Abstract
Keywords
Data Availability
No datasets were generated or analysed during the current study.
Abbreviations
- AA:: Amino acid
- ADF:: Acid detergent fiber
- AMP:: Adenosine monophosphate
- AMPK:: Adenosine monophosphate-activated protein kinase
- AMPK/PGC1α:: AMPK-peroxisome proliferator-activated receptorγ coactivator 1-α
- ATIC/GPAT:: IMP cyclohydrolase/glutamine-PRPP amidotransferase
- ATP5A1:: ATP synthase F1 subunit alpha
- CAT:: Catalase
- C/EBPα:: CCAAT/enhancer-binding protein α
- CF:: Crude fiber
- CLA:: Conjugated linoleic acid
- CP:: Crude protein
- DM:: Dry matter
- EAA:: Essential amino acids
- EE:: Ether extract
- FAA:: Flavor amino acids
- FABP4:: Fatty acid-binding protein 4
- FASN:: Fatty acid synthase
- GADD45a:: Growth arrest and DNA damage
- GSH-Px:: Glutathione peroxidase
- HMP:: 2-Hydroxy-4-methylpentanoic acid
- IBD:: Inflammatory bowel disease
- IMF:: Intramuscular fat
- IMP:: Inosine monophosphate
- LAB:: Lactic acid bacteria
- MDA:: Malondialdehyde
- MUFA:: Monounsaturated fatty acids
- MyHC-I:: Myosin heavy chain-I
- MyHC-IIa:: Myosin heavy chain-IIa
- MyHC-IIx:: Myosin heavy chain-IIx
- MyHC-IIb:: Myosin heavy chain-IIb
- NDF:: Neutral detergent fiber
- NEAA:: Nonessential amino acids
- n-3 PUFA:: n-3 polyunsaturated fatty acids
- NSP:: Non-starch polysaccharide
- PI3K-Akt:: Phosphoinositide 3-kinase- protein kinase
- PPARγ:: Peroxisome proliferator-activated receptor-γ
- PUFA:: Polyunsaturated fatty acids
- SCFAs:: Short-chain fatty acids
- SFA:: Saturated fatty acids
- SOD:: Superoxide dismutase
- SREBP1:: Stearoyl-CoA desaturase enzyme 1
- SSF:: Solid-state fermentation
- TAG:: Triglycerides
- T-AOC:: Total superoxide dismutase
- TCA-SP:: Trichloroacetic acid soluble protein
- UFA:: Unsaturated fatty acids
References
- 1.Chernukha I, Kotenkova E, Pchelkina V, Ilyin N, Utyanov D, Kasimova T, et al. Pork fat and meat.(2023)a balance between consumer expectations and nutrient composition of four pig breeds.Foods.: 690.
- 2.Lebret B, Čandek-Potokar M. Review.(2022)Pork quality attributes from farm to fork. Part I. Carcass and fresh meat.Animal.: 100402.
- 3.Liu YY, Kong XF, Jiang GL, Tan BE, Deng JP, Yang XJ, et al. Effects of dietary protein/energy ratio on growth performance, carcass trait, meat quality, and plasma metabolites in pigs of different genotypes. J Anim Sci Biotechnol. 2015;6.(2015)org/10.1186/s40104-015-0036-x.: 36.
- 4.Chen GS, Sui YN. Production, performance, slaughter characteristics, and meat quality of Ziwuling wild crossbred pigs. Trop Anim Health Prod. 2018;50(2).(2018)org/10.1007/s11250-017-1441-2.: 365.
- 5.Qi KK, Men XM, Wu J, Xu ZW. Rearing pattern alters porcine myofiber type, fat deposition, associated microbial communities and functional capacity. BMC Microbiol. 2019;19.(2019)org/10.1186/s12866-019-1556-x.: 181.
- 6.Lian X, Shi MY, Lin QL, Liang Y, Zhang LY. Research progress of probiotics and fermented feed effects on pork quality. Food Bioeng. 2024;3(1).(2024)1002/fbe2.12082.: 83.
- 7.Zhang S, Huang YQ, Zheng CB, Wang LY, Zhou YB, Chen WT, et al. Leucine improves the growth performance, carcass traits, and lipid nutritional quality of pork in Shaziling pigs. Meat Sci. 2024;210.(2024)109435. https://doi. org/10. 1016/j.meatsci.: 109435.
- 8.Wang LY, Zhang S, Huang YQ, You WJ, Zhou YB, Chen WT, et al. CLA improves the lipo-nutritional quality of pork and regulates the gut microbiota in Heigai pigs. Food Funct. 2022;13(23).(2022)org/10.1039/D2FO02549C.: 12093.
- 9.Liu YY, Peng YL, Chen C, Ren HB, Zhu J, Deng Y, et al. Flavonoids from mulberry leaves inhibit fat production and improve fatty acid distribution in adipose tissue in finishing pigs. Anim Nutr. 2024;16.(2024)11.003.: 147.
- 10.Guo ZY, Chen XL, Huang ZQ, Chen DW, Li MZ, Yu B, et al. Dihydromyricetin improves meat quality and promotes skeletal muscle fiber type transformations via AMPK signaling in growing-finishing pigs. Food Funct. 2022;13(6).(2022)org/10.1039/d1fo03391c.: 3649.
- 11.Sun HX, Chen D, Cai HY, Chang WH, Wang ZD, Liu GH, et al. Effects of fermenting the plant fraction of a complete feed on the growth performance, nutrient utilization, antioxidant functions, meat quality, and intestinal microbiota of broilers. Animals. 2022;12(20).(2022)org/10.3390/ani12202870.: 2870.
- 12.Wang C, Su WF, Zhang Y, Hao LH, Wang FQ, Lu ZQ, et al. Solid-state fermentation of distilled dried grain with solubles with probiotics for degrading lignocellulose and upgrading nutrient utilization. AMB Express. 2018;8(1).(2018)org/10.1186/s13568-018-0715-z.: 188.
- 13.Liu P, Zhao J, Guo P, Lu W, Geng Z, Levesque CL, et al. Dietary corn bran fermented byBacillus subtilisMA139 decreased gut cellulolytic bacteria and microbiota diversity in finishing pigs. Front Cell Infect Microbiol. 2017;7.(2017)526. https://doi. org/10.3389/fcimb.: 526.
