Table 1.
Previously identified stromal cell subpopulations in individual studies.
| Stromal cell | Cancer type | Subpopulation | Markers | Feature | Ref |
|---|---|---|---|---|---|
| CAF | Breast cancer | CAF-S1 | CD29Med FAPHi FSP1Low-Hi αSMAHi PDGFRbMed-Hi CAV1Low |
- High expression of CCL11, CXCL12, 13, 14 - Myofibroblastic subset - Enriched in TNBC - Immunosuppressive - Increase T lymphocyte survival and differentiation - Associated with accumulation of FOXP3 + T lymphocytes - Correlated with CD45+ hematopoietic cells and macrophages - Anti-correlated with CD8 + T lymphocytes - Enhance Treg capacity to inhibit effector T cells |
136 |
| CAF-S2 | CD29Low FAPNeg FSP1Neg-Low αSMANeg PDGFRbNeg CAV1Neg | - Enriched in LumA BC | |||
| CAF-S3 | CD29Med FAPNeg FSP1Med-Hi αSMANeg-Low PDGFRbMed CAV1Neg-Low | - Associated with Juxta-tumor | |||
| CAF-S4 | CD29Hi FAPNeg FSP1Low-Med αSMAHi PDGFRbLow-Med CAV1Neg-Low |
- Enriched in TNBC and HER2 BC - Myofibroblastic subset - Associated with CD8 + T lymphocytes - Anti-correlated with FOXP3 + T lymphocyte |
|||
| Breast cancer |
CAF-S1 ECM-myCAF |
GJB2, LRRC15 |
- ECM-myofibroblastic CAF - Enriched in LumA BC - Involved in collagen synthesis and ECM organization |
38 | |
|
CAF-S1 Detox-iCAF |
ADH1B, GPX3 |
- Detoxification-inflammatory CAF - Enriched in TNBC - Involved in detoxification and inflammatory response |
|||
|
CAF-S1 IL-iCAF |
RGMA, SCARA5 |
- Response to stimuli - Enriched in TNBC - Involved in the response to growth factor, TNF signaling, and IL pathway |
|||
|
CAF-S1 TGFβ-myCAF |
CST1, TGFB1 |
- TGFβ-myofibroblastic CAF - Enriched in LumA BC - Involved in response to TGFβ stimulus and matrisome |
|||
|
CAF-S1 Wound-myCAF |
SEMA3C, SFRP4 |
- Wound healing-myofibroblastic CAF - Enriched in LumA BC - Involved in the assembly of collagen fibrils and wound healing - Correspond to apCAF |
|||
|
CAF-S1 IFNγ-iCAF |
CCL19, CCL5 | - IFNγ and cytokines | |||
|
CAF-S1 IFNαβ-iCAF |
IFIT3, IRF7 | - IFNαβ-inflammatory CAF | |||
|
CAF-S1 Acto-myCAF |
GGH, PLP2 | - Actomyosin-myofibroblastic CAF | |||
| CAF-S2 | FAPNeg CD29Low SMANeg | - Abundant in healthy tissue | |||
| CAF-S3 | FAPNeg CD29Med SMANeg | - Abundant in healthy tissue | |||
| CAF-S4 | FAPNeg SMAHi CD29Hi MCAMHi |
- Restricted to cancer and metastatic lymph nodes - Characterized by a perivascular signature - Pro-metastatic function - Contractile |
|||
| Breast cancer | vCAF | Rgs5 |
- Vascular CAF - Upregulated vascular development and angiogenesis genes - Enriched in tumor core - Localized in proximity to vasculature |
51 | |
| mCAF | Pdgfra, Mfap5, Dcn |
- Matrix CAF - Strong ECM signature - Upregulated ECM, matrisome, and EMT associated genes - Low abundance in the tumor core |
|||
| cCAF | Nuf2, Mki67 |
- Cycling CAF - Upregulated cell-cycle-related genes - Proliferative segments of vCAF |
|||
| dCAF | Scrg1, Sox9, Sox10 |
- Developmental CAF - Upregulated development and morphogenesis of tissue-associated genes - Originate from tumor cells that have undergone EMT |
|||
| PDAC | myCAF | ACTA2, TAGLN, MMP11, MYL9, HOPX, POSTN, TPM1, TPM2, ACTA2high |
