Molec. gen. Genet. 166, 229-231 (1978)
© by Springer-Verlag 1978
Short Communication
Binding of lac Repressor to the Secondary lac Operator
in Escherichia coil
Alain Rambach and Mai Lebastard
Groupe INSERM U. 163, Institut Pasteur; 28, Rue du Dr Roux, Paris 15e, France
Summary. In the lac operon, the existence of a second-
ary repressor binding site, inside Z gene, had been
inferred from in vitro binding studies (Reznikoff et
al., 1974; Gilbert et al., 1975).
A serie of deletions have been constructed from
a lac transducing 2 bacteriophage. Some of those
deleted bacteriophages have still the property of dere-
pressing a chromosomal lac operon, even though they
do not contain any more the lac operator. This phe-
nomenon is an indication that the secondary repressor
binding site is also active in vivo.
Lac repressor acts on lac DNA by binding to a spe-
cific site: the lac operator. The lac operator was first
defined as a genetic locus. Mutations at this locus
suppress repressor binding and hence such mutants
express lac activities constitutively (Jacob and
Monod, 1961). The lac operator was later character-
ized directly by in vitro binding of lac repressor to
DNA fragments (Gilbert and Muller Hill, 1966) and
the relevant DNA fragment has been sequenced (Gil-
bert and Maxam, 1967). The lac operator is located
just next to the structural lac Z gene which codes
for the/%galactosidase.
In addition to this first lac operator, a secondary
lac operator has been described. The secondary lac
operator is located inside the lac Z gene, while the
first operator is outside the structural gene. The sec-
ondary operator has been characterized by experi-
ments involving in vitro binding of lac repressor to
DNA containing various deletions of the lac region
(Reznikoff et al., 1974). We present here data showing
that the lac repressor binds in vivo to this secondary
operator.
To obtain in vivo information on the secondary
lac operator we wanted to compare lac expression
from transducing bacteriophages carrying various re-
gions of the lac genes but not containing the first
operator. We expected some of these bacteriophages
to contain no operator and some to contain the sec-
ond operator. We have constructed transducing 2 bac-
teriophage carrying various distal portions of the Iac
Z gene in the following manner. Barnes et al. (1974),
have constructed a coli strain with a thermoinducible
2 bacteriophage inserted at a site very close to a lac
operon. Among the phages produced after thermoin-
duction of such lysogens, 1% express a lac + pheno-
type, but contain distal portions of the lac Z gene.
These were detected as giving white plaques on
Mac Conkey Lactose plates at 30 ° C on a strain carry-
ing a small deletion of the second half of the lac
Z gene, Hfr 3000 U163, but giving lac + recombinants
with the same Hfr 3000 U163 strain. Approximatively
600 bacteriophages giving white plaques were
screened and 3 were found to recombine with Hfr 3000
U163. Since they recombine with the U163 mutants
these bacteriophages must contain distal regions of
the lac Z genes. They were called 2AR102, ,~AR103
and 2AR104.
Genetic Mapping
The extent of the lac region incorporated in these
transducing bacteriophages has been determined by
genetic recombination with a series of lac Z mutants.
As controls we have also used two other bacterio-
phages picked out of the 600 noted above : a bacterio-
phage giving a red plaque on Mac Conkey Lactose,
designated 2plac and a bacteriophage giving a white
plaque but no lac ÷ recombinanats, designated
2AR105. Another control phage 2placsW4680S carry-
ing a known deletion in the lac Z gene, was used
in this study. The extents of the lac region in the
strains 2AR102 to 105 are shown in Figure 1:
2AR105 contains no lac Z gene, 2AR103 contains
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