Tsitologiya 54(11)

Transcription

1 ｨｬ 54, ﾁ ﾁ ﾁ ｨ 0609ｨ 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; ｨｺｨｰ0806ｨｪｨｪ04ｨｦ ｨ : ｨ 0609ｨ 0508ｨ 09ｨｪｨｨｨｰ0205ｨｹｨｪ04ｨｦ ｨ ｨｪｨ 05ｨｨ04 01ｨ 08ｨ ｨｺｨｰ0208ｨ 0005ｨｮ ｨｪｨｰｨｪ ｨｬ02 ﾂ02ｨｪｨｨ ｨ ｨｪｨｪｨｨ01 05ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ ｨｰｨ ｨｪ04 ｨｪ ｨ ｨｪｨｪ ｨ ｨ 02ｨｪｨｪ0609ｨｰ0701 ｨｨｨｬｨｬｨｮｨｪ060005ｨｮ ｨｪｨｰｨｪ ｨｪｨｨ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ ｨｮ01ｨｺ0502ｨｰ06 ﾂｨｪ ｨｬｨ 08ｨｨ06ｨｪｨ 01 ｨｬ0403ｨｨ 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ 0005ｨｮ ｨｪｨｰｨｪ06ｨｬ02 ﾂ02ｨｪｨｪ0401 ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ 05- ｨｨ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ ｨｮ ﾂ02ｨｪ ｨ ｨｪｨｨ02 ｨ ｨｺｨｰｨｨｨｪｨ 0708ｨｨ ｨｨ09ｨｺｨｮ0909ｨｰ0902ｨｪｨｪ06ｨｬ ｨｨｨｪ00ｨｨｨ ｨｨ080609ｨ ｨｪｨｨｨｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ, ｨ ｨｰｨ ｨｺ ｨｬ02ｨｪｨｰｨ ｨｰｨｨ09ｨｪ ｨ 09ｨｴ ｨｪｨｨ ｨｨ04ｨｮｨ 05ｨｹｨｪ0402 ｨｪｨ ｨ ｨｪｨｨ ｨｪ02ｨｪ ｨ 09ｨｪｨｨｨｰ0205ｨｹｨｪ04ｨｬ ｨｺ0605ｨｨ ﾂ0209ｨｰ0902ｨｪｨｪ04ｨｬ ｨ ｨｪｨ 05ｨｨ0406ｨｬ ｨｨｨｪｨｰ020008ｨ 05ｨｹｨｪ06ｨｦ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｨ 0005ｨｮ ｨｪ02ｨｨｨｨ ｨｪｨ ｨｺ06ｨｪ0006ｨｺｨ 05ｨｹｨｪ0401 ｨｨ0406ｨ 08ｨ 0302ｨｪｨｨ ｨ 02ｨｨ01 05ｨｺ ｨｨｨｬ02ｨｪｨｰｨ 05ｨｹｨｪ ｨｮ0707ｨ 01 ｨｺｨ ｨｺ 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨｨｨｪ00ｨｨｨ ｨｨｨｰ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ, ｨｰｨ ｨｺ ｨｨ ｨ 08ｨ ｨ 06ｨｰｨｺｨｨ 05ｨｬｨ 08ｨｨ06ｨｪ ｨ 0406ｨｦ ｨｪ06 ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｨ 0005ｨｮ ｨｪ02ｨｨｨｨ ｨ 09ｨｪ02ｨｪｨｨ06 09 ｨｺ06ｨｪｨｰ ｨｬ 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ C-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ, ｨｪ06 ｨｪ02 ｨｺ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨ ｨｺｨｰｨｨｨｪｨ ｨｮ ﾂ02ｨｪｨｪ ｨ ｨｪｨｪ ｨｰ ｨｨｨｰｨｹ, ﾂｨｰ06 ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ ｨ ｨｺｨｰｨｨ09ｨｪ ｨｨ ｨｨｨｬｨｬｨｮｨｪ060005ｨｮ ｨｪｨｰｨｪ06ｨｬ ｨｪｨｨｨｨ ｨ ｨｺｨｰｨｨｨｪｨ ｨ ｨ 07ｨｴｨｨ ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ ﾂ ｨ : 0106ｨｨｨｬ0705ｨ ｨｪｨｰｨ 02ｨｨ06ｨｪｨｪ ｨ ｨｨ ｨｬ0403ｨｨ, ｨｪ04ｨｦ ｨ ｨｺｨｰｨｨｨｪ, ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨ07, 05ｨ ｨｪｨ 07 09ｨｺｨ ｨｪｨｨ08ｨｮ06ｨｴｨ 07 ｨｺ06ｨｪ0006ｨｺｨ 05ｨｹｨｪｨ 07 ｨｬｨｨｨｺ080609ｨｺ0607ｨｨ ｨｬ06ｨｰ0807 ｨｪｨ ｨｰ06 ﾂｨｰ ｨ ｨｴ02ｨｪｨｨ ｨｪｨｨｨｨ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ (Clark, Rosenbaum, 1979; Nakayasu, Ueda, 1983; 03ｨ ｨｪ0609, 01ｨ 05ｨ ｨｺｨｰｨｨ06ｨｪ0609, 1987) ｨ 0405ｨｨ 0909ｨｰ0802 ﾂ02ｨｪ ｨｰｨ ｨｰ06 ﾂｨｪ06 09ｨｺ0207ｨｰｨｨ ﾂ0209ｨｺｨｨ, ｨｨ ｨ ｨｪｨｨ ｨｪｨｨ ｨｰｨｨ0502ｨｰｨｨｨｦ ｨｰ ｨｨ05ｨｨ 0708ｨｨ09ｨｮｨｰ09ｨｰ09ｨｨ02 ｨ ｨｺｨｰｨｨｨｪｨ 09 ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｬ ｨｰ060006, ｨｬ06ｨｪ09ｨｰ08ｨｨ080609ｨ ｨｪ06 ｨｮ ﾂｨ 09ｨｰｨｨ02 ｨ ｨｺｨｰｨｨｨｪｨ ｨｬ ｨｪ ｨｮｨｪｨｺ02ｨｨｨｦ, 09ｨｺ0506 ﾂｨ ｨｮ050702ｨｨ ｨｪ0609ｨｪ ｨｰｨ ｨｺ ｨｨｨｨ 0002ｨｪ0609 (09ｨｬ. 06ｨ : Hofmann, 2009; Visa, Percipalle, 2010) ｨ 0502ｨｪｨｨ06, ｨｪｨｨ07 06 ｨｬ0209ｨｰ02 ｨｨ ｨｨ ｨ ｨｺｨｰｨｨｨｪｨ ｨｰ08ｨ ｨｪ09ｨｰ0902ｨｪｨｪ06ｨｦ ｨ ｨｪｨｨ04ｨ 02ｨｨｨｨ ｨｺ0502ｨｰ06 ﾂｨｪ ｨ 0609ｨｰｨ 06ｨｰ0907 ｨｪ02ｨｬｨｪ ﾂｨｨ090502ｨｪｨｪ04ｨｬｨｨ. 02ｨ ｨｲ0709ｨｪ0702ｨｰ ｨｰ06 ｨｰ02ｨｬ, ﾂｨｰ06 ｨ 0605ｨｹ03ｨｨｨｪ09ｨｰ ｨｺ ｨｨｨｬ02ｨｪｨｰｨ 05ｨｹｨｪ ｨ ｨｪｨｪ ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ06ｨｦ ｨｨ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ ｨｮ ﾂ02ｨｪ06 09 ｨｨ ｨｹ040609ｨ ｨｪｨｨ02ｨｬ ｨ ｨｨ0601ｨｨｨｬｨｨ ﾂ0209ｨｺｨｨ01 ｨｨ ｨｬ060502ｨｺｨｮ050708ｨｪ06-ｨ ｨｨ ｨｨ ﾂ0209ｨｺｨｨ01 ｨｬ02ｨｰ0601ｨｨｨｺ, 04ｨ ﾂｨ 09ｨｰｨｮ06 09 ｨ 0209ｨｺ0502ｨｰ06 ﾂｨｪ ｨｨ09ｨｰ02ｨｬｨ 01, ｨｰ060001ｨ ｨｺｨ ｨｺ 09ｨｨ04ｨｮｨ 05ｨｨ04ｨ 02ｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ 09 ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｬ ｨｨ ｨｹ040609ｨ ｨｪｨｨ02ｨｬ ｨｺ05ｨ 0909ｨｨ ﾂ0209ｨｺｨｨ01 ｨｬ ｨｨ ﾂ0209ｨｺｨｨ ｨｮ 0708ｨｨ ﾂｨｨｨｪ 04ｨ ｨｰ08ｨｮ01ｨｪ02ｨｪｨ , 05ｨｰ ｨ ｨｪ06 09 ｨｰ02ｨｬ, ﾂｨｰ06 ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｺ ｨｬ060502ｨｺｨｮ0502 ｨ ｨｺｨｰｨｨｨｪｨ 06ｨ 05ｨ 01ｨ 06ｨｰ ｨｺ06ｨｦ ｨｨ04ｨ ｨｨ08ｨ ｨｰ0205ｨｹｨｪ0609ｨｰｨｹ06 ｨｨ, ｨｨ01ｨｨｨｬ06ｨｬｨｮ, ｨｰ 08ｨ 0405ｨｨ ﾂｨｪ ｨｬ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ (Milankov, De Boni, 1993; Gonsior et al., 1999). 