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NX. E. orton
Report on lac— study Oct. 4, 1956

Preparation of stocks

The plan of this study is to prepare pairs of stocks cohtaining
the same lac- allele, the initial member to carry Cavalli's Hfr, M, and
a UV-induced lac-, the other to be a lac- F= prototroph derived from the
first by recombination with Y10. For chromosome mapping, each Hfr stock
will be modified by selection of Ves a marker closely linked on the left of
lac-l, In future, the Hfr stock will also carry Pe, one locus for which
is reported to lie between lac-l and Ve (Fried, Woe data fit equally
well the order P_ V, _lac-l). an Hfr Pe be atock was obtained by UV
irradiation of W1L&95 and is being tested to determine the location of P-.
Preliminary tests indicate the order P vy lac or VE lac P,

Pending the development of the Pe stock, F~ prototrophs and Hfr

y53 wll2 lac?

M- Pf stocks were prepared far the genes lacl®’, lacy ; jac, 7 2?

316 and for 12 lac derivatives of W195 (1940-51).

wo7
lec, ; 1a W929 lacy
In addition, F- or Fé prototrorhs were prepared for lacy”, lacy, lac, 193,
w3128
and lac (Table 1 and Fig. 1).

A. whe
In the course of this work, we lac- stocks were isolated

which differed in recombination and reversion patterns from the lac- parent:,

W31L59 is a stable isolate from a cross of Y10 with the very highly mutable

W1951, and fails to tecombine with W1951 and all but one of the apparently

single-step lac-1 mutants. W3229 is a spontaneous derivative of W3120

accidentally isolated in serial transfer. It is much more stable than

its 1acy®? ancestars and fails to recombine with any of the recognized lac-1

mutants. At present it is the means by which lac-1 is identified, since the

lac-l pseudoalleles have sufficiently high recombination ratés to be indistinguishable
from unlinked loci in streak tests. W3146 was isolated from a cross of

ouil2

W3129 by W112 in a attempt to introduce lac, into an Hfr stock; it

recombines with W112 and all tested lac-1 mutants and is almost certainly
2
not a derivative of W112, since it remains Ss. gale VE like W3129,. (Of the
stocks in table 1, the Hfr tac? is the only one not yet prepared.) The
origin of the two-step mutants W3229 and W3159 raises questions about the
nature and frequency of spontaneous changes in recombinatio pattern of
lac- mutants.
Streak allelism tests
Cross-streaks of Hfr M- lace end F~ lac- prototrophs on M lac plates
are convenisant tests for allelism, but their interpretation, although elear
in most cases, is in others made difficult by too frequent lacfé reversions,
especially when they occur in the M~ line, and by the relatively bw fertility
of 3H3, W3164, and W3140. Tests with highly fertile Hfr stocks have been
unambiguous.
The lac- stocks tested fall into two groups. The majority fail
to recombine with W3229, and ere therefore designated lac-1 (Table 2). Of
these Y87, Y53, W1950, and W1951 appear to be allelic, but may be separated
by their reversion rates, which are in the order Y53< Y87< W1950 = W1951
when compared as prototrophs. The latter two stocks are exceptionally revertible
and are probably identical, as they were isolated in the same experiment.
Similarly, W1948 and W1949 have not been distinguished by recombination and
revertibility tests. All other apparently single-step lac-1 mutants recombine
with one another. Five lace genes remain unclassified with respect to
locus, since they recombine with lech *”, lace2, 3,455, Fp ahd tac 7128
as well as with each other. The two recently obtained lac~ from W323%6
have not been adequately tested. With chromosome mapping tests, some of these
unclassified genes will probably be found to be pgeadoallelic with known
loci.
Intensive allelism tests
Quantitative recombination tests have been deferred until

ve P- stocks are available. A few intensive allelism tests were carried

out on material at hand, without re-isolation of stocks, so that reversions
-~3-

in the agar stabs over varying tire intervals were confounded with unavoidable
reversions in the Penassay broths in which the cultures were grown up and

on the M lac plates on which they were tested. Colonies were counted at

21, hrs. to minimize reversions on the plates. Despite the erudeness of these
tests, they are of interest in confirming the cross-streak tests and providing
a rough measure of reversion rates (Table 3).

W3128 lace Hist. FY

This stock was received from Borek as a questionable double mutant.

Hist “ reversions on D(O) remain lac-. Lac» prototrophs were obtained from
a cross with W1695. Both hist- and hist # were isolated from lacf/ reversions
on B lac. All the evidence is consistent with independent origin of hist-
and lac-, with hist¢ reversions in some lacf papillae.

