Loyola University ChicagoLoyola eCommons
Dissertations Theses and Dissertations
1953
The Genus Alcaligenes and Related OrganismsRudolph HughLoyola University Chicago
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This work is licensed under a Creative Commons Attribution-Noncommercial-No Derivative Works 3.0 License.Copyright © 1953 Rudolph Hugh
Recommended CitationHugh, Rudolph, "The Genus Alcaligenes and Related Organisms" (1953). Dissertations. Paper 441.http://ecommons.luc.edu/luc_diss/441
Ubr?rY Stritch SctJuu\ oT M3dicine
Loyola Univer~ii.y
'mE GENUS ALCALIOEMiS AND
RiLA TiD ORGAIUSMS
lq'
Rudolph Hup.
M1oroblolol1 Dep&rtJaent
Switch School ot Medlo1ne
A nuaertat10n SubRdtted to tbe Faoul",. ot the Graduate School
ot Loyola Urd.wrsi'Q' in Partial Fu1t1l.lJaent ot
the Requirell8nta tor the Dearee ot
Dootor of PhUoaoplv'
19$)
': '.,: ". > t-~· ~
'.,,;",,-, .,~"::-J
The antin vork waa pertOJ"Md under the
~1ate aupen1a1oa of Prot ... or lUn .. LeU.on, tor
whoa. UDtaU1n& help, eaooura ... nt aM advia. the
au.tbol" would expreaa his .. t a1DCU'8 haarttel t
.... t1t ••
He would alao .xpn.. h1a :1ndebtAtcmea. to
the tJn1ftra1t7 tor .ftont1nc bia t.llonhipa and tbe
opportantv to urr:r out. \hie vorlc. He 18 p'a:tetul
to the 8tamard au ~ ot Indiana who alao
granted t.novshipa.
111
Chapter
1
II HISTORY
III
v
Aloal1aen.. faecal!! A Laa(l u.n.. ;racos. A I ItA i gene. _talc.tig.nee A: OIl -.1 5001 .. 1 ~. LftlitMnea recti AlcaJ.uenel ~i
COMMENTS ON THE HISTORY
MA TERlALS AND KE'l'HOlll
VI DISCUSSION AND CONCLUSIONS
!VALUA TlON OF THE TECHNIQUES ELi.PLOIED
GROUPING OF "l'Hi OOOANlSMS
iv
Pal·
1
2 63 69 72 14 15
76
80
107
135
160
161 165 165 166 172 113 176 119
Chapter
VIr SUMMAI1'
VIII BIBLIOGRAPHY
DEXTROSE OXIDIDas, UNIT S Ethanol Oxidation ?eudOilOn-. =&!!!a PieUdoaonlle ~8a Aii5'bi!!i1 •• p. Bact.llr .... l\rata
DEXTROSE FIRMEIITERS, UNl'T C Ael"OlllOJ1U ep.
y
1Bl 18) 18) 181 189
192
194
191
LIST OF TABLES
Table Pap
I THE CULTtJU COLLECTION 90
II CHARACTERISTICS OF THE OROAIUSMS 109
III GROUP DIfFERENTIATION OF ORGANISMS IN TABU II 130
IV COMPAlU&OIOF KOlfOXIDIZlRS, OlIDIZDS AID rlRMENTERS ON WTROSE MEDIUM l41
V CABBOHlDRATI BASAL KlDIUH CONTROL l44
VI GELATIN MEDIUM COMPARISON 152
VII BETA HEMOLtTIC troNFERMElftIlO GJW1 MEaATIVE ROm 158
VIII PERlTRICHOUS IONOXIDIZERS 162
IX THE RELlTIOWSHIP OF IfITBATI AND SOURCE OF ISOLATION AKONG 2) ALCALIGEJmS BRONCHICAHlS STRAINS 162
X SOURCE OF ISOLATION or PERITRICHOUS NOHOXIDIZERS 164
XI POLAR MULTITRICHOUS DElmOSE NOIOUDIZERS 169
XII LOPHOTRICHO!J5 NOROXIDIZERS 111
XIII SOURCE OF ISOLATION OF LOPHOBACTER 'lECALIS 112
XIV MONOTRICHOUS DEXTROSE NONOXIDlZERS 174
xv NONKOTILE DEXTROSE WOKOXIDIZIP.8 111
XVI SOOReE OF ISOLA. TION OF NOlll1OTlLE NONOXlDUEBS 118
XVII DIF1"ERENTlATIOI OF WTROSi OXIDIZERS (OWIT B) INTO SUBUJUTS 180
XVIII OXIDATION OF ETHANOL 181
Table Page
XIX POLAR MULTITRICHOU5 PSEUDOMOHADS 182
Xl PSE~HAnACEAE 18$
XXI UNIT B SUBUNIT II 188
XXII tmlT B SUBUNIT III 190
XXIII THE OENUSAEROMONAS 193
XXIV nUP1l:RENTIAL CHARACTERISTICS OF THf: ALCALIOEHBS SPECIES 19>
LIST OF P'IOURSB
F!&'ID'N Page
1 Phot.o!ld.CJ'ograph ot A1cal!l!D8. b1"Onch1can1a showing pent:r1oboua tlaplli. 101
2 PhotoJdcrograph of Xant.halonaa u.ltoph111a .p. nov. ehow.t.nc polar IINl t11il&u tli.,.1Ii. 101
3 PhotOllicrograph .t Lophobaotel" taeaal1s &en. at ap. noT. sbovin& lophotr1chous hapm •. 101
4 Pbotad.orolraph of Pseudaaonaa aer2iino~ showing aonotl"1cbous nqelia. 107
Photoa10l"op1lph of PHudOl\\OnA, ap. shov1ng 11 curlT' .onotr1chous flagella. 107
CBAPID. I
Pe'trU0bk7, • 0eNan bactv101OCU\. _ 188, pabl.1ebed • "pen
~ aft 0I'pa1_ ~tad ,.,. epoUed ... 1IIIWtb he M118ftd to be a
h1t.berio ~ t7Pe. In 1896 be ptab11ebld ......... 1 ... repon en .... ,
he ,tated to .. tile __ ..-sa ad ~ l' 1ft ... detaU. lfe ......
_ orpn1a Bulll_ tuoalu !lel!l!!!!!. and .... ita 1cleD\1.t)'Iac
ohal"aoterllJ'1ee. Untanuna'tiel7 ... of ~t. 0\11 .......... I.WI"
tound to be ... ad .... eon"1.,.., .... OWl" .... ~.., atd 'to ...
aten' \be ..,..101..,. of ... "DSa cr1cs..l17 1ao1ated. III Uae ...... of
\_ l' a.._ :I.aoJreu1nl17 01 ... that tbeN ..s..ted ........ of ~Oi-
1eal ..".. of ... .-'_ ha'f1ll& ptv'a101oa1U1 ....... bt1a .,.,1_ \0 \be
cqanUa .. orl1MId __ PetrunhltT.
!Ide 1rrfNUp'ton .............. to ~ _ .~
aorpbol.ocJ ard '* b101oa1oa1 ...... 1ftlu .f .. _S •• plue4 lD tbe
.. ,. 'u!!UI!!!1 b7 00ft~ ~olocuw. 0.. ......... laW t.o
Alea1!p!!! ... elMo nud1ed.
1
CHAPl'U U
,. ...... obJrT (1889) 1mut1aatAtd .... IP011ecl bMr and 1eo1a\ed IUl
erpn1aa tr- _ .pr plate .v.a1ced wittl. the b8er. the OI'Pn1Ia ..
daor:1bed .. a abort __ ... '1" rod "lth 11wJ.7 .,tUl.. Po\ato ~
•• pUow18h ill .S bo ... and later the potato bMa_ brotftdah. He could
41tterenU •• the beer beelliu t ... the tJ'phoid bacm. b7 hie Ii ...
.,. •• li .• wb1eh the t1Pho:l.<l _e111u \uI"Md ... 1d wb1le U. __ baoUlu
turned it al.kal.1De. !be oo1on1ea on apr. of 1Ibe bur ROUlu end ~
tJphoid baom., wwe qv.1te .s.nar, a ... alao \be1l' aoUon 1ft plaUra
_1ob va_ aot llqat1ed.
Guntbw (18,.) 1IIolaW. ea.a Ilbaped bao111a trca 1t01l wh10h
M MIIIM 'parto Wr!&!!!!. It .. iderl\1oe1 1ft 81M and e ... wlth \bit
oholwa Y1br1o. n. orpnUa .. deHribed .. Nina lIO'Ule v1tb • WIt 01
polar tlapUa on both enda. Then ... DO l1qq.taoUoa of ce1at1lh 'fb1 •
• tr1o't. ...... did not tenent ...... and 1Il- • 7dlow18h brow ......... Oft
potato. MUk .. IlO\ ooaculated and the n1troeo-iMol reactlon .... _ .. U ....
PIrtrwIobkI' (1896) deeoribed. an orpn_ .d.oh, aooordlDl to hSa,
.. the ...... that, iaolated b.1 b1a troa spolled ...... (188,) arid ..... 1'bed it
.. • tJphoid-like alJral.i produoel'. Thi.a 01'1&111814 .. found 10 nco1
8peo1Mna. It .. de8Cl"ibed .. Minc graa nep"i_, aotlwl7 _tUB v1 til
2
, parlW.-u nacelle. (Pbot.101'01NPba did not aec~ \b1e ~.)
ColonieS _ gelat1D were sjail .. to ....... 01 the trPhOtd orpns.-. M11k
oalt .... M_ alkaline w1tJ»d aoaplaUoD. 10 au .. proclutId 111 sq ..
...ua (ta. at ... of the npr .. DOt .taW ..... aJWtb1n& .. 1d
IGDelll"D1aa &0141\7). Be oould dUt ... nUate \'be hom_ f!f!!U! ~im!! (n" ... ) baa ~ \JpAoid '*111. '" .... toUcnd.n& ,. .. Uons, •
• ) alpl i ft1\7 1ft I.e houn in 11 ... 1Ibe7, b) 110 agJ.llUraa\1Ga nth tJpbo1d
..... , 0) ~ok laJer of provth Oft potato, with ......... browl1ft& of po_ ••
II1lk .. al .. to\1ftCl 'to beCOM alkallal 14 &.8..... IntnperttonMl. 1a3eot.loa
01 am_l ..... WM'tlon wb1le .~. ~Mt.1on d1d 110'_ '1 ..... (1896) tuorao,rq Uet.ed the -.a1aa .. 8&ow.. t....u.
~f!1!1!DH (PltVu0bk7), and m. .. (1891) til b1a olas.UioaUoa of
"~te. alao .... iDId \he tr1nca1al but. ohanpd \be ..... aDCl &1 ...
the apeU!na \0 p.oteri~ !!e!iIt !!e!!I!!!! (Pe".ohIL7).
r1aoMr (18,,) reported tbe uolatJAm at, autopq of lacUls
t .. oal~ !lE:Y-- Ina var10ua ...... of • ohUd nth _ulea and
tuberculoeu. Ie alao lOUAd the orp.Id _ 18 ..... d1epoMd in eanal -_.
tMIIII_ 8IId _ ... (1m) re.-ci t.M orpnta Bacteria !al.l!!~!!!~
(Pet.naollkT). IIt.p1.a (lSlOO) nIIIM it ~ .. lll. !l!!4JenM (Pev.....,) N1a.
Ford (lJOl), 1n atte.apt1nc the ola.slt1el.\ion of la_tiaal
be.o\w1&, noop1aed a .... or alkali. produo1r.lc ~ wb1c1l ., \b.le t1M
be .. panW 1n\o ,.~ ..... 1, 2. aDd ,. The7 .... ooll.eo'lftlJ'
dellW1bed .... ,....nou .... PI'04\101ni aa alkaline NaoUoB 111 awe v1ibcNt.
a ~ aoidl. and 110 aoaplaUoA. ,.., faU. to 11q-t:r .latta,
.... 1ra 01'
or .t.1OI"OM and 1ndol va. not produ0e4. ('l'bIN is no mention of ..... react.loftt
napU • .orpbololY or Hid produGtlon ~ oarbolV'*ate •• ) 'oN WIt not
cMtJ'1ia1n :Lt II'OlIP 1 01' group 2 va. 14entleal. with Pevusoblq'" 0I1c1na1
ba.mu 0'1 vbnb.er both waN ftI"1et1ea of 'ttdJI bMUlu.a. The PO\lP' an
,. ... bIc,y (19(1) found ».'Ill:.., t......:t;1! alqal,.. .. in pure
oul tlJN in 6 !'OM ape_ on • pat.lent vltb ol1D1G1l \Jpbo1d t...... Ie ... ot
~ op1n101l 1iibat. \bt orpDiaM 1Nlated. .. ller ,... at.oola aut! t..bNa 18olate4
rr. roM qot.s wn 14entloal. He IlAo .tated there ••• -\val tJpbotd
liD diMha not ..... tv tbI t.ypbotd bIoW_ but. appventl.7 by a •• mu
rord (190)) in b1a deta11ed .t.U«!.r of tbe ol ... 1t1oaUoa of 1ibe
intutinal ...... Sa 1n _n foUowcl M1plat. (1900) ~ ueept. tor
1IIft1f .. , aI'I'ON. He ooneldeNd tbe oOlTHt; ... to be "omu 4el! ... . JUcu1a (1900) and not 8&0111_ !!!!Y&e! (Pet.lwohlq') M1pla. He cU. ... ...
~ ftl1d1t7 ot ......-10 .. by vh10h wpm._ ... noop.1aecl and .ta .... \bat
be found ~ on potato .... 118l't18 .in_ potatoe ...... of ocmatant
OOIIpoalt1on. Se po1nted 0'" that. ~ ... orlanl_ 1nooul.ated on two
different potatoft 'MY not .bow ,he ............. 1.\10 powtb and that
s .,.,....... Oft potato 18 not of tax~.~. Ue1nl" Smith
, .... D1IatloD tube with dextroee ~ ,oft dUt1ngu1ehed tbNe fP'OUPII of
.. .una- 1) 01'8.n1aae 1iblob. h .... DO app.-en .ttee' OIl oa.l'bo1\Ydratu, \hew
,rovtb being 11a1W \0 the bulb of the ,_",,-,tea tube .DenI it ex_AdM ..
t • .. tb.e neck 01'117. ~ wotA in tba oloeed an n •• 1n1D1 olar. ·,he
,....uOO 1ft ~ bU.l.b 1a uul.l.7 al1cal1nl. 2) \lao'Wta tdd. .. lenartt the
~we vi* .. prod1aetioa of ..u _\ no au, u. ~ ~ *ouahOU' .. tee. Tbe ..... ot1oft 1ft tM aN 18 &1.,. ao1d. &ad 1ft the
W].b .1~ .. id or albl1ne. ) .. aarn-e Vh1eh.bow tul'b1fl'. aDd .... oua
t.,..nt.at1oD in the ol.oeed .... 1ae1ll.. !\!.!!AM!!!!! .. " __ Ssed ..
belongina to 11"0. _. 'fbu IEI'"P .. ~ nW1Y1ded ..... sa the
table below. "... 11 .... d~ Mtwen ,0JId" table and tdII de.U.s
1 ]
I 11 I .we:
sC if!, .... J Ih , ! "" I )-
= ..,
! U S :I
i t.~ III -:
I I { 1 I '" 1 ... ! 2 ... laem!!! aleYl!-' + + + • ... ... ... ... ... ... • !&O11191 HI" + ... • .. • • ... .. • .. • ~Ulu fl\OI'l • .. + ... • + • .. • .. • 810m ... 01 • • iii r + • + - + + • • • .. • 880m!! ~!!!e£1 • - • .. .. • • • • - •
r:==-~~ -------, 6
tOJ' £!ot.!- 0_1 am boOk:Vl an UaW4 .. pnduoSna an acidl",. ami
a].kal.Wt7, bow .... in t.be dftalled deaW1p\1oft of the indb1.dul OI'pftUu
~ 111 DO .. nt.1on of pnl:la1nar7 aoWv_ (fId.a procSuoUoa of a 1n1Ual
ao1dlt7. pI'Obabl7 frat duVoae 111 the Idl.k. aid .... qGtmt alkal1Dli NaoUoD
1, perbape anala.,. to the ,taU .. ....-iea oo..1Dc vltb the Sb1&a baoUl._
ard Pro __ !'!le!'1I. Mone or tM or.-t.. 111 Vt.e a'bcWe tAle "ow 1Il taw
oloeed .,. 01 t.be Saitb t __ ntaUoA \ub8 ,.t 'ord detlCII"1McI .....
laoult.aUft ... 1" .... )
AlfMbul... (l~) wu of t.be op1alon that. the \Jpboi.d .0111u
oould cbanp to 8aeUlu tea118 !Ys!!WlWt and!!!! .,..H. ~ (l~) u1ng :3 .w", of JacUlo tuoal1e alkal!p_. two of vh1ob ... ,....
Pet.nuscbk.1, __ to the ... oonolul08. :t:Joeben tound ••• roloslea1 en., naeUon bet __ b1a t.hree ,trI.fne and t.M tJphoid bU111 ••
JJeaba- (190S) lII8CI DoeDert', &aa111_ tMoa11e a1elJee!! I
(Strain. II of PetruaoblE7) and found 1\ to be • Idxtun ort 1) plUIIP rod,
v1.t.b perlwloboul napna which ........ l1Ice ante.. 1ft a _ittt. pnpu"ation.
2) .lend .. rode wlth a polar tlapU_ vb10b IIOftd 111 a I.atpt. line.
(Pbo'-iCINpapba did not aeocapaftJ' the I'eport.) ftse two orpna. ....
np&n.t.ed and 1deRt1fled awolo,l.u., and lQ' 11 ... II1lk .. the \Jphoicl
bao1Uua and BaoU1, taeo&ll. aloalM"'~ Napeot.lftlT. Uelnc tbI ,...
O\ll.\un& Bwpau could DOt. tranatON 0118 1IIto the o\bar, nol' could _
...... Vaw &IV' 01"08& aalutinat1oo. BerPa- fOWld aaom_ , ... 118
aloal.1pne. to be a atr1et. ..... , 1f'Ov1ac on1r 1ra \be OpeD pan of the s.1tb
t'l'Mfttatlcm t.ube.
7 ..,
Uled by Al tscbuler and Doeber't, wh1ah tbe7 claimed vere tranatONable 1%1'to \be
tJPho1d bacUlu., originated tl'Oll """"obk.Y. He obtained troa Fleber :in
Berlin tranaplanta ot Petruschk7' s .tHin AI and Poebert.'. .train All. Their
aul turea ...... plated and oolonies were tubed. Subaequent. cul tuna we"
t1P1cal tor a_emus ' .. calia aloal!l!nes and law no aglutlDat10n vlt.b
tJpbold aerum. noebert'. vanatormat,ion exper1Mrl\a .... repeated v1t.b
negative results. To. autAor concluded that BaaW!! taecall. aJ.cal1ee!s 18
a group or OI'Ianl ... rather than • ainale species, &DIi tGat the organ1au
cannot be tl"amtOl'MCt ~ the t.Jpbo1d 'baoUlu.
Conradi. (190S) ."..ad with TrOBUdorf tor he alao totmd Aluchuler'.
cul turea vere oontudnated.
Serpa_ (190S) after exwnei. study of ..... n .wains ot BacUlus • t .. eal~ aJ;oal!g1)q cuae to "lard th1a orprUe. al a nonp1pent producing
.train of Bao1llua fiuore.oens non-lis.faoie.. He o'blerwd the aoWJ.t.y in
hi. oollectlon to be a 11near aoUon, wil1ke the v1gl1l'l1 IIOtioft ot t.Jphoid
bacilli.· Bwghaua could not conliN Petrucbk7" state.nt t.bat laeUlu
t88o&118 aloal1ene. bas peritrioboua tlaplla tor ewn Pek'Uaoblq". strain
II bad 1 to 2 poluo flagella. One ot the .. wral .\raW .tudied had
10pbot.l'lohoUl OJ" polar aml t.1trl0h0ua flaplla. The ZettDow _thod wu used
and. WI lOUD! to be superior to the Loettlv wobD1q_ tor ata1n1De tlapUa.
Growth on potato .. bJ"OVll, and aU were 8tl"1ct aerobes. Pellicle tOl"m8d on
'broth and there ... DO atteot on dextro •• and laotoae. Indol te.t ..
nept1w. Nitrate. vere reduced to Dit.r1te ••
Krenoker (l~) adds notblng to the alread7 .xi.t1ng description
of the Ol'lani ••
8
'lV1:» .... h (1906) 1eola\ed 11& bl_ colonie, tJtoIIl tr1p1,kl..cOJD"adl
a,&I' iMfnIlated with oanal _tel'. fbq .... 1dent1tled b7 Petl'UaetbJr7' a
11-- "7 Md1_ aa Iaoill_ t_~ al.el1A!P!a. P1orkowk1 (1906) concluded that, .. \J'pbo1cl bloUlu m:l1.,.uJ.
,.0&11, a1cal!f!!!! are ditt .... D' OIl tbe bul. of povth OIl plat.:ln. He
.. orib8d colon1aa ot 'bot" orpns... Tr1Dou a_ oua (1901) de8Cll"1bM __ of als...,.,. 1a\oX1oaUoa
pI'N\8AblJ' troa ..... trOll vb1ob .. iaolat.ed 8acUl!! tuoal¥ alca1?:1!~. !be OI',M1_ .. ana poe1\1w aM ".oduoed ...... p1p8n.
Oaehtpna (1907) ualna I fnebl7 1ao1a\ed tqphoid bH:ll1_ fraI ~
blood ot • .,.,1eD\ an. l .vatu of hem_ tuoal.1a I!leal.!f!!!! .. .abl.
to vanet ... tbe ODe orpll1ea ate 'UJI ...... bJ' 11 "liWo _tb.ode. Oeehtpu
aot.loed dill ....... 1n powth of tM _1&1&1_ ora ...... 81 -".1&. He abo
0 .. ,ad \bat. entiMna PI'ftPIIJ'M for .. 1 •• 1ft cUd 110' &Ql.llt1nate aU o\ber
eVa1ne of DM1Uu taeea118 al.!!1k!!!!.
n~ (1901) eon •• ted 22 stra1na ot Bao1l1_ t .... U. &10.11114 • h
"'1& .net 0enaIV', 1Mlud1nc 3 .t.ra1u d:I.Nctl7 ,... Pet.rwlol:dq'. one of
hwuoblq'. IV'a1att produed )"ellov pl,.. OIl aft apr elant. The .. 15
,".ina ....... hl.ll' ~ Vitb 7 of b18 own .vatu 1aolaWd ,... f .....
vine, cd oaDa1 __ of Petropoad. n1aaDko abo .\\Id1ed SaoUl_ I
tluon .... DOII-ll9!!!ao1eu aat a_m .. tlUON .... put1du nvae- It·
appear. olw1ou tl"CII the data preaen'-' tb., \be OJ'pIliaM atudted __ nt.nko
.. SacUl. t-..oal1e .1_111 .... WN of .... \Jpcta. He .... ted that the
...... i •• be d1tterent1awd on t.be ... 18 of pipant and .oid prodllot,1eD tr_
~te._ He Objected to t.be ..... \100 of Bveba .. (1905) that
lfoUlus ,aeeal1a fl .. lle~ be OODaidencl .... acaplgaentM P~!!!I!!!!!e!!.
r,u.enko conoluded that BaoW:u t ... 11 .. at-a!U-- _ polar tlapllaWd.
n1rae11ko (1908) deaor4.becl aae!'J:1!!...s.-e (BOY •• pea.) .. a DW
tr'PI ot a1lr.ali prod..... He ucl:uded Ud.. OJ"Iaa- troa the BaoW.!! t .. oa1,ia
al~wne. &rOUP- The 0I'&&n1aa wu uolated fl-OII t.be apl .. ad blood of ..
p1M& P1l aD4 .. .otU. vi\b pw1triehoU napUa (~p!aI-",
not publ,lIlwc1), uporop~.an4 • __ -..'1,. rod. It pnftaad JlO p1p&nt,
did not U ... ". ,.latin .. rd produoed an alkaline "eaotiaa 1ft .Idl... Ii .. taa
..... not reduud to nitrite. Alkali .. fOlMd 1D 2% ................. td.tb
11taua 1M1oator. JIydropn.;.u1de vas prod1loed.
tatt .... (1908) reponed .. 1aol.atlcD .f S_w..- al.ea1Y!M! trca u. blood of ..... ft'OlY1Dll1lce tIMtDip ~1d t ..... , • ......" ••
11-' vbe7 be ... 1Jl\erlMl3' bl_. (n.pllatiOA'" not "ported.) Then
... t ...... UOn ot _...-.1 ~ .. v1tJl tile ...,,1oA .f laGtoM.
the indol .. t .. Mlati_.
lona (U08) did 801. .. tribe the .... DiM ba\ found DO _\alae. 1ft Baneri_ !J!!l!l!!!!. a.tel ............. , 11& ,..~. s!!p!!!!.
S, •• rl !! fl. (1908) 1eolatM a bao1llue tI'Oa Jelt .. , tap _te
which Nanbled hom_ t56oal~ !1oa:U .. !!, and 1D tb18 pal*' .,... of l'
•• !!. GrOlrt'8llCtF. wna. (1909) nla'beled the ..... n1_ h !tutU!! alGal" •
Th1a bao111wt GaM to their ., .. u. t. l' .. aalut,inated b7 the .....
trOll pran1oall.T .... 1 aa .. of .,S_So ... e""p1nal aen1D&1Ue 1ft Seltaet, •
...
10
of 1a$0. SaoUl_ , •• 0&11. !l!!l1l5!!! (11'81) d1d not. absorb out the ••
lI,luUn1U. !:.. 0F08ftnOl" •• deaeribed. ...... M,at1ve and 110'11.. ,ben
... no liq.taoUoft .ot ,elaUn, no ... (DO .. t1oft of acidiv) in 11 ....
brOth or a'W' other .\18'" or aleobol te .... , .uk ... not coagulated, a taint
traoe of indo1 vu produced, Petruoblq". 11 __ ....,. be_ ta1llU7 alkaline
an4 potato ~ •• brotm.
Md4w (1m) 1eolated Bao1J..lua laeoa1taal.a11pn.u fntI tbe bloocl
of a patieftt v1\b tood potaoDing at ... ba'l"1llg •• ten • "port dlahlt • The
OI"gan1.a .... dea .. 1bed .. be1ft, .'11., no .tfeot. OD aUk, no 11: ........
and 11 __ wheT ..... 8vonc17 alkal1De. The •• tlIoP ,.. 1noUDec1 \0 oona1der
the o .. "an1aa .... poIlItbl. tflr the olWoallT \fp1eal. .... of food ,.s..ord.nc, tor 1 t ._ a.clutinated by the pattenta .....
HaM (UlO) bolated 8 •• BI!! taual18 alea1!ft!!. &CIA taa wtue
IIld peri\oDAl. n.1d 01 a patient. The OI'pl'liM ...... thad .. prod"c" .•
alJral1n1\J' 1ft 11tau wq but. DO ooaplatioD. of 1I1lk. Then .. no ... 1ft
t ...... ta,ta t'abe8 ocmta1n1Jlg du.troae, laot.oa. or .uoroee. (There 18 ao
dat1nate atateMnt concemine aoidi\7 1,. dextroee or tl.apll.er ..,..10.,..)
Hill ..... ' an! 'Aiarotte {1910} npoJ1:ed an epic.te.1o of ,uvo
entel'1t:1a iftftlY1fta " people in .1011 »-0111_ taeoal.i8 alcal!J!le ....
iaolated tJoaa a.pecrt.ed .. t. The author. dtd not cona1d ... tM orpniM
"8POMible tor the ... , poiaoninK outbI"ealc. I' .. <leeor1.bed .. be1nl
,11OtU. and hari.ng the pbJ'atolcc1cal reaottona ot ~ .. Ulue taeoalt. aloal.:lp!!!g
CU_1lani (1910) nported the 1trolat1o~· Oft bil. ealt. ., ... or a
11'_ rwpt1Ye .pu-ill_ f'rOIl at.oolll of fatal bUMD ..... of .rrteroco11t1a.
u ,..nstb _d "pe. s .. 1nd1Y1duala ven 20 to kO :d.0J"0ne 1ft l_rth wlth I \0
4 OOU8 .hUe 0\)111'8 .... roda 01" ... "ped. L1taue aUk • .bowed no ao1d1\J
bat after 8 _ka the IIdlk .. alkal1ne .,d pepMrd.a.. Broth povth .bowel
a .. naral t.urbidI", with pellIcle vb1le arovth QD ..... we vb1t.1ab like that.
of 001011 baoUll. 1"beft ... no lI_taot.toa of .elat.in or ooaplaW •• ,...
b 1ndol tNt. .. De.at.l... The ..... no oblap ira clavo .. , lao\o8. aid
.... 08. 11taae broth.
'''''' (1910) laolaUd an 0I'Id1aa tre. the .. ,pin...,. Vaot ot
clop. 'I'n. orpnUa ".. d"G1"1Md .a • -*' or' n&ft"OW _W. baoWua. (No
.... , .......... '1oIl 01' naaellU' 1IOI'pbolo ... ,1"'1'1.) A plata ..... aVOke
s.a 24 bolIra al»wd aoden. powt.h wblob ... mo1ft, aUa-l'lini. ..,.. aid of
•• t.Icky co-i.IWDo7. PldA apr ,tab II'OVtb vas beat. ftMI" the ... f ....
Potato powUl ... 11aht. tan ool.OI", _UDdIn\, ntaed, _iat._ alia-nina and.
.t.1oJq wlth • deo1cJed odOl" of .t.ale...... 'otato be .... 8l1chtl7 ~ned.
teef1'1v', b~ ...... not. oolored 01' 11qtat1ed. GelaUQ stab aholMd no
11ca-faoUoa. No IU'ta .. growth 1ft bJtot.h, but. the .edlUl1 ......... ~
dowdy. u •• adlk :In 12 ••• be,w to tum blue at. tiYt a .. fa .. , at. f1 ..
ctqa the vbole t.w. .. blU8, at. founen .,. 1lhe ool.ol'd1Appeved fIrca tbe
ttal 01 the t.abe, but no ooalUlatI01l 000UNd. sa1th f.-n"'tlon .....
ttl dextl"08e or otIIer OU"bol\Ydl"aw. ebowd oloudina 1D tIbe open an with no
sible ~ 1ft tM e1oaed..... t.1thel' acid 1101' ... vas produ.oed and tbe
iUll bHaM alkall_. Indol wu not produeed. SW_ troa clop ntt .. 1Dc
«WI diatNlper alva,.. aalut.1Dat.ed 'tibia ora--. J(fao.n (1911) 1ttolaW alii .... 1btKl _ 0I'I.w.. vh1eJl be to\JQd.
t1aaa ln aldMl.. wlth d1a\eaper. His au .. 1 OGI'N
12
rfW't't (1910) and added the 0I'Iara:1s_ ..... ..... nept1... wit.hout apo.N8.
,..,t111 ... rurtV YfII!J' ..,.ked, b1&t tlJapu. .... "..Mnt.. (V.,. poor
~0I'0II"&fJbe show naplla "ftrI ta1rttq. _ ... i.a oluped toaet.ber.)
ft. author oould ION no idaa ot '\be D.UIIber or ~nt of t.be tlapu..
A pell10le va- DOt to.raad 1ft nuw Hd1a. Uta. aUk ... alkal1ne .ttv ...
""a, no coagulat.ion .. obeerwd. The 0l'1an1a 4t4 DOt pror1uoe ao14 OJ"
au 1ft peptone aal.t aedma oonta1ninc &1"' •• , laot.oH, e ..... or otbU'
luted ~.e. He ...... , the Md1_ be..-~ alIcalw. Tbe
OI'Iaa1ea P'ft "*'1ll' ar&aeZ"obloall7 1ft t.be depth of .... 0\Il. t... :1a 1ll1C08tI
..." alll 1D Jh&ebrMn". 'ubaa vlt.h pJr'Opllate ot soda.
'''''' (1911) nport,ed l' ,.. pou1bl.. to :i.Hlata VIe .. p.w. he
duorlbed (1910) ". both tbe .... pintor.1 tno\ and 1tloGd of <lop. Be l\aIIM
.. oraaa18a B. bronobiearda and naW 1\ .. .,.. ne .. t.l .... In 1'12 .. , 1M
I'Hllsed W. .. 110' an appnpriate .... , tor tM OJ"lania oould be 1aolatAtd
troa other w_l., be .. be proposed tta __ Sao1l1_ bl'O"!!!!m'!!!!-..... Da (1911) exu1n.ect a ....... t:ra1na l.abelad BaoUlue t!!!!!!!
a1oal!el" and found t_ al.WJ'8 to be polar, __ victho_ or b1pclar, but
""'1" with parltrlohou tlapllatton .. PeVwtobtq' bad nportect. fba ntbar
used tlaplla na1niI:ac, a aodlt1oatton of Loettler'. t.eolm1que, l'OUt1Dl17 to
Sdent1t7 oraam..-. ,Baaut.U'ul photoa1 ....... pbe ebov the lophov iohcNt!
tl.apllaUon ot 8ao1l1ue t .... U .• alO!Yte •• 1eolaW fI"OII tHea.
01 .... and Haebla (1m) tOWMI 11M' the aorpholOl7 of 1H1l19
tae .. u .. ale1l1!!!e ...... led ebolen nwlo. Tb.e Kral ava1Jl of tb1e
OI"Ian1ea ... tOUftd to haw n ...... p1pent and .... thenton DO' t.Jp1oaJ.
13
alJIO.t 1n pve cul t\1I"8 1n d1an'bMl. ecmdlt.lou in cbelera pa\18"'-. 'lhe7
...... with JC1".nko and n.ot vitb Berl_\Ie "JU'd1ftl the relat.ionship of
,..u.d0ll0llU and Bulll_ t •• oalll alea1'ee!. - Hom ad Huber (1911) oeouSoJMJ1,. to ...... 8 .. 111_ '.0&118 ,
al.u.1p-' in .. f .... of bealttv' COVIJ. ,he t.bNe .tn1nI isolated bad -u.ft17 aotlli\7 I.Dd polar tlaplla, Uluall;y tmipolar. Moderate poowt.b ap .. __
OIl ..... alang, plat1D .. not liquefied, a1llc ....... ,.u .. wt without
ooacul&tton- U'-w wbq be ... blue 1ft 2 to .. 481' •• '10 ... trca dex,""
and laoto.. Ib.tol va .... tlw. 10 aeid 01" .U fI'OII al1arol. 1f1Cb"OIen
.ulfid8 loratioD ... lIOde.te.
Poppe (1911) tound, uona ..,. 0-.. ol'lan1ae, 'aoW. taeoa11s
a1oal1le~ ill the J'Olk and wb1te of noral. beat •• U.. lie ob ...... .-t . ool7 JIOt1ll \taoter1a ,0 tlIroqh tbe .M11 of t.be e._ Iftf'eotion of ... en
ooeta"8 durin, 1" , .. t.1oft ill t.be Ma. rV't.U1Mcl ....... 110ft otten
OCIftt.aaSnaW.
8obottlmll ... (1'11) luted two ..... ot o11n1oal t1Pboid tewr ".
which 8aom_ t!!p11a aloaYe!e! .... lIo1at.ed. tna the blood. The
patient" .... aglat.1:nated the ol'pJd .. bola.... The oqao1_ .re
deaarlbed .. _ ........ _p.t.1_, an1le rocIa, pnduoinc red oelOJd. .. on
Indo'. 1l8Cl1ua. (A1 tanall \b1a OJ'Iania did not. prod.. pa frcIIII da1irOIa
~N 1. no reoord of _ Hid OJ" alkal:lae .... ot1on 1ft dutroal .ell_.)
Ba ... aad Sprtnpr (1912) laola .. BaoUluI t_!!HI alalW-' troa YU"iou 0 .... 19 of a ,..tlent vltJl a",1cal 8J11P'toaI. of trPho1d f .... ,..
TJphoid bacUll ..... t. 1mated boa t.M pat.1ent. but t.bI .... aalu'U..Qawt
\Jpbo1d huUl1 t.o a d1111\1011 of 1.200. 1'lle bola ... llttt1U." -, 11
lk
aloalll-- aalut1na\ed in the pat1eD, ....... to a titer ot 11200. (The -orpniaa .... no' duoribed.)
Baer'tble1n (1912), 1ft a nud7 of cllttennUal Mdia tor cbolara
dbl'io, 1aolatecl rtUMrOU oraam.-. vb10h -~11T .... liD cholera
91brio. !be .. '1PM .... al.ao8\ uolui_lT loaM ill patienu vitA intNt1raal
dbeU8, a. OM &eM an impn •• 1oa of an "lc4oa1e nlatlonab1p of t.he
OJ',an18a to tbe ciUlla... In tbrH .... the ~ .... totmd in U.
$od ot the patianta. ,be Cl"can'S ...... 0\U"ftd on ~" _dJ.a but
rap1dl,y be __ quite .".1"" Oft OtIl\1ftt1aD 011 other.ua MIl .... ..-ata v:itA Mdia taU" \0 cbanp thea back 'to U. c •• .ad tJpe. !be ..... 5_ ....
touad to be aonbeaol.1t1o. *qy MI'o1011oal tJpo ... ,.. tolWd. !be nra1De
produced • nepbel.., or alberr;r like col .. 011 plaUta. OelatlD .. ROt,
Uquet1ed ill 14 Up bf the .. orptd.ae. It. ftI'1ou .... w shoved either
t.ipolar .. wd.polar acmovl0h0_ tlapllaUoll. Baerthl81R thouaht 1t .. U.
to rec\1tJ' ,.tn.seblq'. dhvlpt10n of JIOI'pbo10D" (pol_ tlapUa) alii
pb7e101o11 Cappa_.nee of oolOl'11u on pl._tift).
Poll.ak (19l.2) .... tAt .... Jrt.laUy tn. .... co_lwlS.cu •••
8Mrtb1ela. Ie i_laW eight Ol"lam... lrOIa noolI of patten_ vitA 8n_1o
41eturbanoea. AU oU1~. vee pbtalolopoal1, 1deJlt1oal. AaoDl tile
1Viou obanoter1n1e8 I1ftD, indol va ...... t1.,., aUk DOt ooaculated, 11 ...
Vhe7 bl_ an4 DO I'leaol7a1a of .bGc~ blood. Oft 0l"1a1ral laolatlcm ..
o~ e1'lo1Md 001:W1deJ'ab1e ourn.t.uN. Cul tl .. tioD 0I'l laboratol7 Md:La
.... \IlW 1D .. 10ea ot OV'ftHn and then appeand .wa1abt roda vita 0A17 ..
tew cUl"'Nd 1bdiY1dua1a. Pollak oonoluded ...... t ... .... BaoSfl.- E!!!!U:! a1ftal~ (fbII \JPe of t1apllat1cm _ ROt .... 01 ..... )
15
Cae.Uani (1912) .'taW \bat 9eo111. taecal.1e alMl!l!". of tbe
SA_UDal tract of un 'to 'be aot11e, .,... ..-'lft. caued 11 __ ItUk W
.... alkellM, failed 'to produoe :Indo1 aDd tM Vop.-Proau..- ..... '108,
plat.1D and ooaplatecl .. ,.. Wft Mt llqgatS.e4. 01 ... , .... 1., ltI1lto .. ,
.... 1ft, .-bite, ,alaew .. aDd 1aenlOM .. 1& WN tvmed ao1d.
TOl"N7 ancl Babe (1912-191.') oont1NId the .. _ ... tJ.ona or 'WIIT'T (1910 anf.19U) and M'Gowan (1911). ,. ........... .uond.v ot oultural
reaotione ot ..... 81 huradNd ... as. of _pnt_ 1801ated ,... _1u1a with
41....... All Wft aglut.1ute4 .,. ..". .,..Ua 10'1' •• 1nc1 ..... :1A, ret;
tM7 ...... tect into two poupe 1n ~ to thea aot1on Oft Id....... !he
....,n-..... elovl7 aot.u. in busiJla drop prtrpU'ati.ona. rlapUa ....
10.,. tJwl tbe bod7 ot \be Me111_ aDd ..... t1ft to aix 111 n..... (A
~ ........ ft'I:7 poor17 ata1aed flapUa.) III the 8111. , .... \6.
Uon tube with dextroee, pofth .. _ app ...... in tn. opea .... onlT. A1\bough
!:. ~~ •• not an obl,.te ...... 1t. IN" YVJ' poorlT 1ft the
abHnce of ...,...8. Pl .... 1Doubated .... o1l'1oal17 ... GalT feeble 1J'01f'Ua.
III ~'iA nab poovtJa appeaI'8d ..... the nrtaoe but. .. 1l1.p\ 01" all..", 1rl
the 1_1" pert, of the tube. TheN .. ftC) l~aoUon of ,slaUD, 110 ltdol
prodllO\:1on, anc! pellial. tCl"1at,lon. ". <Nltunl. oharaotem\1oa wn in
eenen1 tM ..... tboa. a1".. 'by both'ltWt7 and S'Oowft. In 11'- IIilk
aud OIl potato \be .... nlon .. skll ... to \hat of SacUl_ fuea11a
alkalte!!t!. !.:. bl"OM¥f!pt1o,U __ te ... aflt1_ heIaol.'rUo .oU. on do"
rabbit or cu1Dea pic ~.
8a1tA (191]) .tud1ed adUlt- ~1Hp'l_ aDd hU .... lptJ.oa
........ ~ \he PYCKf78M-...... Like PJOOrane&, Ba,Ul.,
.... c:th1MPt101J11 18 alao • ,tn., a __ and 1d.tbou:t. acUon on eQPl"8. H1a -
.tra1U of 1:. broaChlne!lou. 1ft a .. ...,. ~, ,... toWl4 too be
_,,,ally tant,loal with ,..\u~d... !:.. !!'!n!!deJ!'1o. ant1aera agl_twt
.. td.8 pert.ua1a .tn1n to tull \1ter. 1\ .. abo ftOttld \bat. tnebb"
s.oJ,at4HI 8watne of tu &lantW"e baOUl. aNI b ~'t1ou ... iJIpON1b1e \0 d1at1~ b7 the ~ .. of If"ow\b on potato.
J'\rib (1f;l:J) reponed a ... vb1c1l appttU'ed o11a1oa11l' .. tnbOld
''''1'. lactert!! ~ .. oal1e ~~ ... 1eolaMcl .rr. tl1e blood al1d etool.
!he orpD1a ... de..nbed .. a ........ t1 ... , aoUle rod (f.1a&ellat1oD ..
aot recorded) 'Iddoh d14 DOt. liquef7 plat.1D, nar produoe indol. 1\ to.r.ed •
-.1. oolol1l' .a Coaractl-Drtaalakt ... 1_ and a pe11101e on broth. It. ..
aglut.inat.ed b7 the pat.lent.'e ..... 1ft a dUuUoA of 1.120. tile au ...
aoUcecl \bat, bie ....... t.ook aVa14 .. not. aglut1nated .. ~ to tbe
oraa.m.- 1aolaW aor ..... paUe.' ......
