publ.vp

Transcription

1 ｨｬ 52, , HSP70, ﾁ ｨｬ02ｨｪｨｺ06, ｨｪｨｰｨｨｨｬ06ｨｪ0609ｨ, ｨｺｨ ｨ, ｨｪｨｨｨｺ, ｨ 0800ｨｮ05ｨｨ09, ｨｮ030609ｨ 1, * 1 06ｨｪ09ｨｰｨｨｨｰｨｮｨｰ 02ｨｨｨｰ ｨｨｨｨ , 05ｨ ｨｪｨｺｨｰ-0302ｨｰ0208ｨ ｨｮ0800, ｨｨ 2 06ｨｨ010606ｨｺ02ｨ ｨｪ09ｨｺｨｨｨｦ ｨｨｨｪ09ｨｰｨｨｨｰｨｮｨｰ ｨ ｨｨ ｨ ｨｪｨｨ ﾂ0209ｨｺ06ｨｦ 01ｨｨｨｬｨｨｨｨ , 0005ｨ 01ｨｨ090609ｨｰ06ｨｺ; *050502ｨｺｨｰ0806ｨｪｨｪ04ｨｦ ｨ : guzhova@mail.cytspb.rssi.ru ｨｬ02ｨｪｨｺ06, ｨｪｨｰｨｨｨｬ06ｨｪ0609ｨ ｨｨ ｨｨｨｪ02ｨｪｨｨ07, ｨ 06ｨｴｨｨ02 05ｨｺ ｨｨ06 03ｨ ｨｪｨ Hsp ｨｺ Hsp70 06ｨ 05ｨ 01ｨ 02ｨｰ 03ｨ ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ06, ｨｨ 05ｨｰ ｨｦ09ｨｰ ｨ ｨｪ ｨ ｨｴｨｨｨｰｨｪ06ｨｦ 00ｨｮｨｪｨｺ02ｨｨ02ｨｦ, ｨｺ06ｨｰ0608ｨ ｨｬ06ｨｪ09ｨｰ08ｨｨ080609ｨ ｨｪｨ ｨｰｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨ02ｨｬ ｨｺ0502ｨｰ06 ﾂｨｪ ｨｨｨｪｨｨｨｦ ｨｨ 03ｨｨ0906ｨｰｨｪ ｨｰ06ｨｬｨｮ ｨ ｨｪｨ - ﾂ02ｨｪｨｨ ｨｨ06ｨ 0802ｨｰｨ 02ｨｰ 0706ｨｨ09ｨｺ 0902ｨｴ0209ｨｰ09, ｨ ｨｪ0401 ｨ ｨｪ04ｨｬ 06ｨ 08ｨ 0406ｨｬ ｨｨｨｰｨｹ ｨ ｨｪｨｨ02 03ｨ ｨｪｨ 09 ｨｰｨｺｨ ｨｪ0701 ｨｨ ｨｺ0502ｨｰｨｺｨ ｨ ｨｪｨｨ04ｨｬｨ ｨｦ 08ｨ ｨ 06ｨｰ02 ｨｬ ｨｨ 09ｨｺ08ｨｨｨｪｨｨｨｪ00 ｨ ｨｨｨｪ02ｨｪｨｨｨｦ ｨｨ ｨｨ05ｨｨ 0109ｨ 0902ｨｴ0209ｨｰ09ｨ, ｨｨ ｨｪ ｨｨｨｺ06ｨｪｨｨｨｪｨ ｨｨ 0501ｨｨｨｪ ｨｬｨ, ｨｺ06ｨｰ ｨｬｨｨｨｺ0806ｨｬ ｨｪ0401 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ ｨ ｨｪ04 ｨｮ090205ｨｨ ﾂｨｨ09ｨ ｨｰｨｹ ｨ ｨｪｨｨ02 03ｨ ｨｪｨ 09 08ｨ 0405ｨｨ ﾂｨｪ0401 ｨｺ0502ｨｰｨｺｨ 01 ﾂ ｨｺｨ ｨｦ09ｨｰ09ｨｨ02ｨｬ 05ｨｰｨｨ ｨｮ ｨｴ0209ｨｰ09 09 ｨｺ0502ｨｰｨｺｨ ｨｨｨｰ ｨｦｨｺ02ｨｬｨｨｨｨ ﾂ ｨｺｨ K562 ｨｰｨ ｨｺ0302 ｨｪｨ ｨ ｨ 02ｨｰ ｨｪｨ ﾂｨｨｨｰ0205ｨｹｨｪ0602 ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 ｨ ｨ ｨｰｨｪ06ｨｦ 03ｨ ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ Hsp70, ﾂｨｰ06 ｨｬ060302ｨｰ ｨｮｨｺｨ ｨ ｨｰｨｹ ｨｪｨ ｨｬ06ｨ ｨｨ05ｨｨ04ｨ 02ｨｨ ｨ ｨｪｨｪ ｨｬ0201ｨ ｨｪｨｨ04ｨｬｨ ｨｺ0502ｨｰｨｺｨｨ ｨｪｨｺｨ ｨ ｨｨ ｨｨ ﾂ0209ｨｺ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ ｨｨｨｪ02ｨｪｨｨｨｦ 0706ｨｺｨ 04ｨ 05ｨ, ﾂｨｰ06 06ｨ 08ｨ ｨ 06ｨｰｨｺｨ ｨｨｨｬｨｨ 0708ｨｨ090601ｨｨｨｰ ｨｺ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ ｨｨ ｨｨ0903ｨｨ01 ｨｺ0502ｨｰ06ｨｺ K ｨｨ 06ｨ 08ｨ ｨ 06ｨｰｨｺ ｨｰｨｺｨｨｨｬ ｨｰ ｨｬ 0306ｨｺ06ｨｬ, ｨｺｨｨ09ｨｹ ｨ ｨｨ 09ｨｰｨ ｨｮ ｨｨｨｪ06ｨｬ ｨｪｨ 20 ｪ50 % 09 04ｨ 09ｨｨ09ｨｨｨｬ0609ｨｰｨｨ 06ｨｰ 02ｨｨｨｰ06ｨｰ06ｨｺ09ｨｨ ﾂ0209ｨｺ ｨ ｨｺｨｰ0608ｨ ｨｮ05ｨｹｨｰｨ ｨｰ ｨｺｨ ｨ 06ｨｰ, ﾂｨｰ ｨｨ ｨｮ ｨｨｨｨ ｨｪ ｨｪ ｨｨｨｬｨｨ ﾂ0209ｨｺｨｨ01 ｨｬ0601ｨｨ00ｨｨｨｺｨ 02ｨｨｨｦ ｨｪｨ 0609ｨｪ ｨ 08ｨ ｨｪｨｪ ｨｴ0209ｨｰ09 ｨｬ0603ｨｪ ｨｮ ﾂｨｨｨｰｨｹ 00ｨ 08ｨｬｨ ｨｺ060002ｨｪｨｪ ｨ 08ｨ ｨｰ04 03ｨｨ0806ｨｺ ｨｨ ｨｦ09ｨｰ09ｨｨ ﾂ ｨ :03ｨ ｨｪ04, ｨｨｨｪ01ｨｮｨｺｨｰ ｨ ｨｪ0609, 03ｨｨｨｺ06ｨｪｨｨｨｪ, 0501ｨｨｨｪ ｨｬ ｨｺｨｨ ｨｰ ｨｺｨ, 09 ﾂｨ 09ｨｰｨｪ0609ｨｰｨｨ 03ｨ ｨｪ Hsp70, ｨｰｨ ｨｰ ｨｬ060502ｨｺｨｮ050708ｨｪ ｨｨ09ｨｰ02ｨｬ04, ｨｪ0206ｨ ｨｨｨｬ ｨｬ 03ｨｨ0904ｨｬ ｨｺ0502ｨｰｨｺｨ ｨｬ ｨｪ0608ｨｬｨ 05ｨｹｨｪ ｨｮｨｪｨｺ02ｨｨ06ｨｪｨｨ080609ｨ ｨｪｨｨ07 ｨｨ ｨ ｨｴｨｨｨｰ04 06ｨｰ ｨｪ02ｨ 05ｨ ｨｨ07ｨｰｨｪ ｨｦ09ｨｰ09ｨｨｨｦ ｨｺｨｮ050708ｨｪ ｨ ｨｪ04 ｨｨ0008ｨ 06ｨｰ ｨｺ0506 ﾂ0209ｨｮ ｨｹ 09 ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｬ 0006ｨｬ020609ｨｰｨ 0402, 06ｨ ﾂｨｨ09ｨ ｨ 09ｨｨ05ｨｹｨｪｨｮ06 ｨｺ06ｨｪ000608ｨｬｨ 02ｨｨ06 ｨ 0205ｨｺ0609, 06ｨｪｨｨ 04ｨ ｨｴｨｨｨｴｨ 06ｨｰ 09ｨｪ0609ｨｹ 09ｨｨｨｪｨｰ0204ｨｨ080609ｨ ｨｪｨｪ ｨｨ070207ｨｰｨｨ ｨｰ ｨｬｨｨ080609ｨ ｨｪｨｨ07 ｨｪ020708ｨ 09ｨｨ05ｨｹｨｪ06ｨｦ 09ｨｰ08ｨｮｨｺｨｰｨｮ0804, 0706ｨｬ0600ｨ 06ｨｰ 0906ｨ ｨｨ08ｨ ｨｰｨｹ ｨｨ 08ｨ 04ｨ ｨｨ08ｨ ｨｰｨｹ ｨｪｨ 01ｨｬ060502ｨｺｨｮ050708ｨｪ0402 ｨｺ06ｨｬ070502ｨｺ0904, 0906ｨ 08ｨ ｨｪｨｪ0402 ｨｨ ｨｪｨｪ0401 ｨ 0205ｨｺ0609 (Hartl, Hayer-Hartl, 2002; Deuerling, Bukau, 2004; Bukau et al., 2006). 01ｨ 08ｨｮ0302ｨｪｨｨ02 ｨｺ06ｨｪ000608ｨｬｨ 02ｨｨｨｨ ｨ 0205ｨｺ0609 ｨｨ ｨｨ01 ｨ ｨ 02ｨｨ ｨｰ ｨｨ ﾂｨｨｨｪ06ｨｦ ｨ 04ｨ ｨ ｨ ｨｪｨｨｨｦ (Perutz, 1999; Chiti, Dobson, 2006). 0507ｨｨ0906ｨｺ ｨｺ06ｨｪ000608ｨｬｨ 02ｨｨ06ｨｪｨｪ0401 ｨ ｨｪ02ｨｦ 09ｨｺ0506 ﾂｨ 02ｨｰ ｨ 07 ｨｰｨ ｨｺｨｨ02 07ｨ ｨｰ ｨｨｨｨ, ｨｺｨ ｨｺ ｨ ｨｪｨｹ 0102ｨｪｨｰｨｨｨｪ00ｨｰ06ｨｪｨ, ｨ ｨｪｨｹ 0802ｨｪｨｪ0201ｨｨ, 0907ｨｨｨｪ ｨ 08ｨ 05ｨｹｨｪ0402 ｨ ｨｰｨ ｨｺ09ｨｨｨｨ, ｨ ｨｪｨｹ 03ｨ 08ｨｺｨｨｨｪ0906ｨｪｨ, ｨ ｨｪｨｹ 0805ｨｹ020002ｨｦｨｬ0208ｨ, ｨ ｨｬｨｨ06ｨｰ080600ｨｨ ﾂ0209ｨｺｨｨｨｦ 05ｨ ｨｰ0208ｨ 05ｨｹｨｪ04ｨｦ 09ｨｺ ｨｨ 0708ｨｨ06ｨｪｨｪ ｨ ｨｰ ｨｨｨｨ. 0308ｨｨ ｨｪｨｨｨｨ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ ｨｬｨｮｨｰｨ ｨｪｨｰｨｪ0401 ｨｨ05ｨｨ ｨｪｨｪ0401 ｨ 0205ｨｺ ｨｺ0502ｨｰｨｺ02 ｨｨ ｨｨ0006ｨｬ ｨｰｨ ｨｪ060907ｨｰ0907 ｨｰ06ｨｺ09ｨｨ ﾂｨｪ04ｨｬｨｨ ｨｨ05ｨｨ 06ｨ 08ｨ 04ｨｮ06ｨｰ ｨ ｨ ｨｰ04, ｨｺ06ｨｰ , ｨｪｨ ｨｹ 09ｨｪｨｮｨｰ08ｨｨ ｨｨ05ｨｨ 09 ｨｪ ｨｰ0902ｨｪｨｪ06ｨｦ ｨ 05ｨｨ040609ｨｰｨｨ ｨｺ ｨｺ0502ｨｰｨｺ02, ｨｬ0600ｨｮｨｰ ｨ ｨｰｨｹ 0202 ｨｨ05ｨｨ ｨｰ0608ｨｬ0604ｨｨｨｰｨｹ 09ｨｪｨｮｨｰ08ｨｨｨｺ0502ｨｰ06 ﾂｨｪ04ｨｦ ｨｰ08ｨ ｨｪ ｨｰ, ｨｬ02ｨｰｨ ｨ 0605ｨｨ04ｨｬ ｨｨ 0108ｨｮ00ｨｨ ｨ ｨｪｨｨ04ｨ 02ｨｨ07 ｨｺ0502ｨｰ06 ﾂｨｪ ｨｨ090601ｨｨｨｰ ｨｺ 00ｨｨｨ 0205ｨｨ ｨｮ050702ｨｨｨｦ ｨｺ0502ｨｰ06ｨｺ ｨｨ (ｨｨ05ｨｨ) ｨｨ01 01ｨｨ0900ｨｮｨｪｨｺ02ｨｨｨｨ. 08ｨ 0301ｨ 07 ｨｨ04 ｨｮｨｺｨ 04ｨ ｨｪｨｪ ｨ ｨｰ ｨｨｨｦ ｨ 03ｨ 02ｨｰ ｨｪｨｪ04ｨｦ ｨｺ05ｨ 0909 ｨｪ02ｨｦ0806ｨｪ0609 ｨｨ ｨｰ0907, ｨｺｨ ｨｺ 0708ｨ 09ｨｨ0506, ｨｨ0506ｨｬ ｨ 09ｨｰ ｨｬ02 ｨｰ060006, ｨｰｨｬｨｨｨｰ06ｨｰｨｨ ﾂ0209ｨｺｨｨ01 ｨｺ0502ｨｰｨｺｨ ｨｨ ｨｦ, ｨｪ02 09ｨｰ08ｨ 01ｨ 06ｨｴｨｨ01 ｨｮｨｺｨ 04ｨ ｨｪｨｪ04ｨｬｨｨ 07ｨ ｨｰ ｨｨ07ｨｬｨｨ, ｨｰｨ ｨｺ0302 ｨｪｨ ｨｺｨ 0705ｨｨ09ｨ 06ｨｰ0907 ｨ 0205ｨｺ ｨ ｨ ｨｰ04, ｨｺ06ｨｰ ｨｨ ｨｰ ｨｺ ｨｪｨ 08ｨｮ0302ｨｪｨｨ06 ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｦ 00ｨｨ04ｨｨ ｨｨｨｨ (Morimoto, Guervo, 2009) ｨ ｨｪｨｨｨｨ ｨｺ0502ｨｰｨｺｨｨ 0209ｨｰｨｹ ｨｪ0209ｨｺ0605ｨｹｨｺ06 04ｨ ｨｴｨｨｨｰｨｪ ｨｨ09ｨｰ02ｨｬ, 0708ｨｨ0409ｨ ｨｪｨｪ ｨｨｨｺ09ｨｨ01ｨｨ080609ｨ ｨｰｨｹ ｨ ｨ ｨｰ04 ﾂ ｨｰ020605ｨｨ ｨｰ0902ｨｰ09ｨｰ09ｨｮ06ｨｴｨｨ01 ｨ 0205ｨｺ0609 ｨｨ05ｨｨ ｨｪ ｨｮ09ｨｺｨ ｨｰｨｹ ｨｨ01 06ｨ 08ｨ ｨ ｨｪｨｨ07 (Wyttenbach, 2004). 