{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T03:08:32Z","timestamp":1791256112914,"version":"4.1.0"},"reference-count":52,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2021,10,26]],"date-time":"2021-10-26T00:00:00Z","timestamp":1635206400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"China Agriculture Research System of MOF and MARA","award":["CARS-40-S20"],"award-info":[{"award-number":["CARS-40-S20"]}]},{"name":"Province Key Research and Development Program of Shaanxi","award":["2021NY-005"],"award-info":[{"award-number":["2021NY-005"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>This study aimed to evaluate the effects of fermented corn\u2013soybean meal mixed feed on intestinal barrier function and cecal microbiota in laying hens. A total of 360 Jingfen No.6 laying hens (22 wk-old) were assigned to 4 dietary treatments, which were offered basal diets (without antibiotics) containing 0, 4, 6 and 8% of fermented mixed feed respectively. The results showed that the pH value and anti-nutritional factor concentrations in fermented mixed feed were lower than those in unfermented feed (p &lt; 0.05). Moreover, fermentation in the feed significantly increased the crude protein content (p &lt; 0.05). Supplementation with fermented feed significantly reduced the crypt depth and increased the villi height:crypt depth ratio of duodenum and jejunum (p &lt; 0.05). Meanwhile, fermented feed increased the secretory immunoglobulin A content and MUC2 mRNA expression of jejunum (p &lt; 0.05). These beneficial effects were exhibited at the addition level \u22656% and microbial composition of caeca in the control, and so 6% fermented feed groups were analyzed. The structure of the gut microbiota was remarkably altered by additions, characterized by increased abundances of some health-promoting bacteria, such as Parasutterella, Butyricicoccus and Erysipelotrichaceae (p &lt; 0.05). In summary, fermented mixed feed modulated cecal flora, subsequently contributing to improvements in intestinal morphology and barrier functions in laying hens.<\/jats:p>","DOI":"10.3390\/ani11113059","type":"journal-article","created":{"date-parts":[[2021,10,26]],"date-time":"2021-10-26T23:49:52Z","timestamp":1635292192000},"page":"3059","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":40,"title":["Fermented Corn\u2013Soybean Meal Mixed Feed Modulates Intestinal Morphology, Barrier Functions and Cecal Microbiota in Laying Hens"],"prefix":"10.3390","volume":"11","author":[{"given":"Yinglu","family":"Liu","sequence":"first","affiliation":[{"name":"College of Animal Science and Technology, Northwest A&F University, Yangling 712100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jia","family":"Feng","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Northwest A&F University, Yangling 712100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yamin","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Northwest A&F University, Yangling 712100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jing","family":"Lv","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Northwest A&F University, Yangling 712100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinghe","family":"Li","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Northwest A&F University, Yangling 712100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lijuan","family":"Guo","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Northwest A&F University, Yangling 712100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuna","family":"Min","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Northwest A&F University, Yangling 712100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,10,26]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1186\/s13568-020-00989-6","article-title":"Probiotics mitigating subclinical necrotic enteritis (SNE) as potential alternatives to antibiotics in poultry","volume":"10","author":"Khalique","year":"2020","journal-title":"AMB Express"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"900","DOI":"10.1016\/S2095-3119(16)61523-X","article-title":"Consuming fermented distillers\u2019 dried grains with solubles (DDGS) feed reveals a shift in the faecal microbiota of growing and fattening pigs using 454 pyrosequencing","volume":"16","author":"Wang","year":"2017","journal-title":"J. Integr. Agric."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1111\/j.1365-2672.2008.03938.x","article-title":"Fermented pig liquid feed: Nutritional, safety and regulatory aspects","volume":"106","year":"2009","journal-title":"J. Appl. Microbiol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1128\/mSystems.00501-19","article-title":"Dynamics of the Physicochemical Characteristics, Microbiota, and Metabolic Functions of Soybean Meal and Corn Mixed Substrates during Two-Stage Solid-State Fermentation","volume":"5","author":"Wang","year":"2020","journal-title":"mSystems"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1080\/00071661003781864","article-title":"Solid state fermented potato pulp can be used as poultry feed","volume":"51","author":"Wang","year":"2010","journal-title":"Br. Poult. Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1586","DOI":"10.3382\/ps.2009-00041","article-title":"Screening of lactic acid bacteria toward their selection as a source of chicken probiotic","volume":"88","author":"Taheri","year":"2009","journal-title":"Poult. Sci."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Jin, W., Zhang, Z., Zhu, K., Xue, Y., Xie, F., and Mao, S. (2019). Comprehensive Understanding of the Bacterial Populations and Metabolites Profile of Fermented Feed by 16S rRNA Gene Sequencing and Liquid Chromatography-Mass Spectrometry. Metabolites, 9.","DOI":"10.3390\/metabo9100239"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1186\/s13567-018-0538-6","article-title":"Biomarkers for monitoring intestinal health in poultry: Present status and future perspectives","volume":"49","author":"Ducatelle","year":"2018","journal-title":"Vet. Res."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Hiippala, K., Jouhten, H., Ronkainen, A., Hartikainen, A., Kainulainen, V., Jalanka, J., and Satokari, R. (2018). The Potential of Gut Commensals in Reinforcing Intestinal Barrier Function and Alleviating Inflammation. Nutrients, 10.","DOI":"10.3390\/nu10080988"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.aninu.2018.11.001","article-title":"Recent advances in fermented feeds towards improved broiler chicken performance, gastrointestinal tract microecology and immune responses: A review","volume":"5","author":"Sugiharto","year":"2019","journal-title":"Anim. Nutr."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.anifeedsci.2011.12.021","article-title":"Fermented liquid feed-Microbial and nutritional aspects and impact on enteric diseases in pigs","volume":"173","author":"Canibe","year":"2012","journal-title":"Anim. Feed Sci. Technol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"228","DOI":"10.1080\/00071660902736722","article-title":"Fermented feed for laying hens: Effects on egg production, egg quality, plumage condition and composition and activity of the intestinal microflora","volume":"50","author":"Engberg","year":"2009","journal-title":"Br. Poult Sci."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Semjon, B., Bartkovsky, M., Marcincakova, D., Klempova, T., Bujnak, L., Hudak, M., Jaduttova, I., Certik, M., and Marcincak, S. (2020). Effect of Solid-State Fermented Wheat Bran Supplemented with Agrimony Extract on Growth Performance, Fatty Acid Profile, and Meat Quality of Broiler Chickens. Animals, 10.","DOI":"10.3390\/ani10060942"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Xie, Y., Liu, J., Wang, H., Luo, J., Chen, T., Xi, Q., Zhang, Y., and Sun, J. (2020). Effects of fermented feeds and ginseng polysaccharides on the intestinal morphology and microbiota composition of Xuefeng black-bone chicken. PLoS ONE, 15.","DOI":"10.1371\/journal.pone.0237357"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"263","DOI":"10.5713\/ajas.2010.90145","article-title":"Effects of Feeding Solid-state Fermented Rapeseed Meal on Performance, Nutrient Digestibility, Intestinal Ecology and Intestinal Morphology of Broiler Chickens","volume":"23","author":"Chiang","year":"2009","journal-title":"Asian Australas. J. Anim. Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"2941","DOI":"10.1007\/s00253-018-8829-4","article-title":"Microbiota in fermented feed and swine gut","volume":"102","author":"Wang","year":"2018","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"868","DOI":"10.3923\/ijps.2006.868.872","article-title":"Feeding of Fermented Soybean Meal on Broiler Performance","volume":"5","author":"Mathivanan","year":"2006","journal-title":"Int. J. Poult. Sci."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1016\/j.anifeedsci.2006.06.003","article-title":"Development of a microbial fermentation process for detoxification of gossypol in cottonseed meal","volume":"135","author":"Zhang","year":"2007","journal-title":"Anim. Feed Sci. Technol."},{"key":"ref_19","first-page":"3996","article-title":"Amino acid and phosphorus digestibility of fermented corn-soybean meal mixed feed with Bacillus subtilis and Enterococcus faecium fed to pigs","volume":"95","author":"Shi","year":"2017","journal-title":"J. Anim. Sci."},{"key":"ref_20","unstructured":"AOAC (2006). Official Methods of Analysis, Association of Official Analytical Chemists. [18th ed.]."