Reynisdottir, G. et al. Structural lung changes and local anti-citrulline immunity are early features of anti citrullinated-proteins antibodies positive rheumatoid arthritis. Arthritis Rheum.http://dx.doi.org/10.1002/art.38201.
Quirke, A. M. et al. Heightened immune response to autocitrullinated Porphyromonas gingivalis peptidylarginine deiminase: a potential mechanism for breaching immunologic tolerance in rheumatoid arthritis. Ann. Rheum. Dis.73, 263–269 (2014). ArticleCASPubMed Google Scholar
Maresz, K. J. et al. Porphyromonas gingivalis facilitates the development and progression of destructive arthritis through its unique bacterial peptidylarginine deiminase (PAD). PLoS Pathog.9, e1003627 (2013). ArticleCASPubMedPubMed Central Google Scholar
Sabeh, F., Fox, D. & Weiss, S. J. Membrane-type I matrix metalloproteinase-dependent regulation of rheumatoid arthritis synoviocyte function. J. Immunol.184, 6396–6406 (2010). ArticleCASPubMed Google Scholar
Miller, M. C. et al. Membrane type 1 matrix metalloproteinase is a crucial promoter of synovial invasion in human rheumatoid arthritis. Arthritis Rheum.60, 686–697 (2009). ArticleCASPubMedPubMed Central Google Scholar
Tolboom, T. C. et al. Invasive properties of fibroblast-like synoviocytes: correlation with growth characteristics and expression of MMP-1, MMP-3, and MMP-10. Ann. Rheum. Dis.61, 975–980 (2002). ArticleCASPubMedPubMed Central Google Scholar
Murphy, G. & Nagase, H. Reappraising metalloproteinases in rheumatoid arthritis and osteoarthritis: destruction or repair? Nat. Clin. Pract. Rheumatol.4, 128–135 (2008). ArticleCASPubMed Google Scholar
Smolen, J. E. et al. L-selectin signaling of neutrophil adhesion and degranulation involves p38 mitogen-activated protein kinase. J. Biol. Chem.275, 15876–15884 (2000). ArticleCASPubMed Google Scholar
Yang, L. et al. ICAM-1 regulates neutrophil adhesion and transcellular migration of TNF-α-activated vascular endothelium under flow. Blood106, 584–592 (2005). ArticleCASPubMedPubMed Central Google Scholar
Woodfin, A., Voisin, M. B. & Nourshargh, S. Recent developments and complexities in neutrophil transmigration. Curr. Opin. Hematol.17, 9–17 (2010). ArticlePubMedPubMed Central Google Scholar
Cross, A., Barnes, T., Bucknall, R. C., Edwards, S. W. & Moots, R. J. Neutrophil apoptosis in rheumatoid arthritis is regulated by local oxygen tensions within joints. J. Leukoc. Biol.80, 521–528 (2006). ArticleCASPubMed Google Scholar
Walmsley, S. R. et al. Hypoxia-induced neutrophil survival is mediated by HIF-1α-dependent NF-κB activity. J. Exp. Med.201, 105–115 (2005). ArticleCASPubMedPubMed Central Google Scholar
Wright, H. L., Thomas, H. B., Moots, R. J. & Edwards, S. W. RNA-seq reveals activation of both common and cytokine-specific pathways following neutrophil priming. PLoS ONE8, e58598 (2013). ArticleCASPubMedPubMed Central Google Scholar
Cassatella, M. A. Neutrophil-derived proteins: selling cytokines by the pound. Adv. Immunol.73, 369–509 (1999). ArticleCASPubMed Google Scholar
Mantovani, A., Cassatella, M. A., Costantini, C. & Jaillon, S. Neutrophils in the activation and regulation of innate and adaptive immunity. Nat. Rev. Immunol.11, 519–531 (2011). ArticleCASPubMed Google Scholar
Fujishima, S. et al. Regulation of neutrophil interleukin 8 gene expression and protein secretion by LPS, TNF-α, and IL-1β. J. Cell Physiol.154, 478–485 (1993). ArticleCASPubMed Google Scholar
Nunes, P., Demaurex, N. & Dinauer, M. C. Regulation of the NADPH oxidase and associated ion fluxes during phagocytosis. Traffic14, 1118–1131 (2013). CASPubMed Google Scholar
