Monocyte subsets across veterinary species

Monocyte subsets across veterinary species

Javier Domínguez
1
,
Loems Ziegler-Heitbrock
2,*
*Correspondence to: Loems Ziegler-Heitbrock, Independent researcher, Monocytomics Research, Herrsching 82211, Germany. E-mail: LZH@monocyte.eu
Myeloid Cells. 2026;1:202601. 10.70401/mc.2026.0004
Received: null , Accepted: May 18, 2026Published: May 18, 2026
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This manuscript is made available in its unedited form to allow early access to the reported findings. Further editing will be completed before final publication. As such, the content may include errors, and standard legal disclaimers are applicable.

Abstract

Monocytes consist of several subsets, which differ in their phenotype and functional capacity. This heterogeneity was first shown in man, and later evidenced in other species. Here, we review the current knowledge on the phenotype and functionality of monocyte populations in species of veterinary interest, in comparison to man. The increasing availability of reagents for leukocyte phenotypic analyses, along with the growing application of -omic technologies, are enabling a better characterization of these subsets, and facilitate comparisons across species. The review demonstrates that classical, intermediate, and non-classical monocytes can be defined in pig, cattle, sheep, camel, buffalo, horse, and dog. However, more phenotypic, functional, and transcriptomic studies are needed in some species to establish an accurate correspondence. A better understanding of shared and species-specific features of these monocyte subsets will pave the way for translational research on strategies selectively targeting these cells to treat different pathological conditions, including infectious, inflammatory, and malignant diseases.

Keywords

Monocyte subsets, veterinary species, phenotype, surface receptors, transcriptomics

References

  • 1. Jakubzick CV, Randolph GJ, Henson PM. Monocyte differentiation and antigen-presenting functions. Nat Rev Immunol. 2017;17(6):349-362.
    [DOI] [PubMed]
  • 2. Mildner A, Kim KW, Yona S. Unravelling monocyte functions: From the guardians of health to the regulators of disease. Discov Immunol. 2024;3(1):kyae014.
    [DOI] [PubMed] [PMC]
  • 3. Serbina NV, Jia T, Hohl TM, Pamer EG. Monocyte-mediated defense against microbial pathogens. Annu Rev Immunol. 2008;26:421-452.
    [DOI] [PubMed] [PMC]
  • 4. Hristov M, Weber C. Monocyte subsets in cardiovascular disease: A biomarker perspective. Thromb Haemost. 2025;125(2):93-96.
    [DOI] [PubMed]
  • 5. Gerhardt T, Ley K. Monocyte trafficking across the vessel wall. Cardiovasc Res. 2015;107(3):321-330.
    [DOI] [PubMed] [PMC]
  • 6. Guan F, Wang R, Yi Z, Luo P, Liu W, Xie Y, et al. Tissue macrophages: Origin, heterogenity, biological functions, diseases and therapeutic targets. Sig Transduct Target Ther. 2025;10(1):93.
    [DOI] [PubMed] [PMC]
  • 7. Wang Z, Wu Z, Wang H, Feng R, Wang G, Li M, et al. An immune cell atlas reveals the dynamics of human macrophage specification during prenatal development. Cell. 2023;186(20):4454-4471.e19.
    [DOI] [PubMed]
  • 8. T’Jonck W, Bain CC. The role of monocyte-derived macrophages in the lung: It’s all about context. Int J Biochem Cell Biol. 2023;159:106421.
    [DOI] [PubMed]
  • 9. Wen Y, Lambrecht J, Ju C, Tacke F. Hepatic macrophages in liver homeostasis and diseases-diversity, plasticity and therapeutic opportunities. Cell Mol Immunol. 2021;18(1):45-56.
    [DOI] [PubMed] [PMC]
  • 10. Shimak MJ, Kim G, Karkache IY, Vu EK, Chavez E, Manser JC, et al. From development, disease, and decline: A review of what defines an osteoclast progenitor. Int J Mol Sci. 2025;26(21):10619.
    [DOI] [PubMed] [PMC]
  • 11. Andoh M, Koyama R. Comparative review of microglia and monocytes in CNS phagocytosis. Cells. 2021;10(10):2555.
    [DOI] [PubMed] [PMC]
  • 12. Kim JS, Trzebanski S, Shin SH, Schori L, Frumer Friedman GR, Ilani NC, et al. Clonal hematopoiesis-associated motoric deficits caused by monocyte-derived microglia accumulating in aging mice. Cell Rep. 2025;44(5):115609.
    [DOI] [PubMed]
  • 13. Guilliams M, Mildner A, Yona S. Developmental and functional heterogeneity of monocytes. Immunity. 2018;49(4):595-613.
    [DOI] [PubMed]
  • 14. Passlick B, Flieger D, Ziegler-Heitbrock HW. Identification and characterization of a novel monocyte subpopulation in human peripheral blood. Blood. 1989;74(7):2527-2534.
