From Monocytes to M1/M2 Macrophages: Phenotypical vs. Functional Differentiation

Matzinger, 2007, Friendly and dangerous signals: is the tissue in control?, Nat Immunol, 8, 11, 10.1038/ni0107-11

Medzhitov, 2008, Origin and physiological roles of inflammation, Nature, 454, 428, 10.1038/nature07201

Nathan, 2002, Points of control in inflammation, Nature, 420, 846, 10.1038/nature01320

van Furth, 1968, The origin and kinetics of mononuclear phagocytes, J Exp Med, 128, 415, 10.1084/jem.128.3.415

Taylor, 2003, Monocytes heterogeneity and innate immunity, Immunity, 19, 2, 10.1016/S1074-7613(03)00178-X

Lin, 2008, Discovery of a cytokine and its receptor by functional screening of the extracellular proteome, Science, 320, 807, 10.1126/science.1154370

Hume, 2012, Therapeutic applications of macrophage colony-stimulating factor-1 (CSF-1) and antagonists of CSF-1 receptor (CSF-1R) signaling, Blood, 119, 1810, 10.1182/blood-2011-09-379214

Geissmann, 2010, Unravelling mononuclear phagocyte heterogeneity, Nat Rev Immunol, 10, 453, 10.1038/nri2784

Jenkins, 2014, Homeostasis in the mononuclear phagocyte system, Trends Immunol, 35, 358, 10.1016/j.it.2014.06.006

Geissmann, 2010, Development of monocytes, macrophages, and dendritic cells, Science, 327, 656, 10.1126/science.1178331

Boltjes, 2014, Human dendritic cell functional specialization in steady-state and inflammation, Front Immunol, 5, 131, 10.3389/fimmu.2014.00131

Schulz, 2012, A lineage of myeloid cells independent of Myb and hematopoietic stem cells, Science, 336, 86, 10.1126/science.1219179

Sieweke, 2013, Beyond stem cells: self-renewal of differentiated macrophages, Science, 342, 1242974, 10.1126/science.1242974

Mills, 2014, M1 and M2 macrophages: the chicken and the egg of immunity, J Innate Immun, 6, 10.1159/000364945

Akashi, 2000, A clonogenic common myeloid progenitor that gives rise to all myeloid lineages, Nature, 404, 193, 10.1038/35004599

Fogg, 2006, A clonogenic bone marrow progenitor specific for macrophages and dendritic cells, Science, 311, 83, 10.1126/science.1117729

Hettinger, 2013, Origin of monocytes and macrophages in a committed progenitor, Nat Immunol, 14, 821, 10.1038/ni.2638

Liu, 2009, In vivo analysis of dendritic cell development and homeostasis, Science, 324, 392, 10.1126/science.1170540

Ajami, 2007, Local self-renewal can sustain CNS microglia maintenance and function throughout adult life, Nat Neurosci, 10, 1538, 10.1038/nn2014

Hamilton, 2008, Colony-stimulating factor in inflammation and autoimmunity, Nat Rev Immunol, 8, 533, 10.1038/nri2356

Bartocci, 1987, Macrophages specifically regulate the concentration of their own growth factor in the circulation, Proc Natl Acad Sci U S A, 84, 6179, 10.1073/pnas.84.17.6179

Tushinski, 1982, Survival of mononuclear phagocytes depends on a lineage-specific growth factor that the differentiated cells selectively destroy, Cell, 28, 71, 10.1016/0092-8674(82)90376-2

Ma, 2012, Structural basis for the dual recognition of helical cytokines IL-34 and CSF-1 by CSF-1R, Structure, 20, 676, 10.1016/j.str.2012.02.010

Wang, 2012, IL-34 is a tissue-restricted ligand of CSF1R required for the development of Langerhans cells and microglia, Nat Immunol, 13, 753, 10.1038/ni.2360

Burgess, 1980, The nature and action of granulocyte-macrophage colony stimulating factors, Blood, 56, 947, 10.1182/blood.V56.6.947.947

Gasson, 1991, Molecular physiology of granulocyte-macrophage colony stimulating factor, Blood, 77, 1131, 10.1182/blood.V77.6.1131.1131

Serbina, 2008, Monocyte-mediated defense against microbial pathogens, Annu Rev Immunol, 26, 421, 10.1146/annurev.immunol.26.021607.090326

Evans, 2009, In vivo activated monocytes from the site of inflammation in humans specifically promote Th17 responses, Proc Natl Acad Sci U S A, 106, 6232, 10.1073/pnas.0808144106

Avraham-Davidi, 2013, On-site education of VEGF-recruited monocytes improves their performance as angiogenic and arteriogenic accessory cells, J Exp Med, 210, 2611, 10.1084/jem.20120690

Passlick, 1989, Identification and characterization of a novel monocyte subpopulation in human peripheral blood, Blood, 74, 2527, 10.1182/blood.V74.7.2527.2527

Ziegler-Heitbrock, 2010, Nomenclature of monocytes and dendritic cells in blood, Blood, 116, e74, 10.1182/blood-2010-02-258558

