th© #«®©ss smlfit# witfe iodin© and then d©t@minlng th© b««nd

pr0pi0«aid®hyd« toy splitting tti# sldstayd® sulf it®' ooapotand tas*

ing 0jad titrating tin© lib©r«t©d sulfite# fh© titration

is tli# ©'«»« «« that. m®©d in tli© laetio aeid datewination

{F3?i®d©as!iii -and leadsll (lff9))« In %h® distillation of th©

propionaldthyd# an ©^s®ss ©f faeOg wat «dd©d to th© «®diw3a and

s distillat®' ©qmiir®l©nt to. two-*tMM« th® original voliun© of

liquid was @©ll©©t®d m«iiig an- i©©d ©ond©n««i',

folatil© »©id£e 'fh© volfttil© asids w@3P© d®t@i»lii©d by

th© a®thod of Oabmrn, Wood and fl©s%««a (1933)# An aliqmot of

•SI*

til# llqwor m %%& y©sidm@a from neutral aia.t illa­

tions mm ®eiiaifl®i te cong© mA with lg^4« eoitoentrated to

150 •«!6#*gOO ©e» m&'then^ sttM distilled* two liters csf-dis-»..

tillst® ••w#r« #oll®0t#d* fh® diatillat® w®» i»efl'ttx©d for on#-

hiilf'ko«s? to ««o^© tti® OOg aad was tlti**8t©d-mad adjusted .to

an .aeidity #f 0«03i maimg wmttf# fh.«s <|ttsntity -of

a®di«ii t© %» diitilled' in •ordim?' t© ©totain B liters of distillat®

ef app.ii?03eiBfttely ^0#0S1 aeld ttf^ngtfe teuld ,«st»ally b® estimat#d

from thd (juamtity ©f f«nii#nt«d augrnTm In rnvtmitk f«*iB«-ntation«

it w8# mmBS&Tf t.o eone#ntr«t® the at««« distlllst@» When

the a$id sti»®iigth wat to@low 0»,015t thi» was dcme hf netitr-aliz-.

iBg th# distillat© with 'StCODgt •wpoyatlftg it to th© p©qai3?©d

^•©Itm®, th«n liheMtiEg th® ^oletil® aeids with ® quantity of

i^ao^ @3ia®tlj «<piT»l«at t© th« Bi^OS)^. pjf®©ifltat®d Sa'SO^

was •r®»©'sr@d fer •dteftttt-ntioa ®nd filtfation* Ui®till®tlon ac­

cording to th® ladthod ®f ©la«t®ad;^it®te#r apd.Baden (19gt)

was w»®d as a method for eoii®®mtr«tliig, h©w@ir®r, iti©a th®

nom^lity of th® dlatillate w®8.0*01S» ©r ahov®, fh® distil­

lat® w*® 'n®wtr«lii«d with »®0S» eir®porat®d to afsproxlmattly

ISO m-0, ®«t«rat«d with moidif i®d with SgSO^ to eonp?

r®d .and st®i® diatille.d* ©n® liter of distillat®' .was aolleot«d*

• partition Method of ©sburn,. Wood and w«rkman {1933) '

is adapttd to th« d®t®rminatloii ©f thr«®. asid aiixtures of

fomiSji «©®tlQ and propionic acids and prof-idos a q^.ialitativ@'-

»@thod for th® dtteotion of oth®r acids* m acid for®i^ to

th® thr®e acid aixtttr® ii d®t®®t®d hy th® diff©r@n6® h®tw®.®n

til# 0©ii#%aiit 13^00 a»i K|^qq

Qatl@ia3.at«d> fli® diff®r®it«#s up&n aold us­

ing puff®- mmp&vmAa wer® fetmd ia all ©as-#® to b« po-sitiv® with

sueti f$mim mM» m hutyrg^, fsltyi© and laetlo' a#ids.

appilotttion ©f tills qtialitatlve ftetliod t© a-eid $olt3i-

tions 0bttiii@d tmm pwopimi& aeld l"#i?»«tttati0fis- tisnallj g»v® .

sMUtll' negative diff«p«tiG#a# fti#se «n«lj'8©s mem thueked by

til® p-ftfHbltion iMthod tising ta^fropi'l ©tliei?- ®s m -s-olirejit 1 -and

hy th@ partitt©n and ^lelgtmsr methMs of Qahfam, Brandt and

W@f^lc®aii Ci®'S4|,- ifeieli &m »4apt9A t© tli® d«t@Wftifi«ti©ii of foui'-

a«id aiJEtmimt of f«wi-e, nettle, propiomle and b«t-fi»i@ acids.

All a@-tli0ds dlitetetd m $M^®rpmt&tlon ©f me «®8n.«

lug ©f til® mg&tlm Mti&mnm ©Maia«d bf tti® qiaslltativ©

s-0h®«©. 4.fi not Imt tli0 asswptioni tte.at tli® ©na-l^^sis

is.-tofi'ttt as ©bti.iii#d ®ip-pt«rs -

fli« f0»i© ii0ld was d®tefiril'ii#d W th@ m^rmria ©lil©rid®

ffldtted @f •Attti'Imek aad I#^lii md w»b Qhe&k&d by th@

ia«i»etti*l© 0xld# a#ttiod &t mimwn, wmd wad w#rteaa«'Ui33)»

l©ji voisitil® a©ids.« Os€l0« in #x@«S8 of that moeBB&rf t© mm Iinwni! -Www. mWiiriinmwiiKDiiMi mmy mm riii»>»iii„miiiin,MMiioiirii ^

r®a©t with th© H^SO^ was- i.-dd®d to th#- i*#ai-d»i@ fro® the distil­

lation of W&e volatil® aeidn* and tli# mlsrfewf© was li#at®d m

th® st®»M festli ftr Jialf m UrnxTm fh® mlxtum was th&n filtered

and til® a@utrftl »©lttti®m was «va|)®fat®d t® a taall v©lwm®»

fh® 0o-a@®iitrat®d li«pid wsi aeidifiad to @®n©a r®d with 50 p@F

mnt hf rolwrn' mt e6ae®iiti*®t®d HgSO^ t«k®ii up with

afteip -st-aadiag at t&r ©a®

was usually dry* It was then worked to a fine powder and

©Xtra0t«d with ©thyl ®fh©r dofttltmoiiily for twelve hours, ffe®

•©th®r'was- €lstlll®d from th« eartraet • after addltiofi of a ^ small

i«6ta.nt of distilled wa%«r* fim rtsidaal solutlG-ii was diluted

to m smitfihlt la&tially 2.00 m* iliipets ©f thi® ©x-

tr*6t wtr# matd f®r the determinatloR of and laotie

eelds# fh« laetio aeid was d©t©rain©d hf th# method of fri©d®»

maim and K®itdail (1909) a® modified l©la:0« (1933) • fhis

wodifiestien. O'Ofislata in .bailing the sample at m\ alkaline pi

for S %•© 10 »iimt#s to hj-drolyaa th« laotid#., a eondensatioa

predict of Isetie aoid _«hish it aot dtt®miii©d by the um©d-

tfi®d mtthodn fh® l»<sti® m±A. in th© hjdroly«®a ®«f>le wes

d#t®r!«ia#d as- d«s@rlb#d im th« origloftl aethod# fh® Bua^inlQ

aeid was d®t«Wiiii#d as th® tsl^itm or the silter salt#' fhs

former mm «s f#ll®wsi-* m ^liquet portion of th#

©•3itraet was' notitraliiid fey h#&tiii-i it on the it«aa b«th with

aa emma of GeO%» fh® n^lMtlon after reaoval ©f the ®x©@ss

amQ-GjS bf filtrfttion,. -wiis to- a tmall volmro and th®

• ml&lvm Mmeifirnt® was pr«eipit»t<it hf additio® ©f ®hom#i 9B

per mn% alsohol t© tj-ring th© • ala©h©l ©©oeeritratloa to

'iO-8S ,p«-r-©-«»%•• Aft®-r i%«iidia-i fo-r eight homr# ia the r«frig-f

•riitor th« p-r@®ipttttt-® was filt«r©d, m€ wa-®hed with 96 p«r mn%

®l©oh!Ol« It mm -4ri®d »t 10©-'® for ©la® hotur and w©i^®d». • Thi^

method la a-ot • i.ppli@®blt t©' e-irtrsots oontaialntg phoaphates or

sulfatts* If th® ©xtraet eontaine-d @ith#r ©f th®s© th@ swo-

einie aold -wms. d©-t#wilii#d as th® -silv©r salt# fh® eartrae-t was

t© wifcli Ba{OH)g «olutl©» mnd the

•imifat® er i&osphat© was- filtered off# Stiffloieat

ailmt© nltris a^td was then ad4®d t© tke tO'aske the

reaotlsii sftd'to ^i% was fclsaa .aad» jttst alka*

lln® to wittoi dilute m «xe«ti of 10

per <5a»t ,4gl©^ a.©lution was addtd' amd after flft@®n. mitmtm

tU® .,sl4'V©r pyeelfltftt© waa fllteyitd SAd wathad# It was di?l©d

at 3.00® itiid. d©t«wiifi«d ,grs.via®t?ieallf # fii© two metbods of

d@tsmlttatlon 'alieglced olostly* fh& total aridity ©f th® «3S-

t^aet was d«tai*»liied 1j|- titration and la %'h.® «li8#.ne8 Cff phos-

plistes Of sulfates was mimalli* fmmd @qn.&l to th® uxm of. th®

dettraiiied ame0iiiio ami latti® a-©ids»

. It aijomld oQtftd glfrnml and per-h-i^s ©thep poll*-

alcohols ®F# @iEtraet«d with ©th^r* f@sts ©n. tti© «th@r #:x-

traets.foi? pslfal#®!!©!® wittot oopper smlfat© ttJid sodium liydrox*

id© weF® aot p9»lti"f» in any .f®m#iitstioa ©3:aained«,'

Ifi sttrtftin. pjrttvl® aeld m • th® O-as©^ additiaii

©ompownd ©©•®tt3?'i»«d • fts icpTOdttet of f®.«®»Jit®tloii# A namb&r of

AXftemnt methods tQT the det^mlastiom of pyi^iivio' aeld w@r0-

t®st«d and th# *i@lamd • iodufawi »©tli.od as »odlfi®d toj- Cook

(1930). wm m tti# one »-0st S'UltaM©.# fymivlo aeld,

^wifled-fey 'faemiam dittiUmtles,, wMoli orystallized la • th® , ie®-

to©3f-*fL© i?*100 pmp mnt tf %%® th«or®tie.&l«.. .fp#»

vioms t0 sfti3Pifi#atioa tM» s«pl® was d#temia«d hf tH© iodo«

fo.r». a®tiM3d.a® ftfpyo^lmat©!^ 8S p#r @«tit pfrmla aoid.

•«tl $, $t » tf * m ® m p» m St « s & s s I s i I 43 -4* .«k ^ 4^ ^ ® fi s o ^ ® sc f4 9 §« ..rt Q U JCS T* • - o r a i a ; S © ® « M * » S% 43 Q # C9: SI 'O l» H O O .®

•tJ » ® |4 »l3 « ^ • 4i> © ®: m m ri U O « u m Q ts ^s> ^ ^ p i ti $4 m G p *3 m O ^ H i 3 4 3

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s©p®i»a.t«lj at 20 11&s» for half m liom:r' aad'mixea at the 'tim

of in©eul»ti0a.»- Bft©to-»y®mftt extract was as a »ouvm. of

nltFOgen in til© Hftjofiti" ©f fementatioiis# A f©w f®i?ffi@ntati0»«

w©i*e 0«yyl®4 otit tiiiag y#««t wat@i»# it was pw.piti'td bj awto-

©laving in w»t»i", .m aaotsiit of api@i oo.aEft©3?el6l yeast «qul¥al@iit

to the Aaslpt.d. pepestitt-g# toy w#lgh.t, snd ramoving the ol©ar

sweMRtant y«ast wat©? Aieh separated otit afttr the matspial

bad stood in th© l6&-box s0V#Pitl daft. Bsato-y-east- ©xt^et is

»0F@ oonvauient to ua« and- rasalts are dteipll©at©d ®asily»

. .fhfta Aiffmrmt typ#®. of baff^y- wmm O-ftCOg,. MaSOOg

an-a a Mlxtur® of. aa# •%P0-4» fh# laSOOg wns prepared ^ •

;St«iplli«iiig - aiiA th»n mrnm^vtitig th.# ®ai»-ba»at® to bi@ari»iiat©

tte-©tt-#i. tft® solution mtil it mm e#l®i»l®s« to

ph@nolphtlii.l«ia# Mfftp, M0 m4 was pr«par@ci by

Bl^ia-s| fch© im^i@«t»-<i pyopo.rtioas of aeifl and eiritt® in wmt@r

with^brm tUyml hi.m »a m inMmtor and amtO'Ola^img at- go lb®*

,fe-»-tiglf ^sn. hoiii'.* fh#-attliod tit mot gite v&r^ &omlBt&nt r®-

Bttffsr® ,pJP®p®re«f wliat ap-p«ar>®4. to b® i6#ntical

eoii«Sitloni often-.is'a-rts-d wld#ly in-pE* Hn. ®ffi0i®at buffer

i^i.-Qla: do&©,,B-©t giir©. off 0&g wouli, siBpllf^* tlie

of ftiialytiSi |Ll.tfeoittg& tills buffer Mitt tli® pi smffie*

i®fitly Q&mtmt to allow good growth, .it iat®Ff®y®d with th®

.gufar a®t«s?«iaatioa and otHer asalysts tlm». yeqttiring a oliaiig®

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• 1. ' |&@ wgmli FootSj. Fred tiid ^ 1&

^Culture received from iafestlgatoi' whom aiae. iU' undei-limta. culfup® from ?ii*tiua®ii» It tfi gftlaetos# *$

therefore, probably is n©% the saia# ftultmr® a® d®sarib@d by Virtanen (19S3)«

#§fM

fUm msulttt fti*# 0&lm%&%e4 m the h&Bi& of jm p&r

.Qtmp^mida fmnd ppewmt ia ti*ae#s ap® ttot in the ©x-*

perimmtal fb.# a©ii*i»®^9iiig is inssmtd fox*

mlmlmtlon to b® li|*drelsfi#d to a ft*sa*»boii sug»T ftnd is ealeu-*

lBt&& ill ©qiiiirsltiit fflM ©f glu©os#.» fh# ©xidatiofi luad I'tdao-

tion «'r@ @ftXowia%@d && fellow#.# An atois Qt' hs^'Amg&n is

«S8ipi®€ » r®d«©tiOtt 'v&3m» *i| tH® oirgsB atea i«t ©xitiittioii

•ralui® of ®f »iyg®ii atoms ia « eiwpwad times

g plm® til® mmbeip ©f Uydi^ogea at@as tiaes ©Qiials th<$ ojxid®*

tlQtk {•*) ®r y®«l«t0tiOE {*1 t'altt® &t tti# eoapomnd, ^Wor exmmple

§0g Urn® an ©xldatioa falw© of 4, ae^tie a©iA

mlthep -.mw a»d propieiiie aeid C.OHgSIgOOOH)

lias ft mitt®'.-of' f« ®j© #;xi#ati©n «®d r®(ta..otio»

balan®# is 4®t®i«i»#i .mltiplfiag tM m @mk produot

by its mspmtlm ©xiAation m mlm and tateraadaing

tl4# sua ®f %im. ©xiaatim walmta and »d«.0tioa -paints*' fb®

h&lmm Is pr«s«iit®d ia tb# tabl©® in fom

8«« of valueg %fm STO velues ^ » 1 •

•Ri© litfidiiig *61ueesi fem®nt)0dj p©r e©iit" it used In

certain tables., fli# aeaning fui»tli©r ©xpleafttion.,

fhe ¥»lti©8 vmdm this blading the per e@nt of th@

originftl gltieose of tb© medium was fermented#

mm*

ihaft@F fw®

Sltr©»en iggm.ty®«eiit8 of th@ pgopioa.ie .^el.d Saeteyia aad ' 15 C^ORiilalSg jil't'erent

MltvGmn arnimm

fh« pfopleaid a©iA feaeteria &ee©f41ng to generl# deaorip-

tloa (g<»plta: ©rganid aitrsg®^ ©eapomads ftur thmlv

growth# If eoB@id#r0tiont is gimn to the s<St«n.@0ii»nt which

has to®@n laad# ree-ently In th« stiady of arowth stlis«l«iits,, ©x-

idatloh-Fftdti&tioa potewtiftls, ©©@ass|wffl, mlmvBl r©c|uir#a©nt.i,

Titiimins, imd ©th®r ©ffeetlv® agent®, it la apparent thet one

may with reason ©oasider that this phase of their ia®t®holiaa

has liot hmn S,fiir«®tigftt#d adtqumttly#

It is apparent thst th« a@taholisai of an organla® groiifl»§

in a a®Aiam of known ahemtml oonstitments say be studied far

mor«. aonmrately than that of on® liiioh is growing in the presenf©

of % @o«pl#x nitrogdij #©^jr®#» It th«r®f©r® wm ad^isahle tiB

initiat® st'least » sursory »tii% of th® nltr©g#a requirements

of 'these organisms# 'fh# author ha® not fomd m meoount in

th« llttratttr® in whieh ©hemleftlly pur# -eoapounds other than

{Rl^O^and l»S0g hair© hmn t#®t©d e® nitrogen sowroes for.th#®®

to«©teria.„ .It app««r®d advisatol# to try' orgaaie nitrogen eon*

tftinlng ooapotjiids .of known ptarlty and oottposition# The media

w@r® «sd# ap in test tmtoes tasing tap water with 0»1 p®r oent

IgHPG^, 3.»0- per e.©nt gltteo®# md 0*3 par e«nt of the nitrogen

oontaininf toMpoiind®* fh@ media w@r« adjusted to pH 7..0 and

mt IS f©x* IS mlnmtes* f®8t Imbsa w©r® inoetx-

lat#d witli 1/3 m* ©f growing ©lilfettr©®# fh® nitrogea

sour©#® aiid r@siil%8 *r» ®Ji.o*fii in fabl$ II •

fli# r#-smlt® iloii- tl»s% in m oas® wer# tli# mm&im adetpat©

t© TOp|w&rt eofitliimed growth, 0|^stiii« tna ertfttln® appeared to

favor gro'wtlt pmTh&ps ttt® to th« fast tlist tliof r©dtt©© the oni**

dati©ii-»r®<iti'6ti03* pot#iitiftl of th© Th® ii®xokiii®s© In-

ttstlgiitloai e# Fromag§@t and fatijm (lf33) »ak« It apparent

that faster® otiior tHaii tli© altrogeia r®qplr©m®nts may hme

ibmm instr«»®iit«l In pmmn%lng growtli.*

It'toas poiat®a ottt la th® -re'S^iew of th@ llt®r»t«r©

t!iat til© reatilt®' tor vm Ii@l, from f«im«iit®tioiis with

w&Tflus momts of ttutolfsat^i ar« aot ao0»pt«fel« as

eonolnslf# of toirtrogen doatttloa %f yeast eoaipomda*

fh# smfgoitftd origin of a®«tle aolA «ai QOg fro® ooapowrMli oo^

emrring iw y#«st l®«ir«s ao«bt r#ltttiw to th# mmT&tSf of his ,

fonaontstiom b®lano«s« It toptsred ess^ntltl to repeat ttios®

®3speri«»*its to d®st#f«iirio if ''thmne r#s(«lts ©owld b« auplioftte-d.'

