J. D. Griffith, L. Comeau, S. Rosenfield, R. M. Stansel, A. Bianchi et al., Mammalian telomeres end in a large duplex loop, Cell, vol.97, pp.503-514, 1999.

A. J. Cesare, N. Quinney, S. Willcox, D. Subramanian, and J. D. Griffith, Telomere looping in P. sativum (common garden pea), Plant J, vol.36, pp.271-279, 2003.

J. L. Munoz-jordan, G. A. Cross, T. De-lange, and J. D. Griffith, ) t-loops at trypanosome telomeres, EMBO J, vol.20, pp.579-588, 2001.

W. Palm and T. De-lange, How shelterin protects mammalian telomeres, Annu. Rev. Genet, vol.42, pp.301-334, 2008.

T. De-lange, How telomeres solve the end-protection problem, Science, vol.326, pp.948-952, 2009.

E. Gilson and V. Geli, How telomeres are replicated, Nat. Rev. Mol. Cell Biol, vol.8, pp.825-838, 2007.

Y. Doksani and T. De-lange, The role of double-strand break repair pathways at functional and dysfunctional telomeres, Cold Spring Harb. Perspect. Biol, vol.6, p.16576, 2014.

M. F. Arlt, B. Xu, S. G. Durkin, A. M. Casper, M. B. Kastan et al., BRCA1 is required for common-fragile-site stability via its G2/M checkpoint function, Mol. Cell. Biol, vol.24, pp.6701-6709, 2004.

K. M. Miller, O. Rog, and J. P. Cooper, Semi-conservative DNA replication through telomeres requires Taz1, Nature, vol.440, pp.824-828, 2006.

A. Sfeir, S. T. Kosiyatrakul, D. Hockemeyer, S. L. Macrae, J. Karlseder et al., Mammalian telomeres resemble fragile sites and require TRF1 for efficient replication, Cell, vol.138, pp.90-103, 2009.

G. Biffi, D. Tannahill, J. Mccafferty, and S. Balasubramanian, Quantitative visualization of DNA G-quadruplex structures in human cells, Nat. Chem, vol.5, pp.182-186, 2013.

M. Tarsounas and M. Tijsterman, Genomes and G-quadruplexes: for better or for worse, J. Mol. Biol, vol.425, pp.4782-4789, 2013.

H. J. Lipps and D. Rhodes, G-quadruplex structures: in vivo evidence and function, Trends Cell Biol, vol.19, pp.414-422, 2009.

J. B. Vannier, V. Pavicic-kaltenbrunner, M. I. Petalcorin, H. Ding, and S. J. Boulton, RTEL1 dismantles T loops and counteracts telomeric G4-DNA to maintain telomere integrity, Cell, vol.149, pp.795-806, 2012.

D. Rhodes and H. J. Lipps, G-quadruplexes and their regulatory roles in biology, Nucleic Acids Res, vol.43, pp.8627-8637, 2015.

M. D. Huber, M. L. Duquette, J. C. Shiels, and N. Maizels, A conserved G4 DNA binding domain in RecQ family helicases, J. Mol. Biol, vol.358, pp.1071-1080, 2006.

P. L. Opresko, W. H. Cheng, C. Von-kobbe, J. A. Harrigan, and V. A. Bohr, Werner syndrome and the function of the Werner protein; what they can teach us about the molecular aging process, Carcinogenesis, vol.24, pp.791-802, 2003.

C. Ribeyre, J. Lopes, J. B. Boule, A. Piazza, A. Guedin et al., The yeast Pif1 helicase prevents genomic instability caused by G-quadruplex-forming CEB1 sequences in vivo, PLoS Genet, vol.5, p.1000475, 2009.
URL : https://hal.archives-ouvertes.fr/hal-02109281

K. Paeschke, S. Juranek, T. Simonsson, A. Hempel, D. Rhodes et al., Telomerase recruitment by the telomere end binding protein-beta facilitates G-quadruplex DNA unfolding in ciliates, Nat. Struct. Mol. Biol, vol.15, pp.598-604, 2008.

A. L. Moye, K. C. Porter, S. B. Cohen, T. Phan, K. G. Zyner et al., Telomeric G-quadruplexes are a substrate and site of localization for human telomerase, Nat. Commun, vol.6, p.7643, 2015.

V. Pfeiffer, J. Crittin, L. Grolimund, and J. Lingner, The THO complex component Thp2 counteracts telomeric R-loops and telomere shortening, EMBO J, vol.32, pp.2861-2871, 2013.

C. M. Azzalin, P. Reichenbach, L. Khoriauli, E. Giulotto, and J. Lingner, Telomeric repeat containing RNA and RNA surveillance factors at mammalian chromosome ends, Science, vol.318, pp.798-801, 2007.

