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Causation was confirmed in mice with targeted deletion of four of six MIDN protein isoforms. MIDN was expressed predominantly in lymphocytes where it augmented proteasome activity. We showed that purified MIDN directly stimulated 26S proteasome activity in vitro in a manner dependent on the ubiquitin-like domain and a C-terminal region. MIDN-deficient B cells displayed aberrant activation of the IRE-1\/XBP-1 pathway of the unfolded protein response. Partial or complete MIDN deficiency strongly suppressed E\u03bc-Myc\u2013driven B cell leukemia and the antiapoptotic effects of E\u03bc-BCL2 on B cells in vivo and induced death of Sp2\/0 hybridoma cells in vitro, but only partially impaired normal lymphocyte development. 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