Uncoupling proteins

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Uncoupling proteins


Uncoupling proteins (UCPs) are mitochondrial anion carrier proteins that can be found in the inner mitochondrial membranes of animals and plants. UCP1 acts as an uncoupler by dissipating the electrochemical proton gradient (mitochondrial membrane potential), generated by the electron transfer pathway by pumping protons from the mitochondrial matrix to the mitochondrial intermembrane space.

Abbreviation: UCP

Communicated by Bufe A 2017-05-04

Uncoupling protein homologues

The gene family of uncoupling proteins (UCP) includes five mitochondrial solute carriers 25 (SLC25), named UCP1 (SLC25A7), UCP2 (SLC25A48), UCP3 (SLC25A9), UCP4 (SLC25A27) and UCP5 (SLC25A14).[1]. These proteins have a tripartite structure and are located in the inner membrane of mitochondria. Presumably all of them contribute to the metabolic regulation elicited by cold exposure, including ROS and lipid metabolism, apoptosis and thermogenesis.[2]
The thermogenic function of UCP1, which was the first uncoupling protein to be discovered in 1978 [3], is already well established, whereas the exact functions of the closely related paralogues UCP2 and UCP3 are yet to be investigated. [4] [5]. UCP4 and UCP5 are primarily expressed in the central nervous system (CNS) where they function as essential uncouplers of oxidative phosphorylation, thereby exerting an important protective function for cells by reducing oxidative stress (ROS).


