Lipid-associated macrophages control metabolic homeostasis in a Trem2-dependent manner
Open accessAbstract
Immune cells residing in white adipose tissue have been highlighted as important factors contributing to the pathogenesis of metabolic diseases, but the molecular regulators that drive adipose tissue immune cell remodeling during obesity remain largely unknown. Using index and transcriptional single-cell sorting, we comprehensively map all adipose tissue immune populations in both mice and humans during obesity. We describe a novel and conserved Trem2+ lipid-associated macrophage (LAM) subset and identify markers, spatial localization, origin, and functional pathways associated with these cells. Genetic ablation of Trem2 in mice globally inhibits the downstream molecular LAM program, leading to adipocyte hypertrophy as well as systemic hypercholesterolemia, body fat accumulation, and glucose intolerance. These findings identify Trem2 signaling as a major pathway by which macrophages respond to loss of tissue-level lipid homeostasis, highlighting Trem2 as a key sensor of metabolic pathologies across multiple tissues and a potential therapeutic target in metabolic diseases.
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MASH is associated with the emergence of LAMs that highly express Trem2 along with Cd9, Gpnmb, and Spp1.
“MASH is associated with the emergence of lipid-associated macrophages (LAM) that highly express Trem2 along with Cd9, Gpnmb, and Spp1 (3–5)”
From Lipid-associated macrophages' promotion of fibrosis resolution during MASH regression requires TREM2 (2024) · cites this paper as [4]
The Results section, “Lipid-associated macrophages are characterized by Trem2 expression in mice and humans” (page not provided), identifies LAMs in obese adipose tissue as expressing Trem2 and Cd9 and reports LAM-like macrophages in mouse liver under high-fat-diet conditions. The paper does not establish their emergence specifically in MASH or identify Gpnmb as part of the reported LAM signature; it mentions Spp1 as differing among macrophage subsets, not as a confirmed high-expression LAM marker. (LLM, conf=0.88)
Experimental — accuracy not yet measuredThe majority of murine studies suggest LAMs/LAM-like KCs play a role in tissue repair, contributing to dead cell and lipid clearance, reducing lipotoxic stress, and preventing uncontrolled inflammation.
“While the majority of murine studies to date have suggested a role for LAMs/LAM-like KCs in tissue repair, contributing to dead cell and lipid clearance, thus reducing lipotoxic stress, and preventing uncontrolled inflammation (36, 108, 122–126)”
From Targeting macrophages in liver fibrosis (2026) · cites this paper as [126]
The Results (page unavailable) show that adipose LAMs have phagocytosis and lipid-metabolism signatures; the Discussion describes their beneficial containment of lipid-laden, cell-death-prone adipocytes. The paper also detects LAM-like macrophages in mouse liver, but does not establish tissue repair or dead-cell clearance by those liver cells. It suggests LAMs may modulate inflammation, rather than showing that they prevent uncontrolled inflammation or reduce lipotoxic stress. (LLM, conf=0.85)
Experimental — accuracy not yet measuredTREM2 ligands include bacterial wall products, Aβ oligomers, lipoproteins, and anionic lipids such as phospholipids and sphingolipids.
“TREM2 ligands include a variety of molecules, such as bacterial wall products, Aβ oligomers, lipoproteins, and anionic lipids, such as phospholipids and sphingolipids29,32–35”
From TREM2 promotes lung fibrosis via controlling alveolar macrophage survival and pro-fibrotic activity (2025) · cites this paper as [33]
The Results section (‘Lipid-associated macrophages are characterized by Trem2 expression in mice and humans’; page unavailable) describes Trem2 as a lipid receptor, supporting the broad lipid-related part of the claim. Our copy does not identify bacterial wall products, Aβ oligomers, lipoproteins, phospholipids, or sphingolipids as TREM2 ligands. (LLM, conf=0.86)
Experimental — accuracy not yet measuredThese included lysosomal enzymes ASAH1, LIPA, CTSA, CTSG, CTSZ, PLA2G15 and PPT1, part of the CLEAR network, as well as PLA2G7.
“Notably, these included the lysosomal enzymes ASAH1, LIPA, CTSA, CTSG, CTSZ, PLA2G15 and PPT1, enzymes part of the Coordinated Lysosomal Expression and Regulation network60, as well as PLA2G7, previously associated with atherosclerotic plaques61 and with the LAM48 and MIMS-foamy microglial states11.”
