{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,4]],"date-time":"2026-03-04T10:46:44Z","timestamp":1772621204633,"version":"3.50.1"},"reference-count":74,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,3,4]],"date-time":"2026-03-04T00:00:00Z","timestamp":1772582400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Vet. Sci."],"abstract":"<jats:p>\n                    Early weaning is a critical stage in pig production, associated with intestinal and immunological stress that negatively impacts gut health and functionality. Given the global restrictions on antibiotic use, natural alternatives have gained increasing attention. This study aimed to evaluate the\n                    <jats:italic>in vivo<\/jats:italic>\n                    effects of short-chain fructooligosaccharides, essential oils and sodium humate on intestinal health in weaned piglets. Four dietary treatments were applied for 15\u202fdays to weaned piglets: a basal diet (control); short-chain fructooligosaccharides (scFOSs group); essential oils of\n                    <jats:italic>L. origanoide<\/jats:italic>\n                    s and\n                    <jats:italic>E. caryophyllata<\/jats:italic>\n                    with sodium humate (EOs-SH group); and a combination of scFOSs and EO of\n                    <jats:italic>Lippia origanoides<\/jats:italic>\n                    (scFOSs-EOLo group). Plasma cortisol and citrulline concentrations, gastrointestinal pH, intestinal histomorphology, intestinal disaccharidase activity, lymphocyte infiltration, goblet cells quantification, mucus bacterial adherence, volatile fatty acids, and microbiota composition were analyzed. No significant differences in cortisol concentrations were found among treatments (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f=\u202f0.514). EOs-SH group showed significantly higher citrulline levels (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f&amp;lt;\u202f0.01), indicating enhanced enterocyte function. The activity of lactase in the proximal jejunum (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f=\u202f0.03) and maltase in the duodenum (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f=\u202f0.045) increased in scFOSs and EOs-SH groups, suggesting improved digestive capacity. Mucus bacterial adherence was also greater in EOs-SH group (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f=\u202f0.039), reflecting enhanced mucosal protective quality. Gastrointestinal pH values were within physiological ranges for the age and intestinal segment evaluated. ScFOSs-EOLo showed a higher gastric pH than the control (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f=\u202f0.033), while no differences among treatments were observed in the remaining gastrointestinal regions (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f&amp;gt;\u202f0.05). No treatment effect was observed for villus height or villus height-to-crypt depth ratio, with shallower ileal crypts in EOs-SH and ScFOSs-EOLo groups (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f=\u202f0.003), indicating improved mucosal integrity. The lowest intraepithelial lymphocyte counts in the ileum were observed in scFOSs and EOs-SH groups (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f=\u202f0.033). Short-chain fatty acid concentrations did not differ statistically among treatments (\n                    <jats:italic>p<\/jats:italic>\n                    \u202f&amp;gt;\u202f0.05). Microbiota analysis revealed that the EOs-SH treatment reduced potentially proinflammatory genera, including\n                    <jats:italic>Clostridium sensu stricto<\/jats:italic>\n                    1,\n                    <jats:italic>Terrisporobacter<\/jats:italic>\n                    ,\n                    <jats:italic>Prevotella<\/jats:italic>\n                    115, and\n                    <jats:italic>Subdoligranulum<\/jats:italic>\n                    . scFOSs supplementation markedly increased\n                    <jats:italic>Lactobacillus<\/jats:italic>\n                    , consistent with its prebiotic effect (Padj&amp;lt;0.05 in all cases). In contrast, ScFOSs-EOLo treatment did not induce any relevant abundance microbial changes (Padj&amp;gt;0.05). All dietary treatments improved some intestinal health parameters, with more consistent responses observed in the treatment with EOs-SH. This supplementation enhanced intestinal integrity and functionality, offering a natural strategy to support gut health and resilience in antibiotic-free production systems.\n                  <\/jats:p>","DOI":"10.3389\/fvets.2026.1771820","type":"journal-article","created":{"date-parts":[[2026,3,4]],"date-time":"2026-03-04T05:42:23Z","timestamp":1772602943000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Dietary modulation of intestinal integrity and functionality in weaned piglets using short-chain fructooligosaccharides, essential oils and sodium humate"],"prefix":"10.3389","volume":"13","author":[{"given":"J. M.","family":"Decundo","sequence":"first","affiliation":[{"name":"Laboratorio de Toxicolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]},{"name":"Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET), Ciudad Aut\u00f3noma de Buenos Aires","place":["Buenos Aires, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"J.","family":"Mozo","sequence":"additional","affiliation":[{"name":"Laboratorio de Toxicolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]},{"name":"Depto. de Producci\u00f3n Animal, Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"D. S.","family":"P\u00e9rez Gaudio","sequence":"additional","affiliation":[{"name":"Laboratorio de Toxicolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]},{"name":"Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET), Ciudad Aut\u00f3noma de Buenos Aires","place":["Buenos Aires, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"G.","family":"Mart\u00ednez","sequence":"additional","affiliation":[{"name":"Laboratorio de Toxicolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]},{"name":"Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET), Ciudad Aut\u00f3noma de Buenos Aires","place":["Buenos Aires, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"S. N.","family":"Dieguez","sequence":"additional","affiliation":[{"name":"Laboratorio de Toxicolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]},{"name":"Comisi\u00f3n de Investigaciones Cient\u00edficas de la Provincia de Buenos Aires (CIC-PBA)","place":["La Plata, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"C. P.","family":"Bianchi","sequence":"additional","affiliation":[{"name":"Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET), Ciudad Aut\u00f3noma de Buenos Aires","place":["Buenos Aires, Argentina"]},{"name":"Laboratorio de Endocrinolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"V. R.","family":"Egu\u00eda","sequence":"additional","affiliation":[{"name":"Laboratorio de Toxicolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"M. L.","family":"Mat\u00e9","sequence":"additional","affiliation":[{"name":"Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET), Ciudad Aut\u00f3noma de Buenos Aires","place":["Buenos Aires, Argentina"]},{"name":"Laboratorio de Farmacolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"J. P.","family":"Lir\u00f3n","sequence":"additional","affiliation":[{"name":"Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET), Ciudad Aut\u00f3noma de Buenos Aires","place":["Buenos Aires, Argentina"]},{"name":"Laboratorio de Farmacolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"F. A.","family":"Amanto","sequence":"additional","affiliation":[{"name":"Depto. de Producci\u00f3n Animal, Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"A. L.","family":"Soraci","sequence":"additional","affiliation":[{"name":"Laboratorio de Toxicolog\u00eda, Depto. de Fisiopatolog\u00eda, Centro de Investigaci\u00f3n Veterinaria de Tandil (CIVETAN), Facultad de Ciencias Veterinarias, Universidad Nacional del Centro de la Provincia de Buenos Aires","place":["Tandil, Argentina"]},{"name":"Consejo Nacional de Investigaciones Cient\u00edficas y T\u00e9cnicas (CONICET), Ciudad Aut\u00f3noma de Buenos Aires","place":["Buenos Aires, Argentina"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2026,3,4]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"186","DOI":"10.54846\/jshap\/875","article-title":"Influence of age at weaning and feeding regimes on the postnatal morphology of the porcine small intestine","volume":"23","author":"Al Masri","year":"2015","journal-title":"J Swine Health Prod"},{"key":"ref2","first-page":"89","article-title":"Effet d\u2019un sevrage \u00e0 l\u2019\u00e2ge de 7 jours et du niveau alimentaire apr\u00e8s le sevrage sur les modifications structurales de l\u2019intestin gr\u00eale chez le porcelet","volume":"34","author":"Marion","year":"2002","journal-title":"J Rech Porc Fr"},{"key":"ref3","doi-asserted-by":"publisher","first-page":"2169","DOI":"10.3390\/ani11082169","article-title":"Effects of weaning age at 21 and 28 days on growth performance, intestinal morphology and redox status in piglets","volume":"11","author":"Ming","year":"2021","journal-title":"Animals"},{"key":"ref4","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1186\/2049-1891-4-19","article-title":"The biological stress of early weaned piglets","volume":"4","author":"Campbell","year":"2013","journal-title":"J Anim Sci Biotechnol"},{"key":"ref5","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1111\/j.1439-0396.2012.01284.x","article-title":"Gastrointestinal health and function in weaned pigs: a review of feeding strategies to 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piglets","volume":"13","author":"Ma","year":"2022","journal-title":"J Anim Sci Biotechnol"},{"key":"ref21","first-page":"33","article-title":"Directive 2010\/63\/EU of the European Parliament and of the council of 22 September 2010 on the protection of animals used for scientific purposes","year":"2010","journal-title":"Off J Eur Union"},{"key":"ref22","volume-title":"Nutrient requirements of swine","year":"2012"},{"key":"ref23","first-page":"21","article-title":"Improvement of productive parameters in nursery piglets by supplementation with sodium humate and oregano and clove essential oils","volume":"25","author":"Mozo","year":"2025","journal-title":"Rev Argent Prod Anim"},{"key":"ref24","doi-asserted-by":"publisher","first-page":"1","DOI":"10.4238\/gmr.15027693","article-title":"Effect of weaning age on cortisol release in piglets","volume":"15","author":"Li","year":"2016","journal-title":"Genet Mol 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Sci"},{"key":"ref28","doi-asserted-by":"publisher","first-page":"328","DOI":"10.1016\/j.clnu.2008.02.005","article-title":"Citrulline as a