  • 15. Ziegler-Heitbrock L, Ancuta P, Crowe S, Dalod M, Grau V, Hart DN, et al. Nomenclature of monocytes and dendritic cells in blood. Blood. 2010;116(16):e74-e80.
    [DOI] [PubMed]
  • 16. Sunderkötter C, Nikolic T, Dillon MJ, van Rooijen N, Stehling M, Drevets DA, et al. Subpopulations of mouse blood monocytes differ in maturation stage and inflammatory response. J Immunol. 2004;172(7):4410-4417.
    [DOI] [PubMed]
  • 17. Geissmann F, Jung S, Littman DR. Blood monocytes consist of two principal subsets with distinct migratory properties. Immunity. 2003;19(1):71-82.
    [DOI] [PubMed]
  • 18. Briseño CG, Haldar M, Kretzer NM, Wu X, Theisen DJ, Kc W, et al. Distinct transcriptional programs control cross-priming in classical and monocyte-derived dendritic cells. Cell Rep. 2016;15(11):2462-2474.
    [DOI] [PubMed] [PMC]
  • 19. Palframan RT, Jung S, Cheng G, Weninger W, Luo Y, Dorf M, et al. Inflammatory chemokine transport and presentation in HEV: A remote control mechanism for monocyte recruitment to lymph nodes in inflamed tissues. J Exp Med. 2001;194(9):1361-1373.
    [DOI] [PubMed] [PMC]
  • 20. Tacke F, Randolph GJ. Migratory fate and differentiation of blood monocyte subsets. Immunobiology. 2006;211(6-8):609-618.
    [DOI] [PubMed]
  • 21. Weber C, Belge KU, von Hundelshausen P, Draude G, Steppich B, Mack M, et al. Differential chemokine receptor expression and function in human monocyte subpopulations. J Leukoc Biol. 2000;67(5):699-704.
    [DOI] [PubMed]
  • 22. Steppich B, Dayyani F, Gruber R, Lorenz R, Mack M, Ziegler-Heitbrock HW. Selective mobilization of CD14+CD16+ monocytes by exercise. Am J Physiol Cell Physiol. 2000;279(3):C578-C586.
    [DOI] [PubMed]
  • 23. Jung S, Aliberti J, Graemmel P, Sunshine MJ, Kreutzberg GW, Sher A, et al. Analysis of fractalkine receptor CX(3)CR1 function by targeted deletion and green fluorescent protein reporter gene insertion. Mol Cell Biol. 2000;20(11):4106-4114.
    [DOI] [PubMed] [PMC]
  • 24. Meghraoui-Kheddar A, Barthelemy S, Boissonnas A, Combadière C. Revising CX3CR1 expression on murine classical and non-classical monocytes. Front Immunol. 2020;11:1117.
    [DOI] [PubMed] [PMC]
  • 25. Sommer K, Garibagaoglu H, Paap EM, Wiendl M, Müller TM, Atreya I, et al. Discrepant phenotyping of monocytes based on CX3CR1 and CCR2 using fluorescent reporters and antibodies. Cells. 2024;13(10):819.
    [DOI] [PubMed] [PMC]
  • 26. Dayyani F, Joeinig A, Ziegler-Heitbrock L, Schmidmaier R, Straka C, Emmerich B, et al. Autologous stem-cell transplantation restores the functional properties of CD14+CD16+ monocytes in patients with myeloma and lymphoma. J Leukoc Biol. 2004;75(2):207-213.
    [DOI] [PubMed]
  • 27. Rogacev KS, Zawada AM, Hundsdorfer J, Achenbach M, Held G, Fliser D, et al. Immunosuppression and monocyte subsets. Nephrol Dial Transplant. 2015;30(1):143-153.
    [DOI] [PubMed]
  • 28. Patel AA, Zhang Y, Fullerton JN, Boelen L, Rongvaux A, Maini AA, et al. The fate and lifespan of human monocyte subsets in steady state and systemic inflammation. J Exp Med. 2017;214(7):1913-1923.
    [DOI] [PubMed] [PMC]
  • 29. Tak T, Drylewicz J, Conemans L, de Boer RJ, Koenderman L, Borghans JAM, et al. Circulatory and maturation kinetics of human monocyte subsets in vivo. Blood. 2017;130(12):1474-1477.
    [DOI] [PubMed]
  • 30. Hettinger J, Richards DM, Hansson J, Barra MM, Joschko AC, Krijgsveld J, et al. Origin of monocytes and macrophages in a committed progenitor. Nat Immunol. 2013;14(8):821-830.
    [DOI]
  • 31. Gamrekelashvili J, Giagnorio R, Jussofie J, Soehnlein O, Duchene J, Briseño CG, et al. Regulation of monocyte cell fate by blood vessels mediated by Notch signalling. Nat Commun. 2016;7:12597.
