Acknowledgement
This work was supported by Major Science and Technology Special Project of the Inner Mongolia Autonomous Region (Project NO. 2021ZD0024-4); National Natural Science Foundation of China (Project NO. 32360842).
References
- Chacon RR, Pacífico C, Ricci S, et al. Prolonged feeding of high-concentrate diet remodels the hindgut microbiome and modulates nutrient degradation in the rumen and the total gastrointestinal tract of cows. J Dairy Sci 2024;107:9235-50. https://doi.org/10.3168/JDS.2024-24919
- Ma J, Li T, Lin L, et al. High-concentrate diet supplemented with hydrolysable tannin improves the slaughter performance, intestinal antioxidant ability and barrier function of fattening lambs. Front Vet Sci 2024;11:1464314. https://doi. org/10.3389/fvets.2024.1464314
- Wang K, Peng X, Lv F, et al. Microbiome-metabolites analysis reveals unhealthy alterations in the gut microbiota but improved meat quality with a high-rice diet challenge in a small ruminant model. Animals 2021;11:2306. https://doi.org/10. 3390/ani11082306 https://doi.org/10.3390/ani11082306
- Tao S, Duanmu Y, Dong H, et al. High concentrate diet induced mucosal injuries by enhancing epithelial apoptosis and inflammatory response in the hindgut of goats. PLOS ONE 2014;9:e111596. https://doi.org/10.1371/journal.pone.0111596
- Mu C, Hao X, Zhang X, Zhao J, Zhang J. Effects of high-concentrate diet supplemented with grape seed procyanidins on the colonic fermentation, colonic morphology, and inflammatory response in lambs. Anim Feed Sci Technol 2021;281: 115118. https://doi.org/10.1016/j.anifeedsci.2021.115118
- An Y, Wang H, Gao A, et al. Effects of Sophora alopecuroides in a high-concentrate diet on the liver immunity and antioxidant function of lambs according to transcriptome analysis. Animals 2024;14:182. https://doi.org/10.3390/ani14020182
- Chen M, Xie W, Zhou S, et al. A high-concentrate diet induces colonic inflammation and barrier damage in Hu sheep. J Dairy Sci 2023;106:9644-62. https://doi.org/10.3168/jds.2023- 23359
- Khafipour E, Krause DO, Plaizier JC. A grain-based subacute ruminal acidosis challenge causes translocation of lipopolysaccharide and triggers inflammation. J Dairy Sci 2009;92: 1060-70. https://doi.org/10.3168/jds.2008-1389
- Suzuki T, Hara H. Quercetin enhances intestinal barrier function through the assembly of zonnula occludens-2, occludin, and claudin-1 and the expression of claudin-4 in Caco-2 cells. J Nutr 2009;139:965-74. https://doi.org/10.3945/ jn.108.100867
- Suzuki T, Tanabe S, Hara H. Kaempferol enhances intestinal barrier function through the cytoskeletal association and expression of tight junction proteins in Caco-2 cells. J Nutr 2011;141:87-94. https://doi.org/10.3945/jn.110.125633
- Mao SY, Huo WJ, Zhu WY. Microbiome–metabolome analysis reveals unhealthy alterations in the composition and metabolism of ruminal microbiota with increasing dietary grain in a goat model. Environ Microbiol 2016;18:525-41. https:// doi.org/10.1111/1462-2920.12724
- Zhang R, Liu J, Jiang L, Wang X, Mao S. The remodeling effects of high-concentrate diets on microbial composition and function in the hindgut of dairy cows. Front Nutr 2022;8: 809406. https://doi.org/10.3389/fnut.2021.809406
- Li Y, Sun YK, Li X, et al. Effects of Acremonium terricola culture on performance, milk composition, rumen fermentation and immune functions in dairy cows. Anim Feed Sci Technol 2018;240:40-51. https://doi.org/10.1016/j.anifeedsci.2018.03. 015
- Li S, Wang H, Li B, et al. Multi-omics analysis reveals the negative effects of high-concentrate diets on the colonic epithelium of dumont lambs. Animals 2025;15:749. https://doi. org/10.3390/ani15050749
