自噬与脂肪组织棕色化及棕色脂肪活性的调节：肥胖干预新靶点

doi:10.12677/pi.2026.156055

期刊菜单

自噬与脂肪组织棕色化及棕色脂肪活性的调节：肥胖干预新靶点
Autophagy and the Regulation of Adipose Tissue Browning and Brown Adipose Activity: A Novel Target for Obesity Intervention

DOI: 10.12677/pi.2026.156055, PDF, 科研立项经费支持
作者: 劳嘉怡, 刘康^*：中国药科大学中药学院中药药理与中医药学系，江苏南京
关键词: 线粒体自噬；脂噬；分子伴侣介导的自噬；脂肪组织；棕色化；肥胖；Mitophagy； Lipophagy； Chaperone-Mediated Autophagy； Adipose Tissue； Browning； Obesity

摘要: 肥胖的核心机制为能量摄入与消耗的长期失衡。促进白色脂肪组织棕色化并维持棕色脂肪组织活性以增强能量消耗，已成为肥胖及相关代谢性疾病干预的重要策略。自噬作为细胞内的重要降解途径，通过线粒体自噬、脂噬以及分子伴侣介导的自噬等形式调控脂肪组织可塑性，相关机制研究近年来受到广泛关注。线粒体自噬主要经由PINK1/Parkin通路以时相依赖性方式调控线粒体质量，影响脂肪棕色化进程与棕色脂肪组织活性维持。脂噬受下丘脑POMC神经元–外周神经轴调控，与溶酶体酸性脂肪酶介导的脂解途径协同促进脂滴降解，为线粒体β-氧化提供底物，支持适应性产热。分子伴侣介导的自噬依赖LAMP2A以温度依赖性方式选择性降解产热调控因子，实现对产热程序的调控。上述三种形式共同构成自噬调控脂肪组织产热的分子网络，为开发靶向自噬的肥胖及相关代谢性疾病精准干预策略提供了理论框架。

Abstract: Obesity results from a chronic imbalance between energy intake and expenditure. Promoting white adipose tissue browning and maintaining brown adipose tissue activity to enhance energy expenditure has emerged as an important therapeutic strategy for obesity and related metabolic diseases. Autophagy, a core intracellular degradation pathway, regulates adipose tissue plasticity through three selective forms: mitophagy, lipophagy, and chaperone-mediated autophagy (CMA), and the underlying mechanisms have attracted considerable attention in recent years. Mitophagy modulates mitochondrial quality mainly via the PINK1/Parkin pathway in a time-dependent manner, thereby influencing white adipose tissue browning and brown adipose tissue activity maintenance. Lipophagy is regulated by the hypothalamic POMC neuron-peripheral neural axis and cooperates with lysosomal acid lipase-mediated lipolytic pathways to promote lipid droplet degradation, providing substrates for mitochondrial β-oxidation to support adaptive thermogenesis. CMA relies on LAMP2A to selectively degrade thermoregulatory factors in a temperature-dependent manner and regulates thermogenic programs. Together, these three selective autophagic forms constitute a molecular network that regulates thermogenesis in adipose tissue, providing a theoretical framework for developing precision therapeutic strategies targeting autophagy in obesity and associated metabolic diseases.

文章引用：劳嘉怡, 刘康. 自噬与脂肪组织棕色化及棕色脂肪活性的调节：肥胖干预新靶点[J]. 药物资讯, 2026, 15(6): 515-529. https://doi.org/10.12677/pi.2026.156055

参考文献

[1]	Chen, K., Shen, Z., Gu, W., Lyu, Z., Qi, X., Mu, Y., et al. (2023) Prevalence of Obesity and Associated Complications in China: A Cross‐Sectional, Real‐World Study in 15.8 Million Adults. Diabetes, Obesity and Metabolism, 25, 3390-3399. https://doi.org/10.1111/dom.15238
[2]	World Health Organization (2025) Obesity and Overweight: Fact Sheet. https://www.who.int/news-room/fact-sheets/detail/obesity-and-overweight
[3]	World Obesity Federation (2025) World Obesity Atlas 2025. World Obesity Federation.
