PMC:6562565 / 22855-32185 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/6562565","sourcedb":"PMC","sourceid":"6562565","source_url":"https://www.ncbi.nlm.nih.gov/pmc/6562565","text":"The Pleiotropic Effects of Adenosine in the Tumor Microenvironment\nAlong with T cells, many other cell types in the TME including other protective or suppressive immune infiltrates, tumor-associated fibroblasts, endothelial cells and cancer cells also express functional ARs (3, 261–266). Here we briefly describe the effects of adenosine-induced signaling on them (Figure 2).\nFigure 2 Overview of the pleiotropic effects of adenosine in the TME. Adenosine enables tumor cells to escape immune-surveillance by limiting the functionality of multiple potentially protective immune infiltrates including T cells, DCs, NK cells, macrophages and neutrophils, while enhancing the activity of immunosuppressive cell-types, such as MDSCs and Tregs. In addition, adenosine not only assists tumor cells in co-opting adjacent fibroblasts for support, but also induces the formation of new blood vessels. Adenosine also affects the capacity of some tumor cell-types to survive, proliferate, migrate and invade neighboring tissues (HPC, bone marrow-derived hematopoietic progenitor cells).\n\nDendritic Cells\nThe biology of DCs, specialized antigen presenting cells (APCs) and critical messengers between the innate and adaptive immune system, can be severely impaired by adenosinergic signaling. For example, it has been reported that adenosine binding to A2BR (267) halts the differentiation of monocytes to DCs (267, 268). In addition, adenosine averts inflammatory stimulus-induced DC activation (269), whereas A2AR (270) and A2BR triggering (267, 271, 272) diminishes the capacity of DCs to prime Th1 immune responses (267, 270, 271) but rather prompts DCs to skew naïve T cell differentiation toward Th2 (267, 271) and Th17 (272) lineages. Adenosine-treated DCs exhibit decreased expression or secretion of TNFα and IL-12 (268–271, 273) and enhanced production of IL-5 (270), IL-10 (267, 268, 270, 273), IL-6 (267, 272) and TGFβ (267). Moreover, such DCs are less motile due to chemokine receptor downregulation (274), and have a tolerogenic effect on the TME due to overexpression of TGFβ (267), IL-10, IDO-1 (267), arginase-2 (267, 275), as well as A2AR-mediated upregulation of PD-L2 (276). Finally, adenosine compels DCs to secrete the proangiogenic factors VEGF (267, 275) in an A2BR-dependent manner as well as IL-8 (267).\n\nMacrophages\nStimulation of adenosine receptors hinders the differentiation of monocytes to macrophages, probably through cAMP accumulation (277). Moreover, by engaging A1R (278), A2AR (278–282), A3R (281, 283) or setting off Gαs-paired ARs (284), adenosine reduces the pro-inflammatory activity of macrophages by dampening their ability to produce IL-12 (279), TNFα (278–280, 282, 283), macrophage inflammatory protein-1α (MIP1α) (281), nitric oxide (278, 285) and superoxide (284). In addition, by triggering A2AR (282, 286, 287), A2BR (288, 289) or unidentified ARs, adenosine promotes an M2 polarization of macrophages by inducing upregulation of arginase-1 (288, 290), IL-10 (279, 286, 289) and VEGF production (282, 287).\n\nNK Cells\nA2AR stimulation by adenosine not only restricts the NK maturation (291), but also their capacity for stimulus-induced CD69 upregulation (292, 293), proliferation (291, 294) as well as IFNγ (292, 293) and TNFα (294, 295) production. Furthermore, largely via A2AR triggering, adenosine diminishes target cell killing by NK cells (292, 294, 296–298).\n\nNeutrophils\nAdenosine exerts a variety of inhibitory effects on neutrophils. For example, triggering of A2AR (299–303), A3R (304), non-specified A2Rs (304–307) or ARs dampens their ability to adhere (299, 305, 308, 309), transmigrate (310), secrete TNFα and MIP1α (300, 306), degranulate (301, 302, 304, 311), perform Fc receptor-mediated phagocytosis (307) and produce superoxide (299, 301–303). Interestingly, others claim that A2AR and A2BR signaling has been shown to suppress VEGF production (310). Finally, A2AR stimulation prompts neutrophils to secrete higher levels of PGE2 (312).\n\nMDSCs\nA2BR-mediated signaling boosts differentiation of bone marrow hematopoietic progenitors toward a tolerogenic myeloid-derived cell subset, the MDSCs (313). Moreover, A2AR activation promotes IL-10 production by MDSCs (314) and treatment with an adenosine analog results in increased expression of CD73 (313). Finally, it has also been shown that A2BR stimulation on MDSCs augments VEGF production (315).