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Structure, function and regulation of p63 and p73 (Review)

M Levrero, V De Laurenzi, A Costanzo, J Gong, G Melino and JYJ Wang


Cell Death and Differentiation (1999) 6, 1146 - 1153


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The p53 tumor suppressor gene is one of the most frequently mutated genes in human cancers.1 p53 is a sequence-specific transcription factor and plays a critical role in activating the expression of genes involved in cell cycle arrest or apoptosis under conditions of genotoxic stress.2,3 For over two decades, p53 was thought to be the only gene of its kind in the vertebrate genomes. This strong conviction, which was widely accepted in the p53 field, has now been proven to be incorrect. Two genes, referred to as p63 and p73, have been found to encode proteins that share a significant amino-acid identity in the transactivation domain, the DNA binding domain, and the oligomerization domainwith p53. In the short period since their cloning, a number of investigators have reported on the structure, the function and the regulation of p63 and p73. This review summarizes the current information on the p63 and the p73 genes, with a focus on the differences between the three members in this newly defined p53-gene family.


1. Hollstein M, Sidransky D and Vogelstein CC (1991) p53 mutations in human cancers. Science 253: 49 - 53

2. Ko LJ and Prives C (1996) p53: puzzle and paradigm. Genes Dev. 10: 1054 - 1072

3. Choisy-Rossi C and Yonish-Rouach E (1998) Apoptosis and the cell cycle: the p53 connection. Cell Death Differ. 5: 129 - 131

4. YangA,KaghadM,WangY, Gillett E, FlemingMD, Dotsch V,AndrewsNC,Caput D, McKeon F (1998) p63, a p53 homolog at 3q27-29, encodes multiple products with transactivating, death-inducing, and dominant-negative activities. Mol. Cell 2: 305 - 316

5. Kaghad M,Bonnet H,YangA, Creacier L,Biscan JC, Valent A, MintyA,ChalonP, Lellas JM, Dumont X, Ferrara P, McKeon F and Caput D (1997) Monoallelically expressed gene related to p53 at 1p36, a region frequentely deleted in neuroblastoma and other human cancers. Cell 90: 809 - 819

6. De Laurenzi V, Costanzo A, Barcaroli D , Terrinoni A, Falco M, Annicchiarico- Petruzzelli M, Levrero M and Melino G (1998) Two new p73 splice variants, gamma and delta,with different transcriptional activity. J. Exp.Med. 188: 1763 -1768

7. De Laurenzi V, Catani MV, Costanzo A, Terrinoni A, Corazzari M, Levrero M, Knight RA and Melino G (1999) Additional complexity in p73: induction by mitogens in lymphoid cells and identification of two new splicing variants epsilon and zeta. Cell Death Differ. 6: 389 - 390

8. Ueda Y, Hijikata M, Takagi S, Chiba T, Shimotohno K (1999) New p73 variants with altered C-terminal structures have varied transcriptional activities. Oncogene 18: 4993 - 4998

9. Zaika AI, Kovalev S, Marchenko ND and Moll UM (1999) Overexpression of the wild typep73 gene in breast cancer tissuesand cell lines.CancerRes. 59: 3257 -3263

10. Casciano I, Ponzoni M, Cunsolo C, Tonini G, Romani M (1999) Different p73 splicing variants are expressed in distinct tumour areas of a multifocal neuroblastoma. Cell Death Differ. 6: 391 - 393

11. Davison TS, Vagner C, KaghadM, Ayed A, Caput D, Arrowsmith CH (1999) p73 and p63 are homotetramers capable of weak heterotypic interactions with each other but not with p53. J. Biol. Chem. 274: 18709 - 18714

12. Di Como CJ, Gaiddon C and Prives C (1999) p73 function is inhibited by tumorderived p53 mutants in mammalian cells. Mol. Cell. Biol. 19: 1438 - 1449

13. Chi S-W, Ayeda A and Arrowsmith CH (1999) Solution structure of a conserved C-terminal domain of p73 with structural homology to the SAM domain.EMBO J. 18: 4438 - 4445

14. Schultz J,PointingCP, HofmanKand BorkP(1997)SAMas a protein interaction domain involved in developmental regulation. Protein Sci. 6: 249 - 253

15. Bork P, Koonin EV (1998) Predicting functions from protein sequences - where are the bottlenecks? Nature Genetics 18: 313 - 318

16. Takada N, OzakiT, IchimiyaS,TodoSandNakagawaraA(1999) Identification of a transactivation activity in theCOOH-terminal region of p73which is impaired in the naturally occurring mutants found in human neuroblastomas. Cancer Res. 59: 2810 - 2814

17. Jost CA, Marin MC and Kaelin Jr WG (1997) p73 is a human p53 related protein that can induce apoptosis. Nature 389: 191 - 194

18. Zhu J, Jiang J, Zhou W, Chen X (1998) The potential tumor suppressor p73 differentially regulates cellular p53 target genes. Cancer Res. 58: 5061 - 5065

19. Lee CW and La Thangue NB (1999) Promoter specificity and stability control of the p53-related protein p73. Oncogene 18: 44171 - 44181

20. Gong JG, Costanzo A,TamgHQ,MelinoG,KaelinWG,LevreroMandWangJYJ (1999) The tyrosine kinase c-Abl regulates p73 inapoptotic response to cisplatininduced DNA damage. Nature 399: 806 - 809

