[CANCER RESEARCH 41, 3556-3565, September 1981]
0008-5472/81/0041-OOOOS02.00
Effects of Murine Leukemia Virus Infection on Long-Term Hematopoiesis
in Vitro Emphasized by Increased Survival of Bone Marrow Cultures
Derived from BALB/Mo Mice1
Joel S. Greenberger,2 Richard K. Shadduck, Rudolf Jaenisch, Abdul Waheed, and Mary Ann Sakakeeny
Joint Center for Radiation Therapy. Department of Radiation Therapy. Harvard Medical School and Sidney Farber Cancer Institute, Boston. Massachusetts 02115
[J. S. G., M. A. S.I; Heinrich Fette Institute. Hamburg. Germany [R. J.]; and Montefiore Hospital, Pittsburgh. Pennsylvania 15213[R. K. S., A. WJ
ABSTRACT
The effect of infection with Moloney murine leukemia virus
(Mo-MuLV) on long-term bone marrow cultures was studied.
Cultures were derived from the bone marrow of BALB/Mo
mice, which carry Mo-MuLV as an endogenous virus, or from
BALB/c, 129/J, or BALB/c x 129/J mice that were infected
with Mo-MuLV in vitro. The following parameters were tested:
longevity of generation of granulocytes; biological properties
of nonadherent cells in colony-forming assays for pluripotential
stem cells and committed granulocyte-macrophage colony-
forming unit culture, erythroid, or metachromasia-positive mast
cell-basophil colony-forming cells; differentiation of nonadher
ent cells following cocultivation with thymic or bone marrow
stromal cells; and generation of WEHI-3 dialyzed conditioned
medium-dependent permanent cell lines. Granulocytes were
generated for 65 weeks in BALB/Mo marrow cultures, 31
weeks for BALB/c, 22 weeks in 129/J, and 28 weeks in
BALB/c x 129/J cultures. Exogenous infection of BALB/c
cultures with Mo-MuLV increased the longevity of hemato-
poiesis to 41 weeks. Granulocyte-macrophage colony-forming
unit cultures were produced for 61 weeks in BALB/Mo cul
tures, 25 to 40 weeks in Mo-MuLV-infected cultures, and 19 to
33 weeks in uninfected control cultures. Nonadherent cells
harvested from BALB/Mo marrow cultures generated cloned
permanent WEHI-3 dialyzed conditioned medium-dependent,
nonleukemogenic granulocyte-macrophage colony-forming
unit culture cell lines at greater efficiency than did Mo-MuLV-
infected or uninfected BALB/c cultures. The cell lines differ
entiated to mature granulocytes following cocultivation with
purified marrow or thymic stromal cells. There was no detect
able differentiation of nonadherent cells to lymphocytes or
mast cells. Thus, Mo-MuLV does not detectably transform
granulocyte progenitor cells in vitro to granulocytic leukemia.
However, Mo-MuLV replication stimulates self-renewal of gran
ulocyte progenitor cells in both primary marrow culture and in
suspension culture in WEHI-3 dialyzed conditioned medium.
INTRODUCTION
Moloney leukemia virus induces thymic T-cell lymphocytic
leukemia in susceptible mouse strains following a latent period
of several months (34, 38). The molecular parameters of leu-
kemogenesis have been studied in animals infected as new-
borns with virus or in mice carrying the virus in their germ line,
the BALB/Mo strain (25). The preleukemic phase in both
situations is characterized by an increase of viral-specific DMA
copies in hematopoietic cells, such as spleen, thymus, and
bone marrow (27, 28). A second increase of viral copies occurs
in tumor cells during the process of malignant transformation.
The available evidence suggests that the increase in viral-
specific DNA sequences is a consequence of superinfection of
susceptible target cells (37), representing a necessary but not
sufficient precondition of malignant transformation.
The development of a long-term culture system for murine
bone marrow has facilitated studies on the effects of Friend,
Rauscher, and Abelson murine leukemia virus infection on
hematopoiesis (7, 12, 13, 15-18, 21-23). Bone marrow cul
tures maintained in horse serum or corticosteroid-supple-
mented PCS3 (11 ) generate predominantly granulocyte colony-
forming cells and mature neutrophilic granulocytes with rela
tively little erythroid, megakaryocytic, or lymphocytic differen
tiation (12, 21 ). However, CFU-T have been identified in murine
marrow cultures, suggesting the presence of available target
cells for thymic lymphocyte leukemogenesis in vitro (30). In
vivo experiments have indicated that factors specific to the
thymic microenvironment are required for T-cell transformation,
implying that induction of T-cell leukemias in bone marrow
cultures might be impossible (3, 31).
The present experiments were carried out to delineate the
effects of Mo-MuLV on hemopoiesis in long-term bone marrow
cultures. We specifically sought to determine whether germ
line integration of the Mo-MuLV genome in the marrow of
BALB/Mo mice produced effects distinct from exogenous in
fection with Mo-MuLV and whether virus-infected hematopoi
etic progenitor cells could be induced to form leukemogenic
cell lines in vitro.
MATERIALS AND METHODS
Mice. AKR/J, C3H/HeJ, BALB/cJ, and 129/J mice were obtained
from The Jackson Laboratory, Bar Harbor, Maine; NIH Swiss mice
were from the animal colonies of the NIH, Bethesda, Md.; Swiss CD-1
mice were from the Charles River Breeding Laboratories, Inc., Wil
mington, Mass. BALB/Mo and control BALB/c x 129/J (BALB/129)
mice, the strain in which BALB/Mo mice were derived (25), were from
the Heinrich Rette Institute, Hamburg, Germany. All mice were 30 to
33 g.
' Supported by National Cancer Institute Grants CA25412-02 and CA26785-
01. by CA-15237-05, a grant from the Deutsche Forschungsgemeinschaft, and
by a research contract from the National Cancer Institute, Bethesda, Md.
1 To whom requests for reprints should be addressed, at Joint Center for
Radiation Therapy, 50 Binney Street, Boston, Mass. 02115.
Received February 23. 1981; accepted June 16, 1981.
3 The abbreviations used are: FCS, fetal calf serum; CFU-T, colony-forming
unit T-lymphocyte; Mo-MuLV, Moloney murine leukemia virus; DMEM, Dul-
becco's modified Eagle's medium; CM, conditioned medium; DCM, dialyzed
conditioned medium; R-MuLV, Rauscher murine leukemia virus; SFFV, spleen
focus-forming virus; GM, granulocyte-macrophage; CFUc, colony-forming unit
culture; CFUe, colony-forming unit erythroid; CFU-L, colony-forming unit B-lym-
phocyte; CFU-meta. colony-forming unit metachromasia; CPUs, colony-forming
unit spleen; CSF, colony-stimulating factor.
3556 CANCER RESEARCH VOL. 41
Research.
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