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Toxicol."],"abstract":"<jats:p>\n                    We previously demonstrated that perinatal exposure to a glyphosate-based herbicide (GBH) or pure glyphosate (Gly) induces intrauterine growth restriction (IUGR) in second-generation (F2) rats. Given the association between IUGR and fetal cardiac remodeling, we investigated whether ancestral exposure programs structural, mitochondrial, and metabolic alterations in F2 fetal hearts. Pregnant F0 rats received GBH or Gly (3.88 and 3.83\u00a0mg glyphosate\/kg\/day, respectively) from gestational day (GD) 9 to F1 weaning. F1 females were subsequently mated and euthanized at GD19 to collect male and female F2 fetal hearts. Both treatments increased relative heart weight in males, while GBH induced left ventricle dilation in both sexes, compatible with a globular phenotype. In males, Gly reduced cardiomyocyte proliferation and\n                    <jats:italic>Dio2<\/jats:italic>\n                    expression, and both treatments increased\n                    <jats:italic>Vcam1<\/jats:italic>\n                    expression. GBH also decreased mitochondrial DNA copy number while upregulating\n                    <jats:italic>Nd1<\/jats:italic>\n                    and mitochondrial biogenesis markers, suggesting a compensatory response. In females, GBH increased\n                    <jats:italic>Pdk4<\/jats:italic>\n                    and\n                    <jats:italic>Pgc1-\u03b1<\/jats:italic>\n                    expression, and Gly increased\n                    <jats:italic>Ppar\u03b1<\/jats:italic>\n                    and\n                    <jats:italic>Acadl<\/jats:italic>\n                    expression, indicating a metabolic reprogramming. Autofluorescence lifetime imaging revealed sex- and treatment-dependent alterations in NAD(P)H and FAD lifetimes. Overall, effects were more pronounced in GBH-exposed lineage and in males, whereas Gly induced subtler alterations. These findings identify the fetal heart as a multigenerational target of herbicide exposure and highlight the contribution of co-formulants to developmental toxicity.\n                  <\/jats:p>","DOI":"10.3389\/ftox.2026.1870011","type":"journal-article","created":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T22:10:52Z","timestamp":1787609452000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Sex-specific cardiac remodeling and metabolic reprogramming in IUGR F2 fetuses following glyphosate or glyphosate-based herbicide exposure"],"prefix":"10.3389","volume":"8","author":[{"given":"Ail\u00edn","family":"Almir\u00f3n","sequence":"first","affiliation":[{"name":"Instituto de Salud y Ambiente del Litoral (ISAL), Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET) \u2013 Universidad Nacional del Litoral (UNL)","place":["Santa Fe, Argentina"]},{"name":"C\u00e1tedra de Fisiolog\u00eda Humana, Facultad de Bioqu\u00edmica y Ciencias Biol\u00f3gicas (FBCB) \u2013 UNL","place":["Santa Fe, Argentina"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Virginia","family":"Lorenz","sequence":"additional","affiliation":[{"name":"Instituto de Salud y Ambiente del Litoral (ISAL), Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET) \u2013 Universidad Nacional del Litoral (UNL)","place":["Santa Fe, Argentina"]},{"name":"C\u00e1tedra de Fisiolog\u00eda Humana, Facultad de Bioqu\u00edmica y Ciencias Biol\u00f3gicas (FBCB) \u2013 UNL","place":["Santa Fe, Argentina"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Javier","family":"Adur","sequence":"additional","affiliation":[{"name":"Instituto de Investigaci\u00f3n y Desarrollo en Bioingenier\u00eda y Bioinform\u00e1tica (IBB), CONICET \u2013 Universidad Nacional de Entre R\u00edos (UNER)","place":["Oro Verde, Entre R\u00edos, Argentina"]},{"name":"Laboratorio de Microscop\u00eda Aplicada a Estudios Moleculares y Celulares (LAMAE), Facultad de Ingenier\u00eda (FI) \u2013 UNER","place":["Oro Verde, Entre R\u00edos, Argentina"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan Ignacio","family":"Etchart","sequence":"additional","affiliation":[{"name":"Instituto de Investigaci\u00f3n y Desarrollo en Bioingenier\u00eda y Bioinform\u00e1tica (IBB), CONICET \u2013 Universidad Nacional de Entre R\u00edos (UNER)","place":["Oro Verde, Entre R\u00edos, Argentina"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Milena","family":"Durando","sequence":"additional","affiliation":[{"name":"Instituto de Salud y Ambiente del Litoral (ISAL), Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET) \u2013 Universidad Nacional del Litoral (UNL)","place":["Santa Fe, Argentina"]},{"name":"C\u00e1tedra de Fisiolog\u00eda Humana, Facultad de Bioqu\u00edmica y Ciencias Biol\u00f3gicas (FBCB) \u2013 UNL","place":["Santa Fe, Argentina"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda Mercedes","family":"Milesi","sequence":"additional","affiliation":[{"name":"Instituto de Salud y Ambiente del Litoral (ISAL), Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET) \u2013 Universidad Nacional del Litoral (UNL)","place":["Santa Fe, Argentina"]},{"name":"C\u00e1tedra de Fisiolog\u00eda Humana, Facultad de Bioqu\u00edmica y Ciencias Biol\u00f3gicas (FBCB) \u2013 UNL","place":["Santa Fe, Argentina"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,8,25]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"590","DOI":"10.3390\/toxics12080590","article-title":"Perinatal exposure to glyphosate or a commercial formulation alters uterine mechanistic pathways associated with implantation failure in rats","volume":"12","author":"Almir\u00f3n","year":"2024","journal-title":"Toxics"},{"key":"B2","doi-asserted-by":"publisher","first-page":"105065","DOI":"10.1016\/J.ETAP.2026.105065","article-title":"Multigenerational effects of perinatal glyphosate exposure in rats: developmental alterations, increased mortality and oxidative imbalance","volume":"125","author":"Almir\u00f3n","year":"2026","journal-title":"Environ. 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