to dr. fred s. oreutt for his may helpful amd atimlsting rfro the nbve short und admittedly...
TRANSCRIPT
SHJDIEEJ IN wa: PEMBIOLOGY OF GENUE BAMLLUSn A
br
Millen B. Coffee\
M
A Thesis Submitted to the Graduate Cmmittee in hrt ia}. Fulfillment
M ot the Requirements for the Begree uf
‘ _’ ?»€Aä'£‘£;8 GF LEACIHIEE ·
§w>T;?RlOLOGY A __
Approved: ·
l' V- V
In Gherge ot Major work Head- ot Department g
Dean 'o? Ägrioulture Applied Science
Virginia Polyteohnio Institute
1948
”AC¥3ä8§’LEDGB¥.$EHT w
The writer wiebee to erprene bis •1¤¤•x·• sppreciatiou M
to Dr. Fred S. Oreutt for his may helpful amd atimlsting r
suggestioan which mde ccmplstioa M this wart e possibilitp
TABLE Ü? 9OH"I'FtNTB E ‘ VE
V E
PageIHTRODUZTIOH••••2••„6•••„A••••••„„•„•••••„•••••••••• 1
}:2LPk1ZI!S!£·ü§’£'A1. ........Q.....Q.............................. 6V V E D1$SCU3$IGN ••••••p••••Vg•••••••”••••••••••p••d•••••••••••••• 184
CONCLUSIQNS •„„•••;.;2••'.•;„•4•.B;•”••2•2••••„•••••••••„••217..220
RKF§IR¤é·T}2;S ......Z.fg.......................Q........ 222 V .
LIST OF TABLESI T
—P¤e¤
I. source und tsorpbological charnatertstics of Bscilli _
Tosted ................................................7II.Gnrbohydrnte Feruentations Using Duplloate Tubes
ofBros:Ihynol Blue Indie-ster • actual pw values .......A 80 4III. Vernentation Reaction: Using Duplioate Tubes of Bros u G
Thyuol Blue ·”Int•rpr•tat1ons ...„........1........;.. 80
IV. Fersrentstion Resotions Using ho Indieators · actuala~-pH Values ............................................ .
V. vermentation Resctions Using Tao Indioators ··Interpretation: ..................................... 108
TQ V
VI. various Phyeiological Heactions of lßacilli Teeted ...188VII.
rroduotion of ao•tyl·¤stbyl·carbino1 and Reaction on BT
Peptons ............................................. 134”·U
VIII. List of abbreviatione Used in Figarss 1 39 .......1441Ix.carbohydrates in Order of fernentability ............V 191
IQX.Rsactions of additional Strains of theQ.terionsGroup on Various Garbobydrates ..............._ 800ÜXI.
Q. uesenterioue Group · Ron•·Biffer•¤tia1 Rsaotione .. 207
GXII.Q. nssenterious Group · Unrsllable Differential c U V
Renetime ............................................ 808;
XIII. Q. uesenterious Group ·· Rsactioas on aesoulin .......810XIV.Q. mesenterlcus Group · Reliebls Differential
Rsnotxons ........................................... zll T
° WuXV, Key tor tbe Dirterestistioa ot Msunbere ot the g, 4 Q
ÄCÜCHÜCÜIUÜC ••••••••••••••••••••••••••••'••••>
XVI, comparison ot Results mtb Various Autborities ,,,, 214 R
2 4LIST 02* 2IOUR;m
1„ M ?"¤6¤
1. jl. albolaatxs ’I••x. .................................. 214522. _§_._£Q_Y__!_Q_'}„°lI• ........................... xu Q .2 Q3. Q. aatbraaie hx. ..............................._.... 147
A
· 4.—= Q. alterrimuejääia. .............................. 148· ‘ · l
149-_ N 6. Q. g_g_;_·_•_up__?>¤1ttxj7ha¤¤ Tax. 150 V.
Q. QQQQQQ .62.. 24.. Q. 151 * 26. Q. 1+. ::4.121d..................
I162
.86.md............ 155 4-
~ 10. Q. caaglane Serlee Wie. ............................ 1544 4 ·
1.1. Q. aahaerans §840 Taz. ........._..................... 8155
12. Qlelleubachaueis $847 Tax. ......°...................15616.Q. 51051;;
1*$*356 Ms. ...............................
15714.Q. Qualen #615-Tax. ............................. 158 4 8
15. Q; @vo¤1m•·:¢r£g¤t‘6i•. ........................... 159 L
.15. Q. .,12,¤,,•§3,9, $818 6.:. ................................. xda { ·17. Q. lactiaola. Q?876 Tax. .............................. 161 Q18. Q. EÜQÜBGPÄHN-TOI• .................................· 162 _ _ ‘
19. Q. maätherium$@404
Ta:. ............................ 165 4
20. Qlmamtherhm 6. E'. 8*% His. ......................... 16421.. Q.-mesautariaua 'Pex. ................................ 165 I 1
22. Q. masentericusä. T. G. M!. ....................... 166 V6 ' ~
23. gg. mgooidas 1. T. G. $89 ‘I‘•:. ..........._............ 167Mg
· 15 24. gg. _g_g§_g_£*1§22ß':‘»1•. .................................. 1651 125. Q. Eraalvd Ta:. ...................................
5169 ‘
26. Q. __1’Q_§_!_1__1_; 2946 Tax. ..................................4_
190
27. Q. *1%:. ............................._......... 171 g
28. Q. ruminatua Tax. .........„......................... 172 189. Q. silvatioua M5? *1%:. ............................. 175 1 1
130. Q. •h¤g1•: {555 Tu. ................................17431.
Q. aubt11i.•ggKo¤h·N¤ty·T¤x. .......................... 175 Q ‘ 1
38. Qpsubtilis h=1arb¤x¥g‘1%x. ............................I 176 g
T7 _$
33. Tax. _....................... 177vg
. 11 Va
· 34.iQ.
subtuis $131 *1%}:. ...............................17855.§_•-m·bt1g11|‘g§jrbur5g‘3«;‘i|. ............................ 179 * .1
2!•6.}“‘ Tgoaßoke $011•g¤ ................... 180 * ‘
37... gg. tuma6•¤• $995 Tb:. .............................. 181 —g 1
58. Q. vu1g;§u• 1%:. ........·...............·............. 18239.lDo Lang S«tx·:1¤ ?‘¤¤t1¤ Fnnwntur Ma. 185
IN TSEJET FMYSIGLGGY O? ämévxä am 11.LUa
INTRODU3Tl0ä p
The importance er the gmane gggggggs has lang been recog•ju)
nined not only because ot the variety ct reections produced by the
group in its natural nnhitat, tne coll, but aleo because ot the ,ability ot the group to term sporee. This letter tecnlty greetly ~
lncreanee the longevlty of this group and coneequently their h¤•e ‘l
portanc•„ This eharacterletic has tor any years been used ae a( primary point er ditrerentiatlon in ettennte to claesitythisvery
large group of organlene. y u ._ ( ,( The earlieet’classltieat1one, nach es that or Mlgula (23),
eere the work of botaniets and ßllßßfl ot other branchen ot biology
ehe were attracted to the noe science or bacteriology end ehe
carrled over many of the ldeae ot cleaslftcatlou from their.prev1•
cue fields of interest. These clnsslftcationeßnatnrelly etroeeed °
eorphclogy, both in major points ct dttrerontietion end in the
g acparatlon ot epeaiee. äoeever, es the science eg baoterlology E
advanced lt became inoreasinnly apparent that aorphologicsl cherneä
terlstics could net alone be used ae the heels ot seperation. This._~,
e(lead to c search for other·nmsns of differetiation more eiteble
to bacterla alone. ‘Besidea cellular morphology, certain other cheracterletlos
A
2of
mlcroorgnnlse have been used in eftorte to bring some erde:out of the vast collection of epeciee in the genue Beci11u••
These lnclude eclony and epore morphology, tye cr epore gerat-
nation, motillty, pigment production and certainnhyeiclogicalr•act1one•The argunente se to the value ot many ot these „
h charecteriettce es e beate tor eeparetloa have continued down to(the present date. FA Chester (3,t) eas probably the first investigator in T
thle contry to attept e classification of the bacllli. He
eennaturellyinfluenced by Euronen sorkers and although he
dtdutllleethe fernentation cr glueoe, eucroee end lactoee in hle
charaeterlsetion ot epeciee, these blochemlcellreaetxone eere not
used ln eeperatlng the major epeeiee ot the group. j g
Ford (9,10) tclloeed up the verk of Chester end addedgreatly to the knoeledge then extent ot this group• In general he e
h made about the name use ot phyelologieel rsectione es did Bheeter•
seen (5), dealing almost enclueively with eoil herne forms, has
also made dietinct contributions to the texcaoey of the group.
üuring all of this time and even more so during the lest fee years,
the value er phyeiology es a basis for the eeeerction ot thin
group lee become increaeingly apperent.
heny of the baeee herstofore need have been attached by
various authoritleet Colony orphology has been ahnen by Lewis(
(21, 22), Flynn end Hettger (ll), äteern end Steern (53, 34) end
5Gordon(12) to very enoraouely with or without alight changes ln ( .
environmental conditions. Cellular norphology, long thought to be ,~
a most conetant quality, has been shown by aettger end Gilleepie
(E5) to be denandent largnly on the condltionr under which the
organiam le cu1tivnted„
_althoughLehmann (18, 19,(ÄO)’believes that the type of
epore germlnation has taxonomic value, Henrici (lb) end Fort (9,
10) came to the conelnelon that this charecternatie_eaa not
·stableenough for classification D¤P¤9¤oe,„Rnay•1 (16) found _F pignsnt production to be entlrelyudependent nnen the obnditlons
ot groeth such an conpoeltion of the media and temperature of in-~
cuhation„ although änayei helievee that notillty ia a eonetant,(
characteristic ct the hacilli, this heller ie not shared byotherIOTXQIÜ
euch es Steorn (53). · _
Lewis (31) in hle work on4§t gzooidse, uoted tht
althugh colony norphology es wall ae other morphologioal characters Ü
varled with elight changes in envlronasntal connitioue or even 4 C
without any such oiterution, no very profound changee occurred in ny
phyelologieal oheracter1etioe• This conetancy of physlologloalN
characterietics has also been conflrned hy Flynn and hettger (11)} e
aa a reeult or recent work by de Sorlano (29, 30)
andFitzgerald(B), the value of bicehemical reaetione in claseiflcatlon
ot auch a heterogenoue group ae the bacilli has been greatly en•
hanced• ßuchanen (2) even none e far aa tc eay that morphclcgy
clone should not he the eole aals of difterentiatxon of genern•
He concluded that n reetriution of freedom in the choice of
crlterla for dlfferentiation er genere ie uneise, and would ln- t
terfere serloualy with a eatletectory natural grouplng of nierenh
t organisch. hell (14) has crttioleed the nee or bebitat es c
nenne er eeperatxon end telieves that phyeioleglcsl reectxons e ·_
should be used forspeciee‘
Recontly Lenanna (19, 20) has endeavored ta uttliseeerologicel reectione tor texononlc purnocee• ‘h1th0gh he claloe
. none success, it will he many year! before a eatietnetory schema
can be worked out for the group es a whole. The factor of ia- .
procticubility also ie important here. Grantlng that e successful u
key wan found on the basis ot eerology, tte use ae far es the
ordinary bacteriologiet ie concerncd would be decldedly llm1ted•A
Fro the nbve short und admittedly inecmplete resume, h
xt is epparent that the phyeiology ot the genus äncillus is being greeorted to more und more to expleln certain dieorepencies in
taxnnoy which have accunlated during the het tes dececee
through the une of variable morphcloglcel und cultural character•
xetios. For example, de öoriano (30) in testing 500 etrclne of
becilli, found that only one half of these could be elaeslficdonthe
basis of already exieting deecrintlons in the literature.
