Pedigrees of Some Strains Escherichia coli K-12 · Thestrain pedigrees are presentedin Charts 1 through 11. The documentation for these diagrams is given in Table 1, underthe strain

BACTMOLoUIcAL REVIEWS, Dec. 1972, p. 525-557Copyright i 1972 American Society for Microbiology

Vol. 36, No. 4Printed in U.S.A.

Pedigrees of Some Mutant Strains ofEscherichia coli K-12

BARBARA J. BACHMANNDepartment of Microbiology, Yale University, School of Medicine, New Haven, Connecticut 06510

INTRODUCTION ................. ............................. 525SOURCES OF THE DATA .............................................. 526PEDIGREE CHARTS ......................... ..................... 526Conventions ............. ................................. 526

Strain designations............................................... 526The genetic symbols............................................... 549Other symbols and abbreviations.......................................... 549The treatment of suppressors ........................................... 550

Comments on Charts ......................... ..................... 550Chart 1. Some early Stanford and Yale strains .......... ................ 550Chart 2. Some derivatives of strain Y1O ............... .................. 550Chart 3. Some derivatives of strain P678 ............. ................... 550Chart 4. Some derivatives of strain AB1 157 ........... .................. 551Chart 5. Some derivatives of strains 58-161 and W6 ........ .............. 551Chart 6. Hfr H thi-, -y- and some of its derivatives ........ ............. 552Chart 7. Some of the early Paris lac- strains ........... ................ 552Chart 8. Other lines derived from wild type ........... .................. 552Chart 9. The derivation of JC12 and JC411 and some of their derivatives . 552Chart 10. Paris strain 200PS and Paris F-lac+..552Chart 11. The Paris F -gal and Frgal ................................... 553Chart 12. Derivation of Garen Pho- and Su+ strains ....... ............. 553Chart 13. The derivations of miscellaneous Hfr strains.................. 553Chart 14. The strains K12S and 112 ........................ 553

DISCUSSION ............. ................................. 553THE CHARTS .............................................. 527LITERATURE CITED ................................ 553

INTRODUCTION

The strain K-12 of Escherichia coli has beenin cultivation in the laboratory for 50 years now.It was isolated from the stool of a convalescentdiphtheria patient in the fall of 1922 (64). Formany years therafter, it served as a standardculture in the bacteriology department of Stan-ford University and was used extensively in theteaching laboratories there (99). That thisvirile strain of E. coli, one of the relatively fewfound to possess significant fertility in thelaboratory (66, 81b, 81c), should have been theone which C. E. Clifton chose to give to E. L.Tatum as the latter set out to produce mutantstrains of bacteria was apparently just a par-ticularly happy accident. K-12 was thought tobe an entirely typical coli culture (but see thework of the Orskovs [81a, 81b, 81c] regardingits antigenic structure.)

Gray and Tatum reported the isolation ofX-ray-induced auxotrophic mutants of bac-teria, including E. coli K-12, in 1944 (35).

These same strains, and others, were used byLederberg and Tatum in their early studies ongenetic recombination in bacteria (69, 70).Since that time, literally thousands of mutantstrains of K-12 have been produced. Theircontribution to the development of molecularbiology is by now well documented and widelyappreciated. In the belief that there is stillmuch to be learned from the study of thismicrobe, we have been involved, over the pastfew years, in setting up a center for thepreservation and dissemination of genetic stockcultures of this organism.When the E. coli Genetic Stock Center was

first set up and we began acquiring strains fromother laboratories, it became apparent that wewere receiving the same mutational event (mu-tant allele) in a variety of strains under avariety of mutant allele designations fromdifferent laboratories. We soon realized that, inorder to assign meaningful and unambiguousmutant allele designations to the mutationscarried in the strains in the collection, it would

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BACHMANN

be necessary to trace the derivation of allincoming strains. The donors of the cultureswere unable' to give us complete derivations fortheir strain6 in most cases, and it proved to beimpossible to trace them completely throughthe published literature. We therefore set outto trace the strains through the unpublishedrecords of the laboratories in which they weremade. This task, which gradually became themajor effort in setting up the Stock Center, hasled finally to a reconstruction of much of thehistory of E. coli K-12. Some of the results ofthis project, which is still going on, are pre-sented here.The pedigrees of K-12 derivatives that are

presented here have been chosen with thefollowing considerations in mind. We havetried to include the derivation of most of thevery early strains of the Stanford, Yale, Wis-consin, and Paris laboratories, which served asancestral stocks for almost all other collectionsand would thus be of the widest possibleinterest. Among later strains, we have includedthose that seem to have been most widely usedas ancestors. And among contemporaneousstrains, we have again chosen those that appearto be widely used in constructing strains forgenetic analysis, based in large part on thefrequency of requests received by the StockCenter. Obvious limitations influencing ourselections have been our lack of experience withstrains from some major collections that wehave not yet explored and, in some cases, ourinability to establish the history of importantstrains due to gaps in laboratory records.

SOURCES OF THE DATAThe strain pedigrees are presented in Charts

1 through 11. The documentation for thesediagrams is given in Table 1, under the straindesignations, listed in alphanumerical order.The ultimate sources of the data were, in

most cases, the laboratory records (strain note-books and strain cards) of the laboratories inwhich the strains were made. I visited theselaboratories and worked with their records inconstructing the strain pedigrees. I wish toassume full responsibility for any misinterpre-tations applied to these data, while acknowl-edging considerable generous assistance fromthe "owners" of the records. In cases of conflictwith reports published in the literature, theselaboratory records were accepted as being cor-rect. These major unpublished sources of dataare given as documentation in Table 1, wherethey are referred to by capital letters, asfollows:

A, Strain notebooks and cards in the labora-tory of J. Lederberg.

B, Strain cards in the laboratory of F. Jacob.C, Strain cards in the laboratory of E. A.

Adelberg.D, Strain notebook of A. L. Taylor.E, Strain records of M. L. Morse.F, Strain list of P. Howard-Flanders.

In other cases, as well, I have called upon theassistance of those who constructed the strains.These investigators, in response to questions,either generously searched their laboratory re-cords and extracted the essential information,or verified pedigrees which I had constructedon the basis of published information andexperience with the strains in question. Theseextensive personal communications are ack-nowledged as sources of data in Table 1, lowercase letters designating data received fromthese investigators, as follows: (a) R. Apple-yard, (b) A. J. Clark, (c) B. D. Davis, (d) A.Garen, (e) W. Hayes, (f) K. B. Low, (g) W.Maas, (h) P. Reeves, (i) P. Treffers (j) N.Willetts, (k) E. Wollman, (1) C. Yanofsky.

In addition to the above unpublished sourcesof data, we have included in Table 1 referencesto published descriptions of strains and theirderivations. These sources are listed underLiterature Cited. In a few cases, where pub-lished reports were sufficiently detailed orwhere we were unable to reach more directsources, citations to the literature are used assole documentation for the pedigrees and straindescriptions.

PEDIGREE CHARTS

ConventionsThe pedigree charts consist of strain descrip-

tions with lines of descent indicated by arrows.The genetic step involved in the production ofmost of the strains was mutation, either spon-taneous or induced. The mutagenic or selectiveagents used are indicated beside the arrows.Relatively few recombinant strains are in-cluded in the charts: in these cases, the selec-tive conditions used in the isolation of recombi-nants (where known) are given beside thearrows indicating these steps. In the caseswhere markers were introduced by transduc-tion, the bacteriophages used and the donorstrains are indicated beside the arrows, e.g., P1from AB1234.

Strain designations. An effort has beenmade to use in the charts, in all cases, theoriginal strain designations assigned by thosewho constructed the strains. Widely used syno-

526 BACTrERIOL. REV.


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PEDIGREES OF SOME E. COLI K12 MUTANT STRAINS

CHART 1. SOYmel - E

mel-isupE57supF68

Y1OF- X-ray 679-8OF+ ( X-ray 679F+thr-i thr-I thr-ileu-6 leu-6thi-isupE44

ITIY46F

Lthr-ileu-6thi-itonA26

679-183F+ X-ray

thr-ipro-37

679-440F X-ray

thr-ipro-38

thr-ileu-6thi-IlacYlunstble

T,sel'n. Y64F

thr-ileu-6thi-llacYllonAt7

uv

EMB-lac Y70Fthr-ileu-6thi-ilacYli.tb1

T,sel'n. ) Y86F

thr-ileu-6thi-ilacYlton-28

Tssel'n. Y94F

thr-ileu-6thi-ilacYIts8-74

T,sel'n. Y100l

thr-ileu-6thi-ilacYIton-29

me early Stanford and Yale strain8E. coli K-12 (X)F+ [wild type]

and

bio-Silu-24

X-ray 58-580F+bio-I

thi-f9

X-ray, 58-593F+bio-ithi-30

X-ray, 58-610F+' bio-i

thi-31

X-ray, 58-741F+' bio-i

his-64

spont. 58-2651F+I bio-i

pro-39

X-ray 58-278F+bio-Iphe-i

Nmustard , Y80F+

Ibio-imetBItonA24glycerol-

Nmustard >Y87F+ spont. W14F+

bio-I bio+metBI metBItonA24 tonAS4lacY4O lacY40

chloro-acetate

sela'n. Y91F+bio-imetBItonA24chloroacetateR

uv ) Y24F+bio-lpAe-icys-47

spont. _58_278MF spont. CS19F+bio-i bio+phe-i phe-imutTI mutTI

VOL. 36, 1972 527

58F+ X-ray + 58-161F+ spont. lW6F+bio-I bio-I > o+

metBI meiBIrel-I

X-ray 58-309F+ sel'n.bio-icys-l Y40F+ spont. W13F+

bio-i-

ho+metBI metBi

X-ray v 58-336F+ tonAs4 tonAS4


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BACHMANN BACTERIOL. REV.