Fossati, G., Bucknall, R. C. & Edwards, S. W. Insoluble and soluble immune complexes activate neutrophils by distinct activation mechanisms: changes in functional responses induced by priming with cytokines. Ann. Rheum. Dis.61, 13–19 (2002). ArticleCASPubMedPubMed Central Google Scholar
Fossati, G., Bucknall, R. C. & Edwards, S. W. Fcγ receptors in autoimmune diseases. Eur. J. Clin. Invest.31, 821–831 (2001). ArticleCASPubMed Google Scholar
Robinson, J., Watson, F., Bucknall, R. C. & Edwards, S. W. Activation of neutrophil reactive-oxidant production by synovial fluid from patients with inflammatory joint disease. Soluble and insoluble immunoglobulin aggregates activate different pathways in primed and unprimed cells. Biochem. J.286 (Pt 2), 345–351 (1992). ArticleCASPubMedPubMed Central Google Scholar
Derouet, M., Thomas, L., Cross, A., Moots, R. J. & Edwards, S. W. Granulocyte macrophage colony-stimulating factor signaling and proteasome inhibition delay neutrophil apoptosis by increasing the stability of Mcl-1. J. Biol. Chem.279, 26915–26921 (2004). ArticleCASPubMed Google Scholar
Derouet, M. et al. Sodium salicylate promotes neutrophil apoptosis by stimulating caspase-dependent turnover of Mcl-1. J. Immunol.176, 957–965 (2006). ArticleCASPubMed Google Scholar
Thomas, L. W., Lam, C. & Edwards, S. W. Mcl-1; the molecular regulation of protein function. FEBS Lett.584, 2981–2989 (2010). ArticleCASPubMed Google Scholar
Savill, J. S. et al. Macrophage phagocytosis of aging neutrophils in inflammation. Programmed cell death in the neutrophil leads to its recognition by macrophages. J. Clin. Invest.83, 865–875 (1989). ArticleCASPubMedPubMed Central Google Scholar
Eggleton, P., Wang, L., Penhallow, J., Crawford, N. & Brown, K. A. Differences in oxidative response of subpopulations of neutrophils from healthy subjects and patients with rheumatoid arthritis. Ann. Rheum. Dis.54, 916–923 (1995). ArticleCASPubMedPubMed Central Google Scholar
Wright, H. L., Chikura, B., Bucknall, R. C., Moots, R. J. & Edwards, S. W. Changes in expression of membrane TNF, NF-κB activation and neutrophil apoptosis during active and resolved inflammation. Ann. Rheum. Dis.70, 537–543 (2011). ArticleCASPubMed Google Scholar
Matsumoto, T. et al. The membrane proteinase 3 expression on neutrophils was downregulated after treatment with infliximab in patients with rheumatoid arthritis. Clin. Appl. Thromb. Hemost.14, 186–192 (2008). ArticleCASPubMed Google Scholar
Wittkowski, H. et al. Effects of intra-articular corticosteroids and anti-TNF therapy on neutrophil activation in rheumatoid arthritis. Ann. Rheum. Dis.66, 1020–1025 (2007). ArticleCASPubMedPubMed Central Google Scholar
Chakravarti, A., Raquil, M. A., Tessier, P. & Poubelle, P. E. Surface RANKL of Toll-like receptor 4-stimulated human neutrophils activates osteoclastic bone resorption. Blood114, 1633–1644 (2009). ArticleCASPubMed Google Scholar
Assi, L. K. et al. Tumor necrosis factor α activates release of B lymphocyte stimulator by neutrophils infiltrating the rheumatoid joint. Arthritis Rheum.56, 1776–1786 (2007). ArticleCASPubMedPubMed Central Google Scholar
Cross, A., Bucknall, R. C., Cassatella, M. A., Edwards, S. W. & Moots, R. J. Synovial fluid neutrophils transcribe and express class II major histocompatibility complex molecules in rheumatoid arthritis. Arthritis Rheum.48, 2796–2806 (2003). ArticleCASPubMed Google Scholar
Lally, F. et al. A novel mechanism of neutrophil recruitment in a coculture model of the rheumatoid synovium. Arthritis Rheum.52, 3460–3469 (2005). ArticleCASPubMedPubMed Central Google Scholar