- Myofibroblastic CAF - Adjacent to cancer cells - Associated with smooth muscle contraction, focal adhesion, ECM organization, and collagen formation |
137 | |
| iCAF | IL6, PDGFRA, CXCL12, CFD, DPT, LMNA, AGTR1, HAS1, CXCL1, CXCL2, CCL2, IL8, ACTA2low Ly6Chigh |
- Inflammatory CAF - High inflammatory mediators: IL6, IL11, LIF - Located in the desmoplastic areas of the tumor - Distant from cancer cells - Associated with the synthesis of hyaluronan and the complement pathway |
|||
| apCAF | H2-Ab1, Cd74, Saa3, Slpi |
- Antigen-presenting CAF - Express MHC class II-related genes - Induce TCR ligation in CD4 + T cells in an antigen-dependent manner |
|||
| Lung cancer | Cluster 1 | COL10A1 |
- Highly enriched in tumor - EMT-related signal - ECM phenotype - HOXB2 and FOXO1 are highly upregulated |
53 | |
| Cluster 2 | COL4A1 |
- The highest expression of ACTA2, a myofibroblast marker - Involved in myogenesis, NOTCH pathway, and angiogenesis - Myogenesis phenotype |
|||
| Cluster 4 | PLA2G2A |
- Similar to Cluster 1 - COL14A1high |
|||
| Cluster 5 | MMP3 |
- Low expression of myogenesis signature - High expression of mTOR signature and glycolysis genes |
|||
| Cluster 6 | FIGF |
- Nonmalignant fibroblast - High expression of elastin - Low expression of some collagens: collagen type I, III, V, and VIII |
|||
| Cluster 7 | CCL2 |
- Present in NSCLC patients - Similar to Cluster 5 but with low expression of glycolysis genes |
|||
| Gastric cancer | myCAF | TPM1, TPM2, MYL9, TAGLN, POSTN |
- Myofibroblastic CAF - Prevalent in intestinal-type GC - Negatively correlated with tumor stemness |
138 | |
| iCAF | IL6, IL11, IL24, CXCL1, CXCL2, CXCL5, CXCL6, MMP1, MMP3, MMP10 |
- Inflammatory CAF - Prevalent in diffuse-type GC - Associated with GC invasion - Promote stemness of tumor cells, high stemness score |
|||
| inCAF | PDGFRA, POSTN, ID1, ID3 |
- Intermediate CAF - Negatively correlated with tumor stemness - inCAF signal is increased with tumor progression from the premalignant state |
|||
| Colorectal cancer | CAF-A | FAP, MMP2, LUM, COL1A2 |
- Involved in ECM remodeling - Intermediate state between NMFs and CAF-B |
139 | |
| CAF-B | ACTA2, TAGLN, PDGFA | - Express cytoskeletal genes known for activated myofibroblast markers | |||
| NMFs | SFRP1/2, MFAP5, DPT, S100A4 | - Normal mucosa fibroblasts | |||
| Pancancer | myoCAF | ACTA2, MYH11 |
- Myofibroblastic CAF - Enriched tumorigenesis and myogenic regulons (TBX2, MEF2C each) |
48 | |
| inflaCAF | MMP11, CTHRC1, FAP, TGFB1 |
- Inflammatory CAF - Associated with dedifferentiation regulon |
|||
| adiCAF | CFD |
- Adipogenic CAF - Associated with EMT regulon |
|||
| EndMT-CAF | RGS5, ACTA2, PLVAP, VWF |
- Endothelial to Mesenchymal Transition CAF - Associated with angiogenesis and endothelial differentiation |
|||
| PN-CAF | MPZ, S100B, LGI4, PLP1 | - Peripheral nerve-like CAF | |||
| apCAF | ACTA2, HLA-DRA, CD74, HLA-DRB1 |
- Antigen-presenting CAF - Enriched in PDAC - Interaction with tumor-infiltrating T-cell clusters |
|||
| TAM | Breast cancer | Stromal Macrophage | CD11b + MRC1+ | - Harbor potent T-cell activation capacity | 140 |
| Hyperplastic Ductal Macrophage | CD11b- |
