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Gene Review

DNA2  -  Dna2p

Saccharomyces cerevisiae S288c

Synonyms: DNA replication ATP-dependent helicase/nuclease DNA2, YHR164C
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High impact information on DNA2

  • Here we show that the endonucleases Dna2 and Fen1 act sequentially to facilitate the complete removal of the primer RNA [1].
  • For example, the DNA-binding domain of the yeast GAL4 protein interacts very poorly with nucleosome cores compared with naked DNA2 (and see below), and binding of other activators is even more strongly inhibited [2].
  • The human Bloom syndrome gene suppresses the DNA replication and repair defects of yeast dna2 mutants [3].
  • The DNA2 gene was cloned by complementation of the dna2ts gene phenotype [4].
  • Specifically, we show that deletion of PIF1 suppresses the lethality of a DNA2-null mutant [5].

Biological context of DNA2

  • This gene is identical to DNA2, encoding a helicase required for DNA replication [6].
  • DNA2 alleles were isolated that cause either temperature sensitivity, sensitivity to alkylation damage, or both [7].
  • Temperature-shift experiments indicate that Dna2p functions during late S phase, although dna2 mutants are not deficient in bulk DNA synthesis [6].
  • Frequencies of mitotic recombination and chromosome loss are elevated in dna2 mutants, also supporting a role for the protein in DNA synthesis [6].
  • As with its role in cell cycle progression, both the N-terminal and C-terminal regions, as well as the kinase domain of Tor1p, are required for rescue of dna2 mutants [6].

Anatomical context of DNA2

  • The ribosomal DNA locus and the nucleolus seem to be particularly sensitive to defects in dna2 mutants, although in dna2 mutants extrachromosomal ribosomal circles do not accumulate during the aging of a mother cell [8].

Associations of DNA2 with chemical compounds

  • The Dna2p is also distributed throughout the nucleus in cells growing in the presence of double-strand-break-inducing agents such as bleomycin [9].
  • Like FEN1, Dna2p can track over substrates with a non-Watson Crick base, such as a biotin, or a missing base within a chain [10].
  • Both the helicase and nuclease activities of Dna2Pho were inhibited by substrates with RNA segments at the 5'-end of flap DNA, whereas the nuclease activity of Dna2 from S. cerevisiae was reported to be stimulated by RNA segments in the 5'-tail (Bae, S.-H., and Seo, Y. S. (2000) J. Biol. Chem. 38022-38031) [11].

Physical interactions of DNA2

  • Dna2 was recently shown to dissociate flap-bound RPA, independent of cleavage [12].

Regulatory relationships of DNA2

  • The Saccharomyces cerevisiae DNA2 gene encodes a DNA-stimulated ATPase and DNA helicase/nuclease essential for DNA replication [13].
  • In agreement with these results, the proposed generation of double strand breaks in pol3-exo(-) rad27 mutants was suppressed by the overexpression of DNA2 [14].

Other interactions of DNA2

  • Full activity of the Dna2 helicase function is therefore not needed for viability, but is required for repairing damage and for tolerating loss of Ctf4 [7].
  • The X-ray sensitivity of dna2 mutants is suppressed by overexpression of a 5' to 3' exonuclease, the yeast FEN-1 structure-specific nuclease, encoded by the RAD27 gene, which also suppresses the growth defect of dna2-ts mutants [13].
  • SGS1 encodes a helicase with similar properties to Dna2 protein [13].
  • In addition, we identified physical and functional interactions between these proteins and found that RPA binds Dna2 predominantly through its large subunit, Rpa1 [15].
  • Arrest of dna2 mutants was RAD9 dependent, but deleting this checkpoint resulted in either no effect or suppression of defects, including the synthetic lethality with ctf4 [7].