Parsonage, G. et al. Prolonged, granulocyte-macrophage colony-stimulating factor-dependent, neutrophil survival following rheumatoid synovial fibroblast activation by IL-17 and TNFα. Arthritis Res. Ther.10, R47–R59 (2008). ArticleCASPubMedPubMed Central Google Scholar
Raza, K. et al. Synovial fluid leukocyte apoptosis is inhibited in patients with very early rheumatoid arthritis. Arthritis Res. Ther.8, R120–R127 (2006). ArticleCASPubMedPubMed Central Google Scholar
Weinmann, P. et al. Delayed neutrophil apoptosis in very early rheumatoid arthritis patients is abrogated by methotrexate therapy. Clin. Exp. Rheumatol.25, 885–887 (2007). CASPubMed Google Scholar
Rollet-Labelle, E. et al. Cross-linking of IgGs bound on circulating neutrophils leads to an activation of endothelial cells: possible role of rheumatoid factor in rheumatoid arthritis-associated vascular dysfunction. J. Inflamm. (Lond.)10, 27 (2013). ArticleCAS Google Scholar
Nurcombe, H. L., Bucknall, R. C. & Edwards, S. W. Neutrophils isolated from the synovial fluid of patients with rheumatoid arthritis: priming and activation in vivo. Ann. Rheum. Dis.50, 147–153 (1991). ArticleCASPubMedPubMed Central Google Scholar
Watson, F., Robinson, J. J., Phelan, M., Bucknall, R. C. & Edwards, S. W. Receptor expression in synovial fluid neutrophils from patients with rheumatoid arthritis. Ann. Rheum. Dis.52, 354–359 (1993). ArticleCASPubMedPubMed Central Google Scholar
Quayle, J. A., Watson, F., Bucknall, R. C. & Edwards, S. W. Neutrophils from the synovial fluid of patients with rheumatoid arthritis express the high affinity immunoglobulin G receptor, FcγRI (CD64): role of immune complexes and cytokines in induction of receptor expression. Immunology91, 266–273 (1997). ArticleCASPubMedPubMed Central Google Scholar
Robinson, J. J., Watson, F., Bucknall, R. C. & Edwards, S. W. Role of Fcγ receptors in the activation of neutrophils by soluble and insoluble immunoglobulin aggregates isolated from the synovial fluid of patients with rheumatoid arthritis. Ann. Rheum. Dis.53, 515–520 (1994). ArticleCASPubMedPubMed Central Google Scholar
Hallett, M. B. & Lloyds, D. Neutrophil priming: the cellular signals that say 'amber' but not 'green'. Immunol. Today16, 264–268 (1995). ArticleCASPubMed Google Scholar
Fossati, G., Moots, R. J., Bucknall, R. C. & Edwards, S. W. Differential role of neutrophil Fcγ receptor IIIB (CD16) in phagocytosis, bacterial killing, and responses to immune complexes. Arthritis Rheum.46, 1351–1361 (2002). ArticleCASPubMed Google Scholar
Lefrancais, E. et al. IL-33 is processed into mature bioactive forms by neutrophil elastase and cathepsin G. Proc. Natl Acad. Sci. USA109, 1673–1678 (2012). ArticleCASPubMedPubMed Central Google Scholar
Van den Steen, P. E. et al. Cleavage of denatured natural collagen type II by neutrophil gelatinase B reveals enzyme specificity, post-translational modifications in the substrate, and the formation of remnant epitopes in rheumatoid arthritis. FASEB J.16, 379–389 (2002). ArticleCASPubMed Google Scholar
Katano, M. et al. Implication of granulocyte-macrophage colony-stimulating factor induced neutrophil gelatinase-associated lipocalin in pathogenesis of rheumatoid arthritis revealed by proteome analysis. Arthritis Res. Ther.11, R3 (2009). ArticleCASPubMedPubMed Central Google Scholar
Elsaid, K. A., Jay, G. D. & Chichester, C. O. Detection of collagen type II and proteoglycans in the synovial fluids of patients diagnosed with non-infectious knee joint synovitis indicates early damage to the articular cartilage matrix. Osteoarthritis Cartilage11, 673–680 (2003). ArticleCASPubMed Google Scholar
