The Tissue and Cell Specific Proteome

All, 20162, human genes are classified according to their expression across all tissue and cell types in the human body. Few of the genes are strictly tissue or cell type specific, however, the genes with an elevated expression in particular tissues or cell types are interesting as a starting point to understand their biology and function, and underlying mechanisms for disease.

Out of the 10930 genes classified as elevated in at least one tissue type, 10109 genes are also classified as elevated in at least one cell type. By comparing the two datasets and gene classifications we can increase the resolution and understadning of tissue and cell specific gene expression. Although the sampled tissues are not a complete overlap, the majority of tissue and cell types are represented.

  • A total of 10930 genes show elevated expression in at least one of the analyzed tissues, included in the Tissue Atlas
  • A total of 15720 genes show elevated expression in at least one of the analyzed tissues, included in the Single Cell Type Atlas


Transcriptome analysis of all main tissue and cell types in the human body can be visualized with regard to specificity and distribution of transcribed mRNA molecules across all 20162 protein coding genes. Specificity illustrates the number of genes with elevated or non-elevated expression in a particular tissue or cell type compared to other tissues and cell types. The analysis includes 10930 genes classified as elevated in at least one tissue type and 15720 genes classified as elevated in at least one cell type. In total, 2826genes are classified as low tissue and cell type specificity, out of which 1887 are detected in all tissue and cell types represented (read more in The housekeeping proteome).

The elevated expression categories

Elevated expression includes three subcategory types of elevated expression:

  • Tissue enriched: At least four-fold higher mRNA level in a particular tissue/cell type compared to any other tissues/cell types.
  • Group enriched: At least four-fold higher average mRNA level in a group of 2-5 tissues or 2-10 cell types compared to any other tissues/cell types.
  • Tissue enhanced: At least four-fold higher mRNA level in a particular tissue/cell type compared to the average level in all other tissues/cell types.

Comparing tissue elevated with cell type elevated

The gene classification strategy defining elevated gene expression is applied to all datasets of the Human Protein Atlas. Here, genes with an elevated expression profile in tissue types and cell types are compared. Cell type specific expression patterns are often overlapping with tissue type elevated expression, however, it is not a rule. Cell types that can be found in all tissue types, such as vascular cells or connective tissue are examples of cell type specific expression that are more general when comparing expression levels between tissue types. In addition, proteins with a general function detected in all cells and tissues can also be associated with a specific cell type with higher activity of that biological process, such is the case with secretion, transportation or signaling molecules that has a general function throuout the body but hihgly expressed selected cell types.


Table 1.The number of genes in the respective categories of elevated expression. Comparing tissue elevated expression based on 36 tissues types with the cell type elevated expression based on 81 cell types (representing 31 tissue types).

Tissue specificity
Tissue enrichedGroup enrichedTissue enhancedLow tissue specificityNot detected Total
Cell type
Cell type enriched 95128561916886 2109
Group enriched 99449988329165 2732
Cell type enhanced 91270142654847154 10879
Low cell type specificity 20721728262 3072
Not detected 2608719148704 1290
Total 31371579617581801011 20082

The single cell type expression profiles are based on 81 cell types grouped into 15 different cell type groups, based on data representing 31 different tissue types. The tissues represented in the Single Cell Resource overlap but not completely cover the 36 tissue types profiled in the Tissue Resource. The tissue types currently lacking representation in the single cell type expression profiles are gallbladder, epididymis, seminal vesicle, urinary bladder, vagina, cervix, adrenal gland, parathyroid gland, thyroid gland and pituitary gland. While the brain representation in the aggregated single cell type overview is limited to the cerebral cortex and the eye is limited to the retina both in the cell type data as well as the tissue bulk data. These details are important to keep in mind when comparing the categories across the data sets.

