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Despite its crucial importance, not all intracellular\n                    <jats:italic>Salmonella<\/jats:italic>\n                    express the SPI-2 T3SS. Using flow cytometry and microscopic analysis of bacteria within host cells, we demonstrate that both the injectisome as well as its effectors exhibit bimodal expression. This bimodality depended on activation of the transcriptional regulator SsrB by sensor kinase SsrA. Within an infected host cell, proliferation of bacteria not expressing the SPI-2 T3SS (SPI-2\n                    <jats:sup>OFF<\/jats:sup>\n                    ) depended on SPI-2 T3SS-expressing bacteria (SPI-2\n                    <jats:sup>ON<\/jats:sup>\n                    ), suggesting that all SPI-2 T3SS effectors necessary for\n                    <jats:italic>Salmonella<\/jats:italic>\n                    intracellular replication can be complemented\n                    <jats:italic>in trans<\/jats:italic>\n                    . SPI-2\n                    <jats:sup>OFF<\/jats:sup>\n                    bacteria had shorter division time\n                    <jats:italic>in vitro<\/jats:italic>\n                    and proliferated faster inside host cells than the SPI-2\n                    <jats:sup>ON<\/jats:sup>\n                    bacteria. SPI-2\n                    <jats:sup>OFF<\/jats:sup>\n                    bacteria egressed more from infected host cells, thus potentially serving as a reservoir for further\n                    <jats:italic>Salmonella<\/jats:italic>\n                    dissemination throughout the host. Collectively, our results suggest that bimodal expression of the SPI-2 T3SS and its effectors represents an adaptive mechanism that might increase\n                    <jats:italic>Salmonella<\/jats:italic>\n                    virulence.\n                  <\/jats:p>","DOI":"10.1371\/journal.ppat.1013728","type":"journal-article","created":{"date-parts":[[2025,12,1]],"date-time":"2025-12-01T18:43:43Z","timestamp":1764614623000},"page":"e1013728","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":7,"title":["Bimodal expression of Type 3 Secretion System 2 enables cooperative virulence among intracellular Salmonella Typhimurium"],"prefix":"10.1371","volume":"21","author":[{"given":"Milada","family":"Posp\u00ed\u0161ilov\u00e1","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alona","family":"Dreus","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paulina","family":"Matheova","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jana","family":"Schmidtova","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Barbora","family":"Pravdova","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michaela","family":"Blazikova","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martin","family":"Capek","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9570-5449","authenticated-orcid":true,"given":"Ondrej","family":"Cerny","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2025,12,1]]},"reference":[{"issue":"11","key":"ppat.1013728.ref001","doi-asserted-by":"crossref","first-page":"716","DOI":"10.1038\/s41579-021-00561-4","article-title":"Salmonella 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