Persistent diplofids

From Hl lacy 93 colonies were isolated which carried Het, as shown
by laev colonies in the cross with W1940. The lac- parents have been stabbed
as N13-2 and the jaev diploids as N13-1.
An attempt was made to test allelism of the lac- segregants of

H271, a diploid lacf which segregates stable and mutable lace. The original
constitution of this stock was rad 3/1ac"*, which was lac- in phenotype.
Unfortunately, the y53 Hfr tester is of low fertility and the wil2 tester

<t
has ‘net. been synthesized, so a conclusive analysis has not yet beenomade.

Interaction of lac, gal- and lacy gale
E. M. Lederberg reported that cross-streaks of dacy- gals

and lac.- gal- gave a bluish color after 48 hrs. on B lac, but that other

1
lace loci are negative or give a less intense color. This has been confirmed,
the color reaction being much clearer on paper prints than on the agar plate.
A gal,- lacf tester should be tried. Cells lysed by T6 on B lac agar

give a blue reaction, but I was not able to cifferentiate lac~l1 from other

loci by this method. In fermentetion tests on EMB agar, read at 24 hrse;
-i-

all the lac- prototrophs in this study (with the exception of the mal~l and

gal-2 stocks) behaved as follows?

Locus mal mt1 gal zylose
2 and 3240 slow slow # 7
3 and 5 0 0 very slow #
ail others 4 4 x Z

Pl transduction

Attempts to grow high titer Fl in L broth were uneuccessful on a

variety of lp® stocks, The Swanstrom- Adams confluent lysis plate method

is now being tried. As soon as good lysates are made, the transduction system

will be explored.
a
Y10 W3128 58—6|
> iN oN
Wil2  Y5SS wieg wos WHS = Wi33 W4s WG
Z werd
W JIN
| W323G
W3224
Source

Moye
y53

Vv wll2
“ wi941
v  w1945
~ w1946
wl948
~~ wl949
w1950
w1951
v’ w3lZ6
w3159
w3229

~ whd
wl08
wo7

wi45

w133

w3128
w1940
wl942
wl943
wl944
wl947

w3237
w3240

 

Boe oe oe oe pe op op

H

NIow FF FP BW NY HE BP HE

Table 1

Hir_ M

ca

W3120

- Lac Stocks

3H3 ind. Hfr (JL)

W3221 ~M+
W1941
W1945
W1946
W1948
W1949
W1950
W1951

W3146 gal5 We Ss NO

W3229
W3164 s™

W3140 Ss”

Hfr M- Pa
W3237
W3240

N6é

NA

oO

Fe prototroph

W3230
W3134
W3089 mal
W3148
W3152

‘1 7

W3153
W174:
W3156
W3157
W3158
W3175 WE
W3159
W3133
W3112
W2243
W2244 Ff
W3127 :
W2245 FY
W3238

o ¥9239 FY
W347
WwaZ9

* W3215
W151

" W3154

N23

N2

N9

N23
N7
Table 2, lac, recombination pattern

Stocks recombine to give lacf if the corresponding bard do not overlap.

3ir¢ 2134 3191 3ue
987, y53, wigsnjy «2222 BBY 51571, 2156

 

3084
wil2 eT
w1941 ai
wl945 2
w3146 3175_
. 35
wl946 ° DID
31S3
3/33 3)33

w3229

 
Table 3 Allelism tests

Exper. 1. O.1 ml. Fe snd 0,1 ml. Hfr from overnight cultures into penassay.
After 4 brs. plate 0.1 of mix on M lac.

F Hfr Me
W3229(3H3 W1941
W3133 0 0 0
W3134 22 23 > 1000
W3148 1 13 0
W3029 oO — > 1000 ‘

Exper. 2. Mix centrifuged, washed with saline, concentrated in saline 1/10.
1.0 ml. of concentrate on M lac.

W3229 W1941
W3133 3 2
W3089 0 =

Exper. 3. 0.1 ml. Fe and 0.1 ml, Hfr in 10 ml. penassay, After 3 hrs. plate
O.1 ml. on M lac.

F- no Hfr allelic Hfr Hfr = W3229
W3133 0 W3229 0 0
W3134 44 3H3 52 50
W3089 0 _ _ 0
W348 0 wi941 0 0
W352 0 W1945 0 0
W3153 1, W1946 15 an
W3174, 0 w1948 0 0
W3156 0 w1949 0 0
W3157 26400 W1950 14200 15400
W3158 29000 wi951 17000 23200
W359 0 W1951 32 0

W3175 0 W316 0 0
Table 3 (cont.)
Exper. he 0.1 ml. Fe and 6.1 mi, Hfr in 10 ml. Penassaye After 24 hrs. plate 0.1 ml.

on M lac.

¥F no Hfr allelic Hfr

W3127 32 W3140 153
W3112 0 W3164 0
W3151 0 WL9A4 0
W3154 1 W1947 1
W3147 3 W1940 2
W3149 1 W1942 0
W3150 1 7 0
W3155 0 " 0
N30

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 
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