    [DOI] [PubMed] [PMC]
  • 32. Gamrekelashvili J, Kapanadze T, Sablotny S, Ratiu C, Dastagir K, Lochner M, et al. Notch and TLR signaling coordinate monocyte cell fate and inflammation. Elife. 2020;9:e57007.
    [DOI] [PubMed] [PMC]
  • 33. Lessard AJ, LeBel M, Egarnes B, Préfontaine P, Thériault P, Droit A, et al. Triggering of NOD2 receptor converts inflammatory Ly6Chigh into Ly6Clow monocytes with patrolling properties. Cell Rep. 2017;20(8):1830-1843.
    [DOI] [PubMed]
  • 34. Talker SC, Barut GT, Lischer HEL, Rufener R, von Münchow L, Bruggmann R, et al. Monocyte biology conserved across species: Functional insights from cattle. Front Immunol. 2022;13:889175.
    [DOI] [PubMed] [PMC]
  • 35. Patel RS, Tomlinson JE, Divers TJ, van de Walle GR, Rosenberg BR. Single-cell resolution landscape of equine peripheral blood mononuclear cells reveals diverse cell types including T-bet+ B cells. BMC Biol. 2021;19(1):13.
    [DOI] [PubMed] [PMC]
  • 36. Vu Manh TP, Elhmouzi-Younes J, Urien C, Ruscanu S, Jouneau L, Bourge M, et al. Defining mononuclear phagocyte subset homology across several distant warm-blooded vertebrates through comparative transcriptomics. Front Immunol. 2015;6:299.
    [DOI] [PubMed] [PMC]
  • 37. Alvarez B, Sánchez C, Bullido R, Marina A, Lunney J, Alonso F, et al. A porcine cell surface receptor identified by monoclonal antibodies to SWC3 is a member of the signal regulatory protein family and associates with protein-tyrosine phosphatase SHP-1. Tissue Antigens. 2000;55(4):342-351.
    [DOI] [PubMed]
  • 38. Moffat L, Rothwell L, Garcia-Morales C, Sauter KA, Kapetanovic R, Gow DJ, et al. Development and characterisation of monoclonal antibodies reactive with porcine CSF1R (CD115). Dev Comp Immunol. 2014;47(1):123-128.
    [DOI] [PubMed]
  • 39. Waddell LA, Lefevre L, Bush SJ, Raper A, Young R, Lisowski ZM, et al. ADGRE1 (EMR1 F4/80) is a rapidly-evolving gene expressed in mammalian monocyte-macrophages. Front Immunol. 2018;9:2246.
    [DOI] [PubMed] [PMC]
  • 40. Ziegler-Heitbrock HW, Appl B, Käfferlein E, Löffler T, Jahn-Henninger H, Gutensohn W, et al. The antibody MY4 recognizes CD14 on porcine monocytes and macrophages. Scand J Immunol. 1994;40(5):509-514.
    [DOI] [PubMed]
  • 41. Sánchez C, Doménech N, Vázquez J, Alonso F, Ezquerra A, Domínguez J. The porcine 2A10 antigen is homologous to human CD163 and related to macrophage differentiation. J Immunol. 1999;162(9):5230-5237.
  • 42. Ondrackova P, Nechvatalova K, Kucerova Z, Leva L, Dominguez J, Faldyna M. Porcine mononuclear phagocyte subpopulations in the lung, blood and bone marrow: Dynamics during inflammation induced by Actinobacillus pleuropneumoniae. Vet Res. 2010;41(5):64.
    [DOI] [PubMed] [PMC]
  • 43. Fairbairn L, Kapetanovic R, Beraldi D, Sester DP, Tuggle CK, Archibald AL, et al. Comparative analysis of monocyte subsets in the pig. J Immunol. 2013;190(12):6389-6396.
    [DOI] [PubMed]
  • 44. Chamorro S, Revilla C, Alvarez B, López-Fuertes L, Ezquerra A, Domínguez J. Phenotypic characterization of monocyte subpopulations in the pig. Immunobiology. 2000;202(1):82-93.
    [DOI] [PubMed]
  • 45. Chamorro S, Revilla C, Alvarez B, Alonso F, Ezquerra A, Domínguez J. Phenotypic and functional heterogeneity of porcine blood monocytes and its relation with maturation. Immunology. 2005;114(1):63-71.
    [DOI] [PubMed] [PMC]
  • 46. Poderoso T, de la Riva PM, Álvarez B, Domínguez J, Ezquerra Á, Revilla C. CD200R family receptors are expressed on porcine monocytes and modulate the production of IL-8 and TNF-α triggered by TLR4 or TLR7 in these cells. Mol Immunol. 2022;144:166-177.
