Telomere- and telomerase-associated proteins and their functions in the plant cell

Frontiers in Plant Science

Volumn 7, Issue , 2016, Pages

  Schrumpfová, Petra Procházková ^a   Schořová, Šárka ^a   Fajkuš, Jirí ^a,b  

^a MASARYK UNIVERSITY   (Czech Republic)

^b INSTITUTE OF BIOPHYSICS   (Czech Republic)

Author keywords

Plant; Shelterin; Telomerase; Telomere; Telomeric proteins; Telomeric repeat binding (TRB)

Indexed keywords

EID: 84976493124     PISSN: None     EISSN: 1664462X     Source Type: Journal    
DOI: 10.3389/fpls.2016.00851     Document Type: Review

References (228)

1. Adams, S. P., Hartman, T. P., Lim, K. Y., Chase, M. W., Bennett, M. D., Leitch, I. J., et al. (2001). Loss and recovery of Arabidopsis-type telomere repeat sequences 50-(TTTAGGG)(n)-30 in the evolution of a major radiation of flowering plants. Proc. Biol. Sci. 268, 1541-1546. doi: 10. 1098/rspb. 2001. 1726
2. Amiard, S., Depeiges, A., Allain, E., White, C. I., and Gallego, M. E. (2011). Arabidopsis ATM and ATR kinases prevent propagation of genome damage caused by telomere dysfunction. Plant Cell 23, 4254-4265. doi: 10. 1105/tpc. 111. 092387
3. Amiard, S., Olivier, M., Allain, E., Choi, K., Smith-Unna, R., Henderson, I. R., et al. (2014). Telomere stability and development of ctc1 mutants are rescued by inhibition of EJ recombination pathways in a telomerasedependent manner. Nucleic Acids Res. 42, 11979-11991. doi: 10. 1093/nar/ gku897
4. Arat, N. O., and Griffith, J. D. (2012). Human Rap1 interacts directly with telomeric DNA and regulates TRF2 localization at the telomere. J. Biol. Chem. 287, 41583-41594. doi: 10. 1074/jbc. M112. 415984
5. Armstrong, S. J., Franklin, F. C., and Jones, G. H. (2001). Nucleolus-associated telomere clustering and pairing precede meiotic chromosome synapsis in Arabidopsis thaliana. J. Cell Sci. 114, 4207-4217.
6. Aronen, T., and Ryynanen, L. (2014). Silver birch telomeres shorten in tissue culture. Tree Genet. Genomes 10, 67-74. doi: 10. 1007/s11295-013-0662-4
7. Bass, H. W., Marshall, W. F., Sedat, J. W., Agard, D. A., and Cande, W. Z. (1997). Telomeres cluster de novo before the initiation of synapsis: a three-dimensional spatial analysis of telomere positions before and during meiotic prophase. J. Cell Biol. 137, 5-18. doi: 10. 1083/jcb. 137. 1. 5
8. Baumann, P., and Cech, T. R. (2001). Pot1, the putative telomere endbinding protein in fission yeast and humans. Science 292, 1171-1175. doi: 10. 1126/science. 1060036
9. Beilstein, M. A., Brinegar, A. E., and Shippen, D. E. (2012). Evolution of the Arabidopsis telomerase RNA. Front. Genet. 3: 188. doi: 10. 3389/fgene. 2012. 00188
10. Beilstein, M. A., Renfrew, K. B., Song, X., Shakirov, E. V., Zanis, M. J., and Shippen, D. E. (2015). Evolution of the telomere-associated protein POT1a in Arabidopsis thaliana is characterized by positive selection to reinforce proteinprotein interaction. Mol. Biol. Evol. 32, 1329-1341. doi: 10. 1093/molbev/ msv025
11. Beneke, S., Cohausz, O., Malanga, M., Boukamp, P., Althaus, F., and Burkle, A. (2008). Rapid regulation of telomere length is mediated by poly(ADPribose) polymerase-1. Nucleic Acids Res. 36, 6309-6317. doi: 10. 1093/nar/ gkn615
12. Bianchi, A., and Shore, D. (2008). How telomerase reaches its end: mechanism of telomerase regulation by the telomeric complex. Mol. Cell. 31, 153-165. doi: 10. 1016/j. molcel. 2008. 06. 013
13. Bilaud, T., Koering, C. E., Binet-Brasselet, E., Ancelin, K., Pollice, A., Gasser, S. M., et al. (1996). The telobox, a Myb-related telomeric DNA binding motif found in proteins from yeast, plants and human. Nucleic Acids Res. 24, 1294-1303. doi: 10. 1093/nar/24. 7. 1294
14. Blackburn, E. H., and Gall, J. G. (1978). A tandemly repeated sequence at the termini of the extrachromosomal ribosomal RNA genes in Tetrahymena. J. Mol. Biol. 120, 33-53.
15. Boltz, K. A., Jasti, M., Townley, J. M., and Shippen, D. E. (2014). Analysis of poly(ADP-Ribose) polymerases in Arabidopsis telomere biology. PLoS ONE 9: e88872. doi: 10. 1371/journal. pone. 0088872
16. Boltz, K. A., Leehy, K., Song, X., Nelson, A. D., and Shippen, D. E. (2012). ATR cooperates with CTC1 and STN1 to maintain telomeres and genome integrity in Arabidopsis. Mol. Biol. Cell 23, 1558-1568. doi: 10. 1091/mbc. E11-12-1002
17. 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. doi: 10. 1038/ng1097-231
18. Bundock, P., and Hooykaas, P. (2002). Severe developmental defects, hypersensitivity to DNA-damaging agents, and lengthened telomeres in Arabidopsis MRE11 mutants. Plant Cell 14, 2451-2462. doi: 10. 1105/tpc. 005959
19. Bundock, P., van Attikum, H., and Hooykaas, P. (2002). Increased telomere length and hypersensitivity to DNA damaging agents in an Arabidopsis KU70 mutant. Nucleic Acids Res. 30, 3395-3400. doi: 10. 1093/nar/gkf445
20. Byun, M. Y., Hong, J. P., and Kim, W. T. (2008). Identification and characterization of three telomere repeat-binding factors in rice. Biochem. Biophys. Res. Commun. 372, 85-90. doi: 10. 1016/j. bbrc. 2008. 04. 181
21. Celli, G. B., and de Lange, T. (2005). DNA processing is not required for ATMmediated telomere damage response after TRF2 deletion. Nat. Cell Biol. 7, 712-718. doi: 10. 1038/ncb1275
22. Cesare, A. J., Quinney, N., Willcox, S., Subramanian, D., and Griffith, J. D. (2003). Telomere looping in P. sativum (common garden pea). Plant J. 36, 271-279. doi: 10. 1046/j. 1365-313X. 2003. 01882. x
23. Chai, W., Ford, L. P., Lenertz, L., Wright, W. E., and Shay, J. W. (2002). Human Ku70/80 associates physically with telomerase through interaction with hTERT. J. Biol. Chem. 277, 47242-47247. doi: 10. 1074/jbc. M208 542200
24. Charbonnel, C., Allain, E., Gallego, M. E., and White, C. I. (2011). Kinetic analysis of DNA double-strand break repair pathways in Arabidopsis. DNA Repair (Amst.) 10, 611-619. doi: 10. 1016/j. dnarep. 2011. 04. 002
25. Charbonnel, C., Gallego, M. E., and White, C. I. (2010). Xrcc1-dependent and Kudependent DNA double-strand break repair kinetics in Arabidopsis plants. Plant J. 64, 280-290. doi: 10. 1111/j. 1365-313X. 2010. 04331. x
26. Chen, C. M., Wang, C. T., and Ho, C. H. (2001). A plant gene encoding a Myblike protein that binds telomeric GGTTTAG repeats in vitro. J. Biol. Chem. 276, 16511-16519. doi: 10. 1074/jbc. M009659200
27. Chen, C. M., Wang, C. T., Kao, Y. H., Chang, G. D., Ho, C. H., Lee, F. M., et al. (2005). Functional redundancy of the duplex telomeric DNA-binding proteins in Arabidopsis. Bot. Bull. Acad. Sin. 46, 315-324.