A. Maicher, L. Kastner, M. Dees, and B. Luke, Deregulated telomere transcription causes replication-dependent telomere shortening and promotes cellular senescence, Nucleic Acids Res, vol.40, pp.6649-6659, 2012.

S. M. Cerritelli and R. J. Crouch, Ribonuclease H: the enzymes in eukaryotes, FEBS J, vol.276, pp.1494-1505, 2009.

B. Balk, A. Maicher, M. Dees, J. Klermund, S. Luke-glaser et al., Telomeric RNA-DNA hybrids affect telomere-length dynamics and senescence, Nat. Struct. Mol. Biol, vol.20, pp.1199-1205, 2013.

R. Arora, Y. Lee, H. Wischnewski, C. M. Brun, T. Schwarz et al., RNaseH1 regulates TERRA-telomeric DNA hybrids and telomere maintenance in ALT tumour cells, Nat. Commun, vol.5, p.5220, 2014.

T. Y. Yu, Y. W. Kao, and J. J. Lin, Telomeric transcripts stimulate telomere recombination to suppress senescence in cells lacking telomerase, Proc. Natl. Acad. Sci. U.S.A, vol.111, pp.3377-3382, 2014.

R. Arora and C. M. Azzalin, Telomere elongation chooses TERRA ALTernatives, RNA Biol, vol.12, pp.938-941, 2015.

I. Draskovic and A. Vallejo, Telomere recombination and alternative telomere lengthening mechanisms, Front. Biosci, vol.18, pp.1-20, 2013.

A. J. Cesare and R. R. Reddel, Alternative lengthening of telomeres: models, mechanisms and implications, Nat. Rev. Genet, vol.11, pp.319-330, 2010.

H. A. Pickett and R. R. Reddel, Molecular mechanisms of activity and derepression of alternative lengthening of telomeres, Nat. Struct. Mol. Biol, vol.22, pp.875-880, 2015.

E. M. Tacconi and M. Tarsounas, How homologous recombination maintains telomere integrity, Chromosoma, vol.124, pp.119-130, 2015.

S. Le, J. K. Moore, J. E. Haber, and C. W. Greider, RAD50 and RAD51 define two pathways that collaborate to maintain telomeres in the absence of telomerase, Genetics, vol.152, pp.143-152, 1999.

M. S. Fitzgerald, K. Riha, F. Gao, S. Ren, T. D. Mcknight et al., Disruption of the telomerase catalytic subunit gene from Arabidopsis inactivates telomerase and leads to a slow loss of telomeric DNA, Proc. Natl. Acad. Sci. U.S.A, vol.96, pp.14813-14818, 1999.

K. Riha, T. D. Mcknight, L. R. Griffing, and D. E. Shippen, Living with genome instability: plant responses to telomere dysfunction, Science, vol.291, pp.1797-1800, 2001.

, Nucleic Acids Research, vol.46, issue.5, p.2445, 2018.

E. Ruckova, J. Friml, P. Prochazkova-schrumpfova, and J. Fajkus, Role of alternative telomere lengthening unmasked in telomerase knock-out mutant plants, Plant Mol. Biol, vol.66, pp.637-646, 2008.

L. Vespa, R. T. Warrington, P. Mokros, J. Siroky, and D. E. Shippen, ATM regulates the length of individual telomere tracts in Arabidopsis, Proc. Natl. Acad. Sci. U.S.A, vol.104, pp.18145-18150, 2007.

O. Da-ines, F. Degroote, C. Goubely, S. Amiard, M. E. Gallego et al., Meiotic recombination in Arabidopsis is catalysed by DMC1, with RAD51 playing a supporting role, PLoS Genet, vol.9, p.1003787, 2013.
URL : https://hal.archives-ouvertes.fr/inserm-01907382

W. Kobayashi, S. Sekine, S. Machida, and H. Kurumizaka, Green fluorescent protein fused to the C terminus of RAD51 specifically interferes with secondary DNA binding by the RAD51-ssDNA complex, Genes Genet. Syst, vol.89, pp.169-179, 2014.

P. Kalhorzadeh, Z. Hu, T. Cools, S. Amiard, E. M. Willing et al., Arabidopsis thaliana RNase H2 deficiency counteracts the needs for the WEE1 checkpoint kinase but triggers genome instability, Plant Cell, vol.26, pp.3680-3692, 2014.
URL : https://hal.archives-ouvertes.fr/inserm-01907361

J. Recker, A. Knoll, and H. Puchta, The Arabidopsis thaliana homolog of the helicase RTEL1 plays multiple roles in preserving genome stability, Plant Cell, vol.26, pp.4889-4902, 2014.