0308ｨｨ 05ｨｰ06ｨｬ 01ｨ ｨｪｨｪｨ 07 ｨｨ04ｨ ｨｨ08ｨ ｨｰ0205ｨｹｨｪ0609ｨｰｨｹ ｨｪｨ ｨ ｨ 02ｨｰ0907 ｨｰ0605ｨｹｨｺ ｨｺ ｨｨｨｬ02ｨｪｨｰｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨ02ｨｬ ｨｪ ｨｬ06ｨｦ ｨｨｨｬｨｬｨｮｨｪ060005ｨｮ ｨｪ02ｨｨｨｨ, ｨｰ060001ｨ ｨｺｨ ｨｺ 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ 0108ｨｮ00ｨｨ01 ｨｬ060502ｨｺｨｮ050708ｨｪ06-ｨ ｨｨ ｨｨ ﾂ0209ｨｺｨｨ01 ｨｬ02ｨｰ0601ｨｨｨｺ (ｨｪｨ 0708ｨｨｨｬ0208, ｨｨｨｬｨｬｨｮｨｪ06ｨ 0506ｨｰｨｨｨｪ00ｨ ) ｨｰ ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｰ ｨｨ ｨｬ04 ｨ ｨｺｨｰｨｨｨｪｨ (Gonsior et al., 1999). 0207ｨｨ09ｨ ｨｪ04 ｨｰｨ ｨｺ ｨｮ ﾂｨ ｨｨ, ｨｺ060001ｨ 08ｨ 04ｨｪ0402 ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｺ ｨ ｨｺｨｰｨｨｨｪｨｮ ｨｮ ｨｪ06 06ｨｺ08ｨ 03ｨｨ09ｨ 06ｨｰ ｨｺ0502ｨｰ06 ﾂｨｪ ｨ, 0601ｨｪｨ ｨｺ06 ｨｪｨ ｨ ｨ 02ｨｬ ｨ ｨｰｨｰ0208ｨｪ04 ｨｬ02 ﾂ02ｨｪｨｨ07 08ｨ 0405ｨｨ ﾂｨ 06ｨｰ0907, ｨｺｨ ｨｺ ｨｮ- ﾂｨ ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｬｨｨ 2G2 ｨｨ 1C7 (Gonsior et al., 1999; Schoenenberger et al., 2005). 01ｨ ｨｨｨ ｨｰｨｪ06ｨｦ 00ｨｨ0706ｨｰ020406ｨｦ, 06ｨ ｨｲ0709ｨｪ0706ｨｴ02ｨｦ 01ｨ ｨｪｨｪｨｮ06 09ｨｨｨｰｨｮｨ 02ｨｨ06, ｨｬ0603ｨｪ06 09 ﾂｨｨｨｰｨ ｨｰｨｹ ｨｬ0603ｨｪ0609ｨｰｨｹ 09ｨｮｨｴ0209ｨｰ090609ｨ ｨｪｨｨ07 ｨｪ0209ｨｺ0605ｨｹｨｺｨｨ01 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｪ06-00ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ0401 ｨｺ06ｨｪ000608ｨｬｨ 02ｨｨｨｦ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ (Gonsior et al., 1999; Pederson, Aebi, 2005). 04ｨ ｨｪ0202 ｨｪｨ ｨｬｨｨ ｨｰｨ ｨｺ0302 ｨ 0405ｨｨ ｨｬ06ｨｪ09ｨｰ08ｨｨ080609ｨ ｨｪ04 09ｨｮｨｴ0209ｨｰ0902ｨｪｨｪ ｨ 0405ｨｨ ﾂｨｨ ｨ ｨｰｨｰ0208ｨｪｨ 01 06ｨｺ08ｨ 03ｨｨ09ｨ ｨｪｨｨ ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205, ｨｮ ﾂ02ｨｪｨｪ0401 ｨｺ C- ｨｨ N-ｨｺ06ｨｪ ｨｬ 0008ｨ 00ｨｬ02ｨｪｨｰｨ ｨｬ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ ( ｨ 0609ｨ, ｨ 0609ｨ, 2009). 01ｨ 0609ｨｪ0609ｨ ｨｪｨｨｨｨ ｨｮ ﾂ02ｨｪｨｪ ｨ ｨｪ ｨｮ05ｨｹｨｰｨ ｨｰ0609 ｨ ｨｨｨｪｨｮｨｰ ｨｪｨｨ02 06 ｨｰ06ｨｬ, ﾂｨｰ06 01ｨ ｨｪｨｪ0402 ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｨ04ｨ ｨｨ08ｨ ｨｰ0205ｨｹｨｪ ｨｰ ｨ 01 05ｨｬｨ 08ｨｨ06ｨｪ ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ06 08ｨ 0405ｨｨ ﾂｨｪ ｨｬ ｨｨ0006ｨｬ0208ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ ｨｹ ｨｪｨ 09ｨｰ0607ｨｴ02ｨｦ 08ｨ ｨ 06ｨｰ04 04ｨ ｨｺ0506 ﾂｨ 05ｨ 09ｨｹ ｨｮ ﾂ02ｨｪｨｨｨｨ ｨｪ ｨ ｨｪｨｪ ｨｬ0603ｨｪ06ｨｬ ﾂ02ｨｪｨｨｨｨ ｨｬ ｨ ｨｪ04ｨｬ 06ｨ 08ｨ 0406ｨｬ 00ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ ｨｬ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｨ ｨｰ08ｨ ｨｪ09ｨｰ0902ｨｪｨｪ06ｨｦ ｨ ｨｪｨｨ04ｨ 02ｨｨｨｨ ｨｬｨ ｨｰｨｨｨｪｨ ｨｰ ｨｬ04 ｨｨ04ｨｮ ﾂｨｨ05ｨｨ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ02 07ｨ ｨｰｨｰ0208ｨｪｨ ｨｨｨｬｨｬｨｮｨｪ060005ｨｮ ｨｪｨｰｨｪ ｨｺ08ｨ 03ｨｨ09ｨ ｨｪｨｨ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ ｨｮ01ｨｺ0502ｨｰ06 ﾂｨｪ ｨｬｨ 08ｨｨ06ｨｪｨ 01 ｨｬ0403ｨｨ 0708ｨｨ ｨｨ09ｨｺｨｮ0909ｨｰ0902ｨｪｨｪ06ｨｬ ｨｨｨｪ00ｨｨｨ ｨｨ080609ｨ ｨｪｨｨｨｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｨ ｨｬ02ｨｪｨｰｨ ｨｰｨｨ09ｨｪ ｨ 09ｨｴ ｨｪｨｨ ｨ ｨｰ0208ｨｨｨ 05 ｨｨ ｨｬ02ｨｰ0601ｨｨｨｺｨ ｨｮ ﾂ02ｨｪｨｨ02 05ｨｬｨ 08ｨｨ06ｨｪ ｨ ｨ 06ｨｰｨ ｨｪ02ｨｪｨ ｨｪｨ ｨｬ0403ｨ 01 BALB/c ｨｨ04 07ｨｨｨｰ06ｨｬｨｪｨｨｨｺｨ ｨ (05ｨ ｨｪｨｺｨｰ-0302ｨｰ0208ｨ ｨｮ0800) ｨｮ ﾂ02ｨｪｨｨ07 03ｨｨ0906ｨｰｨｪ ｨ ｨｰｨｨ080609ｨ ｨｪｨｪ04ｨｬ ｨｺ06ｨｬ ｨ ｨｬ02ｨｪｨｪ0609ｨｰｨｨ ｨｮ 09ｨ ｨｬ06ｨｺ ｨｨｨｪ01ｨｮ02ｨｨ080609ｨ 05ｨｨ 0609ｨｮ050702ｨｨ ｨ ｨｰ0205ｨｹｨｪ04ｨｬｨｨ 0601ｨｪ06ｨｺ08ｨ ｨｰｨｪ04ｨｬｨｨ ｨｨｨｪｨｲ02ｨｺ02ｨｨ07ｨｬｨｨ ｨｰ06 ﾂｨｪ (Folligon, Intervet, ｨ ｨｪ01ｨｨ07) ｨｨ ｨｨ06ｨｪｨｨ- 831