äeule (32) in hle excellent review ot gp eubtllls tlluetrates the g n _ gdieagreemnt ot numeroua authors as to the exact identity er the
5 e
type epeeies cf the genau Bas1l1us• He qnnten inumerable ex-
smplee to show that what has been en1led_§• nubtilin in the past
es well es the present ie shown te very greetly in oellular end
nolonial eorphology and mthode ot epore gereintion• These
veriatiens are so great es to indieate that at least two dietiaotl
spanien are et present call•d_§• euht1l1••lIt
in with a viee towards adding sonewhat to our
knowledge ot the bioehemlntry st thin greup es well an trying tsd
ezplnin nee of the eeeming incempetihilitiee in the phyeiologieal
reactiona already repertea that the present work was uud•rteke¤• T
A an ettemot ue nsede to ynther together n repreeentntlve( group of the members of thegenun hecilhlue. while xt eas reeognlxed " 4
U ne an impoeeloility te obteln oultnree of every npeeiee in ex-U
ietenoe, the authorvbelieven that the group need ie, ee n ehole, ( L
eell repreenntetive of the gonna. A total of 76 onltoree uns •m· U
played conelntlng of 53 froh me University of Taxen, 22 Iren the
University ef ainooonin, und one American Culture (rm)
Roaeoke Co11•g•• Thin group eomprieed 29 different np•eie•• ~A collection, ehileeonteining e exde variety of the epeesee lxeted l
AU
in isergeyw annual (1), in enpeelelly rich in etrelne of
thecommonepeo1ee(•uoh(e• Q. _o__e___rg_g__e_, meäthertg, gg. meenterlogg,U
y _§• gygloidee end
ln mole I is listed the epeoaee name and etrein
claeelfioatton an well ae certain morpbologioel obeervatiomnU U
‘Z‘heae"V1nohlude'”oall dlheneione, eporedixneneions, epore position-
1n eäeee where tt conld be noted, and reeerke es to cell grouping,
prenence (or abeeoee of e hu1g3d( appearanoe atmen still oonttinlug _U
the npore, etc.
Crystal vtelet (13%) for tee minuten een need to stein( (
the celle end moanurements were mda exth an ooulnrmiorometemTheorder of Bergey°e annual (1) eas follovred in giving dimenelonn °
of width before length and all meoeureoonte ure in ·mi¤ro¤n• T (
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T e 15 — T 1 1 1
The orgenieme eere green on plain natrient nger and ezaexined in e gT 1
24 te 36 henre. he many er the eultures failed to yield •aa·••; 1 1s
1in nach a short dneubetion period,”sone sere rsemeeined at periods gextending from tee te tour days.
1 1* 1
'ltxere eas generell: good agreement heteeen the morphe-
g loglcel characters listed in Table I end theee given by her,·:;ey._ ; s1 1
Some eutetanding dieegreemente met be noted, however. The eporee g
of gg. enthracie, gg. ggeveelens, gg. .g._•gj__gg_, ggyleetieels, §_. gjgzg,1 1
1gg. ei_1gglex end gg. tuaeeeens were found te he charter than the geerreepending listings in ?é:ergey'e annual. The epeeeeec gg. ;
1g 1
silvetinue, en the other hand, eere round te be noch longer.;1
The epore dimeneiene of the rather large group
ef_q_g_;e__n_g_oulturee emanined vera eoneietently lens than thosegivenT in the manual. eb. great range in spare eine eas especially noten
g in the nase of gg. geoides, which isin agreemeoteith iäergey. in1
general, it uns found that cell gronplng eos too-vnrinhlogtoany
taxenomie signifieenoe.1 ln cat least one nase, gg. ellenbesheneie;
the; position ef the epore easldianetrieallp oppoaed to that listed.11
g1
g PHYE%lÜlÄ}Ü*IC..°iL\• 'me orgenisme traneferred·_1Aeeveg-el times en
"!>ifco“ Stock Culture äsedium (36) hefore any phyeielogienl tests11
e
were made. lt vue found that this mediaragave exoellent greeth in
the case of these orgeniem diffieult teeultivete.1
Geletin Liguefaetien • Uuplisate tuhee of nutrient hroth containing
A1612;%
gelatin were etebhed and reeulta reeorded after one seek in-
cubation et ?»O°3.e 'ihe culturee were placed in the refrtgerator
for 15 minutes before reading: eere mda. The results are given
tu Table V1. He attenpt eas made to determine the type ot 1lqu••
factien ee this oharaeterietie ie beltaved to be too variable for
taxoncmte purpose:.
0f all 88 epeeiae, only tee eere found to he negattve.
They are Q. style: end gg. egheerieue. also one of the teo streineof Q. __1_j_o__ta__g_e_ failed te ltquefy gelatin.
1
Nltrete Redgctien ·· :’Z~eventy•t«o heur oultures of the baeilll eere g
tested fer nttrata reductloa when green on **Dtfeo'* Fättrets age: (36). T
again tnlthln teet end in all test: following, the culture: ear: in-I
aubated at 30%. It een found that thte temperature eas the epttmum
for meat of the crgantsme end adequate for all. The eulfanilicaotd
dimethyl alphenaphthylamtne test es described lu the S3. A. Ei. 4 .
annual {28) wan employee!. The reeultehf this eork are given in
Table V].f
T V l. A
Bitrttea are produced by the vaet mjerity of the epeoiee
etndied, wenn ergaeteme aotvpreduoing nitrttee are
Q.eohasrene,gg. fueifornia, Q. äreelvei., gg. ruatnetue, gg.ailvetteun, Q. eghaeräeue, end Q. txneeeeeae. Q. yveolene, gg.
megtherium, Q. ezeeenterteus and Q. _1j_e__tg_e;zgg_leere te very in
the ability to reduce nttutee. Une etrain of the large group of T
g. ige}? endg. Lugjgggg orgenieaza een found to be negative es toeitrete reduetion. · A
lv yaeetgl•methll—carb1nol · Therabilaty of the various species to l
produce acetyl•methyl·carbinol from e huffered medium eontsining ·
0.d$ glucoee uns next detcrmined, The medium choeen ess the onerecommended in the S. 1. h. manual (28). Uuplicete tubes nero i¤•
oculnted from fresh culturee and the first one uns exsnlned afterl
two days and the second after four days. this procedure eae
folloeed because of the ability of many organisms to further reduce
scetyl•methyl-carhinol to 8:3 htyleno glycol after several days
inoubeticn even in n well buffered medium; yet for aus slow gre••
ing epecles, tour days lncubaticn 1s necessary before evidence off
seetyl-methyl·oarblnol production is noted,
_ · 9srritt's reagent, containing ¤ä=e1phe naphthol and 40% _ „
ggg, san used. A period of lb minutes in which to allow the lL
rengent tc reset ie recommended before reading the result, This
suggestlon see follosed but an additional 24 hours uns also slloeed
tor both the tec an four days incuhations. The results of thisf
procedure are recorded in Table VII, ehsreae positive and ¤•g«aa«¤
lnterpretatxons are given in ratte VI. _
The various epeoles seem to be rather inceustnnt in their a ”
production of acety1·methyl·cerb1nol, There eas much eertetion,
even among etrains of the same sp•clee• The effieeey of using fourU
‘ days lncubntton in some cases, is sell illuetrated by results ob-
tained slth Q, anthrec1a,_§, mggtherlum, und some verletiee of_§• _
gycoides, This letter spectee, et least sony ct its etretne, also
t 18U )
emply exeaplitlee the value of leaving ünrrltt's reagent overnight.