Pae1notU (1'13) isolated a ftl"1e\J' of ~ 1Do1.u.a, paratJpbo
bao1111 and B.111. t!!!!i!t al.!!lle!!! ,.. 1t.a11au ..... t.ablN. The ..
• ptabl .... otten I,nulled by ...... a11l/1 eatMl ,..w_ He ,olDw ou\ .,
.... tabl .. .,. be·" ___ of Weniolt _, ttae .. orpn1aa.
hftctaU !!'.!&:. (In) obfIwftd • medea. __ t of tree u..n1a
PI"Oducecl 10 .... a- Woth b7 8.0m. aloa~W!!!!. howeftIt tt. \fpboid an4
dJaente17 baoU11 also produeed si.nv ..... ot -.onia 111 .... ,.... brotob
!hen ... ~ vb1ch .... d \0 1nd1oate tM\ 1ft the p ....... of oqpn
',..m-., !!!!1W!!!! can der1 ..... .,. ,.. 11110", or poulbl7 tr_ ._ \lllknovn .. _ ....
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18
~ to' Cou (1899) •• B. 10. 66 •
• traub and Kreis (l9l4) "portH 't.vo 08_ of en_1Us 1ft whlch
JISll- LUOIl18 aloal!eMS wa_ Uolated. cue I ..,u1 .. was 1101a\84
". the blood. Blood a,_ ool.oa1 ...... p1ft head. w.aoJc18h with heaol.yt10
)MIlo. fbe oraan1aa aa:lutina'Md 1ft t.be eenftl.uoen\ .. ra ot CUe II in a
111000 dlluU.on. !he C_ II Ol"laniIa ... touad Sa Ute 1IJ'1:mI. The pattera'·.
__ in • dUllt!.oD of 111000 ..,1.\111& .. ,be leolaW cwPDi.. 1'bI
..-m- leolaW 11'1 t.he .. 2 ........ oou1dwttd YU"iaattJ of BaoUlu
, .. ea1l1 al4Ial!t!-.. .... A thoUCh _ ... 1_ ot 0 ... I 11..,," .. 1&\18. -tM \110 "aaat- .... oa.pand w1trb a'*o\tata11 .. " B.~Ul. ~ .. !.!M! a1e1teel and an ftoorad in the 'table MlovI
.oebata III'Jd MarI'otte (1'14 and 1916) obeeJ"ftd tw patients vs.Ul
patrointaeUMl .,.. .... froa whoa !!tB!- tMOal1e alkal.!M- .. bolaW
trc. the blood. '!be two orpn1_ ...... '" .1-11 ... the paU.m f .....
aglutlA1.t.d ~1d baeW.l 1ft low Uter, 1.50 and 11100, b11\ DOt ••
PU'&V,pboSl..te. Both patient. had ......... 1naW vs.t.h VPhoW MoUll. '1'be
OI',u1_ iaolat8tS wn ~glut1DaHd "'" the ,...,..t1 ... paUtm" ... ra 11& !d.ah
titer, 1.1200 and 111500. The orpld ....... :1.11 ... '\0 a ...... Cue II ud
schottmullerf. orpniam descri"Md. bJ' B_.lIb. (rlap1lation va. IIOt NOONecI.)
.,. OI"I.~ 0l'08' It.,lut1natod to ttter bu.t neither .. auJ,utiu\ed. 'bJ
, _,1taV'})ho1d or lUtt1paratypho1d sera.
SbearMn and MOOl"bead (1916 and In?) ,.ported the ieolat1tul 01
Bao111a !."ali:! al .. l!J!.~ trOll tbe blood of 11 Plott em. 8upee'Md of -.n.-io Weoti.on. AI&lut,1nina tor·tbe ora.""'" UolaW, in .. dilution of
1-200 or hieber, ..... to\lftd 1ft the patie'n,t. "1"\111. ". _.ant .. _"..,ponded vitti t.bAJ orialMl cleeoript1oa of ,.~ with OM or two
....,t1ol18, a ftl"iat1ol'l ,. _t111_, and 1ft .... aU theft .. pepwrd .\loa
of 1i ... mUk. The anthon pointed out that tbe lUlU' ~ 11wa b7
Ca.tellaa1. (1912) for U.eUlu r ... ls.. a1.!Me~ .. in "lI'OI'. HiI'n (1917) 1aolaW l4OUl_ t.'!!lM ea!a31!!!'! in INN CN1\Un
troa the blood ot 12 lndi:riduala. I'harinc lSI16, 622 eaaple • ..,.. __ _
baoteriDlopoallT cd this oraaniea .a 1aola\1t4 2, tJ.au out of 12) ,"IUft
oultuna (l8 • .,.). Captain B. WUab1re, who .. alao 111 Ala:ancII"ta at un.
tSa. lIolaWd th1e o ..... nt. troll tM blood ot three 1ndiY1daala. The table
below 18 Uk_ t .. Hirat', paper. It ebovtt the pr1nc1.pal ......... 18\188
of the Ora!=.. .. Pftn by .... 10.. ....... The Idfte .... 1Da of 811".", Group I .... pl ....... 1., •• peo1a1q at_ repeaW .ubcul.tUN OIl apr. All
\he '''a1fta of \hle II'OUP ..... ftODIIIOWa 1Il prt.17 nltun. OM be_
.0\19817 MUle OD .uboulture and ....... &1 ot \he otbeh be_ l1ua~
lIGUle. All tbe .... 1nII ahowe! n.pua 1IheD .t.a1Md b,. the ae\bod of 110011 •
• m Manx. The \7P1oa1 ........... nt. of tlaplla .. two at ... eIlCI. 'but
1OIIIItSM. \beN _" 81& tlapll. a' tile end. lndol .. net. pl'ocbaoecl, platia
.. not l1quef1ed and aUk ... not peptoa1.... AlUll .. pl'Odaed 1Q
r - 1
Petrusehky Kl1.itlenko Straub " Straub &. Rochaix Castellani Shearwm Hirst subgroup lfraia Kra.1a ,
I strain I stra1n II Marrotte
motUiv + variable ++ ++ ++ + w.riable vpriable
fiagella peri polar polar polar polar
gelatin - - + - - - - -pellicle nocc. turbid turbid sl. turbid turbid
bouillon turbid turbid pellicle pellicle pellicle turbId pellicle pellicle deposit deposit deposit
carb0h7drates - - - - - <text. + - -Ii taus JIlUk blue blue blue blue blue blue blue blue
Idlk clearing clearing clearing clearing sometimes no clear peptonized ing
indol - - - - - - - -potato potato potato
pipent brown brown none none brown
agglutination by 1:1000 patIent's ser_ 1:1000 1:1000 1:2000 1:200 1:50
22
glucose, lactos. and sucro •• broth. Hirst'. second group of organiSlU,
isolated £rom. blood, wre coccoid and no nagella were demonstrable. The.e
strains were physiologically similar to the 9 straine in Group 1. In Hir"t's
opinion the following characteristic. ..ned to identity Bacillus faecali,
allcaligenes' MotUity in hanging drop, gr .. stain, litmus II1Uk, gelatin,
appearance on apr slope, peptone bouillon appearance, lacto.e and gluco.e
in litmus peptone water. The use of the agar slope a'f'Oida contusion with
!!!. Plovaneue and other pipent producer.. The author obtIerved that his
isolates were fairly characteristiC on MaoConkey's lacto.e bile salt agar.
The result. in the table below strol1ll7 sUCgeats that B.cmu faecal18
allcalHenes multiplies in certain abnormal states of the bowel, and 11, 1.
pre,ent in ..all numbers, it at all, in norul individual ••
enteric conYale.cents
dy.entery convalescents
normal .teole
total.
43
51
50
16
29
nil
Thoruon am Hirst (1918) reviewed the work of Shearman and
Moorhead (1916 and 1911) 88 vell as the report by Hirat (1911).
Castellani (1911) found in the Balkan and Adriatic aone.
ease. of rever resembiing enterica which may haft been due to Bacillus
faecali. alkal:;ene.. The author states the cultural reactions ot Bacillus
faecali. alkaligenea Petruschky to be a. tOllOW.1 Aerobic, nonspore-torming
intestinal baOilli, motUe, gram negatift, gelatin negatift, coagulated .erWl.
23
negative, litmus milk alkaline, general turbidity in broth, indol not
produced, Vogea-Proskauer negatiT&, and neither acid nor gas produced trom
glucose, lactos., sucrose, irumnite, maltos., arabinose and man"y other listed
carbohydrate.. 'The typical organi_ produces strong alkalinity in all aucer
broths, but certain strains are said to produce alight acidity in gluoo.e
and ultos.. Some straina peptonize .ilk. strecker (1917) who wrote a diasertation at the Uni .... rsiV of
wuraburg concerning Baoterium alealJj!M8, town they were curved rods e:'h&ilar
to typical oholera vibrio. otten theae cu.rved rode for.ud spirils while
at other times rods tonaed. streeDI' atudied the inlluence of media on the
curved morphology but found no conai.tant correlation between the two. Using
Zetnov'a stain the organiau were bipolar lophotriohous with 1 to 6 fiagella.
strecker waa of the opinion that Patruscblq was mistaken when he found peri
trichou8 flagella, this lIli.talce he thaught was baaed on a culture contaudnat.ed
wi th typhoid baoilli. Strecker exudned BIlany' S~.8 of urine and blood but
never found Bacterium alouigenea. The ol'l&l1i_ was ftry aerobic, as indicat
ed by pellicle tonaation, growth in the upper layer of terantation tubes, and
growth in the upper layer ot shake oultures. There was no liquetaction ot
gelatin and no aoid or poptoniaation of a1l.k. fl. oolon:1 e8 ot the organism
appeared siRdlar to those of typhoid bacilli on Drigal8ki t IS medium. Li taus
mUle t.urned blue, no acid was produced traa glucose, lIW'Ul1te, malte8. or
sucroae.
Ev8.l18 (1918) described. in detail three organisma, Baoterium
abortus, Bacteriwa bronchi8epti0Wl! and Bacteria _litell8a in order that theY'
;qight be oompared. Carbohydrate, gelatin and indol reactiona were negative.
24 .,
!!:. bronchiseeticum could be dist1nguuhed by its motility, abundant growth
and intense alkaline reaction. It. vas concluded that. a serological &tfinit.y
existed among the.. The sarum of a cow b;yper:1auluniaed with Baoteria abortus
was found to agglutinate melitensis and abort.us to a titer ot 1:1280 and
Bacteriwa bronchiaaRticUII to a titer ot 1140. The table below compare. the
tllree orga.nia1u.
Lit4ua urea Aspara- NO) reduced Increase of pH M1lk gin to N02
bronchisepticUl'l elk + + - 2.0
abortus alk + + sl. or neg. 0.8
melitensis alk + taint al. or neg. 0.8
FeJ'T'Y and Noble (1918) found 'that atter repeated subculture
Bacteriua t!J"tuasia had the sanae cultural reaotions aa Baoterium bronoh1.el?ti
~ Bacteria bronohinpticWll anti.erum. agglutinated not only BacteriUlll
bronch1a.eticwa but al80 Bacter1wo E!rtua8ia. Bacterium e!!:tussu antiserum
on t.he other hand agglutinated only the pertussis bacillus, with no ettect on
the au-aina ot Bacterium bronclrl;seEy1cva tested. Ferry and Klix (191B)
conf1nted, in most respects, the 300ft serological relationships using
complell8nt fixation teat.s.
Kraus and Klatten (1918) isolated three .trains of BacteriWll
taecalis alcalipnea from 8\OOls ot patients vi th enteri Us. They are
compared in the table below. Gelatin, ind01 and sugars were negative. It
_s not clear it strain 1821 had one or JU1'l3' polar aagella. The author
m.ak.a much ot colony appearance and states growth on potato was atypical of
- .-
motility
rlagella
aggl. with patient·. 8W'um
aggl. with B. raeo. alo. s.rua
1.921
lively
polar
1:400
11320
blue
sttain nwr.ber
2502
lively
peritriohou.e
1&200
la20
25
2789
-livelT
not demonsttated
1,40
1120
Archibald (1918) isolated trora the 'blood of' a natift ot Sudan with
atypical enter1ca an orcanulIl which bet named B. rlvax. It waa d •• v1bed .e -a gr_ negati,.., actively motUe bacUla 1.6 microna long. (The flagellation
was not recorded.) A pellicle val t'~d 11'1 broth. Gelatin wu not 1iql.l8£ied
Litmus ,mUle was not aoidified but becae alkaline in 48 hour.. Indol aM
Vogea-Proaleauer reactions were negative. Nitrate. were reduced. ACid,
but not ga., was formed in galacto.e and mannitol, while an alkaline reaction
withmlt gas formation ocourred1n gluco •• and other listed carbohrdrates.
The pattent's serum, in dUutions ot 11)0 and 1:60, completely agglutinated
the baoUlws isolated.
Webster (1919) recoverad BacUlue faecalt. alkaligene. trom blood
ot 3 tnf'ants with. enteric rever. 'the orcaniBu were reported a. being gram
negatift and motile with alkali production in sugars and mUk. (Flagella
studies _re not recorded.)
t'!AA+,..11Ani AM l'!hI'l1_p.l1919) defined the dnWl A.1' .... Jlli.".n •• as
r-------------------------------------------------------------~
26
tallon, Gram. negative asporogenous cylindrical long or short oella with
capsule. and which divide in one direction only. Ur()'1&,'th good il'1 ordinary
laboratory .-dia, gelatin not liquefied, aerobe. often f~cultative anaerobes,
no !luorescans or pigment prod action. Do net terment glucose or lactoae and
are characterized by their g .. neral lack of rel"Mntatlve power and by actually
increuing the alkalinity ot the _dill. 1'1Uk is not clotted or rendered
alkaline. llcaltJenes 1'aecal1s (Petruschky, 1896), emendavit Castellani
and ~r8, 1918*. Here also is listed Alca.l!i'::nes Yi"f'U (Archibald, 15'18)
which 'WaS obtained troa the blood ot an individual with inteat1n.al disorder
in the Anglo-Emtian Sudan. The organia. produced acidity in galactose and
mannitol and was characterized by ito lUrked aotUity. 3. ru~ia --- ...... alkal!J!r!!S (Petrusohky) gaft the following negative reactions' Gram,
gelatin, coagulated aerwa, lactose, saccharose, dulcite, fUMite, glucose,
malta •• , dextrin, rattinoe., arabinoM, adonite, inulin, sorbito, galaeto •• ,
lew1ose, ino.ite, salioin, &iq'gdalin, isodulcite, erythrite, glycerine,
indol and Vopa""Proskauer. In broth it producad 8 general turbidity.
iiotUitq wu poeiti," and litmus Jrdlk alkaline. The t;ypical B. taecalia -~kali&~ne. produces a etrong alkaline reaction 1n all auaer broths but
certain strains are laid to produce slight acidity :in glucose and maltose.
50118 strain. peptonin milk.
Cutellani and Chal.Dlers (1919) utle the spelling Alca.l1Jienas and
Alkal!enes, their new genus designation, indiacrwnatly. The J'l8M
... The citation tor 1918 io probably in error, 8S no description or the ienus
can be found .a of this 1.ate, hence it ehould read 1919. This 18 in agreement Wi th Weld.in (1927).
r-------------------------------------------------------------~
27
originally assigned by Petruschky to re:present the beer bacillus was also
giwn a variant spelling by these taxonODliats.
Jl.oQhaix and Blanchard (1920) tabulated oases or l_Bs'_'o_ill_}lS_" a.;."t~..o..;, ... CA .... -1;;;;;i....,s
alcal1&enes inf'eotion and round it. lUoat OC»lmlOn in spring and summer. They -incidence.
Watson (1921) reported the isolation of Bacillus iaecallff ,
!:.llcali,ene! f'l"oa the blood ot foUl" soldiers in India. The patient' Il serf4
showd high agglutination titers with the typhoid Ol"ganiWl aM. low
agglutination titera with Bao1llu~ ~&eoali8 alkaltienes. (A description of
the organism did not accompany the report.)
de Magalhaea and Bettencourt (1921) isolated BacUlu8 te.eoaliB
aloal.!ienes !rOIl the blood or a patient using & bUe .mediwa. The orianie.m
wa, gram negative, no .poreG, ftl"1 motU. with per1trioboul nagella. No
acid from lactose or glucose peptone broth. Milk we not coagulated but
became transparent and yellowish (alkalinization). LitDlus whey was uncbal:1pd.
Indol va. not produced and platin was not. liquefied. The patient t s convales
cent .erma agglutinated the organi8lll isolated in a dUution or 11400.
Raftnel (1921) isolated Baoillu faaoal.i8-al:kaltiene. trOJl the
spleen and intestinal patches ot a cadaTer. The patient had been 111 with
8TdlPtou ot typhoid teYC". r.xtrosa and other carbohydrates listed we,.. not
fermented. Litmus milk remained unchanged in appearance tor 8-10 days, then
it bee&:118 darke .. and showed a slight gelatinous coagulation. (The description
is not cOIllplete as to gram reaction. raot1~i ty and nagellation.) The
organism wa~ considered the causative agent in the production ot the
26 inteatinal leiiona.
Braun and Cabn-Bronner (1921) reported that Alcal!J!!!!s taecal1a
would grow aerobicall.y in a simple aed1Uil1l with ..ani_ lactate &I a source
ot nitrogen and carbon. Growth was poor but irlproved by addition of sultate
ion and aeration. SodiUlll chloride or pota.siUll chloride had to be present
tor growth. (A description of the orgam •• was not included.)
Cooke (1921) reported B.eUlus !!!'2!!aene8 •• the etiologic agent
of ammoniacal excoriation ot the buttocks ot infanta. the orpnisa vaa an
asporogenous gr .. positive bacillus which did not terment aug.,... This
yellow1lllh pignlented organism. bydroliled urea with the production or ammonia.
(This organisDl vas named !lcaliaene. aMOniaenh in Bergey's Manual ot
Detel'llinati ft Bacteriology, )I'd edition 19)0, pap )67. see the report
reviewed in t.his paper b7 Cooke (1926).)
stitzer (1922) isolated troDa water and stools 14 vibrios which were
not aa&lut1nated by anticho1era .erwa. They were recogni.ed a8 BacUlua
fa_calia ~oal*,.ne. by their biocheaical nature. He al.o studied the
intestinal nora ot cholera patients. On. the seeond dq ot illness and
through to the end. ot the disea.e when the stool beoue torlll8d, Baeillu
taecalis alcal'-eP!. W&8 predaainantly round. (A deacriptlon ot the
organiOll va. not giTen.)
Mudd and Warren (1923) isolated Vibrio percolana (new specie.)
trom hay infusion. This gram nagatift, asporogenous organin waa comma shaped
to straight, actively motile with one polar nagellum, though trequently 2 or
3 fla.gella were present. Growth in peptone broth had a pellicle, 1itlaua milk
unchanged, no liquetaction ot gelatin, no :reduction ot nitrate., no indol
r---__ ------------------------------------------------------------1
29 produotion, no fe~ntation ot dextrose, lactose, sucrose and other listed
carbohydrate ••
Khaled (192) in Egypt noted the blood culture from an enterio tqpe
infection reaul toed in the isolation of !:. £aeoalilf. aloaliptne8. (A
description of the organi81ll was not recorded.)
\:8ldin and Levine (1923) published a key to the speoies and
varieti •• of the colon-'Q'phoid group ot organi8lU. UDder the genu.e Bacteri_
was listed the au.b";genua Aloalipnea desoribed as gr&lI nelatift rods growing
well on artificial aedia, uporocenous, nonaotile or motile with peritrichoua
£lacell.. Neither acid nor C .. 11 produced tram glucoa ••
Sargey ~ !!:. (1923) detines Aloal.1swa Castellani and Chal..-rs
.s IIOt118 or noraotUe rode, generall7 occurril'l& in the inteatinal canal ot
normal. an1Jlala. 1')0 not tom acetyl-aethyl-carbinol. Do not teraent arI3' ot
the carbohydrates. The type species, Alcal:ainea tecalis (PetrU8~)
Castellani and Chal.men, ._ daacribed &8 tollow8.
Rods. 0.5 by 1.0 to 2.0 microns, motUe by means ot peritriohous fiagella.
Oram negative.
Gelatin stab. Orq .urtaos growth. No liquetaction.
Agar slant: White, ,listening.
Broth' Turbid, with thin pellicle.
Li traWt aUk, Alkaline.
Potato: scanty to abundant, yellOwish to brovnieb.
Indol not toned.
Ni vatea reduced to ni t.ri tes.
Does not tora acid or gaa in oarbobJdrate media.
r..-----------.
L
)0
Aerobic. facultative.
Habitat. Intestinal canal. Found in endocarditis but generally considered
nonpathogenio.
Alcal!linea bronchisepticua (McGowan) vu also included in this genuau
Rods I 0.4 to 0.$ by 2.0 Microns, motile, po.a •• sing peritriohoue flagalla.
Gram neiative.
Gelatin stabl No liquefact.ion.
Agar alantl Luatroua and mout.
Broth. D1atinctly turbid, with thin, grq pellicle and ropy aeditdnt.
Litmue rdllu Alka.line.
Potato: Fairly abundant, brownish, gl1aten1ng.
Indol not tor..ed.
Nitrates not reduced.
No acid or pa in oarbohTdrate .ad1a.
ADonia formed troa urea and upara&1n.
Aerobic.
Habitat. Bel1eYed to be the C&UM ot diateaper in dop_
Kollath ard Lubinski (1924) reported the isolation ot a gram
negative 1I1OtUe rod with curved shape £rem a patient with diarrhea. The
organi_ wu isolated trOlD the stool &1"l4 at autopa7 from the intestine, liver,
spleen and gall bladder. Flagella stain (Peppler) showed -inlT polar mono
trichous, ranly two flagella. no _pbivicboua or peritr1ohous types were
found. (Photomicrographs were not J;mblished.) In _d1a with dext.r'(,>se,
lacto •• , sucro", mslto •• or umite the litnwa turned blue. Colonies 0:'1
rabbit blood. apr were hemolytio. In the oovae ot study' the original ourved
torm. ot the organism largely disappeared and the organism becaae 8traight.
Anticholera serum did not agglutinate the orianisla nor a stock cal ture ot
Alcalipnes. -Brown (1926) reported a cae. al1n1cal17 11lce whooping caugh caused
by Brucella o/0nchiseptica. The organism 'Was actively motUe but the
nagellar lllC'rphology 1fae not recorded.
Row (1926) reported the isolation ot Bacill~ taeoalie alkalicenes
!rca the stool ot a patient wose general condition lfU that ot vphoid tever.
(A de.cription of the isolated organism was not gi~)
McDonald and Levine (1926) observed.. in a 81llthetia aedt WIt contain-
ing asparagine and mraoni_ lactate as nitrogen and carbon sour.e, vic aoi.d
to be produced by !!:. taecal1s alkal!lenes. Urio acid was also produced in a
peptone JHdiua. In the purine-tree mediUll !::. 2Zomneus alwqs tailed to
produce uric acid, "hUe on peptooe udiUDl it pr~duced uric aoid tnely.
Levine and Soppelanf (1926) in their study' ot bacteria in cre.-rr
was tea isolated ~~~~riUII ~~caliene.~ (Petruschky') Lehmann and Neuunn. 1h8
m.orpholo87 'WU rfescrfbed .a rods, no SPONS, 1I10tUe and VA Mgative. (The
fiagellar morpholoiY was not gtven.)
Dudgeon (1926) reported • .tad,. ot tbe buun intestinal f'lora under
normal and abnomal oonditiou. The Ilut.bor obeerved that B. fa8cal18 ___ I. alka1ienee wu delcribed as a nonMl inhabitant of' the intestinal tract.
HoveTer, trot'll. hil own experienee this baoillus ws "eldOll isolated trOll rece.
on brom-th.Jmol blue lactose agar.
Thompson (1927) ob •• rwd Alcall1t;M8 fecUil in a lHdium containing
0.2% dextrese as caroo!'l source and O.~ glutamiC acid as nitroge!l source.
32
(A description ot t.~e organiam wae not included in the report.)
Cooke (1926) and Cooke and Keith (1927) described Illere tully the
organism which she or1einal17 named Dacillua ~on1aene8 but now suggests
8aoterium amroonia5enes. It was described "$ a rod, nOl\Glotile, asporC'genous,
&TBlll positive, litmls :nilk unohanged except for production ot alight
alkalinity, gelatin not liquefied. and sugars not te:nll6nted. GoraJ. Cunther,
in a communioation to Einar Lei!scn dated l7-VII-51, stated Bacterlw.
awL"llOniageMs 1f68n A.'Tlericlt'.n Type Culture Collection and a .train ahe had
rece~."Nd directly from J. V. Cooke as 13aoterium a:mmoniagenaa to be gram
positive. This is in agreement ".i1th the original description. H. J. ~onn,
who revised the genus AloaltJenes in Bergey' 6 'ianwU ot Detenainattve
-eactertoloQ', 5th edition (1939) continued to list the orgL"1ima as lUcal.1gene.
4:Th'l1onialenes and as gra.'11 negative. The 6th edition of the .:1linual ot
t:eter:rlnative Bacteriology (1948) lists this organism in the J,ppend1x to the
genus AlcaliemUh
\-ljatt (1927) isolated 3aclllus feoalis aloal1pnea froll the blood
and stool of 8. nurse with lIdld t)"phoid-l1ke l)"InptOf!\S. The 88rUlJl a£flut1nated
rthe tew stra.i.n to It titer or 1:80. (A desoription 01' the organism was not
given.)
Weldin (1927) suggested the generic 1''' Alkalipnes, uaed by'
Castellani and Ch.a1:aers (1919), a. variant spelling of Alcalyenes should no
~onger be recognized. Weldin opposed Evans t augeetton that a distinction
~e made between Alcaligenes am Brucell;,a on grounds of pathogenicity, hence
~e included the meli tensia-abortus group in the genus Aloal1JEmes. The
~enue included only those organisms which produced neither acid nor ga.. trom
33
glucose.
teh1nann and Neumann (1927) named the organism under consid.eration
VibriC!. alcal!ienes (Petruschky) L. and N. This vas synoDY'.d1oua with Bacillus -f.ecalis alcaliaenes Petrusohky and Baoterium alcaligenes in their previous -editions. The authors were convinced by the investigation or strecker (1911)
that this organism should be designatel! as Vibrio or Se1rillUlll and was
unrelated to the colon-typhoid group. The organiema were described as
cUJ"'f'8d rods, often distinct norioe and spirillUlll fora. with 1 to 6 polar
flagella. There W&S no liquetaction ot gelatin~ growth on potato was
luxuriant and brownish, on sugar-free JI8<:1'.& it did not tON more alkali
than moat bacteria. It did not produce acid from sugars, and even dextrose
broth became strongly alkaline. The authors recognized that B. marienae -Klimenko (1908) and Vibrio terris!nus Gunther (1894) vere sauehow related to
these organisJu.
Gray and Thornton (1928) described two new species of soU bacteria.
P~eudomonaa dacunhae n. sp.
Rods: 3 by 0.5 to 0.8 microns, cU1"Y8d, motUe. (According to the drawings 1 to 6 polar lophotrichous nagella.) Gelatin stabl N.11head. Agar slant: Pale butt, S'1lOOth, shining growth. Broth: Cloudy. Nitrates not reduced to nitrites. No aoid produced trom carboqydrates. No diastase produced. Phenol was attacked. (Bergey's Manual of Petarminatift Bacteriology, 6th 6diti.nn (19M~) p. 105, retaines this orgsnia.'1l in the genus Psaudoaonu and state. it reduces nitrates to nitrites.)
Vibrio cunea ta n. sp.
Rods: 1 to 3 by 1 microns, l!UM'ed in a short spir31, m.otile. (According t.o the drawings they haft 1 to ) polar ~ultitrichoU8 flagella.)
GraJfl negative. oelatin stab I Liquefied. Agar 8lants Whitish, smooth, shining. Nitrates not reduced to nitrites. No acid produced from carbohydrates. Wo diastase produced. Naphthalene was attacked. (Bergey's Manual of Determinative Bacteriology, 6th edition (1948) p. 199, retaines this organism in the genus Vibrio.)
'rakayanagi (1928) isolated 58 strains ot 8acillus laecalis
alkaliB!nes trom feces and urine ot typhoid patients, and from pus in -appendicitis. Th.s. gram negative organis .. vare poorly flagellated, peri
tr1chous, otten only 2 flagella appeared at both ends (the Manina at this
description is not clear.) Dextrose and other sugars were not utilized,
milk was not changed, alkali formed 10 litmus whey, potato growth was moist
and brown. Antigenically the 58 strains were placed in 15 groups showing
little crossing.
Manias (1928) found that the organisms named Bacteria f.ecalis
alcal!lenes, iaolated by Yarioua authors, could be separated morphologicall1
and physiologically into two groups. The organis_ of the first group, whioh
included Petruacbkyts beer bac11lUIJ, were motUe with peritrichoUB flagella
and fermented·carbohydrates. The second group with polar flagella corre,pond-
ad to the group Pseudomonas, Kigula. He was of the opinion these species
could be called PseudOJllonas· aloal1genes. The 10 organisms in this second
group were gram negative, 150118 straight and some curved with polar flagella,
milk became alkaline Ilnd no acid was produced in carbohydra tea.
Gondolt am stringer (1928) found 10 cases ot !!:.. alcaligenes
urinary tract infections.
Marshall (1928), Daniel and Green (1929) and Danbolt (1929) reported
3$
the aolatlon of B,,,,cillua faecal is alkali&enes fro. stools, blood and gall
,tonet,. (The flaeellar morphology was not recorded.)
weis" (1929) suggested the nama Alcaligenes appendicalia f'or the
organislll he isolated f'rom the blood in three CA .. s of' acute a.ppendici ti ••
It was described a8 a gram negative rod; nonacid-tut; nonmotile without
nagella; no spores, ca.psules or lftetaohromatio granules. There was no
pellicle in broth, no liquetaotion ot gelatin, litmus milk uncbanpd, potato
,howed only soant growth without pigMnt. Blood agar colonies were gre;Y-lIhite
with no heJllOlyais. H7drogen sulfide vu not detected in lead. acetate agar.
Jkm6 of the oODUn~ used laborator)" suprs were fermented.. There W88 no
indol produced, nitrates _re reduced, _tql red and VOles-Proakauer
reactions were negative. Hyperimm.une nrUJlll cI1c1 not agg~ut1nate!:. t.zPl!9swa.
It agglut1nattad the hoJaologoua strain 1&2000, Alcaligenes recali. 11)0,
Alcaliene. bronohi8eptious lc)O, Aloal.!i!ne. ~bortua 1.120, AlcaliJ!n ..
IIlEIlitensis 1:120, and Alcalipnes booker! 11300. The descriptions of specie.
of Alcal.iene8 in Berpy· s Manual (192$) did not tit thia orpnism, MllC8 the
new name was suggested.
lendal.l and Ishikawa (1929) observed that the reduction ot _th71ene
blue by !lresting cells" in the presence ot certain CU"bon compounds was
precisely paralleled by' the tsr.mentation ot the.. nme compounds in culture
media inoculated with identical straw ot prolUerating bacteria. The
organiam8 were not described but BacUlua alcalipnes L"ld Vibrio H/61 did not
ferment t1rv suaerl. '!'he •• two organilms did not reduce methylene blue in
the pnulenoo of a lUlle l1U11aber ot sugars 'but tbere vas reduction vi th ot.ber
comp0un.d9. SOM ot the results are listed in the table below.
Tople,.. and W1lson (1929) state that Brucella broncbi8eptica produces
a beIIOlya1n acting on rabbit red blood cella, no production of hydrogen
.ultlde, grows best under aerobic conditions, no growth under strictly
anaerobic conditions. It 1. motU. b,.. means of 4. to 6 peritrichous flagella.
Gracey (19)0) and Lindemann (19)0) reported several isolations ot
BacillUS !aeca;Lis alcalienes frola blood, gall bladder, urine and testicular
abscess. (Tha flapllar ,uorpbolOU of the organia was not reported.)
Ib'ti (19)0) isolated 6 strains ot Bac1llus faecali. aloaU.enea
Ira blood, teces and stomach contents. All six were gl"ua negative, motile
rods, no spore., no oapsules, 6 to 20 perltricboua nagella. lndol_.
negative, gelatin nogati'ft, alkaline reaction in glucose, lactose, MnnitAl,
mal toae, salicin am sucrose broths, no heaoly'sia ot h'Wlllln blood, tallk not
coagulated, potato aoiat. sl1lq, ,..uow-brown layer. The aut.bor felt thue
organisu should retain Petrusc.hJqt. des:1gnation. He was or the opirdon
that the polar flagellated, alkali produoing rods should be another group
closely related to Bacterium fluor.scens non-li<e!taciena. The lophotrichous
group of Lehmann and Neumann and Pseudomonas of Migula forms one group which
aight be called heudomorr.as alcalipne.. Strecker'. polar nagellated
organisms, named Vibrio alcaligene. by Lehmann and "eu.'Unn, should not be
confused wi tb 'the p9r:!t.richous B.cmu. faecal!. alcaJ.:yenea.
Castellani (1930 and 1930) reported the isolation of Alkal1jenes
37
alk&lotoetidUIJ from tonsUlar crypts. This bacillus was motU., gram -n.gatiYe, no acid produced trom carb~drato., gelatin and coagulated •• rum
were n.ot liquefied, no indol was produced, allcali was produced in litmus sdlk.
Barrow (1931) and Gatewood (1931) isolated. 8acUIUII laeoal1a
IlkAl1J!nes trom patients with conjunctivitia aM Mningit1a, however a -de.cription of t!:e iaolated organism va. not given.
Evans (19)1) described an organi.aa she nued Alcal1enes faecalia
,ubspeci •• radicans. The organ1_ wu obtained from the blood ot a patient
with tewr resembling tj1"phoid. It was described &1 heine £1". negative;
asporogenous) motU. by _ana of peritrichoua flagella, no capsule. winkled,
dull, lIOderate UlOunt of growth appeared on an apr slope in 24 hours} 110
pipentJ liquefaotion of platin began trOll the surtaco. (PhotoJldcroaraph8
showing nagaUa were not publ1shsd.) Litaua aUk p-owth was aoOOllp&nied by
the developMl'lt. of an alkaline reaction which incre .. ed tor a week or -.ore.
No growth appeared on potato and there was no bMol.7eia ot red blood cella.
The organ1 .. vas aerobic. An alkaline reaetien was produced in broth
containing dextrose, lacto.e, 8ueroae, IUIlto .. , anno.e, %1'10", arabinose
and other ~tea. Growth did not occur in synthetiC udia oontaining
inorganio .al t. and oyat1ne, tryptophane or vic acid &II • aovee of nitrogen.
No growth. in Koaer'a oitrate aedium. Acety1 ..... t,byl-carbinol, bjydropn
aultide and indo1 were not produced. Antiradloanh Gena did not agglutinate
!lcal1pnea fucalis or other enteric organ1_. The auth.or stated the
organism isolated bY' Straub and Krais (1914) from blood bolds an intermediate
Position between the species taacalia and the subspecies radioana.
Slobeaiano and Naste (1931) obtained Baoillus tee&lia alou!enee
)6
f'rCl:B abdominal paracentesis nuid of twins with fatal peritonitis. The
Organism wu characterised as being IIlOtUe. gram negative, abundant, dull
gray viscous growth on agar, gelatin not liquefied, broth turbid with a
pellicle, indo1 not produced) 1itau8 J'dlJc becuae blue, unnitol, malto.e,
•• ccharose, lactose, galactose, and arabinose not tel"U18nted. (No record ot
flapllation.)
Asbelowand Mostowa (19)1) isolated 48 strains of Bacillus taecali.
Alcaligenes from the blood ot patient. auspected ot ha:ring t7Pho1d teftr. -'fhq vere gr_ negatift rods, otten curftd and with lift17 lilotU1tT, oolonie.
on Endo t 8 _diua appeared like tho.. of typhoid bacilli, indo! and gelatin
were negatift, .Uk Wb not coagulated, litau8 milk turned blue, no acid
or gas 11':1. Yarious carbohydrat& udia. Sixteen samplea ot normal blood .....
agglutinated about 20% ot the straw. Sera prepared in rabbits against 6
.trains agglutinated all the straw to 111000 or over with the _jority
agglutinating to titer. Nine atraina ot simjlar or,anisma iaolated traa
vater were serologioally heterogenous and showed no antigenio relationship to
the blood strains. (No mention of flagellation was made.)
Andarson (19))) cultured the blood trom two patients exhib1tina a
S;yndroM saulat1ng rhetllUltic po~iti8 and isolated ~r-'~1llU8 al!!YIenea
which vaa gram negatift and lett earbob,ydrate media unchanged. (The organism
vas not deacr10eci further.)
Mason (19)4) isolated a.eUlu i.calis aloallenes from spinal
nuid follOWing spontaneous opening ot a meningocele. 'I'he bacillul was gram
negati"., motUe, produoed no acid or gas in carbohydrate .edia, irdol
negative, lit1m.t.8 milk became alkaline and there vas no liquetaction of gelatin.
39 'j'bere was no agglutination of the organism by' the patients serua. (The
nagellar m.orpholo8'1 was not. recorded.)
stuart !l !!:.. (1934) isolated BaeWu, alkalil!ne! . .t'aecalis troa
the urine and a renal calculus relllOYed tra tM kidney of a caS8 ot chronic
lithiasis. The organiaa was not d8scribed other than that it was urea
rJ8gatiw•
Spra,y and Hawk (19)4) reported a cue of meningitis secon.dar7 to
bilateral otitis I_ia. The patient died and Al,CAlie~s feealis was isolated
from the spinal auid. The acti'V'el1l1lOtUe orgall1.sa was a short, gro
negative rod showing bipolar staining, 1'10 spore. or capsule., gelatin was
liquefied, broth bad a dittuse turbidity with a thin pellicle, lit.us milk
bad no initial acidity and beoue strongly alkaline, carbohydrate broth became
,wongly alkaline to brom."",,"sol-purple, iMol vas not produced, nitrates
WV8 not reduced to nitrites, blackening did not appear on lead acetate peptOl14
iron agar and colonies on Elldo', med1_ were colorle88. The organi_ wu
considered as the specific infecting agent "8poneiol. tor the death of the
patient.
LeU,on (19)S) reported new culture _dia based on sodium
de8ox;ycholate. He obs8J"V8d that Bro~t1cum (peritrichoua) grew fair17
J"8adUy aM Alcal!«!nes (lophotrichous) grew _11 (one strain) on
d.8oqcholate agar. neaoqcholate cit.rat.e agar general.ly inhibited Alcaligene
alaost completly but 80J118 storaina de..,eloped tiny colonie. in 24 hours.
Varioue type. ot mononacellated bacilli including procyaneua grew well on
the madiUDl. The author stated that onl.y polar flagellated bacilli were beilli
included in the genus Alcal.1J!rw ••
40
Saiaawa and Takahaahi (1935) recognized that. Bordet and iIIIIUl7 worker.
onl since described the bacteria causing whooping caugb as being vi thout
flagella. These workers c01'l8t.antly found the Bordet bacUlus to be tlagellat.
'the nagellat.ion was the same aa t.hat seen on Brucella broncilil!Uilpticl.¥ll.
NyMri (19)5) examined 134 organisms labeled !!:. alcaligenef1, 82 ot
en 'Were divided into 2 distinct grOl.lpth
Group AI 71 strains )2 "wage 6 teces ot normal individual.
22 feees of tqphoid and paratyphoid patients 5 urine-pyelitis 6 old laboratory strains X"6ooi.wd trom. Ryti
y were short thick oocooid bacilli easilY mistaken tor Etapgyloooeci. ~o
.s were tormed. They ware mostly nonMotile but in hanging drop some
Zett.novte .taL, showed tn-.y were peritk·ichou8 with 1 to
nagella. seldom 6 to 7. No liquefaction or change ill milkj indol, naethy'l
d, Voge.-Pr08kauer tests were negative; no ac.id trOll 8UlJlirS, however
u1ll0n sugAr Ja&diWll cbanged traa pH 7.45 to 7.35 in one seri •• , in another
aries froa 7.21 to 7.0. irihen the pH roae trOJll 1.45 to 7.75 the orlaniu
d not belong to this group_ Occasionally lophotrichous and trans i tional
orma were encountered which lowered the pH. Mutants and variants werill
ola ted from this group which peptonized milk and iave acidity in xylose and
extrose. This description does not cerraepond to Petrusehkyts original
.scription, but M]berg considered the Bacterium tMcalia alcaligeM8, in
spite of the poor _tlli ty am inore.eeGd acidi ~ of suga:r media.
Group BIll strains S typhoid-paratyphoid rece" which also contained Group I . .3 sewage
'" srowth• Lophotrichous with :3 to 4 fiageUa. These were identical with
Vibrio aloaligenea Lehmann and Neuu.nn. Thi. organiam had been called -Alcalietll1ta raecaliB in the literature but the author called thea Vibrio -aJ,calirn8S. TheY' produced an alkalinization of the dextrose _dium.. -
Group c: S2 straina 2 typhoid-paratyphoid feces So .ewage, incompletJ.y inVestigated
TheY' have characteristics of both Oro~ A. am B. MotUe rods which change
the pH ot augar aed.1a from 7.45 to 6.) or 6.6. (Notice that th18 i. lower
than Group A.) They have about 4 teradnal. flagella, but behave like Group
A on fUny media. The.. _" referred to as Pseudomonas tOl"ll8.
Hasen and. Mortillaro (19)6) described an organism isolated from
the blood in a ca •• with typhoid t .... r 8yaptolU. The organi_ was ref.rred
to as a hi terto und.scribed microorganism ot the Aloal.!I!ne. group and vas
described as follow: Small gram negative aerobic bacilli with no flagella,
capsule. and polar bodies ware not demonstrable. No hemolysis on hors8 blood
infusion agar. Ho liquetaction ot platin. Grovth aoderate and ot ligbt
CHUI. color on potato. Litaua lIdlk was unchanged. In milk containing either
phenol red or broa-oresol-purple, a slight a.lJcal1ne reaction waa obsen'8d.
Indol was not formed. Acetyl-DtlthTl-CU'b1.nol was not demonstrated and the
-thyl red teet va. negat1.... Carbob;ydratea in peptone water aediua were
QOt terRl8nted, how .... a strong alkaline reaction vaa obserm. There ..
no serologioal relationship of the organism to Brucella bronchiseptioa,
abortus or mel1tena1 ••
McInt.yre (19)6) reported tbt isolation ot '.calia al.kalipnee from
the blood of a patient with acute hepatitis and jaundice. The organisa waa
-42
., reported 8S being motile but was not ducribed further.