CHART 2. Some derivatives of strain YIOE. coli K-12 (F) X-ray,,y 679F+ -ray 679-68OF- X-ray Y1OF

thr-J thr-1 thr-1leu-6 leu-6

thi-lsupE44

1ENlB-lac ; sel'ri.IOM Joe I""11'~ U

Y531 W102F- W112F- W133F- W208F- W1394F-thr-I thr-J thr-l thr-I fhr-I (hr-Jleu-6 leu-6 leu-6 leu-6 leu-6 leu-6thi-J thi-Iuhio1 i-a [ethi-h [d thi-JsupE44 tialQO5 laceY' loc-7 8up-49 supE44?lacY)I-'1(1b supE44? supEh44? spE44? lacZ4 1str-110

uv N-mnstardEMB-Iac MB-malUV

W2660FY70F- W IF- Id)

thr-I ,1hr-1 Aleu-60 (u-6

[a thi-l [b) thi-I) Fl fromsuapE44 supE44 W1486lacYlIiabl IolY W208, S F-

inal.4 [x le[ s 2817F+T, str-8 Idseln. T, Tiv,

sel'. sel'ro I-MB-mal Fl!C6F- fron

[al CS100F- W48OF- %'894F- wild fVtonA2J [bl [bh [bl type W2924 Hfr3 W2945 Hfr4analAJ,JXR nialAl,Ja0 enJAlA,JXR Id) [di]Jts-75 JonA galKS(gal,) 2Bna:B5 mal-6

sel' 1. A- A-P04 PC)17

C260F- tiv uv 8ala EMB-gil FMB-arai IMB-lac AB284F-+fonAlAB I I I I 'U [elW566F- W904F- W902lsr-8

[bh [bl (bhuv maJA J,O0R malA t,JiR thr-e uv uvtonAS galK2 leu-6

C600F ( CR34) galTli (gal) ara-14 thi-J AB311 Hfr AB312 Hfr AB313 Hfrla] lacJ [el [el [lCton¢A21 uv uv ttalA 1, XR str-8 str-8 str-8-EMB-xyl EMB-xyl galKO Ad.f X- P013

4 4,t Poll P012amioiopterin Fl from W582- W922F

58-161; [bh lbhsponto malA,JI anialA , XR

oyl-7 galKOCR34, thy-1- J4 Hfr (_ PIO) toctAS ara-14

[a] [a] galTI cylOtonAAI lonA1IthyA6 malB16 uv uv

dra-l x- EMB-ara EMB-mtlX~P018W583F- 945F-

[bh [hbmalA 1,XR malA),OXxyl 7 'aiK2ara-e13 i ara-14tow Af [e yl-galiiT,ml 1-

528


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PEDIGREES OF SOME E. COLI K,2 MUTANT STRAINS

CHART 2. (cont'd)

W583F-

IEMB-mtl

W595F-[b]{malA1 ,XR

[fJy-

[4ara-15mtl-StonASgalTIspont.

W660F[b][fItonASgal+

jEMB-gal

W677F+ W677F-[bh Fl from [bh[f] 58-161 [f ItonAS tonASgal-S(galb) gal-S (gal5)

Flfrom Fl from str58-161 58-161 sel'n.spont. 4 spont.

Reeves 2 Hfr Reeves 3 Hfr W1177F-[b] [b] [b][f [f I [fItonAS tonAS tonASgal-S gal-S gal-SPO110 PO111 str-117

1Fl I UV

CSIIF+ W1603F[b] [b][fI [fI

tonASgal-Sstr-117

A-

spont.

4 f

P676P[b][fItonASgal+

P678P[b][f]tonASgal-6 (galb)

1[Chart 3]

W945F-luvJEMB-malW2914F-

[bhmal+[el

strW2915F- sel'n. W2961F-

[b] [bh[el [elproAS proAMA- str-SO

X from11 01 Hfr4000

AB1103F-[b] AB712F-[el [b]proA2 [elhis-4 proAMx- str-2O

UVuv Plkcfromwild type

AB1115F-[b] AB1859F-[el thr+(proAS leu+

[gi< his-4 thi-IargES lacYl

[elpro+

str 8tr-2Osel'n.F< ~~~~~~T4,,T

AB1133F- sel'n.[b][el AB1621F-[g] thi-1str-51 lacYl

[elstr-SO

T6 tfr-5sel'n. t8x-67

AB1157F[b][el[g]str-31tsx-SSte-"

sup-37

[Chart 41

VOL. 36, 1972 529


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CHART 3. Some derivatives of strain P678P678F- [Chart 2]

thr-ileu-6thi-IlacYI

[h malAi,XRxyl-7ara-iSmtl-StonASgal-6 (gal4)

juv

PA678, X-F-[h]A-

uv

Series PA100-PA125F-[h]+ singleauxotrophicmutation

A-uv

14 r

Series PA200-PA266F-[h]his-I+ singleauxotrophicmutation

X7

uv

PAIOOF- (= P697)[h]his-iA-

uv

PA200F-[h]his-iargHI

I strsel'n.

14 r

PA200,SRF-[h]his-IargHIstr-9_-

str Fl fromsel'n. 58-161

P678, SRF- (- 2001w) P678, F+F+[h] [h]sir-i35A

str.sel'n.

PA100,SRF ( P697,SR)[h]his-istr-17

PA100,SR,Pro-F- (-JW1)[h]hi8-istr-17pro-iAC-

Iuv uv

Series PA300-PA394F- PA351F-[h] [h]his-i his-IargHI argHistr-9 str-9+ single purE4Sauxotrophicmutation

uv uv

Series PA601-640F PA61OF-[h] [hihis-I his-iargHI argHIstr-9 str-9purE43 purE43+ single ly's-26auxotropbic x-mutation_-

uv ) PA641 644F-[hihis-iargHI8tr-9purE43ly8-25+ single auxotrophic mutation

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PEDIGREES OF SOME E. COLI KI2 MUTANT STRAINS

CHART 4. Some derivativeg of strain AB1157[Chart 2]

AB1157FEthr-ileu-6thi-IlacYlgalK2ara-14

mtl-iproA2his-4argE3str-31tsx-S8

sup-S7

AB2462F- NG[i]recAiS

AB2470F- NG[i]recB2i

I TRIMsel'n.

JC5408F-[i]recBRIthyA383

P1 fromJC5474thy+

JC5519F-[ii

thy+recB21recCSS

P1 froiAB247thy+

JC5743F[ilthy+recB21

NA AB1884F-[iluvrCS4

NA AB1885F[i]uvrB5

NA AB1886F--* [ii

uvrA6

uv AB1899F-[illon-i

uv AB2457F-[ilirp-85

I uv

AB1896F-[i]uvrA6lon-6

TRIMsel'Pn AB2495F-

[iitrp-SthyAtO

NG AB2463F- thyRlS- [i]

recA13

TRIM TRIIVsel'n. sel'n.

JC5421F AB2487F-[i] [i]thyASS6 recA13recAlt thyA16

drm-1rRIMsel'n. AB2497F-

[ilthyAltthyR14

rRIMleI'n JC5422F-

[i]thyA825

NG | TRIMI sel'n.

AB2474F- AB2500F-[il li]uvrA6 uvrA6lex-1 thyAl5

drm-S

NG spcselin.