A ©or# soaplst® amalysia of pro^eta wns msi® to ototain- d®-»,

tailed Infomatioii eonoeming tli#se f»M#ntatioas«

faM«8 III oafi, ,If iwaariB® tti® wamlts from two, ®«eli !»-,

¥««tigatioiis*. It i« spp«r«ttt tliat tli© ratio of propioni® aoia

t© «©#tl© aei4 iaoraaaod witli deoraaa® la th® ooaoaatration of ^

yeast wfttar In ©oaforalty witM iraa Siel's r»®«lts« fh® ©artoou

roeoTary i» aot gwtatar tliaa 100 par ©ant la ©as® howavar,:

so that til© asfmptloa is aot J'uatlfi«d thmt produets ^ ara helng

f Ami II

GROWTH 0# PROPIONIC AOID B4GTSHIA I» UmiA. OOmiNING KNOWN OOMPOTODS AS A SOURSI OP NITR00M

I

11 mm mM mii wiss:

iat Transfer ifranafer ft0iar« .} • 72 tmiri . t e • i IS

34- ^aystilays

Wimm

Aspars-gia®

Qystsia#

tr#Sji 0|rstin«

Oreatlne, Aspsreitin©

Wrea* Greatiae

. Asparagine,.

Or®etine. Ofatin®

trr«a, Aspar«gla#, C^stln®

Urea, Oraatinei, Cystin®

trrea, AspRrap5la@i» Oreatin®

Asparagine, 0i»®atin«| Cystine

Urefi, Asparagine, Oystin®# Oroa%in®

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f®sw«A fTpm ©f th® ysfiiBt# fhs oxidation und 3r®dmo»

tt®ia bmltne©! f«fe3.« If aiPt satisfmntofsr «3ceept mhm % per cent

w«s^t48»€, flii» gif«® strong ©tiaenet that th@

y««st wafe®r was not infltt«n®i»g th# ooiirs# ©f th« eartoohy4r«t®

metaboliam bf #.se«ptittg or donnting 'hf Amg&n* fii© otrbon and

o*idmtion reittetion b«lii»<i«,s In tlie f®r«#nt»tion eontain-

lag 1 pw e©nt f®a»t 'wsitftf IMloat# th® ftanJ-i'-tis was inaeottrat®

an^ perhaps tlt*t uiBd®t«w»iiit®i pr©<J«©ts. wem pmnent in the

Beiitm# •§mXf in th@ higi®r ©onoentrtttlon® of :r®fist wat#r did

tli» voittilo' 'aeids approaoli tli® jrlolAs fotind "bf van Il«l« 1®

msed S8 p«r ®®nt |'«ait wAttr in feraontmtlons from Aiefe oarbon

hmlmmB w@r® ©iil«mlat®d» lis mnit.QtmXf hl^k'y'imMu of irola-

til# aold® itoieli ftoeoiant®*! quitntitettiif#!^ for tte® f®m®nt©.d

atagar mmy hm® wmnlt&€ trm tli« tm% tlimt all f®i»®ntcition®

«r©r« pr©-aw»d wder .eonditioni#. fli® rosmlts in fabl© I?

show ooneluiiirolr 'tH# ©onditions md«r-»teioh''f«m©Bt»tiott

mmm latlm.mm ttot disaiailtttiire ehsngtt of tn®, tugar men

to th« ®3irfe#nt that diff#r©n.t ©M pro'tetts. (laotl.e aeld) ar«

prodttoed. fb# sofei©«® of fmraBntrntim d«rl¥#d bf mn li#l for

on® let of oottditiona ©anaot b© ias®d to Inttrpret results

wndor otlior oonditions* fte® f«©ts ©bs«r¥®<l her% support th#

ttaswwption that th» oh»ng@s in th© ratio of propionic neid to

aootio ftoid ar© not to iifdrogen donation or aeooptane® 'hf

ttie ©oapownda' of tli® yoast but rathor to a ohango in th© aetwal

aoohanisffi of dissimilation# %n faot th®s@ rtsialts b®

eonsidarod as diraet ©^idanoa that h^drogan donation hf .the

. 6*

l-esst ©omfomndi ii not %mpQT%mk% ta -pmplmlQ aeid fementft-

tlom. fh« • patios dt«5r®«@@ with lii©f'»a8© in yeast mumntrm*^

ticm wh®r«as on© would ®xpe«t thBm fc© in.er©»s«. with' th« mn»

eentration of fmst ©i» liyiFo^n dietiato-r'S# fh® exp®rim©iits

also suppoft til® h|-|joth©®is that stt-eolialo aeld originates

from th® glttoos© yatli®f tliAn tmm eoapeiandt of th@ |-#®st# If

sueoini® a©id origlaetet tH# f®&st so»© relationsliip

mtg^t b@ #xp®et@d to«tw®®E the f»&&% eono®iiti»«tion and'th®

•^«tttity ot atteeiaia in»@'dtt@®d* flila was mt $h& ©as#,

ffe® ftt^t thiit tli«^ ©Eidtttiea and rtdsieti®!! toalmnees'•mi'© satis-

faoto-f^ whan- g-aesijaie, a®id la inelwdad a» a pyo^ot of sugar

ditsiailation ala© f«pp®i?t« msh a proposal, fli® mPb&a toal-

anoa • !»•• ttie farmtiitatioa eoatalning 10 f#r ©tnt r«»»t watar,

fafel# If, Is ®atisfa#torf.#- la ©thar fawftantatieet®' tha analyals

wa's not acffiiflata pmrnntlmg » ahaek % thia sathod, hewaver,

tha volfttll® aatds did 'not aaaaimt f©r th© sugar farsiaiitadi,

?«« -Hj.#!''® f©r«@iitftti©fi8 in yaast a«t©lysate prodtieai volatile

aaii in axeass of tlia sugar f««eiita4» • It is airidant, thara-

fora.,. that iraE-fial*# raswlts ara not ©Jtaotlr ooaparatola t® •

tit©## ®f t»iis iiiv®sti|iati0a« lawavar it is Imprababla tliat

tlia mitrafan aata'feolisa iafluanaad ©artoohydrat® •distiailatio.ii

aa h0 intarpratad*

•flia praawatloti ©f laetie aald ant tha aon^radualng aatar*

ial will 4isaii»®a4 la iatail ia Ghaptar III, whara tJiasa

rasiilts ara aorralatad wltli othar data#

S©o«ntlj « wiwb@F of patsnti have been. obtalri®d In th@

to. th® preduotlon of ppoplonle aeld

f©w®ntatJ.oa.# indleatlng thftt thla fermentation is ree«iv-

Ing @oi»#reial 0oa»id#rgLtioa« .and WmrTsm&n fltS^ investl-*

gat#d tli@ utiliZfttioR of ft Bmaber ©f agrlifultural by»produ©t8

in propionic aei€ .^®»®fttstioa una reviewed tJi© protolsM .©on-

frenting inAastrial pr©d«®ti®ii of tliis' asid» f!i«y p.0int'0ut

that a €ih@ap i.ftd aatisfaetory sour©® of nitrogen' is @sa@ntial..

fwQ' attregta s-6iir@@», mm' ,glttt#n *#0.1 and flt®#i> water, w@r®

stmdi«d and fomad to •tatisfi' thesa. r.#qmir©a@nts., Gorn gluten

m#«l «R<1 st#.©p wat#r ar« bf.proAiets r®stilti.nft ^froa the eom*

in#reial prodwetion Of .eora .su®ar»' i#sialt« from gluo.os© fer*

aentfttioiia in wliidfe t.h,#s« .«itr0f«ii s©ure#.s haf© b®&ri used .are

tubtilatefl' la fmbl#®..? «ii6. ¥1* fli#a© results ®rt In aaricf r©-

®p®®t» iiall«r,.t-0 thos© ©b%&l»©dl with gltieos©*y»it0t water f®r-^

metttstlons# OQacentration® 0'f S to 10 p©r e«nt st®»p watar

and stpproxl»at®ly 10 p#r 6«.Et gl-ttt#n a©«l app#ar to fulfill

ad#-(|Uttt«3l|- tti@ aitrog©ii ©f fea«it#rljs..,' it i»

lnt©r®itini to netf th.tt « d®6r#*s© la tti#' eoneentration of

nitrogen material m.a\x$Lllf ^ muBeA am inereiks® in the ratio of

propioais. to .sottie, RSit#. m& a d®@r©tts« in th© rm&mrj of

gliisoa® f®iw#iat«d as fslatll# aeida* fli®s® results liwlloat®

% ohattg# in th© aeehsfiism of f»na#iitatioii end suggest non^irol* •

atil® -aoids ms.y hav® fee#a- produ@#d at la tbe f®ria#ntatioR8 witlj.

jeast wmter#

. f 4B1.S ?

BISSIimTIOM ©f QLUGOSI, fAlXABI,.! .GOiaSWfiAflOM OF^SflEP *ifSR* FiRiRSfi® 14 30® ••O#,' immzQ, mo w mbdito

» 500 0©, FiiMi* i gs» o«a%* 10 g»t siitreosi'

- • • • • • • " J'' • '"ff ^ I© •'

* I I . g«g liter . . ^ ^ 100V®fed St0«piQlmm^Isitt@©B®IIpropionieiAt-t'ii'Sf Rati© i ' wa%0i»f t0r** t f#i»- fT0l«:-i . aeii .i a@ii i.py&pioni&f • vola-

smentsd ta«iit®4 itil# t s 'i alelTe^S til® I ,' i;.. • . itteM. I. . ' . ..I , f aeid

P&T I per t m $' m %• m .t i . ' i P©r oenti eeat i ^ ,.|. . , , i ^ . •. i . . , . . . ;j,

O4S i '49^ s.' 81*? llOS*?! • I lt#2 I .4#35 | S8*9 'S ^ S H g

1.0 I BM: t , 04*1'llTliOs IS84S . I 30*5 1 4..gi- t 85*2 . t I I f , I „ , 1 f

5^0 f, 84*0 t 140*0 tmi4Q$ S0i*6 , r 0t«4 s 3*35 t ^•85.,8 .. I S . , S I . t I S

10*0 tv ff*0 I 1«S*0 • t34t*Ss t 84*6 1 SaS'^ 1 97*£ I - 8. j j t I . t .

11 W " ' # i' ' • ' I ' I • 'I '" I • s •

0«S # ©4*0 # 106*7 Ill0*7t , 7.i*g t SS.g | ga4 1 46*4 S • • I i t' t t I

1.0 i @3»0 i 10S,0 ilSS.Si 110,? I 4§»1 I 8.4i s 6?-l I " f I • I , • • f '8 I

5,0 t 8T,4 i im*7 |g38«0| '167,8 1 70,2 t g*3t t 73,8 s s I I s s f

10,0 t -08*i f 1©4,3 fg43,4{ 166.7 | 76*7 f 2,17 t ©6,2 t - ^ . t % s ' . } i t

f Ami m

mssmmmiow ©f gwcosi* GoiimmmfioMB OF 11 w, FtHMT?Wf B:I5 8 BAYS, 30® ABHOBIO, SOO ee. OF ISDUM II

LIT'PR FT ASK, 18 Rt* 1>LTJ00SE, 20 gs* CaQOg

f ^antitit® 'prodncsed ptr'-iitef 01ttt#n 1 fotal 1 proDionii3tAO0ti@ 1 fo»ie t ifttio

1 Volfttil@t aeid' 1 «oi<i ^ ^ asid t SPODionio 1 «Gid 1 1 t

p®r eMti' • Mi f 'isK • • 1 , f iM 1 t 1 f t I

• 1' i loa,8 1 #3.0 s 24 » © 1 19.2 1 g,S7 s s s 1 1

4 1 230. f 1 W&pA f 88.0 t 13#8 s 2.73 s $ t 1 1

12 s 33S.i 1 • m4»e 1 11S»S 1 • g.O s l.Si

• S© f f - 340*0 1 too%o

1 t l3t#S 1 • ?•§

1 1 1.60

1, 1 s f

fh« of f©ffflie seid" in th#3# feTOentstlons m<»

qttifti «oiisid#:ratidii» Fo«aiG «Qid iiamally ia not prodwoed

whm th.®!'® is m abmndan©® of a favorable nitrogen material

pF«s©nt in the a®dim., Imt sometimes ©sow's when th# nitrogen

souro-@ is iaaA@Qtiat«,* tfc# fa®Btion pTOsenta its®if^ does thi»

foraie «0ii originat®'.fro«' tli© glueotsf *tt®3Bpts to-settle

this qutation liave not hmn sueeessful feoeatjs# th© foOTi© aeid

usually 4@#s not o«owr in qaantitl#® la3»-i® enom,gi to »at@riall|p

affeot til® oftjpbon- or o:ii<iation «nd roduetion balane©#. It is

I50«»ilsl® that <i«.ooMpo®ition of the nitrogen materiel, whioh

may to#'aor® setiir® in f©m@at«.tioaa wlier® th® nitrogen a«t©r«

ial is inadequate, l®s.,ds to th® fometion ©f this aeid*

•?c-»

«@l'd baeterta f®il©«l to grow in iMdls eontain-

tng uTO'&p aspas»ffl,gia#, ©ptrntla©, &r Qfstlm or la comblnttfions

©f thdi® aoaipoundt JI)* fli.® po'tsiblllt-if li suggested

that fch®s® altPOi^ii sotirots are not mtllized beeaus© ©tii«r

eenditleus B.m not suita,bl#- tor growtii#,

• , Fementatiom ©f glmo©©© in Alff®3P®nt ,©one©ntratloiis of

y^ast ii»t®r w#rt studied and, a variation in th# pvp&uQtM of

glueose aiisl:fflil,atlon w«s a©ted .Ifsblifa III m& lf)»- It'ls

»ngg®st#ci that th«s# in pi»o4uot® m&VLlt from ohanips

In th# ia©olis.ai»« ©f dissiiiilatioa i?atii«r than from a foiwation

of pFoduets fm» •eompotaisiis ©f the yeast as 8ug,«^e8t©d bf* van

llal« Wid&nm sapportifig hydrogen, {lonatiea ana mm@pt&nm •

bjr th© |*€sa8t oowpdmad® was ftot obtained hf this' m«thod»

fw© nitrogen s©aii»««t foofn glutm meal and st©®p water)

of po»slbi®' QOHsa^reiftl sipilfieane® w#r© studied in glucos#

f#ra®ntaitioiis' |f«ble» ^ imd ¥1). ..F©»i0 mM oeeui*iP@d as a

feiment&tion pi»oduiQt#

yroa^ota • of tht FyQpionic Aeit

0»ly fipapi®atRi^ «tid«3a#« <i3il«t®d «t th# tta@ ©f liiitla-

tion^of tMa ^ liiv®8tig«%i©ii mmewntng th® identity ®f ijit@ip»-

eompotinds oaenfiPiiig ta th® <ti®i»S.all«ti-0ii brotx.ght, ateout.

toy tli€ propionic tt©ia firtanea and lerstrlw (1931)

had iselfttecl a iit®x<}»«-*i8©ft©ph0»^«t@ aad f0dt®, and f©t@i—

sea C 3.930) lied fepeiHfe®# tlj« pTOdttStioa -of laeti'O aold as m.

pm&MQt 0f feMentation# Foot® #t» »!,, did not «ttgg®»t

tliftt th#r eonsld#!'©^ tli® Ia#tl® melfi to be m lfiter®i»dlat®

prediaet, eltliom^ fiiptan®!! (It23) |lt0S| upon th«or#tloiil eon«

liRd ©dn©lttiidi ^at tiiia.aeid ii' iiit®m#diat© In

glaeos© f©Btt®n.tati®f».« fk# Intemsdlat® m®@liaiiS.»fflS preposed toy

•ran *©!»• largely speoulatiTOj nont.of th® int@wa®aiat«

eoiapomds w@-i?©. ifolatsd «nd .l€#ntif|.©<l# In (1933) WTmmgm%

ant fftt«a ttsini tli® 'two @wlfctti»®,a whlcth pE'oda©#A. laotlo sold

In tl»« «;Kp®ri»®ats ®f •peot®,, fy®d and F®t®P8©n d#MonstPat©4

tli« f©watlon et laetio wid* • fhis aeld is Inelttded in tli.©ir

F®tfc «iid Wynn© C19S3) isolated Hdthiyglyoxal thtti

lending ®WDp©rt to flrtanen and ¥an Kiel*# pp©f©s*ls ©f its

TOl® In gltt0o®® fementatlon.t

fli# tttTsstlgmtiQas ©f til® h.mV9 ltd to th« isolation.