    [DOI] [PubMed]
  • 47. Auray G, Keller I, Python S, Gerber M, Bruggmann R, Ruggli N, et al. Characterization and transcriptomic analysis of porcine blood conventional and plasmacytoid dendritic cells reveals striking species-specific differences. J Immunol. 2016;197(12):4791-4806.
    [DOI] [PubMed]
  • 48. Álvarez B, Nieto-Pelegrín E, Martínez de la Riva P, Toki D, Poderoso T, Revilla C, et al. Characterization of the porcine CLEC12A and analysis of its expression on blood dendritic cell subsets. Front Immunol. 2020;11:863.
    [DOI] [PubMed] [PMC]
  • 49. Ondrackova P, Matiasovic J, Volf J, Dominguez J, Faldyna M. Phenotypic characterisation of the monocyte subpopulations in healthy adult pigs and Salmonella-infected piglets by seven-colour flow cytometry. Res Vet Sci. 2013;94(2):240-245.
    [DOI] [PubMed]
  • 50. Blanc F, Prévost-Blondel A, Piton G, Bouguyon E, Leplat JJ, Andréoletti F, et al. The composition of circulating leukocytes varies with age and melanoma onset in the MeLiM pig biomedical model. Front Immunol. 2020;11:291.
    [DOI] [PubMed] [PMC]
  • 51. Cormican S, Griffin MD. Human monocyte subset distinctions and function: Insights from gene expression analysis. Front Immunol. 2020;11:1070.
    [DOI] [PubMed] [PMC]
  • 52. Hillman H, Khan N, Singhania A, Dubelko P, Soldevila F, Tippalagama R, et al. Single-cell profiling reveals distinct subsets of CD14+ monocytes drive blood immune signatures of active tuberculosis. Front Immunol. 2022;13:1087010.
    [DOI] [PubMed] [PMC]
  • 53. Mildner A, Schönheit J, Giladi A, David E, Lara-Astiaso D, Lorenzo-Vivas E, et al. Genomic characterization of murine monocytes reveals C/EBPβ transcription factor dependence of Ly6C- cells. Immunity. 2017;46(5):849-862.e7.
    [DOI] [PubMed]
  • 54. Domínguez J, Ezquerra A, Alonso F, McCullough K, Summerfield A, Bianchi A, et al. Porcine myelomonocytic markers: Summary of the Second International Swine CD Workshop. Vet Immunol Immunopathol. 1998;60(3-4):329-341.
    [DOI] [PubMed]
  • 55. Ezquerra A, Revilla C, Alvarez B, Pérez C, Alonso F, Domínguez J. Porcine myelomonocytic markers and cell populations. Dev Comp Immunol. 2009;33(3):284-298.
    [DOI] [PubMed]
  • 56. Bullido R, Doménech N, Alvarez B, Alonso F, Babín M, Ezquerra A, et al. Characterization of five monoclonal antibodies specific for swine class II major histocompatibility antigens and crossreactivity studies with leukocytes of domestic animals. Dev Comp Immunol. 1997;21(3):311-322.
    [DOI] [PubMed]
  • 57. Weiner LM, Li W, Holmes M, Catalano RB, Dovnarsky M, Padavic K, et al. Phase I trial of recombinant macrophage colony-stimulating factor and recombinant gamma-interferon: Toxicity, monocytosis, and clinical effects. Cancer Res. 1994;54(15):4084-4090.
  • 58. Korkosz M, Bukowska-Strakova K, Sadis S, Grodzicki T, Siedlar M. Monoclonal antibodies against macrophage colony-stimulating factor diminish the number of circulating intermediate and nonclassical (CD14(++)CD16+/CD14+CD16(++)) monocytes in rheumatoid arthritis patient. Blood. 2012;119(22):5329-5330.
    [DOI] [PubMed]
  • 59. Reith W, LeibundGut-Landmann S, Waldburger JM. Regulation of MHC class II gene expression by the class II transactivator. Nat Rev Immunol. 2005;5(10):793-806.
    [DOI] [PubMed]
  • 60. Hofer TP, Zawada AM, Frankenberger M, Skokann K, Satzl AA, Gesierich W, et al. Slan-defined subsets of CD16-positive monocytes: Impact of granulomatous inflammation and M-CSF receptor mutation. Blood. 2015;126(24):2601-2610.
    [DOI] [PubMed]
  • 61. Tamassia N, Bianchetto-Aguilera F, Gasperini S, Grimaldi A, Montaldo C, Calzetti F, et al. The slan antigen identifies the prototypical non-classical CD16+-monocytes in human blood. Front Immunol. 2023;14:1287656.
    [DOI] [PubMed] [PMC]
  • 62. Deloizy C, Bouguyon E, Fossum E, Sebo P, Osicka R, Bole A, et al. Expanding the tools for identifying mononuclear phagocyte subsets in swine: Reagents to porcine CD11c and XCR1. Dev Comp Immunol. 2016;65:31-40.