Baici, A., Salgam, P., Cohen, G., Fehr, K. & Boni, A. Action of collagenase and elastase from human polymorphonuclear leukocytes on human articular cartilage. Rheumatol. Int.2, 11–16 (1982). ArticleCASPubMed Google Scholar
Oseas, R., Yang, H. H., Baehner, R. L. & Boxer, L. A. Lactoferrin: a promoter of polymorphonuclear leukocyte adhesiveness. Blood57, 939–945 (1981). CASPubMed Google Scholar
Sopata, I. et al. Neutrophil gelatinase levels in plasma and synovial fluid of patients with rheumatic diseases. Rheumatol. Int.15, 9–14 (1995). ArticleCASPubMed Google Scholar
Pham, C. T. Neutrophil serine proteases: specific regulators of inflammation. Nat. Rev. Immunol.6, 541–550 (2006). ArticleCASPubMed Google Scholar
Wang, C. H. et al. Expression of CD147 (EMMPRIN) on neutrophils in rheumatoid arthritis enhances chemotaxis, matrix metalloproteinase production and invasiveness of synoviocytes. J. Cell. Mol. Med.15, 850–860 (2011). ArticleCASPubMed Google Scholar
Brinkmann, V. et al. Neutrophil extracellular traps kill bacteria. Science303, 1532–1535 (2004). ArticleCASPubMed Google Scholar
Khandpur, R. et al. NETs are a source of citrullinated autoantigens and stimulate inflammatory responses in rheumatoid arthritis. Sci. Transl. Med.5, 178ra40 (2013). ArticleCASPubMedPubMed Central Google Scholar
Baillet, A. et al. Synovial fluid proteomic fingerprint: S100A8, S100A9 and S100A12 proteins discriminate rheumatoid arthritis from other inflammatory joint diseases. Rheumatology49, 671–682 (2010). ArticleCASPubMed Google Scholar
Simard, J. C., Girard, D. & Tessier, P. A. Induction of neutrophil degranulation by S100A9 via a MAPK-dependent mechanism. J. Leukoc. Biol.87, 905–914 (2010). ArticleCASPubMed Google Scholar
Laurindo, I. M., Mello, S. B. & Cossermelli, W. Influence of low doses of methotrexate on superoxide anion production by polymorphonuclear leukocytes from patients with rheumatoid arthritis. J. Rheumatol.22, 633–638 (1995). CASPubMed Google Scholar
Kraan, M. C. et al. Inhibition of neutrophil migration soon after initiation of treatment with leflunomide or methotrexate in patients with rheumatoid arthritis: findings in a prospective, randomized, double-blind clinical trial in fifteen patients. Arthritis Rheum.43, 1488–1495 (2000). ArticleCASPubMed Google Scholar
Sperling, R. I. et al. Acute and chronic suppression of leukotriene B4 synthesis ex vivo in neutrophils from patients with rheumatoid arthritis beginning treatment with methotrexate. Arthritis Rheum.35, 376–384 (1992). ArticleCASPubMed Google Scholar
Elferink, J. G., Daha, M. R. & de Koster, B. M. A cyclic GMP- and G-kinase-dependent effect of azathioprine on migration by human neutrophils. Cell. Mol. Life Sci.53, 593–599 (1997). ArticleCASPubMed Google Scholar
Akahoshi, T. et al. Rapid induction of neutrophil apoptosis by sulfasalazine: implications of reactive oxygen species in the apoptotic process. J. Leukoc. Biol.62, 817–826 (1997). ArticleCASPubMed Google Scholar
Wandall, J. H. Effects of sulphasalazine and its metabolites on neutrophil chemotaxis, superoxide production, degranulation and translocation of cytochrome _b_-245. Aliment. Pharmacol. Ther.5, 609–619 (1991). ArticleCASPubMed Google Scholar
Cross, A., Moots, R. J. & Edwards, S. W. The dual effects of TNFα on neutrophil apoptosis are mediated via differential effects on expression of Mcl-1 and Bfl-1. Blood111, 878–884 (2008). ArticleCASPubMed Google Scholar
Ginis, I. & Tauber, A. I. Activation mechanisms of adherent human neutrophils. Blood76, 1233–1239 (1990). CASPubMed Google Scholar
Wipke, B. T. & Allen, P. M. Essential role of neutrophils in the initiation and progression of a murine model of rheumatoid arthritis. J. Immunol.167, 1601–1608 (2001). ArticleCASPubMed Google Scholar