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Publications in the MiPMap

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» List of publications: Uncoupling proteins
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 YearReferenceTissue and cellMammal and model
Davis, 2021 Journal of Applied Physiology2021Davis, MS; Barrett, MR (2021) Effect of conditioning and physiology hyperthermia on canine skeletal muscle mitochondrial oxygen consumption. Journal of Applied Physiology Online.Skeletal muscleDog
Friederich-Persson 2020 Am J Physiol Renal Physiol2020Friederich-Persson M, Persson P (2020) Mitochondrial angiotensin II receptors regulate oxygen consumption in kidney mitochondria from healthy and type 1 diabetic rats. Am J Physiol Renal Physiol 318:F683-88.KidneyRat
Cruz 2020 Drug Alcohol Depend2020Cruz JV, Maba IK, Correia D, Kaziuk FD, Cadena SMSC, Zampronio AR (2020) Intermittent binge-like ethanol exposure during adolescence attenuates the febrile response by reducing brown adipose tissue thermogenesis in rats. Drug Alcohol Depend 209:107904.FatRat
Brunetta 2020 J Physiol2020Brunetta HS, Politis-Barber V, Petrick HL, Dennis KMJH, Kirsh AJ, Barbeau PA, Nunes EA, Holloway GP (2020) Nitrate attenuates HFD-induced glucose intolerance in association with reduced epididymal adipose tissue inflammation and mitochondrial ROS emission. J Physiol 598:3357-71.FatMouse
Piel 2020 FEBS J2020Piel MS, Masscheleyn S, Bouillaud F, Moncoq K, Miroux B (2020) Structural models of mitochondrial uncoupling proteins obtained in DPC micelles are not functionally relevant. FEBS J [Epub ahead of print].Saccharomyces cerevisiae
Fink 2019 FASEB J2019Fink BD, Yu L, Sivitz WI (2019) Modulation of complex II-energized respiration in muscle, heart, and brown adipose mitochondria by oxaloacetate and complex I electron flow. FASEB J 33:11696-705.Heart
Skeletal muscle
Oliveira 2019 Mol Nutr Food Res2019Oliveira TE, Castro É, Belchior T, Andrade ML, Chaves-Filho AB, Peixoto AS, Moreno MF, Ortiz Silva M, Moreira RJ, Inague A, Yoshinaga MY, Miyamoto S, Moustaid-Moussa N, Festuccia WT (2019) Fish oil protects wild type and uncoupling protein 1 deficient mice from obesity and glucose intolerance by increasing energy expenditure. Mol Nutr Food Res 63:e1800813.FatMouse
Ramalho 2019 J Leukoc Biol2019Ramalho T, Ramalingam L, Filgueiras L, Festuccia W, Jancar S, Moustaid-Moussa N (2019) Leukotriene-B4 modulates macrophage metabolism and fat loss in type 1 diabetic mice. J Leukoc Biol 106:665-75.Macrophage-derivedMouse
McBride 2019 Arch Biochem Biophys2019McBride S, Wei-LaPierre L, McMurray F, MacFarlane M, Qiu X, Patten DA, Dirksen RT, Harper ME (2019) Skeletal muscle mitoflashes, pH, and the role of uncoupling protein-3. Arch Biochem Biophys 663:239-48.Skeletal muscleMouse
Mayeuf-Louchart 2019 Cell Rep2019Mayeuf-Louchart A, Lancel S, Sebti Y, Pourcet B, Loyens A, Delhaye S, Duhem C, Beauchamp J, Ferri L, Thorel Q, Boulinguiez A, Zecchin M, Dubois-Chevalier J, Eeckhoute J, Vaughn LT, Roach PJ, Dani C, Pederson BA, Vincent SD, Staels B, Duez H (2019) Glycogen dynamics drives lipid droplet biogenesis during brown adipocyte differentiation. Cell Rep 29:1410-18.FatMouse
Girer 2019 Int J Mol Sci2019Girer NG , Carter D, Bhattarai N, Mustafa M, Denner L, Porter C, Elferink CJ (2019) Inducible loss of the aryl hydrocarbon receptor activates perigonadal white fat respiration and brown fat thermogenesis via fibroblast growth factor 21. Int J Mol Sci 20:950.FatMouse
Aguilar 2019 Cell Rep2019Aguilar E, Esteves P, Sancerni T, Lenoir V, Aparicio T, Bouillaud F, Dentin R, Prip-Buus C, Ricquier D, Pecqueur C, Guilmeau S, Alves-Guerra MC (2019) UCP2 deficiency increases colon tumorigenesis by promoting lipid synthesis and depleting NADPH for antioxidant defenses. Cell Rep 28:2306-16.Endothelial;epithelial;mesothelial cellMouse