28. Chen, L. Y., and Lingner, J. (2013). CST for the grand finale of telomere replication. Nucleus 4, 277-282. doi: 10. 4161/nucl. 25701
29. Chen, L. Y., Redon, S., and Lingner, J. (2012). The human CST complex is a terminator of telomerase activity. Nature 488, 540-544. doi: 10. 1038/nature11269
30. Chikashige, Y., Tsutsumi, C., Yamane, M., Okamasa, K., Haraguchi, T., and Hiraoka, Y. (2006). Meiotic proteins bqt1 and bqt2 tether telomeres to form the bouquet arrangement of chromosomes. Cell 125, 59-69. doi: 10. 1016/j. cell. 2006. 01. 048
31. Cifuentes-Rojas, C., Nelson, A. D., Boltz, K. A., Kannan, K., She, X., and Shippen, D. E. (2012). An alternative telomerase RNA in Arabidopsis modulates enzyme activity in response to DNA damage. Genes Dev. 26, 2512-2523. doi: 10. 1101/gad. 202960. 112
32. Cokus, S. J., Feng, S., Zhang, X., Chen, Z., Merriman, B., Haudenschild, C. D., et al. (2008). Shotgun bisulphite sequencing of the Arabidopsis genome reveals DNA methylation patterning. Nature 452, 215-219. doi: 10. 1038/nature06745
33. Comadira, G., Rasool, B., Kaprinska, B., Garcia, B. M., Morris, J., Verrall, S. R., et al. (2015). WHIRLY1 functions in the control of responses to nitrogen deficiency but not aphid infestation in barley. Plant Physiol. 168, 1140-1151. doi: 10. 1104/pp. 15. 00580
34. Cook, B. D., Dynek, J. N., Chang, W., Shostak, G., and Smith, S. (2002). Role for the related poly(ADP-Ribose) polymerases tankyrase 1 and 2 at human telomeres. Mol. Cell. Biol. 22, 332-342. doi: 10. 1128/MCB. 22. 1. 332-342. 2002
35. Corredor, E., and Naranjo, T. (2007). Effect of colchicine and telocentric chromosome conformation on centromere and telomere dynamics at meiotic prophase I in wheat-rye additions. Chromosome Res. 15, 231-245. doi: 10. 1007/s10577-006-1117-7
36. 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. doi: 10. 1038/sj. embor. 7400314
37. Cowan, C. R., Carlton, P. M., and Cande, W. Z. (2001). The polar arrangement of telomeres in interphase and meiosis. Rabl organization and the bouquet. Plant Physiol. 125, 532-538.
38. Cowan, C. R., Carlton, P. M., and Cande, W. Z. (2002). Reorganization and polarization of the meiotic bouquet-stage cell can be uncoupled from telomere clustering. J. Cell Sci. 115, 3757-3766. doi: 10. 1242/jcs. 00054
39. da Costa e Silva, O., Klein, L., Schmelzer, E., Trezzini, G. F., and Hahlbrock, K. (1993). BPF-1, a pathogen-induced DNA-binding protein involved in the plant defense response. Plant J. 4, 125-135.
40. de Lange, T. (2005). Shelterin: the protein complex that shapes and safeguards human telomeres. Genes Dev. 19, 2100-2110. doi: 10. 1101/gad. 1346005
41. Decottignies, A. (2013). Alternative end-joining mechanisms: a historical perspective. Front. Genet. 4: 48. doi: 10. 3389/fgene. 2013. 00048
42. Denchi, E. L., and de Lange, T. (2007). Protection of telomeres through independent control of ATM and ATR by TRF2 and POT1. Nature 448, 1068-1071. doi: 10. 1038/nature06065
43. Deng, W., Buzas, D. M., Ying, H., Robertson, M., Taylor, J., Peacock, W. J., et al. (2013). Arabidopsis Polycomb Repressive Complex 2 binding sites contain putative GAGA factor binding motifs within coding regions of genes. BMC Genomics 14: 593. doi: 10. 1186/1471-2164-14-593
44. Derboven, E., Ekker, H., Kusenda, B., Bulankova, P., and Riha, K. (2014). Role of STN1 and DNA polymerase alpha in telomere stability and genome-wide replication in Arabidopsis. PLoS Genet. 10: e1004682. doi: 10. 1371/journal. pgen. 1004682
45. Desveaux, D., Allard, J., Brisson, N., and Sygusch, J. (2002). A new family of plant transcription factors displays a novel ssDNA-binding surface. Nat. Struct. Biol. 9, 512-517. doi: 10. 1038/nsb814
46. Desveaux, D., Despres, C., Joyeux, A., Subramaniam, R., and Brisson, N. (2000). PBF-2 is a novel single-stranded DNA binding factor implicated in PR-10a gene activation in potato. Plant Cell 12, 1477-1489. doi: 10. 1105/tpc. 12. 8. 1477
47. Dokládal, L., Honys, D., Rana, R., Lee, L.-Y., Gelvin, S., and Sýkorová, E. (2015). cDNA library screening identifies protein interactors potentially involved in non-telomeric roles of Arabidopsis telomerase. Front. Plant Sci. 6: 985. doi: 10. 3389/fpls. 2015. 00985
48. Dong, F., and Jiang, J. (1998). Non-Rabl patterns of centromere and telomere distribution in the interphase nuclei of plant cells. Chromosome Res. 6, 551-558. doi: 10. 1023/A: 1009280425125
49. Du, H., Wang, Y. B., Xie, Y., Liang, Z., Jiang, S. J., Zhang, S. S., et al. (2013). Genome-wide identification and evolutionary and expression analyses of MYBrelated genes in land plants. DNA Res. 20, 437-448. doi: 10. 1093/dnares/dst021
50. Dvořáčková, M., Fojtova, M., and Fajkus, J. (2015). Chromatin dynamics of plant telomeres and ribosomal genes. Plant J. 83, 18-37. doi: 10. 1111/tpj. 12822
51. Dvorácková, M., Rossignol, P., Shaw, P. J., Koroleva, O. A., Doonan, J. H., and Fajkus, J. (2010). AtTRB1, a telomeric DNA-binding protein from Arabidopsis, is concentrated in the nucleolus and shows highly dynamic association with chromatin. Plant J. 61, 637-649. doi: 10. 1111/j. 1365-313X. 2009. 04094. x
52. Fajkus, J., Fulneckova, J., Hulanova, M., Berkova, K., Riha, K., and Matyasek, R. (1998). Plant cells express telomerase activity upon transfer to callus culture, without extensively changing telomere lengths. Mol. Gen. Genet. 260, 470-474. doi: 10. 1007/s004380050918
53. Fajkus, J., Kovarik, A., and Kralovics, R. (1996). Telomerase activity in plant cells. FEBS Lett. 391, 307-309. doi: 10. 1016/0014-5793(96)00757-0
54. Fajkus, J., Kovarik, A., Kralovics, R., and Bezdek, M. (1995). Organization of telomeric and subtelomeric chromatin in the higher plant Nicotiana tabacum. Mol. Gen. Genet. 247, 633-638. doi: 10. 1007/BF00290355
55. Fajkus, J., Sykorova, E., and Leitch, A. R. (2005). Telomeres in evolution and evolution of telomeres. Chromosome Res. 13, 469-479. doi: 10. 1007/s10577-005-0997-2
56. Fajkus, J., and Trifonov, E. N. (2001). Columnar packing of telomeric nucleosomes. Biochem. Biophys. Res. Commun. 280, 961-963. doi: 10. 1006/bbrc. 2000. 4208
57. Fajkus, P., Peska, V., Sitova, Z., Fulnecková, J., Dvorackova, M., Gogela, R., et al. (2016). Allium telomeres unmasked: the unusual telomeric sequence (CTCGGTTATGGG)n is synthesized by telomerase. Plant J. 85, 337-347. doi: 10. 1111/tpj. 13115
