[CANCER RESEARCH 41, 537-545, February 1981]
0008-5472/81 /0041-OOOOS02.00
Purification and Characterization of a Nuclear DMA-binding
Phosphoprotein in Fetal and Tumor Tissues1
Egon Durban, David Roll,2 Gregory Beckner, and Harris Busch
Department of Pharmacology, Baylor College of Medicine. Houston. Texas 77030
ABSTRACT
The basic nonhistone phosphoprotein 110/8.4 (M.W. x
10~3/pl) was found in 0.35 M NaCI nuclear extracts of four
tumor tissues, i.e., fast-growing Novikoff hepatoma, Morris
hepatoma 3924A, HeLa cells, and Namalwa cells; it was also
found in fetal rat liver. This protein was not detected in normal
or regenerating liver and thus may represent an "oncofetal"
protein of potential interest as a cancer "marker." Protein
110/8.4 was purified approximately 4000- to 5000-fold under
nondenaturing condition from 0.35 M NaCI nuclear extracts of
Novikoff hepatoma cells or Namalwa cells by ammonium sulfate
fractionation, calcium phosphate gel treatment, and phospho-
cellulose chromatography. Sodium dodecyl sulfate:polyacryl-
amide gel electrophoresis of the purified native protein revealed
a single polypeptide chain with a molecular weight of approxi
mately 110,000. The pi of the protein was estimated to be 8.4
by nonequilibrium pH gradient electrophoresis in 9 M urea;
accordingly, this protein was designated 110/8.4. Amino acid
analysis showed that Protein 110/8.4 had an acidic:basic
amino acid ratio of 1.25 and a high lysine and serine content;
approximately 20% of the serine residues were found to be
phosphorylated. Hydrazinolysis indicated that the carboxyl-ter-
minal amino acid was serine; the amino terminus appeared to
be blocked. Binding of Protein 110/8.4 to DNA was studied by
the nitrocellulose filter assay. High-affinity binding occurred at
ionic strength equal to or below 0.15 M.
INTRODUCTION
Various lines of evidence indicate that nonhistone phospho-
proteins may be involved in gene regulation (14, 32). Such
proteins have been reported to cause alterations in the rate
and pattern of in vitro gene transcription in many different
systems [references tabulated in a review by Kleinsmith (14)].
In addition, they have been found to bind preferentially to
homologous DNA in vitro (13, 15, 18, 33). Proteins with these
characteristics are present in a phosphoprotein-enriched
subfraction of 0.35 M NaCI chromatin extracts (15).
While extracting chromatin of Novikoff ascites cells (labeled
in vivo with 32Pi) with NaCI solution of the same ionic strength
(0.35 M), a phosphoprotein of an estimated molecular weight
of 110,000 and an estimated pi of 8.4 was found on autoradi-
ograms of 2-dimensional nonequilibrium pH gradient and SDS3:
' These studies were supported by Cancer Research Grant CA 10893, P. 1,
the Taub Foundation, the DeBakey Foundation, and the Pauline Sterne Wolff
Memorial Foundation.
2 Present address: Chemistry Department, Roberts Wesleyan College, Roch
ester, N. Y. 14524.
3 The abbreviations used are: SDS, sodium dodecyl sulfate; NKM, 0.13 M
NaCI:5 rnw KCI: 8 mm MgCI2:0.1 mM PMSF; RSB buffer, 10 mw Tris-HCI (pH
7.4): 10 mM NaCI: 1.5 mM MgCb; PMSF, phenylmethylsulfonyl fluoride: HMG,
high-mobility group; LMG, low-mobility group.
Received May 23, 1980; accepted October 20, 1980.
polyacrylamide gels (41). By ammonium sulfate treatment of
0.35 M NaCI extracts, Protein 110/8.4 was also found by
Coomassie blue staining in rat hepatoma 3924A, in 2 human
tumor lines (HeLa and Namalwa), in a Burkitt's lymphoma-
derived line, and in fetal liver. It could not be detected in normal
or regenerating liver. The present study reports on 2 methods
of purification of this "oncofetal" nuclear phosphoprotein. The
first, in the presence of urea, led to higher yields, which is
useful for chemical characterization. The second method was
carried out under nondenaturing conditions to provide Protein
110/8.4 in native form for studies on its biological function.
MATERIALS AND METHODS
Materials. Proteins used for calibration of molecular weights
were obtained from Bio-Rad Laboratories, Richmond, Calif.
Ampholines were purchased from LKB Instruments, Inc., Rock-
ville, Md. Calcium phosphate gel, Escherichia coli DNA, and B!
nyclease were obtained from Sigma Chemical Co., St. Louis,
Mo., and phosphocellulose P-11 was obtained from Whatman,
Inc., Clifton, N. J. Nonidet P-40 (Shell) was purchased from
Particle Data, Inc., Elmhurst, III. and [3H]dCTP was purchased
from New England Nuclear, Boston, Mass.
Animals and Tumor Cells. Novikoff hepatoma ascites cells
were transplanted i.p. in 200-g male albino rats (Holtzman Co.,
Madison, Wis.). After 6 days, the animals were sacrificed, the
ascites fluid was filtered through cheesecloth, and the cells
were washed 3 times at 4°with NKM (23). Normal, fetal, and
regenerating rat liver and Morris hepatoma 3924A were minced
in NKM and rinsed 3 times in NKM. Fetal livers were obtained
from 19- to 20-day-old fetuses. HeLa S3 cells were grown in
Eagle's medium (Grand Island Biological Co.), washed twice
with phosphate-buffered saline (0.01 M sodium phosphate
buffer, pH 7.2:0.145 M NaCI), and washed once with RSB
buffer. The Namalwa line is a Burkitt tumor line generously
provided by Dr. John Douros and Fred Klein of the Frederick
Cancer Research Center.
Isolation of Nuclei. All tissues or cells, except HeLa and
Namalwa cells, were homogenized in 0.5% Nonidet P-40:0.01
M NaCI:1.5 mM MgCI2:0.1 mM PMSF:10 mM Tris-HCI, pH 7.6
(26), by 15 strokes in a 200-ml Teflon:glass homogenizer with
0.01-inch standard clearance (Glenco Scientific, Inc., Houston,
Texas). The homogenate was centrifuged at 12,000 x g for
10 min. The crude nuclear pellet was resuspended by homog-
enization with a loose-fitting pestle (>0.01-inch clearance) in
1 M sucrose:NKM. Nuclei were collected by centrifugation at
1000 x g for 10 min. The HeLa cells washed once with RSB
buffer were resuspended in RSB buffer containing 0.5% Non
idet P-40 and homogenized in a Dounce homogenizer with 20
up-and-down strokes. After centrifugation at 600 x g for 8
min, pellets were resuspended in 0.25 M sucrose:0.5 mw
FEBRUARY 1981 537
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