Conservation of telomere protein complexes: Shuffling through evolution

Critical Reviews in Biochemistry and Molecular Biology

Volumn 44, Issue 6, 2009, Pages 434-446

  Linger, Benjamin R ^a   Price, Carolyn M ^a  

^a UNIVERSITY OF CINCINNATI COLLEGE OF MEDICINE   (United States)

Author keywords

CST; Gene duplication; Myb repeat; OB fold; POT1; Shelterin; TRF1

Indexed keywords

CHROMOSOME PROTEIN; CONSERVED TELOMERE MAINTENANCE COMPONENT 1 PROTEIN; POT1 PROTEIN; POZ1 PROTEIN; RAP1 PROTEIN; SHELTERIN; STN1 PROTEIN; TELOMERE PROTEIN; TELOMERIC REPEAT BINDING FACTOR 1; TELOMERIC REPEAT BINDING FACTOR 2; TPZ1 PROTEIN; UNCLASSIFIED DRUG;

DNA REPLICATION; GENE DUPLICATION; MOLECULAR EVOLUTION; NONHUMAN; PRIORITY JOURNAL; PROTEIN DNA BINDING; PROTEIN FUNCTION; PROTEIN STRUCTURE; REVIEW; TELOMERE;

ANIMALS; EVOLUTION, MOLECULAR; GENES, DUPLICATE; HUMANS; PROTEIN BINDING; TELOMERE; TELOMERE-BINDING PROTEINS;

EUKARYOTA; MAMMALIA; SACCHAROMYCETALES;

EID: 70350020161     PISSN: 10409238     EISSN: 15497798     Source Type: Journal    
DOI: 10.3109/10409230903307329     Document Type: Review

References (116)