    [DOI] [PubMed]
  • 63. Hussen J, Düvel A, Sandra O, Smith D, Sheldon IM, Zieger P, et al. Phenotypic and functional heterogeneity of bovine blood monocytes. PLoS One. 2013;8(8):e71502.
    [DOI] [PubMed] [PMC]
  • 64. Moreno S, Alvarez B, Poderoso T, Revilla C, Ezquerra A, Alonso F, et al. Porcine monocyte subsets differ in the expression of CCR2 and in their responsiveness to CCL2. Vet Res. 2010;41(5):76.
    [DOI] [PubMed] [PMC]
  • 65. Ondrackova P, Leva L, Kucerova Z, Vicenova M, Mensikova M, Faldyna M. Distribution of porcine monocytes in different lymphoid tissues and the lungs during experimental Actinobacillus pleuropneumoniae infection and the role of chemokines. Vet Res. 2013;44(1):98.
    [DOI] [PubMed] [PMC]
  • 66. Sauter KA, Waddell LA, Lisowski ZM, Young R, Lefevre L, Davis GM, et al. Macrophage colony-stimulating factor (CSF1) controls monocyte production and maturation and the steady-state size of the liver in pigs. Am J Physiol Gastrointest Liver Physiol. 2016;311(3):G533-G547.
    [DOI] [PubMed] [PMC]
  • 67. Glorion M, Pascale F, Huriet M, Estephan J, Gouin C, Urien C, et al. Differential early response of monocyte/macrophage subsets to intra-operative corticosteroid administration in lung transplantation. Front Immunol. 2023;14:1281546.
    [DOI] [PubMed] [PMC]
  • 68. Herrera-Uribe J, Wiarda JE, Sivasankaran SK, Daharsh L, Liu H, Byrne KA, et al. Reference transcriptomes of porcine peripheral immune cells created through bulk and single-cell RNA sequencing. Front Genet. 2021;12:689406.
    [DOI] [PubMed] [PMC]
  • 69. Jaudas F, Bartenschlager F, Shashikadze B, Santamaria G, Reichart D, Schnell A, et al. Perinatal dysfunction of innate immunity in cystic fibrosis. Sci Transl Med. 2025;17(782):eadk9145.
    [DOI]
  • 70. Pernold CPS, Lagumdzic E, Stadler M, Mair KH, Jäckel S, Schmitt MW, et al. Characterization of the immune system of Ellegaard Göttingen Minipigs - An important large animal model in experimental medicine. Front Immunol. 2022;13:1003986.
    [DOI] [PubMed] [PMC]
  • 71. Jarosova R, Ondrackova P, Leva L, Nedbalcova K, Vicenova M, Masek J, et al. Cytokine expression by CD163+ monocytes in healthy and Actinobacillus pleuropneumoniae-infected pigs. Res Vet Sci. 2022;152:1-9.
    [DOI] [PubMed]
  • 72. Frandoloso R, Martínez-Martínez S, Yubero S, Rodríguez-Ferri EF, Gutiérrez-Martín CB. New insights in cellular immune response in colostrum-deprived pigs after immunization with subunit and commercial vaccines against Glässer’s disease. Cell Immunol. 2012;277(1-2):74-82.
    [DOI] [PubMed]
  • 73. Grandoni F, Scatà MC, Martucciello A, de Carlo E, de Matteis G, Hussen J. Comprehensive phenotyping of peripheral blood monocytes in healthy bovine. Cytometry A. 2022;101(2):122-130.
    [DOI] [PubMed]
  • 74. Talker SC, Hope JC, Summerfield A. Phenotype of bovine mononuclear phagocytes- An update. Vet Immunol Immunopathol. 2024;277:110836.
    [DOI] [PubMed]
  • 75. Hussen J, Schuberth HJ. Heterogeneity of bovine peripheral blood monocytes. Front Immunol. 2017;8:1875.
    [DOI] [PubMed] [PMC]
  • 76. Corripio-Miyar Y, Hope J, McInnes CJ, Wattegedera SR, Jensen K, Pang Y, et al. Phenotypic and functional analysis of monocyte populations in cattle peripheral blood identifies a subset with high endocytic and allogeneic T-cell stimulatory capacity. Vet Res. 2015;46:112.
    [DOI] [PubMed] [PMC]
  • 77. Talker SC, Baumann A, Barut GT, Keller I, Bruggmann R, Summerfield A. Precise delineation and transcriptional characterization of bovine blood dendritic-cell and monocyte subsets. Front Immunol. 2018;9:2505.
    [DOI] [PubMed] [PMC]
  • 78. Ingersoll MA, Spanbroek R, Lottaz C, Gautier EL, Frankenberger M, Hoffmann R, et al. Comparison of gene expression profiles between human and mouse monocyte subsets. Blood. 2010;115(3):e10-e19.