01ｨ ｨｨｨ ｨｬ06ｨｴｨｪ06ｨｦ ｨｨ04 ｨｰｨ ｨｺｨｨ01 09ｨｨ09ｨｰ02ｨｬ ｨｰ ｨ ｨｪｨｪ04ｨｦ ｨｬ0201ｨ ｨｪｨｨ04ｨｬ, 0609ｨｪ0609ｨ ｨｪｨｪ04ｨｦ ｨｪｨ ｨ 0205ｨｺｨ 01 ｨｺ05ｨ 0909ｨ Hsp70, 09ｨｺ0506 ﾂｨ 07 ｨｺ0603ｨ ｨｪ04 Hdj1, Bag-1 ｨｨ (00ｨ 0800ｨｮ05ｨｨ09, 01ｨｮ030609ｨ, 2009) ｨｬｨｪ06ｨｬ ｨｺ0605ｨｨ ﾂ0209ｨｰ ｨ ｨ 06ｨｰ 0706ｨｺｨ 04ｨ ｨｪ06, ﾂｨｰ06 ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 05ｨｺ ｨｨｨｨ Hsp70 ｨｨ Hdj1 09 ｨｺ0502ｨｰｨｺｨ ｨｨ090601ｨｨｨｰ ｨｺ 09ｨｪｨｨ0302ｨｪｨｨ06 ﾂｨｨ0905ｨ ｨ ｨ ｨｰ0609 ｨｨ ｨｨ01 08ｨ 04ｨｬ , ｨ ｨｪｨｨ ｨ ｨ ｨｰ0604ｨ ｨｨ ｨｪｨｨ06 03ｨｨ04ｨｪ ｨ ｨｪ0609ｨｰｨｨ (Zhou et al., 2001; Wyttenbach et al., 2001; Sakahira et al., 2002). 0201ｨｪｨ ｨｺ06 09 ｨｺ0502ｨｰｨｺｨ ｨｦ ｨｨ ｨ 09ｨｰｨ ｨｨ0601, ｨｺ060001ｨ ｨｨ ｨｨｨｰ ｨｪｨ 08ｨｮ0302ｨｪｨｨ02 ｨｺ0502ｨｰ06 ﾂｨｪ ｨｬ020609ｨｰｨ 04ｨ ｨｨ ｨｰ0907 ｨ 0205ｨｺ ｨ ｨ ｨｰ04, 09ｨｨｨｪｨｰ ｨ ｨｪ ｨｪｨｨ0302ｨｪ ｨｨ ｨｨ01 ｨｨ ｨｪ0602 ｨｺ0605ｨｨ ﾂ0209ｨｰ ｨｺ0502ｨｰｨｺ02 ｨｪ ｨｺ06 (Soti, Csermely, 2002) ｨｰ06ｨｬｨｮ 0706ｨｨ09ｨｺ 0902ｨｴ0209ｨｰ09, ｨ ｨｪ ｨｨｨｰｨｹ ｨｮ ｨｪｨｹ 05ｨｺ ｨｨｨｨ 03ｨ ｨｪ0609 ｨｨ, ｨｬ0603ｨｪ06, ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ 08ｨ ｨ 06ｨｰ04 03ｨ ｨｪｨｪ ｨｬ0201ｨ ｨｪｨｨ04ｨｬｨ, ｨｰｨ ｨｰ ﾂ02ｨｪｨｹ 09ｨ 03ｨｪ04ｨｬ ｨｺｨ ｨｺ 09 00ｨｮｨｪ01ｨ ｨｬ02ｨｪｨｰｨ 05ｨｹｨｪ06ｨｦ, ｨｰｨ ｨｺ ｨｨ ｨ ｨｺｨｰｨｨ ﾂ0209ｨｺ06ｨｦ ｨｰ06 ﾂｨｺｨｨ ｨｪｨｨ07. 05ｨ 01ｨ ﾂｨｨ ｨｪｨ 09ｨｰ0607ｨｴ ｨｨ ｨ ｨｪｨｨ07 ｪ 09ｨｺ08ｨｨｨｪｨｨｨｪ00 ｨｨ ｨ ｨｪｨ 05ｨｨ ｨｴ0209ｨｰ09, ｨ ｨｪ0401 ｨｨｨｪ01ｨｮ02ｨｨ080609ｨ ｨｰｨｹ 05ｨｺ ｨｨ06 Hsp70, ｨｪ ｨ ｨｨ 05ｨｰ06ｨｬ ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｦ 00ｨｨｨ 0205ｨｨ ｨｨ ｨｹ040609ｨ 05ｨｨ ｨ ｨｨｨ 05ｨｨ06ｨｰ02ｨｺｨｮ ｨｪｨｨ04ｨｺ06ｨｬ060502ｨｺｨｮ050708ｨｪ ｨｴ0209ｨｰ09 06ｨｨ010606ｨｺ02ｨ ｨｪ09ｨｺ

2 ｨｬ02ｨｪｨｺ06, ｨｪｨｰｨｨｨｬ06ｨｪ0609ｨ ｨｨ ｨｨｨｪ09ｨｰｨｨｨｰｨｮｨｰｨ ｨ ｨｨ ｨ ｨｪｨｨ ﾂ0209ｨｺ06ｨｦ 01ｨｨｨｬｨｨｨｨ , 09 ｨｺ06ｨｰ0608ｨｮ ｨｨ ｨｮ06ｨｴｨｨ ｨｮ ｨｴ0209ｨｰ09: ｨｰ08ｨｨｨｰ ｨｪ04 01ｨ ｨｬｨｬｨ 08ｨ ｨｪ ｨ ｨｨ ｨｨ ｨｨｨｺ0604ｨｨ0104; 0002ｨｰ ｨｨｨｺ05ｨｨ ﾂ0209ｨｺｨｨ ｨｨ080601ｨｪ0402 ｨ 05ｨｺｨ 0506ｨｨ0104 ｨｨ04 ｨｬ060809ｨｺｨｨ ｨ ｨｪｨｨ04ｨｬ0609 ｨｨ ｨｨ ｨｨ ｨｪ0402 ｨｨ 0708ｨｨ080601ｨｪ0402 ｨｨ 09ｨｨｨｪｨｰ02ｨｰｨｨ ﾂ0209ｨｺｨｨ02 1, 4-ｨｪｨ 00ｨｰ0601ｨｨｨｪ06ｨｪ04 ｨｨ ｨｨ ｨｨ S-0005ｨｨｨｺ0604ｨｨ ｨ ｨ ｨｪｨ 05ｨｨ04ｨｨ080609ｨ ｨｪ ｨｨｨｪ02ｨｪｨｨｨｦ ｨｨ ｨｪ ｨｴ0209ｨｰ09ｨ, ｨ ｨｪ ｨ ｨｰｨｹ ｨｪｨｨ02 ｨｮ080609ｨｪ07 Hsp70 09 ｨｺ0502ｨｰｨｺｨ 01 ﾂ ｨｺｨ. 0004ｨ 08ｨ ｨｪｨｪ ｨｴ0209ｨｰ09ｨ 06ｨｰｨｪ060907ｨｰ0907 ｨｺ 0008ｨｮ ｨｪｨ 00ｨｰ0601ｨｨｨｪ06ｨｪ0609, ｨｰ06 ﾂｨｪ0202, ｨｺ ｨｨ ｨｪ04ｨｬ 03ｨｨｨｺ06ｨｪｨｨｨｪｨ ｨｨ 0501ｨｨｨｪ ｨｬｨ. 00ｨ ｨｰ0208ｨｨｨ 05 ｨｨ ｨｬ02ｨｰ0601ｨｨｨｺｨ ｨｰ06 ﾂ ｨｪ0402 ｨｺｨｮ05ｨｹｨｰｨｮ ｨｰｨｺｨｨ ｨｪｨｨ ﾂ0209ｨｺ06ｨｦ ｨｬｨｨ ｨｪｨｪ06ｨｦ 0502ｨｦｨｺ02ｨｬｨｨｨｨ ﾂ ｨｺｨ K562 ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ 05ｨｨ ｨｪｨ RPMI-1640, ｨｺ0502ｨｰｨｺｨｨ 0005ｨｨ06ｨ 05ｨ 09ｨｰ06ｨｬ04 ﾂ ｨｺｨ T98G ｪ ｨｪｨ , ｨ ｨｴｨｨ01 10 % 05ｨｬｨ 08ｨｨ06ｨｪｨ 05ｨｹｨｪ06ｨｦ ｨｺ ｨｹ02ｨｦ ｨｰｨｺｨｨ ｨｨ 50 ｨｬｨｺ00/ｨｬ ｨｪｨｰｨ ｨｬｨｨ02ｨｨｨｪｨ, 0708ｨｨ 37 縺05 09 ｨ ｨｰｨｬ % 05O ｨｨｨｪｨｺｨｮｨ ｨ ｨｰ (Sanyo, ｨｪｨｨ07) ｨｰｨｺｨｨ T98G ｨ 05ｨｨ 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ ｨｺ05/ｨｬ05 04ｨ 1 09ｨｮｨｰ ｨｺ ｨｨｨｬ02ｨｪｨｰｨ, ｨｺ0502ｨｰｨｺｨｨ K562 ｪ ｨｪ ｨｰ0902ｨｪｨｪ ｨｺ ｨｨｨｬ02ｨｪｨｰ06ｨｬ 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ ｨｺ05/ｨｬ ｨｰ0609 ｨｨ ｨｹ040609ｨ 05ｨｨ ｨｺｨｮ05ｨｹｨｰｨｮ0804, ｨｬ0208ｨｰ ｨｺ0502ｨｰ06ｨｺ 09 ｨｺ06ｨｰ ｨｪ ｨ 05ｨ 5 %. 0806ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ06 ｨｺ0502ｨｰ06ｨｺ ｨｨ ｨｬ06ｨｴｨｹ06 ｨｺｨ ｨｬ ｨｹ06ｨ ｨ ｨｮ0508ｨ ｨｮ02ｨｬ ｨ 08ｨ ｨｰ04 08ｨ 09ｨｰ ｨｨ ｨｨ ｨｪｨｪ0401 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ ｨｪ0609ｨｨ05ｨｨ 09 ｨｺｨｮ05ｨｹｨｰｨｮ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨ 04 09ｨｪ0609ｨｨ05ｨｨ (1 ｨｬｨｺ05/ｨｬ05) ｨｰ09ｨｨｨｨ ｨｮｨｺｨ 04ｨ ｨｪｨｪ ｨｬ02ｨｪｨｨ ｨｺ0502ｨｰｨｺｨｨ 05ｨｨ04ｨｨ080609ｨ 05ｨｨ 09 ｨ ｨｮ000208ｨｪ06ｨｬ 08ｨ 09ｨｰ RIPA (20 ｨｬM Tris-HCl, ph 7.5, 150 ｨｬM NaCl, 2 ｨｬM EDTA ｨｨ 0.5 % Triton X-100), ｨｨ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ06 ｨ 0205ｨｺｨ 09 05ｨｺ09ｨｰ08ｨ ｨｺｨｰｨ 01 (Bradford, 1976) ｨｨ ｨｬｨｺ00 ｨ 0205ｨｺｨ ｨｨ04 ｨｺｨ ｨｦ ｨ 04 ｨｨ ｨｹ040609ｨ 05ｨｨ ｨｨ0006ｨｰ ｨｪｨｨ ｨｺｨｰ ｨｪ ｨ, ｨ 0609ｨｰｨ 05ｨｹｨｪ0602 ｪ ｨｨ04ｨｬ020802ｨｪｨｨ07 03ｨ ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ Hsp ｨｬ06ｨｴｨｹ ｨｨｨ 05ｨｹｨｪ06ｨｦ ｨｨｨｬｨｬｨｮｨｪ ｨｬ02ｨｪｨｰｨｪ06ｨｦ ｨｰ0209ｨｰ-09ｨｨ09ｨｰ02ｨｬ04 (09ｨｬ. ｨｪｨｨ0302). 06ｨｬｨｬｨｮｨｪ06ｨ 0506ｨｰｨｨｨｪ ｨｺｨｨ 08ｨ ｨｨ ｨｬ02ｨｰ060106ｨｬ SDS ｨｺｨｰ ｨ 09 10%-ｨｪ06ｨｬ ｨｨｨ ｨｺ08ｨｨ05ｨ ｨｬｨｨ01ｨｪ06ｨｬ ｨｪ ｨｺｨｰ ｨｰｨｨ ﾂ0209ｨｺｨｨ 08ｨ ｨｪｨｪ0401 ｨ 0205ｨｺ0609 ｨｪｨ ｨｪｨｨｨｰ ｨｪｨｮ06 ｨｬ02ｨｬｨ 08ｨ ｨｪｨｮ Whatman (Sigma-Aldrich, ) ｨｨ ｨｨ05ｨｨ 0708ｨｨ 0706ｨｬ06ｨｴｨｨ ｨ 0707ｨ 08ｨ ｨｰｨ Trans-Blot (Bio-Rad, ). 0506ｨｪｨｮ ｨ 0205ｨｺｨ Hsp ｨｨ ｨｬ06ｨｴｨｹ ｨｨ00ｨｨ ﾂｨｪ0401 ｨｬ06ｨｪ06ｨｺ0506ｨｪｨ 05ｨｹｨｪ0401 ｨ ｨｪｨｰｨｨｨｰ0205 (ｨｺ0506ｨｪ 3005, ｨｨ ｨｪ ｨ ｨ 0608ｨ ｨｰ0608ｨｨｨｨ) 09 08ｨ ｨｪｨｨｨｨ 1 : 1000 ｨｨ 09ｨｰ ｨ ｨｪｨｰｨｨｨｰ0205, ｨｺ06ｨｪｨｲ0600ｨｨ080609ｨ ｨｪｨｪ ｨｺ09ｨｨ01ｨ 0406ｨｦ ｨｪｨ, 09 08ｨ ｨｪｨｨｨｨ 1 : (Sigma, ) ｨｨ04ｨｮｨ 05ｨｨ04ｨ 02ｨｨｨｨ ｨｺ09ｨｨ01ｨ 04ｨｪ06ｨｦ 0802ｨ ｨｺ02ｨｨｨｨ ｨｨ ｨｹ040609ｨ 05ｨｨ ｨｬ02ｨｰ0601ｨｨｨｺｨｮ ｨｮ09ｨｨ0502ｨｪｨｪ06ｨｦ 0102ｨｬｨｨ0506ｨｬｨｨｨｪ ｨｪ02ｨｨｨｨ ｨ 09ｨｪ06 ｨｨｨｪ09ｨｰ08ｨｮｨｺ02ｨｨｨｨ 00ｨｨ08ｨｬ ｨｨ ｨｨｨｰ (Amersham, ｨｨｨｺ06ｨ 08ｨｨｨｰｨ ｨｪｨｨ07) ｨｪｨｨ02 09ｨｮｨ 09ｨｰ08ｨ ｨｰ ｨ 06ｨｴ02ｨｦ (03ｨ ｨｪｨｪ06ｨｦ) ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ Hsp ｨｨ05ｨｨ ｨｬ06ｨｴｨｹ ｨｬ0601ｨｨ00ｨｨ02ｨｨ080609ｨ ｨｪｨｪ06ｨｬｨｮ ｨｬ ｨ 09ｨｰ0608ｨ ｨｬｨｨ (2004) ｨｬ02ｨｰ0601ｨｮ (Wawrzynow, Zylicz, 1995). 