title: Brown and beige fat in humans: thermogenic adipocytes that control energy and glucose homeostasis.
abstract: Brown adipose tissue (BAT), a specialized fat that dissipates energy to produce heat, plays an important role in the regulation of energy balance. Two types of thermogenic adipocytes with distinct developmental and anatomical features exist in rodents and humans: classical brown adipocytes and beige (also referred to as brite) adipocytes. While classical brown adipocytes are located mainly in dedicated BAT depots of rodents and infants, beige adipocytes sporadically reside with white adipocytes and emerge in response to certain environmental cues, such as chronic cold exposure, a process often referred to as "browning" of white adipose tissue. Recent studies indicate the existence of beige adipocytes in adult humans, making this cell type an attractive therapeutic target for obesity and obesity-related diseases, including type 2 diabetes. This Review aims to cover recent progress in our understanding of the anatomical, developmental, and functional characteristics of brown and beige adipocytes and discuss emerging questions, with a special emphasis on adult human BAT.
authors: Labros
authors: L
authors: Sidossis
authors: Shingo
authors: S
authors: Kajimura
mesh_terms: D002001:Adipose Tissue, Brown / Q000378:metabolism*; D000328:Adult; D000818:Animals; D003924:Diabetes Mellitus, Type 2 / Q000378:metabolism / Q000473:pathology; D004734:Energy Metabolism / Q000502:physiology*; D005947:Glucose / Q000378:metabolism*; D006801:Humans; D009765:Obesity / Q000378:metabolism / Q000473:pathology; D022722:Thermogenesis / Q000502:physiology*
chemical_list: D005947:Glucose
references: Am J Physiol Endocrinol Metab. 2010 Jun;298(6):E1244-53
references: 20354155
references: J Clin Invest. 2013 Aug;123(8):3395-403
references: 23867626
references: Nat Med. 2013 Oct;19(10):1338-44
references: 23995282
references: Nat Med. 2011 Feb;17(2):200-5
references: 21258337
references: J Cell Physiol. 2009 Feb;218(2):444-9
references: 18937285
references: Eur J Appl Physiol Occup Physiol. 1981;46(4):339-45
references: 6266825
references: Cell. 2014 Jun 5;157(6):1292-308
references: 24906148
references: Annu Rev Physiol. 2014;76:225-49
references: 24188710
references: Proc Natl Acad Sci U S A. 2011 Jan 4;108(1):143-8
references: 21173238
references: Front Endocrinol (Lausanne). 2012 Jan 03;2:102
references: 22654837
references: Int J Obes (Lond). 2014 Jun;38(6):812-7
references: 24213309
references: J Appl Physiol Respir Environ Exerc Physiol. 1978 Dec;45(6):908-15
references: 730596
references: Front Endocrinol (Lausanne). 2011 Oct 31;2:64
references: 22654820
references: Nature. 2012 Jan 26;481(7382):463-8
references: 22237023
references: Obesity (Silver Spring). 2011 Jan;19(1):13-6
references: 20448535
references: Cell Metab. 2014 May 6;19(5):810-20
references: 24709624
references: Diabetes. 2014 Dec;63(12):4089-99
references: 25056438
references: J Clin Invest. 2013 Jan;123(1):215-23
references: 23221344
references: N Engl J Med. 2009 Apr 9;360(15):1500-8
references: 19357405
references: Mol Cell Biol. 2012 Jul;32(14):2871-9
references: 22586266
references: Am J Physiol Cell Physiol. 2000 Sep;279(3):C670-81
references: 10942717
references: Diabetes. 2014 Jan;63(1):162-75