    [DOI] [PubMed] [PMC]
  • 79. Belge KU, Dayyani F, Horelt A, Siedlar M, Frankenberger M, Frankenberger B, et al. The proinflammatory CD14+CD16+DR++ monocytes are a major source of TNF. J Immunol. 2002;168(7):3536-3542.
    [DOI] [PubMed]
  • 80. Hussen J, Frank C, Düvel A, Koy M, Schuberth HJ. The chemokine CCL5 induces selective migration of bovine classical monocytes and drives their differentiation into LPS-hyporesponsive macrophages in vitro. Dev Comp Immunol. 2014;47(2):169-177.
    [DOI] [PubMed]
  • 81. Elnaggar MM, Abdellrazeq GS, Mack V, Fry LM, Davis WC, Park KT. Characterization and use of new monoclonal antibodies to CD11c, CD14, and CD163 to analyze the phenotypic complexity of ruminant monocyte subsets. Vet Immunol Immunopathol. 2016;178:57-63.
    [DOI] [PubMed]
  • 82. Park KT, Burnett S, Davis WC. Development and characterization of a monoclonal antibody specific for bovine CD209. Vet Immunol Immunopathol. 2015;163(3-4):216-220.
    [DOI] [PubMed]
  • 83. Zheng W, Wang X, Liu J, Yu X, Li L, Wang H, et al. Single-cell analyses highlight the proinflammatory contribution of C1q-high monocytes to Behçet’s disease. Proc Natl Acad Sci U S A. 2022;119(26):e2204289119.
    [DOI] [PubMed] [PMC]
  • 84. Rigamonti A, Castagna A, Viatore M, Colombo FS, Terzoli S, Peano C, et al. Distinct responses of newly identified monocyte subsets to advanced gastrointestinal cancer and COVID-19. Front Immunol. 2022;13:967737.
    [DOI] [PubMed] [PMC]
  • 85. Wilson A, Alexandre PA, Brice AM, Hine BC, Ingham A, Legrand TPRA, et al. Single-cell transcriptomics uncovers key immune drivers of vaccine efficacy in cattle. BMC Genom. 2025;26(1):750.
    [DOI]
  • 86. Han B, Li H, Zheng W, Zhang Q, Chen A, Zhu S, et al. A multi-tissue single-cell expression atlas in cattle. Nat Genet. 2025;57(10):2546-2561.
    [DOI] [PubMed] [PMC]
  • 87. Eger M, Hussen J, Drong C, Meyer U, von Soosten D, Frahm J, et al. Impacts of parturition and body condition score on glucose uptake capacity of bovine monocyte subsets. Vet Immunol Immunopathol. 2015;166(1-2):33-42.
    [DOI] [PubMed]
  • 88. Pomeroy B, Sipka A, Hussen J, Eger M, Schukken Y, Schuberth HJ. Counts of bovine monocyte subsets prior to calving are predictive for postpartum occurrence of mastitis and metritis. Vet Res. 2017;48(1):13.
    [DOI]
  • 89. Franzoni G, Righi C, de Donato I, Cappelli G, de Matteis G, Scoccia E, et al. Characterisation of cell-mediated immunity against bovine alphaherpesvirus 1 (BoAHV-1) in calves. Vaccines. 2025;13(10):996.
    [DOI] [PubMed] [PMC]
  • 90. Bastos RG, Sears K, Dinkel KD, Knowles DP, Fry LM. Changes in the molecular and functional phenotype of bovine monocytes during Theileria parva infection. Infect Immun. 2019;87(12):e00703-e00719.
    [DOI] [PubMed] [PMC]
  • 91. Pridans C, Davis GM, Sauter KA, Lisowski ZM, Corripio-Miyar Y, Raper A, et al. A Csf1r-EGFP transgene provides a novel marker for monocyte subsets in sheep. J Immunol. 2016;197(6):2297-2305.
    [DOI] [PubMed] [PMC]
  • 92. Ahmed MH, Wilkens MR, Möller B, Ganter M, Breves G, Schuberth HJ. Blood leukocyte composition and function in periparturient ewes kept on different dietary magnesium supply. BMC Vet Res. 2020;16(1):484.
    [DOI] [PubMed] [PMC]
  • 93. Hecker YP, Coronado M, Hurtado-Morillas C, Arranz-Solís D, Sánchez-Sánchez R, Corbí Á, et al. Ovine macrophage identity and plasticity: Novel insights into CSF-driven polarization and species-specific responses. Front Immunol. 2025;16:1680086.
    [DOI] [PubMed] [PMC]
  • 94. Mobley JL, Leininger M, Madore S, Baginski TJ, Renkiewicz R. Genetic evidence of a functional monocyte dichotomy. Inflammation. 2007;30(6):189-197.
