Cited by 530Adipose Tissue and MetabolismAdipokines, Inflammation, and Metabolic DiseasesCardiovascular Disease and Adiposity

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Mapping of human brown adipose tissue in lean and obese young men

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Leitner, Brooks P., Huang, Shan, Brychta, Robert J., Duckworth, Courtney J., Baskin, Alison S., McGehee, Suzanne, Tal, Ilan, Dieckmann, William, Gupta, Garima, Kolodny, Gerald M., Pacak, Karel, Herscovitch, Peter, Cypess, Aaron M., Chen, Kong Y.·2017·Proceedings of the National Academy of Sciences

DOI 10.1073/pnas.1705287114PMC PMC5559032

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Abstract

F-labeled fluorodeoxyglucose positron emission tomography/computerized tomography (PET/CT). Obese men had less activated BAT than lean men (mean, 130 vs. 334 mL) but more fat in BAT-containing depots (mean, 1,646 vs. 855 mL) with a wide range (0.1-71%) in the ratio of activated BAT to inactive fat between individuals. Six anatomic regions had activated BAT-cervical, supraclavicular, axillary, mediastinal, paraspinal, and abdominal-with 67 ± 20% of all activated BAT concentrated in a continuous fascial layer comprising the first three depots in the upper torso. These nonsubcutaneous fat depots amounted to 1.5% of total body mass (4.3% of total fat mass), and up to 90% of each depot could be activated BAT. The amount and activity of BAT was significantly influenced by region of interest selection methods, PET threshold criteria, and PET resolutions. The present study suggests that active BAT can be found in specific adipose depots in adult humans, but less than one-half of the fat in these depots is stimulated by acute cold exposure, demonstrating a previously underappreciated thermogenic potential.

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BackgroundInfluential
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Background
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Background
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Ho, W.-E., Sun, L., Goh, H. J., Tint, M. T., Sun, L., & Leow, M. K. S. (2022) · Adipocyte · doi:10.1080/21623945.2022.2104509 · Cited by 7
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“…differentiated and recruited in adult humans when compared to newborns, likely due to environmental, behavioural and physiological adaptations to large body size and living at thermoneutrality in our species (i.e. preferred temperature range with minimal adaptive thermogenesis requirements) [1,2].”
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Astragalus polysaccharide regulates brown adipocytes differentiation by miR ‐6911 targeting Prdm16
Zhang, S., Song, P., Chen, X., Wang, Y., Gao, X., Liang, L., & Zhao, J. (2021) · Lipids · doi:10.1002/lipd.12328 · Cited by 8
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“Lean adults have more dense brown fat than obese or heavy adults [33].”
Background
Analysis of Factors Associated With Continued Cooling of Core Temperature After Prolonged Cold-Water Swimming
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Background
Irisin reverses high-fat diet-induced metabolic dysfunction via activation of brown adipose tissue in mice
Dai, J., Zhao, Y., Chen, Y., Jiang, Y., Sun, R., Tang, X., Cui, Y., Mao, H. Z., & Peng, X. (2025) · International Journal of Obesity · doi:10.1038/s41366-025-01739-z · Cited by 6
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“BAT is a thermogenic tissue that has a role in controlling body weight by producing energy in the form of heat [16], [17].”
Background
Opioid growth factor receptor promotes adipose tissue thermogenesis via enhancing lipid oxidation
Zhang, S., Chen, J., Li, Q., & Zeng, W. (2023) · Life Metabolism · doi:10.1093/lifemeta/load018 · Cited by 7
“Since most adult humans have at least some BAT or beige fat [74, 75], a controlled increase in tissue activity seems a plausible approach.”
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“The majority of published studies on PVAT’s pro-vs. anti-contractile function are conducted in the abdominal aorta, which is a non-resistance artery with PVAT containing largely brown adipocytes [26].”
Background
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“In some studies, BMI and body fat percentage had a negative association with BAT prevalence, while resting metabolic rate had a strong positive correlation [26,74,75] .”
