From:  Macrophages as predictors and new targets for immunotherapy in colorectal cancer

 Effectiveness of combination therapy based on ICIs with TME inhibitors or plant extracts in pre-clinical animal tumor models.

TargetsCombination therapyIn vivo tumor modelTumor growth and survivalMacrophage activityChanges in the TME
Combination therapy of anti-PD-1/PD-L1 agents and TAM inhibitors
MS4A4A [100]Anti-MS4A4A mAb plus anti-PD-1 mAbMC38-bearing C57BL/6 mice, CT26-bearing BALB/c mice (n = 5 per group)Inhibited tumor growth (monotherapy); suppressed tumor growth and improved survival (combination)An anti-MS4A4A monotherapy: decreased M2-TAM infiltration (F4/80+CD206+); downregulated M2-like markers (CD206) and immunosuppressive molecules (CD39, SIRPα); upregulated M1-like markers (iNOS, MHCII)Reduced T cell exhaustion marker co-expression (PD-L1+LAG3+, PD-L1+TIM3+); enhanced CD8+ T cell effector function and proliferation (IFN-α, Ki67)
TREM2 [98]CAR-T cells, secreted PD-1-TREM2 scFVMC38-bearing C57BL/6J mice (n = 4)Inhibited tumor progression; enhanced survivalDecreased percentages of M2-like TAMs (CD11b+Gr1+; F4/80+CD206+)Increased CD8+ T cell infiltration; decreased MDSCs amount; enhanced perforin and granzyme B release; elevated intratumoral cytokines (IL-2, IL-15, TNF-α, IFN-γ)
SPHK1 [93]SPHK1 inhibitors (PF543 or SKI II) plus anti-PD-1 AbMC38 and CT26 liver metastatic models in immunocompetent mice (n = 5–12 per group)Partially repressed CRLM (monotherapy); enhanced metastasis inhibition; prolonged survival (combination)Reduced total TAMs; decreased p-SPHK1+ TAMsIncreased intermediate exhausted CD8+ T cells with a moderate PD-1 level and PD-1lowCD8+ T cells expressing memory T cell markers (IL-7Ra and Ly6C)
CSF1R [102]PLX3397 plus anti-PD-1 mAb plus anti-CTLA-4 mAbC57BL/6J bearing subcutaneous MC38 miceCompared with the monotherapy, PLX3397 showed a potent synergic effect when it was combined with an anti-PD-1 or an anti-CTLA-4 mAbDecreased expression of macrophage markers CD206 and F4/80 (monotherapy PLX3397)-
Combination therapy of anti-PD-1/PD-L1 agents and TME inhibitors
A20 [95]A20-knockdown plus, anti-PD-1 AbMetastatic BALB/c mouse model bearing A20-knockdown tumor cells (CT-26) (n = 4–8 per group)Inhibited tumor growth in the lung, subcutaneous tumor growth; decreased number of tumor nodules in lung tissue; the overall survival was remarkably longerIncreased F4/80+ macrophagesIncreased CD3+ and CD8+ T cells; increased amount of granzyme B+ immune cells
MIF [96]anti-PD-1 Ab plus anti-MIF AbMC38-bearing (subcutaneously injected); C57BL/6J miceSuperior tumor rejection; reduction of tumor size--
CD73 [97]AB680 plus anti-PD-1 AbCT26-bearing BALB/c mice (n = 3 per group)Tendency to reduce tumor growth compared to monotherapyIncreased the proportion of M1-like macrophages, no effect on M2-like macrophages (combination therapy); decreased the proportion of M2-like macrophages but had no effect on M1 macrophages (PD-1 blockade)Trend towards increased CD8+/CD4+ ratio
Kinases [92]Regorafenib plus anti-PD-1 AbSyngeneic murine MSS CT26 and MSI MC38 CC modelsSustained suppression of liver metastasis even after the end of treatmentReduced F4/80+ and CD206+ macrophages; increased iNOS+ macrophagesReduced number of Tregs under regorafenib treatment; increased intratumoral levels of IFN-γ, a pharmacodynamic marker of cytotoxic T cell activity under an anti-PD-1 therapy
WNT11/CAMKII [112]KN93 plus anti-PD-1 AbC57BL/6 mouse liver metastases model with MC38 cells (n = 5 per group)Decreased liver metastasisReduced CD206+ macrophage infiltration in liver metastasisIncreased infiltration of I-A/I-E+CD8+ T cells in liver metastasis; KN93 induced the expression of CXCL10 and CCL4 and reduced the expression of IL17D
Combination therapy of anti-PD-1/PD-L1 agents and epigenetic modulators
HDAC [110]Tucidinostat plus anti-PD-L1 AbCT26 subcutaneously injected into BALB/c or C57 BL/6 mice (n = 7 per group)Tumor growth suppression; improved mice survivalReduced the number of CD45+CD11b+F4/80+ TAMs; increased the proportion of CD45+CD11b+F4/80+/MHCII+ M1-like macrophagesIncreased CD45+ lymphocytes count, proportion of tumor-infiltrating CD4+ T cells (CD45+CD3+CD4+) cells and CD8+ T cells (CD45+CD3+CD8+) cells; decreased expression of the exhaustion marker PD-1; increased serum level of CCL5 and IFN-γ
