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Ovary |
Department of Neurobiology and Physiology,3 Northwestern University, Evanston, Illinois 60208,
Department of Medicine,4 Feinberg School of Medicine, Northwestern University, Chicago, Illinois 60611,
Robert H. Lurie Comprehensive Cancer Center of Northwestern University,5 Chicago, Illinois 60611
Ovarian follicular development, follicle selection, and the process of ovulation remain poorly understood in most species. Throughout reproductive life, follicle fate is balanced between growth and apoptosis. These opposing forces are controlled by numerous endocrine, paracrine, and autocrine factors, including the ligands represented by the transforming growth factor ß (TGFß) superfamily. TGFß, activin, inhibin, bone morphometric protein (BMP), and growth differentiation factor 9 (GDF-9) are present in the ovary of many animals; however, no comprehensive analysis of the localization of each ligand or its receptors and intracellular signaling molecules during folliculogenesis has been done. The domestic cat is an ideal model for studying ovarian follicle dynamics due to an abundance of all follicle populations, including primordial stage, and the amount of readily available tissue following routine animal spaying. Additionally, knowledge of the factors involved in feline follicular development could make an important impact on in vitro maturation/in vitro fertilization (IVM/IVF) success for endangered feline species. Thus, the presence and position of TGFß superfamily members within the feline ovary have been evaluated in all stages of follicular development by immunolocalization. The cat inhibin 
subunit protein is present in all follicle stages but increases in intensity within the mural granulosa cells in large antral follicles. The inhibin ßA and ßB subunit proteins, in addition to the activin type I (ActRIB) and activin type II receptor (ActRIIB), are produced in primordial and primary follicle granulosa cells. Additionally, inhibin ßA subunit is detected in the theca cells from secondary through large antral follicle size classes. GDF-9 is restricted to the oocyte of preantral and antral follicles, whereas the type II BMP receptor (BMP-RII) protein is predominantly localized to primordial- and primary-stage follicles. TGFß1, 2, and 3 ligand immunoreactivity is observed in both small and large follicles, whereas the TGFß type II receptor (TGFß RII) is detected in the oocyte and granulosa cells of antral follicles. The intracellular signaling proteins Smad2 and Smad4 are present in the granulosa cell cytoplasm of all follicle size classes. Smad3 is detected in the granulosa cell nucleus, the oocyte, and the theca cell nucleus of all follicle size classes. These data suggest that the complete activin signal transduction pathway is present in small follicles and that large follicles primarily produce the inhibins. Our data also suggest that TGFß ligands and receptors are colocalized to large antral follicles. Taken together, the ligands, receptors, and signaling proteins for the TGFß superfamily are present at distinct points throughout feline folliculogenesis, suggesting discrete roles for each of these ligands during follicle maturation.
2 Correspondence: Teresa K. Woodruff, Northwestern University, Department of Neurobiology, O.T. Hogan 4-150, 2205 Tech Drive, Evanston, IL 60208. FAX: 847 491 2224; tkw{at}northwestern.edu
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