Tissue and cell type enriched examples

Comparing gene expression across tissue types, 134 genes are classified as retina enriched, out of which 82 genes are enriched in either Bipolar cells, Horizontal cells, Cone -, or Rod photoreceptor cells of the retina. By comparing the tissue level classification with the cell type enrichment, not only can we determine the genes with retina elevated expression- but also in what retinal cells the genes are expressed. Among mentioned genes, we can for example find the opsin genes, essential for signaling state of light absorption, the three opsin1 genes with short (OPN1SW), medium (OPN1MW) or long (OPN1LW) wave length sensitivity, are all specifically expressed by the cone photoreceptors. Arrestin 3, ARR3, show similar expression profile as opsin 1, and has been detected on protein level using immunohistochemistry, in the Tissue Atlas. Cone-rod homeobox, CRX, on the other hand show group enriched expression profile, expressed and detected on protein level in both rod and cone photoreceptor cells, with lower expression also in bipolar cells. Compared to, Visual system homeobox 2 (VSX2) with enriched expression in bipolar cells and Muller glial cells,
ARR3

CRX

VSX2

ARR3 - Eye

CRX - Eye

VSX2 - Eye

Surfactant protein A1 (SFTPA1) and Surfactant protein B (SFTPB) are both involved in surfactant homeostasis and the defense against respiratory pathogens, they show enriched expression levels in lung compared to other tissue types, and when investigated on cell type level SFTPA1 show enrichment in Alveolar cell type 2, while SFTPB is group enriched, highly expressed by both alveolar cell type 1 and 2. In fact, most of the genes classified as lung enriched are specific for the alveolar cell types, one of the Macrophage scavenger receptor 1 (MSR1)


SFTPA1 - lung

SFTPB - lung

MSR1 - lung

SFTPA1 - Lung

SFTPB - Lung

MSR1 - Lung

Low tissue specificity and cell type elevated

There are 5015 genes classified as low tissue specificity, out of which 168 genes are classified as enriched in a cell type.

Multimerin 1 (MMRN1) is classified as low tissue specificity when comparing bulk RNAseq in tissue types, while on the cell type level is classified as cell type enriched, in lymphatic endothelial cells. MMRN1 is detected on protein level in the endothelial cells, and strong immunolabel is observed in the lymphatic endothelial cells. Similarly, the endothelial cell marker CD34 is classified as low tissue specificity while based on single cell type data shows a selective expression profile, with highest expression in adipocytes, endothelial cells and mensenchymal cells.


MMRN1

MMRN1 - Fallopian tube


CD34

CD34 - Fallopian tube


The Gap junction protein alpha 1(GJA1) is a transporter detected throughout the body, but not in all cell types. It is classified as low tissue specificity, as well as cell type elevated in a group of cell types, such as basal keratinocytes, alveolar cells, endothelial cells and astrocytes. GJA1 in the brain is highly expressed by astrocytes, followed by ependymal cells. This is an example of how combining the tissue specificity with the cell type specificity provides a comprehensive expression profile overview.


GJA1 - Cerebral cortex

GJA1

GJA1 - Fallopian tube

The postsynaptic density scaffolding protein homer scaffold protein 3 (HOMER3) is expressed by and located in Purkinje cells, and interneurons of the cerebellar molecular layer, in the brain. From one perspective it is described as a protein with high specificity, especially when exploring the expression profile in brain cell types. Cell types representing the whole body also tells the story of a rather selective expression profile, with elevated expression in lymphatic endothelial cells, proximal tubular cells, Hofbauer cells and Muller glial cells. Interestingly, when classifying based on tissue bulk data, it shows low tissue specificity, detected in all tissue types. However, high expression is observed in both the cerebellum and spleen (which is dense in lymphatic endothelial cells), and this can also be shown on protein level by immunolabeling.


HOMER3

HOMER3 - Cerebellum

HOMER3

Low cell type specificity and tissue elevated

244 genes are classified as low cell type specificity while also being high lighed as tissue elevated. Out of these, 20 are classified as tissue enriched.