1. Amiard S, Doudeau M, Pinte S, Poulet A, Lenain C, Faivre-Moskalenko C, Angelov D, Hug N, Vindigni A, Bouvet P, Paoletti J, Gilson E, Giraud-Panis MJ 2007. A topological mechanism for TRF2-enhanced strand invasion. Nat Struct Mol Biol 14:147-154.
2. Azzalin CM, Reichenbach P, Khoriauli L, Giulotto E and Lingner J. 2007. Telomeric repeat containing RNA and RNA surveillance factors at mammalian chromosome ends. Science 318:798-801.
3. Baker AM, Fu Q, Hayward W, Lindsay SM and Fletcher TM. 2009. Te Myb/SANT domain of the telomere-binding protein TRF2 alters chromatin structure. Nucleic Acids Res 37:5019-5031.
4. Baumann P and Cech TR. 2001. Pot1, the putative telomere end-binding protein in fssion yeast and humans. Science 292:1171-1175.
5. Bianchi A and Shore D. 2007. Increased association of telomerase with short telomeres in yeast. Genes Dev 21:1726-1730. (Pubitemid 47076472)
6. Bianchi A and Shore D. 2008. How telomerase reaches its end: mechanism of telomerase regulation by the telomeric complex. Mol Cell 31:153-165.
7. Bianchi A, Smith S, Chong L, Elias P and de Lange T. 1997. TRF1 is a dimer and bends telomeric DNA. Embo J 16:1785-1794.
8. Bianchi A, Negrini S and Shore D. 2004. Delivery of yeast telomerase to a DNA break depends on the recruitment functions of Cdc13 and Est1. Mol Cell 16:139-146.
9. Bilaud T, Koering CE, Binet-Brasselet E, Ancelin K, Pollice A, Gasser SM and Gilson E. 1996. Te telobox, a Myb-related telo-meric DNA binding motif found in proteins from yeast, plants and human. Nucleic Acids Res 24:1294-1303.
10. Bilaud T, Brun C, Ancelin K, Koering CE, Laroche T and Gilson E. 1997. Telomeric localization of TRF2, a novel human telobox protein. Nat Genet 17:236-239.
11. Blasco MA. 2007. Te epigenetic regulation of mammalian telom-eres. Nat Rev Genet 8:299-309.
12. Broccoli D, Smogorzewska A, Chong L and de Lange T. 1997. Human telomeres contain two distinct Myb-related proteins, TRF1 and TRF2. Nat Genet 17:231-235.
13. Casteel DE, Zhuang S, Zeng Y, Perrino FW, Boss GR, Goulian M and Pilz RB. 2009. A DNA polymerase-{alpha}{middle dot} primase cofactor with homology to replication protein A-32 regulates DNA replication in mammalian cells. J Biol Chem 284:5807-5818.
14. Celli GB and de Lange T. 2005. DNA processing is not required for ATM-mediated telomere damage response after TRF2 deletion. Nat Cell Biol 7:712-718.
15. Cesare AJ, Quinney N, Willcox S, Subramanian D and Grifth JD. 2003. Telomere looping in P. sativum (common garden pea). Plant J 36:271-279.
16. Cesare AJ, Grof-Vindman C, Compton SA, McEachern MJ and Grifth JD. 2008. Telomere loops and homologous recombination-dependent telomeric circles in a Kluyveromyces lactis tel-omere mutant strain. Mol Cell Biol 28:20-29.
17. Chandra A, Hughes TR, Nugent CI and Lundblad V. 2001. Cdc13 both positively and negatively regulates telomere replication. Genes Dev 15:404-414.
18. Chang M, Arneric M and Lingner J. 2007. Telomerase repeat addition processivity is increased at critically short telomeres in a Tel1-dependent manner in Saccharomyces cerevisiae. Genes Dev 21:2485-2494.
19. Chen Y, Yang Y, van Overbeek M, Donigian JR, Baciu P, de Lange T and Lei M. 2008. A shared docking motif in TRF1 and TRF2 used for diferential recruitment of telomeric proteins. Science 319:1092-1096.
20. Chong L, van Steensel B, Broccoli D, Erdjument-Bromage H, Hanish J, Tempst P and de Lange T. 1995. A human telomeric protein. Science 270:1663-1667.
21. Churikov D and Price CM. 2008. Pot1 and cell cycle progression cooperate in telomere length regulation. Nat Struct Mol Biol 15:79-84.
22. Cooper JP, Nimmo ER, Allshire RC and Cech TR. 1997. Regulation of telomere length and function by a Myb-domain protein in fs-sion yeast. Nature 385:744-747.
23. Court R, Chapman L, Fairall L and Rhodes D. 2005. How the human telomeric proteins TRF1 and TRF2 recognize telomeric DNA: a view from high-resolution crystal structures. EMBO Rep 6:39-45.