- Associated with an advanced tumor stage - Potent phagocytes - Not efficient for activating CD8 T cells - Locally accumulated through the active proliferation |
|||
| Malignant ductal TAM | CD11b + SPP1+ |
- Associated with poor prognosis - Regulate immunosuppressive functions of TAMs of monocytic origin |
|||
| Breast cancer | LAM1:FABP5 TAM | SPP1, FABP5 |
- Similar to lipid-associated macrophages (LAM) - High expression of TREM2 and lipid/fatty acid metabolic genes - Reduced proportion in HER2 + BC - Correlation with worse survival |
141 | |
| LAM2:APOE TAM | APOE |
- Similar to lipid-associated macrophages - High expression of TREM2 and lipid/fatty acid metabolic genes |
|||
| CXCL10 TAM | CXCL10, CXCL11 | - M1-like phenotype | |||
| EGR1, SIGLEC1 TAM | IL2RA, CD209 | - M2-like phenotype | |||
| Gastric cancer | HSP + TAM | HSPA6, HSPA1B, HSPB1 | - Increased HSP-associated genes | 142 | |
| THBS1 + TAM | THBS1 | - N/A | |||
| Chemokine-TAM | CCL3, CCL18, CCL20 | - Increased expression of chemokines | |||
| MMP-TAM | MMP9, MMP12 | - MMP genes | |||
| Complement-TAM | C1QA, C1QB, C1QC | - Complement family | |||
| Cell cycle-TAM | TOP2A, STMN1 | - Cell-cycle regulation genes | |||
| Colon cancer | C1QC + TAM | C1Q genes, TREM2, MERTK, CD80 |
- Derived from IL1B+ Tissue-resident macrophage (TRM) - Expression of MAF/MAFB and JUN/FOS - Increased inflammatory signatures, including complement activation, antigen processing, and presentation pathways |
59 | |
| SPP1 + TAM | SPP1, MARCO, VEGFA |
- Derived from NLRP3 + TRM - Expression of the level of HLA-DRs, CEBPB, and ZEB2 - Angiogenic signatures: enriched tumor angiogenesis, ECM receptor interaction, and tumor vasculature pathways |
|||
| Colorectal cancer | C1QC + MRC1- TAM | C1QC | - Closely related to CD14/CD16 monocytes in blood | 60 | |
| SPP1 + TAM | C1QC, MRC1, STAT1, PPARG |
- Tumor-specific filtration - Originated from THBS1 + TAM - Exhibit shorter progression-free survival |
|||
| THBS1 + TAM | THBS1 |
- Promote malignant migration of cancer - Capable of performing antigen processing and presentation and regulating intestinal immune network for IgA production - Can differentiate into SPP1+ macrophages |
|||
| VCAN + TAM | VCAN | - N/A | |||
| Colorectal cancer | Proinflammatory macrophage | IL1B, IL6, S100A8, S100A9 | - Upregulation of genes associated with cytokines | 143 | |
| SPP1+ macrophage A | SPP1, IL6 |
- Enriched in tumor core and border - Proinflammatory phenotype - Association with CMS type 4 |
|||
| SPP1+ macrophage B | SPP1, CD163, SEPP1, APOE, MAF |
- Enriched in tumor core and border - Anti-inflammatory phenotype |
|||
| Proliferating macrophage | MKI67, KIAA0101 | - Upregulation of genes associated with cell cycle | |||
| Hepatocellular carcinoma | TAM1 | FOLR2, CD163, C1QB, SEPP1, CD163high CD206high |
- Fetal-like TAM - FOLR2 expressing TAM - Exhibit immunosuppressive interactions - Higher expression of immunomodulatory chemokines - Enrichment with TIGIT+ cells |
144 | |
| TAM2 | SPP1, TREM2, FABP5, NUPR1, CD163low CD206low | - SPP1 + TAM | |||
| TAM3 | MT1G, MT2A, MT1X, CD163low CD206low | - MT1G-enriched TAM | |||
| Pancancer | C1QC + TAM | C1QC |