Mobley, 2007, Genetic evidence of a functional monocyte dichotomy, Inflammation, 30, 189, 10.1007/s10753-007-9036-0

Zhao, 2009, Identification of novel functional differences in monocyte subsets using proteomic and transcriptomic methods, J Proteome Res, 8, 4028, 10.1021/pr900364p

Ancuta, 2009, Transcriptional profiling reveals developmental relationship and distinct biological functions of CD16+ and CD16- monocyte subsets, BMC Genomics, 10, 403, 10.1186/1471-2164-10-403

Cros, 2010, Human CD14dim monocytes patrol and sense nucleic acids and viruses via TLR7 and TLR8 receptors, Immunity, 33, 375, 10.1016/j.immuni.2010.08.012

Wong, 2011, Gene expression profiling reveals the defining features of the classical, intermediate, and nonclassical human monocyte subsets, Blood, 118, e16, 10.1182/blood-2010-12-326355

Zawada, 2011, SuperSAGE evidence for CD14++CD16+ monocytes as a third monocyte subset, Blood, 118, e50, 10.1182/blood-2011-01-326827

Frankenberger, 2012, Transcript profiling of CD16-positive monocytes reveals a unique molecular fingerprint, Eur J Immunol, 42, 957, 10.1002/eji.201141907

Barnes, 2005, Experimental comparison and cross-validation of the Affymetrix and Illumina gene expression analysis platforms, Nucleic Acids Res, 33, 5914, 10.1093/nar/gki890

Ziegler-Heitbrock, 2013, Toward a refined definition of monocyte subsets, Front Immunol, 4, 23, 10.3389/fimmu.2013.00023

Wong, 2012, The three human monocyte subsets: implications for health and disease, Immunol Res, 53, 41, 10.1007/s12026-012-8297-3

Ingersoll, 2010, Comparison of gene expression profiles between human and mouse monocyte subsets, Blood, 115, e10, 10.1182/blood-2009-07-235028

Geissmann, 2003, Blood monocytes consist of two principal subsets with distinct migratory properties, Immunity, 19, 71, 10.1016/S1074-7613(03)00174-2

Tsou, 2007, Critical roles for CCR2 and MPC-3 in monocyte mobilization from bone marrow and recruitment to inflammatory sites, J Clin Invest, 117, 902, 10.1172/JCI29919

Auffray, 2007, Monitoring of blood vessels and tissues by a population of monocytes with patrolling behavior, Science, 317, 666, 10.1126/science.1142883

Ziegler-Heitbrock, 2014, Reprint of: monocyte subsets in man and other species, Cell Immunol, 10.1016/j.cellimm.2014.06.008

Sunderkötter, 2004, Subpopulations of mouse blood monocytes differ in maturation stage and inflammatory response, J Immunol, 172, 4410, 10.4049/jimmunol.172.7.4410

Yona, 2013, Fate mapping reveals origins and dynamics of monocytes and tissue macrophages under homeostasis, Immunity, 38, 79, 10.1016/j.immuni.2012.12.001

Varol, 2007, Monocytes give rise to mucosal, but not splenic, conventional dendritic cells, J Exp Med, 204, 171, 10.1084/jem.20061011

MacDonald, 2010, An antibody against the colony-stimulating factor 1 receptor depletes the resident subset of monocytes and tissue- and tumor-associated macrophages but does not inhibit inflammation, Blood, 116, 3955, 10.1182/blood-2010-02-266296

Carlin, 2013, Nr4a1-dependent Ly6C(low) monocytes monitor endothelial cells and orchestrate their disposal, Cell, 153, 362, 10.1016/j.cell.2013.03.010

Hanna, 2011, The transcription factor NR4A1 (Nur77) controls bone marrow differentiation and the survival of Ly6C- monocytes, Nat Immunol, 12, 778, 10.1038/ni.2063

Varga, 2013, Tissue LyC6- macrophages are generated in the absence of circulating LyC6- monocytes and Nur77 in a model of muscle regeneration, J Immunol, 191, 5695, 10.4049/jimmunol.1301445

Shi, 2011, Monocyte recruitment during infection and inflammation, Nat Rev Immunol, 11, 762, 10.1038/nri3070

Swirski, 2009, Identification of splenic reservoir monocytes and their deployment to inflammatory sites, Science, 325, 612, 10.1126/science.1175202

Nguyen, 2013, Circadian gene Bmal1 regulates diurnal oscillations of Ly6C(hi) inflammatory monocytes, Science, 341, 1483, 10.1126/science.1240636

Bain, 2013, Resident and pro-inflammatory macrophages in the colon represent alternative context-dependent fates of the same Ly6Chi monocyte precursors, Mucosal Immunol, 6, 498, 10.1038/mi.2012.89

Rivollier, 2012, Inflammation switches the differentiation program of Ly6Chi monocytes from anti-inflammatory macrophages to inflammatory dendritic cells in the colon, J Exp Med, 209, 139, 10.1084/jem.20101387

