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we next assessed whether these thin projections could mediate cytoplasmic exchange between adjacent spermatogonia. we first used em on over 200 serial sections from p6 testes and observed, at high resolution, apparently continuous cytoplasm connecting neighboring spermatogonia via icbs ( fig.3 a) and cytoplasmic projections ( fig.3 b). we next used fluorescence recovery after photobleaching (frap) to definitively determine whether cytoplasmic projections and icbs represented patent connections between adjacent live spermatogonia. we photobleached id4-egfp+ spermatogonia (to 10% original fluorescence) in various configurations (as, apr and as apparently connected by projections) and quantified fluorescence recovery over a short interval (<12min, fig.3 c). although recovery could theoretically result from synthesis and accumulation of nascent egfp molecules, this generally takes several hours ( kourtis and tavernarakis, 2009 ). we first photobleached as id4-egfpbright spermatogonia exhibiting no apparent connections; they did not regain fluorescence ( fig.3 d-g,p), revealing that was indeed too short an interval for accumulation of detectable egfp. we next photobleached individual id4-egfpbright spermatogonia that were apparently connected to adjacent id4-egfpbright spermatogonia via cytoplasmic projections. these as spermatogonia (not icb-connected) rapidly recovered 40% of their original fluorescence, while their non-photobleached partners lost 40% of their original fluorescence ( fig.3 h-k,p). finally, we photobleached id4-egfpbright spermatogonia in clear apr configurations (connected by icb) and found recovery was rapid (10min), while the non-photobleached partner became 50% less intense ( fig.3 l-p). the linear kinetics of the redistribution of egfp over this brief interval supported rapid diffusion of egfp molecules between cells. the percentage of egfp shared between cells formed the mobile fraction ( fig.3 q). this is the first report, to our knowledge, documenting passage of macromolecules (here, egfp, mw 27kda) between adjacent spermatogonia.
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