- Tumor enriched macrophage - Higher M2 signature and phagocytosis scores |
145 | |
| SPP1 + TAM | SPP1 |
- Tumor enriched macrophage - Higher M2 signature and angiogenesis signature |
|||
| ISG15 + TAM | ISG15 |
- Tumor-enriched macrophage - Upregulated IFN inducible genes - Higher expression of canonical M1 signature |
|||
| FN1 + TAM | FN1 |
- Tumor-enriched macrophage - Mainly present in kidney cancer - Proangiogenic TAM |
|||
| INHBA + TAM | INHBA | - Compensate SPP1 + TAM in stomach cancer with a proangiogenic signature | |||
| VCAN + TAM | VCAN | - Compensate SPP1 + TAM in BC with a proangiogenic signature | |||
| LYVE1 Macrophage | LYVE1 |
- Identified within multiple cancer types - Enriched in noncancer tissue - Tissue-resident interstitial macrophage |
|||
| NLRP3 Macrophage | NLRP3 |
- Enriched in noncancer tissue - Represent proinflammatory TRM (Tissue-resident macrophage) |
|||
| Pancancer | HES1 + TAM | C1QA, C1QB, C1QC, IGF1, CCL3, CCL4 | - Harbor an embryonic signature | 146 | |
| TREM2 + TAM | APOC1, APOE, SPP1, FABP5 |
- Accumulated only in tumor tissue - Involved in metabolic disorders - Potentially immunosuppressive role |
|||
| IL4I1 + TAM |
IL4I1 + PD-L1 + IDO1+ CD38, IDO1, CXCL9, CXCL10, CXCL11 |
- Antigen presentation, interaction with both Th2 and Th1 T cells, T-cell exhaustion, and tryptophan degradation - Suppress T cells and attract Tregs into the tumor by producing chemokine, expressing PD-L1 and PD-L2, and degrading IL4I1/AHR tryptophan - Exhibit immunosuppressive - Promote the entry of Treg into the tumor |
|||
| Proliferating TAM | TOP2A, MKI67, IDO1 | - Accumulated in all cancer types | |||
| TEC | PDAC | Endothelial 1 | IGFBP3, SPP1, CFH, IGLL5, TIMP1 |
- Higher expression of HIF1A - Enriched for ECM organization, regulation of vasculature development, regulation of angiogenesis, cell junction assembly and epithelial cell migration |
147 |
| Endothelial 2 | CLPS, PRSS1, CTRB1, CA4, CELA3A | - Represent normal pancreatic tissue | |||
| Lung cancer | Cluster1 | MT2A | - Normal EC | 53 | |
| Cluster 3 | IGFBP3 |
- Tumor EC - Enrichment of Myc target, nucleotide metabolism, OXPHOS-associated genes - Immune activation-associated genes are downregulated |
|||
| Cluster 4 | SPRY1 |
- Tumor EC - Enrichment of Myc target, nucleotide metabolism, OXPHOS-associated genes - Immune activation-associated genes are downregulated |
|||
| Cluster 5 | EDNRB | - Normal EC | |||
| Cluster 6 | PDPN, PROX1 | - Lymphatic EC | |||
| Gastric cancer | E0 | IGFBP5, STC1, IGFBP3 |
- Influence angiogenic sprouting - Upregulation of mTOR and IGF-1 signaling - Increase the invasion and migration of tumor cells |
148 | |
| E1 | FOXO1, FOXP1, JUN |
- Associated with the regulation of T-cell exhaustion signaling pathway - Suppress immune response |
|||
| E2 | N/A |
- Low activity - Normal endothelial cells |
|||
| E3 | NRP1, FGFR1 |
- VEGF receptor encoding genes are significantly upregulated - White adipose tissue browning pathway and STAT3 pathways are activated - Promote cancer cell invasion and angiogenesis |
|||
| Gastric cancer | Endo1 | COL4A1, COL4A2, PROS1 |