Jaensson, 2008, Small intestinal CD103+ dendritic cells display unique functional properties that are conserved between mice and humans, J Exp Med, 205, 2139, 10.1084/jem.20080414

Tamoutounour, 2013, Origins and functional specialization of macrophages and of conventional and monocyte-derived dendritic cells in mouse skin, Immunity, 39, 925, 10.1016/j.immuni.2013.10.004

Abtin, 2014, Perivascular macrophages mediate neutrophil recruitment during bacterial skin infection, Nat Immunol, 15, 45, 10.1038/ni.2769

Epelman, 2014, Embryonic and adult-derived resident cardiac macrophages are maintained through distinct mechanisms at steady state and during inflammation, Immunity, 40, 91, 10.1016/j.immuni.2013.11.019

Guilliams, 2013, Alveolar macrophages develop from fetal monocytes that differentiate into long-lived cells in the first week of life via GM-CSF, J Exp Med, 210, 1977, 10.1084/jem.20131199

Jakubzick, 2013, Minimal differentiation of classical monocytes as they survey steady-state tissues and transport antigen to lymph nodes, Immunity, 39, 599, 10.1016/j.immuni.2013.08.007

Capulli, 2014, Osteoblast and osteocyte: games without frontiers, Arch Biochem Biophys, 561C, 3, 10.1016/j.abb.2014.05.003

Charles, 2014, Osteoclasts: more than “bone eaters”, Trends Mol Med, 20, 449, 10.1016/j.molmed.2014.06.001

Kikuta, 2013, Osteoclast migration, differentiation and function: novel therapeutic target for rheumatic diseases, Rheumatology (Oxford), 52, 226, 10.1093/rheumatology/kes259

Boyle, 2003, Osteoclast differentiation and activation, Nature, 423, 337, 10.1038/nature01658

Dai, 2002, Targeted disruption of the mouse colony-stimulating factor 1 receptor gene results in osteopetrosis, mononuclear phagocyte deficiency, increased primitive progenitor cell frequencies, and reproductive defects, Blood, 99, 111, 10.1182/blood.V99.1.111

Massey, 1999, Human osteoclasts derive from CD14-positive monocytes, Br J Haematol, 106, 167, 10.1046/j.1365-2141.1999.01491.x

Chen, 2011, The critical role of IL-34 in osteoclastogenesis, PLoS One, 6, e18689, 10.1371/journal.pone.0018689

Matsuzaki, 1998, Osteoclast differentiation factor (ODF) induces osteoclast-like cell formation in human peripheral blood mononuclear cell cultures, Biochem Biophys Res Commun, 246, 199, 10.1006/bbrc.1998.8586

Hayashi, 1998, Commitment and differentiation of stem cells to the osteoclast lineage, Biochem Cell Biol, 76, 911, 10.1139/o98-099

Gautier, 2012, Gene-expression profiles and transcriptional regulatory pathways that underlie the identity and diversity of mouse tissue macrophages, Nat Immunol, 13, 1118, 10.1038/ni.2419

van Furth, 1970, Origin and kinetics of monocytes and macrophages, Semin Hematol, 7, 125

van Furth, 1972, The mononuclear phagocyte system: a new classification of macrophages, monocytes, and their precursor cells, Bull World Health Organ, 46, 845

Volkman, 1970, The origin and fate of the monocyte, Ser Haematol, 3, 69

Volkman, 1968, The function of the monocytes, Bibl Haematol, 29, 86

Epelman, 2014, Origin and function of tissue macrophages, Immunity, 41, 21, 10.1016/j.immuni.2014.06.013

Orkin, 2008, Hematopoiesis: an evolving paradigm for stem cell biology, Cell, 132, 631, 10.1016/j.cell.2008.01.025

Cumano, 2007, Ontogeny of the hematopoietic system, Annu Rev Immunol, 25, 745, 10.1146/annurev.immunol.25.022106.141538

Tavian, 2005, Embryonic development of the human hematopoietic system, Int J Dev Biol, 49, 243, 10.1387/ijdb.041957mt

Enzan, 1986, Electron microscopic studies of macrophages in early human yolk sacs, Acta Pathol Jpn, 36, 49

Migliaccio, 1986, Human embryonic hemopoiesis. Kinetics of progenitors and precursors underlying the yolk sac – liver transition, J Clin Invest, 78, 51, 10.1172/JCI112572

Takahashi, 1989, Differentiation, maturation, and proliferation of macrophages in the mouse yolk sac: a light-microscopic, enzyme-cytochemical, immunohistochemical, and ultrastructural study, J Leukoc Biol, 45, 87, 10.1002/jlb.45.2.87

Hashimoto, 2013, Tissue-resident macrophages self-maintain locally throughout adult life with minimal contribution from circulating monocytes, Immunity, 38, 792, 10.1016/j.immuni.2013.04.004

Ginhoux, 2010, Fate mapping analysis reveals that adult microglia derive from primitive macrophages, Science, 330, 841, 10.1126/science.1194637

