Tsitologiya 55 (6)

Transcription

1 ｨｬ 55, ﾁ ｨ 0108ｨ 01ｨｨｨｬ0609, ｨｮ , ｨｪｨｨ09ｨｨｨｬ ｨ 04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0601ｨｨｨｬｨｨｨｨ ｨｨ ｨ ｨｨ0600ｨｨ04ｨｨｨｺｨｨ 08ｨ , ｨｺｨｰ0806ｨｪｨｪ04ｨｦ ｨ : abdfar@mail.ru ｨ 0108ｨ 01ｨｨｨｬ0609 ｨｨ ｨｨ07ｨｪｨｨ02 00ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨｪｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｪｨｮ ｨ ｨｪｨｨ04ｨ 02ｨｨ06 ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ02ｨｦ ｨｺｨｮｨｺｨｮ08ｨｮ ｨｪｨ 05ｨｨ04ｨｨ080609ｨ 05ｨｨ 0905ｨｨ07ｨｪｨｨ02 00ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ ｪ4 0003ｨ ｨｪｨ ｨｰ ｨｨ ｨ ｨｪｨｨ04ｨ 02ｨｨ06 ｨｺ0502ｨｰ06ｨｺ ｨｰｨｺ0609 ｨｺｨｮｨｺｨｮ08ｨｮ ｨｨ 01ｨ ｨｪｨｨｨｨ ｨ 06ｨｰｨｬ02 ﾂ02ｨｪ ｨｰ0502ｨｪｨｨ02 ｨｬｨ ｨｰ08ｨｨｨｺ09ｨ ｨｬｨｨｨｰ060106ｨｪ0108ｨｨｨｦ, ｨｺ05ｨ 09ｨｰ0208ｨｨ04ｨ 02ｨｨ ｨｬ02ｨｪｨｰ ｨｪ ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺ06ｨｦ 0902ｨｰｨｨ 09 06ｨ 08ｨ ｨ ｨｪｨｨ02ｨｬ ｨｮ ﾂｨ 09ｨｰｨｺ ｨ 09ｨｰ060705ｨ 04ｨｬ04 ｨｨ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ ｨｨｨｰ060705ｨ 04ｨｬ02 ﾂｨｨ0905ｨ ｨｬ0205ｨｺｨｨ01 09ｨ ｨｺｨｮ060502ｨｦ. 01ｨ ｨｰ08ｨ ｨｪ09-09ｨｰ060806ｨｪ02 01ｨｨｨｺｨｰｨｨ060906ｨｬ 04ｨ 00ｨｨｨｺ09ｨｨ080609ｨ ｨｪ06 ｨｪｨ ｨｺ ｨｪｨｨ ｨｨｨｺｨｮ ｨｺｨｰ0806ｨｪｨｪ ｨｰｨｪ04ｨｬ ｨｨｨｬ04ｨｬ, ﾂｨｰ06 ｨｮｨｺｨ ｨ 02ｨｰ ｨｪｨ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ02 ｨｰ08ｨ 00ｨｨｨｺｨ 09 05ｨｪ0106ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪ06ｨｦ 09ｨｨ09ｨｰ02ｨｬ ｨｮｨｺ02ｨｨ ｨｺｨｰ0806ｨｪｨｪ06ｨｦ ｨｰｨｪ0609ｨｰｨｨ ｨｬ02ｨｬｨ 08ｨ ｨｪ04 ｨｰ06ｨｪ060705ｨ 09ｨｰｨ ｨ 05ｨ 09ｨｹ ｨｮｨｬ02ｨｪｨｹ0302ｨｪｨｨ02ｨｬ 09 09ｨ ｨｺｨｮ ｨｺ0605ｨｨ ﾂ0209ｨｰ09ｨ 0609ｨｬｨｨ0600ｨｨ05ｨｹｨｪ ｨｺ0506 ﾂ02ｨｪｨｨｨｦ ｨｪｨｨ02 01ｨ ｨｪｨｨ ｨ 0706ｨｺｨ 04ｨ ｨｪｨｨｨｺｨｪ060902ｨｪｨｨ02 ｨ 05ｨｹｨｰ0208ｨ 02ｨｨｨｦ ｨｰ08ｨｮｨｺｨｰｨｨ09ｨｪ ｨ 08ｨ ｨｺｨｰ0208ｨ 09 ｨｰ06ｨｪ060705ｨ 09ｨｰ02. 01ｨｨ ｨｪ06ｨｬ ｨｨ04 09ｨ 08ｨｨｨ ｨｪｨｰ0609 ｨｪ ｨｹ ｨｪｨ 08ｨｮ0302ｨｪｨｨ ｨｰｨｪ0609ｨｰｨｨ 0705ｨ 04ｨｬｨ 0502ｨｬｨｬ04, ﾂｨｰ ｨｰ ｨ ｨｹ 06ｨｺ08ｨ 09ｨｺ06ｨｦ 0609ｨ ｨｪ0906ｨｬ ｨｮｨ 04ｨｬ. 0308ｨｨ 09ｨ ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ ｨ ｨｰｨｬ ｨｪ ﾂｨ 09ｨｰ06ｨｰｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｪ0401 ｨｪｨ 08ｨｮ0302ｨｪｨｨｨｦ ｨｬ02ｨｬｨ 08ｨ ｨｪ04 ｨｰ06ｨｪ060705ｨ 09ｨｰｨ ｨ 0405ｨ ｨｨ06ｨｪｨ 05ｨｹｨｪｨ 08ｨ 04ｨｪｨｨ ｨ ｨｪｨｨｨｦ: ｨｬｨｨｨｪｨｨｨｬｨ 05ｨｹｨｪ06ｨｦ 0708ｨｨ 09ｨ ｨｰ ｨ ｨｨ ｨｬｨ ｨｺ09ｨｨｨｬｨ 05ｨｹｨｪ06ｨｦ 0708ｨｨ 09ｨ ｨｰ ｨ. 06ｨ ｨｺｨｨｨｬ 06ｨ 08ｨ 0406ｨｬ, ｨｬ ｨ 02ｨｬ ｨｰ06ｨｬ, ﾂｨｰ ｨｪ06ｨｦ ｨｨ04 ｨｬｨｨ0302ｨｪ02ｨｦ 0905ｨｨ07ｨｪｨｨ07 01ｨ ｨｪｨｨ07 ｨｪｨ ｨｺ0502ｨｰｨｺｨｨ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ ｨｰ ｨｪ0106ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪｨ 07 09ｨｨ09ｨｰ02ｨｬｨ ﾂ ｨ :01ｨ ｨｪｨｨ02, 09ｨｰ , 05ｨｪ ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺｨ ｨｰｨｹ, ｨｰｨｺｨｨ ｨｺｨｮｨｺｨｮ08ｨｮ ｨｰｨｪ06, ﾂｨｰ06 09ｨｪ0203ｨｪ ｨ ｨｪｨｨ ｨｨ0702ｨｰ ｨｪｨ 00ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ04ｨｦ 09ｨｰｨ ｨｰｨｮ09 08ｨ 09ｨｰ02ｨｪｨｨｨｦ ｨｨ 03ｨｨ0906ｨｰｨｪ ｨ ｨｺ ｨｰ0208ｨｬ0601ｨｨｨｪｨ ｨｬｨｨ ﾂ0209ｨｺｨｨｨｦ 07ｨ 08ｨ ｨｬ02ｨｰ08 01ｨ ｨｪｨｨ02 09 ｨ ｨｨ ｨｬ05ｨｨ 09ｨ 08ｨｹｨｨ08ｨｮ02ｨｰ 06ｨｰ ｨ ｨｪｨ ｨｰ02 06ｨｺ ｨｺｨｬ 0106 ｨ ﾂ02ｨｬ ｨ 09 06ｨｺ02ｨ ｨｪ09ｨｺｨｨ ｨｮｨ ｨｨｨｪｨ ｨ 09ｨｰ02ｨｪｨｨｨｦ 0706ｨｺｨ 04ｨ ｨｪ06, ﾂｨｰ ｨｰｨｨｨｬｨｮｨｬ ｨ ｨｰｨｬ ｨｪ ｨ ｨｪｨｨ ｨｨｨｰ ｨｪｨ ｨｰｨ ｪ2800 ｨｬ, ｨｨ 06ｨ ｨｪｨ 08ｨｮ03ｨｨ09ｨ 02ｨｰ0907 ｨｨ01 ｨｬｨ ｨｺ09ｨｨｨｬｨ 05ｨｹｨｪｨ ｨｮｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｹ. 05 ﾂｨｨｨｰｨ 02ｨｰ0907, ﾂｨｰ ｨ ｨｪｨｨ02 ｨｰ08ｨ ｨｪ ｨｰｨ, 0006ｨｰ0609ｨｨｨｪｨｰ0204ｨ ｨｨ ｨｰｨ ｨｪｨｨ ｨｮｨ ｨ 05ｨｹ07ｨｨｨｦ09ｨｺ ｨ ｨ ｨｪ06 ｨｨｨｬ02ｨｪｨｪ ｨｪｨｨ02ｨｬ ｨ ｨｰｨｬ ｨｪ ｨ ｨｪｨｨ07 (01ｨ ｨｬｨ 0502ｨｦ, 2004) ｨｬｨ 07 00ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪｨ ｨｹ 01ｨ ｨｪｨｨ07 08ｨ 0909ｨｬｨ ｨｰ08ｨｨ09ｨ 02ｨｰ ｨｰ06 ﾂｨｺｨｨ ｨｪｨｨ ｨｨ03ｨｮｨｴ02ｨｦ 09ｨｨ0504 ｨｬｨ ｨｪ0609ｨ ｨｪ ｨ 09ｨｰ (Knoblauch, Peters, 2010) ｨｰ0209ｨｰ0902ｨｪｨｪ0401 ｨｮ ｨｨ ｨｨｨ 07ｨ 0406ｨｪ ｨｪ ｨｰ02ｨｪ02ｨｨｨ 05ｨ ｨｺ0502ｨｰ06ｨｺ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ 09 04ｨ 09ｨｨ09ｨｨｨｬ0609ｨｰｨｨ 06ｨｰ 09ｨｨ01ｨ ｨｺ060502ｨ 0502ｨｰ ｨｰ ｨC ｨC(4 ｪ5) M03a ｪ 0708ｨｨ 09ｨｨ05ｨｹｨｪ06ｨｬ 06ｨ ｨｨ09ｨ ｨｪｨｨｨｨ (Galm*es et al., 2011). 05ｨｪｨｨ0302ｨｪｨｨ02 01ｨ ｨｪｨｨ07 ｨｪｨｨ ｨｨ04ｨｨ ｨｨ ﾂ0209ｨｺｨｨ01 04ｨｪｨ ﾂ02ｨｪｨｨｨｦ ｨ 02ｨｰ ｨｹ0204ｨｪ04ｨｬｨｨ ｨ 0302ｨｪｨｨ07ｨｬｨｨ ｨｬ02ｨｰｨ ｨ 0605ｨｨ04ｨｬｨ ｨｨ 00ｨｨｨ 0205ｨｹ06 08ｨ 09ｨｰ02ｨｪｨｨｨｦ ｨｬ06ｨｰ0807 ｨｪｨ 0706 ﾂｨｰｨｨ 09ｨｰ060502ｨｰｨｪ0606 ｨｨ09ｨｰ0608ｨｨ06 ｨｨ04ｨｮ ﾂ02ｨｪｨｨ07 01ｨ 05ｨｹｨｪ ｨｰ08ｨ ｨｪ ｨｰｨ 08ｨ 09ｨｰ , ｨ 01ｨｨ02ｨｪｨｰｨ 01 01ｨ ｨｪｨｨｨｦ, ｨｰ0802ｨ ｨｮ06ｨｴｨｨ ｨｪ0609ｨ 08ｨ 09ｨｰ ｨｬ020301ｨｮ ｨ ｨｺ020207ｨｰ060806ｨｬ ｨｨ 0106ｨｪ060806ｨｬ, 0609ｨｰｨ 02ｨｰ0907 ｨｪ ｨｪｨｪ04ｨｬ (Knoblauch, Peters, 2010; Turgeon, 2010). 