    [DOI]
  • 95. Saresella M, Piancone F, Marventano I, Hernis A, Trabattoni D, Invernizzi M, et al. Innate immune responses to three doses of the BNT162b2 mRNA SARS-CoV-2 vaccine. Front Immunol. 2022;13:947320.
    [DOI] [PubMed] [PMC]
  • 96. Hussen J, Schuberth HJ. Recent advances in camel immunology. Front Immunol. 2020;11:614150.
    [DOI] [PubMed] [PMC]
  • 97. Hussen J, Shawaf T, Al-Mubarak AIA, Al Humam NA, Almathen F, Schuberth HJ. Dromedary camel CD14high MHCIIhigh monocytes display inflammatory properties and are reduced in newborn camel calves. BMC Vet Res. 2020;16(1):62.
    [DOI] [PubMed] [PMC]
  • 98. Hussen J, Shawaf T, Al-Mubarak AIA, Humam NAA, Almathen F, Schuberth HJ. Leukocyte populations in peripheral blood of dromedary camels with clinical endometritis. Anim Reprod Sci. 2020;222:106602.
    [DOI] [PubMed]
  • 99. Hussen J. Bacterial species-specific modulatory effects on phenotype and function of camel blood leukocytes. BMC Vet Res. 2021;17(1):241.
    [DOI] [PubMed] [PMC]
  • 100. Hussen J, Althagafi H. Serum Cortisol level as marker of stress in camels: Relationship with immunological profile. Front Vet Sci. 2025;12:1570564.
    [DOI] [PubMed] [PMC]
  • 101. Elnaggar MM, Grandoni F, Abdellrazeq GS, Fry LM, El-Naggar K, Hulubei V, et al. Pattern of CD14, CD16, CD163 and CD172a expression on water buffalo (Bubalus bubalis) leukocytes. Vet Immunol Immunopathol. 2019;211:1-5.
    [DOI] [PubMed]
  • 102. Grandoni F, Fraboni D, Canonico B, Papa S, Buccisano F, Schuberth HJ, et al. Flow cytometric identification and enumeration of monocyte subsets in bovine and water buffalo peripheral blood. Curr Protoc. 2023;3(2):e676.
    [DOI] [PubMed]
  • 103. Grandoni F, Hussen J, Signorelli F, Napolitano F, Scatà MC, de Donato I, et al. Evaluation of hematological profiles and monocyte subpopulations in water buffalo calves after immunization with two different IBR marker vaccines and subsequent infection with Bubaline alphaherpesvirus-1. Vaccines. 2023;11(9):1405.
    [DOI] [PubMed] [PMC]
  • 104. Arunachalam PS, Scott MKD, Hagan T, Li C, Feng Y, Wimmers F, et al. Systems vaccinology of the BNT162b2 mRNA vaccine in humans. Nature. 2021;596(7872):410-416.
    [DOI] [PubMed] [PMC]
  • 105. Mohanty S, Joshi SR, Ueda I, Wilson J, Blevins TP, Siconolfi B, et al. Prolonged proinflammatory cytokine production in monocytes modulated by interleukin 10 after influenza vaccination in older adults. J Infect Dis. 2015;211(7):1174-1184.
    [DOI] [PubMed] [PMC]
  • 106. Scatà MC, De Matteis G, Grandoni F, Di Vuolo G, Cappelli G, Vecchio D. Characterization of cellular immune system at different ages in water buffalo (bubalus bubalis). J Buf Sci. 2024;13:133-139.
    [DOI]
  • 107. Mauel S, Steinbach F, Ludwig H. Monocyte-derived dendritic cells from horses differ from dendritic cells of humans and mice. Immunology. 2006;117(4):463-473.
    [DOI] [PubMed] [PMC]
  • 108. Noronha LE, Harman RM, Wagner B, Antczak DF. Generation and characterization of monoclonal antibodies to equine CD16. Vet Immunol Immunopathol. 2012;146(2):135-142.
    [DOI] [PubMed] [PMC]
  • 109. Gibbons N, Goulart MR, Chang YM, Efstathiou K, Purcell R, Wu Y, et al. Phenotypic heterogeneity of peripheral monocytes in healthy dogs. Vet Immunol Immunopathol. 2017;190:26-30.
    [DOI] [PubMed]
  • 110. Rzepecka A, Żmigrodzka M, Witkowska-Piłaszewicz O, Cywińska A, Winnicka A. CD4 and MHCII phenotypic variability of peripheral blood monocytes in dogs. PLoS One. 2019;14(7):e0219214.
    [DOI] [PubMed] [PMC]
  • 111. McDonald E, Kehoe E, Deines D, McCarthy M, Wright B, Huse S. High-parameter immunophenotyping reveals distinct immune cell profiles in pruritic dogs and cats. Front Vet Sci. 2024;11:1498964.
