Guanghui Liu, Ph.D, Professor

​Principal Investigator

State Key Laboratory of Membrane Biology, Institute of ZoologyCAS

Tel: 010-64807583

Zip code: 100101

Research Areas

1. Decoding the genetic and epigenetic mechanisms of human aging

2. Stem cell-based disease mechanism research and precision medicine on human diseases

3. Development and application of new gene-editing methods and gene therapy technologies


1997-2002, B.S., Peking University Health Science Center

2002-2007, Ph.D., Institute of Biophysics, Chinese Academy of Sciences



2007-2009, Research associate, Dept. of Cell and Molecular Biology, The Scripps Research Institute

2009-2012, Research associate, Gene expression laboratory, The Salk Institute for Biological Studies

2012-2021, Professor, Institute of Biophysics, Chinese Academy of Sciences

2019-, Professor, Institute of Zoology, Chinese Academy of Sciences

Honors & Distinctions

2021, Xplorer Prize

2021, Zhongguancun Award for Outstanding Youth in Beijing

2020, the China's Top 10 Scientific Advances of 2020

2020, the China's Top 10 Advances in Life Sciences of 2020

2020, the Outstanding Scientific and Technological Achievement Award issued by Chinese Academy of Sciences

2020, the State Council Government Special allowances 

2018, the China's Top 10 Advances in Life Sciences of 2018

2018, the China Youth Science and Technology Award Special Award

2018, Tan Jiazhen Prize in Life Sciences Innovation

2017, the first Geriatric Outstanding Contribution Award

2017, Shulan Medical Prize


Selected Publications:

  1. Ma S, Wang S, Ye Y, Ren J, Chen R, Li W, Li J, Zhao L, Zhao Q, Sun G, Jing Y, Zuo Y, Xiong M, Yang Y, Wang Q, Lei J, Sun S, Long X, Song M, Yu S, Chan P, Wang J, Zhou Q, Belmonte JCI, Qu J*, Zhang W*, Liu GH*. Heterochronic parabiosis induces stem cell revitalization and systemic rejuvenation across aged tissues. Cell Stem Cell. 2022. 29(6):990-1005.e10.

  2. Liu Z, Ji Q, Ren J, Yan P, Wu Z, Wang S, Sun L, Wang Z, Li J, Sun G, Liang C, Sun R, Jiang X, Hu J, Ding Y, Wang Q, Bi S, Wei G, Cao G, Zhao G, Wang H, Zhou Q, Belmonte JCI, Qu J*, Zhang W*, Liu GH*. Large-Scale Chromatin Reorganization Reactivates Placenta-Specific Genes that Drive Cellular Aging. Developmental Cell. 2022. S1534-5807(22)00332-X.

  3. Zhao H, Ji Q, Wu Z, Wang S, Ren J, Yan K, Wang Z, Hu J, Chu Q, Hu H, Cai Y, Wang Q, Huang D, Ji Z, Li J, Belmonte JCI, Song M*, Zhang W*, Qu J*, Liu GH*. Destabilizing heterochromatin by APOE mediates senescence. Nature Aging. 2022. 2, 303–316.

  4. Liang C, Ke Q, Liu Z, Ren J, Zhang W, Hu J, Wang Z, Chen H, Xia K, Lai X, Wang Q, Yang K, Li W, Wu Z, Wang C, Yan H, Jiang X, Ji Z, Ma M, Long X, Wang S, Wang H, Sun H, Belmonte JCI, Qu J*, Xiang AP*, Liu GH*. BMAL1 moonlighting as a gatekeeper for LINE1 repression and cellular senescence in primates. Nucleic Acids Research. 2022. 50(6):3323-3347.  

  5. Kang W, Jin T, Zhang T, Ma S, Yan H, Liu Z, Ji Z, Cai Y, Wang S, Song M, Ren J, Hu B, Zhou Q, Zhang W*, Qu J*, Bao Y*, Liu GH*. Regeneration Roadmap: database resources for regenerative biology. Nucleic Acids Research. 2022. 50(D1):D1085-D1090. 

  6. Yan H, Wang R, Ma S, Huang D, Wang S, Ren J, Lu C, Chen X, Zhang W*, Qu J*, Zhou Y*, Liu GH*. Lineage Landscape: A comprehensive database that records lineage commitment across species. Nucleic Acids Research. 2022. doi: 10.1093/nar/gkac951. 