24. Croy JE and Wuttke DS. 2006. Temes in ssDNA recognition by telomere-end protection proteins. Trends Biochem Sci 31:516-525.
25. Crumet N, Carlson RL, Drutman SB and Shampay J. 2006. A truncated acidic domain in Xenopus TRF1. Gene 369:20-26.
26. Dejardin J and Kingston RE. 2009. Purifcation of proteins associated with specifc genomic Loci. Cell 136:175-186.
27. de Lange T. 2004. T-loops and the origin of telomeres. Nat Rev Mol Cell Biol 5:323-329.
28. Denchi EL and de Lange T. 2007. Protection of telomeres through independent control of ATM and ATR by TRF2 and POT1. Nature 448:1068-1071.
29. De Rycker M, Venkatesan RN, Wei C and Price CM. 2003. Vertebrate tankyrase domain structure and sterile alpha motif (SAM)-mediated multimerization. Biochem J 372:87-96.
30. Fairall L, Chapman L, Moss H, de Lange T and Rhodes D. 2001. Structure of the TRFH dimerization domain of the human telo-meric proteins TRF1 and TRF2. Mol Cell 8:351-361.
31. Flory MR, Carson AR, Muller EG and Aebersold R. 2004. An SMC-domain protein in fssion yeast links telomeres to the meiotic centrosome. Mol Cell 16:619-630.
32. Fouche N, Cesare AJ, Willcox S, Ozgur S, Compton SA and Grifth JD. 2006. Te basic domain of TRF2 directs binding to DNA junctions irrespective of the presence of TTAGGG repeats. J Biol Chem 281:37486-37495.
33. Gao H, Cervantes RB, Mandell EK, Otero JH and Lundblad V. 2007. RPA-like proteins mediate yeast telomere function. Nat Struct Mol Biol 14:208-214.
34. Garvik B, Carson M and Hartwell L. 1995. Single-stranded DNA arising at telomeres in cdc13 mutants may constitute a specifc signal for the RAD9 checkpoint [published erratum appears in Mol Cell Biol 1996 16:457]. Mol Cell Biol 15:6128-6138.
35. Gilson E and Geli V. 2007. How telomeres are replicated. Nat Rev Mol Cell Biol 8:825-838.
36. Goudsouzian LK, Tuzon CT and Zakian VA. 2006. S. cerevisiae Tel1p and Mre11p are required for normal levels of Est1p and Est2p telomere association. Mol Cell 24:603-610.
37. Grandin N, Reed SI and Charbonneau M. 1997. Stn1, a new Saccharomyces cerevisiae protein, is implicated in tel-omere size regulation in association with Cdc13. Genes Dev 11:512-527.
38. Grandin N, Damon C and Charbonneau M. 2001. Ten1 functions in telomere end protection and length regulation in association with Stn1 and Cdc13. Embo J 20:1173-1183.
39. Grifth JD, Comeau L, Rosenfeld S, Stansel RM, Bianchi A, Moss H and de Lange T. 1999. Mammalian telomeres end in a large duplex loop. Cell 97:503-514.
40. Grossi S, Puglisi A, Dmitriev PV, Lopes M and Shore D. 2004. Pol12, the B subunit of DNA polymerase alpha, functions in both telomere capping and length regulation. Genes Dev 18:992-1006.
41. Guo X, Deng Y, Lin Y, Cosme-Blanco W, Chan S, He H, Yuan G, Brown EJ and Chang S. 2007. Dysfunctional telomeres activate an ATM-ATR-dependent DNA damage response to suppress tumorigenesis. Embo J 26:4709-4719.
42. Hanaoka S, Nagadoi A, Yoshimura S, Aimoto S, Li B, de Lange T and Nishimura Y. 2001. NMR structure of the hRap1 Myb motif reveals a canonical three-helix bundle lacking the positive surface charge typical of Myb DNA-binding domains. J Mol Biol 312:167-175.
43. Hector RE, Shtofman RL, Ray A, Chen BR, Nyun T, Berkner KL and Runge KW. 2007. Tel1p Preferentially Associates with Short Telomeres to Stimulate Teir Elongation. Mol Cell 27:851-858.
44. Hirano Y, Fukunaga K and Sugimoto K. 2009. Rif1 and rif2 inhibit localization of tel1 to DNA ends. Mol Cell 33:312-322.
45. Hockemeyer D, Daniels JP, Takai H and de Lange T. 2006. Recent expansion of the telomeric complex in rodents: Two distinct POT1 proteins protect mouse telomeres. Cell 126:63-77.
46. Hockemeyer D, Palm W, Else T, Daniels JP, Takai KK, Ye JZ, Keegan CE, de Lange T and Hammer GD. 2007. Telomere protection by mammalian Pot1 requires interaction with Tpp1. Nat Struct Mol Biol 14:754-761.
47. Horvath MP, Schweiker VL, Bevilacqua JM, Ruggles JA and Schultz SC. 1998. Crystal structure of the Oxytricha nova tel-omere end binding protein complexed with single strand DNA. Cell 95:963-974.