There are no clear trends within this list of genes, but there are several examples of generally expressed proteins, such as AAMDC, ADSL, and KPNA4 detected in all cell types, and all tissue types with a higher expression in the skeletal muscle.

Low tissue specificity that is also low cell type specificity are often associated with housekeeping functions, especially in the case where they are detected by all tissue and cell types. Read more about the housekeeping proteome here.

Not detected while elevated expression in the other

As mentioned above, related to Table 1, it is not a 100% overlap between the samples used for profiling, some tissues are missing in the cell type profiling while present in the tissue type sequencing. Such as epididymis, adrenal gland and pituittary gland, all with numerous genes with elevated expression.

Additionally, single cell sequencing facilitates a good profiling also for sparse cells. GJA10 show elevated expression in Horizontal & Bipolar cells, while the expression in tissue bulk retina samples is just below the cut off- thus classified as not detected. The low abundance of expression in retina is no surprise since it is a limited number of cells in the tissue, while the cell type clustering enables a clear cell type expression profile.

Tissue elevated expression

The tissue specific expression profile of all human protein-coding genes can be explored from the perspective of each tissue type. In addition to the specificity category, distribution (visualizes how many genes have, or do not have, detectable levels (nTPM≥1) of transcribed mRNA molecules) is also defined for each tissue type:

  • Detected in single: Detected in a single tissue
  • Detected in some: Detected in more than one but less than one third of tissues
  • Detected in many: Detected in at least a third but not all tissues
  • Detected in all: Detected in all tissues

A. Specificity

B. Distribution

Figure 1. (A) The distribution of all genes across the five categories based on transcript specificity in all 36 analyzed tissues. (B) The distribution of all genes across the six categories based on transcript detection (nTPM≥1) in all 36 analyzed tissues.


Table 2.The number of genes in the subdivided categories of elevated expression in all 36 analyzed tissues.

Distribution in the 36 tissues
Detected in singleDetected in someDetected in manyDetected in all Total
Specificity
Tissue enriched 9241279753194 3150
Group enriched 0853609121 1583
Tissue enhanced 185109330541865 6197
Total 1109322544162180 10930

The amount of tissue elevated genes is highly variable between the analyzed tissue types (see Table 2 below). Testis shows the highest number of tissue enriched genes (n=937), followed by the brain (n=475) and liver (n=263). When taking into consideration all tissue elevated genes, the brain however has a slightly higher number than the testis. A large number of enriched genes in testis is considered to be due to the highly specialized processes occurring during spermatogenesis. Many of these genes likely have a shared expression with oocytes in the female ovaries. Oocytes are however difficult to analyze because of the complex kinetics of female germ cell development, including first rounds of meiosis, which in females occur at the embryonic stage. As expected, tissues that have similar functions and morphology often have higher numbers of shared group enriched genes.


Table 3. The tissue elevated genes for each of the 36 tissue types, shown for the different categories of elevated expression.

Tissue Tissue
enriched
Group
enriched
Tissue
enhanced
Total
elevated
Choroid plexus 33 133 281 447
Brain 475 457 1265 2197
Retina 134 240 411 785
Pituitary gland 25 115 139 279
Thyroid gland 13 27 134 174
Parathyroid gland 29 36 136 201
Adrenal gland 25 49 152 226
Lung 17 42 136 195
Salivary gland 43 80 177 300
Esophagus 20 75 337 432
Tongue 3 226 256 485
Stomach 35 81 205 321
Intestine 128 250 571 949
Liver 263 178 537 978
Gallbladder 3 16 69 88
Pancreas 64 74 173 311
Kidney 58 148 254 460
Urinary bladder 7 37 147 191
Testis 937 296 759 1992
Epididymis 96 73 146 315
Prostate 15 26 85 126
Seminal vesicle 6 13 54 73
Breast 20 37 75 132
Vagina 0 34 113 147
Cervix 0 40 136 176
Endometrium 3 13 73 89
Fallopian tube 19 114 177 310
Ovary 5 28 145 178
Placenta 67 47 179 293
Heart muscle 36 138 245 419
Skeletal muscle 59 269 593 921
Smooth muscle 0 7 43 50
Adipose tissue 4 34 195 233
Skin 184 99 319 602
Bone marrow 115 172 655 942
Lymphoid tissue 209 307 954 1470
Total 3150 1583 6197 10930