Tanaka, D., Kagari, T., Doi, H. & Shimozato, T. Essential role of neutrophils in anti-type II collagen antibody and lipopolysaccharide-induced arthritis. Immunology119, 195–202 (2006). ArticleCASPubMedPubMed Central Google Scholar
Coelho, F. M. et al. The chemokine receptors CXCR1/CXCR2 modulate antigen-induced arthritis by regulating adhesion of neutrophils to the synovial microvasculature. Arthritis Rheum.58, 2329–2337 (2008). ArticlePubMed Google Scholar
Grespan, R. et al. CXCR2-specific chemokines mediate leukotriene B4-dependent recruitment of neutrophils to inflamed joints in mice with antigen-induced arthritis. Arthritis Rheum.58, 2030–2040 (2008). ArticleCASPubMed Google Scholar
Barsante, M. M. et al. Blockade of the chemokine receptor CXCR2 ameliorates adjuvant-induced arthritis in rats. Br. J. Pharmacol.153, 992–1002 (2008). ArticleCASPubMed Google Scholar
Min., S. H. et al. Pharmacological targeting reveals distinct roles for CXCR2/CXCR1 and CCR2 in a mouse model of arthritis. Biochem. Biophys. Res. Commun.391, 1080–1086 (2009). ArticleCASPubMed Google Scholar
Cunha, T. M. et al. Treatment with DF 2162, a non-competitive allosteric inhibitor of CXCR1/2, diminishes neutrophil influx and inflammatory hypernociception in mice. Br. J. Pharmacol.154, 460–470 (2008). ArticleCASPubMedPubMed Central Google Scholar
Grant, E. P. et al. Essential role for the C5a receptor in regulating the effector phase of synovial infiltration and joint destruction in experimental arthritis. J. Exp. Med.196, 1461–1471 (2002). ArticleCASPubMedPubMed Central Google Scholar
Tsuboi, N. et al. Regulation of human neutrophil Fcγ receptor IIa by C5a receptor promotes inflammatory arthritis in mice. Arthritis Rheum.63, 467–478 (2011). ArticleCASPubMedPubMed Central Google Scholar
Sadik, C. D., Kim, N. D., Iwakura, Y. & Luster, A. D. Neutrophils orchestrate their own recruitment in murine arthritis through C5aR and FcγR signaling. Proc. Natl Acad. Sci. USA109, E3177–E3185 (2012). ArticleCASPubMedPubMed Central Google Scholar
Kim, N. D., Chou, R. C., Seung, E., Tager, A. M. & Luster, A. D. A unique requirement for the leukotriene B4 receptor BLT1 for neutrophil recruitment in inflammatory arthritis. J. Exp. Med.203, 829–835 (2006). ArticleCASPubMedPubMed Central Google Scholar
Grevers, L. C. et al. Different amplifying mechanisms of interleukin-17 and interferon-γ in Fcγ receptor-mediated cartilage destruction in murine immune complex-mediated arthritis. Arthritis Rheum.60, 396–407 (2009). ArticleCASPubMed Google Scholar
Sarraj, B., Ludanyi, K., Glant, T. T., Finnegan, A. & Mikecz, K. Expression of CD44 and L-selectin in the innate immune system is required for severe joint inflammation in the proteoglycan-induced murine model of rheumatoid arthritis. J. Immunol.177, 1932–1940 (2006). ArticleCASPubMed Google Scholar
Eyles, J. L. et al. A key role for G-CSF-induced neutrophil production and trafficking during inflammatory arthritis. Blood112, 5193–5201 (2008). ArticleCASPubMed Google Scholar
Williams, A. S. et al. Interferon-γ protects against the development of structural damage in experimental arthritis by regulating polymorphonuclear neutrophil influx into diseased joints. Arthritis Rheum.56, 2244–2254 (2007). ArticleCASPubMed Google Scholar
Kelchtermans, H. et al. Effector mechanisms of interleukin-17 in collagen-induced arthritis in the absence of interferon-γ and counteraction by interferon-γ. Arthritis Res. Ther.11, R122 (2009). ArticleCASPubMedPubMed Central Google Scholar
Davey, M. S. et al. Failure to detect production of IL-10 by activated human neutrophils. Nat. Immunol.12, 1017–1020 (2011). ArticleCASPubMed Google Scholar