58. Feldbrugge, M., Sprenger, M., Hahlbrock, K., and Weisshaar, B. (1997). PcMYB1, a novel plant protein containing a DNA-binding domain with one MYB repeat, interacts in vivo with a light-regulatory promoter unit. Plant J. 11, 1079-1093. doi: 10. 1046/j. 1365-313X. 1997. 11051079. x
59. Fell, V. L., and Schild-Poulter, C. (2015). The Ku heterodimer: function in DNA repair and beyond. Mutat. Res. Rev. Mutat. Res. 763, 15-29. doi: 10. 1016/j. mrrev. 2014. 06. 002
60. Fojtova, M., Peska, V., Dobsakova, Z., Mozgova, I., Fajkus, J., and Sykorova, E. (2011). Molecular analysis of T-DNA insertion mutants identified putative regulatory elements in the AtTERT gene. J. Exp. Bot. 62, 5531-5545. doi: 10. 1093/jxb/err235
61. Fojtova, M., Sykorova, E., Najdekrova, L., Polanska, P., Zachova, D., Vagnerova, R., et al. (2015). Telomere dynamics in the lower plant Physcomitrella patens. Plant Mol. Biol. 87, 591-601. doi: 10. 1007/s11103-015-0299-9
62. Foyer, C. H., Karpinska, B., and Krupinska, K. (2014). The functions of WHIRLY1 and REDOX-RESPONSIVE TRANSCRIPTION FACTOR 1 in cross tolerance responses in plants: a hypothesis. Philos. Trans. R. Soc. Lond. B Biol. Sci. 369: 20130226. doi: 10. 1098/rstb. 2013. 0226
63. Freeling, M. (2009). Bias in plant gene content following different sorts of duplication: tandem, whole-genome, segmental, or by transposition. Annu. Rev. Plant Biol. 60, 433-453. doi: 10. 1146/annurev. arplant. 043008. 092122
64. Fu, D., and Collins, K. (2007). Purification of human telomerase complexes identifies factors involved in telomerase biogenesis and telomere length regulation. Mol. Cell 28, 773-785. doi: 10. 1016/j. molcel. 2007. 09. 023
65. Fulcher, N., and Riha, K. (2016). Using centromere mediated genome elimination to elucidate the funcional redudancy of candidate telomere binding proteins in Arabidopsis thaliana. Front. Genet. 6: 349. doi: 10. 3389/fgene. 2015. 00349
66. Fulneckova, J., and Fajkus, J. (2000). Inhibition of plant telomerase by telomerebinding proteins from nuclei of telomerase-negative tissues. FEBS Lett. 467, 305-310. doi: 10. 1016/S0014-5793(00)01178-9
67. Fulnecková, J., Sevcíková, T., Fajkus, J., Lukesová, A., Lukes, M., Vlcek, C., et al. (2013). A broad phylogenetic survey unveils the diversity and evolution of telomeres in eukaryotes. Genome Biol. Evol. 5, 468-483. doi: 10. 1093/gbe/evt019
68. Fulnečková, J., Ševčikova, T., Lukešová, A., and Sýkorová, E. (2015). Transtitions between the Arabidopsis-type and the human-type telomere sequence in green algae (clade Caudivolvoxa, Chlamydomonadales). Chromosoma. doi: 10. 1007/s00412-015-0557-2 [Epub ahead of print].
69. Gallego, M. E., Jalut, N., and White, C. I. (2003). Telomerase dependence of telomere lengthening in Ku80 mutant Arabidopsis. Plant Cell 15, 782-789. doi: 10. 1105/tpc. 008623
70. Gallego, M. E., and White, C. I. (2001). RAD50 function is essential for telomere maintenance in Arabidopsis. Proc. Natl. Acad. Sci. U. S. A. 98, 1711-1716. doi: 10. 1073/pnas. 98. 4. 1711
71. Garvik, B., Carson, M., and Hartwell, L. (1995). Single-stranded DNA arising at telomeres in cdc13 mutants may constitute a specific signal for the RAD9 checkpoint. Mol. Cell. Biol. 15, 6128-6613. doi: 10. 1128/MCB. 15. 11. 6128
72. Giraud-Panis, M. J., Pisano, S., Benarroch-Popivker, D., Pei, B., Le, Du, M. H., et al. (2013). One identity or more for telomeres? Front. Oncol. 3: 48. doi: 10. 3389/fonc. 2013. 00048
73. Giraud-Panis, M. J., Teixeira, M. T., Geli, V., and Gilson, E. (2010). CST meets shelterin to keep telomeres in check. Mol. Cell. 39, 665-676. doi: 10. 1016/j. molcel. 2010. 08. 024
74. Grabowski, E., Miao, Y., Mulisch, M., and Krupinska, K. (2008). Singlestranded DNA-binding protein Whirly1 in barley leaves is located in plastids and the nucleus of the same cell. Plant Physiol. 147, 1800-1804. doi: 10. 1104/pp. 108. 122796
75. Grandin, N., and Charbonneau, M. (2001). Hsp90 levels affect telomere length in yeast. Mol. Genet. Genom. 265, 126-134. doi: 10. 1007/s004380000398
76. Graumann, K., Runions, J., and Evans, D. E. (2010). Characterization of SUNdomain proteins at the higher plant nuclear envelope. Plant J. 61, 134-144. doi: 10. 1111/j. 1365-313X. 2009. 04038. x
77. Grossi, S., Puglisi, A., Dmitriev, P. V., 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. doi: 10. 1101/gad. 300004
78. Guo, X., Deng, Y., Lin, Y., Cosme-Blanco, W., Chan, S., He, H., et al. (2007). Dysfunctional telomeres activate an ATM-ATR-dependent DNA damage response to suppress tumorigenesis. EMBO J. 26, 4709-4719.
79. Harrington, L., McPhail, T., Mar, V., Zhou, W., Oulton, R., Bass, M. B., et al. (1997). A mammalian telomerase-associated protein. Science 275, 973-977. doi: 10. 1126/science. 275. 5302. 973
80. He, H., Multani, A. S., Cosme-Blanco, W., Tahara, H., Ma, J., Pathak, S., et al. (2006). POT1b protects telomeres from end-to-end chromosomal fusions and aberrant homologous recombination. EMBO J. 25, 5180-5190. doi: 10. 1038/sj. emboj. 7601294
81. He, Q., Chen, L., Xu, Y., and Yu, W. (2013). Identification of centromeric and telomeric DNA-binding proteins in rice. Proteomics 13, 826-832. doi: 10. 1002/pmic. 201100416
82. He, X. L., and Zhang, J. Z. (2005). Gene complexity and gene duplicability. Curr. Biol. 15, 1016-1021. doi: 10. 1016/j. cub. 2005. 04. 035
83. Heller, K., Kilian, A., Piatyszek, M. A., and Kleinhofs, A. (1996). Telomerase activity in plant extracts. Mol. Gen. Genet. 252, 342-345. doi: 10. 1007/BF02 173780
84. Higgins, J. D., Perry, R. M., Barakate, A., Ramsay, L., Waugh, R., Halpin, C., et al. (2012). Spatiotemporal asymmetry of the meiotic program underlies the predominantly distal distribution of meiotic crossovers in barley. Plant Cell 24, 4096-4109. doi: 10. 1105/tpc. 112. 102483
85. Hirata, Y., Suzuki, C., and Sakai, S. (2004). Characterization and gene cloning of telomere-binding protein from tobacco BY-2 cells. Plant Physiol. Biochem. 42, 7-14. doi: 10. 1016/j. plaphy. 2003. 10. 002
86. Hofr, C., Sultesova, P., Zimmermann, M., Mozgova, I., Prochazkova Schrumpfova, P., Wimmerova, M., et al. (2009). Single-Myb-histone proteins from Arabidopsis thaliana: a quantitative study of telomere-binding specificity and kinetics. Biochem. J. 419, 221-228. doi: 10. 1042/BJ20082195, 222 p following 228.
