Exploring the full spectrum of macrophage activation
Open accessAbstract
Macrophages display remarkable plasticity and can change their physiology in response to environmental cues. These changes can give rise to different populations of cells with distinct functions. In this Review we suggest a new grouping of macrophage populations based on three different homeostatic activities - host defence, wound healing and immune regulation. We propose that similarly to primary colours, these three basic macrophage populations can blend into various other 'shades' of activation. We characterize each population and provide examples of macrophages from specific disease states that have the characteristics of one or more of these populations.
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During cancer development, tumor cells and tumor-associated stromal cells continuously change the composition and organization of ECM through deposition/degradation cycles and extensive crosslinking, shaping it more supportive for tumor growth.
“During cancer development, tumor cells and tumor‐associated stromal cells continuously change the composition and organization of ECM through deposition/degradation cycles and extensive crosslinking, shaping it more supportive for tumor growth.[ 33 ]”
From 3D Biomimetic Models to Reconstitute Tumor Microenvironment In Vitro: Spheroids, Organoids, and Tumor‐on‐a‐Chip (2023) · cites this paper as [33]
Split into 2 atoms — 0 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 1 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measuredA more recent model hypothesizes a macrophage spectrum between M1 and M2, regulated through three major homeostatic processes involving host defense, wound healing, and immune modulation.
“A more recent model also hypothesizes the existence of a macrophage spectrum between M1 and M2, regulated through three major homeostatic processes involving the host defense, wound healing, and immune modulation.[ 40 ]”
From Unraveling and Harnessing the Immune Response at the Cell–Biomaterial Interface for Tissue Engineering Purposes (2024) · cites this paper as [40]
The Abstract proposes a spectrum of macrophage populations based on three homeostatic functions: host defence, wound healing and immune regulation. But it describes the linear spectrum between M1 and M2 as an earlier classification that its three-population model seeks to improve, not as the model it proposes. The functions are a basis for classification, rather than processes said to regulate the spectrum. (LLM, conf=0.88)
Experimental — accuracy not yet measuredCommon examples of cytokines that polarize macrophages are IFN-γ and LPS, which skew macrophages toward M1, whereas IL-4 and IL-13 promote a regenerative M2 phenotype.
“Common examples of cytokines that polarize macrophages are IFN‐γ and LPS, which skew macrophages toward the M1 phenotype, whereas cytokines such as IL‐4 and IL‐13 promote a regenerative M2 phenotype (Figure 2 ).[ 40 ]”
From Unraveling and Harnessing the Immune Response at the Cell–Biomaterial Interface for Tissue Engineering Purposes (2024) · cites this paper as [40]
The Classically activated macrophages section links IFNγ and LPS-mediated TLR stimulation to classical activation, while the Wound-healing macrophages section links IL-4 and IL-13 to wound-healing macrophages. But the paper calls LPS a TLR ligand, not a cytokine, and cautions that M2 is broader than the wound-healing phenotype. Our copy provides section labels but no page numbers. (LLM, conf=0.88)
Experimental — accuracy not yet measuredM1-M2 transition is critical for resolution of inflammation and tipping the balance to tissue repair.
“This M1–M2 transition is critical for the resolution of inflammation and tipping the balance to tissue repair [49].”
From Transition from inflammation to proliferation: a critical step during wound healing (2016) · cites this paper as [49]
The Wound-healing macrophages section (page unavailable) says IL-4 can convert resident macrophages to a wound-healing phenotype and that these cells produce few pro-inflammatory cytokines. This supports a shift toward tissue repair, but the paper does not establish that an M1–M2 transition is critical for resolving inflammation. (LLM, conf=0.84)
Experimental — accuracy not yet measuredAβ deposition elicits a robust M1 microglia-mediated inflammatory response contributing to disease pathogenesis.
“Aβ deposition elicits a robust “M1” microglia-mediated inflammatory response contributing to disease pathogenesis (Gordon, 2003; Mosser and Edwards, 2008; Mandrekar-Colucci and Landreth, 2010).”