05ｨｮｨ 09ｨｰ08ｨ ｨｰｨｪ04ｨｦ ｨ ｨｺ-ｨｬｨｨ0302ｨｪｨｹ, ｨｺｨ 08ｨ 06ｨｺ09ｨｨｨｬ02ｨｰｨｨ05ｨｨ080609ｨ ｨｪｨｪ04ｨｦ 05ｨ ｨｺｨｰｨ 05ｨｹｨ ｨｮｨｬｨｨｨｪ ( ), ｨｮ ﾂｨ 05ｨｨ 07ｨｮｨｰ02ｨｬ 0102ｨｪｨ ｨｰｨｮ08ｨ 02ｨｨｨｨ 05ｨ ｨｺｨｰｨ 05ｨｹｨ ｨｮｨｬｨｨｨｪｨ 09800ｨｬ06 ﾂ0209ｨｨｨｪ02, ﾂ ｨ 09ｨｰ ｨ 0205ｨｺｨ 0106ｨ ｨ ｨｨ 01ｨｨｨｰｨｨ06ｨｰ0802ｨｨｨｰ ｨｺ06ｨｪ02 ﾂｨｪ06ｨｦ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 10 ｨｬ00. 02ｨ ｨ ｨｺ02ｨｨ06 ｨ 05ｨｺｨｨ05ｨｨ080609ｨ ｨｪｨｨ07 04ｨ 07ｨｮ09ｨｺｨ 05ｨｨ 09ｨｪ020902ｨｪｨｨ02ｨｬ 09 08ｨ 09ｨｰ ｨ 0205ｨｺｨ ｨｨ060106ｨ 0202ｨｰｨ ｨｬｨｨ01ｨ ｨｨ 0609ｨｰｨ ｨｪｨ 0905ｨｨ09ｨ 05ｨｨ 0202 ﾂ ﾂ 0106ｨ ｨ ｨｪｨｨ02ｨｬ 10 ｨｬ00 01ｨｨｨｰｨｨ06ｨｰ0802ｨｨｨｰ0605ｨ. 0106ｨｰ060904ｨｦ ｨ 04ｨｪ0609ｨｨ05ｨｨ ｨｬｨｺ ｨｮｨｪｨｺｨｨ 96-05ｨｮｨｪ06 ﾂｨｪ ｨ ｨｪ0302ｨｰｨ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 10 ｨｬｨｺ00/ｨｬ ｨ ｨｰｨｪ ｨｬ 08ｨ 09ｨｰ (080504). 05ｨ ｨｦｨｰ04 ｨｪ ｨｨ00ｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨ 0506ｨｺｨｨ080609ｨ 05ｨｨ 0.3%-ｨｪ04ｨｬ 08ｨ 09ｨｰ ｨｬ ｨ 04 ﾂｨｹ ｨｰ06 ﾂｨｪ ｨ 05ｨｹｨ ｨｮｨｬｨｨｨｪｨ (Sigma, ) ｨ ｨｮｨｪｨｺｨｨ 09ｨｪ0609ｨｨ05ｨｨ ｨｺ0502ｨｰ06 ﾂｨｪ ｨｺ09ｨｰ08ｨ ｨｺｨｰ04 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ0701 (ｨｰｨｨｨｰ08ｨ 01) 5 ｪ50 ｨｬｨｺ00/ｨｬ ｨｬｨｺ ｨｺ Hsp70, ｨｴｨｨｨｦ ｨｰｨ 09 ｨｺ0502ｨｰ06 ﾂｨｪ ｨｺ09ｨｰ08ｨ ｨｺｨｰ0609, ｨ ｨｪｨ ｨｰｨｮ08ｨｨ080609ｨ ｨｪｨｪ04ｨｬ ｨ 0205ｨｺ06ｨｬ-ｨｬｨｨ0302ｨｪｨｹ ｨｪｨｨ02 03ｨ ｨｪｨ 0609ｨｮｨｴ0209ｨｰ ｨｨ ｨｬ06ｨｴｨｹ ｨｨ00ｨｨ ﾂ0209ｨｺｨｨ01 ｨ 0000ｨｨｨｪｨｪ0606 ﾂｨｨｨｴ02ｨｪｨｪ ｨｨｨｺ0506ｨｪｨ 05ｨｹｨｪ0401 ｨ ｨｪｨｰｨｨｨｰ0205 RAF ｨｰｨｨ09 Hsp70, ｨｮ ﾂ02ｨｪｨｪ ｨ ｨ 0608ｨ ｨｰ0608ｨｨｨｨ, 09 08ｨ ｨｪｨｨｨｨ 1 : 1000 ｨｨ 09ｨｰ ｨ ｨｪｨｰｨｨｨｰ0205, ｨｺ06ｨｪｨｲ0600ｨｨ080609ｨ ｨｪｨｪ ｨｺ09ｨｨ01ｨ 0406ｨｦ ｨｪｨ, 09 08ｨ ｨｪｨｨｨｨ 1 : (Sigma, ) ｨｬ02ｨｪｨｰｨ ｨｰｨｨ09ｨｪｨｮ ｨ ｨｺ02ｨｨ ｨｨ05ｨｨ 07ｨｮｨｰ02ｨｬ 09ｨｪ020902ｨｪｨｨ ｨｮｨｪｨｺｨｨ 08ｨ 09ｨｰ090608ｨ, ｨ ｨｴ ｨｺｨｨ09ｨｹ ｨ ｨｨ ｨｬ060002ｨｪ ｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ 06ｨｺ08ｨ 09ｨｺｨｨ ｨｨ04ｨｬ ｨｨ 0708ｨｨ 0105ｨｨｨｪ ｨｪ ｨｪｨｬ ｨｬ06ｨｴｨｹ06 ｨ ｨｪｨ 05ｨｨ04ｨ ｨｰ0608ｨ ｨｨｨｬｨｬｨｮｨｪ0601ｨｨｨｬｨｨ ﾂ0209ｨｺｨｨ ｨ ｨｺ02ｨｨｨｦ ｨｰ03 ( ｨ ｨ ｨｪｨ ｨｮ ﾂ0708ｨｨｨ 0608, ｨｨ07) ｨｰｨ ﾂ ｨ 05ｨｨ ｨｰ080201ｨｺ08ｨ ｨｰｨｪ06ｨｦ ｨｬ0409ｨｺ06ｨｦ 05ｨｮｨｪ06ｨｺ; 09 ｨｺｨ ﾂ0209ｨｰ ｨ ｨ 06 ﾂ ｨｨ ｨｬ0409ｨ 06ｨｴ ｨｨ ｨｹ040609ｨ 05ｨｨ ｨ ｨｮ000208ｨｪ04ｨｦ 08ｨ 09ｨｰ (20 ｨｬM Tris-HCl, ph 7.5, 20 ｨｬM NaCl, 10 ｨｬM MgCl 2 ｨｨ 0.05%-ｨｪ04ｨｦ Triton X-100) ｨｪｨｨ02 03ｨｨ04ｨｪ ｨ ｨｪ0609ｨｰｨｨ ｨｺ0502ｨｰ06ｨｺ 0609ｨｮｨｴ0209ｨｰ ｨｨ ｨｬ02ｨｰ060106ｨｬ ｨｰ0902ｨｰ09ｨｰ09ｨｨｨｨ ｨｺ06ｨｬ02ｨｪ01ｨ 02ｨｨ07ｨｬｨｨ 00ｨｨ08ｨｬ ｨｨ ｨｨｨｰ (Molecular Probes, ) ｨｮｨｪｨｺｨｨ 96-05ｨｮｨｪ06 ﾂｨｪ ｨ ｨｪ0302ｨｰｨ 09ｨｪ0609ｨｨ05ｨｨ 0601ｨｪ ｨｬ02ｨｪｨｪ ｨｬｨｺ05 ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｦ 09ｨｮ090702ｨｪ04ｨｨｨｨ 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ02ｨｦ ｨｺ0502ｨｰ06ｨｺ ｨｺ05./ｨｬ05 ｨｨ 08ｨ 09ｨｰ ｨ 0405ｨｨ ﾂｨｪ04ｨｬｨｨ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ07ｨｬｨｨ ｨｨ ｨｮ02ｨｬ ｨｨｨｪ02ｨｪｨｨｨｦ, 09 ｨｺ06ｨｰ060804ｨｬｨｨ ｨｺ0502ｨｰｨｺｨｨ ｨｨｨｪｨｺｨｮｨ ｨｨ080609ｨ 05ｨｨ 09 ｨｰ02 ﾂ02ｨｪｨｨ02 6 ﾂ. 00 ｨｰ ｨｮｨｪｨｺｨｨ ｨｰ09ｨｨｨｨ ｨｮｨｺｨ 04ｨ ｨｪｨｪ ｨｬ02ｨｪｨｨ 09ｨｪ0609ｨｨ05ｨｨ ｨｨｨｪ01ｨｮｨｺｨｰ ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｦ 00ｨｨｨ 0205ｨｨ 09ｨｰｨ ｨｮ ｨｨｨｪ 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 1 ｨｬｨｺ00 ｨｨ05ｨｨ H 2 O 2 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 500 ｨｬｨｺ00 ｨｪｨ 18 ﾂ ｨｨ05ｨｨ ｨ 05ｨｨ ｨｺ0502ｨｰｨｺｨｨ ｨｰｨｺ06ｨｬｨｮ03 ｨｰ ｨｬｨｮ 0306ｨｺｨｮ (45 縺05, 30 ｨｬｨｨｨｪ). 00 ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨ 04 09ｨｪ0609ｨｨ05ｨｨ 1 ｨｬｨｺ05/ｨｬ ｨｮｨｪｨｺｨｨ 0106ｨ ｨ ｨｨ ｨｬｨｺ05 ｨｰ08ｨｨｨ ｨｨ05ｨｰ02ｨｰ08ｨ ｨｨｨｦｨ 0806ｨｨｨｬｨｨ01ｨ 09ｨｨｨｪ (000606), 08ｨ 09ｨｰ ｨｪｨｪ ( /ｨｬ05), ｨｨ ｨｨｨｪｨｺｨｮｨ ｨｨ080609ｨ 05ｨｨ 0708ｨｨ 37 縺05 ｨｨ 5 % ﾂ. 05ｨ ｨｰ02ｨｬ 09 05ｨｮｨｪｨｺｨｨ 09ｨｪ0609ｨｨ05ｨｨ ｨｬｨｺ05 ｨｨ ｨｨ ｨｨ08ｨｰｨ M H05l, ｨｨｨｬ ｨｮｨｪ06ｨｺ ｨｬ0203ｨｨ09ｨ 05ｨｨ ｨ 09ｨｰ ｨｪｨｨ07 06ｨ 08ｨ ｨ ｨ 01ｨｺｨ ｨｬｨ 04ｨ ｨｪｨ ｨｨ ｨｨ04ｨｬ ｨｨ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ 06ｨｺ08ｨ 09ｨｺｨｨ ｨｬ06ｨｴｨｹ06 ｨ ｨｪｨ 05ｨｨ04ｨ ｨｰ0608ｨ ｨｨｨｬｨｬｨｮｨｪ0601ｨｨｨｬｨｨ ﾂ0209ｨｺｨｨ ｨ ｨｺ02ｨｨｨｦ ｨｰ ｨｨ 0105ｨｨｨｪｨ ｨｪ 570 ｨｨ 630 ｨｪｨｬ ｨｮ05ｨｹｨｰｨ ｨｰ04 01ｨ ｨｬ 05ｨｰｨ ｨｬ06ｨｴｨｹ06 ｨｨｨｬｨｬｨｮｨｪ06ｨ 0506ｨｰｨｨｨｪ00ｨ ｨ ｨｨ ｨｪ 09ｨｺ08ｨｨｨｪｨｨｨｪ ｨｨｨｪ02ｨｪｨｨｨｦ ｨｹ06 ｨｪｨ ｨｦｨｰｨｨ 0902ｨｴ0209ｨｰ09ｨ, ｨ ｨｪ ｨ ｨｰｨｹ ｨｪｨｨ02 05ｨｺ ｨｨｨｨ 03ｨ ｨｪｨ Hsp70, ｨｪ02ｨｰ06ｨｺ09ｨｨ ﾂｨｪ ｨｺｨｨ01 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ ｨｺ0502ｨｰ06ｨｺ ｨｨ ﾂｨｪ04ｨｦ 09ｨｺ08ｨｨｨｪｨｨｨｪ ｨｨ05ｨｨ 09 ｨｨ ｨｹ040609ｨ ｨｪｨｨ02ｨｬ ｨｺ0502ｨｰ06ｨｺ 0005ｨｨ06ｨ 05ｨ 09ｨｰ06ｨｬ04 ﾂ ｨｺｨ T98G ｨ ｨｪｨ 05ｨｨ04ｨｨ080609ｨ ｨｪ06 ｨ ｨｨｨｪ02ｨｪｨｨｨｦ, ｨｺｨ ｨｨ04 ｨｺ06ｨｰ ｨｨ ｨｹ040609ｨ 05ｨｨ 09 08ｨ ｨ 06 ﾂ02ｨｦ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 10 ｨｬｨｺ ｨｪ ｨｬ02ｨｪｨｪ ｨｬ06ｨｴｨｹ ｨｪｨｨ09ｨ 05ｨｨ 02ｨｨｨｰ06ｨｰ06ｨｺ09ｨｨ ﾂ0209ｨｺｨｨｨｦ ｨｺｨｰ 0902ｨｴ0209ｨｰ09, ｨｰ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ0701 1, 10 ｨｨ 100 ｨｬｨｺ00 (08ｨｨ09. 1, ｨ ). 01ｨ 08ｨｨ09. 1, ｨ ｨｰｨ ｨｪ ｨｮ05ｨｹｨｰｨ ｨｰ ｨｨｨｬｨｬｨｮｨｪ06ｨ 0506ｨｰｨｨｨｪ00ｨ 05ｨｨ04ｨ ｨｰ0609 ｨｺ0502ｨｰ06ｨｺ T98G, ｨｨｨｪｨｺｨｮｨ ｨｨ080609ｨ ｨｪｨｪ ｨｪ02ｨｺ06ｨｰ060804ｨｬｨｨ ｨｨ04 ｨｨ ｨｮ02ｨｬ ｨ 08ｨ ｨｰ ｨｨｨｪ02ｨｪｨｨｨｦ ｨｮ05ｨｹｨｰｨ ｨｰ ｨｨ ﾂｨｪ ｨｺ08ｨｨｨｪｨｨｨｪ00ｨ ｨ 0405ｨｨ 06ｨｰ06ｨ 08ｨ ｨｪ ｨ ｨ 08ｨ ｨｰｨ ｪ U-133, ｨｪ02ｨｰ06ｨｺ09ｨｨ ﾂｨｪ04ｨｦ ｨｺ0502ｨｰ06ｨｺ ｨｨ ｨ 06ｨｴｨｨｨｦ ｨｮｨｬ020802ｨｪｨｪｨｮ06 05ｨｺ ｨｨ06 Hsp70 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 10 ｨｬｨｺ00, ｨｨ U-103, ｨ ｨｰ06ｨｺ09ｨｨ ﾂｨｪ04ｨｦ 0708ｨｨ ｨｺｨｨ01 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ0701 (08ｨｨ09. 1, ｨ ), ｨｪ ｨ 06ｨｴｨｨｨｦ ｨ 0302ｨｪｨｪ ｨｪｨｨ02 09ｨｨｨｪｨｰ0204ｨ 03ｨ ｨｪｨ ｨｪ ｨ 08ｨ ｨｰｨ U ｨｨｨｰ 0708ｨｨ080601ｨｪ04ｨｦ ｨｪｨ 00ｨｰｨ 04ｨ 08ｨｨｨｪ ｪ ｨｺ08ｨ 09ｨｪ04ｨｦ 07ｨｨ00ｨｬ02ｨｪｨｰ 03ｨｨｨｺ06ｨｪｨｨｨｪ, ｨｪｨｪ04ｨｦ ｨｨ04 ｨｺ0608ｨｪ02ｨｦ