Kierdorf, 2013, Microglia emerge from erythromyeloid precursors via Pu.1- and Irf8-dependent pathways, Nat Neurosci, 16, 273, 10.1038/nn.3318

Hoeffel, 2012, Adult Langerhans cells derive predominantly from embryonic fetal liver monocytes with a minor contribution of yolk sac-derived macrophages, J Exp Med, 209, 1167, 10.1084/jem.20120340

Ginhoux, 2014, Monocytes and macrophages: development pathways and tissue homeostasis, Nat Rev Immunol, 14, 392, 10.1038/nri3671

Wynn, 2013, Macrophages biology in development, homeostasis and disease, Nature, 496, 445, 10.1038/nature12034

Frankenberger, 2013, A defect of CD16-positive monocytes can occur without disease, Immunobiology, 218, 169, 10.1016/j.imbio.2012.02.013

Hambleton, 2011, IRF8 mutations and human dendritic-cell immunodeficiency, N Engl J Med, 365, 127, 10.1056/NEJMoa1100066

Bigley, 2011, The human syndrome of dendritic cell, monocyte, B and NK lymphoid deficiency, J Exp Med, 208, 227, 10.1084/jem.20101459

Randolph, 2013, Macrophages in Marseille, Immunity, 38, 619, 10.1016/j.immuni.2013.04.002

Aziz, 2009, MafB/c-Maf deficiency enables self-renewal of differentiated functional macrophages, Science, 326, 867, 10.1126/science.1176056

Czernielewski, 1987, Further evidence for the self-reproducing capacity of Langerhans cells in human skin, J Invest Dermatol, 88, 17, 10.1111/1523-1747.ep12464659

Kanitakis, 2004, Turnover of epidermal Langerhans’ cells, N Engl J Med, 351, 2661, 10.1056/NEJM200412163512523

Merad, 2002, Langerhans cells renew in the skin throughout life under steady-state conditions, Nat Immunol, 3, 1135, 10.1038/ni852

Davies, 2011, A quantifiable proliferative burst of tissue macrophages restores homeostatic macrophage populations after acute inflammation, Eur J Immunol, 41, 2155, 10.1002/eji.201141817

Coggle, 1984, The proliferation kinetics of pulmonary alveolar macrophages, J Leukoc Biol, 35, 317, 10.1002/jlb.35.3.317

Jenkins, 2011, Local macrophages proliferation, rather than recruitment from the blood, is a signature of TH2 inflammation, Science, 332, 1284, 10.1126/science.1204351

Jenkins, 2013, IL-4 directly signals tissue-resident macrophages to proliferate beyond homeostatic levels controlled by CSF-1, J Exp Med, 210, 2477, 10.1084/jem.20121999

Hibbs, 2007, Mice lacking three myeloid colony-stimulating factors (G-CSF, GM-CSF, and M-CSF) still produce macrophages and granulocytes and mount an inflammatory response in a sterile model of peritonitis, J Immunol, 178, 6435, 10.4049/jimmunol.178.10.6435

Chitu, 2006, Colony-stimulating factor-1 in immunity and inflammation, Curr Opin Immunol, 18, 39, 10.1016/j.coi.2005.11.006

Davies, 2013, Distinct bone marrow-derived and tissue-resident macrophage lineages proliferate at key stages during inflammation, Nat Commun, 4, 1886, 10.1038/ncomms2877

Hume, 2008, Differentiation and heterogeneity in the mononuclear phagocyte system, Mucosal Immunol, 1, 432, 10.1038/mi.2008.36

Tagliani, 2011, Coordinate regulation of tissue macrophage and dendritic cell population dynamics by CSF-1, J Exp Med, 208, 1901, 10.1084/jem.20110866

Ryan, 2001, Rescue of the colony-stimulating factor 1 (CSF-1)-nullizygous mouse (Csf1(op)/Csf1(op)) phenotype with a CSF-1 transgene and identification of sites of local CSF-1 synthesis, Blood, 98, 74, 10.1182/blood.V98.1.74

Shibata, 2001, GM-CSF regulates alveolar macrophage differentiation and innate immunity in the lung through PU.1, Immunity, 15, 557, 10.1016/S1074-7613(01)00218-7

Koenen, 1996, Effect of intraperitoneal administration of granulocyte/macrophage-colony-stimulating factor in rats on omental milky-spot composition and tumoricidal activity in vivo and in vitro, Cancer Immunol Immunother, 42, 310, 10.1007/s002620050288

Lieschke, 1994, Mice lacking granulocyte colony-stimulating factor have chronic neutropenia, granulocyte and macrophage progenitor cell deficiency, and impaired neutrophil mobilization, Blood, 84, 1737, 10.1182/blood.V84.6.1737.1737

Stanley, 1994, Granulocyte/macrophage colony-stimulating factor-deficient mice show no major perturbation of hematopoiesis but develop a characteristic pulmonary pathology, Proc Natl Acad Sci U S A, 91, 5592, 10.1073/pnas.91.12.5592

