1. Gadaleta,M.C. and Noguchi,E. (2017) Regulation of DNA
replication through natural impediments in the eukaryotic genome.
Genes (Basel)., 8, 98.
2. Brewer,B.J. and Fangman,W.L. (1988) A replication fork barrier at
the 3 end of yeast ribosomal RNA genes. Cell, 55, 637–643.
3. Sanchez,J.A., Kim,S.M. and Huberman,J.a. (1998) Ribosomal DNA
replication in the fission yeast, Schizosaccharomyces pombe. Exp. Cell
Res., 238, 220–230.
4. Kobayashi,T. and Horiuchi,T. (1996) A yeast gene product, Fob1
protein, required for both replication fork blocking and
recombinational hotspot activities. Genes Cells, 1, 465–474.
5. Sanchez-Gorostiaga,A., Lopez-Estrano,C., Krimer,D.B.,
Schvartzman,J.B. and Hernandez,P. (2004) Transcription termination
factor reb1p causes two replication fork barriers at its cognate sites in
fission yeast ribosomal DNA in vivo. Mol. Cell. Biol., 24, 398–406.
6. Krings,G. and Bastia,D. (2005) Sap1p binds to Ter1 at the ribosomal
DNA of Schizosaccharomyces pombe and causes polar replication
fork arrest. J. Biol. Chem., 280, 39135–39142.
7. Weitao,T., Budd,M., Hoopes,L.L.M. and Campbell,J.L. (2003) Dna2
helicase/nuclease causes replicative fork stalling and double-strand
breaks in the ribosomal DNA of Saccharomyces cerevisiae. J. Biol.
Chem., 278, 22513–22522.
8. Kobayashi,T., Heck,D.J., Nomura,M. and Horiuchi,T. (1998)
Expansion and contraction of ribosomal DNA repeats in (Fob1)
protein and the role of RNA polymerase I Expansion and
contraction of ribosomal DNA repeats in Saccharomyces cerevisiae:
requirement of replication fork blocking (Fob1) protein and the role.
Genes Dev., 12, 3821–3830.
9. Sasaki,M. and Kobayashi,T. (2017) Ctf4 prevents genome
rearrangements by suppressing DNA double-strand break formation
and its end resection at arrested replication forks. Mol. Cell, 66,
533–545.
10. Hiraoka,Y., Henderson,E. and Blackburn,E.H. (1998) Not so
peculiar: fission yeast telomere repeats. Trends Biochem. Sci., 23, 126.
11. Ishikawa,F. (2013) Portrait of replication stress viewed from
telomeres. Cancer Sci., 104, 790–794.
12. Griffith,J.D., Comeau,L., Rosenfield,S., Stansel,R.M., Bianchi,A.,
Moss,H. and De Lange,T. (1999) Mammalian telomeres end in a
large duplex loop. Cell, 97, 503–514.
13. Parkinson,G.N., Lee,M.P.H. and Neidle,S. (2002) Crystal structure of
parallel quadruplexes from human telomeric DNA. Nature, 417,
876–880.
14. 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, 6128–6138.
15. Grandin,N., Reed,S.I. and Charbonneau,M. (1997) Stn1, a new
Saccharomyces cerevisiae protein, is implicated in telomere size
regulation in association with Cdc13. Genes Dev., 11, 512–527.
16. 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.
Downloaded from https://academic.oup.com/nar/article/49/18/10465/6370253 by Kyoto Daigaku Johogakukenkyuka Tosho user on 25 May 2022
We thank M. Tamura and Y. Watanabe for technical assistance, and A. Katayama, A. Shirabuchi, E. Yamazaki,
S. Fukumura, N. Hayashi, T. Tsuda and Y. Hirata for excellent secretarial work, and J. Hejna for critically reading
the manuscript. This study was conducted through the Joint
Usage/Research Center Program of the Radiation Biology
Center, Kyoto University.
17. 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, 36, 193–206.
18. De Lange,T. (2005) Shelterin: the protein complex that shapes and
safeguards human telomeres. Genes Dev., 19, 2100–2110.
19. Miyoshi,T., Kanoh,J., Saito,M. and Ishikawa,F. (2008) Fission yeast
Pot1-Tpp1 protects telomeres and regulates telomere length. Science,
320, 1341–1344.
20. Mart´ın,V., Du,L.-L., Rozenzhak,S. and Russell,P. (2007) Protection
of telomeres by a conserved Stn1-Ten1 complex. Proc. Natl. Acad.
Sci. U.S.A., 104, 14038–14043.