2 ｨ 0609ｨ 04ｨｨ ｨｺｨ 05ｨｨ04ｨ 02ｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ ｨ ｨｮ01ｨｺ0502ｨｰ06 ﾂｨｪ ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ. 06ｨｬｨ 08ｨｨ06ｨｪ04: ｨ ｪ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ0402, ｨ ｪ 06ｨ 08ｨ ｨ 06ｨｰｨ ｨｪｨｪ ｨ 0406ｨｦ, 09 ｪ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｨ ,6-01ｨｨ b-02-08ｨｨｨ 0600ｨｮ08ｨ ｨｪ0604ｨｨ05ｨ 02ｨｪ04ｨｨｨｬｨｨ01ｨ ｨｬ (DRB), 00 ｪ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｨ07 09 ｨ ｨｺｨｰｨｨｨｪ06ｨｬｨｨ02ｨｨｨｪ06ｨｬ D, 01 ｪ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｨ07 09 ｨ ｨｺｨｰｨｨｨｪ06ｨｬｨｨ02ｨｨｨｪ06ｨｬ D ｨｮ06ｨｴ02ｨｦ 06ｨ 08ｨ ｨ 06ｨｰｨｺ06ｨｦ ｨ 0406ｨｦ. 05ｨｰ080205ｨｺｨ ｨｬｨｨ 0706ｨｺｨ 04ｨ ｨｪ06 ｨｬ02 ﾂ02ｨｪｨｨ02 ｨｪｨ ｨｨ000208ｨｨｨｨ ｨｺ, ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ ｨｺ06ｨｰ ｨ 09ｨｰｨ 02ｨｰ ｨｨ09ｨｺｨｮ0909ｨｰ0902ｨｪｨｪ ｨ ｨｪｨｨ07 ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｨ 09ｨｪｨｨ03ｨ 02ｨｰ ｨ 08ｨ ｨ 06ｨｰｨｺｨｨ 05ｨｬｨ 08ｨｨ06ｨｪ ｨ 0406ｨｦ. ﾂ0209ｨｺ (Chorulon, Intervet, ｨ ｨｪ01ｨｨ07) 0006ｨｪｨ 0106ｨｰ080607ｨｨｨｪ ｪ ｨｪｨ 03ｨｨ0906ｨｰｨｪ0602 c ｨｨｨｪｨｰ020809ｨ 0506ｨｬ ｨｬ020301ｨｮ ｨｨｨｪｨｲ02ｨｺ02ｨｨ07ｨｬｨｨ 44 ｪ48 ﾂ ｨ 09ｨｰ 04ｨ ｨｦ 06ｨｰ09 ﾂｨｨｨｰ0409ｨ 05ｨｨ 06ｨｰ ｨｬ02ｨｪｨｨ ｨｨｨｪｨｲ02ｨｺ02ｨｨｨｨ ｨｨ06ｨｪｨｨ ﾂ0209ｨｺ ｨｪｨ 0106ｨｰ080607ｨｨｨｪｨ. 06ｨｬｨ 08ｨｨ06ｨｪ04 ｨｨ ｨｺｨ 05ｨｨ ｨｨ04 07ｨｦ ﾂ ﾂ ｨｨｨｪｨｲ02ｨｺ02ｨｨｨｨ ｨｨ06ｨｪｨｨ ﾂ0209ｨｺ ｨｪｨ 0106ｨｰ080607ｨｨｨｪｨ ｨｮ F10, ｨ ｨｴｨｮ06 HEPES (25 ｨｬ00, ｪ7.4) ｨ 05ｨｹｨｪ02ｨｦ0302ｨｦ 08ｨ ｨ 06ｨｰ04 ｨｨ ｨｹ040609ｨ 05ｨｨ 04ｨ ｨｨ, ｨｪ02 ｨｨｨｬ020903ｨｨ02 09ｨｨ01ｨｨｨｬ0401 ｨｬ ｨｨ ﾂ0209ｨｺｨｨ ｨｺｨｰ ｨｨｨｪ00ｨｨｨ ｨｨ080609ｨ ｨｪｨｨ07 ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ 05ｨｬｨ 08ｨｨ06ｨｪ04 ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ 05ｨｨ ｨｨｨｪｨｺｨｮｨ ｨ ｨｰ ｨｰｨ ｨｪ01ｨ 08ｨｰｨｪ0401 ｨｮ ｨｨ ｨｰ02 ﾂ02ｨｪｨｨ02 3 ﾂ (Whittingham, 1971), ｨ ｨｴ02ｨｦ 5,6-01ｨｨ b-02-08ｨｨｨ 0600ｨｮ08ｨ ｨｪ0604ｨｨ05ｨ 0204ｨｨｨｬｨｨ01ｨ (DRB; BIOMOL Research Labs) 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 500 ｨｬｨｺM, ｨｨ05ｨｨ 09 ｨｰ02 ﾂ02ｨｪｨｨ02 1 ﾂ , ｨ ｨｴ02ｨｦ ｨ ｨｺｨｰｨｨｨｪ06ｨｬｨｨ02ｨｨｨｪ D (Sigma, ) 09 ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ 5 ｨｬｨｺ00/ｨｬ ｨｬ ｨｬ ｨｮ ｨｪ ｨｮ ｨｪ ｨｰ ｨｪ ｨ 07 ｨｬ ｨｨ ｨｺ ｨｺ ｨｨ ｨｹ040609ｨ 05ｨｨ ｨｨｨｺ0506ｨｪｨ 05ｨｹｨｪ0402 ｨｺ080605ｨｨ ﾂｨｹｨｨ ｨ ｨｪｨｰｨｨｨｰ0205ｨ, ｨ ｨ 06ｨｰｨ ｨｪｨｪ ｨｰｨｨ ｨ 00ｨｬ02ｨｪｨｰｨ C-ｨｺ06ｨｪ ｨｬ02ｨｪｨ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ ( , Sigma, ) (08ｨ ｨｪｨｨ02 1 : 100), 05ｨｨｨ ｨｨｨｺ0506ｨｪｨ 05ｨｹｨｪ0402 ｨｺ080605ｨｨ ﾂｨｹｨｨ ｨ ｨｪｨｰｨｨｨｰ0205ｨ, ｨ ｨ 06ｨｰｨ ｨｪｨｪ ｨｰｨｨ ｨ 00ｨｬ02ｨｪｨｰｨ N-ｨｺ06ｨｪ ｨｬ02ｨｪｨ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ ( , Sigma, ) (08ｨ ｨｪｨｨ02 1 : 200) ｨｨ00ｨｨ ﾂｨｪ0609ｨｰｨｹ ｨ ｨｪｨｨ07 ｨ ｨｪｨｰｨｨｨｰ0205 ｨ 0405ｨ ｨｪｨ 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. 00 ｨｺｨ ﾂ0209ｨｰ ｨｰ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｨ ｨｹ040609ｨ 05ｨｨ ｨｺ0604ｨｹｨｨ ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｰｨｨ09 ｨｨｨｬｨｬｨｮｨｪ ｨ ｨｮ05ｨｨｨｪ0609 ｨｺ080605ｨｨｨｺｨ, ｨｺ06ｨｪｨｲ0600ｨｨ080609ｨ ｨｪｨｪ FITC (Sigma, ). 06ｨｬｨ 08ｨｨ06ｨｪ04 00ｨｨｨｺ09ｨｨ080609ｨ 05ｨｨ 09 ｨｰ02 ﾂ02ｨｪｨｨ02 40 ｪ60 ｨｬｨｨｨｪ 09 4%-ｨｪ06ｨｬ 08ｨ 09ｨｰ ｨ 08ｨ ｨｬｨ 05ｨｹ010200ｨｨ01ｨ ｨｪｨ PBS 0708ｨｨ ｨｺ06ｨｬｨｪｨ ｨｰｨｪ06ｨｦ ｨｰ02ｨｬ070208ｨ ｨｰｨｮ0802, ﾂ ｨｰｨｬ0409ｨ 05ｨｨ 09 ｨｰ ｨｨ0701 PBS. 02ｨ ｨｺ0502ｨｰｨｺｨｨ ｨｬ02ｨ ｨ ｨｨ05ｨｨ04ｨｨ080609ｨ 05ｨｨ %-ｨｪ06ｨｬ 08ｨ 09ｨｰ ｨｨｨｰ06ｨｪｨ X-100 ｨｪｨ PBS 09 ｨｰ02 ﾂ02ｨｪｨｨ02 10 ｨｬｨｨｨｪ ｨｮ06ｨｴ02ｨｦ ｨｰ080201ｨｺ08ｨ ｨｰｨｪ06ｨｦ 06ｨｰｨｬ0409ｨｺ06ｨｦ 09 PBS ｨｨ 10-ｨｬｨｨｨｪｨｮｨｰｨｪ06ｨｦ ｨｨｨｪｨｺｨｮｨ ｨ 02ｨｨ02ｨｦ 09 10%-ｨｪ06ｨｬ 08ｨ 09ｨｰ ｨｬｨ 08ｨｨ06ｨｪｨ 05ｨｹｨｪ06ｨｦ ｨｰｨｺｨｨ ｨｪｨ PBS ｨｰ0908ｨ ｨｴ02ｨｪｨｨ07 ｨｪ ｨｨ00ｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨ ｨｪｨｰｨｨｨｰ ｨ ｨｰ02ｨｬ 05ｨｬｨ 08ｨｨ06ｨｪ ｨｬ02ｨｴｨ 05ｨｨ 09 08ｨ 09ｨｰ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｨ ｨｨｨｪｨｺｨｮｨ ｨｨ080609ｨ 05ｨｨ 0708ｨｨ 4 縺05 ｨｪ02 ｨｬ02ｨｪ ﾂ ｨｰｨｬ0409ｨｺｨｨ 09 PBS ｨｨ 10-ｨｬｨｨｨｪｨｮｨｰｨｪ06ｨｦ ｨｨｨｪｨｺｨｮｨ ｨ 02ｨｨｨｨ 09 05ｨｬｨ 08ｨｨ06ｨｪｨ 05ｨｹｨｪ06ｨｦ ｨｰｨｺ02 04ｨ ｨｨ 0706ｨｬ02ｨｴｨ 05ｨｨ 09 08ｨ 09ｨｰ ｨｰ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｪｨ 60 ｨｬｨｨｨｪ 0708ｨｨ ｨｺ06ｨｬｨｪｨ ｨｰｨｪ06ｨｦ ｨｰ02ｨｬ070208ｨ ｨｰｨｮ ｨｰｨｬ0409ｨｺｨｨ 09 PBS 05ｨｬｨ 08ｨｨ06ｨｪ04 04ｨ ｨｺ0506 ﾂｨ 05ｨｨ ｨｮ VectaShield (Vector Laboratories, ). 