Background
Experimental studies on the correlations of male respiratory parameters with their body composition
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“According to its functional role, phenotype and profile of gene expression, adipose tissue can be further divided by two as WAT and BAT [24].”
BackgroundInfluential
Automatic segmentation of human supraclavicular adipose tissue using high-resolution T2-weighted magnetic resonance imaging
Wu, B., Cheng, C., Qi, Y., Zhou, H., Peng, H., Wan, Q., Liu, X., Zheng, H., Zhang, H., & Zou, C. (2022) · Magnetic Resonance Materials in Physics Biology and Medicine · doi:10.1007/s10334-022-01056-w · Cited by 5
“Accurate and efficient segmentation of BAT is important for BAT identification and the subsequent function assessment to better evaluate its therapeutic potential [14, 15].”
BackgroundResult
Human brown adipose tissue function: insights from current in vivo techniques
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Energy Expenditure and Macronutrient Oxidation in Response to an Individualized Nonshivering Cooling Protocol
Sánchez‐Delgado, G., Alcántara, J. M. A., Acosta, F. M., Martínez‐Téllez, B., Amaro‐Gahete, F. J., Merchán-Ramírez, E., Löf, M., Labayen, I., Ravussin, É., & Ruiz, J. R. (2020) · Obesity · doi:10.1002/oby.22972 · Cited by 5
“Although other methods have been proposed, such as skin temperature clamping (18) or cold perception adjustment (19), the lowest tolerable temperature above the shivering threshold has been broadly accepted and used as a valid approach to induce BAT activation and CIT (17,20-23).”
Method
Fetal programming of adipose tissue function by gestational chronodisruption
Halabí, D., Ehrenfeld, P., Méndez, N., Hans, G. H. R., & Torres‐Farfan, C. (2020) · Sleep Science · doi:10.5935/1984-0063.20200014 · Cited by 8
Allobaculum plays a predominant role in the activation of brown fat tissue by diallyl trisulfide
Hu, Y., Xu, Y., Huangfu, B., Zhang, R., Xu, J., Gao, R., Wang, T., Wang, J., Huang, K., Luo, Y., et al. (2025) · Food Frontiers · doi:10.1002/fft2.456 · Cited by 4
Brown adipose tissue, anthropometric traits, and metabolic profiles in women with normal weight obesity
Dojo, K., Ishida, K., Fuse-Hamaoka, S., Aoki, M., Tsunoda, M., Yamamoto, M., Ota, N., & Hamaoka, T. (2025) · Scientific Reports · doi:10.1038/s41598-025-10583-0 · Cited by 5
The role of immune responses and microbiota in adipose tissue homeostasis
Azar, S., & Rehermann, B. (2025) · The Journal of Physiology · doi:10.1113/jp286422 · Cited by 5
Cross-Talk Between Signaling and Transcriptional Networks Regulating Thermogenesis—Insights into Canonical and Non-Canonical Regulatory Pathways
Simka-Lampa, K. (2026) · International Journal of Molecular Sciences · doi:10.3390/ijms27020754 · Cited by 4
Vitamin A Derivatives and Adipose Tissue Differentiation: Molecular Pathways Driving Browning and Anti-Obesity Effects
Aygun, B. B., Basturk, B., & Ayaz, A. (2026) · Current Obesity Reports · doi:10.1007/s13679-025-00684-2 · Cited by 7
“The amount of BAT in adults is relatively limited; some studies have reported that BAT depots constitute approximately 1.5% of total body weight and 4.3% of total fat mass [28 ].”
Influential
Transferrin receptor 1‐mediated iron uptake supports thermogenic activation in human cervical‐derived adipocytes
Alrifai, R., Seo, M., Karadsheh, G., Mahendra, F. R., Demény, M. Á., Győry, F., Mótyán, J. A., Fésüs, L., Kristóf, E., & Arianti, R. (2026) · FEBS Letters · doi:10.1002/1873-3468.70312 · Cited by 4
“[6] observed that brown or brownable adipose tissue can be found in DeepC, supraclavicular, axillary, mediastinal, para-spinal, and abdominal regions.”