references: 24062245
references: Cell Metab. 2014 Apr 1;19(4):593-604
references: 24703692
references: Dev Biol. 2006 Aug 1;296(1):164-76
references: 16730693
references: Nat Med. 2012 Jun;18(6):918-25
references: 22561685
references: Cell. 2014 Jun 5;157(6):1279-91
references: 24906147
references: J Anat. 1972 May;112(Pt 1):35-9
references: 5086212
references: Proc Natl Acad Sci U S A. 2013 Feb 26;110(9):E798-807
references: 23388637
references: Diabetes. 2012 Jul;61(7):1801-13
references: 22522611
references: J Anat Physiol. 1908 Oct;43(Pt 1):43-58
references: 17232790
references: J Anat. 2009 Jan;214(1):171-8
references: 19018882
references: Mol Cell Biol. 2005 Sep;25(18):8311-22
references: 16135818
references: Genes Dev. 2008 May 15;22(10):1397-409
references: 18483224
references: N Engl J Med. 2009 Apr 9;360(15):1509-17
references: 19357406
references: Am J Physiol Endocrinol Metab. 2010 Oct;299(4):E601-6
references: 20606075
references: PLoS One. 2014;9(1):e84910
references: 24416310
references: Clin Nucl Med. 2006 Apr;31(4):193-6
references: 16550009
references: Proc Natl Acad Sci U S A. 2007 Mar 13;104(11):4401-6
references: 17360536
references: Genes Dev. 2012 Feb 1;26(3):271-81
references: 22302939
references: Diabetes. 2010 Jul;59(7):1789-93
references: 20357363
references: Genesis. 2010 Jul;48(7):424-36
references: 20641127
references: J Clin Invest. 2014 Feb;124(2):515-27
references: 24401271
references: Diabetes. 2009 Jul;58(7):1526-31
references: 19401428
references: Obesity (Silver Spring). 2013 Nov;21(11):2279-87
references: 23554353
references: Cell Metab. 2009 Oct;10(4):324-35
references: 19808025
references: J Nucl Med. 2003 Nov;44(11):1789-96
references: 14602861
references: Nat Med. 2013 May;19(5):635-9
references: 23603815
references: Cell. 2012 Jul 20;150(2):366-76
references: 22796012
references: AJR Am J Roentgenol. 2004 Oct;183(4):1127-32
references: 15385319
references: Cell Metab. 2011 Jul 6;14(1):67-79
references: 21723505
references: J Clin Invest. 2012 Feb;122(2):545-52
references: 22269323
references: Cell Metab. 2014 Jan 7;19(1):96-108
references: 24411942
references: Diabetes. 2014 Nov;63(11):3686-98
references: 24954193
references: Eur J Nucl Med Mol Imaging. 2007 Jul;34(7):1018-22
references: 17225118
references: Proc Nutr Soc. 1989 Jul;48(2):243-56
references: 2678120
references: Nature. 2008 Aug 21;454(7207):961-7
references: 18719582
references: J Clin Invest. 2011 Jan;121(1):96-105
references: 21123942
references: Biochim Biophys Acta. 2013 May;1831(5):950-9
references: 23454374
references: J Clin Invest. 2012 Mar;122(3):1022-36
references: 22307324
references: Endocrinology. 2012 Jul;153(7):3133-46
references: 22549226
references: Eur J Appl Physiol Occup Physiol. 1990;60(6):418-24
references: 2167842
references: Eur J Nucl Med Mol Imaging. 2002 Oct;29(10):1393-8
references: 12271425
references: Nature. 2009 Aug 27;460(7259):1154-8
references: 19641492
references: Cell Rep. 2013 Dec 12;5(5):1196-203
references: 24290753
references: Mol Syndromol. 2012 Apr;2(3-5):202-212
references: 22670141
references: Cell Metab. 2009 Feb;9(2):203-9
references: 19187776
references: Nat Med. 2013 Oct;19(10):1252-63
references: 24100998
references: Cell Metab. 2013 May 7;17(5):798-805
references: 23663743
