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Since VSRs are crucial for directing cargo proteins to protein storage vacuoles (PSVs) during seed development and mediating the targeting of newly synthesized proteinases to PSVs for seed germination 45 , we observed significant defects in the seeds of the snx1 mutant and the snx1/snx2a/snx2b triple mutant, as well as GEF mutants of RAB5 and RAB7, such as vps9a-2 and mon1/ccz1ab