Examples of tissue elevated proteins

The comprehensive analysis presented here has identified 10930 human genes that display a tissue elevated expression pattern across the human body. By combining the analysis with antibody-based protein profiling using immunohistochemistry, the exact location of the corresponding protein expression pattern at a cellular and subcellular level can be provided. Examples of protein expression patterns of tissue elevated genes are presented below.

Brain

  • GFAP (Glial fibrillary acidic protein) - astrocyte intermediate filament protein
  • MBP (Myelin basic protein) - a major constituent of the myelin sheath
  • ELAVL3 (ELAV like RNA binding protein 3) - neural-specific RNA-binding protein


GFAP - cerebral cortex

MBP - hippocampus

ELAVL3 - cerebral cortex

Endocrine tissues

  • FSHB (Follicle stimulating hormone beta subunit) – hormone inducing egg and sperm production
  • TG (Thyroglobulin) - substrate for the synthesis of thyroid hormones
  • HSD3B2 (Hydroxy-delta-5-steroid dehydrogenase, 3 beta- and steroid delta-isomerase 2) - involved in the biosynthesis of hormonal steroids


FSHB - pituitary gland

TG - thyroid gland

HSD3B2 - adrenal gland

Gastrointestinal tract

  • PGA4 (Pepsinogen 4, group I (pepsinogen A)) - enzyme for digestion of dietary proteins
  • DEFA5 (Defensin alpha 5) - antimicrobial and cytotoxic peptide involved in host defense
  • KRT20 (Keratin 20) - maintains keratin filament organization in intestinal epithelia


PGA4 - stomach

DEFA5 - duodenum

KRT20 - colon

Liver & gallbladder

  • ALB (Albumin) - plasma protein
  • CYP2A13 (Cytochrome P450 member) - involved in drug metabolism, cholesterol and steroid synthesis
  • CHST4 (Carbohydrate sulfotransferase 4) - an enzyme involved in the modification of glycan structures


ALB - liver

CYP2A13 - liver

CHST4 - gallbladder

Pancreas

  • AMY2A (Amylase, alpha 2A) - an enzyme that digests carbohydrates, secreted by exocrine cells
  • INS (Insulin) - involved in lowering of blood glucose, secreted by beta cells
  • GCG (Glucagon) - involved in the elevation of blood glucose, secreted by alpha cells


AMY2A - pancreas

INS - pancreas

GCG - pancreas

Kidney & urinary bladder

  • SLC22A13 (Solute carrier family 22 member 13) - membrane-bound organic anion transporter
  • NPHS2 (Podocin) - involved in the regulation of glomerular permeability
  • UPK2 (Uroplakin 2) - membrane protein preventing cell rupture during bladder distention


SLC22A13 - kidney

NPHS2 - kidney

UPK2 - urinary bladder

Male tissues

  • DMRT1 (Doublesex- and mab-3-related transcription factor 1) - involved in meiosis
  • SEMG1 (Semenogelin I) - a predominant protein in semen
  • KLK3 (Kallikrein related peptidase 3) - also called PSA, is used clinically to diagnose prostate cancer


DMRT1 - testis

SEMG1 - seminal vesicle

KLK3 - prostate

Female tissues

  • CSH1 (Chorionic somatomammotropin hormone 1 ) - hormone important for growth control during pregnancy
  • OVGP1 (Oviductal glycoprotein 1) - mucus protein important in mucociliary transport of the fertilized ovum
  • PWWP3B (PWWP domain containing 3B) - a protein with a mutated melanoma-associated antigen 1 domain, associated with cancer


