BATOKİNLER

Agueda-Oyarzabal, M., & Emanuelli, B. (2022). Immune cells in thermogenic adipose depots: The essential but complex relationship. Frontiers in Endocrinology, 13, 839360. https://doi.org/10.3389/fendo.2022.839360

Cataldo, L. R., Gao, Q., Argemi-Muntadas, L., Hodek, O., Cowan, E., Hladkou, S., Gheibi, S., Spégel, P., Prasad, R. B., Eliasson, L., Scheele, C., Fex, M., Mulder, H., & Moritz, T. (2022). The human batokine EPDR1 regulates β-cell metabolism and function. Molecular Metabolism, 66, 101629. doi:10.1016/j.molmet.2022.101629

Cereijo, R., Gavaldà-Navarro, A., Cairó, M., Quesada-López, T., Villarroya, J., Morón-Ros, S., Sánchez-Infantes, D., Peyrou, M., Iglesias, R., Mampel, T., Turatsinze, J.-V., Eizirik, D. L., Giralt, M., & Villarroya, F. (2018). CXCL14, a brown adipokine that mediates brown-fat-to-macrophage communication in thermogenic adaptation. Cell Metabolism, 28(5), 750-763.e6. doi:10.1016/j.cmet.2018.07.015

Chen, Y., Hu, Q., Wang, C., & Wang, T. (2023). The crosstalk between BAT thermogenesis and skeletal muscle dysfunction. Frontiers in Physiology, 14, 1132830. doi:10.3389/fphys.2023.1132830

Coassolo, L., Dannieskiold-Samsøe, N. B., Zhao, M., Allen, H., & Svensson, K. J. (2022). New players of the adipose secretome: Therapeutic opportunities and challenges. Current Opinion in Pharmacology, 67, 102302. doi:10.1016/j.coph.2022.102302

da Silva, C. P. V., Hernandez-Saavedra, D., White, J. D., & Stanford, K. I. (2019). Cold and exercise: Therapeutic tools to activate brown adipose tissue and combat obesity. Biology, 8(1), 9. https://doi.org/10.3390/biology8010009

Diez-Roda, P., Perez-Navarro, E., & Garcia-Martin, R. (2024). Adipose tissue as a major launch spot for circulating extracellular vesicle-carried microRNAs coordinating tissue and systemic metabolism. International Journal of Molecular Sciences, 25(24), 13488. doi:10.3390/ijms252413488

Fisher, F. M., Kleiner, S., Douris, N., Fox, E. C., Mepani, R. J., Verdeguer, F., Wu, J., Kharitonenkov, A., Flier, J. S., Maratos-Flier, E., & Spiegelman, B. M. (2012). FGF21 regulates PGC-1α and browning of white adipose tissues in adaptive thermogenesis. Genes & Development, 26(3), 271-281. doi:10.1101/gad.177857.111

Gaspar, R. C., Pauli, J. R., Shulman, G. I., & Munoz, V. R. (2021). An update on brown adipose tissue biology: A discussion of recent findings. American Journal of Physiology-Endocrinology and Metabolism, 320(3), E488-E495. https://doi.org/10.1152/ajpendo.00310.2020

Gavaldà-Navarro, A., Villarroya, J., Cereijo, R., Giralt, M., & Villarroya, F. (2022). The endocrine role of brown adipose tissue: An update on actors and actions. Reviews in Endocrine and Metabolic Disorders, 23(1), 31-41. doi:10.1007/s11154-021-09640-6

Ghesmati, Z., Rashid, M., Fayezi, S., Gieseler, F., Alizadeh, E., & Darabi, M. (2024). An update on the secretory functions of brown, white, and beige adipose tissue: Towards therapeutic applications. Reviews in Endocrine and Metabolic Disorders, 25(2), 279-308. doi:10.1007/s11154-023-09850-0

Gilani, A., Stoll, L., Homan, E. A., & Lo, J. C. (2024). Adipose signals regulating distal organ health and disease. Diabetes, 73(2), 169-177. doi:10.2337/dbi23-0005

Ghosh, A., Chénier, I., Leung, Y. H., Oppong, A. K., Peyot, M.-L., Madiraju, S. R. M., Al-Khairi, I., Abubaker, J., Al-Mulla, F., Prentki, M., & Abu-Farha, M. (2024). Adipocyte Angptl8 deletion improves glucose and energy metabolism and obesity associated inflammation in mice. iScience, 27(12), 111292. doi:10.1016/j.isci.2024.111292

Guerrero, J. J. G., del Rosario, M. A. S., Encarnacion, P. C., Lin, C.-S., Chuang, L.-T., Notarte, K. I., Zhou, J., Wang, C.-H., Chang, C.-W., & Li, W.-C. (2026). Adipose tissue browning in MASLD and its molecular mechanisms and metabolic crosstalk. Current Obesity Reports, 15(1). doi:10.1007/s13679-026-00746-z