Adipose tissue catecholamine resistance: a metabolic safeguard that challenges weight control
Guardia, L. D., & Shin, A. C. (2026) · Metabolism · doi:10.1016/j.metabol.2026.156590 · Cited by 7
PCPE-1 promotes cardiac fibrosis with aging and obesity
Hsiao, Y., Yoshida, Y., Tsuchimochi, H., Tang, J., Aung, T. M., Chang, C.-H., Barinda, A. J., Li, Z., Othman, N. S. B., Yoshizaki, T., et al. (2026) · JCI Insight · doi:10.1172/jci.insight.195508 · Cited by 4
Two-step regulation by matrix Gla protein in brown adipose cell differentiation
Zhang, L., Cai, X., Ma, F., Qiao, X., Ji, J., Ma, J. A., Vergnes, L., Zhao, Y., Yao, Y., Wu, X., et al. (2024) · Molecular Metabolism · doi:10.1016/j.molmet.2024.101870 · Cited by 4
“BAT is abundant in the interscapular and perirenal locations in human infants, whereas adult human BAT is found mainly in the cervical, supraclavicular and paravertebral regions [3,9].”
Background
Stimulation of non-shivering thermogenesis by bioactive compounds: A focus on gut microbiota-mediated mechanisms
Alhamoud, Y., Abudumijiti, T., Wu, J., Lu, L., Zhao, M., Luo, X., Feng, F., & Wang, J. (2024) · Trends in Food Science & Technology · doi:10.1016/j.tifs.2024.104779 · Cited by 5
Thermogenic Activation of Adipose Tissue by Caffeine During Strenuous Exercising and Recovery: A Double-Blind Crossover Study
Soto, D. A. S., Pérez, D. I. V., de Souza, M. R., Reis, M. D. P., Almeida, N. R., Miarka, B., Aedo–Muñoz, E., Oyarzo, A. I. A., Sillero‐Quintana, M., Queiroz, A. C. C., et al. (2025) · Metabolites · doi:10.3390/metabo15080517 · Cited by 4
“On the other hand, Infrared thermography (IRT) presents a non-invasive, low-cost alternative by measuring skin temperature in BAT-rich regions, such as the supraclavicular area, where superficial deposits are abundant [8,9].”
Local large arterial perivascular adipose tissue metabolic and anatomical features are associated with hypertensive clinical outcomes: a PET/CT-based study
Long, H.-Y., Chen, G., Liu, J., Cheng, K., Yu, W.-J., & Chen, B.-Q. (2025) · Annals of Medicine · doi:10.1080/07853890.2025.2578724 · Cited by 4
“CT values were constrained to the range of −300 to −10 Hounsfield unit (HU) [26].”
Extraction and Characterization of Human Adipose Tissue-Derived Collagen: Toward Xeno-Free Tissue Engineering
Kim, M., Yeo, M., Lee, K., Park, M.-J., Han, G., Lee, C., Park, J., & Jung, B. (2023) · Tissue Engineering and Regenerative Medicine · doi:10.1007/s13770-023-00612-2 · Cited by 4
“Adipose tissue is widely distributed throughout the human body and serves the functions of energy storage and body temperature control [31].”
Sex-specific response of the human plasma lipidome to short-term cold exposure
Höring, M., Brunner, S. P., Scheiber, J., Honecker, J., Liebisch, G., Seeliger, C., Schinhammer, L., Claussnitzer, M., Burkhardt, R., Hauner, H., et al. (2024) · Biochimica et Biophysica Acta (BBA) - Molecular and Cell Biology of Lipids · doi:10.1016/j.bbalip.2024.159567 · Cited by 4
Semi-automated analysis of supraclavicular thermal images increases speed of brown adipose tissue analysis without increasing variation in results
Law, J., Morris, D. E., Robinson, L., Symonds, M. E., & Budge, H. (2021) · Current Research in Physiology · doi:10.1016/j.crphys.2021.07.001 · Cited by 4
“Due to the heat-generating property of BAT, infrared thermography is a natural alternative to PET-CT for measuring BAT activity, especially given the relatively superficial location of the supraclavicular depot which is the largest BAT depot in humans (Leitner et al., 2017).”