Johnson 2019 Mol Metab2019Johnson JM, Verkerke ARP, Maschek JA, Ferrara PJ, Lin CT, Kew KA, Neufer PD, Lodhi IJ, Cox JE, Funai K (2019) Alternative splicing of UCP1 by non-cell-autonomous action of PEMT. Mol Metab 31:55-66.FatMouse
Crescenzo 2019 Mol Neurobiol2019Crescenzo R, Spagnuolo MS, Cancelliere R, Iannotta L, Mazzoli A, Gatto C, Iossa S, Cigliano L (2019) Effect of initial aging and high-fat/high-fructose diet on mitochondrial bioenergetics and oxidative status in rat brain. Mol Neurobiol 56:7651-63.Nervous systemRat
Nacarelli 2018 Geroscience2018Nacarelli T, Azar A, Altinok O, Orynbayeva Z, Sell C (2018) Rapamycin increases oxidative metabolism and enhances metabolic flexibility in human cardiac fibroblasts. Geroscience 40:243-56.Heart
Rossignoli 2018 J Nutr Biochem2018Rossignoli CP, Dechandt CRP, Souza AO, Sampaio IH, Vicentini TM, Teodoro BG, Neto MPC, Ferrari GD, Couto-Lima CA, Alberici LC (2018) Effects of intermittent dietary supplementation with conjugated linoleic acid and fish oil (EPA/DHA) on body metabolism and mitochondrial energetics in mice. J Nutr Biochem 60:16-23.Skeletal muscle
Nervous system
Friederich-Persson 2018 Acta Physiol (Oxf)2018Friederich-Persson M, Persson P, Hansell P, Palm F (2018) Deletion of Uncoupling Protein-2 reduces renal mitochondrial leak respiration, intrarenal hypoxia and proteinuria in a mouse model of type 1 diabetes. Acta Physiol (Oxf) 223:e13058.KidneyMouse
Rocha 2018 J Physiol Biochem2018Rocha C, Koury OH, Scheede-Bergdahl C, Bergdahl A (2018) Cardiac mitochondrial respiration following a low-carbohydrate, high-fat diet in apolipoprotein E-deficient mice. J Physiol Biochem 75:65-72.HeartMouse
Boutoual 2018 Scientific Reports2018Boutoual, R., Meseguer, S., Villarroya, M., Martin-Hernandez, E., Errami, M., Martin, M. A., Casado, M., and Armengod, M. E. (2018) Defects in the mitochondrial-tRNA modification enzymes MTO1 and GTPBP3 promote different metabolic reprogramming through a HIF-PPARgamma-UCP2-AMPK axis. Scientific reports 8, 1163Other cell lines
Schiffer 2018 PLoS One2018Schiffer TA, Christensen M, Gustafsson H, Palm F (2018) The effect of inactin on kidney mitochondrial function and production of reactive oxygen species. PLoS One 13:e0207728.KidneyRat
Jezek 2018 Antioxid Redox Signal2018Jezek P, Holendova B, Garlid KD, Jaburek M (2018) Mitochondrial uncoupling proteins: subtle regulators of cellular redox signaling. Antioxid Redox Signal 29:667-714.
Horscroft 2017 Proc Natl Acad Sci U S A2017Horscroft JA, Kotwica AO, Laner V, West JA, Hennis PJ, Levett DZH, Howard DJ, Fernandez BO, Burgess SL, Ament Z, Gilbert-Kawai ET, Vercueil A, Landis BD, Mitchell K, Mythen MG, Branco C, Johnson RS, Feelisch M, Montgomery HE, Griffin JL, Grocott MPW, Gnaiger E, Martin DS, Murray AJ (2017) Metabolic basis to Sherpa altitude adaptation. Proc Natl Acad Sci U S A 114:6382–7.Skeletal muscleHuman
Porter 2017 Adipocyte2017Porter C (2017) Quantification of UCP1 function in human brown adipose tissue. Adipocyte 6:167-74.Fat
Smith 2016 Diabetes2016Smith BK, Ford RJ, Desjardins EM, Green AE, Hughes MC, Houde VP, Day EA, Marcinko K, Crane JD, Mottillo EP, Perry CG, Kemp BE, Tarnopolsky MA, Steinberg GR (2016) Salsalate (Salicylate) uncouples mitochondria, improves glucose homeostasis, and reduces liver lipids independent of AMPK β1. Diabetes 65:3352-61.Skeletal muscle
Chondronikola 2016 Cell Metab2016Chondronikola M, Volpi E, Børsheim E, Porter C, Saraf MK, Annamalai P, Yfanti C, Chao T, Wong D, Shinoda K, Labbė SM, Hurren NM, Cesani F, Kajimura S, Sidossis LS. (2016) Brown adipose tissue activation is linked to distinct systemic effects on lipid metabolism in humans. Cell Metab 23:1200-6.Human