©1? d@t®®tioa @f m o-f otIit*» poaaitol# int®«®dlat© mm»

ponnds# f!|yi pf^^e^dnres and ©xptrimtntal data, by #iieh tn©s«

«Qai>.©mnds mem ar# • Im tills eliapt®i»» ffli#

06ri»®latton ©f ' wsnlts tut© a 'Siagl® 'seli©®® of dlssialla*

%i.6n wliiGfei fits %h® &Lft9mnt h&lm.ms is ©ongid-*

®i»®d la th® mx%

a.»--lloa-wtmeiiig

m^fitigatloa Qt 8la#« f#,m#iitstions %

dwrtog til© eourse mf III®, gltaeet® disslail)fcti©n led to the d®»

of 00ap©mad,^ it ms f©md'tlist tti© oarton of the

g'3.m@as# feirai^nted wm »&t- s0ft®wt&d foy hj foiatlld aoidSg

smeeiai© and ImtlQ' aelds' mt ani" tlm® pi*#vlous t©' the 80ttpl«te

©f tlia 0f tM® a«dliaift# After' the glmeos®

wft® ®«tip«ly "d#<ioa|>0»#d. tisutr® was « ©ii|itlnti«d f®T»©iitfttioat

li©w®¥®i*, -wltH til® tQVjmttm Qt «j®- umml pa?©teots,' fUi» gave

% fiiiftl &f' ©ftiPfeoft t©- appr©,3ii«iit0l.|r' ©a® hundred

I'esmlti' amppert th® ©f the iiit0r-

aiedlat® f&vmtioti ©f a i»®®@rv© f©oa mutti'lal not leading It-'

self, to • d®t«»iaBtlQii hf th# mimal »«tli©ds ©aployed. toalysii-

vm md« fop tion-redwelBg aattFtal, m& data in fabl# ?ii

Htayis®' th® retults ottt:»tn®d fp#a « f®.»@iitstlott In whieh thl®

analfsis mm la©lttd»d« f|i» s«tai» was pmtti'talli' 'e^aipletely

f«3«®iit®d after Ftllewing this •tlsd^g.O aai of

glia0.©s© w&m f©i®®Rt«d «jtd 20*9 m of mlmtiU aeid mmve pro-

duQ«d* fh® aoftljsit for ttOR-^feaaeliig 0s3*te©hydrftte showed an

laer®®®# in thi® a®t®rial mtil th© ©Ighth d«r vhen It d®-

ey@®,®®d siaultati#©u8ly with th® i»op@«s® in velatil® adids.

31»q© t&s ©figiaal g&i?a no in we^uQlngMmtmut"

ial sfter .h^'drolyais it Is ap0iipa»t that tii® •non-p#auoiiig mat®!**-

ial .was pmAimd hj th® fer®«ntatton* M&vmin that the non*

rediieing material is hf&Tolfm^ t© « 6-c«yboa smgar and that

g.«0 sH of aoifi are ppodiieefi from ©aoh ^ of sugar

feraented tti@n 19#6 &f tti© i?0«9 im of 'rolatil© aelA® lAich

w®i?s pr©diie@d aft#r th® ©igjitfe 'daj fir® assaemted foi? bj th®

gl-acote and iio»»i»®dtaiei»g ffiatai-ial femented aft®? this tint®#

fhis l#Ma tireet impport t© th@ abeir® mssumptioa of tli«' €•

carboti' tiatttiw, ;#£ tk© ttydfelyied

fms fll

DISSIlIMflOl Of aWQOSl, M t, 3tf® i*, «4IB0BIQ X##,Pia Clif 0LI}OOSK', 2.4S FEE Slit I«HQ©g, 0.iS FSR CSlf liSASf IXtHAaf

• • I T ^ mitntli"lii~lTb^ua»<l' 1"

PeTiods I jvola-spionios Aceticj reciaoingt of sfJlijcosesSlncosestile t acid : ©old jmaterisl:!

Hatio

incu-1 fer- s fsT" jaeid s s s as J ®c«ti0 bation:ra0nt©d jmented $ S .r s p:lucoge i,. «©id'

6.awBttmT omits wM j: . tils . ' .1 s Tf$i 1 , .| 1 f 1 1 1

S } 42.4 1 36,0 1 i0*8| 37.g| 13# # 1 1 ,g«73, S 1 i ' s f f

S, 1 ma t S4.#S } SSirit 61t©l t6»6 f 4'a i B:*55 t f 1 t 1 t

6 t 78 ,4 : 6t»g ?7.i| St.4 t , 5.3 f g.39 t f f s 1 I

8 t 97.7 t ss*o is.ft 3@,f t 7,8 : 2,5i f s 1 ' 1 f :

• M,: f 1.00 *0 1 S7*0 tlSS.is 107.41 4©.t t 1.7 t g.3S ! . S $' 1 t 1

13: t 100,0 1 87»0 iMi*Ss I0i*ii 47,f 1 fpaw s i.33 • • i L, S f s 1 t

.»f4-

flig'SMlitS' of thM proplomlQ sold'toaeterls to ajntlieslr,®

a non*3?©sfMeliig eoapotmsS fro® » aonoaseeliapldt 1® a feettir© of

tlisli* aetafeollsm w!ileli, has not hnBti polinted ©ut pMirlotisly.

It la Qiilt© posalbl© tliat the aj-mtlie-iis of this ppoduot Is a

impoTtAnt pfim« Im th,t »«t«lJ©lis» of thme organlsats.*

fht non-yodtacing Qmpmn^ «»f b#-,, of mxirm, ot no direot im-

pertanee, fdr exampl® stTfiRf as a rsitrt® aaterlBl foi* the

Ijacterta^ It Is pdssitol®, Yiowmm^ that this eoapotiM foras

an Indispeasabl© piitj*t of tlia fewemtatloti as a prede.oessof of

.filial • pr©toots of dlatlallmtloa. liidted, la-th® pTOplomlc

iicl4 aisslmllfttleii of dl^ti«a© eompounds there flMt may b® ®

gyntfefsls to a noE-re^melttg ©smpomd i^teh is ©osaaon to the

•picQploalc f@«5.«iit8tlen» .fhls ,iion-r©<ttt.eliig eompotimd «ey In tuirii „

be fe3fia®at©d to. ireletll® aolds, SOg, and stieelnic acid.

In ,elos« i»©lstlea' to this specmlatton. mre some iuggtestlons

bj ?irtaft#ii ,tM t«,ipati»Sm. {ItSlK In fsraeatrntlons of pr^^uvlc

meld bj pfoploalt aeld oi^gaais*® these Investigfttors tk&VB

foimd smoctule mid Qm ©f the ®iii-pTOdiiet«, Thef suggest

th»|^ th« seli is first iynthetlsed to « he:KOse aol®ciul<J «

whleh l» .in tnm. br@k®n. towm a 2* md i-oai^bon ©leaveg® to

s^efiialc aeiA tafl -a 2»0arboii eoapswid. In of th© restalts

yepo3»t«d hBTBf. th# srnthdfis aay^ b# t© & fom more ©oaple.at than

ill# ehaia jjostaltted by firtanett and KaTStrom*

AnfeljaiB ©f'washed ^ells of th® f@tta#iitetiori dlselosed

.that a portion of th®, iiQii-r®aii0in.« aatdyl®! was pi»©i.®nt tn th©

<S0lls.of> had been iiiolttded with them In th© fovm ot en. insoltibl©

preelpitate# leweTer., largest i>'Oi»ti©n of the iion»r®duQiiig

siBtarial was • wetei^-solwhl©. ,

fhe aon-reAtelng wefcertsl has been deteeted in fermenta-

tlons of gl^eerol and laetls aeld aM for this mmon it is

rapMstntsd as origlnfttlng ffOB a triose by synth#slg in .th«

feyroftiitatlon s©h©m©»

Th« non-redijolng iB»fc6.rlal has been fotrnd to be produced

bjr QV&Tf c\iltui*e of pfopionio aoid bacsttris studied in this

inTestliiatlon* fmof will aot be tabulated h®?© btjt this fast

la shown hy tables throti^.otit th« theglf* Oulta.i't 11 w (Tablea

If En4 lo^' eoae^ntretlons of pyodmoed pertlemlarlf

InwgQ aaomits ©f this ©ompewtiAi 3t#3 p@r mnt of th# ©.arbon of

the gltitest f©i»@iiteft was ittemitttd in this form in on® fer*

laentstiOE. As f®t th©^ ehtntical aetiiTO of this oompawnd has

not b00H iavsstigat®^ biit caltwp® 11 W showld prov® valtiabl©

fo.r tufth a study#

aeeti©Mt g» Iia®t40 • aeid#

Although thi®. iisid has b««n fmn^ as ®n ®u4. pFQduet of

f€ra«Et6tion, fftttoips mv mndt^lmu impen*tant in eaiasing an

gieotttwlfttioh of this ©asilf &mlA h»ir@, not been d©t©i^

mlh0d» liiieh lacti# «@ia trom glucos®

in e3cp#i»im@iita Foot#, et# aJ» 119303 w«s •««©<! und»r oendi-

tions b@ll«^®d, t# to© eosparabl® Imt the aelfi wa® not ppodttCdd.*

In atu%in.g th« alti?©g®R r«<plr®»tnt» ©f tai@a® bii0t@rl® it was

fetmd, Ohapt®!' fwo,, Seiation 2, fatol® I?, thst mltVLit& 11 ,w

pFoAneM l&etis meld in raaflla eontalning low ©onoeEtrations

of yeatt, Sxp«Plm@nt:s stawafia#^ In'fable ?|II w©r© flt'^ised

to.eataMlsh whathciJ* qt not this *as a general phenomenon of

United, to this ep#<iiea, A of the i»®swlfcs shows that

11 W is th# mlf. cultwe mmintne6. whleh ppodua^a. lactic meld

in substantial qttsiititi#s under these eoaditlons# Partlmilar

care wa$ taken to ©stfibllsh. the ptai»ity of eultur© 11 .if b©fop©

®nd eftsi* ftmantfttioa and fclxs a«tlio2» is eeafidfent that the

f©i*ment®tlons were not eontimlnated, ffe# fset thst this .ciil-

ttire irtien t2*aiiS'f®rr#A t'C laeclla ©ontainlng o0iie'#ntj?atloii®

©f ysMt pTQdnm& f««a®iitatiOBS wMch yielded prtdomi-nately

TOlatilo, »eias with llttl# ei-'ii'O la©tl© a-eld fmrthdy .sutostam-

ttated thla vt#'W# ,ftis feREtat-stioa % 11 W In 0#g per eent

^©ast Is. remarkable, laatle mlA and tlie non-i^ediiolng mftttflal

aceomt for 95:»3 per m-nt &f tb® earboa of the. gluoos© fei»-

mentBd, gn explftnatioii of tlies# resmlts Is not apipertnt at

tbis timt, howeV'®!*., th# eon^ltions Ina'^e been ©steblished. which,

eaus© tills p.'si^tleislti' sp^el^s to produce lastle. aoia#

taetie t-el-d haa ».lm bs'sn found m a femtntation prodwot

wh&n m0di» wmm f@ni«at@d w&iefe. oontsiii#d IgO- ©.n-d'H^PO^ as

tmffer (falsi®® II aatf X:)« m^dla oontalned tli@ usual eon-

©ontration Qt jr©tst ©sctfact. .«ad ©th©!* than 11 W p3?o»

dsi0#«i tfe# Insults* fk@ imv»atlgftti0» of th©s© fementatloas

toas' R©t hmn ©xteailf® feut th® pipodu^tioa ©f laotle -aeld wa» •

deflnlt©# mmTTmm of laatio maid In gli-e©!*©! f©m©iat»-

tlon 1» of Agsim th# eatts© ©f this eliaagt In

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disslmilfttioti is mot mdsrgteod# flie pE ehenges were verj

slailer to those tn fcsmentetlons eontel.ning CaOOgf starting

,&t approximately and dropping to ebout 5#0 at the conclu­

sion of f©m©iitation* It wotild fed of interest to determina

tlie magn#si-am 02» |iiospk&.t© ioa in comMnation with

other ioas produces this pl%@mm0mm It sho^ild b© mentioned

that si thorn#! the sufai* aiial^ais mmn not aecurat® it appeared

V0Ty pfobabl® that th© carhon m^oveTf was In these fer-

«©ntfttloms indta'&ting the foi?iaatS.Qii M a oonsideratol© amount

ofan unaaaly^ifed'-

f AS.B I

FKHM'ZNtATIOKa Wifl 1^, %fO^ BUfFSRS^

^feltarei'''' '" '''''"'^aailflil®!'"'pyjlSo llfaF" g ^ ' " ^ ifitio

f eraen-: vola-X Fropionio $ Aset ic $ Fomie ilisot ie.i SU0e inles Pirogion^ tation stil© i acid s acids aaifi i aoid i acid |'~'Ie®fTF* »©• saoid J 1 1 1 ! s

1 lal I S®''' ISi i s " a#"' 1 f 1 f f •s r 't !

: 34 W, 1 SO»Ef f iB,*r 1 1.9 1 52«3 1 1.5 t S,"?g t 1 t 1 •t 1 S

49 Wg ll40,0s 124,3 t iStT 1 0.0 t i4,4 1 23*4 f 7.9g f 1 t 1 1 1 t

34 W, tE3.9*6| ii4,g s S..4 # 0»0 s 7.9 t 5»8 s 39,6 X s i S :«

fatol#,'lX fe? 60iidltioM 0f •xp®riB®nt*

fh© -of th#s© i««swlts should to# tonisider-ei in

Felatlon'to th© po-salW.® of Isetie aeid a®, an iittemadi-

ate# -t-adtis is F©a€ilj, .femftiit^d by all »p«ei®s of pro*

pionio «,ei<!. t3S0t#yie, theydfop# it is prohaM© thet trnder th«

proper eonditions it nmf pl«,T « pmrt in the int«no«diat#

fli® a0l€ amst to© pP34m©®d fro® gluom® mom rap*

idlr th«n It is mntmfeA mhm i% is <let®et©i t® a produet of

fersentation# tt 1» fmwf taHlife®l|*, howe'ver, that the feraenta.--^

tiofi ©f ttii l» #»tir©ly eliminated by conditions wtilofe

l«a4 to its aecTWilfition in th,® f®pa©iit»tioiR., fli©r©for© It Is .