Henriksen (1937) reported, in a dusertation on the bacterial
flora of the respiratory t:ract, the isolation of a nonmotile, helllOl,-t.ic
organis.'I. he ruuaod Alcaligenes heIlol.laane. It vas isolated trom elewn
,pecimens ot sputum of patients with chronic bronChitis, 8stmatic bronchitis
and other conditione. This erganip was very svol'JllT beIaoJ.7t,ic and the
baOlYsin vas filtrable and antigenic. The straiDa grew vigorously in broth,
toraing a pellicle and a SOl.tll8what rlscowa dtlposit. s.. straw also grew
protuae17 below the surtace with tbe .. diUll beOOl'l1nc denaely' ",VOid. Milk
110",ly !orMd a 80tt ooagulUll which later peptoniaed, but reu1ned alkaline.
Gelatin wae liquetied and coaplated S.l'WI vas not liquefied. Nitrate. vere
not reduced. Fementation ot dextrose, lactose, sucrose, malto .. and i4U1I'litol
did not occur. All .trains were unif'ora blocheaical13 and .el"ologioall7.
Bradford and Slavin (19)7) reported that the morphological, b10-
cheJllical am t.unolorical oharacterutica ot Ha_o2hil~ p!rtussis, Bl"UOftlla
bronchi8!2ticua and the p!!:aJ?!tlussu bacUlus lIIW'e aiDIilal".
Eldering and Kendrick (1937 and 19)8) observed the close relation
.hip or BacUlus 2!rtU8sia and S,,;c1l1us bronch1ael?tic~ to B4CB1" P!!"a
e:rtussia on tbe buis of morphology, growth on variOU8 ud1a, biological
characteristics, .erologioal reactions including &Iglutinin aDsorption teste
and certain reactions in an1Dials. Although thes. organiau vve siaUar
J!l!pertU$l!d .. was identical. with neither of the other two. '1'be7 point ou.t
that Bacillus parapertussis un not. be placed in t.b8 genwa Haeao:e!!llua since
it grows readily' on ordinary blood tree aedia.
Goldberg (1938) presented a ca.. wbich c1.ini~lT appeared a8
43 .. nl!lC:ttifl • 'he bloorl culture on 2 occasions resulted in tht~ :\.tJolation of
"hat he cal1ad i3acl1;~ aloall~enes. It was gra."a negative, C'tUt'hl'l litmus
aUk to b$oome a.1kalino and Russel's nte11um to sholi no gM tnt slight aeid
wa" present 'tl1th an alkaline !Slant. (~o further description was Given.)
watanabe (l9~8) bel::'eved that BAcillus !;!!!"tussis changes !nto a
aotile animal ~ EacilluB pertussis. These two organisms res&~le
B. bronch18eEtiou1!l in morphology, cultural behavl.or, ant.1cenicity and -biochemical characteristics as well as in the t.oxic1ty mld intra.cutaneous
reactions ot the toxic substance obtained from the bacterial water extract.
The author was of the opinion that the change £rOll Bacillus p!ri.us8is to the
80tile animal type Bacillus p!rtussis was a mutation and not a variation,
,inca he bad oOa.rftd a human case naturally Wected with the an_I tn>e
am the DlOt1lit7 waa not reversible. The author also lias ot the opinion
that the reason Europeans a..'1d Am.ericans had not recognized the experimental
whooping cough of dogs vas that this bacillus was not regarded as a variant
ot the Bordet bacillus, but mi.talum for ~ bronchisepticus.
Berthelot and J\rmoureux (19)8) reported the isolation of Bacterium
alcaliJ!n!. trOll purulent knee exudate.
BPe and Hartraann (193A) reported the isolation of BacteriUll
alealigene. trOll purulent knee exudate.
Evan!; and Maitland (1939) described the toxin ot Brucella
brO.nchueptica and found it sillilar to the toxin of ilaeaoi!hi;l.ua J?!rt.U8si'h
leither toxin was entiaeme and both were heat labile. They alao round that
.era from ca.es of vhooping cough talled to neutralise p!!;tuss18 toxin. The
relationship ot the •• erganis. was evident since thq produced the saIH kind.
-'" of »cutc If:t-ions in the lungs ofax.periinentally Wec:ted mice. Antigenic
relat,ior.S}li;' 1.as df.;ncr.strf).t~;j by i.llfill:ni.atic~, precip!tat.ion, cO:ilplSlIisnt
t'iXAt..ion, f!tEllltir.ation and ag~~lut1nation absorption tests. 11;,$ cultl.::ral
and ~orphological 8imilariti~a were also described.
i3rllci::ner 2nd !~van(l (19J9) studied the toxin ot HaeBlO2hilua pertu •• i8
Brucella bronchisept1ca and Baoillus p!rapertu8.ia. All thr.e produced ... ,Wlar lesions in litxperi.msntal an1mal... 'their tOJdna were separable from
~. bacterial antiisns and were alike in their reactiona to heat., tor4li'Uin,
filtration and lack ot anti&enici~. The results ot agglutination,
aompl~uent fixation and precipitation tests ~ in agreement with those of
Ildering and Kendrick: and show that the three OI'iardalu are c1.o.ell' related
antiEemicslly. The authors concluded tb4i.t the data justify placing the
~.e orianisms in the same genus.
Conn at ale (1940) state tr.at. the original description of Alca.l.iene
taecal1JJ (a.aeUlus faecalu aloal!ien •• Petruschky) .ale •• it clear that it ia
• peritrichoUB, non-spore-foTldng rod ot intestinal origin, turning aUk
alkaline and prodilces no acid in Dugar bl"otha. Conn recognized that many
workers described it. as polar fiagell61ted. None or the above oharactel"ist.ics,
Conn points out., serves to distinguish it f'roa Rhiaobiua, Pb~nas,
Bacteriwn r&diobacter, ChromobacteriWtl and AchrOlAobaoter. COM collected
22 cult.ures labled Alcaligenes taeoa.lis. All agreed with Petruachky'.
original description of the organism except for three strains, one of which
vas nonmot.ile 1.1 .. other two were polar flagellated. The ot.her 19 had what
t.he author called ttdegenerate peritriobous aagallation. 1t All but a 5JUll.
~ority ot the cultures had the following cbaracteristice. They produced
4S '" aJ,kalinity without peptoniaation in litmus .uk. grew poorly in synthetic
rlUC088 media but growth was greatly stilaulated b)" tlut add.ition of a amall
aaount of 18&81;. extract; there was no evidence ot any acid rrom sugar except
CO.' but 1901118 or the cul tv.,s produced this in abundance in a glucose _diwa
,uitable to their grOW'th. About half the culturea studied produced nitl"ites
trOltl nitrate.. He concluded there was a well recognised organia to which
tb8 name Aloal1J!nes taeoal1s could be .uigned. Conn, a eoU bacteriologut,
cautioned that all intestinal organis .. , producing a.l.kalinity in litaua mUk
and no acid in gluco •• , ru::; not M Alcalyenes raecal1s since otber types ot
organ18lU also have tbe.e characteristics. Or,ani... isolated fl"Oll tfmllent-
1ng sauerkl"aut and f'r0ll& a Cuban .ugar .ill proved impossible to d1atingu1ah
trOll the typical AloaJ.!&ftne. fae04lie. Ma.nI' 80ll bacteria aimilar in
~.iology were found to nave the same type ot degerllrate flagellation as
llcal1,en.. fuoalis. (Photomicrographs did not accompany the paper "but
drawings were included to represent the t)"pea of flagellation of the yuioWl
organisms. )
Barron and Friede-.nn (1941) oba.ned that Alcal;1lenes faecali.
did not oxidise ,lucos. but did oxidize hexose lIlOnophosphate and hexo ••
diphosphate. This r.UUN to oxidize glucose vu stated to be due to
1na.bUity to ph08phoJ71ate. (The organi .. was not described.)
Pob! and Raymond (1941) described a oase with abscess on the neck in
whioh the etiolOgic apnt vas bel~.e'Md to be Bacillus 1'"0&11$ elcal!enes.
The d1tterentiation of this organiam. froa Sh!iella alkal •• cellll and ~
!lJcalesc.ns vas .aid to be indetendnat.t!. (The bacter1010gio.l defjcription
or the isolated or,anism was not included in tl. report.)
r--------..
l
46 .,
R1dering (1941 and 1942) reported the isolation of a protein tree
polysaccharide trOll Brucella oronohiaeptica" Haem9Ehilwt. pertussis and the
parapertussis bacillus. iTotection teste with the polysacoharide tract.ion
and whole killed organisme emphasiMd the close relationship of thes8 three
organisu.
Voorhies and Wilen (1942) recorded a case of bacteremia oy
Aloal.yenes taecalia. The baatereJda and. tewr lasted tour months. Interest
ing manUe.tations lIVe a po17arthritia and _nine1tis. '!'he organism was
charaoterised .. slightly liquefying gelatin, no t<31"'J1entation of luted
carbohydratu. Litllua milk becae alkaline in 4 days. The orcani_ was 8
slightly motU. gr_ n.gati .... rod. No aglut1nation ooourred with the
patients .er_ and the isolated organi8a.
!had (1942) observed Baoillus taeoalia aloalienes 1n. oat.heri.ed
urine ot a patient with C1'8titia. It. P'h' out on MacConkey's asU' and was a
short, JIlOtUe, non-spore-bearing rod. Glucose and other carbobydrates listed
vere not fermented but alkali wa. tonaed. L1 au lI1lk showd prolftsai ve
alkali formation. Vogea-Proakauer, irdol and -tb71 :red. tests were negative.
Liquetaction ot gelatin or reduotion of nitratu did not. occur. Convalescent
serua agglutinated the iaolated organ181l in a dUution ot 1.200.
Conn (1942) attAtmpted to learn wbethAtr certain Boll wgani8lU
belonged to the genus Alcaligenes. When the t.7P8 specie. ~1caliI8ne. taecaliB
Castellani and Ch.alMrs (Bacillus taecaliB &lealippe. Petruschlcy) vas
investigated, it was tound that the type species wa. not well characterized
and, the genus was 1ll-detined. The study shoved t.hat at least thr.e species
vera being distributed as Alcal!&enes ta.calia,' 1) a peritrichous rod
47 requiring organic n1 trogen, 2) a lophotrichous rod requiring organic n1 trogen,
.I'd 3) a peritrichoua rod using nitrate or ammonium 8alta as lole Bources of
nitrogen. It seemed probable that the first vas the organism. ori&itIall7
described b7 Petruschky. Th18 organia appee.red ver:r similar to Baoteriua
ltronchi.epticua. The second was excluded trom. consideration .s the Petruachq ... u
baOUlu8 because it had lophotrichous nagella. The third agreed with
Petrua~'s description and had been distributed b7 type cult.ure collections.
Its fecal origin was doubtful and since it can grow on synthet.ic Mdia it.
probablT belongs to a different. genua. He concluded that strains reco'Y8l"ed
trom Boil which did not produce acid or gas from. carbohydrate media were
related to the third and not to the first group.
Conn (1942)reviewed the cl.aaa1tication ot PetruscbJq'. orsan1a
through se.,..&! editions ot serpy'. Manual ot Detendnatift Baeteriologr.
Of the various cultures which he accUllUlated frGlll the A.lllvican 'l':f.pe Culture
Collect1on, the National Collection ot 'l)'pe Cul turea, C. Nyberg in Helainki,
Pribra's colloction in Chicago, E. o. D. Murra,.. in Montreal and ot.herB,
IODl8 were lophotrichous and obviously belong elaewhere in the taxonomic
Icheme. He recogni.ed that bacteriologists generall7 were regarding &l\Y
nontermenting &1'- negatift rod ot tecal. origin &II Alcal!rne,s !aecalia. In
COIIlparing the .oil forma with the intestinal fonu he found a .bdlar
aarphology, and aoid and gas tests on ordinal7 media were al._ya negatift.
Production or nitrite £rem nitrate proved a ...... 1able oharaoter. Two feature.
were said t.o be useful tor making d1at1notional 1) ability to gl"OW on
l1nt.hetio Dl8dia, 2) produotion of CO2 .troa glucose. The following two ltedia
-.re usedl
48
Mediwa I
(NH4)2HP04 1.0 g.
Kel .2
MgS~·7H20 .2
yeast extract .2
agar lS.O
waur 1000.0
Med1UBl II
sa. u above with the addition of glueo"e.
'the soU t01"JU showed scanty growth on Med1um I, but very vigorous growth
011 Medium II. Only rift of the eu.ltures received as Alcaligenes taeoal.1a
be.bawd a:1JllUar17. The l"emaini."lg str9.in8 of Alcal:!,ienf!f! ~a.e.~al13 ohm ... d poor
OJ" n<' growth in either _1wa. Then was no evidence that tJ.Ce.li§()l'!E:I' f'aec&J.l~
could grow without organic nitrogen. CI')rlrl eoneluded that Qbund~nt CO2 w.aa
produced b7 the soil forms but not by typical Alcalie~! ~aecal.i~. Based
on this erid.nee COM questicJfls it aUTone knows juat what C0fl3t1 tutos t-n
Alcal.!lene8, 8ince it could be either those organisms which groil en his
Iynthetic media or those vh10h ~quire organic nitrogen. Soth types wer6
being distributed under the same nmna.
Pirot and Bour2a1."l (1942) reported ~berrant oreh1ti:s in ,trl.l):;t;!. pigs
III being due to Bacillus ill!11!.£enes f!.8c?J11,£!, var. !!!1!:2!!.
Leit8o~ls textbook (1942, page 178) shows distinctly, for the
first ~ime, the peritrichous flagellation of Brucella bronchisepticum. He
also suggests that this organism may be out of place in the genus Brucella
and should, perhaps, more properly be placed in the genus Alcaligenes. The
49
1J'1COns1.8t.ancy" regarding the basic characteristics of the genua Alkaligenes
..... recogniBed (page 181). The Alkala.,enes aoat cOlllllOnl;,r encountered in
,toOls have very characteristic polar flagella, (photomicrographs show
10pbOtrlchoW!l tong) quite different !ro. tbe aagalla ot most bacteria. The
peritrichous type was stated to be inhibited to a P'Ut extent b7 bile salts.
Anatomically and physiologically the latter seeaed ident1cal with Brucella
brOnch1.8epti~. -Weintraub and !'fetal" (194) reported a ca •• ot baoteremia coap11cat-
inI diabetes melitue in which Bacillus aloalienel! va8 isolated 3 tiM. trOll
the blood. No apecitic agglutinins vere found during the 1ll:ne... Growth
ot the organi8ll on plain agar was abUDfant. Dextro.e and other luted
carbohydrates WN not turned acid. L'ldol vu not produoed and gelatin ••
not liquefied. Conn exandned the strain and stated it required or,an1c
nitrogen to &:rOW. (Morphological studies weN not recorded.)
Raeburn (1944) described) ca ••• ot enteritis occuring in EQPt.
which appeared to be due to an organism like Nyberg t. Bacteriua alkal!:l!ne ••
serum of the pat.1ente agglut.inated the isolated oraani_ in 1110 or la2$
dilution. The oraani.811 isolated va a gram. negative rod, feebly' IJiOtUe.
Dextrose ani other listed carbohydrates were not termented. Indol was not
produced and litmus aUk became slightly alkaline. In the l1tmua a1l.k:
reaction tb organiul diflered trQlJl H;yberg'a BAOter1U11l alkal!ene~. The
author cOUIII'lented that N)berg' 8 propoaed separation provided no clue a. to
Which, it either, ot his groupe or ol"ganiPlU were responsible tor the
1ntectiona reoorded in the literature.
Cbaudur1 (1941.) iaolated BacteriUlll aloal.!enes and Bacteri~
so .. ta1calilenes from 6 patients, 80M or which bad specific alglutinins in -ib8 serum. (There waa no d •• cription ot the isolated organism.)
Shulman and Johnson (1944) reported the "cowry ot a a1aroorganism
frOiI. blood and spinal fluid ot a preature intant. It. wu a .hort, straigbt,
lI'am negatift baoillus. Flagella, endospores, capsules and m.etachromat.io
,.nul.,. were absent. There was no hem.ol7si8 on rabbit. blood agar; platin
va. liquefied. growth va. slightl,. yellow on &181" alant. The or,aniaa vaa
.. robic and growth va. prevented by bile salta. Litmus aUk vaa alkaline,
reduoed, peptonized and ooagulated. Indol, acety1-m.ethy'1-carbinol and
!qdrogen sulfide were not produced. Nitrates vere not reduced. Methyl red
and citrate apr teats we" MgatiTch Ho aoid or gaa as produced in
peptone-carbo~ate broths, but 1ft a 81Dtbetic oarboh1'drate _diUII lB&n.Y
lugu. were t.,..nted. Th18 baoillus ... 1aUar to the soU orpa1au
.tooted by Corm.
Wut1 (1945) reported tour cae •• ot enterio rever in which blood
oulturea showed Baoteriwa faecalis alka}..!Enes and the patients una
contained qglut1nine tor the uolated oreani.. in t1 tel'. vU"'11n& tr'ODl 1 a $0
to 1:1$00. The author alao round 81lht blood culture. positift tor this
organi_ in which aulutinins were l .. s than lase or had no t1ter at all.
(The OJ"gllllliaa vu not described.)
Alderson (194S) de.oribed a nn MIIIbV ot the genus Alealipnee
which differed cul turall7 and plV'aiolog1oall,.. troa other described species
of the genus. The outatand~i.ng charaot.eristi0 of thu organi811 1IU ita
pronounced acid tolerance. It was a ..u gr_ negatift rod, which produced
a aparce growth on the oOtaOn culture aedia as compand vi th Aloaltenea
51 f.calith GroWth on nutrient agar .... poor. Broth growth was sparce vith a -ganular precipitate, the supernatent liquid being clear. Gelatin was not
liquefied. lit.mua milk was rendered al.1cal1ne, withou.t coagulation, and. vas
.,.rt slowly peptoniaedj no growth on potato. Carbohydrates and citrate wr.
not ut.ilized} n1 trates vere not reduced, indol and pipent was not produced
and blood was not altered. ,No mention 18 -.de ot lIOt:Uit7 or flagellation.)
A1tem.eier (1945) reported the marked 8ene1tb1.ty or tift strains
of Bac1llUB alkalienes ,aecalis to penicillin. (Io d .. cripUon ot the
organl8tU vas given.)
Fulton et &1. (1946) reported that the tindings ot Altemeier could --not. be confirmed. Diepit.e the tact that the genus Aloal.!ienes vas .eparated
from the Enterobacteriaceae and grouped vi ttl other genera in the t&lrl1l.7
Rhilobiaceae, it occurred. with, and re .. JIlb1ed pn,-aiologioally, the a8lIbers ot
the fOl"Mr t&l1117. The r.aMb1ance includes iD8ensltiv1ty to penicillin.
Oeding (1946) isolated Alcsl.1e1l!.!. beao1z!ans trOll the .,... ot a
DlW born. The child .uttered trOll a aoderate purulent conjuncUv1tl. in
both eyes. The strain was de8cribed am found to be in accordance with
Henriksents (l937) description. In addition to the O&.e described above the
author found 90 ca.e. in which Alcali;eaes vas isolated from clinical uterial
and entered in the laboratOl7 diarie. durin, the preceeding 7.S years.
Organi_ rout1nel1 .... re classified as ~lcalilene~ if they were toum to be
gram. negatift rods, whioh did not teraaent suprs or torm pipaent. The author
POinted out the possibility ot ra1stak1ng wale pipent producir..g Pseudomonas
etrains as A1Mligene ••
Dick (1946) tound Baeter1U11l faeoal18 alkal.Ysenes in the blood in
S2 '"
ti.e o&ses in which enteric group orgaDiems were also isolated fro. blood,
urine or teces. In three other CAses. w10h wre clinically enteric fe ... ,
Baoterium la.calis alkalil!nes vas iaolated tra. the blood. (The or,an1am vas -not. described.)
Wilson and Miles (1946, page 786) ob$eM'ed that. :l.n the 5th .dition
of Bergey' a Manual (1939), aacterita bronchi.eRtleu .. included in tbe
~lla group, who •• type speci •• , Br~a ~i tens 1. 1t re •• mbled 1a -JIOJ1)bology and carbohydrate lementation. This waa in aoooa-dance with :bans
(1918) • WUson and MUe. point out tba antigenic relationship between
• bronch1aeeticua and BaemoEhllua pertuas1e and sucgest reclas.ification.
tigenic siJa1larit7 alone does not necessarily indicate a taxorlOlldo relation
th1p, but in th18 oa_ other resemblanoes are suffioiently good to warrent
visional 1nclusion ot h bronchisel?ticus in the ... group as HUJIt.02hUua
tussis. The cuI tural relationships betwen ~~e!!11us p'!rtussi8 and
rucella bronch1aeRtiCU8 are as clo.e a. tho.e between B~oella bronchi.eDttc
~!ucella .. liteneis.
Cos .. r,r (19h6) isolated Bact:. raeo&lis aloal.!lenee tram blood ot "
t1ent with enteric tever. (The O.I"ganiea vu not described,,)
Idruan1 and s.etbaraan (1946 and 1947) lound ~aoter1W1l eloal.!.l!nea
dead duck _bryce. It vas a notaotUe, P"D negative rod, on agar colonies
loped le8lOft-yellow color J no liquetAotion of gelatin; no chanp in glucoa.,
eto •• or sucrose broth; Voges-Proskauer, _thyl red and indol reactions
re negati.ftJ no reduction of ni.trate. litmus ndlk va. render~ alkaline,
drog8n sulfide va_ produced. 't'he7 suggested this orgama was Bacteria
-S3
..,
Spanedda and Aru (1946) studied )6 s'b"aina ot motile and nonmotUe
Alcaligenes isolated trOll feces. -Rush (1947) in a dissertation presented to the faculty at Purdue
univeraiv, on a taxonomic study of the .sophllic Achromobacter, observed
that S serological grou.ps ot his collection of 40.5 organilJlU did not change
litmu8 m.ilk and did not produce acid in dextrose llediUIJ.. All were said to
baft been stained 0,. Grq' 8 method for flagella and fouftd. to have peri'b"icboua
aagellation.
Terry !1 !!.:. (1941) isolated Alcalienes faeeal,ia tour t1us trom
.pinal tluid and once frau blood of a patient which se8Ul.Gd to establish
bacteremia and acute _ning1 tis due to this organism. 'ine or,mU.II was a
_tUe graa negatlT8 rod which did not produce acid or gas in carbob7drate
_dia, 1itm\18 Bdlk bec8.lll9 alkaline, nitrates were reduced to nitrites and
indol was negative. It was identified on this basis as Alcaligenes taecal"U.
Specific agglutinins for this organism were no'ttoum in tho blood at an;r
time.
Dyer (1947) studied microorganisms .frau Atlantio ood. She isolated
tift strains of llkaligenes aaon!!enes and tive strains of Alkal.ijenes
I118tallcaligezwa. They were assigned the,.s. names on the bu,si$ oJ: th.
descriptions giwn in the Sth edition or BergeT's ~1anual.. (The Ol"ianisu
vere not described in the paper.)
Shrewsbury and Barson (1947) reported the isolation or 2 groups
o! organ18llla which appeared to be Alcal1genea. They were isolated trOll
!rion ~ (black slug). The mot11it,- v.aried but flagellar studies were not
recorded.
S!& ;·iont.estruc !! !!.:. (191,,7) reported they found. Alealienes faecal!.
-!n dysentery stoole. (The organ1sme were not debcribe,ci.)
Bieebor!' !! !!:.. (1948 and 1948) reported the isolation ot
~aliJ!ne8 reoalia from 2 cues of baeter_ia .00 meningiti8 in infants. The -orga.niSDl was described I • .. gram negative aotUe rod. Gelatin was not
liquefied; indol, methyl red and Voges-Proakauer teata were negative. Dextr08
and other listed carbohydrates gave a negatift reaction; litmus !IliL1( was
.lowly alkalinizlt(f.
Saturm and Bourbon (1948) studied both a&cteri~ alcal;e_. tftcalia
and t~ae!9EhU\up !3r.~,poh~!'.e~icus. Thes. orlanisru had the $;1lM morphological
and biochemical properties. They were ::IlOtlle with lophotrichous fiagell&tion
(with the exception of tour etrains)J grR!Jl neptlve with no spores or capsule.
gelatin 'AS not liquefied; dextrose and other lute~ csrbor..;;r:"..rntsa wers not
fermented, aerobicJ hydrogen sulfide, indol, methyl red and Voges-Pl'"oskauer
tests wve negatlft, nona were hemolytic tor horse :red blood cells, litaus
IIlilk was always turned blue. (The nagellatlon was not substantiated with
photomicrographs.)
ShrenbU1'7 and Barson (1948) reported the isolation of orgii;rri.~.ms
Ibdlar to Baeteri1ll! .:leal il!ne" frOID. Periplaneta aruricaM J the American
cockroach.
Moustata !! !!:.' (1948) isolated !!:.. t .. calb a.lk .. liene~ while
studying tood poisoning in Emt. (The organia was not described.)
Qu1ntos and Buensueeso (1948) reported the isolation of several
strains ot Allcal1jtenes during 4 study ot diarrhea DOng Filipino Wants.
(The organisu were n,ot described.)
SS Sexiey's KaIlua! ot Detenrdnative Baoteriology, 6th edition (1948)
de'cribes Alcaligenes f •• oalis •• tpllowsl
Bod': 0., by 1.0 to 2.0 1l1icrons, motUe with perit.richou8 flagella. OJ'811 negative. oelatin stabs No liquefaotion. Apr slanta white, glistening, opalescent, undulate uaargin. 8J'Otb l Turbid, with thin pelliole, am moid sediment. Gi ..... oft MalOnia. Litmus milka Alkaline. Potato' Scant.)" to abumant, yellowish to brownieh 1J"OWt,b. Indol not torated. 11trite production trom nitrates variable. 10 aoid or gas trOll carbohydrate _dia. ,.rObie, facultative. Habitat: Intestinal canal. Generally considered nonpathopnio.
AlOaliJenes taeoalis var. radicans 18 luted as a platin liquetying. .in.
Pulvirent1 (1949) reported that. !:. f •• calis alca1!encs produc ••
h1drogen sulfide. however the other characteri8tics of the orgamea wre not
published.
Roland and Bourdon (1949), recorded that Alcaligenes fecal1e giTeS
• variable Ferguson-Hook urea reaotion.
Hall and Oan'an (1949) reported a fatal ca .. ot septicemia due to
Bacteriu. faecali. alkalilenes. Tbs motile gram negati.a bacillus was
1.o1ated trom the blood in a tebrile illnes. lastinc tor .are than three
aonths. Dextros. and other listed carbohydrates WN not fermented, indol
vae ~v,,·~;. produced, Ii taus milk became alkalh:f·. No agglutinins tor the
organism isolated were found --, 1 blr.-od or the patient. (The flagellar
IIlOrphology was not reoorded.)
Bassler and Peters (1949), without listing ret!U'8nces, state
_cal1genee taecal18 i. found in about 20% of nonaal. stools, with the percent
ge rising in intestinal intla:rwation. Vibrio aloal;!.p1l88 appears norully
in the int. t
'" ~, they are capable at 't1.aes ot pnduo:1nc an enterltia, and can be
~red troa the blood in "8001&t1ca 'II1t.b SalaoneU~.
Banerjee and Sarkai' (1949) ... ported t.b.e isolation of Baeteri_
, .. calia ~!r!!s frca blood in 21 .... , t .... aselUe nuid in 2 caMS' -troa abae ... pull in 2 .... , bw& knee joint. fiu1d 1n 1 case, am frca call-bladder bUe 1ft 1 .... fbere were a t." ~s of n.onapl\lt1M.tlon ot
the 1.Iolated etrain with the patients ..,.., bownv attAar repeated a\&bc:ndtun
tbe OJ"1",nau agluUnated. (A duor1pUon ot the orgard.ne 1aolated ..
DOt g1ven.)
01_. (19l.9) reported in his d1eeert.aUon the 18ola"* of
Hellconttll& £I'0Il 6 ohUdl-en. Thu paoUU... oraanlaa could onlJ be 1solatec:l ... under anaerobio CODd1tlona. The *'I"Pb010IT pre •• nta a 9V7 he~O\18
pioture, troa roda wbiob ...... sva1gb" to tIloae lIhiob were ap1l"ocbete-11ke.
V1br10 tons 8a well .. ooccoid bodiea ... _n. The gr&JI reacrUOI'l ...
wriable. The eultUI'M soon after iaolatlon be __ laoultatlft .... robe ••
Mot111 V .. 118 ...... en in arv ot the toru and naplla could not De
cl-..onstrated. Agar alant growth va. thin, wh1tl8b, nat, glistening and
undulate. Gelat1ll .. no" It.-tied. bouillon beoa. alight.l7 turb:Ld, ...
svaina produced a little prec1pitate and rina. No growth Oft pou.t.o. H11k
.. uncl1a.rlpdJ OM svain i 8 .... a posit!ft hfdroaen sulfide teat .ue the
l'8at van _Cat1ft, indol, urea, n1vate, -tbJ'1 red and Voga-PJooskauar teste
,ware necaUve. Heaol181a ot red blood oella did not occur. Olucoee, suoroae
and laow .. gave no acid or gas to..-t1ora. The 878teut10 position of the
OJ"@ani_ .. considered t.o be very 1ndttt1n1te.
Adauon (1949) reported he bad 1.ol.a\ed Aloal • faeoalia at
S7 _or0p87 f'r'o,a "'a cl1n1c&l. case of ~1d fe..,.. The organiB .. pre.ent in
... abdoadnall1Jlph nod .. , spl .. n, vaohM, bl"onoh1, b11w, ax1l1U"J" and
1agU1n&1 l.ylaph nod... Salaonella !irJ!!!~ .. BOt loud in any part. exam:1Decl.
(Tile orpn1aa .... not. descr1bed.)
Sever1 (1949) uolated an. orgard.aa ...... 11n& Vibr10 al!!Men:ea
".. a thigh phlegmon. It. .. a 11'. nept.1,.. rod, "17 aot.1"17 .,me,
.11«htb' ~ with pointed enda, and polar napu.. arotb crcwth Vb
hoIIOpmtows with a peUiole and lediMnt.. The orienta. we ... obl0' .latin
not 11qut1tledJ no beaolya1a an blood aPl" .. noted. C~te brot.be
beO&'Qe al.kal1ne, except a17oe1"1ne, slue .. , laevul._ and duloltolvtd.oh
raa1nad unchanpd. Nitrat ...... ndu.oed to n1trlteeJ litama tdlk be ...
alkaline and reduced. fIy'dropn eul.t1da .. produced. The autbol" tbink. a
e1a •• U'1cat,lon of 'the ahow baOteriUII i. not. :ret poe.1ble and thenf ... 1t 1.
OOI'l1'8llient, to _1nta1n the tma V:1bri~ aloal1(!ne. to 1M1 .. te Al!!Menae
l1ke bacteria which are cbal"act.eriaed by V1br1o aorpIloloa and are no' ....... an
identical. with AI.£!!!!O_rlua.
Blt_ and 'W1Uiau (1.91.9) nparted \he 18olat1oo of Hveral 8Walna
of Alcaly .... trca the hind gut contents of Pel"!£l.aneta u.ericana, t.be
'-rlcan eookI"oaeh. OrI~n1.sas, h; en did not fVllluJt dextroes, laotols or
8UCro.8 were plaeed in the genua ~loal!l!_.,. (The_ OI"Ianiau .... INt
d •• onDed 110ft ful17.)
caz.r.ou !!: ale (191&9) pUbl1ahed & report abow1.ng tbat Aloal!lenea
,v.ina responded vvioual7 to .tn~ and aureaqc1n, with ... swaine
tenstti ... , otbera resistant. Their .... ults would 1D'ileata that aUl'eOiqC1n 18
eft.ott .... 1n Aloal?:i!M8 Ul"tnu"7 tract. 'llf<i~(ltion ••
SB simitonda and P'ruton (19~) cteacriDed an OI"CarUsa (sr) which
aU).iSed peptide. and was 1dent1tled .. Al!!Y.Enu. The OJ'Ianiaa .. a
.bOrt., JlOMOtUe, p". neg.ti .... rod. 1n 1l\l'U'1en:\ lWotb conta1n1ne alucoae,
If.G''Ose or sucro .. 1t did not oaue t.ba r __ 'lOB ot acid 01' auJ attar 1
." .uch t.Nt. IJo1ut.iona bad .. pH ot 8 or hicber. uta. aUk beu.
alkaline and gelatin waa liquefied. S1aaonds am m\oft (195'1) 1I1th aOM
.uau;eatlona 'by Einar fAit.on dNcr1bed the OI"IaraiM .... .... t\l117. Strain
., .. JIlOt1le, polar n.gallated, and predoa1nan't.l7 aoDOv1oboua. Tba
_tillty .. beet Ob8U'ftd 1n culture. 1ncNb&t.ed a' about 2QOC.J GUlt ....
poovn at. 3TOe. were pnctloally' nOJa)tUe. In ..,1t. of the alkaline reaction
1ft peptone oarbob1*at.e Mdi8, .. sTUtheUc Meli_ OOJIp08ed or UCII'p.n1o salta,
lnoludL'1i phosphate am _old\R tona, the baner1a utUised gluoo .. as ..
oarbon source. strain SF .. acre 10l1oaUT placed in t.bI pllwa ,..~.
rat.her than Aloal,1:I." a. auaeatett earlier.
W81nbv, (19$0) etudied rltu.1n ... quil"eanta of dwarf oolon.r j
twianta of Al~!.I!~' ~uoal1a. (The pbploloc1oal and aorpbQl.O&1oal
oba:racterlatiu of the OI'Can1s1 .. Dot publiahed.)
M1lu (19S0) found baowi_ atkal.!I!!!!! naponelble for red lea 1a
fl"oga. '!'he oraaniM lias $ poaa naptt.,., nonaoid-taet rod, tMbll' _t11e ntb
peri v1chO'u tlaplla. Horse blood a.e oolon1e. wre ,..Uow1ah and nonblllo
lyticJ it tuiled to 1 .... 1'1t Il vide nng~ of o.IJ'bob7dr.tea. Cttrate, Wol,
.. thyl red, VC; " .. ~~-Pro.k_uel", _tal ... , aaaoo1a and to'dropn 1Nlt1de teaw
.... neptt... Gelatin vu not l1qutltled; nitr •• _ .... not l"8duoed to
nitrites. 11tau ndlk be .. alkaline. the Gr,_llia .. not agluUMted to'
ant1aerua too authent.ic swaine of ~il\l. broncb1aeeticu.
s, wrinkle !l !l:. (1950) found Aloal1pnea to be one of the principal
. ..--- ot ba.eteria in paeteu:r1sed and ~teUl1sed _gg_ a..'ld eg, produ.c ....
Ely-an aDd i4u8tat .. (19S0) in £cypt leolaWd. tl'CD _ter1al8 obtained
cluJ'ins food pol.ord.ng, !!. fae~~ alDl¥f,H,!! 01\ l'1Uoo S. s. agar •
.,lirta (1950). wh11. in 0.11'0. EQ'I)t dw1ftc Werld War 11 reported
tile iDol_tion of !!!. ,!'!.~~ !!o81!ene'!, f'roa nta •• p\ .... in •• s ball ••
Cott.,. !!. !!:. (19S0) and GUW7 !1!!!. (19S0) NpOl"\ed studi •• n\h
obloruphen1oo1 ooncel"Jline !! Yit.ro rasutance and tnatMMi or UI'1DaJ.7 wact.
Wections b7 ~loal!J!n~.
X1rb1 at &1 .. (l'SO) reported \Jw 1801atioll ot S .vatu .t Baeten_ ................ . alkal.wnea in a st\ld7 ot neonatal d1anhh and YOII1 Una ..
Castellanos !l !!:. (19S0) reported the isolat,ion ot ~e&lHe~"
along with t ..... 1ch1a and Proteus tr_ .senteric ~ nodes of a ohUd who , 3 • f'
0 ... !!. ~ (1950) t80.lat.ed A1oal!l!!!! tecall~ froca Wut. ••
Muhin (19$0) studied gadro .... nter1t1a 1n a ch11dJWl t s boapltal 111
MIlbourne. .Aaong other .. p.ni ... B~""'l'P. aloal!enea taecalta 'WU laolat.ecl
Wice .rro. abnoral .~cl.. OM a.ain .. isolated 1ft uaoc1at1on with
Proteuslli:l'abUU am the other OM _S the o!dy nonlaeto .. -tenaenteJ' P"Mft' ,
in the fecal epectt.n.. Both stra1u were aaUwl¥ motUe on uo1a'1on.
Signorine (19S1) ude 6S .tool oult .... ot brest ted Wanta with
enteric dist.urbances. A Sh1:Jella .. ieolated once, .. S,!'lmonella t:.hne \1ae.
and '.oal~ alka1!J!!I!'! 12 t1id8.
Weinstein am W ••• anum (19Sl) pr ... nted ..... with inteoUou
atnlOnucleoei. ard 8aowr:1ua alctll1g$U8 bact8rai..a oom.pl1cated by tuo-
60 .,
8PiroObetal an~. The isolated or,ania Vb de.cribed as a fP"Ul _sat1".
rod ~fh ftlll(tf;uh IIOtUi'tg. It produced no beaolTad .• , no indol. litaus milk
beC3l84l alkali,., and & r1ark brown pia-nt .. produced on potato. nextroae
and otheJt 1 tated carbohydratn wn not tenwnted. Ucl1.ng a. an\1pn tM
isolated strain, the S.1"Wll titer or the patient rose as follow& 1()2
(1lldad,.'81on), la128 (10 days later), lt$12 (4 weJr. after onset of bact.ereada)..
LeU.on (1951) published photoJlicrographa ot ..... ral strains or
Alcall16M,8 ahowing the perltr1chou and lophotriohou tlapllaUon verT
clearlT·
~ !1 ~-;. (1952) ~ the :laolAUon of A1!S!l!ne! from
cl0ud7 .rebroap1nal nuid .ap1rat.ed froa a tbrM .,... old. cbUd at necro1'87-
The OJ'1w" .. a motU. gram Mgatift rod which raUed to t ... nt glucoae,
lact.oH or BUOl'08e. Hydrogen sultide produot1on and ltelat1a 11quetaetion
testa .... ra,a\1:".. This or,aniaa ... &llo iaolated traa tDa blood a.nd
colon. ~~el1a R!!!&s.ntAtr1ae II va_ also isolated fr<a ~ colon.
Jensen (1951) studied) N-~!ene. sVains, isolated freta
RrasUian soU, in ropeR to nitrification of ox1ae COIJIpOunds.
Eldering and Kendrick' (19S2) differentiaW !f!!op!lUua ~ua.~.
the parapertunia baeill us. and ~r~OfIlla ~no!E:!!2t1- b.r the charaoteristics
6ti'ftn in the table 'below. Since t.htt •• organ1Ba ..... aiatita in the genu
~U!1" _net ant1genioally related it .. I .... ted they be ~
wgetber in HII8 other ,enus.
Sbnwbury and Barson (19S2) 180lated troa a ho •• 8puTOV an
.. pnin. vblch ... d.YOid ot any ncobarolJ't10 actbi.ty undV the taat
oond1tiolUJ.
Sir Aldo CutellAni (personal •• Iunication 19S2) p.I"OpOIIed \be
fallowing tuOftOllio acheme tor the new ed1Uon of tb.el'tanual ot Troploal
Hedicine (1919) ftot ,-et publ1ahed.
OErJ5 ALCALIOii;NEB caetelluS and Chal .... , 1919.
Det1nit1oru i'bu'tbeae which do not t __ nt gluon or laotoa., and .....
characterised b7 their pnenJ. laok ot t.naentat,ive powr, producd.DI H1tber
acidi t7 nor aaa, am b7 actuall7 incna81ng the alkal1nl ty ot the aed1a.
MUle 18 act clotted, ard i8 nl'lCiend alkal1ne.
'l)pel A1!!Mi!nee fuoali. (Petruoblq, 1896)' eaendav1t Caatellani and
Chal.men, 1918.
Cl.8s1fioatlcxu The •• ftl"ioue orpni8u oan be d1tferentiated .e toUeMU
62
cutellani'. SeirW.- a!llanioua (1910) .s placed in the PJlWl V1briotbrix.
CQtellani .tatea tbat orpnieu belong1D& to th1a genua ..... ot\4m m1staken
tor lloal!rJ!! ••
tn.r1ch am Reedhu (19S:J> ... portecI the dUteJ"entatioD of Brw.lla J
~;}].ch1a.et:1cue and Alcal.!I .... taecal.1a by b:1ocbuliea1 and nutritional
_thoda. (The fiagellar morpho1og of the orpnisu at.udied .. DOt recorded
bT the author.. It can be seen t'roa the data pnnnted that. Ulrich aDd
leedhaa atraina 212, 21&9, 282, 4141 and 8149 00IT88pond .... pect.1wl¥ ~ BH
'''aina 1)7, 430, 431, )77 and 175. The .. atra1ne can not be coaeidenct ..
\J'pical A1oal!t!_. t • ...,a1ia tor OM or JIIOl"e of the foUow1n& ..... ona, 1)
production of acddity !rca oarboh7dJ'ate., 2) polar ault.itr1choua tlapllat.ion,
3) n01'DO'111tq_ )
AdaMta (lS89) isolated, trca V1enu brook aWl', a capsulated
orpn18a which turned &Uk utav'. Tbe oapnle 1ureaeecI 1ft sue when the
OI'IanlMa va8 &l"0WIl in .Uk. It waa a alight1y lIIOt.Ue I"Od OCOUl"l':in& 81na17
and occu1onall.T in t.woe. The colCl'l7 on. 81111" aedia appeU'ed el.1a7. HUk
oultur •• allo be __ .11Jq, ropy aDd sl1iht.l7 alkaUne without. odor or
pl'8Cipi tat,ioft ot cuein. SliM was produced ill IM'Ptorae Ial." .W. The
cgogan1_ VN urobl0 and .s nam.ed BaeUl. laot1a rucoswp. (!'beN 1& no
record ot s1M, P''' reaotion, fiapllatlon or action on ~atea.)
Ad_ta (1891) ".',ed t.h:1a .... orpn1a in !IIUk trca Sv1'-rlaDd.
The organia waa traoed to weU vatar. It .. _'11e, without. spores, did
not 11quetr gelatin stab but cauaed lI1lk to beocM I'OPY. sucroee and. lac"a
were not. .ttaoked and no ps waa produced in .:Uk. lfAnball (l.896) in & paper Oft l"opilw.. in .... and aUlc: duOl"1Md
1ft orpn1aa ..,. tbtt bead1na "The JiJac1UWI C.-1nc tbe n-ouble.- (1'0 DaM
.. giYel1..) The orpn1_ 111&. a short, thick, nontItOtU. bacillus 2 b7 l.as IIlerona. No.,... WI'8 detected (aru reacn:ioD .. not recorded). QelatiD
•• not liquefied. The culture on qar slant ... vhl't18b, al1lq and row. Potato arowth appeared JellowUh white. Bouillon at.ruug out when the AMdla
lias applied. Ao1dl ty ot a1lk dec.naaed slich'UT but no QdOl" .. d,1acem1ble.