  7. Liu ZP, Li W, Geng L, Sun L, Wang Q, Yu Y, Yan P, Liang C, Ren J, Song M, Zhao Q, Lei J, Cai Y, Li J, Yan K, Wu Z, Chu Q, Li J, Wang S, Li C, Han JD, Hernandez-Benitez R, Shyh-Chang Ng, Izpisua Belmonte JC, Zhang WQ*, Qu J*, Liu GH*. Cross-species metabolomic analysis identifies uridine as a potent regeneration promoting factor. Cell Discovery. 2022. 1;8(1):6.

  8. Shan H, Geng L, Jiang X, Song M, Wang J, Liu Z, Zhuo X, Wu Z, Ji Z, Wang S, Chan P, Qu J*, Zhang W*, Liu GH*. Large-Scale Chemical Screen Identifies Gallic Acid as a Geroprotector for Human Stem Cells. Protein Cell. 2022. doi: 10.1007/s13238-021-00872-5.

  9. Yuan B, Zhou X, Suzuki K, Ramos-Mandujano G, Wang M, Tehseen M, Cortés-Medina LV, Moresco JJ, Dunn S, Hernandez-Benitez R, Hishida T, Kim NY, Andijani MM, Bi C, Ku M, Takahashi Y, Xu J, Qiu J, Huang L, Benner C, Aizawa E, Qu J, Liu GH, Li Z, Yi F, Ghosheh Y, Shao C, Shokhirev M, Comoli P, Frassoni F, Yates JR 3rd, Fu XD, Esteban CR, Hamdan S, Li M, Izpisua Belmonte JC. Wiskott-Aldrich syndrome protein forms nuclear condensates and regulates alternative splicing. Nature Communications. 2022. 13(1):3646.

  10. Hishida T, Yamamoto M, Hishida-Nozaki Y, Shao C, Huang L, Wang C, Shojima K, Xue Y, Hang Y, Shokhirev M, Memczak S, Sahu SK, Hatanaka F, Ros RR, Maxwell MB, Chavez J, Shao Y, Liao HK, Martinez-Redondo P, Guillen-Guillen I, Hernandez-Benitez R, Esteban CR, Qu J, Holmes MC, Yi F, Hickey RD, Garcia PG, Delicado EN, Castells A, Campistol JM, Yu Y, Hargreaves DC, Asai A, Reddy P, Liu GH, Izpisua Belmonte JC. In vivo partial cellular reprogramming enhances liver plasticity and regeneration. Cell Reports. 2022. 39(4):110730.

  11. Wang W, Zheng Y, Sun S, Li W, Song M, Ji Q, Wu Z, Liu Z, Fan Y, Liu F, Li J, Esteban CR, Wang S, Zhou Q, Belmonte JCI, Zhang W*, Qu J*, Tang F*, Liu GH*. A Genome-wide CRISPR-Based Screen Identifies KAT7 as a Senescence Driver. Science Translational Medicine. 2021.13, Jan 6;13(575):eabd2655.

  12. Zou Z, Long X, Zhao Q, Zheng Y, Song M, Ma S, Wang S, He Y, Esteban CR, Huang J, Yu N, Chan P, Belmonte JCI, Zhang W*, Qu J*, Liu GH*. A single-cell transcriptomic atlas of human skin aging. Developmental Cell. 2021. 56(3):383-397 (Cover story)

  13. Liang C, Liu Z, Song M, Li W, Wu Z, Wang Z, Wang Q, Wang S, Yan K, Sun L, Hishida T, Cai Y, Izpisua Belmonte JC, Guillen P, Chan P, Zhou Q, Zhang W*, Qu J*, Liu GH*. Stabilization of Heterochromatin by CLOCK Promotes Stem Cell Rejuvenation and Cartilage Regeneration. Cell Research. 2021. 31(2):187-205.

