FREE SHIPPING over $40
OVER 10000 FIVE-STAR REVIEWS
5% OFF ALL SUBSCRIPTIONS
100% HAPPINESS GUARANTEE
Analyze Diet
0
International journal of molecular sciences2026; 27(18); 8015; doi: 10.3390/ijms27188015

Gut Microbiota as a Molecular Regulator of Mineral Homeostasis in Horses: Mechanisms and Future Perspectives.

Abstract: Mineral homeostasis in horses is traditionally attributed to the coordinated regulation of the intestine, kidneys, skeleton, and endocrine system. Emerging evidence suggests the gut microbiota also shapes this process by modifying the intestinal environment where mineral absorption occurs. Despite the growing interest in host-microbiota interactions, the mechanisms linking the equine gut microbiota to mineral metabolism remain poorly understood and have not been comprehensively synthesized. This review integrates current knowledge of equine gastrointestinal physiology, gut microbial ecology, and mineral metabolism to examine how the gut microbiota influences mineral homeostasis, focusing on microbial fermentation, bioactive metabolite production, luminal pH regulation, epithelial barrier integrity, modulation of intestinal mineral transporters, and interactions with endocrine and immune pathways. The assessment of macro minerals such as calcium, phosphorus, and magnesium, along with trace minerals like iron, zinc, copper, selenium, and manganese, was conducted with a critical approach. This evaluation differentiates between results derived from equine research and those inferred from studies on other species. However, direct mechanistic evidence in horses remains limited, highlighting the need for future research. Advancing equine microbiome research through integrated microbiome profiling, metabolomics, molecular biology, and mineral balance studies will improve our understanding of host-microbiota interactions and support microbiome-informed nutritional strategies to enhance mineral utilization, gastrointestinal health, skeletal function, and long-term equine health and performance.
Get Full Text
Publication Date: 2026-09-09 PubMed ID: 42794449PubMed Central: PMC13607830DOI: 10.3390/ijms27188015Google Scholar: Lookup
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
LinkedInCopy
  • Journal Article
  • Review

Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

Overview

  • This review article explores how the gut microbiota in horses influences the regulation of mineral balance in the body, integrating current knowledge about gut physiology, microbiology, and mineral metabolism.
  • It highlights the potential roles of gut microbes in mineral absorption and metabolism, identifies gaps in understanding specific to horses, and suggests directions for future research.

Introduction to Mineral Homeostasis in Horses

  • Traditional understanding attributes mineral balance primarily to coordinated functions of the intestine, kidneys, skeleton, and endocrine system.
  • The gut microbiota is now recognized as an important molecular regulator that modifies the intestinal environment where mineral absorption occurs.
  • Emerging research suggests microbial populations influence mineral availability and uptake, highlighting a complex host-microbe interaction influencing equine mineral homeostasis.

Mechanisms by Which Gut Microbiota Influence Mineral Metabolism

  • Microbial Fermentation: Microorganisms ferment dietary fibers producing short-chain fatty acids (SCFAs) that can impact mineral solubility and absorption.
  • Bioactive Metabolite Production: Microbes synthesize metabolites influencing host physiology, potentially modulating mineral transport and utilization.
  • Luminal pH Regulation: Microbial activity alters the pH within the gut lumen, affecting mineral solubility and ionic state critical for absorption.
  • Epithelial Barrier Integrity: The gut microbiota supports the health and function of the intestinal lining, which is essential for optimal mineral transport across epithelial cells.
  • Modulation of Intestinal Mineral Transporters: Microbial signals may regulate expression or activity of specific mineral transporter proteins in gut epithelial cells.
  • Interactions with Endocrine and Immune Pathways: The microbiota influences host hormonal and immune responses that can indirectly regulate mineral metabolism and distribution.

Minerals Assessed and Species-Specific Considerations

  • Macrominerals: Calcium, phosphorus, magnesium — critical for skeletal health, enzymatic processes, and cellular functions.
  • Trace Minerals: Iron, zinc, copper, selenium, manganese — essential cofactors for enzymes, antioxidants, and immune function.
  • The review differentiates between findings derived directly from equine studies and those extrapolated from other species, emphasizing that direct mechanistic evidence in horses remains scarce.

Knowledge Gaps and Future Research Directions

  • Current evidence linking gut microbial communities and mineral balance in horses is largely correlative or inferred from other species, indicating a lack of direct mechanistic studies.
  • Future research should employ integrated approaches combining:
    • Microbiome profiling to chart microbial populations.
    • Metabolomics to identify microbial metabolites influencing mineral metabolism.
    • Molecular biology to study gene regulation of mineral transporters and host responses.
    • Mineral balance studies to assess physiological outcomes of microbiota changes.
  • Understanding these complex interactions will enable development of microbiome-informed nutritional strategies aimed at improving mineral uptake, gastrointestinal and skeletal health, and ultimately equine performance and longevity.

Implications for Equine Health and Nutrition

  • Enhanced knowledge of microbiota-mineral interactions can inform feed formulation to optimize mineral bioavailability.
  • Improved gut health supported by favorable microbial populations could enhance mineral absorption and reduce deficiencies or imbalances.
  • Protection of skeletal integrity and function through optimized mineral homeostasis may reduce diseases related to mineral deficiency or excess.
  • Integration of microbiome science in equine care could lead to personalized nutrition and management practices boosting overall performance and well-being.