[4]	Kerr, J.A., Patton, G.C., Cini, K.I., Abate, Y.H., Abbas, N., Abd Al Magied, A.H.A., et al. (2025) Global, Regional, and National Prevalence of Child and Adolescent Overweight and Obesity, 1990-2021, with Forecasts to 2050: A Forecasting Study for the Global Burden of Disease Study 2021. The Lancet, 405, 785-812. https://doi.org/10.1016/s0140-6736(25)00397-6
[5]	Xiong, S., Yu, S., Wang, K., Xiong, X., Xia, M., Zeng, G., et al. (2023) Dietary Apigenin Relieves Body Weight and Glycolipid Metabolic Disturbance via Pro‐Browning of White Adipose Mediated by Autophagy Inhibition. Molecular Nutrition & Food Research, 67, e2200763. https://doi.org/10.1002/mnfr.202200763
[6]	Martinez-Lopez, N., Garcia-Macia, M., Sahu, S., Athonvarangkul, D., Liebling, E., Merlo, P., et al. (2016) Autophagy in the CNS and Periphery Coordinate Lipophagy and Lipolysis in the Brown Adipose Tissue and Liver. Cell Metabolism, 23, 113-127. https://doi.org/10.1016/j.cmet.2015.10.008
[7]	Lu, Y., Fujioka, H., Joshi, D., Li, Q., Sangwung, P., Hsieh, P., et al. (2018) Mitophagy Is Required for Brown Adipose Tissue Mitochondrial Homeostasis during Cold Challenge. Scientific Reports, 8, Article No. 8251. https://doi.org/10.1038/s41598-018-26394-5
[8]	Wu, J., Boström, P., Sparks, L.M., Ye, L., Choi, J.H., Giang, A., et al. (2012) Beige Adipocytes Are a Distinct Type of Thermogenic Fat Cell in Mouse and Human. Cell, 150, 366-376. https://doi.org/10.1016/j.cell.2012.05.016
[9]	Ricquier, D. and Bouillaud, F. (2000) Mitochondrial Uncoupling Proteins: From Mitochondria to the Regulation of Energy Balance. The Journal of Physiology, 529, 3-10. https://doi.org/10.1111/j.1469-7793.2000.00003.x
[10]	Harms, M. and Seale, P. (2013) Brown and Beige Fat: Development, Function and Therapeutic Potential. Nature Medicine, 19, 1252-1263. https://doi.org/10.1038/nm.3361
[11]	Seale, P., Bjork, B., Yang, W., Kajimura, S., Chin, S., Kuang, S., et al. (2008) PRDM16 Controls a Brown Fat/Skeletal Muscle Switch. Nature, 454, 961-967. https://doi.org/10.1038/nature07182
[12]	Scheja, L. and Heeren, J. (2016) Metabolic Interplay between White, Beige, Brown Adipocytes and the Liver. Journal of Hepatology, 64, 1176-1186. https://doi.org/10.1016/j.jhep.2016.01.025
[13]	Cairó, M. and Villarroya, J. (2020) The Role of Autophagy in Brown and Beige Adipose Tissue Plasticity. Journal of Physiology and Biochemistry, 76, 213-226. https://doi.org/10.1007/s13105-019-00708-1
[14]	Jiang, J., Zhou, D., Zhang, A., Yu, W., Du, L., Yuan, H., et al. (2023) Thermogenic Adipocyte-Derived Zinc Promotes Sympathetic Innervation in Male Mice. Nature Metabolism, 5, 481-494. https://doi.org/10.1038/s42255-023-00751-9
[15]	Raun, S.H., Henriquez-Olguín, C., Karavaeva, I., Ali, M., Møller, L.L.V., Kot, W., et al. (2020) Housing Temperature Influences Exercise Training Adaptations in Mice. Nature Communications, 11, Article No. 1560. https://doi.org/10.1038/s41467-020-15311-y
[16]	Safdar, A., Saleem, A. and Tarnopolsky, M.A. (2016) The Potential of Endurance Exercise-Derived Exosomes to Treat Metabolic Diseases. Nature Reviews Endocrinology, 12, 504-517. https://doi.org/10.1038/nrendo.2016.76
[17]	Ringholm, S., Grunnet Knudsen, J., Leick, L., Lundgaard, A., Munk Nielsen, M. and Pilegaard, H. (2013) PGC-1alpha Is Required for Exercise-and Exercise Training-Induced UCP1 Up-Regulation in Mouse White Adipose Tissue. PLOS ONE, 8, e64123. https://doi.org/10.1371/journal.pone.0064123