Hondares, E., Iglesias, R., Giralt, A., Gonzalez, F. J., Giralt, M., Mampel, T., & Villarroya, F. (2011). Thermogenic activation induces FGF21 expression and release in brown adipose tissue. Journal of Biological Chemistry, 286(15), 12983-12990. doi:10.1074/jbc.M110.215889

Klepac, K., Georgiadi, A., Tschöp, M., & Herzig, S. (2019). The role of brown and beige adipose tissue in glycaemic control. Molecular Aspects of Medicine, 68, 90-100. doi:10.1016/j.mam.2019.07.001

Kong, X., Yao, T., Zhou, P., Kazak, L., Tenen, D., Lyubetskaya, A., Dawes, B. A., Tsai, L., Kahn, B. B., Spiegelman, B. M., Liu, T., & Rosen, E. D. (2018). Brown adipose tissue controls skeletal muscle function via the secretion of myostatin. Cell Metabolism, 28(4), 631-643.e3. doi:10.1016/j.cmet.2018.07.004

Le Lay, S., & Scherer, P. E. (2025). Exploring adipose tissue-derived extracellular vesicles in inter-organ crosstalk: Implications for metabolic regulation and adipose tissue function. Cell Reports, 44(6), 115732. doi:10.1016/j.celrep.2025.115732

Lino, M., Palermo-Ruiz, G., Efthymiou, V., Rodrigues, A., Bhonsle, S., Nawaz, A., Ghosh, A., Brandão, B. B., Watanabe, Y., Barrera Salgueiro, R., Yu, Y., & Kahn, C. R. (2026). Brown fat-specific RNA labelling reveals a network of inter-organ communication by secreted microRNAs. Nature Metabolism, 8(7), 1583-1602. doi:10.1038/s42255-026-01558-0

Lynes, M. D., Leiria, L. O., Lundh, M., Bartelt, A., Shamsi, F., Huang, T. L., . . . Tseng, Y.-H. (2017). The cold-induced lipokine 12,13-diHOME promotes fatty acid transport into brown adipose tissue. Nature Medicine, 23(5), 631-637. doi:10.1038/nm.4297

Malicka, A., Ali, A., MacCannell, A. D. V., & Roberts, L. D. (2025). Brown and beige adipose tissue-derived metabokine and lipokine inter-organ signalling in health and disease. Experimental Physiology, 110(7), 918-935. doi:10.1113/EP092008

Martins, F. F., Martins, B. C., Teixeira, A. V. S., Ajackson, M., Souza-Mello, V., & Daleprane, J. B. (2024). Brown adipose tissue, batokines, and bioactive compounds in foods: An update. Molecular Nutrition & Food Research, 68(6), 2300634. doi:10.1002/mnfr.202300634

Mcleod, K., Datta, V., & Fuller, S. (2025). Adipokines as cardioprotective factors: BAT steps up to the plate. Biomedicines, 13(3), 710. doi:10.3390/biomedicines13030710

Mendez-Gutierrez, A., Aguilera, C. M., Cereijo, R., Osuna-Prieto, F. J., Martinez-Tellez, B., Rico, M. C., . . . Sanchez-Delgado, G. (2024). Cold exposure modulates potential brown adipokines in humans, but only FGF21 is associated with brown adipose tissue volume. Obesity, 32(3), 560-570. doi:10.1002/oby.23970

Mirmiranpour, H., & Barber, T. M. (2026). The role of batokines in metabolism and their therapeutic potential. Expert Review of Endocrinology & Metabolism, 21(2), 143-151. doi:10.1080/17446651.2026.2624400

Negroiu, C. E., Tudorascu, I., Bezna, C. M., Godeanu, S., Diaconu, M., Danoiu, R., & Danoiu, S. (2024). Beyond the cold: Activating brown adipose tissue as an approach to combat obesity. Journal of Clinical Medicine, 13(7), 1973. doi:10.3390/jcm13071973

Ni, B., Farrar, J. S., Vaitkus, J. A., & Celi, F. S. (2015). Metabolic effects of FGF-21: Thermoregulation and beyond. Frontiers in Endocrinology, 6, 148. doi:10.3389/fendo.2015.00148

Niemann, B., Haufs-Brusberg, S., Puetz, L., Feickert, M., Jaeckstein, M. Y., Hoffmann, A., Zurkovic, J., Heine, M., Trautmann, E.-M., Müller, C. E., Tönjes, A., Schlein, C., Jafari, A., Eltzschig, H. K., Gnad, T., Blüher, M., Krahmer, N., Kovacs, P., Heeren, J., & Pfeifer, A. (2022). Apoptotic brown adipocytes enhance energy expenditure via extracellular inosine. Nature, 609(7926), 361-368. doi:10.1038/s41586-022-05041-0