3 ｨｨｨｪ02ｨｪｨｨ07, ｨ 06ｨｴｨｨ02 05ｨｺ ｨｨ06 03ｨ ｨｪｨ Hsp ｨｨ Hsp70-ｨｨｨｪ01ｨｮ02ｨｨ08ｨｮ06ｨｴｨ ｨ ｨｪ0609ｨｰｨｹ 01ｨｨｨｪ06ｨｨ01ｨｪ ｨｨｨｪ02ｨｪｨｨｨｦ. ｨ ｪ ｨ ｨｪｨ 05ｨｨ04 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ 0902ｨｴ0209ｨｰ09 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 10 ｨｬｨｺ ﾂｨ ｨｦ ｨｨｨｪｨｺｨｮｨ ｨ 02ｨｨｨｨ 09 ｨｪｨｨｨｬｨｨ ｨｺ0502ｨｰ06ｨｺ 0005ｨｨ06ｨ 05ｨ 09ｨｰ06ｨｬ04 ﾂ ｨｺｨ T98G ｨｬ02ｨｰ060106ｨｬ ｨｨｨｬｨｬｨｮｨｪ06ｨ 0506ｨｰｨｨｨｪ00ｨ ; ｨ ｪ ｨ ｨｪｨ 05ｨｨ04 02ｨｨｨｰ06ｨｰ06ｨｺ09ｨｨ ﾂ0209ｨｺ ｨｦ09ｨｰ09ｨｨ ｨ 08ｨ ｨｰ0609, 0706ｨｺｨ 04ｨ 0903ｨｨ01 03ｨ ｨｪｨｨｨｪ01ｨｮ02ｨｨ08ｨｮ06ｨｴｨｮ06 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ, ｨｬ06ｨｴｨｹ06 ｨｬ02ｨｰ0601ｨ ; 09 ｪ 01ｨｨｨｬｨｨ ﾂ0209ｨｺｨｨ ｨｬｨｮ ｨｴ0209ｨｰ09, 0706ｨｺｨ 04ｨ 0903ｨｨ01 Hsp70-ｨｨｨｪ01ｨｮ02ｨｨ08ｨｮ06ｨｴｨｮ06 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ. 08 ｪ ｨｺ06ｨｪｨｰ080605ｨｹ ｨ ｨ ｨ ｨｪｨｨ ｨｴ0209ｨｰ09. ｨｨ ｨｹ04ｨｮ02ｨｬ ｨｰ08ｨ 01ｨｨ02ｨｨ06ｨｪｨｪ06ｨｦ ｨｰ06 ﾂｨｪ06ｨｦ ｨｬ0201ｨｨ02ｨｨｨｪ02 08ｨ 09ｨｰ02ｨｪｨｨ07 Lithospermum erithrorizon Sieb. et Zucc. 01ｨ 0609ｨｪ ｨｰ ｨｨｨｺ06ｨｪｨｨｨｪｨ ｨ ｨｮ ﾂ02ｨｪ ｨｨ ｨｪ ｨｴ0209ｨｰ0906 ｪ ｨｰ02ｨｰ08ｨ -02-ｨ 0202ｨｰｨｨ05-b-D ｨｺ0607ｨｨ08ｨ ｨｪ0604ｨｨ ｨ 06ｨｺ060906ｨｦ ｨｨ ｪ U ｨｪ0609ｨｮ ｨ 08ｨ ｨｰｨ U ｨｰｨ ｨｰ 0501ｨｨｨｪ ｨｬ, 0609ｨｪ0609ｨｪ06ｨｦ 07ｨｨ00ｨｬ02ｨｪｨｰ ｨｬ060809ｨｺｨｨ ｨｦ. 0601ｨｨｨｪ ｨｬ ｨｨｨｬ0202ｨｰ 08ｨ 04ｨｪ0606ｨ 08ｨ 04ｨｪｨｮ06 ｨ ｨｨ ｨｨ ﾂ0209ｨｺｨｮ06 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ, 09 ﾂｨ 09ｨｰｨｪ0609ｨｰｨｨ 06ｨｪ ｨｰ ｨｺｨｰｨｨ09ｨｪ04ｨｬ ｨ ｨｪｨｰｨｨ06ｨｺ09ｨｨ01ｨ ｨｪｨｰ06ｨｬ. 0002ｨｴ0209ｨｰ0906, ｨｰ0602 ｨｪｨ ｨｬｨｨ ｨｨ ｨ ｨｪｨｨ07, ｨｰｨ ｨ 06ｨｦ ｨ 0202ｨｰｨｨ05ｨｨ080609ｨ ｨｪｨｪ04ｨｦ ｨｰ08ｨｨ ｨｺ0604ｨｨ ｨｨｨｪ ｨｬｨ (000608ｨｬｨｮ ｨｰｨ ｨｪ04 ｨｪｨ 08ｨｨ09. 1, 09) ｨｺ0605ｨｹｨｺｨｮ ｨ 08ｨ ｨｰ U ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ ｨｬｨｺ ｨ 05 ｨｺ0502ｨｰ06 ﾂｨｪｨｮ06 00ｨｨｨ 0205ｨｹ, ｨ 08ｨ 04ｨｪ06 ｨ ｨｨ ｨ ｨｰｨｹ, ｨ 06ｨｰ 05ｨｨ ｨ ｨｪｨｨ04ｨｺｨｨ02, ｨ 05ｨｨ04ｨｺｨｨ02 ｨｺ 00ｨ 08ｨｬｨ ｨｺ ｨｨ ﾂ0209ｨｺｨｨｨｬ, ｨ 06ｨｨ ｨ 08ｨ ｨｰ0609 ｨｨｨｪ01ｨｮｨｺ02ｨｨ06 Hsp70. 01ｨ 08ｨｨ09. 2, ｨ 09ｨｨ01ｨｪ06, ﾂｨｰ ｨ 08ｨ ｨｰ U ｨ 02ｨｰ 04ｨｪｨ ﾂｨｨｨｰ0205ｨｹｨｪ0602 ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 05ｨｺ ｨｨｨｨ 03ｨ ｨｪｨ 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 0.5 ｨｬｨｺ00 09 ｨｺ0502ｨｰｨｺｨ ｨｨｨｰ ｨｦｨｺ02ｨｬｨｨｨｨ K562 ﾂ ﾂ ｨｨｨｪｨｺｨｮｨ ｨ 02ｨｨｨｨ ｨｴ0209ｨｰ0906ｨｬ, ｨ ｨ 08ｨ ｨｰ U ｨ 05ｨ 01ｨ 02ｨｰ 03ｨ ｨｪｨｨｨｪ01ｨｮ02ｨｨ08ｨｮ06ｨｴ02ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ06 ｨｮ ｨｨ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 0.05 ｨｬｨｺ00 ﾂ ﾂ ｨｨｨｪｨｺｨｮｨ ｨ 02ｨｨｨｨ ｨ 08ｨ ｨｰ04 09 ｨｮｨｺｨ 04ｨ ｨｪｨｪ0401 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ0701 ｨｪ02ｨｰ06ｨｺ09ｨｨ ﾂｨｪ ｨｺ0502ｨｰ06ｨｺ ｨｮ02ｨｰ 06ｨｰｨｬ02ｨｰｨｨｨｰｨｹ, ﾂｨｰ06 06ｨ ｨ 0902ｨｴ0209ｨｰ09ｨ ｨ 06ｨｰ ｨｪｨｨ02 ｨｮ080609ｨｪ07 03ｨ ｨｪｨ Hsp ｨｺ0502ｨｰ06 ﾂｨｪ ｨｨｨｪｨｨ0701, ｨｺ06ｨｰ ｨｬ04 ｨｨ ｨｹ040609ｨ 05ｨｨ ｨ ｨｰｨ 05ｨｹｨｪ ｨ ｨｪｨ 05ｨｨ04ｨ : 0005ｨｨ06ｨ 05ｨ 09ｨｰ06ｨｬｨ ﾂ ｨｺｨ T98G, ｨｬｨｨ020506ｨｨ01ｨｪｨ ｨｦｨｺ02ｨｬｨｨ07 ﾂ ｨｺｨ U-937, ｨｪ02ｨｦ0806ｨ 05ｨ 09ｨｰ06ｨｬｨ ﾂ ｨｺｨ SH-SY-5Y ｨｨ 00ｨｨｨ 0806ｨ 05ｨ 09ｨｰ04 ｨｬ0403ｨｨ 3T3 (01ｨ ｨｪｨｪ0402 ｨｪ ｨｺｨ 04ｨ ｨｪ04). 0209ｨｪ0609ｨｪ06ｨｦ 01ｨ 08ｨ ｨｺｨｰ0208ｨｨ09ｨｰｨｨｨｺ06ｨｦ Hsp ｨｰ ｨ ｨｪｨｪｨ 07 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ. 06ｨｬ02ｨｪｨｪ06 09 ｨｪ02ｨｦ ｨ 06ｨｰ ｨｺ0502ｨｰ06 ﾂｨｪ ｨｮｨｪｨｺ02ｨｨｨｨ ｨ 0205ｨｺｨ, 09 ﾂｨ 09ｨｰｨｪ0609ｨｰｨｨ ｨ ｨｴｨｨｨｰｨｪｨｮ06 00ｨｮｨｪｨｺ02ｨｨ ｨｰ ﾂｨｰ06ｨ ｨｨｨｰｨｹ, ｨｪｨ 09ｨｺ0605ｨｹｨｺ06 Hsp70, 05ｨｺ ｨｨ07 ｨｺ06ｨｰ ｨ 09ｨｰｨ 02ｨｰ 09 ｨｺ0502ｨｰｨｺｨ ｨｦ09ｨｰ09ｨｨ02ｨｬ ｨｨ04ｨｮ ﾂｨ 02ｨｬ ｨ 08ｨ ｨｰ0609, ｨ ｨｺｨｰｨｨ0902ｨｪ ｨｺｨ ｨｺ 03ｨ ｨｪ, ｨｬ04 ｨｨ ｨｹ040609ｨ 05ｨｨ ｨｰ0209ｨｰ, ｨｬ0601ｨｨ00ｨｨ02ｨｨ080609ｨ ｨｪｨｪ04ｨｦ 09 ｨｪｨ 0302ｨｦ 05ｨ ｨ 0608ｨ ｨｰ0608ｨｨｨｨ ( ｨｨ 0108., 2004), ｨｺ06ｨｰ ｨｰ ｨ ｨｪ0609ｨｰｨｹ 03ｨ ｨｪｨ ｨ ｨｰｨｹ 0102ｨｪｨ ｨｰｨｮ08ｨｨ080609ｨ ｨｪｨｪ04ｨｦ ｨ ｨｺ. 00 ｨｺｨ ﾂ0209ｨｰ ｨｮｨ 09ｨｰ08ｨ ｨｰｨ 0802ｨ ｨｺ02ｨｨｨｨ ｨｨ ｨｹ040609ｨ ｨｰｨ ｨｪ ｨｪｨｪ04ｨｦ ｨｺｨ 08ｨ 06ｨｺ09ｨｨｨｬ02ｨｰｨｨ05ｨｨ080609ｨ ｨｪｨｪ04ｨｦ 05ｨ ｨｺｨｰｨ 05ｨｹｨ ｨｮｨｬｨｨｨｪ, ｨｨｨｬｨｬ06ｨ ｨｨ05ｨｨ040609ｨ ｨｪｨｪ04ｨｦ ｨｪｨ 96-05ｨｮｨｪ06 ﾂｨｪ ｨ ｨｪ0302ｨｰｨ ｨ 0903ｨｨｨｦ ｨｮｨ 09ｨｰ08ｨ ｨｰ06ｨｬ 03ｨ -

4 ｨｬ02ｨｪｨｺ06, ｨｪｨｰｨｨｨｬ06ｨｪ0609ｨ ｨｨ ｨｨ ｨｨ04ｨｺ06ｨｬ060502ｨｺｨｮ050708ｨｪ ｨ 08ｨ ｨｰ04 03ｨｨｨｺ06ｨｪｨｨｨｪ U-103 ｨｨ 0501ｨｨｨｪ ｨｬ U ｨ 06ｨｰ 09 ｨｪｨｨ04ｨｺｨｨ01 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ ｨｨｨｪｨｰ0204 ｨ ｨｺｨｰｨｨ09ｨｪ Hsp70. ｨ ｪ ｨｺ0502ｨｰｨｺｨｨ 0508ｨｨｨｰ0806ｨ 05ｨ 09ｨｰ06ｨｬ04 ﾂ ｨｺｨ K562 ｨｨｨｪｨｺｨｮｨ ｨｨ080609ｨ 05ｨｨ ｨｴ0209ｨｰ09ｨ ｨｬｨｨ U-103 ｨｨ U ｨｮｨｺｨ 04ｨ ｨｪｨｪ0401 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ ｨｰ02 ﾂ02ｨｪｨｨ02 3,6ｨｨ24 ﾂ, ﾂ ｨｺ0502ｨｰ06 ﾂｨｪ ｨｨ04ｨ ｨｰ ｨ 05ｨｨ ｨ ｨｪｨ 05ｨｨ04ｨｮ ｨｬ06ｨｴｨｹ06 ｨｨｨｬｨｬｨｮｨｪ06ｨ 0506ｨｰｨｨｨｪ00ｨ 09 ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｬｨｨ ｨｰｨｨ09 Hsp70; ｨ ｪ ｨ ｨｪｨ 05ｨｨ04 03ｨ ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ ｨｬ02ｨｰ060106ｨｬ ｪ ｨｺ06ｨｪｨｰ080605ｨｹ ｨｪ ｨｨ ｨｬ06ｨｴｨｹ ｨｨ00ｨｨ ﾂ0209ｨｺｨｨ ｨｨｨｺ0506ｨｪｨ 05ｨｹｨｪ0401 ｨ ｨｪｨｰｨｨｨｰ0205 ｨｨ 09ｨｰ ｨ ｨｪｨｰｨｨｨｰ ｨｬ02ｨｪｨｰｨ ｨｰｨｨ09ｨｪ06ｨｦ ｨｬ02ｨｰｨｺ06ｨｦ. 01ｨ 08ｨｨ09. 