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Fig. 2. ATM is strictly required for chromatin-wide phosphorylation of H2AX during leptotene. (A) Leptotene Atm^-/- spermatocytes are devoid of ${gamma}$ H2AX. Quantitative analysis of Atm^-/- surface-spread preparations shows that 100% of the Atm^-/- spermatocytes undergoing leptonema are devoid of ${gamma}$ H2AX (n=64 nuclei, two animals). (B) Leptotene ATM-deficient spermatocytes can not phosphorylate H2AX in response to radiation-induced DSBs: Spo11^-/- (a,b) and Atm^-/- Spo11^-/- (c,d) double-mutant mice were irradiated (3 Gy) or mock treated. 1 hour after irradiation, mice were sacrificed and their testes used to prepare structurally preserved nuclei, which were subsequently immunolabeled with antibodies against SCP3 (in red) and ${gamma}$ H2AX (in green). 50 nuclei were scored for each condition. Irradiation-induced DSBs can restore H2AX phosphorylation in Spo11^-/- spermatocytes undergoing leptotene (b), whereas, in the absence of functional ATM, no alternative PIKK can phosphorylate H2AX in leptotene spermatocytes in response to radiation-induced DSBs (d). The bright red spots represent nucleolar SCP3 aggregates.

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