[18]	Fisher, F.M., Kleiner, S., Douris, N., Fox, E.C., Mepani, R.J., Verdeguer, F., et al. (2012) FGF21 Regulates PGC-1alpha and Browning of White Adipose Tissues in Adaptive Thermogenesis. Genes & Development, 26, 271-281. https://doi.org/10.1101/gad.177857.111
[19]	Kyung, D.S., Lee, E., Chae, S., Son, Y., Moon, Y., Hwang, D., et al. (2024) Single-Cell Transcriptomic Analysis Reveals Dynamic Activation of Cellular Signaling Pathways Regulating Beige Adipogenesis. Experimental & Molecular Medicine, 56, 2309-2322. https://doi.org/10.1038/s12276-024-01252-9
[20]	Ohno, H., Shinoda, K., Spiegelman, B.M. and Kajimura, S. (2012) PPARgamma Agonists Induce a White-to-Brown Fat Conversion through Stabilization of PRDM16 Protein. Cell Metabolism, 15, 395-404. https://doi.org/10.1016/j.cmet.2012.01.019
[21]	Kotzbeck, P., Giordano, A., Mondini, E., Murano, I., Severi, I., Venema, W., et al. (2018) Brown Adipose Tissue Whitening Leads to Brown Adipocyte Death and Adipose Tissue Inflammation. Journal of Lipid Research, 59, 784-794. https://doi.org/10.1194/jlr.m079665
[22]	Cairó, M., Villarroya, J., Cereijo, R., Campderrós, L., Giralt, M. and Villarroya, F. (2016) Thermogenic Activation Represses Autophagy in Brown Adipose Tissue. International Journal of Obesity, 40, 1591-1599. https://doi.org/10.1038/ijo.2016.115
[23]	Kim, J., Kundu, M., Viollet, B. and Guan, K. (2011) AMPK and mTOR Regulate Autophagy through Direct Phosphorylation of Ulk1. Nature Cell Biology, 13, 132-141. https://doi.org/10.1038/ncb2152
[24]	Villarroya, J., Franco-Bordés, C., Blasco-Roset, A. and Villarroya, F. (2026) Brown Adipose Tissue Decline in Aging, a Role for Autophagy? Geromedicine, 2, Article ID: 202517. https://doi.org/10.70401/geromedicine.2026.0013
[25]	Tanida, I., Tanida-Miyake, E., Ueno, T. and Kominami, E. (2001) The Human Homolog of Saccharomyces Cerevisiae Apg7p Is a Protein-Activating Enzyme for Multiple Substrates Including Human Apg12p, GATE-16, GABARAP, and MAP-LC3. Journal of Biological Chemistry, 276, 1701-1706. https://doi.org/10.1074/jbc.c000752200
[26]	Ichimura, Y., Kirisako, T., Takao, T., Satomi, Y., Shimonishi, Y., Ishihara, N., et al. (2000) A Ubiquitin-Like System Mediates Protein Lipidation. Nature, 408, 488-492. https://doi.org/10.1038/35044114
[27]	Mizushima, N., Yamamoto, A., Hatano, M., Kobayashi, Y., Kabeya, Y., Suzuki, K., et al. (2001) Dissection of Autophagosome Formation Using Apg5-Deficient Mouse Embryonic Stem Cells. The Journal of Cell Biology, 152, 657-668. https://doi.org/10.1083/jcb.152.4.657
[28]	Komatsu, M., Waguri, S., Ueno, T., Iwata, J., Murata, S., Tanida, I., et al. (2005) Impairment of Starvation-Induced and Constitutive Autophagy in Atg7-Deficient Mice. The Journal of Cell Biology, 169, 425-434. https://doi.org/10.1083/jcb.200412022
[29]	Singh, R., Xiang, Y., Wang, Y., Baikati, K., Cuervo, A.M., Luu, Y.K., et al. (2009) Autophagy Regulates Adipose Mass and Differentiation in Mice. Journal of Clinical Investigation, 119, 3329-3339. https://doi.org/10.1172/jci39228
[30]	Zhang, Y., Goldman, S., Baerga, R., Zhao, Y., Komatsu, M. and Jin, S. (2009) Adipose-Specific Deletion of Autophagy-Related Gene 7 (Atg7) in Mice Reveals a Role in Adipogenesis. Proceedings of the National Academy of Sciences, 106, 19860-19865. https://doi.org/10.1073/pnas.0906048106
[31]	Taylor, D. and Gottlieb, R.A. (2017) Parkin‐Mediated Mitophagy Is Downregulated in Browning of White Adipose Tissue. Obesity, 25, 704-712. https://doi.org/10.1002/oby.21786