AbUi t7 to produce rop1neu .. loat when clll tllNd on laboratory I8tiUlll.
6)
(A~t.).
64 'luAe (1696) luted tbi. orpn1_ as Bao1llue 'riSOoa118 lact1e
Cbeat.,. (1897), in a clanitica\1oa of the 8chisOIqcetel,
~1.ed DaoMriuaa 1111008_ 1ao1.18 (Adaau> .. nomaotUe, capsulated,
aapGrOpDOU8 am i1'U posi".... The organia _a not 1ttrietJ.¥ aerobic and
did not liquetJ platina M1l.k HOQa v1acous, pepton1aaUon and clearing
0C0\DTed with no odor. Apr colonies.,.. d1r\7 whit.e &ad alUV-. Habitat
aUk. (~ate reaotlon wa. not recorded.)
LebaM and Jetaann (189') oIw'aot.eriHd Baotvll11 laoUs 'I1aooaua
(Adneu) .. !lOl8OtUe, p"u positive, ClI~u1a't4d.Orape".q&r ~ 11111(
sugar were not te,..nted. 11t\la indol and no b7drogen sulfide we pnduoed.
The authors atl.te that thea eu3 .... , obtatnld trOll bal, no1th1n& ... to be
••• n .of the spore to.r'JUltlon which 2:1anenunn ala1lled to have ... n. 1'bta
organi_ .. diaOOftred bj" Adat.ts .. an 1Btportant eDllDG" of the butter
industry. f.~1".i1IAml found thi. or,an1aa 10 vatar.
Conn (1899) in h1a CllN.U1oat1on ot dair;y bacteria dsacr1betJ
!! ria~ •• ~!!!¢. II (n. ep.). It .. a rod tour t!au u lol'li q broad
(aiR not. reponed), .urroWlded b7 a .... olel... It did not 1ique1)'
gelat.1n. A, ... artnrtb was al1a,yJ potato s;rowtll .. P'¢ab brown. M1lJc
be.- .lilq and allcal1ne vi th q odor rauind1l1l one of .trona c.t.ue. Th1e
baoUlWl, tba author .tated was pb)teiologioal.l.7' aiudlal" to !:. via!!!¥. lagMf
ot AdMe1.., but. ita aorpholou vas quite dttt.-ct. Wh1l. 1:. ... v1..,._008U .......... waa
near17 .. broad a8 long, th18 OJ"laniM" 10na am slender. (there .. no
lIeftt10n of 11'_ ataln, JIlOtUlty. tlqeUat,1<m .. CU'bob;vdrate .... aoUon.)
WU'd (1899) reported on rop1ne.. :1D aUk am ere... He vaa of the
65 opinion. t.b.at \he orcani8a dea01"ibM b7 _ahal1, vh10h was responaible for
tbtt. row a1lk ou.tbreak 1n Micb1can, 001Ild *" be poaltiwl1' ident1tied trca
the brief de.cripUon available, but then ... n&8Oft to suspect it .. iclenti.
with SacUlu lap'1,. "IiItcoau. He ct..crib" an 0I"pa1aa widell vu uolated
Ire. __ r and aUk aDd probab17 1dct.icral with aa0!M!! l,f.Ot1a visco ••
Adalleta (1889). fbi. ratbel' CCltPlet deaonption _taw ta. orCani- to be
V" poslUft_
M1pla (1900) u.ed the DaM i:l~~_ &~!2e! t. the organiul
BaatH- lao'll. ~co._ A~., but WIed tbtI d •• or1pt1on J"eCIOI"ded b7
Zi_r_ma. The ... _ po.iti .... , noraotile crew_ •• de.cribecl .. tol'aiDl
sperea which had not been obMned to pra1nate. gelatin .. 1\0\ l1ql18t1ect.
Xipla aho deaer1bed Bacteri_ 'If1aO"_ (rr .... ) 1141&- It. ... Hd 1 _
2.S to h adcrou, nonaotUe. Oelatin vaa liquefied ad IltJq P'~ ooo1lJftd
on CU"J'Ot alicea.
Conn at al. (1906) pointed out that t.be OZ'Can1au produc1n& alJ.iQ" --aUk "do not rona a cl ••• tv'thaaelft., lor thia ~ of rendering II1lk
au.;, __ to be loWld v1de17 ._t.\eNd aaoI'&I t,ta bacwl'1alt •
8uobanarl and IiaIIIIar (191S) in their .\\117 of 81.,. and 1'01>7 aUk
lie' 8.o~rlUJ ... 1&..,0...,.\18 ... v1aeoama (Adaeta) I.e-'- am. :NeUilllDft .. IJ.JTlOfJI'lIOUI
with.
Thea. autbon con.ider the orpn1ea to be a a:na negati .. , nonaotU.,
a.porocenoue rod. The orpn1.M did not pr'oduce acid or iU trom. ...
66
carbohydrate.'" They propoe. the DI/I.M BaG ... 1. yuco-cocc:01d1_ ep. DOV.,
aM g1.,. it ... ~us w1th Group IV, 'ftI"1eti •• ~ and 10 or 1ian'1aont •
,11. tdlk bacteria, tor an organl .. wb10h produoad acid trca dextroae,
laow. and Ml to .. and lor~ sH.- in allk. (Baowz.l_ YillJco-ooocoidiUlJ1 18
still retained in Bergft7'_ Manual of Detend.Datift 8_ter101017, 6th edition
(1948) as a e1ftOJVJl. tor AloalteM! 'rieoo_ .... )
AeOOl"ding to Mapluaaon (191S), a oultUN 01 • aupr fermenting
Ol"pn1_ naaed. a.eWu 140\18 Yi.eoau Amu.t., and 68awed to be aaoter1wa ,
froa tiMt Kral collection.
Weldin and r.eYine (1923) nauaed the orcan1a Bactvl-. ~OO8" arad
plaoed 1t 1n the 8dlgem:&8 Alcal!l!!m!. It. •• a _paulated, UporOlClBO_,
gr_ n.,ati .... rod, nonm.otUe or IIiOtUe by perlt.r1cbou tlaplla. We acid 01"
gu ... produced troa gluco... ~Uk was "ndend slhq' and ptat1D wae QO\
liquefied.
ner •• yt. Manual of Detena1natlw Sao\eriolol7 (192) used tbe __
_ La .... o .... tob ........ a_~_,i .. ll_ua_r rl.ooe~ (Ad_ta) with probable qrl0!VJl81 BacUlue ruoHU
1&ot1l, Conn; Bacillus rl.OOSUlll 1act~ II. S,o_iwa l.oUl aoid1, ~J
BacwrlW1 vUoo-coCQfl1d1_. ~uchanan aM Haaaer. It is de_cribed .a baY1n&
no action on carbob,ydrate8. nO!'ROtUe and __ .. ,a'1...881"P7 aleo WIld thIt
1laJ.I8 A1eH:I1nea Mtalcal1J1nea (A.dMlts) aI¥!l c1tea .ldaat,ta work of 1891.
It ... da.or1bed .. a V" po81t1ve, indo.! produo1n& orpniea.
BerP7t 8 iiamaal ot Datera1nat1. 8&:o1Ar1ol.oer. 2nd edition (192$)
carrie. the nM8 ot tlWl oraaniaa as AobI'oJIo'baoter '9'1a... (Aduek) vi t.b k 1
the .... duCU'iption whicb appeand 1D the prerioue ed1 t10n.
67
Weldin (1927) 1nolud88 Alcal!l!n.es visC08\1Dl (aduetz) oomb. nov. in
!de taxonmq· It vas synonymous with.
Ba.oter1ua viacosUlll (Ad_ta) Weldin and Levine, 192) Baotr1ua lac!!. ViSCOSU8 Adaaeta, 1889 aacn:lu~ ;racosis lictis (AdU19tS) Krause, in F1ugge, 1896 £tieterlua v1scosu.t Iictla (Adaaets) Ctt.e.ter, 1897 B&cterlua LiciLJ n.co8UII (Ad_te) Leblaann and Neumann 1901 tictobaorlIus Y1s008u8 (Adueta) Berge,. et al. 192) I&CiillOGaotei' Visco.. (Adame',) Bar..,. it at. 1925 --
Weldin statea that Ad,,"'a considered tili. organ1 •• to be 11'- negati"1fe, but
ac" resistant to deco102'iHtion tban related speoie ••
Lehrumn and tfeuurm (1927) continued to list the oraaniaa .s
Baoteria lacti. viaooeum (AdaMts). - . -.----~ tong and RPMI' (193)) reported the isolation ot tat-splitting
[bacteria in not"fHl. am abnormal (but not rom) milk.. -ere .. and other similar
products. Th~ lix ot the.. culture. showed the charaoteristic. ot
!Ucal1.gene. v18008., iDClud1ng rop1ne8a in JIlilk. Carbohy-drates in bouUlon
rwere attacked only slightly 1.t' at all. A few of the oultures produced !loid,
[but no gae, trom certain ot the sugars but later rewrsed t,M reaction. They
~e aerobic, noraotU., asporogenous rods, 0.6 to 1 b:r 0.8 to 2.6 microns.
They ~ e8.ential.ly gr .. negative but in certain cultures there wer~ a tew
cella showina a tendenay to retain the grata stain. Capsules vere produced.
. in milk cultures. Agar growth was white. aelatin liaS not li'luefied. Litmus
~ilk became ropy, alkaline, without coagulation or proteol:rsis. The following
reacti0n8 were negatlft: indol, nitrate reduction, lVdroge!l sulfide
production, naeth:rl red. and VOg8.-Proskauer. 'lhve were 2S cultures which were
.. ike Alcal:!,.genes viscesus, 8XCflpt tor the failure to produce repiness aM were
censidered to be nonropy .trains of Aloalipnes 'ViIC08US. The,. were designated
1l1~1Rene8 Yi8COSUli var. d1aa1lll1lie. ,......
68
. Druce and Aynsl.,. (1949) aade .~ observations on row milk in
lLanCaswrshire, Ellf.land• Cultures isolated war. tound to correspond in IUllT
!rdpect.s to ~4;l;enes vi8cosu.a (SerpY' 1948). This organi_ dilfers lrOll
~tl'ge,.'. d.8cription 1n the type ot growth on broth and apr.
Abd ... 1.....al..k (1952) round that. Alcal;a ... s !iaco.us showed lipolytio
~t1v1ty. This conf1nled the :report b1 Long and HallMr (19)S). Sam. ot the
.trains studied '1:>7 Abd...a""'flUllek reduced nit.1"ate. Citrate oxidation aM sliMe
~uction did not appear to De 1dentif11ng characteri81iic8 of Alca1~ne8
rnsCOSU8.
Castellani and Chal.ftlers (1919) listed an organism in their taxonomic
,cheme which was differentiated from Alcal1pnea f.ecalis (Petru8chky 1896) by
ita nonmotility. The organism. was Ihrted 4S Alca.lieJl8s metalkal!ienes from
the organism named l!:. ~ a1kalijen&! described by Castell~n:l. This organism.
was described 8.S being nonmotile, gr_ negative, gelatin and coagulated serUlfl
not liquefied. Alkaline reactions were observed in glucose, maltose, lactose
and sucrose. Indol was variable and broth showed a general turbidiV.
Bergey's Manual of netermiMtive Bacteriology (1923) lists this
organiSM as follows: Alca1Ysines .IIetalca11cines (Ad8JlUtts). SynolVJlOus with
.BeUlus :netalca1il1Ms and Bacterium lactis viscosum. Adaetz (1691). Rods 0.5
!bY 1.0 to 1., microns, occurring singly and in pairs, nonaotile, gru positive,
itncapsulated. Growth on agar slant mOist, 8JIlooth, spreading, sl1rqJ broth
~eooill.es turbid, with sliIq sediment} litmus srllkbecol48s thick, u11lq, but
~t coagulated. Potato growth 18 gra,1, ao1st, spreading, sl1lQ'i ind01 is
~orll1edJ acid in dextrose. Not pathogenic. Aerobic. Found in slimy !aUk.
wtlldin and LeTine (1923) in their key to the .peelss and varieties
of intest.inal bacteria list this organtsl'll 4' Bacteria _talcal1pne~ in the
~ubgtlnU8 Jq.cali,enes. This organism was stated to be nOfL1lOtile and not
!Liquefying gelatin, not fOl'llling capsule. in milk nor turning it sl:lmy.
Bergey's Manual of Daterainative Bacteriology, 2nd edition (1925)
70
lilted the organism. as Achromobacter metalcal1§enes.
LaTine and Soppeland (1926) isolated Bacterium me tal cal igenes
castellani and Chalmers, from creamery wastes, and which had the tollowing
characteristics) gram negatiT., asporogenous, nonmotile, aerobic rods. HeaT,1
pellicle in broth, no liquetaction ot gelatin or Loetfier r S Berum, milk becomes
alkaline without coagulation or peptonization, lit.us reduced in 10 days,
nitrate reduction variable, no gas. Hydrogen sulfide and indol was not
produced. No acid or gas trom glucose, lactose, sucrose or glyoerol, peptone
Weldin (1927), in his taxonomic scheme listed this organism as
gCaligenes metalca11gene. Castellani and ehal,uers (1919).
Bergey's Manual ot Detendnative Bacteriology, ,3rd edition (1930)
removed the Adametz reterence frCll'l t he description of the organi8ll for it
appeared as Alcaligenes metalcal!Jenes castellani and Chal.Ilers, (19l9). (There
i8 no reference as to the source of the description.)
Bergey's Manual or Determinative Bacteriology, 6th edition (1948)
describes Alcaligenes metalcaliJenes Caatellani and Chal.at.ers as follows:
Rods: 0.6 by 1.5 microns, with rounded ends, occurring singly and in pairs. Nonmotile. Ore nagatiTe. Gelatin stabl No liquefaction. Agar colonies; Circular, raised, Mooth, .. nwrpnous, entire, gray. ~roth: Membranous pellicle with heavy sediment. Litmus Dlillu Alkaline. Potato: Scanty, glistening, smooth, sometimes taint pink. Indol not formed. Nitrite production from nitrates variable. Starch not hydrolyzed. ~lood serum not liquefied. No action on carbohydrates. ~erobic, facultative. Optimum temperature 22°C. ~bitatl Intestinal canal.
71 Allen and Brooks (1949) reported they reco'Nred Bacterium
Iietalkalig'=1n8e (Levin.e and Soppeland. 1926) tram chlorine treated 8t1wage - Elliker !!!!h (19~2) nported Alcaligenes metalcaligenes produces
• laarked gelatinous rilm around the curds ot cottage cheese at low teMperature.
(The organi&11 was not described.)
Booker (1887) was the firat to describe an organism which has sinoe
jbeen included 1."'1 the genus Al:al4iienes. He reported too isolu t10n of BacUla
~ troa alvine discharge of ehlldren std'£erin(; 'With cbolora infantum. '1'he
IOrganis,$} ~ actively 5lotile in baniing drop, m.Uk became coagulated, alkaline
land peptonized, it liquetied gelatin a.."ld gr$W on pota'to \o11th a brown color.
(TheJ'8 wa. no mention of oarbohy'drate reactions, ar- stain, or aagellar
p.o:rphology'. ) l)ooker (189('» recogn12ed tha. t Bacillus A 11Ul1' have been Proteus
IvuJ,garis. sternbe.rgts (189) Manual of Bacteriology included Sooker's
~.soription ot Bacillus A.
FON (190» isolated and .tudied an organism, troM tl':e stomach of a
Itoundlinf,:, which he oonsidered to be the sue as BaeUlus A and. l'WI'led it
a.eUlu. bookeri, Ford. lie added the following charaeterht1c. to Booker t •
~.8cript1on. Rods with no spores, no aoid or ga.e produced from dextrose,
p.lI.ctose or 8110x'ose. In 8 £illith tube oontai.nL"'lg dextrose broth growth WAS
",im.1ted to the open bulb. (No gram reaotion or nagellar morphology vas
I'-neluded.) Indol was not produoed. Abundant yellow or yellow-brown growth
~ppeared on <!:gar slant. Litmus lllilk beC3llle alkaline and reduced but no
~oabula.tion occurred. (It should be noted that tile milk reaction and pigmenta
~ion in this report were variOUdly given.) Nitrates were not reduced to
~itr·ite8. Oelatin, casein aM coagulated blood serum. "Were lique:tied.
12
73
LevL"ie an~l 0o,ppsl':lnd (1926) when studyine btlcteria of creamery wastes
,dded characteristics to thn organ.18m. they called ~!leterium booker! (Ford).
This organism iII-aS gram. rtegst,ivfl, potato (l"o'Wth 'Wu abundant in 48 hour. but
with pale oltTs color; sli$rht t"duetion of nitrates to 1'11 tr1t"s with no ¥as
production ooserYe1. 'l~e na.'Ile BllotfJrium. ~-8!! 'ftr. 11J1!lI.()bilu~, nov. yar., was
coined for the. nortllct11e variant.
Bergey', Manual of D~rm1native Bacteriology (192) li.te~ this
org·tmism IUS I.lcaltprntl bOOkeri (Booker), motU. by means of per1trichou.
naplla, gram negative and nitra.te. not reduced.
Weldin'" (1921) tEXonomic seheme li.ts this orge.n1u as ~lcal!i~n168
tbookeri (Ford) Bergey!!!h.. (1923).
Kutscher (19)1) isolated from spinal fluid an organisn ~iob be
!called Alca1i£sr.es bookeri. It WAS .. small, lI10tUe FM negatl'ft rod. Growth
jon plain &car was abumantJ gl"O',,:th on potato vas abundant and brown; f,elatin
!was liquefied. litm'll8 Yd.l.k: was reduced and alkAline in 24 hours with clearing
~n the upper portion. No fermentation ot ~ carbohydrates including ¥luco.e,
~nnitol am xy-lose, which becfU'!1e alkaline. Indol, _thyl red and V'ogea
Proskauer testa were ner,at1ve and nitrates were not reduced.
Sergey's Manual ot Deterainat198 B&cterlo1og1, 6th edition (19hB)
~ont1rmes to desoribe Alca.lieMa bookeri as being motile w1.th per1.trichous
naP-811ft. Nt tratetc were not reduced to nitrite ••
Alcali;lenes reot1
Ford (1903) in a report on the claaa1.tioatioll and d1at.ribut1on or
intestinal baoteria in lI8Il, described Bacillus recti, (n. sp.). This organ1aa
vas an actively lIIOtUe, upcrcgenoua rod 0.5 bl' 1.S to 2.0 lII1crona, gelatin
wu liquefied. Dutro •• , lactose and sucros. 'were not tel:'Mnt.ed. In a SJlith
tube 00l1ta1n1na dextrose broth growth 1IU l1m.ited to the open bulb. Nitrate.
_r8 reduced to nitrites, indol vas Act produced, litillua ndlk bee .. allcal1ne.
Bercel". Manual ot Deterrdnative BactericloQ', 6th edition (1948)
continue. to list tbi. org4nism a8 Alcalig.ne. recti (Ford) Berp7 !! ~ The description ,i,"l1 by Ford 1IU uaed with the addition ot two tundalumtal
oharacteristics, peritrlchoUi fi.gellation and gru react1on., which were not
recorded by Ford or .ubHquent vonan.
74
Marshall (l904) reported the isolation of .. peptonising baoteriwa
trom milk, but did not describe the organie:a.
Conn !l. !!:. (1906) published a claaa1tioaUon of.' dairy bacteria
and stated t.bey bad atudied the organism. which Marahall iaolatecl. It vas
a gram negatift asporogenous rod without capsule. (There is no mention of
tl&gellation or .IllOtllit}".) aelatin was liquefied) no acidiv, &&. or closed
ana growth in sBdth ferraentation tube. (carbohydrates tested were not listed).
Agar streak growth va8 "Iiacid, uootb, creQ-vb1te becoming leaon-reUow.
MUk became alilq, alkaline and dip.tad without coagulation. Potato growth
wu 1ellon-yellow. (There was no _ntion of indol or nitrate reactions.)
Tbe authors vere of the opinion that e.8ent.ial.l.7 the sue orlania vae found
in Connecticut am lev York exoept it ditfered in that aUk .. not d1geeted.
Buchanan and H.... (191$) in a report on .11ay and row milk luted
Bacterium marshalli Conn, with all the character1etics reported by Conn et ale --(1906)' l:n1t recorded there .. no motUity or acid produced troa atV' 8\11ara.
Serpy's Manual of Determinatift Bacteriology, 6th edition (1948)
recorded that Alca1~"8 marshalli Berpy !!.!!:.. waa nonraot1le, no acid from
any sugars, indol not formed, and nitratea not reduced.
7$
CHAPTER III
C(MMENTS ON m HlSTORt
It the genus Aleal!ielW8 is to be cbaraoterized as having
peritricboua nagella ..... ral specie. (booker1 and reoti) nov recognized
in the genus Uou!lenea in BergeT' a Manual of I)atena1na~ive :a.cteriology, 6th
edition (1948) should be deleted. Ne1 tb.er ~be or1;1nal description, nor &'If¥
lub.equent descriptions, or these oJ"gan1au haft included tba nature of the
nagellatton. Actuall7 the 11"17 motility reoorded tor bookeri am rect1 are
more sUIPatt ft of polar than peri tz-1choua flagellation.
Cul tUNS of p.cal!le!¥,~ r~ot1 and P.l.e!il!ene~ urebal.l;1 are not
tound in the Amer1can 'l'ype Culture Collection, the National Coll!«tnt1on or Type Cultures, nor could they be obta1nad 1'1"0iIl private collections. Two
cultures labeled A~caliene,s b?~~ri wre stud1ed (RH 21 and 229) and both
were found to 'be polU' fiageUated.
COnYinc1ng .'Vidence hu not been otfered wh1ch ind1cates ti'.at
Alcal!&en.es lIUlZ'Jlhall1 should be MMOtUe. Buchanan and daswer (1915) state
thatB _a_c_t ..... r_1_um_,,~11~1 is nOl'lllOtUe, howe".,. the description given was
that of Conn et al. (1906), to which va. added a atate_nt regarding non---aotUity. It 18 augseated that Alcal.!I!!!!~ aarahall1 no longer be recognized,
linee it hal not been adequatly descr1bed. It 1. suggested that the n&ale8
Aleu1S.,MS bookeri, %'ect1 and I1I8l"shalli becoae known as nOl111na dub1a .inee
76
77
t,beir application is uncertain.
castellani and Cbal.-rs (1919) d .. cribed !lealieenee utalkalisent?8
and stated the na_ lUle taken t'roa h !!!:! alkal!l!I8s which appeared in a
paper by C.st.ellani (1915). The orilirw.l .... terence bas not been found.
Doctor Robert Breed (personal correspondence 19S2) .... likewi.. unable to
locate the original publication. Sir.Alclo Castellani (penonal correspondence
19S2) w ... uuble \0 gift the UA-et docuauntation since hie reprints were
lost during the war, how_I" be thought the OI"ganiaa was tiret ant10ned
and its characteristics giwn in a lecture be delivered at Taranto, Italy,
in 1915. H4t tbought the leoture and/or a report ot the orpniaa waa published
111 Hwral Italian medical journals.
The description by Adamets ot aaeUlus lact1s visCOSH18 does not .. ,lUI
include the gJ:"8JI ruct,1on. S ..... ral t.axonoai.ta such as Chester (1897),
Lll'iwln and leuma.nn (1899), and Migula (1900) considered this OJ'I&n1aa to be
p'1UIl positive. ward'. (1899) exper1ena. 1nd1oated the organiall to be srpoliU.,.. 8I'ld recogn1Hd. the earlier d.soription of 13.elllu. laot18 vi.COS118
I
Adamets was incomplete tor precise re1dentit1cation. Buchanan and ~
(1915) ""' ot the opinion that lIl8I1T of the oraaniau described earlier,
suoh .s BacUlu. laott. viscosus, AdUlet., Bacillus vUcosua l&otis. Conn, atxt , ".. 1 • I
BaoUlua 'riaooeue laet.ia II, Conn, though 1acldn& at. atatement of t.he "'_
reaotion in the or1&inal description, wre gJ'Ul negattve. Generaliaationa aa
the .. can only bte aooepted, at beat, with considerable reservation.
Kore recently Long ard. Halalaer (193S) st.udied ul\Y orpniaaa capable
of producing rapinea" 1n milk. AltboUlh t.hese orsan1au ware INa neg&tift
and considered to be Alcaligenes 'V1SCOSUB, the cOIIUI8nts ooncerning their
78
eftect on dextrose bouillon indicate. they were capable of' oxidizing
carbOhydrates. All this can onlJ' _ean that h1a strains included in
Aloaliene:s as viacosus WU'e tar fr(lfll beiug boMOB;ttn~ous. -AdaMta (1891) who i. general.lJ' oredited with the original
chaJ"&Cterization of A1oal!&eMI!l V!seosu8 found it to be .lightly tIOtUe.
workers since bay. conaiatantl:r reported this en"gani .. a8 b4;ing norwotlla.
,lbd-e1""11Ullek lugseata that tbia indicatea that Adameta wae in eft'OZ". Berg.,..' 8
Manual ot T»tendnatl'ft Bacteriology (1948) continues to refer to this report.
concerning aaotU1t7.
As notMd in the review of the literature, sewral spellings flJ"fJ
ued to elenote the genus Alkaf!l!M$. The spelling :!~c.lig!ne. i8 p.robably
correot.. The GeJ'.maxla in transliterating f'rCla the Oree¥. !reqlWntl.7 use a "kll ,
.. in kokkul, but 1n taxonoodc work all rut_san 14tinize-i aM .henld be
transliterated as • cit • Fort.unately Petrwschky' in hiB f'irst proposal of the
I'lUIIB ot the specie. HS;u-ded ae the type, used the spelling B4ciflu8 f'ae(.t,alia
alealij •• a. Castellani L"ld Cb.al.aera (1919) ued both s,Pell1ngs
irxiiscr1m1nately.
The probl_ of .,bether or not bronch1oanis Ferry (19ll) should be .. substituted forbr __ o_D_o_h_111_ . ..!.. .... nt.ioti '8'1!f'y' (1912) 18 rJPt 81llPle. It raises the
question or whether t.lla author of' an unfort.unate ll8l:De tJtis the right to correct
hb error. Under tbe older Botan1c~1 Rules, usage would be followed rathar
than prlor1t.1'. Doctor Breed, editor 0:£ Bergezr's l:4anual of f.:et.endnative
Bacteriolou, haa preterred to use the al.1lOat un! vel'sal.ly aceeptsct narae to
avoid changes in llOJIl&nclature. This appears to be good judgem.ent while wait
ing for the tonrulation of a 'POlicy aocept.'\ol'" ton the International Judicial
79
C~i88ion on Bacteriological Nomenclature. Be this as it may, Chapter III,
• seotion 1, Article 16 of' the International. Rul .. of Botanical Nomenclature,
.s accepted by' the Fit-at International Microbiological Congress in Pari.
19)0, reads as follows: An organiam Moan bear onl¥ one valid l'I&IIle, the
earliest that is in accordance with the Rule. of tfoaaenclatu:re". Chapter III,
section 12, Article 59 read. aa tollow: -A !'lUll or epithet must not be
rejected, changed or modified merely' 'becauae it 1e badly ohoaen, or
disagreeable, or becaulM another 18 preferable or bet.ter known."
Bued on the aboft historioal study' the author would recOlaIInd tha:t:
1. The designatlona Alcalienes utalcal.!lenes, bookeri, recti and marshalll
be rejected since their application 18 uncertain (nomina du.bia). I
2. The designation Alca1t&!ntts viacosus be l1a1ted to graa negatlft rods
which do not cx1di •• or ferant oarbo~tea.
J. The epecies d.s1rnation broncbiseptioa be rejected tor the epitbot
bronchioan1a, Ferry (1911), which haa priorit,...
4. The spelling Alcalti!n8s beCOltB universally accepted .. the genus de.iana-
tion.
CHAPTER IV
MATERIALS AND l.m.THOtS Bacterial Cul. tures
Two sets of stock cultures were maintained throughout the study.
one .et wa. kept at 6°c while the second set, with rubber stoppers, was kept
at room temperature. 'I'he t'ollowing semisolid llled1ua was used,
beet extract tryptone pH 6.9
The meciiUlll was dispensed in )-4 ml. Mlounta in l)XlOO M. tubes and
autoclaved at 121°C t'or 1$ m.1nutes. The swot cultures were transf'erred
every 3-4 :montha, incubated at )Ooe UlltU good growth appeared and then
replaced in storace.
The tu.porature ot' incubation ot the inoculated media vas at 300e.
Oeneral17 there wu better growth at ~C than at :37°e for the majority of
the organis.. studied.
Morphology
ORA:i STAIN
I. Amm.onium Oxalate Crystal Violet (Hucker")
Solution A Solution B ~/stal violet (90% dye content) 2 g. 8lSI1oniwa oxalate 0.8 g.
distilled water 80 ml et~l alcohol (9S%) 20 al.
Mix solutions A and B.
80
-Iodine Solution (Kopalott and Beerman'l)
iodine normal NaOH (!to g. per liter)
2 g. 10 ti.
81
Atter iodine 18 dislolved.. make up to 100 Ill. wit.h d1atUled water.
III. Deco1oriaer
Equal volumes of 95% e~l alcohol and acetone.
IV. Counterstain (.8 urke f 8)
salranin 0 (as!' dye content) distilled water
2 g. 1001'41.
1. fJry th1nl)r spread t1lu in air without heat. 2. Flood with Ol7stal violet. Allow to stand 1 minute. J. 'Waah in tap vater. Shake to HiIIlOW "oesa water. 4. Cover with iodine for 2 ainutel. ,. Walh in tap water. Shake to re.ove excess vater. no not blot or allow
to dry. 6. Add decolori"8r to the slide drop by drop untU the decolorizer £lows
color18.1 from. the slide. 7. Wash in tap water. Shake to 1'_0," excess water. S. Countel'8tain. 9. Wash in tap water and dr;r.
Smears tor gram staining were m.ade trOll. 18-24 hour agar slant growth ..
ruCrELU STAll
The technique tor studying nagella was outlined by I.eif'aon (19$1).
It is believed the taxonomic cri tenon of significance should bca the tqpe
ot flagellation or aOlence ot flagella rather than motili~ or nonmotilitT
since nonmotile flagellated organi8lU have been reported.
Pb7ai01ogy
CARBOHYDNATE tT'1'ILlZATIO]i -pancre8.tic digest of casein agar earbohydra te
(Bacto-Casitone) O.s% 0.2
bromthymol blue (1.6% aqueous solution) tinal pH 1.1
1.0 0.2 m1.
82
Ten percent earbo~drate was Seits filtered and added aseptioally to
the 1181 ted baae which had been autoclaved at 1210 C for 1$ minutes. The
medium was then clispenBed aseptioal.l.7 in 4 ill. IUlOUnts into sterUe 131100 tam.
tubes. All ~1a prepared in this manner were tested tor sterUit)" b7 allow
ing them to /IItand owmight at room temperature. Eaoh organiaa was inoculated
br stabbing into duplicate tubes ot the 'base m.ediwa without carboh1drate and
duplicate tubas ot the carbo~rate mediua. One ot eaoh pair ot duplicate
tubes was la,..red with sterile lUlted petr01atlll to a depth of 2 011. This
was found to be the most efrioientpract1cal method of' excluding air. 'l'he
tubes were checked for changes or indicator e"17 24 hours tor 1 df'~s.
pancreatiC d.lgest or oasein (Bacto-casitone) 1.($ pH 1.1
The broth va81.i8pensa~ in 1,31100 M. tubes, 4 nil. per tube, and
autoelaved a,t 1210 C'tor 15 minutes. 'the medium was inoculated in duplicate
and tested tor indol production atter 24 and 48 hours. ~od1ti.d Kovacs'
reagent was used to test for indol production.
paradift.t~laminob.~~ald.brde 18oa~1 alcohol (B8Ker and Adamson 1210) h1drochloric acid, concentrated
S g. 15 Ill. 2S ml.
Aerobacter aeroenes and Escherichia ~ wre used al control organiama. -liE'l'BYL ,m~r: AN):: VOCES .. PROi::,Y,AUER TE:.l1S -
Methyl red teat: Four ml. ot broth was dispensed in 1)1100 mm.
tubes and autoclaved at 121°C tor 1, minutes. Atter inoculation and S darB
incubation the reaction or the !lle'!ium wa8 tested with 5 drops or nlethyl red
solution (0.1 g. methy'l red in 300 rule 95% ethyl alcohol and diluted to 500
ml. 'With distilled water).
VOg8s-Proskauer test; One ;Ill. 0: the broth was dispensed in 13);~OO
lWll. tubes and autoclaved at l2loC for 15 minutes. Atter inoculation and 2
daTs incubation, 0.6 nU.. ot ,% alpha naphthol in absolute ethyl alcohol and
0.2 ml. of saturated sodium hydroxide WrG added to determine the presence
ot acetylMthylcarbinol. Aerobacter aerog.;u»s and ~.8cher:tohia ~ were
used as control organisms tor th~~e test ••
CITRATE
Bacto Shimon. Citrate Agar va. uaed.
Three ml. of the I'48diUtfl vas disperu,,~d in l)llOO WI. tubes and
autoclaved at 12loe for 15 minutes. The tubes were slanted, lightly'
inoculated and indicator cr.ange recorded a£ter 24 a.nd 48 hours. Escherichia
.202!!. and AerobacteT aerogenes were USl€ld a8 control organisms.
UP.EA -Bacto Urea B;roth Concentrate was used.
This mediwa its prepared acoordi.'lg to the fomula of itwltigian and
84
stuart (1941) and stuart, van Stratum and Rustigian (194$). The conoentrat.
was brought to volume with distilled 'Water and aseptically dispensed in 2 ml.
allounts in steril. 13Xl00 Mm. tubes. Heavy inoculation was .tUilde from an
18-24 hour agar slant culture. Resulte were recorded atter 24 and 48 hours
incubation. Proteus Moreni and Escherichia ~ were uHd as control
organisiwh
GELATIN (Four media 1I8.N compared.)
Medium 1 gelatin pH 6.8
12.0%
the .-d1wa was dispensed in S m. quantities in 131100 M. tubes
and autoclaved at. 121°C tor 15 minutes. The mediUll wu stabbed, incubated
and tested tor liquetaotion each day, beginning with the second daT, tor 15
days by inwrt1ng the tube after cooling at aOe for one hour.
Medium. 2 beet extract pancreatic digeat ot casein (Bacto-cas1tone) platin pH 7.0
'!'hie medium was handled 8S gelatin medium 1 above.
Medi_ .3 yeaat extract. tz7ptone plat-in pH 6.9
0.,3% 0.5 12.0
O. '3 1.0
12.0
This JHdiWl was handled as gelatin Medita 1 above. The re.ul ta
recorded in Table II are based on this Melia.
Medi\ll1 4 Bactc stone Extract aelatin Agar
gelatin beet extraot
3.0% .0
agar pH 6.B
8S
The mediwa, descr1bed by Stone (19.33) was dispeneed 1n 2 al.
q,uantities in 131100 lIR. tubes and autoolavecl at. 121°C tor lS rdnute8. The
tubes were .lanted without butts. The _diu. •• inoculated by streaking and
incubated tor 48 hours. Saturated (rOOll wlIP4trature) 8IM&OniUll'l sul.!ate vas
then added to cover the slant. Results were reoordtld a8 positive U' the
medium remained clear atter exposure to the umsoni'Ula sultate. Proteu
wlgaris and Salaonella tmbos& wre used •• control organiau in thes. -gelatin teats.
HYDROGEN SOLP'InE
Baoto Kligler Iron Agar va. u •• d.
'Ihe medium was dispensed in 4 JIll. quantities in 1.3XlOO mm.. tube.
and autoc1aved at l2loC tor 1$ minutes. Tubes were slanted with a deep butt.
The lUdiWll vaa inoculated tr01Il a broth culture by 8tabbing and etnald.n,.
Proteus moraani, Saltaonella Whoa .. and Escherichia .22!!. 'tIJV1t used a8 control
organiUl8.
N1TRATE REDUCnON 10 NITRITE
[<fedi_ 1 yeut. extract. 0.,)% panoreati~ digest of casein (e4cto-Cuitone) O.S alar 1.S pH 7.0
The mediuaa vu dispensed in 8 ml. quantities in JSXl2S .. tubes,
autoe1aved at 121°C for 1, minute. and then slanted.
Mediwn 2 nereatic d et ot OB.aein Bllcto-Casitone
potassiUl'rl nitrate pH 1.0
86
i'4edium 2 was dispensed. in one ml.. quanti tiel in 10175 llIl.Tl.
unsterllized tubes. The 18-24 hour growth trom ~.dium 1 was he.v1l1 inoculat
ed into 11ed1W1l 2 and allowed to incubate in a )7°C water bath tor 1 hour. The
presence or nitrite was detected oy adding 3 drops each or 8% d1.lJlethy1a1pba
naphthylruaine in SN acetic acid, and S% sulfanilic acid in SN acetio acid.
The negative tubes were tested with linC dust tor the presence of unreduced
nitrate.
NITRATE REDUCTION TO GAS •
pancreatic digest or casein yeast extract potaas1ura nitrate pH 7.0
(Bacto-Cui tone) 1.0% 0.) 0.2
The medium va. dispe.nsed in 8 ml. quantities in lsn2S mul. tubes
with inverted nals. to collect evo1ftd gas. The medium vaa a1.ltoc1aved at
1210 C tor 15 minutes. Oas wa.a recorded after 1, 2 and ) days incubation.
NITRATE AS A SUBSTITUTE FOR OXYGEN FOR GROWTH
Mediwa number pancreatic digest of ca.ein potaeaiWl1 nitrate pH 1.1 petro1a tum cover
1 (Bacto-Cuit.one) 1%
o
+
2 1% o
-
J 1% 0.1
+
The i118diwa was disp6118ed in S ml. quanti tie. in 13XlOO m:a. tubes
and autoelaved at 121°C tor 1S Llinutes. Macroscopic relative growth in the
above media was contpared at the end ot 18 hours growth.
MANOAN'E:~E PSEUDOMONAS prG:~ENT l-tCT)IUM -pancreatic digest ot casein yeast extract agar II'tanganese sulfate N82HP04·l2H20
Na.tf2POL• H20
pH 7.2
(Bacto-Casltone) 0.5% 0.1 0.3
.01 0.32
.08
67
'I'he medium was dispensed in 4 ml. quantities in 1)1100 _. tub~8
and autoclaved at 1210C tor IS Dlinu:t.es. The medium vas stabbed. ' .. Jhen the
medium showed vater soluble pipent it was extracted with chloroform.
pancreatiC digest ot casein yeast extract agar pH 1.0
(B~eto-Casitone) 0.51 0.3 1.5
The m.edium WIlS dispensed in 15 1111. quantities in tubes then
autoelave1 at 121°C tor 15 minutes. One al. ot li~U8 milk was 8septicall1
added to each tube. The mediUM was then poured into a petri plate. After
solidification the medium was inoculated in the center ot the dish with a
needle for ~;iant colony development. Clearing ot' the agar about the giant
colony was recorded atter 24 and 48 hours.
BLOOD AGAR
yeast extract pL~creatic digest of casein (Bact~-Casitone) sodium chloride a~:ar pH 7.1 defibrinated rabbit blood
The mediUiIl was disoonsed in l~ BIl. ouantities and autoclaved at
88
12100 tor IS minute.. The liquid adi_ va_ cooled to 450C and defibrinated
rabbit blood and proper inoculum added. The poured plates were incubated
tor .L8 hours then retrigerated at 8°e for 12 hours and ilelUOlysis studied wit.h
the aid ot a 35x lens SY8t~ Eoth deep and surface hemolysis was recorded.
B>i,cto L1 taus Milk "1U used.
'I':~6 mediuoa was dispensed in 5 r.Ll. runounts in 13X1oo mm. t..mes and
sterilized rq steam at 100°C :for 45 M.i~tes on three sucoessive days. 'I'h8
,oodiUllt f'o11ow...ng inoculation and inoubation was observed for ch.anges after
1, 2. ), 4, S, 6, 7, 10 and 15 da7s. The pH after 15 days was determdnad
with the glass electrode.
GRffi.,"l'H ON DEsnXYCHOLA'I'E AGAR
Laitson's desoxycholate agar (19.35) was used. pancreatic rl1gest ot casein (Bacto-Casitone) agar sodium. chloride dipotassium phosphate lact.ose ferric ammonium citrate so(iium d~("~x~rcholate (HaltiJllore Biol. Lab.) MutriU r'0,1 (1" aqueous solution)
1.0% 1.7 0.5 0.2 1.0 0.2 0.1 0.33 ml.
Two oJ"gt<llimllS wre streaked lightly on each plate. Growth on the ,
mediUlll was recorded after 18-24 hours incubation .e it ooaparsd with growt.h
on three other ooncentrations of sodium deaoxy-cholate, ll&IIMtly ~, 0.2% and
0.5%.
Table 1 lists the organi~ studied &iving the source of isolation
and rionor. The blank under the heading RECEIVED AS indicates no name was
.. signed at the initiation ot the study. The source of uolation ot JIIU'V'
,trains is probably unknown, hence the blank apace under the heading SOURCE.
The toll owing abbreviations were used to iIrlica te the source of
the cultures:
A tiS ATee ISP5L MSH NCTC NIH NRRL SSM
Academy of Natural Sciences, PhUadelph1a, Pennqlvan1a. American Type Culture Colleotion, wubington, D. C. lllinoil State Public Health L&boratori •• , Chicago, Illinoi •• Mount Sinai Hospit31, C!lieago, lllino18 .. National Collection ot T,ype Culture., London. England. National llUltituta ot Iiealth, Bethesda, MaTyland. Northern Regional Re •• ~ch taboratory, Peoria, Illino1e. St:ritch School ot lied1cine or Loyola tlniversity, Chicago, Illinois.
TABLE I
THE CULTURE COLIECTIOI
lUfl RECEmD AS souacr~ DOlOR
1 Psaudoaonas aeru£iDOSa 'Univ. Ul. Ccll.eg_ 14ed.
2 Alcal¥!re .train Sf dipeptide solution s. Siatsonds .. Yale Univ.
) Pseudoaonas ae!!Jinoaa wll vater it.. Hugh. SSM
5 U~Dh tucaliB U'nb'. m. College Ked.
6 troc tank vater R. Hugh, ss."f
7 trog tank vater R. ifu&h. SSM
9 F. Friever. lSPBL 157lB
10 AJc;.al~ bemol.7.au spinal fillid P. 0ed1I1g. Uni v. Serpa, fIorva7 lOS8
u Alca:J.~ne.a non-heaol.ytic spirwl. Quid P. Oeding, Uniy. Bergen, lo~
12 Alcal1aenH ap. soU t. Fred.~ Purdue Um:w. S5s
13 U~.8P. soU t. Fredarl.ck, Purd\1e Univ. 118
lh Alcal.¥!re raecal i. E. ~, Url1 •• otl.a..