21. Takikawa,M., Tarumoto,Y. and Ishikawa,F. (2017) Fission yeast Stn1
is crucial for semi-conservative replication at telomeres and
subtelomeres. Nucleic Acids Res., 45, 1255–1269.
22. Matmati,S., Vaurs,M., Escandell,J.M., Maestroni,L.,
Nakamura,T.M., Ferreira,M.G., G´eli,V. and Coulon,S. (2018) The
fission yeast Stn1-Ten1 complex limits telomerase activity via its
SUMO-interacting motif and promotes telomeres replication. Sci.
Adv., 4, eaar2740.
23. Miller,K.M., Rog,O. and Cooper,J.P. (2006) Semi-conservative DNA
replication through telomeres requires Taz1. Nature, 440, 824–828.
24. Oizumi,Y., Kaji,T., Tashiro,S., Takeshita,Y., Date,Y. and Kanoh,J.
(2021) Complete sequences of Schizosaccharomyces pombe
subtelomeres reveal multiple patterns of genome variation. Nat.
Commun., 12, 3–8.
25. Waga,S. and Stillman,B. (1994) Anatomy of a DNA replication fork
revealed by reconstitution of SV40 DNA replication in vitro. Nature,
369, 207–212.
26. 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.
27. Escandell,J.M., Carvalho,E.S., Gallo-Fernandez,M., Reis,C.C.,
Matmati,S., Lu´ıs,I.M., Abreu,I.A., Coulon,S. and Ferreira,M.G.
(2019) Ssu72 phosphatase is a conserved telomere replication
terminator. EMBO J., 38, e100476.
28. Ganduri,S. and Lue,N.F. (2017) STN1-POLA2 interaction provides a
basis for primase-pol ␣ stimulation by human STN1. Nucleic Acids
Res., 45, 9455–9466.
29. Lue,N.F., Chan,J., Wright,W.E. and Hurwitz,J. (2014) The
CDC13-STN1-TEN1 complex stimulates Pol ␣ activity by promoting
RNA priming and primase-to-polymerase switch. Nat. Commun., 5,
5762.
30. Nakaoka,H., Nishiyama,A., Saito,M. and Ishikawa,F. (2012)
Xenopus laevis Ctc1-Stn1-Ten1 (xCST) protein complex is involved in
priming DNA synthesis on single-stranded DNA template in Xenopus
egg extract. J. Biol. Chem., 287, 619–627.
31. Casteel,D.E., Zhuang,S., Zeng,Y., Perrino,F.W., Boss,G.R.,
Goulian,M. and Pilz,R.B. (2009) A DNA polymerase-␣·primase
cofactor with homology to replication protein A-32 regulates DNA
replication in mammalian cells. J. Biol. Chem., 284, 5807–5818.
32. Stewart,J., Wang,F., Chaiken,M.F., Kasbek,C., Chastain,P.D.,
Wright,W.E. and Price,C.M. (2012) Human CST promotes telomere
duplex replication and general replication restart after fork stalling.
EMBO J., 31, 3537–3549.
33. Chastain,M., Zhou,Q., Shiva,O., Whitmore,L., Jia,P., Dai,X.,
Huang,C., Fadri-Moskwik,M., Ye,P. and Chai,W. (2016) Human CST
facilitates genome-wide RAD51 recruitment to GC-rich repetitive
sequences in response to replication stress. Cell Rep., 16, 1300–1314.
34. Alfa,C., Fantes,P., Hyams,J., McLeod,M. and Warbrick,E. (1993) In:
Experiments with Fission Yeast: A Laboratory Course Manual. Cold
Spring Harbor Laboratory Press, NY
35. Sutani,T., Sakata,T., Nakato,R., Masuda,K., Ishibashi,M.,
Yamashita,D., Suzuki,Y., Hirano,T., Bando,M. and Shirahige,K.
(2015) Condensin targets and reduces unwound DNA structures
associated with transcription in mitotic chromosome condensation.
Nat. Commun., 6, 7815.
36. Miyoshi,T., Kanoh,J. and Ishikawa,F. (2009) Fission yeast Ku protein
is required for recovery from DNA replication stress. Genes Cells, 14,
1091–1103.
37. Kaiser,C., Michaelis,S. and Mitchell,A. (1994) In: Methods in Yeast
Genetics: A Cold Spring Harbor Laboratory Course Manual. Cold
Spring Harbor Laboratory Press, NY.