- 14.Yang YZ, Yan GH, Meng XH, Wang X, Zhao ZQ, Zhou SG, et al. Effects ofLactobacillus plantarumandPediococcus acidilacticico-fermented feed on growth performance and gut microbiota of nursery pigs. Front Vet Sci. 2022;9.(2022)1076906. https://doi. org/10.3389/fvets.: 1076906.
- 15.Xu F, Wu HZ, Xie JJ, Zeng T, Hao LJ, Xu WW, et al. The effects of fermented feed on the growth performance, antioxidant activity, immune function, intestinal digestive enzyme activity, morphology, and microflora of yellow-feather chickens. Animals. 2023;13(22).(2023)org/10.3390/ani13223545.: 3545.
- 16.Zhu JJ, Gao MX, Zhang RL, Sun ZJ, Wang CM, Yang FF, et al. Effects of soybean meal fermented byL. plantarum,B. subtilisandS. cerevisieaeon growth, immune function and intestinal morphology in weaned piglets. Microb Cell Fact. 2017;16(1).(2017)org/10.1186/s12934-017-0809-3.: 191.
- 17.Liu SQ, Du M, Tu YA, You WJ, Chen WT, Liu GL, et al. Fermented mixed feed alters growth performance, carcass traits, meat quality and muscle fatty acid and amino acid profiles in finishing pigs. Anim Nutr. 2023;12.(2023)09.003.: 87.
- 18.Rahman M, Bora JR, Sarma AK, Roychoudhury R, Borgohain A. Effect of deep litter housing and fermented feed on carcass characteristics and meat quality of crossbred Hampshire pigs. Vet World. 2015;8(7).(2015)881–7. https://doi. org/10.14202/vetworld.: 881.
- 19.Tian ZM, Deng D, Cui YY, Chen WD, Yu M, Ma XY. Diet supplemented with fermented okara improved growth performance, meat quality, and amino acid profiles in growing pigs. Food Sci Nutr. 2020;8(10).(2020)1002/fsn3.1857.: 5650.
- 20.Zheng Z, Liu ZQ, Li N, Mu SQ, Liang SY, Liu ZH, et al. Effects of fermented bamboo powder on growth performance, apparent digestibility, carcass traits, and meat quality in growing-finishing pigs. Livest Sci. 2023;277.(2023)105358. https://doi. org/10. 1016/j.livsci.: 105358.
- 21.Qiu YQ, Li KB, Zhao XC, Liu SL, Wang L, Yang XF, et al. Fermented feed modulates meat quality and promotes the growth of longissimus thoracis of late-finishing pigs. Animals. 2020;10(9).(2020)org/10.3390/ani10091682.: 1682.
- 22.Chu GM, Park BK. Effects of fermented carrot by-product diets on growth performances, carcass characteristics and meat quality in fattening pigs. Acta Agr Scand, A Anim Sci. 2023;72(1–2).(2023)40–8. https://doi. org/10.1080/09064702.: 40.
- 23.Ding XQ, Li HY, Wen ZW, Hou Y, Wang GL, Fan JH, et al. Effects of fermented tea residue on fattening performance, meat quality, digestive performance, serum antioxidant capacity, and intestinal morphology in fatteners. Animals. 2020;10(2).(2020)org/10.3390/ani10020185.: 185.
- 24.Hao LH, Su WF, Zhang Y, Wang C, Xu BC, Jiang ZP, et al. Effects of supplementing with fermented mixed feed on the performance and meat quality in finishing pigs. Anim Feed Sci Technol. 2020.https.(2020)//doi. org/10. 1016/j.anifeedsci.
- 25.Lu JF, Han QC, Wang SY, Wang ZL, Li X, Hu JH, et al. Effect of fermented corn–soybean meal on carcass and meat quality of grower-finisher pigs. J Anim Physiol Anim Nutr. 2020;105(4).(2020)1111/jpn.13444.: 693.
- 26.Jiang HJ, Azad MAK, Zhu Q, Ni HJ, Kong XF. Fermented cassava residue meal improves meat quality by regulating muscle fiber and enhancing lipid metabolism in Huanjiang mini-pigs. Animals. 2025;15(2).(2025)org/10.3390/ani15020177.: 177.
- 27.Zhang Y, Wang C, Su WF, Jiang ZP, He H, Gong T, et al. Co-fermented yellow wine lees byBacillus subtilisandEnterococcus faeciumregulates growth performance and gut microbiota in finishing pigs. Front Microbiol. 2022;13.(2022)1003498. https://doi. org/10.3389/fmicb.: 1003498.
- 28.Ahmed ST, Mun HS, Islam MM, Ko SY, Yang CJ. Effects of dietary natural and fermented herb combination on growth performance, carcass traits and meat quality in grower-finisher pigs. Meat Sci. 2016;122.(2016)7–15. https://doi. org/10. 1016/j.meatsci.: 7.
- 29.Lei XJ, Yun HM, Kim IH. Effects of dietary supplementation of natural and fermented herbs on growth performance, nutrient digestibility, blood parameters, meat quality and fatty acid composition in growing-finishing pigs. Ital J Anim Sci. 2018;17(4).(2018)984–93. https://doi. org/10.1080/1828051x.: 984.
- 30.Xu CH, Xiong PW, Song WJ, Song QL, Hu Y, Song TX, et al. Effects of fermented navel orange pulp on growth performance, carcass characteristics, meat quality, meat nutritional value, and serum biochemical indicators of finishing Tibetan pigs. Foods. 2024;13(12).(2024)org/10.3390/foods13121910.: 1910.
- 31.Xu X, Li LM, Li B, Guo WJ, Ding XL, Xu FZ. Effect of fermented biogas residue on growth performance, serum biochemical parameters, and meat quality in pigs. Asian-Australas J Anim Sci. 2017;30(10).(2017)16.0777.: 1464.
- 32.Hati S, Patel M, Mishra BK, Das S. Short-chain fatty acid and vitamin production potentials ofLactobacillusisolated from fermented foods of Khasi Tribes, Meghalaya. India Ann Microbiol. 2019;69(11).(2019)org/10.1007/s13213-019-01500-8.: 1191.
- 33.Li SY, Zhao YJ, Zhang L, Zhang X, Huang L, Li D, et al. Antioxidant activity ofLactobacillus plantarumstrains isolated from traditional Chinese fermented foods. Food Chem. 2012;135(3).(2012)1914–9. https://doi. org/10. 1016/j.foodchem.: 1914.
