{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T13:51:21Z","timestamp":1791294681380,"version":"4.1.0"},"reference-count":69,"publisher":"American Society for Microbiology","issue":"11","license":[{"start":{"date-parts":[[2007,11,1]],"date-time":"2007-11-01T00:00:00Z","timestamp":1193875200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/journals.asm.org\/non-commercial-tdm-license"}],"content-domain":{"domain":["journals.asm.org"],"crossmark-restriction":true},"short-container-title":["Infect Immun"],"published-print":{"date-parts":[[2007,11]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    The intracellular pathogen\n                    <jats:italic>Listeria monocytogenes<\/jats:italic>\n                    escapes from a phagosomal compartment into the cytosol by secreting the pore-forming cytolysin listeriolysin O (LLO). During the proliferation of\n                    <jats:italic>L. monocytogenes<\/jats:italic>\n                    bacteria in the mammalian cell cytosol, the secreted LLO is targeted for degradation by the ubiquitin system. We report here that LLO is a substrate of the ubiquitin-dependent N-end rule pathway, which recognizes LLO through its N-terminal Lys residue. Specifically, we demonstrated by reverse-genetic and pharmacological methods that LLO was targeted for degradation by the N-end rule pathway in reticulocyte extracts and mouse NIH 3T3 cells and after its secretion by intracellular bacteria into the mouse cell cytosol. Replacing the N-terminal Lys of LLO with a stabilizing residue such as Val increased the in vivo half-life of LLO but did not strongly affect the intracellular growth or virulence of\n                    <jats:italic>L. monocytogenes<\/jats:italic>\n                    . Nevertheless, this replacement decreased the virulence of\n                    <jats:italic>L. monocytogenes<\/jats:italic>\n                    by nearly twofold, suggesting that a destabilizing N-terminal residue of LLO may stem from positive selection during the evolution of this and related bacteria. A double mutant strain of\n                    <jats:italic>L. monocytogenes<\/jats:italic>\n                    in which upregulated secretion of LLO was combined with a stabilizing N-terminal residue was severely toxic to infected mammalian cells, resulting in reduced intracellular growth of bacteria and an \u223c100-fold-lower level of virulence. In summary, we showed that LLO is degraded by the N-end rule pathway and that the degradation of LLO can reduce the toxicity of\n                    <jats:italic>L. monocytogenes<\/jats:italic>\n                    during infection, a property of LLO that may have been selected for its positive effects on fitness during the evolution of\n                    <jats:italic>L. monocytogenes<\/jats:italic>\n                    .\n                  <\/jats:p>","DOI":"10.1128\/iai.00164-07","type":"journal-article","created":{"date-parts":[[2007,8,6]],"date-time":"2007-08-06T20:54:18Z","timestamp":1186433658000},"page":"5135-5147","update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":55,"title":["Listeriolysin O Secreted by\n                    <i>Listeria monocytogenes<\/i>\n                    into the Host Cell Cytosol Is Degraded by the N-End Rule Pathway"],"prefix":"10.1128","volume":"75","author":[{"given":"Pamela","family":"Schnupf","sequence":"first","affiliation":[{"name":"Graduate Group in Microbiology"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianmin","family":"Zhou","sequence":"additional","affiliation":[{"name":"Division of Biology, California Institute of Technology, Pasadena, California 91125"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexander","family":"Varshavsky","sequence":"additional","affiliation":[{"name":"Division of Biology, California Institute of Technology, Pasadena, California 91125"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel A.","family":"Portnoy","sequence":"additional","affiliation":[{"name":"Graduate Group in Microbiology"},{"name":"Department of Molecular and Cellular Biology"},{"name":"School of Public Health, University of California, Berkeley, California 94720-3202"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"235","reference":[{"key":"e_1_3_2_2_2","first-page":"1","volume":"257","year":"2001","unstructured":"Alouf, J. 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