08ｨｪｨ 05ｨｨ ｨ 08ｨ ｨｰ ｨｨ05ｨｨ ｨｬ06ｨｴｨｹ06 05ｨ ｨｪ ｨｺｨ ｨｪｨｨ08ｨｮ06ｨｴ ｨｺ06ｨｪ0006ｨｺｨ 05ｨｹｨｪ ｨｬｨｨｨｺ080609ｨｺ0607ｨ Leica TSC SL. 00 ｨｺｨ ﾂ0209ｨｰ0902 ｨｺ06ｨｪｨｰ080605ｨｹｨｪ0401 ｨｨ ｨｹ040609ｨ 05ｨｨ 05ｨｬｨ 08ｨｨ06ｨｪ04, 06ｨ 08ｨ ｨ 06ｨｰｨ ｨｪｨｪ0402 ｨｰ0605ｨｹｨｺ06 09ｨｰ060804ｨｬｨｨ ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｬｨｨ. 02 ｨ 08 ｨ ｨ 06 ｨｰ ｨｺ ｨｮ ｨ ｨｦ ( /ｨｬｨｺ05, Sigma, ) ｨｨ05ｨｨ 09 ｨｰ02 ﾂ02ｨｪｨｨ02 1 ﾂ 0708ｨｨ 37 縺05. 06ｨｪｨｰ020008ｨ 05ｨｹｨｪｨｮ06 ｨｨ ｨｪ ｨｰ 02 ｨｪ 09 ｨｨ 09 ｨｪ ｨｰ ｨｹ ｨｮ ｨｪ 02 ｨｨ ｨｨ ｨｪｨ ｨｺ06ｨｪ0006ｨｺｨ 05ｨｹｨｪ0401 ｨｨ0406ｨ 08ｨ 0302ｨｪｨｨ ｨｨ ｨｬ06ｨｴｨｹ ｨ ｨｬｨｬ04 ImageJ. 01ｨ ｨｺｨ ｨｬ 0607ｨｰｨｨ ﾂ0209ｨｺ06ｨｬ ｨｨ ｨｨ05ｨｨ 04ｨ ｨｬ ｨｪｨ ｨ 00ｨｬ02ｨｪｨｰｨ ｨ ｨｨ ｨ 00ｨｬ02ｨｪｨｰｨ ｨｪｨ 09ｨｪ02 ｨｺ0502ｨｰｨｺｨｨ ｨｺｨ ｨｦ ｨｨｨｨ ｨｺ06ｨｪ0006ｨｺｨ 05ｨｹｨｪ0401 ｨｨ0406ｨ 08ｨ 0302ｨｪｨｨｨｦ ｨ ｨｪ04ｨｬ 06ｨ 08ｨ 0406ｨｬ ｨ ｨｪｨ 05ｨｨ04ｨｨ080609ｨ 05ｨｨ ｨｰｨｨ ﾂ0209ｨｺｨｨ ｨ ｨｰ02ｨｬ 01ｨ ｨｪｨｪ0402 ｨｮ ｨｪ0705ｨｨ ｨｨ 08ｨ 0909 ﾂｨｨｨｰ0409ｨ 05ｨｨ 06ｨｰｨｪ060302ｨｪｨｨ02 ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｨ 0005ｨｮ ｨｪ02ｨｨｨｨ ｨ ｨｺ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｨ 0006ｨｪｨ ｨｺｨ ｨｦ 05ｨｺ ｨｨｨｬ02ｨｪｨｰｨ 05ｨｹｨｪ06ｨｦ

3 0508ｨ 09ｨｪｨｨｨｰ0205ｨｹｨｪ04ｨｦ ｨ ｨｪｨ 05ｨｨ04 01ｨ 08ｨ ｨｺｨｰ0208ｨ 0005ｨｮ ｨｪｨｰｨｪ ｨｬ02 ﾂ02ｨｪｨｨ ｨ ｨｪｨｪｨｨ01 05ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ ｨｨ ｨｺｨ 05ｨｨ04ｨ 02ｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ ｨ ｨｮ01ｨｺ0502ｨｰ06 ﾂｨｪ ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ. 06ｨｬｨ 08ｨｨ06ｨｪ04: ｨ ｪ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ0402, ｨ ｪ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｪ ﾂ in vitro, 09 ｪ 06ｨ 08ｨ ｨ 06ｨｰｨ ｨｪｨｪ ｨ 0406ｨｦ, 00 ｪ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｨ07 09 DRB, 01 ｪ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｨ07 09 ｨ ｨｺｨｰｨｨｨｪ06ｨｬｨｨ02ｨｨｨｪ06ｨｬ D. 05ｨｰ080205ｨｺｨ ｨｬｨｨ 0706ｨｺｨ 04ｨ ｨｪ06 ｨｬ02 ﾂ02ｨｪｨｨ02 06ｨｺ ｨｪ06ｨｦ 0406ｨｪ04 02ｨｨｨｰ060705ｨ 04ｨｬ04, ｨｪｨｨｨｺｨ 06ｨｴ ｨｨ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｨｨｨ 05ｨｬｨ 08ｨｨ06ｨｪ0609 in vitro. 0008ｨｮ ｨ ｨｪｨ 05ｨｨ04ｨｨ080609ｨ 05ｨｨ ｨｪ02 ｨｬ02ｨｪ ｨｬｨ 08ｨｨ06ｨｪ ｨｰ ｨｪｨｨ ｨｰ ｨｪ0609ｨｰｨｨ 08ｨ 04ｨｪｨｨ0204 ｨｬ020301ｨｮ 0904ｨ 0608ｨｺｨ ｨｬｨｨ ｨｨ ｨｹ040609ｨ 05ｨｨ t-ｨｺ08ｨｨｨｰ0208ｨｨｨｦ 05ｨｰｨｹ060102ｨｪｨｰｨ ｨｮ05ｨｹｨｰｨ ｨｰ04 08ｨ ｨｺ 0607ｨｨ09ｨ ｨｪ06 08ｨ ｨｪ0202 ( ｨ 0609ｨ, ｨ 0609ｨ, 2009; ｨ 0609ｨ, 03ｨ ｨｪ0609, 2012), 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ 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 03ｨｨ09ｨ ｨｪｨｨ ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ , ﾂ02ｨｬ 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 03ｨｨ09ｨ ｨｪｨｨ07 02ｨｨｨｰ060705ｨ 04ｨｬ04, 0708ｨｨ 05ｨｰ06ｨｬ 06ｨ 08ｨ ｨｴｨ 02ｨｰ ｨｪｨ 0902ｨ 07 09ｨｪｨｨｨｬｨ ｨｪｨｨ02 ｨ ｨｨｨ 02ｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ ｨｨ000208ｨｨ02ｨｦ ｨｺ (08ｨｨ09. 1, ｨ ). 0308ｨｨ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｨｨｨ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨｬｨ 08ｨｨ06ｨｪ ｨｰ02 ﾂ02ｨｪｨｨ02 3 ﾂ in vitro 01ｨ 08ｨ ｨｺｨｰ0208 ｨｬ02 ﾂ02ｨｪｨｨ07 ｨｪ02 ｨｨ04ｨｬ02ｨｪ0702ｨｰ0907 (ｨｪ ｨｺｨ 04ｨ ｨｪ06). 0308ｨｨ 06ｨ 08ｨ ｨ 06ｨｰｨｺ02 05ｨｬｨ 08ｨｨ06ｨｪ ｨ 0406ｨｦ 06ｨｺ08ｨ 03ｨｨ09ｨ ｨｪｨｨ ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｬｨｨ ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ ｨ ｨｪ0702ｨｰ0907, 0601ｨｪｨ ｨｺ06 01ｨ 08ｨ ｨｺｨｰ ｨｪ0209ｨｺ0605ｨｹｨｺ06 09ｨｨ0106ｨｨ04ｨｬ02ｨｪ0702ｨｰ0907 (08ｨｨ09. 1, ｨ ) , 04ｨｪｨ ﾂｨｨｨｰ0205ｨｹｨｪ06 09ｨｪｨｨ03ｨ 02ｨｰ0907 ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ ｨｬ02 ﾂ02ｨｪｨｨ07 ｨｪｨ ｨｨ000208ｨｨｨｨ ｨｺ. 04ｨ ｨｪｨｨ02 ｨ ｨｺｨｰｨｨｨｪｨ 09 ｨｪｨｮｨｺ ｨ 04ｨｬ ｨｨ06ｨ 0802ｨｰｨ 02ｨｰ ｨｬ02ｨｪ ｨｨ0000ｨｮ04ｨｪ04ｨｦ 01ｨ 08ｨ ｨｺｨｰ0208, ｨｮ ﾂｨ 0209 ｨｬ0603ｨｪ06 ｨｪｨ ｨ ｨ ｨｰｨｹ ｨｪｨｨ02 06ｨｰ010205ｨｹｨｪ ｨ 05ｨ 09ｨｰ02ｨｦ ｨ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ ｨｬ02 ﾂ02ｨｪｨｨ ｨｮ02ｨｰ ｨｰｨ ｨｺ ｨｰｨｬ02ｨｰｨｨｨｰｨｹ, ﾂｨｰ06 09 ｨ 0605ｨｹ03ｨｨｨｪ09ｨｰ ｨｮ ﾂｨ ｨ 05ｨｬｨ 08ｨｨ06ｨｪ0609, 06ｨ 08ｨ ｨ 06ｨｰｨ ｨｪｨｪ ｨ 0406ｨｦ, ｨｰ ｨｰ 06ｨｺ08ｨｮ0005ｨｮ ｨｬｨｮ (08ｨｨ09. 