Nishio, M., & Saeki, K. (2020). The remaining mysteries about brown adipose tissues. Cells, 9(11), 2449. doi:10.3390/cells9112449

Noriega, L., Yang, C.-Y., & Wang, C.-H. (2023). Brown fat and nutrition: Implications for nutritional interventions. Nutrients, 15(18), 4072. doi:10.3390/nu15184072

Ooijen, A. M. J. C., Westerterp, K. R., Wouters, L., Schoffelen, P. F. M., Van Steenhoven, A. A., & Van Lichtenbelt, W. D. M. (2006). Heat production and body temperature during cooling and rewarming in overweight and lean men. Obesity, 14(11), 1914-1920. doi:10.1038/oby.2006.223

Ortiz, G. U., & De Freitas, E. C. (2023). Physical activity and batokines. American Journal of Physiology-Endocrinology and Metabolism, 325(5), E610-E620. doi:10.1152/ajpendo.00160.2023

Pan, R., & Chen, Y. (2021). Management of oxidative stress: Crosstalk between brown/beige adipose tissues and skeletal muscles. Frontiers in Physiology, 12, 712372. doi:10.3389/fphys.2021.712372

Pfeifer, A., Mikhael, M., & Niemann, B. (2024). Inosine: Novel activator of brown adipose tissue and energy homeostasis. Trends in Cell Biology, 34(1), 72-82. doi:10.1016/j.tcb.2023.04.007

Pilkington, A.-C., Paz, H. A., & Wankhade, U. D. (2021). Beige adipose tissue identification and marker specificity—Overview. Frontiers in Endocrinology, 12, 599134. doi:10.3389/fendo.2021.599134

Pinckard, K. M., & Stanford, K. I. (2022). The heartwarming effect of brown adipose tissue. Molecular Pharmacology, 102(1), 39-50. doi:10.1124/molpharm.121.000328

Pinhal, D., Gonçalves, L. D. B., Campos, V. F., & Patton, J. G. (2025). Decoding microRNA arm switching: A key to evolutionary innovation and gene regulation. Cellular and Molecular Life Sciences, 82(1), 197. doi:10.1007/s00018-025-05663-3

Planavila, A., Redondo, I., Hondares, E., Vinciguerra, M., Munts, C., Iglesias, R., Gabrielli, L. A., Sitges, M., Giralt, M., Van Bilsen, M., & Villarroya, F. (2013). Fibroblast growth factor 21 protects against cardiac hypertrophy in mice. Nature Communications, 4, 2019. doi:10.1038/ncomms3019

Rial-Pensado, E., Freire-Agulleiro, O., Ríos, M., Guo, D. F., Contreras, C., Seoane-Collazo, P., Tovar, S., Nogueiras, R., Diéguez, C., Rahmouni, K., & López, M. (2022). Obesity induces resistance to central action of BMP8B through a mechanism involving the BBSome. Molecular Metabolism, 59, 101465. doi:10.1016/j.molmet.2022.101465

Ruocco, C., Malavazos, A. E., Ragni, M., Carruba, M. O., Valerio, A., Iacobellis, G., & Nisoli, E. (2023). Amino acids contribute to adaptive thermogenesis: New insights into the mechanisms of action of recent drugs for metabolic disorders are emerging. Pharmacological Research, 195, 106892. doi:10.1016/j.phrs.2023.106892

Singh, R., Barrios, A., Dirakvand, G., & Pervin, S. (2021). Human brown adipose tissue and metabolic health: Potential for therapeutic avenues. Cells, 10(11), 3030. doi:10.3390/cells10113030

Sponton, C. H., Hosono, T., Taura, J., Jedrychowski, M. P., Yoneshiro, T., Wang, Q., . . . Kajimura, S. (2020). The regulation of glucose and lipid homeostasis via PLTP as a mediator of BAT–liver communication. EMBO Reports, 21(9), e49828. doi:10.15252/embr.201949828

Suchacki, K. J., & Stimson, R. H. (2021). Nutritional regulation of human brown adipose tissue. Nutrients, 13(6), 1748. doi:10.3390/nu13061748

Svensson, K. J., Long, J. Z., Jedrychowski, M. P., Cohen, P., Lo, J. C., Serag, S., Kir, S., Shinoda, K., Tartaglia, J. A., Rao, R. R., Chédotal, A., Kajimura, S., Gygi, S. P., & Spiegelman, B. M. (2016). A secreted Slit2 fragment regulates adipose tissue thermogenesis and metabolic function. Cell Metabolism, 23(3), 454-466. doi:10.1016/j.cmet.2016.01.008