- 34.Ding WR, Wang LN, Zhang JA, Ke WC, Zhou JW, Zhu JX, et al. Characterization of antioxidant properties of lactic acid bacteria isolated from spontaneously fermented yak milk in the Tibetan Plateau. J Funct Foods. 2017;35.(2017)481–8. https://doi. org/10. 1016/j.jff.: 481.
- 35.Garbacz K. Anticancer activity of lactic acid bacteria. Semin Cancer Biol. 2022;86(Pt 3).(2022)12.013.: 356.
- 36.Li FH, Ding ZT, Adesogan AT, Ke WC, Jiang Y, Bai J, et al. Effects of class IIa bacteriocin-producingLactobacillusspecies on fermentation quality and aerobic stability of alfalfa silage. Animals. 2020;10(9).(2020)org/10.3390/ani10091575.: 1575.
- 37.Londero A, León Peláez MA, Diosma G, De Antoni GL, Abraham AG, Garrote GL. Fermented whey as poultry feed additive to prevent fungal contamination. J Sci Food Agric. 2014;94(15).(2014)1002/jsfa.6669.: 3189.
- 38.Qian JY, Wang YZ, Hu ZJ, Shi TQ, Wang YT, Ye C, et al.Bacillussp. as a microbial cell factory.(2023)advancements and future prospects.Biotechnol Adv.: 108278.
- 39.Xu RP, Tian T, Hu B, Zhang ZQ, Liu J, Yu DH, et al. Effect of solid-state fermentation withBacillus pumiluson the nutritional value, anti-nutritional factors and antioxidant activity of faba bean (Vicia fabaL.) meal. Lwt. 2023;185.(2023)115117. https://doi. org/10. 1016/j.lwt.: 115117.
- 40.Zhang MM, Huang YW, Zhao HC, Wang TF, Xie CQ, Zhang DY, et al. Solid-state fermentation ofMoringa oleiferaleaf meal usingBacillus pumilusCICC 10440. J Chem Technol Biotechnol. 2017;92(8).(2017)1002/jctb.5203.: 2083.
- 41.Liu SQ, Cai PR, You WJ, Yang MS, Tu YA, Zhou YB, et al. Enhancement of gut barrier integrity by a Bacillus subtilis secreted metabolite through the GADD45A‐Wnt/β‐catenin pathway. Imeta. 2025;4(2).(2025)1002/imt2.70005.
- 42.Chuang WY, Lin WC, Hsieh YC, Huang CM, Chang SC, Lee TT. Evaluation of the combined use ofSaccharomyces cerevisiaeandAspergillus oryzaewith phytase fermentation products on growth, inflammatory, and intestinal morphology in broilers. Animals. 2019;9(12).(2019)org/10.3390/ani9121051.: 1051.
- 43.Rai AK, Pandey A, Sahoo D. Biotechnological potential of yeasts in functional food industry. Trends Food Sci Tech. 2019;83.(2019)11.016.: 129.
- 44.Rai AK, Kumari R, Sanjukta S, Sahoo D. Production of bioactive protein hydrolysate using the yeasts isolated from soft chhurpi. Bioresour Technol. 2016;219.(2016)239–45. https://doi. org/10. 1016/j.biortech.: 239.
- 45.Aktar J, Islam KMS, Chowdhury R, Debi MR, Emon AI. Effect of baker’s yeast fermented moist feed on the growth and bone mineralization in broiler. J Adv Vet Anim Res. 2024;11(1).(2024)78–84. https://doi. org/10.5455/javar.: 78.
- 46.Azrinnahar M, Islam N, Shuvo AAS, Kabir AKMA, Islam KMS. Effect of feeding fermented (Saccharomyces cerevisiae) de-oiled rice bran in broiler growth and bone mineralization. J Saudi Soc Agric Sci. 2021;20(7).(2021)476–81. https://doi. org/10. 1016/j.jssas.: 476.
- 47.Hammod A. Effect of corn fermented bySaccharomyces cerevisiaein diets on production performance of broiler chicks. J Pure Appl Microbiol. 2019;13(2).(2019)2.40.: 1025.
- 48.Nocek JE, Holt MG, Oppy J. Effects of supplementation with yeast culture and enzymatically hydrolyzed yeast on performance of early lactation dairy cattle. J Dairy Sci. 2011;94(8).(2011)4046–56. https://doi. org/10.3168/jds.: 4046.
- 49.Vailati-Riboni M, Coleman DN, Lopreiato V, Alharthi A, Bucktrout RE, Abdel-Hamied E, et al. Feeding aSaccharomyces cerevisiaefermentation product improves udder health and immune response to aStreptococcus uberismastitis challenge in mid-lactation dairy cows. J Anim Sci Biotechnol. 2021;12.(2021)org/10.1186/s40104-021-00560-8.: 62.
- 50.Lim DH, Han MH, Ki KS, Kim TI, Park SM, Kim DH, et al. Changes in milk production and blood metabolism of lactating dairy cows fedSaccharomyces cerevisiaeculture fluid under heat stress. J Anim Sci Technol. 2021;63(6).(2021)1433–42. https://doi. org/10.5187/jast.: 1433.
- 51.Espitia-Hernandez P, Ruelas-Chacon X, Chavez-Gonzalez ML, Ascacio-Valdes JA, Flores-Naveda A, Sepulveda-Torre L. Solid-state fermentation of sorghum byAspergillus oryzaeandAspergillus niger.(2022)effects on tannin content, phenolic profile, and antioxidant activity.Foods.: 3121.
- 52.Colla LM, Ficanha AM, Rizzardi J, Bertolin TE, Reinehr CO, Costa JAV. Production and characterization of lipases by two new isolates ofAspergillusthrough solid-state and submerged fermentation. BioMed Res Int. 2015;2015.(2015)725959. https://doi. org/10.1155/.: 725959.
- 53.Falony G, Armas JC, Mendoza JCD, Hernandez JLM. Production of extracellular lipase from Aspergillus niger by solid-state fermentation. Food Technol Biotech. 2006;44(4).(2006)235–40.: 235.