Cite This Article

APA
Sajid QUA, Asghar MU, Wróblewska P, Król B, Korczyński M. (2026). Gut Microbiota as a Molecular Regulator of Mineral Homeostasis in Horses: Mechanisms and Future Perspectives. Int J Mol Sci, 27(18), 8015. https://doi.org/10.3390/ijms27188015

Publication

International journal of molecular sciences
ISSN: 1422-0067
NlmUniqueID: 101092791
Country: Switzerland
Language: English
Volume: 27
Issue: 18
PII: 8015

Researcher Affiliations

Sajid, Qurat Ul Ain
  • Department of Animal Nutrition and Feed Sciences, Wroclaw University of Environmental and Life Sciences, 25 C.K. Norwida St., 51-630 Wrocław, Poland.
Asghar, Muhammad Umair
  • Department of Animal Nutrition and Feed Sciences, Wroclaw University of Environmental and Life Sciences, 25 C.K. Norwida St., 51-630 Wrocław, Poland.
Wróblewska, Patrycja
  • Department of Animal Nutrition and Feed Sciences, Wroclaw University of Environmental and Life Sciences, 25 C.K. Norwida St., 51-630 Wrocław, Poland.
Król, Barbara
  • Department of Animal Nutrition and Feed Sciences, Wroclaw University of Environmental and Life Sciences, 25 C.K. Norwida St., 51-630 Wrocław, Poland.
Korczyński, Mariusz
  • Department of Animal Nutrition and Feed Sciences, Wroclaw University of Environmental and Life Sciences, 25 C.K. Norwida St., 51-630 Wrocław, Poland.

MeSH Terms

  • Animals
  • Horses / microbiology
  • Horses / metabolism
  • Homeostasis
  • Gastrointestinal Microbiome / physiology
  • Minerals / metabolism
  • Trace Elements / metabolism

Conflict of Interest Statement

The authors declare no conflicts of interest.