- Wang Y, Xu L, Liu J, Zhu W, Mao S. A high grain diet dynamically shifted the composition of mucosa-associated microbiota and induced mucosal injuries in the colon of sheep. Front Microbiol 2017;8:2080. https://doi.org/10.3389/ fmicb.2017.02080
- Ye H, Liu J, Feng P, Zhu W, Mao S. Grain-rich diets altered the colonic fermentation and mucosa-associated bacterial communities and induced mucosal injuries in goats. Sci Rep 2016;6:20329. https://doi.org/10.1038/srep20329
- Sharma V, Kaur R, Bhatnagar A, Kaur J. Low-pH-induced apoptosis: role of endoplasmic reticulum stress-induced calcium permeability and mitochondria-dependent signaling. Cell Stress Chaperones 2015;20:431-40. https://doi.org/10. 1007/s12192-014-0568-6 https://doi.org/10.1007/s12192-014-0568-6
- Yan A, Ding H, Liu J, et al. Black Lycium barbarum polysaccharide attenuates LPS-induced intestine damage via regulation gut microbiota. Front Microbiol 2023;13:1080922. https: //doi.org/10.3389/fmicb.2022.1080922
- Tao S, Luo Y, He B, et al. Paraoxonase 2 modulates a proapoptotic function in LS174T cells in response to quorum sensing molecule N-(3-oxododecanoyl)-L-homoserine lactone. Sci Rep 2016;6:28778. https://doi.org/10.1038/srep28778
- Tao S, Tian P, Luo Y, et al. Microbiome-metabolome responses to a high-grain diet associated with the hind-gut health of goats. Front Microbiol 2017;8:1764. https://doi.org/10.3389/ fmicb.2017.01764
- Plaizier JC, Danscher AM, Azevedo PA, Derakhshani H, Andersen PH, Khafipour E. A grain-based sara challenge affects the composition of epimural and mucosa-associated bacterial communities throughout the digestive tract of dairy cows. Animals 2021;11:1658. https://doi.org/10.3390/ani1106 1658
- Zhu C, Wu Y, Jiang Z, et al. Dietary soy isoflavone attenuated growth performance and intestinal barrier functions in weaned piglets challenged with lipopolysaccharide. Int Immunopharmacol 2015;28:288-94. https://doi.org/10.1016/j. intimp.2015.04.054
- Zou Y, Wei HK, Xiang QH, Wang J, Zhou YF, Peng J. Protective effect of quercetin on pig intestinal integrity after transport stress is associated with regulation oxidative status and inflammation. J Vet Med Sci 2016;78:1487-94. https://doi.org/10.1292/jvms.16-0090
- Verediano TA, Martino HSD, Dias Paes MC, Tako E. Effects of anthocyanin on intestinal health: a systematic review. Nutrients 2021;13:1331. https://doi.org/10.3390/nu13041331
- Zheng C, Li F, Li F, et al. Effects of tannic acid addition in milk replacer on development of gastrointestinal tract of 7 to 28 days old Hu lambs. Sci Agric Sin 2019;52:3924-33. https://doi.org/10.3864/j.issn.0578-1752.2019.21.019
- Li Y, Meng XY, Ling Y, et al. Effects of tea saponin on intestinal mucosa morphology and intestinal flora of Anhui white goat. J Yangzhou Univ (Agric Life Sci Ed) 2021;42:25-32. https://doi.org/10.16872/j.cnki.1671-4652.2021.05.005
- Zhao MD, Di LF, Tang ZY, Jiang W, Li CY. Effect of tannins and cellulase on growth performance, nutrients digestibility, blood profiles, intestinal morphology and carcass characteristics in Hu sheep. Asian-Australas J Anim Sci 2019;32:1540-7. https://doi.org/10.5713/ajas.18.0901
- Xu Y, Yin F, Wang J, et al. Effect of tea polyphenols on intestinal barrier and immune function in weaned lambs. Front Vet Sci 2024;11:1361507. https://doi.org/10.3389/fvets.2024.1361507
- Tolentino MLDL, Tolentino LHO, Gonçalves LMF, et al. Ruminal and intestinal morphometric parameters of lambs fed diets containing Mimosa tenuiflora (Willd.) hay replacing Buffel grass hay. Small Rumin Res 2025;247:107493. https://doi.org/10.1016/j.smallrumres.2025.107493