Mestas, J. & Hughes, C. C. Of mice and not men: differences between mouse and human immunology. J. Immunol.172, 2731–2738 (2004). ArticleCASPubMed Google Scholar
Branzk, N. & Papayannopoulos, V. Molecular mechanisms regulating NETosis in infection and disease. Semin. Immunopathol.35, 513–530 (2013). ArticleCASPubMedPubMed Central Google Scholar
Cooper, P. R., Palmer, L. J. & Chapple, I. L. Neutrophil extracellular traps as a new paradigm in innate immunity: friend or foe? Periodontol. 200063, 165–197 (2013). ArticlePubMed Google Scholar
Yousefi, S., Mihalache, C., Kozlowski, E., Schmid, I. & Simon, H. U. Viable neutrophils release mitochondrial DNA to form neutrophil extracellular traps. Cell Death Differ.16, 1438–1444 (2009). ArticleCASPubMed Google Scholar
Wang, Y. et al. Histone hypercitrullination mediates chromatin decondensation and neutrophil extracellular trap formation. J. Cell Biol.184, 205–213 (2009). ArticleCASPubMedPubMed Central Google Scholar
Knight, J. S., Carmona-Rivera, C. & Kaplan, M. J. Proteins derived from neutrophil extracellular traps may serve as self-antigens and mediate organ damage in autoimmune diseases. Front. Immunol.3, 380 (2012). ArticlePubMedPubMed Central Google Scholar
Garcia-Romo, G. S. et al. Netting neutrophils are major inducers of type I IFN production in pediatric systemic lupus erythematosus. Sci. Transl. Med.3, 73ra20 (2011). ArticlePubMedPubMed Central Google Scholar
Lande, R. et al. Neutrophils activate plasmacytoid dendritic cells by releasing self–DNA–peptide complexes in systemic lupus erythematosus. Sci. Transl. Med.3, 73ra19 (2011). ArticlePubMedPubMed Central Google Scholar
Villanueva, E. et al. Netting neutrophils induce endothelial damage, infiltrate tissues, and expose immunostimulatory molecules in systemic lupus erythematosus. J. Immunol.187, 538–552 (2011). ArticleCASPubMed Google Scholar
Midgley, A., McLaren, Z., Moots, R. J., Edwards, S. W. & Beresford, M. W. The role of neutrophil apoptosis in juvenile-onset systemic lupus erythematosus. Arthritis Rheum.60, 2390–2401 (2009). ArticleCASPubMed Google Scholar
Hacbarth, E. & Kajdacsy-Balla, A. Low density neutrophils in patients with systemic lupus erythematosus, rheumatoid arthritis, and acute rheumatic fever. Arthritis Rheum.29, 1334–1342 (1986). ArticleCASPubMed Google Scholar
Denny, M. F. et al. A distinct subset of proinflammatory neutrophils isolated from patients with systemic lupus erythematosus induces vascular damage and synthesizes type I IFNs. J. Immunol.184, 3284–3297 (2010). ArticleCASPubMed Google Scholar
Theilgaard-Monch, K. et al. The transcriptional program of terminal granulocytic differentiation. Blood105, 1785–1796 (2005). ArticleCASPubMed Google Scholar
Hakkim, A. et al. Impairment of neutrophil extracellular trap degradation is associated with lupus nephritis. Proc. Natl Acad. Sci. USA107, 9813–9818 (2010). ArticleCASPubMedPubMed Central Google Scholar
Romero, V. et al. Immune-mediated pore-forming pathways induce cellular hypercitrullination and generate citrullinated autoantigens in rheumatoid arthritis. Sci. Transl. Med.5, 209ra150 (2013). ArticleCASPubMedPubMed Central Google Scholar
Scally, S. W. et al. A molecular basis for the association of the HLA-DRB1 locus, citrullination, and rheumatoid arthritis. J. Exp. Med.210, 2569–2582 (2013). ArticleCASPubMedPubMed Central Google Scholar
Pratesi, F. et al. Antibodies from patients with rheumatoid arthritis target citrullinated histone 4 contained in neutrophils extracellular traps. Ann. Rheum. Dis.http://dx.doi.org/10.1136/annrheumdis-2012-202765.