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Atlas tissue group Human Homo sapiens

brain

The Human Protein Atlas classes 2,227 genes as elevated in brain (475 tissue enriched, 439 group enriched, 1,313 tissue enhanced), in Human Protein Atlas version 25.1 (release 2026-05-25); entry tag 25, read 2026-09-09. The values are the atlas's own nTPM, which is not TPM; nothing on this page is compared with a GTEx value.

The atlas's own pages

The elevated genes came from the atlas's search field for the identical expression the link below carries, so the page it opens lists the rows this one holds.

The same search at the atlas

01The elevated genes

Genes the atlas classes as elevated in brain

What this tells you

The genes are the atlas's own specificity classification for this tissue group, read from its search field on 2026-09-09 and built into this site. The atlas release is version 25.1 (release 2026-05-25); entry tag 25: the entry tag is read from the one machine-readable statement the atlas makes of its version, the entry tag of its per-gene record, in the same build, and the version number and release date beside it are the atlas's release history page's [R30], quoted below. The atlas's consensus table lists this group's sub-tissues separately and carries no row under the group's own name, so no consensus card is shown.

The categories, as the atlas defines them [R29]. Enriched: nTPM in a particular tissue/region/cell type at least four times any other tissue/region/cell type [R29]. Group enriched: nTPM in a group (of 2-5 tissues, brain regions, single cell types or cell lines, or 2-10 immune cell types) at least four times any other tissue/region/cell line/immune cell type/cell type [R29]. Enhanced: nTPM in a one or several tissues, brain regions, cell lines, immune cell types or single cell types that has at least four times the mean of all tissue/region/cell types [R29]. The score beside an enriched or group-enriched gene: TS/CS-score is calculated as the fold change from the tissue/cell line with highest RNA to the tissue/cell line with second highest RNA. [R29] The atlas publishes none for a tissue-enhanced gene, and the table shows none.

The unit is the atlas's own: all TPM values of all samples within each data source (HPA + GTEx human tissues, HPA immune cell types, HPA cell lines) were normalized separately using Trimmed mean of M values (TMM) to allow for between-sample comparisons. The resulting normalized transcript expression values, denoted nTPM, were calculated for each gene in every sample. nTPM values below 0.1 are not visualized on the Atlas sections. [R29] nTPM is not TPM, and no value here is put beside a GTEx value from this site's GTEx pages. The consensus value is a maximum, never an average: The consensus nTPM value for each gene and tissue type represents the maximum nTPM value based on HPA and GTEx. For tissues with multiple sub-tissues (brain regions, immune cells, lymphoid tissues and intestine) the maximum of all sub-tissues is used for the tissue type and the total number of tissue types in the human tissue consensus set is 37. [R29]

The release: Protein Atlas version 25.1. Release date: 2026.05.25. Ensembl version: 109. [R30] Over the whole atlas, its tissue resource says all putative 20162 protein coding genes have been classified with regard to abundance and distribution of transcribed mRNA molecules, including 11035 proteins showing a significantly elevated level of expression in a particular tissue or a group of related tissues and 8813 proteins detected in all organs and tissues [R31], and its specificity counts on that page are 3132 tissue enriched, 1547 group enriched, 6356 tissue enhanced, 8096 of low tissue specificity and 1031 not detected [R31]. The atlas is licensed under the Creative Commons Attribution 4.0 International License for all copyrightable parts of our database [R28] and asks a website to cite the source in a manner that is clear, accurate and easily discoverable and link to the source [R28], which the provenance line under each card does, naming its primary publication [R07] and the versioned address the data came from.

An elevated gene here is one the atlas classes as elevated in this group by its own thresholds over its own consensus values, of which the GTEx column is GTEx v8 folded into nTPM; a value on this page and a TPM on this site's GTEx pages are two measurements of two sample sets in two units.