Iogi0al t© matm® tliat laatie a@ii aay «cit as mn lRt®ra©diat®

• @@mp0md in glueos© ferment at ioa# m®Qliftfii«»s leading

t© th« foiwfttioii lmti& mlA and pyniirie aeld fro* gltteos®

«r«: pr®s«iit#d in Ssetioa 4, Fi^r© SI* Tk® m#th©4s #f foraa--

tion ©f tli<is# tw® mnpmxiM «r® olos®ly r#l«t®4«

, S«dtioa rnspi©ftald«hy<S@*

fh# ,ijit©3«#aiet« Tern&timm pmm^n§ th# profiuetion, of

prepioni§ aeit tef# r®o«i'S'«d relatiirely littl© attention,'

firtan®fi |lSt3) aad (1.9gS) sttggftatftd that glumm i« <iistimllat»d

t© t«etie teid whioh is to prepionie aeld and oxidis®€

t© ft«s«ticj a®i€ and fh# widen®#' supporting th© lnt®r«©d-

iat® oeeurreu®®- of Imtia ««id has l}®«ii aumiiiarlsed in s#ctioii

g ©f this •^hjipt»r and it is apparent that suah a oomir©r®ion

must to® eoa8i€©r®a profeabl®, flu© aeohaiiisa ©f propio»lo aeld

fowui,tio» proposed by ^mn »1«1 iiaa m© experimental ©vl<l®ne@ to

support it and ifiua. Ii«3. offers littl® apocfulatioii to substariti*

at® it.. fh@ fewitiitiition ©f gly@@r©l ©ff«r« an opporfuiiity of

itudying tlil« pha®© of th® •disslMilmtiOia.* in thii r#a&rkabi®

• ferfflfntsition, fm Bitl found that' glyoerol i» oontertod prao-

• tiemlly qu.ftntltatifoly tmder anaerobic toaiitions- to propionic

S'Sid and no ga# i« fti® glyeerol. femantatlon was for

this reason ehoaea, fm ttttif#

f® 4®%#®% • iitt®»®€ia%« ©eapemas th® following aedim

was m«®di la®%®-|'®«st a^t^aefe 0#§ p#3? mn%, S#0 pep

east aad ©altiw ©aftodtiat# f'#0 pai* ©ant# Salaitia sulfita 1.0

©aat was Imalm^ad ia th© aadiim as a fisting faaga-nt# • th®

f®w«initetl®iijs war# msMally 'earriad otit ia 800 #©• flasks with

300.,®#* '©f ^aadiaa^ fh® taaparatmire^ &f inemtoaticm waa 30® 0#

Qmltwaa 54 W and 4i l prodwad th« sipat aatif® fernantation#

Aftar i ®i? .i <lay« %nmh&%ion. tfease famaatationa war# ta®t®d

for aMfhyia® -or katonaa fey tli® additian ©f £, 4-dittitro«phanrl-

h^draiiu#' diaa'alvad ia gl 161# Aa aMndant fellm prasipitata

was ©totaiiiad* fha following axaapla ia tfpieal of tha aathod

ttsad in idantifying th® mmpmn&m fha aadita waa dlatili«d to

half Tolwa usiag a» itad eatidaniar* fha mitooim mrhQUft

compo'ttiid irolatilizad m& wai raaavarad im. tha dlatillata# ipha ^

laetarial waa aa»©a.iitratad fey radistillatiom and darivati^as

wara praparad «»d than purifiad tei* e-ryatalliiatioa fram- aloohol#

fabla XI givas tha saltiaf points abtaiaad#

M

iiWfiFiOAfsti OF fiofi.©iALDifflroi

^ ' '' ' '''^al'b'ing'poiRta^ f tttTOWtttFropion-fliji^ it,itera^mra'

gariirati'^a $ faldahjdai .i t I s 't

2, 4*dfcitra^Ry3iir<3^©«®l iS4»® i isg*s t 1S5.0 f 15§*0 I s I

p-mitrophanylhydrazana § lti*0 $ 124*5 $ lg4»§ t 1SS#0 I I t t

ptgedoB damatiya s IMM s l&M i.

8tl«© -lifts b@@n fix#d and in

gljeefol tmmmmtmtlm % a .ie-osnd »0%h©dt flie following ia@€.-'

ins wa® f©»tat#ai ymB% mtmet -O.S f®i* @©st, flye»i?ol '2^.0

p&r iftHOtg 0-»§ f®,r ©•#»% fto# <li»@-a©n. (i|ji»tli|fi-diliy4r©-

r0S0i?flin0l) 0-I.S3- per eeEt# fb® fefatatetioni w#r# iB®M.bfit»d

at 30® 0*, -atr-obisslli* ia BOO m$ tl»akB* Attor -ft'pproxiia«t«ly m wmk a pr»0ipit-afet eetild 'to© d®t©@te€ on th-© siaes of th®

fliia&s. fli® w«y« tnali-atd fey 'th© following

-Af%®-2»'3 to-4 iwQubattoii %h% iup©waa%ant liquid

was-^ »nd th® piweipitat# -was di«-»olv#d ira. hot 98 per

0#iit mlmh&X* It wm mpm^lplt&ted by adfiifeioE ©f water and

purified by ©ryst-ttilisinn froai ale^li©!# fh® «®ltiiig -poiat and-

»i3f#d'Mltiag p#iiit ld®atifi©d til® ©ca»p®ufid m tli® dim#d©»

dtri^atif® 0f pr@pi©-»ftld«liyd®<» ' fropidttftld#liyd« has b«®ii, ideii*

tifi-ed by this a#tliod iu glye®rol f&-r««tt%atioii using-©ultures

49 W, 11 *0 M W, and 48 *# It -should b® m«n%im@d that

dia.0doii ii an seidi© gpupeimd and that it do«s not pr@eipltat«

ald©hyd®a in neutral toluti©n» fh® laiSQg wm «dd®d- t© mak«

th# p»'®f th« aiediiM fair-orebl© for grewth. Is th# prgaEiis»8

f@i»®«t®d th© gly@«rol th® mmll qusntity ©f biearbpnat# was

daeeaipos&d and th® Msdiu» was sooa suffieiantly a©id t-o alle-w

th® pr@-0ipltata t© fom#.

^ -fha pr©pi©m«ld#hyd« pr©-due-ad im gly®®rPl fixation axpar-

iaanta.was' d«t-«rBin®d fuantitatliraly by atrial" aniiySas during

th« ©ours© -of %m faidant at ion®# - fit# results ar® "'giT©-n is

fabla XII. fha madia »®iitained 3*0 par,- @®at glyearol,; 1^0 par

<•811# 1

0«.S©g mnd 0.#5 mnt jmmt ftj© c?n#

tl©a ^wft® tmffered «it!i 8 gas#. &nA 40„ @e« of 4 Si i^osphorle-

fehft other with, 3,6 gms, Oa0Og#, light hi«wlip®d ce* of ai«d»,

tmik wej«©-feireiemtsd, in lltei* flask® at j, • and • toy ©tiltixre ,

M W» In th®m mxp^Tlmmn%B th® ftfrntnfeetlQii trntiemd mlth.

Ig0 gw® tli« lartist ®f pi^pi@iiald®hya©» ,

m#0«ffi»la%i:oa ®f »d- smfes^qwnt €@8tinietion of

th® lii til© teffertd with. OftQtg wa#' ofef-

s#»@4 fysftttiitiF' In fh# ©©•

in apl%® ©f the fast thtt tli® glya®!*©! feraentatlQii wms.

not •#sap2.#t»# ttls ..t# tliat p2«opieiiaia®iii|^<l® •

mn b® fey tli# proploai©^ «#S.d toaetdPlm and alao tliftt

imdtp Q«r%«te mn.Al%tmw will aot tin tldsliyd®' of t&»

•tftia that Uss fe®@ii fix###

fimi 111

iOTliOL fmiEKTATlOK, FIXATIOI Of FlOflOSALDlMDl M SftS0.«, FROPIOKALBSim)! » « fEl tlfll ^

t t 1 i • Pays tii©wbit:t#t | s f f -t 11 1 1® t m

t 1 ,1 1 Hgo • II,»04 , 1 s»s 1 li»8 t

1 IS.S t n,7 Hgo • II,»04 ,

f 1 t 1 $

OkOO. < i*© i &*2 t 4.S 1 o,s 1 0*1 i 1 t f

WymvM aelt wm& d#t#et®d in »mftll eQii0«iiti'atS.ofia along

with proplofi»14s!i|-4« ia f^mem&tima fey mltum

34 W* . imthMa ot will to© ttso'Uis®i In the

s®©ti®ii on pfravia aeid.

ft

^84-

•• Attsnpts -to,detect p?oplan&l€@li,yd« lia g1.ticos®i lactle »0l4

and ssanltol fewtntatioBS msliig flxatieu mothofls ha^#

te©©ii TOBtieeetsftil.#. Jn-gaitho"i?*i ©plalon ttils is not eon*

olu®iir« ®"fiti#n0» thftt pi»opl©n&ld«lij*d® *i0t an int#m®diat®

in til® llissimilfttlon ©f ^#s® ©oapomds# An mmplB hm to®en

giwn ail&oif# .illtntrftttng that tmlfit© is not ®in effeotiw fix-

sti¥® md«r »11 eoinaitions# fh# ooaeentpatlon ©f frem »l'd®%d»

md«bt#dlj affeets the m%e of fliation. In ©©rtain. ftwentm-

tiont tli@- «?on0«ati!»fttion of ald«li|'d» may not' b® suffiolante to

bfing ebottt ffeelfitetlon ©t flxationi in for exmplm

during glfesKJl fe«#nt®;tldn tli# Roemroalation of ppopion*

a®!- hm twffieient» fhls suggestion is substantiated

l>y th,0 fsot that propionald«lis-d© wa-i not d®t«et@d in certain

of th© aotiv® gl.ye©i»ol f#fp®Bt,ation8« In the atoasno® of

mn%mr$- @¥td«n@© th«r#.i® littl® reason to aasum© tliat th®

int«w«diat@ pr©.#aot0 l#«ding to propioni© aeid B.m markedly

diff@r#nt in tlie glyoarol and glaoos© fermentation® •

in glr®@rol and gluoos# fewentations a trios# phosphmte

of or glireerol aldehyde probsfelr is, foraed in

tti©' first piias® of f#natnt»ti©n« The sah^m in fipir® I

gests « «©#!i«nism for 'th© fomation of prop-ionald^hyd® and pro*

pioni® soid from tMa oospomd#

flie oonponnd® in^'oli?«d in this s@ri#® of r«.a©tions «liieh

luiv® %e«n identified in tli® propionie aold fermentation ar©

ai®t}iylgl|'©3f»l, pr©pi©n»ld©lird« and propionio soid« klbhm^

alpatft*lifdr03^-propion®ld(iti|'d® li»s not fe®#n id®ntifi®d> th®

•ss#.

f©wiik%l©ti ©f 3.a©tle aald ®tigg#8t® th® ©eaurreiiG©

©f thli afffl»p©t3tiii.# It 4» fosiltol© that • th© laotle aeld ii

forai®€ la thes® f#*«®«tatioiia • tmm «lpfeia-ls|-<lipo-xy-p3?opional(t®-

liyd© toy inyiipatloa «aa t®liydy«gtaatioii» fto® faet that tfe©

pi»dpl©iil©'aiSia ©ewert laetie aaif Calpha-^'lii'ai'ojcif

propiottid miA) t© pmplmi&' a#l<l swbetantlating

th# pr«p®8®^ 4ls»iMil&ti6n-0'f .nlphs-Hyteea^-p-ropiottsiaoliyd® t©-

H«0 - t-04 ^ * «• » 1

• tfios® ph©tphftt®

SS- •CHg-•OH

propioni<i acii

a#thrig1.re3Kal

.zm. »0H ^

pr©pioa«l diehyda liydafat©

01

<3 » •|f I

alf)iia-hrto©3Ey- ppoplon&ldehf^

t ga

I pi»opional«i®liyd@

figttw 1'

M®©M-attlt« ©f tH® fPios® fhospliftt® t© fPOfi©»«ld«lif'd® sirMi'f3?©pl©nle AeM

•fH«;am^©r it l«\F®«s©naM® t©'a©®®pt that

plotti© a@it i® f©wi®€ fer »©i»® t^an ©n®. ®«ri@» of raaotioE®,

fm «x®»pl®, %hpmL0h s©id and pi'otl®»ftl^liyd®# fli®s®

r®a®tioiBS will 'h@ ff»®»#nt®d in a @0«pl®t® s®li®«e ©f iltieo®®

aisiimilfttioE'(figiAF® If Slisptei* Fotti*)#

fh%' •mmnt of BiM«a (1^33) and leyerhof

C193S) liRt^© ®«pliRslae<i the importsne# of phospho^gly©®ric aeid

la Qftfbo^ipsite »0t«bollsa of y@«st md musel# tlss-at*; fli©

se!i#i»e» %hm^ fai»' .propostd miiiig'>li0spii©gl|^e®.rle ®oid as an- in*

t^i^tdiat# lisv#' ,lii©l«d®A ©lily m0 ald©ljjd©, a

ti?i©s© -pli^spliat® of (©•i» €iliyirox|^-a00t0ne)

m& Hal?# «xelud#d m®feto.f gl|f©3r«il ©tJier tlire® eai*toon sld®»

hy<S«s fyom tsli# pi?©pos©4.ffi®eli-Bfitsa dissiailatiani,. ffe® 1»©»'

latien.tf ^^plouald^teyi® impert&at miAmm fa'^'orlng

m@ mmtipmm ®f » pha®# of pm^imtQ: mid iwhieh

does aot Imvolv#'^©si^eglya'®?i®'• ftfid# fli« sek«itt«s- proposed

by iSaiMen and Q^h&m iadieat# ttoat the trioa© is

oonv©rt#d direetly to a eoaipomd mntBlnlmg a e®rb©^3tyl gt'oiip*

la: @3!»d#.ip- t« ©totain pTOpioaaldelii'd# • f©lX©wiiig sueli a 0OOT©i»sioa

if wottld li® nmmmTf^ to mrho^l gramp. t^s-s oaf^

•boB|fl'grottp* It't® ir«»f impi»obatol® tiiat th«. pFopiomi© baeteria

earn pfodae# t flj.® pp©p®s®d sohsme involving

»«%li|'lgl|'®3c.al trnm^mT prnrn^m kaowlodga th® aios.t log-

i@.«l «»«»» ®f ®^t*i«d»g pi?©pi©nald»liyd##

Seetiem #*. f|-*%vie. »@id» "

fiFt!Rii®ai ClfS3) .©festCT-ed tiifet pfwrniQ mM'la ©asily f#i^

seated femf ®0'aelnd®d ftliat «fid I# net- m inttritedlat® in

,g|ti©0a# • diisiailafciott# lawreT'(Itg?) and v®a Ii®l C1®S8)

poatttlated. that pyfutle meld i» fo.r«®d duicing tli© dis-simllittiQa

O'f gluoosQ the kete-aeid wai not id®ntifi©d aa an ia*'

prodmet# fan Il#l p:ropo«sd 'thst ^frmi9 mid hj**

dfat®' i» disaiaiilated momding t© fh© following ©quatlon and -/

that th® mm&mion i&#3 m% imQlm ste-©tal€®hyd®i

/@S • ^ iigQoei QOg • m,^

fhie aflstiapti®!! mm pi»©apt«<l b^r his f&tlmm to

th@ ^imanfeltstlire ooiiv@r»l©» of lat© aeeti© aeld

er th® pp«»mo# of »a®taM«h|^a# its»if durlag %h® f©wetttatloii ••

of gltioos# tlthowfh mmM b® » nomal. ppodtaot of

&mmTPho'XfMtlon ©f py-iniirlo aof4». 0th®]p wasoiits h«s©d tipou.

sp»oml»%loa were off©i»«a for the aoh®me hut the ©vldeii©®, al«

thouah attp«©%l¥«|. was m mmm 9ub$tmti&% proof. Aoot*

aldehyde might will b« an pyodast taid mt h® ©on-

verted quantittttlv®!!- to a@®tie toid'-apoii its addition to a

femontstiott# m&h & mm@mlon m^lms & hrdrog^fi ©ooopto'i?

which mB.f oy.»«r h« ft-railmhl® Im ®iiffioi#iit ^satitsr to

oxitis# 0«potiiids ©the3» than thos© arising fi»oa th® •sMgai?,

proof of th# iiit#»©diat@ fowation of pymflo sold fro» '

•Rlnoost woiald l«iiA, i-mfpopt to Tin lisl»s th©©i^ assuming

ismth ,p»of it .is ossentiftl to pmm that ifto«tftld0hyd@ i® not

pro.d«o0d:,.f»om th® p.yrttirl« -ROld In ©yd#!* to establish oomplete

•#v4d@iiOfr for th# thmTft, ?li?t«ii#ii ®rtd, Ifepstrla -(19311 fouad

that d3?y p«ipi©iiio" sold fet®t#rlg do not doooapos© pymavlc moid

taildss phoapiat© 'Is sddod and polatsd out that these haotoria

•do not ooatftia

fli# fimtloa appealed t© most promising'

M^aas of ituijiiig fcto.is pi^felem#. 0ai?to©3Qrl.as« is imported

letil>«rg aM R®lafmrtto |19tO) feo a^fe mpoa tii# Msuljphit©' a.<i4i»

'tloa QoauemMi of pymfie inei'd t© fom •fchi# tolaiil'-'

pMfc# aoapeandl* In tht« pp#««aee ©f #a3?teoxyla»0., fissafeien ©f

pyMVic aeltJ bj stilfit® thotild mt pi>#v®iit sla«it«a©0M® foma-

tion of ae#fcald«liyd®» Qn the othap hand a fiction of pyruvie

eei<i ima4coBipa»i®d 'toy the fomation of aoataldahyd® wowld of'-

f®r••strong indication fcliBt tfee^ toaet-aria do not petsas® th® an--

gyrae earfeoxi'la##.

faMeEtattaas mm out in a aaditm ©f •©•S par eant

Baeto I'aasJ axtifaat, 2«0 par ©ant tlnoos# and 2*0 par mn%

CaO0^« . tt^aa? -sMi-aii felsmlpliit® (€!•§ par aant) ©r ea3Laii«

gulf It® {3.«0-0#fi pa®^ aawtl was inalulte# In tha saatim# A ;

niOTtoai' af SOO ©a# flaa&a aaali' aaistaining 300 m* of medium •

mm pmp»m4* Faiwaatatloaa wmm at S#' ffrawtH waa lux*

vtwimnt wi-tls iftosfc ©ttltwast altiiom#i •©••rtslm ml^ureM graw.

poorlf (S'f W .a»^ 48-l|f IgS toaing pi»a4ti'©®4 iii thasa farmenta-

tions# fha Msiilts wa^-a substantially tlmilai' with the twa

fixatives, flia fallowirtg axpariaafit C3^ W) i® t^'piaal* fiia

eialtura... . • was iaambstad tail days aad •than .i?aaa'vaa foi» anal-

Srsia# fast^s for following dlstillatian of th@

aatiwa afaaMiag ta tlia matliaci ©f and prad iXQm) 'war#

nagatiira* Upwa addition ®f ti4<».dlBity0'^amylJi^di*asi»a to tfea

»®dtw».aa almataat praoipitata wa-i ©'tetalaa'S «felah wa« dtssalvat

im satitta ea^bamata aolution mA i»apraaipitataii liy aeidlfleatiaa*

kftmT float loft of ei4-ainlti»opli,@Eylhydi»asliie ppecipl" .

tat©j the melting point was a »l:xtui'0 •with ewthentl©

pyra^le a<sid 2i4»ilnitTOpli#:aylliLydp©.2orit (if^p^ S13® (ef« Alleii,>.

imO)) melted at

A was likewise pi'epiarfid from tin®

femsntstloa solutieuf It hai ttm l#f* 218#5® alone and gl8.#5®'

#i®ii Bised with autli#ntle a«i4 p-nitropheni-lhs^drazoii®

(MtP.

ffrmtQ^ was fixsA and idtatified in glme^s® f@m©nta»

tioas- hf ©iiltttre.a- 34 l, 4i^t and 11- i| th® r®sulta nega-

tiw.-'with 5!? w aad 48 *» fMt ft0l4 ha® else lb®®!! t«i@ntified

1« maanitel, laetl# aftid and gl|"®eTOl f»ra®i3ta%lo«s tasing

ctiltmr# S4 f# •^isntltetiir® also.war© m&de in

lona«b®s* ijf f@m«iitatl.ons* ftia rasuiti er© shown in.fabl#s

xiii and tm* •

. .fiiia 'identlfieation ©f primal0 aeld supports the soh©m®

of glmeo*® disfiailation ,p»fO€©d by iran Ii®l# fk© fomation

of p|-«irl0 "aeia mfit©©apftni#<l hf ae®t©l<l«liyd0 is apparently

til# Qt %ti©, InsMlitJ ©f th# pf^pionic aeid b.a®tsei*ifi to

«s®a«3?li03isrl»t«' tliii. pfvml^ • aeifi «ttlfIt© empmind liiisli its@lf

i.s..n©t. siil»J#0t t® tofdfation end stibstq^.cnt 4®ii|-ar©-gsniition#

. «qp::«iititative aim* that, a donsldsfabl# gaoimt of

psrinw-i© •««i€ .1.0 f©»i4# fh# fi^sati.©a ©f pfrmiQ ®'Si.d. wmM tm.