- 54.Lima LGR, Gonçalves MMM, Couri S, Melo VF, Sant’Ana GCF, Costa ACAD. Lipase production byAspergillus nigerC by submerged fermentation. Braz Arch Biol Technol. 2019.https.(2019)//doi. org/10.1590/1678-4324-.
- 55.Yang LJ, Zeng XF, Qiao SY. Advances in research on solid-state fermented feed and its utilization.(2021)The pioneer of private customization for intestinal microorganisms.Anim Nutr.: 905.
- 56.Couto SR, Sanromán MÁ. Application of solid-state fermentation to food industry—a review. J Food Eng. 2006;76(3).(2006)291–302.: 291.
- 57.Sugiharto S, Ranjitkar S. Recent advances in fermented feeds towards improved broiler chicken performance, gastrointestinal tract microecology and immune responses.(2019)a review.Anim Nutr.: 1.
- 58.Ranjan A, Sahu NP, Deo AD, Kumar S. Solid state fermentation of de-oiled rice bran.(2019)effect on in vitro protein digestibility, fatty acid profile and anti-nutritional factors.Food Res Int.: 1.
- 59.Su WF, Jiang ZP, Wang C, Zhang Y, Gong T, Wang FQ, et al. Co-fermented defatted rice bran alters gut microbiota and improves growth performance, antioxidant capacity, immune status and intestinal permeability of finishing pigs. Anim Nutr. 2022;11.(2022)413–24. https://doi. org/10. 1016/j.aninu.: 413.
- 60.de Castro RJS, Ohara A, Nishide TG, Bagagli MP, Gonçalves Dias FF, Sato HH. A versatile system based on substrate formulation using agroindustrial wastes for protease production byAspergillus nigerunder solid state fermentation. Biocatal Agric Biotechnol. 2015;4(4).(2015)678–84. https://doi. org/10. 1016/j.bcab.: 678.
- 61.Cerda A, El-Bakry M, Gea T, Sánchez A. Long term enhanced solid-state fermentation.(2016)inoculation strategies for amylase production from soy and bread wastes byThermomycessp. in a sequential batch operation.J Environ Chem Eng.: 2394.
- 62.Qureshi AS, Khushk I, Ali CH, Chisti Y, Ahmad A, Majeed H. Coproduction of protease and amylase by thermophilicBacillussp. BBXS-2 using open solid-state fermentation of lignocellulosic biomass. Biocatal Agric Biotechnol. 2016;8.(2016)146–51. https://doi. org/10. 1016/j.bcab.: 146.
- 63.Ferrarezi AL, Hideyuki Kobe Ohe T, Borges JP, Brito RR, Siqueira MR, Vendramini PH, et al. Production and characterization of lipases and immobilization of whole cell of the thermophilicThermomucor indicae seudaticaeN31 for transesterification reaction. J Mol Catal B-Enzym. 2014;107.(2014)106–13. https://doi. org/10. 1016/j.molcatb.: 106.
- 64.Pandey AK, Edgard G, Negi S. Optimization of concomitant production of cellulase and xylanase fromRhizopus oryzaeSN5 through EVOP-factorial design technique and application in Sorghum Stover based bioethanol production. Renew Energy. 2016;98.(2016)51–6. https://doi. org/10. 1016/j.renene.: 51.
- 65.Yang LJ, Yang ZB, Yang WR, Li HR, Zhang CY, Jiang SZ, et al. Conventional solid fermentation alters mycotoxin contents and microbial diversity analyzed by high-throughput sequencing of aFusariummycotoxin-contaminated diet. Can J Anim Sci. 2018;98(2).(2018)org/10.1139/cjas-2017-0093.: 354.
- 66.Cullen JT, Lawlor PG, Cormican P, Gardiner GE. Microbial quality of liquid feed for pigs and its impact on the porcine gut microbiome. Animals. 2021;11(10).(2021)org/10.3390/ani11102983.: 2983.
- 67.Missotten JAM, Michiels J, Ovyn A, De Smet S, Dierick NA. Fermented liquid feed for pigs. Arch Anim Nutr. 2010;64(6).(2010)437–66. https://doi. org/10.1080/1745039x.: 437.
- 68.Lawlor PG, Lynach PB, Gardiner GE, Caffrey PJ, O’Doherty JV. Effect of liquid feeding weaned pigs on growth performance to harvest. J Anim Sci. 2002;80(7).(2002)1725–35. https://doi. org/10.2527/.: 1725.
- 69.Missotten JA, Michiels J, Degroote J, De Smet S. Fermented liquid feed for pigs.(2015)an ancient technique for the future.J Anim Sci Biotechnol.: 4.
- 70.Plumed-Ferrer C, von Wright A. Antimicrobial activity of weak acids in liquid feed fermentations, and its effects on yeasts and lactic acid bacteria. J Sci Food Agric. 2011;91(6).(2011)1002/jsfa.4278.: 1032.
- 71.Canibe N, Jensen BB. Fermented and nonfermented liquid feed to growing pigs.(2003)effect on aspects of gastrointestinal ecology and growth performance.J Anim Sci.: 2019.
- 72.O’ Meara FM, Gardiner GE, O’ Doherty JV, Clarke D, Cummins W, Lawlor PG. Effect of wet/dry, fresh liquid, fermented whole diet liquid, and fermented cereal liquid feeding on feed microbial quality and growth in grow-finisher pigs. J Anim Sci. 2020;98(6).(2020)org/10.1093/jas/skaa166.: 1.
- 73.Canibe N, Hojberg O, Badsberg JH, Jensen BB. Effect of feeding fermented liquid feed and fermented grain on gastrointestinal ecology and growth performance in piglets. J Anim Sci. 2007;85(11).(2007)2527/jas.2006-744.: 2959.
- 74.Moran CA, Scholten RH, Tricarico JM, Brooks PH, Verstegen MW. Fermentation of wheat.(2006)effects of backslopping different proportions of pre-fermented wheat on the microbial and chemical composition.Arch Anim Nutr.: 158.
- 75.Scholten RHJ, van der Peet-Schwering CMC, den Hartog LA, Balk M, Schrama JW, Verstegen M W. Fermented wheat in liquid diets.(2002)Effects on gastrointestinal characteristics in weanling piglets.J Anim Sci.: 1179.