AB3022F- KL252F-[ii [i]trp-85 trp-85IthyASO thyA2OthyRIS thyRSOrecC22 spc-lIl

P1 fromAB1157thyA+

JCU74F-[i]trp-85thyA+recC22thyRIS

P1 fromJC5474thy+

JC5489F-thy+YecCR2

P1 fromAB2463cys+

11 r

JC2926F-[ilc1e+recAIS

I

VOL. 36, 1972 531

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K.12 (Xl)F+ [wild type]

X>-rav

58F+bio -i

I X-ray

58-161 F+ spontbio -I

metBI

,,v yW45 F+T biolI

IacZl

Y40F+ spont. W131+to-1 - bio+ntelBBi metBtorA.4 tonAR4

uv W67F+N bio l

mtnstard enetBiIonA24oacZS9

Y87F+ spont. W14F+bto-1 bio0nelBI netBIlonA24 lonAf4lcy40.'t"1. lacY40"bl.oe

W5161F+ EMBlacnoetBliIonAR4lascY40""pur -49

UVW5181 + EMBloo

netBitonA24lacY40"t'tgaIT1S(gal5)X

uvW7501+ EMBlacmetBiionAR4lacY40galTff(gal,l)

uv

W11631'+4 EMBlacsnetBitonAR4lacl'40"'b

JEMBlno

W1210P+metBhtouA24lac Y4OgaLK8(gal,)

CHART 5. Some derivaties of strains 58-161 and IT6

PI Hfr Sp(metBI < -rel-lP0103

P2 Hfr sptoetBI <

rel-lP0106

P5 Hfr SPCmetB]rel-lP0107

P6 Hfr sptmetBi (rel-lP0108

P8 Hfr spoocelBi

rel-lP0109

P13 Hfr sp<enetBI +rel-lP0104

P72 Hfr spometBi <rel-lP0102

Reeves I Hfr opometBi 4

rel -l

Reeves 4 Hfr 0pOmetBlrel-IPOIOO

CS21- 4ntetBirel-l

W43541- AmetBirel-lAR

CSO1I Hfr spont. KIO HfrmetBi niej-+rel-l rcl-IlonfPf tonAEOT,R T2RPO2A P02A

1T., T2

N stable sel'n.W6F+ mustar(l Cavalli Hfr cloboe W'1895 Hfr

bio- > todD) toelBtI _ lv0v0toietBI rel-1 eel-1 EMBomat lrel-1 P02A PO2A

spont. P3 Hfr ( 4000) W2252-Hfr W3236 Hfr'ont. neelBI tetBI melBI

rel-l el-i rel-IP021B - pro-o

P02A PO2Aspont. P4X Hfr

ont. MneIBI T.rtel-i sel'n.P0'3

moztilitA str. atzi W3787 Hfragar 58-161F- Spicer sel' s. 58-161F-,SOF- sel'n. 58-161F-,SR,AziR'- MeiBI

ont. MeeB] 6otetBi 6eeetBI rel-Iel- lrel- rel-l pro-S

sor-100 slr-100 l8x-76,,v W3208P' azi-7 PO2A

' eeelBl Ifont. rel-l Fl froeo Series

P056 F8 W6 oflac-

(I W3213 Hfre3 58-161F+ SR, AziRl'+ mutants

Uri t.

jilt

)II t

*nit

nt

.0L

eteetBitol-lP057

Wtc'3201 F'

rel-lP060 F15

EMBilasc W1655F+etBli

rel -l

Tesel'n. W4161e+

otnetBirell-tol -30

uv W435Fe+etelBi

r'el-I

lac-45teV

FMBgal W620F+metRlrell-galKP4

motilityagar W220701-

metB1tel-lenal -31

tetBeIrel-lstrle 00

ozi-7

I spot)t.Hayes Hfr

plietBlAO W1655, F- F- rel-l.metBi slr-100

rel-l azi-7AR 'A- P() I

spoilt. W2071F+' I> teetBI K15F+

rel-l metBleel-s33 rel-l- oesr-100

azi-7e,v W3807 Hfrs

rtietBIrel-lnl ut -2P008

W1655F+ ,X,'I spolet. Broda 1-12 Hfr'sneelBi > oteetD

rel-l rel-l

Motilitynigar W3135F-

>metBIrel--IA-

532

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[Chart 51 Hayes HfrmetBIret-I

aIr-lOO

azo-7(phenocopy)

CoutT 6. Hfr H thi-, X- andsome of its derivatiwesW677F+ [Chart 21 X-raytvr-Iisu4-lAi-I

lacYl

malAl,XIR uv

xl-7

ara-ls

gal-S NtonAl mustarc

4, (3000)W3634Hfr uv Hfr H,thi- uv Hfr H thi-,A- uv

thi-i th-l lhi-lrel-I rel-l ret-lA- POI XA-POI POI

phenylspont. galactosii

TRIM sol'n.KL161 Hfr sel'n. KL16 Hfr uv KL20F+

Ul Ul UlTAyAS4 P045drm-3 -v4KL14 HfrP045 EMS uL

nal P068 NGKI166 Hfr sel'n. JC4474 Hfr<) Ul [vKL99 Hfr

thyAR4 1kr-OO Udvm-s P045 lac-42salAIS P042P045 EMSL84K

P1 from rKL96 Hfr

KLI68 Hfr AB2470 JC4490 Hfr lil(i Ul P044

dnrM4 thr-300receBSl ilv-318 EMSt1kV P045P045 KL174 Hfr

P1 from ape UlKLI69 Hfr AB3058 sel'n. his-57

ai] P044

drm-S JC5429 HfrrecCt P1 fromtlhy+ 52&rsOO M107P045 plO8

phenocopy pe-OO KL173 HfrKL16-9 Hfr xHfr7-4 045 Vl

thi-l?t supD43rot-It NO TRIM P044recAI sel'n.drm4fA- JC5088 Hfr JC5401 HfrP045 [PiIIi

recAS6 thVA29TRIM PO45 Pl45sel 'n.

KL162 Hfr P1 from P1 from

Ire-If JC5SI AB2470

vetl1?,recAI JC5491 Hfr X JC5421F- X JC5412 Hfr JCtA PI lillch trt4 [1 1drm- or thyRlO lA1' llkyAsU lAhy' tA- recBfo T'y+ recA13 Thy+ recBSi r

P045 reeCf I8IL'j [SRIP045 I'

JCSS47F-

vocAlSveeilveeCitSrwcSttrwCn

4',JC5495F

IiivoeAlSrvAIS

Paris "X" series; e.g. 3000X74UlfacA74PO1

Paris "U" series; e.g. Hfr H U 355U1lys-POI

Paris "YA" series; e.g. YA149 HfrU1pyrF40POI

30SOHfr uv 30S0U1-7 Hfr'slacZ4 lacZ43P01 series of pyv

mutationsri POIido

3300 Hfr uv 3310 Hfr

lacf(Waci,) lacIff(lac,,)POI lacZ46

POIAT705F+ uv 3320 Hfr

rbs-I lacIst(glt7I? lacZlS(lac,,)

fk Kgad8It PO1IldRi? uv 3340 ITTr

uv A¶715 Hfr U17 [l laclot

1j hlacZ47(lac.,)rbh-l POIP054

ov Paris "M" seriesFA76Hfr

fkJrbs-IP055

P1 fJC4

C5426IIIthy+reCtS

rrom474

Hfr X JC5421F-hil lchart 41tJiyASWS

Thy rvcA 1S

JC554'F-lii

vecC IS

VOL 36, 1972 533

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CHART 7. Some of the early Paris lac- strains[Chart 6] 3000 Hfr

thi-irel-IA-PO1

I30SOHfr X

thi-Irel-1lacZ43P-POl

3000 1thi-1rel-IP-POI

"X" X [Chart 6] 3300 Hfrthi-irel-IlacISS

w ~~x-2300 F

thi-ilaciSSstr-1i3A-

Hlfr x 24

I2000F-

thi-1str-135A-

P678,SRF- [Chart 3]thr-Ileu-6thi-1lacYi

[h] malAI,Rxyt-7

ara-13mit-S

tonA2gal-6str-135A-

'0SOF-thi-1lacZ43malA1 ,XRxyl-7mtl-2ara-13str-135

P678 ,SRF[h]str-135A-

I UV

'- X 3310 Hthi-1rel-1lacISSlacZ46x-P01

2310F-thi-ilacIlSlacZ46mtlA1, XRxyl-7ara-13mtl-Sstr-135A-

[fr X PA351F-[hihis-1argHi

I 8tsr-9purE43xf

2310eFthi-1lacIS2lacZ46malAi,ARxyt-7ara-13mtl-Shis-1argHipurE43str-9A-

I3320 Hfr X P678,SRF-

thi-i [h]rel-1 str-135lacISSlacZ13

POl

2320F-thi-IlacISSlacZ13malAI ,XRxyl-7ara-13mtt-Sstr-135A-

Uv

3340 Hthi-Irel-1lacIRAlacZ4P-POl

[fr X 2001dF-[PA351derivative]

2340eF-thi-ilaclfllacZ47malAI ,XRxyl-7ara-13mtl-2argHipurE4Sstr-9A-

534

I I


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CHART 8. Other line8 derived from wild typeE. coli K-12 (X)F+ [wild type]