\n\nStromal Cells\nAdenosine, along with critically contributing to the establishment of a tolerogenic TME, also enables tumors to subvert fibroblasts into supporting them and to induce formation of new blood vessels, processes essential to their growth and dissemination. CAFs, for example, are stromal cells that support tumors by secreting the pro-metastatic and angiogenic (316) chemokine CXCL12 (317), as well as the mitogenic (318) fibroblast growth factor 2 (FGF2) (319). Triggering of A2BR on the surface of CAFs boosts expression of both CXCL12 and FGF2 (320) whereas A2AR-induced signaling stimulates their proliferation (54). As previously mentioned, adenosine can stimulate VEGF secretion by multiple cell types found within the TME, which in turn promotes angiogenesis by supporting the survival, migration and proliferation of endothelial cells (321, 322). It has also been shown that A2AR (323) and A2BR (66) stimulation diminishes production of the anti-angiogenic factor thrombospondin-1 by endothelial cells. Furthermore, adenosine not only augments the rate of intra-tumoral nutrient delivery by inducing vasodilatation (324), but also hinders leukocyte extravasation (325) through downregulation of adhesion molecules, such as E-selectin (326, 327) VCAM-1 (326, 327) and ICAM-1 (327, 328) on the surface of endothelial cells, as well as by limiting vascular permeability (325, 327, 329–331) through A2BR activation (329–331). Finally, signaling initiated by triggering of A2AR (332, 333), A2BR (334–336) or non-specified ARs prompts endothelial cells to overexpress CD73 (334) as well as the proangiogenic factors VEGF (332, 333, 335, 336), IL-8 (335) and basic fibroblast growth factor (bFGF) (335, 336).\n\nTumor Cells\nAdenosine binding to ARs on the surface of cancer cells has a profound impact on their biology. For example, the triggering of A1R (337, 338), A2AR (54, 339, 340), and A3R (339, 341–343) induces a variety of cellular responses that augment cancer cell survival such as AKT and ERK1/2 stimulation, as well as Bad inactivation (342). Additional responses to AR signaling contributing to bolstered cancer cell survival include upregulation of Bcl2 (343), downregulation of p53 (338) and Bax (343) as well as aversion of caspase-9 (343) and caspase-3 (54, 343, 344) activation. Paradoxically, extracellular adenosine has also been demonstrated to cause cancer cell death either by setting off A1R (345, 346), A2AR (341, 347), A2BR (348, 349), and A3R (339, 350–354) or via induction of AMPK activation upon its cellular uptake and subsequent conversion to AMP (345).\nMoreover, A1R (337, 355), A2AR (341, 356), A2BR (344, 357–359), and A3R (343, 360) stimulation augments cancer cell proliferation through activation of PLC (356), protein kinase C-delta (PKC-δ) (356), AKT (356, 357), ERK1/2 (356–360), JNK (356, 358), and p38 (358). Furthermore, triggering of the ARs leads to upregulation of cyclins A (343), B (358), D (343, 358), E (337, 343, 358), estrogen receptor-α (355) as well as downregulation of the cell-cycle inhibitors p27 (337) and p21 (343, 358). Surprisingly, though, activation of A2BR (349) and A3R (341, 350, 353, 361–363) has also been reported to result in a potent cytostatic effect.\nMotility (358, 359, 364–369) and invasiveness (358, 359, 367, 370) are additional features of cancer cells that are boosted upon engagement of A1R (364, 365), A2AR (366), A2BR (358, 359, 367, 368), and A3R (369, 370). In terms of mechanisms, signaling initiated by these receptors promotes filopodia formation (367) as well as expression of matrix metalloproteases (MMPs) (358, 359, 370) and FXYD5 (359), a cell membrane glycoprotein known to drive metastasis by reducing cell adhesion (371). In contrast, others claim that A3R triggering hinders the motility and invasiveness of cancer cells (372, 373). Finally, A2AR (374), A2BR (369, 375), and A3R (369, 375–377) stimulation on the surface of cancer cells promotes angiogenesis by boosting secretion of the pro-angiogenic factors VEGF (369, 375, 377), IL-8 (369, 375), angiopoietin 2 (376), and erythropoietin (374).\nThe contrasting consequences of triggering particular ARs, on the survival, proliferation or migration and invasiveness of tumor cells most probably occur due to the heterogeneity between cells and/or experimental settings employed to assess them. For instance, two different cancer cell lines of distinct tissue origin could have profoundly diverse AR expression profiles as well as different ability to transmit/terminate signaling initiated by these receptors. Moreover, they might have different capacity to produce adenosine, which once released into the medium can trigger ARs in an autocrine fashion. Finally, different concentrations used between experiments, as well as limited specificity of the AR agonists/antagonists, probably constitute additional factors contributing to the observed 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