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The Cisuralian Epoch of the Permian Period
299 to 271 million years ago


The Cisuralian epoch, a period of well over 20 million years, was - as far as the terrestrial ecosystem went - a fairly stable period ruled over by a diverse selection of , including the large finbacks such as the carnivorous , Ctenospondylus, and Secodontosaurus, all of which attained a lengths of up to 3 meters, as well as semi-aquatic and the big Eryops and smaller eel-like Archeria. This was a biome strongly tied to water and to a plant-arthropod-fish food chain; Diadectes and Edaphosaurus were the only herbivores. In this respect it continued the pattern of the great coal swamps. The drier upland was inhabited by a different fauna, mostly smaller insectivores with the herbivorous and proto- Tetraceratops as significant newcomers. Both in the uplands and the lowlands continued to represent an astonishing diversity of forms.

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PeriodEpochAgeWhen beganDuration (Ma)
PermianGuadalupianRoadian270.62.6
CisuralianKungurian275.65.0
Artinskian284.48.8
Sakmarian294.610.2
Asselian299.04.4
CarboniferousPennsylvanianGzhelian303.94.9




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To test this hypothesis, tumor-derived CD11b+ MDSCs were isolated from C26GM-bearing BALB/c mice. MDSC suppressive activity was determined by admixing MDSCs with CFSE-labeled HA-specific CD8+ (clone 4) or CD4+ (6.5) T cells pulsed with their relevant peptide in the presence or absence of sildenafil (Fig. 7, A and B). Although the addition of tumor-derived MDSCs significantly impaired antigen-specific T cell proliferation as demonstrated by the low percentage of CFSElow clonotypic T cells, sildenafil almost completely restored both CD4+ and CD8+ responsiveness of these antigen-specific T cells. The absence of sildenafil-mediated enhancement in T cell function in the groups lacking CD11b+ cells underscores the targeted role of sildenafil on the MDSC population. Because in a Th1 cell–prone environment MDSC suppression is only NOS2 dependent (34), we examined the role of PDE5 in MDSCs in a C57BL/6 background where NOS2–/– mice are also available. CD11b+ MDSCs were isolated from either C57BL/6-NOS2+/+ or B16GM-bearing C57BL/6-NOS2–/– B16GM melanoma-bearing mice. A suppression assay was performed by stimulating OVA-specific CD4+ T cells with the relevant peptide in the presence or absence of MDSCs obtained from either NOS2+/+ or NOS2–/– tumor-bearing mice (Fig. 7 C). Although the addition of C57BL/6-NOS2+/+ MDSCs induced considerable T cell suppression, no suppression was observed with MDSCs from NOS2–/– mice. Furthermore, although PDE5 inhibition reversed MDSC suppression in NOS2+/+ mice, sildenafil failed to augment T cell responsiveness in the NOS2–/–-derived MDSC suppression assay. These results confirm the role of NOS2 in MDSC-mediated T cell suppression (Fig. 7 C) and underscore the ability of PDE5 inhibition to reverse the two major suppressive pathways in MDSCs (ARG1 and NOS2). Yohimbine is a safe agent with few known adverse effects. It is administered daily in a dose of 5.4 mg (1 tab) 3 times/d. 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