es e coacrete example, Q. eubtilie Marburg ue. {gives a negative
reaetion after tee days lnoubatlon, a elight reaction after four gdaye incubation when harr1tt'e reagent ie alloeed to act rer 15
0
minutes and a etrongly positive reaction when the culture ie
allowed to ctand overnight. t
Pegtone · The reeetion of all apecies cr bactlli used eas tested
oe 1% "d1roo” peptone. Brom thymol blue und bro creol purple
(1 ml. per liter of medium) nee used as en iudieator er hydrogen
t ion concentration und the medium see adjuetcd tc an initial pH orU
6.8 to 7.0. pH reading: were made et intervale ot twenty·four,
torty~elght, eeventy•t•o hours und one week. The results obteiued
are recorded in TableVII.Fromthis table tt may be round that derinite evidences
er acid production on peptone exist': for Q. elholaetie, Q. _g_l__v_g_}_,
Q. anthracis, Q. ellentvachensia, und Q. ggraalvei. The iurlueace
er this phenomencn on interpretation or rarioute rerazentation0 0
reeutiona elll be dleouseed later.I) S ln l U
Carbohzgrate Fermetstione • The ability of these organime to
tement carbohydratea auch ee pentoeea, hexoeee, nolyhydrio al- Qcohole, dlsaecharidee, glucoeidee, pelysecoharldes and other
fementable eubstancaa aaa determined. me medium consieted
otheptcmeand 0.5% or the carbohydrute and vue adjusted to M1
19
initial pn ot 6.8 to 7.0. sending: were ende after t:enty•four, Vfortyretght, •evanty•tuo hours ßßd one seek. an no asrehacllll
w were nmnng the culture: exeained, nu use ess nude of
Durbanfsrnentatiotubes ss lndicators of gas production. w
In the first experiment, the R3 oultures obtained fron
the University of Tlseonsin sure run in duplicate on various carte-,
1hydrntesuelsg hren thymol blue (1 mlr per liter) es an indieator
of seid preduction. the results of this study ars given in tabley, ll es epproxlmute ph valuee.1 ln a res cases where th: pn reading:
or the duplicate tubes ers not in agreeent, both values are given,
side by side, ln the table.l
V A‘1n Table Ill ure given lnterpretstsoaa for the ph reading:
of Tkblß ll. Any reading below 6.4 sse taken as strongly positive g Vend indiceted by two pluses. Slxgtly positive reeetions, around
6.6, are shown by a single plus und in a tes cases, eher:
acidproductioneas so sllght es to be in doubt, a plus and a minus is
rooorded. lf the orguntsm ourried the ph to 7.0 or above in 24 _
hours, e negative interpretation 1: given.Free u study of these results, the fact beeaeelnoreaslngly
1
obvious that many ot the organxsme were able to produce acid ln
suftioient qusntity to carry the na loser than 6.0, ehioh ls the
lower ließt of bfüß thymol blue.1
—
Therefore, all 76 of the oultures ser: tested on u much
larger variety of carbobydrates than sure eneloyed in experiment 1,e
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27
and three pü indioetore sera used in an effort te determine the
extcnt of both acid end elaali production free the carbehydrate·
peptone mdiun. The indicatore and their ph rnnges_ere an folloeez rBree thymol blue V 6.0 to 7.6
ß—
5Bra creeol purple 5.2 to
6.8Ere¤·cresolgreenl
3.8 to 5.4_ l IA
r After several prelininary runs, brem cresol green·•se
diseerdsd ae it ses obeerved that fee if any ef the bacilli wereV
_l
able to lower the ph by acid production from the csrbohydrstee ueed
to a point below about 5.0. Aa this region conetitutee the upper _
range of hren oreeol green, it eas found to be of little value endbrem creeol purple, which ie rather distinetive in its loser range,
uns taken ae sufficient for pB'• in the region of 5.4.5
The results of testing all 76 becilli on 36 differentearhohydretes are given in Täblß IV se actual ph reluee. Headings
f
were taken es in the previous experiment at one, tee, three und
seven days. ln a·nu¤her of inetancee, the ph in·the two tube;
containing the tee indioetorn eas not the anne. there this unsthe eeee, the reading of the tube containing broithyeol blue ie
nleeye given first, and directly beside it und to the right isl
r given the reading of the hren cresel purple tube.Ql
T
atteupts eere made in several inetanoes to check ths _
results onteinnd hy duplioating the experiment end running check
renctions. thsnever thle see done, the reading for the check
88 J _Table IV ·J Feruentatlon Reaction: Using Two lndicetore • actual ph Values
J Two lndlcatore were need, bron thymol blue end brem creeolv
4 purple. then the ph reading: in the duplioate tubee were in agree-
ment, only one value le recorded. where dieegreement eee neted, the
ph reading of the tube containing brom thynol blue 1: always given 4 T
tiret and dlrectly beelde lt and tc the right 1: given the reading
or the bron oresol purple tube. .
In thcee cases where check reectione eere run, these
values tor the correeoonding iucubation period: are listed in the ~epace directly beneeth the reading: for the original reection. The
explanation above for the tec indlcatore eppllee here also.
The following example will aid in reading the table:
IoU-O2J J g_ J?Orggnlem eg ~• „Be l1°'·i 1. l
Tßxe J Geß2 6.4 6.2 ‘J 6.4 dance creeol purple value: dieagreeing
J 3 6.2 ith.corre:ponding bro thynol blue —J
l 6.l 6.3 values.
J rße ·Ö;‘Ü4 „a Orlgincl Reaction ‘ Check Reaction
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za
reeetion ie given directly henenth the original reection reading
fer the ccrreepending lucubeticn time. hhenever difference:—”
occurred in the two iudicatcr tube: in the check reeetion, the ·
reading of the brom thymol blue tube, es in the ccee cf the original,
in always given first end beeide it end tc the right ie the nl
ccrreeponding reading ef the brom creecl purple tube. where thei ‘_gä ee shown by the tee indicctore, within the range where they
cverlepped, eas idcnticel, only one value iereccrded.Inthe cuee of come of the ¤rgeex«ee't¤e«• see epperent
t inhihiticn due to the becteriestetie action cf either of the twoindicetcre used. This wan purticulcrly noticeeble in the fermen•
tetlone ct Q,_QQ!g;_end_§, eethrncie. Becterioeteeie appears ini i
come ceeee tc he due tc the brom thymol blue end in others tc the
tree creeel purple. Thin inhibiticn did not, hoeever, eccur in
any other eeeee except fer Q, ggranlvei, which ie clcaely relnted‘
te Q, giga}, AB exneinetion cf the tehle of pH reading: for theae
crghnieun will lllustrete this inhlhiticn. __ 7encther difficulty eroee in the ability ct some epeciee
tc decompoee the indiceter. ihie occurred meh more frequently
for brcm thyeol blue, but caeee whereie hren creecl prple uns in- A
volved den else he found. The following crunniene decoepoeenbrom
thymcl blue in e week cr leee ned also hren creeol purple tc e
certain extentz Q. _c__g·_ey_g_ A."E‘.¥72. hie., Q. gzcoidee hrißxt hie.,_Q. gghveclehe mright hie., end_Q. tumeucene ?995 Tex. ühie
ui
A10?f 0
deooepoeition see especially noticeeble when the orgnnians were
gruen on eucrcee. The dye particlee seem to be adeorbed hy theA
baoteria and carried down into the aedlment, for when the tub te
ehnkon, often the yellow or blue color of brom thymol blue returns.
dtill another difficulty wan noted in the oase of such
pigmnt oroducere an Q, ethrrlmue and Q, glggg, where this· ffcheracteriatic tondec to interfere eith oolorinetric nä reading:. ‘
lhowever, the interference wne never eo great that e mistake ot more
’ then two or three tantbs of e H unit would be involved.,
xa Table V are lieted iuterpretetions ae positive erl
fnegative based on the ph values given in Table IV and taking0 0cogniaance ot the reection ot the organism on peptone alone. In-
terpretatione of these results are somewhat different from the
preoeeding table ot interpretation:. In all cases, any ph below
6.0 ie coeidered etrongly positive, any ph between 6.0 and 6.4 is
- considered positive except rer thoee orgnnieme that produce detinite
acid from peptone alone. Flus—¤lnus interpretatione indlcate acid
production just elightly below the pd reading for the correepondingl V
inoutatien period on peptone alone, i.e. cases ebere there eas doubt
ae to whether any acid in exeeee ot that produced free peptone alone
had been derived from the oarhohydrate.
hegntlve reaotions are given for inctancea where the i
atrains carried the ph to 7.0 or above during the riret 48 hour’
incuhation period. This again does not apply to those organisns
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8. vulutu 2 7 18 nu. 1- 1 7.2TO!. 84 Ill'!. + 2 7.3‘ 15 Ixßn + 8 '•5 2 l
84 un. + 1 V ·_ 7.6 7
D• Inq 8 15 • 1 7.5 ·_Putin !•:••¤t•1· 1 84 hm. ·· 2 7.2
13“m•
"‘ v
v•‘I
145thatproduoe acid from peptone‘elo¤e• In theee oaeee, e negative 4
reaetio le given if the phproduced ehen the organiame aregreenon
the oarbohydrnte folloee within the limits of experimental errorQ
# de ' the ph eben the orgeniane are green on peptone alone. e i [
Tb fureher illnetrnte the renotioae of the ornniems 6 y e e .8
the various oeroohydretes and to bring into even ehnrper toene thef
e
diverse factors involved, repreeentative grmphe of all spaciee are _
given. there one graph of a etrain of n certain speoieaunel
illuetrative of the epecies greup ae e whole, it eds deemed nh- · _
necessary to further elaborate the point with additional grnphl¤g•“la
In many canon, however, the diseimilatien of earhohydratee by the#
fvarious etraine of c epeoieo eas so msrkedly different that graphe · e
were prepered tor the nueerene variante of the parent •p•oie•• m
On every graph, in the form of a dotted line, ie given the reeetion’v”
ot the orgnniem 0 neptone alone. In Table VIII are the abbrev1·atiene used in uerking the graph••
Four other caraohydrntee, omely nrnblnie acid,
Ca-levnllnate,meliziteee and B~methy1•d·gyloside, eere also tested 2
in edditio to those listed in Thblh VIII. hoeever, es none of theetraine of eaoilli need shoeed any ability to produoe aeldfreeT
then, they were not included ln the graphs, It eas also deeeed #g
nnneceeaary to prepare gropho tor ga tueiforeie §539 hex•,_§• # o
fneitornie Wright Mu, g. _1_j_o_ge;__n_e_.tI1e•, and Q. sphaerieue #348 ftex. ae none of theee four organiewe indloeted the elighteettendeney to produoe held from any of ßhß 256 different enrbohydratee.
V G L ma. #11: p V L
GListor Ahbraiiations Urea in figuren 1 to 39
Graph IV
Graph II G
G Adoaitcl Adoa.G
V Dulcitol ~ Dai.G
--d·Arab1no•• d•Arab. Erythritol I Ery. G G1•Arab1n0•• 1·Arab. d·Ga1actoa• Gal. G
G _’C•11oh1oae Gsllo. Glucose Glu.Dextria V Da:. ca—g1uco¤at•
Graph IIIG
Graph IV L V
a—msthy1-d a·¤et.·d·g1u. Laotoea p Lac.glucoaids Levuloae Lev.Glyosrol Glycarol Haltoae G Hal.Glyoogan Glyoogaa deännitcl uauattol1·Incs1t¤1 Iuoa. d-Maaaoaa p üaaacaa s 1Iaulta Iaal. a•¤•thy1•d•° a·m•t.-V manaosida _ d•man¤. „
Graph V Graph VI V
V Malibiosa ßal. Storch Sta. 'Ratfiaooa Rar!. Sacrasa Sao.1—Rha¤¤¤¤a Khan. Trahaloae Tra.Salioia Sal. d•Xy1oa• d-xy.d-sorbitol Sorb. 1•Xy1o•• 1-xy. G
I T DISGUGSIOH d T _'
I Seas of the eorphclogical and physiclogical exesinatlons
« crdinarily ueed in clsssifylng the genue Becillus have been inves-t 4.
tigated. In addition, the ability of theae crganiass to feraent $6 4
different cerbohydretse has been deternlned. The lqsortant pointsT
I of these expsrinents will be diecuseed with- a view tcserd enphas1n· t
ing those reactlons which are suitable for prlnry differentiationd
of the genue and an attenpt will be nede to present reesone tor thed
easy cases of dissgreenent in the literature concerning the earbo- t g
T hydrete fersentations of this group.’