Tadokoro, 2001, Bone marrow-derived macrophages grown in GM-CSF or M-CSF differ in their ability to produce IL-12 and to induce IFN-gamma production after stimulation with Trypanosoma cruzi antigens, Immunol Lett, 77, 31, 10.1016/S0165-2478(01)00197-3

El-Behi, 2011, The encephalitogenicity of T(H)17 cells is dependent on IL-1- and IL-23-induced production of the cytokine GM-CSF, Nat Immunol, 12, 568, 10.1038/ni.2031

Fleetwood, 2009, GM-CSF- and M-CSF-dependent macrophage phenotypes display differential dependence on type I interferon signaling, J Leukoc Biol, 86, 411, 10.1189/jlb.1108702

Fleetwood, 2007, Granulocyte-macrophage colony-stimulating factor (CSF) and macrophage CSF-dependent macrophage phenotypes display differences in cytokine profiles and transcription factor activities: implications for CSF blockade in inflammation, J Immunol, 178, 5245, 10.4049/jimmunol.178.8.5245

Verreck, 2004, Human IL-23-producing type 1 macrophages promote but IL-10-producing type 2 macrophages subvert immunity to (myco)bacteria, Proc Natl Acad Sci U S A, 101, 4560, 10.1073/pnas.0400983101

Martinez, 2006, Transcriptional profiling of the human monocyte-to-macrophage differentiation and polarization: new molecules and patterns of gene expression, J Immunol, 177, 7303, 10.4049/jimmunol.177.10.7303

Ghigo, 2013, Multicolor fate mapping of Langerhans cell homeostasis, J Exp Med, 210, 1657, 10.1084/jem.20130403

Paolicelli, 2011, Synaptic pruning by microglia is necessary for normal brain development, Science, 333, 1456, 10.1126/science.1202529

London, 2013, Microglia and monocyte-derived macrophages: functionally distinct populations that act in concert in CNS plasticity and repair, Front Cell Neurosci, 7, 34, 10.3389/fncel.2013.00034

Bogie, 2014, Macrophages subsets and microglia in multiple sclerosis, Acta Neurophatol, 128, 191, 10.1007/s00401-014-1310-2

Pollard, 2009, Trophic macrophages in development and disease, Nat Rev Immunol, 9, 259, 10.1038/nri2528

Blin-Wakkach, 2014, Roles of osteoclasts in the control of medullary hematopoietic niches, Arch Biochem Biophys, 561C, 29, 10.1016/j.abb.2014.06.032

Kleyer, 2014, Arthritis and bone loss: a hen and egg story, Curr Opin Rheumatol, 26, 80, 10.1097/BOR.0000000000000007

Randolph, 2014, Mechanisms that regulate macrophage burden in atherosclerosis, Circ Res, 114, 1757, 10.116/CIRCRESAHA.114.301174

Klein, 2007, Kupffer cell heterogeneity: functional properties of bone marrow derived and sessile hepatic macrophages, Blood, 110, 4077, 10.1182/blood-2007-02-073841

Ganz, 2012, Macrophages and systemic iron homeostasis, J Innate Immun, 4, 446, 10.1159/000336423

Sánchez-Valle, 2012, Role of oxidative stress and molecular changes in liver fibrosis: a review, Curr Med Chem, 19, 4850, 10.2174/092986712803341520

Davies, 2013, Tissue-resident macrophages, Nat Immunol, 14, 986, 10.1038/ni.2705

Maus, 2002, Role of resident alveolar macrophages in leukocyte traffic into the alveolar air space of intact mice, Am J Physiol Lung Cell Mol Physiol, 282, L1245, 10.1152/ajplung.00453.2001

Carey, 2010, The molecular basis of pulmonary alveolar proteinosis, Clin Immunol, 135, 223, 10.1016/j.clin.2010.02.017

Nguyen, 2011, Alternatively activated macrophages produce catecholamines to sustain adaptive thermogenesis, Nature, 480, 104, 10.1038/nature10653

Zigmond, 2013, Intestinal macrophages: well educated exceptions from the rule, Trends Immunol, 34, 162, 10.1016/j.it.2013.02.001

Wallace, 2014, Immunopathology of inflammatory bowel disease, World J Gastroenterol, 20, 6, 10.3748/wjg.v20.i1.6

Chorro, 2010, Development and homeostasis of “resident” myeloid cells: the case of the Langerhans cell, Trends Immunol, 31, 438, 10.1016/j.it.2010.09.003

Novak, 2013, Phenotypic transitions of macrophages orchestrate tissue repair, Am J Pathol, 183, 1352, 10.1016/j.ajpath.2013.06.034

den Haan, 2012, Innate immune functions of macrophages subpopulations in the spleen, J Innate Immun, 4, 437, 10.1159/000335216

Kohyama, 2009, Role of Spi-C in the development of red pulp macrophages and splenic iron homeostasis, Nature, 457, 318, 10.1038/nature07472

Labonte, 2014, The role of macrophages polarization in infectious and inflammatory diseases, Mol Cells, 37, 275, 10.14348/molcells.2014.2374

Mantovani, 2013, Macrophage plasticity and polarixation in tissue repair and remodelling, J Pathol, 229, 176, 10.1002/path.4133