Background
Electrocortical temporal complexity during wakefulness and sleep: an updated account
González, J., Cavelli, M., Mondino, A., Pascovich, C., Castro‐Zaballa, S., Rubido, N., & Torterolo, P. (2020) · Sleep Science · doi:10.5935/1984-0063.20200013 · Cited by 6
CFTR regulates brown adipocyte thermogenesis via the cAMP/PKA signaling pathway
Choi, K.-M., Cho, S.-H., Kim, J. H., Kim, A.-R. L., Kong, X., & Yoon, J. C. (2022) · Journal of Cystic Fibrosis · doi:10.1016/j.jcf.2022.08.012 · Cited by 4
“adipocytes are much more extensive in the human body than previously realized and can significantly impact cardiometabolic health [26,27].”
Background
Adipose Tissue Plasticity and Insulin Signaling in the Pathogenesis of Type 2 Diabetes
Sakaguchi, M. (2023) · Diabetology International · doi:10.1007/s13340-023-00676-4 · Cited by 5
“In adult humans, brown adipocytes are primarily found in regions such as the cervical, supraclavicular, paravertebral, and perirenal–adrenal areas [7].”
Neck Circumference and Cerebral Gray Matter Volume
Shin, C., Kim, R. E. Y., THOMAS, R. J., Yun, C., Lee, S. K., & Abbott, R. D. (2020) · Alzheimer Disease & Associated Disorders · doi:10.1097/wad.0000000000000386 · Cited by 5
Sleep and maternal behavior in the postpartum rat after haloperidol and midazolam treatments
Peña, F., Rivas, M., González, J., Schwarzkopf, N., Torterolo, P., Ferreira, A., & Benedetto, L. (2020) · Sleep Science · doi:10.5935/1984-0063.20200019 · Cited by 4
Social modulation on daily variability in electric behavior
Gascue, V., da Silva, A. C. R., & Migliaro, A. (2020) · Sleep Science · doi:10.5935/1984-0063.20200012 · Cited by 6
Yellow adipocytes comprise a new adipocyte sub-type present in human bone marrowPreprint
Attané, C., Estève, D., Chaoui, K., Iacovoni, J. S., Corre, J., Moutahir, M., Valet, P., Burlet‐Schiltz, O., Reina, N., & Muller, C. (2019) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/641886 · Cited by 4
Nutrients Lowering Obesity-Linked Chemokines Blamable for Metastasis
Ion, G., Bostan, M., Hardman, W. E., McFarland, M. P., Bleoţu, C., Radu, N., Diaconu, C. C., Mihaila, M. A., Caramihai, M. D., & Hotnog, C. M. (2025) · International Journal of Molecular Sciences · doi:10.3390/ijms26052275 · Cited by 4
“The most common classification is by localization (e.g., upper body, lower body, subcutaneous, visceral, and ectopic) [104–107].”
Cold-induced phosphatidylethanolamine synthesis in liver and brown adipose tissue of mice
Hidrobo, M. S., Höring, M., Brunner, S. P., Liebisch, G., Schweizer, S., Klingenspor, M., Schreiber, R., Zechner, R., Burkhardt, R., & Ecker, J. (2024) · Biochimica et Biophysica Acta (BBA) - Molecular and Cell Biology of Lipids · doi:10.1016/j.bbalip.2024.159562 · Cited by 3
“However, the amount of BAT in humans is negligible compared to the total volume of adipose tissue in the body [18,19].”
Background
Brown Adipose Tissue Activity and Childhood Exposure to Cold Are Associated With Hot Flashes at Menopause
Sievert, L. L., Shreyer, S., Witkowski, S., & Brown, D. E. (2024) · American Journal of Human Biology · doi:10.1002/ajhb.24148 · Cited by 3
“Higher BMI— the supraclavicular subcutaneous fat layer in particular— may have decreased the likelihood of detecting BAT (Gatidis et al. 2016; Jimenez-Pavon et al. 2019; Leitner et al. 2017).”
