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Macrophage polarization
Macrophage polarization is a process when macrophage expresses different functional programs in response to microenvironmental signals.〔Mantovani, Alberto, et al. "Macrophage polarization: tumour-associated macrophages as a paradigm for polarized M2 mononuclear phagocytes." Trends in immunology 23.11 (2002): 549-555.〕 There are lots of functional states of macrophage polarization and they can be fully polarized and acquire specific phenotype like M1 (classically activated macrophages) or M2 (alternatively activated macrophages).〔Hamilton, T. A. (2014). "Myeloid Colony Stimulating Factors as Regulators of Macrophage Polarization." Frontiers in Immunology〕 These specific phenotypes depend on the tissue and specific microenvironment where macrophages are.〔Lawrence, Toby, and Gioacchino Natoli. "Transcriptional regulation of macrophage polarization: enabling diversity with identity." Nature reviews immunology 11.11 (2011): 750-761.〕 On one hand macrophage polarization is very important for host defense against pathogen, but on the other hand it is essential for maintenance of homeostasis.〔Hamilton, Thomas A., et al. "Myeloid colony-stimulating factors as regulators of macrophage polarization." Frontiers in immunology 5 (2014). 〕 Prolonged M1 type of macrophages is harmful for the organism and that is why tissue repair and restoration is necessary. M2 macrophages are responsible for that although they are connected with chronic infectious diseases.〔Benoit, Marie, Benoît Desnues, and Jean-Louis Mege. "Macrophage polarization in bacterial infections." The Journal of Immunology 181.6 (2008): 3733-3739. 〕
== M1 macrophages ==
Classically activated macrophages was named by Mackaness in the 1960s.〔Mackaness GB: Cellular resistance to infection. J Exp Med 1962,116:381-406. 〕 They express transcription factors like interferon-regulatory factor (IRF5), nuclear factor of kappa light polypeptide gene enhancer (NF-kB), activator protein 1 (AP-1) and STAT1.〔 Krausgruber, Thomas, et al. "IRF5 promotes inflammatory macrophage polarization and TH1-TH17 responses." Nature immunology 12.3 (2011): 231-238.〕 Typical are pro – inflammatory molecules (e.g. IFN-γ, IL-12, IL-23, TNF, IL-6, IL-1, specific chemokines and antigen presentation molecules).
M1 macrophages are irreplaceable during acute infectious diseases and provide host protection against intracellular bacteria or viruses by production of nitric oxide (NO) or reactive oxygen intermediates (ROI).〔Sica, Antonio, et al. "Macrophage polarization in tumour progression."Seminars in cancer biology. Vol. 18. No. 5. Academic Press, 2008. 〕〔Martinez, Fernando O., and Siamon Gordon. "The M1 and M2 paradigm of macrophage activation: time for reassessment." F1000prime reports 6 (2014). 〕 IFN-γ produced by Th1 lymphocytes is the most important cytokine which is responsible for classical macrophage activation. Natural killer (NK) cells and macrophages themselves produce IFN-γ as well. This cytokine regulates gene expression programs of macrophages like cytokine receptors, cell activation markers or cell adhesion molecules. For classical macrophage activation is necessary also lipopolysaccharide (LPS) – typical for gram negative bacteria – which is mainly recognised by TLR4, or lipoteichoic acid (LTA) – typical for gram positive bacteria. Granulocyte – macrophage colony stimulation factor (GM-CSF) stimulates M1 too.〔Martinez, Fernando O., and Siamon Gordon. "The M1 and M2 paradigm of macrophage activation: time for reassessment." F1000prime reports 6 (2014). 〕
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