Villarroya, F., Cereijo, R., Villarroya, J., & Giralt, M. (2017). Brown adipose tissue as a secretory organ. Nature Reviews Endocrinology, 13(1), 26-35. doi:10.1038/nrendo.2016.136

Virtanen, K. A., Lidell, M. E., Orava, J., Heglind, M., Westergren, R., Niemi, T., Taittonen, M., Laine, J., Savisto, N.-J., Enerbäck, S., & Nuutila, P. (2009). Functional brown adipose tissue in healthy adults. New England Journal of Medicine, 360(15), 1518-1525. doi:10.1056/NEJMoa0808949

Walker, M. E., Kodani, S. D., Mena, H. A., Tseng, Y.-H., Cypess, A. M., & Spite, M. (2024). Brown adipose tissue activation in humans increases plasma levels of lipid mediators. The Journal of Clinical Endocrinology & Metabolism, 109(7), 1837-1849. doi:10.1210/clinem/dgae016

Wang, H., Guo, S., Gao, H., Ding, J., Li, H., Kong, X., Zhang, S., He, M., Feng, Y., Wu, W., Xu, K., Chen, Y., Zhang, H., Liu, T., & Kong, X. (2024). Myostatin regulates energy homeostasis through autocrine- and paracrine-mediated microenvironment communication. Journal of Clinical Investigation, 134(16), e178303. doi:10.1172/JCI178303

White, J. D., Dewal, R. S., & Stanford, K. I. (2019). The beneficial effects of brown adipose tissue transplantation. Molecular Aspects of Medicine, 68, 74-81. doi:10.1016/j.mam.2019.06.004

Whitehead, A., Krause, F. N., Moran, A., MacCannell, A. D. V., Scragg, J. L., McNally, B. D., Boateng, E., Murfitt, S. A., Virtue, S., Wright, J., Garnham, J., Davies, G. R., Dodgson, J., Schneider, J. E., Murray, A. J., Church, C., Vidal-Puig, A., Witte, K. K., Griffin, J. L., & Roberts, L. D. (2021). Brown and beige adipose tissue regulate systemic metabolism through a metabolite interorgan signaling axis. Nature Communications, 12(1), 1905. doi:10.1038/s41467-021-22272-3

Xu, J., Cui, L., Wang, J., Zheng, S., Zhang, H., Ke, S., Cao, X., Shi, Y., Li, J., Zen, K., Vidal-Puig, A., Zhang, C.-Y., Li, L., & Jiang, X. (2023). Cold-activated brown fat-derived extracellular vesicle-miR-378a-3p stimulates hepatic gluconeogenesis in male mice. Nature Communications, 14(1), 5480. doi:10.1038/s41467-023-41160-6

Yan, C., Burley, G., Gao, H., & Shi, Y.-C. (2025). Emerging insights into brown adipose tissue crosstalk with pancreatic β-cells in metabolic regulation. Endocrinology, 166(9). doi:10.1210/endocr/bqaf118

Yang, F. T., & Stanford, K. I. (2022). Batokines: Mediators of inter-tissue communication (A mini-review). Current Obesity Reports, 11(1), 1-9. doi:10.1007/s13679-021-00465-7

Yoneshiro, T., Matsushita, M., Hibi, M., Tone, H., Takeshita, M., Yasunaga, K., Katsuragi, Y., Kameya, T., Sugie, H., & Saito, M. (2017). Tea catechin and caffeine activate brown adipose tissue and increase cold-induced thermogenic capacity in humans. The American Journal of Clinical Nutrition, 105(4), 873-881. doi:10.3945/ajcn.116.144972

Zhang, X., Xiao, J., Jiang, M., Phillips, C. J. C., & Shi, B. (2025). Thermogenesis and energy metabolism in brown adipose tissue in animals experiencing cold stress. International Journal of Molecular Sciences, 26(7), 3233. doi:10.3390/ijms26073233

Zhang, Y., Shen, W., Qiu, S., Yang, P., Dempsey, G., Zhao, L., Zhou, Q., Hao, X., Dong, D., Stahl, A., Kraemer, F. B., Leung, L. L., & Morser, J. (2021). Chemerin regulates formation and function of brown adipose tissue: Ablation results in increased insulin resistance with high fat challenge and aging. The FASEB Journal, 35(7), e21687. doi:10.1096/fj.202100156R

Ziqubu, K., Dludla, P. V., Mabhida, S. E., Jack, B. U., Keipert, S., Jastroch, M., & Mazibuko-Mbeje, S. E. (2024). Brown adipose tissue-derived metabolites and their role in regulating metabolism. Metabolism, 150, 155709. doi:10.1016/j.metabol.2023.155709