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Fig. 10. Proposed model by which the paracrine network of BMP/GDF-9 growth factors and their binding proteins interact to regulate apoptosis in the COC microenvironment. Oocyte-secreted BMP-15 and BMP-6 signal through the cumulus cell receptor BMPR-II to actively prevent cumulus cell death. Oocyte-secreted GDF-9, also acting through BMPR-II but using a different type-I receptor to the BMPs, does not prevent cumulus cell apoptosis but induces cumulus cell gremlin expression. GDF-9-stimulated gremlin expression may in turn block theca and granulosa cell-derived BMPs from competing with BMP-15, BMP-6 and GDF-9 for BMPR-II binding. In addition, follistatin produced by mural granulosa cells and present in follicular fluid, may limit the anti-apoptotic effects of oocyte BMP-15 specifically to the COC microenvironment.





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