From Foamy microglia link oxylipins to disease progression in multiple sclerosis (2026) · cites this paper as [48]
The Results section, “Lipid-associated macrophages are characterized by Trem2 expression in mice and humans” (page unavailable), identifies and characterizes the LAM state. But the passages in our copy do not identify PLA2G7 as associated with LAM; a search for PLA2G7/Pla2g7 found no such passage. This supports the LAM identification, not the specific PLA2G7 association attributed to citation 48. (LLM, conf=0.81)
Experimental — accuracy not yet measuredThe foamy, GPNMB+ microglial phenotype in MS lesions bears striking transcriptional similarity to LAMs in obesity, amyloid-responsive microglia in Alzheimer's disease and foam cells in atherosclerosis.
“The foamy, GPNMB+ microglial phenotype in MS lesions bears striking transcriptional similarity to LAMs in obesity, amyloid-responsive microglia in Alzheimer’s disease and foam cells in atherosclerosis18,46,48,50,51.”
From Foamy microglia link oxylipins to disease progression in multiple sclerosis (2026) · cites this paper as [48]
The Results section reports that obesity-associated LAMs have a highly similar gene profile to Alzheimer’s disease-associated microglia, and the Discussion also describes similarity to aortic macrophages in atherosclerosis (page unavailable in our copy). The paper does not examine foamy GPNMB+ microglia in MS lesions or compare their transcriptional profile with those cell types. (LLM, conf=0.88)
Experimental — accuracy not yet measured
The disease-associated microglial phenotype is shared across aging and neurological and non-neurological diseases with different causes, including Alzheimer's disease, amyotrophic lateral sclerosis, atherosclerosis and obesity.
“The disease-associated phenotype of microglia seems to be shared by aging, as well as neurological and non-neurological diseases with different etiology, e.g. AD and amyotrophic lateral sclerosis (Holtman et al., 2015; Keren-Shaul et al., 2017), atherosclerosis and obesity, characterized by lipid overload (Jaitin et al., 2019; Martinet et al., 2019).”
From Alzheimer's-associated PU.1 expression levels regulate microglial inflammatory response (2021) · cites this paper as Jaitin et al., 2019
There was considerable overlap between the Micro_D profile and previously reported microglial states, such as MIMS-foamy, amyloid-associated microglia in Alzheimer's disease and lipid-associated macrophages in obesity.
“There was considerable overlap between the Micro_D profile and previously reported microglial states, such as the MIMS-foamy state reported in ref. 11, amyloid-associated microglia in Alzheimer’s disease43,46,47 and lipid-associated macrophages (LAMs) in obesity48 (Extended Data Fig. 8).”
From Foamy microglia link oxylipins to disease progression in multiple sclerosis (2026) · cites this paper as [48]
In adipose tissue, TREM2 sustains a population of lipid-associated macrophages (LAMs) that prevent dysmetabolism engendered by a high-fat diet.
“In the adipose tissue, TREM2 sustains the presence of a population of lipid-associated macrophages (LAMs) that prevent the dysmetabolism engendered by a high-fat diet (Jaitin et al., 2019).”
From TREM2 Modulation Remodels the Tumor Myeloid Landscape Enhancing Anti-PD-1 Immunotherapy (2020) · cites this paper as Jaitin et al., 2019
Lipid-associated macrophages have recently been identified in mouse and human adipose tissue, where TREM2 expression was found to regulate lipid homeostasis.
“lipid-associated macrophages have recently been identified in mouse and human adipose tissue, where TREM2 expression was found to regulate lipid homeostasis [57].”
From TREM2/PLCγ2 signalling in immune cells: function, structural insight, and potential therapeutic modulation (2021) · cites this paper as [57]
These lipid-associated macrophages had a similar transcriptional profile to DAM.
“These cells had a similar transcriptional profile to DAM [57]”
From TREM2/PLCγ2 signalling in immune cells: function, structural insight, and potential therapeutic modulation (2021) · cites this paper as [57]
The mouse disease-associated microglia transcriptome profile may not be fully present in Alzheimer's disease brain, because human microglia show a distinct Alzheimer's-related gene signature.