References

This article includes 119 references
  1. Blaine J., Chonchol M., Levi M.. Renal Control of Calcium, Phosphate, and Magnesium Homeostasis. Clin. J. Am. Soc. Nephrol. 2015;10:1257–1272.
    doi: 10.2215/CJN.09750913pmc: PMC4491294pubmed: 25287933google scholar: lookup
  2. Council N.R.. Nutrient Requirements of Horses. 6th ed. National Academies Press; Washington, DC, USA: 2007.
  3. Geor R.J., Harris P.A., Coenen M.. Equine Applied and Clinical Nutrition: Health, Welfare and Performance. Saunders Elsevier; Edinburgh, UK: 2013.
  4. Hartman M.-L., Razzaque M.S.. Editorial: Endocrine Regulation of Mineral Ions and Their Relevance to Metabolic Diseases. Front. Endocrinol. 2024;15:1391096.
    doi: 10.3389/fendo.2024.1391096pmc: PMC11021768pubmed: 38633752google scholar: lookup
  5. Ralston S.L.. Evidence-Based Equine Nutrition. Vet. Clin. N. Am. Equine Pract. 2007;23:365–384.
    doi: 10.1016/j.cveq.2007.03.007pubmed: 17616318google scholar: lookup
  6. Frape D.E.N.. Feeding. 4th ed. Wiley-Blackwell; Chichester, UK: 2010.
  7. Chaucheyras-Durand F., Sacy A., Karges K., Apper E.. Gastro-Intestinal Microbiota in Equines and Its Role in Health and Disease: The Black Box Opens. Microorganisms 2022;10:2517.
    doi: 10.3390/microorganisms10122517pmc: PMC9783266pubmed: 36557769google scholar: lookup
  8. Costa M.C., Arroyo L.G., Allen-Vercoe E., Stämpfli H.R., Kim P.T., Sturgeon A., Weese J.S.. Comparison of the Fecal Microbiota of Healthy Horses and Horses with Colitis by High Throughput Sequencing of the V3–V5 Region of the 16S rRNA Gene. PLoS ONE 2012;7:41484.
    doi: 10.1371/journal.pone.0041484pmc: PMC3409227pubmed: 22859989google scholar: lookup
  9. Dougal K., Fuente G., Harris P.A., Girdwood S.E., Pinloche E., Newbold C.J.. Identification of a Core Bacterial Community within the Large Intestine of the Horse. PLoS ONE 2013;8:77660.
    doi: 10.1371/journal.pone.0077660pmc: PMC3812009pubmed: 24204908google scholar: lookup
  10. Koh A., Vadder F., Kovatcheva-Datchary P., Bäckhed F.. From Dietary Fiber to Host Physiology: Short-Chain Fatty Acids as Key Bacterial Metabolites. Cell 2016;165:1332–1345.
    doi: 10.1016/j.cell.2016.05.041pubmed: 27259147google scholar: lookup
  11. Louis P., Flint H.J.. Formation of Propionate and Butyrate by the Human Colonic Microbiota. Environ. Microbiol. 2017;19:29–41.
    doi: 10.1111/1462-2920.13589pubmed: 27928878google scholar: lookup
  12. Parada Venegas D., Fuente M.K., Landskron G.. Short Chain Fatty Acids (SCFAs)-Mediated Gut Epithelial and Immune Regulation and Its Relevance for Inflammatory Bowel Diseases. Front. Immunol. 2019;10:277.
    doi: 10.3389/fimmu.2019.00277pmc: PMC6421268pubmed: 30915065google scholar: lookup
  13. Donohoe D.R., Collins L.B., Wali A., Bigler R., Sun W., Bultman S.J.. The Warburg Effect Dictates the Mechanism of Butyrate-Mediated Histone Acetylation and Cell Proliferation. Mol. Cell. 2012;48:612–626.
    doi: 10.1016/j.molcel.2012.08.033pmc: PMC3513569pubmed: 23063526google scholar: lookup
  14. Furusawa Y., Obata Y., Fukuda S., Endo T.A., Nakato G., Takahashi D.. Commensal Microbe-Derived Butyrate Induces the Differentiation of Colonic Regulatory T Cells. Nature 2013;504:446–450.
    doi: 10.1038/nature12721pubmed: 24226770google scholar: lookup
  15. Smith P.M., Howitt M.R., Panikov N., Michaud M., Gallini C.A., Bohlooly-Y M., Glickman J.N., Garrett W.S.. The Microbial Metabolites, Short-Chain Fatty Acids, Regulate Colonic Treg Cell Homeostasis. Science 2013;341:569–573.
    doi: 10.1126/science.1241165pmc: PMC3807819pubmed: 23828891google scholar: lookup
  16. Costa M.C., Silva G., Ramos R.V., Staempfli H.R., Arroyo L.G., Kim P., Weese J.S.. Characterization and Comparison of the Bacterial Microbiota in Different Gastrointestinal Tract Compartments in Horses. Vet. J. 2015;205:74–80.
    doi: 10.1016/j.tvjl.2015.03.018pubmed: 25975855google scholar: lookup
  17. Underwood E.J., Suttle N.F.. The Mineral Nutrition of Livestock. 3rd ed. CABI Publishing; Wallingford, UK: 1999.
  18. Soetan K.O., Olaiya C.O., Oyewole O.E.. The Importance of Mineral Elements for Humans, Domestic Animals and Plants: A Review. Afr. J. Food Sci. 2010;4:200–222.
  19. Bronner F.. Mechanisms of intestinal calcium absorption. J. Cell. Biochem. 2003;88:387–393.
    doi: 10.1002/jcb.10330pubmed: 12520541google scholar: lookup
  20. Christakos S., Dhawan P., Verstuyf A., Verlinden L., Carmeliet G.. Vitamin D: Metabolism, Molecular Mechanism of Action, and Pleiotropic Effects. Physiol. Rev. 2016;96:365–408.
    doi: 10.1152/physrev.00014.2015pmc: PMC4839493pubmed: 26681795google scholar: lookup
  21. Baaij J.H.F., Hoenderop J.G.J., Bindels R.J.M.. Magnesium in Man: Implications for Health and Disease. Physiol. Rev. 2015;95:1–46.
    doi: 10.1152/physrev.00012.2014pubmed: 25540137google scholar: lookup
  22. Razzaque M.S.. The FGF23–Klotho axis: Endocrine regulation of phosphate homeostasis. Nat. Rev. Endocrinol. 2009;5:611–619.
    doi: 10.1038/nrendo.2009.196pmc: PMC3107967pubmed: 19844248google scholar: lookup
