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Division of Reproductive Biology, Department of Gynecology and Obstetrics, Stanford University School of Medicine, Stanford, California 94305-5317
Address all correspondence and requests for reprints to: Aaron J. W. Hsueh, Ph.D., Stanford University School of Medicine, Department of Obstetrics and Gynecology, 300 Pasteur Drive, Stanford, California 94305-5317.
Progression of preantral follicle development is essential to further
follicle maturation and ovulation, but there are few models for
studying the regulation of preantral follicle survival and growth. We
have evaluated preantral follicle survival in vivo and
in vitro, and have developed a serum-free rat follicle
culture system that can be used to characterize the regulation of
preantral follicle growth and differentiation. Analysis of ovarian cell
DNA fragmentation during the first wave of follicle growth in the
infantile rat indicated negligible apoptosis up to day 16 of age.
However, a major increase in apoptosis was found by day 18, a time
point associated with the appearance of large antral follicles.
In situ analysis confirmed that apoptotic DNA fragments
were limited to antral follicles. Culture of individual preantral
follicles mechanically dissected from ovaries of 12- or 14-day-old rats
in serum-free conditions led to major increases in follicle cell
apoptosis, similar to that seen in cultures of antral and preovulatory
follicles. In contrast to antral and preovulatory follicles, treatment
of preantral follicles with gonadotropins or cAMP analogs did not
prevent apoptosis. However, treatment with 8-bromo-cGMP or 10% serum
suppressed apoptosis by 75% in cultured preantral follicles. In
situ analysis identified granulosa cells as the cell type
susceptible to apoptosis regulation. Taking advantage of the ability of
the cGMP analog to suppress apoptosis, we evaluated the potential of
FSH as a growth factor. In the absence of serum, FSH treatment for
48 h did not affect follicle size compared to controls; however,
treatment with the cGMP analog together with FSH increased follicle
diameter (13%; P < 0.01) and viable cells
(2.4-fold; P < 0.01) compared to control values.
Immunoblot analysis further indicated that the inhibin-
content of
the cultured follicles was increased by treatment with the combination
of FSH and 8-bromo-cGMP, demonstrating the induction of follicle cell
differentiation during culture. Therefore, we demonstrated that
activation of the cGMP pathway promotes the survival of cultured
preantral follicles and that in the presence of a cGMP analog, FSH is a
growth and differentiation factor for preantral follicles. The present
serum-free follicle culture model system will be useful in further
evaluation of the regulation of growth and differentiation of preantral
follicles.
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