
Медицинская вирусология / Букринская А.Г., Жданов В.М. Молекулярные основы патогенности вирусов
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al., 1989]. B JIHMrpouHTax, Cb!B0J)0TKe Kp0BH II Cf!HHH0· M03fOBOEi }Kl!/l,K0CTH C II0M0!Ub!0 MOJICKYJIHpHoii r116pH,ll,l!33L1HH 6wrn o6ttapy)KeHbl reHCTJ1tICCKlle II0CJICJl.0BaTeJibH0CTH BHpyca rprnma. B Ka l lCCTBC [1H)ll10ilKTillllll,J., :mu;wn 6hIJIII HCII0.flb30!laHbl K,Jiom1poBaHHbIC B 11.naaMH/1,ax reHbI NP, M, Hl II H3.
140