®xp#0t©i • to pmMm m tmmmm- in the Mtl© ©f pro-pi©nis #©ia

to a.0®ti© This i»«tio is-also in.fla#n©#a by otlmr faetors

and tli0 VBmlts may ^faFifttol®*

tmiM iiii

mSSIMILATIOH OF GLUCOSE, 11 W, 50°C., S.0 Cfflf SKGOBl, £.0 FIB GEIIT C&CO«. 0,7S Plii CFi^^T 1

-w •^Matlties - npoaaeei pay lito-least jP^iod of ttlai.€©i»tilm@#®«i f#t«3. iri'opionlctlceticiForslcjI^oticjpyravie «tfTOttia«gia»tioa8f«iv • « Smt-' tmlmb&mi aeid i eeidt acM i aeM s acid

Eatio pTOoicatie

•t •s ^ i : t i J 1 t 1 f rj^ e«-|t. tet s •®st .-f, •rfi ' i aM ,•§ sS f. ^ 1 ifi I ^ s itf • » ,

' - s • t f • s ? $ 1 t • 1 s t*S- f m t i ss^4 s ma t - m*4 1 1S»S| §».0 » lt,S 1 50.7 I • .8.84 •

1 t 1 • t 1 t s s • S • 1 m f •69,0 t im*S t ist.s t im*s 1 • mat 4*Q 1 Sf.S 1 S.SS-. J- t $ 1 'it • s •« f. • i

^^tKsciaie MiiS„ CX)g, «»i' a#»-r«ia^af mtmriA mm m% ictewtaM.

f A M M at

FlllflOI Of Wtmm AQW» GLUOOSB flMWlflOI, AlAIROBIG, 50^ 0. ®lSf M,tl, 3*0 PSK OSMf, OaSO^ g.O FIR asif

.,:.,,,w^y,.mRAC?) 0,5 mn G|MT t t f F#rld<i of I ffrmM mM

Culfcujp# f. FixfttiTii ito-ettbation i aoi I

m p&T iit«r

1 t 1 t 49 W 1 #lM©08« t Gaa©^ f m f !§•§

f 1 ^ # 1 1 • 1 IsQ^ f t

m t. 1 ®5.B0O#« 1 S«», i i§ 1 f 1 t 1 t 0.5^ i 1

m w f laanitol f MaHSO« 1 80 1 ss.i s t 1 f 1 t 0.5'| 1 1

3 4 w il^ftstie «eM| laHSO^ 1 io 1 IS.8 1 t •| 1

fli« witli' 0«M per mm% fmmt a^mitFieed In

fabl® XIIX mm t© iat®»#dlat# e«powid® in*

volved is %ij« ©f 3.a®tl.© 'aeid* l®tliylgl|-©3Ciil and

alpl:ia"liydF©:3ey*f'ir©pit3iiald®li^-i« ar# posslfet# iat»i«®diat«s aat

w#l,3. .m pymrtQ «td* itloa® wai ^and

evidence wts by tliia Metliod tov tli#. iiit#w®diat® *

oeourrence ®f ti« ©,ttot©p eoapemad®#

With %©• th® mo4& of faiwatlen @f pyrttvio «oid, th« •

foilowintg 3P©aeti#i:it • i» Fl«:u3pe II s®v#ral po®sitol®

meohaniema#

gt pmmnt. tUBipm is laeithep QvMmnm. for or agelMt^ th©

o0©iiFi?tni©® ' of plio»ph©glye®pio aeid in th& pmpionic seid ,fai^

awatttioB# flid stro-iig pMof Qf it» iMp&TtmQ^ in th® ©art>©-

dl»siailati©iit fei»ou#i% &hm% by ttssu® ©nisysaes, h(mev%r$

•"ft-*

1" IS '•• K®® ^ ••®M 1 |,Q_* ., ... §*a»» i«:|

'#8.

» • •twlmrn

•pli^ogphtte

OH

trios® -pliosphut# •hfdjr'at©

•1 .0 •* * oh

phos^©gl|'©ei'i© ••acid

-«S^'4

T 5 .0

GH^*0»C!!' ^3 H

MBthflglfmml

OS • • • ©

# S«# A .OH

® *on

»ethylgljo:^sl hydyat©

0 0 •#

4 1^0

alpha-hydroxj-propionel dehjrdo

-M,> 0%-c.o, ei

Pl'i'iiirle aoi€

f21

k

®5-£' s oh

o;H «i

alpha-hydroxy-proploneldehyd®

liydrat©

OH 0 — S 1 . * G * 0 * *

oh

laetle «©id

ftgttPt II

i«0hani«.'.tf Bl8alml3.st»l0fi #f fi?i:6t© fhoipMat© to fymirl,© ^ an<| A®i4»

*95*.

that it. siiQuM to© given oouiii{i®i»at;ioa. flitcss|>li0gl|'0»j»i®

aeld is ineludtti in tli® iofotia© to show how it. may funofeion in

tills • f©wt©ntstion«

four dlff«i»#iit sei»i@s of reaetions era- presented in

Figtii*# II itil-Qh l®ad to th© fomatioii of Isotio and p^rmiQ

aeids. fh« autUor is of t!ie opinion that im tlx# ppopionie aeiA

femontatton, it is tmlikely tli&t oalj en® series ©f i*®aoticnis

is iwolved in t.h«, fomatioa of tlae ®iid pyodtaofcs, it s©»ms

peaaonnW© t.o mBvme tto&t %h& mn4X%tm« in 'rtiicli tk#

tion ii f»>odtie#d will govern to @ larg© tlx® r®aettonis

whisli'will pr^Aoaittate. - fit® pr©pi©3aie aoid-tefttterie. mvf

prohBhtf liftfe m ®cpip»etit Oftpabl# -of bringing about

TOoit of t'h© ©liitiig#® • i?«p»s©i3.t®A in tte© s#feda«» fMs i» up-

pafeat fi*oa the foliawiag ft@ts* Thm propionie »©id hmtrnv-lrn

a-ftB y®du.0« ffTOVie ftsii, to protplonie. gcid, tli# reawetion pre-

Ibafelj' pmMm th^eugh laetic .a©id; tli©|- mn mm&^t lactio aoli

to pyimvl© ttoid. as pmmn ' isolatiom throuRh f i^tatioai -mnd

th©!* 06« #oaf©i»t a»tli|'lgl|-0!jrsl to Imtio bqM (W&ub@rg ai»4 iopi*'

•(19gg))* mMmm favwiag, tfe« ©e.emipiron®® of sl^a-iiydroxy-'

,pmpion&14«hyM in pmpionl® mid f&rmMmtim h&B already beea

giT®ii| •©«» pif©babl^ • 0.©iiv©i*t this . to laeti®-

meld-* aeiil -kat'@"iiof b@#n

4,»t^dstii»tei ms rtt# tMs «v,i4®a6@ t&r th®

Hate 0oaf®T«ioiia ©f the©# ©©apewads is ba»i» fof atsmiag .

tto-gy MMf b® pr0dm®®€ tv&m tli® amgar sine® tli® tter©@ k#r ©oapcmB«lS|i

l^l-ravie igdtie mM and ai»tli|"1lgl|r©3iitl kmir© to®®n; iselat^d

from pt^oploRle ««ild fh® feet® jmst gl¥en oon-

stltiita atpeng p^s*©-f&f of th# ©{Sd-aFpeaoe ©f thmm Peaotlons,

fh® f«a©tloii ifhleli will pf^domlnate will tipon the eon-

aitlons .that nm dptiwain ,f©r til© fiigfiii®8. Stec© tiie optluram

conditions ftTO aot ths stts® f&r &mh. enzfrne a dtiange of eondl-

tloit will ?#stilt In ft ^.anfl-e in the pfeAoffllnmnt. TOtotlon# |,4*

dl.ti©ail eirld.ena® sunneytiRP tMs hj-potlissls will be pi»esented

in, 3»«sult~p to. t)9 <lts0ii:ss«€ later In tills tlieels.

S«et;i©ii S* Ae#%la «n-d sussJiil© aelfis*

iuealai© aei<i teeea irseo^gnizta as ®. pfedmet of plant

smd anlaal «#,tal39llsa t&r a imafeer ©f r@aFs sM mr@ml th©opi#8

hrnm •itavtfie@a for its f©matIon aad fiiii©tl®ii»

( w m ) wi#l«nd (1922) Sttig0i%«4 tim fomatioa of

•TO<i0lale ®#ia fei' %Ii® a@liy€r©g#nAtien of two mmlmmlm& of aa©tto,

ROli .itnd pr©po8#d %hA% .imeaimie aeid was In ttim <llt«iiiiilat«'iil

.as/ in flgmr# III* .

-iaooi 1

mm V

mm ,• 1

..5§« j

0001 1 o«e 1

a§o« 1

. eso I

-i i. t L.II ""a", OHdH

^ 1

0001 1 o«e 1

a§o« 1

. eso

01, J 3 I ^

»"0H, ^ 1

«g ft 3

mou OOOH • «0«. • iooi @©0*'

. Smolnle .^aiaarl® l&li® .©jfalaaetle pyruvic aeld «#.td • aeld . ft-eld aeld #

dioxid®

III .

. fh# liM wi©lan« t®aQ-feioa, i«rl©«

•

a«e#at' ln-srastigations of mold, jeast ausel# metabollsa

kav® ©ff#p®d amppart to this Butktwitseli (1929)C103O|

Isolftt®^ aumlnlQ anil fmai-io 'followimg'tlie-'dlsslallatiC!®

of aeetis aeid bi*. fumgi* fisland."ind s<3nd«rlioff (1932) like-

•wit® d«oi%sti?ato€ th® foKistloji of aiaseinie aeid from settle

mM ^In the »el;ab©.13.s» of mus^l® all tHe steps of

til© aQh.m& In Figa^© III iia¥<© reseived ^xpmriment^l support

witH^ %lm mmption #f tk# #Qiiir©i«sioE. ©f aMtle'lnto -suocinio

»®14 li3g|.,*

In iri®w of mmnt ©bstr^ation ©n ftingl and y&asf

' aetabolieiBt the rtsttlfes oht&imA anelfsi® of ®

• f®m®ntatlen wltli eulttt^© S4 W {fabl# If) f&m of pmrtlcwlay'.

interest• A mrv^f of tlie data tn tHia tmbl© dlBOlo©®® tto®«

faets of sf«9l^l iMpoi^%mmm (i| ftie q«e«tity of acetic •aeld

QQmrrlng in the inereasud aurlng tlie ^a.T%f p»]pt of tto«

f©xw«atatloii teit this wfe® then fojloweci hf & d&sp0m0» (8) fte#,

qpftiitity ©f auceinlQ asld likewli© tn0'i*©ased and then d«eP€aa«il»

(3) fh® i*«tl#.«>f pmpi&nie t© .noetic acid iiieTOftsed ©on- ,

thJ?0u#i©iAt til® f#fts®at&ti0n» if#.smlts show

that th« fto»%.ie •aei^® way© pm^mA and In. %uto"

mm «ting ms ccapomn^s, fh® inei»«a«®.

.... in the ©f, pmplmlQ. aeid t® a@id stigg®»%.8 strongly

• tlint til# dl®slBi3.®tl.0a ©f 8@#tle. seii mm o®©wring •.©Ten toefojN&'

it' was ttstmalli' sh^w& ^ a <a®#r®as« In the quantitj of tMs

80id» th® a®©tl@ mM and swesinto mlA show^ a <i«or®fts«

m%T at ® •time wfe®n w#r« <ltaoMp«8t'd more rapiaiy tifttftn

imM X?

M^iaiL&Tiof m SLicosi, u.Mt so® c., e lities of wwi, s*e pm ew tLocosi, .o».4 fm cm Km®, pa est

.M<Wi»iii».wmmi.iii»iiiiii I Liiiluu.niiii'f iirimiiimr iT. ww»«Miai«iniwiw>i-nwiiji' iiiM'itwtfu'H'i! i.ii ij." in" I'iniuii i ii wt wiii i»inuiiiitiiiriiiMirw>iij)Mii;iiiiiii>niiwiiiiwMiKu.ii iiii>iiiii-'iiniiMiiir • t' t t .flttsstitles aro^ieed. per liter t 4 C yecoir-

Periesd oft Slucose s Slucoge j Total :FropioaicjAc®tic:Succi3aiciLacticjioa-r«- i Istio t ered in iacaxte tio Bj&risebted s f er® ea i yoI® tile i &eid t aciJ.: aeii : acid :ducing 8»oolottie8a3 Mlysetf-

t f 1 •ar % . • « :m&teriait ae#ttc

Days jTser c®it f • 1 ott 1 ^ t sM f ^ : sM : itser e«ot i t »

• 1 i J i i s § « .t 't s t * , 0,i 1 0,i 1 0«O 1 »

t 1 t S " t t • s- f t s' - S * S0.3 't 4t,4- f sa»i t 4S 1 .1 isat :• 4.1 s • 1*1 t 1 t»SS : t. iS,4

|. i 1 s t t s s 1 f ? • | 4S.S • t ?4»f s 104^ t 7S*4 s 2S..It g.,0 1 1,S> f t ?E,f

t i •s i 1 1 s « • 1 i" t ' 4 .S%»4 s to»s f its^s s lOl.f s , 9,l i •t»0^ 5 f.f, 1 • S»?S • s

t t s t: J }. •s * I u 1 S4,f: t 10S*8 f iso,« . S Itf.S t tfar s*.a < t I*S.. $ 4..ae ?S.?.

t ' t t s 1 1 1 i s i • IS t fo,i- 1 s if?.t t 140.4 - s f*s' t t.f s t sas- 1 fS»S

s 1 : « • . s 1 s s t 1

IS 1 t its#s f i 164.$ s . 'S»6 t i»-0 t 1...S i t.m t 7»,$ i- 1 !• t i 1 s s s 1

Eg t •?t,g « 1ES,4 s- id?,o 1 ma 1 tO.Ss 8»t 1 1»S s sa t t fs.a. 1: £ 1 .S i s s * • s

aot

.th®y mm produmA, fhli appefcmtly oomarpad only In the

later part of t!^© feratntatlon# If thla femeEtatioii is oom-

pared, with tha om detsr.lfeod .in fable ?ll, .geetion 1 of this

oliapt@r it is ©iriieat that th,® two ar® w&ry different and that

th« me®haBl.sMa .of ^issimiliitlon eould not haw bean the .aama.

fhe anaeroMe distimilatloa of a««ti0 acid s'uggests the pos*

sitoilitrf that smeoihie aeid is ijtifhasizsd from aeetie aeidi,

If tMf omnrmA ttit sweeiaie aesid as fomB& waa deoompoged,

for its,' aoemffialatiom was aaatll-. even s% th# time of aaxiiaum

yield. . It is tppftr©iit that th© proof th«s-© smggestioma r«*<.

quires more eonertt© ©'fideao# than hm hmn- given by th®s@- r®-.

stilts* fh« ftstftblishmsRt of stteh a.. pro©f is eat of the pur-

p«s©a of the isveatination r&portad in th® nm% eh.spter of

this t!i0..«,i.8i!