- Predominantly enriched in tumor - Downregulated MHC class II genes - Limited antigen presentation function - Strong activation of TNF, VEGF, PDGF, PGF, and Notch signaling - Involved in angiogenesis |
149 | |
| Endo2, Endo3, Endo4 | N/A | - N/A | |||
| Colorectal cancer | Tip-like EC | RGCC, RAMP3 |
- Overrepresentation of regulators of angiogenesis in tumor - Overrepresentation of antigen processing and presentation in normal |
143 | |
| Stalk-like EC | ACKR1, SELP | - Associated with apoptosis inhibition and proliferation | |||
| Proliferative EC | BIRC5, CKS1B | - Overexpression of BIRC5 and CKS1B | |||
| Lymphatic EC | LYVE, PROX1 | - Found both in normal and tumor | |||
| Hepatocellular carcinoma | PLPP3 + TEC | PLPP3 | - Enriched in tumor tissue | 144 | |
| PLVAP + TEC | PLVAP, HLA-DRA, |
- Enriched in tumor tissue - Facilitate the emergence of fetal-like macrophages - Mainly enriched in fetal and tumor tissues - Major subset expressing the receptor for VEGF |
|||
| IGFBP3 + TEC | IGFBP3 | - Enriched in tumor tissue | |||
| Clear cell renal cell carcinoma | AVR1 TEC | PLVAP, FLT1, KDR, FLT4, EDNRB, VWF, HSPG2 |
- Higher expression of VEGF receptor - Upregulation of genes involved in hemostasis, angiogenesis, and stimulation of endothelial growth and regeneration |
63 | |
| AVR2 TEC | ACKR1, SELP | - Evade angiogenesis inhibitors | |||
| Glioblastoma | Pe1 EC | KLF2, TIMP3, SLC2A1, SLCO1A2, TSC22D1, DEGS2, CAVIN2 |
- Quiescent endothelial cells derived from nonmalignant tissue - Associated with vascular integrity and BBB function |
150 | |
| Co1 EC | COL4A1, COL4A2, HSPG2, INSR, KDR |
- Derived from tumor core - Angiogenic phenotype - Associated with developmental and tumor angiogenesis, vascular basement membrane remodeling, cytoskeletal rearrangements, angiogenic sprouting, and endothelial tip cell formation |
|||
| Co2 EC | TMSB4X, RPLP2, RPL39, GAPDH, VIM, ACTB |
- Derived from tumor core - Intermediate phenotype - Associated with cytoskeletal and ribosomal protein expression |
|||
| Pe2 EC | CCL3, CCL4, CCL4L2, HLA-DRB1, HLA-DRA, HLA-DPA1, HLA-DPB1, HLA-DQB1 |
- Immune-activated phenotype derived from nonmalignant tissue - Expression of inflammatory cytokines and MHC II-mediated antigen presentation genes |
|||
| Co3 EC | NR4A3, IL1B, IL1R1, SELE, SELP, VACM1 |
- Derived from tumor core - Upregulation of immune-activated genes - Associated with inflammation and immune cell recruitment |
|||
| Pancancer | ESM1 tip EC | ESM1, NID2 |
- Only resided in malignant tissue - Upregulation of glycolysis and OXPHOS |
37 | |
| ACKR1high HEV and venous EC | ACKR1, SELP |
- ACKR1high endothelial venules and venous EC - Enriched in tumor |
|||
| CA4 capillary EC | CA4, CD36 | - Characterized by PLVAP and IGFBP7 | |||
| FBLN5 arterial EC | FBLN5, GJA5 | - Upregulated fatty acid biosynthesis | |||
| PROX1 lymphatic EC | PROX1, PDPN | - Increased fatty acid oxidation | |||
| TECs | PLVAP, IGFBP7 |
- Activation of HOXB pathways - Reduced carbonic acid metabolism |