IS UcalteDh riscoaus E. w"nne, tiniy. Okla.
16 Alcalip ... sp. stool it(. Ful. ton, SSi<t 66 ! 17 Alcal1&en&& sp. diarrheal stool M. Fut ton, S5.i<l $39
18 frog tarE vater It. Hugh, SSM
~ :~ 19 frog tank water R. Hugh, SStt
21
22
2)
24
2S
26
27
28
29
)l
32
3$
)6
37
)8
1&2
43
4h
UcaliJenea bookeri
Alc.al1J!nea taecal.1s 'WU'". r&dIcans ,
Alesl.:1geMS viacosus
Alca1Wnea ap.
Alealipnn sp.
A;le;.lipnea ap.
AlealiJenea sp.
Alc.a11ene8 sp.
AlealEMs ap.
Alcaligenes ape
llca1!1ene. sp.
Aloalk!ne8 T18cosua
Alcal.yenea vi8coaU8
Alcal¥!g!s sp.
Brucella Ml1\eus1a . . ..
Brucella Mlit.ensie
Brucella abortus
beet
blood
blood
diarrheal stool
dia.rrbeal stool
diarrheal stool
old stock eul ture
diarrheal stool
water
dead. trog' ~ ~
J. Edwards, ATCC9128, same as NCTC 6$)$
A. gva.na, ATCC 4741. see RH 226 and 317 ,
B. Hammer, ATeC 90)6
\tie Randall. ATCC 10lS3
M. Fulton, SSM: 19Sh
M. Ful'ton, $Sit 22$
!'4. Fulton. SSM 22S8
K. Fulton, SS:i 2294
M. Fulton, SSM 2)92
./1(. Fulton, 5S.11 2b4S
M. Fulton. SSM JOh6
lova S tate Colle.. 1
Iowa Sate Cella. 2
M. 'ul teD, SSM 2686
R. Hugh, SSM
Univ. 1ll. College Med.
Im!a8
lIIB 4S6
~ t: .... g a [ its I ~
45 Bruoella. au1a triR 48) -46 Brucella ~eptiCl1 dog J. Rq. iebraHa 107
47 Brucalla ~i?I!~iJ.pt1cll dog J. Ray, Nebrulca)465
4B Brucella ~ptloa pig J. Rq, lfebn.ska )67:C
49 Brucella brorachiael>~~ pig J. Rtt1'. Nebr~ska 41c:tl
SO Bncalla ~c::hiaeptlC& plg J. Rq, Nebruka Sl26
51 Bncel.la bro~p~ca. pig J. .a.v, Jfebruka S22k
S2 Bncella ~~1 .. pt1ca dog J. RaT. r.iebruka 5246 5) ~~pticta dog J. Rq, ~&aka 6025
S4 Brucella ~ncm~fJeptica dog J. i.a7, Sabnska Wi32
55 Pseudc:aoQaa a~noA Chieago *d. ColJttae 1.
56 Paeudoaonas ... ~ Chicago iiIed. Colle. 2
I S1 Paeudoao .. ~a Chioa.go~. CollI3p :3 ... S8 ~ .. ~sa Ch:1cago .W. College
t 59 ~ nuores08llS Ch1eqo Hod. College 1
60 llcal.i«e __ rae-u. ChicalO ~. College
61 Paeudcaorau tlv.oreacena Chicago t4ed. Collep 2 f= 62 Belicon.a ap. t ... o£ child E. Ol.aen, Delllllll'k 1 I 6) Halloo,.. ap. t ... ot ch1lrl &. Olsen. Del'lllAl'k 2 I
64
6$
66
67
66
69
70
71
83
69
9$
107
Hel.iconau. ap.
Hel.lconelM, sp.
AleYen!! ap.
Brucella abonua
Brucella abortus
Brucella au1a -Brucella ;';;;;;";;;;;;;;;.;;:;;;0 ... !!!!. Brucella melitensis
A.lcal1e!!!! f'aecalis
no Alcal1e!!!! tucalia
III
ill
113
114
US
117
Al~?-&enaa faecalia
PHudOllOM,J a.Nino ..
Pae~ 8:!!j!Iinosa
BJ'UC8Ua Mlit.ensia
Brucella lJIi8li tensia
Brucella .alltansia
feces of child
fecea of cblld
old stock strain
Chicago tap water
Ch1c~ tap_tar
Wheaton tap vater
stool
a\OOl
E.. Olsen. Denaark .3
E. Olsen. TlenIUric 4
14. Fulton, SSM 3130
s. 0arR. Laredo, Te.x:.aa
s. Garza, I..aJ"edo, '1' ... &
5. Oe.raa, Laredo, TfQtU
s. Garu, J.uoedc, Texas
s. aaru, Laredo, 'fexu
E. Weinberg, Iln1v. Chic.
R. Hugh. SSM
R. Hugh, SSM
R. 1luP. SSM
M. Goldin, HSH A.
ft. Goldht. MSB 8
M. OoltHn, ~
H. 00ld1n, MSH
S. GarK, Laredo, Texas ,3021
S. Garza, !.a:I"edo, 1axu 4801
s. llana, Laredo, 1uu 5063
, ... I
I i~
i 3
118 Brucella .IHlite.naia s. Oar ... Laredo,. Texas S2Sl
119 Brucella JAel.itenais s. Garza, Laredo, Texas S37$
120 Brucella .el1tenaia s. Oan&, .i.eredo, T6DS
122 PaaudOJ:iOnas aeruainosa K. HottJiiS,M, Ohio State Univ. .3)0
123 Pseudoaonas ~8a Ne!il lo:'k SottUlie&l. Gardena "Donlf
12.4 AlcaJ.i;I!nas SPa stool M. Fulton. SSM 2120
125 Ucal1en!! f'aecalia blood ;:. Lane. England 1
126 A1ealicenea taeoalia blood w. Lane, I;ngland 2
127 t. Frederick, Purdue uniT. 378
128 vater M. Goldin. MSH
129 uri~ M. Goldin, MSR tA64
1)0 urine M. 001d1 .. 1, }{sH 48)4 "" ! 131 atool !-f. u.w.d:1n, MSB W.J&h M
132 bl.00d 11. GQldin, MSH U~n
I 113 stool rea Gol.cl1n. MSR 4519
13S ~l~1g.li8a tucal1a H. C~ l~C 0150
136 Alcal~~~ I .. calia a tool O. Felaente1d, ATCC 9220 1-137 .u~1aenes t •• calie E. Jordan, L TCC 212 I 1)8 UCUll1cenea 'YisC08. ropy aUk L. Regen, A'fCC 331 J ... as ! 1C'l'C 3233
96 T
able I continued tlu
I ,9
0
~ i i ~
j i § ~ ~ a
! . ~
I ; I , ; ; f f f f f f f f f f li , » ~ ~ ~ ~ ~ ~ ~ ~
J J
J J ~
• ~ 01 ~ ~ I
J J J )
« « I 1 I I I I J J I J I I I I I I
I J
J j J = III
r'~ l
219 Alkalipnea I_calia lD"'ine
222 Bacterium alcal!enes feces
22) Bacteria alcaliJ!nes
224 Bacteriua alcal1cenes urine
225 Bacteria alcallcenes guinea pig lung
226 B.cteri. alcal1&!nes
Bacteria alealigeneB blood 221
228 I.cteri .. ' ... cal1& a1caligenes blood Yar. radicana
229
2)1
Bact.eriWll bookeri
232 Brucella bronchisept1ca
242 Baeter1UDl alcal.!,rnes
245 Alcali&ene. taeCalis
246 Avobacteriwa tUlletaciens
241 Vibrio~
248
249
Vibrio cuneatua
PBeudol1lO1l&S saccharophi1ia
pleural fluid?
egg
throat of dog
ot1tis lHdia
haT infusion
soil
st.agnant water
Allin, Fort \<iUliau MSS1-Sl
lere 4lS
HCTC 655
S. Dyke, ICTC 1200
JlCTCl219
ICTC 1341
JCTe 1101t, .a. as Rbaen 18
/1, ~
b'g. ~$ _ :;r,.... c::8 ET -_ ..... :=:.0 C» :is ....,.. '<2 iI.~ .;tg,.
~: a
A. EftDa. MeTC )Sl2. see RH 22 and 317
NCTC 6572
F. Fr1ever, ISPHL 6789A
L. LeBeau. Uni.,.. ID. Col. Med. 11-1 ... s. Cowan, England "L" !
CD
Midwest Culture Serrlce, Terre Hautey Indiana g
M1dveat Cl1l. t .... Service, TerreHautei Indiana I Mudd and riarnn, It. Tee 8461, Bee RU t1 260 and 261 B ~ B. Grq, !'fCC 6972 i M. Doudoroff, A'l'CC 9114 ~
2$0 Pseudomonas iehWosmia spoiled adlk M. Speck, !TOe 7966. sau as HCTe 8049
251 PseudOllOnas hydrophila M. Speck, A1'CC 1965, sau as NCTe 7812
252 PseudOllOnu ~roeUa frog aeptieaia. w. Kulp, !TCC 90n
25) Aeromonaa liquetaciens A. U\q'fV, liolland L4l1
254 Aero.onas liquetaeiena canal water A. KlUTNr, Holland L4lB, see l'm 271
2SS Aeroaaonaa hldrol&Ua 1'roI aepticea1a A. Kl~r, Hollam UW1
251 !lC8.l1genes sp. stool ,. 'riner, lSPHL 273)~
2S8 Alcal!J!n!s ap. stool ,. Fr1awr, !SPliL 2133
259 ,U.cal~enas ap. otitis media S. Cowan, England, "P"
260 Vibrio p!rcolans ..... 1ant of RH 241 R. Hugh, SSM
261 Vibrio p!!'colans 'fV1ant ot Rl! 241 R. Hugh, SSM
262 Acetobact.er aceti ATeC 8)0) ... 26) Acet.obacter oxydans ATCC 941) ~ .,
H 266 P.eudOlllOll&a sp. paracentesis fluid F. 'rawer, lSPHL .3369 (')
8 268 strum Keller, AJrS 49.3 U/2 ct'
t; 269 sVe_ 'eUer, AIS SI7 IJ./4 I
Sl.
2n Aeroaonu liquetaciens canal water E. tUles. England Lbl8, see RH 254 :rg i
272 Chl"cmobacteriua ~~\~ Hopk1na Marine sta., Calif., .1. G. ! 2.1 • 273 SpirUlWil TirginianUtll Honkins kiarine sta •• Calif •• 0.1.1 ~
274 Rhodospirillum rubrua Hopkins Marine st.&... Calif.. ATR. 1.1.1
275 R~~~~Uti9JllOtl&~ ~llStr1!t Hopkins .Marine Sta., CalU., ATR. 2.1.1
277 Acetobacter rancens liopkins Marine Sta., Calif'.. M. A. 7. f 280 atreara Keller, AHS SO tJ./)
283 st.rau Keller, ANS 12 lJ./2
284 a1i.l'8aa Keller, ANS 100 IJ./2
287 ave .. KeUer, AlIS 154 IJ./2
289 at.reaa Keller, ANS 156 IJ./3
292 atreu& KeUer, AHS 169 IJ./3
294 streUl Keller, ANS 186 u./2
29$ atrea Keller. ANS 196 U/2
296 stor_ Keller, .lIS 199 IJ./2 tof
=-298 atre8lll Keller, ANS 220 IJ./2 tit H
300 stream Keller, .us 22S IJ./2 0
)02 str'eaa (eller, .lIS 229 U/3 ! b
304 atzeam 'eller, AJS 2)4 IJ./2 I )06 atreua Keller, ANS 239 U/2 i~ )07 atreu Keller I AJIS 247 IJ./2 i
': )08 .toreara Keller, Al'iS 2SS IJ./2 (g
)09 aweg Keller, AliS 262 IJ./2
)10 streaa Kelle1", AliS 265 U/2
)ll .tre_ KeUer, ANS 269 U/2
)12 stream Keller, Iu~ 210 Ll/2
313 stream KeUer t AHS 281 IJ./2
)lh s1ireu Keller. .lIS 286 IJ./2
318 stream te11er6 ANS 309 U/2
)20 aWeUl Keller, ANS )28 IJ./2
22 at.reaa Keller, ANS 3.32 IJ.j2
24 stz"ea1a Keller, ANS 334 IJ/2
30 stream. KeUer, Ali"S 339 IJ/2
i333 streu Keller, ANS ~O IJ/2 & 133S Keller, ANS 341 IJ./2
.., stream •
a-t
stream Keller, ANS )60 IJ./2 I (eller, 11S )61 U/2 strealla I stream KeUer. .lIS )62 Ll/4
streaa Keller, ,lNS )66 U/2 is straaa KeUer, ANS )67 Ll/2 i
0: stream leller, AlfS 408 IJ./2 ~
~ stream Keller, AN!) 410 U/2
347 aVe_ relier, AN;;, 433 IJ./2
349 str .. Y.eller. A..'1S 434 u./2
35) stre .. r.eller, A1ts 473 Ll/2
)S5 stream Keller I A.lJS 4B2 U/2
)60 blood of frog vi th R. Hugh, f:s'lI{ red leg
363 blood of frog wi ttl R. Hugh,. SSM red leg
364 blood of frog with R. Hugh, SSM red leg
)70 parapwt.uas1a baolllus G. !lde.ring, Mich. tept. Health 21-)53
37) parapertuaau bacillus a. Eldering, Mich. Iept.. health 2l-6S6
374 parapertussis bacUlus o. Eldering, ftich. Dept. Health i1 ~ 23--456 •
)75 Pf'eifterella paeudoullel Glq'1 Ne'I'C 7431 ..... g
376 U~1&ene. _ sp. !TOe 213, received thru J. Ulrich ~
i of *70 Cl1111c, Minnesota ~
371 Alq .. U.p~~ .p- blood A. Iwms, ATCC 4141. reoeiftd thru -i b J. Ulrich of Kayo CliIlic, Minnesota \,.>J
see IH 22 and 228 1
)76 .A.l.caligenu t_calls ATee 81ldl, receiwd tbn J. trlrlch , of Mayo Clinic, Minn.esota, .. lUI 174 ~
., 319 ~l~~_~~~ ape tecea ATCC 9220, received thru J. Ulrich or
Mqo Clinic, 8M lUi 1)6
)80 Alcal1J!l!!8 sp. old stock eul. tUN J. Ulrich, Mqo Clinie UlO
)8) orange E. !Ailson, SSM
)84 PseudOlaOn&8 ap. dysentel7 stool M. lul ton, SSM 3108
)85 Alcaligenes ap. abdOlldnal nab K. )"ulton, SSM 2781
)87 ~~J:.li\ typhoaa Pan.aaa 58 atrain
JSS SalJaonella para:2;phl Unlv. Ul. College Mad.
)89 Salaone~ achottauelleri o. Fel.senteld, Belctoen Inst., Chicago
390 salaonella enteridi t1s o. Felsenteld, Hektoen Inst., Chicago
398 Proteus lIOrganii M. j."ul ton, SSM 2181
401 Proteu rettgeri M. Fulton, SSM %7 to!
40S Proteus '9'Ul~is a. Fulton, SSM 3488 ! • 406 Escherichia coli Univ .. Ill. CoJ.la&e Ked. H - 0
407 Aerobacwr aerogene. UniT. ill. College M6d. 0 ::s c+
409 Vibrio ~ch.ol.ra o. lelaenteld, aelctoen 1n8~, C.bie.1
JUo Bactariua aniwa twa w. Ferguson,liich. 1»pt. Health 5h2 ~ f-'
411 BacteriUlll anitrat.a w. Ferguson, !tich. 'Cept. Health Sw) i i-i
412 V:ibr1o CQIIbI& opva o. Felaenf'eld, Hektoen Inat. Chic. ~
413 Vibrio coaa inaba o. Felsenteld, Hektoen lnst. Chic. ~ i ~
r ~
lUll Shfjella er!dzsent.er1ae IV
1117 PseudOllOn&s ap. M. Fulton, SSM 2661
418 PseudQlllonas SPa urine M. l ulton, SSM 2689
U9 Pseudoraonas sp. ur1ne M. F'ul t.on, SS-"t 2803
420 .Pseudomonas sp. M. Fulton, $Si'{ 280$
421 Pseudom.onu ap. Ii. fulton, SSM 2820
422 Paaudcaonas spa M. Ftuton, SSM 30$1
423 Pn11doaonu ap. , l.. Fl!l. t.on. SSM 3099
424 PaeudOJlOn&8 ap. M. Fulton, ss:.t )100
42S Paeudoaonaa ape 1(. 'ul ton, SSM )123
426 heudOlaOn&8 ap • i;4. Ful ton, s..~ Jl24 . 427 heudcaonu spa i't. Fulton, SSM 3l4O ...
I-428 P .. udaoU8 ape i4. Fulton, SSM 3154
.., • to-!
429 heudCtllOl1U ap. r;f. Fult.on, SSM )162 ~ 0
4)0 !lcalk~~ IJp. J. Ulrich, Mayo Clinic 249 :l
i 4)1 Alcaligenes ape J. Ulrich. }tqo Clinic 282 Ot
432 anaeropnic paracolon bacUla Enduoeba biatol. 'ria Ba.l..aI1uth, Nortbwatern t1n1y. Ii IRS CUlture
I 433 paracolon bac1llua stool F. Fr1ever, lSPHL 66)01'
b)6 ~lla aaadMpenei8 old stock culture F. Frlewer, lSPHL 229 !
106 0
Table I
oontinued trQA
l ))&
88
90 f CJ
..... ... n .. • +3 ~
~ J at
..::t .lIIC
0 0
~ ~
'E I ~
.. i '0"
.. 'iI
J i '''l
~~
i ..
~
~
l! :
~
• •
• •
f>I:. 0
lJQ
p~
H
C <ER V
"~m'.M. )ATA Phow, ~ ero .phs .)ho .ing Bactor1al r'lage 1
· C •
•
•
. Figure) Figure 4 --- '----- ------o")j
-'-'
107
108
Figure 1. Alunes bro.'1Ch1canU a1lov1Dc t.yp1oal perivielloua tlqellation. Pho crograph i45l)b.-
Figure 2. Ianthcuon&s ul~l11a 8p. 1lOY. ahowlng Wi-cal polar aulti&ionoua l1ageiIAt 00.- Photca1cro .... ph l4OOO.
Figure). Lophobaeter fucal1a gen. et .p. nO'¥'. ahO'W1nc t1Pioal. lophotrIohOus lIigerlat.1on. Photca.1vopaph~.
Figure 4. P .. udaaonas a~1nola ehOldng tTPioa1 RlOnOtriobou tlaaenat1on. AiotoiIorograp llt'RXS.
KEY TO THE TABUS
peri polar multi lopho mono ? -+ 2 ... (0
:t blank A B b r nc r
perltrichous polar JIlul ti triohoua lophotrichous tIlOnotr1cnous type of polar nageUat.lon aq be quee tioned. nonmotU., Mgati'ft test, no hemolJ"s1& positi.... test. or turbidity' degree of tl1t'biditq greater than ... poeitiw test tor nitrite and gas, acid and gas poor growth teat not pi rtoraed alpha hem.olytlis beta h8Jlol",1s blue color or litmus red color of litaus no change of indicator carboh1drate termented
.....
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I 51
g
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ell
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c+
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+
,
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tose
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'"" '""
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l.I'\ l.I'\
l.I'\
Table II contd. from page 109 1% 'b1oOli litmu~ , casitone pigment agar milk
hemol.
J.. § Q) +l ;:j ~ "' Q) .$ s:: C1I r-I .-i
0 .!I: +l 0 Q) .g "' .r! '0 III ~ "' !1 r-I r-I J.. +l Q) CD 0 r-I .g r-I ~
t 0 Q) "' Q) ." J.. M +l 0 Q) 0 ..c:
~ r-I I/) Q) Q) Q) Q) 0 0.- Q) .~ "' ~ 0. r-I til tID Q) 0 r-I 0 III III I/) I/) CD .~ i
I +l r-I 0 C1I 0 ~ IV ~ ~ 0 0 0 I/) r-f I/) "' .., 0 8. ~ I/) C""\ M C1I M
s:: tID J.. .., s:: 0 ~ 0 Q) .fj "' C1I M r-I 0
~ 0. c.... 0 0
"' >< 0 0 r-I
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.., o oro a u r-f IV 9 r-I III
;§ d ~ "' :::l ~ ~ s:: ~ g .r! 9 CD N ::x: 0 IV 0 ::x: Q) r-I I/) .r! 0 tID ::x: Z 0. s:: :z: ::x: u 0 E! '0 I/) 0 0. '0
54 peri - - - - - - - - - - + + - - + - + - - - - - b 806 + -55 mono + - - - + + + - - - + - - - (f - + 2+ + - + - + b 8.0 +
56 mono + - - - + + + - - - + - - - i - + 2+ + - ... - + b 7.9 +
57 mono + - - - + + + - - - + - - - i - + 2+ + + + - + b 7.9 +
58 mono + - - - + + + - - - + - - - i - + 2+ + - + - + b 7.9 +
59 mono + - - - + + + - - - + - - - i - + 2+ + - + - + b 8.1 +
60 peri nc - - - - + - - - - + - - - i - + 2+ - - - - b 8.5 + -61 - + - - - + + + - - - + - + - If - + 2+ + + - + b 8.2 +
62 - nc nc nc nc nc nc nc - - - - - - - - - + - - - - - ! r -63 - - - - - - - - - - - - - - - - - + - - - - - nc -64 - - - - - - - - - - - - - - - - - + - - - - - nc -65 - nc nc nc nc nc nc nc - - - - - - - - - + - - - - - b -66 mono + - - + + + - - - - + - - - " - + 2+ - - - + b8.) +
83 peri nc - - - - + - - - - + - - - If - + 2+ - - - - b 8.4 +
.... ....
.... ....
N
N
N
.....
.....
N
N
N
N
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Um
ber
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:;3
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:;3
:;3
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:;3
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0 0
0 0
0 0
0 0
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ag
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+
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+
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tro
se
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s la
cto
se
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+
I su
cro
se
IJQ
CD
mal
tose
.....
0 \0
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nose
+
+
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lose
+
+
+
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ol
met
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Vog
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+
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ate
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rea
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tin
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pet
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01
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pet
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.....
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+
+
+
+
+
+
+
+
+
t'llQ
. N
N
I\
.)
N
N
N
N
NOJ
& p
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+
+
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+
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0'
0'
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J
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CXl
C
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CXl
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J
CXl
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. .
. .
. .
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. pH
0
0 I-
' 0
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I-'
0 0
-J
N
0 '"
+
+
+
+
+
+
+
+
+
+
deso
xych
olat
e
~
,
Table II con td. from page 109 , 1% casitone I pigment
M 3 ~ ~ ~ e '" Q)
I:: "'~,.; ,.; o ..!J4 +' OQ).o • .-f 'tj 10 III '" M"; .~
M +' G) Q) 0 :;!,.; +'.0"; M It ~ '" Q) 10 Jot M +' 0 Q>.=' 0 aI
~ ,.; 10 G)Q) Q)G) 0 o..~ c '" M 0.."; II) b.O ~ ....... 0 lOll 1010 ~ ,.; f~.;.t ,.;+' 0 aI (.) Q) MOO 00 Il'~";p,. lIlal +' OQ) al!jlll (Y'\ MaiM
I:: bD +' +'M +'~ 0.0 o..tf ~M alai Mo.. ,.; 0 ..!J4 0..ft.4 0 aI >< U 0 ,.; aI 'tj+' bD+' ~,.; Cfl +' 0 u,.; ~ ....... ,.; ~ aI =' r;j ~ M c ~ 0..-1 M G) _N 0 Q) 0 0 N:I: 0 ~ (I) ~ 0 '-4 '0"; 11.1 it >{aI orl/:1 ~ 0 =' bD ..... :z 0..1:::Z:::C: 0 0 III 'tj 10 U
?mono 1 + + + + + 1 + - b 8.2
+ + + + + + + - - r 1+
lopho I - + - 2+ + 1- - Ib 6.7 1+
+ + - A + b B.o I +
lopho I - + - 2+ + 1- 1 - - Ib 8.1 I +
peri + + - A + b 8.41 +
peri I nc - + + i + 2+ - b 8.4 +
per i nc - + ,,- + - i + 2 + - b 804 I +
peri 1- + + - b 8.4 I :l:
no nc no no no no nc - + - nc
peri I - + + + I - + - b 8.5 I +
mono I + + + + + i I - + 2+ • + + b 7.7 I +
mono I + + + + + - i I - + 2+ • + + b 7.9 I +
mono I + + + + + i I - + 2+ I + 1+ 1 - + 1 b 7.8 I +
Table II contd. from page 109 1% 1 blood 1 litmus' , casitone pigment agar milk
hemal.
H § ~ ~
§ '" (I) Q) ~ '" r-4 r-f ~ 0 .!:t: ~ 0 Q) .g II! ..... 't1 CI) 3 II! !l r-f r-f H +l Q) Q) 0 r-f 11 r-4 ~
t 0 Q) co Q) III H H ~ 0 8. 0 ...c: ~
r-4 III Q) (I) Q) (I) 0 0.. Q) .~ II! b r-4 III b.O Q) U r-4 0 CI) I/) III CI) Q) .~ r-4 I ~ r-f 0 CIS u ~ (I) b 0 0 0 0 III rl
~ I/) '" ~ 0 Q) ~ III f""\ ~ '" H ~ :: ~ H ~ .~ o .D 0 ~ !1 CIS '" ~6 b 0. cr rl ..!i4 0. ft..t 0 I< u U r-f rl CIS "0 (I) r-4 o (.) r-f
~ Q) a 'd :x= III ::t: cl (I) CIS ;::! m ~ ~ .~ ~ 0 .....
~ (I) ~ 0 (II :x= 0 (I) Q)
~ 't1 rl I/) ;::. u b.O :x= z s:: z :::r:: u u "0 CI) U 0. 't1
~44 mono + .. - - + + + - - - + - + - ~ - + 2+ + - + - + b 8.4 +
~45 mono + .. - .. + + + - - - + .. + - i - + 2+ + - + - + b 8.3 +
~46 mono + - .. - + + + - - - + - + .. ~ .. + 2+ - + - + b 8.1 +
~47 mono + - - - + + + - - - + .. + - If .. +2+ + - + .. + b 802 +
148 mono + - .. - + + + .. .... - + .. + - " .. + 2+ + - + .. + b 8.1 +
149 mono + - .. - + + + .. .. - + - - - i - + 2+ + .. + - + b 7.6 +
150 mono + - .. - + + + .. - - + - + - i - + 2+ + - + .. + b B.o +
151 mono + - - .. + + + - - - + - + .. i .. + 2+ + - + - + b 8.2 +
153 polarmulti + .. - .. + + + .. .. .. + .. .. - - .. + - + - .. - - b +
154 ?polar multi + + .. + + + + .. .. .. + .. - .. - - + .. + .. - .. .. r +
155 mono + + - + + + + .. .. .. + .. .. .. .. - + - .. - - - b 6.4 +
156 mono + - .. .. + + + .. - .. + - + - Ii - + 2+ + - + .. + b 8.1 +
157 mono + .. - - + + + .. .. - .. .. - - " .. + .. .. - - - b 7.1 +
159 - + + - - + + + - .. - + - .. .. .. - + - - - .. .. r .. I
ep +
+
+
+
+
+
+
+
+
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r-i r-I
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r-I r-I
r-i r-I
r-I r-I
r-I r-I
Table II contd. from page 109
J..4 Q)
1 ga
190 mono
191 mono
192 mono
5 -M ~ cd
~ ell bil cd q
193 polar multi
194
195 mono
196 mono
191 polar multi
19B mono
199 polar multi
200 polar multi
201
202 mono
203 mono
Q) III C) Q) o ~ III ~ 0 0 +3 .jJ ~ ~ CJ CJ ell cd ::; '0 r-4 ~
+
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+ 2+ I +
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brown
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b +
B +lbB.11+
+lbB.21+
+Ib +
B + I b +
b 8.3 I +
b B.2 I +
- I b 8.) I +
A ... lbB.OI+
+lb8.31+
- • b
Table II contd. from page 109 , 1% casi tone\ pigment
~ ~
~ i'II ~ Cd Q) Q) rl rl ~
0 ~ ~ 0 Q) .g ClI .r-! "0 CIl ~ ClI b rl rl H ~ Q) Q) 0 rl .g rl ~ t 0
Q) cd Q) CIl H 6:: ~ 0 Q) 0 ..c: .0 rl CIl Q) Q) Q) Q) 0 $ ~ ClI b 0.. rl CIl ~ (I) 0
~ rl 0 CIl CIl CIl CIl Q) ~ rl I .r-! rl 0 ClI 0 ~ Q) H 0 0 0 0 CIl .r-! rl E CIl Cd ~ 0 Q) otoJ III C"'"'I H ClI H ~ bD ~ ~ H ~ ~ 0 .0 0 ~ b cd cd Jj 0.. rl 0 ~ o..fo..t 0 0
i cd >< t? Y. ...-1 r:-I ClI ~ ~ Q) rl til .!"" _('f"' 0 U rl rl (I) H rl CIl rl Q)
-
~ '0
204 mono + + + + ... i + 2+ - b 8.3 I ...
205 polar multi + + + - 2+ + yellow b 8.0
207 7lopho + ... ... + ... i + + yellow b 8.1 1+
208 Inono + ... + + ... i + 2+ ... ... ... b 7.5 1 +
209 mono ... ... ... ... + + i + + + + + b 7.5 1+
210 mono + + + + + + i + + + + B + b 8.4 1+
211 + + - b 800 I +
212 7lopho + + b 6.9 1+
213 polar multi ... + ... + + + + b 1+
214 peri + + + b 8.6 1 +
216 I - + + -1- ... -I- I- I B +lb7.71+
217 mono + + + + + + " + + + + + + b7.71+
218 mono + + + + + + " + 2+ + + + + b 7.8 I +
219 + + + + + - b7.91+
al-eTO
l{OA
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Table II contd. from page 109 \ 1% casitgne\ pigment
H 3 ~ +>
I=l 3 ell Q) ell rl rl
0 ~ +> 0 Q) -§ ·rl -0 U) § ell H rl ~I I H +> Q) J) 0 rl +-' -§ rl &
Q) ell Q) U) H H +> 0 Q) 0
] rl U) Q) Q) Q) Q) 0 0.. Q) .~ ell b P... rl en bD rl 0 en U) U) U) Q) .~ rl i +> rl 0 ell Q) 13 0 0 0 0 U) rl >. U) '" +> 0 Q) ~ U) CV"\ H
I=l bD +> H +> § 0 .g 0 .G ~ b ell rd b P... rl 0 ell >< 0 0 rl rl -0 +> Q) rl C/) 0 0 rl
::rl rl Q) ell ~ ~ ~ ~ ~ .~ Q) o ·rl 9 Q) N 0 Q) 0 0 N ::r: 0 ~ ~ -0 rl U) E! P- o bO ;:c :z: 0- c: :z: ::r: 0 0
263 + + + Ib 7.0
266 .nono + + + + + + + + ~ + 2+ b 7.2
268 mono ~ peri + + + b 7.1
269 yellow-
Ib 6.7 mono ~ peri + + orange I + I
271 peri
272 mono + + + + + + + + + b 7.5
273 lopho + + + c 7.1
274 lopho + red c 6.5
275 mono + red b 6.9
277 + • +
280 polar multi + + + + + + + b 8.2
283 peri + + + + + + + b 7.8
284 mono + + + + + + + + -1- + -I~ Ib ;.7 287 polar multi + + + - - + yellow + b 8.3
'vJ
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W
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III
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III
0 0
., .,
., fl
ag
ell
ati
on
H
i3
a a
(')
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~
~
0 f-
-' :J
c+
c+
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c+
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'-f-
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<,
to<,
to<,
to<,
H
) +
-+
-+ -+
-+ IE
> -+
.. ..
-+ -+
IE>
IE>
dex
tro
se
'1
0
+
+
.. ..
.. I la
cto
se
S +
+
'0
+
I su
cro
se
III
()Q
CD
+
.. +
..
+
+
-+ I m
alto
se
~
0
+
+
+
+
.. -+
lman
nose
\0
+
+
..
+
+
+
+
+
+
-+ +
I x
ylo
se
+
-+ ..
+
-+ +
+
..
I ara
bin
ose
ind
ol
met
hy
l re
d
Vo
ges
-Pro
skau
er
+
+
.. -+
-+
-+ +
+
+
I cit
rate
+
+
+
u
rea
+
+
I gela
tin
(') III
(Jl
+
+
+
Ino
p
etro
latu
m I f-
'-...
.. +
+
..
+
+
.. c+~
0
tro
latu
rn
::s CD
+
+
+
IH20
solu
ble
CH
C13
solu
ble
"d
...,- (1tl
a CD
_col
or
::s c+
\
+
+
Imll
k ag
ar
deep
surf
ace
0-
0-
0-
0-
0-
0-
0-
0-
0-
0-
colo
r -.
.J
-.J
-.J
-..J
(X
l (X
l ex
> -.
J
-.J
(Xl
-.
. .
. .
• .
. .
pH
\J\
0 N
'v
J I\
)
0 'v
J \J
\ 0
'vJ
des
ox
ych
ola
te
.......
....:::t ....:::t
\I\ o
• .
r-r-
\,()
,D
.relle +
+
+
~
0 ~
.10
TO::> r-t
C
r-t Q
) ~
~
tT:JH
8 .,-1
aTqnT
os p..
aTqnT
os O~H
+
+
Q)
wn~:eTO.I+9d ~
ON
+
+
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s:: 0 um
lET
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ou +
+
+
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~
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N
N
en
wnleT
o.I+ad
+
+
+
ON S~H
U"neTa~
+
ea.rn +
9le.I+1::> +
+
+
+
.Iene}{so..Id-sa~o
pa.I TA
lf+aru
TOPU
1
asoU1qe.Ie
+
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+
asoutnroJ +
+
+
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+
+
+
9S0.I::>ns +
asol::>-eT +
.,
aS0.I+
xep +
+
+
+
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U01leTTa~eTJ
0 0
0 0
.,-1
c s::
c s::
... 0
0 0
0 Q
) s:;
s:; S
S 0
-
r-t N
1
"'\ ....:::t
co .Iequm
u H~
r-t r-t
r-t r-t
r-t C
""\ C
""\ 1
"'\ C
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"'\
Lf\ . +
:. 0 rl
r-t
~
+
+
+
+
N +
0
+
~
+
+
~
~
~
0 0
.,-1 0
s:: 0
s:: ...
s:: 0
s:: 0
CI> 0
l"! 0
l"! 0
-S
C'.
S
0 N
....:::t
0 1
"'\ N
N
N
1
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+
+ +
l!I ~
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s::
... 0
CI> e:
0-
\I\ '-0
1
"'\ 1
"'\ C
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+
+ +
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-
cc 1
"'\ 1
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,D
+
+
0 s:: 0 l"!
~
C""\
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\.
.)
\..)
\.
.)
\..)
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.)
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.)
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\.
.)
\..)
IRH
num
ber
-J
-J ~
0-
3'
V1.
\n
r:
-r:
-g::
r:
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1.
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()
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Ib
0-
S f-
J P
....,
a s
s '0
s
a en
0 '0
0
0 0
m
0 0
0 ::::
l 0
::::l
::::l
::::l
'1
::::l
::::l
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H
0 I-
-' 0
0 0
~
0 0
0 H
Ib
fl
ag
ell
ati
on
'1
("
) 0
~ !:
j c+- o..
c+-
o .....
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<)
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....,
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dex
tro
se
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+
+
+
+
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Cf)
+
~
+
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tose
'0
ill
I su
cro
se
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+
en
+
+
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tose
f-
J 0 '-0
+
+
+
+
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nose
+
+
+
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lose
+
+
+
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bin
ose
ind
ol
met
hy
l re
d
Vo
ges
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skau
er
+
+
+
cit
rate
+
+
+
ure
a , I
+
gela
tin
+
+
+
+
IH2S
NO
IPet
rola
tum
("
)
+
Ib
til
N
N
1-'-
t-'
+
+
+
+
+
+
+
+
+
+
no
petr
olat
um"'
" c+~
0
+
+
INa
) &
pet
rola
tum
::::
l en
+
+
+
+
+
+
IH20
solu
ble
CH
Cl)
solu
ble
'0
I-
<)
.... - O'Q
0"
0-
0-
nl-
-'
~
f£ '1
'1
'1
o
m
0 0
0 1-
-'(1)
~
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§ §
00
-l-
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lor
c+
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~
p..
+
Imilk a
gar
deep
su
rfac
e
0'
0-
0"
colo
r ..
co
-J
-J
• 0
0 pH
I\)
\,()
co
des
ox
ych
ola
te
r---
\.oJ
\.oJ
VJ
\.oJ
VJ
VJ
VJ
\.oJ
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\.oJ
V
J V
J V
J w
IR
R nu
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r '-
0
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en
en
en
C
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en
CD
C
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.... '3
CD
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D
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D
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'0
'0
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(1)
CD
CD
CD
CD
('8
0 0
CD
('8
CD
CD
CD
0 "1
"1
"1
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'i
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......
'i
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'i
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......
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~
~
1-'-
1-'-
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\l)
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1-'-
1-'-
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f1 H
0
'i
flag
ell
ati
on
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n
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("t-
("t-
("t-
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1-'-
~
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) H
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) H
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) d
extr
ose
:(
~
q;)
~
~
+
+
+
+
+
§ +
+
Il
acto
se
'0
III
sucr
ose
OQ
+
(1
)
mal
tose
.....
. +
0 '0
+
+
man
nose
+
+
+
+
+
Ixy
lose
+
+
lara
bin
ose
ind
ol
met
hyl
red
Vo
ges
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skau
er
+
+
+
+
+
+
+
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rate
ure
a
+
Igela
tin
H2S
+
~
~
+
q;)
IN
O pet
rola
tum
n III
ti
l
no
pet
rola
tum
1-
'-1-'
+
+
+
+
+
+
+
+
c+-~
0 ~
~
~
NO
) &
pet
rola
tum
::l
+
+
+
CD
+
R 20
solu
ble
ReI
) so
lub
le
'0
I-'-
(lQ
i3 ro ::l
("t-
'"
ho
late
·rp~tot{oAxosap
Hd
., .I°TO
O
aO"eJ.Ins daap
.Ia~'e :>tUUI
~
.I°tOO
I tt~H~
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tos
+
+
atqn
tos OZH
+
+
+
+
a> run 'll~ to
.I,+ +
+
+
s::
"i
N
N
0 ,~~
rl
-,-1 +
+
+
+
+
+
CJ)
It! ()
$ $
$ $
SZH
trnata~
+
+
+
+
aam
a'la.I'+ 10 +
+
+
+
+
+
..rana:>tso..rd-sa~oA
pa.I tAt{'l
t op u1
asoU1qa.za
+
+
+
+
+
+
9S0t AX
+
+
+
+
+
+
9S +
+
+
+
+
+
0
\ 0
aso'+t aW
r-i
Q)
QD
It!
aso..rons 0..
aso'lo-et $
~
+
+
+
+
S ~
~
~
~
0 f..t aso.I'lxap
+
~
+
+
+
+
+
+
+
+
+
'+-i ~
~
~
~
. "0
+> s:: 0 ()
u01+atta~1l1J
H
.,-1 ..-t
.,-1 0
0 0
0 0
0 0
H
f..t f..t
f..t s::
s:: s::
s:: s::
s:: s::
Q)
Q)
Q)
Q)
0 0
0 0
0 0
0 r-i
0.. 0..