  14. Wang S, Yao X, Ma S, Ping Y, Fan Y, Sun S, He Z, Shi Y, Sun L, Xiao S, Song M, Cai J, Li J, Tang R, Zhao L, Wang C, Wang Q, Zhao L, Hu H, Liu X, Sun G, Chen L, Pan G, Chen H, Li Q, Zhang P, Xu Y, Feng H, Zhao GG, Wen T, Yang Y, Huang X, Li W, Liu Z, Wang H, Wu H, Hu B, Ren Y, Zhou Q, Qu J*, Zhang W*, Liu GH*, Bian XW*. A single-cell transcriptomic landscape of the lungs of patients with COVID-19. Nature Cell Biology. 2021. 23(12):1314-1328 (Leading contact)

  15. Zhang H, Li J, Ren J, Sun S, Ma S, Zhang W, Yu Y, Cai Y, Yan K, Li W, Hu B, Chan P, Zhao GG, Belmonte JC, Zhou Q, Qu J*, Wang S*, Liu GH*. Single-nucleus transcriptomic landscape of primate hippocampal aging. Protein Cell. 2021. 12(9):695-716. (Cover story)

  16. Aging Atlas Consortium, Zhang W*, Jing Qu*, Bao Y*, Liu GH*. Aging Atlas: a multi-omics database for aging biology. Nucleic Acids Research. 2021. 8;49(D1):D825-D830.

  17. Li H, Wei X, Zhou L, Zhang W, Wang C, Guo Y, Li D, Chen J, Liu T, Zhang Y, Ma S, Wang C, Tan F, Xu J, Liu Y, Yuan Y, Chen L, Wang Q, Qu J, Shen Y, Liu S, Fan G, Liu L, Liu X, Hou Y, Liu GH*, Gu Y*, Xu X*. Dynamic cell transition and immune response landscapes of axolotl limb regeneration revealed by single-cell analysis. Protein Cell.  2021. 12(1):57-66.

  18. Li CJ, Xiao Y, Sun YC, He WZ, Liu L, Huang M, He C, Huang M, Chen KC, Hou J, Feng X, Su T, Guo Q, Huang Y, Peng H, Yang M, Liu GH, Luo XH. Senescent immune cells release Grancalcin to promote skeletal aging. Cell Metabolism. 2021. 33(10):1957-1973.

  19. Wang S, Zheng Y, Li J, Yu Y, Zhang W, Song M, Liu Z, Min Z, Hu H, Jing Y, He X, Sun L, Esteban C, Chan P, Qiao J, Zhou Q, Belmonte JC*, Jing Qu*, Tang F*, Liu GH*. Single-Cell Transcriptomic Atlas of Primate Ovarian Aging. Cell. 2020. 180(3):585-600. (Cover story)

  20. Ma S, Sun S, Geng L, Song M, Wang W, Ye Y, Ji Q, Zou Z, Wang S, He X, Li W, Rodriguez Esteban C, Long X, Guo G, Chan P, Zhou Q, Belmonte JC*, Zhang W*, Qu J*, Liu GH*. Caloric Restriction Reprograms the Single-Cell Transcriptional Landscape of Rattus Norvegicus Aging. Cell. 2020. 180(5):984-1001.

  21. Ma S, Sun S, Li J, Fan Y, Qu J, Sun L, Wang S, Zhang Y, Yang S, Liu Z, Wu Z, Zhang S, Wang Q, Zheng A, Duo S, Yu Y, Belmonte JCI, Chan P, Zhou Q, Song M*, Zhang W*, Liu GH*. Single-Cell Transcriptomic Atlas of Primate Cardiopulmonary Aging. Cell Research. 2020. 10;1-18.

  22. Yan P, Liu Z, Song M, Wu Z, Xu W, Li K, Ji Q, Wang S, Liu X, Yan K, Esteban CR, Ci W, Izpisua Belmonte JC, Xie W, Ren J, Zhang W*, Sun Q*, Qu J*, Liu GH*. Genome-wide R-loop landscapes during cell differentiation and reprogramming. Cell Reports. 2020. 32(1):107870

  23. Wang F, Zhang W, Yang Q, Kang Y, Fan Y, Wei J, Liu Z, Dai S, Li H, Li Z, Xu L, Chu C, Qu J, Si C, Ji W*, Liu GH*, Long C*, Niu Y*. Generation of a Hutchinson-Gilford progeria syndrome monkey model by base editing. Protein Cell. 2020. 11(11):809-824. (Cover story)

  24. Zheng Y, Liu X, Le W, Xie L, Li H, Wen W, Wang S, Ma S, Huang Z, Ye J, Shi W, Ye Y, Liu Z, Song M, Zhang W, Belmonte JCI, Xiao C, Qu J*, Wang H*, Liu GH*, Su W*. A human circulating immune cell landscape in aging and COVID-19. Protein Cell. 2020. 192:111357.