. ~~~ ~~~~~~~~~~~~~~~~~~~~I.uv

J5F+pro-22

uvI blood agar

V1485F+supE42_-

W

uv juv3F+ J4F+lac-28 J5-3F+ J5-1OF+DD60 trp-30 pro-2 pro-f2

met-63 his-66 KL25 Hfr uv

P046

J4-5F+ Ra-2 Hfr & spont. Ra-I Hfr uvlac-28 mal-28,AR -* mal-28,AXRhi8-51 8fa-4 sfa-5trp-3O (atP,052) (at P048)

A- A- )uv P048 P052

J6-2F+lac-28his-51tIp-30proC23

uvW2070F+ EMB-gal W1673F+ uv? W1654F+ uv?

ser-26 8er-26 ser-26pro-40 pro-40 A-galTf3(gal,)A-

uv lIV

KL98 Hfr W1872 F+P053 A-

motilityagar W2637F- EMB-gal W3110F-

A- AsupE42?galwake X

W3091F-galT22 XHFT

W3101F- fromgalT2 W2346A-

W3092F-galK2 XHFT

gal,W3102F- fromgalK2 W3160A-

W3096F-galT725 XHFT

W3106F- fromgalT23 W3066A-

W3097F-galTi XHFT

gal7W3107F- fromgaiT1 W3067A4-

W3099F-galE9 XHFT

gal.W3109F- fromgalE9 W3285A-

W3108F- uv W3098F- AHFTgalK8 galK8 galsA- from

W3218

VOL. 36, 1972 535

xV' uv

sel 'n.

CR67F+ JI-0XR lac-28

1CR6

8uz W3100F-IXHFT

gal4 fromW2431

W3104F-gal T12A- Ix

W3094F-galTl2

W3110,thy-F-thyA36A4-

jNG

p3478F-thyAS6polAl


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CHART 9. The derivation of JC12 and JC411 and 8ome of their derivatives[Chart 2]

AB312 Hfr [Chart 5] P4X Hfr X P697/T6F- [Chart 3]thr-l metBI thr-1leu-6 rel-1 leu-6thi-l P03 thi-IlacZ4 lac Ylstr-8 malA I,XRx- [h[ xyl-P012 ara-13\ | ~~~~~~~~~~~~~~~mtl-2\ v Thr+Leti+ tonA2\T61 gal-6

his-1\~~~~~~~~~~~~~~~~~sIIte\ | ~~~~~~~~~~~~A-

[Chart 6]3300Hfr AB352Hfr X AB314F-

thi-1 thr-I metBIrel-1 leu-6 lacYllacI22 thi-1 xyl-7P01 lacZ4 His+ mul-

str-8 [Thr+Leu+J gal-6purFI his-I

U1V x- tonA2P012 tex-1

A-

JC158Hfr X JC12Hfr X PA265F- [Chart 3]thi-I rmetBl [h]rel-1 purFI his-1lacISS lacYl or Z4 argG6serA6

[ml yl-7 Ax-~~ ~ ~ ~~Imul-2

POl gal-6tonAS

Ser itsx-1 Thi+[Met] A- [Ade']

P012

EMS

JC182Hfr KL211Hfr JC355F-purFI [ml metBIthi-1 argGS4 leu-6P01 A- his8-P012 P012 aorgG6(Double [nl 4 lacYl or Z4male) malA1AXR

xyl-7Mtl-f.gal-6A-

strsel 'n.

JC411F-[n,str-104sup -69

X JC182 jAmino JEMS {NG NG Jspont.

JC590F' JC500F- JC1552F- JC1553F- JC1554F- JC1557F-[n] [n] [n] [n] [nl [nl, exceptstr-104 str-104 str-i 04 str-104 str-104 mal+,A'F20(P012) thyAS trp-31 recA1 recA2 sir-104A- k- sup-56 A- A- 8Up-69

spont. TRIM Fl fromsel'n. 58-161

JC1569F- KL11OF- Hfr7-4Hfr[nl except [n] [n]mat+, X str-104 str-104

sir-104 recA1 recA1recA I ihyASS P050A- A- A-

536


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PEDIGREES OF SOME E. COLI K1, MUTANT STRAINS

CHART 10. Paris strain 20OPS and the Paris F-lac+P804Hfr X 234OeF- [Chart 7] [Chart 6] 3000Hfr X P678, SRF- [Chart 3]

thi-1 'h-l thi-i thr-1AdOR lacIf rel-I leu-6P065 lacZ47 A- thi-1

malA1,XR PO1 lacYl[o xyl-7 malA1,XR

ara-13 ara-13mtl-R mtl-Bstr-9 tonA2purE43 gal-6argHl 8tr-185

[Chart 5]2340e F-lac+F' X 234OpF- P4XHfr X 200PF-

[ol [?J metBI thi-1F42 rel-1 lacYl

P03 malAl,XRMal+ xyl-7Met+ ara-13

mtl-28tr-185

2340p F-lac+F' X 200PSF-[?J thi-1F42 (P065) lacYl

200PS F-lac+F'thi-1lacYl(-F42 (P065)

VOL. 36, 1972 537


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BACTERIOL. REV.

CHART 11. The Paris Fl-gal and F2-gal[Chart 5] 4000Hfr X Gal1-2-(Xhd)38OF

metBI [?]rel-1PO2B

Gal2r-(Xhds)380 Fl-gal F' X PA200,SRF- [Chart 3][?] [h] Chart 3F100 (PO2B) his-i

argHIstr-9

PA200,SR Fi-gal F'[h]his-iargHistr-9

F100 (PO2B)

[Chart 5] 4000HfrmetBIrel-iPO2B

X PA309F- [Chart 3][h] [Chart 3]hi8-1arHl8tr-9trp-1x_

PA309 F2-gal F'[h]his-iargHI8tr-9trp-i

F152 (PO2B)

W3101 F2-gal F'galTSf

F152 (PO2B)

X W3101F- [Chart 8]galTfx_

538 BACHMANN


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CHART 12. Derivation of Garen pho- and Su+ strains

[Chart 5] T1,T2Cavatli Hfr sel'n.

,metBIrel-iPO2A

CSlOlHfr spont. KlOHfrmetBI - et+rel-1 rel-1tonAR2 [p] tonAftT2R J2RPO2A PO2A

IIEMS

S26Hfr[P]phoA4PO2A

IX-ray

E15Hfr[PIphoA8(DEIO)PO2A

spont. S26RleHfr- [PI

phoA48upDS2(Sul+)PO2A

spont. S26RldHfr[-[P

phoA4supE3S(Su2+)PO2A

P1 from S26,Su6+Hfr[PIphoA48up-51 (Su6+)PO2A

I NH2OH

H12Hfr[PI

I phoA6PO2A

I NGC9OHfr

[PIphoS9PO2A

IEMS

SlOHfr[P1phoA 11PO2A

IUv

UllHfr[PIphoA6PO2A

|tsPont.

UllRldHfr[P1phoA6supG46 (Su5+)PO2A

1NG

G24OHfr[PIphoA7phoAlOPO2A

JNG

G24OR4Hfr[PIphoA 7phoA IOsup-6f(Su9+)PO2A

spont. H12R7AHfrI [Pi

phoA6supC47(Su4+)PO2A

spont. H12RBAHfr[P1phoA5supFS4(Su3+)PO2A

" . X9 I I

539VOL. 36, 1972

I


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BACTERIOL. REV.

CHART 13. The derivations ojHfr strains

R5 (Reeve58-161:metBIrel-1

s5F+ X W67

I thrleuthilacma

R5Hfrthi-llacYlmalAl ,XRxyl-7mtl-R*gal-3P047

r7F*-1t-6

-Y1zlAI ,XR

f miscellaneous

xyl-7ara-1Smtl-2*gal-3tonA2

P802, P804 and P808Y1OF-

thr-1leu-6thi-1

lUV58F+ X P22F-

bio-1 thr-1leu-6thi-Iton.Xd22

P25F+ Xthi-1Xd 22

A

P678F-(X26)thr-1 mtl-2leu-6 xyl-7thi-l ara-1SlacYl gal-6malAl,XR tonAP.

X26

P802Hfr P804Hfrthi-i thi-1XdD xdflP069 P065

i021.1b2

P804GHfrthi-IX7P065

P808Hfrthi-IlacYlxyl-7mtl-StonA2Xd 5P0105

BACHMANN540


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VOL. 36, 1972 PEDIGREES OF SOME E. COLI K,2 MUTANT STRAINS 541

CHART 14. The 8train8 K12S and 112[Chart 5] W416F+ X Y64F- [Chart 1]

metBI thr-lrel-i leu-6ton-30 thi-I

lacYltonAh 7

sel'n.[Chart 5] W750F+ X W588F- FMB-iac W477F- segregation W465 (_ Het-1)

metBI thr-i thr-l heterozygotetonA24 leu-6 leu-6 (see Comments)lacY4O thi-l thi-lgalTSR lac+ lacYl

41tonA27 tonA27

[Chart 51 W518F+ X W888F- segregation "X" heterosygotemetBI thr-l (see Comments)tonA24 leu-6lacY4O thi-ltgalTtI galTSS

tlonA24 or *7

W1213F+lacY4OgalT**tonA24 or 27thi-l?

spont.