- 76.Zhang AR, Yang YY, Li Y, Zheng YF, Wang HM, Cui HX, et al. Effects of wheat-based fermented liquid feed on growth performance, nutrient digestibility, gut microbiota, intestinal morphology, and barrier function in grower-finisher pigs. J Anim Sci. 2024.https.(2024)org/10.1093/jas/skae229.
- 77.Plumed-Ferrer C, Llopis M, Hyvönen P, Wright AV. Characterization of the microbial community and its changes in liquid piglet feed formulations. J Sci Food Agric. 2004;84(11).(2004)1002/jsfa.1818.: 1315.
- 78.Engberg RM, Hammershoj M, Johansen NF, Abousekken MS, Steenfeldt S, Jensen BB. Fermented feed for laying hens.(2009)effects on egg production, egg quality, plumage condition and composition and activity of the intestinal microflora.Br Poult Sci.: 228.
- 79.Missotten JA, Michiels J, Dierick N, Ovyn A, Akbarian A, De Smet S. Effect of fermented moist feed on performance, gut bacteria and gut histo-morphology in broilers. Br Poult Sci. 2013;54(5).(2013)627–34. https://doi. org/10.1080/00071668.: 627.
- 80.Xin HL, Wang MY, Xia Z, Yu B, He J, Yu J, et al. Fermented diet liquid feeding improves growth performance and intestinal function of pigs. Animals. 2021;11(5).(2021)org/10.3390/ani11051452.: 1452.
- 81.Shuai CY, Chen DW, Yu B, Luo YH, Zheng P, Huang ZQ, et al. Effect of fermented rapeseed meal on growth performance, nutrient digestibility, and intestinal health in growing pigs. Anim Nutr. 2023;15.(2023)420–9. https://doi. org/10. 1016/j.aninu.: 420.
- 82.Tie Y, Li L, Liu J, Liu CL, Fu JJ, Xiao XJ, et al. Two-step biological approach for treatment of rapeseed meal. J Food Sci. 2020;85(2).(2020)1111/1750-3841.15011.: 340.
- 83.Zhu Xf, Wang LY, Zhang Z, Ding LR, Hang SQ. Combination of fiber-degrading enzymatic hydrolysis and lactobacilli fermentation enhances utilization of fiber and protein in rapeseed meal as revealed in simulated pig digestion and fermentation in vitro. Anim Feed Sci Tech. 2021;278.https.(2021)//doi. org/10. 1016/j.anifeedsci.
- 84.Hu YQ, He YY, Gao S, Liao ZQ, Lai T, Zhou HM, et al. The effect of a diet based on rice straw co-fermented with probiotics and enzymes versus a fresh corn stover-based diet on the rumen bacterial community and metabolites of beef cattle. Sci Rep. 2020;10.(2020)org/10.1038/s41598-020-67716-w.: 10721.
- 85.Lin BS, Yan JB, Zhong ZL, Zheng XT. A study on the preparation of microbial and nonstarch polysaccharide enzyme synergistic fermented maize cob feed and its feeding efficiency in finishing pigs. BioMed Res Int. 2020;2020.(2020)8839148. https://doi. org/10.1155/.: 8839148.
- 86.Sebothoma P, Hlatini VA, Ncobela CN, Thomas RS, Chimonyo M. Effect of liquid-fermented potato hash, with or without exogenous enzymes, on carcass characteristics of growing pigs. S Afr J Anim Sci. 2024;53(5).(2024)v53i5.07.: 678.
- 87.Liu Y, Liu Y, Cao YH, Wang CL. Pretreatment of palm kernel cake by enzyme-bacteria and its effects on growth performance in broilers. Animals. 2025;15(2).(2025)org/10.3390/ani15020116.: 116.
- 88.Ji MT, Rong XY, Wu YF, Li HN, Zhao XL, Zhao Y, et al. Effects of fermented liquid feed with compound probiotics on growth performance, meat quality, and fecal microbiota of growing pigs. Animals. 2025;15(5).(2025)org/10.3390/ani15050733.: 733.
- 89.Xie KH, Dai YQ, Zhang AR, Yu B, Luo YH, Li H, et al. Effects of fermented soybean meal on growth performance, meat quality, and antioxidant capacity in finishing pigs. J Funct Foods. 2022;94.(2022)105128. https://doi. org/10. 1016/j.jff.: 105128.
- 90.Lu JF, Zhang XY, Liu YH, Cao HG, Han QC, Xie BC, et al. Effect of fermented corn-soybean meal on serum immunity, the expression of genes related to gut immunity, gut microbiota, and bacterial metabolites in grower-finisher pigs. Front Microbiol. 2019.https.(2019)//doi. org/10.3389/fmicb.
- 91.Shi CY, He J, Yu J, Yu B, Mao XB, Zheng P, et al. Physicochemical properties analysis and secretome ofAspergillus nigerin fermented rapeseed meal. PLoS ONE. 2016;11(4).(2016)pone.0153230.
- 92.Sun H, Qian ZC, Wu YF, Tang JW, Shen Q, Li JH, et al. Effects of fermented broccoli stem and leaf residue on growth performance, serum characteristics and meat quality of growing pigs. J Anim Physiol Anim Nutr (Berl). 2023;107(4).(2023)1111/jpn.13804.: 1035.
- 93.Su WF, Jiang ZP, Wang C, Xu BC, Lu ZQ, Wang FQ, et al. Dynamics of defatted rice bran in physicochemical characteristics, microbiota and metabolic functions during two-stage co-fermentation. Int J Food Microbiol. 2022;362.(2022)2021.109489.: 109489.
- 94.Wang C, Shi CY, Su WF, Jin ML, Xu Bc, Hao LH, et al. Dynamics of the physicochemical characteristics, microbiota, and metabolic functions of soybean meal and corn mixed substrates during two-stage solid-state fermentation. mSystems. 2020.https.(2020)1128/mSystems.00501-19.
- 95.Zheng L, Li D, Li ZL, Kang LN, Jiang YY, Liu XY, et al. Effects ofBacillusfermentation on the protein microstructure and anti-nutritional factors of soybean meal. Lett Appl Microbiol. 2017;65(6).(2017)1111/lam.12806.: 520.
- 96.Olukomaiya OO, Fernando WC, Mereddy R, Li X, Sultanbawa Y. Solid-state fermentation of canola meal withAspergillus sojae,Aspergillus ficuumand their co-cultures.(2020)effects on physicochemical, microbiological and functional properties.LWT.