C. Charbonnel, M. E. Gallego, and C. I. White, Xrcc1-dependent and Ku-dependent DNA double-strand break repair kinetics in Arabidopsis plants, Plant J, vol.64, pp.280-290, 2010.
URL : https://hal.archives-ouvertes.fr/inserm-00595784

M. T. Kurzbauer, C. Uanschou, D. Chen, and P. Schlogelhofer, The recombinases DMC1 and RAD51 are functionally and spatially separated during meiosis in Arabidopsis, Plant Cell, vol.24, pp.2058-2070, 2012.

S. Amiard, C. Charbonnel, E. Allain, A. Depeiges, C. I. White et al., Distinct roles of the ATR kinase and the Mre11-Rad50-Nbs1 complex in the maintenance of chromosomal stability in Arabidopsis, Plant Cell, vol.22, pp.3020-3033, 2010.
URL : https://hal.archives-ouvertes.fr/inserm-00595794

I. Schubert, P. F. Fransz, J. Fuchs, and J. H. De-jong, Chromosome painting in plants, Methods Cell Sci, vol.23, pp.57-69, 2001.
DOI : 10.1007/978-94-010-0330-8_7

J. B. Vannier, A. Depeiges, C. White, and M. E. Gallego, Two roles for Rad50 in telomere maintenance, EMBO J, vol.25, pp.4577-4585, 2006.
URL : https://hal.archives-ouvertes.fr/inserm-00595800

M. E. Gallego and C. I. White, RAD50 function is essential for telomere maintenance in Arabidopsis, Proc. Natl. Acad. Sci. U.S.A, vol.98, pp.1711-1716, 2001.
DOI : 10.1073/pnas.98.4.1711

URL : https://hal.archives-ouvertes.fr/inserm-00595834

M. Heacock, E. Spangler, K. Riha, J. Puizina, and D. E. Shippen, Molecular analysis of telomere fusions in Arabidopsis: multiple pathways for chromosome end-joining, EMBO J, vol.23, pp.2304-2313, 2004.

J. Y. Bleuyard and C. I. White, The Arabidopsis homologue of Xrcc3 plays an essential role in meiosis, EMBO J, vol.23, pp.439-449, 2004.
URL : https://hal.archives-ouvertes.fr/inserm-00595813

J. B. Vannier, A. Depeiges, C. White, and M. E. Gallego, ERCC1/XPF protects short telomeres from homologous recombination in Arabidopsis thaliana, PLoS Genet, vol.5, p.1000380, 2009.
DOI : 10.1371/journal.pgen.1000380

URL : https://hal.archives-ouvertes.fr/inserm-00595799

H. Gaillard and A. Aguilera, Transcription as a threat to genome integrity, Annu. Rev. Biochem, vol.85, pp.291-317, 2016.
DOI : 10.1146/annurev-biochem-060815-014908

URL : https://digital.csic.es/bitstream/10261/165338/1/accesoRestringido.pdf

E. Majerova, M. Fojtova, I. Mozgova, M. Bittova, and J. Fajkus, Hypomethylating drugs efficiently decrease cytosine methylation in telomeric DNA and activate telomerase without affecting telomere lengths in tobacco cells, Plant Mol. Biol, vol.77, pp.371-380, 2011.

J. Vrbsky, S. Akimcheva, J. M. Watson, T. L. Turner, L. Daxinger et al., ) siRNA-mediated methylation of Arabidopsis telomeres, PLoS Genet, vol.6, p.1000986, 2010.

K. Rippe and B. Luke, TERRA and the state of the telomere, Nat. Struct. Mol. Biol, vol.22, pp.853-858, 2015.

L. J. Barber, J. L. Youds, J. D. Ward, M. J. Mcilwraith, N. J. O'neil et al., RTEL1 maintains genomic stability by suppressing homologous recombination, vol.135, pp.261-271, 2008.
DOI : 10.1016/j.cell.2008.08.016

URL : https://doi.org/10.1016/j.cell.2008.08.016

B. Khadaroo, M. T. Teixeira, P. Luciano, N. Eckert-boulet, S. M. Germann et al., The DNA damage response at eroded telomeres and tethering to the nuclear pore complex, Nat. Cell Biol, vol.11, pp.980-987, 2009.

Z. Xie, K. A. Jay, D. L. Smith, Y. Zhang, Z. Liu et al., Early telomerase inactivation accelerates aging independently of telomere length, Cell, vol.160, pp.928-939, 2015.

K. A. Jay, D. L. Smith, and E. H. Blackburn, Early loss of telomerase action in yeast creates a dependence on the DNA damage response adaptor proteins, Mol. Cell. Biol, vol.36, pp.1908-1919, 2016.