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.jcs.2013.11.006","article-title":"The influence of fermentation processes and cereal grains in wholegrain bread on reducing phytate content","volume":"59","author":"Buddrick","year":"2014","journal-title":"J. Cereal. Sci."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1089\/jmf.2004.7.430","article-title":"Aspergillus oryzae GB-107 fermentation improves nutritional quality of food soybeans and feed soybean meals","volume":"7","author":"Hong","year":"2004","journal-title":"J. Med. Food"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"140","DOI":"10.1016\/j.ceb.2004.01.005","article-title":"Setting up a selective barrier at the apical junction complex","volume":"16","author":"Anderson","year":"2004","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1038\/nri3608","article-title":"Intestinal epithelial cells: Regulators of barrier function and immune homeostasis","volume":"14","author":"Peterson","year":"2014","journal-title":"Nat. Rev. Immunol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1048","DOI":"10.3390\/ijms11031048","article-title":"Effect of soybean variety on anti-nutritional factors content, and growth performance and nutrients metabolism in rat","volume":"11","author":"Gu","year":"2010","journal-title":"Int. J. Mol. Sci."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1016\/j.lwt.2015.12.002","article-title":"Improvement of bioactivity of soybean meal by solid-state fermentation with Bacillus amyloliquefaciens versus Lactobacillus spp. and Saccharomyces cerevisiae","volume":"68","author":"Chi","year":"2016","journal-title":"LWT Food Sci. Technol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"3402","DOI":"10.1007\/s12010-014-0779-4","article-title":"Control of spoilage fungi by protective lactic acid bacteria displaying probiotic properties","volume":"172","author":"Varsha","year":"2014","journal-title":"Appl. Biochem. Biotechnol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1186\/s40104-017-0184-2","article-title":"Solid-state fermentation of corn-soybean meal mixed feed with Bacillus subtilis and Enterococcus faecium for degrading antinutritional factors and enhancing nutritional value","volume":"8","author":"Shi","year":"2017","journal-title":"J. Anim. Sci. Biotechnol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2608","DOI":"10.1007\/s13197-012-0787-8","article-title":"Effect of fermentation on antinutrients, and total and extractable minerals of high and low phytate corn genotypes","volume":"51","author":"Sokrab","year":"2014","journal-title":"J. Food Sci. Technol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) Method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1635","DOI":"10.5713\/ajas.2008.80032","article-title":"Characteristics of Solid-state Fermented Feed and its Effects on Performance and Nutrient Digestibility in Growing-finishing Pigs","volume":"21","author":"Hu","year":"2008","journal-title":"Asian Australs. J. Anim. Sci."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1136\/gut.20.4.291","article-title":"Comparison of oral feeding of peptide and amino acid meals to normal human subjects","volume":"20","author":"Silk","year":"1979","journal-title":"Gut"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1080\/00071668.2013.811718","article-title":"Effect of fermented moist feed on performance, gut bacteria and gut histo-morphology in broilers","volume":"54","author":"Missotten","year":"2013","journal-title":"Br. Poult. Sci."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1080\/00071668.2020.1801988","article-title":"Probiotic fermented feed improved the production, health and nutrient utilisation of yellow-feathered broilers reared in high altitude in Tibet","volume":"61","author":"Li","year":"2020","journal-title":"Br. Poult. Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"299S","DOI":"10.1093\/ajcn\/55.1.299s","article-title":"Physiology and pathophysiology of intestinal absorption","volume":"55","author":"Caspary","year":"1992","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1030","DOI":"10.1093\/ps\/82.6.1030","article-title":"Effects of dietary fructooligosaccharide on digestive enzyme activities, intestinal microflora and morphology of male broilers","volume":"82","author":"Xu","year":"2003","journal-title":"Poult. Sci."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2034","DOI":"10.3382\/ps\/pey035","article-title":"Effects of Pediococcus acidilactici, mannan-oligosaccharide, butyric acid and their combination on growth performance and intestinal health in young broiler chickens challenged with Salmonella Typhimurium","volume":"97","author":"Jazi","year":"2018","journal-title":"Poult. Sci."