- 97.Shi CY, He J, Yu J, Yu B, Huang ZQ, Mao XB, et al. Solid state fermentation of rapeseed cake withAspergillus nigerfor degrading glucosinolates and upgrading nutritional value. J Anim Sci Biotechnol. 2015;6.(2015)org/10.1186/s40104-015-0015-2.: 13.
- 98.Mukherjee R, Chakraborty R, Dutta A. Role of fermentation in improving nutritional quality of soybean meal - A Review. Asian Austral J Anim. 2016;29(11).(2016)15.0627.: 1523.
- 99.Mok WK, Tan YX, Lee J, Kim J, Chen WN. A metabolomic approach to understand the solid-state fermentation of okara usingBacillus subtilisWX-17 for enhanced nutritional profile. AMB Express. 2019;9(1).(2019)org/10.1186/s13568-019-0786-5.: 60.
- 100.Feng HY, Qu H, Liu Y, Shi YH, Wu SL, Bao WB. Effect of fermented soybean meal supplementation on some growth performance, blood chemical parameters, and fecal microflora of finishing pigs. Rev Bras Zootec. 2020.https.(2020)org/10.37496/rbz4920190096.
- 101.Song J, Wang X, Cao Y, He Y, Yang Y. Effects of corn-soybean meal-based fermented feed supplementation on growth performance, meat quality, fatty acid profiles, nutritional values, and gut microbiota of lean-type finishing pigs. Foods. 2025;14(15).(2025)org/10.3390/foods14152641.: 2641.
- 102.Han G, Liu S, Zhao C, Lei L, Yi R, Ma Z, et al. Effects of fermented liquid feed on growth performance, meat quality, and intestinal microbiota of Yuedong Black Pigs. Animals. 2025;15(18).(2025)org/10.3390/ani15182657.: 2657.
- 103.Zhang A, Zheng Y, Yin W, Pu X, Yu A, Wang H, et al. Impact of fermented liquid feed supplementation on grower-fattening pigs production.(2025)insights into growth performance, carcass traits, meat quality, and metabolite profiles.Food Chem Mol Sci.: 100268.
- 104.Ma W, Ma Z, Mao P, Zhang X, Wu X, Gao M, et al. Effects of feed supplemented with fermented pine needles (Pinus ponderosa) on carcass quality, meat quality, and antioxidant capacity of growing-finishing pigs. Foods. 2025;14(12).(2025)org/10.3390/foods14122046.: 2046.
- 105.Li XY, Zhang L, Zhang YM, Luo X, Yu J, Ren SF, et al. Effects of dietaryInonotus obliquusfermentation products supplementation on meat quality and antioxidant capacity of finishing pigs. Meat Sci. 2025;224.(2025)109789. https://doi. org/10. 1016/j.meatsci.: 109789.
- 106.Zhao ZW, Wu J, Yao XH, Sun H, Wu YF, Zhou HH, et al. Influence of fermented broccoli residues on fattening performance, nutrient utilization, and meat properties of finishing pigs. Animals. 2024.https.(2024)org/10.3390/ani14131987.
- 107.Niu JK, Liu X, Xu JY, Li F, Wang JC, Zhang XX, et al. Effects of silage diet on meat quality through shaping gut microbiota in finishing pigs. Microbiol Spectr. 2023;11(1).(2023)1128/spectrum.02416-22.
- 108.Chaiwang N, Bunmee T, Arjin C, Wattanakul W, Krutthai N, Mekchay S, et al. Effect of deep bedding floor and fermented feed supplement on productive performance, carcase, meat quality and fatty acid profile of crossbred pigs. Ital J Anim Sci. 2021;20(1).(2021)479–88. https://doi. org/10.1080/1828051x.: 479.
- 109.Fang JC, Cao Y, Matsuzaki M, Suzuki H, Kimura H. Effects of apple pomace-mixed silage on growth performance and meat quality in finishing pigs. Anim Sci J. 2016;87(12).(2016)1111/asj.12601.: 1516.
- 110.Chu GM, Kang SN, Nam JM, Kim HY, Ha JH, Ibrahim RIH, et al. Effects of dietary fermented persimmon diet on the meat quality of fattening pigs. Korean J Food Sci Anim Resour. 2012;32(5).(2012)604–11. https://doi. org/10.5851/kosfa.: 604.
- 111.Chu GM, Kang SN, Yang JM, Kim HY, Song YM. Effects of a dietary fermented mushroom (Flammulina velutipes) by-product diet on pork meat quality in growing-fattening Berkshire pigs. J Anim Sci Technol. 2012;54(3).(2012)199–207. https://doi. org/10.5187/jast.: 199.
- 112.Yan L, Kim IH. Effect of dietary grape pomace fermented bySaccharomyces boulardIIon the growth performance, nutrient digestibility and meat quality in finishing pigs. Asian Austral J Anim. 2011;24(12).(2011)1763–70. https://doi. org/10.5713/ajas.: 1763.
- 113.Yan L, Meng QW, Kim IH. Effects of fermented garlic powder supplementation on growth performance, nutrient digestibility, blood characteristics and meat quality in growing-finishing pigs. Anim Sci J. 2011;83(5).(2011)411–7. https://doi. org/10. 1111/j.1740-0929.: 411.
- 114.Ao X, Meng QW, Kim IH. Effects of fermented red ginseng supplementation on growth performance, apparent nutrient digestibility, blood hematology and meat quality in finishing pigs. Asian-Australas J Anim Sci. 2011;24(4).(2011)525–31. https://doi. org/10.5713/ajas.: 525.
- 115.Lee SD, Kim HY, Jung HJ, Ji SY, Chowdappa R, Ha JH, et al. The effect of fermented apple diet supplementation on the growth performance and meat quality in finishing pigs. Anim Sci J. 2009;80(1).(2009)00598.x.: 79.
- 116.Sasaki K, Nishioka T, Ishizuka Y, Saeki M, Kawashima T, Irie M, et al. Comparison of sensory traits and preferences between food co-product fermented liquid (FCFL)-fed and formula-fed pork loin. Asian-Australas J Anim Sci. 2007;20(8).(2007)1272–7. https://doi. org/10.5713/ajas.: 1272.