“although the mouse DAM transcriptome profile may not be fully present in AD brain, as human microglia show a distinct AD-related gene signature (Jaitin et al., 2019; Mathys et al., 2019).”
From Alzheimer's-associated PU.1 expression levels regulate microglial inflammatory response (2021) · cites this paper as Jaitin et al., 2019
The Results and Discussion (page not provided) describe a DAM-like signature shared across macrophages and say it is highly similar to signatures in the brains of mice and humans with neurodegenerative disease. Our copy does not assess whether the mouse DAM transcriptome is fully present in Alzheimer’s disease brain or report a distinct Alzheimer’s-related signature in human microglia (searched for those comparisons). High similarity does not rule out differences, so this is not a contradiction. (LLM, conf=0.73)
Experimental — accuracy not yet measuredIn experimental liver fibrosis in murine models, TREM2-deficient macrophages failed to acquire reparative phenotypes, showed impaired efferocytosis, and exacerbated fibrogenesis.
“However, in experimental liver fibrosis in murine models, TREM2-deficient macrophages failed to acquire reparative phenotypes, showed impaired efferocytosis, and exacerbated fibrogenesis (36, 122–126).”
From Targeting macrophages in liver fibrosis (2026) · cites this paper as [126]
The Results section reports that Trem2-deficient macrophages failed to acquire a lipid-associated program in obese adipose tissue. The copy does not address experimental liver fibrosis, reparative macrophage phenotypes in that setting, efferocytosis, or fibrogenesis (searched: liver fibrosis, fibrogenesis, reparative macrophages, efferocytosis). (LLM, conf=0.78)
Experimental — accuracy not yet measuredCultured progranulin-deficient macrophages exhibited increased foam cell formation compared to wild type macrophages.
“The finding of lipid droplet containing microglia in GRN−/− mice is corroborated by a recent study which showed that cultured progranulin-deficient macrophages exhibited increased foam cell formation, i.e. lipid droplet accumulation, compared to wild type macrophages49.”
From Lipid-droplet-accumulating microglia represent a dysfunctional and proinflammatory state in the aging brain (2020) · cites this paper as [49]
The Results section on Trem2-dependent macrophage remodeling reports reduced lipid uptake and storage in Trem2-deficient macrophages, not increased foam-cell formation in progranulin-deficient cultures. Searches for progranulin, GRN, and foam cells found no passage in our copy comparing cultured progranulin-deficient macrophages with wild-type macrophages; page numbers were not available. (LLM, conf=0.76)
Experimental — accuracy not yet measuredMonocyte-derived/bone marrow-dependent macrophages in NASH livers share striking similarities with macrophages in related extrahepatic compartments such as the bone marrow or adipose tissue, including a subset of lipid-associated macrophages with a unique metabolically activated phenotype.
“These monocyte-derived/bone marrow-dependent macrophages in NASH livers share striking similarities with macrophages in related extrahepatic compartments such as the bone marrow or adipose tissue,72,73 including a subset of “lipid-associated macrophages” with a unique metabolically activated phenotype.74”
From Understanding the cellular interactome of non-alcoholic fatty liver disease (2022) · cites this paper as [73]
Monocyte-derived macrophages replace Kupffer cells that normally clear iron, causing iron overload, oxidative stress, and activation of HSCs into myofibroblasts.
“Second, monocyte-derived macrophages replace liver resident Kupffer cells that normally clear iron, resulting in iron overload, oxidative stress, and activation of HSCs into myofibroblasts (70, 74-77).”
From Found in translation—Fibrosis in metabolic dysfunction–associated steatohepatitis (MASH) (2023) · cites this paper as [76]
In fatty liver, lipid-associated macrophages have a unique ability to metabolize lipids, but their contribution to fibrosis is unknown.
“In fatty liver, a novel subset of lipid-associated macrophages (LAMs) display a unique ability to metabolize lipids, but their contribution to fibrosis is unknown (70, 76).”
From Found in translation—Fibrosis in metabolic dysfunction–associated steatohepatitis (MASH) (2023) · cites this paper as [76]
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- Authors
- Jaitin, D. A., Adlung, L., Thaiss, C. A., Weiner, A., Li, B., Descamps, H., Lundgren, P., Bleriot, C., Liu, Z., Deczkowska, A., Keren-Shaul, H., David, E., Zmora, N., Eldar, S. M., Lubezky, N., Shibolet, O., Hill, D. A., Lazar, M. A., Colonna, M., Ginhoux, F., Shapiro, H., Elinav, E., & Amit, I.