CSH1 - placenta

OVGP1 - fallopian tube

PWWP3B - ovary

Muscle tissues

  • TNNI3 (Troponin I3, cardiac type) - mediates muscle relaxation
  • TNNT2 (Troponin T2, cardiac type) - mediates muscle contraction
  • MYH7 (Myosin heavy chain 7) - expressed in slow type I muscle fibers


TNNI3 - heart muscle

TNNT2 - heart muscle

MYH7 - skeletal muscle

Skin

  • KRT1 (Keratin 1) - involved in squamous differentiation and skin barrier function
  • KRT27 (Keratin 27) - plays a role in hair formation
  • CASP14 (Caspase 14) - involved in keratinocyte differentiation and cornification


KRT1 - skin

KRT27 - hair

CASP14 - skin

Bone marrow & lymphoid tissues

  • MPO (Myeloperoxidase) - major component of neutrophil azurophilic granules
  • CD8B (CD8b molecule) - plays a critical role in thymic selection of CD8+ T-cells
  • CD22 (CD22 molecule) - mediates interactions between B-cells


MPO - bone marrow

CD8B - thymus

CD22 - lymph node

Cell type elevated expression

The Single Cell type specific expression profile of all human protein-coding genes can be explored from the perspective of each grouped cell type. In addition to the specificity category, distribution (visualizes how many genes have, or do not have, detectable levels (nTPM≥1) of transcribed mRNA molecules) is also defined for each single cell type:

  • Detected in single: Detected in a single cell type
  • Detected in some: Detected in more than one but less than one third of cell types
  • Detected in many: Detected in at least a third but not all cell types
  • Detected in all: Detected in all cell types

A. Specificity

{

B. Distribution

Figure 2. (A) The distribution of all genes across the five categories based on transcript specificity in all 81 analyzed cell types, grouped into 15 cell type groups and representing 31 tissue types. (B) The distribution of all genes across the six categories based on transcript detection (nTPM≥1) in all 81 analyzed cell types.


Table 4.The number of genes in the subdivided categories of elevated expression in all 81 analyzed tissues.

Distribution in the 81 cell types
Detected in singleDetected in someDetected in manyDetected in all Total
Specificity
Cell type enriched 2241292483110 2109
Group enriched 01728851153 2732
Cell type enhanced 141194161462651 10879
Total 365496174802914 15720

Similarly to the tissue elevated expression profiles, the germ cells show a distinct expression compared to other tissues and cells, here indicated by the highest number of elevated genes found in the group of germ cells, where Testis specific cell types are responsible for 4786 genes out of the 5400 genes with an elevated expression in germ cells.

Table 5. The tissue elevated genes for each of the 15 grouped cell types (from 31 different tissue types) representing 81 different tissue cell types, shown for the different categories of elevated expression.

Cell type group EnrichedGroup enrichedEnhancedTotal elevated
Glandular epithelial cells 24541328973555
Squamous epithelial cells 6198755914
Specialized epithelial cells 24432624242994
Endocrine cells 25101652778
Neuronal cells 13063230333795
Glial cells 17053430543758
Germ cells 575128435415400
Trophoblast cells 7812112071406
Endothelial cells 1073471554
Muscle cells 13616810901394
Adipocytes 1162412485
Pigment cells 1830212260
Mesenchymal cells 1078813901
Undifferentiated cells 317206226
Blood & immune cells 39333525063234

Examples of single cell type elevated proteins

The comprehensive analysis presented here has identified 10930 human genes that display a cell type elevated expression pattern. By combining the analysis with antibody-based protein profiling using immunohistochemistry, the exact location of the corresponding protein expression pattern at a cellular and subcellular level can be provided. Examples of protein expression patterns of cell type elevated genes are presented below.

Glandular epithelial cells

  • FOXJ1 (forkhead box J1) - transcription factor required for the production of motile cilia.
  • LPO (lactoperoxidase) - an enzyme that activates antimicrobial agents in the saliva.
  • MYLK (myosin light chain kinase) - involved in smooth muscle contraction in breast myoepithelial cells.