A Self-archived copy in
Kyoto University Research Information Repository
https://repository.kulib.kyoto-u.ac.jp
10476 Nucleic Acids Research, 2021, Vol. 49, No. 18
56. Noguchi,E., Noguchi,C., McDonald,W.H., Yates,J.R. and Russell,P.
(2004) Swi1 and Swi3 are components of a replication fork protection
complex in fission yeast. Mol. Cell. Biol., 24, 8342–8355.
57. Noguchi,E., Noguchi,C., Du,L.-L. and Russell,P. (2003) Swi1
prevents replication fork collapse and controls checkpoint kinase
Cds1. Mol. Cell. Biol., 23, 7861–7874.
58. Gadaleta,M.C., Das,M.M., Tanizawa,H., Chang,Y.-T., Noma,K.,
Nakamura,T.M. and Noguchi,E. (2016) Swi1Timeless prevents repeat
instability at fission yeast telomeres. PLoS Genet., 12, e1005943.
59. Cooper,J.P., Nimmo,E.R., Allshire,R.C. and Cech,T.R. (1997)
Regulation of telomere length and function by a Myb-domain protein
in fission yeast. Nature, 385, 744–747.
60. Kanoh,J., Sadaie,M., Urano,T. and Ishikawa,F. (2005) Telomere
binding protein Taz1 establishes Swi6 heterochromatin independently
of RNAi at telomeres. Curr. Biol., 15, 1808–1819.
61. Wang,Y. and Chai,W. (2018) Pathogenic CTC1 mutations cause
global genome instabilities under replication stress. Nucleic Acids
Res., 46, 3981–3992.
62. Durkin,S.G. and Glover,T.W. (2007) Chromosome fragile sites. Annu.
Rev. Genet., 41, 169–192.
63. Ide,S., Miyazaki,T., Maki,H. and Kobayashi,T. (2010) Abundance of
ribosomal RNA gene copies maintains genome integrity. Science,
327, 693–696.
64. Houseley,J. and Tollervey,D. (2011) Repeat expansion in the budding
yeast ribosomal DNA can occur independently of the canonical
homologous recombination machinery. Nucleic Acids Res., 39,
8778–8791.
65. Ganley,A.R.D., Ide,S., Saka,K. and Kobayashi,T. (2009) The effect
of replication initiation on gene amplification in the rDNA and its
relationship to aging. Mol. Cell, 35, 683–693.
66. Rustchenko,E.P., Curran,T.M. and Sherman,F. (1993) Variations in
the number of ribosomal DNA units in morphological mutants and
normal strains of Candida albicans and in normal strains of
Saccharomyces cerevisiae. J. Bacteriol., 175, 7189–7199.
67. Bentsen,I.B., Nielsen,I., Lisby,M., Nielsen,H.B., Gupta,S. Sen,
Mundbjerg,K., Andersen,A.H. and Bjergbaek,L. (2013) MRX
protects fork integrity at protein-DNA barriers, and its absence
causes checkpoint activation dependent on chromatin context.
Nucleic Acids Res., 41, 3173–3189.
68. Carneiro,T., Khair,L., Reis,C.C., Borges,V., Moser,B.,
Nakamura,T.M. and Ferreira,M.G. (2010) Telomeres avoid end
detection by severing the checkpoint signal transduction pathway.
Nature, 467, 228–232.
69. Audry,J., Wang,J., Eisenstatt,J.R., Berkner,K.L. and Runge,K.W.
(2018) The inhibition of checkpoint activation by telomeres does not
involve exclusion of dimethylation of histone h4 lysine 20
(H4k20me2). F1000Research, 7, 1027.
70. Wang,Y., Brady,K.S., Caiello,B.P., Ackerson,S.M. and Stewart,J.A.
(2019) Human CST suppresses origin licensing and promotes
AND-1/Ctf4 chromatin association. Life Sci. Alliance, 2, e201800270.
71. Tanaka,H., Katou,Y., Yagura,M., Saitoh,K., Itoh,T., Araki,H.,
Bando,M. and Shirahige,K. (2009) Ctf4 coordinates the progression
of helicase and DNA polymerase ␣. Genes Cells, 14, 807–820.
72. Berthiau,A.S., Yankulov,K., Bah,A., Revardel,E., Luciano,P.,
Wellinger,R.J., G´eli,V. and Gilson,E. (2006) Subtelomeric proteins
negatively regulate telomere elongation in budding yeast. EMBO J.,
25, 846–856.
73. Pasquier,E. and Wellinger,R.J. (2020) In vivo chromatin organization
on native yeast telomeric regions is independent of a cis-telomere
loopback conformation. Epigenetics Chromatin, 13, 23.