48. Houghtaling BR, Cuttonaro L, Chang W and Smith S. 2004. A dynamic molecular link between the telomere length regulator TRF1 and the chromosome end protector TRF2. Curr Biol 14:1621-1631.
49. Jacob NK, Lescasse R, Linger BR and Price CM. 2007. Tetrahymena POT1a regulates telomere length and prevents activation of a cell cycle checkpoint. Mol Cell Biol 27:1592-1601.
50. Kanoh J and Ishikawa F. 2001. spRap1 and spRif1, recruited to telom-eres by Taz1, are essential for telomere function in fssion yeast. Curr Biol 11:1624-1630.
51. Kanoh J and Ishikawa F. 2003. Composition and conservation of the telomeric complex. Cell Mol Life Sci 60:2295-2302.
52. Karamysheva ZN, Surovtseva YV, Vespa L, Shakirov EV and Shippen DE. 2004. A C-terminal Myb extension domain defnes a novel family of double-strand telomeric DNA-binding proteins in Arabidopsis. J Biol Chem 279:47799-47807.
53. Kim H, Lee OH, Xin H, Chen LY, Qin J, Chae HK, Lin SY, Safari A, Liu D and Songyang Z. 2009. TRF2 functions as a protein hub and regulates telomere maintenance by recognizing specifc peptide motifs. Nat Struct Mol Biol 16:372-379.
54. Kim SH, Davalos AR, Heo SJ, Rodier F, Zou Y, Beausejour C, Kaminker P, Yannone SM and Campisi J. 2008. Telomere dysfunction and cell survival: roles for distinct TIN2-containing complexes. J Cell Biol 181:447-460.
55. Klempnauer KH and Sippel AE. 1987. Te highly conserved amino-terminal region of the protein encoded by the v-myb oncogene functions as a DNA-binding domain. EMBO J 6:2719-2725.
56. Konig P and Rhodes D. 1997. Recognition of telomeric DNA. Trends Biochem Sci 22:43-47.
57. Konishi A and de Lange T. 2008. Cell cycle control of telomere protection and NHEJ revealed by a ts mutation in the DNA-binding domain of TRF2. Genes Dev 22:1221-1230.
58. Lee J, Mandell EK, Tucey TM, Morris DK and Lundblad V. 2008. Te Est3 protein associates with yeast telomerase through an OB-fold domain. Nat Struct Mol Biol 15:990-997.
59. Lei M, Podell ER, Baumann P and Cech TR. 2003. DNA self-recognition in the structure of Pot1 bound to telomeric single-stranded DNA. Nature 426:198-203.
60. Lei M, Podell ER and Cech TR. 2004. Structure of human POT1 bound to telomeric single-stranded DNA provides a model for chromosome end-protection. Nat Struct Mol Biol 11:1223-1229.
61. Lei M, Zaug AJ, Podell ER and Cech TR. 2005. Switching human tel-omerase on and of with hPOT1 protein in vitro. J Biol Chem 280:20449-20456.
62. Li B, Oestreich S and de Lange T. 2000. Identifcation of human Rap1: implications for telomere evolution. Cell 101:471-483.
63. Li B, Espinal A and Cross GA. 2005. Trypanosome telomeres are protected by a homologue of mammalian TRF2. Mol Cell Biol 25:5011-5021.
64. Li S, Makovets S, Matsuguchi T, Blethrow JD, Shokat KM and Blackburn EH. 2009. Cdk1-dependent phosphorylation of Cdc13 coordinates telomere elongation during cell-cycle progression. Cell 136:50-61.
65. Liu D, O'Connor MS, Qin J and Songyang Z. 2004a. Telosome-A mammalian telomere associated complex formed by multiple telomeric proteins. J Biol Chem 279:51338-51342.
66. Liu D, Safari A, O'Connor MS, Chan DW, Laegeler A, Qin J and Songyang Z. 2004b. PTOP interacts with POT1 and regulates its localization to telomeres. Nat Cell Biol 6:673-680.
67. Loayza D and De Lange T. 2003. POT1 as a terminal transducer of TRF1 telomere length control. Nature 243:1013-1018.
68. Lundblad V. 2006. Budding yeast telomeres, pp. 345-386. In: de Lange T, Lundblad V, Blackburn E, eds, Telomeres, Cold Spring Harbor Laboratory Press. New York, USA.
69. Marcand S, Gilson E and Shore D. 1997. A protein-counting mechanism for telomere length regulation in yeast [see comments]. Science 275:986-990.
70. Martin V, Du LL, Rozenzhak S and Russell P. 2007. Protection of tel-omeres by a conserved Stn1-Ten1 complex. Proc Natl Acad Sci USA 104:14038-14043.
71. Miller KM, Rog O and Cooper JP. 2006. Semi-conservative DNA replication through telomeres requires Taz1. Nature 440:824-828.