2, ｨ ｨｰｨ ｨｪ ｨｨ05ｨｹ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ 03ｨ ｨｪｨ Hsp70 09 ｨｺ0502ｨｰｨｺｨ 01 K562, ｨｨｨｪｨｺｨｮｨ ｨｨ080609ｨ ｨｪｨｪ ｨ 08ｨ ｨｰｨ ｨｬｨｨ 09 ｨｰ02 ﾂ02ｨｪｨｨ02 ｨｮｨｺｨ 04ｨ ｨｪｨｪ ｨｬ02ｨｪｨｨ. 01ｨ ｨｪｨｪｨ 07 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ ｨ 09ｨｰｨ 02ｨｰ ｨｬ ｨｨ06ｨｪｨ 05ｨｹｨｪ06 09 ｨｺ0605ｨｨ ﾂ0209ｨｰ0906ｨｬ ｨ 0205ｨｺｨ Hsp70, ｨｬ ｨｬ02ｨｰ060106ｨｬ ｨｨｨｬｨｬｨｮｨｪ06ｨ 0506ｨｰｨｨｨｪ00ｨ 09 ｨｰ ｨｺ0502ｨｰｨｺｨ 01 (08ｨｨ09. 2, ｨ ). 02ｨ ｨ ｨ 08ｨ ｨｰｨ ｨ 05ｨｨ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 ｨｺ0605ｨｨ ﾂ0209ｨｰ09ｨ ｨ ｨｺｨｰｨｨ09ｨｪ060006, ｨｰ ｨ ｨｪ ｨ ｨｰｨｹ 0102ｨｪｨ ｨｰｨｮ08ｨｨ080609ｨ ｨｪｨｪ04ｨｦ ｨ ｨｺ-09ｨｮｨ 09ｨｰ08ｨ ｨｰ , 0708ｨｨ ﾂ02ｨｬ 09ｨｰ020702ｨｪｨｹ 05ｨｰ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ07 04ｨ 09ｨｨ090205ｨ 06ｨｰ ｨｬ02ｨｪｨｨ ｨｦ09ｨｰ09ｨｨ07, ｨ ｨｬｨ ｨｺ09ｨｨｨｬｨｮｨｬ ｨｨ ﾂｨｨｨｪ04 ｨ ｨ ｨｰｨｪ06ｨｦ 03ｨ ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ ｨｪｨ ｨ ｨ ﾂ ﾂ ｨｪｨ ﾂｨ 05ｨ ｨｨｨｪｨｺｨｮｨ ｨ 02ｨｨｨｨ ｨ 08ｨ ｨｰｨ ｨｬｨｨ (08ｨｨ09. 2, ｨ ). 05ｨ ｨｴｨｨｨｰｨｪ ｨｦ09ｨｰ09ｨｨ02 03ｨ ｨｪｨ 09 ｨｺ0502ｨｰｨｺｨ 01 K ｨｦ09ｨｰ09ｨｨ ｨ 08ｨ ｨｰｨ ｨｬｨｨ ｨｨ04ｨｮ ﾂｨ 05ｨｨ ｨｬ06ｨｴｨｹ ｨｬｨ ｨｰ ｨｮｨｪ06 ﾂｨｪ ｨ ｨｰ ｨ 08ｨ ｨｰ04 09ｨｪ0609ｨｨ05ｨｨ 09 05ｨｮｨｪｨｺｨｨ 96-05ｨｮｨｪ06 ﾂｨｪ06ｨｦ 0705ｨ ｨｰ04 04ｨ 6 ﾂ010609ｨｪ020902ｨｪｨｨ07 ｨｨｨｪ01ｨｮｨｺｨｰ ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｦ 00ｨｨｨ 0205ｨｨ ｨｨ05ｨｨ 0106 ｨｰ060006, ｨｺｨ ｨｺ ｨｺ0502ｨｰｨｺｨｨ ｨ 05ｨｨ ｨｰｨｺ06ｨｬｨｮ03 ｨｰ ｨｬｨｮ 0306ｨｺｨｮ (45 縺05, 30 ｨｬｨｨｨｪ). 02ｨ ｨｪｨｪ0402, ｨｰｨ ｨｪｨｪ0402 ｨｪｨ 08ｨｨ09. 3, 0102ｨｬ06ｨｪ09ｨｰ08ｨｨ08ｨｮ06ｨｰ, ﾂｨｰ ｨ 08ｨｨｨｰ0205ｨｹｨｪｨ 07 ｨｨｨｪ01ｨｮｨｺ02ｨｨ07 09ｨｨｨｪｨｰ0204ｨ 03ｨ ｨｪｨ 09 ｨｺ0502ｨｰｨｺｨ 01 K ｨｬ06ｨｴｨｹ ｨ 08ｨ ｨｰ0609 U-133 ｨｨ U-103 ｨｮ090205ｨｨ ﾂｨｨ09ｨ 05ｨ ｨｺ0605ｨｨ ﾂ0209ｨｰ ｨｨ ｨｺ0502ｨｰ06ｨｺ ｨｦ09ｨｰ09ｨｨ07 ｨｨｨｪ01ｨｮｨｺｨｰ060806ｨｬ ｨ ｨｰ0604ｨ 09ｨｰｨ ｨｮ ｨｨｨｪ06ｨｬ (1 ｨｬｨｺ00) c 65 ﾀ ﾀ 4.8 ｨｨ 77.2 ﾀ 3.1 % ｨｰ0902ｨｰ09ｨｰ0902ｨｪｨｪ06 (08ｨｨ09. 3, ｨ ) ｨ 08ｨ ｨｰ U ｨ 05 ｨｺ0605ｨｨ ﾂ0209ｨｰ ｨｨ04ｨｪ ｨ ｨｪ0401 ｨｺ0502ｨｰ06ｨｺ ｨｦ09ｨｰ09ｨｨ07 H 2 O 2 (500 ｨｬｨｺ00) ﾀ ﾀ 4.1 % (08ｨｨ09. 3, ｨ ), ｨ ｨ 08ｨ ｨｰ U-103 ｪ 0708ｨｨ ｨｰｨｺ06ｨｬ ｨｰ ｨｬ 0306ｨｺ02 (45 縺05, 30 ｨｬｨｨｨｪ) ｨ 05 03ｨｨ04ｨｪ ｨ ｨｪ0609ｨｰｨｹ ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｦ ｨｮ050702ｨｨｨｨ c 38 ﾀ ﾀ 2.6 % (08ｨｨ09. 3, 09). 02ｨ 09ｨｮ030102ｨｪｨｨ ｨｨ09ｨｺ 00ｨ ｨｺｨｰ , ｨ ｨｪ ｨ ｨｰｨｹ ｨｮ ｨｪｨｹ 05ｨｺ ｨｨｨｨ 03ｨ ｨｪｨ Hsp70, ｨｰ0907 ｨｮ0302 ｨ ｨｰ ｨｪｨ - ﾂ02ｨｪｨｨ02, ｨｺ06ｨｰ ｨｬｨｮ 0708ｨｨ01ｨ 02ｨｰ0907, ｨ ｨｪ ｨｮ06 06 ﾂ ｨｹ 09 ｨｮｨｪｨｨｨｺｨ 05ｨｹｨｪ04ｨｬｨｨ ｨｦ09ｨｰ09ｨ ｨｬｨｨ 03ｨ ｨｪｨ. 0604ｨｬ02ｨｪ02ｨｪｨｨ02 ｨｮ080609ｨｪ07 Hsp70 09 ｨｺ0502ｨｰｨｺｨ 01, ｨ ｨｴｨｨ01 ｨｬｨｮｨｰｨ ｨｪｨｰｨｪ0402 ｨ 0205ｨｺｨｨ 09 ｨｪｨ 08ｨｮ0302ｨｪｨｪ06ｨｦ ｨｺ06ｨｪ000608ｨｬｨ 02ｨｨ02ｨｦ, ｨｺ06ｨｰ ｨｰ ｨｨ ﾂｨｨｨｪ06ｨｦ ｨ ｨｺ06ｨｪ000608ｨｬｨ 02ｨｨ06ｨｪｨｪ ｨ ｨ ｨ ｨｪｨｨｨｦ, 0708ｨｨ090601ｨｨｨｰ ｨｺ 09ｨｪｨｨ0302ｨｪｨｨ06 ｨｺ0605ｨｨ ﾂ0209ｨｰ09ｨ 06ｨ 08ｨ 04ｨｮ06ｨｴｨｨ ｨ ｨ ｨｰ0609 ｨｨ, ｨｰｨ ｨｺｨｨｨｬ 06ｨ 08ｨ 0406ｨｬ, 0907ｨ 09ｨ 02ｨｰ ｨｪ020809ｨｪ0402 ｨｺ0502ｨｰｨｺｨｨ 06ｨｰ 00ｨｨｨ 0205ｨｨ ｨ ｨｪ02ｨｪｨｨ02 ｨｺ0502ｨｰ06 ﾂｨｪ ｨｬ020609ｨｰｨ 04ｨ ｨｨ 0708ｨ 09ｨｨ05ｨｹｨｪ ｨ 0205ｨｺ ｨｨｨｪ00ｨ ｪ ｨｪ0206ｨ ｨｨｨｬ0602 ｨｮ ｨｨ02 09ｨｮｨｴ0209ｨｰ090609ｨ ｨｪｨｨ ｨ 06ｨｦ ｨｺ0502ｨｰｨｺｨｨ, 01ｨ 0302 ｨｪ ｨ ｨｴ02ｨｦ ｨｬｨｮｨｰｨ ｨｪｨｰｨｪ0401 ｨ 0205ｨｺ ｨｺ0502ｨｰｨｺｨ ｨｨ ｨｦ ｨｨ ｨｨｨｰ ｨｪｨ ｨｺ ｨｪｨｨ02 ｨ 0205ｨｺ ｨ ｨ ｨｰ0609, ｨ ｨｺ0605ｨｨ ﾂ0209ｨｰ ｨｨｨｪｨｰ0204ｨｨ08ｨｮ02ｨｬ ｨ ｨｪｨ 0706ｨｪｨｨ03ｨ 02ｨｰ0907, ｨｮ05ｨｹｨｰｨ ｨｰ02 ﾂ ｨｪｨｨ03ｨ 02ｨｰ0907 ｨｺ06ｨｪｨｰ080605ｨｹ 04ｨ ｨｺｨ ﾂ0209ｨｰ0906ｨｬ ｨｺ0502ｨｰ06 ﾂｨｪ0401 ｨ 0205ｨｺ0609 (Morimoto, 2008) ｨｪｨｨ02 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ 06ｨ ｨｴ ｨ 0605ｨｹ03ｨｨｨｪ09ｨｰ09ｨ 0002ｨｪ ｨ ｨｪ0609 ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨ06ｨｪｨｪ ｨ ｨｺｨｰ0608ｨ HSF1 0708ｨｨ ｨ ｨｪｨｨ ﾂ02ｨｪｨｨｨｨ 01ｨｨ02ｨｰ04 ｨｮ ｨｪ02ｨｬｨ ｨｰ0601 C. elegans 04ｨｪｨ ﾂｨｨｨｰ0205ｨｹｨｪ ｨ 02ｨｰ ｨｨｨｰ0205ｨｹｨｪ0609ｨｰｨｹ 03ｨｨ04-

5 ｨｨｨｪ02ｨｪｨｨ07, ｨ 06ｨｴｨｨ02 05ｨｺ ｨｨ06 03ｨ ｨｪｨ Hsp ｨｨ ｨ 08ｨ ｨ 06ｨｰｨｺｨ ｨｺ0502ｨｰ06ｨｺ K ｨ 08ｨ ｨｰｨ ｨｬｨｨ 03ｨｨｨｺ06ｨｪｨｨｨｪｨ U-103 ｨｨ 0501ｨｨｨｪ ｨｬｨ U ｨｨ090601ｨｨｨｰ ｨｺ ｨｪｨｨ06 ｨｮ09ｨｰ06ｨｦ ﾂｨｨ090609ｨｰｨｨ ｨｺ0502ｨｰ06ｨｺ K562 ｨｺ 0102ｨｦ09ｨｰ09ｨｨ06 09ｨｰｨ ｨｮ ｨｨｨｪｨ (ｨ ), ｨｺｨｨ09ｨｨ ｨ (ｨ ) ｨｨ ｨｰｨｺ ｨｰ ｨｺｨ (09). STA ｪ 09ｨｰｨ ｨｮ ｨｨｨｪ, HS ｪ ｨｰ ｨｦ 0306ｨｺ ｨｺｨ ｨｮ ﾂｨ ｨｰｨ ｨｪ04 01ｨ ｨｪｨｪ ｨｨ04 3 ｨｪ0204ｨ 09ｨｨ09ｨｨｨｬ ｨｺ ｨｨｨｬ02ｨｪｨｰ ｨｨ ｨｪ ｨｨ 0603ｨｨｨ ｨｺｨｨ ｨｪ K ｪ ｨｺ06ｨｪｨｰ080605ｨｹ. ｨｪｨｨ, ｨ ｨ ｨｪｨｨ ｨｮｨｪｨｺ02ｨｨｨｨ, ｨｪｨ ｨｰｨｨ09, ｨｮｨｺ0608ｨ ﾂｨｨ09ｨ 02ｨｰ (Hsu et al., 2003) ｨｰｨ 0802ｨｪｨｨ ｨｨ ｨ ｨｪｨｨ04ｨｬ ｨｺ ｨｨ07 HSF ｨｬ02ｨｪ02ｨｬ ｨ 01ｨｨ08ｨｮ02ｨｰ, ｨ 07 ｨｺ0502ｨｰｨｺｨｨ ｨｬ02ｨｪ ｨｨ09ｨｰ02ｨｪｨｰｨｪ04ｨｬｨｨ ｨｺ ｨｺ0502ｨｰ06 ﾂｨｪ04ｨｬ 06ｨｺ09ｨｨ01ｨ ｨｪｨｰｨ ｨｬ (Calderwood et al., 2009) ｨ ｨｪ0609ｨｰｨｹ ｨｺ 09ｨｨｨｪｨｰ0204ｨｮ Hsp ｨ 09ｨｰ06ｨｬ ｨ ｨ 0209ｨ 02ｨｰ ｨｨ 09 ｨｬ ﾂｨｪ06ｨｦ ｨｰｨｺｨ ｨｪｨｨ. 00 ｨｪ0608ｨｬ02 05ｨｪ020800ｨｨ ﾂｨｪ ｨｺ08ｨ ｨｴ02ｨｪｨｨ07 ｨｬ ｨｨ ｨｰ ｨｺ 09ｨｨｨｪｨｰ0204ｨｮ ｨ 0205ｨｺ0609 ｨｰ ｨｺｨ, ｨｪ06 09 ｨｬ ﾂｨｪ06ｨｦ ｨｰｨｺｨ ｨｪｨｨ 09ｨｰｨ ｨｨ0906ｨｰｨｪ0401 ｨｨ ｨｨ ｨｦ 05ｨｰ06ｨｰ ｨ ｨｪ. 03ｨ 0102ｨｪｨｨ02 ｨｬ ﾂｨｪ06ｨｦ ｨｬｨ ｨｨ ｨｮｨｰ08ｨ ｨｰｨｮ ｨｬ ﾂｨｪ06ｨｦ 09ｨｨ ｨ 06ｨｰ ｨ ｨ 0502ｨｪｨｨ02ｨｬ ｨ ｨｪ0609ｨｰｨｨ ｨｺ 09ｨｨｨｪｨｰ0204ｨｮ Hsp ｨｦ09ｨｰ09ｨｨｨｰ0205ｨｹｨｪ06, ｨｰ0607ｨｪｨｪｨ 07 05ｨｺ ｨｨ07 Hsp70 ｨｮ ｨｰ08ｨ ｨｪ090002ｨｪｨｪ0401 ｨｬ040302ｨｦ ﾂｨ 09ｨｰｨｨ ﾂｨｪ06 06ｨｰｨｬ02ｨｪ0702ｨｰ 05ｨｰｨｨ ｨｺｨｰ04 (Kayani et al., 2008; Calderwood et al., 2009) ｨｨ ﾂｨｨ090502ｨｪｨｪ ｨ ｨｪｨｪ0401 ｨｬ0201ｨｨ02ｨｨｨｪ09ｨｺｨｨ02 09ｨｰ08ｨ ｨｰ0200ｨｨｨｨ, ｨｪｨ 0708ｨ ｨｪｨｪ0402 ｨｪｨ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 ｨｮ080609ｨｪ07 03ｨ ｨｪｨ 09 ｨｺ0502ｨｰｨｺｨ 01, ｨｰｨ ｨｰ ｨｹｨｬｨ ｨｺｨｰｨｨ09ｨｪ04ｨｬｨｨ. 00 ｨｪｨ 09ｨｰ0607ｨｴ ｨｬ07 09ｨｮｨｴ0209ｨｰ09ｨｮ02ｨｰ ｨｪ0209ｨｺ0605ｨｹｨｺ ｨｺ 0708ｨｨｨｬ02ｨｪ02ｨｪｨｨ06 03ｨ ｨｪ ｨｰ0208ｨ 07ｨｨｨｨ ｨｺ06ｨｪ000608ｨｬｨ 02ｨｨ06ｨｪｨｪ ｨ ｨｰ ｨｨｨｦ. 04ｨ 0408ｨ ｨ ｨ ｨｰ0409ｨ 06ｨｰ0907 ｨｰｨ ｨｺ ｨｪｨ ｨ 02ｨｬ ｨｨｨｬｨｨ ﾂ0209ｨｺｨｨ02 03ｨ ｨｪ04, ｨｰ. 02. ｨｬｨ ｨｬ060502ｨｺｨｮ0504, ﾂｨ ｨｴ ｨｰｨ ｨｴｨｨ ｨ 06ｨｦ ｨｨ09ｨ 01ｨ 08ｨｨ0104 ｨｨ05ｨｨ 0609ｨｬ0605ｨｨｨｰ04, ｨｪｨ 0708ｨｨｨｬ ｨｺｨｰ06ｨｨｨｪ ｨｨ05ｨｨ ｨ 02ｨｰｨ ｨｨｨｪ (Furusho et al., 2005). 