[32]	Lu, X., Altshuler-Keylin, S., Wang, Q., Chen, Y., Henrique Sponton, C., Ikeda, K., et al. (2018) Mitophagy Controls Beige Adipocyte Maintenance through a Parkin-Dependent and UCP1-Independent Mechanism. Science Signaling, 11, eaap8526. https://doi.org/10.1126/scisignal.aap8526
[33]	Altshuler-Keylin, S., Shinoda, K., Hasegawa, Y., Ikeda, K., Hong, H., Kang, Q., et al. (2016) Beige Adipocyte Maintenance Is Regulated by Autophagy-Induced Mitochondrial Clearance. Cell Metabolism, 24, 402-419. https://doi.org/10.1016/j.cmet.2016.08.002
[34]	Moore, T.M., Cheng, L., Wolf, D.M., Ngo, J., Segawa, M., Zhu, X., et al. (2022) Parkin Regulates Adiposity by Coordinating Mitophagy with Mitochondrial Biogenesis in White Adipocytes. Nature Communications, 13, Article No. 6661. https://doi.org/10.1038/s41467-022-34468-2
[35]	Ji, X., Zhang, X., Zhang, T., Xue, Y., He, M., Li, C., et al. (2025) PNPLA7 Mediates Parkin-Mitochondrial Recruitment in Adipose Tissue for Mitophagy and Inhibits Browning. Nature Communications, 16, Article No. 6651. https://doi.org/10.1038/s41467-025-61904-w
[36]	Cairó, M., Campderrós, L., Gavaldà‐Navarro, A., Cereijo, R., Delgado‐Anglés, A., Quesada‐López, T., et al. (2019) Parkin Controls Brown Adipose Tissue Plasticity in Response to Adaptive Thermogenesis. The EMBO Reports, 20, e46832. https://doi.org/10.15252/embr.201846832
[37]	Yau, W.W., Wong, K.A., Zhou, J., Thimmukonda, N.K., Wu, Y., Bay, B., et al. (2021) Chronic Cold Exposure Induces Autophagy to Promote Fatty Acid Oxidation, Mitochondrial Turnover, and Thermogenesis in Brown Adipose Tissue. iScience, 24, Article ID: 102434. https://doi.org/10.1016/j.isci.2021.102434
[38]	Yau, W.W., Singh, B.K., Lesmana, R., Zhou, J., Sinha, R.A., Wong, K.A., et al. (2019) Thyroid Hormone (T₃) Stimulates Brown Adipose Tissue Activation via Mitochondrial Biogenesis and MTOR-Mediated Mitophagy. Autophagy, 15, 131-150. https://doi.org/10.1080/15548627.2018.1511263
[39]	Dobre, M., Virgolici, B. and Timnea, O. (2025) Key Roles of Brown, Subcutaneous, and Visceral Adipose Tissues in Obesity and Insulin Resistance. Current Issues in Molecular Biology, 47, Article No. 343. https://doi.org/10.3390/cimb47050343
[40]	Guo, Y., Cordes, K.R., Farese, R.V. and Walther, T.C. (2009) Lipid Droplets at a Glance. Journal of Cell Science, 122, 749-752. https://doi.org/10.1242/jcs.037630
[41]	Singh, R., Kaushik, S., Wang, Y., Xiang, Y., Novak, I., Komatsu, M., et al. (2009) Autophagy Regulates Lipid Metabolism. Nature, 458, 1131-1135. https://doi.org/10.1038/nature07976
[42]	Jais, A. and Brüning, J.C. (2017) Hypothalamic Inflammation in Obesity and Metabolic Disease. Journal of Clinical Investigation, 127, 24-32. https://doi.org/10.1172/jci88878
[43]	Enriori, P.J., Evans, A.E., Sinnayah, P., Jobst, E.E., Tonelli-Lemos, L., Billes, S.K., et al. (2007) Diet-Induced Obesity Causes Severe but Reversible Leptin Resistance in Arcuate Melanocortin Neurons. Cell Metabolism, 5, 181-194. https://doi.org/10.1016/j.cmet.2007.02.004
[44]	Madden, C.J. and Morrison, S.F. (2016) A High-Fat Diet Impairs Cooling-Evoked Brown Adipose Tissue Activation via a Vagal Afferent Mechanism. American Journal of Physiology-Endocrinology and Metabolism, 311, E287-E292. https://doi.org/10.1152/ajpendo.00081.2016
[45]	Li, F. and Zhang, H. (2019) Lysosomal Acid Lipase in Lipid Metabolism and Beyond. Arteriosclerosis, Thrombosis, and Vascular Biology, 39, 850-856. https://doi.org/10.1161/atvbaha.119.312136