references: J Clin Endocrinol Metab. 2011 Jan;96(1):192-9
references: 20943785
references: Nature. 2014 Sep 4;513(7516):100-4
references: 25043053
references: J Clin Invest. 2006 Mar;116(3):615-22
references: 16511594
references: Nat Cell Biol. 2013 Jun;15(6):659-67
references: 23624403
references: J Lipid Res. 2007 Jan;48(1):41-51
references: 17041251
references: Cell. 2012 May 11;149(4):871-85
references: 22579288
references: Mol Imaging Biol. 2011 Dec;13(6):1278-83
references: 21140233
references: Cell. 2014 Jan 16;156(1-2):304-16
references: 24439384
references: Diabetologia. 2013 Jan;56(1):147-55
references: 23064293
references: Cell Metab. 2014 Feb 4;19(2):302-9
references: 24506871
references: Science. 2010 May 28;328(5982):1158-61
references: 20448152
references: Cell Metab. 2012 Apr 4;15(4):480-91
references: 22482730
references: Am J Physiol Endocrinol Metab. 2007 Aug;293(2):E444-52
references: 17473055
references: Diabetes. 2014 Oct;63(10):3253-65
references: 24789919
references: Cell Metab. 2012 Sep 5;16(3):348-62
references: 22940198
references: Cell Metab. 2010 Apr 7;11(4):257-62
references: 20374957
references: Nature. 2011 Dec 1;480(7375):104-8
references: 22101429
references: Cell Metab. 2011 Sep 7;14(3):324-38
references: 21907139
references: PLoS One. 2010;5(6):e11391
references: 20613988
references: PLoS One. 2012;7(11):e49452
references: 23166672
references: Nat Med. 2014 Aug;20(8):911-8
references: 25038826
references: Cell Metab. 2014 Sep 2;20(3):408-15
references: 25185947
references: Nature. 2008 Aug 21;454(7207):1000-4
references: 18719589
references: Clin Sci (Lond). 1984 Feb;66(2):179-86
references: 6692652
references: Int J Obes (Lond). 2013 Jul;37(7):993-8
references: 23032405
references: Am J Physiol Regul Integr Comp Physiol. 2011 May;300(5):R1115-25
references: 21368268
references: J Med Genet. 2003 Apr;40(4):300-3
references: 12676904
references: J Clin Invest. 2013 Aug;123(8):3404-8
references: 23867622
references: Cell Metab. 2011 Aug 3;14(2):272-9
references: 21803297
references: N Engl J Med. 2009 Apr 9;360(15):1518-25
references: 19357407
references: Genes Dev. 2013 Feb 1;27(3):234-50
references: 23388824
references: Mol Imaging Biol. 2006 Jan-Feb;8(1):24-9
references: 16333529
references: Endocrinology. 2002 May;143(5):1741-7
references: 11956156
references: J Clin Endocrinol Metab. 2013 Jul;98(7):E1218-23
references: 23780370
references: Cell Rep. 2014 Dec 11;9(5):1584-93
references: 25466254
references: Endocrinology. 2004 Jun;145(6):2594-603
references: 14976145
references: Cell Metab. 2013 Apr 2;17(4):562-74
references: 23499423
references: Nature. 2013 Dec 5;504(7478):163-7
references: 24196706
references: Nat Cell Biol. 2013 Dec;15(12):1455-63
references: 24212090
references: Proc Natl Acad Sci U S A. 2012 Apr 10;109(15):5874-9
references: 22451920
references: FASEB J. 2009 Sep;23(9):3113-20
references: 19417078
references: Int J Obes (Lond). 2011 Aug;35(8):1050-5
references: 21189472
references: Diabetologia. 1998 Mar;41(3):357-61
references: 9541178
references: J Biol Chem. 2010 Mar 5;285(10):7153-64
references: 20028987
references: Cell Metab. 2014 Sep 2;20(3):433-47
references: 25043816
references: Cell Metab. 2012 Mar 7;15(3):395-404
references: 22405074
references: Nat Med. 2013 May;19(5):631-4
references: 23603813