FOXJ1 - bronchus

LPO - salivary gland

MYLK - breast

Stratified squamous epithelial cells

  • KRT5 (keratin 5) - a type II cytokeratin which belongs to the keratin family of proteins and is involved in keeping the structural integrity of the basal cells together with keratin 14.
  • CASP14 (caspase 14) - play a role in keratinocyte differentiation and required for cornification
  • TP73 (Tumor protein p73) - a transcription factor participating in the apototic response to DNA damage.


KRT5 - skin

CASP14 - skin

TP73 - esophagus

Specialized epithelial cells

  • AQP4 (aquaporin 4) - a cell membrane-bound channel that regulates water homeostasis of the fluid lining the lung.
  • NOS1 (nitric oxide synthase 1) - a nitric oxide generating enzyme that has, among other functions, antimicrobial activity.
  • SLC12A1 (solute carrier family 12 member 1) - a transporters essential for regulating the contents and volume of urine


AQP4 - lung

NOS1 - salivary gland

SLC12A1 - kidney

Endocrine cells

  • PYY (peptide YY) - inhibits the secretion of digestive enzymes from the pancreas and peristaltic movements in the jejunum and colon
  • IAPP (islet amyloid polypeptide) - a hormone that regulates glucose metabolism and acts as a satiation signal.
  • INSL3 (insulin like 3) - a hormone which plays a role in reproductive tissue development.


PYY - rectum

IAPP - pancreas

INSL3 - testis

Germ cells

  • TSPY2 (testis-specific protein, Y-linked 2) - a transcription factor that plays a key role in male sex determination and differentiation by controlling testis development and male germ cell proliferation.
  • SPACA4 (sperm acrosome associated 4) - a sperm surface membrane protein that may be involved in sperm-egg plasma membrane adhesion and fusion during fertilization.
  • ZP3 (zona pellucida glycoprotein 3) - a component of the zona pellucida, a structure that is important for subsequent fertilization by a sperm cell.


TSPY2 - testis

SPACA4 - testis

ZP3 - ovary

Trophoblast cells

  • PEG10 (paternally expressed 10) - expressed by cytotrophoblasts and involved in cell proliferation, differentiation and apoptosis.
  • CSH2 (chorionic somatomammotropin hormone 2) - a hormone only produced during pregnancy, involved in stimulating lactation, fetal growth, and metabolism.
  • PAPPA2 (pregnancy and pappalysin 2) - a metalloproteinase thought to be a local regulator of insulin-like growth factor.


PEG10 - placenta

CSH2 - placenta

PAPPA2 - placenta

Muscle cells

  • ATP2A2 (ATPase sarcoplasmic/endoplasmic reticulum Ca2+ transporting 2) - an enzyme catalyzing the hydrolysis of ATP coupled with the translocation of calcium from the cytosol to the sarcoplasmic reticulum lumen.
  • MYH2 (myosin heavy chain 2) - expressed in fast twitch muscle fibers.
  • CNN1 (calponin 1) - a thin filament-associated protein that is implicated in the regulation and modulation of smooth muscle contraction.


ATP2A2 - heart muscle

MYH2 - skeletal muscle

CNN1 - prostate

Mesenchymal cells

  • CD248 (CD248 molecule) - a glycoprotein that is implicated in the fibrotic process.
  • ACTA2 (smooth muscle alpha actin) - a protein highly expressed in the peritubular structure.
  • HOXA11 (homeobox A11) - play an important role in proliferation, differentiation and receptivity of the tissue.


CD248

ACTA2 - testis

HOXA11 - endometrium

Blood and immune cells

  • THEMIS (Thymocyte Selection Associated) - a protein involved in the late phases of T-cell development.
  • MS4A1 (Membrane Spanning 4-Domains A1) - expressed on the surface of B-cells during maturation and absent in early pro-B cells.
  • MS4A2 (Membrane spanning 4-domains A2) - a membrane receptor in basophils and mast cells that binds to the FC region of IgE antibodies in order to enable the degranulation response where enzymes and other pro-inflammatory mediators are released to fight infection.