IORPHOIDGIGAL.4 The results of the aorphologicel investigationd iudicste that the present use of both colony structure and cell ·
T tore and diesnstcns forprinery differentiaticn should be eodified.‘ These charecteristics tend to very too nuchveith slight changes in
environesntal conditions tc retain their hltherto lnportant place
in texoncsy. whlle certain of these charecteristlce are undoubted·· ‘
ly of inportence, the replecenent of those that are variable by
conetent physiolcgical reectlons would seen to he s logical pro-d
cedure.PHISIOLOGIGAL. The physiologicel reecticns of the group of organ-
iene exnnined, especially the fernentetlon of carbohydrstes, showsf
definlte prcaise of being e very useful tool ln the differentiation
185of
th various epecles of the genus Baclllue. Several new concepts
of the epplloetlon cf these fernentatlons to texoneey wlll be pre- *
eented, especially the effect of the basel aediun on the reeetlon
obtelned.Gelatln
Llguefactlon · Se few of the hacllll fall to llquafy gels-
tln that thls charecterlstlc has little lf any valu ln classlflca-
tion. In those cases where the orgenlae ls unable to produce
gelatlnase, thle feet ls eertslnly lnportant and sets thespeclesoff
free the rast of the group. However, thls ls such a rare
occurrenoe for bacllll ln general, tht only in a very few oaeesl u
does lt have any slgnlflcence. v
Hltrata Reductlon · „Ihe senbers of th genen Beclllus aere found)
to exhlblt narked lnconstsncy ln their ablllty to reduce nltratee.
Varlsblllty anong the stralns of a nueber of specles was quite
netleeable. Also dlstlnct dlsagreaaent was noted when the results
of-these experlnents were ooapared with the results cbtelned by
Fitzgerald (8), Sorlane (89), und those llsted ln Bergsy•s Inusl
(1). In the case of the letter euthorlty an especlal lack of con-1flraetlon was noted ln the case of several speclee for whichposltlve reectlons were obtalnd by the author. At best this teet
can only be ueed for purposes of charecterlzatlon after a partic-
ular apecles has been classlfled on the basis of sera coneletant (
reectlons.
Acetgl-n•tggl·cerhino1 - Here egein the etreine exenined veried _·greetly, even nere eo then in the eaee of nitrete rednction. Thin l.
teet ie not recerded generelly ee being of texonic valne in
p eleeeifying the heeilli ent it hee been propoeed hy nene eorkere. ·8ign1fioent°diegr•enent nee noted when eoeperieon nee ende eith„
the findinge ot other eorhere, especially Sorieno (29). Thevalue ef thin teet even fer cheracterietion pupceee le question-eble. It we ehonn thet e positive teet depened very eneh on the
P length of ineuhatlon.tnleo
it ee found that fer one ot the ooenonnethode er testing for production cf ecetyl-eethyl·cerhinol
(
(Brritte reegent), the length cf reection tlne ie ef lnpcrtencein errtving et e poeitive or negative interpretation.
cerhogggggte Fernentatione • The reenlte of te action ot 76( t
etreine of heeilll conprielng 28 different epeelee on 36 cerho-hydretee ie best llluetreted by the eeriee of grephe·in fignree p1 to 39. Thea graphe eerve to ehoetvery_eel1 the exteneive
veriebllity in both qualitative en qnentitetive production cacid tree the eerhohydretee need. while the grephs •¤ee••le••
tt are in e eenee eelf-explenetoryt certain eellenttfeetnree nerit ‘ ‘.
further dineneeton. N ttt···-·- ~ ·4;cwhn the line repreeenting the action of the orgenlen
on e oerteln enger folloee the dotted line repreeenting the eetion
on peptone aloe, within the linite ot experleentel error, no p
187 d _difficult! le experienced in recording a negative interpretation.
lor ls there any reason tor heeltetlon ln deolcring an orgenlen ‘ '
ae etrongly positive ln its ability to fernent e certain carbo-
hydrata when lt la able to lower the pH to 5.2, which ia the loeer
1ln1t_o£ the hren oreecl purple indloator.
In those ceeaa where the ga produced ln the preeenee ctd
the sugar ia only allghtly loner thn the raaction on peptona „
alone, or·eher• acid production reachaa a level cf nbbut 6.0
andreeelnathere tor a week, the netter ct interpretation heconea 2
. nero difficult. Alao ot lnportance la the dalnyod ternentatlon
ot certain auhatancee, a perttculcrly good exnnple being the action
I ct a nunbcr ot organlnne cn glycerol. 2 ph
Dlttlculttes ot interpretation are ancountercd also ln
caeea where the organlaa’cerrlee the p ot the ndin to the palknllne elde tor the first few days, but after a week has elapaed,
acid production Iron the carhohydrete han lcwored the pH to neu-
trality or alxghtly below. there la obvlously fcrnentation here
ae lllustratcd hy acid production to the cxtent ot cverconlng theV d
initial aenonla production trca peptoneg yet the flnl an ot the
ned1un.aftcr one week la neutral cr only allghtly acid. Such ad
reactlon would bo claaaltled aa negative hut wald not truly repro-
oent the tcota of the oaac. _
The action o!_g, gggggglßhdd Tex. (tig; 7) turnlahea a
good example ot the case just cltad ln its reactlon on d-gelactose, 2
lu o e
ca-gluconato,}§h~glucon¤te‘and sucroae. ABDÜHBP excellent exaaple— la turnlehed by the netto of_g, ßßllßßütigüß Tex. (Fig. 21) on,
‘ ‘ p Jd·galactoee, 1-lnonitol und both d- und l-xyloee. After a periodct cnc neck the graphe show that theec erneutes on the particularcrbohydratea are productng acid in ever increeein quantity.
Therefore, to deuotc auch a roacticn an negative ls to dieregardthe true situation. .
In nac caaea it eas oted that an crganta would ahoe s
allght alkali production on peptcne alone, but when certain carbo-hydretee were also preeent, the rate of elkali production aaa in-
”
creesad oe well an the amount produced. Thin pheoaonon ie shownbp·gh cgggglann {Fig. 10). An cxplantlon cf this reoctlon or itsalgnlfioanoe le looking. It night be that chen both carbohydretaund peptone are present, Intermediate producte are fcraed froa theaction cf the organiea cn the tec eubatancea; theee lnteraedlateathen coablnlng in aoae nanner ao that the orgeniaa la evcntally _.teraentig thin coapound rather then the two orlglnlly present.
Qualitative diffcrencee in acid production froa a carbo-hydrete can be acecunted for on the beate that the erneuten dcea
or does not produce the enayaa neoeaaary to fcrant the carbo-
hydrcte. Quantitative difterencee, however, cannot be dealt with
ao easily. The fact that acid production from the carbohydroteand alkali production froa the peptone are in direct cppoeltion A
is in host nasee the underlylng raaaon for divargent, delayed
orahcrrantraacticna.Thun tt becceee lncreaalngly clear that a anaher of p
c carbchydrate farnentations attrtbuted to certain epentea of hanilll
in the past are only nenlfeatations of acid production froa te
protatne or peptone of the basel nediu. On the other hand, the
rapidity with which eany organiaaa produoe aeeonia froa peptone is
undoubtedly a source cf nany false negative reactlona. This ia
especially true in the case cf those organisee that prodnce hut al
sllght aeount of acid froa a particular carbohydrate. conaequantly,
the reaction of the crganien on peptone alone anat be eecertained
efora a proper interpretation can he ende of the action of the a
organiae cn any narhohydrate. ·
Alkäll production free pepton cannot he ueed to fu1ly°
explain the failure cf certain crganteas to carry the nu denn tc
g 5.8 chen green in the preaence of cuhetanoea that they apparently
fernent. Por exaaple, the reactton cn peptone alone le identical
for g. aterrtana §353 lie. (1*13. 4) and gl. ggveclene #615 rex. d a
(Fig. 14); yet in the case cf the ferner, the pn on any carte-)
hydrate ts never taken lover than about 6.0 ehlle the letter orga·
nie producer enough acid free thoee eubatancee that lt fereente A
to carry the pH to et leaet 5.8.