[46]	Duta-Mare, M., Sachdev, V., Leopold, C., Kolb, D., Vujic, N., Korbelius, M., et al. (2018) Lysosomal Acid Lipase Regulates Fatty Acid Channeling in Brown Adipose Tissue to Maintain Thermogenesis. Biochimica et Biophysica Acta (BBA)—Molecular and Cell Biology of Lipids, 1863, 467-478. https://doi.org/10.1016/j.bbalip.2018.01.011
[47]	Yeh, Y., Evans, T.D., Iwase, M., Jeong, S., Zhang, X., Liu, Z., et al. (2025) Identification of Lysosomal Lipolysis as an Essential Noncanonical Mediator of Adipocyte Fasting and Cold-Induced Lipolysis. Journal of Clinical Investigation, 135, e185340. https://doi.org/10.1172/jci185340
[48]	Kaushik, S. and Cuervo, A.M. (2018) The Coming of Age of Chaperone-Mediated Autophagy. Nature Reviews Molecular Cell Biology, 19, 365-381. https://doi.org/10.1038/s41580-018-0001-6
[49]	Mestres-Arenas, A., Quesada-López, T., Blasco-Roset, A., Gavaldà-Navarro, A., Godoy-Nieto, F.J., Cereijo, R., et al. (2025) Chaperone-Mediated Autophagy Controls Brown Adipose Tissue Thermogenic Activity. Science Advances, 11, eady0415. https://doi.org/10.1126/sciadv.ady0415
[50]	Zhuang, Y., Zhang, X., Zhang, S., Sun, Y., Wang, H., Chen, Y., et al. (2025) Chaperone-Mediated Autophagy Manipulates PGC1alpha Stability and Governs Energy Metabolism under Thermal Stress. Nature Communications, 16, Article No. 4455. https://doi.org/10.1038/s41467-025-59618-0
[51]	Rodriguez-Navarro, J.A., Kaushik, S., Koga, H., Dall’Armi, C., Shui, G., Wenk, M.R., et al. (2012) Inhibitory Effect of Dietary Lipids on Chaperone-Mediated Autophagy. Proceedings of the National Academy of Sciences, 109, E705-E714. https://doi.org/10.1073/pnas.1113036109
[52]	Kaushik, S., Juste, Y.R., Lindenau, K., Dong, S., Macho-González, A., Santiago-Fernández, O., et al. (2022) Chaperone-mediated Autophagy Regulates Adipocyte Differentiation. Science Advances, 8, eabq2733. https://doi.org/10.1126/sciadv.abq2733
[53]	Jansen, H.J., van Essen, P., Koenen, T., Joosten, L.A.B., Netea, M.G., Tack, C.J., et al. (2012) Autophagy Activity Is Up-Regulated in Adipose Tissue of Obese Individuals and Modulates Proinflammatory Cytokine Expression. Endocrinology, 153, 5866-5874. https://doi.org/10.1210/en.2012-1625
[54]	Yang, K., Cai, J., Pan, M., Sun, Q. and Sun, C. (2020) Mark4 Inhibited the Browning of White Adipose Tissue by Promoting Adipocytes Autophagy in Mice. International Journal of Molecular Sciences, 21, Article No. 2752. https://doi.org/10.3390/ijms21082752
[55]	Mauthe, M., Orhon, I., Rocchi, C., Zhou, X., Luhr, M., Hijlkema, K., et al. (2018) Chloroquine Inhibits Autophagic Flux by Decreasing Autophagosome-Lysosome Fusion. Autophagy, 14, 1435-1455. https://doi.org/10.1080/15548627.2018.1474314
[56]	Parray, H.A. and Yun, J.W. (2017) Combined Inhibition of Autophagy Protein 5 and Galectin‐1 by Thiodigalactoside Reduces Diet‐Induced Obesity through Induction of White Fat Browning. IUBMB Life, 69, 510-521. https://doi.org/10.1002/iub.1634
[57]	Li, Y., Zheng, W., Li, X., Lue, Z., Liu, Y., Wu, J., et al. (2024) The Autophagic Regulation of Rosiglitazone-Promoted Adipocyte Browning. Frontiers in Pharmacology, 15, Article ID: 1412520. https://doi.org/10.3389/fphar.2024.1412520
[58]	Armani, A., Cinti, F., Marzolla, V., Morgan, J., Cranston, G.A., Antelmi, A., et al. (2014) Mineralocorticoid Receptor Antagonism Induces Browning of White Adipose Tissue through Impairment of Autophagy and Prevents Adipocyte Dysfunction in High-Fat-Diet-Fed Mice. The FASEB Journal, 28, 3745-3757. https://doi.org/10.1096/fj.13-245415
[59]	安冬, 梁永林, 高艳奎, 等. 大黄黄连泻心汤加味对肥胖2型糖尿病大鼠内脏脂肪线粒体自噬及棕色化影响[J]. 中国实验方剂学杂志, 2024, 30(24): 131-140.