87. Holt, S. E., Aisner, D. L., Baur, J., Tesmer, V. M., Dy, M., Ouellette, M., et al. (1999). Functional requirement of p23 and Hsp90 in telomerase complexes. Genes Dev. 13, 817-826. doi: 10. 1101/gad. 13. 7. 817
88. Hong, J. P., Byun, M. Y., An, K., Yang, S. J., An, G., and Kim, W. T. (2010). OsKu70 is associated with developmental growth and genome stability in rice. Plant Physiol. 152, 374-387. doi: 10. 1104/pp. 109. 150391
89. Hong, J. P., Byun, M. Y., Koo, D. H., An, K., Bang, J. W., Chung, I. K., et al. (2007). Suppression of RICE TELOMERE BINDING PROTEIN 1 results in severe and gradual developmental defects accompanied by genome instability in rice. Plant Cell 19, 1770-1781. doi: 10. 1105/tpc. 107. 051953
90. Hrdlickova, R., Nehyba, J., Liss, A. S., and Bose, H. R. Jr. (2006). Mechanism of telomerase activation by v-Rel and its contribution to transformation. J. Virol. 80, 281-295. doi: 10. 1128/JVI. 80. 1. 281-295. 2006
91. Hwang, M. G., Chung, I. K., Kang, B. G., and Cho, M. H. (2001). Sequence-specific binding property of Arabidopsis thaliana telomeric DNA binding protein 1 (AtTBP1). FEBS Lett. 503, 35-40. doi: 10. 1016/S0014-5793(01)02685-0
92. Hwang, M. G., Kim, K., Lee, W. K., and Cho, M. H. (2005). AtTBP2 and AtTRP2 in Arabidopsis encode proteins that bind plant telomeric DNA and induce DNA bending in vitro. Mol. Gen. Genom. 273, 66-75. doi: 10. 1007/s00438-004-1096-3
93. Idziak, D., Robaszkiewicz, E., and Hasterok, R. (2015). Spatial distribution of centromeres and telomeres at interphase varies among Brachypodium species. J. Exp. Bot. 66, 6623-6634. doi: 10. 1093/jxb/erv369
94. Janouskova, E., Necasova, I., Pavlouskova, J., Zimmermann, M., Hluchy, M., Marini, V., et al. (2015). Human Rap1 modulates TRF2 attraction to telomeric DNA. Nucleic Acids Res. 43, 2691-2700. doi: 10. 1093/nar/gkv097
95. Kaminker, P. G., Kim, S. H., Desprez, P. Y., and Campisi, J. (2009). A novel form of the telomere-associated protein TIN2 localizes to the nuclear matrix. Cell Cycle 8, 931-939. doi: 10. 4161/cc. 8. 6. 7941
96. Kang, S. S., Kwon, T., Kwon, D. Y., and Do, S. I. (1999). Akt protein kinase enhances human telomerase activity through phosphorylation of telomerase reverse transcriptase subunit. J. Biol. Chem. 274, 13085-13090. doi: 10. 1074/jbc. 274. 19. 13085
97. Kannan, K., Nelson, A. D., and Shippen, D. E. (2008). Dyskerin is a component of the Arabidopsis telomerase RNP required for telomere maintenance. Mol. Cell. Biol. 28, 2332-2341. doi: 10. 1128/MCB. 01490-07
98. Karamysheva, Z. N., Surovtseva, Y. V., Vespa, L., Shakirov, E. V., and Shippen, D. E. (2004). A C-terminal Myb extension domain defines a novel family of double-strand telomeric DNA-binding proteins in Arabidopsis. J. Biol. Chem. 279, 47799-47807. doi: 10. 1074/jbc. M407938200
99. Kasbek, C., Wang, F., and Price, C. M. (2013). Human TEN1 maintains telomere integrity and functions in genome-wide replication restart. J. Biol. Chem. 288, 30139-30150. doi: 10. 1074/jbc. M113. 493478
100. Kazda, A., Zellinger, B., Rossler, M., Derboven, E., Kusenda, B., and Riha, K. (2012). Chromosome end protection by blunt-ended telomeres. Genes Dev. 26, 1703-1713. doi: 10. 1101/gad. 194944. 112
101. Kim, J. H., Kim, W. T., and Chung, I. K. (1998). Rice proteins that bind single-stranded G-rich telomere DNA. Plant Mol. Biol. 36, 661-672. doi: 10. 1023/A: 1005994719175
102. Kiss, T., Fayet-Lebaron, E., and Jady, B. E. (2010). Box H/ACA small ribonucleoproteins. Mol. Cell. 37, 597-606. doi: 10. 1016/j. molcel. 2010. 01. 032
103. Ko, S., Jun, S. H., Bae, H., Byun, J. S., Han, W., Park, H., et al. (2008). Structure of the DNA-binding domain of NgTRF1 reveals unique features of plant telomerebinding proteins. Nucleic Acids Res. 36, 2739-2755. doi: 10. 1093/nar/gkn030
104. Koonin, E. V. (2010). Preview. The incredible expanding ancestor of eukaryotes. Cell 140, 606-608. doi: 10. 1016/j. cell. 2010. 02. 022
105. Kovarik, A., Fajkus, J., Koukalova, B., and Bezdek, M. (1996). Species-specific evolution of telomeric and rDNA repeats in the tobacco composite genome. Theor. Appl. Genet. 92, 1108-1111. doi: 10. 1007/BF00224057
106. Krause, K., Kilbienski, I., Mulisch, M., Rodiger, A., Schafer, A., and Krupinska, K. (2005). DNA-binding proteins of the Whirly family in Arabidopsis thaliana are targeted to the organelles. FEBS Lett. 579, 3707-3712. doi: 10. 1016/j. febslet. 2005. 05. 059
107. Krutilina, R. I., Oei, S., Buchlow, G., Yau, P. M., Zalensky, A. O., Zalenskaya, I. A., et al. (2001). A negative regulator of telomere-length protein trf1 is associated with interstitial (TTAGGG)n blocks in immortal Chinese hamster ovary cells. Biochem. Biophys. Res. Commun. 280, 471-475. doi: 10. 1006/bbrc. 2000. 4143
108. Kuchar, M., and Fajkus, J. (2004). Interactions of putative telomere-binding proteins in Arabidopsis thaliana: identification of functional TRF2 homolog in plants. FEBS Lett. 578, 311-315. doi: 10. 1016/j. febslet. 2004. 11. 021
109. Kwon, C., and Chung, I. K. (2004). Interaction of an Arabidopsis RNA-binding protein with plant single-stranded telomeric DNA modulates telomerase activity. J. Biol. Chem. 279, 12812-12818. doi: 10. 1074/jbc. M312011200
110. Kwon, C., Kwon, K., Chung, I. K., Kim, S. Y., Cho, M. H., and Kang, B. G. (2004). Characterization of single stranded telomeric DNA-binding proteins in cultured soybean (Glycine max) cells. Mol. Cells 17, 503-508.