StMSftWs HP WW'" W.'-M!'™ ,

fh® ®f .th® diasimilstive chsngs® prodmced

*.h@ pr©pl#mie' a«i<J l9»®t#ria hw b»0ii stttdied fei* iaolating ©r

dtt.e^ting iatftraadllat# ©o»p©md® of the foMientation., fhi$

^h«a fe@#B -don# lar th©. f©Hewing method®®

(1) €#ai0as%r.a%i«g m iiier®aae »iii atilJs®«|ttent daereao®

Im ©ertftin oowpetnids threwgh »a»l|'s©® mad®'iiit®iMitt-©ntls'..

th0 feiwsntstions ,a«riog the eoura® of the. diaaiMi.l»tion.

CS).B|' varying the. 0o«.diti«?iis #f th# f©•mentation, th«»

eawsing in «00iiiiitt.l8tl0tt of cc«ipomnds preirio^igly n.©t found to

©ectir-as'sM'preatiets of the mvmmt».t%Qn»

(3) ,Br flying with WalSOg, OaSOg? OJ* diiae<4on, al«l®hydes

and t&tomB proAie©d lfitei*ii#dlat#XT tefing Ih# dlsslmilationi,

fhes® methods hnm l#d to obserfttions whicsh support th@

concliiiloii that the ,fo3Llowin.g eotepomAs ftinetloii i!at©y«tedlat®ly'

In ftwemtatlon® hf th© pfoploale mM. hB©%®ria«

1* A no!Gf»F«tiietog

t* taettc aeld

'3, proplofialdehi-d#

4* fj'TtiviQ'mM

.5, acetic aeid

a,# SnmlniB aoid#

?h® probaihl.# waehanism ©f the psaetions ImAlnw, to the

tioa aM dlssiailatiea of.these e<»p<5md..s, is 4n0e«ss@d ana

th® TOSiilt? aF@ ®©nsld#Fo4 i» 3?®latloii to a sehem© i»®ps'@s©nt«

ing. the Qoaiplet® ffl#0hatii®«s of gliioes© dlssinilatloa.

-it*

fh® Umhmlam &t tfei® ©isi-iallatiir® Slifta«®s ppodtioed tay til# fyijionAe AtH. ia ,Muc©ae

fh© mmhmi'm &t %h» mnmmt&mB Irn&Aim to tii# formation

©f propionic • aaid, g>@«t3.«i sold »na OO-g feav© hmn oonsidered In

Ohgipt©j? fhme tetit til® liip©rtaiit of iseomtlng for tii#

®it0®liilo ad id ifeleh ©eotira «s a f®i?«©nt«tlon pi»0'<ltiet urnst to®

in¥#stig«t®d hetor® m eoapl#t® aedliattlsm eta b© given for th®

glm0®s© dlssinllfttloiftn fh# mnfu&ion In th® liter'attir© r©!®*

ti^« t© tit# »©d© 0f fomatlom of smeeinlt »©id has fe@©n pointed

®mt In ?«rt 1# Wmn il®! @®a®lmd®d tliat th# lueQini© aold 1«

net ft ©rlgln tet ia f0m#d from ©csttpowntds of

th® j®sst ®^tra«t, itiil® Virt«ii®n well as Foot®, Fr®d,' and

f®t«r®on liav« r«p®rt®d tliat t!i® sia«ieinlei aoid is of oarljohy*

drat® origin# lirid®ne® swpportlni tli® latter iri#w was obtain®d

te|- til® ftutiior Mw-img a study of th® n4trog®ii r®(pir®miita of

th®a® Meteria C<3.J»apt®r fw©)« fliaa® data kav® t>»@n pr®s®nted

in falil® -:IV» Wood and warkaan (WB^ U»r& polntad out that

th® f^amlta obtainad by flrtanan d®«oa«tratlng th® formation

of S'ueoinio aoid by a 4- and g- oarbon olaaTag® of th® glmoos©

molaoul® aOGording to th© following aquatloni

Vl2®6 0.001«01g»01giGOOa 'OHgOlO 4 HgO

o-gomot 'b® @oraiid»r«d a« oo»pl®t® for a@®tald®feyd® or ita oon-

•varaion pro^toot i» not aooomntad for ?lrtan®n»s data* fh&

probl®ffl r®latif# to tH® deoomposltion of ao®tlo aoid i«d ano*

elnl© aeid ©to'serfecl to th® f©i:^©i3.l;®.'feloii in fiib3.« W

p#^lr®8 Ift to# interpreted

aii-i he im ». sohewe#

S%. Rppttred pfcifeftfel® thtt ttj# ioltttlon ef th®#© proteltms

e^mld he attaimM ffloafc aeeumtaly tlnTOtigh tli« qtmntitfttlw

i«t«.iwiiiatlo?i 0f i.11 f«w©nt.atioa pr©ia<ita *nd th© tetting up

©£ Jf®w®fi%iifti©a Wf this. »®tliod th# iurittlysl# ©ould

b©. ,eli«ek®i it$T rnmwtmf thr©tiii4 tl»# mThSfU aai ©nidation and

P«dm©tlo» .'^and %li« p^ssiMlitf of arrors fe«. ,

%& m aiiiiw»»

Ass^ittg ^astisftte%.#iy, $&!•&©», ani iia<l

fealaaeas ara alfeaiaad it: l« »vlt©»t tHat a scteam® ©an b@ da*

¥ls#d whioti will satiafj th© ©f %h.@ iata# ftiis,

Howafar, i©«« not pfov® tti# mh^met mwmQtg for tti© iufcaraad-

iate mmhmiBm in arror and yet satisfy tha absa^rvad

qttantifeias ®f and products# Hawawr if th# aohaaa utilizes

ia%®»adiata eawpomds wljish ha^a toaan idantifiad than it a®^

toa @i«Mildarad a^arimaafeall,? astablishad .,f©r it ehaelcs

all faats • 'ralativ© t® th® faimantation aaehaaiaa# fha a:spar-.

i»ants af -this aaation mm sat up t® aitafellaii if posaibla

amah a sahaaa:#

fha i«t©rast ia tha »©fhaEla« of sttoeiaio aoid produotioa

made it pirtiaulRrli- l«p©i*tant to liii?®fltig«te th© eause af the

dluarapftnoias la th® rastilts ©f diffarant Iwastigators with

raspaet to this aaid and ta datarmiha tha eonditians whieh

iaad t© tha praiuotiou af^^larga Mamats af smaeiais aeid#.' An

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fmmmt

wmmzmmw or tsiiKifioii, laitxjg mmm

* t ' s, s t. M^tog tleXwm ofj folwe of feilturs'jTesper&tuxe: Perioi ofi Qx^gm s Age of t §i»»s6sBaHCQ« t Baeto- « asedtum iferiRentatioii

iSo,'®' t of tlncufeatiottt t«asi« liacciiluaj s " % yeast iferaenteds flasi: siaeal^ticii $ 1, s. t sextaraet •

•*

t 1 t €mm ii3#r it$»r eeatsotr eeatj t , m» t " A s S • 1 t « s i J 1 ••

«4% t .i M S t t*S. ^4 t 0,,S • * • f 1000 t I • 1 . s •1 t

M wg t: s® 1 i.«i»erobie* •f i i t '•0.*S • * i 600®

t I f®-

J : t/ 1 ' t f i' f " t

' I f®- i a t. *«foMe. t 4 f #. 1:,8 » s 1 soo t s " , • ; ^ s • J i s t • |

49 *4 1 If® * U ' t®ia«poM.e-.s ! g.s i £ ,0 ' s § , s ' I im 1 1000

t -

^ • 1 S 1 S • ; s S •• t " s

49 % i ^ • s tssaerobici i 1 » 1,S . s O.S % mm » • 2000 < • i. « s s •1- I- s • t

« %, 1 1 m t-s®s«robie* S i • t t 0.S- • 1 urn I

> J . . . . , , S- '• t • s 1. % ' J -,;l,

a®«lga«t® amber'of iate fftseatatios*

tms xfii

mmiMmmm m mMmm m tmnmm acid mwwm.

Cultaxe * t .feastiti«a cscrodae^i oer : t £ : asd Gluoos®tGte©o®ei Pro- s£,ceticjCS% t Ion- sRst^io i Istlo iC .rewr-sS recov-jOxidstioa

fer-aea- t fsi^ jplaaie« ««id -1 . leisiefraSee-: 0>« iproplomi-^&'eA in smt&& m t fslue tatios a©at®«i-:'s«i^^ s acid i t s aeidj iag : to ; to s*i&ly zedi f ucciaics reduction

Mo* ' s ^ s .1 i t J' iiater-s acetic: acetie sproductsJ acid s value t !• 1 , f. « « «

*• . * acid : seM • -« : X 100

©er««sti ^ s ^ s ^ 1 J ^ .s wM t • i®er c®atSB©r eent: s s s , s • 1 s t J i :• m-*•

S4 Wi 100.0 J 1S9.7 s lS9,4t 4S*« 1 s 10*1 8 «• S.fii ' : 32.S 6*0' t t t . t t 1 t : : t • s I

S4 1g 37.2 : 111.8 f m*ti ^,0 .sSS»®i S»St t l.CK)i 2.4t •f 88*8 $• S.i' -f ' « • 8f«§ t i £ s 1 s •» m t s : !

49-% 100.0 t 98.2 M. lf?»4# 1 1 1 ?• S,90 i ; 88»1 s S..1 * • »

t • t s. t 1 • « t t s -•

49 W4 .100,0 t iS-9*S » t f 9»&t- na s J S.SS 1 m»$ t 4i»S f s t s t i f 'S : t: i t

m ®s 10©«0. t tf.S I. lQS,ff ^*4 sSt^ Q,fi44 s- 8S,«- 1. S,«f. t -mt s ''1 . i i « J : : s.' J f.

48 *g l£30»«..-t S ::.8fi..,6: 54;.8 s40.8: 6.2: ,14.« : 1.1^: 2.5r r ®t,S- t. 4.f t _ §?•§. s ^ .. t . t JL .,J t -t «

• 'I ..A MI £m omiiMms. ®f expearijwat®.'

o

i $

ir ' u

«w tJ o ®

III I? «H

Q ^ (G

ss

w (to

«a o o o

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8 Q rH f~4 3^ § § •»*•«• .. «» •« *« ^ . «» - «• »*' Ml' •«

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§ I i i § I 1 § § §

M| itfs .h© e» O C J O O ' O ' O O O o o

J© » t iH t. r4 <#

•». r4 • t4 o ' «

4 & O

J "3 # &

«l m **• • M t* . «t «• •• •*•• •» Wf ## Mt •» » »# • * » » » *

# O m

© • »

O ». m .©^

m o a

« « to d * o

m "«SP

•##. m». W •»' m m ** •** ##. ## «• •* •» ••# «• •• '•* *0 #• «9 1M»

%ei .ify m m 1^3 m

S ** ' 8 '

t© $1 ^ «0 to

•«. •#. «* #«

fi» s I 3 1

I . m m m • m- m .m m -m *9

a s s s s s s a s s a a

I % I °s I I; % 15 % "8 I I

•» •# •* «« .40. ** #•.. m • •#•• ««' »»• 4Ni' «r ##«#«#.«* Ml «

.» I S B t 3 i- $• •! t M - u ^ u n

f iUBfcl XII

Di5siMiL«ioi Of si^^si m ms. PK5PI0IIC Acis MCfaiA.* mmm

i i tetastitiea arodaoM per liter i iati#s .latio sS *«S0o*^'S0 reeov-sOsliitt'ite® mA sGliicoscsfllacosejPro- sl«etie«C5Sg t 'OSg spro-doaias ered ,i» sered as ; value

ferstea-s fer» s far- tpioaiej acids " s cinxe: diacieg i .^a ,t to jasalyaeds succ Inle jredxictioa tatioa i fflfKBteflssmted j seid : s t sei(a:ffl&teiv t».©feti«s acetic : products t acid i vsltie

m» s" . . f I t I t I l&l t &&!€ % acid t i s g: lOQ ii^pceatt ia8 t m 1 1# S lA t 1^ s : .. • » tw&s" Q<m%%mr cents t t t . s $ - J 1 1 • s s •

54 s 97.4 t 14S.S s tOg»:gr' ES.tt9f.5t IS.Ss 4.0 » 5.6S s 7*84 ' . I 9f...5 s S.8 101,f: • s t t t r 1 i «

« J S' I s 54 Wg t •as.g s 10S.5 s i®S»,8« H^ltSa.St . 8.6s 4a $ S.26 < • t f7.4 S.g • 90.f 54 Wg

: • t s » 1 s s . * » • I

49 Iq s 100.0 s 1SS.S t 4.4 $ $.1S s •• 7.70 .-S ,8S.O ? 14.1 s • «

* « t « '« « »' • i S: 1 s t

t 100.0 s 156.S s im*st SS.6ie6.7s S7. 4 J 4.S t 1*8? • .&.1S. /S Sf.4 s is*a s D5.4 1 { 3 s i t : t s J •

•

m t l{»-0 J 169.7 i t07..it. g0.6s70*5« 44*8« t,4 %• 5.41, t 10»11 1 f 17.4 I 88.8 m s I • J t t # • / ' S •s ; S ^ t s '

«

48«ig s 46.9 • ?e.s 1 Sf St t»st • t t4»-5 1 ' f,S i '.S8 4 48«ig .1 ^ - t 1 " s •I s. s s s t s #

««lf s fl»0 i 10$.®^t 4S«li • i $ s J.92 .t f7.»t f i ««lf .1 £ •1 t ••s '$ s i i i s

8t% t St»4 I ms s. iia>«i». ei»StiS,4s 20»f1, 4»i 1 0»4l s l.fS ..8 »6.g s IS-S t S4*g 8t% 1 t $ t f • i 1 : 3 S s 1

St I S«.,9 1 104»,S f ltO*:0S to##!. 1 !©»§? t s. £•96 f iO.© s is«f' t St • t %

• • I. s •s 1 • 1 s s i 1 s ' '

St% t 60.9 1 91.f S. lll.ft 1 l£.4i' S.t s : sao t 88,7 t t#,d 1 St% # i s f 1 1 t i t t

11 f 100»0 .t im,Q « 15g.Si fl*Si6S»gt lS*fi 11.6 t f*lt • 1 lOO.S t 4.1 t. lOO.l 11 t s' s t i s J s I { s s

il»18 1 100»0 1 MSwO S igs.is f5.flSS»l# 10.*8t i 0»8t s. »•!£ $ fl.S i S,4 f. ®s*s il»18 J- 1 1 .t t . s • *

. • *

-JU. « *

Cf» Win for ©cmditioas of €ai}»is®®t»

I*B1E II .a*

DissiKL&fiQi m QwmsE^ m wmfmmmmms mmimmm, mmBmrn. wmton imimM

ferio^i f' • I C^satllAfeS PgoSaci^''per'lite ' ' s Ratio: $€ i^eo-r"tC recoy-t^^lSatli^' of iSlaooseiQliaco®®? Pro- jJtcetier ; Sa&-iL»ctiej loa- i COg $ f«red ia sered as t TOX^

incu~ s fer- : fer- jpionics aeid s®g :ciaicj acid sreduc-j to spionietanalysed:voiatile;reductioa bationssienteiS jseated t acid ; s ;aeM ?. : isg rsc^ties to j^odiietsiacid a»ds v&lt»

5 t t:' s s ?aater-i mW saeetict s 00^ t x 100.^ ; .» t • t I • t t i ' s i&I t ' I aeld 8 !___ j_

. §mm . ^ i s itf £ s siM .1 "t ?. • t tmr •em.tsrmr -©eats t »

• i t • t 5 * i t - 1 t r •• t 5 1 Sl.S s Sl.S t ^fi»si:-is,s $w*m sa.t i»o I •| 1.17 f, 5.SS t 71.6 4 66.6 1 7e»t

5 •J t 1 S S • s i t t J. J «

l£ 1 61.1 : 98.g f 110«ft M.S 4.1 t f.S £.t 't l.ts s S.#S s 1 76.6 $ §§•0 t : ^ • s s s s s s. s t

tt 1 78.2 i 12E.S 1 160,6 f £9.4 tfS.ft • 5,f $ US t l.&. "? -s fi^iS s Si,»l s 6S.3 « « •M.4

: \ i •1 1 t s- . t • *

:« * »• • •

» •« i • tt

•»'

fecto-i?mst ejtfe psi St fei*Beatati«a^ 1 i i % t '' i t i • t • s 1 t

80 f s t i t 1 C}<»5 -t- Q,*!^ t 0.0 ; t- s S s. 1 t i i i . t ^ •

• ; 3 •t 1 t

•K- Total volatile acid = 3,5 m.m.