[60]	张晖, 孙立亚, 王庆峰, 等. 益糖康对db/db小鼠白色脂肪棕色化及PINK1/Parkin通路的影响[J]. 中国中医药信息杂志, 2024, 31(12): 91-98.
[61]	杨丽娟, 洪逸莲, 林怡, 等. 人参皂苷Rb2通过抑制自噬促进肥胖小鼠白色脂肪棕色化[J]. 中华内分泌代谢杂志, 2020, 36(12): 1055-1061.
[62]	Leu, S.Y., Tsai, Y.C., Chen, W.C., Hsu, C., Lee, Y. and Cheng, P. (2018) Raspberry Ketone Induces Brown-Like Adipocyte Formation through Suppression of Autophagy in Adipocytes and Adipose Tissue. The Journal of Nutritional Biochemistry, 56, 116-125. https://doi.org/10.1016/j.jnutbio.2018.01.017
[63]	Wang, Z., Yang, T., Zeng, M., Wang, Z., Chen, Q., Chen, J., et al. (2023) Miquelianin in Folium Nelumbinis Extract Promotes White-to-Beige Fat Conversion via Blocking AMPK/DRP1/Mitophagy and Modulating Gut Microbiota in HFD-Fed Mice. Food and Chemical Toxicology, 181, Article ID: 114089. https://doi.org/10.1016/j.fct.2023.114089
[64]	Cheng, S., Ni, X., Yao, Y., Sun, Y., Yu, X., Xia, D., et al. (2023) Hyperoside Prevents High-Fat Diet-Induced Obesity by Increasing White Fat Browning and Lipophagy via CDK6-TFEB Pathway. Journal of Ethnopharmacology, 307, Article ID: 116259. https://doi.org/10.1016/j.jep.2023.116259
[65]	Kim, S.N., Kwon, H.J., Akindehin, S., Jeong, H. and Lee, Y. (2017) Effects of Epigallocatechin-3-Gallate on Autophagic Lipolysis in Adipocytes. Nutrients, 9, Article No. 680. https://doi.org/10.3390/nu9070680
[66]	Foster, D.O. and Frydman, M.L. (1979) Tissue Distribution of Cold-Induced Thermogenesis in Conscious Warm-or Cold-Acclimated Rats Reevaluated from Changes in Tissue Blood Flow: The Dominant Role of Brown Adipose Tissue in the Replacement of Shivering by Nonshivering Thermogenesis. Canadian Journal of Physiology and Pharmacology, 57, 257-270. https://doi.org/10.1139/y79-039
[67]	Muzik, O., Mangner, T.J., Leonard, W.R., et al. (2012) Assessment of Oxidative Metabolism in Brown Fat Using PET Imaging. Molecular Imaging and Biology, 14, 347-357.
[68]	Blondin, D.P., Labbé, S.M., Noll, C., Kunach, M., Phoenix, S., Guérin, B., et al. (2015) Selective Impairment of Glucose but Not Fatty Acid or Oxidative Metabolism in Brown Adipose Tissue of Subjects with Type 2 Diabetes. Diabetes, 64, 2388-2397. https://doi.org/10.2337/db14-1651
[69]	Carpentier, A.C. and Blondin, D.P. (2024) Is Stimulation of Browning of Human Adipose Tissue a Relevant Therapeutic Target? Annales d’Endocrinologie, 85, 184-189. https://doi.org/10.1016/j.ando.2024.05.006
[70]	Perez, L.C., Perez, L.T., Nene, Y., Umpierrez, G.E., Davis, G.M. and Pasquel, F.J. (2022) Interventions Associated with Brown Adipose Tissue Activation and the Impact on Energy Expenditure and Weight Loss: A Systematic Review. Frontiers in Endocrinology, 13, Article ID: 1037458. https://doi.org/10.3389/fendo.2022.1037458

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