W1293F+lacY4Ogal+thi-l ttonA24 or 27

I spont.

W1294F+ (-K12S Pasadena) AKl2SF+ (Paris)lac+ tonA24 or 27

thi-l?tonAf4 or 27

P1F+gal- (gal)tonA24 or 27

IPllOF+

gal-SCys-23

(a C112) P112-12F+ reisolation P112F+ (a 112)gal-5 gal-Scye-*S Cys-*Shi8-49 his-49A7 X7_


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TABLE 1

Strain designation Sources of' Published references Chart n Synonyms and comments~~~data

E. coli K-12 35, 64, 98, 99 WG1; see text.(A) 57,58,60,61F' I 17, 67

AB250 C See PAIOOAB253 C See W208, SAB254 C See W208, S" FAB257 C See 4000 Hfr; not Cavalli Hfr!AB258 C See W58.3AB259 C See Hayes Hfr, Thi-, AAB262 C See PA100, S' Pro; JW1AB264 C E. coliK-12 wild type; ara due to the

presence of phage mu-1 in the aracistron.

AB265 C See W2915AB266 C See W2961AB280 C "58-161", actually W6AB281 C Hayes Hfr, Thi-, A-; see 3000AB284 C 101 2 W208, S' F+AB311 C, D 101 2 AT11AB312 C, D 101 2, 9 AT12AB313 C, D 101 2 AT1:3AB314 C, D 9 AT14AB352 C, D 9 AT52AB673 C See P10 Hfr (J4)AB674 C See Reeves 1 HfrAB712 C 2AB781 C See W677AB808 C See 3300AB815 C See JC12AB856 C See JC158AB862 C See JC 182, "double male"AB869 C See JC355AB1103 C 2AB1115 C 2AB1133 C 2AB1157 C 45 2AB1621 C 2 2AB1859 C 2AB1884 F 44 4AB1885 F 44 4AB1886 F 43, 44, 45 4AB1896 F 4AB1899 F 43, 45 4AB2301 See 3.300AB2457 F 4AB2462 F 46 4AB2463 F 43, 44, 46 4AB2470 F 28, 43 4AB2474 F 43 4AB2487 F 109 4AB2495 F 28 4AB2497 F 28, 44 4AB2500 F 44 4AB3004 C See C600 (CR34)AB3022 F 28 4AB3591 C See CR63AB3642 C See CR34, ThyAT11 D See AB311AT12 D See AB312AT13 D See AB313AT14 D See AB314

542


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TABLE 1-Continued

Strain designation udratcasf Published references j Chart Synonyms and comments

AT52AB705AT715AT716B1-B12Broda 1-Broda 12,

Hfr'sC6C60C90C112C600sup E44

Cavalli Hfr

CR34CR34, ThyCR63sup D60

CR67CS2CSilCS19CS100Cs1o0E15G240G240R4H12H12R7AH12R8AHayes Hfr

Hayes Hfr, Thi-Hayes Hfr, Thi-, X-

(Paris)Hayes Hfr, Thi-, X-

(Wisconsin)Hfr1 (Wisconsin)Hfr2 (Wisconsin)Hfrs (Wisconsin)Hfr4 (Wisconsin)Hfr4 (Paris)Hfr6 (Wisconsin)Hfr. (Wisconsin)Hfr13 (Wisconsin)Hfr,5 (Wisconsin)Hfr 7-4JIF+J4F+J4-5F+J5F+J5-3F+J5-10F+J6-2F+JlHfrJ2HfrJ3HfrJ4HfrJ5HfrJ6Hfr

DDDD

d

a

a

dddddde, A, B

e, k, A, Be, k, A, B, C

A

AAA

BAAAAgcccccccBBBBBB

12,40,41,77

44313a427,9116

480,81511, 27, 1135939292939337, 56191930, 32, 10530, 3232, 10639

5050

50

26a

242424242424

666

5

2212

2

5,12

228

852125, 121212121212125,6

66,7,10

6

98888888

See AB352

See Broda 1-12 Hfr's

See P112CR34 std. A indicator

Hfr C, W1895, Hfr, (Lederberg) notAB257

See C600

X host range indicator

Avir. indicator

HfrH; W2323Hfr2 (Lederberg); Hfr2 or Hfr2 (Paris)Hfr4 or Hfr4 (Paris)HfrH, AB259,HfrC, 3000 and Hfr H (Paris)See W3634

See Cavalli HfrSee Hayes HfrSee W2924See W2945See Hayes Hfr ThiSee W3807See W3208See W3213See W3201MA1048

J45

J53J510J62See P1See P4XSee P3, 4000See P10See P72See P13

.1.

543


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See P808See J4-5See J5-3See J6-2See J5-10AB815AB856AB862, "double male'AB869

See PAIOO, S', Pro-

See first entrySee W1294

Mutants of Hfr3300 (Paris)

TABLE 1-Continued

J7HfrJ45F+J53F+J62F+J510F+JC 12JC 158JC 182JC355JC411

sup-59JC500JC590JC 1552JC 1553JC 1554JC 1557JC 1569JC2915JC2926JC4474JC4490JC5029JC5088JC5401JC5408JC5412JC5421JC5422JC5426JC5474JC5489JC5491JC5495JC5519JC5544JC5547JC5743JW1K10K12, wild typeK12SK15KL14KL16KL16-99KL20KL25KL96KL98KL99KL110KL161KL162KL166KL168KL169KL173KL174KL211KL252M series

202020, 22

22,2336

22

21, 2321, 231414108

21, 10921, 109

10821, 108, 110108108108, 11010861081081081081086

56, 93

4, 71a, 106b

7473, 7473

73, 7473, 747374

544


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TABLE 1-Continued

Straindesignation |Sources of | Published references Chart SynonymsandcommentsStrandeignaion data ___________ yoysadcmet

P1P1, HfrP2P3

P4XP5P6P8P10P13P21P22P25P31P72Pl10P112P112-12P676

P678

P678, SRP678, F+P678, A-P697P697, SRP802P804P804GP808P3478PA100PA100, SRPA100, S , ProPA100-PA125PA200PA200-PA266PA265PA200, SRPA200, SRF,-galPA300-PA394PA309PA309 FrgalPA351PA601-PA640PA610PA641-PA644R1-R5Ra-1Ra-2Reeves 1Reeves 2Reeves 3Reeves 4Reeves 5S10S26S26RldS26R1e

52, 54 (p. 162)

49, 52, 54 (p. 162)

49, 52, 54 (p. 162)

49, 52, 54 (p. 162)

49, 54 (p. 162)

27

49, 52, 54 (p. 162)

50, 54 (p. 60),

11254 (p. 60)111, 112

54 (p. 60), 112

49, 54 (p. 162)

26

55, 72

55, 72, 7340, 41, 83838340, 41, 838330, 79, 9732, 33, 9732, 3332

14555,11

5,9,1055525

1313

51414142

2,3, 13

3,7,1033

1310, 13131383,93333393,1111311113, 7333

8852251312121212

HfrPl, Hfr Type 1, HfrJ1HfrP2, HfrP21HfrP3, HfrP31, Hfr Type 3, 4000,

HfrJ3, AB257HfrP4X, Hfr Type 2, HfrJ2HfrP5HfrP6HfrP8HfrP1O, Hfr Type 4, HfrJ4, AB673HfrPl3, Hfr Type 6See P2

See P3Hfr P72, Hfr Type 5

2001w

See PA100See PA100, SR

Hfr P808, Hfr Type 7

P697P697, SRJW1

Series of auxotrophic mutants




Series of auxotrophic mutantsSee Reeves 1-5

HfrRlHfrR2HfrR3HfrR4HfrR5

545

BBBB

BBBBBBBBBBBBBB, kB

B

BBBBBBBBB

BBBBBBBBBBBBBBBBhffhhhhhdddd


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TABLE 1-Continued

Strain designation Sourceso f 1 Published references T Chart Synonyms and comments~~~~data

See 58-278MUV mutants of HfrH 3000 (Paris)

58-161, bio-; AB280

Y40, bio+Y87, bio'

AB253

Unstable heterozygote

AB258

AB781

W677, SR

Hfr2, stable clone selected from Ca-valli Hfr

S26, Su6+T94AU seriesUllUlIRldWisupE44

W6biotrel

W13W14W45W67W102W112W133W208W208, S5W416W435W465W477W480W516W518W566W582W583W588W595W620W660W677

gal-3W677, F+W750W888W894W902W904W922W945W1163W1177W1210W 1213W1293W1294W1394W1485supE42

W1603W1654W1655W1655, FW1655, F+, XRW1673W1872W1895

diBddA

A

AAAAAAAACA

AAA

AAAAAAAAA

eAAAAAAAAAAAAAAA

AAA

AAA

30.30

.33, 106

.34, 67:34, 67, 89, 1043, 9, 10, 9634.34

65, 681075959

6162a

34, 5934, 59, 61, 78

78

18, 63, 6678

59, 7859

59, 78

104

63, 6678

61, 102102, 11361

6189a12

61

12

612122

1, 5

1,51, 555222225, 1451414255, 14222142522,6, 13

2,65, 14142222252514141428

2855588

I.