•10?-

Thlm ms® ©f a uwjaber of taall f#m®»tatioris Instead of on®

Iftfg# f®M®iit®tioii was Isy thm €iffi-e'altiy ©»« ©a-

eownt®i*s in i?«a©irliig aptpreaentativ® saMples of OaSOg for th#

^OOg ftHiljsii» fli® aedim ®®mt,iiiiie4 J*0 p#y «©nt. glueos#, 1,4

peT-,j$m% GaSOs aiatt Q#4.p#3* ©t»t, #3cti»iaot» fh® ter*

aentttiona. ©ondu^tsd ftnterpMealli- at'30 0# A feRsent®-

tioa <l©8lgri#d to <Ietewi.ii« the quimtity ©f p?©a«iits wtiieh ©sn

fe® pm^mA fmrn th©.. feast BXtmst was t«t up in m meittm con-

taining 1*4' i>©i», ©©at 0«00g jind 0'#4 p«3? ©#iit B®©t0-y®B®t ©x- '

traet.#;, result® &m reported ia fatel® M.*

A mr^ey .©f; fttol# Ifll.sfeews. that ia the pmsmrn

HaHGOg ta«ff®r f«Mi®.atati0Ss ,s»@s«lt©d. whioM ar® ©©Mparabl® for ^

til#, Atff«r«iit @mltares atrnfiied «ad la »«!• r®8f#©ts • similar

to resultS' ,r»p©rt#.4.. by ©tlitr iiiir«atig«t©ri. fii® ratio of GOg

to a$«tl@ afid, it ftpprO'Xiiittt^li' on®# tlb^e ratio# of propionic.

mM to ««tio. »#id, m» .within. tM l.lmit.8 0fei®nr«d toy van ii®l

md the .qtamiit.lti.®@'Of • sm@Q.imt0 aeit ^«,r® sot. larg®* F®rm®nt«»

tloas iiif.O«0Og Mff#rj,., MXt ©.-a tht eoatrary gave results

varjiiig msrke-dlf .with th.® {Mltww-s# the ratios of propionio

aei-d, tO/.»0®ti0 ftoM 'vmief from 1«78 to th® ratios of OOg \

to «««tio ftOii froa Q»Sg {aoTOraei' of 0,41 is qii«stioiiabl©) to

©•gS •.sM th® •ftmooini.o m&i.d frea 4«1 p®r mnt to 18#g p&r mnt

of th«. § of th® f@ra®Btei smger*. fh®i® rttnltB itr# •ualilt®

thos® reported is th# llttrmtwr® anfi .s0h««®s liiioh h«v# h®®n

8mit®st@a thme far, by iavestigatora of th® propioaio m@ld f®r-

«®nt®tloit ®r@ not Mdoqumto t© is®®t the r®^lr@a©iits of th®®«

data# With smell i»«-smlt.a tlt« m&mm&y tti® m&ljma mlipit

b« qw0,ttloii«d but ©liteMs bf th®. eai»b0n aiid oscidation and r®*

Aiotlon b«l«ne#s showtliftt most of ,the an&lfBBS are smtlsfao*

t<5?y» fli# p-ttrlti- Qf til®' 0wlttti»es Is liaMly t© fee qia@stion#«l

f©j? tli« f®»tEt,»tt©iia W'tr# tildfUR%(®ly cheeked t® «2.isiE®t0 en^

•eoatsBBlniit!®^ Mepwen&nted m&ng th@ ©wlt«ip«s. ai?#.

these fro* thp®® difftTOat iiiv®®tigatei!'8» ill •©«1-

•tmrea hme .l3Nt#n &li#ek®d toy a ny&t&m&tM study ®ith©i»- toy werkmoa

snd ©y simm ttet tia® Iweatlgators In this

• With »giiyd t©. th@ ©aubeii'feitlaaet-s, tli# eapben P©eof@?y

la ,©f'eeartt mt Kioaplet# ia- cms# ail pm^&ts wmm not d®t#r-..

• The -f'ssialts s*«ms.riss®d in ftbl# -IX ar® not wholly sat*

isfa^story in •.gooemting. fop th« f©«©nt®d stigar. f^ty indieat®

tJiat •tmd»t«twlii®d |>i»oda0t8'were. pr©s»iit ^fartlemlarly dwlag'

©srly ..phs#«s^t^' fittlbly tfela Is aoeowntM

f©r ©y-. til© .'fftet;'that ©@11 atiltlplleatiais'was gp®ftt#at at ttels

%Sa® -ttiid ft %&¥§&r. pmwmn^&g& ©f thm mrhm ®f tli® aufaf- f#j>

WHS' 1» aahjlng ©ell .8til>st»»©©». ©mek am #ff®0t

»l^t b« •»fid«.ttt when tli« qaantity ©f is'saall#

.»id»,ti©ii. and 3?»dti@ti.©ii ^al&ae# la f#«B@Rtattom® In iteloli

t^i© oxiilised «Bd F0itto«d pyodneti-.n^- mt bala.iie®d" show, iti'

every cas© sn @3C6®i8 &t p»du©®d'.prodweta,-posslfely, indioatlug

a donation 'nf hydrops toy ©©apomds of th® y«a®t «Jttra@t as

siig^st#^ toy -fsm Si«l (ltS8), and .food, and w^Tfmmi' (1®34^)«

fh# p©.a®IMlity of errors in th® analysis w»t ©f doura# t3@

eoiiald#r#d with »«eli

Qh©ii«]p«lly til® »eh«M®s ©f glue©®# iiasiallation i»®p'i»©s@nt

th# hex©s©' Qtiaia m Ming split into t«© 3- omrtoon aol^oul®®#

seli®«# ®f tli® 4* and 2- «arb©ii eliavag© is ©n.@ of

th# «j?otptloii0» fli® csftFteon a&k&mm require® that every g-

€Sftrfe©n predttot to© a®oc®pmiiied toy a !• Qarteon ewpomnd# It is

a-^idtnt (fatol® XIX) that th« QQ^ is mot @qmal to th« aooti©

aoid'in a mmlmT of th« f©«@ntatlons« fh® large'#xo@ss of

SOg found in tli@»® f«m«ntfttioni smg^sts th@ ooourrene® of a

g« ©arbon intem&filat® oompoani whioh is oonfertod fey syntheala

into a ©s^apound ©f a hi:^®!* ma»b©r ©f otrbons# Sw®®inio aoid •

way •orlginat# by suoli a syatheals# fh® • results in Sootion 5

Chapter ?hr®# suggest its foraation frcm aootis aoid. In this

eas# for ©aoh aoioottl® of sueeinio acid-prodiae»d tw© aol®ottl«s

of eOg m ®oa® other !•» earbon oompotmd will to® • formed. In

oortain f«w@mt»tion» th® ©ataMiahtd quantities of GOg ar®

gr®at«r and in o-l^or® l®s® than that r@qalr®d by this hypo*

th®»i#:». fh® ©oowrone® of 0O.g in qmaatities gr®at®i* than can

fe®' aooomtod for toy th® formation of sttooinie, aold throii.#! a

syntheais frora aO®tis aeid tndi©at«,s th®r® it- another' souro®

offh® oh®Mio«l atrmotwr® of'th® non-^rtdtteing compound' •

isvnot' known bmt th«r© 1® apparently no oorrtlation b®tw®®n it

and th® quantities of ©Og prodtt®«d» fh® smgiiestion is mad®

that- 0Og- originat®.® 'in th® foiraation of propionio aoid. A do-

oarboaiylatioii of smeoinie aoid satisfi®# ameh a s®h«»e and th®

proposal is anpportad by th® faot ttiat swoeinie aeid has b®@n

foitind to aet as »ii intowadiat® oompomd (Seotion S. Qhapt®r

«ir®®)»

*110-

mm lts®lf of aeeetintlug for %im

ri#ia« of 'QOii' ^leh- ar® l@s» ^%hm tli® eal-eultt^d,

siieislnle is asayft## ta hm f&Tm& W synthesi#,* imeois^por-

atlon into- th# , stsli#*. tli# 4* 2** mrhon el®«va|^ ot .

¥lrtaii«tt fomlng suesliiia a.cl€ -wltliemt dirsBtly Ittirolving a

1* mmpornid .%ppmTB m a liteiili' aoltJtidn#, • •

'A s«ih#a® ®f glme©s« <l.i®sl»llatlo» bmmA upon the f<j-r©go­

ing proposal# is pi»©«#nted in Figuf# lf» It ineliia®« (1) th#

4- and 2*. earbon el#sfage ©f til® glmoO'S© aelfstal# as stiggeetsd

by'VirtsasiK:, '12) tli« bqI# •of pjp«v1q -a^ii'.as tuggestei by van

Kiel and sMb0t»»%ift%«<l hf Wm& m& 13.) tli®

intermediafc® mmtwmm of pFoptoiialAttoyij®^ • (4) m Ec»*r@A«oifig .-

eo«poiitti> C5-) %%% sfn%h®w%M ©f smetini© atid from a««tie aeld

ftud {6)' -tli® 4#<l<»po»iti©n ©f sii^olnie t© GOg:,aRd.propif^ie

fesid^

•fh®- ttatlj©'!? hi«s i33.dieitt»«J th&t f©»t«nfcati©a sohemea pi*«-»

s©a,t#d, by oth'^j/* liiT®stigftto?i are mot wholly .sattsfaetory* ,

fhin is «5rid©mt f©s> it is imposslfcl# fc© siibstltttt© fstes#i»¥ed

values ff<m th®- ilfttft ©bt#l«#4 In th&BB exprniments into nmh •

fermentatiea -seteta## aad #btai» ® WlmiQW* tti®. s0h#ra@ presemtti.

in Figure IV mtill««s .tstablialiM iateipaediat® pro^dtaeta Temt t©

pBO'f® that til® a^kmmm fits all tii.® «;gp0i*im®iital dat#^ it was

by' STab«tit»ti©ii iiit© tlia sehaffli' @f 0bs@rvt'A.

Fomentation 34 W7, fmble HX lined) aa-i f®«mentii«

tl<m 11 W„, T«M. XIX (n™.ber« tn p.renth«..«) h.r« besn u.^

.f®t» tliig ptiff®,!®.

'Ill*

.i*6 (13.*^)

(OgSxoOs)). .»©a*r«da«iBf

a&Ssrlel

(tig.S)

• gtg , :* ^ Cf^ ^ 'iSi&l i4f^s;

a trios# phospJi&tt'

pl»0.8|^li.t# OsHisOe Slttco#®.

msS. (150.8)

6, CMO.®| S»4 (H.-C-O. • H.W

ae'W^lgl^oxid

ii.t 0 0 (68.4) ^ hsi>04

IftttMf a©M

•^IgO •

Ilil l«-4) OH ,0

V$'C' ® OH *oa

^ruvic ftcU Ijydrate

SsS-'

Cii»f)

(11.6

;.i»a»B (sio.s)

.c. ® a 'h

al|4wi-ls|"i3»»y~ pro|3ioaitldel^d«

m ch,*6*c

•«

ras>.t C6S,4) (s)c, ch«* w0fl

acetic acM

•a

(liO.8) 0

I9fl>8

mA*K ^ h k

popopioiiaMtii^#

4%0

•SI

p»ofto®lo *®M

(lS9-»i) jai (o»ol

{15.7)

J

« « sii ,, (SQS.S) , (50X,«)

fifaa?# If • $mme oi^mMrnrn

Jii'malting tli.0ae tmbstltu^iona Into tlie sohea®(Fl.®ay® I f )

-mrtmln reaotioaa hmv'& not hmn ttsed thtts sliaplifylng th® e-al-

eulatloia. fhls €o®a not imply that tiiast reactions do not 00-

o«i» In til© f©«»6nt&tlo.ii Of tliat the re actions used In tli© eal-

eulfttloji take plme ifl the indleated pi'opoFtloii, la aalcing

%li« oli©ol£ til© ' tf- final pm§m%B tormA thi*ou#i, a glmn

«#ri#s, of r«a@ti©Ms m&f h& -aiad also- th© serim -by whi&ii

th® cHangos • oscur. loag as-tb©- qu-antitl®® ©f end'pifoduGts

equal that of ttie ofe©@i»ve-4 mimes#

The qu.nWt=l«s of 00^. .oette eetd .uoolnlc .aia have

l&een um-d .m & atmrtlng b#sis-*- Fewtentatlou 34 .w,^ .fable- ,"KI3C

Biay m an exa«^l® ©f.on® In will ell th« gltiees© mmf

been di$»|.»llat©t toy* a 3- oliifitraf®, • Aastaailng

only a 3- mTbm (5l«®t¥«g#, thf ,fGm&tidn of 8S«8- * of, aoetle

ecid and'10#g tti mf gives rli© t©-§4*8 ,niM'of 00^.

fmn t f t i gh t gn i - f c i j i '®# a* -o f GO^ awi . . t here fore , t i nae -

domtei. f©F* ^.lii'the fomatlo-ft of propleialc acid tram stieelni©

«eM 3*0 wM of GO- • ar# fo-med, for «aeh lallliaol of propiotiis S8 3

mtiA —••"'• OP t»i rtK ©f -pipdpionl© aislA imst, to# o pmam&S. tmm .thm sweelwi#', aotd t-© .gif® •%&# <si'bs@i»v«fi 'S'sIm# of

•e,Qg'» Fifty stTeti mmd fomi?'tenths « of a®®tlo aoid ai*© m*

•qmifm-d for th@ pio-iMcstloa of 19#S rail of sweelnle add -and i.S

mf of pfOpioMo fh© 'Wl; of pyrwio aeld hydrat®

muM b« -AlsilmllatoA iistowoAl^toly,, th&mfor&g is tqpial to

4 gs»8 88 8p.#g# fh® m.&i twim® phOBphmt& QorreateA for

,th® non-reduoiiig aateylal minus Sa*g gtfes tli@ <|ttantity of

pyopionlc «:Old t}ii*oii|0 iEath5*lglyo»l, TRii c|u®ntlty

of propionts sold 119^^$} plma th© cpantity produead from sue*

oinle R0ld glTOs the quaatlfef ©f propionic .aeId

20t.3# fh@ -v&lm tQT pr@pioaio aeid is •202.. Sine«

the ca^tson T&m-WQTf -does aot mmh. 100 per sent- tti© e8lati3.ttt#d

and •©bsQW^d valtisa aot ftot oaleulssited qaantlttds

of •l.-and -i ©toowa at th9 bottom of 'th® seli®a®#

F#-i?»®ntfttion 11 Wj^,^ fabls - XIX. may b« &ltmd as m ©xampl®

in-whieh it is nmmmvy te msxm® •a-4- an4 2- eertoon olea'^iig®.

If iill Qf the mM and .sudolni® isit mm proaueed fol- •

lowing s 3- Qiftoon ole.tfag# Qf # sjcos® moleoul® th©F© womlt

to® fomgd 10S*S of OOg# flu© o"fea#Ff.ed "ffslia# it 68#§• I»<$h

»cj1@©ii2.@ of 'glumm f©w©»t®d to smeoiaic and. soetle aold hf

a 4» and 2^ caFbon ol^avagt %h« .quaatity ©f GOg wfeieU

.will b© p»dtt®#d % 3 a©l®0ul#8» 103»2 «isu® 68»§- dlirld«a bjr •

3 glv## 11»$, tfe® small®®!; ©f glmeos© wMs^ ean uiid©i»» -

^•go a 4"* »iid< 2» mrhoti' elf a.vag« and glv& a oalemlat#^ qmnntlti-

of aOg ^qual to th® ob.erwa. . Below this value the quantity

iOf •guotlait aeid. wfeieij must b©; fsrs®#' si'iitlitsl© is so

great as to •eaus® m 0©g pi?<?4ttetlo» abOT© th® 0bs©i:*v®4» 08iag

;il.«i al. s« th#.gltt©®.s@ by & 4- and $* earboa cl#airag#,

•stifesti tut ions .ttm& been 3tiat« in 'tii® as desef'lb^ci above#

It is, ®Tli«»t tlftt tt seli-ems of fewn^ntation hmB hmn'd®-.

whieh »®«t0 the »^«tP0aents of the data of these expei—

:im«titt# i»©smlt8 together with i*©t.ult8 obtained in ,th©

atudr of th® • int©fa®«li&t.® »«tiibolissi support a 4- and e- earboii

eXsavage of tlie .cluoea® rao3.:«eml©, %hM fortaatlon of auQClni®

acid hf s^iathesls from .ftsetle aeid, and the d^eoiaposltloii of

stieolnlc acid to^ propionic aeifi anfi CG^# a ntimtoer of

til® ,.f®OT0iitation8 do aot t^v& m good bfti«jie@s m the. two ii*

Itistratlons chosen, thej In gemrml stttostantiat"® the soheia© &b

given, «

fh® geh©a« of feraentation as pr#»mted suggests five

sex^les of Mactlon® laading to the forraatlon of propionic aeld

and few to acetic sold, Th® author b©ll©¥©s that ohanges of

condltlGn.s of fomentation an.d the spocies usefl will taping dif»

feTmmt' phases of the fomostatlon Into pi^dosinane## • this i«

supported hf th© fact that In the pi»©s@no© of laHOOg ..(fafel©

Mil) th® ppoduction of nuoelaie mid. Is aatex'iallf reduced

sna the dlasimllation is not eomparahl® to that in utiloh G:&SOg ..