  1. [R07] Uhlén M, Fagerberg L, Hallström BM, Lindskog C, Oksvold P, Mardinoglu A, et al. (2015). Tissue-based map of the human proteome. Science 347:1260419. PMID 25613900, doi 10.1126/science.1260419.
  2. [R28] The Human Protein Atlas, proteinatlas.org. Licence & Citation. https://www.proteinatlas.org/about/licence, read 2026-09-09.
  3. [R29] The Human Protein Atlas, proteinatlas.org. The human proteome, Methods summary, Transcriptomics. https://www.proteinatlas.org/humanproteome/tissue/method/transcriptomics, read 2026-09-09.
  4. [R30] The Human Protein Atlas, proteinatlas.org. Release history. https://www.proteinatlas.org/about/releases, read 2026-09-09.
  5. [R31] The Human Protein Atlas, proteinatlas.org. Tissue resource, Tissue-based map of the human proteome. https://www.proteinatlas.org/humanproteome/tissue, read 2026-09-09.

The atlas classes 439 genes as group enriched in brain; showing 301 to 400 in pages of 100, in the atlas's own order. The nTPM is the atlas's own unit.

Genes the Human Protein Atlas classes as group enriched in brain, page 4 of 5
GenePPEF1EnsemblENSG00000086717TS-score6nTPM8.9Elevated inbrain 8.9; breast 3.9; testis 13.1
GenePPFIA2EnsemblENSG00000139220TS-score5nTPM34.3Elevated inbrain 34.3; retina 46.4
GenePPM1EEnsemblENSG00000175175TS-score7nTPM11.7Elevated inadrenal gland 5.1; brain 11.7; retina 20.7; testis 6.2
GenePPP1R1BEnsemblENSG00000131771TS-score4nTPM1,368.7Elevated inbrain 1,368.7; choroid plexus 1,463.9
GenePPP2R2BEnsemblENSG00000156475TS-score5nTPM59.7Elevated inbrain 59.7; retina 39.7; testis 17.6
GenePRR36EnsemblENSG00000183248TS-score8nTPM46Elevated inbrain 46; pituitary gland 27.8
GenePRRT3EnsemblENSG00000163704TS-score4nTPM13.9Elevated inbrain 13.9; choroid plexus 7; fallopian tube 7.1; parathyroid gland 14.3; pituitary gland 10.4
GenePSRC1EnsemblENSG00000134222TS-score5nTPM91.4Elevated inbrain 91.4; retina 38.7
GenePTPRNEnsemblENSG00000054356TS-score5nTPM157.2Elevated inbrain 157.2; pituitary gland 178
GenePYDC1EnsemblENSG00000169900TS-score13nTPM25.5Elevated inbrain 25.5; skin 1 22
GeneQDPREnsemblENSG00000151552TS-score5nTPM782Elevated inbrain 782; liver 209.2
GeneRAPGEF5EnsemblENSG00000136237TS-score4nTPM158.3Elevated inbrain 158.3; parathyroid gland 364.8
GeneRBFOX3EnsemblENSG00000167281TS-score5nTPM273.5Elevated inbrain 273.5; intestine 81.6; urinary bladder 78.6
GeneRELNEnsemblENSG00000189056TS-score7nTPM26.2Elevated inbrain 26.2; liver 12.6
GeneRFPL2EnsemblENSG00000128253TS-score5nTPM4.3Elevated inbrain 4.3; prostate 3.4
GeneRFX4EnsemblENSG00000111783TS-score5nTPM22.2Elevated inbrain 22.2; testis 70.2
GeneRGS11EnsemblENSG00000076344TS-score5nTPM164Elevated inbrain 164; pituitary gland 52.6
GeneRGS8EnsemblENSG00000135824TS-score4nTPM26.7Elevated inbrain 26.7; retina 28.2
GeneRIMS2EnsemblENSG00000176406TS-score5nTPM41.6Elevated inadrenal gland 45.5; brain 41.6; retina 72.6
GeneRIPPLY2EnsemblENSG00000203877TS-score5nTPM32.4Elevated inbrain 32.4; pituitary gland 10.6
GeneRNF165EnsemblENSG00000141622TS-score4nTPM21.7Elevated inbrain 21.7; retina 16.9
GeneRNF220EnsemblENSG00000187147TS-score5nTPM85.5Elevated inbrain 85.5; skeletal muscle 22.8
GeneRNFT2EnsemblENSG00000135119TS-score5nTPM10.3Elevated inbrain 10.3; choroid plexus 4.7; pituitary gland 4.9; retina 4.6; testis 6.5
GeneRTN4RL2EnsemblENSG00000186907TS-score8nTPM34Elevated inbrain 34; liver 37.8; thyroid gland 28.5
GeneRUNDC3AEnsemblENSG00000108309TS-score7nTPM297.6Elevated inbrain 297.6; pituitary gland 94.7
GeneS100A1EnsemblENSG00000160678TS-score5nTPM1,266.4Elevated inbrain 1,266.4; heart muscle 2,882.1; skeletal muscle 2,868.7; tongue 4,577.6