Hume, 2012, The complexity of constitutive and inducible gene expression in mononuclear phagocytes, J Leukoc Biol, 92, 433, 10.1189/jlb.0312166

Lech, 2012, Tissue use resident dendritic cells and macrophages to maintain homeostasis and to regain homeostasis upon tissue injury: the immunoregulatory role of changing tissue environments, Mediators Inflamm, 2012, 951390, 10.1155/2012/951390

Mills, 2012, M1 and M2 macrophages: oracles of health and disease, Crit Rev Immunol, 32, 463, 10.1615/CritRevImmunol.v32.i6.10

Barth, 1995, Review of the macrophage disappearance reaction, J Leukoc Biol, 57, 361, 10.1002/jlb.57.3.361

Struyf, 1998, Synergistic induction of MCP-1 and -2 by IL-1β and interferons in fibroblasts and epithelial cells, J Leukoc Biol, 36, 364, 10.1002/jlb.63.3.364

Shi, 2011, Bone marrow mesenchymal stem and progenitor cells induce monocyte emigration in response to circulating toll-like receptor ligands, Immunity, 34, 590, 10.1016/j.immuni.2011.02.016

Serbina, 2006, Monocyte emigration from bone marrow during bacterial infection requires signals mediated by chemokine receptor CCR2, Nat Immunol, 7, 311, 10.1038/ni1309

Nahrendorf, 2007, The healing myocardium sequentially mobilizes two monocyte subsets with divergent and complementary functions, J Exp Med, 204, 3037, 10.1084/jem.20070885

Willenborg, 2012, CCR2 recruits an inflammatory macrophage subpopulation critical for angiogenesis in tissue repair, Blood, 120, 613, 10.1182/blood-2012-01-403386

Ishida, 2008, Chemokine receptor CX3CR1 mediates skin wound healing by promoting macrophage and fibroblast accumulation and function, J Immunol, 180, 569, 10.4049/jimmunol.180.1.569

White, 2012, Fractalkine: a survivor’s guide: chemokines as antiapoptotic mediators, Arterioscler Thromb Vasc Biol, 32, 589, 10.1161/ATVBAHA.111.237412

Wang, 2009, CCR2 and CXCR4 regulate peripheral blood monocyte pharmacodynamics and link to efficacy in experimental autoimmune encephalomyelitis, J Inflamm (Lond), 6, 32, 10.1186/1476-9255-6-32

Debien, 2013, S1PR5 is pivotal for the homeostasis of patrolling monocytes, Eur J Immunol, 43, 1667, 10.1002/eji.201343312

Arnold, 2007, Inflammatory monocytes recruited after skeletal muscle injury switch into antiinflammatory macrophages to support myogenesis, J Exp Med, 204, 1057, 10.1084/jem.20070075

Crane, 2014, The monocyte to macrophage transition in the murine sterile wound, PLoS One, 9, e86660, 10.1371/journal.pone.0086660

Gautier, 2009, Regulation of the migration and survival of monocyte subsets by chemokine receptors and its relevance to atherosclerosis, Arterioscler Thromb Vasc Biol, 29, 1412, 10.1161/ATVBAHA.108.180505

Robbins, 2013, Local proliferation dominates lesional macrophage accumulation in atherosclerosis, Nat Med, 19, 1166, 10.1038/nm.3258

Chorro, 2009, Langerhans cell (LC) proliferation mediates neonatal development, homeostasis, and inflammation-associated expansion of the epidermal LC network, J Exp Med, 206, 3089, 10.1084/jem.20091586

Ajami, 2011, Infiltrating monocytes trigger EAE progression, but do not contribute to the resident microglia pool, Nat Neurosci, 14, 1142, 10.1038/nn.2887

Tymoszuk, 2014, In situ proliferation contributes to accumulation of tumor-associated macrophages in spontaneous mammary tumors, Eur J Immunol, 44, 2247, 10.1002/eji.201344304

Amano, 2014, Local proliferation of macrophages contributes to obesity-associated adipose tissue inflammation, Cell Metab, 19, 162, 10.1016/j.cmet.2013.11.017

Mills, 2001, Macrophage arginine metabolism to ornithine/urea or nitric oxide/citrulline: a life or death issue, Crit Rev Immunol, 21, 399, 10.1615/CritRevImmunol.v21.i5.10

Zigmond, 2012, Ly6Chi monocytes in the inflamed colon give rise to proinflammatory effector cells and migratory antigen-presenting cells, Immunity, 37, 1076, 10.1016/j.immuni.2012.08.026

Mills, 2000, M-1/M-2 macrophages and the Th1/Th2 paradigm, J Immunol, 164, 6166, 10.4049/jimmunol.164.12.6166

Pesce, 2009, Arginase-1-expressing macrophages suppress Th2 cytokine-driven inflammation and fibrosis, PLoS Pathog, 5, e1000371, 10.1371/journal.ppat.1000371