  25. Wang S, Min Z, Ji Q, Geng L, Su Y, Liu Z, Hu H, Wang L, Zhang W, Suzuiki K, Huang Y,Zhang P, Tang T, Qu J*, Yu Y*, Liu GH*, Qiao J*. Rescue of Premature Aging Defects in Cockayne Syndrome Stem Cells by CRISPR/Cas9-Mediated Gene Correction. Protein Cell. 2020. 11(1):1-22.

  26. Yan P, Li Q, Wang L, Lu P, Suzuki K, Liu Z, Lei J, Li W, Ren R, He X, Wang S, Ding J, Chan P, Zhang W, Song M, Belmonte JC, Qu J*, Tang F*, Liu GH*. FOXO3-Engineered Human ESC-Derived Vascular Cells Promote Vascular Protection and Regeneration. Cell Stem Cell. 2019. 24(3), 447-461. (Cover story)

  27. Deng L, Ren R, Liu Z, Song M, Li J, Wu Z, Ren X, Fu L, Li W, Zhang W, Guillen P, Belmonte J, Chan P, Qu J*, Liu GH*. Stabilizing Heterochromatin by DGCR8 Alleviates Senescence and Osteoarthritis. Nature Communications. 2019.10(1):3329.

  28. Cheng F, Wang S, Song M, Liu Z, Liu P, Chan P, Wang Y, Wang L, Zhang W*, Qu J*, Liu GH*. DJ-1 is Dispensable for Human Stem Cell Homeostasis. Protein Cell. 2019. 10(11):846-853.

  29. Fu L, Hu Y, Song M, Liu Z, Zhang W, Yu FX, Wu J, Wang S, Belmonte JC, Chan P, Qu J*, Tang F*, Liu GH*. Upregulation of FOXD1 by YAP Alleviates Senescence and Osteoarthritis. PLoS Biology. 2019. 17(4):e3000201.

  30. Ling C, Liu Z, Song M, Zhang W, Wang S, Liu X, Ma S, Sun S, Fu L, Chu Q, Belmonte JC, Wang Z, Qu J*, Yuan Y*, Liu GH*. Modeling CADASIL Vascular Pathologies with Patient-Derived Induced Pluripotent Stem Cells. Protein Cell. 2019. 10(4):249-271.

  31. Ren X, Hu B, Song M, Ding Z, Dang Y, Liu Z, Zhang W, Ji Q, Ren R, Ding J, Chan P, Jiang C, Ye K, Qu J*, Tang F*, Liu GH*. Maintenance of Nucleolar Homeostasis by CBX4 Alleviates Senescence and Osteoarthritis. Cell Reports. 2019. 26(13):3643-3656.

  32. Geng L, Liu Z, Zhang W*, Li W, Wu Z, Wang W, Ren R, Su Y, Wang P, Ju Z, Chan P, Song M*, Qu J*, Liu GH*. Chemical Screen Identifies a Geroprotective Role of Quercetin in Premature Aging. Protein Cell. 2019. 10(6):417-435.

  33. Zhang W, Wan H, Feng G, Qu J, Wang J, Jing Y, Ren R, Liu Z, Zhang L, Chen Z, Wang S, Zhao Y, Wang Z, Yuan Y, Zhou Q, Li W*, Liu GH*, Hu B*. SIRT6 Deficiency Results in Developmental Retardation in Cynomolgus Monkeys. Nature. 2018. 560(7720):661-665. (Leading contact) 

  34. Wang S, Hu B, Ding Z, Dang Y, Wu J, Li D, Liu X, Xiao B, Zhang W, Ren R, Lei J, Hu H, Chen C, Chan P, Li D, Qu J*, Tang F*, Liu GH*. ATF6 Safeguards Organelle Homeostasis and Cellular Aging in Human Mesenchymal Stem Cells. Cell Discovery. 2018. Jan 5;4:2. 

  35. Fang J, Yang J, Wu X, Zhang G, Li T, Zhang H, Wang CC, Liu GH*, Wang L*. Metformin Alleviates Human Cellular Aging by Upregulating the ER Peroxidase GPx7. Aging Cell. 2018.17(4):e12765. 