  23. Bäckhed F., Ley R.E., Sonnenburg J.L., Peterson D.A., Gordon J.I.. Host-Bacterial Mutualism in the Human Intestine. Science 2005;307:1915–1920.
    doi: 10.1126/science.1104816pubmed: 15790844google scholar: lookup
  24. Tremaroli V., Bäckhed F.. Functional Interactions Between the Gut Microbiota and Host Metabolism. Nature 2012;489:242–249.
    doi: 10.1038/nature11552pubmed: 22972297google scholar: lookup
  25. Rowland I., Gibson G., Heinken A., Scott K., Swann J., Thiele I., Tuohy K.. Gut Microbiota Functions: Metabolism of Nutrients and Other Food Components. Eur. J. Nutr. 2018;57:1–24.
    doi: 10.1007/s00394-017-1445-8pmc: PMC5847071pubmed: 28393285google scholar: lookup
  26. Scholz-Ahrens K.E., Ade P., Marten B., Weber P., Timm W., Açil Y., Glüer C.-C., Schrezenmeir J.. Prebiotics, probiotics, and synbiotics affect mineral absorption, bone mineral content, and bone structure. J. Nutr. 2007;137:838–846.
    doi: 10.1093/jn/137.3.838spubmed: 17311984google scholar: lookup
  27. Valdes A.M., Walter J., Segal E., Spector T.D.. Role of the Gut Microbiota in Nutrition and Health. BMJ 2018;361:2179.
    doi: 10.1136/bmj.k2179pmc: PMC6000740pubmed: 29899036google scholar: lookup
  28. Coleman SJ, Moss C, Wilson A, Reed K, Jennings K, Kunz I. The ‘Horse-Side’of Host-Microbe Interactions and Gastrointestinal Homeostasis in the Equine Hindgut. J. Anim. Sci. 2024;102:216.
    doi: 10.1093/jas/skae234.253google scholar: lookup
  29. Muhonen S, Sadet-Bourgeteau S, Julliand V. Effects of Differences in Fibre Composition and Maturity of Forage-Based Diets on the Microbial Ecosystem and Its Activity in Equine Caecum and Colon Digesta and Faeces. Animals 2021;11:2337.
    doi: 10.3390/ani11082337pmc: PMC8388671pubmed: 34438794google scholar: lookup
  30. Julliand V, Grimm P. Horse Species Symposium: The Microbiome of the Horse Hindgut: History and Current Knowledge. J. Anim. Sci. 2016;94:2262–2274.
    doi: 10.2527/jas.2015-0198pubmed: 27285903google scholar: lookup
  31. Harris PA, Geor RJ. Nutrition for the Equine Athlete: Nutrient Requirements and Key Principles in Ration Design. In: Hinchcliff K.W., Kaneps A.J., Geor R.J., editors. Equine Sports Medicine and Surgery. Saunders Elsevier; New York, NY, USA: 2014. pp. 797–818.
  32. Costa MC, Weese JS. Understanding the Intestinal Microbiome in Health and Disease. Vet. Clin. N. Am. Equine Pract. 2018;34:1–12.
    doi: 10.1016/j.cveq.2017.11.005pubmed: 29402480google scholar: lookup
  33. Stewart RD, Auffret MD, Warr A, Walker AW, Roehe R, Watson M. Compendium of 4,941 Rumen Metagenome-Assembled Genomes for Rumen Microbiome Biology and Enzyme Discovery. Nat. Biotechnol. 2019;37:953–961.
    doi: 10.1038/s41587-019-0202-3pmc: PMC6785717pubmed: 31375809google scholar: lookup
  34. Dougal K, Fuente G, Harris PA, Girdwood SE, Pinloche E, Geor RJ, Nielsen BD, Schott HC, Elzinga S, Newbold CJ. Characterisation of the Faecal Bacterial Community in Adult and Elderly Horses Fed a High Fibre, High Oil or High Starch Diet Using 454 Pyrosequencing. PLoS ONE 2014;9:87424.
    doi: 10.1371/journal.pone.0087424pmc: PMC3913607pubmed: 24504261google scholar: lookup
  35. Ericsson AC, Johnson PJ, Lopes MA, Perry SC, Lanter HR. A Microbiological Map of the Healthy Equine Gastrointestinal Tract. PLoS ONE 2016;11:0166523.
    doi: 10.1371/journal.pone.0166523pmc: PMC5112786pubmed: 27846295google scholar: lookup
  36. Wunderlich G, Bull M, Ross T, Rose M, Chapman B. Understanding the Microbial Fibre Degrading Communities & Processes in the Equine Gut. Anim. Microbiome 2023;5:3.
    doi: 10.1186/s42523-022-00224-6pmc: PMC9837927pubmed: 36635784google scholar: lookup
  37. Li F, Kong X, Khan MZ. Gut Microbiome Regulation in Equine Animals: Current Understanding and Future Perspectives. Front. Microbiol. 2025;16:1602258.
    doi: 10.3389/fmicb.2025.1602258pmc: PMC12504203pubmed: 41070119google scholar: lookup
  38. McKinney CA, Oliveira BC, Bedenice D, Paradis M-R, Mazan M, Sage S, Sanchez A, Widmer G. The Fecal Microbiota of Healthy Donor Horses and Geriatric Recipients Undergoing Fecal Microbial Transplantation for the Treatment of Diarrhea. PLoS ONE 2020;15:0230148.
    doi: 10.1371/journal.pone.0230148pmc: PMC7064224pubmed: 32155205google scholar: lookup
  39. Raspa F, Chessa S, Bergero D, Sacchi P, Ferrocino I, Cocolin L, Corvaglia MR, Moretti R, Cavallini D, Valle E. Microbiota Characterization Throughout the Digestive Tract of Horses Fed a High-Fiber vs. a High-Starch Diet. Front. Vet. Sci. 2024;11:1386135.
    doi: 10.3389/fvets.2024.1386135pmc: PMC11130486pubmed: 38807937google scholar: lookup
  40. Julliand V, Fombelle A, Drogoul C, Jacotot E. Feeding and Microbial Disorders in Horses: Part 3—Effects of Three Hay:Grain Ratios on Microbial Profile and Activities. J. Equine Vet. Sci. 2001;21:543–546.
    doi: 10.1016/s0737-0806(01)70159-1google scholar: lookup
  41. Edwards JE, Shetty SA, Den Berg P, Burden F, Doorn DA, Pellikaan WF, Smidt H. Multi-Kingdom Characterization of the Core Equine Fecal Microbiota Based on Multiple Equine (Sub) Species. Anim. Microbiome 2020;2:6.