111. Lamarche, B. J., Orazio, N. I., and Weitzman, M. D. (2010). The MRN complex in double-strand break repair and telomere maintenance. FEBS Lett. 584, 3682-3695. doi: 10. 1016/j. febslet. 2010. 07. 029
112. Lazzerini-Denchi, E., and Sfeir, A., (2016). Stop pulling my strings-what telomeres taught us about the DNA damage response. Nat. Rev. Mol. Cell. Biol. 17, 364-378. doi: 10. 1038/nrm. 2016. 43
113. Lee, J. H., Kim, J. H., Kim, W. T., Kang, B. G., and Chung, I. K. (2000). Characterization and developmental expression of single-stranded telomeric DNA-binding proteins from mung bean (Vigna radiata). Plant Mol. Biol. 42, 547-557. doi: 10. 1023/A: 1006373917321
114. Lee, L. Y., Wu, F. H., Hsu, C. T., Shen, S. C., Yeh, H. Y., Liao, D. C., et al. (2012). Screening a cDNA library for protein-protein interactions directly in planta. Plant Cell 24, 1746-1759. doi: 10. 1105/tpc. 112. 097998
115. Lee, W. K., and Cho, M. H. (2016). Telomere-binding protein regulates the chromosome ends through the interaction with histone deacetylases in Arabidopsis thaliana. Nucl. Acids Res. 44, 4610-4624. doi: 10. 1093/nar/gkw067
116. Lee, W. K., Yun, J. H., Lee, W., and Cho, M. H. (2012). DNA-binding domain of AtTRB2 reveals unique features of a single Myb histone protein family that binds to both Arabidopsis-and human-type telomeric DNA sequences. Mol. Plant 5, 1406-1408. doi: 10. 1093/mp/sss063
117. Lee, Y. W., and Kim, W. T. (2010). Tobacco GTBP1, a homolog of human heterogeneous nuclear ribonucleoprotein, protects telomeres from aberrant homologous recombination. Plant Cell 22, 2781-2795. doi: 10. 1105/tpc. 110. 076778
118. Lee, Y. W., and Kim, W. T. (2013). Telomerase-dependent 30 G-strand overhang maintenance facilitates GTBP1-mediated telomere protection from misplaced homologous recombination. Plant Cell 25, 1329-1342. doi: 10. 1105/tpc. 112. 107573
119. Leehy, K. A., Lee, J. R., Song, X., Renfrew, K. B., and Shippen, D. E. (2013). MERISTEM DISORGANIZATION1 encodes TEN1, an essential telomere protein that modulates telomerase processivity in Arabidopsis. Plant Cell 25, 1343-1354. doi: 10. 1105/tpc. 112. 107425
120. Lei, M., Baumann, P., and Cech, T. R. (2002). Cooperative binding of single-stranded telomeric DNA by the Pot1 protein of Schizosaccharomyces pombe. Biochemistry 41, 14560-14568. doi: 10. 1021/ bi026674z
121. Lermontova, I., Schubert, V., Bornke, F., Macas, J., and Schubert, I. (2007). Arabidopsis CBF5 interacts with the H/ACA snoRNP assembly factor NAF1. Plant Mol. Biol. 65, 615-626. doi: 10. 1007/s11103-007-9226-z
122. Linger, B. R., and Price, C. M. (2009). Conservation of telomere protein complexes: shuffling through evolution. Crit. Rev. Biochem. Mol. Biol. 44, 434-446. doi: 10. 3109/10409230903307329
123. Lugert, T., and Werr, W. (1994). A novel DNA-binding domain in the Shrunken initiator-binding protein (IBP1). Plant Mol. Biol. 25, 493-506. doi: 10. 1007/BF00043877
124. Maï, M. E., Wagner, K. D., Michiels, J. F., Ambrosetti, D., Borderie, A., Destree, S., et al. (2014). The telomeric protein TRF2 regulates Angiogenesis by binding and activating the PDGFRβ promoter. Cell Rep. 9, 1047-1060. doi: 10. 1016/j. celrep. 2014. 09. 038
125. Maillet, G., White, C. I., and Gallego, M. E. (2006). Telomere-length regulation in inter-ecotype crosses of Arabidopsis. Plant Mol. Biol. 62, 859-866. doi: 10. 1007/s11103-006-9061-7
126. Majerova, E., Fojtova, M., Mozgova, I., Bittova, M., and Fajkus, J. (2011). Hypomethylating drugs efficiently decrease cytosine methylation in telomeric DNA and activate telomerase without affecting telomere lengths in tobacco cells. Plant Mol. Biol. 77, 371-380. doi: 10. 1007/s11103-011-9816-7
127. Mandakova, T., and Lysak, M. A. (2008). Chromosomal phylogeny and karyotype evolution in x = 7 crucifer species (Brassicaceae). Plant Cell 20, 2559-2570. doi: 10. 1105/tpc. 108. 062166
128. Marian, C. O., and Bass, H. W. (2005). The terminal acidic SANT 1 (Tacs1) gene of maize is expressed in tissues containing meristems and encodes an acidic SANT domain similar to some chromatin-remodeling complex proteins. Biochim. Biophys. Acta 1727, 81-86. doi: 10. 1016/j. bbaexp. 2004. 12. 010
129. Marian, C. O., Bordoli, S. J., Goltz, M., Santarella, R. A., Jackson, L. P., Danilevskaya, O., et al. (2003). The maize single myb histone 1 gene, Smh1, belongs to a novel gene family and encodes a protein that binds telomere DNA repeats in vitro. Plant Physiol. 133, 1336-1350.
130. Martinez, P., Thanasoula, M., Carlos, A. R., Gomez-Lopez, G., Tejera, A. M., Schoeftner, S., et al. (2010). Mammalian Rap1 controls telomere function and gene expression through binding to telomeric and extratelomeric sites. Nat. Cell Biol. 12, 768-780. doi: 10. 1038/ncb2081
131. Martinez-Perez, E., Shaw, P., Reader, S., Aragon-Alcaide, L., Miller, T., and Moore, G. (1999). Homologous chromosome pairing in wheat. J. Cell Sci. 112, 1761-1769.
132. Mignon-Ravix, C., Depetris, D., Delobel, B., Croquette, M. F., and Mattei, M. G. (2002). A human interstitial telomere associates in vivo with specific TRF2 and TIN2 proteins. Eur. J. Hum. Genet. 10, 107-112. doi: 10. 1038/sj. ejhg. 5200775
133. Mimori, T., Akizuki, M., Yamagata, H., Inada, S., Yoshida, S., and Homma, M. (1981). Characterization of a high molecular weight acidic nuclear protein recognized by autoantibodies in sera from patients with polymyositisscleroderma overlap. J. Clin. Invest. 68, 611-620. doi: 10. 1172/JCI110295
134. Moore, J. (2009). Investigating the DNA Binding Properties of the Initiator Binding Protein 2 (IBP2) in Maize (Zea Mays). Master thesis, Florida State University, Tallahassee.
135. Moriguchi, R., Kanahama, K., and Kanayama, Y. (2006). Characterization and expression analysis of the tomato telomere-binding protein LeTBP1. Plant Sci. 171, 166-174. doi: 10. 1016/j. plantsci. 2006. 03. 010
136. Morse, R. H. (2000). RAP, RAP, open up! New wrinkles for RAP1 in yeast. Trends Genet. 16, 51-53.
137. Mozgova, I., Schrumpfova, P. P., Hofr, C., and Fajkus, J. (2008). Functional characterization of domains in AtTRB1, a putative telomere-binding protein in Arabidopsis thaliana. Phytochemistry 69, 1814-1819. doi: 10. 1016/j. phytochem. 2008. 04. 001
138. Najdekrova, L., and Siroky, J. (2012). NBS1 plays a synergistic role with telomerase in the maintenance of telomeres in Arabidopsis thaliana. BMC Plant Biol. 12: 167. doi: 10. 1186/1471-2229-12-167