    [DOI] [PubMed] [PMC]
  • 112. Ammons DT, Harris RA, Hopkins LS, Kurihara J, Weishaar K, Dow S. A single-cell RNA sequencing atlas of circulating leukocytes from healthy and osteosarcoma affected dogs. Front Immunol. 2023;14:1162700.
    [DOI] [PubMed] [PMC]
  • 113. Kim MC, Gu T, Seo H, Moon Y, Borcherding N, Kolb R, et al. High-resolution single-cell RNA sequencing using canFam4 reveals novel immune subsets and checkpoint programs in healthy dogs. Front Immunol. 2025;16:1680437.
    [DOI] [PubMed] [PMC]
  • 114. Garcia-Morales C, Rothwell L, Moffat L, Garceau V, Balic A, Sang HM, et al. Production and characterisation of a monoclonal antibody that recognises the chicken CSF1 receptor and confirms that expression is restricted to macrophage-lineage cells. Dev Comp Immunol. 2014;42(2):278-285.
    [DOI] [PubMed]
  • 115. Härtle S, Sutton K, Vervelde L, Dalgaard TS. Delineation of chicken immune markers in the era of omics and multicolor flow cytometry. Front Vet Sci. 2024;11:1385400.
    [DOI] [PubMed] [PMC]
  • 116. Staines K, Hunt LG, Young JR, Butter C. Evolution of an expanded mannose receptor gene family. PLoS One. 2014;9(11):e110330.
    [DOI] [PubMed] [PMC]
  • 117. Hu T, Wu Z, Bush SJ, Freem L, Vervelde L, Summers KM, et al. Characterization of subpopulations of chicken mononuclear phagocytes that express TIM4 and CSF1R. J Immunol. 2019;202(4):1186-1199.
    [DOI] [PubMed] [PMC]
  • 118. Maxwell M, Söderlund R, Härtle S, Wattrang E. Single-cell RNA-seq mapping of chicken peripheral blood leukocytes. BMC Genomics. 2024;25(1):124.
    [DOI] [PubMed] [PMC]
  • 119. O’Connor KW, Liu T, Kim S, Briseño CG, Georgopoulos K, Murphy TL, et al. Bcl6, Irf2, and Notch2 promote nonclassical monocyte development. Proc Natl Acad Sci U S A. 2023;120(35):e2220853120.
    [DOI] [PubMed] [PMC]
  • 120. Hanna RN, Carlin LM, Hubbeling HG, Nackiewicz D, Green AM, Punt JA, et al. The transcription factor NR4A1 (Nur77) controls bone marrow differentiation and the survival of Ly6C- monocytes. Nat Immunol. 2011;12(8):778-785.
    [DOI] [PubMed] [PMC]
  • 121. Hume DA, Wollscheid-Lengeling E, Rojo R, Pridans C. The evolution of the macrophage-specific enhancer (Fms intronic regulatory element) within the CSF1R locus of vertebrates. Sci Rep. 2017;7(1):17115.
    [DOI] [PubMed] [PMC]
  • 122. Entrican G, Lunney JK, Wattegedera SR, Mwangi W, Hope JC, Hammond JA. The veterinary immunological toolbox: Past, present, and future. Front Immunol. 2020;11:1651.
    [DOI] [PubMed] [PMC]
  • 123. Mwangi W, Maccari G, Hope JC, Entrican G, Hammond JA. The UK Veterinary Immunological Toolbox Website: Promoting vaccine research by facilitating communication and removing reagent barriers. Immunology. 2020;161(1):25-27.
    [DOI] [PubMed] [PMC]
  • 124. Banstola A, Reynolds JNJ. The sheep as a large animal model for the investigation and treatment of human disorders. Biology. 2022;11(9):1251.
    [DOI] [PubMed] [PMC]
  • 125. Lunney JK, van Goor A, Walker KE, Hailstock T, Franklin J, Dai C. Importance of the pig as a human biomedical model. Sci Transl Med. 2021;13(621):eabd5758.
    [DOI] [PubMed]
  • 126. van Rhijn I, Godfroid J, Michel A, Rutten V. Bovine tuberculosis as a model for human tuberculosis: Advantages over small animal models. Microbes Infect. 2008;10(7):711-715.
    [DOI] [PubMed]
  • 127. Guerra-Maupome M, Palmer MV, McGill JL, Sacco RE. Utility of the neonatal calf model for testing vaccines and intervention strategies for use against human RSV infection. Vaccines. 2019;7(1):7.
    [DOI] [PubMed] [PMC]

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Domínguez J, Ziegler-Heitbrock L. Monocyte subsets across veterinary species. Myeloid Cells. 2026;1:202601. https://doi.org/10.70401/mc.2026.0004

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