    doi: 10.1186/s42523-020-0023-1pmc: PMC7807809pubmed: 33499982google scholar: lookup
  42. Wunderlich G, Bull M, McGilchrist N, Zhao C, Ross T, Rose M, Chapman B. The Horse Gut Bacteriome and Anaerobic Mycobiome Are Influenced by Seasonal Forages and Small Intestinal Starch Digestibility. J. Appl. Microbiol. 2025;136:203.
    doi: 10.1093/jambio/lxaf203pubmed: 40802476google scholar: lookup
  43. Boucher L, Leduc L, Leclere M, Costa M C. Current Understanding of Equine Gut Dysbiosis and Microbiota Manipulation Techniques: Comparison with Current Knowledge in Other Species.. Animals 2024;14:758.
    doi: 10.3390/ani14050758pmc: PMC10931082pubmed: 38473143google scholar: lookup
  44. Daly K, Stewart C S, Flint H J, Shirazi-Beechey S P. Bacterial Diversity within the Equine Large Intestine as Revealed by Molecular Analysis of Cloned 16S rRNA Genes.. FEMS Microbiol. Ecol. 2001;38:141–151.
    doi: 10.1111/j.1574-6941.2001.tb00892.xgoogle scholar: lookup
  45. Grimm P, Combes S, Pascal G, Cauquil L, Julliand V. Dietary Composition and Yeast/Microalgae Combination Supplementation Modulate the Microbial Ecosystem in the Caecum, Colon and Faeces of Horses.. Br. J. Nutr. 2020;123:372–382.
    doi: 10.1017/s0007114519002824pubmed: 31690358google scholar: lookup
  46. Salem S E, Maddox T W, Berg A, Antczak P, Ketley J M, Williams N J. Variation in Faecal Microbiota in a Group of Horses Managed at Pasture over a 12-Month Period.. Sci. Rep. 2018;8:8510.
    doi: 10.1038/s41598-018-26930-3pmc: PMC5981443pubmed: 29855517google scholar: lookup
  47. Whitfield-Cargile C M, Chung H C, Coleman M C. Integrated Analysis of Gut Metabolome, Microbiome, and Exfoliome Data in an Equine Model of Intestinal Injury.. Microbiome 2024;12:74.
    doi: 10.1186/s40168-024-01785-1pmc: PMC11017594pubmed: 38622632google scholar: lookup
  48. Lozupone C A, Stombaugh J I, Gordon J I, Jansson J K, Knight R. Diversity, Stability and Resilience of the Human Gut Microbiota.. Nature 2012;489:220–230.
    doi: 10.1038/nature11550pmc: PMC3577372pubmed: 22972295google scholar: lookup
  49. Flint H J, Scott K P, Duncan S H, Louis P, Forano E. Microbial Degradation of Complex Carbohydrates in the Gut.. Gut Microbes 2012;3:289–306.
    doi: 10.4161/gmic.19897pmc: PMC3463488pubmed: 22572875google scholar: lookup
  50. Derrien M, Belzer C, Vos W M. Akkermansia muciniphila and Its Role in Regulating Host Functions.. Microb. Pathog. 2017;106:171–181.
    doi: 10.1016/j.micpath.2016.02.005pubmed: 26875998google scholar: lookup
  51. Cani P D, Vos W M. Next-Generation Beneficial Microbes: The Case of Akkermansia muciniphila.. Front. Microbiol. 2017;8:1765.
    doi: 10.3389/fmicb.2017.01765pmc: PMC5614963pubmed: 29018410google scholar: lookup
  52. Janssen P H, Kirs M. Structure of the Archaeal Community of the Rumen.. Appl. Environ. Microbiol. 2008;74:3619–3625.
    doi: 10.1128/aem.02812-07pmc: PMC2446570pubmed: 18424540google scholar: lookup
  53. Gruninger R J, Puniya A K, Callaghan T M, Edwards J E, Youssef N, Dagar S S, Fliegerova K, Griffith G W, Forster R, Tsang A. Anaerobic Fungi (Phylum Neocallimastigomycota): Advances in Understanding Their Taxonomy, Life Cycle, Ecology, Role and Biotechnological Potential.. FEMS Microbiol. Ecol. 2014;90:1–17.
    doi: 10.1111/1574-6941.12383pubmed: 25046344google scholar: lookup
  54. Solomon K V, Haitjema C H, Henske J K, Gilmore S P, Borges-Rivera D. Early-Branching Gut Fungi Possess a Large, Comprehensive Array of Biomass-Degrading Enzymes.. Science 2016;351:1192–1195.
    doi: 10.1126/science.aad1431pmc: PMC5098331pubmed: 26912365google scholar: lookup
  55. Liggenstoffer A S, Youssef N H, Couger M B, Elshahed M S. Phylogenetic Diversity and Community Structure of Anaerobic Gut Fungi (Phylum Neocallimastigomycota) in Ruminant and Non-Ruminant Herbivores.. ISME J. 2010;4:1225–1235.
    doi: 10.1038/ismej.2010.49pubmed: 20410935google scholar: lookup
  56. Golomidova A, Kulikov E, Isaeva A, Manykin A, Letarov A. The Diversity of Coliphages and Coliforms in Horse Feces Reveals a Complex Pattern of Ecological Interactions.. Appl. Environ. Microbiol. 2007;73:5975–5981.
    doi: 10.1128/aem.01145-07pmc: PMC2075005pubmed: 17704275google scholar: lookup
  57. Kulikov E. Biodiversity and Dynamics of Bacteriophages in Equine Feces.. Mikrobiologiia 2007;76:271–278.
    pubmed: 17583225
  58. Flint HJ, Bayer EA, Rincon MT, Lamed R, White BA. Polysaccharide Utilization by Gut Bacteria: Potential for New Insights from Genomic Analysis. Nat. Rev. Microbiol. 2008;6:121–131.
    doi: 10.1038/nrmicro1817pubmed: 18180751google scholar: lookup
  59. Gilroy R. Metagenomic Investigation of the Equine Faecal Microbiome Reveals Extensive Taxonomic and Functional Diversity. PeerJ 2022;10:13084.
    doi: 10.7717/peerj.13084pmc: PMC8957277pubmed: 35345588google scholar: lookup
  60. Zhang W, Guo R, Sulayman A, Sun Y, Liu S. Research Progress on Influencing Factors of Gastrointestinal Microbial Diversity in Equine. Vet. Med. Sci. 2025;11:e70271.