- Li X, Meng M, Shi H, Zhou S, Ma N, Shen X. Dietary supplementation of yeast polysaccharides enhance lamb growth performance by improving immune and intestinal barrier function and the abundance of cecal microbiota. Int J Biol Macromol 2025;309:142849. https://doi.org/10.1016/j.ijbiomac.2025.142849
- Yang W, Guo G, Chen J, et al. Effects of dietary fucoidan supplementation on serum biochemical parameters, small intestinal barrier function, and cecal microbiota of weaned goat kids. Animals 2022;12:1591. https://doi.org/10.3390/ani12121591
- Hu QY, Man JJ, Luo J, et al. Early-life supplementation with mannan-rich fraction to regulate rumen microbiota, gut health, immunity, and growth performance in dairy goat kids. J Dairy Sci 2024;107:9322-33. https://doi.org/10.3168/jds.2024-24903
- Yuan C, Wang S, Gebeyew K, et al. A low-carbon high inulin diet improves intestinal mucosal barrier function and immunity against infectious diseases in goats. Front Vet Sci 2023;9: 1098651. https://doi.org/10.3389/fvets.2022.1098651
- Bujňáková D, Kucková K, Váradyová Z, et al. Effects of dietary zinc and/or an herbal mixture on intestinal microbiota and barrier integrity in lambs. Agriculture 2023;13:1819. https://doi.org/10.3390/agriculture13091819
- Liu M, Liu W, Zhang W, Yao J, Mo X. Ultrasound-assisted extraction of boulardii yeast cell wall polysaccharides: characterization and its biological functions on early-weaned lambs. Food Sci Nutr 2021;9:3617-30. https://doi.org/10.1002/fsn3.2318
- Guo S, Xing Y, Xu Y, Jin X, Yan S, Shi B. Progress of studies on plant-derived polysaccharides affecting intestinal barrier function in poultry. Animals 2022;12:3205. https://doi.org/10.3390/ani12223205
- Zhuang S, Ming K, Ma N, et al. Portulaca oleracea L. polysaccharide ameliorates lipopolysaccharide-induced inflammatory responses and barrier dysfunction in porcine intestinal epithelial monolayers. J Funct Foods 2022;91:104997. https://doi.org/10.1016/j.jff.2022.104997
- Xu C, Wang K, Ding YH, Li WJ, Ding L. Claudin-7 gene knockout causes destruction of intestinal structure and animal death in mice. World J Gastroenterol 2019;25:584-99. https://doi.org/10.3748/wjg.v25.i5.584
- Bruning EE, Coller JK, Wardill HR, Bowen JM. Site-specific contribution of toll-like receptor 4 to intestinal homeostasis and inflammatory disease. J Cell Physiol 2020;236:877-88. https://doi.org/10.1002/jcp.29976
- Tang X, Liu H, Yang S, Li Z, Zhong J, Fang R. Epidermal growth factor and intestinal barrier function. Med Inflamm 2016;2016:1927348. https://doi.org/10.1155/2016/1927348
- Csernus B, Biró S, Babinszky L, et al. Effect of carotenoids, oligosaccharides and anthocyanins on growth performance, immunological parameters and intestinal morphology in broiler chickens challenged with escherichia coli lipopolysaccharide. Animals 2020;10:347. https://doi.org/10.3390/ani10020347
- Ren Z, Guo C, Yu S, et al. Progress in mycotoxins affecting intestinal mucosal barrier function. Int J Mol Sci 2019;20:2777. https://doi.org/10.3390/ijms20112777
- Mei H, Li YF, Ma X, Yu M. Effects and regulatory mechanisms of quercetin on animal intestinal mucosal barrier function. Chin J Anim Nutr 2022;34:5475-88. https://doi.org/10.3969/j.issn.1006-267x.2022.09.004
- Zhang K, Xu Y, Zheng Y, et al. Bifidobacterium pseudolongum-derived bile acid from dietary carvacrol and thymol sup-plementation attenuates colitis via cGMP-PKG-mTORC1 pathway. Adv Sci 2024;11:2406917. https://doi.org/10.1002/advs.202406917
- Volstatova T, Marchica A, Hroncova Z, Bernardi R, Doskocil I, Havlik J. Effects of chlorogenic acid, epicatechin gallate, and quercetin on mucin expression and secretion in the Caco-2/HT29-MTX cell model. Food Sci Nutr 2019;7:492-8. https://doi.org/10.1002/fsn3.818