g Here then ia an exaaple ot an apparently xnherent ‘, g
oharactaristic of a speciea that aeeae to llait the aaount of acid
VVV VV 190 ‘ 1yVVVVV
V cauV_produoe_ free eaxfbohydrotost tthle would indlcate not only-
M8 value, but the ueeeaslty of using two lndtoatere auch ae hrenV
thynol oreeol purple La oeteralnlng the produeuez;V
ot acid freu: eorbohyarotce. V V
The relative eaee ot ternentation ot the differenthydretee by the various orgenlaae le above in Table II. the actual
VV V1
munter ot epeclea produoieg acid rraa the carbohydrete la given at 1V
1 1 the right in each case. V V VV
In the oase er aleeeeharldee end triaaceharldee, the Vability of the orpnlan to break the original end elao V
V V
° - the esse with ahloh the orgenua een utllize the products ot this
VV
mittel farneutation are ot · tnpertaneu tor instance, cellohioeeV
(Gluoopyrenoee-4-ß-gluoopyrano·s1de), which contains two glueoee
loleouloß, le attached by twelve speciee, uhila leetoee (clucopy-
V‘ re.¤oae·4--B-galsctopyranoalde) auch contains one glucoee noleoule c 0
‘ d and one galaetoae noleoule, ie ferueutad by only nix epeciem Bath ·V· earbohydretes contain the hate linkege, therefore, the difference
in tarnentahility appears te he the result ot the dltterenoe in
the Qnoloeulee of which the ortglnal nolaeole ia oenpoeed. 1V
Bieeooharidae eueh ae naltoae (ülucapyraneee-4-a—glnco- V
pyranoaloai , sooroee (a·Glueopyrenoe•·l•B-truotofurenoeioe) , und - ·
trehaloue (vgluoopyranoee-l—e•glucopyranoeid•) era readlly hydra-
lyzed hy aoat orgenleaa, ueraae nellbxese (0lucopyranoae—6·a· ‘
7 1911 1 1 1 1
- wublc IX1
71Garbchydr¤t•• in Ordcr ct F•rm•ntab11ity 1
Gluccsc 25 6·Xyl¤• 91L•vu1c•• 24 H••51¤ccaat• 8 7 7 1 1
Trchalosc 34 Rartiacsc > 7”
1Glyccrcl 25 l•xy1c•• 7 7 7
Starck 20 Lactcne 61 171
Dcxtrin 19 Ma11bic¤• 6
Maltou 7 19 1-Rhumo•• 5 1 1 1 1 1
Gucrcac 1 19 1¤u11¤ 7 1417
Galicia 16 · Adcnitcl 2 7 7 07 6•Arab1¤¤•c '157 2 Arabiaic Ac1d1 2 7 7
clyccn 7 15 77 7 a~a•th11•d•g1ucc•16• 2 6 1
d-äaanitcl 151
Exythritcl 7 11 1
C¤l1cb1cs• 12 i·1ac•1tc11
11
6·0a1actc¤•1
712- Dulcitol1
0 __
1—Arabi¤c•• 10 0••lcvu11a•t• 01
dßuaanonn 7 10 7 ••¤•thy1·6·¤a¤ao•1d•07
- d·Sork1to1 10 M•11z1to•• 01
7 Ca•g1ucoautc91
7 B~¤nthy1•d•x71o•!d6 0
1924
1 4
gelectopyranoalde) le tereented by onlycl:. Here again lt eeeae
that those earbchydratee noet easily broken down by the bacllliare ende ep ot ncleculee of engere which are reedllyattacked by ‘4the
orgenlene. where teo nolecclee ot glaeoee er one er glucoee4
4and one er levuloae (tructoee) coaprlce the dleaecheride ae¤1•e¤1¢, 4
ha B 4
teraeatablllty ia high, but when galaotone le one or tec4
_
aoleculen eeking up the dieaceharide, the ability ot ue orgenleae4 ‘
to utlline the eerbohydrete le nerhedly reducedt ·9‘ro• Table IX ,4
e lt will be round that only twelve orgenleae poeceee theabllltyto
utillae gelaetoae ehereea glocoee ie feraentedyby teenty··t1ye Ey
and levuloae by teenty-tour”orgen1eae. ‘!'he4he4 facts tend to con-
txrn the point already ende.4
_—AA 44 44 l „
In the caee ot the trieaceherldee,4;nanely rartinoee B
(2-·B·Fructoturenoee·l·a·glacoae·8-e·galectos1de) and nellzltoee _
(l·a-0lacopyrancee·}2·B·trnctot1xrenoee·6~a-gl¤copyranoaide) , the
line ot reeaonlng eapleyed above apperently cannot be need. Ratt-
lnoee, containing galaetoee, ln utlllaed by seven orgenlaae while4
nelinltcae, containing tec glucoee aa1•ea1•¤ anc one levuloee nole- =
oule, ie attached by none ct the orgenlnne. However, ln the caee B
ct noch nore couple: linkagee, the ablllty ot organlena to break
the original llnkagee le probably aore lnportant end depende upon
their ability to prodnce rather couple: enzyaea.4Bote
anat also be ande ot the dltterencee ln the degree e «
62 193
6H 6
ot sold produeed uhere both the dextro and love rotetory roms ‘ ’
ot a sugar are used. Both torno were tested ln the oase o1'Aara· 6A
A blnoee sad xyloae. Strlklng enuplee ot aeld production tree 2A ·
d•arab1nooe but not l—erablnoee era Q. eohoerene (F1:. ll, „6
Q. global! (Pig. 15), Q. _r9_b__u__1; (Fig. 26) and Q. sage: (1*13. 650), AA 2
shoreae Q. aterrinue (Pig. 4) and the De Ina; etrain of peetln A
. toreentor (Fig. 59) utlllze the lovo rora but uotftho dextro tern.6
Q. coheerena (Fig. ll) utlllzee d·:y1oee bat not. l·zyloe•.6
A8 e general rule, noet organlue that set on both ferne
A . A ot xylooo utlllao the dextro tore auch eore readlly than the lovo A6
tern. 'rho seat extreno exaeple er this gonerellzetlon Als round in6 A6
A the graph ot Q. lvol (F13. 28).6 6 e
A-» Heroln, therefore, lies a probeblo source ot eoao ot the 6 6
dloagreeeenta in the literature aa to the ablllty ot an organlae to6
A tornont auch engere as arablnoee and 6xylo•e. It one author uses6
a t 6
the dextro for: ned another uses the levo fern but nelther aakos A
note ot hle eholoe, then tor eaeee slnllar to those listed above,6
2
dleegreenent ls to be expected. A 6
several eospounda oloeoly related to glueoeo ure tested Q6
6te esoertaln the relative ability ot the organlsns tc 1'ernent then. AResults obteined show that ehereae all orgunlena eere able to ter-
6 6
nent glueoao that ehovod any aotlon on earbohydratee et all, these 2 6=
nene organlane were often unable to utlllao the gluooee xeoleeule6 6
A
H 194 V
when ellght changes were nude in the structure such ae eubetltutlon
ot a eethyl group tor the hydrogen ot the hydroxyl group attached 1
to the number one carbon. __9
Substitution ot e aethyl group nade the glucose soleoule
inpervioue to attack in all except three caeee, nenely, Q.’
(Fig. 2), Q. neeenterioue Tax. (Fig. 21) und Q. eubtllle Harburg
Tex. (Fig. 32). Soee other oesee urhsre elight acid production was
uoted can also be nentioned, but these are not very nuueroue nor
do they appear detinito enough tc be lnportant.I
The action ot the etrcina used was aleo tested on d-nennoee
und e-aethyl-d-nennocide. Hhereee ten epeciee were able to utlllze 9
d-nennoae, only one etrein, Q. _g_e_r_•___u_g wie. (Fig. 9), gave detinlte
acid production on e-nethyl•d-nennoeide. The results ot coepsriaon
ot both dextro and levo roms ct xylcee with B-euethyl-d·xylos1d•
were even more dtanetricel. Here no orgenleee ehcued any inel1na··
tion to utilizs the eltered peltosu.
A eoaewhat larger nunber of crgenisns eppeared tc poeseae 9
the ability to prcduee the euzyne necessary to break doen the cel-
eiua and eodlun ealte ot glyconlc acid. Ueually the organieue that
could utlllae the one could utillue the other. Aa would be expected,
there are exceptlons tc this genereliaation, one being Q. __eQ_r•_g_e_ A
A. T. C:. Me. (Fig. 8). A reearkeble illustration of dlvergence in
ttm action ot a epeclee on these two carbohydretee is etforded by
h 1 e 195 h1
~ ‘_
‘ g. ___:·u_e1__;¤;_g_¤£{vig1; 261. e {h · 9 51A eouewhetAdeleftd.{ernentat1ont¤es neted tor—the two p
9 ferne Jeet nentioned in everel eeses. It appears that theorge—nienneeds tina in which to elehorete the enzynee necessary to 1 9A
Abreek dene theae rather uneonnen earbohydretee, er to proéueever1¤¤t9ee1on1ee which peeeeee the ab1l1ty to eleherate the neces-sary eneynas. h 1l
This lose er th ehillty er celle tc ntilise ae•rte1usuhetaneethat they erdinerlly use, when sltghtqehanea exe ende ·
1in the naleeulea, 16 nt rare le the nnnle ot bielogy, hut appear;11tehe nere er lena e neuen oo¤¤rr•¤ee• hhenerer it ie £¤¤nd~thnthaeilll Pöseeee the ehiltty to {erneut any et these enbetituted
1
1eerhohydretee, they wtll be ueetul in d1t{erentiet1¤g the pnrt1·e 9
9 9 “ eeu1¤r·epe¤19ee, beeauee11tha‘•ejor1ty·et orguniens seen teleek thle »·
ah111ty•I
1 E 1h
n4 ·
‘ hhen ee ehgllty et the hec1111 te {erneut certain 1
hydrexy e1eeha1•—wee eenparad with theirjehillty to {erneut theeorrespending enger, etr1k1ng hut ineneietent resultswereed.
Whereea 25 epeelee fßhhßhelü glueoee, enly ten were able
to“ter¤entthe eerreepondihg hyärexy eleohel, s¤rh1to1.e Even nere14
hetrlking uns the difference in ternentehility between galaetose
its d•r1vet1we,dn1e1tol. Tnelxa apeelee re-1eet1one on gelaetoee while net e eingle one appeared able to
_ ut111ze duleitol.9
19e ’_ T 9 _ Q
In the case ot nennitol, the hydroxy alcohol ot nennoee, A
enhanced ternentability tollowed the reduction of thesldehyde toe
Qthe alcohol group. Aoiäj production eas exhlbited by only ten or• 4
ganiens when in the preeence ot scnnoae wherees thirteeh9
Q , e deeonetrated this ability nnnnitol. 1•*x·o¤.theee results lt lsL eseen that there is no correletion shetscever es to the use bYbeellli ot a sugar and tte hydroxy alcohol ee e source ot energy. T
y Taxononio Sggitlcenoevot Results - In easy cases the grephe9 e 9
druvn tor one etrein ot e epeelea eere tound to be representetive
ot the group ot etreine es e ehole. This ia the oase tor the 9Q E
Q tolleelng organiena: Q. _e_l_g_e_}_, Q. enthreole, Q.eterrime,Q.glcblgii, Q. Qgcoides and Q. _n_gg_:;. tor certainggroupa et _ T
streins euch es Q. _g_e_ge__¤re;, Q. gaveclens, Q. negtherlun, T R
Q. sesenterlcus and Q. eubtllie, where verieblllty eas ncted amongI
the various stralne, separate grapha are given to lllustrete theeee ’
ditterences. Steller lnstancee ot verlatlon snong these speeies A
was also noted by Ritzgerald (8). A discussion ot these eignltl-e
cent ditterenceetolloee.Eight
streina ot Q. _g_•_r_•__gg_ eere eaplcyed end the repa•e—e
sentetlve grephe tor then are resae in Fige. 5 to 9. Fig. 5 re-
presents not only Q. g_g__r•__u_Q Tex., but also Q. Qe__r_•_;_u_e_ ßtook 'tex.,
Q. _g_•£_g_1_g_ {non-notlle) Tex. and Q. _c_e___r_e_u_g_ clerk Tex. This group
ot stralns eas found to agree so cloeely es to carbohydrete „er t
’_ J 197‘
- 1 ( t
te:-ncntatioas that one graph could he drawn to repreaent allotthen.
The other strains varied sera or less and thue separate·
gsphs are included.‘ The majority ot atrains ot Q. _q_gr_qg_g_ seen to prcduee (
eebundantacid from those oarbohydrates that they de ternent. ThisV
generalizatxon ia not adhered to by Q. Smith-Thon and, al- ‘
though to a leeeer extent, by Q. _o_o__r_e_u_e_ die. All etralns exnsined(
(producedacid tree cellcbiose, glucoee, levulcse, ealioin, etarch, »A
—
sueroee, and trehalose although theamount verled in smc cases.
gAllexcept one strsin tereented dextrin end d-nnnttol. Q. _q_¤__r_•_;_¤_g_
Salth-·The¤ appears to he an sberrent etreln but agree: withtheother
seven strsins tor the most part except tor tereentisg norede
aware but produoisg less acid from then. ° Q
aargeys manuel (1) lists Q. ellenbachensia (Fig. 1.*3) ( _
aa a syacaya tor Q. _c_e__r_e_u__g_. It agreee etrikiagly in its physio- (
logical reactions with the majorlty ot Q. _e_g_gQu_g_ strains tested,(
but veries meihly in produciag definite acid free peptcne alone.)