    doi: 10.1002/vms3.70271pmc: PMC11948667pubmed: 40145999google scholar: lookup
  61. Bergman EN. Energy Contributions of Volatile Fatty Acids from the Gastrointestinal Tract in Various Species. Physiol. Rev. 1990;70:567–590.
    doi: 10.1152/physrev.1990.70.2.567pubmed: 2181501google scholar: lookup
  62. Buffie CG, Pamer EG. Microbiota-mediated colonization resistance against intestinal pathogens. Nat. Rev. Immunol. 2013;13:790–801.
    doi: 10.1038/nri3535pmc: PMC4194195pubmed: 24096337google scholar: lookup
  63. Pickard JM, Zeng MY, Caruso R, Núñez G. Gut Microbiota: Role in Pathogen Colonization, Immune Responses, and Inflammatory Disease. Immunol. Rev. 2017;279:70–89.
    doi: 10.1111/imr.12567pmc: PMC5657496pubmed: 28856738google scholar: lookup
  64. Schank N, Cottone A, Wulf M, Seiter K, Thomas B, Miller LMJ, Anderson SL, Sahyoun A, Abidi AH, Kassan M. The Role of Short-Chain Fatty Acids (SCFAs) in Colic and Anti-Inflammatory Pathways in Horses. Animals 2025;15:3482.
    doi: 10.3390/ani15233482pmc: PMC12691112pubmed: 41375540google scholar: lookup
  65. Panek M, Čipčić Paljetak H, Barešić A, Perić M, Matijašić M, Lojkić I. Methodology Challenges in Studying Human Gut Microbiota–Effects of Collection, Storage, DNA Extraction and Next Generation Sequencing Technologies. Sci. Rep. 2018;8:5143.
    doi: 10.1038/s41598-018-23296-4pmc: PMC5865204pubmed: 29572539google scholar: lookup
  66. Alou MT, Naud S, Khelaifia S, Bonnet M, Lagier J-C, Raoult D. State of the Art in the Culture of the Human Microbiota: New Interests and Strategies. Clin. Microbiol. Rev. 2020;34:e00129-19.
    doi: 10.1128/CMR.00129-19pmc: PMC7605308pubmed: 33115723google scholar: lookup
  67. Lagier J-C, Dubourg G, Million M, Cadoret F, Bilen M, Fenollar F, Levasseur A, Rolain J-M, Fournier P-E, Raoult D. Culturing the human microbiota and culturomics. Nat. Rev. Microbiol. 2018;16:540–550.
    doi: 10.1038/s41579-018-0041-0pubmed: 29937540google scholar: lookup
  68. Reuter JA, Spacek DV, Snyder MP. High-throughput sequencing technologies. Mol. Cell. 2015;58:586–597.
    doi: 10.1016/j.molcel.2015.05.004pmc: PMC4494749pubmed: 26000844google scholar: lookup
  69. Kauter A, Epping L, Semmler T, Antao EM, Kannapin D, Stoeckle SD, Walther B. The Gut Microbiome of Horses: Current Research on Equine Enteral Microbiota and Future Perspectives. Anim. Microb. 2019;1:14.
    doi: 10.1186/s42523-019-0013-3pmc: PMC7807895pubmed: 33499951google scholar: lookup
  70. DeSantis TZ, Hugenholtz P, Larsen N, Rojas M, Brodie EL, Keller K. Greengenes, a Chimera-Checked 16S rRNA Gene Database and Workbench Compatible with ARB. Appl. Environ. Microb. 2006;72:5069–5072.
    doi: 10.1128/aem.03006-05pmc: PMC1489311pubmed: 16820507google scholar: lookup
  71. Quast C, Pruesse E, Yilmaz P, Gerken J, Schweer T, Yarza P. The SILVA Ribosomal RNA Gene Database Project: Improved Data Processing and Web Based Tools. Nucleic Acids Res. 2013;41:D590–D596.
    doi: 10.1093/nar/gks1219pmc: PMC3531112pubmed: 23193283google scholar: lookup
  72. Janda JM, Abbott SL. 16S rRNA Gene Sequencing for Bacterial Identification in the Diagnostic Laboratory: Pluses, Perils, and Pitfalls. J. Clin. Microbiol. 2007;45:2761–2764.
    doi: 10.1128/jcm.01228-07pmc: PMC2045242pubmed: 17626177google scholar: lookup
  73. Klappenbach JA, Saxman PR, Cole JR, Schmidt TM. Rrndb: The Ribosomal RNA Operon Copy Number Database. Nucleic Acids Res 2001;29:181–184.
    doi: 10.1093/nar/29.1.181pmc: PMC29826pubmed: 11125085google scholar: lookup
  74. Vetrovsky T, Baldrian P. The Variability of the 16S rRNA Gene in Bacterial Genomes and Its Consequences for Bacterial Community Analyses. PLoS ONE 2013;8:e57923.
    doi: 10.1371/journal.pone.0057923pmc: PMC3583900pubmed: 23460914google scholar: lookup
  75. Klindworth A, Pruesse E, Schweer T, Peplies J, Quast C, Horn M. Evaluation of General 16S Ribosomal RNA Gene PCR Primers for Classical and next Generation Sequencing-Based Diversity Studies. Nucleic Acids Res 2013;41:e1.
    doi: 10.1093/nar/gks808pmc: PMC3592464pubmed: 22933715google scholar: lookup
  76. Álvarez Narváez S, Beaudry MS, Norris CG, Bartlett PB, Glenn TC, Sanchez S. Improved Equine Fecal Microbiome Characterization Using Target Enrichment by Hybridization Capture. Animals 2024;14:445.
    doi: 10.3390/ani14030445pmc: PMC10854943pubmed: 38338088google scholar: lookup
  77. Quince C, Walker AW, Simpson JT, Loman NJ, Segata N. Shotgun Metagenomics, from Sampling to Analysis. Nat. Biotechnol. 2017;35:833–844.
    doi: 10.1038/nbt.3935pubmed: 28898207google scholar: lookup
  78. Zou Y, Xue W, Luo G, Deng Z, Qin P, Guo R. 1520 Reference Genomes from Cultivated Human Gut Bacteria Enable Functional Microbiome Analyses. Nat. Biotech. 2019;37:179–185.
    doi: 10.1038/s41587-018-0008-8pmc: PMC6784896pubmed: 30718868google scholar: lookup
  79. Caporaso JG, Kuczynski J, Stombaugh J, Bittinger K, Bushman FD, Costello EK. QIIME Allows Analysis of High-Throughput Community Sequencing Data. Nat. Meth. 2010;7:335–336.