Physical and Physiological Properties of Fat
Sugii, S., & Velan, S. S. (2020) · Advances in magnetic resonance technology and applications · doi:10.1016/b978-0-12-817057-1.00027-5 · Cited by 3
Effectiveness of Leininger's Transcultural-based GATHER Counseling Model on Exclusive Breastfeeding
Karnasih, I. G. A. (2021) · Health Notions · doi:10.33846/hn50503 · Cited by 3
“Babies with brown fat could maintain their body temperature in normal conditions so that on the first day the baby did not need excess calories from outside the body (12).”
Background
Association Between Adipose Tissue Proton Density Fat Fraction, Resting Metabolic Rate and FTO Genotype in Humans
Drabsch, T., Junker, D., Bayer, S., Wu, M., Held, C., Karampinos, D. C., Hauner, H., & Holzapfel, C. (2022) · Frontiers in Endocrinology · doi:10.3389/fendo.2022.804874 · Cited by 3
An Improved Thermogram Dataset for Obesity Detection Research
Leo, H., Saddami, K., Roslidar, R., Muharar, R., Munadi, K., & Arnia, F. (2023) · International Workshop on Artificial Intelligence and Image Processing (IWAIIP) · doi:10.1109/iwaiip58158.2023.10462838 · Cited by 3
“However, a study [20] reported that BAT depots could also be found in other body parts apart from the SCV region.”
Background
GPCR-mediated regulation of beige adipocyte formation: Implications for obesity and metabolic health
Cui, Y., Auclair, H., He, R., & Zhang, Q. (2024) · Gene · doi:10.1016/j.gene.2024.148421 · Cited by 3
Integrating Magnetic Resonance Chemical Shift Imaging for Localized Prostate Cancer Risk Stratification on the Basis of the Impact of Periprostatic Brown Adipocytes Within Tumor Microenvironment
Shen, C.-Y., Pan, J.-K., Lin, W.-D., Cheng, H., Hu, C., Wu, K., Kuo, Y.-M., Shieh, G.-S., & Lu, P. (2025) · Annals of Surgical Oncology · doi:10.1245/s10434-025-17512-5 · Cited by 3
Fifty shades of brown: The functions, diverse regulation and evolution of brown adipose tissue
Speakman, J. R. (2019) · Molecular Aspects of Medicine · doi:10.1016/j.mam.2019.07.006 · Cited by 3
Fattening the role of Ca2+ cycling in adaptive thermogenesis
Gamu, D., & Tupling, A. R. (2017) · Nature Medicine · doi:10.1038/nm.4457 · Cited by 3
The Mediating Role of Brown Fat and Skeletal Muscle Measured by 18F‐Fluorodeoxyglucose in the Thermoregulatory System in Young Adults
Martínez‐Téllez, B., Adelantado‐Renau, M., Acosta, F. M., Sánchez‐Delgado, G., Martínez-Nicolás, A., Boon, M. R., Elvira, J. M. L., Martínez‐Vizcaíno, V., & Ruiz, J. R. (2019) · Obesity · doi:10.1002/oby.22461 · Cited by 3
“Furthermore, we repeated the analyses using BAT-related outcomes as well as skeletal muscle activity multiplied by lean body mass percentage (29), and the results remained unchanged (data not shown).”
Method
Protocolo de lipoaspiração mecânica para redução de gordura submentoniana e submandibular pela técnica TLA: Relato de caso
Fernandes, A. T. R., Agnol, E. D., de Oliveira, R. C. G., & Rizzatti‐Barbosa, C. M. (2022) · AESTHETIC OROFACIAL SCIENCE · doi:10.51670/aos.v3i1.75 · Cited by 2
Glucose sensing and homeostasis by adipocyte GPCR
Hasan, N., & Chhabra, K. H. (2025) · Frontiers in Endocrinology · doi:10.3389/fendo.2025.1657747 · Cited by 3
“Development of metabolic syndromes, including insulin resistance due to this excess visceral fat deposition is well known (24, 25) and leads to cardiovascular diseases and type 2 diabetes (26 – 28).”