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/j.anifeedsci.2004.09.009","article-title":"Influence of processing of full fat soya beans included in diets for piglets","volume":"118","author":"Zarkadas","year":"2005","journal-title":"Anim. Feed. Sci. Technol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1008","DOI":"10.1126\/science.aaz5807","article-title":"Structure of the secretory immunoglobulin A core","volume":"367","author":"Kumar","year":"2020","journal-title":"Science"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1007\/s004410050909","article-title":"Mucosal surface area in chicken small intestine during development","volume":"290","author":"Mitjans","year":"1997","journal-title":"Cell Tissue Res."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"C1213","DOI":"10.1152\/ajpcell.00558.2003","article-title":"The tight junction: A multifunctional complex","volume":"286","author":"Schneeberger","year":"2004","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Awad, W.A., Hess, C., and Hess, M. (2017). Enteric Pathogens and Their Toxin-Induced Disruption of the Intestinal Barrier through Alteration of Tight Junctions in Chickens. Toxins, 9.","DOI":"10.3390\/toxins9020060"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"4659","DOI":"10.1073\/pnas.1006451107","article-title":"The two mucus layers of colon are organized by the MUC2 mucin, whereas the outer layer is a legislator of host-microbial interactions","volume":"108","author":"Johansson","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1186\/s40104-020-00508-4","article-title":"Modulations of genes related to gut integrity, apoptosis, and immunity underlie the beneficial effects of Bacillus amyloliquefaciens CECT 5940 in broilers fed diets with different protein levels in a necrotic enteritis challenge model","volume":"11","author":"Doranalli","year":"2020","journal-title":"J. Anim. Sci. Biotechnol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1186\/s40104-019-0421-y","article-title":"Dietary encapsulated essential oils and organic acids mixture improves gut health in broiler chickens challenged with necrotic enteritis","volume":"11","author":"Pham","year":"2020","journal-title":"J. Anim. Sci. Biotechnol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1520","DOI":"10.1038\/s41396-019-0364-5","article-title":"Defining the role of Parasutterella, a previously uncharacterized member of the core gut microbiota","volume":"13","author":"Ju","year":"2019","journal-title":"ISME J."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1416","DOI":"10.3389\/fmicb.2016.01416","article-title":"The Probiotic Butyricicoccus pullicaecorum Reduces Feed Conversion and Protects from Potentially Harmful Intestinal Microorganisms and Necrotic Enteritis in Broilers","volume":"7","author":"Eeckhaut","year":"2016","journal-title":"Front. Microbiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1361","DOI":"10.1007\/s00253-011-3847-5","article-title":"Intestinal microbiota associated with differential feed conversion efficiency in chickens","volume":"96","author":"Stanley","year":"2012","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/03079450802577848","article-title":"Tissue distribution of haemolytic Gallibacterium anatis isolates in laying birds with reproductive disorders","volume":"38","author":"Neubauer","year":"2009","journal-title":"Avian Pathol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1214","DOI":"10.1099\/ijs.0.041376-0","article-title":"Description of Alloprevotella rava gen. nov., sp. nov., isolated from the human oral cavity, and reclassification of Prevotella tannerae Moore et al. 1994 as Alloprevotella tannerae gen. nov., comb. nov.","volume":"63","author":"Downes","year":"2013","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1186\/s40104-021-00600-3","article-title":"Dietary oregano essential oil supplementation improves intestinal functions and alters gut microbiota in late-phase laying hens","volume":"12","author":"Feng","year":"2021","journal-title":"J. Anim. Sci. Biotechnol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1178","DOI":"10.1016\/j.psj.2020.10.052","article-title":"Comparison of intestinal permeability, morphology, and ileal microbial communities of commercial hens housed in conventional cages and cage-free housing systems","volume":"100","author":"Wiersema","year":"2021","journal-title":"Poult. Sci."}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/11\/11\/3059\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T07:24:15Z","timestamp":1760167455000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/11\/11\/3059"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,10,26]]},"references-count":52,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2021,11]]}},"alternative-id":["ani11113059"],"URL":"https:\/\/doi.org\/10.3390\/ani11113059","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,10,26]]}}}