Q. albolaotis (Fig. 1) ie listed by Bergey's Manuel (1) es a ·
variant of Q. _c_g_r_9_¤}_§_. Here, however, extensive dieagreeaent was ‘
noted in the ability to dissinllate various engere, etc. In pro-
ducing acid from peptone, Q. slbolectis resenbles Q. ellenbachenels, (
but not in regards to physlologlcal reactions cn carbohydratee. lt
Q. albolactis in a variant ot Q. _c_e_r_gQg, the eharacterletlcs in
193 Q ‘ Q
which tt dtffera are noe eo meeroue that tt ratee the spectee
etetue tt had tn the previous edttton of aergeyw lenuel. (
Tee etretns of Q. ggveolens were used, one fron Ms-
oonain and one free 'Poxee, end they eere found to dteegreeeppre-ctably
tn their cerhohydrete fernentetlone. This te ln eloee ( e
agreeeent wlth e etetlar ettuetton found by Fttzgerald (8) tor tec
Q. gveolens etretne that he used. Q. Eeveolene le conetdered
Q a variant of Q. tuneecene tn B•rgey*e Manuel (1) end excellent Q „
egreenent wee found between the reections ot the Texas etraln of* Q. greveolene end the etretn of Q. tvneeeena exenined.
Q. laottoola te conetdered tdenttcel etth Q. byBergey•e Hensel (1) end etth the exceptton of eltght eotd produc-
A tion froe eeltcln and ecrbttol by Q. lecttcole, the results cb- ~ ptetned conftre this aesueptton.
(
A Pour etretne of Q. Qggtherlue were teated, two obtetned · I
free the Unlverstty of Texee end tec fron the University of wie-
conein. 'the two Texas atretne agree very clceely end the tec
vteeonstn etralne agree very eleeely, hat the two petra do not
agree in e greet neny tnetsncee when coeperee. The reeotiona
oftheteo wteconatn etralns egreed so oloeely that only onegraphue
prepered for both etrelns. Fttsgereld (8), using three etrelne A
cf Q. eegthertue, elao records eeverel eaeee of dtsegreeeent.(
A soneehet eietler attention tee-(notedfor Q. eeeenterteue.
4199 . b 1
One streln froo Texas and two Iron Ulaoonotaooro onoalnod, Al-
though agroooont aould. bs notod gooorally, on several ougars4
1 oxootly opposito rosottoao ooro obtslnod.
Too strolsa of p_. _r_o__gg_n_o_ tore used, ooe beim theor131•nal
strain lsolatod by Roberto (34) in 1955 at tho University of4„
'toxoa, and the other a Ilooonsla strato. Aooordtng to Roberts (24)
and Borg•y•s Ianual (1), the organloa does not fornaat any of the4
b
oarbohydrateo. This statoaont vos substantlatod by tho strato froa bXisooasla. However, the strato froo Texas showod dolayod fornonta·· 4
tion of praottcally ovory earbohydrato upon which it uns tested.4
1'bls delay was probably duo to the foot that the orgaaloa ns a‘
4 rather slow grooor and oloudtnooo nos often not oboorvod untll 48 1 9
hours had olspsod after lnooulstlon. This oxtroao dtvorsity
1roaotloaoby the Texas straln ls at4'preso¤t unaooonatod for. This
ot:-ain also dlfforod in produoisg an orange plgoont after several ~44
4days growth. _
4
g_. go_r_a;o__lng__l, shloh lo considered very olosolyb relatedto_§_.both so to norphology and pathogonloity, oss found to 1
41
ayoo also as regards phyeiology. 1'hls strlkuag olailarlty aan bo4 4 4 ‘
easily oaan by a oouporison of fig:. 2 and 25.4
4 S1: stroine ot _g_. subtllls nero oxoninod and ouoh a oldo V 1
variation in physlology eas aotod that grapho vors proporod tor all4
14 1
straina. This extensive varlablllty ls not so startliug ln view of4
1
the oontrovorsy still raglng ao to the oxaot identity ofa§_.subtll1o. 1
Thls subject le revleeed in the papers of $0ale (50) and conn (6).
I Although the varloue_g, aubtllleetraln• de net agree es to all o
femeentatlons, nne of then leeer the Q3 nnen below 6.0 ln theé
oase of those earbohydratea that they do attaek._JAbl1lty to pro-
duee large anoants of eeeenle free peptene brings nest poeltlve
reaetlons beck te neutral or la none easee to the elkellnesldeafter
the lapee ef severaldays.,
B. ggooldee Group · Some of the neet interesting results eere ob-
talned rath the large groups of_g. ggooldee etralne. In all,H
nlneteen different varlante eere exanlned. seventeen of these eaesfreu Texas and teo froawlaeoneln. The eelluler and eoloalal ner-phelogy of the nlneteen etralns fron Taxae has been exanlnad ln I Jdetall by Leele (20, 21). For the neet part the reeotlone of all, J
etrelne eere qulte eoaetant. when a ugar eas hydrolysed, deflnlte I
eeld prodaetlo eas neted, the pa belng taken doen te 5.2, und llke— J
else, non-fereentetlon-eee alearly deneted by extenelve elkallpro-.
daetlon. ° y , gDeflnlte laek ef eorrelatlen ana nete, heeever, ln the I
case of galaetose, selleln, eaeroee and trehalose. Jäere both _strongly posltlve sad etrongly negatlve reeetlone ere reeorded.
f
Check reeetlona eere run ln s nanber of eeeee end lt eas fond thatthe etralne that gave held free saerose originally, failed te do so A
_ a fer the duplleate reeotione on the sugar. Reoheehe ea eellein agreed in all
caeee with theoriginal.Iocorrelation or egreeeantuae te the tour segare uponwhich Q. lcoldee etreine varted eo could be fhund. That
rie, the aeae organise did not exhibit eieilar reactionc on galac-
1.
y tose, eeltcin, eucrcee and trehaloee. It appears that variatien re 5 winrthe ability of the ctreine to either produce cr not to produce
r
the ensyaae necessary to disaieilate the tour earbchydratee te due Qto a separate end dietlnet diescciatton in each caee. eQ
rThe unexpected ability of one etrain of Q. coidee,
”
neeely #450 Tex., to etrongly feraent lactose eas co escepticnal a
Q a u to the action of the other ctghteen Vntrelne that the reaction waa
checked three tinee. In all cacee the in the preeence of
lactoee wae carried down to around 5.2 to 5.4. Why this particu-lar variant should ec etrogly fcrnent lactcee while all othere a g rr
«r
ehoeed no evidence ot acid production unexplained. 1r
g4 lewin (21), in hle thorcugh exaalnation ot thece etrainecf Q. ocidee noted a tendency tceerde the production of variant
4 er eeccndary coloniae. Thin eecondary pheee of groeth ie due to Q
the depletion ot preferred nutrients end eubeequent utilizaticn of Q u er u uuused eubetances. The ability to attack unuaed nutrients ie Qacquired through varlatlon due to the specific etlmlue exerted by · A
the euhetencec concerned. sucrcae end peptone es well ae certain u
protein fractione eere found eepecially active in prcsotlng thle
seoondary growth. a _
202~
p(
The subraces established freu secondary oolontee show anenhanced capacity for utillaatlon of the- coupcund to whlch verla·
ticn occurred and de net again prcduce secondary colonlee ln tte .
preeence. These variante appear relatlvely stable. 3
4 This nlght well be the explanetlcn for the deflnlte post- _tive er negative results cbtetned wlth etralna cf Q. ßggctdes en 2 3
engere other than sucrcee, naeely, galactcee, eallcln and trehalose.
The feet that check reactlons cn eellcla fereantetlons ccnfireed 3the original reedlnge would ludlcete stable seeondary colcny forea-
tlon. failure to coufirn the posltlve reactlons on eucrose night
be taken es evidence of the production of adaptive enzynearatherthan
actual secondary cclonlee. ‘
B. aeseatericus tig • The Q. neeenterleus group es listedlng
Bergey•s Ianual (l), coneietc of a umher of different crganiaue
3 which were listed es separate speclee until the publication cf the”
5th edition. Included era auch canon heollll es Q. aterrluuc,Q. globigii, Q. and Q. vgßatus. The cheraoterlstics usedin the past to separate these organlcua are no longer considered
4 (
of sufflcieut lnportenoe to afford then the status of speclcs. Hence
they are given as vcriatlee cf Q. necenterlcua. _ (
( Although Ba!*8¤f°• Manual (1) nekes an ettipt to show that
the crgealaee ecuprislng the group do dlffer in their phyetcleglcal
_ 3 reactlona, only four carbohydrates are llsted and cn two of these,
205 vde
g
all aeehere of the group are positive. tree thegraphe showing
the feraentation reactioes of these orgenisaa, it ie clear that
g they are et variance ln nore then s tee inatancem In ecae eaeeeh
where neabera of the group agree qualitetively in their reectlon_
on s particular carbohydrete, they dieeyee quentitatively. A,
This group hee been exteneively inveetipted by Flynn
and Rettger (ll), Rettger and Gilleepie (25), and cordon (12). AllI
of these workere stress the tehdeney ot the various neabere of theh
a p~oup to give rlse to variant oolcniee which often de not poeseee
aany or the cherecteriettcs pecullarto the parent etrain. $oae·· a
tiaes a galn an the ability to utilize a certain euhetance is
p · noted, in others the orpsin loeee the power er utllizing e h e
particular eutetance.h t W
hIt appears that although there ie ecne evidence tor
grouplng these einilar orgauisas into one rather heterogenoue _ hN
group, there ie also a definite need torah
declsive aethod ot {_
tdifferentlatingthe tour variante as euch, because of their tra-t
t quent occurrence und distinctlve eppeerano•„ As it was felt that
ln sone inotencee, the repreaentatlvec ot the group were feeh
in nuaber to draw definite eonelueione, two addltloul etrains er
g. lvgätue and one streln cf g. Qobgiiwere eeoured. These threeorganieae eere teeted on certain iaportant cerbohydratee where it
t
uns felt additional valuable infornatlon night be cbtaiaed. a
2O‘«,pj° "__„„„g.HlQ6'
The reeultavcf thieeork are gtvenaeih ‘1•able~X.' actual ·ph values are reeorded tollowed hy interpretetlone se to positiveer negetive.4 Here? as ln previous cases where interpretatieee were e wmade, the reactten ot the organlsae an peptcae alone aaa eoneidered-
Th•·eerbahydrat• reectleae ot all eeehera er thls groupwere thoroaghly etueled wlth the idee ef Qlllllßilhß all variablereaetlene and also all cases in ehteh e eartchydret•·eaeeitherdtereented
er not tereented by all five members ot the greap. In t
p T 7 Table II are listed these teraeatahle eutetaaeee upon which the u
organisss were all either posltlve er negative.In Table XII are given the esrhchydrates tree ehloh vari-
able reaetions were ehtaled, the vertehll£ty·b•1¤g notes hy e ’ 7
eepltol V in each case. It can he seen that the group le more thene little lneenetent le their reaetiene en several eeaaon carte- °hgdratee. These tladings appear to he eorrohoreted when it le re-
celled tc elsa the nueher er authors that have reeorded the eeeetng-ly inete tendeney of this sroap toeerd variatlon ln both eelluler
T and celonlal aerpholegy. —7
Free the experhaete of e previous author} Fitzgerald (8);lt aae neted that the aeahere cf this group ahowed etgnltleent p ~devlatlone la the feraentetlen ot aesculln. Theretere, all aeebereef this group eere aleo tested on this sugar in addition to the 36earbohydretee used ln the cae ot all 76 hee1lll• Thea results
I _‘ ‘ . 205
O · I$0E 2 2 2 2 +6; *°°‘¥T"*?»? 2 0 0 0 0 .I.