06ｨ ｨｺｨｨｨｬｨｨ ｨｬ060502ｨｺｨｮ05ｨ ｨｬｨｨ ｨ 00ｨ 02ｨｰ0907 ｨｮ09ｨｨ05ｨｨｨｰｨｹ 03ｨ ｨｪｨｪ0402 ｨｬ0201ｨ ｨｪｨｨ04ｨｬ04 09 ｨｺ0502ｨｰｨｺｨ 01, ｨｨ 0109ｨ ｨｪｨｨ ｨｴ0209ｨｰ09ｨ 0706ｨｺｨ 04ｨ 05ｨｨ , 07ｨｮ09ｨｰｨｹ ｨｪ02ｨ 0605ｨｹ03ｨｮ06, ｨ ｨｪｨｰｨｨｨ ｨ ｨｰｨｪｨｮ06 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ 09 ｨｺ0502ｨｰｨｺｨ 01, ｨｬ ｨｨ08ｨｮ06ｨｴｨｨ01 ｨ ｨｪｨｹ ｪ00ｨ ﾂｨ 0106, 0601ｨｪｨｮ ｨｨ04 ｨｰｨ ｨｺ ｨｪｨ ｨ 02ｨｬ ｨｨ0005ｨｮｨｰｨ ｨｬｨｨｨｪ ｨ ｨｰ ｨｨｨｦ (Furusho et al., 2005) ｨｬ02 ｨｰ060006, ｨｪｨｨ050609ｨｹ, ﾂｨｰ06 ｨｨｨｪ00ｨｨｨ ｨｨｨｰ ｨ 09ｨｨ09ｨｨｨｬ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ ｨｺ0502ｨｰ06 ﾂｨｪ ｨ ｨｪｨ Hsp ｨ ｨｪｨ ｨｬｨｨ02ｨｨｨｪ, ｨｨ ｨｪ0402 ｨｺ06ｨｰ ｨｰ0907 ｨｪｨ ｨｪ0703ｨｪｨｨｨｦ 0102ｨｪｨｹ ｨｪｨ ｨｨｨ ｨｺｨｰｨｨ09ｨｪ04ｨｬｨｨ ｨｰｨｨ ｨｮ ｨｬｨｨ ｨ 08ｨ ｨｰｨ ｨｬｨｨ, ｨｨｨｪ01ｨｮ02ｨｨ08ｨｮ02ｨｰ 05ｨｺ ｨｨ06 Hsp70 ｨｨ 04ｨ ｨｴｨｨｨｴｨ 02ｨｰ ｨｺ0502ｨｰｨｺｨｨ ｨｬ060400ｨ 06ｨｰ 09ｨｰｨｨｨｬｨｮ0507ｨｰ0608ｨ 09ｨｨｨｬ07ｨｰ06ｨｬ0609 ｨ ｨｪｨｨ 03ｨ 08ｨｺｨｨｨｪ0906ｨｪｨ, 00ｨ ｨｺｨｰ0608ｨ (Shen et al., 2005) ｪ ｨ ｨｮｨ 05ｨｨｨｺ0609ｨ ｨｪ ｨ ｨ 06ｨｰ, 0102ｨｬ06ｨｪ09ｨｰ08ｨｨ08ｨｮ06ｨｴｨｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ ｨ 08ｨ ｨｰ0609, ｨｮ ﾂ02ｨｪｨｪ0401 ｨｨ04 ｨｪ02ｨｺ06ｨｰ ｨ 09ｨｰ02ｨｪｨｨｨｦ, 09 ｨ 01ｨ 07ｨｰ060002ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ06 09 ｨｬ0601ｨｮ050702ｨｨｨｨ 05ｨｺ ｨｨｨｨ Hsp ｨｰｨ ｨｺｨｨｨｬ ｨ 08ｨ ｨｰｨ ｨｬ 06ｨｰｨｪ0609ｨｨｨｰ ｨ 09ｨｰ080605, 06ｨ 08ｨ ｨ 06ｨｰｨｺｨ ｨｺ06ｨｰ060804ｨｬ 0708ｨｨ090601ｨｨｨｰ ｨｺ 09ｨｨｨｪｨｰ0204ｨｮ Hsp70 ｨｨ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ06 ｨｮ09ｨｰ06ｨｦ ﾂｨｨ090609ｨｰｨｨ ｨｺ0502ｨｰ06ｨｺ ｨｺ 02ｨｨｨｰ06ｨｰ06ｨｺ09ｨｨ ﾂ0209ｨｺｨｨｨｬ 00ｨ ｨｺｨｰ0608ｨ ｨｬ (Westerheide et al., 2004). 06ｨｰｨｨ ｨ 08ｨ ｨｰ04, ｨｬ0402 ｨｨ04 08ｨ 09ｨｰ02ｨｪｨｨｨｦ, ｨｺ06ｨｰ ｨｨ ｨｹ04ｨｮ06ｨｰ ｨｰ08ｨ 01ｨｨ02ｨｨ06ｨｪｨｪ06ｨｦ ｨｺｨｨｨｰｨ ｨｦ09ｨｺ06ｨｦ ｨｬ0201ｨｨ02ｨｨｨｪ02, 06ｨｰ05ｨｨ ﾂｨ 06ｨｰ ｨ ｨｪ0609ｨｰｨｹ06 ｨｺ ｨ ｨｺｨｰｨｨ09ｨ 02ｨｨｨｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨ06ｨｪｨｪ ｨ ｨｺｨｰ0608ｨ ｨｰ ｨｺｨ HSF1, ｨｮ0507ｨｰ0608ｨ 0706 ﾂｨｰｨｨ ｨｨｨｪ01ｨｮ02ｨｨｨ 0205ｨｹｨｪ ｨｪ0609 (Wes-

6 ｨｬ02ｨｪｨｺ06, ｨｪｨｰｨｨｨｬ06ｨｪ0609ｨ ｨｨ terheide, Morimoto, 2005). ﾁｨｰ06 09ｨｮｨｴ0209ｨｰ0902ｨｪｨｪ06, ｨ 09ｨｰ ｨｨ ｨ ｨｪ ｨｬｨｮ 0902ｨｴ0209ｨｰ09ｨ, 06ｨ ｨｪｨ 08ｨｮ0302ｨｪｨｪ ｨ 0405ｨｨ ﾂｨｪ ｨｺｨ 0809ｨｰ0902ｨｪｨｪ0401 ｨｰ08ｨ 09ｨ 01, ｨｮ0302 ｨｨ ｨｹ04ｨｮ06ｨｰ ｨｰ0208ｨ 07ｨｨｨｨ ｨｬｨｪ0600ｨｨ01 07ｨ ｨｰ ｨｨｨｦ, 09 ｨｰ06ｨｬ ﾂｨｨ ｨ ｨｪｨｪ ｨ ｨｨ080609ｨ ｨｪｨｨ02ｨｬ ｨｬｨｮｨｰｨ ｨｪｨｰｨｪ0401 ｨ 0205ｨｺ ｨｪ02ｨｦ0806ｨｪｨ ｨｮ02ｨｰ 09ｨｺｨ 04ｨ ｨｰｨｹ, ﾂｨｰ ｨｨ09ｨ ｨｪｨｪ ｨｨｨｪ01ｨｮｨｺｨｰ ｨ ｨｪ ｨ 05ｨ 01ｨ 06ｨｰ ｨｨｨｬｨｨ ｨｰ06ｨｨｨｪ09ｨｰ09ｨ ｨｬｨｨ ｨｨ ｨｪ ｨｰｨ ｨｰｨｺｨ ｨｬｨｨ. 01ｨ 0708ｨｨｨｬ0208, ｨ 09ｨｰ080605, ｨｪｨ ｨｺ06ｨｰ060804ｨｦ ｨ 00ｨ 02ｨｰ0907 ｨｬｨｪ ｨｪｨ , 0706ｨｺｨ 04ｨ ｨｹｨｪ06 09ｨｺ0806ｨｬｨｪｨｮ ｨ ｨｪ0609ｨｰｨｹ ｨｺ ｨｨｨｪ01ｨｮｨｺ02ｨｨｨｨ Hsp70 ｨｨ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ06 03ｨｨ04ｨｪ ｨ ｨｪ0609ｨｰｨｨ ｨｬ06ｨｰ0608ｨｪ0401 ｨｪ02ｨｦ0806ｨｪ ｨｺｨｮ05ｨｹｨｰｨｮ ｨｰ060006, 09ｨ ｨｬ ｨ ｨ 08ｨ ｨｰ 06ｨ 05ｨ 01ｨ ｨ 0302ｨｪｨｪ06ｨｦ ｨｪ02ｨｦ0806ｨｰ06ｨｺ09ｨｨ ﾂｨｪ0609ｨｰｨｹ06 (Kalmar, Greensmeeth, 2009). 0809ｨｰ ｨｪ02ｨｦ 08ｨ ｨ 06ｨｰ04 06ｨｰ01ｨ 06ｨｰ ﾂｨｰ02ｨｪｨｨ ｨｮ0006ｨｬｨｮ ｨｨｨｪ01ｨｮｨｺｨｰ0608ｨｮ Hsp70 ｪ ｨ 08ｨｨｨｬ06ｨｺ0506ｨｬ0605ｨｮ, ｨｺ06ｨｰ060804ｨｦ 0102ｨｬ06ｨｪ09ｨｰ08ｨｨ08ｨｮ02ｨｰ ｨｰ0208ｨ ｨｰｨｨ ﾂ0209ｨｺｨｨｨｦ ｨｺｨｰ 09 ｨｬ0403ｨｨｨｪ06ｨｦ ｨｬ ｨｨ ｨ ｨｬｨｨ06ｨｰ080600ｨｨ ﾂ0209ｨｺ ｨ ｨｰ0208ｨ 05ｨｹｨｪ ｨｺ ｨ (ALM). 0808ｨｨｨｬ06ｨｺ0506ｨｬ0605 ｨｪ02 ｨｰ0605ｨｹｨｺ06 ｨｮ090205ｨｨ ﾂｨｨ09ｨ 05 05ｨｺ ｨｨ06 Hsp70 09 ｨｪ020809ｨｪ06ｨｦ ｨｰｨｺｨ ｨｪｨｨ ｨｰ08ｨ ｨｪ090002ｨｪｨｪ0401 ｨｬ040302ｨｦ SOD G93A, ｨｪ06 ｨｨ ｨ 05 03ｨｨ04ｨｪ ｨ ｨｪ0609ｨｰｨｹ ｨｬ06ｨｰ0608ｨｪ0401 ｨｪ02ｨｦ0806ｨｪ0609, 06ｨ ｨｴｨｮ ｨｨｨｰ0205ｨｹｨｪ0609ｨｰｨｹ 03ｨｨ04ｨｪｨｨ ｨｬ040302ｨｦ ｨｨ 0708ｨｨ090601ｨｨ05 ｨｺ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ06 ｨｬ ﾂｨｪ06ｨｦ 09ｨｨ0504 (Kalmar et al., 2008). 0201ｨｪｨ ｨｺ06 ｨｨ ｨｮ ｨ 08ｨｨｨｬ06ｨｺ0506ｨｬ0605ｨ, 0706ｨｺｨ 04ｨ ｨ ｨｬ02 ﾂｨ ｨｰ0205ｨｹｨｪ04ｨｦ ｨｺｨｰ 09 ｨ ｨｨ ｨｨ ﾂ0209ｨｺｨｨ01 05ｨｺ ｨｨｨｬ02ｨｪｨｰｨ 01, ｨｪｨ 09ｨｰ060806ｨｦ 09ｨｰｨ 01ｨｨｨｨ ｨｺ05ｨｨｨｪｨｨ ﾂ0209ｨｺｨｨ01 ｨｨ090704ｨｰｨ ｨｪｨｨｨｦ, 0706 ｨｮ09ｨｰｨｪ04ｨｬ ｨ ｨｴ02ｨｪｨｨ07ｨｬ ｨｨｨ 05ｨｨ09ｨｰ0609, ｨ 0405ｨｨ 06ｨ ｨｪｨ 08ｨｮ0302ｨｪ ｨ 06 ﾂｨｪ ｨｺｨｰ ｨｰ ｨｮ02ｨｰ, ﾂｨｰ ｨｨ09ｨｺ ｨｨｨｪ01ｨｮｨｺｨｰ ｨ ｨｪ0609, ｨｺ06ｨｰ ｨｬ0603ｨｪ06 ｨ ｨ 04 ｨｨ ｨｹ040609ｨ ｨｰｨｹ 09 ｨｺ05ｨｨｨｪｨｨｨｺ02, ｨｪ02 04ｨ ｨｺ06ｨｪ ﾂ02ｨｪ ｨｨ 09 05ｨｰｨｨｨｬ ｨｬ04 ｨｪｨ ﾂｨ 05ｨｨ ｨ ｨｪｨ 05ｨｨ04 ｨ ｨｨｨ 05ｨｨ06ｨｰ02ｨｺｨｨ 0902ｨｴ0209ｨｰ09, ｨ ｨｪｨｪ06ｨｦ , ｨｺ06ｨｰ0608ｨ ｨ 05ｨ ｨｰ08ｨｨｨｰ ｨｪ04 01ｨ ｨｬｨｬｨ 08ｨ ｨｪ ｨ, 0002ｨｰ ｨｨｨｺ05ｨｨ ﾂ0209ｨｺｨｨ ｨｨ080601ｨｪ0402 ｨ 05ｨｺｨ 0506ｨｨ0104 ｨｨ04 ｨｬ060809ｨｺｨｨ ｨ ｨｪｨｨ04ｨｬ0609, ｨ ｨｰｨ ｨｺ ｨｨ080601ｨｪ0402 ｨｨ 09ｨｨｨｪｨｰ02ｨｰｨｨ- ﾂ0209ｨｺｨｨ02 1, 4-ｨｪｨ 00ｨｰ0601ｨｨｨｪ06ｨｪ04 ｨｨ ｨｨ ｨｨ S-0005ｨｨｨｺ0604ｨｨ ｨ ｨｬ ｨｮ01ｨ ｨｹ 06ｨ ｨｪｨ 08ｨｮ03ｨｨｨｰｨｹ 0109ｨ 0902ｨｴ0209ｨｰ09ｨ, 03ｨｨｨｺ06ｨｪｨｨｨｪ U-103 ｨｨ 0501ｨｨｨｪ ｨｬ U-133, ｨｺ06ｨｰ ｨ 08ｨｬｨ ｨｺ ｨｨ ﾂ0209ｨｺｨｨ01 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ0701 ｨ ｨｺｨｰｨｨ09ｨｨ080609ｨ 05ｨｨ 09ｨｨｨｪｨｰ0204 Hsp70 09 ｨｺ0502ｨｰｨｺｨ ｨｨ04ｨｮ- ﾂ02ｨｪｨｪ0401 ｨｺ0502ｨｰ06 ﾂｨｪ ｨｨｨｪｨｨｨｦ ｨｪｨｹ ｨｪｨ ｨｺ ｨｪｨｪ ｨｰ02 ﾂ02ｨｪｨｨ02 12 ﾂ ｨ 0205ｨｺｨ Hsp ｨ 05 ｨｺ06ｨｪｨｰ080605ｨｹｨｪ04ｨｦ 09 08ｨ 0404, 0708ｨｨ- ﾂ02ｨｬ ｨｪｨｨ02 ｨ ｨ ｨｨ09ｨｰ08ｨｨ080609ｨ ｨｪ ｨｰｨ ｨｺ0502ｨｰｨｺｨ ｨｬｨｨ 08ｨ 0405ｨｨ ﾂｨｪ ｨｨ ｨｪｨｨ ｨ ｨｪｨ ｨｨｨｨ ｨ 09ｨｰ ｨｬ ｨｰｨ ｨｰ ｨｪ ｨｰｨｪ04ｨｬ, ﾂｨｰ06 ｨｮｨｺｨ 04ｨ ｨｪｨｪ ｨｴ0209ｨｰ09ｨ ｨ ｨｮ01ｨｮｨｰ ｨｨｨｬ02ｨｰｨｹ Hsp70-ｨｨｨｪ01ｨｮ02ｨｨ08ｨｮ06ｨｴｨｮ06 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ ｨ ｨｪｨｨ04ｨｬ02. 