AVR aortic valve replacement, BBB brain‒blood barrier, CMS consensus molecular subtype, EMT epithelial–mesenchymal transition, GC gastric cancer, HER2, BC Her2-positive breast cancer, LumA, BC luminal A breast cancer, NMF normal mucosa fibroblasts, NSCLC non-small cell lung cancer, OXPHOS oxidative phosphorylation, PDAC pancreatic ductal adenocarcinoma, TNBC triple-negative breast cancer, TRM tissue-resident macrophage. ACKR1 atypical chemokine receptor 1 (Duffy blood group), ACTA2 actin alpha 2, smooth muscle, ACTB actin beta, ADH1B alcohol dehydrogenase 1B (class I), beta polypeptide, AGTR1 angiotensin II receptor type 1, AHR aryl hydrocarbon receptor, APOC1 apolipoprotein C1, APOE apolipoprotein E, ASMA actin alpha 1, skeletal muscle, BIRC5 baculoviral IAP repeat containing 5, C1QC complement C1q C chain, CA4 carbonic anhydrase 4, CAV1 caveolin 1, CAVIN2 caveolae associated protein 2, CCL2 C-C motif chemokine ligand 2, CCL4L2 C-C motif chemokine ligand 4 like 2, CD4 CD4 molecule, CEBPB CCAAT enhancer binding protein beta, CELA3A chymotrypsin like elastase 3A, CFD complement Factor D, CFH complement Factor H, CKS1B CDC28 protein kinase regulatory subunit 1B, CLPS colipase, COL10A1 collagen type X alpha 1 chain, COL14A1 collagen type XIV alpha 1 chain, COL4A1 collagen type IV alpha 1 chain, COL4A2 collagen type IV alpha 2 chain, CST1 cystatin SN, CTHRC1 collagen triple helix repeat containing 1, CTRB1 chymotrypsinogen B1, CXCL1 C-X-C motif chemokine ligand 1, Dcn Decorin, DEGS2 delta 4-desaturase, sphingolipid 2, DPT dermatopontin, EDNRB endothelin receptor type B, EGR1 early growth response 1, ESM1 endothelial cell specific molecule 1, FABP5 fatty acid binding protein 5, FAP fibroblast activation protein alpha, FBLN5 fibulin 5, FGFR1 fibroblast growth factor receptor 1, FIGF vascular endothelial growth factor D, FLT1 Fms related receptor tyrosine kinase 1, FN1 fibronectin 1, FOLR2 folate receptor beta, FOS Fos proto-oncogene, AP-1 transcription factor subunit, FOXO1 forkhead Box O1, FOXP1 Forkhead Box the P1, FSP1 S100 calcium binding protein A4, GAPDH glyceraldehyde-3-phosphate dehydrogenase, GGH gamma-glutamyl hydrolase, GJA5 gap junction protein alpha 5, GJB2 gap junction protein beta 2, GPX3 glutathione peroxidase 3, H2AB1 H2A. B variant histone 1, HAS1 hyaluronan synthase 1, HER2 Erb-b2 receptor tyrosine kinase 2, HES1 Hes family bHLH transcription Factor 1, HIF1A hypoxia inducible factor 1 subunit alpha, HLA-DPB1 major histocompatibility complex, class II, DP beta 1, HLA-DQB1 major histocompatibility complex, class II, DQ beta 1, HLA-DR human leukocyte antigen - DR isotype, HLA-DRA major histocompatibility complex, class II, DR alpha, HLA-DRB1 major histocompatibility complex, class II, DR beta 1, HOPX HOP homeobox, HOXB homeobox B, HOXB2 Homeobox B2, HSPG2 Heparan sulfate proteoglycan 2, ID1 Inhibitor of DNA binding 1, IDO1 indoleamine 2,3-dioxygenase 1, IFIT3 interferon induced protein with tetratricopeptide repeats 3, IGF1 insulin like growth factor 1, IGFBP3 insulin like growth factor binding protein 3, IGLL5 immunoglobulin lambda like polypeptide 5, IL1B interleukin 1 beta, IL1R1 interleukin 1 receptor type 1, IL2RA interleukin 2 receptor subunit alpha, IL4I1 interleukin 4 induced 1, INHBA inhibin subunit beta A, INSR insulin receptor, IRF7 interferon regulatory factor 7, ISG15 ISG15 ubiquitin like modifier, JUN Jun