Class I HDACis (1, 2, 3, and 10), VEGFR [103]Chidamide plus cabozantinib/regorafenib plus anti-PD-1 AbCT26-Bearing (subcutaneously injected) BALB/c miceInhibited tumor growth; increased survival rate; tumor shrinkage after discontinued treatmentDecreased amount of CD11b+ TAMs in tumorsUpregulated enrichment of IFN pathway gene signature and downregulated enrichment of the T cell gene signatures; reduced levels of polymorphonuclear-MDSCs in the tumor
Class I HDACis (1, 2, 3, and 10), VEGFR [103]Chidamide plus cabozantinib/regorafenib plus anti-CTLA-4 AbCT26-Bearing (subcutaneously injected) BALB/c miceInhibited tumor growth; enhanced ORRSuppressed macrophage/monocyte gene signaturesEnhanced T cell cytotoxicity
HDAC [111]AVS100 plus anti-PD-1 AbCT26-bearing BALB/c mice (n = 6–7 per group)An increase in responders from 35% in anti-PD-1 treatment alone to 80% in combination therapy; did not relapse after termination of the treatment; resistant to a subsequent tumor challengeIncreased IFN-TAMs; decreased regulatory and angiogenic TAMsUp-regulation of genes involved in T cell activation, T cell effector function, and NF-κB signaling; increased expression of genes controlling IFN and NF-κB signaling (Stat4, Nfkb1, Rel, and Il12rb2), the inositol-3-phosphate/Akt pathway, and immune infiltration (Hcst, Itpr1, and Itgal1) with AVS100 treatment, independent of anti-PD-1 therapy
Combination therapy of anti-PD-1/PD-L1 agents and plant extract components
[104]Lobeline plus anti-PD-1 AbC57BL/6 mice bearing MC38 tumors (subcutaneously injected) (n = 6 per group)Reduced tumor growth; upregulated SLURP1 expression in tumors; suppressed cancer cell proliferationIncrease in the number of intratumor CD45+CD11b+F4/80+ M1-like macrophages and decrease in the number of CD45+CD11b+CD206+ M2-like macrophagesDecreased Foxp3 protein levels; increased granzyme B levels
[113]Astragaloside IV plus anti-PD-1 AbBALB/c mice bearing CT26 tumors (injected into the axillary fat pad) (n = 8)Suppressed tumor growth more effectively than monotherapyAstragaloside monotherapy: altered M2-like macrophage polarization toward M1-like (decreased CD11b+F4/80+CD206hi M2-like macrophages; increased CD11b+F4/80+MHCIIhi M1-like macrophages)Increased the number of cytotoxic T cells; increased IFN-γ and IL12p70 secretion; decreased TGF-β secretion
[105]SFIH plus anti-PD-1 AbMC38-bearing (subcutaneously injected), C57BL/6 mice (n = 7–8 per group)Significant reduction in tumor volume compared with an anti-PD-1 Ab; no effect in the SFIH groupIncreased CD11b+, F4/80+, MHCII+, CD206 M1-like macrophage infiltration; no changes in the CD11b+, F4/80+, MHCII, CD206+ TAM populationIncreased CD3+ and CD8+ T cells, release of granzyme B, expression of IFN-γ and CXCL9; decreased population of CD11b+, GR+ MDSCs, decreased expression of TGF-β1, TGF-β2, VEGFA, and IDO2; downregulated TGF-β signaling
[114]CYP plus anti-PD-1 mAbXenograft models, subcutaneously injected: MC38-bearing C57BL/6 mice (n = 6) and CT26-bearing BALB/c mice (n = 3)Combination therapy was more effective in reducing tumor volume and size (CYP alone had no significant anti-tumor effect); increased survival in miceDecreased number of infiltrating macrophages, mainly CD206+ TAMsIncreased number of CD8+ T cells in the tumor; increased expression of granzyme B

Ab: antibody; CAR-T: chimeric antigen receptor T; CCL: C-C motif chemokine ligand; CRLM: colorectal cancer liver metastasis; CTLA-4: cytotoxic T lymphocyte-associated antigen-4; CXCL: C-X-C motif chemokine ligand; CYP: Chinese yam polysaccharide; HDAC: histone deacetylase; ICIs: immune checkpoint inhibitors; IFN: interferon; IL: interleukin; mAb: monoclonal Ab; MDSCs: myeloid-derived suppressor cells; MHCII: major histocompatibility complex II; MIF: migration inhibitory factor; MSI: microsatellite instability; MSS: microsatellite stable; ORR: objective response rate; PD-1: programmed cell death-1; PD-L1: programmed cell death ligand-1; scFv: single-chain variable fragment; SFIH: sesquiterpene lactones derived from Inula helenium L.; SPHK1: sphingosine kinase; TAM: tumor-associated macrophage; TGF: transforming growth factor; TME: tumor microenvironment; TNF: tumor necrosis factor; Tregs: regulatory T cells; VEGF: vascular endothelial growth factor.