Macrophage activation and polarization: nomenclature and experimental guidelines
Open accessAbstract
Description of macrophage activation is currently contentious and confusing. Like the biblical Tower of Babel, macrophage activation encompasses a panoply of descriptors used in different ways. The lack of consensus on how to define macrophage activation in experiments in vitro and in vivo impedes progress in multiple ways, including the fact that many researchers still consider there to be only two types of activated macrophages, often termed M1 and M2. Here, we describe a set of standards encompassing three principles-the source of macrophages, definition of the activators, and a consensus collection of markers to describe macrophage activation-with the goal of unifying experimental standards for diverse experimental scenarios. Collectively, we propose a common framework for macrophage-activation nomenclature.
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iNOS2 has been associated with immunostimulation and immunosuppression.
“iNOS2, which has been associated with immunostimulation (Hibbs et al., 1987; Murray et al., 2014; Stuehr and Nathan, 1989), as well as immunosuppression (Shi et al., 2001).”
From TREM2 Modulation Remodels the Tumor Myeloid Landscape Enhancing Anti-PD-1 Immunotherapy (2020) · cites this paper as Murray et al., 2014
The Background to the problem (p. 4) discusses macrophage activation and arginine metabolism, but does not link iNOS2 to immunostimulation. Searches for iNOS, NOS2, and nitric oxide did not locate a passage in our copy making that association. In the citing sentence, immunosuppression is attributed to Shi et al. (2001), not to this source. (LLM, conf=0.71)
Experimental — accuracy not yet measuredMacrophage polarization is instructed by biomechanical properties of the fibrotic microenvironment, including matrix stiffness, fluid shear stress, and intercellular tension, through integrin/FAK, Piezo1/TRPV4, YAP/TAZ, NF-kB, and ROCK signaling.
“Beyond biochemical cues, the biomechanical properties of the fibrotic microenvironment-notably increased matrix stiffness, fluid shear stress, and intercellular tension-are now recognized as active instructors of macrophage polarization (6,74,75).”
From The mechanotransduction-immune axis in organ fibrosis: dual regulatory mechanisms and translational therapeutic perspectives (2025) · cites this paper as [75]
A pro-wound healing macrophage phenotype correlates with upregulation of Dcir, Stab1, CCL22, YM1, and Arg1.
“A pro-wound healing phenotype of macrophage correlates with up regulation in expression of Dendritic Cell Immune Receptor (Dcir), Stabilin 1 (Stab1), chemokine (C-C motif) ligand 22 (CCL22), Chitinase-like 3 (YM1), and Arginase 1 (Arg1)44, 45, 46.”
From Size- and shape-dependent foreign body immune response to materials implanted in rodents and non-human primates (2015) · cites this paper as [45]
A pro-inflammatory macrophage phenotype correlates with elevated expression of TNFα, IL1, and IRF5.
“A pro-inflammatory macrophage phenotype correlates with elevated expression of Tumor Necrosis Factor α (TNF α), Interleukin 1 (IL1), and Interferon Regulatory Factor 5 (IRF5)44, 45, 46.”
From Size- and shape-dependent foreign body immune response to materials implanted in rodents and non-human primates (2015) · cites this paper as [45]
A regulatory macrophage phenotype correlates with elevated expression of IL10, Light, and Sphk1.
“Meanwhile, a regulatory macrophage phenotype has been characterized to correlate with elevated expression of Interleukin 10 (IL10), TNF superfamily member 14 (Light), and Sphingosine Kinase 1 (Sphk1)44, 45, 46.”
From Size- and shape-dependent foreign body immune response to materials implanted in rodents and non-human primates (2015) · cites this paper as [45]
In vivo, macrophages and monocytes show mixed phenotypes and are not fully polarised as the M1/M2 classification suggests.
“In vivo, however, macrophages (and monocytes) display mixed phenotypes and are not completely polarised as described by the in vitro classification of M1/M2 (42,43).”
From The impact of ageing on monocytes and macrophages (2021) · cites this paper as [43]
The M1/M2 nomenclature for activated microglia is now deemed inaccurate as accumulating evidence has revealed a broad diversity of macrophage activation statuses and remarkable plasticity to change phenotypes in response to environmental cues.
“However, this nomenclature is now deemed inaccurate as accumulating evidence has revealed a broad diversity of macrophage activation statuses and shown that macrophages show remarkable plasticity to change phenotypes in response to environmental cues25,26.”
