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Gene Review

PHO2  -  Pho2p

Saccharomyces cerevisiae S288c

Synonyms: BAS2, D2350, GRF10, General regulatory factor 10, Regulatory protein PHO2, ...
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High impact information on PHO2

  • The yeast regulatory gene PHO2 encodes a homeo box [1].
  • Similarly, we find that low levels of transcription by Bas1/Bas2 are TATA-independent, whereas high levels are TATA-dependent [2].
  • Phosphorylation of a fourth site blocks the interaction of Pho4 with the transcription factor Pho2 [3].
  • Induction of the yeast gene PHO5 is mediated by the transcription factors PHO2 and PHO4 [4].
  • BAS1 has a Myb motif and activates HIS4 transcription only in combination with BAS2 [5].

Biological context of PHO2


Anatomical context of PHO2

  • The vacuole inheritance defect in vac5-1 cells is dependent on the presence of the Pho85 kinase and its targets Pho4p and Pho2p [11].

Associations of PHO2 with chemical compounds

  • These results suggest that glycine affects Bas1p activation of transcription rather than DNA binding and that Bas2p is not required for this affect [12].
  • The transcription factors BAS1 and BAS2/PHO2, which are also regulators of the histidine pathway, participate in the regulation of the purine biosynthetic pathway [13].
  • The basal level transcription of the HIS4 gene is under the control of two genes, BAS1 and BAS2, which are also required for the control of purine biosynthesis [6].
  • In contrast, a LexA-Bas1p fusion activated the lexAop reporter in a BAS2-dependent and adenine-regulated fashion [14].
  • Nevertheless, a Bas2p cysteine-free mutant was not sufficient to confer resistance to oxidative stress [15].

Physical interactions of PHO2

  • Grf10 protein purified from a bacterial expression system binds DNA cooperatively with Swi5 in vitro [10].
  • The Swi5 zinc-finger and Grf10 homeodomain proteins bind DNA cooperatively at the yeast HO promoter [10].
  • Gel retardation assays with the PHO2 protein revealed a binding region that lay between the two PHO4-binding sites [16].
  • Direct biochemical analysis shows that the BAS2 gene encodes a protein that binds to both the HIS4 and PHO5 promoters [6].
  • The other element shows strong sequence identity with the Bas2p binding site and appears to be involved in basal and phosphate-mediated regulation of LEU4 [17].

Regulatory relationships of PHO2

  • A Bas1p-VP16 fusion protein activated expression in a bas1bas2 strain but no glycine effect was observed while a Bas1p-Bas2p fusion protein activated expression to a lesser extent with a slight stimulation by glycine [12].
  • We show here that Pho4 and the homeobox protein Pho2 interact in vivo and act cooperatively to activate the PHO5 UAS, with interaction being regulated by the phosphate switch [18].
  • Previous studies have shown that GIT1 expression is regulated by inorganic phosphate (P(i)) availability through the transcription factors Pho2p and Pho4p [19].
  • We conclude that efficient transcription of yeast AMP biosynthesis genes requires interaction between Bas1p and Bas2p which is promoted in the presence of a metabolic intermediate whose synthesis is controlled by feedback inhibition of Ade4p acting as the purine nucleotide sensor within the cell [20].

Other interactions of PHO2


Analytical, diagnostic and therapeutic context of PHO2

  • This fragment was used to evict the genomic copy and with appropriate genetic crosses we proved, that the cloned gene is PHO2 [25].
  • Final purification of the 83-kDa GRF10 protein was achieved by cooperative interaction-based DNA affinity chromatography [26].
  • These data combined with LacZ fusions and northern blot analysis allow us to show that synthesis of enzymes for all 10 steps involved in purine de novo synthesis is repressed in the presence of adenine and requires BAS1 and BAS2 for optimal expression [27].
  • Immunoprecipitation experiments showed that Pho4p and Pho2p form a complex on a 36-bp sequence bearing an upstream activation site (UAS) and protein binding assays indicated that these proteins interact directly [28].