  36. Wu Z, Zhang W, Song M, Wang W, Wei G, Li W, Lei J, Huang Y, Sang Y, Chan P, Chen C, Qu J*, Suzuki K*, Belmonte JC*, Liu GH*. Differential Stem Cell Aging Kinetics in Hutchinson–Gilford Progeria Syndrome and Werner Syndrome. Protein Cell. 2018. 9(4):333-350. (Cover story)   

  37. Ren R, Deng L, Xue Y, Suzuki K, Zhang W, Yu Y, Wu J, Sun L, Gong X, Luan H, Yang F, Ju Z, Ren X, Wang S, Tang H, Geng L, Zhang W, Li J, Qiao J, Xu T*, Qu J*Liu GH*. Visualization of Aging-associated Chromatin Alterations with an Engineered Tale System. Cell Research. 2017. 27(4):483-504. (Cover story)

  38. Yang J, Li J, Suzuki K, Liu X, Wu J, Zhang W, Ren R, Zhang W, Chan P, Belmonte JC, Qu J*, Tang F*, Liu GH*. Genetic Enhancement in Cultured Human Adult Stem Cells Conferred by a Single Nucleotide Recoding. Cell Research. 2017. 27(9):1178-1181.

  39. Wang L, Yi F, Fu L, Yang J, Wang S, Wang Z, Suzuki K, Sun L, Xu X, Yu Y, Qiao J, Belmonte JC, Yang Z, Yuan Y*, Qu J*, Liu GH*. CRISPR/Cas9-Mediated Targeted Gene Correction in Amyotrophic Lateral Sclerosis Patient iPSCs. Protein Cell. 2017. 8(5):365-378. (Cover story) 

  40. Kubben N, Zhang W, Wang L, Voss T, Yang J, Qu J, Liu GH*, Misteli T*, Repression of the Antioxidant NRF2 Pathway in Premature Aging. Cell. 2016. 165(6):1361-1374.

  41. Pan H, Guan D, Liu X, Li J, Wang L, Wu J, Zhou J, Zhang W, Ren R, Zhang W, Li Y, Yang J, Hao Y, Yuan T, Yuan G, Wang H, Ju Z, Mao Z, Li J, Qu J*, Tang F*, Liu GH*. SIRT6 Safeguards Human Mesenchymal Stem Cells From Oxidative Stress by Coactivating NRF2. Cell Research. 2016. 26(2):190-205.

  42. Suzuki K, Tsunekawa Y, Hernandez-Benitez R, Wu J, Zhu J, Kim EJ, Hatanaka F, Yamamoto M, Araoka T, Li Z, Kurita M, Hishida T, Li M, Aizawa E, Guo S, Chen S, Goebl A, Soligalla RD, Qu J, Jiang T, Fu X, Jafari M, Esteban CR, Berggren WT, Lajara J, Nuñez-Delicado E, Guillen P, Campistol JM, Matsuzaki F, Liu GH, Magistretti P, Zhang K, Callaway EM, Zhang K, Belmonte JC. In Vivo Genome Editing via CRISPR/Cas9 Mediated Homology-Independent Targeted Integration. Nature. 2016. 540(7631):144-149.

  43. Fu L, Xu X, Ren R, Wu J, Zhang W, Yang J, Ren X, Wang S, Zhao Y, Sun L, Yu Y, Wang Z, Yang Z, Yuan Y, Qiao J, Belmonte JC*, Qu J*, Liu GH*. Modeling Xeroderma Pigmentosum Associated Neurological Pathologies with Patients-Derived iPSCs. Protein Cell. 2016.7(3):210-221.

  44. Wang H, Diao D, Shi Z, Zhu X, Gao Y, Gao S, Liu X, Wu Y, Rudolph KL, Liu GH, Li T, Ju Z. SIRT6 Controls Hematopoietic Stem Cell Homeostasis through Epigenetic Regulation of Wnt Signaling. Cell Stem Cell. 2016. 7;18(4):495-507.