    doi: 10.1038/nmeth.f.303pmc: PMC3156573pubmed: 20383131google scholar: lookup
  80. Schloss PD, Westcott SL, Ryabin T, Hall JR, Hartmann M, Hollister EB, Lesniewski RA, Oakley BB, Parks DH, Robinson CJ. Introducing Mothur: Open-Source, Platform-Independent, Community Supported Software for Describing and Comparing Microbial Communities. Appl. Environ. Microbiol. 2009;75:7537–7541.
    doi: 10.1128/aem.01541-09pmc: PMC2786419pubmed: 19801464google scholar: lookup
  81. Ludwig JA, Reynolds JF. Statistical Ecology—A Primer on Methods and Computing. .
  82. Simpson EH. Measurement of diversity. Nature 1949;163:688.
    doi: 10.1038/163688a0google scholar: lookup
  83. Chao A. Nonparametric Estimation of the Number of Classes in a Population. Scand. J. Stat. 1984;11:265–270.
  84. Maier I, Kienzle E. A Meta-Analysis on Quantitative Calcium, Phosphorus and Magnesium Metabolism in Horses and Ponies. Animals 2024;14:2765.
    doi: 10.3390/ani14192765pmc: PMC11475699pubmed: 39409714google scholar: lookup
  85. Etemadi F, Tabatabaei Naeini A, Aminlari M. Assessment of calcium, phosphorus, magnesium, vitamin D and PTH levels in sera of lame horses. Vet. Med. Sci. 2023;9:2070–2077.
    doi: 10.1002/vms3.1198pmc: PMC10508493pubmed: 37466035google scholar: lookup
  86. Selle PH, Ravindran V. Microbial Phytase in Poultry Nutrition. Anim. Feed Sci. Technol. 2007;135:1–41.
    doi: 10.1016/j.anifeedsci.2006.06.010google scholar: lookup
  87. Ganz T. Systemic iron homeostasis. Physiol. Rev. 2013;93:1721–1741.
    doi: 10.1152/physrev.00008.2013pubmed: 24137020google scholar: lookup
  88. Anderson GJ, McLaren GD. Iron Physiology and Pathophysiology in Humans. Humana Press; New York, NY, USA: 2012.
  89. Cousins RJ, Liuzzi JP, Lichten LA. Mammalian zinc transport, trafficking, and signals. J. Biol. Chem. 2006;281:24085–24089.
    doi: 10.1074/jbc.r600011200pubmed: 16793761google scholar: lookup
  90. Prohaska JR. Impact of Copper Limitation on Expression and Function of Multicopper Oxidases (Ferroxidases). Adv. Nutr. 2014;5:476–479.
    pmc: PMC3065751pubmed: 22332037
  91. Rayman MP. Selenium and human health. Lancet 2012;379:1256–1268.
    doi: 10.1016/s0140-6736(11)61452-9pubmed: 22381456google scholar: lookup
  92. Tinggi U. Selenium: Its Role as Antioxidant in Human Health. Environ. Health Prev. Med. 2008;13:102–108.
    doi: 10.1007/s12199-007-0019-4pmc: PMC2698273pubmed: 19568888google scholar: lookup
  93. Aschner M, Erikson KM. Manganese. Adv. Nutr. 2017;8:520–521.
    doi: 10.3945/an.117.015305pmc: PMC5421128pubmed: 28507016google scholar: lookup
  94. Sprekeler N, Müller T, Kowalewski MP, Liesegang A, Boos A. Expression patterns of intestinal calcium transport factors and ex-vivo absorption of calcium in horses. BMC Vet. Res. 2011;7:65.
    doi: 10.1186/1746-6148-7-65pmc: PMC3221622pubmed: 22017756google scholar: lookup
  95. Wilkens MR, Marholt L, Eigendorf N, Muscher-Banse AS, Feige K, Schroder B, Breves G, Cehak A. Trans- and Paracellular Calcium Transport along the Small and Large Intestine in Horses. Comp. Biochem. Physiol. Part A Mol. Integr. Physiol. 2017;204:157–163.
    doi: 10.1016/j.cbpa.2016.11.020pubmed: 27915149google scholar: lookup
  96. Cehak A, Wilkens MR, Guschlbauer M, Mrochen N, Schröder B, Feige K, Breves G. In Vitro Studies on Intestinal Calcium and Phosphate Transport in Horses. Comp. Biochem. Physiol. Part A Mol. Integr. Physiol. 2012;161:259–264.
    doi: 10.1016/j.cbpa.2011.11.005pubmed: 22119207google scholar: lookup
  97. Weaver CM, Heaney RP. Calcium. In: Shils ME, Shike M, Ross AC, Caballero B, Cousins RJ, editors. Modern Nutrition in Health and Disease. Lippincott Williams & Wilkins; Philadelphia, PA, USA: 2006.
  98. Salim R, Nehvi IB, Mir RA, Tyagi A, Ali S, Bhat OM. A Review on Anti-Nutritional Factors: Unraveling the Natural Gateways to Human Health. Front. Nutr. 2023;10:1215873.
    doi: 10.3389/fnut.2023.1215873pmc: PMC10501406pubmed: 37720376google scholar: lookup
  99. Gibson GR. Dietary Prebiotics: Current Status and New Definition. Food Sci. Technol. Bull. Funct. Foods. 2010;7:1–19.
    doi: 10.1616/1476-2137.15880google scholar: lookup
  100. Brinkley-Bissinger K, Cersosimo LM, Sullivan KE, Livingston SE, Bobel JM, Valdes EV, Warren LK. Phytate Supplementation Has Minimal Impact on Mineral Digestibility in Horses. J. Anim. Sci. 2020;98:89.
    doi: 10.1093/jas/skaa278.163google scholar: lookup
  101. Plancade S, Clark A, Philippe C, Helbling J-C, Moisan M-P, Esquerré D, Le Moyec L, Robert C, Barrey E, Mach N. Unraveling the Effects of the Gut Microbiota Composition and Function on Horse Endurance Physiology. Sci. Rep. 2019;9:9620.
    doi: 10.1038/s41598-019-46118-7pmc: PMC6610142pubmed: 31270376google scholar: lookup
  102. Schoster A, Mosing M, Jalali M, Staempfli HR, Weese JS. Effects of Transport, Fasting and Anaesthesia on the Faecal Microbiota of Healthy Adult Horses. Equine Vet. J. 2016;48:595–602.
    doi: 10.1111/evj.12479pubmed: 26122549google scholar: lookup