  1. The yeast regulatory gene PHO2 encodes a homeo box. Bürglin, T.R. Cell (1988) [Pubmed]
  2. TATA-dependent and TATA-independent transcription at the HIS4 gene of yeast. Pellman, D., McLaughlin, M.E., Fink, G.R. Nature (1990) [Pubmed]
  3. Roles of phosphorylation sites in regulating activity of the transcription factor Pho4. Komeili, A., O'Shea, E.K. Science (1999) [Pubmed]
  4. Phosphorylation of the transcription factor PHO4 by a cyclin-CDK complex, PHO80-PHO85. Kaffman, A., Herskowitz, I., Tjian, R., O'Shea, E.K. Science (1994) [Pubmed]
  5. BAS1 has a Myb motif and activates HIS4 transcription only in combination with BAS2. Tice-Baldwin, K., Fink, G.R., Arndt, K.T. Science (1989) [Pubmed]
  6. Multiple global regulators control HIS4 transcription in yeast. Arndt, K.T., Styles, C., Fink, G.R. Science (1987) [Pubmed]
  7. Long-range interactions at the HO promoter. McBride, H.J., Brazas, R.M., Yu, Y., Nasmyth, K., Stillman, D.J. Mol. Cell. Biol. (1997) [Pubmed]
  8. Mutations in the pho2 (bas2) transcription factor that differentially affect activation with its partner proteins bas1, pho4, and swi5. Bhoite, L.T., Allen, J.M., Garcia, E., Thomas, L.R., Gregory, I.D., Voth, W.P., Whelihan, K., Rolfes, R.J., Stillman, D.J. J. Biol. Chem. (2002) [Pubmed]
  9. Determining the requirements for cooperative DNA binding by Swi5p and Pho2p (Grf10p/Bas2p) at the HO promoter. Brazas, R.M., Bhoite, L.T., Murphy, M.D., Yu, Y., Chen, Y., Neklason, D.W., Stillman, D.J. J. Biol. Chem. (1995) [Pubmed]
  10. The Swi5 zinc-finger and Grf10 homeodomain proteins bind DNA cooperatively at the yeast HO promoter. Brazas, R.M., Stillman, D.J. Proc. Natl. Acad. Sci. U.S.A. (1993) [Pubmed]
  11. A truncated form of the Pho80 cyclin redirects the Pho85 kinase to disrupt vacuole inheritance in S. cerevisiae. Nicolson, T.A., Weisman, L.S., Payne, G.S., Wickner, W.T. J. Cell Biol. (1995) [Pubmed]
  12. Transcriptional regulation of the one-carbon metabolism regulon in Saccharomyces cerevisiae by Bas1p. Subramanian, M., Qiao, W.B., Khanam, N., Wilkins, O., Der, S.D., Lalich, J.D., Bognar, A.L. Mol. Microbiol. (2005) [Pubmed]
  13. Coregulation of purine and histidine biosynthesis by the transcriptional activators BAS1 and BAS2. Daignan-Fornier, B., Fink, G.R. Proc. Natl. Acad. Sci. U.S.A. (1992) [Pubmed]
  14. Evidence that complex formation by Bas1p and Bas2p (Pho2p) unmasks the activation function of Bas1p in an adenine-repressible step of ADE gene transcription. Zhang, F., Kirouac, M., Zhu, N., Hinnebusch, A.G., Rolfes, R.J. Mol. Cell. Biol. (1997) [Pubmed]
  15. Redox regulation of AMP synthesis in yeast: a role of the Bas1p and Bas2p transcription factors. Pinson, B., Gabrielsen, O.S., Daignan-Fornier, B. Mol. Microbiol. (2000) [Pubmed]
  16. The two positively acting regulatory proteins PHO2 and PHO4 physically interact with PHO5 upstream activation regions. Vogel, K., Hörz, W., Hinnen, A. Mol. Cell. Biol. (1989) [Pubmed]
  17. Additive activation of yeast LEU4 transcription by multiple cis elements. Hu, Y., Kohlhaw, G.B. J. Biol. Chem. (1995) [Pubmed]
  18. The transcription factor, the Cdk, its cyclin and their regulator: directing the transcriptional response to a nutritional signal. Hirst, K., Fisher, F., McAndrew, P.C., Goding, C.R. EMBO J. (1994) [Pubmed]
  19. Posttranscriptional regulation of Git1p, the glycerophosphoinositol/glycerophosphocholine transporter of Saccharomyces cerevisiae. Almaguer, C., Fisher, E., Patton-Vogt, J. Curr. Genet. (2006) [Pubmed]
  20. Yeast AMP pathway genes respond to adenine through regulated synthesis of a metabolic intermediate. Rébora, K., Desmoucelles, C., Borne, F., Pinson, B., Daignan-Fornier, B. Mol. Cell. Biol. (2001) [Pubmed]
  21. Possible cross-regulation of phosphate and sulfate metabolism in Saccharomyces cerevisiae. O'Connell, K.F., Baker, R.E. Genetics (1992) [Pubmed]
  22. Function of the PHO regulatory genes for repressible acid phosphatase synthesis in Saccharomyces cerevisiae. Yoshida, K., Ogawa, N., Oshima, Y. Mol. Gen. Genet. (1989) [Pubmed]
  23. A 28-bp segment of the Saccharomyces cerevisiae PHO5 upstream activator sequence confers phosphate control to the CYC1-lacZ gene fusion. Sengstag, C., Hinnen, A. Gene (1988) [Pubmed]
  24. Sequence of the PHO2-POL3 (CDC2) region of chromosome IV of Saccharomyces cerevisiae. Simon, M., Bénit, P., Vassal, A., Dubois, C., Faye, G. Yeast (1994) [Pubmed]
  25. The sequence of the Saccharomyces cerevisiae gene PHO2 codes for a regulatory protein with unusual aminoacid composition. Sengstag, C., Hinnen, A. Nucleic Acids Res. (1987) [Pubmed]
  26. Identification and purification of a protein that binds DNA cooperatively with the yeast SWI5 protein. Brazas, R.M., Stillman, D.J. Mol. Cell. Biol. (1993) [Pubmed]
  27. Role of the myb-like protein bas1p in Saccharomyces cerevisiae: a proteome analysis. Denis, V., Boucherie, H., Monribot, C., Daignan-Fornier, B. Mol. Microbiol. (1998) [Pubmed]
  28. The transcriptional activators of the PHO regulon, Pho4p and Pho2p, interact directly with each other and with components of the basal transcription machinery in Saccharomyces cerevisiae. Magbanua, J.P., Ogawa, N., Harashima, S., Oshima, Y. J. Biochem. (1997) [Pubmed]
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