THEMIS - thymus

MS4A1 - lymph node

MS4A2 - tonsil

Among the blood and immune cells, there are tissue specific macrophages, assigned to specific tissues.

  • Hofbauer cells, CD68 - a transmembrane glycoprotein that is highly expressed by human monocytes and tissue macrophages.
  • Kupffer cells, CD209 - a C-type lectin receptor that binds to pathogen associated molecular patterns on various microorganisms and this binding activates phagocytosis.
  • Langerhans cells, CD1A - a transmembrane glycoprotein with structural similarities to MHC class I.


CD68 - placenta

CD209 - liver

CD1A - skin

Exploring the list of immune cells and the elevated genes across tissue types, there are numerous examples of immune cell specific expression that show low tissue specificity, due to being infiltrating cells found in most tissue types throughout the body. Above are examples of immune cells found in tissues but also being elevated on tissue level. The expression of tryptase beta 2 (TPSB2) show low tissue specificity, but a strong association to immune cells in different peripheral tissue types, such as the intestine.


TPSB2

TPSB2 - Small intestine


Relevant links and publications

Tissue specific

Uhlén M et al., Tissue-based map of the human proteome. Science (2015)
PubMed: 25613900 DOI: 10.1126/science.1260419

Bergman J et al., The human adrenal gland proteome defined by transcriptomics and antibody-based profiling. Endocrinology. (2016)
PubMed: 27901589 DOI: 10.1210/en.2016-1758

Edqvist PH et al., Expression of human skin-specific genes defined by transcriptomics and antibody-based profiling. J Histochem Cytochem. (2015)
PubMed: 25411189 DOI: 10.1369/0022155414562646

Lindskog C et al., The human cardiac and skeletal muscle proteomes defined by transcriptomics and antibody-based profiling. BMC Genomics. (2015)
PubMed: 26109061 DOI: 10.1186/s12864-015-1686-y

Sjöstedt E et al., Defining the Human Brain Proteome Using Transcriptomics and Antibody-Based Profiling with a Focus on the Cerebral Cortex. PLoS One. (2015)
PubMed: 26076492 DOI: 10.1371/journal.pone.0130028

Zieba A et al., The Human Endometrium-Specific Proteome Defined by Transcriptomics and Antibody-Based Profiling. OMICS. (2015)
PubMed: 26488136 DOI: 10.1089/omi.2015.0115

O'Hurley G et al., Analysis of the Human Prostate-Specific Proteome Defined by Transcriptomics and Antibody-Based Profiling Identifies TMEM79 and ACOXL as Two Putative, Diagnostic Markers in Prostate Cancer. PLoS One. (2015)
PubMed: 26237329 DOI: 10.1371/journal.pone.0133449

Habuka M et al., The Urinary Bladder Transcriptome and Proteome Defined by Transcriptomics and Antibody-Based Profiling. PLoS One. (2015)
PubMed: 26694548 DOI: 10.1371/journal.pone.0145301

Andersson S et al., The transcriptomic and proteomic landscapes of bone marrow and secondary lymphoid tissues. PLoS One. (2014)
PubMed: 25541736 DOI: 10.1371/journal.pone.0115911

Habuka M et al., The kidney transcriptome and proteome defined by transcriptomics and antibody-based profiling. PLoS One. (2014)
PubMed: 25551756 DOI: 10.1371/journal.pone.0116125

Mardinoglu A et al., Defining the human adipose tissue proteome to reveal metabolic alterations in obesity. J Proteome Res. (2014)
PubMed: 25219818 DOI: 10.1021/pr500586e

Kampf C et al., Defining the human gallbladder proteome by transcriptomics and affinity proteomics. Proteomics. (2014)
PubMed: 25175928 DOI: 10.1002/pmic.201400201