- 117.Song YM, Lee SD, Chowdappa R, Kim HY, Jin SK, Kim IS. Effects of fermented oyster mushroom (Pleurotus ostreats) by-product supplementation on growth performance, blood parameters and meat quality in finishing Berkshire pigs. Animal. 2007;1(2).(2007)org/10.1017/S1751731107683785.: 301.
- 118.Tang XP, Liu XG, Zhang K. Effects of microbial fermented feed on serum biochemical profile, carcass traits, meat amino acid and fatty acid profile, and gut microbiome composition of finishing pigs. Front Vet Sci. 2021;8.(2021)744630. https://doi. org/10.3389/fvets.: 744630.
- 119.Mancini RA, Hunt MC. Current research in meat color. Meat Sci. 2005;71(1).(2005)100–21. https://doi. org/10. 1016/j.meatsci.: 100.
- 120.Viljoen HF, de Kock HL, Webb EC. Consumer acceptability of dark, firm and dry (DFD) and normal pH beef steaks. Meat Sci. 2002;61(2).(2002)org/10.1016/s0309-1740(01)00183-8.: 181.
- 121.Zdunczyk W, Tkacz K, Modzelewska-Kapitula M. The effect of superficial oregano essential oil application on the quality of modified atmosphere-packed pork loin. Foods. 2023;12(10).(2023)org/10.3390/foods12102013.: 2013.
- 122.Lindahl G, Lundstrom K, Tornberg E. Contribution of pigment content, myoglobin forms and internal reflectance to the colour of pork loin and ham from pure breed pigs. Meat Sci. 2001;59(2).(2001)org/10.1016/s0309-1740(01)00064-x.: 141.
- 123.Hughes J, Clarke F, Li Y, Purslow P, Warner R. Differences in light scattering between pale and dark beef longissimus thoracis muscles are primarily caused by differences in the myofilament lattice, myofibril and muscle fibre transverse spacings. Meat Sci. 2019;149.(2019)11.006.: 96.
- 124.Hughes J, Clarke F, Purslow P, Warner R. High pH in beef longissimus thoracis reduces muscle fibre transverse shrinkage and light scattering which contributes to the dark colour. Food Res Int. 2017;101.(2017)228–38. https://doi. org/10. 1016/j.foodres.: 228.
- 125.Moeller SJ, Miller RK, Edwards KK, Zerby HN, Logan KE, Aldredge TL, et al. Consumer perceptions of pork eating quality as affected by pork quality attributes and end-point cooked temperature. Meat Sci. 2010;84(1).(2010)06.023.: 14.
- 126.Liu SQ, Tu YA, Sun JB, Cai PR, Zhou YB, Huang YQ, et al. Fermented mixed feed regulates intestinal microbial community and metabolism and alters pork flavor and umami. Meat Sci. 2023.https.(2023)//doi. org/10. 1016/j.meatsci.
- 127.Frank D, Joo ST, Warner R. Consumer acceptability of intramuscular fat. Korean J Food Sci Anim Resour. 2016;36(6).(2016)699–708. https://doi. org/10.5851/kosfa.: 699.
- 128.Hocquette JF, Gondret F, Baeza E, Medale F, Jurie C, Pethick DW. Intramuscular fat content in meat-producing animals.(2010)development, genetic and nutritional control, and identification of putative markers.Animal.: 303.
- 129.Fortin A, Robertson WM, Tong AKW. The eating quality of Canadian pork and its relationship with intramuscular fat. Meat Sci. 2005;69(2).(2005)07.011.: 297.
- 130.Fernandez X, Monin G, Talmant A, Mourot J, Lebret B. Influence of intramuscular fat content on the quality of pig meat - 2. Consumer acceptability ofm. longissimus lumborum. Meat Sci. 1999;53(1).(1999)org/10.1016/s0309-1740(99)00038-8.: 67.
- 131.Font-i-Furnols M, Tous N, Esteve-Garcia E, Gispert M. Do all the consumers accept marbling in the same way? The relationship between eating and visual acceptability of pork with different intramuscular fat content. Meat Sci. 2012;91(4).(2012)448–53. https://doi. org/10. 1016/j.meatsci.: 448.
- 132.Hou JJ, Ji X, Chu XR, Shi ZY, Wang BJ, Sun KL, et al. Comprehensive lipidomic analysis revealed the effects of fermentedMorus albaL. intake on lipid profile in backfat and muscle tissue of Yuxi black pigs. J Anim Physiol Anim Nutr (Berl). 2024;108(3).(2024)1111/jpn.13932.: 764.
- 133.Lenighan YM, McNulty BA, Roche HM. Dietary fat composition.(2019)replacement of saturated fatty acids with PUFA as a public health strategy, with an emphasis on α-linolenic acid.Proc Nutr Soc.: 234.
- 134.Tortosa-Caparrós E, Navas-Carrillo D, Marín F, Orenes-Piñero E. Anti-inflammatory effects of omega 3 and omega 6 polyunsaturated fatty acids in cardiovascular disease and metabolic syndrome. Crit Rev Food Sci Nutr. 2017;57(16).(2017)2015.1126549.: 3421.
- 135.Wood JD, Richardson RI, Nute GR, Fisher AV, Campo MM, Kasapidou E, et al. Effects of fatty acids on meat quality.(2004)a review.Meat Sci.: 21.
- 136.Yi WZ, Huang QX, Wang YZ, Shan TZ. Lipo-nutritional quality of pork.(2023)the lipid composition, regulation, and molecular mechanisms of fatty acid deposition.Anim Nutr.: 373.
- 137.Frayn KN, Arner P, Yki-Jarvinen H. Fatty acid metabolism in adipose tissue, muscle and liver in health and disease. Essays Biochem. 2006;42.(2006)org/10.1042/bse0420089.: 89.
- 138.Zhang ZQ, Zang MW, Zhang KH, Wang SW, Li D, Li XM. Effects of phospholipids and reheating treatment on volatile compounds in phospholipid-xylose-cysteine reaction systems. Food Res Int. 2021;139.(2021)2020.109918.: 109918.
- 139.Liu SQ, Du M, Sun JB, Tu YA, Gu X, Cai PR, et al.Bacillus subtilisandEnterococcus faeciumco-fermented feed alters antioxidant capacity, muscle fibre characteristics and lipid profiles of finishing pigs. Br J Nutr. 2024;131(8).(2024)org/10.1017/S000711452300291X.: 1298.
