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This phenomenon occurs even in the absence of infection when pathogen mimetics trigger paternal immune activation (PIA). While this implicates a shared component of the immune response, the specific mechanisms responsible remain unknown. Cytokines are a key component of innate immunity and are highly upregulated following exposure to both pathogens and their mimetics. Here we investigate whether paternal administration of the pro-inflammatory cytokines interleukin-1\u03b2 (IL-1\u03b2) and tumor necrosis factor-\u03b1 (TNF-\u03b1) recapitulates alterations to sperm small ncRNAs and offspring phenotypic changes from other models of PIA. C57BL\/6J male mice were administered a single dose of IL-1\u03b2, TNF-\u03b1, or vehicle, prior to mating with na\u00efve female mice, to produce offspring. Offspring from IL-1\u03b2-treated fathers exhibited altered fasting responses and female-specific alterations to stress-coping behavior. In response to paternal TNF-\u03b1, male offspring showed increased anxiety-like behavior. Analysis of paternal sperm small ncRNA revealed that IL-1\u03b2 significantly downregulated several transfer RNA-derived small RNAs (tsRNAs) and P-element-induced wimpy testis (PIWI)-interacting RNA (piRNA) clusters. Surprisingly, paternal TNF-\u03b1 only slightly altered small ncRNAs, downregulating a single piRNA cluster. These results partially recapitulate offspring phenotypic changes following paternal exposure to severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2, in a COVID-19 mouse model) and polyinosinic:polycytidylic acid (poly I:C, a viral mimetic). Ultimately, we provide the first evidence that cytokines have intergenerational effects on brain and behavior, and that they contribute to altered offspring phenotypes in viral-like PIA. This study demonstrates that paternal cytokines are\u00a0causally\u00a0involved in a mechanism of epigenetic inheritance following PIA, and highlights the\u00a0importance\u00a0of PIA as a pre-conceptual factor that may modulate offspring health, including risk of neuropsychiatric disorders.<\/jats:p>","DOI":"10.1038\/s41380-026-03674-5","type":"journal-article","created":{"date-parts":[[2026,6,18]],"date-time":"2026-06-18T23:02:26Z","timestamp":1781823746000},"page":"6183-6201","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Paternal cytokine administration alters sperm small ncRNAs and offspring brain and behavior"],"prefix":"10.1038","volume":"31","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5350-3360","authenticated-orcid":false,"given":"Pamudika","family":"Kiridena","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4525-0949","authenticated-orcid":false,"given":"Nicholas","family":"van de Garde","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bethany A.","family":"Masson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8078-1390","authenticated-orcid":false,"given":"Carolina","family":"Gubert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9242-6249","authenticated-orcid":false,"given":"Sonali N.","family":"Reisinger","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7532-8922","authenticated-orcid":false,"given":"Anthony J.","family":"Hannan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,6,19]]},"reference":[{"key":"3674_CR1","doi-asserted-by":"publisher","first-page":"1560","DOI":"10.3945\/ajcn.2009.28838","volume":"91","author":"KL Whitaker","year":"2010","unstructured":"Whitaker KL, Jarvis MJ, Beeken RJ, Boniface D, Wardle J. 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