1, ｨ ). 0308ｨｨ ｨｨｨｪ00ｨｨｨ ｨｨ080609ｨ ｨｪｨｨｨｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｺｨ ｨｺ ｨｬ06ｨｴｨｹ06 DRB (08ｨｨ09. 1, 09), ｨｰｨ ｨｺ ｨｨ ｨｬ06ｨｴｨｹ06 ｨ ｨｺｨｰｨｨｨｪ06ｨｬｨｨ02ｨｨｨｪｨ D (08ｨｨ09. 1, 00) 06ｨ 08ｨ ｨｴｨ 02ｨｰ ｨｪｨ 0902ｨ 07 09ｨｪｨｨｨｬｨ ｨｪｨｨ02 ｨｮ09ｨｨ0502ｨｪｨｨ ｨｮ ｨｪ02ｨｨｨｨ ｨｪｨ ｨｨ000208ｨｨｨｨ ｨｺ. 0308ｨｨ 05ｨｰ06ｨｬ ｨｬ02ｨｪ0702ｨｰ0907 ｨｪ02 ｨｰ0605ｨｹｨｺ ｨｨ04ｨｮｨ 05ｨｹｨｪｨ 07 ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ, ｨｪ06 ｨｨ 09ｨ ｨｬ 01ｨ 08ｨ ｨｺｨｰ0208 ｨｬ02 ﾂ02ｨｪｨｨ07. 00ｨ 05ｨｨ04ｨｨ ｨｺ ｨｪｨ ﾂｨｨｨｪｨ 06ｨｰ ｨｰｨｹ0907 ｨｬｨｪ ﾂｨｨ090502ｨｪｨｪ ｨｰｨｪ ｨｺ08ｨｮ ｨ 05ｨ 09ｨｰｨｨ ｨ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ06ｨｦ 0005ｨｮ ｨｪ02ｨｨｨｨ (08ｨｨ09. 1, 09, 00) ｨ 08ｨ ｨ 06ｨｰｨｺｨｨ ｨ 0406ｨｦ ｨｺ0605ｨｨ ﾂ0209ｨｰ ｨ ｨｪ ｨｰ08ｨｮｨｺｨｰｨｮ08 ｨｨ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ ｨｨ ｨｮ ｨｪ02ｨｨｨｨ ｨｮｨｬ02ｨｪｨｹ03ｨ 06ｨｰ0907 (08ｨｨ09. 1, 01) ｨｪ ｨｬ02 ｨｨ 08ｨ 04ｨｬ0208ｨ ｨｬ 09ｨｰ08ｨｮｨｺｨｰｨｮ ｨ 0605ｨｹ0306ｨｬ ｨｺ0605ｨｨ ﾂ0209ｨｰ ｨｰ0802 ﾂｨ 06ｨｰ0907 ｨｨ ｨｪｨｰ08ｨ 05ｨｹｨｪ ｨ 05ｨ 09ｨｰ0701 ｨｪｨｮｨｺ ｨ 04ｨｬ04 (08ｨｨ09. 1, 01), ｨｺ06ｨｰ0608ｨ ｨｪ0609ｨｰｨｹ06 ｨｰ ｨｰ 01ｨｨ0000ｨｮ04ｨｪ04ｨｦ 01ｨ 08ｨ ｨｺｨｰ0208 ｨｬ02 ﾂ02ｨｪｨｨ07, ｨｪｨ ｨ ｨ 02ｨｬ04ｨｦ 09 ｨｺ06ｨｪｨｰ (08ｨｨ09. 1, ｨ ) ｨｺ08ｨ 03ｨｨ09ｨ ｨｪｨｨ07 ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨｬｨ 08ｨｨ06ｨｪ0609 ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｬｨｨ, ｨ ｨ 06ｨｰｨ ｨｪｨｪ04ｨｬｨｨ ｨｺ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ (08ｨｨ09. 2, ｨ ), ｨｪｨ ｨｰｨｨ09, ｨｺｨ ｨｺ ｨｮ ｨ ｨｴｨ ｨｹ, ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ 0005ｨｮ ｨｪ02ｨｨｨｨ 02ｨｨｨｰ060705ｨ 04ｨｬ , ﾂ02ｨｬ ｨ ( ｨ 0609ｨ, ｨ 0609ｨ, 2009) ﾂ02ｨｪｨｨ02 ｨｪｨｮｨｺ ｨ 04ｨｬ04 ｨｨｨｬ0202ｨｰ ｨｬ0205ｨｺ0601ｨｨ ｨｪ04ｨｦ 01ｨ 08ｨ ｨｺｨｰ0208, ｨ ｨｨｨ 02ｨｨ ｨｬ ｨ ｨｺｨｰｨｨｨｪｨ ｨｨ000208ｨｨ02ｨｦ ｨｺ ｨｪ ｨ 0302ｨｪｨ. 0308ｨｨ ｨｺｨｮ05ｨｹｨｰｨｨ09ｨｨ080609ｨ ｨｪｨｨｨｨ 05ｨｬｨ 08ｨｨ06ｨｪ ｨｰ02 ﾂ02ｨｪｨｨ02 3 ﾂ in vitro 07ｨ ｨｰｨｰ0208ｨｪ ｨｬ02 ﾂ02ｨｪｨｨ07 ｨｪｨｮｨｺ ｨ 04ｨｬ04 ｨｪ02 ｨｬ02ｨｪ0702ｨｰ0907, 0601ｨｪｨ ｨｺ ｨ 05ｨ 09ｨｰｨｨ 02ｨｨｨｰ060705ｨ 04ｨｬ04, ｨ ｨｪｨｨ ﾂｨｪ06ｨｦ ｨｬ, ｨｬ02 ﾂ02ｨｪｨｨ02 09ｨｰｨ ｨｪ0609ｨｨｨｰ0907 ｨ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ04ｨｬ (08ｨｨ09. 2, ｨ ), ﾂ ｨｪｨｨｨｺ060001ｨ ｨｪ02 ｨｪｨ ｨ ｨ 02ｨｰ0907 ｨｮ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨｬｨ 08ｨｨ06ｨｪ0609, 08ｨ 0409ｨｨ09ｨ 06ｨｴｨｨ in vivo. 0308ｨｨ 06ｨ 08ｨ ｨ 06ｨｰｨｺ02 05ｨｬｨ 08ｨｨ06ｨｪ ｨ 0406ｨｦ ｨｰｨ ｨｺ ｨｰｨｬ02 ﾂｨ 02ｨｰ0907 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ ｨｬ , 0601ｨｪｨ ｨｺ06 ｨｺｨ ｨｺ ｨｨｨ 06 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07 07ｨ ｨｰｨｰ0208ｨｪｨ 06ｨｺ08ｨ 03ｨｨ09ｨ ｨｪｨｨ07 ｨｪｨ ｨｬｨｨ ｨｪ ｨｪ06 (08ｨｨ09. 2, 09). 0308ｨｨ ｨｨ09ｨｺｨｮ0909ｨｰ0902ｨｪｨｪ06ｨｬ ｨ ｨｪｨｨｨｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨ06ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ ｨｪ0204ｨ -

4 ｨ 0609ｨ 04ｨｨ ｨｬ02ｨｪ02ｨｪｨｨ02 ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｨ 0005ｨｮ ｨｪ02ｨｨｨｨ ｨｮ01ｨｺ0502ｨｰ06 ﾂｨｪ ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ ｨｪｨ ｨｺ06ｨｪ0006ｨｺｨ 05ｨｹｨｪ0401 ｨｨ0406ｨ 08ｨ 0302ｨｪｨｨ ｨｦ09ｨｰ09ｨｨ07 DRB ｨｨ ｨ ｨ ｪ ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ, ｨ ｪ ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｺ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ ｨｰｨｨｨｺｨ 05ｨｹｨｪ ｨｰ080204ｨｺｨｨ ｨｰ0902ｨｰ09ｨｰ09ｨｮ06ｨｰ 0008ｨ ｨｪｨｨ02ｨ ｨｬ ｨｨｨｰ0205ｨｹｨｪ0401 ｨｨｨｪｨｰ020809ｨ ｨｮ080609ｨｪ07 04ｨｪｨ ﾂｨｨｨｬ0609ｨｰｨｨ ﾂｨｺｨ ｨｬｨｨ 06ｨ 0604ｨｪｨ ﾂ02ｨｪ04 04ｨｪｨ ﾂ02ｨｪｨｨ07, ｨｰ ｨｪ06 06ｨｰ05ｨｨ ﾂｨ 06ｨｴｨｨ ｨｰ ｨｺ06ｨｪｨｰ ｨｨ09ｨｨｨｬ06 06ｨｰ ｨｨ ｨｹ040609ｨ ｨｪｨｪ ｨｨｨｪ00ｨｨｨ ｨｨｨｰ0608ｨ ｨｰｨ ｨｺ0302 ｨｪ02 ｨｪｨ ｨ ｨ 02ｨｰ0907 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07 01ｨ 08ｨ ｨｺｨｰ0208ｨ 0005ｨｮ ｨｪｨｰｨｪ ｨｬ02 ﾂ02ｨｪｨｨ07 ｨｪｨｮｨｺ ｨ 04ｨｬ04 (08ｨｨ09. 