—€
„ 0 0 Op • • • 20 0 II0**2 9 "2 "2 *2 I 9 "2 "2 9 I 9 *2 *2 9 + 2
6 6 6 6 I6 6 .6 _ 0 0 0 2 0‘
I 9 9 9 9 *2, 9 9 9 9 I *2 9 9 9 I·~:· S °T°‘“'°¤I°¥6 0 0 6 0 0 6 6 0 0- 0 0 I„ *2 9 ” I:3 0 0 .
·· 2. 0 0 0 2b 0 0 0 D-0 2 _ . . - .
E 9 9 9 *2 9 9 9 9 I Iux g
¤°9°°‘tgß6 6 6 I 6 6 6 6 I
I Q 0v-4 Q • ·,¤· ig , 0ä 5, 2 2 2 2 + I
Ié_ *'¤1°¤'[*‘l“1•P 0 0 an 0
-
.° , *2 9 9 9 ‘ .0 0 0 6 I· 0 I
~ •, 0u"‘ ,„„m„_„29 9 9 '2 I 9 9 9 *2 I 9 9 'E 9 ‘I I I
aä 0 I 0 0 b 0 0 0 t~· 0 0 I ,0 0 1*
•~·• . .•¤• . 2 ITVS! OU? w Ü
',_ '__ 2
ss · I£¤2{I•-·• ·_¢z.—·· an 6 _ 2«-1 ma va 6 rt ca 0 6
B I IÜ r A Ü ‘ I
"‘ 2
8 2 3• • • —— ätaß B O0 ‘ gn FI I. I rf • G·I § Ep. · E :5 üb .
3 :5 225 :5 :5 5 2 .
» 4 206
eo2 6„ KX Q Q Q Q 4 Q **1 Q Q g 1* Q 1* *2+'QT 6 6 6 6 6 _ 6 6 6 0 uu us an 4- 4
4 4Q Q Q '1-4 Q Q 1 Q 4°‘°‘°“$?¤ ux 0 «¤ 4- nn wa 0 an 4-
·—: ··: 2 2 +q¤.t¤$$ _ n ua an so -|—
. GJ •-4 rr 0 m N N 0 ‘ ‘ ·.-. T°“‘*·‘°$•V 6 J 6 ef‘
6 el ef 6'
43 2 4Q 4 .g •¤¤¤¤1Iqg•( é es L:—‘¢ ··: -2M ‘ 0 rsÄ nscu ~ Q Q Q Q I Q Q Q Q 4 Q Q Q Q4n ¥3•»‘*H Ü 6 6 6 6 6 6 6 0 na en ¤ —|—
4 vz V Q4 0 0 0 2°*¤$'WTT€'?€ Ä Änr;o
0 4 0 4 ca 4 46 no2 °*°¤¤*?*f·P 5 5 6 J4 6 6 Äefll!}A N vi 6 •··• an va 6 •-• zu so- 6
E E -‘ EE 2 • gd 2 · °·,; ··* °Ö . ,2 QE 6 E av ¤os~· .Q • • • • • • ·Ö If! -4 IR TD QV! "*·¢
W 207Y Y" Y
Z WY raue xx Y Y Y
W g. meeenterxcue G1·oup·§Q¤·¤1tt¤re¤tia1 Reactieu ‘W
.Y
Y Y- ·Gz·o¤g FernauY
GlucoseY Y
= Y Y ;WZ _ YL•vu1o•• Z Y Y Y,Y Y · dääuuuitel
Y-F Y Y
._ „ W * YW'rx·•h.a1o••
YF
Y Groug Do-ee Net Fernuni WYY
Adouitol Y B-¤•thy1•é-Xyloside V
ArabinicA¤1dY Y'
Duleitol
Ca•g}.u¤¤¤at¢ i·I¤o•1to1 ZY
, Cadevulimte Lactoee ; Y Z
Hm-glucounte uelxbiese WY
· F eq··¤et¤y1·¢-gluooeide Melxzitoee Y 1 ·· _
a•¤ethy1·d-manucaide RatfinoeeY 1•Rben¤o••
Y
ace Z ’ 1 ·Table
XII_g•uuantartcua 6x·oup·¥U¤z~•11a§1e 1äJ1ff•x·¢x11;1a1 äzaaeuons Z
• T;2 •• •-• S an 1 ·•• 2 1Z 2 3 8 ¤ ä 3 S'; E 2 ••• •-·• 0 v«·• Q ca M 2 U _S 1 0 H __} 8 •·• 3g sn 2: 2 ·? -·· ··· . Z1 „.Z ’ 1 ·¤ G H ·¤ Ä •¤ M
21;- atarrivme ·• 1- ·-1- V 1- 1 -1- 1- -1- 1-
B• 510111311I • V -1* • -1- ·· -1- -1- -1- •
Fh msauzericus V V V · 1- V -1- V + V1 B• ntger · ~ · -1- e · 1- •· V V
S • • Z Z ·B; vulptus V 1 • V V V -1- 1- V V V
are- presented in Table XIII. ,‘ „ 7 g g'T Q Q
. When the entire neas of data ot date aequlred for thisv group eos sltted to ellnlnateall cases Ihereia the organisse need gv q
‘ v
varied er hall or rgeotlons of all saubere of the group g,”
p vers ideatioal, it waslfound that three oarbohydrates atood out es y g ,T being of taxonoalc eignificanes. Those were eesculln, dextrin sad ‘
starch. The reaetions of the flveorganlsns ou these three earbo··»
pT
hydrates arelietsd in fehle Xlv. It is seen that g. aterrleus ’ sfernents all three, g. ggg fereenta none and the reactlons of ~
dthe others are auch es to readlly lead theaeelvcs to a separetion‘of
the various saubere of the group. g g d
Using these sugars, a key for the identification of the g V,five aeebers of the g. nesenterioue group has been prspared und
isgivenin Table XV. This key hae the advantage of heiß short, con-‘ ciee, and clear-cut in its differentiatioa of the fiveorganisee. T
TIhetherthe reederchoosee to accept Bergey•s classification and precognlze the five orgenisas es ecnstltuting a single speciee elthseveral variante, or chooses teregard all five haoilli aedisiuctspecies,
the value of the key ls ln no dininiahad.°_,
The procedure folloeed in finally arriviag et e satin-
factory eeparation of this particular group of organieee could
probably he applied to other groupsor even to the geaus es n n
whole. It eas felt that in the case of other groups studied, the
_ 210 A7.
6 Thblo XIII 6 A
7 B. mosantartcus 0ro¤p·R•¤¤t10¤• on Aa•¤¤11¤AB. aturrtwus 6.6 6. ¤•••¤t¤r1¤u• 6.6 A_ 7 6.0 6.2 . 5.8 A
7 ' ·6.4 · ” 7 5.8 6 · _7 ‘°‘ 6 +A B. atarrimun 6.6 7 B. masuntaricua 6.7 A
P•••k wie. 5.9 Frost ätrnin 6.0A 6.0 S10. 5.8· 0•8 5•Ü A%— 7 —P
B. at•rr1¤¤• 6.6 6. ntgur 5226 6.8R. Ü. W1!. 5•8 Ü1$• ?•Ü .alo vlo
·
A
+ •B. glubigii 6.8 5. nignr G16 6.8 A§356 w1•. 7.0 w1•. ßtrain 7.0 7
?•Ü w1I• 7.ÜA 7.0 „ 7.0
§• ßißlf 3 6•° AVbgpl Strnln 7.0 Win. 7.0win. 7.0 X ' 7.0 _7•° 76. mnsautnricus 6.6 A 6. vulgmtus 7.2 _
A0.8
K
211
'1\¤b1• XIV3
_g<1_• uasaatcricua é2x·oup·R•1i¤b1• Eitfarautial Eieact ion: 6 Q *
Gt*6 ··* 6: §s'43 U2 1B. atarrinus + + +F6. globigxx · +•B•
IIBGBCOPÄQUI + ° "
Es. niger — •-
3
ß• vulgntua ·• 4- 4-
212'lublo XV
Key tor the Läittereutiatioa of haben
G of the j_3_• meeeutertoue Group G
A• Dextrin-1— -Q8. st•¤·¤h+ 3
G• A••ou11¤+ §_•sterrinunCC.
aesoulin · Q. vulgtusStanz: -
° 3C• Aeeoulin • _§• gobigii
am DextriaB.
sstaroh · . 3
C. Aesou11¤—1— g. useuterieue 3
cc. heuculizx · Q.
repreeentetion of the aenbers in the list of organieee tested unenot large enough ae that valid eonelueiene could be draen to the
*extentof eetully working out a key.
l Gorrelation with Previous Authors - An ettnpt uns ende to nume-rize the reaetions of ell epeoiee of beeilli teeted and eoepere theresults eith thoee of auch previous Iorknra es Bergey (l}, «
Fitzgerald (8) and Soriene (29). mie evmery is preeented in Table
XVI. High peroentegee of agreenent eith hergey in the nase of ellreaetions is indleeted. For the noet part, resotione for organieasere only reeordsd in Bergey after they have been eheeked thoroughlyby e nunber of inveetigetore. A t X t- A
very high pereentagee oft sgreeeenteere eleo noted where ‘A_negative reeetione eere eonpered with äoriano. In the leaee of ‘positive rsactione, however, the nunber egreeing van epproxinetelyebelaneedby the number dieegreeing. A very plensible explenation Xof this diseordaney is the feet that Seriano need Andrade•s indien-
ter in her experteents. Thie indientor in prepared by dleeolvingeoid fuehein in l I lIe0H.