P.
e s
s E!
s e
s .0
q
j M
\.I\
'-0
r-0
-0
rl
N
1""\ --=:t
r-eo
0-
0 E-<
..raqnmu
0 0
0 0
0 rl
.3 rl
.3 .3
rl
M
rl
N
--=:t --=:t
--=:t --=:t
--=:t --=:t
--=:t --=:t
--=:t --=:t
--=:t
Table II contd. from page 109 , 1% casi.tone\ pigment
S +l
;3 I1S Q)
t:::: I1S rl rl 0 ..\f:: +l 0 Q) .g • .-1 "0 rn S ~ .!3 rl
J..I I +l Q) Q) 0 .g rl t I1S Q) rn J.. J.. +l 0 Q) 0 I1S rl til Q) Q) Q) (l) 0 Il.. Q) .~ I1S J.. 0- rl rn QD rl 0 til til rn rn ~ .~ rl I +l rl +l 0 I1S
J.. 0 0 0 0 'dE rn I1S +l 0 Q) al!1 rn 1"'\ J.. +l +l J.. +l t:::: - 0 ,.C ~!l I'd I1S J.. 0- rl 0 >< 0 0 r;d ~ rl I1S -g +l Q) rl CIl +l 0 ~ rl Q) lIS ;:j ~ ~ Q) 0 ·rl S Q) NO Q) 0 0 (\I 0 "0 rl til S • .-1 I> 0 QD ::x: Z 0- t:::: Z ::x: u 0
421 mono + + + + + + i + 2+ + +
422 mono I + + + + + + (JI - + 2+ I + +
423 mono I + + + + + + il - + 2+ I + +
424 mono I + + + + + + il - + 2+ I +
425 mono I + + + + + + i I - + 2+ I + +
426 mono I + + + + + 1- il - + 2+ 1+ +
427 mono I + + + + + + \j I - + 2+ 1+
428 mono + + + + + + i + 2+ +
429 mono + + + + + i + 2+ ,
430 weak
polar multi + + + 1- yellow weak
431 polar multi + <I- + + yellow
432 + + + + + + +
433 peri Ifi +
f+
~
g: ::-
~ IRH num
ber ~
VI
~
VI
'0
t-3 III 0
-i3~
i3 ~
..... 81='.:
0 0
Cll
5 '0
o~
::r H
0
H
Cll fla
gella
tion
::s
0 c+-
o 0
::l s::
c+ en
P-o ~
+
+
+
lOext.rose
Ii 0 i3
""d ill
+
+
Isucro
se I)Q
Cll
+
+
Im:.!.l t.n
l'll'! f-' 0 '0
+
+
+
+
+
+
og
es-Pro
skau
er itra
te
+
+
j 0 III en
... +
+
Ino
oetro.Latum
1--"
I-' c+
"lA
0
&
petro
latum
I ~
lub
le
solu
ble
'd
1--" '1
I)Q
S Cll
Cll 0
-::s c+-
-oJ
-oJ
I""""" TABLE III
GROUP DIFFERENTIATION OF THE ORGANISMS IN TABLE II •
i""-" pigm't
I:: Q,)
r-I 0 Q,) .g ..-I r-f C/l ~ .g r-f M
OJ m 0 OJ -t> C/l Q,) Q) r-f $ .~ r-f C/l
~ .,-i 0 C/l rn r-f 0 ~ § g. M 0 0 Q) CIS ~ rn ..., ~ ~ M bD !3 CIS CIS I:: .,-i -g 0 >< 0 0 m Q) r-I 0 ~ § M OJ C1I :::J r-f ..-I g OJ (II
iE rn QD '0 r-f rn ~ I:: bD ::r: ::r:
A 1 - - - - - - - - 135, 223, 258, 376, 378
- - - - - + - - 222, 336, 338
- - - ~ - + - + 322 yellow
- - :~ - + - - - 48, 49, 50, 51, 183, 184, 214
~ - + - - - ~ 138, 260
- - - + + - ~ - 29, 46, 47, 52, 53, 54, 140, 171, 172, 173, 180, 181, 182, 225, 227, 232
- - - Ij ~ - - - 12, 13, 60, 83, 136, 137, 379, 380
2 - - - - - + - - 294
- - - - - + - + 162, 287, 430, 431 yellow
nc - - ~ + - - - - 7
- - - 't - + - - 229
- - - + - + - + 205 yellow
3 - - - - - - - + 274, 439 red
- - - - - + - - 273
- - - 1 + - - - - 17, 31, 32, 37, 124, 131, 133, 212, 226 261, 385
- - - + + - - - 185
contd. pigmt 130 ., 131
r-t ., ., -§ r-t 0)
-§ r-t 1-4 0 .,
r-i 0)
~ 0 .~ 0)
s::
- - - + -- - - - - + - rose-black
- - - + - - 21, 27, 28, 125, 126
- - - + 275 red
- - - j + - - + - 25, 26 brown
- - - 0 - - -- .. - + - + - pink-orange
- - - + + .. - -- - + \I - - - + yellow-brown
5 - - - - - - 14, 18, 19, 23, 35, 36, 1)2, 63, 64, 110, 127, 128, 132, 139, 160, 175,
, 194, 211, 245
- - - - - .. + - brown
- - - 0 + - .. , 165, 166, 167, IG~, 170, 216
- - - - + - - -- - - - + - + - 373, 374 brown
- - +
+ 69 ye11ow-organge
+ ... - - -
Table II contd. pigm,tt from pa 130
s::: II» 132: r-i ., 0 II» -§ ..-i rl 10 ...,
-§ r-i J.o Q) aI 0 ., ~ 10 Q) II» r-i Q)
~ rl ~ ~ ori 0 10 10 r-i ..., 0
§ §' J.o 0 0 Q) CIS ..., 10 ori ~
..., ..., J.o bil .!3 CIS C1I ori .g 0 >< 0 u cd Q) rl 0 0 s:::
§ J.o Q) CIS ~ rl ..-i a Q) N N ~ 10 b.O "0 rl C/l r... s::: b.O :z: :z:
B I 1 + - -)(. - 262
2 + - - '¥'---+ - 5, 153, 19), 191, 21), 2ao, 298, )02, )$5, 384
+ - -
3 + - - l5
4 + - -
+ - -
+ - -
+ - -
+ - -
+ - -
+ - -
+ - -
+ - -
+ - -
+ - -
+ - -
~ ~ ~ - - 99, 200, 257
i - - - + 201 yellow
- - - - - 63
+ - - 203, 198
- - 66, 89, 95
+ - ~ 129
- - + - + 364 flesh colored
- - - + - 2, 313, 314
+ + - )00, 341
i - - - - 151, 204, 429
@ - - + - 55, 56, 57, 58, 59, 111, 112, 113, 123, 141, 142, 14), 149, 187, 189, 190, 208
~ - + + - 1, 3, 122, 144, 145, 147, 148, 150, 151, 156, 161, 188, 191, 192, 195, 196, 202, 209, 210, 217, 218, 417, 418, 419, 420, 421, 422, 423, 424, 425, 426, 427, 428
~ - + - - 146
9 - + - - 242
contd. pigmtt 130
~ Q) 133 r4 .. 0 Q) -g ~ r4 II)
~ .g r4 M Q) lIS 0 Q) ~ II) Q) Q) rl Q) ;i rl ~ ] °8 0 til til rl ~ 0
§' M 0 0 Q) lIS ~ til ~
~ ~ ~ ~ M bD !l ~ ro ~ § 0 >< u 0 ell Q) rl 0
M Q) ro ~ q ~ ~ Q) (II
~ II) bD "U rl II) ~ bD ::r::
5 + - 219
+ - + 360 yellow
+ 0 263, 277
+ + -377
+ {j + + - 61
II 1 + + + + 318 yellow
+ +
*: + - - - 283
, + + - - - - 246 r 177 + + - - ~
+ + + + ~ - 176
2 + + ~+ + + 248
3 + - + C>o- + - - - - 443, 444, 272
4 + + 0 + - 363
III 1 + + ~- + - 154
2 + + - 155, 295, 296, 312
+ + 6 + - 284, 308, 309, 310, 311, 259
+ + {j + 266
3 + + - 11, 16, 24, 130, 159, 410, 411, 432
+ + - 0 + - 168
+ + + 10
IV 1 + + + ~{j - + - 375
2 + + + 0-- - 307
I"""'"
Table II contd. pigm't from pa ~e 130
§ I» 134 ...-t .. .., .g .,-i r-i U)
+' .g r-i M Q) Q\ 0 Q)
+' U) Q) Q) r-i Q) .~ ...-t U)
~ .,-i 0 It) U) ...-t +' 0 ~
§ §' M 0 0 Q) I'll +' U) ...t .., +' +' M ;t !l Q\ I'll ~ .n -§ 0 >< 0 0 Q) ...-t 0 0
§ M Q) Q\ ;::$ ...-t ...t S Q) N N
~ U) QD 1:1 ...-t U) fH ~ bO ::z:: ::z::
C I f+ - peri 387, 401
f+ - mono 306, 324, 333, 409
f+ - - 414
II f+ + peri 383
f+ f+ - 436, 437
f+ f+ mono 412, 413
III f(j 174, 224
f~ 228
f~ - peri 254, 271, 388, 389, 390, 398, 405
f~ - mono 250, 251, 252, 253, 255, 289, 292, 320, 335, 343, 349, 353
IV" ftf + peri 304, 344, 433
V f(j f~ peri 406
f~ f~ - 401
Note For brevity reactions which were recorded as nc in Table II appear as - in Table III.
L
DISCUSSION AND CONCLUSIONS
Of the var10u oharacteriat.loa vh1ch haft bMrl ued in bacterial
taxon_ BOMB 11ft undoubtedl1' aore bu1oa.ll7 iIIport.aftt tIutn otherS. No
.1n,J,e cbaracteriatic seems 1mv1abla but sa. appear .... var1abl.e or un
,table than othel'S. Pl'act1eal col181deratlone haw larcel7 d10tated the
character1at.1oa wh1ch are choMn to separate bacteria into YU"ioua iJ"OUp.h
The enteric ,..oup 18 .. par.ted b7 t.be1r aetion on laot.o.. ..... tb1a a' one
t.ime ..... d to oonelate nth the pr8ftll1nc idea of vlJat oanaUtut.ed •
pathocen or BOnpatbopn. The diaOO"fV,Y ot patbopn1c lano .. t .... ntera
has tended to a1n1id.. \be ftlue of tb1a nacUorl. The aloll aooept.ance ot
pat.b.opn1c acmlacto .. t81"1G8nt.tng bacUll is 181' .. 17 due to the belW t.bat all
1aot.oa. t.ra __ n _re nonpatllo&eD1c. Bactel'iolo,au DOW ..... all7 &aree
t.hat patholenic1\7 1. not a .table charaoteriatic and should occuP1 a 141.8
dominant role 1n taxonoaio .".tau. Baoter1oloa1au"t. d1aaoc1ate tbauel
trOll the diotatea ot speo1al1aed &J"Ol.IPa auob .... dloSne, 1B:!u.aVJ' and
&gricul tUN and bu1l.d a taxonca:f.o 8,.,. ba .. d upon biolog10al afflnitie ••
An at.,,, haa been III&de ben to emute the taxoJlOlllic iaportance
or various charaater1atics. The present d1aouulon 1a concvnad with crMsati .. red8 which are able to 11"011 1n simple peptone or ,.. .. t extract. in the
II
136 presence ot air.
CARBOHYDRATE UTILIZATION
Man.Y t,1P88 ot ada haft been c:le1'1aed ard are in .e tor the
detection of reaidual acidity trOlil the _tabolla of carbob,Jdratea. The
usual bual l18diwa tor the Enterobacteriaceae couieta of 1$ peptone, O.s%
sodium chloride, with or without 0.1 to 0.)% beet extnot. So.t1aM)'HIft
extraot 18 included in the tol".llUl.ae for the aore ta8t1dioua orpn1au.
Various iM1oatora ..... been aployed. titlmlB or Andrade's indioator wu
used during early t_8. Now one encounMn more trequentlT a eulphoDphalein
indioator such aa 'bromcrellol purple, phenol red alld brcattl9'aol blue. titaua
and Andrade t II indioatora haw not. witbatood tha teat ot t:iM tor they do not
gi .... aocurate ruult.a in t.eru or b;ydrcgen ion concentration.
Scae peeudoacnada produce a transient surtace chance of indicator to
the aoid oolor in 8QM ot the aboTe media. Howver, lIlOat media nov in WI8
8how no suoh changes or, at 'best, only ftl"Y' alight traui.nt obange. ot
reaction. ,
Man7 ".porte in 'the literature, as well as the author'. frequent
ob8enation, indicates that the CO!'I1poaition ot the bual aed1UDl contain1ns
organic n1:t.l"opn U7 greatly 1l:ttluence the results obtained in a atl.ldT or
oarbob1dnte deptadation b:r orpn,1au oonoenwd .....
P •• udcwmas Pl!Cl!!!", studied b7 DIB~ (~) in • Mdiua con'\a1n
ing 2% peptone, M/20 phoaphate ani U glucose, utUiled all the sua" but
did not produce an acid reaction. Sean arYl Gourley J.!928) tolBld that when
Pseudcaonaa aeru..ainoaa was P"O'WJl in 0.1 to S% peptone vater. it 'broUght &bout
1)1
a pH ot 8.8 in 6 days at whioh pomt the reaction remained stationery. The ..
authOn found, ae did Elrod and Braun ~l~). \bat the ni tragen _tabo1i_
resulted in the aocumulation ot alkaline aubatanaea surticifIDt to keep the
acid products ot sugar _tabo1i .. neutralised. Th1a vas not true in Md.ia
containing ....u quantitie. ot peptone, tor her., a arlee<! r18e in acidity
resulted.
produced acid end products trom various carbobydra tea. 5al.m and Ltnd.I
(~Ml studied what they called termentation or P •• udOllCftU on an OJ"&an1c
nitrogen aediwa (peptom 1~, beet extract O.l~, sodiuza chlori4e O.S~ with
phenol red indioator) in cOIJlpU'ison to a synthetic modiU8l conta1l11na the
carbob7drate. He concluded, a8 did Bender am Levine (191$1), Stein (1942), --_.-,._- - ._---stein at ala (1942), Sender (1942) and tiu (19S2) that P.eud014On&8 t4ll'lleftta-
- - - <;;.. ,~-> - "~'--"
tion can beat be studied 1n .. synthetic aed1wn. Liu (1952) vas ot the
opinion that the ta:Uure to produce an acid reaction in peptone-ce.J"boh7d;nte
broth 18 due to the preference ot the orcan1ar& to utili •• l"lPtone as a
aourco ot carbon .s well .s the source ot ni tropn. The "stal ta at
Paeudcaonu carbo~ate studies haft been reported b7 UD1' other workers
including Jordan (1903), Saleen and ftark (1943), OabT (1946) am Munos et
ale (1949). -Conn (1942) treated th18 topic ot fenaentation tbeont.1cal.l7. He
stated there i8 no question ot t.nnntati .... abUiW when an OJ"pni,ul produces
acid in a peptone aaediua. Howwr J it no aoid 18 produced the orlani_ 'lU'1'
stm be a t .... nter. Actually it uy be (1) unable to \18. the eupr, (2)
Us
reaction to the aediua except CO.-
It .. found ftI7 eU'ly in the e\wV that III8I\Y ot tl1t 01"ga.rdaB
in tbe collect1oft vould not grow well 10 .. ainal. s1Jltbet1c med1ua devis.d.
An attempt vas IUd. to dnue a peptone-~te .-4i_ vh10b would
'UPport sood growth and still De capaDl.e ot 1nd1oat1Dc acid end products. Tlw
organic aediua derued baa theadvant&ce of 00IIpU"iD& aU t.he oqmUu on
t.b8 .ame baa i.,. 'the orgaDi8lU respond _11 with growth wbather the reaction
under ob •• nat1oD .. positi,. or n.atift. Vera (USO) pointed out the care
one .1W8t obaerw in choosing constituents tor aed1a sinoe t __ mabl •
• ubst.ances U'8 contaiDed in som.e pept.orae., 78 •• t autoly.ates and beet extract.
The ettect ot ,....t extract, niuatN, phosphatee, agar, peptone arid dextro ..
concentration o.n the change ot indiaat.o:r in carbohydrate Iledia .. inftstiaat
ed. The tol101dng conol.ueiona are baaed on this study with the nontena.entina
&roup ot orpn1eu.
1. As the peptone ooncentration .. increased trOll 0.5 to S!& the
final aoidi V be.. 1noreuingl7 leee. Pananatic digests ot 801' bean au1
and gelat.in vve le •• aat1atact.ol7 tor incorporation in carboh;rdrate Mdia.
Tbe aOl peptone oontained substances micb the orpniau were able to con_"
to acid end pl"Oduna. Oxidet!.,. acidiv vaa in JWV' 1natancea cOllPletely
.sked wban t.r7Ptio dips' ot gelat.in was uad.
2. Ntt ... :t... when reduced. to n1 vi tee 01" ni tI"ogen incn&aed the
a.l.kalinit7 ot the aKluIIl.
3. Phosphatee atabU1aed tbe pH.
4. Yeut extract had an .ttect s1ll11ar to that of pepwne. It
contains 8ubstances which 'tI.&'1' be ooaftrtAd to acid em produots by the
139
oxidizers.
S. The optimwu concentrat.ion ot 4 ......... and otbar car'boh7dratu
is abOllt 1%. A concentration ot 0.', destroH OJ" le .. was det1nite17
interior to U dextrose. A dextrose concentrat1on ot l% or hi.her part1all7
inhibited the iJ"Owt.h ot 80M ot the orgarrl.ae tesW.
6. Alar at about o. ~ prevented oonvection o1U7ents and aided in
intenaif)i.ng and stabU1a1na the 1n:ilcator color eM ...
Of the -1\1 OOAbinat1ona tried 'tbe toraula below appeved JI08t
aat1afaotor,y.
pancreatic digest ot casein (Bacto-G.a1tone) 0.5J a~ 0.2 ~te (Seita fUtered) 1.0 final pH 7.1
Uhen those orglllnilmas which produced a surface aci4i ty trca clesV'oN
are laT8red wi.th petrolatum there is g8nanll.7 no growth or &Oid prod.uct;1on.
This 18 8tl'ik1n~ ditt.rent trOll the bahaYior of aRIbers of tbe taadq
Enterobacteriao.... The Etttel"obacneriaceae Il"OV well and produce a h1&h
aciditY' under the petrolatua. Baaed on t.lWI teo.bD.1que two types of
oarboh7drate metabol1a. 'IIA:f be d1st.inguiabed nueq, (1) aerobic (ox1d&t1on)
and (2) aserob1c (!.,mentation). a.striction of the wm ttteJ'llj&ntat1on"
to anurob1,c :18\&1>011_ of carbohydrates i8 in aocordance with the ori&1nal
Wlap ot Pa8teur. \ibUe studying carboh:(drat.e ut1l1aation by ,...8t., Pasteur
discovered there WU'e {'ants ot lite which do not. require ataoapbario oqpn,
and stated, "La termentat1nn eat 1. vie sana air". '.,..ntation then, in .... , ............ - ............................... its original meani.D.g, 18 a proce.. wbereb,. the oqan1aJl, in the abe.nee ot
aa, obtains energy troll earbohydl"8tee.
L
lhO
Table IV CODlpar •• erowth and WioatoJ' cbMp8 or nonoxjdi.el"ll,
oxid1 •• ra and t ... nter8 on tbe dexVcta. --.:s..oUd Mdi_ desvibed. The
aediwa waa inoculated in duplicate with OM of t.be tubes layeHd. vit.h
petrolatUll.
The nontermantel"ll an uMble to pow in the dax.troM-peptone Jll8d1ua
when air 18 excluded with a petrolatUII layer. ifOW'9W, the renantera grow
rlgorouelJ in • 8Wlar enriro~nt. The .. obaerw.tiona OOf'Nlate with the
indicator chan,... The l'umtemente:"'8, being unable to ptow, do not. ohange the
indicator. wh1le the renan'ten p~ change the indicator yellow eYen 1n
the abaence or air.
Nontel'llltmtent can be recognised by indioator cbanpa at tl1e .urface
ot the Md1_ Qp08ed tn "..be alr. Tn.y are diYided into two ,roupe, tile (A) ~AM. : ± " ~-.4-c
nonox1dlsen and (B) oxM~ .. r'. The nonox1di.fn"eA produce a ,.11011 svla.
color change which alt&r 1&-1 MfII "':1' extend to the bottoa ot the tube.
The tenllenWN quic1rlr prodl~_ acidity t.hroughout the ent11"e tube in
the pre •• nee 01 air. j\l$\ a. wu ob ...... d under petrolatUlle ramil:tatift
anaerobu can now be detirted moM detinitively U orlani_ whioh can p-ov
vell with ~g.n end can grow in the -baence ot ~,. proYided tv-Dtable
C8l"bob7drates ... present which supply enerO' tor arowth- 'Wbeneftr the teN
tacultat1ft anaerobe 18 used 'tM compound or _diUII which allow the orga:niut
to VOW in the absence or Idr 8h~uld be sttlted. It. appear. that aU the
organiaau in the tudly EntArobaoteroiaceae t.runt dext.roee undw pet.rolatua
ant therefore U'e tacultatift.
Baotel"U lIay then be •• parated into three units on tbe bub ot
aerobio OJ" anaerobio _tabol1a ot du •
I"""""
., TABLE IV
COMPARISON OF NONOXIDIZERS, OXIDIZERS AND FERMENTERS ON DEXTROSE MEDIUM
I""'" dextrose type or with dextrose
[petrola tum J.4
dextrose ~a a .,
rg~ ~ .Q
o~ .Q utilization as as
~ ... 0 ~ ... 0 a I) 0 ..... 0 .....
~ 0 r-f't:l 0 '(j]
~ ... o ~ b, bO 0..-1 0 .....
12 A1ca1i~enes sp. - nc + It> surf
46 A1ca1isenes bronchicanis nonoxidizers - nc + b surf
125 Pseudo.oTlaS sp. - nc + ib surf
2 Pseudomonas sp. - nc + ~ surf
176 Agrobacterium radiobacter - nc + ~ surf 0?Cidizers
218 Pseudomonas sp. - nc + r;t surf
410 Bacterium anitratum 5w2 - nc + ~surf
412 Vibrio comma + y + y
25) Aeromonas liquefaciens + y + Y
406 Escherichia coli + y + y - fermenters 407 Aerobacter aerogenes + y + y
414 Shi~11a Earadlsenteriae IV + Y + Y
387 Salmonella typhosa + y + y
KEY:
b surf deep blue surface, later becomes blue throughout the tube y surf yellow surface (oxidation) y yellow throughout the tube before 48 hours (fermentation)
" 141
B. Orld1181'8 a. .c:ae PeeudOllOn&S b. XanthOJlOnaa o. ffi"lOfiiiGi:!iUII d. 1£!&oWlua
a. S.laoMU.
:: Ea d. Xii=O&iCt.er •• le&r1061& t. '!.brIo 00lIIIIa g. X.J"CIIIIOD&8
. Orpn1a_ in U. P.eudol&oDaa-Al!4!e9!~ group are pnerall7 mable
to grow 1ft the aba4Inoe of air eftn it a utUlsabl. carbohJ'd;rat.e 1 • .,...._t in the aedt., hence should be reterm to aa aerobio. Selle ot the orpn1e_
in th1l croup can grow anaerobioally it n1 tn.. or e:JaU.,. eoaapouada are
pneent whioh oan accept hydro,. ~ an ox1d1sabl.e aub.trata. Sola
orCanieu in the 8I"oup obtain energy anald"Obieall7 tl"Oll aa1Do Hids and auat
be clueU1.ed as tacultati .... in thie reepect. The .. haft not Deen atlld1ed
in d.taU.
Frca 'table IV 1. t appears that the pnu A.eraaonae 11 plve1.o1oc1.oall.7
unrelated to the genua PeeudCllonaa, and Bacterlua ani.atva 18 pbp101ogioall7
unrelated to the Enterobaoteria.... It would be wU to note too that
V1DrI0 001IIII& 18 oapable of ter_DUng davoa. .. d.o the Enterobaoteriaceae
hence ahould not be considered aerobic u 18 so otten done. It grove
14)
anaerobically in a peptone _diu when dextl"oae 1. pre •• nt. Physiologically
Vibrio comma appears to be out ot place in the taa1l.y' Peeudoaonadaoe.e where -the aerobic oxidizers u ~seudOl&ona •• Xant.honaonas, Aoet.obaoter and SEir1llum
are clas.iliad. Vibrio coma shows a J)hTs1ologioal and Jlorphological
relationship to the genua ~_e_r_OMO_na_ ••
The ntaotiens recorded in Table IV were controlled bY'duplicate
inoculation in bual _diu (lacking added carbohydrate). One of the
duplicate. was layered with petrolatUf4. The results are recorded in Table
v. The colif'onu, typhoid and dy •• nte1'7 organisms prodUM JIIOderate
amounts ot alkali trOll peptone that is s1rlUar in det,TH to that produced by
Bacillus alcal!eenes and PseudOttonaa aocording to Kendall (191) am Uu
(19.52). Table V shows that the r .... n1'.er. p.roduce no change ot reaotion in
the open tube.. This probably means t.here were traces ot termentable sub
stances in the peptone which vera converted to aoid end products, hence
Illasking alkali production 1'rom peptone metabolu.. In the pre.ence ot air
these aoid products were either tvtber utU1sed to neutral end products or
were neutralised by the inoreas1ng quantity ot alkaline end products ot
peptone metabolu., possibly both OCCl11" s1m.ul:taneouely. Thi. wu :f'vther
brought out by the reaction Been when the 8ue _diUII va_ cO'VVed vi th
petrolatUDl to exclude air. Here the aoids produced were not neutralized or
utUized, but the acidity W&8 permanent. The growth in the closed tube vu
releti vel,. poor •• compared to that in the open tube. Probabl;r the only
energy avaUable anaerobicall.7 tor moat Enterobacteriaceae is the fermentable
substances in the peptone. The peptone urxler petrolatum did not serve a8
",-
.. TABLE V
CARBOHYDRATE BASAL MEDIUM CONTROL to-' basal basal
type of medium with medium petrolatum
... carbohydra te M M It Co-4 0 ~ 0
'§ O~ O~
utilization .£: C1I
~ :"d
~ M 0 M 0 ~ CD ;. OoM OoM
~ 0 'd'S 0 '3"8 :J:; Q, M ~ OoM btl o 'M
12 Alcali~enes spo - nc + b surf
46 Alcalisenes bronchicanis - nc + b surf
125 Pseudomonas spo - nc + b surf
2 Pseudomonas spo nonfermenters - nc + b surf
176 Agrobacterium radiobacter - nc + b surf
218 Pseudomonas sp 0 - nc + b surf
410 Bacterium anitratum 5W2 - nc + b surf
412 Vibrio conma - wk y + nc
253 Aeromonas liquefaciens - wk y + nc
406 Escherichia coli - wk y + nc -- fermenters 407 Aerobacter aero genes - wk y + nc
414 Shisella Earadlsenteriae IV - wky + nc
387 Salmonella tlEhosa - wk y + nc
KEY:
wk y weak yellow
-- 144
~.------------. l4$
an anera source, since growth was le.8 vithout air than with air. Most
Enterobaoteriaceae apparentl.y are taoultatift cm.l1' in the presence of a
t~n~ble o~~aw.
It M7 be noted heN that this teobaique, ot atudying peptone.
under pevolatwa with bl"ODlthyaol blue indicator, 18 all excellent •• \hod of
determ.Wng the pres.nce or tementable sub.tances in peptone ••
MORPHOLOGY I
Consideration here i8 l1m1ted to bacteria wbich are araa negatift
and rod shaped. Both ot these cbancteriatica IfI!I.T be indefinite and at times
it appears impossible to cassity an OI"lam. aa a rod or a cooem8, a. gr_
negat! .... or gr. poslti.,.. /fo SUSle.tlona on this probl •• are presented and
!ortunatctlT it is not vel'7 common. The 81 .. ot the rod 18 not a vel7
aign1tlcant characteristic and ls subjeot to oonsiderable variation in
different Md1a and d1fterent growth phas.a. The shape ot the :rod 18 BO_What
1101"8 baportant, particularly the 0\U'"ft ture.
The pronounced OUl"'f'&tUN round in the spirochaete. and typical
spirilla appears to be ot taxonona1c iJaportance. Sli&ht curvature IIIWJt be
treated cautioualy. SCM OI"Canias show 8OIJI8 cUZ'fttun in certain _dia but
no ourftture in other lIledta. A str1king exuaple i, Vibrio 00IIII& which when
cultivated in the laboratory otten ~ocaes perfectly straigbt. A sligbt
CUl"ftture 18 alao typical ot the lophotrioboua I"oda but the cbarao1ierut,io
is not su.ttlolently pronounced to be 1IlOJ"e t.ban ot .1nor taxoncalc UIportance.
Considered togetbar wit.h napllat1on, the Ddno:r oU1"'f'&~u:re help. to indicate
posslble relationship ••
. ,
~-------------. 146
It one exaadnes several species ot the COIIWOnly' recognised spirilla
one finds that the llcellular" units are otten relatively short, slightly
curved rods. A comparison of the morphology ot Spil"illua virSini&n'W\l and the
lophotrichous Aloalijenes shows a striking s1m.Uar1t7 in nagellation and
body curvature. Lehmann and Neumann (1927) proposed the name Vibrio alkali
ienes tor the organism wh1ch nov appears to be morphologionll7 related to
the sp1rUla. Pronounced pleomorphism, with tendel107 to torm biaarre shapes
and long fUuwnt., does not appear to baft lI1.uch taxonOlaic 1Iaport.anoe in this
group. The morphological characteristic of major importance appears to be
flagella tion. The various nagallar types were described bY' Leitson (1951)
and haw been demonstrated by pl-..otora1crographs in the presentation of the
experimental data. The ir.cjor signif'icant categories are a8 tollova&
1. peritrichcus 2.. polar l'Ilul ti trichoua 3. lophotrichous 4. monotrichoU8 5. nontlagellated
Although the aboft arrangements are characteristic aM stable
properties of bacteria they are subject to changes. Laitson and Hugh (1953)
described an oreanism (RH 247) which mutated b'ca lophotrichous to per1tr1cho
with intermed1ar)" tonu haYing both types ot nagellar arrangemenu. Another
organla vas alao described which varied trOll aonot.richou8 to peritrichous
with intenae<iiary t,ypee haTing both kind. of flagella. J'ust how CQlUllOft th1a
ph-enomenon is cannot be stated at. pre.ent. but it. 18 probably .ate to say that
it is somewhat unusual. It does _pba81 .. hoveftr, the importance ot not
relying too much on art{ single characteristic in the classitication of
bacteria. The recognition of nonpolar fiagella on predOlI'11nantly polar
147 organisms was facU1tated 07 the aore Itcurl.,y'f' shape of the lateral flagella
as compared to the polar. It is pos8ible that with aOll&8 bacteria the flagella
of the mutants If!J1A.'1' have the sante shape, and loIould thus be JI.Ol"e diffioult to
recognize. An odd fiagellum whioh appears to COIIIle frODl the aide of a polar
type 'IIl&7 be a polar flagellum which baa been bent undv the ol'lanl8. The
formation of multicellular t1l.aments by a polar flagellate 18 another cause
for contusion, Oray and Smith (1950), LeU.on (1951 page 387 figurea 23 and
24). The filaments appear to be peri trichously fiagellated. The shorter
units, which are usually present in a~ culture, estab11sbes tbe flaaellat1on.
The shape of bacterial flagella is generall7 stable and character
istic. The usual curvature is like that. found. on orgam .. such u Salaonel1aJ
PseudO\llon&s and Proteus. This shape appears remarkably constant and &IV'
great deviation i.laMdiately DeCOMa apparent.. The curvature ot the tlagellum
however is not free f'roIn mutational change, Leifson (1951, page 381 figures
$ and 6). These studies indicate that the usual type ot flagella tor
Salmonella can mutate to the less stable more tightly curved flagella.
A strain ot physiological17 si1dl.ar' IIOnotriehoua organi8l1l8 (rut 339,
342 and )47) whioh had an unusual17 greater flagellar curvature was ieolated
from water. Theae flagella are 8'tirikingl;y dilterent from the usual
Pseudomonas types (a .. figure 5 under Experhlental Data).
The fiagella of the lophotrichoua type usually baa only one or two
curves. 1'hi8 group is very liistinct1ve in appearance a11.1 868m3 to torm a
natural group when considered together with the tend.ency· ot the bacterial
body to curve.
The number ot flagella, whether polar or nonpolar, appears by itself
148 to be of relatively minor taxonom1c ilaportance. More emphasis m.ay be t:iven
to the nwuber of flagella in the polar group than in the nonpolar. The
lophotrichous types usually have two or more flagella. The polar multi
trichous organisms also produce two or more flagella. ~h.n the polar multi
trichous group is considered together with its ~iological characteristics,
the nwnber of rla~ella a~p6ars to ha.e so~ taxonomic .alue.
A consideration ot the mutation from flagellated to non!lagellated
types, and flagellated motile to nagal.lated normotile, tria_r and Le1fson
(1952), ..... vuranted. The later type vas not round alllOrlg the nontermenters
studied am it is not very important from the taxonomic standpoint provided
the presence of !'lagella is not overlooked. Nonflagellated variants or mut
ants are fairly common and such Mutation occurs frequently when handling
cultures in the laboratory, particularly with tile peritrict~us types.
The organisms studied in this work which were found to have flagella
were motile. SOl'I18 of the ol'"gani8ll1s were nonn\otile at 3"fOC but were motUe
when incubated at lower temperatures.
Tittsler and 5andholzer (1936) have proposed the use or a stab
culture in semisolid agar tor demonstration oi.' motility'. Hettie organisms
show a diftuse aone of growth spreading rrca the liM ot inooulation, non
motile cul tur.. do not. This test is a good oheck on the hanging drop method
but mainly applicable to organisms which are able to grow facultatiYely as
the Enterobacteriaceae.
1 mOL PHorUC'HON
The production of indol from. tryptophane is of cOl'widel'able
149 but apparently not for the nonf'enrantere. All the nontermentere tatted gave
a negative indol teat.
i1ETHn IU~D
All the strains of nontermenter8 tested gaft a negative laethyl
red test.
VOG.r~S-PROSU UER
Approxiu:tely halt the ow. tva. in the collection were tested tor
acetylme~1carbinol. All nontermenter8 were negative.
CITRATE
The nontermenters gave both positive and negative reactione on
Sim.'1tOn's citrate agar. Typioal pigmented P.eudoraonas strains gaft an
unequivocal positive citrate test. Howeftr, Ulong other groaps the teat
vas often difficult to interpret. This was especially true ot the peri
triohous and nonmotile nonoxidi.era. The 01 tra te reaction 1s or partiro.,.
value in identifying nonp1gm.ented or weakly pigmented Pseudoaonas straw.
UREA
The hydrolys1s of urea appears to be ot considerable taxonomic
value. stuar(, van Stratum, and Rust1g1an (1945) reported that Proteua in a
strongly buttered med1um could be d1lterent1ated trQl. all other ItleIf1bera of
the enteric group by its action on urea. This is in agreement vi til Rust1gian
and stuart (1941 aM 1943) and Ferguson and Hook (1943>-
Organiau racei ved 8S Brucella abortus, Bruoella suia. Brucella
I!l!litensis, Brucella (or HaemophUua) bronohiaeptioa (20 strains) and
1$0
parapertussis baoilli (3 stra1na) law a po.iti .... urease test. Elder1ng and
Kendrick (19$2) reported that occasionally parapert.uaaia bacilli were
encountered whioh ga .... a negati.,.. urease tes"'. TM Ul'e&8e teat. alight serve
a8 an aid in the differentiation ot the peritr1choua respiratory' tract
organisms, bronofl1cania, frOll. other Alcalienes specie ••
Excluding the organi81U labled Brucella and broncbiseptica the
following 18 nonter.entera also gave a positive urea.e test in 48 hours.
number or strains peri tricboua S lophotrichous 1 monotrichous 1 nonmotile $
RH nwabera 29, 116, 22$, 227, 28) lSS 22, 259, 284, )08, )09, )10, 311 10, lS, )10, )7), 374
The ftlue ot the urea.. reaction for species differentiation ot
organisu concerned here, outside ot the peri trichous nonoxidi"ers, 1s not
yet clear. Ho .... , the reaotion may al.o "M'8 to separate a specie.
aJIlong the normaotU. nonoxidbers.
Strains ot bacteria sometimes are isolated which lique.t'y gelat.in at
first but lose this property upon oontinued oultivation. The rate and degre.
a! liquetaction 'IIItlI:1 be highly variable. Wi th sOIII.e strains the liquefaction
is barely detectable and various techniques give conflicting naul. ta. This
type ot teat is of limited value as a means ot ldetltT1ng baoteria •.
Lerine and Carpenter (192) pOinted out that the viscosity of
autoclaved gelatin change8 little tor two or three da18, but attar thia time
it incresses markedly tor a week to ten days. A ,elatinol1t1c organiSlll mAT
bring about liquefaction quite rapidly when inoculated into freshly prepared
~...-------------. I ISl : gelatin but more slowly if inoculated into the ... mediWfl ten days or t.wo
weeks old. 1'1ore of the gel in the latter instance must be trarustortned to
the sol state to make liquetaction evident. The •• worker. divided bacteria
into three groups with respect to their action on gelatina
1. Gelatin not hydrolyzed; no liquefaction.
2. Gelatin partially nydrolyzed and liquefied. Decomposition
slow. if' arq.
3. C-elatin more completely hTdroly"sed, arrl liqt18fied. DeCOJlpoai
ticn rapid. accompanied by marked increase in formol titration.
The tour gelatin media, described in 14aterials and Methods, vere
compared si.''I1ultaneously by the inoculation of seleoted organism... Table
VI shows that the detection ot digesticn of gelatin is a function of ~ime
and the Mdi ... eIlPI07ed. There is a wide range over which an organi_ can be
considered positive or negative, since there 18 little agreeaent as to time
of incuba ticn or what media should be employed. Frasier (1926) suggested
that gelatin stab liquetactlon teat baa not been given a prominent place
in classUication because ot its slowess and un.reliability. However he
suggested that this technique might. be used if only those organiaIu that
showed liquefaction atter a week ware oonsidered positive.
" 152
TABLE VI
GELATlN ,1~DIOM COMPARISON
days 2 5 7 14 21 28 49
.RH number medium nwaber
1 .. - - - - .. -57 2 - .. - - .. + + 3 .. - - - + + + 4 - -1 - - - + + + +
122 2 - - - + + + + 3 - - + + + + + 4 + +
1 - - - - - .. -10 2 - - - - + + + -) - - - .. + + +
1& - -HYDROOEN SULFIDE
Many indicator. have been suggested tor use in deteoting hydrogen
sulfide namel.T lead, iron, binw.th, nickel and cobalt, aU of which produce
a dark color or the sull'ide ot these _tala. To o'ftrcorae the toxic eftect
ot heavy metals in media it has been suggested that a test strip ot rUter
paper izrlpreanated with a .metalie salt be placed in the l'l1OUth of the tube
during incubation.
It is generally agreed that the paper strip method. i8 far .e
sensitive than many ot the tests which incorporate the toxic indicator system
in the lIledia. The value ot the paper strip method beCOlll8S limited since it
gives litO many positive reactions, for eftn such organisu ae Shigella
1Y".senteriae give a positive wst. The test is influenced b7 the nature of
L
153 the peptone, and such Bubstances &a c;ratine and thiosulfate Which may
increase the number ot positive reactions.
All the organisms studied were tested on JCligler'8 iron a,ar. The
non£ermentera are generally unable to grow in the agar stab hence are unable
to produce blackening in the butt. Some pseudomonada on prolonged incubation
may show blackening in the line of growth on the .. lant. 1'h1a 18 confirmed by
f\Ozansky !! !!:. (1950). Pulrlrent1 (19h9) alao obaernd HaS production b7
l!!. Plooianeum. Howver, it was found that -any typic.al pigmented PseudOllOnas
strains do not give a p081ti~ H2S test even with the paper strip technique.
The bJdrogen sulfide test seems of little taxonomic value tor d1tferentiation
ot the majorit.y ot nontermenters.
Two unusual monotrichous organ1aRl8, rut 231 and )64, produced H~S
in the butt ot Kligler's iron agar. Both of these nontementers probably
obtain energy tor anaerobic growth !rom andno acids.
NITRATE TO GAS
The reduction ot nitrate has been widely used as a differential teat
Uitrate uy be decOIIlpo.ed beyond the nitrite stage and in lUl'.'\Y instanoes it is
converted to molecular nitrogen. other reduction products are apparently
produced, but nitrites am nitrogen gas are moat read1lT detected. The
reduction of nitrates to nitrogen gas by bacteria ia called denitr1tication.
:-ficrobial. den1tr1.f'ication apparently is a process equivalent to aerobic
oxidation with 'nitrate replacing oxygen as the ultimate oxidizing agent. A
concentration of 0.2 to 0.5% nitrate in the mediua is satisfactory for
obse~~ evolution ot gas. The gas may be trapped in a Durham tubo in
154 liquid medium. 'l'he formation of nitrogen bubbles in seDd.aolid Mdiwa ie not
alvays reliable. Experi.ll18nts conducted by Wen and van Hiel (1952) with the
denitrity'ing bacterium. PseudOlllOna. stutser1 ehowd that neither udno acids,
amide. nor nit;:'ogenous constituents or bacteria in the Mdi_ contribute to
the nitrogen evolved during reduction of nitrate.
Nitrate reduction to gas was not frequentl.7 encountered 8l1IOng the
nonoxidiz.rs. The nonoxidizing denitriliers were RH 12 and 13 (peritrichoUlt)
and RH 330 (monotrichous).
AMong the dextrose oxidizers, nitrate reduotion to gas 1s of
taxonomic importance in the monotrichous group- Ni trogen gas produotion is
particularly valuable in identifying pigmented pseudonaonads &a well as
apyoc1anogenic and afluorescigenic strains.
NI'l'RA TE TO NITRITE
The lUteen lIlin';lt8 period ot incubation as reCOllWended by Brough
(1950) tor the determination of nitrate reduction does not SMIIl adequate when
studying nonterm.enting o1"gani8I'U. An extention of the time to one hour was
far more adequate since it gave more positive and rever doubtful reactions.
ANAEROBIC GRO~~H WITH NITRATE
Many organ18lll8 are capable or oxidizing organic substances in the
absence of molecular oxygen by 8~,u1taneously reducing nitrate. Most
compounds which serve a8 an oxidizable substrate under aerobic conditions
are suitable enera sources in nitrate containing mediwn in the absence of
air.
Nitrate in a O.l'~ concentration supports excellant anaerobic growth
155 in peptone water tor those aerobic orlanu. which convert nitrate to gaa.
The anaerobic growth ot denitritiars 1n the pre.ence of nitrate was JIlO" rapid and profuse than that .een in media without nitrate expo.ed to the air.
The anaerobic growth with nitrate i. generall,y aoccapard.ed b7 copious gas
production. Thia i. well exemplified b7 oraani_ RH 12 and 13. With these
Ol"ganistu there i8 no visible anaerobic growth in peptone vater in 18 hours
but anaerobic crovth vi til nitrate i. he.rier than that ... n in nitrate tre.
peptone exposed to the air.
5a. ot tbt organisu which conftrt n1 trate only aa far .. nitr1 t.e
show L'I&8robic poovth in the preeence or nitrate. This growth boweftr 18
slover aDl1ess abundant than that described tor true denitr1t1en. Thie
type ot reaot1on i. frequently .een ~ the lophotrichous C1"aani8.llUh
PIGMENTATION
P1pent produotion i8 ot coneiderable diagnostic importance. The
benefit derived tra. pigments b7 the organisms i. not clear with the exception
of those pigments which Bre •• aential to photosyntheaj_s. The placing of
axcesaiTa emphasie on pigmentation in taxo~ otten rai.e. a question ot
proper n01ll8nclature tor nonpigftlented straina derived trom. pigmented orpnisu.
Pigment production is not i.ntrequentl;r dependent upon the compos! tion ot the
mediua, tAbperature of incubation and oxygen. Aaong the ta1ntl7 pipented
organisms color characterization i8 ditficult. The factors e.8ential tor
pigment production V'lU"ie8 with the orcan18a am no pneral aediua can 'be
deVised which 1IUl be opt1arlum in all O&Be •• · These factor. indioa te the
disadvantages entailed in emphasising pigmentation a. an identit,ying character
lS6
i,tie of some groups ot' organiaJu.
A study of the water soluble pip.enta of PaeudOlllOnas baa been
made by Leifaon (unpublished) and was .found to btt conatderablT enhanced in
raany cu •• by adding O.Ol~ lUnganes8 salts to a a.uolid nutrient agal'. The
magnesium salta were 1 ... etfect1.,. though alao stiJllulat0Z7. :1oV8ftr. even
with t..he manganeae mediUlll a nUlilber ot strains having all the other character
lstics of Pseudomonas aeNinosa stUl failed to produce a water soluble
pigment. Several papers haft recentJ.7 been published on the P.eudomonas
pigments, Sel.en and stark (194). Burton !! ale (1947 am 1946), King!!!!:..
(1948)J lio.ansley !! !!!. (19S0), Harris (19$0), Haynes (19$1), Hellinger
(19$1), Aft.ltu (19$1), and Totter and lio.aley (1953).
Roz&nsky at ale (19$0), who apparently were working with highly --pigru.ented strains of pseudOlllonads, observed pigment to be abundantly produced
on toetner' 8 J'Ilediwa. Many times Loefnar',. medillIll showed no pigment, when
obviously pipent.ed strain. were employed. In three instance. RH 167, 186 am
189 pigmentation va. obsened on Loeffler'. media but not on IAl'II' other
_diUlll tried. toetner's l18diulll has an 1nherant d1aadvantap since it 18 otten
rapidly liquefied betore pigment develops on the slant.
MILK AGAR CLEARING . Clearing ot milk agar is perhaps more sensi tift and reliable than
litmus mUle as an :1.ndication ot proteol1's1a. T here is no assurance 1n this
test that casein is cleared only by prot.eOlT8U. Clearing ot the agar was
observed atter 2h and h8 h~urs.
