JOURNAL OF BACrERIOLOGY, Mar., 1966
Copyright © 1966 American Society for Microbiology
Vol. 91, No. 3
Printed in U.S.A.
Cellular Site in Bacillus subtilis of a Nuclease Which
Preferentially Degrades Single-Stranded Nucleic
Acids
H. C. BIRNBOIM
Department of Biochemistry, Albert Einstein College of Medicine, New York, New York
Received for publication 8 November 1965
ABSTRACT
BIRNBOIM, H. C. (Albert Einstein College of Medicine, New York, N.Y.). Cellular
site in Bacillus subtilis of a nuclease which preferentially degrades single-stranded
nucleic acids. J. Bacteriol. 91:1004-1011. 1966.-A nuclease, identified by a marked
preference for single-stranded nucleic acids, has been demonstrated in extracts of
Bacillus subtilis. The enzyme was associated with the cell wall-membrane fraction
of mechanically disrupted cells and was released from cells which had been con-
verted to protoplasts by lysozyme. The nuclease activity prepared by the latter
procedure was found to be activated and solubilized by treatment with trypsin. The
enzyme had about 2% activity on native deoxyribonucleic acid (DNA) as com-
pared with denatured DNA. By use of CsCl analytical density gradient ultracen-
trifugation, this preparation was shown to degrade denatured DNA selectively in
mixtures of native and denatured DNA.
The cellular location of certain enzymes in
bacteria has been the subject of several recent
investigations. In Escherichia coli, several phos-
phatases and nucleases (6, 10, 11, 12) have been
shown to be released from bacterial cells when
converted to spheroplasts. The localization of
these enzymes at or near the cell surface has been
proposed. In Bacillus licheniformis (previously
classified as B. subtilis), a penicillinase has been
found attached to the protoplast membrane
(3, 4). In B. subtilis, Momose et al. (8) demon-
strated a 5'-nucleotidase, the major portion of
which could be released from cells upon conver-
sion to protoplasts with lysozyme. In strains of
Mycoplasma, several enzymes, including a ribo-
nuclease and a deoxyribonuclease have been
found associated with cell membranes (13).
In this paper, we present evidence for the exist-
ence in B. subtilis of a nuclease characterized by
a marked preference for single-stranded deoxy-
ribonucleic acid (DNA), as compared with native,
double-stranded DNA, as a substrate. In mechan-
ically disrupted cells, about one-half of the
enzyme was found associated with the "cell wall-
membrane" pellet. When cells were converted to
protoplasts by treatment with lysozyme, the en-
zyme was released and recovered in the super-
natant fraction. The association of the nuclease
with the cell wall-membrane of B. subtilis was
studied, and the partially purified enzyme was
characterized with respect to substrate and re-
requirements for optimal activity.
MATERIALS AND METHODS
Bacterial strain. B. subtilis 168-2, a transformable
strain requiring tryptophan and leucine, was used for
studies on enzyme distribution and purification. B.
subtilis 1681>, which requires thymine and trypto-
phan, was used for the preparation of radioactive
DNA.
Media and growth conditions. V-Y broth [Difco
Veal Infusion broth, 2.5% (w/v); Difco yeast extract,
0.5% (w/v)] was the routine medium used for grow-
ing B. subtilis for enzyme isolation. Volumes of 1
liter were inoculated from a single colony and incu-
bated in 2-liter flasks at 37 C in a New Brunswick
gyrotory floor shaker for 16 hr. Under these condi-
tions, the culture reached the early stationary phase
of growth, about 3 g (wet weight) of bacteria per liter.
Reagents. Lysozyme (egg white), three-times
crystallized, B grade, was a product of Calbiochem.
Tris(hydroxymethyl)aminomethane (Tris) was Trizma
of the Sigma Chemical Co., St. Louis, Mo. Analytical
reagent-grade sucrose was obtained from Mallinckrodt
Chemical Works, New York, N.Y. Trypsin (Worth-
ington Biochemical Corp., Freehold, N.J.) was used
without further purification. Diethylaminoethyl
(DEAE) cellulose (Carl Schleicher and Schuell Co.,
Keene, N.H.) was prepared for use by washing with 1
N NaOH, water, 1 N HCI, 1 N NaOH, and water, and
1004
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