09ｨ 08ｨ ｨｺｨｰ0208ｨｪ06ｨｦ ｨ 02ｨｪｨｪ0609ｨｰｨｹ06 Hsp ｨｰ ｨ ｨｪｨｪｨ 07 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ, 09 ｨｺ06ｨｰ060806ｨｦ ｨ 06ｨｰ ｨ ｨｨ ｨｨ ﾂ0209ｨｺｨｨ02 00ｨｮｨｪｨｺ02ｨｨｨｨ 05ｨｰ ｨ 0205ｨｺｨ, ｨｨ ｨ ｨｮｨｴ0209ｨｰ0902ｨｪｨｪ ｨｨｨｰｨｹ ｨｺ0502ｨｰｨｺｨ 01, 06ｨ 08ｨ ｨ 06ｨｰｨ ｨｪｨｪ ｨ 06ｨｨｨｬｨｨ ｨｨｨｪ02ｨｪｨｨ07ｨｬｨｨ ｨｨ04ｨｬ020802ｨｪｨｨ07 ｨｬ04 ｨｨ ｨｹ040609ｨ 05ｨｨ ｨｬ0601ｨｨ00ｨｨ02ｨｨ080609ｨ ｨｪｨｪｨｮ06 ｨｰ0209ｨｰ-09ｨｨ09ｨｰ02ｨｬｨｮ, ｨｨ ｨｬ06ｨｴｨｹ06 ｨｮ01ｨ ｨｹ ｨｪｨｨｨｰｨｹ, ﾂｨｰ ｨｨｨｪ02ｨｪｨｨ07 U-103 ｨｨ U-133 ｨｮ090205ｨｨ ﾂｨｨ09ｨ 06ｨｰ 03ｨ ｨｪｨｪｨｮ06 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ ｨｰｨ ｨｺ ｨ ｨ ｨｺ 08ｨ ｨｪ0202 ｨｪｨ ｨｬｨｨ ｨ ｨｮ09ｨｰｨ ｨｪ ｨｪ06, ｨｰｨ ｨｺ0602 ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 03ｨ ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ Hsp70 09 ｨｺ0502ｨｰｨｺｨ 01, 06ｨ 08ｨ ｨ 06ｨｰｨ ｨｪｨｪ0401 ｨｰ ｨｬ 0306ｨｺ06ｨｬ, ｨｨ ｨｨｨｰ ｨ 05ｨ ｨ 0807 ｨｪ02 ｨｰ0605ｨｹｨｺ ｨｰｨｮ ｨｺ0605ｨｨ ﾂ0209ｨｰ09ｨ, ｨｪ06 ｨｨ ｨｪｨｨ06 ｨｮ080609ｨｪ07 ｨｺ0603ｨ ｨｪｨ Hdj1/Hsp40 ( ｨｨ 0108., 2005) ｨｰ06ｨｬｨｮ ｨｬ ｨ 00ｨ 02ｨｬ, ﾂｨｰ06 08ｨ 0909ｨｬｨ ｨｰ08ｨｨ09ｨ 02ｨｬ ｨｹ 0902ｨｴ0209ｨｰ09ｨ ｨ 06ｨｰ 09ｨｨｨｪｨｰ ｨ 06ｨｨ01 03ｨ ｨｪ0609, ｨｺ06ｨｰ ｨ ｨ 06ｨｰｨ 06ｨｰ ｨｺ02 ｨｨ 04ｨ ｨｴｨｨｨｴｨ 06ｨｰ ｨｺ0502ｨｰｨｺｨｨ ｨｬ060400ｨ 06ｨｰ ｨｬｨｪ ﾂｨｨ0905ｨ 07ｨ ｨｰ060002ｨｪ0609, 09 ﾂｨ 09ｨｰｨｪ0609ｨｰｨｨ 06ｨｰ ｨ 0205ｨｺ ｨｨ0005ｨｮｨｰｨ ｨｬｨｨｨｪ060904ｨｬｨｨ ｨｰ0608ｨ ｨｬｨｨ, 06ｨ 08ｨ 04ｨｮ06ｨｴｨｨ01 ｨ ｨ ｨｰ04 (Zhou et al., 2001) ｨｪｨｨ02 02ｨｨｨｰ ｨｰ02ｨｺｨｰ0608ｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ 0904ｨ 08ｨ ｨｪｨｪ0401 ｨｪｨ ｨｬｨｨ 0902ｨｴ0209ｨｰ ｨｺｨ 04ｨ 0506, ﾂｨｰ06 06ｨ 08ｨ ｨ 06ｨｰｨｺｨ ｨｨｨｬｨｨ ｨｺ0502ｨｰ06ｨｺ 0708ｨｨ090601ｨｨｨｰ ｨｺ 09ｨｪｨｨ0302ｨｪｨｨ06 ｨｮ080609ｨｪ07 09ｨｬ0208ｨｰｨｪ0609ｨｰｨｨ 06ｨｰ ｨｰ ｨ ｨｺｨｰ ｪ 0502ｨｰｨ 05ｨｹｨｪ ｨｰ ｨｺｨ, ｨｺｨｨ09ｨｨ ｨ ｨｨ 09ｨｰｨ ｨｮ ｨｨｨｪｨ. 06ｨｰ06 09ｨｪｨｨ0302ｨｪｨｨ02 ｨ ｨ 02ｨｪｨｪ06 06ｨｴｨｮｨｰｨｨｨｬ04ｨｬ ｨｴ0209ｨｰ09ｨ U-103, 06ｨ 08ｨ ｨ 06ｨｰｨｺｨ ｨｺ06ｨｰ060804ｨｬ 0708ｨｨ090601ｨｨ05ｨ ｨｺ ｨｪｨｨ06 03ｨｨ04ｨｪ ｨ ｨｪ0609ｨｰｨｨ ｨｺ0502ｨｰ06ｨｺ K ｨｨ ｨｰ ｨｬ 0306ｨｺ ｨｨ 45 縺 ｨ ｨｺｨｰｨｨ ﾂ0209ｨｺｨｨ ｨ 04ｨ. 06ｨ ｨｺｨｨｨｬ 06ｨ 08ｨ 0406ｨｬ, ｨｬ ｨｺｨ 04ｨ 05ｨｨ, ﾂｨｰ ｨ 0904ｨ 08ｨ ｨｪｨｪ0401 ｨｪｨ ｨｬｨｨ ｨｨｨｪ02ｨｪｨｨ07 09ｨｰｨｨｨｬｨｮ05ｨｨ08ｨｮ06ｨｰ 09ｨｨｨｪｨｰ0204 Hsp70, ｨｮ090205ｨｨ ﾂｨｨ09ｨ 06ｨｰ ｨ ｨｪｨｪｨｮ06 ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ ｨｨ ｨｰ0902ｨｰ09ｨｰ0902ｨｪｨｪ ｨ 06ｨｰ 04ｨ ｨｴｨｨｨｰｨｪ04ｨｦ 0706ｨｰ02ｨｪ02ｨｨｨ 05 ｨｺ0502ｨｰｨｺｨｨ. 06ｨｪ04ｨｬｨｨ ｨ ｨｬｨｨ, 06ｨ ｨ ｨ 08ｨ ｨｰｨ 0102ｨｬ06ｨｪ09ｨｰ08ｨｨ08ｨｮ06ｨｰ 00ｨ 08ｨｬｨ ｨｺ ｨｨ ﾂ0209ｨｺｨｨｨｦ 0706ｨｰ02ｨｪ02ｨｨｨ 05 ｨｨ, ｨｪ0209ｨｬ06ｨｰ0807 ｨｪｨ ｨｪ ｨｰｨ ｨｰｨｺｨｨ ｪ ｨ ｨｪｨｨ ﾂ02ｨｪｨｪｨｮ06 08ｨ 09ｨｰ090608ｨｨｨｬ0609ｨｰｨｹ ｨｪ06ｨｦ ｨｨ ｨｮｨｬ020802ｨｪｨｪｨｮ06 ｨｰ06ｨｺ09ｨｨ ﾂｨｪ0609ｨｰｨｹ, ｨｬ0600ｨｮｨｰ ｨ 04ｨｰｨｹ ｨ ｨ 06ｨｰｨ ｨｪ04. 06ｨｬ02ｨｪｨｪ06 ｨｰｨ ｨｺｨ 07 ｨｺ ｨｺ02ｨｨ07 01ｨｨｨｬｨｨ ﾂ0209ｨｺ06ｨｦ 09ｨｰ08ｨｮｨｺｨｰｨｮ ｨｴ0209ｨｰ ｨｨｨｰ ｨｪｨ 09ｨｰ0607ｨｴ ｨｬ ｨｰ ｨ 05ｨ ｨ 0807ｨｰ 04ｨ ｨｦ09ｨｰ09ｨｨ ｨ ｨｬ06ｨｬ ｨｪｨ ﾂｨ ｨ ｨ 06ｨｰ 0906ｨｰ08ｨｮ01ｨｪｨｨｨｺ ｨｨ010606ｨｺ02ｨ ｨｪ09ｨｺ ｨｨｨｪ09ｨｰｨｨｨｰｨｮｨｰｨ ｨ ｨｨ ｨ ｨｪｨｨ- ﾂ0209ｨｺ06ｨｦ 01ｨｨｨｬｨｨｨｨ ｨｨ 01ｨｨ0802ｨｺｨｰ0608ｨ ｨｨｨｪ09ｨｰｨｨｨｰｨｮｨｰｨ ｨｰ06ｨｪｨｨｨｺｨ. 04ｨ ｨ 06ｨｰｨ ｨｪ02ｨｪｨ 0708ｨｨ 00ｨｨｨｪｨ ｨｪ ｨｦ ｨｺ ｨｨｨｦ09ｨｺ ｨｪ01ｨ 00ｨｮｨｪ01ｨ ｨｬ02ｨｪｨｰｨ 05ｨｹｨｪ0401 ｨｨ ｨ ｨｪｨｨｨｦ ( ｨｺｨｰ ) ｨｨ ｨ 05ｨｹｨｪ ｨｪｨｰ08ｨ ｨｪｨ ｨｮ ﾂｨｪ06-ｨｰ0201ｨｪｨｨ ﾂ0209ｨｺｨｨ ｨ ｨｬｨｬ ( ). 0507ｨｨ0906ｨｺ 05ｨｨｨｰ0208ｨ ｨｰｨｮ ｨ 0800ｨｮ05ｨｨ , 01ｨｮ030609ｨ ｨｦｨｪｨ ｨｹ 03ｨ ｨｪ ｨｰ0902ｨｰ02 ｨｺ0502ｨｰｨｺｨｨ ｨｨ ｨ ｨｪｨｨ04ｨｬｨ ｨｪｨ 09ｨｰ ｨｨｨｰ ｨｨ (3) : 219 ｪ , ｨ , ｨｺｨ ｨ , 00ｨ 0800ｨｮ05ｨｨ , 01ｨｮ030609ｨ ｨｨｨｪｨ ｨｬｨｨｨｺｨ 03ｨ ｨｪｨｪ0401 ｨｺ06ｨｬ070502ｨｺ Hsp70 09 ｨ 0205ｨｺｨ ｨｬｨｨ-0706ｨｬ06ｨｴｨｪｨｨｨｺｨ ｨｬｨｨ Hdj1 ｨｨ Bag ｨ ｨｺ02ｨｨｨｨ ｨｺ0502ｨｰ06ｨｺ 0508ｨｨｨｰ ｨｦｨｺ02ｨｬｨｨｨｨ ﾂ ｨｺｨ K562 ｨｪｨ ｨｰ ｨｦ 09ｨｰ ｨｨｨｰ ｨｨ (7) : 620 ｪ627. Bradford M A rapid and sensitive method for quantitation of microgramm quantities of protein utilizing the principle of protein-dye binding. Anal. Boichem. 72 : 248 ｪ254. Bukau B., Weissman J., Horwich A Molecular chaperones and protein quality control. Cell. 125 : 443 ｪ451. Calderwood S. K., Murshid A., Prince T The shock of aging: molecular chaperones and the heat shock response in longevity and aging ｪ a mini-review. Gerontology. 55 : 550 ｪ558. Chiti F., Dobson C. M Protein misfolding, functional amyloid, and human disease. Annu. Rev. Biochem. 75 : 333 ｪ366. Deuerling E., Bukau B Chaperone-assisted folding of newly synthesized proteins in the cytosol. Crit. Rev. Biochem. Mol. Biol. 39 : 261 ｪ277. Fewell S. W., Smith C. M., Lyon M. A., Dumitrescu T. P., Wipf P., Day B. W., Brodsky J. L Small molecule modulators of endogenous and co-chaperone-stimulated Hsp70 ATPase activity. J. Biol. Chem. 279 : ｪ Furusho K., Yoshizawa T., Shoji S Ectoine alters subcellular localization of inclusions and reduces apoptotic cell death induced by the truncated Machado ｪJoseph disease gene product with an expanded polyglutamine stretch. Neurobiol. Dis. 20 : 170 ｪ 178. Hartl F. U., Hayer-Hartl M Molecular chaperones in the cytosol: from nascent chain to folded protein. Science. 