•was ms«d as a huif^r*

• 'fht 3?«.smlt« in.fsfel# XX indioftte that th© quantity of GOg •

inci*ea®»s .©3 th© aootie .-acia iooreasas at woBld he ©xpeotefi

from the acheii.© ©f ferm®ntatlo».# ' •

Mdltlonal support ms to th© intemddlat© .behft^^^lor of

eoetio and auoeliile s-oids has hoen obtsiiitd in itudids using

th# • Baroroft*WftrtaJ»g t®ohiiic and by d^ttminatlon of aoti^atiom

thro'Ui#! oxidatlon-reatactioia potentials. fh«»© studias will b@

reporte.d. in •a^pmmte eonBnuiloations#

fh© eoiit#F@loii of aueoinio aold to prepionie nold is p7*e»

mnt0A la th© iohsa© »a a simple d»@ai»h03qrlati0ii. the author

wishes, to male® it that perhaps »ii iiit#w@diat® aeohanism

-lis--

Is ia¥ol^@d In tliis, eonTerslon slmllfti" in mtriy mnpBQts to th©:

serits of yetstlons propdati fey fhmixirf.# .

fhQTB Is *3.0 expei*t»eBtal mlA&nm to the iltermture coii«

ceynlns- the intewtdiate mtoJianisa ©f th« 4» and. 2- carbo.n

elsavage* It Is ©vldent that iweh s eo®pJ.&te change In stimotiim

f2»0B .thst of the glucose sol#m:i.le G&mot eocur without so®# In*

temodiftt© msshanlsm. fTo^t is lacking that th© g*. cartoon

clemTage passes throw.,aedtaldehyd##

Wood and w#rte».6n .13.934^3 fointed ©mt that if amecinie

mcid aiii sestie acli mm toy » 4- and ^mrtoon ol®avftg®j, .

th@ ratio of GO2 to .ntdtis tcii will toe Itss tha.n 1 .and th©

rstio of propionie aolA to atstic acid will b# lass than 2*

fhay • c0Bslil©r0d sueoinic acii and «estio a@id to be final pro-

duets of fer«0mtstlon« In vim of th© results reported hore

their e-oneltjsicjns • are. not rigdrmiily aeaurst®# It is ©fid#nt|,

however, th«t fara®atg.tioa.s whleh into If e the formRtion of sue*

Clnis mM pr#a©ainately by a 4- and g- Cftrtoon cl@a¥.afi;e will

uppresoh th@s® rtsialtsji th« AmmrhoMflmtlQU of ffuaoiaio .a.cid

to fem propi-oni-e a«id will hmm&t raise the mlum. of th®

ratios soatwhst#

fh« fomation of .sueeiaie asitl toy a ®ynth®iis from mmttQ

aeid will sutostsntially in©r®»se the ratios, fh® OOg prodmo-

tion is gres-teri, th® ftoetio sold is .destroyed end, th© dehy'dro*

gtnatien ef a<i«tie aei.4 oamies an iner®a.t#A femation of pre-

pionid a-oi'd* Wxm th@ resialts sr@ O0.nsid©r«d in .this, re-speet

the .®xp©rimental istta in futol# Hi. support th© eonelusion that

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aeld from the s©iire@ of tiltmgen Is @xelt3.<l#d,

fli# ^s«iiiilati©» ©f glu@#s© If proptoni® ba<it#i'i»

bfts been inwstigat^d audi €fit«iir® pi«#s#nt@d mhldh s'tipp-ort tli«

•fdllowlrag eoneteslent*

1* S©ti#aes of pFOfioniQ mM femmnt^tion whleli h«v® been

gmggestefl toy pi»#iriott® investigators ax*® BOt sttitstol® for the

data in this ehftpter^ A ao'r® oomplet© ®o}i®«® of f®r-

.aentatioa is pfts^nted tPigai^ If)»

2, . 0®rtaia • 0«ltmi»»s ©f propioni© asid teaeteria ppodti©@d

sw©Gliii«$' a®id in gimme fe-maatatioas® ajnttiBtimllf froa aeeti®

•aeid*

a, 8«0©ini0 aeit i® d®®<»p©s®d as an iat&f®®diat« aetalso**

lit© in th® dia»i«ilati©n ©f glmeot® to pj?opi©Bie a@id. and 00q«. Hi 4* fh® 4» a»A g-' mrhm ©leavag® ©f gl«@©s® as proposed

hf-WlTtrnm feat hmm s«taMisli#d in femtatation# with a awittb®i»

®f propioni© afid lift@t«ria#

:$m' flu® .«@ehanism of tta,© profioni® aeid f®w©atatioii "Var-

i®a' witli til® environatiital ©oaditions and with the ap®©l©s»

QowQhmmMQ

1# fli«- atelianlsm of tlite dls»i»tl«ti'ir© ehengts protoeed

l5y pfoplonle «eld baeterlt Is intlnmnmA hj th# somro® of

n.l%i*©gen ta the «®dliia«

t, h.m 1j#«n pi«aeiit#d ttiowlng tli® lnt@m@«ilat;©

of th# follewl'iig ^oapemni® t» th« pi*opi©iile a®ld

ftm«iita%i©iif

«) A a©a-F®4m©lng aaterial

to) liftsti# aeia

e) ?i*opi©aa3.d®li|r^

a) fywE'rie adiA

f) Stiseiaie aeii

g) Mtttiylglyossftl an<S «lplia-li|'ai'oicy-pi'©plomeld«h|-d0.»

3» of proplenie «oid f©wi«ntfttion wfcileh hev«

8«-ggfst#d "by prei'icm® invest I iaftsrs ar© mot

fh,« @li®n^s mmlA ^ In

n) ffe® f«f»sti©ii ®f stt©0lBl© aeia toy syntliesls from

eeidf

%) fh@ f©»atiou ©f s"ut@©tiile a®i€ and a©l«l

by a 4* safl i- earboa eltmira^ ®f g3.«©®t®.

©) fh© dlasiailatleu of suetinl© aeid to proplonie

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f. • JtSO* idtntifitatioii of eoapeunas toy mm •©f 2i i* i»#Ff efti#a» S©e« If,

gn@Thmk, Wm ami .1# ^ list# i®it»ag® mm mwoMni i»r A»®i0®as«tty«. Z»

Steftia, |0f,

ia43*'' m'ber dlt i@r G^lieail.# m# «lJEi#Fii3.s>gi« ti0»3§#

ireekaniim, i« 0#, and W«ytoan, O# 1# 1933, B#t®TOin®tlon of g, 3- bu%yl#ii® glyeoX Ifi fer-in©ntalit©it»# Ind» Enp;. , iliial* id* S, g06»7*

liitk®witsoh, la, S.# M« W» „ 3.929, ib#r €i®- -Wwaltrng d©r l#sig®|m's*® dmi*^ Mu®oip-&%Qlmli9T ia md P«aar8«ai»@ tt»d das f#i^ fiaaip@» «tty.fi*®iiwijig md qfrnaatitfttlireR i@stiwi«iag'

• di#a@r Bmtmt Mimhm* Z» gQ?« 302-318.

m%'kmnw@h9 m* i» ma f«d03?off, i. i. 1930, di® VfftilltBls®® 8wis0li®tt lislg*-, Bernstein-1, ptamar-, md ia d#a v®n iteeor stolonljt®ip md ®tnig»n ant®rn pilssea# Bioohtm# !• 219, 87-100*

Oo®k, S, F., •'MiS0« a®ii in %aet.«Fi«l «@t«to©lS.s»» Bloohea,

.J, |4, ligi-3?.

Otmas, Malftg^tt #t. I.«b3.an«, f-# li4ft t«s s®li®s ii®%a0®t©niq^9 «t "butyro-fts^fcifa®. Q&mptp 3?®nd»' Aa&d* d» s©* S5, TO1*7S4#

SaMen,. §•, iwitlolce!, ,H* J« and Kyaft;., ItSS#' TO®!" di® vm^mg l3i®l ©lykolys® in def wti»lm'lat«r##. llin« mmhB&hr* 3£, 213-15#

Fit«, I. „ IBl&m 1f®to®y Sifalfcpll2giAim»«@n« d« d«u%* 0®s. 11,

Wt%%$ A» tt ' 'Wf# Wmw B©y# <!• d®mt, ®@®. iW-4Sl*

fltl^ • . 188<3# fbtr SpaltptlSi&Magta. l@i»« fi« d»ut# Oh®a« 0#s, If, iao9»

f o o t ® , * * » ! • B . e n d F © t ® r s © % * * » • . 1930# fh® fementatlon ©# p«at#«®i fef o^rtaia p3?Qt

•. plonlc aold bsetoria. Zm%r§ M»k%» II 8t, 3?9-3Btt-

•fi?®«d®ja-f®l®h, S,».v®a md 0rla*^$tt8®n, |» 1906* di® in Bwe«tal®rleas# i%attfiiid#iid®. fs*©-

^ pionsau:p®#l»iig* •Bairt# II Abt» SSt*§464

ffl#d#aann, f. 1, and-E»adallj, I* 1# ' 19S9. The d@te«iiiiati@ii ©f laetie aeid# Biol#

88, 23*$8» - .

Fy©»ag®ot, 0. \|nd Tatw, 1,# %* . .

1933, Xfber ®itt@a .ifctifato?. d®ft • Stoffwathstls d®? fro-, pioasaur®ba3£t®ri®iii , Bi«fh®»# i* giT», 36t»I^S#

(|ottli®b, J. 1844. liwwiflmng V0II «©liia®l,»®iid®ij. laltoydpat auf Rolnp«ia@k@r, Snami, Sierlwi@hl mad Maimitii d®r

;0h®a,; Si,. • .

Maim, A» irsa# ^ , 193g. ub®r ©tJ^isriftf^gsfOygaag® i« Mmskel. E* f» Blbl. m, 3S§*3#i»

iitthnfr, S*l. 1938# A oultui*al «twwl|' of fe® prepienl® acid baoteri®., •f* E®©t» 40*41»

Mm,. S# 1863* On the deodwp®#ltioia ©f eitrat® of llm® in eoii« taot with putrefying QWd« ^art» ®fe«ra# See# l,ond0fi

•^©tea©a# M* ir»* p®tera0R, W». H# end Frsd, S# 1» i931» Oxidation tftd i»eductlon relations b®tw®©ii sub* strat® and product# ia th® ao®ton®«bttt3rl aleohol fer­mentation# 3* Biol# 8h®m# 91, 569-591#

laurar, 1« ' ' " » ' B®@ba€flatung®n mb«r di® zu®k«r»paltung Aireh da®

•"la^teri^ propionieua# li@iii®i0i# 2# 191» 83»87#

, o» 1S3S«' pi?©dttfts last ©f etrlJihydrata 'breakdowm' Itt th©"'»@t«bolis» of aw®®!# md in altoholic f«wi#atati©a» Satwf© IJg, Sdf^SiO,

©•#• mad ttrnmlim, w« lf33« • m®w dlt phosphoipyll«i*t@a Swls®litnpi«odMlEt© md a.«t«t«in I%ii«#«i d®r al^ohQlisah«n Blo®li®ni 2»

S13»48#'

:!,• S« aad f» H. liOS# fh@ ©f pedtitiag amimi? methods • • 3,

Ch©m» €i3»86#

f®!®©!!, M.R. 1933. l«tij;©ds fmt d«temliilag lit®ti© aold »nd their iipplio.iiM®«i in Oft f#i?»@iitati©B« Unpublished fiiesls, |©iwm Stat,® #©ll©®tr '

»#tlb®rg, C. und mWf ^ 1925. Ub02» ,4m Mll^slurebildang b«i B®kt«Fi®ii» 'SS.#fli««# Z:» lif» 48g*4S?#

StMberg, C. tmd Reinftirtii, s. 1920» troer di@ Verglrbark#lt der ii*«iistPtuto@nsiur« mt«*" d®n B«dir»piiig@ii d«s Abf*®Jagv@rfeli.i?«»%«. Ber# d0iitsoh» Ohem, Gas. 55^ lOSS-Sg.

itmberg, 0# tmd Wlndls®h,' f*' • • ' • i9S5. iSber dl® lisigjsamn^s: ^d dl®. ®h#ailseh@n I,®iatwiig«ii

dar Kssigb»-kt#t»l#ii# %•» 166, 4S4~3l*

J* . •Ii4f, Uebep TOlJi®a'Pelnstftin #iitat@ii@nd«.# . ©igenth'omlieh® Sumrft Ab*i» d@r Ohm* 343-351#

lil©l, 0, Bt wn, 19S8. fH® propionl® iioid baeteri®. i* f, Ifitgeirersaaalc,

W. teiisevaln »d §©• lasrl®®,

wSHii®rs 0:« |S41t M® F8®ttd0-ls|Sgil«r«# i|u« »ti8 d©r

• ^i&rssetawiis &»T^ rnimmim dareh #li-i«uag ter^rorgsgitng®!!, • - mti» d®3?. Ohem, 38, 299«307,

oi»st«fid, w» 1#, whitmmg *• »• «nd a, w» 1999, st®tti distilli-tioia ®f th# low#? ¥©latil@ fatty aolds from m imlt «®luti0m» !• iiol« Oii®a» 85, 109-14#

i*, .A* Q* itSl* k Mlmplmg d#li@a%@ aai i apid of doteet* iag tit® formafi©«t ©f as«ty2ji#tliyle«i*Ma©l hj hmtmTlm fdwenting oaFto0liy4r«t«s« f# r»tli« »a®t# M, 401*406*

0. %,*$ MTmn4M WwlewBit ©• !• If34,» • iw-lmitted f©.^ pib3.1@«tioii. lai* Qli@a» liial#

0»te«im, 0» I*#, W©®#i M* »# 'isii'd' *@3?hMa», E« • 1933, ©»t«Minatior3i. of ,f0wii«, and pwpionle a0ia» in •» miMtum* 'Itoi# ®ig# Aaal# §», g47*.5©.

f«t®P8®a,- w* !• am€ fired, 1» 8# .ISSO, fh© |»9?©A».3iti«>tt ®f' certain, pen*)-t©serfeiw®»t4iig It'tt©!'!.*# Si®lf QhwSw.M, 2«»4S,

f®%t, X.# B« end l^nne, 4« *• 1933-. Studies ©a pi©8^ftta«®s* J* RI, 1660-.71,

I,# B» and Wfnmg 4« M# • 1933vj^» Tho metabolism of p»pi©iii6 Held tosetePlttt !• The degradation of ohosphi&ri® teld by Pf®-plonJ^'baoterliiM iTensenil (va.fi frans# E®3r» secn-Canada S7, 119-22,

Pett, h* B« «nd Wfime, k* 1« 1934. The infIrnem©®,. of mmem%0 «ftd artonit® on the eneymid breakdowt-#f ito#«fli®ipi6 e®id esters# ilos&ea#

|8, ,

iedtenbaoher, 1846# Setts tt^er tine mm Uitetefemiifsiiifelse de^r ,«eta» eetenaa«re» Aiwi# der dlwrn# gf, I,f4*f7» „

R.« eEd.^emanit f« M« 19g3, The produetlon of tolatile fatty s.etds and ©ai^ bon dio3cide by propionic aeid liROtoria, wltli speeial referenee to tlieii* eatioa ia efeeete# Bai:i^ i®, €, msfOQrn . - ,

Siefwan, 1« 19g6, ®»» produotion of oatala## % an. ana®i?oble ©!?• ganisa, B«0t._]U, 417*|li*

t-, and Shaw, l#-!* 19 1* Assooifttlf© feaete lal aotion in tli® propioni©.. aeid faraientstlon* I"*'Hen, fliytlol# 3, 657-58#

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i# 1. and Kandall, i# I* lfS0f Physiology and classification @f tti,® pfopitnio m&t'A tesdteria. J. Baot. 19, 18»

W®^k«an, 0. H» and Kendall, S» B. 1951, The propionic acid "bn^terie I* , ilftssifiefttiosi • and noraenclatur#, lowo Stat# Ooll* Itt#

teitti#?, 1« ©« and Stisiiaan, f* M» 19S3» proDionic s.cia and ^®ts«i®s f tm vhrn fm IM# Bitg«

, Shem* lf>, Sf?9-.31,

B* '!*• »« and llbmSg *# S, 1924• ' fhit y»t®s ©f fomentation ©f hj the pro*

.. pionio Xnd# ^Iteg. ist#

Wi®I«ad, H. ^ 1922* tn3©r den Mechanismis der Ox^dati©n»vopglng@* Srg«tei«0® FiiTflal# 477-S18#

Wieland, H#-and tond#i^off, , 193a, tber den Meohanlsms d©r Oxjrdafcionsirorgange Di© ©nzsnaafcisah© Oxydafcion von SssigsSui^ duroh K®f@# ton# , dtr Ch®m« 499» glS*8i:,,

Wil®©n, ?• t,, Fr@d, !• S.. and f®t©i»son, W# 1# 1930• Bildung mid Xdentifizienmg der von v#f|ehi©d#n®a Stifflaen von Fropionilurebakterien g®bild»t«n mvLmn»

t* t8t» fe71*S0*

WoJtki®wi62', A. von* ^ « 1923. Propionsauregaruni^ is imimentaler las® ims#ieh#r Fabrikatlon. Oentr, BsVct* tt Abt, 333-»6»

f Op •• , „

1912, Sau@runf?sbalct@^i@n, insonderheit Mil^Wttr®l®ng» sftabchen imd Fropionsatirebildn©! in Molker®ip*NB>dalEt«n| speaiell in d®n versohiedenen Kas^sa^ten, lakt. II Abt. M, 494-S40*

Wedd^ H* #• iud w©i?lauinji 0« H* 1934 • Th# pTOpionifi acid bacteria on th# ii©Glianiia ©f gluoBs© dissimilation. J» Biol. Ch«ia» |,0S» §3-72*

wood# 1# ci» and werlEBusn, ©• H» 1934. • pyruvic acid in the di««iailation of glttoos® by th® propiont© aeid tei80t#ria# lJ.#eh®M#. J. §8, 74S»'f47#

•1^6-

Wfofi, H. (!• and werkman, 0. H. 1934«» Intemediete products of the fipepitni® atM . f@PBi®ntatlon» Pi?0e» Sdo» ®xp« Biol* *#A» 33.j §38-940#

Wood, !• §• iittd Werlmttn, 0, H# 1934^. Accepted for pnbllsatloft b|r t# Agl*« t©8»«i?0}i» Oct* lt34.