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Activation of p. 38 MAPK signaling by overexpression of constitutively active upstream kinases such as MKK3 and MKK6 is sufficient to trigger a G2M checkpoint in cultured cells Garner et al., 2. Wang et al., 2. 00. By use of a transgene expressing active MKK6 under control of the Lck promoter, which specifically targets gene expression to T lymphocytes, it has been shown that active MKK6 causes G2 arrest in vivo, with resultant deregulated differentiation and accumulation of immature thymocytes Diehl et al., 2. In the case of stress mediated cell cycle arrest, p. MAPK contributes to G2 arrest after various types of stress. It is noteworthy that activation of p. MAPK has been primarily described for non IR ionizing radiation types of stress, and a direct connection between damaged DNA and p. Liu et al., 1. 99. However, recent evidence supports the argument that activation of a G2M checkpoint after genotoxic stress, such as that caused by double strand breaks after IR Dent et al., 2. Wang et al., 2. 00. DNA alkylating agents Katayama et al., 2. MAPK. Thus, similar to the well known regulator of the cell cycle, Chk. MAPK seems to play an important role in controlling the G2M transition under a variety of conditions Pearce and Humphrey, 2. Another physiological example for p. MAPK mediated G2M arrest is osmotic stress in renal epithelial cells. Survival of these proliferating cells under continuous osmotic stress requires functional checkpoints. Inactivation of p. MAPK allows cells that have not reached the G2M border to progress into mitosis prematurely, causing significant cell death under conditions of osmotic stress Dmitrieva et al., 2. There are several potential mechanisms to explain how p. MAPK controls G2M progression, including the ability of p. MAPK to maintain inhibitory phosphorylation of human Cdc. B phosphatase at serine 3. Cdc. 25. C at serine 2. Bulavin et al., 2. Katayama et al., 2. Tarn et al., 2. 00. It has also been suggested that an inhibitory effect of p. MAPK on Cdc. 25. C phosphatase can be mediated through regulation of another stress kinase, Chk. Wang et al., 2. 00. Katayama et al., 2. It is interesting that p. MAPK phosphorylates the same sites in Cdc. B and Cdc. 25. C as Chk. Bulavin et al., 2. Phosphorylation of these sites triggers sequestration by 1. Cdc. 25. B and Cdc. C from removing inhibitory phosphates on Cdc. Cdk. 1 to allow activation of this key kinase that drives the G2M transition. Cdc. 25. A phosphatase, which has a central role in promoting the G1S transition, has been found to be critical for mitotic entry as well Molinari et al., 2. Zhao et al., 2. 00. The ability of Cdc. A to regulate cell cycle progression from G2 to M phase can be explained by at least two mechanisms. Phosphorylation of Cdc. A at serine 1. 78 and threonine 5. Chk. 1 kinase facilitates the recruitment of 1. Cdc. 25 phosphatases Chen et al., 2. Cdc. 2 kinase, and thus Cdc. Cdc. 25. A were available. As p. 38 MAPK and Chk. Cdc. 25 phosphatases, the assumption can be made that under certain conditions p. MAPK may be involved in the phosphorylation of Cdc. A at S1. 78 and T5. Cdc. 25. A and Cdc. B1 complex. However, this requires further in vivo confirmation, as Chk. Chk. 1, does not phosphorylate these two sites Chen et al., 2. In addition, Cdk. Cdc. 25. A phosphatase during the G1S transition, which in turn can be negatively regulated by p. MAPK previously discussed, has been recently found to be critical in phosphorylation of threonine 1. Cdc. 25. C in Xenopus laevis Margolis et al., 2. This phosphorylation may further facilitate the release of 1. Cdc. 25. C phosphatases at the end of interphase, thus allowing Cdc. C to access Cdc. 2 kinase and promote entry into mitosis. Notably, phosphorylation of Cdc. MAPK is somewhat unusual, as p. MAPK almost invariably phosphorylate serine or threonine residues that are followed by a proline. However, there have been at least two reports that p. MAPK phosphorylates serine and threonine residues that are not followed by proline in the microtubule associated protein Tau Reynolds et al., 2. Tab. 1 protein Cheung et al., 2. These observations indicate that potential residues in substrates phosphorylated by p. MAPK cannot always be predicted based on the amino acid sequence of a protein. Therefore, there may be additional p. 4 Square Saltine Crackers. MAPK substrates that have roles in cell cycle regulation.