Targeting p21Cip1 highly expressing cells in adipose tissue alleviates insulin resistance in obesity
Open accessAbstract
Insulin resistance is a pathological state often associated with obesity, representing a major risk factor for type 2 diabetes. Limited mechanism-based strategies exist to alleviate insulin resistance. Here, using single-cell transcriptomics, we identify a small, critically important, but previously unexamined cell population, p21 Cip1 highly expressing (p21 high) cells, which accumulate in adipose tissue with obesity. By leveraging a p21-Cre mouse model, we demonstrate that intermittent clearance of p21 high cells can both prevent and alleviate insulin resistance in obese mice. Exclusive inactivation of the NF-κB pathway within p21 high cells, without killing them, attenuates insulin resistance. Moreover, fat transplantation experiments establish that p21 high cells within fat are sufficient to cause insulin resistance in vivo. Importantly, a senolytic cocktail, dasatinib plus quercetin, eliminates p21 high cells in human fat ex vivo and mitigates insulin resistance following xenotransplantation into immuno-deficient mice. Our findings lay the foundation for pursuing the targeting of p21 high cells as a new therapy to alleviate insulin resistance.
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Senescent cells in human and mouse adipose tissue express CDKN2A and CDKN1A, have higher SA-β-gal activity, DNA damage foci, proliferation arrest (MKI67), and SASP factors including IL-6, CXCL8, IL-1β, TNF, CCL2, SERPINE1, MMP3, and MMP12.
“Senescent cells in both human and mouse adipose tissue have been characterized by expression of CDKN2A and CDKN1A, higher SA-β-gal activity, DNA damage (γH2AX) foci and arrest of cell proliferation (MKI67), and by expression of SASP factors including IL-6, CXCL8 (also known as IL-8), IL-1β, TNF, CCL2, SERPINE1, MMP3 and MMP12 (refs. 119–122).”
From SenNet recommendations for detecting senescent cells in different tissues (2024) · cites this paper as [122]
The Introduction and Discussion describe p21-high cells in mouse adipose tissue as having higher SA-β-gal activity, reduced proliferation and a SASP; the paper also examines γH2AX-positive telomere-associated foci. It treats p21-high and p16-high cells as partly distinct populations, rather than establishing that senescent cells express both markers. Although it uses human adipose tissue for transplantation, the cited passages do not establish the full marker profile in human tissue or substantiate the entire listed SASP panel or MKI67 specifically. Page numbers were not available in our copy. (LLM, conf=0.87)
Experimental — accuracy not yet measuredCdkn1ahigh and Cdkn2ahigh cells exist as two distinct cell populations in adipose tissue, indicating heterogeneity and complexity of senescent cell populations.
“Importantly, Cdkn1ahigh and Cdkn2ahigh cells exist as two distinct cell populations122,125, indicating the heterogeneity and complexity of senescent cell populations in adipose tissue.”
From SenNet recommendations for detecting senescent cells in different tissues (2024) · cites this paper as [122]
The Results section reports that p21-high (Cdkn1a-high) and p16-high (Cdkn2a-high) cells were distinguishable in mouse visceral adipose tissue after 10 months of high-fat feeding, despite some overlap. The Discussion calls them distinct populations, but explicitly limits that conclusion to at least the context of obesity; the citing sentence does not retain that qualifier. (LLM, conf=0.84)
Experimental — accuracy not yet measuredThe accumulation of p21+ cells has been identified in aged tissues in vivo, and clearance of this population in rodents has been shown to both extend lifespan and improve healthspan.
“The accumulation of p21 positive (p21+) cells has been identified in aged tissues in vivo, and clearance of this population in rodents has been shown to both extend lifespan and improve healthspan [7,8,9].”
From Fluorescence-Guided Spatial Mapping of p21-Expressing Senescent Cells in Aged Liver at Single-Cell Resolution (2026) · cites this paper as [9]
The Introduction says senescent cells accumulate with aging, but does not establish that the accumulating cells are specifically p21-positive. The Results and Discussion report that clearing p21-high cells improves metabolic function in obese mice; they do not report extended lifespan or improved healthspan in aged rodents. Page numbers are unavailable in our copy. (LLM, conf=0.86)
Experimental — accuracy not yet measuredDecondensation occurs in essentially all centromeric satellites in vitro, but only a fraction of centromeric satellites undergo unwinding in vivo.
“Moreover, while decondensation occurs in essentially all centromeric satellites in vitro24, only a fraction of centromeric satellites undergo unwinding in vivo25,26 (Fig. 1 F).”
From Guidelines for minimal information on cellular senescence experimentation in vivo (2024) · cites this paper as [26]
The STAR METHODS passages describe telomere staining and other assays in adipose tissue, not centromeric satellites. Searches for centromeric-satellite unwinding in vivo and decondensation in vitro found no passage addressing the claim; page numbers were not supplied. (LLM, conf=0.78)
Experimental — accuracy not yet measuredSenescence markers beyond p16 include p21, p19, uPAR, and GPNMB.
“Studies have begun to reveal the remarkable heterogeneity of cellular senescence in different tissues, highlighting the importance of cellular senescence markers beyond p16 – including Cdkn1a (p21) [7–10], Cdkn2d (p19) [11], uPAR [12], and glycoprotein non-metastatic melanoma protein B (GPNMB) [13] – and emphasizing the urgent need for a better understanding of senescent cell heterogeneity.”