From Mechanisms Underlying the Rapid Peroxisome Proliferator-Activated Receptor-γ-Mediated Amyloid Clearance and Reversal of Cognitive Deficits in a Murine Model of Alzheimer's Disease (2012) · cites this paper as Mosser and Edwards, 2008
The ‘Classically activated macrophages’ section says their pro-inflammatory cytokines can damage the host and contribute to inflammatory pathology. The paper also mentions CNS microglia in ‘The origin and maintenance of macrophages,’ but does not say that Aβ deposition triggers an M1 microglial response or that such a response contributes to disease pathogenesis. No page numbers are available in our copy. (LLM, conf=0.85)
Experimental — accuracy not yet measured
The biochemical composition of the microenvironment is especially important for determination of macrophage phenotypes.
“The biochemical composition of the microenvironment is especially important for the determination of macrophage phenotypes.”
From Unraveling and Harnessing the Immune Response at the Cell–Biomaterial Interface for Tissue Engineering Purposes (2024) · cites this paper as [40]
Macrophages/microglia are capable of exhibiting a spectrum of phenotypic activation states which dictate their activities.
“Macrophages/microglia are capable of exhibiting a spectrum of phenotypic activation states which dictate their activities (Gordon, 2003; Mosser and Edwards, 2008; Mandrekar-Colucci and Landreth, 2010).”
From Mechanisms Underlying the Rapid Peroxisome Proliferator-Activated Receptor-γ-Mediated Amyloid Clearance and Reversal of Cognitive Deficits in a Murine Model of Alzheimer's Disease (2012) · cites this paper as Mosser and Edwards, 2008
The observed heterogeneity of macrophage shapes (elongated and round) reflects the diverse plasticity of macrophage phenotypes.
“the heterogeneity of cell morphologies observed (elongated and round cells) reflect the diverse plasticity of macrophage cells that define its myriad of phenotypes [56].”
From Characterization of topographical effects on macrophage behavior in a foreign body response model (2010) · cites this paper as [56]
Macrophages, as part of their myeloid lineage, do not possess stress fibers, possibly because their interaction with surrounding matrix when not migrating is not a key part of their innate function.
“It has been shown that macrophages, as part of their myeloid lineage, do not possess stress fibers,58 which may in large part be due to the fact that their interaction with their surrounding matrix, when not migrating, is not a key part of their innate function.29”
From Student award winner in the undergraduate category for the society of biomaterials 9th World Biomaterials Congress, Chengdu, China, June 1–5, 2012: The effects of substrate stiffness on the <i>in vitro</i> activation of macrophages and <i>in vivo</i> host response to poly(ethylene glycol)‐based hydrogels (2012) · cites this paper as [29]
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 [44]
Macrophages can be categorized into three activation states: classical/pro-inflammatory, alternative/pro-healing, and regulatory.
“There is a broad spectrum of macrophage activations which can be categorized into three states namely, classical activation (pro-inflammatory, MClassic), alternative activation (pro-healing, MWound), and regulatory activation (MReg)44.”
From Size- and shape-dependent foreign body immune response to materials implanted in rodents and non-human primates (2015) · cites this paper as [44]
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 [44]
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 [44]
The M2c subset is induced by IL-10 and plays a major role in tissue remodeling and suppression of inflammatory immune reactions by secreting TGF-b and IL-10.
“Alternatively, the M2c subset is induced by IL-10 and plays a major role in tissue remodeling and suppression of inflammatory immune reactions by secreting transforming growth factor-b (TGF-b) and IL-10 [1,23].”
From Biomaterial based modulation of macrophage polarization: a review and suggested design principles (2015) · cites this paper as [23]
Macrophage phenotype occurs along a continuum between M1 and M2 extremes.
“Rather, macrophage phenotype has been described as occurring along a continuum between the M1 and M2 extremes [12].”
From Macrophage polarization: an opportunity for improved outcomes in biomaterials and regenerative medicine (2012) · cites this paper as [12]
M2 macrophages include three sub-populations (M2a, M2b, M2c) with distinct inducers, markers, and functions.
“While the term ‘M2’ encompasses all non-classically activated macrophages, M2 macrophages have been shown to include three well recognized sub-populations (M2a, M2b, and M2c), each with its own distinct inducers, markers, and functions.”
From Macrophage polarization: an opportunity for improved outcomes in biomaterials and regenerative medicine (2012) · cites this paper as [12]
Macrophages are a heterogeneous cell population with different markers and functions.
“Macrophages, like the monocytes from which they derive, are a heterogeneous cell population each with different markers and functions [1,8–12].”