An ultrasound-guided biopsy technique for obtaining supraclavicular brown fat biopsies and preadipocytes
Andersen, E. S., Jespersen, N. Z., Henriksen, T. I., Lorentsen, K. M., Palani, N. P., Pedersen, B. K., Schéele, C., Nielsen, M. B., & Nielsen, S. L. (2025) · Scientific Reports · doi:10.1038/s41598-025-24183-5 · Cited by 2
Oxylipin Responses during the Yukon Arctic Ultra: The Longest and Coldest Ultramarathon in the World
Coker, R. H., Tsuji, T., Thomas, R. L., Coker, M. S., Aristizabal‐Henao, J. J., Kiebish, M. A., Kienest, C., Schalt, A., Tseng, Y., & Steinach, M. (2025) · Military Medicine · doi:10.1093/milmed/usaf622 · Cited by 2
SEL1L–HRD1 ERAD–autophagy interplay maintains mitochondrial homeostasis in brown adipocytes
Chen, X., Wang, S., Torres, M., Hao, S., Sun, S., & Qi, L. (2026) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2529914123 · Cited by 2
“As BAT plays a central role in energy expenditure and thermo genesis in humans (78, 79), and considering that mitochondrial function and dynamics are to the pathogenesis of various diseases (80, 81), our findings have significant implications for met abolic disorders such as obesity and type 2…”
Imaging Brown Adipose Tissue with TSPO PET Tracers in Preclinical Animal Studies
Zhu, B., Liang, S. H., & Ran, C. (2023) · Methods in molecular biology · doi:10.1007/978-1-0716-3167-6_13 · Cited by 2

Authors: Leitner, B. P., Huang, S., Brychta, R. J., Duckworth, C. J., Baskin, A. S., McGehee, S., Tal, I., Dieckmann, W., Gupta, G., Kolodny, G. M., Pacak, K., Herscovitch, P., Cypess, A. M., & Chen, K. Y.
Affiliations: National Institutes of Health (US)
Beth Israel Deaconess Medical Center (US)
Eunice Kennedy Shriver National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, MD 20892
ORCID: Leitner, Brooks P.
Huang, Shan
Brychta, Robert J.
Duckworth, Courtney J.
Baskin, Alison S.
Gupta, Garima
Cypess, Aaron M.
Chen, Kong Y.
Venue: Proceedings of the National Academy of Sciences
Publisher: National Academy of Sciences
Published: 2017-07-24
Volume: 114
Issue: 32
Pages: 8649-8654
ISSN: 0027-8424 (ISSN-L)
1091-6490
DOI: 10.1073/pnas.1705287114
PMC: PMC5559032
OpenAlex: W2736913806
Cited by: 530
References: 43
Citations by year: 2026: 36 · 2025: 45 · 2024: 51 · 2023: 68 · 2022: 73 · 2021: 82 · 2020: 73 · 2019: 45 · 2018: 46 · 2017: 9
FWCI: 34.99
Type: journal article
Language: en
Open access: green
www.ncbi.nlm.nih.gov
Licence: www.pnas.org/site/misc/userlicense.xhtml (vor)
Locations: Proceedings of the National Academy of Sciences (publishedVersion)
PubMed Central (publishedVersion)
PubMed (publishedVersion)
Funders: HHS | NIH | National Institute of Diabetes and Digestive and Kidney Diseases (Z01 DK071014, Z01 DK075116)
Topics: Health Sciences › Medicine › Physiology (medical) › Adipose Tissue and Metabolism
Health Sciences › Medicine › Endocrinology, Diabetes and Metabolism › Adipokines, Inflammation, and Metabolic Diseases
Health Sciences › Medicine › Cardiology and Cardiovascular Medicine › Cardiovascular Disease and Adiposity
Keywords: brown adipose tissue, obesity, cold exposure, thermogenesis, BAT activation
MeSH terms: Adult, Adipose Tissue, Brown, Humans, Male, Obesity, Tomography, X-Ray Computed, Body Mass Index, Glucose-6-Phosphate, Positron-Emission Tomography, Adiposity

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Mapping of human brown adipose tissue in lean and obese young men

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