Supplementary Table 1 | Summary of rodent brown, beige and white markers

Marker class / Markers identified from studies in adipocytes in vitro and whole tissue
White / *DPT19,21,31, *HOXC819,31,58, *IGFBP321,31, INHBB19,31, LEP125, RESISTIN20,46, *TCF2119,21,31
*DPT and HOXC8 are also expressed in iWAT, though their expression is downregulated in iWAT after cold exposure19,58
*IGFBP3 does not show differential expression in different AT depots19
*TCF21 is also expressed in iWAT19
Beige / CD13718, CITED120, *HOXC919–21, SHOX219, TBX118, TMEM2618
*HOXC9 is also expressed in white adipocytes20,21,31 and in WAT19
Brown / EBF318, EVA118,46, FBXO3118, LHX819–21,31,46, *miR-20619,21,38, miR-13332,59,60, *MEOX221,31, MYLPF19, *MYOGENIN21,31,32, *TBX1519,31, ZIC119–21,31,46
*miR-20619,21 and miR-13332,59,60 were detected in brown preadipocytes
*MEOX2 was detected in brown preadipocytes31, but does not show differential expression in different AT depots19
*MYOGENIN was detected in brown preadipocytes21,31,32
*TBX15 is also expressed in iWAT19

Experimental design of major studies: Ref. 18 compared brown adipocytes, beige adipocytes, and white adipocytes, all of which were differentiated from immortalized precursors. Ref. 19 compared multiple BAT and WAT depots, with inguinal WAT (iWAT) as a representative depot enriched with beige cells, especially after cold-exposure. Refs 20 and 21 compared brown adipocytes, white adipocytes, and beige adipocytes. Brown adipocytes were differentiated from BAT adipocyte precursors, white adipocytes were differentiated from WAT adipocyte precursors, and beige adipocytes were differentiated from WAT adipocyte precursors treated with PPARg agonists. Ref. 31 compared brown adipocytes to white adipocytes. Brown adipocytes were differentiated from BAT adipocyte precursors, and white adipocytes were differentiated from WAT adipocyte precursors. Ref. 46 compared BAT to WAT, and compared white to brown adipocytes which were differentiated from immortalized precursors.


Several studies have identified brown-, beige- or white-specific markers using gene expression analysis. Only markers which are discussed in more than one publication or which have been characterized in human BAT are included in this summary. See Fig.4 of the paper.

Supplementary Table 2 | Enrichment of brown, beige and white rodent markers in comparative studies of human AT

Human AT depot / Rodent markers found to be enriched in each AT depot
Subcutaneous WAT (ScWAT) / White markers: LEP105, *HOXC8103
Beige markers: *HOX9103,105
Brown markers: *EVA1105, *EBF3105, *FBXO31105
*HOXC8 and HOX9 were found to be enriched in supBAT compared to ScWAT in a different study20.
*EVA1103, EBF3103, and FBXO3118,103 did not show differential expression between supBAT and ScWAT in other studies.
Intermediate layers of neck fat / Beige markers: CD137105, SHOX2105, TMEM26105
Periadrenal BAT / Beige markers: HOXC9104
Brown markers: EVA1104
Supraclavicular BAT (supBAT) / Beige markers: *CD13718,20,105, CITED120, SHOX2104,105, TBX118,20,103,104, *TMEM2618,20,103,105
Brown markers: *LHX8103, miR-206103, miR-133103, *ZIC1103
*CD137 did not show differential expression between supBAT and ScWAT in a different study103.
*TMEM26 did not show differential expression between supBAT, periadrenal BAT, and infant iBAT depots in a different study104.
*LHX8 and ZIC1 were not detected in this depot in a different study20.
Deep layers of neck fat / Brown markers: LHX8105, ZIC1105
Infant interscapular BAT (iBAT) / Brown markers: ZIC1104

Experimental design of major studies: Refs 18 and 103 compared adult human supBAT to ScWAT. Ref. 20 compared multiple adult human BAT depots to ScWAT. Ref. 104 compared infant human iBAT, adult human periadrenal BAT, and adult human supBAT. Ref. 105 compared different layers of adult human neck fat, from subcutaneous to intermediate to deep. Results from factor analysis of gene expression are reported.


Several studies have compared the gene expression levels of rodent brown-, beige-, and white-specific markers in multiple human AT depots. See Fig. 4 of the paper.