GeneSALL3EnsemblENSG00000256463TS-score4nTPM8.5Elevated inbrain 8.5; cervix 2.2; kidney 2.8; prostate 5.2; vagina 6.8
GeneSATB2EnsemblENSG00000119042TS-score5nTPM13.1Elevated inbrain 13.1; intestine 30.8
GeneSBK1EnsemblENSG00000188322TS-score4nTPM43Elevated inbrain 43; skeletal muscle 41.1
GeneSCG2EnsemblENSG00000171951TS-score7nTPM391.8Elevated inadrenal gland 340; brain 391.8; pituitary gland 267
GeneSCHIP1EnsemblENSG00000151967TS-score8nTPM76.6Elevated inbrain 76.6; heart muscle 74.5; skeletal muscle 92.5; tongue 86.3
GeneSCN8AEnsemblENSG00000196876TS-score4nTPM16.9Elevated inbrain 16.9; pituitary gland 6.3; retina 7.3
GeneSCRT2EnsemblENSG00000215397TS-score30nTPM3.9Elevated inbrain 3.9; retina 3.6
GeneSEC14L5EnsemblENSG00000103184TS-score4nTPM35.1Elevated inbrain 35.1; tongue 10.2
GeneSEPTIN4EnsemblENSG00000108387TS-score5nTPM1,097.2Elevated inbrain 1,097.2; retina 375.4
GeneSERPINI1EnsemblENSG00000163536TS-score5nTPM303.7Elevated inbrain 303.7; retina 78.2
GeneSH3GL2EnsemblENSG00000107295TS-score6nTPM227.8Elevated inbrain 227.8; retina 69.5
GeneSH3GL3EnsemblENSG00000140600TS-score5nTPM56.9Elevated inbrain 56.9; testis 24.8
GeneSH3TC2EnsemblENSG00000169247TS-score7nTPM4.5Elevated inbrain 4.5; retina 3.6; testis 1.4
GeneSHDEnsemblENSG00000105251TS-score7nTPM12.7Elevated inbrain 12.7; heart muscle 21.3
GeneSHISAL2BEnsemblENSG00000145642TS-score9nTPM3.3Elevated inbrain 3.3; pancreas 4.6; stomach 1 2.7
GeneSIRT2EnsemblENSG00000068903TS-score4nTPM492.9Elevated inbrain 492.9; skeletal muscle 306.9; tongue 203.7
GeneSLC17A7EnsemblENSG00000104888TS-score32nTPM385.5Elevated inbrain 385.5; retina 292.9
GeneSLC1A3EnsemblENSG00000079215TS-score8nTPM527.4Elevated inbrain 527.4; retina 204.7
GeneSLC22A17EnsemblENSG00000092096TS-score5nTPM276.8Elevated inbrain 276.8; choroid plexus 234.8; pituitary gland 178.3
GeneSLC25A18EnsemblENSG00000182902TS-score6nTPM139Elevated inbrain 139; liver 106.7
GeneSLC25A34EnsemblENSG00000162461TS-score4nTPM15.3Elevated inbrain 15.3; heart muscle 13.4; intestine 13; skeletal muscle 36.5; tongue 43.5
GeneSLC25A48EnsemblENSG00000145832TS-score4nTPM14.8Elevated inbrain 14.8; kidney 6.9; skeletal muscle 6.3
GeneSLC30A3EnsemblENSG00000115194TS-score6nTPM30.3Elevated inbrain 30.3; epididymis 64.6; testis 32
GeneSLC35F4EnsemblENSG00000151812TS-score6nTPM13.2Elevated inbrain 13.2; retina 6; seminal vesicle 3.3
GeneSLC4A10EnsemblENSG00000144290TS-score21nTPM32.6Elevated inbrain 32.6; choroid plexus 54; retina 31.8
GeneSLC4A8EnsemblENSG00000050438TS-score4nTPM12Elevated inbrain 12; pituitary gland 16.9; retina 10.3; testis 11
GeneSLC6A1EnsemblENSG00000157103TS-score4nTPM85.6Elevated inbrain 85.6; liver 82.4; retina 34.6
GeneSLC6A17EnsemblENSG00000197106TS-score5nTPM53.7Elevated inbrain 53.7; pituitary gland 23.8
GeneSLC7A10EnsemblENSG00000130876TS-score14nTPM20.1Elevated inadipose tissue 30.2; brain 20.1; breast 13
GeneSLC7A14EnsemblENSG00000013293TS-score4nTPM15Elevated inbrain 15; pituitary gland 6.7; retina 5.7
GeneSLC8A3EnsemblENSG00000100678TS-score10nTPM8.6Elevated inbrain 8.6; retina 9.4; skeletal muscle 19; tongue 8.7
GeneSLC9A5EnsemblENSG00000135740TS-score5nTPM15.1Elevated inbrain 15.1; lymphoid tissue 17.8
GeneSLCO1C1EnsemblENSG00000139155TS-score6nTPM19.6Elevated inbrain 19.6; choroid plexus 16.3; pituitary gland 7.7
GeneSLIT1EnsemblENSG00000187122TS-score9nTPM32.7Elevated inbrain 32.7; pituitary gland 12.6
GeneSLITRK2EnsemblENSG00000185985TS-score4nTPM11.5Elevated inbrain 11.5; retina 30.4
GeneSMIM10L2AEnsemblENSG00000178947TS-score5nTPM38.6Elevated inadrenal gland 28.8; brain 38.6; epididymis 13.6