Porter 2016 Cell Metab2016Porter C, Herndon DN, Chondronikola M, Chao T, Annamalai P, Bhattarai N, Saraf MK, Capek KD, Reidy PT, Daquinag AC, Kolonin MG, Rasmussen BB, Borsheim E, Toliver-Kinsky T, Sidossis LS (2016) Human and mouse brown adipose tissue mitochondria have comparable UCP1 function. Cell Metab 24:246-55.Skeletal muscle
McFarlane 2016 Thesis2016McFarlane SV (2016) Temperature-dependent alterations of brown adipose tissue metabolism during hibernation in the thirteen-lined ground squirrel, Ictidomys tridecemlineatus. Dissertation p74.FatOther mammals
Chondronikola 2015 Int J Obes (Lond)2015Chondronikola M, Annamalai P, Chao T, Porter C, Saraf MK, Cesani F, Sidossis LS (2015) A percutaneous needle biopsy technique for sampling the supraclavicular brown adipose tissue depot of humans.. Int J Obes (Lond) 39(10):1561-4.FatHuman
Schrauwen-Hinderling 2015 Antioxid Redox Signal2015Schrauwen-Hinderling VB, Kooi ME, Schrauwen P (2015) Mitochondrial function and diabetes; consequences for skeletal and cardiac muscle metabolism. Antioxid Redox Signal 24:39-51.Heart
Skeletal muscle
Jezek 2015 Antioxid Redox Signal2015Ježek J, Dlasková A, Zelenka J, Jabůrek M, Ježek P (2015) H2O2-activated mitochondrial phospholipase iPLA2γ prevents lipotoxic oxidative stress in synergy with UCP2, amplifies signaling via G-protein-coupled receptor GPR40, and regulates insulin secretion in pancreatic β-cells. Antioxid Redox Signal 23:958-72.Islet cell;pancreas;thymus
Petruzzelli 2014 Cell Metab2014Petruzzelli M, Schweiger M, Schreiber R, Campos-Olivas R, Tsoli M, Allen J, Swarbrick M, Rose-John S, Rincon M, Robertson G, Zechner R, Wagner EF (2014) A switch from white to brown fat increases energy expenditure in cancer-associated cachexia. Cell Metab 20:433-47.FatMouse
Chondronikola 20142014Chondronikola M, Volpi E, Børsheim E, Porter C, Annamalai P, Enerbäck S, Lidell ME, Saraf MK, Labbe SM, Hurren NM, Yfanti 7, Chao T, Andersen CR, Cesani F, Hawkins H, Sidossis LS. (2014) Brown adipose tissue improves whole-body glucose homeostasis and insulin sensitivity in humans. Diabetes 63(12):4089-99.FatHuman
Conley 2013 J Clin Endocrinol Metab2013Conley KE, Amara CE, Bajpeyi S, Costford SR, Murray K, Jubrias SA, Arakaki L, Marcinek DJ, Smith SR (2013) Higher mitochondrial respiration and uncoupling with reduced electron transport chain content in vivo in muscle of sedentary versus active subjects. J Clin Endocrinol Metab 98:129-36.Skeletal muscleHuman
Jaburek 2013 Int J Biochem Cell Biol2013Jabůrek M, Ježek J, Zelenka J, Ježek P (2013) Antioxidant activity by a synergy of redox-sensitive mitochondrial phospholipase A2 and uncoupling protein-2 in lung and spleen. Int J Biochem Cell Biol 45:816-25.Mouse
Hals 2012 Biochem Biophys Res Commun2012Hals IK, Ogata H, Pettersen E, Ma Z, Björklund A, Skorpen F, Egeberg KW, Grill V (2012) Marked over expression of uncoupling protein-2 in beta cells exerts minor effects on mitochondrial metabolism. Biochem Biophys Res Commun 423:259-64.Islet cell;pancreas;thymus
Dlaskova 2010 Biochim Biophys Acta2010Dlasková A, Clarke KJ, Porter RK (2010) The role of UCP 1 in production of reactive oxygen species by mitochondria isolated from brown adipose tissue. Biochim Biophys Acta 1797:1470-6.FatMouse
Namgaladze 2010 Atherosclerosis2010Namgaladze D, Preiss S, Dröse S, Brandt U, Brüne B (2010) Phospholipase A2-modified low density lipoprotein induces mitochondrial uncoupling and lowers reactive oxygen species in phagocytes. Atherosclerosis 208:142-7.Blood cellsHuman