From The heterogeneity of cellular senescence: insights at the single-cell level (2023) · cites this paper as [10]
p16 is not a specific or sensitive marker for cellular senescence because some p16high cells are not senescent and some senescent cells do not express p16.
“However, p16 might not be a specific or sensitive marker for cellular senescence since not all p16 highly expressing (p16high) cells are necessarily senescent [4] and some senescent cells do not express p16 [5,6].”
From The heterogeneity of cellular senescence: insights at the single-cell level (2023) · cites this paper as [6]
Clearance of p21high cells reduces senescence markers, improves physical function in aged mice, and alleviates metabolic dysfunction in obese mice.
“Remarkably, clearance of p21high cells reduces senescence markers, improves physical function in aged mice [9], and alleviates metabolic dysfunction in obese mice [10].”
From The heterogeneity of cellular senescence: insights at the single-cell level (2023) · cites this paper as [10]
Adipose p21high cells showed key senescence features in the contexts of both aging and obesity.
“Notably, adipose p21high cells showed a number of key senescence features (assessed by imaging flow cytometry) in the contexts of both aging and obesity.”
From The heterogeneity of cellular senescence: insights at the single-cell level (2023) · cites this paper as [10]
Imaging flow cytometry allows protein-level detection of several senescence markers or reporters along with assessment of several senescence features at the single-cell level in a relatively high-throughput manner.
“Another useful technology for examining senescent cells is imaging flow cytometry, which has been used to examine senescent cells in vivo [9,10,36]. The major advantage of this method is that it allows for protein-level detection of several senescence markers or reporters along with assessment of several senescence features at the single-cell level in a relatively high-throughput manner.”
From The heterogeneity of cellular senescence: insights at the single-cell level (2023) · cites this paper as [10]
In adipose tissue, p21high and p16high cells are two distinct cell populations in terms of cell types, tissue location, accumulation kinetics, and physiological roles.
“They showed that p21high and p16high cells are two distinct cell populations in terms of cell types, tissue location, accumulation kinetics, and physiological roles.”
From The heterogeneity of cellular senescence: insights at the single-cell level (2023) · cites this paper as [10]
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- Authors
- Wang, L., Wang, B., Gasek, N. S., Zhou, Y., Cohn, R. L., Martin, D. E., Zuo, W., Flynn, W. F., Guo, C., Jellison, E. R., Kim, T., Prata, L. G. P. L., Palmer, A. K., Li, M., Inman, C. L., Barber, L. S., Al-Naggar, I. M. A., Zhou, Y., Du, W., Kshitiz, Kuchel, G. A., Meves, A., Tchkonia, T., Kirkland, J. L., Robson, P., & Xu, M.
- Affiliations
- University of Connecticut (US)
- Central South University (CN)
- The Jackson Laboratory for Genomic Medicine (US)
- UConn Health (US)
- Mayo Clinic (US)
- Corresponding author
- Xu, Ming
- ORCID
- Venue
- Cell Metabolism
- Publisher
- Elsevier BV
- Published
- 2022-01
- Volume
- 34
- Issue
- 1
- Pages
- 75-89.e8
- ISSN
- 1550-4131 (ISSN-L)
- PMC
- PMC8732323
- OpenAlex
- W3216561293
- Cited by
- 163
- References
- 41
- Citations by year
- 2026: 22 · 2025: 31 · 2024: 42 · 2023: 41 · 2022: 27
- FWCI
- 19.79
- Type
- journal article
- Language
- en
- Update notices
- erratum (2022-01-04)
- Open access
- bronze
- doi.org
- Licence
- www.elsevier.com/tdm/userlicense/1.0/ (tdm)
- www.elsevier.com/legal/tdmrep-license (tdm)
- www.elsevier.com/open-access/userlicense/1.0/ (vor, 368 d)
- doi.org/10.15223/policy-017 (stm-asf)
- doi.org/10.15223/policy-037 (stm-asf)
- doi.org/10.15223/policy-012 (stm-asf)
- doi.org/10.15223/policy-029 (stm-asf)
- doi.org/10.15223/policy-004 (stm-asf)
- Locations
- Cell Metabolism (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- The Mouseion at the JAXlibrary (Jackson Laboratory) (submittedVersion)
- Funders
- American Foundation for Aging Research
- University of Connecticut (T32HG010463)
- National Institutes of Health (K08CA215105, P01AG062413, P30AG067988, R33AG061456, R37AG013925)
- Noaber Foundation
- NIA (AG063528, AG066679, AG068860)
- Mayo Clinic
- Glenn Foundation for Medical Research
- Topics
- Health Sciences › Medicine › Epidemiology › Adipokines, Inflammation, and Metabolic Diseases
- Health Sciences › Medicine › Physiology › Adipose Tissue and Metabolism
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Peroxisome Proliferator-Activated Receptors
- Keywords
- p21, insulin resistance, obesity, adipose tissue, senolytic therapy, dasatinib, quercetin, NF-κB signaling, type 2 diabetes, fat transplantation, single-cell transcriptomics
- MeSH terms
- Adipose Tissue, Animals, Diabetes Mellitus, Type 2, Insulin Resistance, Mice, Inbred C57BL, Obesity, Cellular Senescence, Mice, Diet, High-Fat
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