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. 2019 Nov 14;10:2612. doi: 10.3389/fimmu.2019.02612

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Copyright © 2019 Lai, Wang, Fagenson, Sun, Saredy, Lu, Nanayakkara, Yang, Yu, Shao, Drummer, Johnson, Saaoud, Zhang, Yang, Xu, Mastascusa, Cueto, Fu, Wu, Sun, Zhu, Qin, Yu, Fan, Shen, Sun, Rogers, Choi, Wang and Yang.

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

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(A) Macrophages (Mφ) can be polarized into 10 (potentially more) different subsets, and the markers and main functions of 10 Mφ subsets are different (PMID: 24998279; 25319333; 25973901). M1, M4, and Mox are proinflammatory while the rest of the Mφ subsets are anti-inflammatory. (B) Mechanism I: The 10 macrophage (Mφ) subset markers (30) are differentially expressed in macrophages from various tissues, and lung, liver, spleen, and small intestine upregulate more M1 Mφ markers than M2 Mφ markers in the physiologic condition in comparison to lean ATMφ. (1) Retnla, CD163, and MRC1 are relatively adipose tissue-specific Mφ (ATMφ) markers; (2) STAB1, NFE2L2, and SRXN1 are relatively bone marrow-specific Mφ markers; (3) ARG1 is a relatively specific Mφ marker for peritoneum, Chil4 is a relatively specific Mφ marker for lung, IL1B is a relatively specific Mφ marker for liver, PDE4DIP and HMOX1 are relatively specific Mφ markers for spleen, and CXCL9 is a relatively specific Mφ marker for small intestine. (C) Mφ subset markers are differentially expressed in various tissues. (D) Tissue Mφ have different compositions of Mφ subsets as judged by the expressions of Mφ subset markers. (1) The genes of macrophage subtypes such as Mhb, Mhem, and HA-mac are relatively adipose tissue-specific. (2) The genes of Mox are bone marrow macrophage-specific. (3) Retnla is a M2a subset marker in peripheral tissues. (4) ILIB and CXCL9 are specific markers for small intestine M1 macrophages. (5) ARG1 is a specific marker for peritoneal M2a and M2c macrophages.