- Affiliations
- Weizmann Institute of Science (IL)
- University of Pennsylvania (US)
- Shanghai Jiao Tong University (CN)
- Tel Aviv University (IL)
- Tel Aviv Sourasky Medical Center (IL)
- Children's Hospital of Philadelphia (US)
- Washington University in St. Louis (US)
- Agency for Science, Technology and Research (SG)
- Corresponding author
- Elinav, Eran
- Amit, Ido
- ORCID
- Jaitin, Diego Adhemar
- Adlung, Lorenz
- Thaiss, Christoph A.
- Weiner, Assaf
- Li, Baoguo
- Descamps, Hélène
- Lundgren, Patrick
- Liu, Zhaoyuan
- Deczkowska, Aleksandra
- David, Eyal
- Zmora, Niv
- Eldar, Shai Meron
- Lubezky, Nir
- Shibolet, Oren
- Hill, David A.
- Lazar, Mitchell A.
- Colonna, Marco
- Ginhoux, Florent
- Shapiro, Hagit
- Elinav, Eran
- Amit, Ido
- Venue
- Cell
- Publisher
- Elsevier BV
- Published
- 2019-07
- Volume
- 178
- Issue
- 3
- Pages
- 686-698.e14
- ISSN
- 0092-8674 (ISSN-L)
- PMC
- PMC7068689
- OpenAlex
- W2955261360
- Cited by
- 1,377
- References
- 61
- Citations by year
- 2026: 243 · 2025: 244 · 2024: 229 · 2023: 218 · 2022: 170 · 2021: 152 · 2020: 97 · 2019: 20 · 2018: 1
- FWCI
- 83.54
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- Cell (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- Funders
- Chan Zuckerberg Initiative
- HHMI
- European Research Council (724471-HemTree2.0)
- Thompson Family Foundation
- MRA (509044)
- Foundation (703/15)
- Ernest and Bonnie Beutler Research Program for Excellence in Genomic Medicine
- Helen and Martin Kimmel
- NeuroMac DFG/Transregional Collaborative Research Center
- International Progressive MS Alliance
- NMSS (PA-1604-08459)
- Adelis Foundation
- Wolfson Foundation
- NIH (DK116668, DK49780)
- CHOP
- The Center for Applied Genomics
- Children's Hospital of Philadelphia
- Human Metabolic Tissue Bank
- Penn Diabetes Research Center (P30 NIH DK19525, P30-DK-019525)
- Edward Mallinckrodt, Jr. Foundation
- Agilent
- Global Probiotics Council
- PennCHOP Microbiome Program
- Penn Institute for Immunology
- Penn Center for Molecular Studies in Digestive and Liver Diseases (P30-DK-050306)
- Penn Skin Biology and Diseases Resource-based Center (P30-AR-069589)
- Gilead Sciences International Research Scholars Program in Liver Disease
- The V.R. Schwartz Research Fellow Chair
- Y. and R. Ungar
- Leona M. and Harry B. Helmsley Charitable Trust
- J.L. and V. Schwartz
- D.L. Schwarz
- Israel Science Foundation
- Helmholtz Foundation
- Bill and Melinda Gates Foundation
- Topics
- Health Sciences › Medicine › Epidemiology › Adipokines, Inflammation, and Metabolic Diseases
- Health Sciences › Medicine › Physiology › Adipose Tissue and Metabolism
- Life Sciences › Immunology and Microbiology › Immunology › Immune cells in cancer
- Keywords
- TREM2, lipid-associated macrophages, white adipose tissue, obesity, adipose tissue macrophages, metabolic homeostasis, adipocyte hypertrophy, hypercholesterolemia, impaired glucose tolerance, single-cell transcriptomics, macrophage origin, metabolic disorders
- MeSH terms
- Animals, Humans, Lipids, Macrophages, Membrane Glycoproteins, Mice, Inbred C57BL, Monocytes, Obesity, Receptors, Immunologic, Signal Transduction, Glucose Intolerance, Mice, Knockout, Intra-Abdominal Fat, Lipid Metabolism, Mice, Adipose Tissue, White, Single-Cell Analysis, Diet, High-Fat
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