From Interplay between innate immunity and Alzheimer disease: APOE and TREM2 in the spotlight (2018) · cites this paper as [26]
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- Authors
- Murray, P. J., Allen, J. E., Biswas, S. K., Fisher, E. A., Gilroy, D. W., Goerdt, S., Gordon, S., Hamilton, J. A., Ivashkiv, L. B., Lawrence, T., Locati, M., Mantovani, A., Martinez, F. O., Mege, J.-L., Mosser, D. M., Natoli, G., Saeij, J. P., Schultze, J. L., Shirey, K. A., Sica, A., Suttles, J., Udalova, I., van Ginderachter, J. A., Vogel, S. N., & Wynn, T. A.
- Affiliations
- St. Jude Children's Research Hospital (US)
- Centre for Immunity, Infection and Evolution (GB)
- Agency for Science, Technology and Research (SG)
- New York University (US)
- University College London (GB)
- Heidelberg University (DE)
- University of Oxford (GB)
- The Royal Melbourne Hospital (AU)
- Hospital for Special Surgery (US)
- Centre d’Immunologie de Marseille-Luminy (FR)
- University of Milan (IT)
- University of Maryland, College Park (US)
- European Institute of Oncology (IT)
- Massachusetts Institute of Technology (US)
- University of Bonn (DE)
- University of Maryland, Baltimore (US)
- Università degli Studi del Piemonte Orientale “Amedeo Avogadro” (IT)
- University of Louisville (US)
- National Institutes of Health (US)
- Corresponding author
- Murray, Peter J.
- ORCID
- Murray, Peter J.
- Allen, Judith E.
- Fisher, Edward A.
- Gilroy, Derek W.
- Goerdt, Sergij
- Gordon, Siamon
- Hamilton, John A.
- Ivashkiv, Lionel B.
- Lawrence, Toby
- Locati, Massimo
- Mantovani, Alberto
- Mosser, David M.
- Natoli, Gioacchino
- Saeij, Jeroen P.
- Schultze, Joachim L.
- Shirey, Kari Ann
- Sica, Antonio
- Udalova, Irina
- Vogel, Stefanie N.
- Wynn, Thomas A.
- Venue
- Immunity
- Publisher
- Elsevier BV
- Published
- 2014-07
- Volume
- 41
- Issue
- 1
- Pages
- 14-20
- ISSN
- 1074-7613 (ISSN-L)
- PMC
- PMC4123412
- OpenAlex
- W2159560463
- Cited by
- 6,337
- References
- 50
- Citations by year
- 2026: 590 · 2025: 519 · 2024: 585 · 2023: 540 · 2022: 653 · 2021: 637 · 2020: 635 · 2019: 489 · 2018: 439 · 2017: 477 · 2016: 408 · 2015: 294 · 2014: 54 · 2012: 2
- FWCI
- 186.58
- Type
- journal article
- Language
- en
- Update notices
- erratum (2014-08-21)
- Open access
- bronze
- doi.org
- Licence
- Locations
- Immunity (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- Open Research (University of Surrey) (submittedVersion)
- UCL Discovery (University College London) (submittedVersion)
- eScholarship (California Digital Library) (submittedVersion)
- Research Portal (King's College London) (submittedVersion, other-oa)
- Research Explorer (The University of Manchester) (submittedVersion)
- VUBIR (Vrije Universiteit Brussel) (submittedVersion)
- Funders
- Wellcome Trust (095831, 087520)
- National Institutes of Health (2r01ai046712-26)
- Medical Research Council (MR/J001899/1, MR/K01207X/1)
- National Heart, Lung, and Blood Institute (R01HL084312, HL-084312)
- National Cancer Institute (P30CA021765)
- National Institute of General Medical Sciences (R01GM102589)
- National Institute of Allergy and Infectious Diseases (R56AI018797, AI-18797, R21AI097485, AI080621, R37AI018797, AI062921, R01-AI046712, R01AI018797, R01AI080621, R01AI062921, R01AI044938)
- Topics
- Life Sciences › Immunology and Microbiology › Immunology › Immune cells in cancer
- Life Sciences › Immunology and Microbiology › Immunology › Immune responses and vaccinations
- Life Sciences › Immunology and Microbiology › Immunology › Immune Response and Inflammation
- Keywords
- macrophage activation, macrophage polarization, M1/M2 macrophages, nomenclature, standardization, macrophage markers
- MeSH terms
- Animals, Humans, Macrophage Activation, Macrophages, Terminology as Topic, Research, Macrophage Colony-Stimulating Factor, Granulocyte-Macrophage Colony-Stimulating Factor, Guidelines as Topic, Mice
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