72. Mitton-Fry RM, Anderson EM, Hughes TR, Lundblad V and Wuttke DS. 2002. Conserved structure for single-stranded telo-meric DNA recognition. Science 296:145-147.
73. Miyake Y, Nakamura M, Nabetani A, Shimamura S, Tamura M, Yonehara S, Saito M and Ishikawa F. 2009. RPA-like mammalian CTC1-Stn1-Ten1 complex binds to single-stranded DNA and protects telomeres independently of the POT1 pathway. Mol Cell (in press).
74. Miyoshi T, Kanoh J, Saito M and Ishikawa F. 2008. Fission yeast Pot1-Tpp1 protects telomeres and regulates telomere length. Science 320:1341-1344.
75. Moser BA, Subramanian L, Chang YT, Noguchi C, Noguchi E and Nakamura TM. 2009. Diferential arrival of leading and lagging strand DNA polymerases at fssion yeast telomeres. EMBO J 28:810-820.
76. Murzin AG. 1993. OB (oligonucleotide/oligosaccharide binding)-fold: common structural and functional solution for non-homologous sequences. Embo J 12:861-867.
77. Ogata K, Morikawa S, Nakamura H, Sekikawa A, Inoue T, Kanai H, Sarai A, Ishii S and Nishimura Y. 1994. Solution structure of a specifc DNA complex of the Myb DNA-binding domain with cooperative recognition helices. Cell 79:639-648.
78. Paeschke K, Juranek S, Simonsson T, Hempel A, Rhodes D and Lipps HJ. 2008. Telomerase recruitment by the telomere end binding protein-beta facilitates G-quadruplex DNA unfolding in ciliates. Nat Struct Mol Biol 15:598-604.
79. Palm W and de Lange T. 2008. How shelterin protects mammalian telomeres. Annu Rev Genet 42:301-334.
80. Pennock E, Buckley K and Lundblad V. 2001. Cdc13 delivers separate complexes to the telomere for end protection and replication. Cell 104:387-396.
81. Petreaca RC, Chiu HC, Eckelhoefer HA, Chuang C, Xu L and Nugent CI. 2006. Chromosome end protection plasticity revealed by Stn1p and Ten1p bypass of Cdc13p. Nat Cell Biol 8:748-755.
82. Poulet A, Buisson R, Faivre-Moskalenko C, Koelblen M, Amiard S, Montel F, Cuesta-Lopez S, Bornet O, Guerlesquin F, Godet T, Moukhtar J, Argoul F, Déclais AC, Lilley DM, Ip SC, West SC, Gilson E, Giraud-Panis MJ. 2009. TRF2 promotes, remodels and protects telomeric Holliday junctions. EMBO J 28:641-651.
83. Price CM. 2009. Antics at the telomere: uncoupled polymerases solve the end replication problem. Embo J 28:795-796.
84. Puglisi A, Bianchi A, Lemmens L, Damay P and Shore D. 2008. Distinct roles for yeast Stn1 in telomere capping and telomerase inhibition. Embo J 27:2328-3239.
85. Qi H and Zakian VA. 2000. Te Saccharomyces telomere-binding protein Cdc13p interacts with both the catalytic subunit of DNA polymerase alpha and the telomerase-associated est1 protein. Genes Dev 14:1777-1788.
86. Raices M, Verdun RE, Compton SA, Haggblom CI, Grifth JD, Dillin A and Karlseder J. 2008. C. elegans telomeres contain G-strand and C-strand overhangs that are bound by distinct proteins. Cell 132:745-757.
87. Rhodes D, Fairall L, Simonsson T, Court R and Chapman L. 2002. Telomere architecture. EMBO Rep 3:1139-1145.
88. Riha K, Heacock ML and Shippen DE. 2006. Te role of the nonho-mologous end-joining DNA double-strand break repair pathway in telomere biology. Annu Rev Genet 40:237-277.
89. Rossignol P, Collier S, Bush M, Shaw P and Doonan JH. 2007. Arabidopsis POT1A interacts with TERT-V(I8), an N-terminal splicing variant of telomerase. J Cell Sci 120:3678-3687.
90. Sabourin M, Tuzon CT and Zakian VA. 2007. Telomerase and Tel1p preferentially associate with short telomeres in S. cerevisiae. Mol Cell 27:550-561.
91. Schrumpfova PP, Kuchar M, Palecek J and Fajkus J. 2008. Mapping of interaction domains of putative telomere-binding proteins AtTRB1 and AtPOT1b from Arabidopsis thaliana. FEBS Lett 582:1400-1406.
92. Sfeir A, Kosiyatrakul ST, Hockemeyer D, MacRae SL, Karlseder J, Schildkraut CL and de Lange T. 2009. Mammalian telomeres resemble fragile sites and require TRF1 for efcient replication. Cell 138:90-103.
93. Shakirov EV, Surovtseva YV, Osbun N and Shippen DE. 2005. Te Arabidopsis Pot1 and Pot2 proteins function in telomere length homeostasis and chromosome end protection. Mol Cell Biol 25:7725-7733.