From Macrophage polarization: an opportunity for improved outcomes in biomaterials and regenerative medicine (2012) · cites this paper as [12]
Macrophages serve three primary functions: host defense, wound healing, and immune regulation.
“Indeed, macrophages serve three primary functions in the body: host defense, wound healing, and immune regulation.95”
From Biocompatible Materials for Continuous Glucose Monitoring Devices (2013) · cites this paper as [95]
Individual macrophages can dynamically switch phenotype, altering their functional properties.
“Moreover, individual macrophages hold the potential for a dynamic phenotype switch, which affects the functional properties ofthe macrophage155,156.”
From Inflammation and its resolution in atherosclerosis: mediators and therapeutic opportunities (2019) · cites this paper as [155]
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 [25]
Transitioning from a pro-inflammatory macrophage-dominant wound to an anti-inflammatory macrophage-dominant milieu is essential in resolving inflammation and preparing the wound for effective repair.
“Transitioning from a pro-inflammatory macrophage-dominant wound to an anti-inflammatory macrophage-dominant milieu is essential in resolving inflammation and preparing the wound for effective repair [72].”
From Immunology of Wound Healing (2018) · cites this paper as [72]
In the late stages of the inflammatory phase of wound healing, macrophages switch from a proinflammatory to an alternatively activated phenotype.
“In the late stages of the inflammatory phase of wound healing, macrophages are found to switch from a proinflammatory to an alternatively activated phenotype, which has been assumed to be requisite for the switch from the inflammatory to the proliferative stages in wound healing as discussed below in more detail.46”
From The Innate Immune System in Acute and Chronic Wounds (2016) · cites this paper as [46]
M1 and M2a/b/c are not discrete Mp differentiation states but represent convenient but limited and arbitrary in vitro-defined reference points on a time-dependent and multidimensional continuum of possible Mp phenotypes and functions.
“Thus, available evidence indicates that M1 and M2a/b/c are not discrete Mp differentiation states but instead represent convenient but limited and arbitrary in vitro-defined reference points on a time-dependent and multidimensional continuum of possible Mp phenotypes and functions [9].”
From Macrophage phenotypes during tissue repair (2013) · cites this paper as [9]
Rodent and human macrophages do not exhibit entirely similar responses to in vitro activation.
“Finally and perhaps most importantly, rodent and human Mp do not exhibit entirely similar responses to in vitro activation [9,34,66,80], and it remains to be determined whether there are also species-dependent differences in Mp function during in vivo tissue repair.”
From Macrophage phenotypes during tissue repair (2013) · cites this paper as [9]
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- Authors
- Mosser, D. M., & Edwards, J. P.
- Affiliations
- University of Maryland, College Park (US)
- Corresponding author
- Mosser, David M.
- Venue
- Nature Reviews Immunology
- Published
- 2008-12
- Volume
- 8
- Issue
- 12
- Pages
- 958-969
- ISSN
- 1474-1733 (ISSN-L)
- 1474-1741
- PMC
- PMC2724991
- OpenAlex
- W2054472394
- Cited by
- 9,144
- References
- 121
- Citations by year
- 2026: 308 · 2025: 470 · 2024: 582 · 2023: 589 · 2022: 661 · 2021: 706 · 2020: 680 · 2019: 592 · 2018: 534 · 2017: 592 · 2016: 549 · 2015: 587 · 2014: 586 · 2013: 534 · 2012: 461
- FWCI
- 17.23
- Type
- journal article
- Language
- en
- Update notices
- erratum (2010-06-01)
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Nature reviews. Immunology (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- Funders
- National Institutes of Health
- National Institute of Allergy and Infectious Diseases (R21AI055576, R01AI055576, AI 55576, R01AI049388, AI49388, R01AI049383)
- Topics
- Life Sciences › Immunology and Microbiology › Immunology › Immune cells in cancer
- Life Sciences › Immunology and Microbiology › Immunology › Phagocytosis and Immune Regulation
- Health Sciences › Medicine › Cardiology and Cardiovascular Medicine › Cardiac Fibrosis and Remodeling
- Keywords
- macrophage activation, macrophage plasticity, homeostasis, host defense, wound healing, immune regulation, macrophage heterogeneity
- MeSH terms
- Animals, Humans, Immunity, Innate, Inflammation, Macrophage Activation, Macrophages, Wound Healing, Cytokines
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