biomarker of intestinal failure due to enterocyte mass reduction","volume":"27","author":"Crenn","year":"2008","journal-title":"Clin Nutr"},{"key":"ref29","doi-asserted-by":"publisher","first-page":"858","DOI":"10.2460\/javma.22.12.0543","article-title":"Citrullinemia is a suitable biomarker for post-weaning performance in piglets under intensive farming","volume":"261","author":"Soraci","year":"2023","journal-title":"J Am Vet Med Assoc"},{"key":"ref30","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1016\/S0076-6879(07)00810-5","article-title":"Analysis of citrulline, arginine, and methylarginines using high-performance liquid chromatography","volume":"440","author":"Wu","year":"2008","journal-title":"Methods Enzymol"},{"key":"ref31","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1007\/s102380200018","article-title":"A simple new method to calculate small intestine absorptive surface in the rat","volume":"2","author":"Kisielinski","year":"2002","journal-title":"Clin Exp Med"},{"key":"ref32","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1186\/s40104-021-00601-2","article-title":"Supplementation of a lacto-fermented rapeseed\u2013seaweed blend promotes gut microbial and gut-immune modulation in weaner piglets","volume":"12","author":"Hui","year":"2021","journal-title":"J Anim Sci Biotechnol"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1016\/0003-2697(64)90115-0","article-title":"Method for assay of intestinal disaccharidases","volume":"7","author":"Dahlqvist","year":"1964","journal-title":"Anal Biochem"},{"key":"ref34","first-page":"449","article-title":"Inhibitory effects of intestinal mucus on bacterial adherence to cultured intestinal epithelial cells after surface burns","volume":"113","author":"Bai","year":"2000","journal-title":"Chin Med J"},{"key":"ref35","doi-asserted-by":"publisher","first-page":"D590","DOI":"10.1093\/nar\/gks1219","article-title":"The SILVA ribosomal RNA gene database project: improved data processing and web-based tools","volume":"41","author":"Quast","year":"2013","journal-title":"Nucleic Acids Res"},{"key":"ref36","doi-asserted-by":"publisher","first-page":"e61217","DOI":"10.1371\/journal.pone.0061217","article-title":"Phyloseq: an R package for reproducible interactive analysis and graphics of microbiome census data","volume":"8","author":"McMurdie","year":"2013","journal-title":"PLoS One"},{"key":"ref37","first-page":"131","article-title":"Volatile fatty acid and alcohol determination in digestive contents, silage juices, bacterial cultures and anaerobic fermentor contents","volume":"2","author":"Jouany","year":"1982","journal-title":"Sci Aliment"},{"key":"ref38","volume-title":"RStudio: Integrated development for R","year":"2015"},{"key":"ref39","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol"},{"key":"ref40","doi-asserted-by":"publisher","first-page":"82","DOI":"10.17221\/8731-CJAS","article-title":"Transient effects of weaning on the health of newly weaned piglets","volume":"61","author":"Tao","year":"2016","journal-title":"Czeh J Anim Sci"},{"key":"ref41","doi-asserted-by":"publisher","first-page":"4879","DOI":"10.1128\/AEM.00770-14","article-title":"Lactulose and Lactobacillus plantarum, a potential complementary synbiotic to control postweaning colibacillosis in piglets","volume":"80","author":"Guerra-Ordaz","year":"2014","journal-title":"Appl Environ Microbiol"},{"key":"ref42","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1111\/jpn.12666","article-title":"Review on the effects of potential prebiotics on controlling intestinal enteropathogens in piglets","volume":"102","author":"Tran","year":"2018","journal-title":"J Anim Physiol Anim Nutr"},{"key":"ref43","doi-asserted-by":"publisher","first-page":"327","DOI":"10.1002\/ffj.1967","article-title":"Essential oils and aromatic plants in animal feeding: a European perspective","volume":"25","author":"Franz","year":"2010","journal-title":"Flavour Fragr J"},{"key":"ref44","doi-asserted-by":"publisher","first-page":"1617","DOI":"10.5713\/ajas.2012.12292","article-title":"Effects of adding essential oil to the diet of weaned pigs on performance, nutrient utilization, immune response, and intestinal health","volume":"25","author":"Li","year":"2012","journal-title":"Asian Australas J Anim Sci"},{"key":"ref45","doi-asserted-by":"publisher","first-page":"1717","DOI":"10.3390\/molecules23071717","article-title":"Essential oils as feed additives: future perspectives","volume":"23","author":"Stevanovi\u0107","year":"2018","journal-title":"Molecules"},{"key":"ref46","doi-asserted-by":"publisher","first-page":"143","DOI":"10.2527\/jas.2005-206","article-title":"Effects of dietary humic substances on pig growth performance, nutrient digestibility, and fecal microbial concentration","volume":"105","author":"Ji","year":"2006","journal-title":"Livest Sci"},{"key":"ref47","first-page":"1","article-title":"Effects of humic substances on gastrointestinal functionality and health in pigs and poultry","volume":"9","author":"Koch","year":"2019","journal-title":"Animals"},{"key":"ref48","doi-asserted-by":"publisher","first-page":"2080","DOI":"10.3390\/ani11072080","article-title":"Application of creep feed and phytase super-dosing as tools to support digestive adaption and feed efficiency in piglets at weaning","volume":"11","author":"Lee","year":"2021","journal-title":"Animals: Open Access J 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