546


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TABLE 1-Continued

See HfrH; Hfr,See HfrH, Thi-, A+

Hfr3Hfr4AB266

See textSee textHfr13

HfrH, Thi-, A-, not Paris strain 3000

Hfr6, MA1040

X-ray mutants of HfrH 3000 (Paris)

W2070W2071W2207W2252W2323W2324W2637W2660W2817W2914W2915W2924W2945W2961W3091W3092W3094W3096W3097W3098W3099W3100W3101W3101 F2-galW3102W3104W3106W3107W3108W3109W3110W3110, ThyW3135W3201W3208W3213W3236W3634W3787W3807W4354"X" seriesY10supE44

Y24Y40

bio+Y46Y53Y64Y70Y80Y86Y87

bio+Y91Y94Y100YmelsupE57supF58

78

8787

85, 86, 8784

87

42, 77, 9513, 77

25

859,62,70,992762, 70, 9959, 62346259, 626259626259, 623462626288, 10211391

547


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BACHMANN

TABLE 1-Continued

BACTERIOL. REV.

Strain designation 1 Sources of Published references I Chart Synonyms and comments~~~data

"YA" series

5858- 161

Reversion to bio+Mutation to rel

58-161F-58-161F-, SR58-161, F-, SR AZ"58-161, F+, SR, AzR58-27858-278M58-30958-33658-58058-59358-61058-74158-2651112112-12200P200PS200PS F-lac+679679-183679-440679-680

F-20002001w2001d20SO230023102310e23202340e2340p3000

3000X7430SO30SOU1-U733003310332033404000

7

35,62,98,9962, 98, 9934, 67, 89, 1043, 9, 10, 9638, 39, 9438, 3938, 3938, 396210398989898989898

494935,62,98,9998, 999862,98,996729

15, 29, 76

11, 2929,7629,7629,76,82

7,767, 1529,76,8229,82828282

6

1,5, 131,5, 13

5555

1

11111111

10

10

10

1, 21

1

1,2

7

7777

7

7

7, 10106, 7, 10

66, 766,7,96, 76, 76, 75, 11

N-mustard mutants of HfrH 3000(Paris)

W3301See also W6

58-161F- Spicer; W6 derivative58-161F- Hayes; W6 derivativeW6 derivativeW6 derivative

T94A; Treffers mutator strain

See P112See P112-12

See P678, SR

HfrH, Thi-, A-, Hfr"C" (Paris),AB259, HfrH

Series of pyr- mutants

P3, P31, Hfr Type 3, AB257

nyms are given in Table 1 and are cross-

indexed to the original designations. The al-phabetical prefixes used in the original straindesignations may, in addition to identifyingthe strains, indicate the laboratories in whichthe strains were made and sometimes conveyother information, as well. Some of the prefixesused in the charts, and the laboratories in

which they were assigned, are as follows. ABwas used by E. A. Adelberg and his collabora-tors at the University of California at Berkeleyand later at Yale University for K-12 strains;non-K-12 strains were designated by AC num-

bers. P. Howard-Flanders A. J. Pittard, A. L.Taylor and others have had AB "numberblocks" and designated their strains and mu-

548

B

BB, kBBB

B

BBBB

B

B

BBBBBBB


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tant alleles according to this system. Now,however, Howard-Flanders and Taylor usetheir own systems. The AT prefix is used by A.L. Taylor, University of Colorado MedicalCenter. The C prefix was used at the CaliforniaInstitute of Technology. CR was used by R.Appleyard after he moved from the last-namedinstitution to Chalk River, Canada. CS wasused by P. D. Skaar, and others, for strainsisolated in the Cold Spring Harbor Laboratoryof Quantitative Biology.The prefix J was used by B. D. Davis,

Cornell University Medical College, for someK-12 derivatives. However, Davis worked ex-tensively with the "W," or Waksman, strain ofE. coli [American Type Culture Collectionstrain number 9637], not to be confused withthe Wisconsin W strains of K12.) J is alsoapplied to some of the Hfr strains isolated by F.Jacob and E. Wollman at the Pasteur Institutein Paris. JC is used by A. J. Clark, Universityof California, Berkeley. KL is used by K. B.Low of Yale University (formerly at New YorkUniversitv School of Medicine).P was used by Jacob, Wollman, and others at

the Pasteur Institute for F+ and Hfr strains,while PA was used to designate their F-straints (with the exception of a few early F-strains that had P designations). The largenumber of synonymous strain designations ap-plied to some of the Paris Hfr's has led toconsiderable confusion, which we have at-tempted to resolve in Table 2.The W prefix was used by J. Lederberg and

collaborators at the University of Wisconsin todesignate mutant strains. They used the prefixWG (Wisconsin Genetics) to designate wild-type strains: E. coli K-12 was designated"WG1," and all other WG numbers referred tonon-K-12 strains in their system. The prefix Ywas used in the laboratory of E. L. Tatum atYale University in the 1940s. The very earlieststrains, those produced by Gray and Tatum atStanford University, were given only numberdesignations.The genetic symbols. The genetic symbols

used throughout are those of Taylor (100), withthe following exceptions and additions. Theolder symbols malA and malB are used forMalh mutations originally mapped in terms ofthese loci. The symbol thyR is used to refer tothe mutation involved in producing the pheno-type "thymine low requirement" when it is notknown whether the locus affected is the drm ordra locus. The symbol PO is used to designatethe points or origin of Hfr strains, each indi-vidual mutation to the Hfr state being assigneda unique number; the Hfr descendants of, as

well as the episomes derived from, an Hfrstrain are assumed to have inherited the pointof origin of their Hfr ancestor. The symbol sfa isused to designate sex factor affinity sites asdefined by Adelberg and Burns (1). The pheno-typic symbols T,2 chloroacetate", andglycerol - are used in the descriptions of some ofthe early strains to designate resistance tobacteriophage T2 and chloroacetate, and theinability to ferment glycerol, respectively.The mutant allele designations used

throughout are those assigned for used in the E.coli Genetic Stock Center and do not neces-sarily correspond to those used in any otherlaboratory. The Stock Center numberingsystem is built on the system employed by E.A. Adelberg in his strain records; some of thesemutant allele designations have thus been inuse for many years and may be recognizablefrom their frequent appearance in the litera-ture. There are three sets of mutant alleledesignations that are sufficiently widely recog-nized that we have felt it necessary to includethem alongside the Stock Center designationsin the strain descriptions. These are the galdesignations of the Wisconsin laboratory (alongwith the galb designation assigned in the Parislaboratory), some of the lac designations of theParis laboratory, and the Su designations usedby Garen and co-workers for suppressor alleles.The Wisconsin gal designation if given for agal- allele the first time it appears on apedigree chart and in the descriptions of the XHFT gal lysates in Chart 8. The gal, symbol isindicated on Charts 2 and 3 in the descriptionsof strain P678. A few of the Paris lac- designa-tions are given in Charts 6 and 7.The symbol Fl is used throughout to refer to

the wild-type F factor of E. coli K-12.The symbol X- is used to indicate the ab-

sence of bacteriophage A. The presence of X isnot noted in strain descriptions as this is thewild-type state. Resistance to A is symbolizedby A' without any indication as to the locusinvolved because the locus of this resistance isnot known for many of the early strains.The symbol SK occurs in some of the early

strain designations, where it indicates resist-ance to streptomycin.Other symbols and abbreviations. The fol-

lowing symbols and abbreviations are used todesignate mutagenic agents: AO = acridineorange; EMS = ethyl methane sulfonate; irrad.= radiation of unknown character; NA =nitrous acid; NG = N-methyl-N, -nitro-N-nitrosoguanidine; N-mustard = riitrogenmustard; spont. = mutation occurring inabsence of deliberate mutagenic treatment;

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UV = ultraviolet irradiation; X-ray = X-irra-diation.The following symbols and abbreviations are

used to designate selective (sel'n.) agents; APT= aminopterin; azi = azide; blood agar =selection for lysis on blood agar plates; EMB-lac = selection for or against utilization of theindicated sugar (lactose here) on eosin-methylene blue agar plates; X"" = selectionwith indicated type of bacteriophage X; mo-tility agar = selection on basis of motility insemisolid agar; nal = nalidixic acid; spc =

spectinomycin; str = streptomycin; T1, T2, T6= bacteriophages T,, T2, and T6, respectively;TRIM = trimethoprim. The standard ab-breviations for sugars are used: ara = arabi-nose; gal = galactose; lac = lactose; mal =maltose; mtl = mannitol; and xyl = xylose.The treatment of suppressors. In our ex-

perience it has proved very difficult to tracksuppressors through the strain pedigrees. Thisis true not only because the presence or absenceof suppressors was seldom noted knowledgea-bly for the early strains, but also because theirexpression seems to be affected by so manyother factors. For these reasons we have in mostcases noted the presence of suppressors only inthose strains in which they have been reportedand have note noted the likelihood of theirpresence in ancestors or offspring of thesestrains. The one exception is the supE- allele ofthe Y10 line, which will be discussed in thecomments on Charts 1, 2, 3 and 4.