- 140.Shi M, Yang YN, Hu XS, Zhang ZY. Effect of ultrasonic extraction conditions on antioxidative and immunomodulatory activities of aGanoderma lucidumpolysaccharide originated from fermented soybean curd residue. Food Chem. 2014;155.(2014)50–6. https://doi. org/10. 1016/j.foodchem.: 50.
- 141.Akbari M, Razavi SH, Khodaiyan F, Blesa J, Esteve MJ. Fermented corn bran.(2023)a by-product with improved total phenolic content and antioxidant activity.LWT.
- 142.Sawangwan T, Porncharoennop C, Nimraksa H. Antioxidant compounds from rice bran fermentation by lactic acid bacteria. AIMS Agric Food. 2021;6(2).(2021)578–87. https://doi. org/10.3934/agrfood.: 578.
- 143.Meng QW, Yan L, Ao X, Zhou TX, Wang JP, Lee JH, et al. Influence of probiotics in different energy and nutrient density diets on growth performance, nutrient digestibility, meat quality, and blood characteristics in growing-finishing pigs. J Anim Sci. 2010;88(10).(2010)2527/jas.2009-2308.: 3320.
- 144.Xie CL, Teng JY, Wang XK, Xu BY, Niu YR, Ma LB, et al. Multi-omics analysis reveals gut microbiota-induced intramuscular fat deposition via regulating expression of lipogenesis-associated genes. Anim Nutr. 2022;9.(2022)10.010.: 84.
- 145.Xie CL, Zhu XY, Xu BY, Niu YR, Zhang XL, Ma LB, et al. Integrated analysis of multi-tissues lipidome and gut microbiome reveals microbiota-induced shifts on lipid metabolism in pigs. Anim Nutr. 2022;10.(2022)280–93. https://doi. org/10. 1016/j.aninu.: 280.
- 146.Fang SM, Xiong XW, Su Y, Huang LS, Chen CY. 16S rRNA gene-based association study identified microbial taxa associated with pork intramuscular fat content in feces and cecum lumen. BMC Microbiol. 2017;17.(2017)org/10.1186/s12866-017-1055-x.: 162.
- 147.You WJ, Liu SQ, Ji JF, Ling DF, Tu YA, Zhou YB, et al. Growth arrest and DNA damage-inducible alpha regulates muscle repair and fat infiltration through ATP synthase F1 subunit alpha. J Cachexia Sarcopenia Muscle. 2023;14(1).(2023)1002/jcsm.13134.: 326.
- 148.You WJ, Xu ZY, Sun Y, Valencak TG, Wang YZ, Shan TZ. GADD45α drives brown adipose tissue formation through upregulating PPARgamma in mice. Cell Death Dis. 2020;11(7).(2020)org/10.1038/s41419-020-02802-5.: 585.
- 149.Duan YH, Li FN, Wang WL, Guo QP, Wen CY, Yin YL. Alteration of muscle fiber characteristics and the AMPK-SIRT1-PGC-1α axis in skeletal muscle of growing pigs fed low-protein diets with varying branched-chain amino acid ratios. Oncotarget. 2017;8(63).(2017)18632/oncotarget.22205.: 107011.
- 150.Choi YM, Ryu YC, Kim BC. Influence of myosin heavy- and light chain isoforms on early postmortem glycolytic rate and pork quality. Meat Sci. 2007;76(2).(2007)11.009.: 281.
- 151.Li BJ, Yin D, Li PH, Zhang ZK, Zhang XY, Li HQ, et al. Profiling and functional analysis of circular RNAs in porcine fast and slow muscles. Front Cell Dev Biol. 2020;8.(2020)322. https://doi. org/10.3389/fcell.: 322.
- 152.Hwang YH, Kim GD, Jeong JY, Hur SJ, Joo ST. The relationship between muscle fiber characteristics and meat quality traits of highly marbled Hanwoo (Korean native cattle) steers. Meat Sci. 2010;86(2).(2010)456–61. https://doi. org/10. 1016/j.meatsci.: 456.
- 153.Luo YL, Ju N, Chang J, Ge RX, Zhao YY, Zhang GJ. Dietary alpha-lipoic acid supplementation improves postmortem color stability of the lamb muscles through changing muscle fiber types and antioxidative status. Meat Sci. 2022;193.(2022)108945. https://doi. org/10. 1016/j.meatsci.: 108945.
- 154.Mizunoya W, Iwamoto Y, Shirouchi B, Sato M, Komiya Y, Razin FR, et al. Dietary fat influences the expression of contractile and metabolic genes in rat skeletal muscle. PLoS ONE. 2013;8(11).(2013)pone.0080152.
- 155.Nakazato K, Song H. Increased oxidative properties of gastrocnemius in rats fed on a high-protein diet. J Nutr Biochem. 2008;19(1).(2008)12.019.: 26.
- 156.Zhang WG, Xiao S, Ahn DU. Protein oxidation.(2013)basic principles and implications for meat quality.Crit Rev Food Sci Nutr.: 1191.
- 157.Chen J, Tian M, Guan W, Wen T, Yang F, Chen F, et al. Increasing selenium supplementation to a moderately-reduced energy and protein diet improves antioxidant status and meat quality without affecting growth performance in finishing pigs. J Trace Elem Med Biol. 2019;56.(2019)38–45. https://doi. org/10. 1016/j.jtemb.: 38.
- 158.Simonetti A, Perna A, Gambacorta E. Comparison of antioxidant compounds in pig meat from Italian autochthonous pig Suino Nero Lucano and a modern crossbred pig before and after cooking. Food Chem. 2019;292.(2019)108–12. https://doi. org/10. 1016/j.foodchem.: 108.
Acknowledgements
Not applicable.
Funding
We thank members of the Shan Laboratory for comments and this work was partially supported by the National Natural Science Foundation of China (32272887), the Natural Science Foundation of Zhejiang Province (LZ22C170003) to TZS.
Ethics Declaration
Ethics approval and consent to participate
Not applicable.
Consent for publication
Not applicable.
Competing interests
The authors declare no competing interests.
Rights and Permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/ . The Creative Commons Public Domain Dedication waiver ( http://creativecommons.org/publicdomain/zero/1.0/ ) applies to the data made available in this article, unless otherwise stated in a credit line to the data. Reprints and permissions