Granulosal and thecal expression of bone morphogenetic protein- and activin-binding protein mRNA transcripts during bovine follicle development and factors modulating their expression in vitro

Evidence supports local roles for TGFβ superfamily members including activins and bone morphogenetic proteins (BMP) in follicle development. Access of these ligands to signaling receptors is likely modulated by extracellular binding proteins (BP). In this study we compared expression of four BPs (chordin, gremlin, noggin, follistatin) in granulosal (GC) and theca interna (TC) compartments of developing bovine antral follicles (1-18mm). Effects of FSH and IGF on BMP and BP expression by cultured GC, and effects of LH and BMPs on BP expression by cultured TC were also examined. Follicular expression of all four BP transcripts was higher in GC than TC compartments (P<0.001) a finding confirmed by immunohistochemistry. Follicle category affected (P<0.01) gremlin and follistatin mRNA abundance, with a significant cell-type x follicle category interaction for chordin, follistatin and noggin. Noggin transcript abundance was lower (P<0.05) in GC of large 'E-active' than 'E-inactive' follicles while follistatin mRNA level was higher (P<0.01). FSH enhanced CYP19, FSHR, INHBA and follistatin by GC without affecting BMP or BMP-BP expression. IGF increased CYP19 and follistatin, reduced BMP4, noggin and gremlin but did not affect chordin or FSHR mRNA levels. LH increased TC androgen secretion but had no effect on BMP or BP expression. BMPs uniformly suppressed TC androgen production whilst increasing chordin, noggin, and gremlin mRNA levels up to 20-fold (P<0.01). These findings support the hypothesis that extracellular BP, mostly from GC, contribute to the regulation of intrafollicular BMP/activin signaling. Enhancement of thecal BP expression by BMP implies an autoregulatory feedback role to prevent excessive signaling.

Relationship of dayside auroral precipitations to the open-closed separatrix and the pattern of convective flow

The implications are discussed of acceleration of magnetospheric ions by reflection off two magnetopause Alfvén waves, launched by the reconnection site into the inflow regions on both sides of the boundary. The effects of these waves on the ion populations, predicted using the model described by Lockwood et al. [1996], offer a physical interpretation of all the various widely used classifications of precipitation into the dayside ionosphere, namely, central plasma sheet, dayside boundary plasma sheet (BPS), void, low-latitude boundary layer (LLBL), cusp, mantle, and polar cap. The location of the open-closed boundary and the form of the convection flow pattern are discussed in relation to the regions in which these various precipitations are typically found. Specifically, the model predicts that both the LLBL and the dayside BPS precipitations are on newly opened field lines and places the convection reversal within the LLBL, as is often observed. It is shown that this offers solutions to a number of paradoxes and problems that arise if the LLBL and BPS precipitations are thought of as being on closed field lines. This model is also used to make quantitive predictions of the longitudinal extent and latitudinal width of the cusp, as a function of solar wind density.