i
It han been ehoen by Fitzgerald (8), that Andradvs
indieator reeeins eolorleee ee far dem the pH seele es pa 5.4 and „ae tar up the eeale es pn 7.5. Free the present etudy of the beeilli,it nee preven that snny of the orgenisne produeing acid free oerbo—
hydretee rail te lower the ph to e point below 6.0. Therefors, 6
2149 ”
V I 1'ubh XVI9
1
Comparison ot Results with Various Luthsritxss
POSITIVE1-
Garbchgdrste Vsrmsutsttcus‘—··
Agrssmsut Dissgrssmmt Agreement Disagrsemsnt Agresmaxt DisagrsemsutBerge; Beggz Yxtgggrsld Fitzgrsld Sorisno Sorisno
40 0 54. ss ( 140 4 4(9745) (8-9%)
· gest;).-m•tbg1•csrb1¤o1, ¤•l•t1¤· ·N1t1·st•
83 6 ( 26 2 29 3(79•5$) (80-7%) (9ß•7$) (7-%) (99/PS) (9-%)
ummxvx aßmsrxois9)’
V ¤s1¥p__¤g1§‘il ta !‘s1·m¤tstgM¤es_)
Ü ·
89 _ 95 72 19 „_ 9 110_ 60 4(85.4%) (14.55) (Bam!) · (·64„'??;·)„ ($5.3%) g
ÄGBt!Ä'l1§$hZ1•6I1’ÜÄß0l.” ¤•1sti¤„ mtrsts V
_ B 0 1 5 6 *7In
, 4(100%) (05%) (l6•'?$) (93•#) (65•5i$) (36•5‘$)
' 815 e A —-
orgenlcne which could not produce enough acid to lower the ul ot6
the ncdtun below 5.4 would be listed es gneptlve for that partl-6
eulsr sugar. Thot this nighthevehgooourred. ct Sor1nno•e
reactions scene highlyprobsble.Acc
es um the«„t‘ect A¤drad••• teexcatar usexrrxe v „elkellne enough to reise the pH ot the nedlun in which tt is en;
ployed et least several tentheguot a pa degree end the reason tor
auch a lack ot ncorrelatlon beoonee norc epperent. ne the hydroul‘_ tone or the solvcnt thue reduce the enount of tree cold in the6eedtun,the aecuracy ot thte nethod ot testing tor acid production g
6
ts oonsequently leeeened. Ihen these sane renotlons are cxenined
using a ncre suitable indieetor meh se broethynol blue, these
diecrepenelee are brought to light.- The course ot the reactlone
te nude even noredletlnct chen e conblnetton ot lndtoatore ie6
enployed such ac bron thynol blue end bron cresol purple, ehlohV· 6
oonbined, cover adequstely a px range Iron 5.8 to 7.6.V
7 6'When
the results obtelned were conpered with __thoee^ ofV 5
Fitzgerald (8), elnoet one hundred peroent egreenont uns found tor 7_
those orgsniene producing acid tzen the cerbohydretee used by both
ot ue. on the other hand, dtetlnctly elgnlticent dleagreeecnt wce _
noted in the case ot those. orpntens taillng to produce acid.
’Allltäßlltitngerald recorded that eone ot the organtena6
e
he worked with prccucec acid froh peptonc alone, he scene to have a 6g
316 a i
aegleeted to anke use of thie·faot in reording his reacticne es
_ · positive or negative; Hi aeployed hrue thynol hdue ae an ind1ca~d
.y; d L
tor of acid production end liste only interpretatione, ot actual d
pa values, of the eneuat of·ecid produced by the hecilli he studied.As s result of this procedure, it could eeee that in e number of ,
inetences where he decleree an ornanlen epeble of produclng acid "
free • carhohydrate, the acid ie actually derived frun hwdrolysie y p
„ of the peptone contained in the beel medion, ‘
the point has been reieed hy Nedun (35) that ln sone
aaes the failure cf an organiee to produce acid free a particular pcarbohydrate does not neceeearily prove that the cerhohydrate uns t
not utilised. It is certainly true that in sone inetances
theproductsof carbohydrate eetaholiaa era not aeidie in nture, hatlß
ney coneiet of alcohola, etc. However, this feet in ne way leesans
the value of acid indicator methode in the classification of
nicroorgenisna. ,_A
The fact that Wed: (Sb) found acidindicetcr·nethodstrusteorthy
in seat inetancee eould indicate that the najority of
organina produce acid as s result of cerhohydrate„netahol1ee•
When the crltaria of acid production and not nerely oarbohydrate
uttllzatica ia taken aa the standard, perfeetly rellahle nethods
of aeparaticn can he shcon to exist.
e h217GDHUEBSIUIB
p 1. The lapnrtane of aerphology ln the claeelflcatlo of Vthe gcnus ggglllue anat be revleeed ln the llght of eork deannetret·
1
lng the extensive varlatlon oeourrlag ln thle characterlstlc of Vo
I
aloroorgenleae. It appears that both eellular and colnnlal aorphology
anat be relegeted to a leee laportant posltlon in the taxonnay of thle 2
genne, especially ln cases where oonetent phyelolcgleal reactlone can_
be eoen to exlct.I I o
V2. Such recotlone ae nltrete reduetlon end ao•tyl·aethyl- V
carhlnl production, ehlch have ben need ln the past for prlanry
eeeretlon, have been ahnen to very too greetly—aaeng dlfferent atrelne
er the aaae epeclec tc be retalned ln thelr preeent laportent'poaltlon
ln tcxonoay. I V·
3. In the aatter of gelatln llquefectlon, eo fee bacllllo
fall to glve e poeltlve renctlo that the teat le of value only for h
charccterlzetlon aftr the orgenleaeae·b•n plaeed ln lte proper
place by other·anre dlfferentlal”reectlone, V‘J"* 3
V4. It hac_h•en clearly deaonetrated that th! Rlctnre of
acid productlon free oarhohydretea le fer fra being e alaple one.”
leny dlverce factore are lneolved end anat ue taken into ooneldera—o
V tion ln draelng onnelnelon froa the reaulte obtelned.5. It eae found tet the action cf the organlaae on the t
I peptcne of the basel aedlua could glve rlae to both falee poaltlve h
818
V and false negative results lf this action te not taken into acconnt.t n
6. The use cf e estee of tndteatcre oonelsttng of et
least two which edequately cover a pH renge free 5.2 to 7.6 is adve-
cated. Only when thle procedure te follceed an well es the nee of _
control tests on the banal nedlnn alone can any lcglcel concluslons
be drawn. 6 e
V 7. A perold of tncnbetton of et least one week le edvoca•‘ted tn deerntntng acld production fron earbohydratcs by bectllt.
This ta to allow the appearance of acid fron those carbohydrates
which are bet sloely fereented.v
B. The 36 dlfferent carhohydrstes tested exhlbtted rather p —
erprlslng degree: of ferenteblltty. In the case cf ecne poly- t
eaccharldes, the netter of ltnkagea and the eonosaceherlde eeleculce4
coepostng the pclyeacohartde see tc gevern-the ability of the
orgentne tc fernent te sbetanee. Steller correletton was not found a
between nany engere and thetr·hydroxy elcchele or ncthyl ebetltuted .n
derivetivee, except that the nejortty of the letter were unfernentable.
9. crgenteee were found to dtffer in their abtltty to e
utillze the dextro er the levo forn of certain sugars. This d1ffer· V
ence ls advanced es an explenatln for eone of the dleegreenents
existing tn the literature with respect tc these particular engere u
where authors have failed to state which f¤:n.wae used.
l¤. A nneher of reter unonnon oarbchydrete: were tested
· 1219 e
~l
and the value of sone of then for texonoele purpeeee eeeas eeteb-llehed. others have definltely been ruled out beoeuee too teeorganiena are able to produoe ecld froh the. a p
U
ll. The present olaeelfloetion of the gene ggelllue ee epresented in the ßth edition of Bergey•a Iennel (1) hee been re- gviewed and eznnplee of eonflrnation have been noted es eel1—aeinetencee where organieas that have been gronped together are de-finltely not related to the ertent lndleated by the grouplng.
12. Nlneteen etretne of_g, ggggtdee eere ezenlned end thereeetlone of these etreine are dleeueeed at length elth poaelbleerplanntione given for their aeenlng lnoeneletenoy ln thefernenta-ttte of eeverel oarhohydretea. other epeoiee are alao dieeuesed, 2
the lnportanee of the oonluslone drewn bein relatively proportionals to the nber of etraine need in the experinnte.
13. The_g, neeenterloue group hae be•n.•xeelned ln the es 1llght of their oerhohydrete fereentetions and e deflnlte eyetee ofoleeelfloetion hee been devleed. This procedure le suggested ee a
* pattern for future work with elnller groupe end eventuelly with the. genue ee e whole. ‘„ In · f . a e
A 14. It le the opinion of the author that lt the proednreneed in these experlnantn le followed for a repreeentetlve Hülbßf
of etralna of all epoelee of haollll, a oleeront, eorkable key forthe eleaeifieetlon ot the gene eould be evolved.
220 l
a ‘!*¤enty—e1ght different epectee cf the genue Becillue havebeen exenlned both ee to norphological characterietice und physio- „logical reactlons. The phyelologicel reacticne tested includegela-tin
llquefacttona, acetyl-eethyl-carblncl production, nitrate reduc—(
tion and 36 different carbohydrate fernentations.A
The action cf' all 78 etretne cf the 28 epeeiee usa also(
deterntned on peptone and grephe have been prepared conparlng thle
action with that of the orgenleee on thecerbohydretes.Definlte
explenetlone are given for eany of the1nccne1s·-tenciee
existing- in the literature for the fernentation reectlone oftea aaaixlx. an es
„. s —Ä The results-ebtained are conpared wtththoee recorded by (N
three previous eorkers, naesly Ültzgerald (8), Sorlenc (29) and ~ ABergey•e Manual (1). Reaeone have been advanced tc explaln caeee
ofdieegreement.are concerned nalnly with the indicntor used endthe failure te take into account the action of the crgenleneunder
investigation upon the banal eedlun.
where it was felt that a repreaentative nnnber of straine
were exanined, definlte conoluaione ae to the ability of the orgnn-·
inne to dleeinilete various carbohydratee are dream.l-
A key for the classification cf the g. eesentericue grouphas been foreulated based antirely cn cerbohydrate fereentaticns.
t The proeedure telloued appears to be lvpliceble to other groups
eswelles to the entire gehue. A ._ h
822 ·, E
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