MacMicking, 1997, Nitric oxide and macrophage function, Annu Rev Immunol, 15, 323, 10.1146/annurev.immunol.15.1.323

Van den Bossche, 2012, Pivotal advance: arginase-1-independent polyamine production stimulates the expression of IL-4-induced alternatively activated macrophage markers while inhibiting LPS-induced expression of inflammatory genes, J Leukoc Biol, 91, 685, 10.1189/jlb.0911453

Murray, 2011, Protective and pathogenic functions of macrophage subsets, Nat Rev Immunol, 11, 723, 10.1038/nri3073

Mantovani, 2004, The chemokine system in diverse forms of macrophage activation and polarization, Trends Immunol, 25, 677, 10.1016/j.it.2004.09.015

Biswas, 2012, Orchestration of metabolism by macrophages, Cell Metab, 15, 432, 10.1016/j.cmet.2011.11.013

Rodríguez-Prados, 2010, Substrate fate in activated macrophages: a comparison between innate, classic, and alternative activation, J Immunol, 185, 605, 10.4049/jimmunol.0901698

Janeway, 2002, Innate immune recognition, Annu Rev Immunol, 20, 197, 10.1146/annurev.immunol.20.083001.084359

Kawai, 2011, Toll-like receptors and their crosstalk with other innate receptors in infection and immunity, Immunity, 34, 637, 10.1016/j.immuni.2011.05.006

Unanue, 1984, Antigen-presenting function of the macrophage, Annu Rev Immunol, 2, 395, 10.1146/annurev.iy.02.040184.002143

Gordon, 2005, Monocyte and macrophage heterogeneity, Nat Rev Immunol, 5, 953, 10.1038/nri1733

Gordon, 2003, Alternative activation of macrophages, Nat Rev Immunol, 3, 23, 10.1038/nri978

Martinez, 2008, Macrophage activation and polarization, Front Biosci, 13, 453, 10.2741/2692

Sica, 2012, Macrophage plasticity and polarization: in vivo veritas, J Clin Invest, 122, 787, 10.1172/JCI59643

Mosser, 2008, Exploring the full spectrum of macrophage activation, Nat Rev Immunol, 8, 958, 10.1038/nri2448

Mantovani, 2002, Macrophage polarization: tumor-associated macrophages as a paradigm for polarized M2 mononuclear phagocytes, Trends Immunol, 23, 549, 10.1016/S1471-4906(02)02302-5

Fleming, 2011, Regulatory macrophages: setting the threshold for therapy, Eur J Immunol, 41, 2498, 10.1002/eji.201141717

Manjili, 2014, Evolution of our understanding of myeloid regulatory cells: from MDSCs to Mregs, Front Immunol, 5, 303, 10.3389/fimmu.2014.00303

Martinez, 2014, The M1 and M2 paradigm of macrophage activation: time for reassessment, F1000Prime Rep, 6, 13, 10.12703/P6-13

Okabe, 2014, Tissue-specific signals control reversible program of localization and functional polarization of macrophages, Cell, 157, 832, 10.1016/j.cell.2014.04.016

Pettersen, 2011, Tumor-associated macrophages in the cutaneous SCC microenvironment are heterogeneously activated, J Invest Dermatol, 131, 1322, 10.103/jid.2011.9

Vogel, 2013, Macrophages in inflammatory multiple sclerosis lesions have an intermediate activation status, J Neuroinflammation, 10, 35, 10.1186/1742-2094-10-35

Lawrence, 2011, Transcriptional regulation of macrophage polarization: enabling diversity with identity, Nat Rev Immunol, 11, 750, 10.1038/nri3088

Lin, 2009, Bone marrow Ly6Chigh monocytes are selectively recruited to injured kidney and differentiate into functionally distinct populations, J Immunol, 183, 6733, 10.4049/jimmunol.0901473

Italiani, 2014, Transcriptomic profiling of the development of the inflammatory response in human monocytes in vitro, PLoS One, 9, e87680, 10.1371/journal.pone.0087680

Mylonas, 2009, Alternatively activated macrophages elicited by helminth infection can be reprogrammed to enable microbial killing, J Immunol, 182, 3084, 10.4049/jimmunol.0803463

Stout, 2005, Macrophages sequentially change their functional phenotypes in response to changes in microenvironmental influences, J Immunol, 175, 342, 10.4049/jimmunol.175.1.342

Egawa, 2013, Inflammatory monocytes recruited to allergic skin acquire an anti-inflammatory M2 phenotype via basophil-derived interleukin-4, Immunity, 38, 570, 10.1016/j.immuni.2012.11.014

Gundra, 2014, Alternatively activated macrophages derived from monocytes and tissue macrophages are phenotypically and functionally distinct, Blood, 123, e110, 10.1182/blood-2013-08-520619

Albina, 1989, Regulation of macrophages functions by l-arginine, J Exp Med, 169, 1021, 10.1084/jem.169.3.1021

Morris, 2009, Recent advances in arginine metabolism: roles and regulation of the arginases, Br J Pharmacol, 157, 922, 10.1111/ji.1476-5381.2009.00278