157
BLOOD AGAR
Henriksen (1937) drew attention to heJlOl7tic svains ot nonmotile
Alcaligenes. The Alcl'lliJe f18, !!!,mP,ll8~ de8cribed produced a soluble,
antigenic hemolysin which was inactivated at SOOc in i: hour. Se'VVal strains
ot nonmotile organismu called Aloalifen8s £:!!oll8ans wre receiftd trOlll
Henriksen and Oeding in Norway (rui 10, 164, 165, 166, 161, 168, 169 and. 170).
These eight strains produced a beta helllOl.7si8 about subsurface colonies, u
well .s hemolytic lones about the surtace colonies. RH 10 and 168 oxidized
carbohydrates, while the others were nonoxidizen. In addit.ion to the above
strains rive other organise with the same type of 11_01781s were encountered
in the study. The various beta hemolytic strains are listed in Table VII
with some ot their characteriStics.
Deep colonies ot all the remaining organiSM studied ... " nou
hemolYtic with the following exceptions. The peritrichoue organisms RH 135 a
22) produced an alpha hemolysis. The monotrichous organisu RH 188 and 231
produced an alpha hemolysis. Alpha hemo17sis was also ob.er'Wd &IIong the
nODftotlle organisms.
~
.,
-TABLE VII
BETA HEMOLYTIC NONFElli~NTING GRAM NEGATIVE RODS
. {/J S CIS ..-4
J.. ~ II 1:1 CIS II
1 ~ .-t II .~ QJ III II ,.; ~ +l 0 ..... u CD CII +I CIS !J CD ~ tID M CIS !J u CIS +l ,.; >< ga CD 0 q ..... CD ..... II M II U tID I:: 1:1
10 Alcali~enes hemollsans spinal fluid - + - - +
164 Alcali~nes hemollsans throat - + + .. -165 Alcal~enes helllollsans throat - + + - -166 Alcali~enes hemollsans throat - + + - -161 Alcaliienes hemollsans throat - + + - -168 Alcalisenes hemollsans throat - + + - +
169 Alcalisenes hemollsans throat - + + - -110 Alcalisenes hemollsans throat - + + - -186 stool - - - - -194 stool - + - - -191 stool polar lIlulti + - - +
210 blood lIlono + + i + .
216 Alka11senes faecal 18 stool - + + - -
-158
1S9
LITMUS MILK -, Milk hu been used tor the .t.ud;r ot ld.croorganulU since the
beginning of bacteriology. The useMne •• or JIUk wu increased bY' the addi
tion of' Ii suitable il'1'iicat.or su.ch &s litmus. Litmus has an adftl'\tage over
mal17 indicators for it is reduced 0Y' some bacteria. It 18, howeftr, not
too aons1tiv. to slight changes of reaction.
As earl,. as 1889 Petrusc:h.ky recognised the li.utations ot
ditterentiat1ng bacteria on the oasis of milk reactions. He used a litmus
vhey _dium which was tree trom casein for tho d1tf'erentiation of' 9aoUlus . taecalis alOalil!n8B and the typhoid bacillus.
A,-era and Johnson (1913) and Ford (1903) observed that in 11taua
milk often enough alkali was produced by organilu to diesol ft the ca.ein.
Thi. happened on prolonged periods of incubation. Ayer8 _t al. (1919) belaTed --the alkaline reaction to be due al..Iaost entire17 to formation ot alkaline
carbonate. troJa the oxidation or the salts of' organic aCids, presUlU.b17 citric
acid. This is in agreement with the earlier lIOrk ot Seiffert and W~
(1912).
At least S or 6 reactions mq occur in litlnus nd.l.k. TheY' are
dextrose, lactoM and citrate metabolism, reduction of 11_U8, caseolys1.,
rennet and scid coagulation ot casein. Simpler _dia haft been deviled tor
the study ot carboh7drate metabolism without contusing this reantion with
other activities which also may be taking place in the same tube
simultaneous17·
The AlcaliJ!Ms bronohicani. strains and other nontermenters
reduce an intense alkaline reactio
160 appearance ot the _diu was not \1I'Jl.1ke that s .. n with proteolytic organiau.
'the 11taus aUk rea.ction does not ... to add to the ident.1t71ng
cbaracter1atic8 ot the orga.nlSlU when it 18 under.tood that the alk411
production ariee. trona citrate utilisatloD or ni\ro,en _tabelia. SQU' ot
the.e reactton. were studied indi'ri.dual.l7 with a acre eu1t.abl. Md1wa.
DESOIYCHOLA TE
The photoaicrographa published by ruhllllUnll (1911) .owe that be
was able to iaolate only lophotrichous orprWuas from tecal _tv1al. LeUson
(19)S) stated he vas unable to isolate, troa fecal _t.er1&l .treaked on bUe
apr, the peritrichou8 Aloaligenes but lophotrichous organl8lU WN enoounter
ed. These reports se. to indicate that it 1. the lophotrichous -Al.-l!Jeneen
wh10h is enoountered 1n tecal .tudies wdng bUe udJa.
Growth on 0.1% desox,yobolate &aU" ... studied to tol"llUlate an ide.
ot the natural habitat ot nontenaenting or,ani ... ~ a •• WIling tailure to grow ..
an indication of nornnteatinal origin. The tollov1n.g generali.ations could
be made,
1. The lopbotricboua IfAlcal.yenes- grft weU.
2. The bronohloan1a straina Vew VW7 poorl1', if at all, however, 6 peritriChO"U. nonoxid1 •• n (M 12, 13, 222, 223 am 232) arew well.
3. P1pi8nted pseudomonads .... w well. Sa. ot the nonox1d.il1ng JaOnovichou8 or«anilHU did not II'OV.
OROUPD~O OF TiE ORGANlSKS
The orlani.. concerned in thi. treatl., tor Q'8tematio considc'a
tion. vere divided into three units on the buis ot abUity to un dexvo .. ,
Un1 t A noncxid1aera (120 orpni8118) Unit B ox1diaera(12) 0l"Ian1au) Unit C termenten (39 orpn1au)
Unit A •• further divided into t1 ... groupe.
Oroup 1 peri trichoua Group 2 polar mul \1 trichou Group 3 lopbotr1choU8 Group 4 monotriohous Group 5 nonmotile
Table III shows the separation ot the oraanis.. luted in Table II
into vieldT units, subuni\S and (p."Ol1p8, which will be considered 1ndiv1dua1.l7.
Group 1 ot Unit A va. compriHd chiefl7 of .. ,ard •• l"8oelwd
laOeled Brucella bronobieeetica or HaeaoeM,lua bronchi •• ptlou (the first
20 orlani ... listed in Table VIII). It va. obaened that 1I1thout ucept10n
all 20 strains labeled in thi. wq were aorpbologicall7 .S.11.... All ....
a poe1tlw un .... teet. Pb7alo10g1cal1y'the7 dUfered in their action on
nitrate. The charaoteri.tice of the 20 swaine va in ac:re-nt with the
delineation ot Brucella bronchis!pt1ca •• worked out '07 Elder1ng and
Kendrick (19$2).
The detailed obaraotar18tica ot the peritrichous nonoxidl.ers are
shown in TaOle VIII. In addition to the 20 broncbioan1. etra1ne, :3 other
or,anisM .... encountered which appear to belong to the bronch1can1a &rOUP
(the firet 23 organia.e listed in Table VIII). It aiaht be well to draw
special attention to the organism received labeled Banez-iUll aloal;enee,
RH 225, isolated troa t.he lungs ot a guinea p1&- 'lhu organiua has all the
morphololical and physiological charaoteriet1ca ot the bronchioanil strains.
but the citrate reaction is nepti.... Should wob an ol"aania be thrown out
ot tbe bronohicania group because ot the citrate reaction? Or,an18u RH 29
TABLE VIII PERITRICHOUS NOHOXIDIZERS
48, 49, 50, 51, 183, 184, 214
46, 47, 52, 53, 54, 140, 171, 172, 173, lBO, 181, 182, 232
29, 225, 227
338, 336
322 yellow water insoluble pigment
222
135, 223, 258, 376, 378
138, 260
12, 13
379, 380
60, 83, 1)6, 137
- - - - - - - - - - + - + - - - -
- - - ~ - - - - - ~ + + + - - - -
- - - - ~ - - - - - + + - - - - -
- - - - - - - - - - - ~ + - - +
- - - ~ - - - - - - - - + - - +
- - - - - - - - - - - + + - - ~
- - - - - - - - - - - - + - - - -
- - - - - - - - - - - + - - - - -
- - - - - - - ~ - - - ~ + - - - -
- - - - - + - - - - i + - - -
nc - - - - + - - - - - , + - - - -
TABLE IX
THE RELATIONSHIP OF NITRATE AND SOURCE OF ISOLATION AMONG 23 ALCALIGENES BROHCHICAHIS STRAINS
RH nUJIber source nitrate RH nUlllber source nitrate
29 + 172 guinea pig + 46 dog + 173 guinea pig + 47 dog + 180 + 48 pig - 181 hwaan + 49 pig - 182 murine + 50 pig - 183 canine -51 pig - 184 -52 dog + 21.4 dog -53 dog + 225 guinea pig + 54 dog + 227 human blood + 140 + 232 dog + 171 guinea pig +
16)
and 221 thougb of ditferent origin are "'17 s1JUlar to lUi 22S.
Siaons am loser oitrate reaotlona, tor the peritrichoua non
oxidizera. in -1lT instances were not a det1n1te positi.,. or napti... The ••
citrate reactiou le.ve MUch rOOIl tor interpretation, hence probab17 are not
a sound beau tor species difterentiation.
The relat.ionahip of nitzate reduction to the 80wee of iaolation
of theae orpn1eu is shown in Table IX. Oenerall7 at;roaina 1I0lated frail the
dog gaft a positi.,. nitrite teat whUe those trcm the pig ..... raepti ....
The vv1at1on in the oitrate and nitrate reaot.lone ahould probably
be coneidered as variationa within a apecies. The nitrate reaotion baa been
recognised b,. T0JTe7 am Rahe (1912-1913) as being variable aaong the
bronchicani •• tzaina. hence this variation within the speci •• 18 not a MW
departure in .,..tema tica.
There 18 li~tl. Justification tor the Hparat10n of ,aMra on the
baais or source of uolatlon. All nonoxld1s1ng peritr1ohous17 nagellated
oraa.n1alu raa:r be clas.ified as Aloal!Je,... It appears that the reaaon tor
not ooneider1n& broncb1oanu as belong1nc to the genua Al!!Heraea W&8 the
undue silnifioanoe g1 Yen to the source of isolation. Dv1.rlc the t1ae when
bronchioaaie waa tint being described it did not seea 10l10al 1.0 include
ol"lan:isu iaolat.ed tJoca the respirat0J7 tract in the ... pnus as Aloalipnea,
whioh vas erronaoual.7 thought to bave an uo1ua1w teoal bal:l1tat.. Furthermore
JtI8ftT baCt.erio1oli8't.1 at that U. col'UJidwed l.oUlus f.eoal1e alcal!lenea
to l)e polar rlaae1lated, Bergbaus (190$), Kluinko (1901), K'tIhnuann (1911),
Glasser am Hacbla (1911), Horn and Huber (19ll), and Baerthlein (1912).
The aouroee of uolat1cn of the perivichoua nonoxidizing OI'mll"l1. __
164 in Unit A Group 1, exclusive or tho .. urea positiTe organisms considered to
De Alcal1lel1la bronchicanis, are giftn in Table X.
TABLE X
SOrmCE ~' ISOLATION OF PERITRICHOUS NONOIIDIZEHS
RlJ nwrtben source reee1wd as
12 and 1) soU Aloali!ne. .p. 13> and 378 Aim .... t"08l18 1.38 row tdlk IlealI!; •• fuC08U. 222 tece. UaClir_ Alllnea 22) Sacterlua it nes 258 teces ~!in •• ape 260 variant ot 247 o percolana 322· aWe_ 3)6 and 3)8 stream 376 Alcal1pnea 8p.
*' produce. a water 1naoluble yellow pipent..
The author reCOllUMs that the OJ"laniau in Table X, sinoe they
haft the .... morphology and .Wlar pb;rs1oloQ, should be placed in the
s._ genu. togethor with bronchicarda in'eapeotive ot the .ource ot 1.ola't.1on.
The .. organisM witb difterent sources ot isolation, whether 1t be soU,
re.pirator.y tract or teeo., .hould 'not be placed in difterent Cenera unle ••
then exi.w taxonoadcall:r significant dift.rene... Su.oh \axOnoIdo dUterences
haTe not been demonstrated.
The genus Fla'YObacterium hu been created tor water insoluble
yellow perif.richoua nontermenters. orCan1a an 322, shown in Table Vill, by
t.be above definition belong. to tbe genua FlayoDactariUll1.. P1gR18ntatlon as
considered earlier i8 otten an unstable charaoteristic and nonp1.gmanted
variants are read1lY isolated from niuIllAntad .tr ... btJII_ A.1.da :f'J"ftItI "'1 ___ "6" lnn
165 the characteristics of RH )22, both anat.ca.1oa1 and plv'aio10gic&l, corre.pond
to the Aloa1;pnes faecalis strains .3)6 and ))8.
Organ.1sra 1m 260 vas part,1culU"17 1Dte .... t1Dc s1nce Leiteon and
Hugh (1953) reported it'8 isolation fro. 1Ul 261 wb10b had lopho'trioboue
nagalla. The lophotrichous organialll KH 261 .... :leo1ated rre. Clrl8l'daa au
247 vbich was desoribed as Vibrio P!!'col&D8 by MUdd and WarND (1923).
The second croup ot 11 orgm iau in Table VIII can becOII8 known
u Alca1!J!ne~ t.eoalia (lUi 338, )6, 322, 222, l)S, 223, as8, 376, )76, 1)8
and 260). Alca1!e!!s laecalis can be d.scribed to haft tbe toUov1zIa
character1eticsl Ora neeati ... , peritrioholl8l.y' aageUated, aapozoopnoua
aerobic rods. noea not oxidi •• or t .... nt carbohydrates. Indol, __ 1 red,
VO£ •• -Proak:auer and vea testa negat1Yfb liwates 1ltIl7 be reduced to nitrites
but not to gu. The citrate and plat1n react:1ona .,.. 'ft1"1able. 1be
hydrogen sulfide te.t 18 neg. tift on nigler's iron agar. The author 18 !lOt
aYers. to including yellow vater ineoluble p~nted organis .. as RH 322,
with the above characteristics in the species faecalia.
T'wo organ1as isolated trOll soil, RH 12 and 1), could. read1l.7 be
d.1atinguisbed trom. aU other per1v1choua nonoxidiaers since theY'reduced
nitrates to nitrogen gas. The name Alcalipnes den1trUac1.eNf, ap. new. is
proposed tor tbo .. true den1tr1tiera.
The six orgamau, RH 60, 6), 136, 137, )79, )80, luted at the
botte. of Table VII! are considered separatelr fora ~~e other peritricbou.
nonoxidi.era,. Thea. organiau wre unuaual in that t.he7 produced oxidative
aciditY' from qlo.8. It u:r not be out of place to speculate that all 6
strains vere of the same origin, since til. old.at culture, Rli 137, was de.
166 posi ted in the A_rican 'l'ype eul ture Collection 1J1 192$ and the or1a1n of the
· remaining strains is not certain. The l>h1's101oa of the orl.niau RH 60, 83.
136 and 137 appears ditfercnt tr.la.n the zaajorit,y ot the dextro .. nonoxidiHr8.
Interpretation of the dextrose reAction OT the teohnique ~plo7ed wae
difficult for the organisms did not produce the usual abundance ut allcali
e.haracter1stic of nonoxld1;.crtll but lett tbe reaction ot tho Mdiua uncbanged
or only very faintly acid.
The YBlidity or the meager description ot Bacillus l.ecalia J •
alcal±ienes lett b.1 Petru8chky can be doubted tor the following reaaonsa
(1) Peuuachky recorded only gas production trom carboh)"ctrate media studies.
(2) ?roof by Petruschky's contemporaries that atrains of Petruschk;r'a BaeUlu
i'aecalu Alcaligenes were cont.aJUnated with t1J)hoid bacUli, Pseudomonas and
other water insoluble pigmented organi_s. 5hould one aocept PetrWlchky' , 8
characteristics ot Bacillus faeca1its alca111enes without question ot validity,
one CL"1 jUltU'iablT'oubt that the charaoteristics reoorded by i'et.1:"ilsoll.lc;y vere
complete enough to re1dentit7 his organism. 'l'his 18 in agreement l>'1th Conn
(19hZ) s:l'QCe h~ too 'Was not certain that th1a organis. could be recognized
from tbe liescription gi van b7 Petru.schky.
Atter caretul consideration of the literature it appears that the
oilaracterutics ot B.aoUlus la.calis alcal!enes 68 given b7 Petruschlq (1889
and 1896) also fit t.~ description ot the hoaogeneoua group ot organis_ being
called Brucella broncnisept1ca. Torre7 and Rahe (1912-1913) recognized that
the Ii taws illk and potato reactions vere siail&r tor both bronchiseytio! am
BacUlas fe:acali! alkal1S6nea.
,,'USOA and Miles (I946), and SJUth and Conant (19,2), place the
167
£!'onehiaepUca organia in the pnue HaeaoJ!!1l!! b4toaun ot OOlllaOD ant1aene &.
found by Eldering and !(endr1ok (19)1 and 19)8), Huddle.on (1939), Floadort
!!!!!. (1941), Parfentjev !!!!:. (1947); common toxine aa found by Eurus
(19LtO) j a!1d the typo of "apirawrl" dia ..... produced in ani .. la and man ••
found by Brown (1926). Pathogenloity and antigen1clt7 ha .... a greater
taxonOll1o signiticance than l'lorpho1oa an1 phye101oQ' when b'r'onchi •• ptJ.ea is
placed in the genua. ~;l'l~.:.hUWJ.
Manom.etr1e studies, by MoCullough and Be.l (19S1) and Altenbenl
and Housewright (1952) haw cont1rMd earlier reports 'by McUp1na and Slaneta
(1928) and McCullough and Dick (1943) on cuoohydrate utUiAtion b7 at.raina
ot Brucella 4bortus, _Br .... u_oe ...... l ...... l_~ .!!!!! and Brucella .e1itensis. Usin& t.ba Md1wa
and technique defined above £itteen strains (Rli 42, 43, 44, 45, 67, 68, 69,
70, 71, 114, 115, 117, 118, 119 and 120) ct ~bortU8, !!!! and m.e11tena1a were
found to produce oxidative acidi~ trom carbohydrates. There exi.ts then a
.narked dltterence in the ph)rsiology of theae three Brucella species and tM
bronchiseptica strains, which do not utabolize carbobydrates. Thia
phTaiolog1cal dU'terence indicates that b:;onchue.etica should no longer be
retained in the genus Brucella in spite o.f the shdlarity ot the nucleoprotein.
Eldering and Kendrick (1952) realizing the antigenic relatio".ah1p
of the parapertussu bacillus to both HaeaoeUua ertunis and Brucella
bronchi.eptiea. ellJ)haoiaed that th ••• three organieu are rtdaf'ita in the genua
HHaOP!UUB and suggest that they 'be placed together in a ditterent genua.
Anderson (1952) oonfirmed the serological relationship 8lIIOni these three
organisms b,- ercss-ttbsorpt1on testa using 0 and K antigens and eron-neutral
isation testa with toxin-antitoxin. The.e orcaniams were identical with
~~--------------~ 168
regard t.o the theJ'lilOS table c011Jfl.on 0 antigen, and baemorrhagia toxin, whereas
each of them possesses its own specitic K antigen and additional oY8rlapping
K antigens.
It appears justifiable t~ rec~~r.d that bronohiseptica be placed,
as in Sergey's Manual ot tetermtnative naot8riolou, 4th edition (19)4j, in
the ,enus .U~aliiiene8 and becOf48 known 8S ~r:onohican1s. since this na.me has
priority. Consideration should be given to making bronohicanis Ferry (1910, ,
1911 and 1912) and M'GOyan (1911), A very ~oi8el1 described orlanis., the
type species of the ,enus .Alcal~ene8. there are no discrepancies w1th the
description of the probably unidentifiable ~acU1U8 taecal~s alcalijenes,
Petruschk,r and the description of bronchicanis.
UNIT A GROUP 2 -----'l'h1e polar Ilultitrichous group, Table II, show a clos8 relationship
to both Pseudomonas and Ianthomonas. The existing de.cription of the •• two
genera are quite vague to be certain which genus should be assigned. A
speci.s distinction in one of these genera is warranted for the 6 strains
not utilising dextrose. These organisms did not produce a water soluble
pigment out tour produced a weak or .trona; water inaoluo1e yellow pipent
whUe two produced no pipent. The resemblance to lanthoawJUl.$ 8uage8\8 1 teel!'
because ot the yellow pigment, however Xant..tlam.olUlas 18 descrioed u being
predOldnantly IlOnotrichows.
It 18 recommended th9.t the polar lIultitrichous dextrose nonoxidizers,
RH 294, 287, 430, 4)1, 229 and 2OS, become known a8 lantilolRona8 tllaltophilia
sp. nov. They can be described as having the following characteristics'
169
TABLE XI
POLAn AULTITRICHOtS DEXTROSE NONOXIDl.ZERS
294
287*
I - - - + + - - - -
~ - - + + - - - -
4)0, 4)1* - - - + + - - - -
- - + ~ - - +
~ - + - - - +
- - + - - ...
229 - - - + - - - - - - - - + - + - ~
205* ~ ~ ~ + - - - - - - - - + - + - -
162* nc - - + + - - - - - - - + - - - +
1 ftC - - + + ~ ~ - - - - - - ~ + - +
*produces Ii yellow water insoluble pigment
Gram negative, asporogenous, aerobic rods with polar multitriohous flagellation
Do not oxidize or ferment dextrose, lactose, sucrose, xylose or arabinos ••
1xidative acidity is produced trom ~ltose. iiannOS9 may or may not be ox1rl1z-
ed. Indol, methyl red, Voges-i'roskauer, citrate and urea teats are negative.
Nitrates m.q 0. reduced to nitrites but not to gas. Gelatin 18 generaJ.l7
liquefied. 'l'he hydrogen sulfide test is negati .... on Kligler's irOll agar. . .~.
yellow water insoluble pigaent r~ be produced. These organisms were isolated
.t.l-om stream and stool cultures. One etrain may ruW'8 boen isolated i'rom pleural
170
nuid.
Among the polar mul. ti tricboua orlanisu listed in Table Xl are
RH 7 and 162. The.e unusual organi8m.s ,aft no change ot indicator or
possibly a faint trace of acid in the dextrose med1u.. When either sodi~
chloride or phosphate uere incorporated in the same medium, organil5r4 RH 162
uruaistakably oxidized dextro8e. ,)rganislll !t.>! 7 could not be SilJli1arly
encouraged.
UN1T A GROUP..l
The lophotrichous nonoxidizers are listed in Table XlI. Two
strains, ill{ 274 and 439 .. of the photosynthetic organism Rhod081?1rUlUia rubrwa
were included in the table for comparative purposes. 1h$ remaining strains
in the tabl& are fairly hOdlogeneoua. This group has long been considered as
Ale.licenes t .. eal~. Leifson (1942 atd 19S1)we.s in agre8ll1ent with KuhnerlMfI
(1910 and 1911) for both pub1ishec! exclJllant photomicrographs of similar
organisms ot iecal origin. These organiBme were in general lu. sensitive to
bUe salta t.han the perit.richous Al~81!l.nes. They wen la.beled Vibrio
alcal1pnes by LehrWln and Neumann (1927) after the york of strecker (1917)
Nyberg (1935), his Group 3, and Sewri (1949) also eneountered Vibrio alcali
genes. The genus designation Vibrio, is unsuited for these organiRs s:tnce
they do not ferment dextrose.
S,eir1llU!ll virfinianum, llii 273, 1s included here Mong the lopho
trichous nonoxidizers since there is similar flagellar and 8011& tic llorpbology
H well H aim1lar physiology.
., 171
TABLE Xll
LOP'dJTRlCHOUS NONOXIDIZERS
t: , .,.,
I '! p.,
Ci • ~ f:') t) !4' ~ ~ 0 J t%
~ i 0 Ii) 8 Ii) liD .;1 '" 0 j 5 ~ b .s 8 'Cf b 1: b +i> .g ; ~ ~ ~ 0 'I i r~ ] ~
i ~" 0 ...
.!t :$ 5 I '" ~ &! fit .., .. (;) =: :a
274, 439 - - - - - - - - - - - - - - -
213 - - - - - - - - - - - + - - - +
~7, 3124 32, ~7, 1 ,1)1, - - - - - - - - - - - - - + - -~)3, 212, 226, 261, )8S
- - -~BS - - - - - - + + - - + - -1247* - - - - - - - - - - - - - + -'*PERITRICHOUS Attn LOPHOTRICHOUS see RH 260 and 261
Or..,.. and Thornton (1928) d.escribed Pseudoaonu dacunh&e n. sp. It
~hich did not produce acid trOlil earbob;;'drates (see uiaw17, Gray and Thornton
~928). The figure. they- drew doscribing the lIlOrpholo~ show curved rods with
~ophotrlchot1S .flagella. Contrary to 3erpy's l..w.nual ot tletel'ld.native
~acteriolo&y, 6th edition (1948 page 10$) the gl"aa stain was not. recorded auld
tl1trates were not reducro to nitrites.
Tatarott (1891) de3cribed Bacillus aguat11is sulcatl!~ (rieichselbeum)
i8 a motile sporulating rod ).5 to 5 ~erons long, producing a water soluble
112
blue-green p1pent. Migula (1900 pap 93.3 am Table XIII tigure S) printed a
photomicrograph showing the lophotrichous fiagellation of P.eudoaonaa
ag.uatUise Bergey'. Manual of Detera1natiw Baoterioloo. 6th edition (1948
page 146) carries PaeudollOr1U .quatUia in Appendix I of the pllUS
Pseudomonas. The author hal neftI' encountered a lophotrichoUIJ water soluble
pigment producer.
Although. aOOOl'ding to the I1t.r&ture~ the 1ophotrioboua svaina
are predOllinantly of fecal origin there II'" to be S0118 indication that the7
mJq be ubiquitous. Table XlII shows the source ct isolation of the straw
studied.
RH nUllbers
11. 31, 124
)2
31
131. 1))
212
226
273
261
38S
TABLE XIII
SOURCE OF IsnUTIOI or LOPHOB!CTER FAECALIS
source of 1aolatlon
stool
vater
stool
blood
variant of 241
abdominal awab
Alcalip11!8 ape
Alcali,enes ap.
Aloalipne8 ap.
BacteriUll alcal!enes
5p11"111_ Y1I"p.n1anUil
Vibrio percolana
Alcalienea ap.
The name Lophobacter' I.ecala gen. et ap. 1'lOY. i. suggeated tor the
.' 173 follov1ng tvelve etn.1na, RH 273, 17, )1, 32, 37, 124, 131. 11), 212, 226,
261 and )8S, lated 1n Tabl •• XII and XIII. It 1a described a. tollow.
Gram negatift, asporogenous, aerobic rod. with lophotrichous tlagellat10n.
Does not oxidise or ferment dextro •• , lacto •• , auoron. aalto •• , .&annO.e, xylo •• or arabinose. Indol, -th71 red, Vopa-Proakaaer, oitrate and UI'H
testa Mgative. Nitrates are aeneral17 reduoed to nitrites but not to gu.
oelatin 'IIfII1'1 be liquefied. The hTdrogen sulfide teat is ne.ati" on Kl1gl.er'.
iron agar.
UNIT A OROUP 4
Frequently nonoxidiling BIOnovicboua Ol"ianiBlU U"e oonsidered to
be meaben ot the genus Alcaligenes. Table XIV above the reacticma of 1:he
monotriohoua dextrose nonoxidizers. A photosynthetiC JUiodOl?!eudOllOnaa,
RH 27S, .. included in the table tor coaparati" pUl"poses. N011l ot the.e
organislU, with the exception of RH 25 and 26 produced a water soluble
p1pent.. Mon1as (1928) .UC, •• t.ed that aonotr1cboua organisu which did not
produce acid !rca carb0h7drataa should be oaUed ' •• udoaona. aloal!&enea.
However, it did not appear that he dUterent1ated the lophotrichous Vibrio
alcal!l!D88 ot Lahunn and tlel.1iMmk The Mi. Pa.udomonu aloalipne. • ....
well suited tor orlani ... RH 6, 21, 22, 27, 28, las, 126, 330, 2S and 26.
In addition to the above nonoxidizing monotrichous paewlOlllOnads
three strainl were tmOountered, RH 339, )42 and )47, which were like the
above in BlOat retspecta .aft tor the .bape of the iIOno\r1chowt fiagellua. The
8ingle polar tlapllta vas strikingly d1tte"'nt. than that •• en on the usual
Paeudaraon&a tor it va. tto1ll"11"_ The vaft length. of the fiagellWll va.
- TABLE XIV
MONOTRICHOUS DEXTROSE NONOXIDIZERS* .. -~
~ s:: Q)
~ ;::s ell bD ~ ..-l
Ol '0 Ol 0. ~ fl) fl) 0 ;, fl) Q) Ol ~ ~ • ~
CJ) CI> CI> CI> CI> 0 Q) s:: .s ...-l bD 0 Ol CJ) Ol CJ) CI> .~ rl I ~ -rl 0 C\I ~ 0 0 0 0 Ol S Ol ell ~ AI I/) s:: ~ ~ ~ ~ § 0 .D CI> 13 ell C\I b ..!l: >< 0 0 ...-l r-i C\I bO CI> r-i Cfl 0 ...-l ::r:: CI> C\I ;::s C\I ~ ~ ~ 0 -rl 8 Q) (II oM (II oM
~ '"0 r-i I/) e; e; > 0 tID ::x:: s:: ::x:: e;
~75a - - - - - - - - - - - -
~ - - - - - - - + - - - - - -r->l - - - - - - - - + - - - + - -22 - - - - - - - - - + + - - + - -27, 28, 125, - - - - - - - - - - - - + - -126
330 - - - - - - - - - - - 0 - -25, 2~ - - - - - - - - - - - - - + + -
339, 342, 347c - - - - - - - - - - + - - - -
38d - - - + - - - - - - - + - - - +
231e - - - - - + - - - - + + + - -249 - - + - + - - - - - - i - -268
f - - - - - - - - - - + - - - +
269fg - - - - - - - - - - + - - - +
* The indol test was negative for all the organisms listed. a RhodoEseudomonas b brown water soluble pigment c curly flagella d rose-black water insoluble pigment e pink-orange water insoluble pigment f monotrichous and peritrichous g yellow-or 'bge water insoluble pigment
175
oonsiderably shorter in comparieon to the .... wnal type. The shape
variation can be .. en b7 coaparing the pho'torliOZ'OVapha, Figure- 4 and S,
on page 107. Perhaps this ph.n~non !Iliaht 'be related to the tlageUar
variant or Salaonella ~oh.ta photop'aphed b7 LaUson (19S1 tigve 6 Pl,.
380). A specie. distinction in the genus P~udOllOnu ..... juat.1tied.
Three other monotrichous organisms wer. encountved which M7 be
more closely related to the dextrose oxidis.ra. RS)8 ox1d.ised w to.. and
produced a vater ineoluble r08e-blaolc picMnt. RB 231 Olddiud arabinoee but
11kewi .. did. not oxidi •• dextrose. PeeudOllOnas s.ocharoP!B1a, RH 249,
which oxidized sucros. and llUUlnOee, 18 better know by the work or Doudorott (1940).
Or,an1uuJ RH 268 and 269 were nonoxidizing organ1_ ditticw. t to
place in anr genua because ot their flagellar JIlOrphology. The,. are
predOll1nant17 lIlOl1otrichoua organia_ with scattered peritricboue cells show
ing IIlOJ"e convoluted lateral nagalla. They have been photographed b,. LeUson
and Hugb (19S). Pure per1tr1choU8 or monotrichous strains could not be
maintained in culture.
Berge,.. s Manual of Detena1na t1 ve Bacterioloa, 6th edi tlon (1948)
d.scribe. Hftral specie. ot Pseudomonas which appear to have a ... relation
ship to the JIOD.otricMUS nonoxidi •• rs in Table XIV. 801M ot the" area
pace 103 pap 135 page 139 pap 140 page l40 pap 143 pap 144
Although Bergey's Manual. of DOt.erm1natift B_ctAarioloa, 6th edit-ion
176 (1948) describes the monotriohous pnua Iantboaonas u producing acid troa
Monosaccharide. and disaccharides, the speci •• alfalta' (page 165), pharmicala
(page 1(6) and niVOIUculans (page 166) are described &8 not utilising
carbohydrates. Thes. too then "1' be related to the orlaniau l1ated 1n
Table XIV.
UNIT A OROUP S
The last group ot OI"lan1uas to be considered in Unit A are the
nOflIIOtUe dextrose nonox1diaers. One can recognize the 11m1:taUona ot
assigning nonmotile nonoxidizing or,ani .. to one ot the tour tlagellated
dextrose nonoxidizing groups tor thq u'1 haft evol ftd trOll &rv ot then
groups. Perhaps Il serological study ot somatic antigens could otter S0ia8
help in a.signing these nol'llllotUe orgarli..;;;_ to t.he proper tlapllated &roup.
Table XV showe the characteristics ot the nonmotUe dextrose nonondi ... e.
Table XVI showe their source of i.olation.
Four authentic strains ot nonmotile Alcal!l!nea visco.us, lUI lS, 23,
3S and )6, were studied and listed in Table xv. Organiaa aolated tr_
tece., blood and. water _1"8 wry s1adlar to these strains ot Aloaliene!
lina nonoxidizing beta heaolTtic strains ot Alcalienea were studied
Six ot thes. straw, RH 164, 165, 166, 167, 169 and 170, isolated trom the
throat, were receiftd as Alcal:.enes heaokenB. The author haa chosen to recoillin the tint 2S orlaniSIU of Table
XV .. Alcal1&!nel YiscosWh Considerable 811n1!icanoe i. placed on the tind
ing ot the beta hellOl.ytic fecal organism. RH 194. which appears to be aiDLUar
~
.., TABLE XV
NO~OTILE DEXTROSE NONOXIDIZERS * 1-
~ ~ c: .,
i ., ~
" ., ; ~ ., G) to ~ taO • ~
10 t) t) ., 0 8 , ~ taO at r-t 0 10 to ro 10 Q) CIt 0 b 0 0 0 8 II) .... at ~ '8 8) c: ~ " ~ 0 .0 b at at
~ ~ 0 0 i c: ~
at 5 r-t 0 0 :t: t) GIl :;j I ; on t) r-t N a: ~ r-I 10 0 ~ .0 ~
14, 18, 63, 64 - - - - - - - - - - - - -186 - - - - - - - - - - - B -245 - - - - - - - - - - - -19, 23, 35, 36, 127, - - - - - - - + - - - - -211
9, 110, 128, 132, 160 - - - - - - - + - - - A -194 - - - - - - - + - - - B -216 - - - - - - - + - + - B -1164, 165, 1169, 170
166, 167, - - - - - - - + - + + B -370, 373, 374 - - - - - - - - + - + brown
115 - - - - - - - + + - - - -201 - - - - - - - + - - - A + brown
107 nc - + - - - - - - - - - -62, 65, 139, 175 nc nc nc nc nc nc nc - - - - - -
* Indo1, me~l red, Voges-Proskauer, hydrogen sulfide am nitrate tests were negative.
~
177
178
to the authentic Alcaligenes mco8ua ani ~.lcal1ene. helROtz!!9! strains
studied. The entire group 'Warranta further inwstigat10n betore hemolTtic
aU-ai.ns are separated from. Alcaliene. tiecoI'WI a. a new epecies. Alca.11enes
viscosus can be described a8 haYing the following cha.l"act.erut.iC8 s Gram
negative, asporogenous, aerobic, non.'IOtUe rod. without. nagell.. Does not
oxidise or tenwnt dextroN, lactose, lucrose, ul:to •• , MMO", xylose, or
arabinose. Indol, IIl8thyl red, Vogea-Proslcauer, urea, h1dro,en aul.t'1de and
nitrate tests negatift. Citrate and gelatin te.ts variable. Blood agar deep
colonie. tIIIlT show an alpha, beta or a .. heao17.ia.
'1'ABLlt XVI
SOURCE Or' ISOLATION fI NONMOTILE NOlfOXIDIZiRS
9, 127
14, 1)9, 17S, 24S
IS, 23, 3S, 36
18, 19
62, 6), 64, 6, 107, 128
UO,216
1)2, au 160, 186, 194, 201
164, 16$. 166, 161, 169, 170
370, 313, 314
source
reces
feces
blood
reces
throat
lloali&!neB f.ecalia
Alcalienes v1s~u
aeliconema sp.
AIO&l!&!nes taecalia
parapertusia bacillus
179 The Alcal.!ene. viacosws atralne etudled by Long am HaDav (19)5)
and tho.e liated in Table XV gaft negatiw ftlt.rate teata. Abd-el-alek and
Oib.on (19$2) found nitrate positi" strains. !be _jority ot cultures
examined bY' Abd-el-malelc and Gibson did TlCt produce .. J'OP1 condition in aUk.
Viscosity ie described 8. an indefinite propertY'ftt1'ing 11'1 degree f'rQll stra1n
to strain and among coloni •• isolated fraa a s::1ngle culture. The oxidation
of Citrate, and hence the production ot alkalinity in lita118 aUk, was alao
shown to be slow and erratic.
Four etrains, R1i 201, :-'10, 373 and 37h produae • brown _tv soluble
pipent. The latter three strains an parapertussis bacUli lihUe RH 201 1&
of tecal origin. RH 15 and 201 are sia1lar to the parapertuau bac1llWh
'I'he.. .v.ina 1U.7 beeo.M k%lovn .s Aloaliena. er&E!rtussls. £lder1ng and
Kendrick (1952) recorded a positive citrate reaction tor parapertua81a
bacUli. The strains RH )10, 373 and )74 ga.". a ne,ative Shlllon'8 citrate
reaction.
Orca.n1.Ml RH 62, 6;, 107, 139 and 175 produced no change ot
lMlcator or onlY' • wry taint. trace of acid in the dextrose mediua. Thie
was quite unusual since the _jor1tq ot organ1eM produced an acid or al.lcaline
reaction in thi8 aed1um.
The paraaount objeot1 ... of this d.1Hertat1on vaa not the ult1m.ate
oream.ation ot the carbohydrate oxidi.ers. Howewr, pnerall7 lIJ'INOOgn1zed
signitioant. differences weJ"it enoountend wb1ch cannot be ldn1aized wben
or,anisation 11'1 this group 1s to be cona:1dend. Tbe ox1d1aera were 11'1ftsU
gated tor (1) orientation, (2) oonfidence .s to what should or should not be
called Alcal1£!ne •• (3) what organi_ are concerl'l8d when Alcalipnea and
160 related generic names have been applied. Unit a vas cOlIprised ot 123 organ-
iau which oxidiaed dextrose aM was diYided into IV 5ubuni tat on the baai.
of oxidation of lactose and sucrose. Table XVII shows how Unit 3 was di rlded
into the IV Subunits. The subunits were furth .. separated into groups on the
basis of flagella as was done in t1ni t A.
TABLE XVII
DLFFE.RENTIATION OF DEXTROSE OXIDIZERS (UNIT B) INTO SUBUNITS
subunit dextrose lactose sucrose
I + - ...
II + ... +
III + + ...
IV + + +
WIT B SUBUNIT I OROUP 1
0n.l7 one peritr1chous organism was found in Group 1. Thu w.. ton.
organia Acetobacter !leet1, lUi 262, apparently a JIL'1anoaer tor the entire
genua is considered b.1 Vaughn (1943) to be lIOI1Otr'ichous it aotU.. Photo
micrographe Moldng the napllation of Acetobaowr an ,e,.ral.l1' laokjng,
thoae published b.1 Vaughn (1942) and Zeidler (1898) are unaonv1nc.\r1g.
Aocording to VaUCM (1942 and 1943) ther. ex1at.a a YW'7 cl08.
relationship betwen Acet.obacter And PHUdOJlOnas. The two pnera probablT
ditfer in their utilitarian aignitioanoe. Acetic aoid bact.eria can grow at an
extreaely low pH and can oxidis. 6% alcohol to acetic acid.
Two Acetobacter strains, RlI 26) and 277, vve COlIlp&r8d siaultan-
"""
;,
TABLE XVIII
OXIDATION OF ETHANOL
ethanol RH number received as flagellation dextrose 1% 3% 5%
1 Pseudomonas aeruginosa mono + +2 +2 +2
3 Pseudomonas aeruginosa mono + +2 +2 .. +2
5 Alcaligenes faecalis polar multi + + + +
9 - - + +3 + 3
21 Alcali~enes bookeri mono - + +2 +2
28 Alcali~enes sp. mono - +2 +2 +2
35 Alcalis€nes viscosus - - + + +2
66 Alcaligenes sp. mono + \ \ +4
122 Pseudomonas aeruginosa mono + +4 +4 +4
127 - - - - -177 Agrobacterium t~~efaciens peri + +2 +2 +2
262 Acetobacter aceti peri + + +2 +2
263 Acetobacter oxyrians - + +2 +2 +2
277 Acetobacter rancens - +2 + + +2
339 mono - + + +
KEY:
- nonmotile or no change of indicator + acid reaction of indicator in one day The subscripts indicate the number of days required to change the indicator.
181
182
eoualy in aloohol media vi th the peri tl"ichoua AoetobacteJ" lUi 262 and other
ftrioualy flagellated organimu in the collection. The carbohydrate base
d..scribed a'boft vas used, to 'Which vu added ethanol to g1 .... a tinal
concentration ot 1, ) and S% b7 velUM. The Nault. aN 801111 in Table IVIII.
The criterion tor the .eparation ot Acetobaeter tr. "eudaaonas •
\)7 acid production in high concentration ot ethanol 18 not tan1q\a8 tor it
appe8l"s that other genera are also able to produce acidity in tbe above mediUII
UNITS SUBWiIT I GROUP 2
i'he second Croup ot Subunit I was .cOloprised or 13 polar JIlUltitr1choUi
orlanisJu. The u.jor1ty" produced a vatoer soluble pigment. A fev ot the
pipented strains produced a chlorotona aoluble pipent. T!1ble XIX above the
reactioM of then polar Jlultitrichoua 'aeudoaonas strains.
TABLE XIX
POun MUL'l'ITRICBOUS PSEOOOMONAOO*
~ .. c::
'-t J i 10 0.
I tJ • ., ~ ~ .a Ull I • I I 8 .s ! 0 ; :
~ 'f'1 b fl ~ s:: ~ ~ 0 1 &
s..
~ i 'i $ :& 0
~ -3 ~ or4 ft 10 <I ==
199, 200, 2S7 + - - - + + + + - - - - -S, 1S3. 193. 197, 213 , + - - - + + + + - - - + -280, 298, )02, )SS, )84
*The irldol, -th71 red alii Vogsa-PJ"oskauer testa were nelati ....