139. Nelson, A. D. J. (2012). Telomerase regulation in Arabidopsis thaliana. Ph. D. thesis, A&M University, Texas.
140. Nelson, A. D. J., Forsythe, E. S., Gan, X., Tsiantis, M., and Beilstein, M. A. (2014). Extending the model of Arabidopsis telomere length and composition across Brassicaceae. Chromosome Res. 22, 153-166. doi: 10. 1007/s10577-014-9423-y
141. Oguchi, K., Tamura, K., and Takahashi, H. (2004). Characterization of Oryza sativa telomerase reverse transcriptase and possible role of its phosphorylation in the control of telomerase activity. Gene 342, 57-66. doi: 10. 1016/j. gene. 2004. 07. 011
142. Olivier, M., Da Ines, O., Amiard, S., Serra, H., Goubely, C., White, C., et al. (2016). The structure-specific endonucleases MUS81 and SEND1 ere essential for telomere stability in Arabidopsis. Plant Cell 28, 74-86. doi: 10. 1105/tpc. 15. 00898
143. Ottaviani, A., Gilson, E., and Magdinier, F. (2008). Telomeric position effect: from the yeast paradigm to human pathologies? Biochimie 90, 93-107. doi: 10. 1016/j. biochi. 2007. 07. 022
144. Peska, V., Fajkus, P., Fojtova, M., Dvorackova, M., Hapala, J., Dvoracek, V., et al. (2015). Characterisation of an unusual telomere motif (TTTTTTAGGG)n in the plant Cestrum elegans (Solanaceae), a species with a large genome. Plant J. 82, 644-654. doi: 10. 1111/tpj. 12839
145. Peska, V., Schrumpfova, P. P., and Fajkus, J. (2011). Using the telobox to search for plant telomere binding proteins. Curr. Protein Pept. Sci. 12, 75-83. doi: 10. 2174/138920311795684968
146. Peska, V., Sykorova, E., and Fajkus, J. (2008). Two faces of Solanaceae telomeres: a comparison between Nicotiana and Cestrum telomeres and telomere-binding proteins. Cytogenet. Genome. Res. 122, 380-387. doi: 10. 1159/000167826
147. Phillips, D., Nibau, C., Wnetrzak, J., and Jenkins, G. (2012). High resolution analysis of meiotic chromosome structure and behaviour in barley (Hordeum vulgare L.). PLoS ONE 7: e39539. doi: 10. 1371/journal. pone. 0039539
148. Pisano, S., Galati, A., and Cacchione, S. (2008). Telomeric nucleosomes: forgotten players at chromosome ends. Cell Mol. Life. Sci. 65, 3553-3563. doi: 10. 1007/s00018-008-8307-8
149. Podlevsky, J. D., Bley, C. J., Omana, R. V., Qi, X., and Chen, J. J. (2008). The telomerase database. Nucleic Acids Res. 36, D339-D343. doi: 10. 1093/nar/gkm700
150. Price, C. M., Boltz, K. A., Chaiken, M. F., Stewart, J. A., Beilstein, M. A., and Shippen, D. E. (2010). Evolution of CST function in telomere maintenance. Cell Cycle 9, 3157-3165. doi: 10. 4161/cc. 9. 16. 12547
151. Price, C. M., and Cech, T. R. (1987). Telomeric DNA-protein interactions of Oxytricha macronuclear DNA. Genes Dev. 1, 783-793. doi: 10. 1101/gad. 1. 8. 783
152. Qi, H., and Zakian, V. A. (2000). The 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.
153. Rabl, C. (1885). Uber Zelltheilung. Morphologisches Jahrbuch 10, 214-330.
154. Recker, J., Knoll, A., and Puchta, H. (2014). The Arabidopsis thaliana homolog of the helicase RTEL1 plays multiple roles in preserving genome stability. Plant Cell 26, 4889-4902. doi: 10. 1105/tpc. 114. 132472
155. Ren, S., Johnston, J. S., Shippen, D. E., and McKnight, T. D. (2004). TELOMERASE ACTIVATOR1 induces telomerase activity and potentiates responses to auxin in Arabidopsis. Plant Cell 16, 2910-2922. doi: 10. 1105/tpc. 104. 025072
156. Ren, S., Mandadi, K. K., Boedeker, A. L., Rathore, K. S., and McKnight, T. D. (2007). Regulation of telomerase in Arabidopsis by BT2, an apparent target of TELOMERASE ACTIVATOR1. Plant Cell 19, 23-31. doi: 10. 1105/tpc. 106. 044321
157. Renfrew, K. B., Song, X., Lee, J. R., Arora, A., and Shippen, D. E. (2014). POT1a and components of CST engage telomerase and regulate its activity in Arabidopsis. PLoS Genet. 10: e1004738. doi: 10. 1371/journal. pgen. 1004738
158. Richards, E. J., and Ausubel, F. M. (1988). Isolation of a higher eukaryotic telomere from Arabidopsis thaliana. Cell 53, 127-136. doi: 10. 1016/0092-8674(88)90494-1
159. Riha, K., Fajkus, J., Siroky, J., and Vyskot, B. (1998). Developmental control of telomere lengths and telomerase activity in plants. Plant Cell 10, 1691-1698. doi: 10. 2307/3870766
160. Riha, K., McKnight, T. D., Fajkus, J., Vyskot, B., and Shippen, D. E. (2000). Analysis of the G-overhang structures on plant telomeres: evidence for two distinct telomere architectures. Plant J. 23, 633-641. doi: 10. 1046/j. 1365-313x. 2000. 00831. x
161. Riha, K., Watson, J. M., Parkey, J., and Shippen, D. E. (2002). Telomere length deregulation and enhanced sensitivity to genotoxic stress in Arabidopsis mutants deficient in Ku70. EMBO J. 21, 2819-2826. doi: 10. 1093/emboj/21. 11. 2819
162. Rossignol, P., Collier, S., Bush, M., Shaw, P., and Doonan, J. H. (2007). Arabidopsis POT1A interacts with TERT-V(I8), an N-terminal splicing variant of telomerase. J. Cell Sci. 120, 3678-3687. doi: 10. 1242/jcs. 004119
163. Rotkova, G., Sykorova, E., and Fajkus, J. (2009). Protect and regulate: Recent findings on plant POT1-like proteins. Biol. Plant. 53, 1-4. doi: 10. 1007/s10535-009-0001-7
164. Ruckova, E., Friml, J., Prochazkova Schrumpfova, P., and Fajkus, J. (2008). Role of alternative telomere lengthening unmasked in telomerase knock-out mutant plants. Plant Mol. Biol. 66, 637-646. doi: 10. 1007/s11103-008-9295-7
165. Schmidt, J. C., and Cech, T. R. (2015). Human telomerase: biogenesis, trafficking, recruitment, and activation. Genes Dev. 29, 1095-1105. doi: 10. 1101/gad. 263863. 115
166. Schober, H., Ferreira, H., Kalck, V., Gehlen, L. R., and Gasser, S. M. (2009). Yeast telomerase and the SUN domain protein Mps3 anchor telomeres and repress subtelomeric recombination. Genes Dev. 23, 928-938. doi: 10. 1101/gad. 1787509
167. Schrumpfova, P., Kuchar, M., Mikova, G., Skrisovska, L., Kubicarova, T., and Fajkus, J. (2004). Characterization of two Arabidopsis thaliana myb-like proteins showing affinity to telomeric DNA sequence. Genome 47, 316-324. doi: 10. 1139/g03-136
168. Schrumpfova, P. P., 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. doi: 10. 1016/j. febslet. 2008. 03. 034
169. Schrumpfova, P. P., Vychodilova, I., Dvorackova, M., Majerska, J., Dokladal, L., Schorova, S., et al. (2014). Telomere repeat binding proteins are functional components of Arabidopsis telomeres and interact with telomerase. Plant J. 77, 770-781. doi: 10. 1111/tpj. 12428
170. Schrumpfova, P. P., Vychodilova, I., Hapala, J., Schorova, S., Dvoracek, V., and Fajkus, J. (2016). Telomere binding protein TRB1 is associated with promoters of translation machinery genes in vivo. Plant. Mol. Biol. 90, 189-206. doi: 10. 1007/s11103-015-0409-8