21. Agami R, Blandino G, OrenM, Shaul Y (1999) Interaction of c-Abl and p73a and their collaboration to induce apoptosis. Nature 399: 809 - 813

22. Yuan ZM, Shioya H, Ishiko T, Sun X, Gu J, Huang YY, Lu H, Kharbanda S, Weichselbaum R and Kufe D (1999) p73 is regulated by tyrosine kinase c-Abl in the apoptotic response to DNA damage. Nature 399: 814 - 817

23. Liu ZG, Baskaran R, Lea-Chou ET, Wood LD, Chen Y, Karin M and Wang JY (1996) Three distinct signalling responses by murine fibroblasts to genotoxic stress. Nature 384: 273 - 276

24. Baskaran R,Wood LD, Whitaker LL, Canman CE, Morgan SE, Xu Y, Barlow C, Baltimore D, Wynshaw-Boris A, Kastan MB and Wang JY (1997) Ataxia telangiectasia mutant protein activates c-Abl tyrosine kinase in response to ionizing radiation. Nature 387: 516 - 519

25. Balint E, Bates S, Vousden KH (1999) Mdm2 binds p73 alpha without targeting degradation. Oncogene 18: 3923 - 3929

26. Ongkeko WM, Wang XQ, Siu WY, Lau AW, Yamashita K, Harris AL, Cox LS, PoonRY(1999)MDM2andMDMXbindandstabilize thep53-relatedproteinp73. Curr. Biol. 12: 829 - 832

27. Zeng X, Chen L, Jost CA,MayaR,KellerD,WangX,Kaelin JrWG, Oren M, Chen J, Lu H (1999) MDM2 suppresses p73 function without promoting p73 degradation. Mol. Cell. Biol. 5, 3257 - 3266

28. Prives C and Hall PA. (1999) The p53 pathway. J. Pathol. 187: 112 - 126

29. Marin MC, Jost CA, Irwin MS, DeCaprio JA, Caput D and Kaelin Jr WG (1998) Viral oncoproteins discriminatebetween p53 and thep53homologp73. Mol. Cell. Biol. 18: 6316 - 6324

30. Prabhu NS, Somasundaram K, Satyamoorthy K, Herlyn M, El-Deiry WS (1998) p73beta, unlike p53, suppresses growth and induces apoptosis of human papillomavirus E6-expressing cancer cells Int. J. Oncol. 13: 5 - 9

31. DobbelsteinMandRoth J (1998) The large Tantigen of simian virus 40 binds and inactivates p53 but not p73. J. Gen. Virol. 79: 3079 - 3083

32. Higashino F, Pipas JM and Shenk T (1998) Adenovirus E4orf6 oncoprotein modulates the functionof thep53-related protein,p73. Proc.Natl. Acad.Sci.USA 95: 15683 - 15687

33. Steegenga WT, Shvarts A, Riteco N, Bos JL, Jochemsen AG (1999) Distinct regulation of p53 and p73 activity by adenovirus E1A, E1B, and E4orf6 proteins. Mol. Cell. Biol. 19: 3885 - 3894

34. Dobner T, Horikoshi N, Rubenwolf S, Shenk T (1996) Blockage by adenovirus E4orf6 of transcriptional activation by the p53 tumor suppressor. Science 272: 1470 - 1473

35. Tao W and Levine AJ (1999) P19(ARF) stabilizes p53 by blocking nucleocytoplasmic shuttling of Mdm2. Proc. Natl. Acad. Sci. USA 96: 6937 - 6941

36. ZhangYandXiongY(1999)Mutations in humanARFexon 2 disrupt its nucleolar localization and impair its ability to block nuclear export of MDM2 and p53. Mol. Cell 5: 579 - 591

37. Nimura Y, Mihara M, Ichimiya S, Sakiyama S, Seki N, Ohira M, Nomura N, Fujimori M, Adachi W, Amano J,HeM,PingYMand Nakagawara A(1998) p73, a gene related to p53, is notmutated in esophageal carcinomas. Int. J. Cancer 78: 437 - 440

38. Nomoto S, Haruki N, Kondo M, Konishi H, Takahashi T, Takahashi T and Takahashi T (1998) Search for mutations and examination of allelic expression imbalance of the p73 gene at 1p36.33 in human lung cancers. Cancer Res. 58, 1380 - 1383

39. Tsao H, Zhang X, Majewski P and Haluska F (1999) Mutational and expression analysis of the p73 gene in melanoma cell lines. Cancer Res. 59: 172 - 174

40. Ikawa S, Nakagawara A, Ikawa Y (1999) p53 family genes: structural comparison, expression and mutation. Cell Death Differ. 6: 1154 - 1161

41. Mills AA,ZhengB,WangXJ,Vogel H,RoopDRandBradleyA(1999) p63is ap53 homologue required for limb and epidermal morphogenesis. Nature 398: 708 -713

42. Yang A, Kaghad M, Wang Y, Gillett E, Fleming MD, Doetsch V, Andrews NC, Caput D and McKeon F (1999) p63 is essential for regenerative proliferation in limb, craniofacial and epithelial development. Nature 398: 714 - 718

43. Brodeur GM, SekhonG and GoldesteinMN (1977) Chromosomal aberrations in human neuroblastomas. Cancer 40: 2256 - 2263

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