GeneSMIM10L2BEnsemblENSG00000196972TS-score4nTPM38.2Elevated inbrain 38.2; heart muscle 10.2
GeneSMIM18EnsemblENSG00000253457TS-score6nTPM8.8Elevated inbrain 8.8; pituitary gland 5; retina 3.4
GeneSNCAEnsemblENSG00000145335TS-score6nTPM263.1Elevated inbone marrow 257.8; brain 263.1
GeneSNCBEnsemblENSG00000074317TS-score25nTPM562.7Elevated inbrain 562.7; retina 169.1
GeneSNORCEnsemblENSG00000182600TS-score4nTPM88.1Elevated inbrain 88.1; skin 1 31.3
GeneSOCS7EnsemblENSG00000274211TS-score4nTPM42.7Elevated inbrain 42.7; testis 69.2
GeneSOHLH1EnsemblENSG00000165643TS-score8nTPM46.2Elevated inbrain 46.2; testis 23.9
GeneSOWAHAEnsemblENSG00000198944TS-score15nTPM142.5Elevated inbrain 142.5; intestine 37
GeneSOX10EnsemblENSG00000100146TS-score5nTPM151.8Elevated inbrain 151.8; salivary gland 105.2
GeneSOX21EnsemblENSG00000125285TS-score6nTPM12.1Elevated inbrain 12.1; esophagus 9.6; skin 1 15; stomach 1 33.1
GeneSPEF1EnsemblENSG00000101222TS-score6nTPM27.6Elevated inbrain 27.6; choroid plexus 23.5; fallopian tube 9.3; testis 11.9
GeneSPHKAPEnsemblENSG00000153820TS-score12nTPM33.6Elevated inbrain 33.6; heart muscle 27.3
GeneSPRNEnsemblENSG00000203772TS-score6nTPM12.6Elevated inbrain 12.6; liver 7.3
GeneSPTBN4EnsemblENSG00000160460TS-score5nTPM24.8Elevated inbrain 24.8; pituitary gland 7.4
GeneSPTBN5EnsemblENSG00000137877TS-score6nTPM7.1Elevated inbrain 7.1; retina 25
GeneSRGAP2BEnsemblENSG00000196369TS-score4nTPM55.9Elevated inbrain 55.9; skin 1 15.1
GeneSRRM3EnsemblENSG00000177679TS-score6nTPM77.8Elevated inbrain 77.8; retina 25.2
GeneSRRM4EnsemblENSG00000139767TS-score26nTPM44.3Elevated inbrain 44.3; retina 27
GeneSSTEnsemblENSG00000157005TS-score12nTPM622Elevated inbrain 622; intestine 363.7; pancreas 479.7; stomach 1 416.6
GeneST18EnsemblENSG00000147488TS-score6nTPM21.9Elevated inbrain 21.9; retina 7.5
GeneST6GAL2EnsemblENSG00000144057TS-score5nTPM12.5Elevated inblood vessel 33.5; brain 12.5; thyroid gland 27.2
GeneST8SIA5EnsemblENSG00000101638TS-score5nTPM23.2Elevated inadrenal gland 5.9; brain 23.2; parathyroid gland 5.8
GeneSTUMEnsemblENSG00000203685TS-score4nTPM47.5Elevated inbrain 47.5; intestine 15.2
GeneSTX1AEnsemblENSG00000106089TS-score12nTPM169.5Elevated inbrain 169.5; pituitary gland 94.1
GeneSV2BEnsemblENSG00000185518TS-score23nTPM27.3Elevated inbrain 27.3; retina 52.2
GeneSVOPEnsemblENSG00000166111TS-score10nTPM25.2Elevated inbrain 25.2; pituitary gland 7.2; retina 12.1
GeneSYCE1EnsemblENSG00000171772TS-score5nTPM17.7Elevated inbrain 17.7; testis 69.5
GeneSYT1EnsemblENSG00000067715TS-score10nTPM280.3Elevated inbrain 280.3; retina 187.2
GeneSYT12EnsemblENSG00000173227TS-score5nTPM49.4Elevated inbrain 49.4; liver 21.4; parathyroid gland 60.1
GeneSYT5EnsemblENSG00000129990TS-score9nTPM106Elevated inbrain 106; pituitary gland 55.4
GeneTACR3EnsemblENSG00000169836TS-score8nTPM1.2Elevated inbrain 1.2; retina 2.5; urinary bladder 1.7
GeneTAFA4EnsemblENSG00000163377TS-score8nTPM6.9Elevated inadrenal gland 14.1; brain 6.9; retina 10.3; stomach 1 4.6
GeneTBC1D3HEnsemblENSG00000274226TS-score5nTPM1.7Elevated inbrain 1.7; testis 4.4
GeneTCEAL6EnsemblENSG00000204071TS-score4nTPM95.6Elevated inbrain 95.6; pituitary gland 67.7; testis 25.1
GeneTCTE1EnsemblENSG00000146221TS-score9nTPM7.9Elevated inbrain 7.9; fallopian tube 13.6; testis 28.4
GeneTESPA1EnsemblENSG00000135426TS-score5nTPM79.7Elevated inbrain 79.7; lymphoid tissue 70
GeneTFAP2EEnsemblENSG00000116819TS-score4nTPM9.7Elevated inbrain 9.7; retina 3.2
GeneTHEnsemblENSG00000180176TS-score50nTPM129.8Elevated inadrenal gland 185.6; brain 129.8