Analytical, diagnostic and therapeutic context of DNA2


  1. RPA governs endonuclease switching during processing of Okazaki fragments in eukaryotes. Bae, S.H., Bae, K.H., Kim, J.A., Seo, Y.S. Nature (2001) [Pubmed]
  2. Nucleosome disruption and enhancement of activator binding by a human SW1/SNF complex. Kwon, H., Imbalzano, A.N., Khavari, P.A., Kingston, R.E., Green, M.R. Nature (1994) [Pubmed]
  3. The human Bloom syndrome gene suppresses the DNA replication and repair defects of yeast dna2 mutants. Imamura, O., Campbell, J.L. Proc. Natl. Acad. Sci. U.S.A. (2003) [Pubmed]
  4. A yeast gene required for DNA replication encodes a protein with homology to DNA helicases. Budd, M.E., Campbell, J.L. Proc. Natl. Acad. Sci. U.S.A. (1995) [Pubmed]
  5. Evidence suggesting that Pif1 helicase functions in DNA replication with the Dna2 helicase/nuclease and DNA polymerase delta. Budd, M.E., Reis, C.C., Smith, S., Myung, K., Campbell, J.L. Mol. Cell. Biol. (2006) [Pubmed]
  6. Characterization of Saccharomyces cerevisiae dna2 mutants suggests a role for the helicase late in S phase. Fiorentino, D.F., Crabtree, G.R. Mol. Biol. Cell (1997) [Pubmed]
  7. Dna2 mutants reveal interactions with Dna polymerase alpha and Ctf4, a Pol alpha accessory factor, and show that full Dna2 helicase activity is not essential for growth. Formosa, T., Nittis, T. Genetics (1999) [Pubmed]
  8. Mutations in DNA replication genes reduce yeast life span. Hoopes, L.L., Budd, M., Choe, W., Weitao, T., Campbell, J.L. Mol. Cell. Biol. (2002) [Pubmed]
  9. Dynamic localization of an Okazaki fragment processing protein suggests a novel role in telomere replication. Choe, W., Budd, M., Imamura, O., Hoopes, L., Campbell, J.L. Mol. Cell. Biol. (2002) [Pubmed]
  10. Dna2p helicase/nuclease is a tracking protein, like FEN1, for flap cleavage during Okazaki fragment maturation. Kao, H.I., Campbell, J.L., Bambara, R.A. J. Biol. Chem. (2004) [Pubmed]
  11. Helicase and nuclease activities of hyperthermophile Pyrococcus horikoshii Dna2 inhibited by substrates with RNA segments at 5'-end. Higashibata, H., Kikuchi, H., Kawarabayasi, Y., Matsui, I. J. Biol. Chem. (2003) [Pubmed]
  12. Significance of the dissociation of Dna2 by flap endonuclease 1 to Okazaki fragment processing in Saccharomyces cerevisiae. Stewart, J.A., Campbell, J.L., Bambara, R.A. J. Biol. Chem. (2009) [Pubmed]
  13. The pattern of sensitivity of yeast dna2 mutants to DNA damaging agents suggests a role in DSB and postreplication repair pathways. Budd, M.E., Campbell, J.L. Mutat. Res. (2000) [Pubmed]
  14. Okazaki fragment maturation in yeast. II. Cooperation between the polymerase and 3'-5'-exonuclease activities of Pol delta in the creation of a ligatable nick. Jin, Y.H., Ayyagari, R., Resnick, M.A., Gordenin, D.A., Burgers, P.M. J. Biol. Chem. (2003) [Pubmed]
  15. Bimodal interaction between replication-protein A and Dna2 is critical for Dna2 function both in vivo and in vitro. Bae, K.H., Kim, H.S., Bae, S.H., Kang, H.Y., Brill, S., Seo, Y.S. Nucleic Acids Res. (2003) [Pubmed]
  16. Genetic analyses of Schizosaccharomyces pombe dna2(+) reveal that dna2 plays an essential role in Okazaki fragment metabolism. Kang, H.Y., Choi, E., Bae, S.H., Lee, K.H., Gim, B.S., Kim, H.D., Park, C., MacNeill, S.A., Seo, Y.S. Genetics (2000) [Pubmed]
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