171. Schwacke, R., Fischer, K., Ketelsen, B., Krupinska, K., and Krause, K. (2007). Comparative survey of plastid and mitochondrial targeting properties of transcription factors in Arabidopsis and rice. Mol. Genet. Genom. 277, 631-646. doi: 10. 1007/s00438-007-0214-4
172. Sfeir, A. (2012). Telomeres at a glance. J. Cell Sci. 125, 4173-4178. doi: 10. 1242/jcs. 106831
173. Shakirov, E. V., McKnight, T. D., and Shippen, D. E. (2009a). POT1-independent single-strand telomeric DNA binding activities in Brassicaceae. Plant J. 58, 1004-1015. doi: 10. 1111/j. 1365-313X. 2009. 03837. x
174. Shakirov, E. V., Perroud, P. F., Nelson, A. D., Cannell, M. E., Quatrano, R. S., and Shippen, D. E. (2010). Protection of telomeres 1 is required for telomere integrity in the moss Physcomitrella patens. Plant Cell 22, 1838-1848. doi: 10. 1105/tpc. 110. 075846
175. Shakirov, E. V., Song, X., Joseph, J. A., and Shippen, D. E. (2009b). POT1 proteins in green algae and land plants: DNA-binding properties and evidence of co-evolution with telomeric DNA. Nucleic Acids Res. 37, 7455-7467. doi: 10. 1093/nar/gkp785
176. Shakirov, E. V., and Shippen, D. E. (2004). Length regulation and dynamics of individual telomere tracts in wild-type Arabidopsis. Plant Cell 16, 1959-1967. doi: 10. 1105/tpc. 104. 023093
177. Shakirov, E. V., Surovtseva, Y. V., Osbun, N., and Shippen, D. E. (2005). The Arabidopsis Pot1 and Pot2 proteins function in telomere length homeostasis and chromosome end protection. Mol. Cell. Biol. 25, 7725-7733. doi: 10. 1128/MCB. 25. 17. 7725-7733. 2005
178. Simonet, T., Zaragosi, L. E., Philippe, C., Lebrigand, K., Schouteden, C., Augereau, A., et al. (2011). The human TTAGGG repeat factors 1 and 2 bind to a subset of interstitial telomeric sequences and satellite repeats. Cell Res. 21, 1028-1038. doi: 10. 1038/cr. 2011. 40
179. Smith, S., Giriat, I., Schmitt, A., and de Lange, T. (1998). Tankyrase, a poly(ADP-ribose) polymerase at human telomeres. Science 282, 1484-1487. doi: 10. 1126/science. 282. 5393. 1484
180. Smogorzewska, A., van Steensel, B., Bianchi, A., Oelmann, S., Schaefer, M. R., Schnapp, G., et al. (2000). Control of human telomere length by TRF1 and TRF2. Mol. Cell. Biol. 20, 1659-1668. doi: 10. 1128/MCB. 20. 5. 1659-1668. 2000
181. Song, X., Leehy, K., Warrington, R. T., Lamb, J. C., Surovtseva, Y. V., and Shippen, D. E. (2008). STN1 protects chromosome ends in Arabidopsis thaliana. Proc. Natl. Acad. Sci. U. S. A. 105, 19815-19820. doi: 10. 1073/pnas. 08078 67105
182. Starr, D. A., Hermann, G. J., Malone, C. J., Fixsen, W., Priess, J. R., Horvitz, H. R., et al. (2001). unc-83 encodes a novel component of the nuclear envelope and is essential for proper nuclear migration. Development 128, 5039-5050.
183. Stewart, J. A., Wang, F., Chaiken, M. F., Kasbek, C., Chastain, P. D. II, Wright, W. E., et al. (2012). Human CST promotes telomere duplex replication and general replication restart after fork stalling. EMBO J. 31, 3537-3549. doi: 10. 1038/emboj. 2012. 215
184. Surovtseva, Y. V., Churikov, D., Boltz, K. A., Song, X., Lamb, J. C., Warrington, R., et al. (2009). Conserved telomere maintenance component 1 interacts with STN1 and maintains chromosome ends in higher eukaryotes. Mol. Cell. 36, 207-218. doi: 10. 1016/j. molcel. 2009. 09. 017
185. Surovtseva, Y. V., Shakirov, E. V., Vespa, L., Osbun, N., Song, X., and Shippen, D. E. (2007). Arabidopsis POT1 associates with the telomerase RNP and is required for telomere maintenance. EMBO J. 26, 3653-3661. doi: 10. 1038/sj. emboj. 7601792
186. Sykorova, E., and Fajkus, J. (2009). Structure-function relationships in telomerase genes. Biol. Cell 101, 375-392. doi: 10. 1042/BC20080205.
187. Sykorova, E., Fulneckova, J., Mokros, P., Fajkus, J., Fojtova, M., and Peska, V. (2012). Three TERT genes in Nicotiana tabacum. Chromosome Res. 20, 381-394. doi: 10. 1007/s10577-012-9282-3
188. Sykorova, E., Leitch, A. R., and Fajkus, J. (2006). Asparagales telomerases which synthesize the human type of telomeres. Plant Mol. Biol. 60, 633-646. doi: 10. 1007/s11103-005-5091-9
189. Sykorova, E., Lim, K. Y., Kunicka, Z., Chase, M. W., Bennett, M. D., Fajkus, J., et al. (2003). Telomere variability in the monocotyledonous plant order Asparagales. Proc. Biol. Sci. 270, 1893-1904. doi: 10. 1098/rspb. 2003. 2446
190. Tamura, K., Goto, C., and Hara-Nishimura, I. (2015). Recent advances in understanding plant nuclear envelope proteins involved in nuclear morphology. J. Exp. Bot. 66, 1641-1647. doi: 10. 1093/jxb/erv036
191. Tamura, K., Liu, H., and Takahashi, H. (1999). Auxin induction of cell cycle regulated activity of tobacco telomerase. J. Biol. Chem. 274, 20997-21002. doi: 10. 1074/jbc. 274. 30. 20997
192. Tani, A., and Murata, M. (2005). Alternative splicing of Pot1 (Protection of telomere)-like genes in Arabidopsis thaliana. Genes Genet. Syst. 80, 41-48. doi: 10. 1266/ggs. 80. 41
193. Taylor, J. S., and Raes, J. (2004). Duplication and divergence: the evolution of new genes and old ideas. Annu. Rev. Genet. 38, 615-643. doi: 10. 1146/annurev. genet. 38. 072902. 092831
194. Tiang, C. L., He, Y., and Pawlowski, W. P. (2012). Chromosome organization and dynamics during interphase, mitosis, and meiosis in plants. Plant Physiol. 158, 26-34. doi: 10. 1104/pp. 111. 187161
195. Ting, N. S., Yu, Y., Pohorelic, B., Lees-Miller, S. P., and Beattie, T. L. (2005). Human Ku70/80 interacts directly with hTR, the RNA component of human telomerase. Nucleic Acids Res. 33, 2090-2098. doi: 10. 1093/nar/gki342
196. To, T. K., Kim, J. M., Matsui, A., Kurihara, Y., Morosawa, T., Ishida, J., et al. (2011). Arabidopsis HDA6 regulates locus-directed heterochromatin silencing in cooperation with MET1. PLoS Genet. 7: e1002055. doi: 10. 1371/journal. pgen. 1002055
197. Tomaska, L., Nosek, J., Kramara, J., and Griffith, J. D. (2009). Telomeric circles: universal players in telomere maintenance? Nat. Struct. Mol. Biol. 16, 1010-1015. doi: 10. 1038/nsmb. 1660
198. Tran, D. T., Cao, H. X., Jovtchev, G., Neumann, P., Novak, P., Fojtova, M., et al. (2015). Centromere and telomere sequence alterations reflect the rapid genome evolution within the carnivorous plant genus Genlisea. Plant J. 84, 1087-1099. doi: 10. 1111/tpj. 13058
199. Uringa, E. J., Youds, J. L., Lisaingo, K., Lansdorp, P. M., and Boulton, S. J. (2011). RTEL1: an essential helicase for telomere maintenance and the regulation of homologous recombination. Nucleic Acids Res. 39, 1647-1655. doi: 10. 1093/nar/gkq1045
200. van der Fits, L., Zhang, H., Menke, F. L., Deneka, M., and Memelink, J. (2000). A Catharanthus roseus BPF-1 homologue interacts with an elicitor-responsive region of the secondary metabolite biosynthetic gene Str and is induced by elicitor via a JA-independent signal transduction pathway. Plant Mol. Biol. 44, 675-685. doi: 10. 1023/A: 1026526522555
