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Sildenafil and tadalafil significantly delayed tumor outgrowth by 50 to 70% in immune-competent mice, although all mice ultimately died (Fig. S1, available at http://www.jem.org/cgi/content/full/jem.20061104/DC1). Similar results were obtained even if sildenafil treatment was started on day 7 after tumor challenge in the CT26WT model (Fig. S2). The fact that no difference in tumor outgrowth was seen between early versus late administration of sildenafil suggests that PDE5 inhibition does not appreciably affect the early phases of tumor uptake but rather influences the later stages of tumor outgrowth. Because the addition of sildenafil to cultured CT26WT cells did not increase their apoptosis or affect their doubling time (unpublished data), we conclude that sildenafil does not have a direct antitumor effect but rather interferes in hosttumor interactions. 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Almost immediately after the discovery of cAMP by Sutherland and colleagues, cyclic nucleotide PDE activity was described (Butcher and Sutherland, 1962). With the subsequent discovery of cGMP, it was found that both cAMP and cGMP could be hydrolyzed by the same type of activity, i.e., hydrolysis of the 3? cyclic phosphate bond. On the basis of substrate competition studies, it was clear that at least some of these activities must have the same catalytic site. In fact, many of the early studies on cyclic nucleotides were directed toward understanding PDE activity since at that time it was much easier to measure PDE activity than either cAMP or cGMP themselves or the enzymes that catalyzed their synthesis. With the advent of assays using radioactive substrate, it became clear that there were likely to be multiple forms of PDEs with different kinetic and regulatory properties (Thompson et al., 1979; Beavo et al., 1982). 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