Dikov 2010 Exp Gerontol2010Dikov D, Aulbach A, Muster B, Dröse S, Jendrach M, Bereiter-Hahn J (2010) Do UCP2 and mild uncoupling improve longevity? Exp Gerontol 45:586-95.HeLa
Guillet 2010 Neurogenetics2010Guillet V, Gueguen N, Verny C, Ferre M, Homedan C, Loiseau D, Procaccio V, Amati-Bonneau P, Bonneau D, Reynier P, Chevrollier A (2010) Adenine nucleotide translocase is involved in a mitochondrial coupling defect in MFN2-related Charcot-Marie-Tooth type 2A disease. Neurogenetics 11:127-33.Nervous systemHuman
Jones 2010 Biochim Biophys Acta2010Jones TT, Brewer GJ (2010) Age-related deficiencies in Complex I endogenous substrate availability and reserve capacity of Complex IV in cortical neuron electron transport. Biochim Biophys Acta 1797:167-76.Nervous systemRat
De Meis 2010 PLoS One2010De Meis L, Ketzer LA, da Costa RM, de Andrade IR, Benchimol M (2010) Fusion of the endoplasmic reticulum and mitochondrial outer membrane in rats brown adipose tissue: activation of thermogenesis by Ca2+. PLoS One 5:e9439.FatRat
Romanatto 2009 J Biol Chem2009Romanatto T, Roman EA, Arruda AP, Denis RG, Solon C, Milanski M, Moraes JC, Bonfleur ML, Degasperi GR, Picardi PK, Hirabara S, Boschero AC, Curi R, Velloso LA (2009) Deletion of tumor necrosis factor-alpha-receptor 1 (TNFR1) protects against diet-induced obesity by means of increased thermogenesis. J Biol Chem 284:36213-22.Skeletal muscleRat
Guachalla 2009 Aging (Albany NY)2009Guachalla LM, Ju Z, Koziel R, Figura G, Song Z, Fusser M, Epe B, Jansen-Dürr P, Rudolph KL (2009) SOD2 haploinsufficiency does not accelerate aging of telomere dysfunctional mice. Aging (Albany NY) 1:303-15.Nervous systemMouse
Anderson 2009 J Clin Invest2009Anderson EJ, Lustig ME, Boyle KE, Woodlief TL, Kane DA, Lin C-T, Price JW, Kang L, Rabinovitch PS, Szeto HH, Houmard JA, Cortright RN, Wasserman DH, Neufer PD (2009) Mitochondrial H2O2 emission and cellular redox state link excess fat intake to insulin resistance in both rodents and humans. J Clin Invest 119:573-81.Skeletal muscleHuman
Nabben 2008 FEBS Lett2008Nabben M, Hoeks J, Briedé JJ, Glatz JF, Moonen-Kornips E, Hesselink MK, Schrauwen P (2008) The effect of UCP3 overexpression on mitochondrial ROS production in skeletal muscle of young versus aged mice. FEBS Lett 582:4147-52.Skeletal muscleMouse
Crisan 2008 Stem Cells2008Crisan M, Casteilla L, Lehr L, Carmona OMC, Paoloni-Giacobino A, Yap S, Sun B, Leger B, Logar A, Penicaud L, Schrauwen P, Cameron-Smith D, Russell AP, Peault B, Giacobino JP (2008) A reservoir of brown adipocyte progenitors in human skeletal muscle. Stem Cells 26:2425-33.Endothelial;epithelial;mesothelial cellHuman
Rabinovich 2007 Eur Respir J2007Rabinovich RA, Bastos R, Ardite E, Llinàs L, Orozco-Levi M, Gea J, Vilaró J, Barberà JA, Rodríguez-Roisin R, Fernández-Checa JC, Roca J (2007) Mitochondrial dysfunction in COPD patients with low body mass index. Eur Respir J 29:643-50.Skeletal muscleHuman
Breen 2006 J Biol Chem2006Breen EP, Gouin SG, Murphy AF, Haines LR, Jackson AM, Pearson TW, Murphy PV, Porter RK (2006) On the mechanism of mitochondrial uncoupling protein 1 function. J Biol Chem 281:2114-9.Rat
Dejean 2001 Biochim Biophys Acta2001Dejean L, Beauvoit B, Bunoust O, Fleury B, Guérin B, Rigoulet M (2001) The calorimetric-respirometric ratio is an on-line marker of enthalpy efficiency of yeast cells growing on a non-fermentable carbon source. Biochim Biophys Acta 1503:329-40.Fungi
Savagner 2001 J Clin Endocrinol Metabol2001Savagner F, Franc B, Guyetant S, Rodien P, Reynier P, Malthiery Y (2001) Defective mitochondrial ATP synthesis in oxyphilic thyroid tumors. J Clin Endocrinol Metabol 86:4920-25.Human
Publications in the MiPMap