  103. Skrypnik K, Suliburska J. Association Between the Gut Microbiota and Mineral Metabolism. J. Sci. Food Agric. 2018;98:2449–2460.
    doi: 10.1002/jsfa.8724pubmed: 28991359google scholar: lookup
  104. Dinu L-D, Avram I, Pelinescu D-R, Vamanu E. Mineral-Enriched Postbiotics: A New Perspective for Microbial Therapy to Prevent and Treat Gut Dysbiosis. Biomedicines 2022;10:2392.
    doi: 10.3390/biomedicines10102392pmc: PMC9599024pubmed: 36289654google scholar: lookup
  105. Peterson L.W, Artis D. Intestinal Epithelial Cells: Regulators of Barrier Function and Immune Homeostasis. Nat. Rev. Immunol. 2014;14:141–153.
    doi: 10.1038/nri3608pubmed: 24566914google scholar: lookup
  106. Pajarillo E.A.B, Lee E, Kang D-K. Trace Metals and Animal Health: Interplay of the Gut Microbiota with Iron, Manganese, Zinc, and Copper. Anim. Nutr. 2021;7:750–761.
    doi: 10.1016/j.aninu.2021.03.005pmc: PMC8379138pubmed: 34466679google scholar: lookup
  107. Kortman G.A.M, Raffatellu M, Swinkels D.W, Tjalsma H. Nutritional Iron Turned inside out: Intestinal Stress from a Gut Microbial Perspective. FEMS Microbiol. Rev. 2014;38:1202–1234.
    doi: 10.1111/1574-6976.12086pubmed: 25205464google scholar: lookup
  108. Sun Y.F, Han Z.X, Yao X.K, Meng J, Ren W.L, Wang C.K, Wang J.W. Effects of Different Stages of Training on the Intestinal Microbes of Yili Horses Analyzed Using Metagenomics. Genes 2025;16:504.
    doi: 10.3390/genes16050504pmc: PMC12111061pubmed: 40428326google scholar: lookup
  109. Culhuac E.B, Elghandour M.M.M.Y, Adegbeye M.J, Barbabosa-Pliego A, Salem A.Z.M. Influence of Dietary Selenium on the Oxidative Stress in Horses. Biol. Trace Elem. Res. 2022;201:1695–1703.
    doi: 10.1007/s12011-022-03270-ypubmed: 35526205google scholar: lookup
  110. Mach N, Midoux C, Leclercq S, Pennarun S, Le Moyec L, Rué O, Robert C, Sallé G, Barrey E. Mining the Equine Gut Metagenome: Poorly-Characterized Taxa Associated with Cardiovascular Fitness in Endurance Athletes. Commun. Biol. 2022;5:1032.
    doi: 10.1038/s42003-022-03977-7pmc: PMC9529974pubmed: 36192523google scholar: lookup
  111. Yur F, Dede S, Deger Y, Kilicalp D. Effects of Vitamin E and Selenium on Serum Trace and Major Elements in Horses. Biol. Trace Elem. Res. 2008;125:223–228.
    doi: 10.1007/s12011-008-8109-8pubmed: 18797827google scholar: lookup
  112. Ingala M.R, Simmons N.B, Wultsch C, Krampis K, Speer K.A, Perkins S.L. Comparing Microbiome Sampling Methods in a Wild Mammal: Fecal and Intestinal Samples Record Different Signals of Host Ecology, Evolution. Front. Microbiol. 2018;9:803.
    doi: 10.3389/fmicb.2018.00803pmc: PMC5938605pubmed: 29765359google scholar: lookup
  113. Brandi L.A, Nunes A.T, Faleiros C.A, Poleti M.D, Oliveira E.C.D.M, Schmidt N.T, Brandi R.A. Dietary Energy Sources Affect Cecal and Fecal Microbiota of Healthy Horses. Animals 2024;14:3494.
    doi: 10.3390/ani14233494pmc: PMC11639918pubmed: 39682460google scholar: lookup
  114. Mach N, Foury A, Kittelmann S, Reigner F, Moroldo M, Ballester M, Esquerré D, Rivière J, Sallé G, Gérard P. The Effects of Weaning Methods on Gut Microbiota Composition and Horse Physiology. Front. Physiol. 2017;8:535.
    doi: 10.3389/fphys.2017.00535pmc: PMC5524898pubmed: 28790932google scholar: lookup
  115. Mura E, Edwards J, Kittelmann S, Kaerger K, Voigt K, Mrázek J, Moniello G, Fliegerova K. Anaerobic Fungal Communities Differ Along the Horse Digestive Tract. Fungal Biol. 2019;123:240–246.
    doi: 10.1016/j.funbio.2018.12.004pubmed: 30798879google scholar: lookup
  116. François A.C, Taminiau B, Renaud B, Gonza-Quito I.E, Massey C, Hyde C, Votion D.M. In Vitro Investigation of Equine Gut Microbiota Alterations During Hypoglycin A Exposure. Animals 2025;15:3343.
    doi: 10.3390/ani15223343pmc: PMC12649563pubmed: 41302050google scholar: lookup
  117. Duvallet C, Gibbons S.M, Gurry T, Irizarry R.A, Alm E.J. Meta-Analysis of Gut Microbiome Studies Identifies Disease-Specific and Shared Responses. Nat. Comm. 2017;8:1784.
    doi: 10.1038/s41467-017-01973-8pmc: PMC5716994pubmed: 29209090google scholar: lookup
  118. Wei J, Wei L, Ullah A, Geng M, Zhang X, Wang C, Khan MZ, Wang C, Zhang Z. Metagenomic Applications to Herbivore Gut Microbiomes: A Comprehensive Review of Microbial Diversity and Host Interactions.. Animals 2025;15:2938.
    doi: 10.3390/ani15202938pmc: PMC12560936pubmed: 41153865google scholar: lookup
  119. McEwan NR, Nava-Morales GM, Morrison PK, Reis de Souza T, Escobar-García K, Bautista-Marín SE, Mariscal-Ladín G, Silva-Jarquín JC, Gomez-Soto JG, Olvera-Ramírez AM. Using Gut Microbiota and Gut Metabolites as Biomarkers for Diseases in Monogastric Farm Animals.. Animals 2026;16:2247.
    doi: 10.3390/ani16142247pmc: PMC13403961pubmed: 42511129google scholar: lookup

Citations

This article has been cited 0 times.
Subscribe to our newsletter
Join 107,147 horse owners like you who receive our email updates on horse health and nutrition.
OmegaPrime+ Extra-Strength Omega-3 Oil - Mad Barn
NEW PRODUCT
OmegaPrime+
Extra-Strength Omega-3 Oil
Shop Now
×