295 : 1852 ｪ1858. Hsu A. L., Murphy C. T., Kenyon C Regulation of aging and age-related disease by DAF-16 and heat-shock factor. Science. 300 : 1142 ｪ1145. Kalmar B., Greensmith L Activation of the heat shock response in a primary cellular model of motoneuron neurodegeneration-evidence for neuroprotective and neurotoxic effects. Cell Mol. Biol. Lett. 14 : 319 ｪ335. Kalmar B., Novoselov S., Gray A., Cheetham M. E., Margulis B., Greensmith L Late stage treatment with arimoclomol delays disease progression and prevents protein aggregation in the SOD1 mouse model of ALS. J. Neurochem. 107 : 339 ｪ350. Kayani A. C., Close G. L., Broome C. S., Jackson M. J., McArdle A Enhanced recovery from contraction-induced damage

7 ｨｨｨｪ02ｨｪｨｨ07, ｨ 06ｨｴｨｨ02 05ｨｺ ｨｨ06 03ｨ ｨｪｨ Hsp in skeletal muscles of old mice following treatment with the heat shock protein inducer 17-(allylamino)-17-demethoxygeldanamycin. Rejuvenation Res. 11 : 1021 ｪ1030. Morimoto R. I Proteotoxic stress and inducible chaperone networks in neurodegenerative disease and aging. Genes Develop. 22 : 1427 ｪ1438. Morimoto R. I., Cuervo A. M Protein homeostasis and aging: taking care of proteins from the cradle to the grave. J. Gerontol. Biol. Sci. Med. Sci. 64 : 167 ｪ170. Perutz M. F Glutamine repeats and neurodegenerative diseases. Molecular aspects. Trend Biochem. Sci. 24 : 58 ｪ63. Sakahira H., Breuer P., Hayer-Hartl M. K., Hartl F. U Molecular chaperones as modulators of polyglutamine protein aggregation and toxicity. Proc. Nat. Acad. Sci. USA. 99 : ｪ Shen H. Y., He J. C., Wang Y., Huang Q. Y., Chen J. F Geldanamycin induces heat shock protein 70 and protects against MPTP-induced dopaminergic neurotoxicity in mice. J. Biol. Chem. 280 : ｪ Soti C., Csermely P Chaperones come of age. Cell Stress Chaperones. 7 : 186 ｪ190. Wawrzynow A., Zylicz M Divergent effects of ATP on the binding of the DnaK and DnaJ chaperones to each other, or to their various native and denatured protein substrates. J. Biol. Chem. 270 : ｪ Westerheide S. D., Bosman J. D., Mbadugha B. N., Kawahara T. L., Matsumoto G., Kim S., Gu W., Devlin J. P., Silverman R. B., Morimoto R. I Celastrols as inducers of the heat shock response and cytoprotection. J. Biol. Chem. 279 : ｪ Westerheide S. D., Morimoto R. I Heat shock response modulators as therapeutic tools for diseases of protein conformation. J. Biol. Chem. 280 : ｪ Wyttenbach A Role of heat shock proteins during polyglutamine neurodegeneration: mechanisms and hypothesis. J. Mol. Neurosci. 23 : 69 ｪ96. Wyttenbach A., Swartz J., Kita H., Thykjaer T., Carmichael J., Bradley J., Brown R., Maxwell M., Schapira A., Orntoft T. F., Kato K., Rubinsztein D. C Polyglutamine expansions cause decreased CRE-mediated transcription and early gene expression changes prior to cell death in an inducible cell model of Huntington ｯs disease. Hum. Mol. Genet. 10 : 1829 ｪ1845. Zhou H., Li S. H., Li X. J Chaperone suppression of cellular toxicity of huntingtin is independent of polyglutamine aggregation. J. Biol. Chem. 276 : ｪ ｨｰｨｮ07ｨｨ05ｨ 9 XI 2009 NOVEL COMPOUNDS INCREASING CHAPERONE HSP70 EXPRESSION AND THEIR BIOLOGICAL ACTIVITY E. M. Eremenko, 1 O. I. Antimonova, 1 O. G. Shekalova, 1 S. G. Polonik, 2 B. A. Margulis, 1 I. V. Guzhova 1, * 1 Institute of Cytology RAS, St. Petersburg and 2 Pacific Institute of Bioorganic Chemistry RAS, Vladivostok; * guzhova@mail.cytspb.rssi.ru Hsp70 possesses chaperonic activity, the property associated with the protective function that was demonstrated in experiments on a great number of cell and animal models. Therefore, it seems important to search for the substances able to innocuously elevate the chaperone concentration in an organism cells and tissues. In our work, we screened of more that 60 compounds and found two chemicals, derivatives of shikonin and echinochrome that able to increase the chaperone level in a variety of human cells. It was shown that in human erythroleukemia K562 cells treated with the both substances concomitantly with elevation of Hsp70 level the absolute chaperonic activity was also increased; this can indicate mobilization of the whole cellular chaperonic machinery by above mentioned compounds. Estimating biological activity of the two substances, we demonstrated that treatments of cells by them prior to hard heat stress, hydrogen peroxide or staurosporine reduced cell mortality by 20 ｪ50 % depending on a cytotoxic factor. The results show that after simple chemical modifications these compounds might be taken as a basis of pharmaceuticals for therapy of wide range of disorders. K e y w o r d s: chaperone, chaperone inducer, shikonin, echinochrome.