The relationship of the.e polar IIWltit.richoua pseudoaonada to
18)
p •• udoaonaa urua1n08. 18 very stzo1king. The P •• ud0Jl0na8 l1!!)¥iinoaa straina
reviewed in Table XX were true denitrU1era where .. the polar lIlult1trichoua
pseudomonads do not reduce nitrates.
PseudOllOllaa ezneranea (Ehrenberg) Migula. (Vibrio yncl!!!ua,
Khrenberg), va. described b7 M1gula (1881) a. a _ter 801Uble pipaent
producer. A drawing !'$L~0V8 polar au1titrichoua napUatioa. 10 sU!.teJIl8nt
is ude concerning carbohydrate utUuation. This name i8 retained in
BergeTts Manual of tetorainatiw aaoteriolol1. 6th edition (19~8 pap 92).
!pilT B SUBUNIT I GROUP J
Thi8 third Group ill represented by but one organielll, RH 207. It I. nageUation was so;.aevhat unusual tar it haa short nagella with a elicht
curvature resembling lophotrichous torMS.
UNIT B SUBUNIT I GROUP 4
The fourth group includes the better' known pseudomonad.. The blue
green vater and cblorotOl"ll soluble p1pent.ed P .. udoaonas aerg1nosa i8 euily
identitied. Jordan (1699) did not exclude apyoqanopn1c .trains Il"OII
conaideration .. f:. "mino... He eucp8ted, without ottering conrlnoi.n&
eTidenee. that the dUterenoe between PHudOll&OJ'}.U fiuonscene and ap7OC7&!10-
genic etra1na ot !:. "!j1Ilno" vae one ot 'Maperature opt1M. Many workers,
Sandiford (1937), aaby (1946), and ehrittie (1948), .ince Jordan'. report
haw continued to consider P. nUONsoena a modified orde,enerate torm ot -------!:.. aer!llino.a.
There haft been .... ral aerololioa.l reporta, Christie (:.t.9h8),
Mayr-Harting (1948), Munos et ale (1949) and van den tilde (1952), concerned --
184 vi th the study of pseudOllonads but the taxollOlll1o significance ot the work 18
diff'icul t to interpret.
Seleen and stark (1943) and Haynes (19$1) appear to agree that
E:.. aerus;inosa w1llgrow at high temperatures, )7-42oC, wbUe !.:. fiuoreaoene
and most otber nuorescent bacteria are unable to do 80. Hqnes tound that
the abUity or f.:. aer)liinosa to gro'W at 420 C correlated with oxidation ot
potassium gluconate anti alae production.
The tirat Sh orgmullu listed in Table XX are considered to be
PseudollOn&a aer91no8~. Ito attempt VIla DUilde to dift.rentiat. aer~sa tl"oal
tluorescena. Theae organis .. have the toll01l1ng oonalatant ph7s1olopoal
pattern I
dextrose ... citrate ... (rarely -) laotose - indol -lucro .. - -tb71red -maltole - Vogea-f'roskauer -manno •• + urea -xylose + nitrate • arabinose +
The carboh1drate pattern 18 in agreefdnt _"1 th the recent report by
Uu (1953). 'the production ot a vater soluble pigment and liquefaotion of'
gelatin are h1ghl.y' variable charaoteriatioa.
Th1a ~ioloeical. Pllttern w1ll be recogn1Md as bav1n& considerable
value in identity:1ng Pseudomonas UrsUlO.. whet.her these oriania:u appear
with or without a water 801uble p1pant. The sign1tioance ot tbe abOft
carboh7dr&te pattam tor ?eeudomonaa will bec .. appreciated when it 18
realiled that it is a reaponee in an Ol'lanic nitrogen m.ed1wa. 'l'he carboh¥d
rate pattern J"t;8ponse established above wa. "producible tor it 1n'.8 repeated
with -D¥ .trains and with new bat.ches of _dia m&n,1 times. The med.iUPl
,,-TABLE XX
PSEUDOMONADACEAE "'""
.. +' E! QI)
10 OM
s... ~ Q) Q)
Q) 10 .
~ 17) Q) G> Q) Q) 0 Q) Q) ...-t o~ 0 til til 10 til G> .~ +' +' 0 !l 0 0 0 8 10 cd cd til +' l=: +' s... +' 0 .0 s... !l lIS X 0 0 ...-t l=: ...-t cd +' tI) 0 ...-t ::r: Q) lIS ;:::l cd
£1 ~ s... oM (II OM (II Q) ~ "U rl til a as 0 :x: l=: :x: QI)
55, 56, 57, 58, 59, 111, 112, 113, 12), 141, 142, 14), 149, + - - - + + + + - ~ + -187, 189, 190, 208
1, 3, 122, 14h, 145, 147, 148, 150, 151, 156, 161, 188, 191, 192, 195, 196, 202, 209, 210, + - - - + + + + - ~ + + 217, 218, 417, 418, 419, 420, 421, 422, 42), 424, 425, 426, 427, 428
146, 2h2 + - - - + + + + - ~ - +
429 + - - - + + + + - ~ - -157 + - - - + + + - - ~ - -2* + - - - + + + + - - + -341 + - - - + + + + - - + +
66, 204 + - ~ + + + - + - ~ - -. , 203 + - - + + + - + - + - -129 + - - + + - - - - - - +
364 (flesh colored colonies) + - - + - - - - + - - +
198 + - - + - - - + - + - -163 + - - - - - - + - - - -313, )14 + - - - - - - + - - + -300 + - - - - - - + - - + +
89, 95 + - - - - - - - - " - -Note: The indol, methyl red, Voges-Proskauer and urea tests were negative.
*Biochemical pattern sam ~ as polar multitrichous organisms in Table XIX.
186
introduced here. in oontrut to the iDorl&nic aedll111, alao has the distinct
!ldvan1.a.gfJ ot different1a.ting oxidiurs tro. tel"lll8nters .0 that there can be no
mistake of plaoing a polar flagellated term8nter or e..a ottwr terment8rs in
The cltrate reaction was ot greater value in thl. JaOnotrichous group
than in the ~cal!iel~8 group tor it gaw a wry deBoiai," change ot indicator.
nitrate 11&, 'f'e'I!Y charaot-etrutically' reduced to nitrogen CaB tor the
PeeudO!!ltOnlle •• !!Iinosa strains. Oas production was cont1naed b7 anaerobio
growth .tudiee. This phenoaenon vas elaborated earlier in the disoussion.
The latter 13 orlsnieu in Table IX could be assigned 8peole.
epi theta in the genus P •• udoaona. but should perhaps ava! t f'u.:rtb.v atud7
and isolation ot Blare strains betore their orientation beoOMa olear. Wo\lld
it be JIlOre OOl'l"8ct to assign orpniam. RU J64 to the genus Xanthoaonas rather
than Pseudomonas? No attempt was made to eatabl18h relationship of the.e
organ18IU to previ.ously de.oribed organi:.u. IUn;r of theae unndatakable
psoutlOlllOflads did not reduce nitrate to gu but appeared to stop at nitrite
or' could not reduce ni we te at all.
UNIT B SUBUNIT I GROUP 2 The tifth and lut Group or Subvni t I concerns aix ftCnIaOtUe
organisUla. Two wen labeled Acetobaoter (aB 26) and m). Or,aniaa Rli )60
!With yellow vater insoluble p1pent oould probab17 be olaa.it1ed, according
~o Berge,." Manual. of DeteJ"Jl1natiw Baoteriolou. 6th edition (1949). a. a
nonmotUe organiSM in the genua , •• udca.onaa, Xanthoaonu or Flavobaoterlu..
:rhe nonmotUe organiaa R.li 61 va.a "Ie1:7 interesting to haft encountered since
.... ~.+ ... -~... .•
187 and flagellation.
UNIT B SUBUHIT II
The ten organiau in Subunit II an listed in Table XXI. TbeT are
recognised 'by their ability to oxidise dextro .. and 8\lCI"Ose but not lacto.e.
The peritl'ichoua Group 1 in this Subunit II -1' be 1MtlIlbera of the genua
AEobacteriua. hOlMver Rii 318 _y be a Flavobacterit1ll ainoft it produces a
yellow pigoiant. Tho remaining nonm.otile and polar nag.llated organ1.tu mal'
become eucro" positive epecies in the genua PseudOMOnas.
Vibri.o cuneata n. ap. was d •• crib'Jd by Gray and Thornton (l928) a.
a curftd rod with 1 to , polar flagella. Drawings indicate polar aul tit.rioho
tlapllation. sarae,,'. Manual of' DetAlnainatift Bacterio1oQ, 6th edition
(1948 pap 199) retains this organUm :in the genua Vibrio. Troia orpniul
was dhcribed (... Hutory. Grq and Thornton, 1928) as not producing acid
from carbohydrates. '·1brio cu.r.t.eatWl, RH 248, deposited by Grq and Thornton
in the ,..-ioan Type Culture Collection, had polar aul t1 trichous flaplla
but. produ.oed oxidatift acidity in dextrc:Je and other carbobfdratea as shown
in Table XlI.
WIT B SUBUNIT III II
OrpnisJU in Subunit III oxidise dUtroH and lactos. but not .
suoros.. The orlani... and their reactioM ar. luted in Table XXII. the
entire subunit mght be included in the ,.nus P •• udoacna8.
The urea reaction tor the monotricboua orgam..u .. ceneral17 ver.r
weak. .The positive urea reactlone in the table 1M1cate chan,.s of indicator
seen in 48 bours. Han,y times a positive reaction .... en in three da7a.
'" TABLE IXI
UNIT B SUBUNIT II*'
~ s:: 1 0 oM
..-4 Q, r.. ~ Q) I) CIS IV Vl • ~
M Vl IV IV IV I) 0 .$ I) M ;l M 0 Vl Vl Vl Vl IV ;l ..., 0 I) !J 0 0 0 0 III CIS CIS 10 ~ s:: ~ ~ r.. ~
~ 0 ~ !J CIS r.. cd
>I (,) (,) M
'Sa ~ ~ 0 M
~ M ~ cd ~ ~ .~ ..... "E 'JJ" fa. ~ M (,)
Group 1
318a _peri trichous + - + + + - - - - - -- +
2~3 peritrichous + - + + + + + - + .. - -246 peritrichous + - + + + + + - - + - -171 peritrichous + - + + + + + - - - - -176 peritrichous + - + + + + + - + + - -
Groups 2, 3, and 4
248 polar multi + - + - + + + + - + + +
443, 444 monotrichous + - + + + + + - - + - -272 monotrichous + - + + + + - + - + - -363 - + - + - + + + + - - + -
KEY:
*The indol, methyl red, Voges-Proskauer and hydrogen sulfide tests were negative
a yellow water insoluble pigment .
188
18' Group 3 1nclude. the nonaotUe Baoter1_ anlwata d.aor1bed b;r
Sohaub !! ~ (1948), st.uart !!!!:.. (1949), 'ft'CWtOft !!. !!.:. (1950) and Brooke
(19$1). Th1e 18 a 'NrY bom.ogenous ... 0010 croup not. to be oontued with the
para colon bacilli which .rerMnt dexwo... Sotallb (1948) obeerftd t.bat
carboh,ydrate. vere ox.1diaed only in the presence ot au and tbat th1s differed.
t'roat t.he f'ern!umta tlon seen _ong the Enterobaoteriaceae. Jfurt.bermore a 10%
lactos. agar "lant 11 hot essential to demol'l5trate acid produotion since it 18
readUy .een on the U lactose ~1wa described. BrookQts (19$1) negative
lacto.e tinding empJuasi ... again tb8 iIaportance of using a more suitable
Contrary to Schaub'. (1948) :tinding, a positive Mtb11 red test va.
not obaerved. Brooke (1951) found 84 or hie 66 atraine ga"" a napti_ -thTl
Balaauth'e (l9Sl) organism NRS-ll, RM 432, should not be considered
an ttanaeropnic paracolon baoillus". Strain Rh 10 was unusual sinoe it pn
• positive BWJtlgian-Stuart urea teat. Schaub round _Ba_c....,te_r_i ......... , anitrat!,l!l!
g1..,.. a positive Christensen's urea agar test. Brooke alISO found urea
])Ositl ... straiNh Oelatin was posiUve in but one 1natanoe, RH 168, howeftr
RH 10 vas originall.7 described as giT1ng a positi .. gelatin reaction. The
svains RH 16 and 168 cleared milk agar. The tollow1n& ten etra1n8 listed
in Group 3 of Table XlII are cons1de.r1td to be BacteriUl'l anivat_, RH 11, 16,
24, 1)0, lS9, 14.10, W, 432, 168 and 10.
UNIT B SUBtJllT IV fIM
1.'h18 last Subunit- of Unit B va •• stablished. on the basis ot having
., TABLE XXII
UNIT B SUBUNIT III • ~ -s:: tb 0
·ri ·ri H ~ 0-~ ro Q)
Q)
~ r-l II) Q) Q) Q)
II) . Q) 0 .$ Q) r-l .~ r-l 0 II) II) II) II) Q) ;i ~ 0 Q)
~ 0 0 0 0 II) QI ro III ~ s::: bO ~ ,.. ~ s::: 0 .0 ~ C1I H ro
:z: ro >< () () r-l
~ ~ C1I Q) ~ 0 r-l 1'""1 Q) ro ~ m fa -ri H ·ri (II Q) CJ:: fo...4 "0 r-l II) () ~ s::: :z: till
Group 1
154 polar multi + + - + + + + + - - + -Group 2
155, 295, 296, 312 mono + + - + + + + + - - - -284, 308, 309, 310 mono + + - + + + + + + - - -311
259 ?mono + + - - + + + + + - - -266 mono + + - + + + + + - @ - +
Group 3
11, 16, 24, 130, - + + - - + + + + - - - -159, 410, 411, 432
168 - + + - - + + + + - - - +
10 - + + - - + + + + + - - -
Note: The indol, methyl red, Voges-Proskauer and hydrogen sQlfide tests were negative.
-·190
191
studied two organiau which oxidized dextrose, lactoae and sucrose. These
polar flagellated organisu oan be siftn specie. recognition in. the genus
Pseudomonas. The reactions ot these two Ol'g&niAa, 1ft )01 and )7S, oan be
seen in Tables II and Ill.
UNIT C
The 39 org~ni~~s in Unit C are quite difterent than the previouslY
considered Units A am B. Unit C is eOMorised ot taeultatiw organisms wtdeh
tV1!l8nt dextrose, i.e. the;r metabolize dextr'ose in the presence or absence of
air. The or,anil'!118 of this Unit ax-a divideti tnt.a five Subunits Oft the buis
ot response in dextrose and lactose. The o&s1s tor tr_ differentiation 1s
seen in Table 111. It vas not the purpoS\l of this work to consider this
Unit in d.etaU. however cursorY' obaerntione were JUde to elucidate the
r .... ntat1ve am oxidative Spectrunl ot OrganiAUU.
Drganiem RH 383 terments dextrose without gas production and
oxidi.e. lact08e. The taxonomio 8 ta tus ot this organism is not certain but
perhaps there 1s sou zoela tionsh1p aaong the anaeragen10 paracolon bacil11.
Two Sh!iella, RH 4)6 and 437, ~si. and sonne reapeot1.a11,
.... re studied which fermented both dextrose ani lactose.
Crimi.8U Rli 174, 224 and 228 were included he,.. only because the)"
j:; were .ent to the laboratory for study labeled as Alcalienea, s.n obY1oua .,j
Ddsnom.er since they terment dextrose. It is indeed unfort.unate that the
Arlerican Type Cult.ure Colleetion and the National Collection ot Type eul t.UJ"ElS
continue to disaeaine, te and carry these organil'llU as Aloalilenes.
Three per1trlchoU8 organulIl8, RH )04. )lW. and 433, one of which
rr-----------, 192
vas an authentic pan.oolon baclllua, were studied which t4ll'Mnted dextroae
with gas and oxidised laotose.
AIlong the genera included in t7ni t C vh1ch t ..... nted dextro.. with
acid and gas vere salJaonella:, Proteus, anti Aeroaonas. Aerc.onu 18 nadUy
distinguuhed b7 ita monotriohous flagellation. The deta1led ,..aotions of the
Aeroaonas strains studied are g1 wn in Tabl. run. The.. oraan1au are
includ.d in the genus Pseudcaon&S in Bergey'. t.fanual of 'De~t1ft
Bactv1ologr 6th ed. (1948), but since they'terMnt dextro.e the .. nus
AWCIIflonu .eau more suitable. The validity ot thi. genua 18 r ..... 1a.d
b1 MU.s and :1U .. (19$1). PeritriGhouelJ flagellated indirldl:lllU an shown
to occur, LeUson and Hugh (19))), in early growth phase. with onl)r sinal.
polar nagella in later stages. Aerom.onu ligU8tac1ena strain .L4l8 was sent
by J. Klurver (RR 2S4)and Ellen H. Mile. (RH 271). Both sources of th1a
strain indioated the culture to be IlOnOtr10h0ua, .e. 1'41le. and Mile. (19$1).
In our laborator)" this organ is. was found to be peritrichoua. More recently
t11le. and KIU1"l8l" haft cont1rMcl the per1tr1choua na,ellation (personal.
cQU&Ul11cat.ion 1952). The genua A81"OlllOnu appears to be related to the
peritr1cboualy fiagellated tam.ily Enterobacteriaceae since Aercaso1"&&8 termenu
carboh7drate. and produce. peritr1cbous flaplla in 10Wlg cul t.UNa.
.., TABLE XXIII
THE GENUS AEROMOHAS
M
c: ~ 0 ~
·ri "0 0) M -t3 CD f 0 CD IS CD 0)
~ j r-f 0) 8) CD CD 8) 0 CD 3 .;; r-f 0 0) 10 10 0) CD .;; i I -t3 CD !J 0 0 0
I C7)
~ 10 .. If -t3 c: tID ~ M -t3 0 ~ CD ~ ~ ~ .. >< u U r-f r-f tID
~ r-f CD If ~ I ~ fa I 0 -ri -ri CD CO-4 "0 r-f C7) -ri P u c: bO
250, 251, 289, 335, 343, .349, mono i - + + + - + + - + + + + 353
252 IIlono i - + + + - + + - + - + +
253 mono i - + + + - - + - + + + +
255 mono @ - + + + - + - - + + + +
292 IIlono @ - - + + - + + - + + + +
320 IIlOno i - - + + - - - - + + + +
254* PERI i - + + + + + - + + + + +
271* PERI t - + + + + + - + + + + +
KEY:
* The peritrichous organisms RH 254 and 271 are included here only because Miles and Miles (1951) considered the. authentic strains of AeraJlonas. nuynr (personal cODlDlunication, 1952) has contirmed the peritrichous fiagellation of RH 254.
The urease test for these organisms was negative. Hydrogen sulfide tests were negative on IO..ig1ers Iron Agar. Water soluble pigment was not produced.
193
CHAmR VII
SU:t4ARY
1. The bistory of the genWl Alcal1pne. baa been rev1ewd and an
extenaift bibliography presented.
2. Strains of Alcalipnea and related organie .. vve :laolated and
accwaulated trOlL lIl&D7 laboratori.a and coUections. Their JIIOI"pbological. and
physiological characteri.tica are recorded.
3. A technique i. presented which dUferentiate. oraan1au on the
basis of carbohydrate utilisation. Three uni ta were fonaed, uone the organ ...
i_ studied, on the basi, ot metabo1iua ot dextroee, they' are. (A) no
acidiv produced, (B) oxidatift acidity produced and (C) termentativa acidit7
produced. F.J"Mntation 18 described .s a proce.s whereb,. an organts. can
obtain .nerd from carbohydrates in the abaence ot olQ"gen.
4. One hundred and twenv organie_ were .tudied which did not
produoa acidity in dextrose _di-. Bacteriologi.ts have o.tm inclined. to
place I&Ost ot these organi ... in the genua Alcalijene. ewn thoUih the7 are
not hOllOgeneoua in respect to morpbolol7 and phy.ioloa. Th... orlanulII8
were d:Lrlded 1nto aroup. on the balta ot naaellar aorphology.
S. Appl.ying the above cone.pta, the genus Alcal1pnea i. limited
and detined as tollowsl Or .. negative rode. Per1t.richou. or noraotile.
194
19S Growth on agar slanta nonchromogen1c to l'tUov. The pigMnt doe. llOt diftuse
,through the agar. (Rldering and Kendrick, 1952, do not describe PFapertuasi.
as producing a pi,raent but preter to call it a -browning ot peptone _dium".)
qlose or arab:L"lOSG. IJldol~ mathyl red and VOCGs .... ProsKauer test. negative.
Hydrogen sul1'1de test negative on Y.lj'i1er's iron agar. The speoies are
d1tt.rantiated in Table XXIV.
TABI..E: XXIV
(I)
+> cd
cd M (I) +> ~ ·M
s::
A. broncllicnnis + + •. l -+ %
A. den1tritaciona + $ -A. v:1scosus ------... ..
*occaslonally negative
6. Threo groups of oraani8lll8 Rre nov clearly excltlded troa the
genua AloaliJe..... They are placed in three separate giF\era. (A) Xanthomonas
naaltophUia ap. nov. was created tor polar lll:illtitrichoua dextrose nonoxidizers
which produce oxidative acidity trOll altose. (13) The designation Loeobacter
fa.calia gen. et ap. nov. is proposed for lophotrichou8 nonoxidizing &r1.l1l
196
negative rod~. These organisms have been shown to have a morphological and
physiological relationship to S,e1rUlum 't'irg!.nianwa. (C) Pscudcmonu
alcal1senes 18 retained for the monotrichous nonox1diBing gram negative rods.
7. There a.rs fOUl' f:roups of d~x.tro8e oxidizers which warrant Bin ...
phasis here. (A) There appears to be no objection to the use of Pseudomona.g
sl!lganea to represent the polar fIlultitl-iobous water soluble pigment producers
listed in Table Xn.. (B) ?seuflamonas aeruginosa represents strains Nith arrr
combination of pyocyanine or fluorescein or even ld.thout pig..nt. Thes.
oraanisu are described as having Ii. very definite lJWrpholo&ical and phy'sio
logical pattern. (C) The genus designatiM AebacteriUl4 should probably be
reserved for peritrichous rods whioh oxidise dextrose and sucrose but not
laotose. (::;) The biochemical activity of Baoterium anitratuw is shown to - r • •
be a hcmlogenous group of aerobio orianis..'1lS when liai tat! to dextl'ose and
lactose oxidizers a8 shown in group .l in Table XXII.
8. The genus designation AerOlllOnae i8 a.ppropriate lor monotrichous
gram negatift rods which ferment dextrOse with the production ot acid and gas.
Tbe.1 are organisms ~lich uho'W a .~rphological relationship between the tam-
Uies Ps.udoaonad.ac~ae and Enterobacter1&oeae.
9. The designa:tioolll Alealienefi bookeri, reoti, £IlA~hall1 and
~taleal;lle!...e~ should becom.e nomina dub~ 8ince the application ot' these
epithets is uncertain.
I I
1
'Ii :1 Ii
l
CHAPTER VIII
BlflLlOOP.APHI
Abd-el-malek, Y. and T. Oibson 19$2 studies in the Bacter101017 ot aUk.
Ad_ta,
Ad&IUta,
Adaraeta,
Ahad, w.
Adaluon,
IV. The gram negative roda of .Uk. J. Dairy Re •• arch 191294-301. -L. 1889 Ueber einen Err.ger del' achleim1gen Milch (Bacillus 1aetia visco.us). MUch Zeitung 18,941-943_ -
L. 1891 Untersuchungen Uber Bacillus laot18,.~ ~inan we1 tverbrei teten milchtdrthschirnlChen S&(u1I1,. Landlr1l"thschaf'tlicbe Jabrbucber 20118S-207. -
L. 1891 Untersuchungen uber Bacillus l.otis vi.oosua, .iRan veitverbreiteten Milch wirtheC&tUlcnen sCliad1lng. Zentr. But. Par.aitenk., Crig. 9.698-700. -1942 The role of Bacillus ta_calia alcal~M8 as a pathogen in C7stitis ot Ul"iMr7 blaacler. Iridian Mid. nett. 77:530-532. -
C. A. 19149 A bactenologioal studT ot lymph noda' A.nal.7sia ot post&ortwa specimens w1th particular reterence to clinical serological and histo-patbo1ogiea1 .findings. Acta iied. Scan. Supple 227:1-172. -
Allen, L. A. and E. Brooks 1949 Destruction ot bacteria in sawage and other liquids b7 chlorine and by" ""anogen chloride. ~r. Hyg. 47 & )20-336. -
Allen, M.. B. and C. B. van Nlie1 1952 Experiments on baeterial denitrification. J. Bact. 64'397-412. -
Alte.1ar, W. A. 1945 lnactifttion ot penicillin by various gram negative bacteria. Surg. Omec. Obstet. 611379-38). -.
Altenbern, R. A. and R. D. Housewright 19$2 Carbohydrate oxidation and citric acid synthesis by IJIOOth Brucella abortus Strain 19. Arch. B1ochel1l. Biopqs. l2.aJ4S-3S6.
A1 tschuler, E. 190U t1ber die Bez1ehunpn des Bacillus tocalis alka11iene. au den TJphuabaaillen. Munchen _d. W&&_ ll.'BM-870.
197
t I
I
Il
198 Andersen, E. K. 19$2 Serological. studie8 on H. pel"tua81a. B.~tue.i8
and H. bJ"onchisepticua. Prel1a1na17 -report. leta 'Pi It croble!. Sca~ j2~>4-5a.
Ander.on, J. A. 1945 A new member of the genua A~Cal~8 and ita qabi08ie with acid tonaing organiU18. J. Bact. ".204- mi. -
Anderson, ,"I. K. 1933 Two cues of S.eUlus alcal!enea intections. Practitioner 1311102-10S. -
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20)
Blderin" G. and P. Kendrick 1952 Incidence or parapertussu in the Orand Rapids area as indicated by 16 year. t experience with diagnostic cultures. Amer. J. Pub. Health 42.27-.31. -
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20$
faecalis alcalienea und des BaoUlua nuoreaoeM !!!!! l1gufac1ena. tanG. Baitt. Paras!tenk., I Abt. ('$11. noai7i-M.
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206
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Kuhnemann, o. 1910 M1Xrophotograph1sobe neaoutnt1on.n ubV diAl Oelueln del' 'f:i-phuabaa1llen gruppe. M\mchen mad. woohenacbr. 1!.219.
ICul'lneIunn, o. 1911 7.v Ident1tlaleruna d .. Baoillua taeoalis aloal!&!!!ea. Zentr. Sakt. Paraaitenk., I Abt. OJ"t;. 57'469=471. -
Kutaohfl', G. W. 1931 Allca11:1ines bookeri mening1tis. Ren.v ot the literature on ar __ negatiw SioUlua lI8ftingitia. Arch. Pediat. &a610-616.
Lattorpa, M. 1'.«16 Quelques remarques a propoa d'wa BaoUla ~p dana une infection typhoide. COJapt. rend. aoc. biOi. §II ... •
l.ebItaM, K. B. urd R. KeU1lUWl 1899-1927 Atlaa unci OrwldrU8 der BakWrioSOI~ W'ld Lehrbuch del' a~e1ell.n SikterIoJ.;OI.!0;i 15Iiaiji.Ulc. • .~, :4unCli8ri. na: ea., lB~ J 'iJi ." 7 •
LeUaon, E. 19)5 1fav oulture _dia. baaed on .odiwa duoqoholate tor the v 1aolat1on ot intestinal pathocel'l8 and tor the enUllU'at1on ot colon bacUli in .Uk and water. J. Path. Bact. hOI 561-599. -
LeUson, E. 19~2 Bacterio10tI; Paul. B. 8oebe1'" Inc., l4edioal Book Dept. ot Harper & Broib8rs, York.
IAifaon, E. 19$1 sta1n1ng, ehape am arrana_nt of baot.rial naplla. J. Bact. 62.)77-389. -
208
LeUson, E. and R. Hugh 19$) Variation in shape and erranp_nt ot bacter1al flagella. J. Bact. 6)126)-271. -
Le'f'1ne, M. and r. C. Carpenter 192) Gelatin liqataction b7 'baoteria. J. Bact. 8,297-306. -
Lenn., M. and L. Soppeland 1926 Baoteria in creaaaer.r ... tee. Iowa State Oollege ot Agriculture and Mechanio AI'te, Ames. Engwer1nc Expv1lllent [:tation. Bulletin 77, ~11-72.
Lindeann, o. 19)0 Ueber die Patbogenitat de. Baclllus taecal1. alcali;emuh Zentr. But. Par.sitenlc., I Abt. 01'1&. mej2a-Jj).
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!Wahall, C. E. 1896 Ropinea. in cre ... or lIlUk. Bulletin mmlber 140 H10hiaan State Agricultural College Exper1ment Stat10n pages 97-108.
Marshall, c. E. 19~ A prel1.m1nar7 note on the '&asociati" action of bacteria in the souring ot .ilk. Zentr. But. Paraaitenk., II Abt. 11:739-71&4. -
Marshall, M. Y. 1928 Pathogenic1ty of Bacillus recalia alkal!l~m8 •• Kentucky Mad. J. 26:89-89. -
Mason, R. J. 1934 Acute aen1llgitls due to Bacillua recalis aJ.calienes. J. Pediat. ~IS14-$16.
Marr-Harting, A. 1948 The "1'01017 or Pseudomonas PIONanea. J. Uen. rticrobiol. 2.)1-39. -
McAlpine, f1. o. and C. A. S1aneta 1928 Studies on the meuboli_ at the abortus-raelitensie group. III. Glucose utilizat1on. J. Int. Die. 42:73-78. -
McCullough, N. B. and G. A. Seal 1951 Growth and JIUUlometrio studie. on carboh7drate utUization of B""""r .... UGe ......... l .... l .... !. J. Int. DU. ~1266-271.
McCullough, N. B. and L. A. Dick 1943 Growth ot Brucella in a .imple chendcally defined lled1.ua. Proc. Soc. Exper. \:1101. tied. ,21 )10-)11.
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t .. oDonald, J. F. and V. E. Levine 1926 studies in uric acid _tabol1.aa The produotion ot uric aoid b7 bacteria. !Mr. J. PhTaiol. 78,4)7-448. -
!~clnt::rre, C. A. 1936 A cue of blood at.ream intection vi tb Feoal18 a.loal1pnea and laarked hepatitis. M1l1t.al7 Surgeon lfai4O'-143.
MtGowan, J. P. 1911 Some observations on a lahorator,y epldeaic, principally among dOl. and oats, in whioh the aniJula affected pre.anted the aymptoaa of the dide.s8 called It(l1~peru. J. Path. Bact. 1$. )72-426. -
:-ligula, \i. 1887 Schi'JII~cete8 (Bact.ran). Diti tiaturliehen pnanzentudl1en, begrundet A. :Engler unci K. Prantl. ""tiips!g, 1 ten, 10 .St. A. (1900) pages 2-1).
Migula, w. 1900 SZ.tea!!!!: ~aktenen. Zveit8r Band. Gustav'iacher, Jena.
MUe .. , E. Ii. 19S0 Red-leg in tree-trogs caused b7 Bueteri_ ~al!ene •• J. Gen. i"1wobiol. 414)4-4)6. -
Miles, E. M. and A. A. ~Ule9 19$1 The ident.ity of Proteus ~b1l\lS 3ergey !l ale and Proteus aelanO!OJ!nes MUes ana a , aiil their relation to~. genus leroilOnaa ~UYVU' Il.nd van Niel. J. Qen. Microblol. as 296-)06.
Moni .. , E. L. 1928 Clusif'icat1on ot Bacwl"iUlll alcal11.."}nes, I?lo!l!!!!- and ~uon8c.n!.. J. lnt, Dis. ~ajSO:j34.
MontestrllC, E., 11:. lagusin at P. Cal.ibet 1947 Etiologie de. dyeent.erles • 1a t4artineque. aula 500. Path. !~xot. !!Q.s)27-132.
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~udd, S. and S. warren 1923 A reMily cultivable vibrio, filterable through berketeld .. ,rtf candles, Vibrio peroolans (new specie.). J. Bact. 8.447-458. -
Munoz, J., 1<. ~cherago and R. l-i.~..jGaftr 1949 A .orolol1c81 studT ot _Jabera ot the Pseudomonas pnus. J. Baot. $112$-216. -
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Olsen, E. 1949 studies on the intestinal nora of Wants. Ph. D. the.is. Ejn.... Munksgaard, Copenhapn.
Oppenhe1nlar, R. 1913 Die pyelitis. Zettsohr. t. ttro1og. Ch1ru.g. 1:17-u).
Pactnotti, G. 1913 Anguillulo8i ed anchUoatoa1a81 d1ttuaa a1 part di alouns iItAlatt1e !ntetti .... per alitdntaalort.e di orulli crud1.. Zentr. 8alct. raras1t.enk., 1 ~bt. Ret.rate '71l42. -
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Petruacbky, J. 1896 B_cUlwa faeca1ia al!!Henes (n. ap.). ZentZ'. But. Paras1tenk., 1 Abt. 191IS1-1~1. " -
Petrwsoblq', J. 1902 V"Buche sur speo1t1.chen Bebandlung d.. 'l)'phu abdoudnalia. Zeit-scbr. t. Hy'g. 40:561-594, (see pap 57). -
Piorkowski 1906 Zur D1tt.rer~ierung des Typhus bacillus und Bacillus taecaU... Zentr. But. Paraaitenk., I Abt. Orig. 40:4)'=4)'. -
P1rot, R. et M. Bourgain 1942 Orohite a Bacillus alkali!;r18. faecalia chez le Cobal., au cour. d' un pa.sage d. ~1PJiU8 M'ur : Nouwn. Cause d'elT.v dans l'interpretation du test d. NeUl-Mooaer. Arch. Inst. Pasteur Tunis ~.126-1)1.
Pohl, C. and S. W. Ra)"tIond 1941 Bacillus recalia aloal.!Jenes absce ••• J. Internat. Coll. E:urg. 4'541 .. $41. p -
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Poppe 1911 Zur Frage del' Ubertragung von Drankhe1 'taerre,ern durch Huhnereier Zentr. Bakt. Parasitenk., 1 Abt. Reterate 49d.4$-146. -
.Pulv:l.rent,1, 0. d. 1949 La Produsione Dell t Idrogeno sollorato cia Parte z:.i Batteri. Revista di Bio1ogia 41'507-,09. -
Quintoa, F. and o. Buensueeao 1948 A bacteriological study of infantUe diarrhea among Filipino infants. Acta Medica Ph1lippina 2.,13-18.
Raeburn, C. 1944 An enteric-like Weotion due to !!:. faecalia alkalienee. 'BDy. Anq Med. Corps., J. ~'lSl .. 1$2.
Raftne1, tie P. 1921 Lesions ot typhoid teft%' produced b7 Bacl11ua taecal!.-
211
!lkaligCn8s. J. j,mer. Med. Assoc. 76;720-12.1. -Ridder 1909 Beitrag aur Fraga der Aetiolog1e dar FleiachYergittungen.
Flerlill8;' Klin. \;~chnachr. 22:2232-22)4.
Rocha.1x, A. at A. Blanchard 1920 BacUlua taecalis a1cal!rne., Agent Pa.thosena son Role en Pe.t.1iOlogle Hwaaliii. Inii8l •• a. Medec1ne 6:113-184. -
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Rochaix, A. at H. t1arrotte 1916 Sacillus taecalu aloal.!senas, agent pathogene. Compt. Rend. aoo. bIoI. I2:dI~-3I9.
Ro1ard, F. et D. Bourdon 1949 Techniqi18 Dt identiticat.ion Rapid. des Enterobacteriaceae. Ann. !nat. l>asteur 12.,346-3h9.
Row, Y. s. 1926 A eaae of rever due to '3aolllus luc,li. alkali,eMS. Indian Med. Galette .2!.1289-290.
ROlansky, R. at ale 1950 Production of hydrogen sul.tide by P.eudOll1onas &clljij'iDosa. ..\m.er. J. Cl:in Path. ~'l090-1091.
Rush, J. M. 1947 A taxonomic study ot the lHsophilic AchrOlllObacte:..-. Ph. D. thosis, Purdue Universi1;r. •
Rustig1an R. and C. A. Stuart 1941 Decomposition ot Urea by ?roteua. froc. Soc. Exper. 9iol. Ned. 471108-112. -
Rustigian !t. am C. A. Etuart 194) Taxonomic relationships in the ,enua Pro~ua. !'roc. Soc. Exper. :8iol. !I4ed. 2J.1241-243.
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Salvin, S. 9. and ~f. L. Levie 1946 External otitis, with additional studies on the genu. PseudoaoMlh J. Bact. 2!z495-506.
Sandiford, P. R. 1937 Oilservatiorus on :?seudoIllonas pY0!:lanea. J. Path. :9&ot. 44,$67-$72. -
Schaub, I. 0. and F. ~. Hauber 1948 A bioohemical and serological study ot • group ot identical unidentifiable ar ___ negat1ve bacilli tram human scurces. J. Bact. 561379-365. -
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SchottJsull.r, H. 1911 D1e typho.en Erkrankunpn. (Found in the text Bandbuch der Inneren Med1s1n, ereter Sand, Wektionskrankhei ten. a.riG, Verlag von JuIlue Springer, pap 572, chart pages 576-577.)
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Sa1een, W. A. and C. N. Stark 1943 SCM character1at1ca ot green-fluorescent pipallnt-producil'll baoteria. J. Bact. ~'491-500.
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Shearman, C. H. and '!'. G. Moorhea1 ~~aeQalis alcaligene~.
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Shearman, C. H. and T. G. Moornead 1917 Baoi11aud.a due to infection with !!. laecal!!, alcalien95,. H01u1 Anq Hed. Corps., J. ~:104-112.
Shippen, L. P. 1914 Principal Types or micro-orrenisms in Baltiaore .adlk. Johns :iopkins Hospital. 13ulletin &,122-128.
ShrevsbUl7t J.P. D. and G. J. r_son 1947 A contribution to the study of the bacterial nora of arion at,:,. .?roe. 50c. Applied Baot. 2.70 .. 16. -
Shrevaburr, J. F. n. and G. J. darson 1948 It. stud,. of the intestinal ilor .. of Slatell. amer1cmna. J. Path. Bact. 60,$06-$09 • ............... ........"..... ...... .....
Shrewsbur,r, J. F. ~. and o. J. Barson 1952 A bacteriological stud,y ot the Aoua8 sperrow, Passer dOMst1cNe d0fl88tious. J. Path. Bact. ~1605-618. ·
Shulman, B. H. and ft. S. Johnson 194U A case ot .uningitis in a pre_tun infant due to a proteolytic gram-nolj;ative bacillus. fl. Lab. C11r~. Ked. ~1500 ... 501.
Signorine, L. F. 1951 ~·enta.t1vi <.11 ieol ... nto d1 shigdUe (B. diasenwrici) dalle teei di lattanti attetti de. s1ndrwu enterit1ohe. Riv. eli:l. Ped1at. 49IUl1-424. -
Si;lUJlOnds, s. and .1. s. l'"'rutou 1949 A l41C1~gan1sm exhib1ting & growth requirement for peptides. Scienoe !Q2:561-~62.
S1Daonds, s. a.nd J. S .. Frutoll 1951 The lict10n or Penicillin on the growth
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responae of a gro-negatiw baoillu8 to udno acid. and. peptide •• Yale J. Blol. Med .. ll:401-lt26.
Slobo.iano, H. at M. Nasto 1931 Peritonite toetale • Sao1llua tecalia aloallgenes. Compt. rend. 800. BioI. !E!;8S1<:8$i.
Smith, D. T. and W. F. Conant 1952 Zinsnr'8 Textbook ot Baoter101oll. 10th ed. App1eton-Gentury-Crofis, fnc., fi.v Yon.
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Spanedda, A •• L. Aru 1946 Risult.a:t.i di In.da€1ni eaocul1.urall in malati dl forme tifoidi .. si."l'l.Utlt'oidi. Giorn. di B.attoriol. a lMun. J!' 251-210.
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Straub, H. and ~. Krais 1914 Bacillus taeca11a alcal.i,enea ala K'ra.nkheitserr.ger. Deutsche .d. ~Cliii8Chr. fio,)8l)-jB~. -
Strecker, J. 1917 Unter Suohungen uber oacterium alcal;pnea L und N (nacll1u.a taecalis alcaltcenee f1etruachkY j. !naUe,'""Ul"cl Dissertation a i t:'nlveraIl1 0'1 k'urz6urg.
Stuart, C. A., E. 'ftl1 strat'Wll and R. Ruat1ailm 1945 l'Ul"tHr studie. 011 urea. .. p~uot10n b7 Proteu. a.M Nlated orBani.... J. Bact. h2a431-444.
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Taka7anag1, o. 1928 Agglutinatorische EinteUung von aacUlus laecalli alkal!i.ne~h Zentr. Rut. Parasitenk., 1 Abt. arti. !2!t'3S'-~87.
Tataro!t, D. 1891 1'l1e '!)orpnwr Wasserbacterien. Inaugural Dissertation Uni"lVsitat au norpat.
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ThOOlPson, L. 192" Synthetic :ied1ums in the identification of tbe typhoid paratyphoid bacteria. .1. Int. Disease ~'16-20.
Thouon, Jt> G. &too l~. ;'. Hirst 1918 Reports trona. the pathologiOAl. labOl"at.or-itt¥ ot ;010. Gttneral Hospital, Aluandria. La."l.cet 1.$66-567. - -
Titts1er, R. P. and 1. A. f:andholser 19:;6 The Wle ot 8ent.i-eol1d apl' for the detection of bacterial motility. J. Bact. J!:31S .. )80.
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Traudort, R. 1905 TJphuab&z1llua lUld Baa1l.lW!l t&eoalis .~en.~7V81 nicht vervandte Speaies. Munchen. iiiid. \10&;:'&. ii-1 .
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~t:ld1es on '1aeillus pert.u8,18. ,lat->. J. Expel', Bed. 16. -~atson, o. P. 1921 J;. note on tour C8ses of infection with BQcillus raeca1i8
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alkal1;&enes. Royal.4.rIq t~ed. Corps, J • .ll114.3-145.
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..
APPROl'AL SHEET .
The dissertation submitted bY' Rudolph Hugh has been rand A."1d
approved bY' t.hree members of the Department of ~icl"obioloU and tour faculty
.flbers outa1de of the tteputraent of Hierobiology.
The tinal copies have 'been examined by the director of the
dissertation and the signature which appears below verities the tact that
an;y necesaary changes haw been incorpora.tfid, and that tile t.hesis is now
given final approyal with reterence to content, forn, and mechanical
aecur"CT_
The thesis is tr.erefore accepted i.n partial fult1lbent of the
requirements tor th4t Tiegree of Doctor of Philosophy.
u
Date Signature of SponaOl'
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