2, 00, 01). 02ｨｰｨｬ02 ﾂｨ 02ｨｰ0907 ｨｰ0605ｨｹｨｺ06 ｨｮ ｨｨ09ｨ ｨｪｨｪｨ ｨｮ ｨｪ02ｨｨ07 06ｨｺ ｨｪ06ｨｦ 06ｨ 05ｨ 09ｨｰｨｨ 02ｨｨｨｰ060705ｨ 04ｨｬ04, ﾂｨｰ06, ｨｺｨ ｨｺ 0706ｨｺｨ ｨ 06ｨｰ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨｺ ｨｨｨｬ02ｨｪｨｰ04, ｨ ｨｪ06 09 ｨｮ ｨｨ07ｨｬｨｨ in vitro, ｨ ｨｪ ｨｦ09ｨｰ09ｨｨ02ｨｬ ｨｨｨｪ00ｨｨｨ ｨｨｨｰ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｪ02ｨｪｨｨ02 ｨｺ 09ｨｨ04ｨｮｨ 05ｨｹｨｪ04ｨｬ ｨｪｨ ｨ ｨｪｨｨ07ｨｬ ｨｪｨ ｨｬｨｨ ｨ ｨｪ 0908ｨ 09ｨｪｨｨｨｰ0205ｨｹｨｪ04ｨｦ ｨ ｨｪｨ 05ｨｨ04 ｨｨｨｪｨｰ020008ｨ 05ｨｹｨｪ06ｨｦ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｨ 0005ｨｮ ｨｪ02ｨｨｨｨ ｨｪｨ ｨｺ06ｨｪ0006ｨｺｨ 05ｨｹｨｪ0401 ｨｨ0406ｨ 08ｨ 0302ｨｪｨｨ0701 (08ｨｨ09. 3) ｨｨｨｪ00ｨｨｨ ｨｨ080609ｨ ｨｪｨｨ07 ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｬ06ｨｴｨｹ06 DRB ｨｮ ﾂｨ 02 ｨｨ ｨｹ040609ｨ ｨｪｨｨ07 ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨｨｦ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｨ 0005ｨｮ ｨｪ02ｨｨｨｨ ｨｬｨ 08ｨｨ06ｨｪ ｨ 09ｨｪ02ｨｪｨｨ06 09 ｨｺ06ｨｪｨｰ ｨｬ ｨｪ ｨｪ06 (4.69 ﾀ ｨｨ 4.89 ﾀ ｨｰ0902ｨｰ09ｨｰ0902ｨｪｨｪ06). 0308ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ C-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｪ06 09ｨｮｨｴ0209ｨｰ0902ｨｪｨｪ0602 ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｨ 0005ｨｮ ｨｪ02ｨｨｨｨ ｨ 09ｨｪ02ｨｪｨｨ06 09 ｨｺ06ｨｪｨｰ ｨｬ (6.23 ﾀ 0.66 ｨｨ 4.10 ﾀ ｨｰ0902ｨｰ09ｨｰ0902ｨｪｨｪ06). 06ｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ 0005ｨｮ ｨｪ02ｨｨｨｨ , ｨｬ02 ﾂ02ｨｪｨｪ ｨｬ06ｨｴｨｹ06 ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ, ｨｪ02 ｨｬ02ｨｪ0705ｨ 09ｨｹ ｨ 08ｨ ｨ 06ｨｰｨｺｨｨ 05ｨｬｨ 08ｨｨ06ｨｪ ｨ 0406ｨｦ (4.42 ﾀ ｨｰｨｨ ﾀ ｨｺ06ｨｪｨｰ ). 01ｨ ｨｰｨｨ09, ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ 0005ｨｮ ｨｪ02ｨｨｨｨ , 06ｨｺ08ｨ 0302ｨｪｨｪ ｨｬ06ｨｴｨｹ06 ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ, ｨｮ090205ｨｨ ﾂｨｨ09ｨ 05ｨ 09ｨｹ ｨ 08ｨ ｨ 06ｨｰｨｺｨｨ ｨ 0406ｨｦ ｨ 09ｨｪ02ｨｪｨｨ06 09 ｨｺ06ｨｪｨｰ ｨｬ 0706 ﾂｨｰｨｨ ｨ 04ｨ (8.62 ﾀ 0.98 ｨｨ 4.10 ﾀ ｨｰ0902ｨｰ09ｨｰ0902ｨｪｨｪ06). 02ｨ 09ｨｮ030102ｨｪｨｨ ｨｮ02ｨｰ 06ｨｰｨｬ02ｨｰｨｨｨｰｨｹ, ﾂｨｰ ｨ ｨｴｨ 07 ﾂｨ 09ｨｰｨｹ ｨｨ ｨ ｨｪｨｨｨｦ, ｨｪｨｰ ｨｪｨｨｨｬｨ ｨｪｨｨ07 ｨｺ06ｨｰ ｨｪｨ ｨｨｨｰ ｨｪ04ｨｦ ｨ ｨｺｨｰｨｨｨｪ, ｨｨｨｰ0907 ｨｪｨ ｨｺｨｮ05ｨｹｨｰｨｮ08ｨ ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺｨｨ01 ｨｺ0502ｨｰ06ｨｺ ｨｨｨｬ02ｨｪ02ｨｪｨｨ02ｨｬ ｨｮ06 06 ﾂ ｨｹ ｨ ｨｨ0601ｨｨｨｬｨｨ ﾂ0209ｨｺｨｨ01 ｨｨ ｨｬ060502ｨｺｨｮ050708ｨｪ06-ｨ ｨｨ ｨｨ ﾂ0209ｨｺｨｨ01 ｨｬ02ｨｰ0601ｨｨｨｺ ｨｰ06ｨｬｨｮ, ｨｪ0209ｨｬ06ｨｰ0807 ｨｪｨ ｨ ｨｺｨｰｨｨ09ｨｪ0402 ｨｨ ｨ ｨｪｨｨ ｨｨ ｨ ｨｺｨｰｨｨｨｪｨ ｨ ｨｪ ｨｬ02ｨｰｨ ｨ 0605ｨｨ04ｨｬｨ, 01ｨ ｨｪｨｪ ｨ ｨｪｨｨｨｨ 05ｨｰ ｨ 0205ｨｺｨ 09 09ｨｨ09ｨｰ02ｨｬ02 ｨｰ080201ｨｬ0208ｨｪ06ｨｦ ｨｪ06ｨｦ ｨ 0801ｨｨｨｰ02ｨｺｨｰ06ｨｪｨｨｨｺｨｨ 05ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｨ05ｨｨ ｨｨｨｰ0609 ｨｬ0502ｨｺ0607ｨｨｨｰｨ 06ｨｴｨｨ ｨｰｨ 06ｨｰ0907 ｨｺ08ｨ ｨｦｨｪ02 09ｨｺｨｮ01ｨｪ04ｨｬｨｨ ｨｨ 08ｨ ｨｪ02ｨｪｨｪ04ｨｬｨｨ (Nguyen et al., 1998; Johnson et al., 2003; Bohnsack et al., 2006; ｨ 0609ｨ, ｨ 0609ｨ, 2009; ｨ 0609ｨ, 03ｨ ｨｪ0609, 2012) ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺｨｨ01 ｨｺ0502ｨｰ06ｨｺ ｨｬ0502ｨｺ0607ｨｨｨｰｨ 06ｨｴｨｨ01 ｨｮｨ 0201ｨｨｨｰ0205ｨｹｨｪ ｨｬ06ｨｪ09ｨｰ08ｨｨ080609ｨ ｨｪ06 ｨｮ ﾂｨ 09ｨｰｨｨ02 ｨ ｨｺｨｰｨｨｨｪｨ ｨ 01 ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ, 09 ｨｰ06ｨｬ ﾂｨｨ ｨｮ ﾂｨ 09ｨｰｨｨ02ｨｬ ｨｨｨｬ0208ｨ 0404 II (Miralles, Visa, 2006; Obrdlik et al., 2007; Pederson, 2008; Hofmann 2009; Visa, Percipalle, 2010). ﾁｨｰ06 ｨｺｨ 09ｨ 02ｨｰ ｨｪｨｨｨｦ ｨ ｨｪｨｨｨｨ ｨ ｨｺｨｰｨｨｨｪｨ 09 ｨｮ ｨｨ ｨｪｨｨ0302ｨｪｨｨ07 ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨ06ｨｪｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ, ｨｰ06 06ｨｪｨｨ ｨｺ08ｨ ｨｦｨｪ02 ｨｪ02ｨｬｨｪ ﾂｨｨ090502ｨｪｨｪ ﾂｨ 09ｨｰｨｪ0609ｨｰｨｨ, 0706ｨｺｨ 04ｨ ｨｪ ｨｪｨｨ ｨｰｨｨ ｨ ｨｺｨｰｨｨｨｪ ｨｨ05ｨ ｨｬ02ｨｪｨｰ ｨｨ ｨｨ09ｨｺｨｮ0909ｨｰ0902ｨｪｨｪ06ｨｬ ｨ ｨｪｨｨｨｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｨｨｰｨ 01 ｨ ｨｬ00ｨｨｨ ｨｨｨｦ (Scheer et al., 1984). 