- Shi L, Xun W, Peng W, Hu H, Cao T, Hou G. Effect of the single and combined use of curcumin and piperine on growth performance, intestinal barrier function, and antioxidant capacity of weaned wuzhishan piglets. Front Vet Sci 2020;7:418. https://doi.org/10.3389/fvets.2020.00418
- Jia L, Wu J, Lei Y, et al. Oregano essential oils mediated intestinal microbiota and metabolites and improved growth performance and intestinal barrier function in sheep. Front Immunol 2022;13:908015. https://doi.org/10.3389/fimmu.2022.908015
- Cheng Y, Liu Y, Chen D, et al. Dual effects of quercetin on protein digestion and absorption in the digestive tract. Food Chem 2021;358:129891. https://doi.org/10.1016/j.foodchem.2021.129891
- Tong Z, Lei F, Liu L, Wang F, Guo A. Effects of Plotytarya strohilacea Sieb. et zuce tannin on the growth performance, oxidation resistance, intestinal morphology and cecal microbial composition of broilers. Front Vet Sci 2022;8:806105. https://doi.org/10.3389/fvets.2021.806105
- Pelegrin-Valls J, Álvarez-Rodríguez J, Martín-Alonso MJ, et al. Effect of maternal dietary condensed tannins from sainfoin (Onobrychis viciifolia) on gut health and antioxidantimmune crosstalk in suckling lambs. Agriculture 2022;12:1694. https://doi.org/10.3390/agriculture12101694
- Xiao Y, Chen L, Xu Y, He X, Gan S, Yin F. The effects of tea polyphenols in feed on the immunity, antioxidant capacity, and gut microbiota of weaned goat kids. Animals 2025;15:467. https://doi.org/10.3390/ani15040467
- Zhao X, Song JL, Yi R, et al. Comparison of antioxidative effects of insect tea and its raw tea (kuding tea) polyphenols in kunming mice. Molecules 2018;23:204. https://doi.org/10.3390/molecules23010204
- Imam H, Zhang KQ, Wu H, Xiao JS, Huang MQ, Cao YP. Epigallocatechin gallate (EGCG) inhibits lipopolysaccharideinduced inflammation in RAW 264.7 macrophage cells via modulating nuclear factor kappa-light-chain enhancer of activated B cells (NF-κB) signaling pathway. Food Sci Nutr 2023;11:4634-50. https://doi.org/10.1002/fsn3.3427
- Singh R, Chandrashekharappa S, Bodduluri SR, et al. Enhancement of the gut barrier integrity by a microbial metabolite through the Nrf2 pathway. Nat Commun 2019;10:89. https://doi.org/10.1038/s41467-018-07859-7
- Mu CT, Wei H, Xie YZ, Zhang JX, Hao XY. Effect of grape seed proanthocyanidins on colon epithelial barrier of sheep fed a high-concentrate diet and its mechanism. China Anim Husb Vet Med 2024;51:4270-81. https://doi.org/10.16431/j.cnki.1671-7236.2024.10.008
- Li H, Gu Y, Jin R, He Q, Zhou Y. Effects of dietary rutin supplementation on the intestinal morphology, antioxidant capacity, immunity, and microbiota of aged laying hens. Antioxidants 2022;11:1843. https://doi.org/10.3390/antiox11091843
- Heidebrecht HJ, Kulozik U. Fractionation of casein micelles and minor proteins by microfiltration in diafiltration mode. Study of the transmission and yield of the immunoglobulins IgG, IgA and IgM. Int Dairy J 2019;93:1-10. https://doi.org/10.1016/j.idairyj.2019.01.009
- Lv X, Chen L, Zhou CS, et al. Dietary tea tree (Melaleuca alternifolia) oil supplementation enhances the expressions of amino acid transporters in goat ileal mucosa and improves intestinal immunity. Food Sci Nutr 2022;10:3749-58. https://doi.org/10.1002/fsn3.2972
- Lu Y, Li X, Liu S, Zhang Y, Zhang D. Toll-like receptors and inflammatory bowel disease. Front Immunol 2018;9:72. https://doi.org/10.3389/fimmu.2018.00072
- Shao X, Sun C, Tang X, et al. Anti-inflammatory and intestinal microbiota modulation properties of jinxiang garlic (Allium sativum L.) polysaccharides toward dextran sodium sulfate-induced colitis. J Agric Food Chem 2020;68:12295-309. https://doi.org/10.1021/acs.jafc.0c04773
