Table 3

Cytokines that human neutrophils can potentially express and/or produce

Cytokine familySpecific cytokines
C-X-C chemokines CXCL1, CXCL2, CXCL3, CXCL4*, CXCL5, CXCL6, CXCL8, CXCL9, CXCL10, CXCL11, CXCL12*, CXCL13* 
C-C chemokines CCL2, CCL3, CCL4, CCL17, CCL18, CCL19, CCL20, CCL22 
Proinflammatory cytokines IL-1α, IL-1β, IL-6, IL-7, IL-9(?), IL-16(?), IL-17A, IL-17B, IL-18, MIF 
Anti-inflammatory cytokines IL-1ra, IL-4(?), TGFβ1, TGFβ2 
Immunoregulatory cytokines IFNα(?), IFNβ*, IFNγ(?), IL-12, IL-21, IL-23, IL-27, IL-18BP 
TNF superfamily members TNFα, TNFSF6/FasL, TNFSF8/CD30L, TNFSF10/TRAIL, TNFSF14/LIGHT*, TNFSF3/Lymphotoxin-β*, TNFSF13/APRIL, TNFSF13B/BAFF/BLyS, TNFSF5/CD40L, TNFSF11/RANKL 
Colony-stimulating factors G-CSF, M-CSF(?), GM-CSF(?), IL-3(?), SCF* 
Angiogenic and fibrogenic factors VEGFs, BV8 (prokineticin 2), HB-EGF, FGF-2, TGFα, HGF, angiopoietin-1 
Other cytokines NGF*, BDNF*, NT4*, PBEF (visfatin/NAMPT), amphiregulin, midkine, oncostatin M, activin A, endothelin 
Cytokine familySpecific cytokines
C-X-C chemokines CXCL1, CXCL2, CXCL3, CXCL4*, CXCL5, CXCL6, CXCL8, CXCL9, CXCL10, CXCL11, CXCL12*, CXCL13* 
C-C chemokines CCL2, CCL3, CCL4, CCL17, CCL18, CCL19, CCL20, CCL22 
Proinflammatory cytokines IL-1α, IL-1β, IL-6, IL-7, IL-9(?), IL-16(?), IL-17A, IL-17B, IL-18, MIF 
Anti-inflammatory cytokines IL-1ra, IL-4(?), TGFβ1, TGFβ2 
Immunoregulatory cytokines IFNα(?), IFNβ*, IFNγ(?), IL-12, IL-21, IL-23, IL-27, IL-18BP 
TNF superfamily members TNFα, TNFSF6/FasL, TNFSF8/CD30L, TNFSF10/TRAIL, TNFSF14/LIGHT*, TNFSF3/Lymphotoxin-β*, TNFSF13/APRIL, TNFSF13B/BAFF/BLyS, TNFSF5/CD40L, TNFSF11/RANKL 
Colony-stimulating factors G-CSF, M-CSF(?), GM-CSF(?), IL-3(?), SCF* 
Angiogenic and fibrogenic factors VEGFs, BV8 (prokineticin 2), HB-EGF, FGF-2, TGFα, HGF, angiopoietin-1 
Other cytokines NGF*, BDNF*, NT4*, PBEF (visfatin/NAMPT), amphiregulin, midkine, oncostatin M, activin A, endothelin 

The expression and/or production of most of these cytokines (updated from Mantovani et al) has been validated in human neutrophils by gene expression techniques, immunohistochemistry, enzyme-linked immunosorbent assay, or biological assays. *, studies performed at the mRNA level only; (?), controversial data; APRIL, proliferation-inducing ligand; BAFF/BLyS, B-cell activating factor/B lymphocyte stimulator; BDNF, brain-derived neurotrophic factor; CCL, CC chemokine ligand; CXCL, CXC chemokine ligand; FasL, Fas ligand; FGF, fibroblast growth factor; HB-EGF, heparin binding-like epidermal growth factor; HGF, hepatocyte growth factor; IFN, interferon; IL-1ra, IL-1 receptor antagonist; IL18BP, IL-18 binding protein; LIGHT, homologous to lymphotoxins; M-CSF, macrophage-colony stimulating factor; MIF, macrophage inhibitory factor; NAMPT, nicotinamide phosphoribosyl transferase; NGF, nerve growth factor; NT4, neurotrophin-4; PBEF, pre-B-cell colony-enhancing factor; RANKL, receptor activator of nuclear factor k-B ligand; SCF, stem cell factor; TGF, transforming growth factor; TNFSF, TNF super family; TRAIL, TNF-related apoptosis-inducing ligand; VEGF, vascular endothelial growth factor.

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