06ｨｬ0206ｨｰ ｨｪｨｨ07 06 ｨｪｨ ｨｺ ｨｪｨｨｨｨ ｨ ｨｺｨｰｨｨｨｪｨ 09 ｨｺ05ｨ 09ｨｰ0208ｨ 01 ｨｨｨｪｨｰ ｨｬｨ ｨｰｨｨｨｪ ｨ ｨｪｨｮ05 09 ｨ ｨｪｨ ｨｨ ﾂｨｪ0401 ｨｮ ｨｨ0701 (Wang et al., 2006). 00 ｨｪｨ 0302ｨｦ 08ｨ ｨ 06ｨｰ02 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ02 09ｨｪｨｮｨｰ08ｨｨ ｨｪ ｨ ｨｰｨｰ0208ｨｪｨ 08ｨ ｨｪｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ 0708ｨｨ ｨｨ09ｨｺｨｮ0909ｨｰ0902ｨｪｨｪ06ｨｬ ｨ ｨｪｨｨｨｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｬ0603ｨｪ06 ｨ ｨｪｨ ｨ ｨ ｨｰｨｹ 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ, ｨｪ06 ｨｪ02 ｨｺ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ. 0308ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ DRB ｨｨ ｨ ｨｺｨｰｨｨｨｪ06ｨｬｨｨ02ｨｨｨｪｨ D ｪ ｨｨｨｪ00ｨｨｨ ｨｨｨｰ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ 09 08ｨ 04ｨｪ04ｨｬｨｨ ｨｬ0201ｨ ｨｪｨｨ04ｨｬｨ ｨｬｨｨ 0102ｨｦ09ｨｰ09ｨｨ07 (Perry, Kelley, 1970; Sehgal et al., 1976; Zandomeni, Weinmann, 1984) ｪ 08ｨ ｨｪｨｨ02 ｨ ｨｺｨｰｨｨｨｪｨ ｨ 01 05ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｨ04ｨｬ02ｨｪ ｨｹ ｨｪ04ｨｬ 06ｨ 08ｨ 0406ｨｬ ｨ 06ｨｨ ｨｮ ﾂｨ ｨｰｨｬ02 ﾂｨ 05ｨ 09ｨｹ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ ｨｪｨ 01, ｨ ｨｨｨｨ080609ｨ ｨｪｨｪ ｨｨ000208ｨｨ02ｨｦ ｨｺ. 01ｨ 0906ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺｨｨ01 ｨｺ0502ｨｰｨｺｨ 01 ｨｬ0502ｨｺ0607ｨｨｨｰｨ 06ｨｴｨｨ01 ｨ ｨｺｨ 04ｨ ｨｪ06, ﾂｨｰ ｨｨ 0209ｨｰ0209ｨｰ0902ｨｪｨｪ06ｨｬ 04ｨ ｨｬ ｨｪｨｨｨｨ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ( ｨｰ0607ｨｪｨｨｨｨ 04ｨｨｨｬｨｪ02ｨｦ ﾂｨｺｨｨ) ｨｨｨｬ0202ｨｰ ｨｬ0209ｨｰ06 ｨｰ0209ｨｪ ｨ ｨｨｨｬ060102ｨｦ09ｨｰ09ｨｨ02 ｨｬ020301ｨｮ ｨｰ0205ｨｹ02ｨ ｨｬｨｨ 08ｨ 01ｨ 05ｨ ｨｨ ｨｺｨ ｨｬｨｨ (Malatesta et al., 1994) ｨｺｨｨ ｪ ｨ 08ｨ 04ｨｪ ｨｰ08ｨｮｨｺｨｰｨｮ0804, 01ｨ 08ｨ ｨｺｨｰ0208ｨｪ ｨ ｨｪｨｪｨｨ01 09ｨｰｨ 01ｨｨｨｦ 08ｨ 0409ｨｨｨｰｨｨ07 ｨｬ0502ｨｺ0607ｨｨｨｰｨ 06ｨｴｨｨ01, ｨｰ09ｨｨｨｨ 01ｨ 06ｨｴｨｨ02 ｨｪｨ ﾂｨ 0506 ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨ06ｨｪｨｪ06 ｨ ｨｺｨｰｨｨ09ｨｪ04ｨｬ ｨｺｨ ｨｬ (Fl*echon, Kopecny, 1998) ｨｬ02 ｨｰ060006, ｨｬ06ｨｪ09ｨｰ08ｨｨ080609ｨ ｨｪ06 ｨｨ01 ｨｮ ﾂｨ 09ｨｰｨｨ ｨ 01 ｨ ｨｨ060002ｨｪ0204ｨ ｨｰ ｨ 01ｨ 05ｨ (Ferreira, Carmo-Fonseca, 1995) ｨｨ09ｨｺ0506 ﾂ02ｨｪ06, ﾂｨｰ ｨｺｨｰ08 00ｨｮｨｪｨｺ02ｨｨｨｦ 05ｨｰｨｨ ｨｪ ｨｬ02ｨｪ ｨｪｨ ﾂｨｨｨｰ0205ｨｹｨｪ06 03ｨｨ0802, ﾂ02ｨｬ ｨ 00ｨ 02ｨｰ ｨｪ ｨｪ ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ ｨ ｨｨｨｬ060102ｨｦ09ｨｰ09ｨｨ02 ｨ ｨｺｨｰｨｨｨｪｨ ｨｬｨ ｨｰｨｨｨｪ06ｨｬ ｨｬ06ｨｪ09ｨｰ08ｨｨ080609ｨ ｨｪ ｨｬ ｨ ｨ 06ｨｰ, ｨｪｨ 0906ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺｨｨ01 ｨｺ0502ｨｰｨｺｨ 01 ｨｬ0502ｨｺ0607ｨｨｨｰｨ 06ｨｴｨｨ01 (Papoulas et al., 1998; Zhao et al., 1998; Shen et al., 2000; Hofmann, 2009; Visa, Percipalle, 2010) ｨｪ06 ｨ ｨｨ01ｨ ｨｰｨｹ, ﾂｨｰ ｨｨ 06ｨ 08ｨ ｨ 06ｨｰｨｺ ｨ 08ｨ ｨｰ ｨ 0406ｨｦ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ ｨｬ02 ﾂ02ｨｪｨｨ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ ｨ ｨｮ0102ｨｰ 09ｨｪｨｨ03ｨ ｨｰｨｹ ｨ 09 ﾂ02ｨｰ ｨｮ01ｨ 0502ｨｪｨｨ07 07ｨｮ05ｨ ｨｬ060502ｨｺｨｮ05, ｨｪｨ ｨｴｨｨ ｨｬ06ｨｦ ｨ ｨｨｨ 02ｨｨｨｨ ｨｪｨ ｨｺ06 ｨｬ04 ｨｪｨ ｨ ｨ 05ｨｨ ｨｬ ｨｰｨｨ ｨｪ04ｨｦ ｨｮ05ｨｹｨｰｨ ｨｰ: ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ ｨｬ02 ﾂ02ｨｪｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ 0708ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ0504 ｨｮ090205ｨｨ ﾂｨｨ09ｨ 05ｨ 09ｨｹ. 02ｨ ｨｪｨｪ0602 ｨｪｨ ｨ ｨｪｨｨ02 ｨｬ0603ｨｪ06 06ｨ ｨｲ0709ｨｪｨｨｨｰｨｹ ｨｰ02ｨｬ, ﾂｨｰ06 09 ｨｪｨ ｨｰｨｨ09ｨｪ06ｨｬ ｨｰ0607ｨｪｨｨｨｨ ﾂｨ 09ｨｰｨｹ ｨｬ060502ｨｺｨｮ05 ｨ ｨｺｨｰｨｨｨｪｨ, ｨｪｨ ｨｴｨｨ ｨｬ06ｨｬ 0904ｨ ｨｨｨｬ060102ｨｦ09ｨｰ09ｨｨｨｨ ｨｪ , ｨ 09 ｨ 0205ｨｺｨ ｨｬｨｨ ｨｬｨ ｨｰｨｨｨｪｨ, ｨｬ060302ｨｰ ｨ 04ｨｰｨｹ 04ｨ ｨｺ0804ｨｰｨ ｨ ｨｨｨｬ060102ｨｦ09ｨｰ09ｨｨ07 09 ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｬｨｨ. 0308ｨｨ ｨｮ01ｨ 0502ｨｪｨｨｨｨ ｨ ｨｦｨｰ ｨ ｨｪｨｨ07 09ｨｰｨ ｨｪ060907ｨｰ0907 ｨ ｨｰｨｮ07ｨｪ04ｨｬｨｨ ｨｦ09ｨｰ09ｨｨ07 ｨ ｨｪｨｰｨｨｨｰ0205, ｨｨ ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ ｨｬ02 ﾂ02ｨｪｨｨ ｨ 09ｨｰｨ 02ｨｰ ｨ ｨｺ0506 ﾂ02ｨｪｨｨ ｨｮ02ｨｰ 06ｨｰｨｬ02ｨｰｨｨｨｰｨｹ, ﾂｨｰ ｨｪｨ ｨ ｨ 02ｨｬ0402 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07 08ｨ ｨｪｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ ｨ 01 05ｨｬｨ 08ｨｨ06ｨｪ ｨｰｨｬ02 ﾂｨ 05ｨｨ09ｨｹ ｨｰ0605ｨｹｨｺ ｨｨ ｨｨ ｨｹ040609ｨ ｨｪｨｨｨｨ ｨ ｨｪｨｰｨｨｨｰ0205 ｨｺ 05-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ ｨｬ060502ｨｺｨｮ ｨ ｨｪ0202 ｨｬ ｨ 00ｨ 05ｨｨ, ﾂｨｰ06 ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｺ 05- ｨｨ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨ ｨｬ ｨｬ060502ｨｺｨｮ0504 ｨ ｨｺｨｰｨｨｨｪｨ ｨｰ 08ｨ 04ｨｪ ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ ｨｬ04 ｨ ｨｺｨｰｨｨｨｪｨ, ｨｪｨ ｨｴｨｨ ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ ｨ ｨｨｨｬ ｨｬｨ ｨｰｨｨｨｪ06ｨｬ ｨｨ 09 ｨ 0707ｨ 08ｨ ｨｰ06ｨｬ ｨｰ08ｨ ｨｪ09ｨｺ08ｨｨ0702ｨｨｨｨ ｨｰ0902ｨｰ09ｨｰ0902ｨｪｨｪ06 ( ｨ 0609ｨ, ｨ 0609ｨ, 2009). 02ｨ ｨｪｨｪ0402, ｨｮ ﾂ02ｨｪｨｪ ｨｪｨ 09ｨｰ0607ｨｴ02ｨｦ

5 130508ｨ 09ｨｪｨｨｨｰ0205ｨｹｨｪ04ｨｦ ｨ ｨｪｨ 05ｨｨ04 01ｨ 08ｨ ｨｺｨｰ0208ｨ 0005ｨｮ ｨｪｨｰｨｪ ｨｬ02 ﾂ02ｨｪｨｨ ｨ ｨｪｨｪｨｨ01 05ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ ｨ ｨ 06ｨｰ02, 0909ｨｨ0102ｨｰ0205ｨｹ09ｨｰ09ｨｮ06ｨｰ ｨｰｨｨ09 01ｨ ｨｪｨｪ ｨｪｨｨ ｨｰｨｪ06, ﾂｨｰ06, ｨｪ0209ｨｬ06ｨｰ0807 ｨｪｨ ｨｪｨｮ06 ｨｨｨｪｨｰ02ｨｪ09ｨｨ09ｨｪ0609ｨｰｨｹ ｨ ｨｪｨｨ07 01ｨ ｨｪｨｪ0401 ｨ ｨｪｨｰｨｨｨｰ ｨ ｨｺｨｰｨｨｨｪ06ｨｬ ｨｪｨ ｨｨｨｬｨｬｨｮｨｪ06ｨ 0506ｨｰｨ 01 ( ｨ 0609ｨ, 03ｨ ｨｪ0609, 2012), ｨ ｨｪｨｰｨｨｨｰ0205ｨ ｨｺ N-ｨｺ06ｨｪ ｨｬｨｮ 0106ｨｬ02ｨｪｨｮ 09 09ｨｨ05ｨｮ ｨｨ01 00ｨｨ04ｨｨｨｺ06-01ｨｨｨｬｨｨ ﾂ0209ｨｺｨｨ ｨ 02ｨｪｨｪ0609ｨｰ02ｨｦ 06ｨ 05ｨ 01ｨ 06ｨｰ ｨｬ02ｨｪｨｹ0302ｨｦ ｨ ｨｪ0609ｨｰｨｹ ｨ ｨｰｨｹ ｨｪ04ｨｬ ｨ ｨｺｨｰｨｨｨｪ06ｨｬ 0708ｨｨ ｨｬ02 ﾂ02ｨｪｨｨｨｨ ｨｰ06ｨｰｨ 05ｨｹｨｪ ｨ 08ｨ ｨｰ ｨｬｨ 08ｨｨ06ｨｪ0609 ｨｬ0403ｨｨ. 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 ｨｬｨｬ ｨｺｨｮ050708ｨｪｨ 07 ｨｨ ｨｺ0502ｨｰ06 ﾂｨｪｨ 07 ｨ ｨｨ ｨｨ ｨｨ0906ｨｺ 05ｨｨｨｰ0208ｨ ｨｰｨｮ ｨ 0609ｨ , ｨ 0609ｨ ｨｺｨ 05ｨｨ04ｨ 02ｨｨ07 ｨ ｨｺｨｰｨｨｨｪｨ ｨ ｨｮ01ｨｺ0502ｨｰ06 ﾂｨｪ ｨ ｨｦ ｨｬ0403ｨｨ. 00ｨｨｨｰ ｨｨ (8) : 663 ｪ ｨ 0609ｨ , 03ｨ ｨｪ ｨ 02ｨｪｨｪ0609ｨｰｨｨ ｨｨｨｬｨｬｨｮｨｪ060005ｨｮ ｨｪｨｰｨｪ ｨｪｨｨ ｨｪ ｨ ｨｺｨｰｨｨｨｪｨ 09 08ｨ ｨｪｨｪｨｨ01 05ｨｬｨ 08ｨｨ06ｨｪｨ 01 ｨｬ0403ｨｨ. 00ｨｨｨｰ ｨｨ (7) : 541 ｪ ｨ ｨｪ , 01ｨ 05ｨ ｨｺｨｰｨｨ06ｨｪ ｨｪｨｮｨｰ08ｨｨ ｨｪ0402 ｨ ｨｺｨｰｨｨｨｪ ｨｬｨｨｨｺ080600ｨｨ05ｨ ｨｬ02ｨｪｨｰ ｨｨｨｰｨ 01 ｨｰ08ｨ 0907ｨｪ06ｨｦ ｨｮ03ｨｺｨｨ. 00ｨｨｨｰ ｨｨ07. 29(2) : 142 ｪ149. 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Chem. 259 : ｪ Zhao K., Wang W., Rando O. J., Xue Y., Swiderek K., Kuo A., Cradtree G. R Rapid and phosphoinositol-dependent binding of the SWI/SNF-like BAF complex to chromatin after T-lymphocyte receptor signalling. Cell. 95 : 625 ｪ ｨｰｨｮ07ｨｨ05ｨ 2 VII 2012 A COMPARATIVE STUDY OF FLUORESCENT PATTERNS IN THE NUCLEI OF EARLY MOUSE EMBRYOS USING ANTIBODIES AGAINST DIFFERENT DOMAINS OF THE ACTIN MOLECULE I. O. Bogolyubova Institute of Cytology RAS, St. Petersburg; mail: ibogol@mail.ru In this work, new data on the peculiarities of actin immunofluorescent detection in 2-cell mouse embryos using antibodies against C- and N-terminal domains are presented. We studied the distribution of nuclear actin after artificial suppression of transcription and after enzymatic digestion of DNA. The visual observations were

6 ｨ 0609ｨ supplemented with morphometric analyses of confocal images. In both experimental groups (treated by transcription inhibitors or DNase) the reliable increasing of fluorescence intensity is revealed when antibody against C-, but not against N-terminal domain of actin was used. These finding allows to suppose that antibody against C-terminal domain of actin is suitable with a high efficiency for immunofluorescent studies on the nuclei of cleavage mouse embryo. K e y w o r d s: preimplantation mouse embryos, nuclear actin, transcription, laser scanning confocal microscopy.