- Niu JH, Yuan J, Wei R, et al. Protective effect and mechanism study of jujube polysaccharides on intestinal immune barrier in mice. Sci Tech Food Ind 2021;42:295-300. https://doi.org/10.13386/j.issn1002-0306.2020060068
- Qu J, Huang P, Zhang L, et al. Hepatoprotective effect of plant polysaccharides from natural resources: a review of the mechanisms and structure-activity relationship. Int J Biol Macromol 2020;161:24-34. https://doi.org/10.1016/j.ijbiomac.2020.05.196
- Mao SH, Feng DD, Wang X, et al. Magnolol protects against acute gastrointestinal injury in sepsis by down-regulating regulated on activation, normal T-cell expressed and secreted. World J Clin Cases 2021;9:10451-63. https://doi.org/10.12998/wjcc.v9.i34.10451
- Yang C, Cheng Y, Li X, et al. Effects of dietary Macleaya cordata extract inclusion on transcriptomes and inflammatory response in the lower gut of early weaned goats. Anim Feed Sci Technol 2021;272:114792. https://doi.org/10.1016/j.anifeedsci.2020.114792
- Ye J, Ye C, Huang Y, Zhang N, Zhang X, Xiao M. Ginkgo biloba sarcotesta polysaccharide inhibits inflammatory responses through suppressing both NF-κB and MAPK signaling pathway. J Sci Food Agric 2018;99:2329-39. https://doi.org/10.1002/jsfa.9431
- Sun L, Xu G, Dong Y, Li M, Yang L, Lu W. Quercetin protects against lipopolysaccharide-induced intestinal oxidative stress in broiler chickens through activation of Nrf2 pathway. Molecules 2020;25:1053. https://doi.org/10.3390/molecules25051053
- Ma G, Kimatu BM, Yang W, et al. Preparation of newly identified polysaccharide from Pleurotus eryngii and its anti-inflammation activities potential. J Food Sci 2020;85:2822-31. https://doi.org/10.1111/1750-3841.15375
- Sokol H. Probiotics and antibiotics in IBD. Dig Dis 2014;32: 10-7. https://doi.org/10.1159/000367820
- Lai PK, Roy J. Antimicrobial and chemopreventive properties of herbs and spices. Curr Med Chem 2004;11:1451-60. https://doi.org/10.2174/0929867043365107
- Lee HC, Jenner AM, Low CS, Lee YK. Effect of tea phenolics and their aromatic fecal bacterial metabolites on intestinal microbiota. Res Microbiol 2006;157:876-84. https://doi.org/10.1016/j.resmic.2006.07.004
- Tremaroli V, Bäckhed F. Functional interactions between the gut microbiota and host metabolism. Nature 2012;489:242-9. https://doi.org/10.1038/nature11552
- Cheng X, Du X, Liang Y, et al. Effect of grape pomace supplement on growth performance, gastrointestinal microbiota, and methane production in Tan lambs. Front Microbiol 2023;14:1264840. https://doi.org/10.3389/fmicb.2023.1264840
- Chen K, Liu Y, Cheng Y, et al. Supplementation of Lactobacillus plantarum or Macleaya cordata extract alleviates oxidative damage induced by weaning in the lower gut of young goats. Animals 2020;10:548. https://doi.org/10.3390/ani10040548
- Inupala SN, Pande P, Uzzaman MR, Worku M. 121 Evaluation of effects of garlic extract on health parameters and gut health in st. croix sheep. J Anim Sci 2023;101:99-100. https://doi.org/10.1093/jas/skad068.118
- Ding S, Jiang H, Fang J, Liu G. Regulatory effect of resveratrol on inflammation induced by lipopolysaccharides via reprograming intestinal microbes and ameliorating serum metabolism profiles. Front Immunol 2021;12:777159. https://doi.org/10.3389/fimmu.2021.777159
- Wang Y, Sun W, Wu E, et al. Polysaccharides from Abrus cantoniensis Hance modulate intestinal microflora and improve intestinal mucosal barrier and liver oxidative damage induced by heat stress. Front Vet Sci 2022;9:868433. https://doi.org/10.3389/fvets.2022.868433
- Sichert A, Corzett CH, Schechter MS, et al. Verrucomicrobia use hundreds of enzymes to digest the algal polysaccharide fucoidan. Nat Microbiol 2020;5:1026-39. https://doi.org/10.1038/s41564-020-0720-2