Uncoupling protein - Abstracts

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Porter 2018 MiP2018b
Skeletal muscle mitochondrial thermogenesis in hypermetabolic humans.
Garipi 2018 MiPschool Tromso E2Phenotyping mitochondrial metabolism in Barrett’s metaplasia-dysplasia-adenocarcinoma sequence: respiratory capacity, extracelular proton flux and ROS productionHumanEndothelial;epithelial;mesothelial cell
Other cell lines
Porter 2018 MiP2018a
Evidence of functional uncoupling protein 1 in the white adipose tissue of hypermetabolic humans.
Paes 2014 Abstract IOC 2014-04 SchroeckenPaes MC, Saraiva FMS, Nogueira NP, Laranja GAT, Coelho MGP, Oliveira MF (2014) Heme increases mitochondrial membrane potential and ROS production in Trypanosoma cruzi epimastigotes. Mitochondr Physiol Network 19.02.Protists
Andersen 2014 Abstract IOC 2014-04 SchroeckenAndersen M, Vienberg SG, Novod JB, Brandauer J, Holst B, Treebak JT (2014) Nicotinamide phosphoribosyltransferase knockdown impairs mitochondrial function in mouse myoblasts. Mitochondr Physiol Network 19.02.MouseSkeletal muscle
Other cell lines
Jaburek 2014 Abstract MiP2014The role of phospholipase A2γ in the regulation of mitochondrial uncoupling protein 2–dependent antioxidant function.Islet cell;pancreas;thymus
Gnaiger 2013 Abstract Mito2013Gnaiger E, Søndergaard H, Munch-Andersen T, Damsgaard R, Hagen C, Díez-Sánchez C, Ara I, Wright-Paradis C, Schrauwen P, Hesselink M, Calbet J, Christiansen M, Helge JW, Saltin B, Boushel R (2013) Conserved mitochondrial coupling efficiency and adaptive substrate control in Inuit and Caucasians: adaptation and acclimatization to the Arctic winter.HumanSkeletal muscle
Chicco 2013 Abstract MiP2013 PosterChicco AJ, Claiborne MS, Le CH, Mulligan CM (2013) Remodeling of skeletal muscle mitochondria in response to exercise training in taz shRNA mouse model of human Barth Syndrome. Mitochondr Physiol Network 18.08.MouseSkeletal muscle
Palm 2013 Abstract MiP2013Palm F, Hansell P, Friederich-Persson M (2013) Role of mitochondrial function for the onset and progression of kidney disease. Mitochondr Physiol Network 18.08.Mouse
Shabalina 2013 Abstract MiP2013Shabalina IG, Petrovic N, Kalinovich AV, Cannon B, Nedergaard J (2013) Comparative study of brown and white adipose tissue mitochondria in mice upon cold acclimation. Mitochondr Physiol Network 18.08.Fat
Vercesi 2013 Abstract MiP2013Vercesi AE (2013) Mitochondrial respiration, Ca2+ transport and oxidative stress: Fungi, trypanosomes, plants and mammals. Mitochondr Physiol Network 18.08.Rat
Other invertebrates
Gnaiger 2013 Abstract MiP2013Gnaiger E, Søndergaard H, Munch-Andersen T, Damsgaard R, Hagen C, Díez-Sánchez C, Ara I, Wright-Paradis C, Schrauwen P, Hesselink M, Calbet J, Christiansen M, Helge JW, Saltin B, Boushel R (2013) Biochemical coupling efficiency in permeabilized fibers from arm and leg muscle in Inuit versus Caucasians: A functional test of the uncoupling hypothesis in Greenland. Mitochondr Physiol Network 18.08.HumanSkeletal muscle
Kressig 2013 Abstract IOC79Kressig F (2013) et al. Mechanism of increased pulmonary vascular remodeling in UCP2 k/o mice. Mitochondr Physiol Network 18.07MouseLung;gill
Other cell lines
Jezek 2013 Abstract MiP2013Ježek P, Ježek J, Zelenka J, Dlasková A, Jabůrek M (2013) Antioxidant synergy of mitochondrial uncoupling protein UCP2 and phospholipase iPLA2γ. Mitochondr Physiol Network 18.08.MouseLung;gill
Islet cell;pancreas;thymus


  1. Ramsden DB, Ho PW-L, Ho JW-M, Liu HF, So DHF, Tse HM, Chan KH, Ho SL (2012) Human neuronal uncoupling proteins 4 and 5 (UCP4 and UCP5): structural properties, regulation, and physiological role in protection against oxidative stress and mitochondrial dysfunction. Brain and Behavior 2:468–78.
  2. Criscuolo F, Gonzalez‐Barroso MdM, Bouillaud F, Ricquier D, Miroux B, Sorci G (2005) Mitochondrial uncoupling proteins: New perspectives for evolutionary ecologists. Amer Naturalist 166:686-99.
  3. Nicholls DG, Bernson VSM, Heaton GM (1978) The identification of the component in the inner membrane of brown adipose tissue mitochondria responsible for regulating energy dissipation. In: Girardier L, Seydoux J, eds. Effectors of thermogenesis: Proceedings of a symposium held at Geneva (Switzerland) on 14 to 16 July 1977. Basel: Birkhäuser Basel. p 89-93.
  4. Cannon B, Nedergaard J (2004). Brown adipose tissue: Function and physiological significance. Physiol Rev 84:277-359.
  5. Ricquier D, Bouillaud F (2000) The uncoupling protein homologues: UCP1, UCP2, UCP3, STUCP and ATUCP. Biochem J 345:161-79.

  6. MitoPedia topics: Uncoupler