Lindskog C et al., The lung-specific proteome defined by integration of transcriptomics and antibody-based profiling. FASEB J. (2014)
PubMed: 25169055 DOI: 10.1096/fj.14-254862

Gremel G et al., The human gastrointestinal tract-specific transcriptome and proteome as defined by RNA sequencing and antibody-based profiling. J Gastroenterol. (2015)
PubMed: 24789573 DOI: 10.1007/s00535-014-0958-7

Kampf C et al., The human liver-specific proteome defined by transcriptomics and antibody-based profiling. FASEB J. (2014)
PubMed: 24648543 DOI: 10.1096/fj.14-250555

Djureinovic D et al., The human testis-specific proteome defined by transcriptomics and antibody-based profiling. Mol Hum Reprod. (2014)
PubMed: 24598113 DOI: 10.1093/molehr/gau018

Fagerberg L et al., Analysis of the human tissue-specific expression by genome-wide integration of transcriptomics and antibody-based proteomics. Mol Cell Proteomics. (2014)
PubMed: 24309898 DOI: 10.1074/mcp.M113.035600

Danielsson A et al., The human pancreas proteome defined by transcriptomics and antibody-based profiling. PLoS One. (2014)
PubMed: 25546435 DOI: 10.1371/journal.pone.0115421

Uhlén M et al., Tissue-based map of the human proteome. Science (2015)
PubMed: 25613900 DOI: 10.1126/science.1260419

Bergman J et al., The human adrenal gland proteome defined by transcriptomics and antibody-based profiling. Endocrinology. (2016)
PubMed: 27901589 DOI: 10.1210/en.2016-1758

Edqvist PH et al., Expression of human skin-specific genes defined by transcriptomics and antibody-based profiling. J Histochem Cytochem. (2015)
PubMed: 25411189 DOI: 10.1369/0022155414562646

Lindskog C et al., The human cardiac and skeletal muscle proteomes defined by transcriptomics and antibody-based profiling. BMC Genomics. (2015)
PubMed: 26109061 DOI: 10.1186/s12864-015-1686-y

Sjöstedt E et al., Defining the Human Brain Proteome Using Transcriptomics and Antibody-Based Profiling with a Focus on the Cerebral Cortex. PLoS One. (2015)
PubMed: 26076492 DOI: 10.1371/journal.pone.0130028

Zieba A et al., The Human Endometrium-Specific Proteome Defined by Transcriptomics and Antibody-Based Profiling. OMICS. (2015)
PubMed: 26488136 DOI: 10.1089/omi.2015.0115

O'Hurley G et al., Analysis of the Human Prostate-Specific Proteome Defined by Transcriptomics and Antibody-Based Profiling Identifies TMEM79 and ACOXL as Two Putative, Diagnostic Markers in Prostate Cancer. PLoS One. (2015)
PubMed: 26237329 DOI: 10.1371/journal.pone.0133449

Habuka M et al., The Urinary Bladder Transcriptome and Proteome Defined by Transcriptomics and Antibody-Based Profiling. PLoS One. (2015)
PubMed: 26694548 DOI: 10.1371/journal.pone.0145301

Andersson S et al., The transcriptomic and proteomic landscapes of bone marrow and secondary lymphoid tissues. PLoS One. (2014)
PubMed: 25541736 DOI: 10.1371/journal.pone.0115911

Habuka M et al., The kidney transcriptome and proteome defined by transcriptomics and antibody-based profiling. PLoS One. (2014)
PubMed: 25551756 DOI: 10.1371/journal.pone.0116125

Mardinoglu A et al., Defining the human adipose tissue proteome to reveal metabolic alterations in obesity. J Proteome Res. (2014)
PubMed: 25219818 DOI: 10.1021/pr500586e

Kampf C et al., Defining the human gallbladder proteome by transcriptomics and affinity proteomics. Proteomics. (2014)
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