Comments on ChartsChart 1. Some early Stanford and Yale

strains. As can be seen from this chart, manyof the early strains were isolated after therather drastic treatment of X-irradiation. Anappreciation of this fact has led to their beingabandoned by many in later years as ance- )rsfor the construction of new stocks. Neverthe-less, the majority, by far, of the strains thathave come to our attention can be traced backto these early lines.The strains W6, W13, W17, and CS19 are

included here to emphasize the instability ofthe bio-1 mutation, which reverted very earlyon in several important ancestral stocks (34).This allele will be discussed at more length inthe comments on Chart 5.The suppressor mutation supE44 was de-

tected in strain Y10 in 1966 (27) and may bepresent in most or all of its direct descendants(i.e., strains descended by mutational ratherthan recombinational events).Chart 2. Some derivatives of strain Y10.

supE- mutations have been reported in strainsY10 (27), C600 (91), and Wi (106). We haveassumed that these are all the same mutantallele (designated supE44 by us) which resultedfrom a mutation in strain Y10 or one of itsancestors and which may be in all of itsdescendants. The uncharacterized amber mu-tation in strain W208 (H. Hoffman-Berling,Dersonal communication) may be supE44, butwe have assigned it the unique mutant alleledesignation sup-49 until more is known aboutit.The strain designations CR34 and C600 are

synonymous. The strain C600 was "reisolated"from a single colony by R. Appleyard, after hemoved from the California Institute of Tech-nology to Chalk River, and was rechristenedCR34 at that time (R. Appleyard, personalcommunication). Considerable confusion hasarisen from this renaming and from the unfor-tunate fact that when Okada, Yanagisawa, andRyan (80, 81) made a Thy derivative of thisstrain they did not give this derivative a newstrain designation. The only name for the latteris now CR34, Thy-.Another source of confusion has been the

gal- markers in the line from W677 to the Parisstrain P678 and its descendants. As shown inthe chart, the sequence of events involved threegal- mutations and two reversions, whichmay have been due to suppressor mutations.Morse, Lederberg, and Lederberg in 1953 (78)recognized that their gal5 (our gal-3) marker inW677 was complex. The galb marker (our gal-6)in P678 and its derivatives have been observedto give a variety of Gal phenotypes upon recom-bination (E. A. Adelberg, personal communica-tion) and may even involve a chromosomerearrangement (R. Curtiss, III, personal com-munication).The malB5 mutation in the Wisconsin Hfr3

(W2924) and the mutation malB16 in the ParisHfr J4 (P10) both involve the integration of thesex factor in the malB locus (Hfr3 [85, 86, 87];J4 [90]).The strain W208SR, ancestor of AB283, was

acquired by E. A. Adelberg in the Paris labora-tory and is probably not identical with theWisconsin strain W2325, which is also a SRderivative of W208.Chart 3. Some derivatives of strain

P678. The widespread early derivatives of theParis strain P678 consist, for the most part, offive series of auxotrophic strains, each seriesbeing produced from a single strain from theprevious series. All were descended from P678cured of bacteriophage lambda. The strainPA100 was, in Adelberg's collection, called

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P697, a designation used only briefly in Paris.Similarly, the designation JW1 for the StrH,Pro- derivative of PA100 has been used in theAdelberg collection for some years but wasapparently evanescent in Paris. The galb con-stellation is in all of these strains. The suppres-sor from the Y10 line may be here also.Chart 4. Some derivatives of strain

AB1157. After discovering the complexity ofthe galb marker, Adelberg used the Wisconsinstrain W2915 in the construction of many of hisstrains. AB1157, a derivative of W2915, wasthen used extensively by P. Howard-Flandersand A. J. Clark, as is shown in this chart.Again, the suppressor from strain Y10 may bein some or all of these stocks.Chart 5. Some derivatives of strains

58-161 and W6. Most of the widely used Hfr'sare to be found on this chart. They werethought to be derived from 58-161 (bio-1,metBl) at the time they were made, but it waslater noted (34, 67, 89, 104) that the bio-1marker had reverted, apparently rather shortlyafter 58-161 was made. Later still, it was dis-covered that the spontaneous mutation rel-I(to the relaxed state with respect to RNA syn-thesis) had appeared in the strain, also veryearly in its history (3, 9, 10, 96). The result isthat most of the well-known Hfr's are bio+ andcarry the rel-1 marker.

The bio-1 marker reverted in the strains Y40and Y87 also. The strains that are described inthe literature as having come from Y87 mostlikely came from the revertant, designated W14(34).The Hayes Hfr is a StrH, Azi H derivative of

W6. The derivation of the widely used Thi-derivative, a recombinant, is given in Chart 6.So many synonymous strain designations

have been used to refer to the more widely usedParis Hfr's that considerable confusion hasarisen concerning their nature and derivations.Table 2 is designed to clarify some of thesepoints, as a supplement to the charts. Concern-ing the Paris Hfr P3 (more widely known asHfr4000), it should be pointed out that this isnot the Cavalli Hfr, as has been erroneouslybelieved by many workers in America. TheCavalli Hfr was isolated by L. L. Cavalli (16)after treatment of "58-161" (actually W6) withnitrogen mustard. The Hfr P3 (which we shallcall 4000 and which is called Hfr Type 3 in thereferences immediately following) arose spon-taneously from 58-161 (W6) in the hands ofJacob and Wollman (49, 51, 52; 54, p. 162).This strain was assumed by E. A. Adelberg tobe the Cavalli Hfr. It was designated AB257 inhis collection and has been widely dis-seminated as "the Cavalli Hfr" or "HfrC,"which it is not (82). The point of origin of this

TABLE 2. The Paris Hfr's

Typea Synonyms Point of origin Derivation Genotype ] Comments

P01 thi thr

PO1 thi thr

P01 thi hr

P0103 leu azi

P03 pro lac

PO2B purE lip

P018 thi malB thr

PO102 ilv met

PO104 mtl ilv

P0105 tonA pro

PO106 lac purE

P065 pro lac

a As listed in references 49, 51, 52, 53; 54, p. 162.

Spont. from 58-161 F+,SR, AziR

Recombination: HfrH x

W677 F+Hfr 4, cured of XI by UV.

Spont. from 58-161 (W6)


metBi, rel-1, str-100, azi-7

thi-1, rel- I

thi-1, rel-l, A-

metBI, rel-l

metBI, rel-1

Spont. from 58-161 (W6) ImetBI, rel-l

Spont. from C6000 F+


Spont. from 112, SR

Spont. from cross: P25F+x P678


Spont. from cross: P25F'x P678

thr-1, leu-6, thi-1, lacYl,tonA21, malB16, A

metBI, rel-I

his-49, cys-23, gal-5,str-58, A T', T3'

Thi-1, lacYl, xyl-7, mtl-2,tonA2

metBI, rel-I

thi-I

Lost very early

Source of F,-gal and F2-gal of Paris (F100 andF152)

F is integrated in malBlocus

Source of F-lac of Jacoband Adelberg, (F42)

L1-I

Hfr H

Type 1

Type 2

Type 3

Type 4

Type 5

Type 6

Type 7

Hfr 2; originalHayes Hfr

Hfr 4; Hfr4

Hfr 3000; Hfr"C"

