Importins transport proteins into the nucleus to effect gene transcription changes; they also sequester proteins to intracellular domains and mediate BMP and WNT signalling [1, 2, 3]. Dynamic IPO5 expression and subcellular localisation are functionally linked with key male germline developmental steps, as germline-specific Ipo5 deletion in fetal (>E15.5) or newborn (PND3) testes causes germ cell depletion before/during the first round of spermatogenesis [1, 4, 5]. Here we examine IPO5 function in spermatogonia to assess its importance in adult male fertility.
In vivo analyses were conducted using our inducible Ipo5xUBC-cre knockout model (Ipo5TAM-KO). Control and Ipo5TAM-KO adult males received tamoxifen for 2 days, and testes harvested 3d (day), 1w (week), 2w, 4w, and 6w post-injection (n=3-5/timepoint). Cell type-specific markers were analysed using section immunofluorescence. RNAseq was performed on 3d, 1w, and 2w post-injection testes to identify transcriptional changes that mediate later phenotypes. Finally, potential IPO5 binding partners were identified in cultured adult mouse undifferentiated spermatogonia using immunoprecipitation and mass-spectrometry (n=3).
Ipo5TAM-KO mouse testes at 1w & 2w had more CC3/9+ cells (apoptosis, p<0.01), and fewer PLZF+ (undifferentiated spermatogonia) and GILZ+ cells (spermatogonia/spermatocytes) (p<0.05). Testis weights were progressively reduced at 4w and 6w (p<0.0001), and Sertoli cell-only tubules were abundant by 6w. RNAseq identified 87 DEGs between Control and Ipo5TAM-KO samples, most exclusive to spermatogonia/spermatocytes and relating to meiotic cell cycle and DNA repair. In agreement, IPA upstream regulator analysis implicated inhibition of STRA8, necessary for meiosis [6]. Amongst the 91 IPO5 binding proteins were histone H2A.X and SALL4, essential for undifferentiated spermatogonia maintenance [7]. Functional confirmation assessing subcellular localisation following IPO5 depletion is underway.
These investigations identify IPO5 as the first importin essential to sustaining the mouse spermatogonial/SSC population. Analysis of IPO5 binding partners will delineate regulatory proteins that underpin male germ cell biology and are required for male fertility.