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Table 1.

Efficacy of a single injection of entolimod in increasing 40-day survival of lethally irradiated NHPs when administered at different dose levels within 1–48 hours after TBI.

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Fig 1.

Improved survival of non-human primates (NHPs) injected with entolimod 1–48 hours after lethal irradiation.

Kaplan-Meier plots of non-human primate (NHP) survival over the 40 days following exposure to LD50/40 –LD75/40 doses of total body irradiation (TBI) are shown. Time frame of entolimod efficacy (panels A, B) was evaluated in studies Rs-03 (treatment at 1 h after LD75/40 TBI; N = 10) and Rs-06 (treatment at 16, 25, or 48 h after LD75/40 TBI; N = 8–12). Dose-dependence of entolimod efficacy (panels C, D) was tested in studies Rs-09 (treatment at 1 h after LD50/40 TBI; N = 18) and Rs-14 (treatment at 25 h after LD50/40 TBI; N = 10).

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Fig 2.

Accelerated hematological recovery of peripheral blood in NHPs injected with entolimod 16–48 hours post-irradiation.

NHPs were treated with a single injection of entolimod at 16–48 hours after LD50/40 or LD75/40 TBI. A, C, E, G: Effect of 40 μg/kg entolimod administered at different time points (16, 25 or 48 hours) after 6.5 Gy TBI (LD75/40; study Rs-06; N = 8–12). B, D, F, H: Effect of different entolimod doses (10 or 40 μg/kg) administered at 25 hours after 6.75 Gy TBI (LD75/40; study Rs-14; N = 10). Cytopenia/anemia thresholds: dotted lines—Grade 3 (platelets <50,000/μL; neutrophils <1,000/μL; hemoglobin <80 g/L); dashed lines—Grade 4 (platelets <10,000/μL; neutrophils <500/μL; hemoglobin <65 g/L). Error bars represent standard errors.

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Table 2.

Mean nadir values of neutrophils, platelets and hemoglobin in peripheral blood following total body irradiation and vehicle or entolimod treatment.

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Fig 3.

Enhanced morphological recovery of hematopoietic and lymphoid organs in NHPs treated with entolimod post-irradiation.

NHPs were treated with a single injection of 40 μg/kg entolimod 16, 25 or 48 hours after LD75/40 total body irradiation (TBI). Tissue morphology was assessed 40 days post-irradiation and compared to that in control NHPs treated with vehicle 16 hours after LD75/40 TBI. Representative histological images (hematoxylin-eosin staining) of sternum bone marrow sections, thymuses, spleens and mesenteric lymph nodes of animals that survived to study termination on Day 40 post-TBI (study Rs-06) are shown. Scale bars: 100 μm for bone marrow, 200 μm for thymus, spleen, and lymph node.

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Table 3.

Histological evaluation of hematopoietic/lymphoid organs from NHPs that survived to day 40 after 6.5 Gy TBI and vehicle or entolimod treatment (study Rs-06)

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Fig 4.

Entolimod treatment ameliorates radiation damage in the gastrointestinal (GI) tract.

A, B. Small intestine sections from NHPs 8 hours after exposure to 6.5 Gy TBI and treatment with vehicle or 40 μg/kg entolimod 1 h later (study Rs-04). Blue—DAPI nuclear staining, red—smooth muscle actin immunostaining. A. TUNEL staining showing fewer apoptotic cells (green) in GI crypts of entolimod-treated NHPs (scale bar 100 μm); B. SOD2 immunostaining (green) showing more positive cells in GI villi (arrowheads) and lamina propria (arrows) of entolimod-treated NHPs (scale bar 50 μm). C, D. Small intestine sections of NHPs 7 days after exposure to 11 Gy TBI and treatment with vehicle or 40 μg/kg entolimod 4 h later (study Rs-22). C. Visualization of proliferating cells in the jejunum crypts: EdU (10 mg/kg i.v. 1 h before euthanasia) inclusion in replicating DNA (green) and phosphohistone 3 immunostaining of mitotic cells (red) showing more intensive proliferation of GI crypts in entolimod-treated NHPs (scale bar– 200 μm). D. H&E staining of ileum sections: upper panels—low magnification (scale bar– 200 μm), lower panels—high magnification (scale bar– 50 μm).

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Table 4.

Histological evaluation of GI tract segments on day 7 after 11 Gy TBI and vehicle or 40 μg/kg entolimod treatment at +4 hours (study Rs-22, N = 4/group).

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Fig 5.

Effect of entolimod treatment on G-CSF and IL-6 levels in peripheral blood of irradiated NHPs.

A, B: Effect of different entolimod doses administered 1 h after LD50/40 TBI (6.75 Gy; study Rs-09; N = 18). C, D: Effect of different entolimod doses administered 25 h after LD50/40 TBI (6.75 Gy; study Rs-14; N = 10). E, F: Comparison of dose-dependence of background-adjusted Area Under the Curve (AUC0-24) values for G-CSF and IL-6 after entolimod treatment given 1 h versus 25 h after LD50/40 TBI (with dashed log-linear regression lines). Error bars represent standard errors.

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Fig 6.

Schematic presentation of mechanism(s) underlying anti-acute radiation syndrome (ARS) effects of entolimod.

Entolimod binding to Toll-like receptor 5 (TLR5) initiates a cascade of events, all merging at attenuation of major pathological processes—leading causes of death in ARS: damage to hematopoietic (HP) and gastrointestinal (GI) systems resulting in bleeding and sepsis. The immediate TLR5-dependent effectors include anti-oxidants (e.g., SOD2), anti-apoptotic factors (both NF-κB-dependent (i.e., IAP and Bcl family members [6770]), and NF-κB-independent (i.e., PI3K/AKT, MKP7 and STAT3 [54, 7173]), hematopoietic cytokines (e.g., G-CSF and IL-6 [49]), anti-infective factors [54, 9095] and processes (e.g., neutrophil mobilization). In addition, stimulation of TLR5 is expected to inhibit radiation-induced aseptic inflammation involved in secondary tissue damage [64] e.g. via induction of an anti-inflammatory cytokine IL-10, IL-1β antagonist (IL-1βa) [76] and stimulation of mesenchymal stem cells (MSC) known to express TLR5 [77, 78] and to have anti-inflammatory properties [79]. Together with fibroblasts that can be induced to proliferate via TLR5 stimulation [104], MSC may also contribute to wound-healing processes [79]. Dashed lines show all molecular connections downstream of TLR5 that are not directly established for entolimod, but are extrapolated from published data on TLR5-dependent effects of flagellin.

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