proto-oncogene, AP-1 transcription factor subunit, KDR kinase insert domain receptor, KIAA0101 PCNA clamp associated factor, KLF2 KLF transcription factor 2, LGI4 leucine rich repeat LGI family member 4, LIF LIF interleukin 6 family cytokine, LMNA lamin A/C, LRRC15 leucine rich repeat containing 15, Ly6C lymphocyte antigen 6 family member C 1, LYVE1 lymphatic vessel endothelial hyaluronan receptor 1, MAF MAF bZIP transcription factor, MAFB MAF bZIP transcription factor B, MARCO macrophage receptor with collagenous structure, MEF2C myocyte enhancer Factor 2C, MERTK MER proto-oncogene, tyrosine kinase, Mfap5 microfibril associated protein 5, Mki67 antigen identified by monoclonal antibody Ki 67, MMP1 matrix metallopeptidase 1, MPZ myelin protein zero, MRC1 mannose receptor C-type 1, MT1B metallothionein 1B, MT1X metallothionein 1X, MT2A metallothionein 2A, MYH11 myosin heavy chain 11, MYL9 myosin light chain 9, NID2 nidogen 2, NLRP3 NLR family pyrin domain containing 3, NR4A3 nuclear receptor subfamily 4 group A member 3, NRP1 neuropilin 1, Nuf2 NUF2, NDC80 kinetochore complex component, NUPR1 Nuclear protein 1, transcriptional regulator, PDGF platelet-derived growth factor, Pdgfra platelet derived growth factor receptor alpha, PDGFRB platelet derived growth factor receptor beta, PD-L1 CD274 molecule, PDPN podoplanin, PGF placental growth factor, PLA2G2A phospholipase A2 group IIA, PLP1 proteolipid protein 1, PLPP3 phospholipid phosphatase 3, PLVAP plasmalemma vesicle-associated protein, POSTN periostin, PPARG peroxisome proliferator activated receptor gamma, PROS1 protein S, PROX1 prospero homeobox 1, PRSS1 serine protease 1, RAMP3 receptor activity modifying protein 3, RGCC regulator of cell cycle, RGMA repulsive guidance molecule BMP coreceptor a, Rgs5 regulator of G-protein signaling 5, RPLP2 Ribosomal protein lateral stalk subunit the P2, S100A8 S100 calcium binding protein A8, S100B S100 calcium binding protein B, Saa3 serum amyloid A 3, SCARA5 scavenger receptor class A member 5, Scrg1 scrapie responsive gene 1, SELE selectin E, SELP selectin P, SEMA3C semaphorin 3C, SEPP1 selenoprotein P, SFRP4 secreted frizzled related protein 4, SIGLEC1 sialic acid binding Ig like lectin 1, SLC2A1 solute carrier family 2 member 1, SLCO1A2 solute carrier organic anion transporter family member 1A2, Slpi secretory leukocyte peptidase inhibitor, Sox9 SRY (sex determining region Y)-Box 9, SPP1 CXXC finger protein 1, SPRY1 Sprouty RTK signaling antagonist 1, STAT1 signal transducer and activator of transcription 1, STC1 stanniocalcin 1, TAGLN transgelin, TBX2 T-box transcription factor 2, TFF3 Trefoil factor 3, TGFB1 transforming growth factor beta 1, THBS1 thrombospondin 1, TIGIT T-cell immunoreceptor with Ig and ITIM domains, TIMP1 TIMP metallopeptidase inhibitor 1, TMSB4X thymosin beta 4 X-linked, TNF tumor necrosis factor, TOP2A DNA topoisomerase II alpha, TPM1 tropomyosin 1, TREM2 triggering receptor expressed on myeloid cells 2, TSC22D1 TSC22 domain family member 1, VACM1 Cullin 5, VCAN versican, VEGFA vascular endothelial growth Factor A, VIM Vimentin, VWF Von Willebrand factor, ZEB2 Zinc finger E-box binding homeobox 2.