Z. Xu, E. Fallet, C. Paoletti, S. Fehrmann, G. Charvin et al., Two routes to senescence revealed by real-time analysis of telomerase-negative single lineages, Nat. Commun, vol.6, p.7680, 2015.
URL : https://hal.archives-ouvertes.fr/hal-01308895

S. Badie, J. M. Escandell, P. Bouwman, A. R. Carlos, M. Thanasoula et al., BRCA2 acts as a RAD51 loader to facilitate telomere replication and capping, Nat. Struct. Mol. Biol, vol.17, pp.1461-1469, 2010.

A. A. Neumann, C. M. Watson, J. R. Noble, H. A. Pickett, P. P. Tam et al., Alternative lengthening of telomeres in normal mammalian somatic cells, Genes Dev, vol.27, pp.18-23, 2013.

M. A. Blasco, H. W. Lee, M. P. Hande, E. Samper, P. M. Lansdorp et al., Telomere shortening and tumor formation by mouse cells lacking telomerase RNA, Cell, vol.91, pp.25-34, 1997.
DOI : 10.1016/s0092-8674(01)80006-4

URL : https://doi.org/10.1016/s0092-8674(01)80006-4

J. Min, W. E. Wright, and J. W. Shay, Alternative lengthening of telomeres mediated by mitotic DNA synthesis engages break-induced replication processes, Mol. Cell. Biol, vol.37, pp.226-243, 2017.

F. M. Roumelioti, S. K. Sotiriou, V. Katsini, M. Chiourea, T. D. Halazonetis et al., Alternative lengthening of human telomeres is a conservative DNA replication process with features of break-induced replication, EMBO Rep, vol.17, pp.1731-1737, 2016.

N. W. Cho, R. L. Dilley, M. A. Lampson, and R. A. Greenberg, Interchromosomal homology searches drive directional ALT telomere movement and synapsis, Cell, vol.159, pp.108-121, 2014.
DOI : 10.1016/j.cell.2014.08.030

URL : https://doi.org/10.1016/j.cell.2014.08.030

R. L. Dilley, P. Verma, N. W. Cho, H. D. Winters, A. R. Wondisford et al., Break-induced telomere synthesis underlies alternative telomere maintenance, Nature, vol.539, pp.54-58, 2016.
DOI : 10.1038/nature20099

URL : http://europepmc.org/articles/pmc5384111?pdf=render

C. Y. Lin, H. H. Chang, K. J. Wu, S. F. Tseng, C. C. Lin et al., Extrachromosomal telomeric circles contribute to Rad52-, Rad50-, and polymerase delta-mediated telomere-telomere recombination in Saccharomyces cerevisiae, Eukaryot. Cell, vol.4, pp.327-336, 2005.
DOI : 10.1128/ec.4.2.327-336.2005

URL : http://europepmc.org/articles/pmc549320?pdf=render

M. Larrivee and R. J. Wellinger, Telomerase-and capping-independent yeast survivors with alternate telomere states, Nat. Cell Biol, vol.8, pp.741-747, 2006.

E. Y. Basenko, A. J. Cesare, S. Iyer, J. D. Griffith, and M. J. Mceachern, Telomeric circles are abundant in the stn1-M1 mutant that maintains its telomeres through recombination, Nucleic Acids Res, vol.38, pp.182-189, 2010.

J. D. Henson, Y. Cao, L. I. Huschtscha, A. C. Chang, A. Y. Au et al., DNA C-circles are specific and quantifiable markers of alternative-lengthening-of-telomeres activity, Nat. Biotechnol, vol.27, pp.1181-1185, 2009.

B. Zellinger, S. Akimcheva, J. Puizina, M. Schirato, and K. Riha, Ku suppresses formation of telomeric circles and alternative telomere lengthening in Arabidopsis, Mol. Cell, vol.27, pp.163-169, 2007.

C. I. Nugent, G. Bosco, L. O. Ross, S. K. Evans, A. P. Salinger et al., Telomere maintenance is dependent on activities required for end repair of double-strand breaks, Curr. Biol, vol.8, pp.657-660, 1998.

M. J. Mceachern and J. E. Haber, Break-induced replication and recombinational telomere elongation in yeast, Annu. Rev. Biochem, vol.75, pp.111-135, 2006.

C. J. Sakofsky and A. Malkova, Break induced replication in eukaryotes: mechanisms, functions, and consequences, Crit. Rev. Biochem. Mol. Biol, vol.52, pp.395-413, 2017.

A. P. Sobinoff, J. A. Allen, A. A. Neumann, S. F. Yang, M. E. Walsh et al., BLM and SLX4 play opposing roles in recombination-dependent replication at human telomeres, EMBO J, vol.36, pp.2907-2919, 2017.