P1

P4 x 6;J2

P3; P31, 4000

P10; J4

P72; J5

P13

P808

P2; P21

P804

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strain was designated P02, the number as-signed to the point of origin of the Cavalli Hfr.The Stock Center inherited this nomenclature,unfortunately, and has been perpetuating thiserror until quite recently. No one has found, toour knowledge, any difference between thepoints of origin of strain 4000 and the CavalliHfr. However, it would not be surprising ifthese two strains differed in genetic back-ground considering the different treatment theyreceived. We have assigned them distinctivePO designations, as follows. The Cavalli Hfrnow has PO2A, in our nomenclature, and thepoint of origin of Hfr 4000 is PO2B. In caseswhere it is impossible to decide which of thetwo strains is involved, the point of origin willbe called simply P02.The parent of Cook and Lederberg's (25)

extensive series of several hundred lac- mu-tants (W3787) is given on this chart.The Wisconsin strains W3208 and W3201,

sometimes referred to as Hfr8 and Hfr,5, re-spectively, require special comment. When thestrains W3208 and W3201 were isolated, it wasassumed that they were Hfr's because of theirability to transfer chromosomal markers athigh frequency. When these two strains wereexamined in greater detail later on, it wasfound that both were F' strains, harboring F8and F15, respectively (42, 47, 95). Thus nothingis known of the Hfr strains that may have beenthe immediate ancestors of these two F' strains(J. Lederberg, personal communication).Chart 6. HfrH Thi-, A- and some of its

derivatives. The most widely used version ofthe Hayes Hfr, HfrH Thi-, A-, is sometimesthought to be the original strain isolated byHayes. It, is, however, a recombinant strain,resulting from a cross between a phenocopy ofHfrH and the heavily marked Wisconsin strainW677, F+. The strain isolated from this crosswas Thi- and carried bacteriophage lambda.Oddly enough, considering its ancestry, it ap-pears to be suppressor-free. Both the Paris andWisconsin laboratories then cured this strain oflambda phage. The Paris HfrH, Thi-, A- is theone that is widely used: it is equally well knownas strain 3000. Some confusion has resultedfrom the fact that it was also called "HfrC"within the Pasteur Institute (see Table 2).

Several important series of mutant strainswere produced from strain 3000 in the Parislaboratory. The series produced by X-irradia-tion included some of the widely used lacdeletions, such as those in strains 3000 X74and 3000 Xlli. Another series was isolated af-ter U1-U488, etc. A third series, isolated aftertreatment with nitrogen mustard, consisted

of some 297 auxotrophic mutants calledYAl-YA297. (It is important to note thatstrains bearing higher YA numbers arose in adifferent manner.) From the strain 30SO, alacZ- derivative of strain 3000, there wereisolated, after UV-irradiation, seven pyrimi-dine-requiring auxotrophs, designated 30SOU1-7, respectively.From the strain 3300, a lacI- derivative of

strain 3000, were isolated three widely usedstrains carrying lacZ mutations, 3310, 3320,and 3340, and a series of strains carryingauxotrophic mutations which were designatedas HfrH "M" strains 1-30.From F+ revertants of strain 3000, K. B. Low

and A. L. Taylor produced some of their widelyused Hfr's, and Taylor produced several seriesof auxotrophic strains used in his mappingstudies. (The cluster of markers gltS7, gadSl,and gadR2 may or may not be present inAT705. They were found by Lupo and Halpern(75) in a recombinational derivative of AT705,and it is thought that they may have arisen,along with the rbs-1 mutation, when strain3000 was treated with nitrosoguanidine toproduce strain AT705). A number of widelyused rec- Hfr strains have been produced fromHfr KL16 by Low and A. J. Clark.Chart 7. Some of the early Paris lac-

strains. This chart is designed to show therather complex relationships between thewidely used 3300 series and 2300 series ofstrains from the Paris laboratory. There is,however, some doubt about the parentage ofstrain 2300, which was made by J. Monod.Chart 8. Other lines derived from wild

type. The strains W1485 and W3110 have beenused extensively as ancestral stocks in an effortto get away from the heavily mutagenized earlyStanford strains. The amber suppressor,supE42, found in W1485 is not in W3110, whichis either suppressor-free or carries a weak ochresuppressor (C. Yanofsky, personalcommunication).The genetic step involved in the isolation of

W3110 was the selection of a strongly ferment-ing colony on EMB-gal, as W2367 appeared tobe "a weak gal fermenter."Chart 9. The derivation of JC12 and

JC411 and some of their derivatives. Theserecombinant strains are included here becausethey have been used widely for the constructionof stocks. Note that the gal-6 (galb) constella-tion is in most of these strains.Chart 10. Paris strain 200PS and Paris

F-lac+. There is a widespread misconceptionthat the designation 200PS refers to a particu-lar F' strain, carrying a particular episomal

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element. This is not the case. 200PS is an F-strain, which is the host strain to a number ofepisomes in various Parisian F' strains. In thischart it is the host to the Jacob and AdelbergF-lac+ episome (48), our F42, derived fromParis Hfr P804 (Chart 13).Chart 11. The Paris Fl-gal and F gal.

These two widely used episomes arose from theHfr 4000, not the Hfr Cavalli. The point oforigin is, however, similar to, if not the same as,that of the Cavalli Hfr (see notes to Chart 5).The F,-gal is also called the "long" F-gal orF-gal, Aat', bio. At least one widely used versionof this episome (in strain M57 of Meselson)carries a mutant suppressor allele, probably atthe supE- locus (M. Meselson, personal com-munication).Chart 12. Derivation of Garen Pho- and

Su+ strains. This chart gives the derivation ofthe set of strains used most often as sources ofthese pho- and sup- (Su+) markers. Strain C90produces alkaline phosphatase constitutively.Strain E10 carries a phoA deletion.Chart 13. The derivations of miscellane-

ous Hfr strains. The Hfr strain known as R5or Reeves 5 arose spontaneously during across between 58-161F+ (W6) and W677F-(P. Reeves, personal communication). It is arecombinant of these two strains.The Hfr P804 is the source of the Paris F-lac+

episome (our F42: Chart 10).Chart 14. The strains K12S and 112. One

of the first AX (i.e., X+) strains isolated by theLederbergs was the strain W518 (61). A deriva-tive of W518, designated W1294, was made bythe Lederbergs as shown in Chart 14. Thestrain W465 which was involved in this pedi-gree, and which was described by J. Lederbergin 1949 (62a) as "H-1", is still not fully under-stood. It was a heterozygote, an unstable dip-loid, which segregated out large portions ofthe genomes of both of the parent strains. Itcannot be explained by simply assuming that itwas an F-prime strain. The strain W477 was astable segregant from W465. The strain "X,"indicated in this pedigree, which was neithersaved nor given a "W" number, was also aheterozygote, and strain W888 was a segregantfrom this unstable diploid.The strain W1294, in which the presence of

the thi-1 marker was questionable, was sent toJ. Weigle at the California Institute of Tech-nology in Pasadena by E. Lederberg in 1950(106a; and J. Lederberg, personal communica-tion). This strain was called in Pasadena sim-ply "S" (106a) or, later, K12S (4, 71a, 106b).This strain was sent by Weigle to the Parislaboratory, where it was found not to require

thiamine (llOa).Monod and Wollman induced in strain K12S

the gala mutation (our gal-5) (llOa), creatingstrain P1. From strain P1, Wollman producedstrain 112 (llOa), later called P112, by a seriesof UV-induced auxotrophic mutations. A reiso-late of strain 112, picked by Wollman on oneoccasion from a colony #12, led to the strainsbeing called 112-12 (later P112-12) in Paris (E.Wollman, personal communication). Thisstrain was then sent to Pasadena, where it wascalled C112 (3a).The data books of the late J. J. Weigle were

not consulted by the author. The above recon-struction of the pedigree of strain 112 is themost plausible one that could be reached on thebasis of information supplied by the sourcescited above. Some question arises due to thefact that Weigle apparently referred to morethan one strain as K12S (106b).

DISCUSSIONIt is now possible to trace the derivation of

almost all of the strains held by the StockCenter and to apply a uniform and unambigu-ous system of strain designations and mutantallele designations to our stocks. Unfortu-nately, almost all of the strain descriptions thathave been drawn up (and sent out) by theStock Center previously must now be revisedand corrected in the light of information gainedthrough tracing the pedigrees.Once this task is completed, we hope to be

able to provide for investigators in the field ofE. coli genetics strain descriptions and pedi-grees that will make it possible to compare thegenetic backgrounds of most of the importantlines of mutant strains now in use.

ACKNOWLEDGMENTSThis work was supported by National Science Foundation

grants GB7515 and GB22866 in the laboratory of E. A.Adelberg.

I wish to express my gratitude to all of those whogenerously contributed information concerning strain deri-vations and, in particular, to those who made their laborato-ries and laboratory records available to me. In addition tothose acknowledged in the text, R. C. Clowes, E. Lederberg,M. Meselson, E. E. M. Moody, and P. H. A. Sneath gavevaluable assistance.

Particular gratitude is due to E. A. Adelberg and K. B.Low for their constant advice, criticism and encouragementthroughout this work, as they patiently taught me about E.coli.

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1965. Optimal conditions for mutagenesisby N-methyl-N'-nitro-N-nitrosoguanidine inEscherichia coli K12. Biochem. Biophys. Res.Commun. 18:788-795.

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Documents

Pedigrees of Some Strains Escherichia coli K-12 · Thestrain pedigrees are presentedin Charts 1 through 11. The documentation for these diagrams is given in Table 1, underthe strain