201. Vannier, J. B., Depeiges, A., White, C., and Gallego, M. E. (2009). ERCC1/XPF protects short telomeres from homologous recombination in Arabidopsis thaliana. PLoS Genet. 5: e1000380. doi: 10. 1371/journal. pgen. 1000380
202. Vannier, J. B., Pavicic-Kaltenbrunner, V., Petalcorin, M. I., Ding, H., and Boulton, S. J. (2012). RTEL1 dismantles T loops and counteracts telomeric G4-DNA to maintain telomere integrity. Cell 149, 795-806. doi: 10. 1016/j. cell. 2012. 03. 030
203. Venteicher, A. S., Meng, Z., Mason, P. J., Veenstra, T. D., and Artandi, S. E. (2008). Identification of ATPases pontin and reptin as telomerase components essential for holoenzyme assembly. Cell 132, 945-957. doi: 10. 1016/j. cell. 2008. 01. 019
204. Vespa, L., Warrington, R. T., Mokros, P., Siroky, J., and Shippen, D. E. (2007). ATM regulates the length of individual telomere tracts in Arabidopsis. Proc. Natl. Acad. Sci. U. S. A. 104, 18145-18150. doi: 10. 1073/pnas. 070 4466104
205. Vrbsky, J., Akimcheva, S., Watson, J. M., Turner, T. L., Daxinger, L., Vyskot, B., et al. (2010). siRNA-mediated methylation of Arabidopsis telomeres. PLoS Genet. 6: e1000986. doi: 10. 1371/journal. pgen. 1000986
206. Wang, H., Liu, C., Cheng, J., Liu, J., Zhang, L., He, C., et al. (2016). Arabidopsis flower and embryo developmental genes are repressed in seedlings by different combinations of polycomb group proteins in association with distinct sets of cis-regulatory elements. PLoS Genet. 12: e1005771. doi: 10. 1371/journal. pgen. 1005771
207. Wang, Y., Ghosh, G., and Hendrickson, E. A. (2009). Ku86 represses lethal telomere deletion events in human somatic cells. Proc. Natl. Acad. Sci. U. S. A. 106, 12430-12435. doi: 10. 1073/pnas. 0903362106
208. Watson, J. M., and Shippen, D. E. (2007). Telomere rapid deletion regulates telomere length in Arabidopsis thaliana. Mol. Cell. Biol. 27, 1706-1715. doi: 10. 1128/MCB. 02059-06
209. Wei, C., and Price, C. M. (2004). Cell cycle localization, dimerization, and binding domain architecture of the telomere protein cPot1. Mol. Cell. Biol. 24, 2091-2102. doi: 10. 1128/MCB. 24. 5. 2091-2102. 2004
210. Weiss, H., and Scherthan, H. (2002). Aloe spp.-plants with vertebratelike telomeric sequences. Chromosome Res. 10, 155-164. doi: 10. 1023/A: 1014905319557
211. Wellinger, R. J., and Zakian, V. A. (2012). Everything you ever wanted to know about Saccharomyces cerevisiae telomeres: beginning to end. Genetics 191, 1073-1105. doi: 10. 1534/genetics. 111. 137851
212. Wen, R., Moore, G., and Shaw, P. J. (2012). Centromeres cluster de novo at the beginning of meiosis in Brachypodium distachyon. PLoS ONE 7: e44681. doi: 10. 1371/journal. pone. 0044681
213. Wu, L., Multani, A. S., He, H., Cosme-Blanco, W., Deng, Y., Deng, J. M., et al. (2006). Pot1 deficiency initiates DNA damage checkpoint activation and aberrant homologous recombination at telomeres. Cell 126, 49-62. doi: 10. 1016/j. cell. 2006. 05. 037
214. Yang, D., Xiong, Y., Kim, H., He, Q., Li, Y., Chen, R., et al. (2011). Human telomeric proteins occupy selective interstitial sites. Cell Res. 21, 1013-1027. doi: 10. 1038/cr. 2011. 39
215. Yang, S. W., Jin, E., Chung, I. K., and Kim, W. T. (2002). Cell cycledependent regulation of telomerase activity by auxin, abscisic acid and protein phosphorylation in tobacco BY-2 suspension culture cells. Plant J. 29, 617-626. doi: 10. 1046/j. 0960-7412. 2001. 01244. x
216. Yang, S. W., Kim, D. H., Lee, J. J., Chun, Y. J., Lee, J. H., Kim, Y. J., et al. (2003). Expression of the telomeric repeat binding factor gene NgTRF1 is closely coordinated with the cell division program in tobacco BY-2 suspension culture cells. J. Biol. Chem. 278, 21395-21407. doi: 10. 1074/jbc. M209973200
217. Yang, S. W., Kim, S. K., and Kim, W. T. (2004). Perturbation of NgTRF1 expression induces apoptosis-like cell death in tobacco BY-2 cells and implicates NgTRF1 in the control of telomere length and stability. Plant Cell 16, 3370-3385. doi: 10. 1105/tpc. 104. 026278
218. Ye, J., Renault, V. M., Jamet, K., and Gilson, E. (2014). Transcriptional outcome of telomere signalling. Nat. Rev. Genet. 15, 491-503. doi: 10. 1038/nrg3743
219. Yoo, H. H., Kwon, C., Lee, M. M., and Chung, I. K. (2007). Single-stranded DNA binding factor AtWHY1 modulates telomere length homeostasis in Arabidopsis. Plant J. 49, 442-451. doi: 10. 1111/j. 1365-313X. 2006. 02974. x
220. Yu, E. Y., Kim, S. E., Kim, J. H., Ko, J. H., Cho, M. H., and Chung, I. K. (2000). Sequence-specific DNA recognition by the Myb-like domain of plant telomeric protein RTBP1. J. Biol. Chem. 275, 24208-24214. doi: 10. 1074/jbc. M003250200
221. Yun, J. H., Lee, W. K., Kim, H., Kim, E., Cheong, C., Cho, M. H., et al. (2014). Solution structure of telomere binding domain of AtTRB2 derived from Arabidopsis thaliana. Biochem. Biophys. Res. Commun. 452, 436-442. doi: 10. 1016/j. bbrc. 2014. 08. 095
222. Zachova, D., Fojtova, M., Dvorackova, M., Mozgova, I., Lermontova, I., Peska, V., et al. (2013). Structure-function relationships during transgenic telomerase expression in Arabidopsis. Physiol. Plant. 149, 114-126. doi: 10. 1111/ppl. 12021
223. Zellinger, B., Akimcheva, S., Puizina, J., Schirato, M., and Riha, K. (2007). Ku suppresses formation of telomeric circles and alternative telomere lengthening in Arabidopsis. Mol. Cell. 27, 163-169. doi: 10. 1016/j. molcel. 2007. 05. 025
224. Zentgraf, U. (1995). Telomere-binding proteins of Arabidopsis thaliana. Plant Mol. Biol. 27, 467-475. doi: 10. 1007/BF00019314
225. Zhang, P., Pazin, M. J., Schwartz, C. M., Becker, K. G., Wersto, R. P., Dilley, C. M., et al. (2008). Nontelomeric TRF2-REST interaction modulates neuronal gene silencing and fate of tumor and stem cells. Curr. Biol. 18, 1489-1494. doi: 10. 1016/j. cub. 2008. 08. 048
226. Zhou, X., Graumann, K., Wirthmueller, L., Jones, J. D., and Meier, I. (2014). Identification of unique SUN-interacting nuclear envelope proteins with diverse functions in plants. J. Cell Biol. 205, 677-692. doi: 10. 1083/jcb. 201401138
227. Zhou, Y., Hartwig, B., James, G. V., Schneeberger, K., and Turck, F. (2016). Complementary activities of TELOMERE REPEAT BINDING proteins and polycomb group complexes in transcriptional regulation of target genes. Plant Cell 28, 87-101. doi: 10. 1105/tpc. 15. 00787
228. Zhu, Q. H., Ramm, K., Shivakkumar, R., Dennis, E. S., and Upadhyaya, N. M. (2004). The ANTHER INDEHISCENCE1 gene encoding a single MYB domain protein is involved in anther development in rice. Plant Physiol. 135, 1514-1525. doi: 10. 1104/pp. 104. 041459