title: Macrophage activation and polarization: nomenclature and experimental guidelines.
abstract: Description of macrophage activation is currently contentious and confusing. Like the biblical Tower of Babel, macrophage activation encompasses a panoply of descriptors used in different ways. The lack of consensus on how to define macrophage activation in experiments in vitro and in vivo impedes progress in multiple ways, including the fact that many researchers still consider there to be only two types of activated macrophages, often termed M1 and M2. Here, we describe a set of standards encompassing three principles-the source of macrophages, definition of the activators, and a consensus collection of markers to describe macrophage activation-with the goal of unifying experimental standards for diverse experimental scenarios. Collectively, we propose a common framework for macrophage-activation nomenclature.
authors.affiliation: Departments of Infectious Diseases and Immunology, St. Jude Children's Research Hospital, Memphis, TN 38105, USA. Electronic address: peter.murray@stjude.org.
authors.forename: Peter J
authors.initials: PJ
authors.lastname: Murray
authors.affiliation: Centre for Immunity, Infection, and Evolution, School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3JR, UK.
authors.forename: Judith E
authors.initials: JE
authors.lastname: Allen
authors.affiliation: Singapore Immunology Network, A(∗)STAR, 8A Biomedical Grove, Immunos Building, Level 4, Singapore138648, Singapore.
authors.forename: Subhra K
authors.initials: SK
authors.lastname: Biswas
authors.affiliation: Center for the Prevention of Cardiovascular Disease, New York University School of Medicine, Smilow 7, 522 First Avenue, New York, NY, USA.
authors.forename: Edward A
authors.initials: EA
authors.lastname: Fisher
authors.affiliation: Division of Medicine, Rayne Institute, University College London, 5 University Street, London WC1 6JJ, UK.
authors.forename: Derek W
authors.initials: DW
authors.lastname: Gilroy
authors.affiliation: Department Dermatology, University Medical Center Mannheim, University of Heidelberg, 68167 Mannheim, Germany.
authors.forename: Sergij
authors.initials: S
authors.lastname: Goerdt
authors.affiliation: Sir William Dunn School of Pathology, University of Oxford, Headington, Oxford, OX1 3RE, UK.
authors.forename: Siamon
authors.initials: S
authors.lastname: Gordon
authors.affiliation: Department of Medicine, University of Melbourne and Royal Melbourne Hospital, Parkville, VIC 3050, Australia.
authors.forename: John A
authors.initials: JA
authors.lastname: Hamilton
authors.affiliation: Hospital for Special Surgery and Weill Medical College, Cornell University, 535 East 70(th) Street, New York, NY 10021, USA.
authors.forename: Lionel B
authors.initials: LB
authors.lastname: Ivashkiv
authors.affiliation: Centre d'Immunologie de Marseille-Luminy, 13009 Marseille, France.
authors.forename: Toby
authors.initials: T
authors.lastname: Lawrence
authors.affiliation: University of Milan School of Medicine, Istituto Clinico Humanitas, Via Manzoni 56, 20089 Rozzano, Milan, Italy.
authors.forename: Massimo
authors.initials: M
authors.lastname: Locati
authors.affiliation: University of Milan School of Medicine, Istituto Clinico Humanitas, Via Manzoni 56, 20089 Rozzano, Milan, Italy.
authors.forename: Alberto
authors.initials: A
authors.lastname: Mantovani
authors.affiliation: Botnar Research Centre, Nuffield Department of Orthopaedics, Rheumatology, and Musculoskeletal Sciences, University of Oxford, Headington, Oxford OX3 7LD, UK.
authors.forename: Fernando O
authors.initials: FO
authors.lastname: Martinez
authors.affiliation: Infectious Diseases, Aix Marseille University, 27 Boulevard Jean Moulin, 13285 Marseille, France.
authors.forename: Jean-Louis
authors.initials: JL
authors.lastname: Mege
authors.affiliation: Department of Cell Biology, University of Maryland, College Park, MD 20742, USA.
authors.forename: David M
authors.initials: DM
authors.lastname: Mosser
authors.affiliation: Department of Experimental Oncology, European Institute of Oncology, Via Adamello 16, 20146 Milan, Italy.
authors.forename: Gioacchino
authors.initials: G
authors.lastname: Natoli
authors.affiliation: Department of Biology, Massachusetts Institute of Technology, Cambridge, MA 02139, USA.
authors.forename: Jeroen P
authors.initials: JP
authors.lastname: Saeij
authors.affiliation: Genomics & Immunoregulation, LIMES-Institute, University of Bonn, 32115 Bonn, Germany.
authors.forename: Joachim L
authors.initials: JL
authors.lastname: Schultze
authors.affiliation: Department of Microbiology and Immunology, University of Maryland School of Medicine, Baltimore, MD 21201, USA.
authors.forename: Kari Ann
authors.initials: KA
authors.lastname: Shirey
authors.affiliation: Istituto Clinico Humanitas, Via Manzoni 56, 20089 Rozzano, Milan, Italy; Department of Pharmaceutical Sciences, Università degli Studi del Piemonte Orientale "Amedeo Avogadro," Via Bovio 6, 28100 Novara, Italy.
authors.forename: Antonio
authors.initials: A
authors.lastname: Sica
authors.affiliation: Microbiology & Immunology, University of Louisville School of Medicine, 319 Abraham Flexner Way, Louisville, KY 40292, USA.
authors.forename: Jill
authors.initials: J
authors.lastname: Suttles
authors.affiliation: Kennedy Institute of Rheumatology, University of Oxford, Headington, Oxford, OX3 7FY, UK.
authors.forename: Irina
authors.initials: I
authors.lastname: Udalova
authors.affiliation: Laboratory of Cellular and Molecular Immunology, Vrije Universiteit Brussel, Pleinlaan 2, 1050 Brussels, Belgium; Laboratory of Myeloid Cell Immunology, VIB, Pleinlaan 2, 1050 Brussels, Belgium.
authors.forename: Jo A
authors.initials: JA
authors.lastname: van Ginderachter
authors.affiliation: Department of Microbiology and Immunology, University of Maryland School of Medicine, Baltimore, MD 21201, USA.
authors.forename: Stefanie N
authors.initials: SN
authors.lastname: Vogel
authors.affiliation: Laboratory of Parasitic Diseases, National Institute of Allergy and Infectious Diseases, National Institutes of Health, Bethesda, MD 20892, USA.
authors.forename: Thomas A
authors.initials: TA
authors.lastname: Wynn
mesh_terms: D000818:Animals; D016178:Granulocyte-Macrophage Colony-Stimulating Factor / Q000276:immunology; D017408:Guidelines as Topic; D006801:Humans; D008262:Macrophage Activation / Q000276:immunology*; D016173:Macrophage Colony-Stimulating Factor / Q000276:immunology; D008264:Macrophages / Q000276:immunology*; D051379:Mice; D012106:Research; D009626:Terminology as Topic
chemical_list: D016173:Macrophage Colony-Stimulating Factor; D016178:Granulocyte-Macrophage Colony-Stimulating Factor