94. Shakirov EV, McKnight TD and Shippen DE. 2009. POT1-independent single-strand telomeric DNA-binding activities in Brassicaceae. Plant J 58:1004-1015.
95. Skopp R, Wang W and Price C. 1996. rTP: a candidate telomere protein that is associated with DNA replication. Chromosoma 105:82-91.
96. Smogorzewska A and de Lange T. 2004. Regulation of telomerase by telomeric proteins. Annu Rev Biochem 73:177-208.
97. Song X, Leehy K, Warrington RT, Lamb JC, Surovtseva YV and Shippen DE. 2008. STN1 protects chromosome ends in Arabidopsis thaliana. Proc Natl Acad Sci USA 105:19815-19820.
98. Spink KG, Evans RJ and Chambers A. 2000. Sequence-specifc binding of Taz1p dimers to fssion yeast telomeric DNA. Nucleic Acids Res 28:527-533.
99. Stansel RM, de Lange T and Grifth JD. 2001. T-loop assembly in vitro involves binding of TRF2 near the 3′ telomeric overhang. Embo J 20:5532-5540.
100. Sugiyama T, Cam HP, Sugiyama R, Noma K, Zofall M, Kobayashi R and Grewal SI. 2007. SHREC, an efector complex for hetero-chromatic transcriptional silencing. Cell 128:491-504.
101. Surovtseva YV, Shakirov EV, Vespa L, Osbun N, Song X and Shippen DE. 2007. Arabidopsis POT1 associates with the telom-erase RNP and is required for telomere maintenance. Embo J 26:3653-3661.
102. Surovtseva Y, Churikov D, Bolts K, Song X, Lamb J, Warrington R, Leehy K, Heacock M, Price C and Shippen D. 2009. Conserved Telomere maintenance Component 1 interacts with STN1 and maintains chromosome ends in higher eukaryotes. Mol Cell (in press).
103. Teixeira MT, Arneric M, Sperisen P and Lingner J. 2004. Telomere length homeostasis is achieved via a switch between telomerase-extendible and-nonextendible states. Cell 117:323-335.
104. Teobald DL and Wuttke DS. 2004. Prediction of multiple tandem OB-fold domains in telomere end-binding proteins Pot1 and Cdc13. Structure (Camb) 12:1877-1879.
105. Teobald DL, Mitton-Fry RM and Wuttke DS. 2003. Nucleic acid recognition by OB-fold proteins. Annu Rev Biophys Biomol Struct 32:115-133.
106. Tomita K and Cooper JP. 2008. Fission yeast Ccq1 is telomer-ase recruiter and local checkpoint controller. Genes Dev 22:3461-3474.
107. Tseng SF, Lin JJ and Teng SC. 2006. Te telomerase-recruitment domain of the telomere binding protein Cdc13 is regulated by Mec1p/Tel1p-dependent phosphorylation. Nucleic Acids Res 34:6327-6336.
108. Verdun RE and Karlseder J. 2007. Replication and protection of tel-omeres. Nature 447:924-931.
109. Viscardi V, Bonetti D, Cartagena-Lirola H, Lucchini G and Longhese MP. 2007. MRX-dependent DNA damage response to short telomeres. Mol Biol Cell 18:3047-3058.
110. Wang F, Podell ER, Zaug AJ, Yang Y, Baciu P, Cech TR and Lei M. 2007. Te POT1-TPP1 telomere complex is a telomerase processivity factor. Nature 445:506-510.
111. Wang W, Skopp R, Scofeld M and Price C. 1992. Euplotes crassus has genes encoding telomere-binding proteins and telomere-binding protein homologs. Nucleic Acids Res 20:6621-6629.
112. Wu Y, Xiao S and Zhu XD. 2007. MRE11-RAD50-NBS1 and ATM function as co-mediators of TRF1 in telomere length control. Nat Struct Mol Biol 14:832-840.
113. Xin H, Liu D, Wan M, Safari A, Kim H, Sun W, O'Connor MS and Songyang Z. 2007. TPP1 is a homologue of ciliate TEBP-beta and interacts with POT1 to recruit telomerase. Nature 445:559-562.
114. Ye JZ, Donigian JR, van Overbeek M, Loayza D, Luo Y, Krutchinsky AN, Chait BT and de Lange T. 2004a. TIN2 Binds TRF1 and TRF2 Simultaneously and Stabilizes the TRF2 Complex on Telomeres. J Biol Chem 279:47264-47271.
115. Ye JZ, Hockemeyer D, Krutchinsky AN, Loayza D, Hooper SM, Chait BT and de Lange T. 2004b. POT1-interacting protein PIP1: a telomere length regulator that recruits POT1 to the TIN2/TRF1 complex. Genes Dev 18:1649-1654.
116. Yu EY, Wang F, Lei M and Lue NF. 2008. A proposed OB-fold with a protein-interaction surface in Candida albicans telomerase protein Est3. Nat Struct Mol Biol 15:985-989.