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J. Biol. Chem., Vol. 276, Issue 20, 17092-17100, May 18, 2001
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and
From the Laboratoire de Génétique Moléculaire,
Ecole Normale Supérieure, 46 rue d'Ulm,
75230 Paris Cedex 05, France
DNA replication origins are located at random
with respect to DNA sequence in Xenopus early embryos and
on DNA replicated in Xenopus egg extracts. We have recently
shown that origins fire throughout the S phase in Xenopus
egg extracts. To study the temporal regulation of origin firing, we
have analyzed origin activation in sperm nuclei treated with the DNA
polymerase inhibitor aphidicolin. Sperm chromatin was incubated in
Xenopus egg extracts in the presence of aphidicolin and
transferred to a fresh extract, and digoxigenin-dUTP and biotin-dUTP
were added at various times after aphidicolin release to selectively
label early and late replicating DNA. Molecular combing analysis of
single DNA fibers showed that only a fraction of potential origins were
able to initiate in the presence of aphidicolin. After release from
aphidicolin, the remaining origins fired asynchronously throughout the
S phase. Therefore, initiation during the S phase depends on the normal
progression of replication forks assembled at earlier activated
origins. Caffeine, an inhibitor of the checkpoint kinases ATR and ATM,
did not relieve the aphidicolin-induced block to origin firing. We
conclude that a caffeine-insensitive intra-S phase checkpoint regulates
origin activation when DNA synthesis is inhibited in
Xenopus egg extracts.
Supported by fellowships from the Association pour la
Recherche sur le Cancer and the Fondation pour la Recherche
Médicale and by the CNRS.
§
To whom correspondence should be addressed. Tel.:
33-1-44-32-39-20; Fax: 33-1-44-32-39-41; E-mail:
hyrien@wotan.ens.fr.
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