Downloaded from https://academic.oup.com/nar/article/49/18/10465/6370253 by Kyoto Daigaku Johogakukenkyuka Tosho user on 25 May 2022
38. Sugawara,N.F. (1988) DNA Sequences at the Telomeres of the
Fission Yeast S. Pombe. Ph.D. dissertation. Harvard University,
Cambridge, MA.
39. Skoog,L. and Nordenskjold,B.
(1971) Effects of hydroxyurea and
1-beta-D-arabinofuranosyl-cytosine on deoxyribonucleotide pools in
mouse embryo cells. Eur. J. Biochem., 19, 81–89.
40. Naito,T., Matsuura,a and Ishikawa,F. (1998) Circular chromosome
formation in a fission yeast mutant defective in two ATM
homologues. Nat. Genet., 20, 203–206.
41. Nakamura,T.M., Cooper,J.P. and Cech,T.R. (1998) Two modes of
survival of fission yeast without telomerase. Science, 282, 493–496.
42. Sadaie,M., Naito,T. and Ishikawa,F. (2003) Stable inheritance of
telomere chromatin structure and function in the absence of telomeric
repeats. Genes Dev., 17, 2271–2282.
43. Jain,D., Hebden,A.K., Nakamura,T.M., Miller,K.M. and Cooper,J.P.
(2010) HAATI survivors replace canonical telomeres with blocks of
generic heterochromatin. Nature, 467, 223–227.
44. Tashiro,S., Nishihara,Y., Kugou,K., Ohta,K. and Kanoh,J. (2017)
NAR breakthrough article: subtelomeres constitute a safeguard for
gene expression and chromosome homeostasis. Nucleic Acids Res.,
45, 10333–10349.
45. Toda,T., Nakaseko,Y., Niwa,O. and Yanagida,M. (1984) Mapping of
rRNA genes by integration of hybrid plasmids in
Schizosaccharomyces pombe. Curr. Genet., 8, 93–97.
46. Maleszka,R. and Clark-Walker,G.D. (1993) Yeasts have a four-fold
variation in ribosomal DNA copy number. Yeast, 9, 53–58.
47. Krings,G. and Bastia,D. (2004) swi1- and swi3-dependent and
independent replication fork arrest at the ribosomal DNA of
Schizosaccharomyces pombe. Proc. Natl. Acad. Sci. U.S.A., 101,
14085–14090.
48. Hayashi,M., Katou,Y., Itoh,T., Tazumi,A., Tazumi,M., Yamada,Y.,
Takahashi,T., Nakagawa,T., Shirahige,K. and Masukata,H. (2007)
Genome-wide localization of pre-RC sites and identification of
replication origins in fission yeast. EMBO J., 26, 1327–1339.
49. Fan,J., Grothues,D. and Smith,C.L. (1991) Alignment of Sfi I sites
with the Not I restriction map of Schizosaccharomyces pombe
genome. Nucleic Acids Res., 19, 6289–6294.
50. Lisby,M., Mortensen,U.H. and Rothstein,R. (2003) Colocalization of
multiple DNA double-strand breaks at a single Rad52 repair center.
Nat. Cell Biol., 5, 572–577.
51. Nakamura,T.M., Du,L., Redon,C. and Russell,P. (2004) Histone
H2A phosphorylation controls Crb2 recruitment at DNA breaks,
maintains checkpoint arrest, and influences DNA repair in fission
yeast. Mol. Cell. Biol., 24, 7820–7820.
52. Rogakou,E.P., Pilch,D.R., Orr,A.H., Ivanova,V.S. and Bonner,W.M.
(1998) Double-stranded breaks induce histone H2AX
phosphorylation on serine 139. J. Biol. Chem., 273, 5858–5868.
53. Rozenzhak,S., Mej´ıa-Ram´ırez,E., Williams,J.S., Schaffer,L.,
Hammond,J., Head,S.R. and Russell,P. (2010) Rad3 decorates critical
chromosomal domains with gammaH2A to protect genome integrity
during S-Phase in fission yeast. PLoS Genet., 6, e1001032.
54. Boddy,M.N., Gaillard,P.H.L., McDonald,W.H., Shanahan,P.,
Yates,J.R. and Russell,P. (2001) Mus81-Eme1 are essential
components of a Holliday junction resolvase. Cell, 107, 537–548.
55. Ii,M., Ii,T. and Brill,S.J. (2007) Mus81 functions in the quality
control of replication forks at the rDNA and is involved in the
maintenance of rDNA repeat number in Saccharomyces cerevisiae.
Mutat. Res. - Fundam. Mol. Mech. Mutagen., 625, 1–19.
...
論文の公開元へ