  45. Li Y, Zhang W, Chang L, Han Y, Sun L, Guo X, Tang H, Liu Z, Deng H, Ye Y, Ren R, Wang Y, Li J, Qiao J, Qu J*, Zhang W*, Liu GH*. Vitamin C Alleviates Aging Defects in a Stem Cell Model for Werner Syndrome. Protein Cell. 2016. 7(7):478-488. (Cover story)

  46. Zhang W, Li J, Suzuki K, Qu J, Wang P, Zhou J, Liu X, Ren R, Xu X, Ocampo A, Yuan T, Yang J, Li Y, Shi L, Guan D, Pan H, Duan S, Ding Z, Li M, Yi F, Bai R, Wang Y, Chen C, Yang F, Li X, Wang Z, Aizawa E, Goebl A, Soligalla RD, Reddy P, Esteban CR, Tang F*, Liu GH*, Belmonte JC*. A Werner Syndrome Stem Cell Model Unveils Heterochromatin Alterations as a Driver of Human Aging. Science. 2015; 348(6239):1160-1163.

  47. Reddy P, Ocampo A, Suzuki K, Luo J, Bacman SR, Williams SL, Sugawara A, Okamura D, Tsunekawa Y, Wu J, Lam D, Xiong X, Montserrat N, Esteban CR, Liu GH, Sancho-Martinez I, Manau D, Civico S, Cardellach F, Del Mar O'Callaghan M, Campistol J, Zhao H, Campistol JM, Moraes CT, Izpisua Belmonte JC. Selective Elimination of Mitochondrial Mutations in the Germline by Genome Editing. Cell. 2015; 161(3):459-469.

  48. Duan S, Yuan G, Liu X, Ren R, Li J, Zhang W, Wu J, Xu X, Fu L, Li Y, Yang J, Zhang W, Bai R, Yi F, Suzuki K, Gao H, Esteban CR, Zhang C, Belmonte JC, Chen Z, Wang X, Jiang T, Qu J*, Tang F*, Liu GH*. PTEN Deficiency Reprograms Human Neural Stem Cells Towards a Glioblastoma Stem Cell-Like Phenotype. Nature Communications. 2015. 6:10068.


  1. Longevity secret - a pluripotent superpower. Cell Metabolism. 2022. DOI: 10.1016/jNaNet.2022.05.001.

  2. A Stem Cell Aging Framework, from Mechanisms to Interventions. Cell Reports. 2022. 41(3):111451.

  3. The landscape of aging. Science China Life Science. 2022. 2:1-101.

  4. Opening up the black box of human cell plasticity. Innovation. 2022. 3(5):100276.

  5. Epigenetic regulation of aging: implications for interventions of aging and diseases. Signal Transduction and Targeted Therapy. 2022. DOI: 10.1038/s41392-022-01211-8.

  6. Emerging role of RNA m6A modification in aging regulation. Current Medicine. 2022. DOI:10.1007/s44194-022-00009-8.

  7. Potential Approaches for Intervening Aging. National Science Open. 2022. DOI: 10.1360/nso/20220021.

  8. Deciphering aging at three-dimensional genomic resolution. Cell Insight. 2022. DOI:10.1016/j.cellin.2022.100034.

  9. Rejuvenation of Tissue Stem Cells by Intrinsic and Extrinsic Factors. Stem Cells Translational Medicine. 2022: 11(3):231-238.

  10. Mitochondrial sirtuins, metabolism, and aging. J Genet Genomics. 2022. 49(4):287-298.

  11. The Aging Epigenome and its Rejuvenation. Nature Reviews Molecular Cell Biology. 2020. 21(3):137-150.

  12. Engineering Tissues and Organs: The Road to the Clinic. Cell. 2020. 181(1):22-23. (Leading Edge Voices)

  13. Single-Cell “OMICS” in Aging: A Young Growing Field. Nature Metabolism. 2020. 2(4): 293-302.

  14. A Beta-Galactosidase Kiss of Death for Senescent Cells. Cell Research. 2020. 30(7):556-557.

  15. Treating Osteoarthritis via Gene Therapy with Rejuvenation Factors. Gene Therapy. 2020. 27(7-8):309-311. 

  16. Protein Quality Control of Cell Stemness. Cell Regeneration. 2020. 12;9(1):22.

  17. Age-Related Cardiopathies Gene Editing. Aging. 2019. 11(5):1327-1328.

  18. Epigenetic Modifications in Cardiovascular Aging and Diseases. Circulation Research. 2018. 123(7):773-786.

  19. Regulation of Stem Cell Aging by Metabolism and Epigenetic Factors. Cell Metabolism. 2017. 26(3):460-474.