Vodovoz, 1993, Mechanisms of suppression of macrophages nitric oxide release by transforming growth factor beta, J Exp Med, 178, 605, 10.1084/jem.178.2.605

Biswas, 2009, Endotoxin tolerance: new mechanisms, molecules and clinical significance, Trends Immunol, 30, 475, 10.1016/j.it.2009.07.009

Bastard, 2006, Recent advances in the relationship between obesity, inflammation, and insulin resistance, Eur Cytokine Netw, 17, 4

Martín-Fuentes, 2007, Individual variation of scavenger receptor expression in human macrophages with oxidized low-density lipoprotein is associated with a differential inflammatory response, J Immunol, 179, 3242, 10.4049/jimmunol.179.5.3242

Swann, 2008, Demonstration of inflammation-induced cancer and cancer immunoediting during primary tumorigenesis, Proc Natl Acad Sci U S A, 105, 652, 10.1073/pnas.0708594105

Biswas, 2010, Macrophage plasticity and interaction with lymphocyte subsets: cancer as a paradigm, Nat Immunol, 11, 889, 10.1038/ni.1937

Stout, 2004, Functional plasticity of macrophages: reversible adaptation to changing microenvironments, J Leukoc Biol, 76, 509, 10.1189/jlb.0504272

Biswas, 2012, Macrophage polarization and plasticity in health and disease, Immunol Res, 53, 11, 10.1007/s12026-012-8291-9

Medzhitov, 2009, Transcriptional control of the inflammatory response, Nat Rev Immunol, 9, 692, 10.1038/nri2634

Smale, 2010, Selective transcription in response to an inflammatory stimulus, Cell, 140, 833, 10.1016/j.cell.2010.01.037

Ley, 2014, The second touch hypothesis: T cell activation, homing and polarization, F1000Res, 3, 37, 10.12688/f1000research.3_37.v02

Kawakami, 2005, Live imaging of effector cell trafficking and autoantigen recognition within the unfolding autoimmune encephalomyelitis lesion, J Exp Med, 201, 1805, 10.1084/jem.20050011

Wakim, 2008, Dendritic cell-induced memory T cell activation in nonlymphoid tissues, Science, 319, 198, 10.1126/science.1151869

Muller, 2012, CD4+ T Cells rely on a cytokine gradient to control intracellular pathogens beyond sites of antigen presentation, Immunity, 37, 147, 10.1016/j.immuni.2012.05.015

Choi, 2009, Identification of antigen-presenting dendritic cells in mouse aorta and cardiac valves, J Exp Med, 206, 497, 10.1084/jem.20082129

Choi, 2011, Flt3 signaling-dependent dendritic cells protect against atherosclerosis, Immunity, 35, 819, 10.1016/j.immuni.2011.09.014

Koltsova, 2012, Dynamic T cell-APC interactions sustain chronic inflammation in atherosclerosis, J Clin Invest, 122, 3114, 10.1172/JCI61758

Lindquist, 2004, Visualizing dendritic cell networks in vivo, Nat Immunol, 5, 1243, 10.1038/ni1139

Ifrim, 2014, Trained immunity or tolerance: opposing functional programs induced in human monocytes after engagement of various pattern recognition receptors, Clin Vaccine Immunol, 21, 534, 10.1128/CVI.00688-13

Janssen, 2011, Fas determines differential fates of resident and recruited macrophages during resolution of acute lung injury, Am J Respir Crit Care Med, 184, 547, 10.1164/rccm.201011-1891OC

Boraschi, 1979, Defective tumoridical activity of macrophages of A/J mice. II. Comparison of the macrophages cytotoxic defect of A7J mice with that of lipid A-unresponsive C3H/HeJ mice, J Immunol, 122, 1592, 10.4049/jimmunol.122.4.1592

Netea, 2011, Trained immunity: a memory for innate host defense, Cell Host Microbe, 9, 355, 10.1016/j.chom.2011.04.006

Quintin, 2012, Candida albicans infection affords protection against reinfection via functional reprogramming of monocytes, Cell Host Microbe, 12, 223, 10.10167/j.chom.2012.06.006

Bowdish, 2007, Macrophages receptor implicated in the “adaptive” form of innate immunity, Microbe Infect, 9, 1680, 10.1016/j.micinf.2007.09.002

Vivier, 2011, Innate or adaptive immunity? The example of natural killer cells, Science, 331, 44, 10.1126/science.1198687

Netea, 2013, Training innate immunity: the changing concept of immunological memory in innate host defence, Eur J Clin Invest, 43, 881, 10.1111/eci.12132

Quintin, 2014, Innate immune memory: towards a better understanding of host defense mechanisms, Curr Opin Immunol, 29, 1, 10.1016/j.coi.2014.02.006

Ostuni, 2013, Latent enhancers activated by stimulation in differentiated cells, Cell, 152, 157, 10.1016/j.cell.2012.12.018

Murray, 2014, Macrophage activation and polarization: nomenclature and experimental guidelines, Immunity, 41, 14, 10.1016/j.immuni.2014.06.008