TS-score is the atlas's tissue specificity score, published for enriched and group-enriched genes and for no tissue-enhanced gene; none published is the atlas's absence, not a zero. The nTPM column is the value the atlas labels brain; a group-enriched gene lists every group the atlas names for it, with the atlas's own labels.

Pages of 100 are this site's own cut of the atlas's answer, which came whole; the categories are the three elevated categories the atlas defines, and its other two, low tissue specificity and not detected, are not tissue lists and are not shown.

  • Human Protein Atlas, the tissue specificity field for brain · Human Protein Atlas version 25.1 (release 2026-05-25); entry tag 25 · read · the same search at the atlasHuman Protein Atlas version 25.1 (release 2026-05-25); entry tag 25, proteinatlas.org; data available from v25.proteinatlas.org/search/tissue_category_rna:brain;tissue+enriched,group+enriched,tissue+enhanced (Uhlén M et al. Science 2015). CC BY 4.0.

02The consensus values

The atlas's consensus table

The atlas's consensus table carries no row named brain. In the atlas's own words, for tissues with multiple sub-tissues the maximum of all sub-tissues is used for the tissue type, and its table lists those sub-tissues under their own names rather than under this group's. No consensus value is shown here, and none is composed from the sub-tissues by this site.

The consensus nTPM value for each gene and tissue type represents the maximum nTPM value based on HPA and GTEx. For tissues with multiple sub-tissues (brain regions, immune cells, lymphoid tissues and intestine) the maximum of all sub-tissues is used for the tissue type and the total number of tissue types in the human tissue consensus set is 37.