0604ｨｬ02ｨｪ02ｨｪｨｨ02 01ｨ ｨｪｨｨ07 09 ｨｺ0502ｨｰ06 ﾂｨｪ0401 ｨｺ06ｨｬ07ｨ 08ｨｰｨｬ02ｨｪｨｰｨ ｨｨｨｰ ｨｪ ｨｮ050702ｨｨｨｨ ｨｬｨｪ ｨｰ09ｨ 00ｨｨ04ｨｨ ｨｨ ﾂ0209ｨｺｨｨ : ｨｮ09ｨｰｨｹｨｨ ﾂｨｪ ｨｨ0302ｨｪｨｨｨｦ (Ache et al., 2010), ｨｰｨ (Abe, Horikoshi, 1995), ｨｪｨ 09ｨｰｨｨｨｦ (00ｨ ｨｦｨｪｨ 08, 1987) ｨｨ ｨｰ ｨｪ ｨｬ07 09ｨｪｨｨｨｬｨ ｨｪｨｨ02 ｨｺ 01ｨ ｨｪｨｨ06 08ｨ 09ｨｰ02ｨｰ ｨｨ 09 00ｨｨ0706ｨｰ020406ｨｦ 06 09ｨｨ00ｨｪｨ 05ｨｹｨｪ06ｨｦ 00ｨｮｨｪｨｺ02ｨｨｨｨ ｨｪ 01ｨ ｨｪｨｨ ｨｨ0108ｨ 0905ｨｨ ﾂ0209ｨｺｨ 07 00ｨｨ0706ｨｰ0204ｨ ｨ 00ｨ 02ｨｰ, ﾂｨｰ06 ｨ 0409ｨｰ ｨ ｨｰ08ｨ ｨｪ0706ｨｴｨｨ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07 00ｨｨ ｨｨｨｪｨ ｨｬｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨｬ0600ｨｮｨｰ ｨ 04ｨｰｨｹ ｨｪ0609 ﾂｨｨｨｺ06ｨｬ 09ｨｨ00ｨｪｨ 05ｨｹｨｪ06ｨｦ ｨｨｨｪ000608ｨｬｨ 02ｨｨｨｨ 09 ｨ ｨｨ ｨｨ ﾂ0209ｨｺｨｨ01 06ｨ ｨｲ02ｨｺｨｰｨ 01 (Dziubinska, 2003) ｨ ｨ 0502ｨｬ ｨｨ07ｨｪｨｨ07 01ｨ ｨｪｨｨ07 ｨｪｨ 03ｨｨ ｨ ｨｪｨｨ04ｨｬ04 ｨｪｨ ｨｨｨ ｨｨ04ｨｮ ﾂ02ｨｪ ｨｺｨｰ ｨｺ ｨ ｨｪｨｨ07, ｨｨ01ｨｨｨｬ06ｨｬｨｮ 0706ｨｰ06ｨｬｨｮ, ﾂｨｰ ｨｺ ｨ ｨｪｨｨ02 03ｨｨ0806ｨｺ ｨｨｨｬ02ｨｪ0702ｨｰ ｨ ｨｨ06ｨｰ0201ｨｪ ｨｨｨｨ ｨｰ0208ｨｨ05ｨｨ04ｨ 02ｨｨｨｨ 09ｨ ｨｺ02ｨｨｨｪ ｨｨ ｨｺ06ｨｪ ｨ 02ｨｨｨｨ 07ｨｨｨｴ ｨｮｨｺｨｰ0609 (Rothschild, Mancinelli, 2001). 00 ｨｬｨｪ ﾂｨｨ090502ｨｪｨｪ ｨ ｨ 06ｨｰｨ 01, ｨｪ02ｨｪｨｪ ｨｨ 01ｨ ｨｪｨｨｨｨ 40 ｪ ｨ ｨｨ , 09 06ｨ 05ｨ 09ｨｰ0701, ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ0402 ｨｪｨ 08ｨｮ0302ｨｪｨｨ07 09ｨｰｨ ｨｪ060907ｨｰ ﾂ0209ｨｨ01ｨｪ04ｨｬｨｨ, ｨ 0405ｨｨ ｨｪ04 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ ｨｨ07ｨｨ01ｨｪ ｨｬ02ｨｬｨ 08ｨ ｨｪ, 01ｨｨ ｨｨｨ 02ｨｨ ｨｨ0006ｨｬ0208ｨｪ0401 ｨ 0205ｨｺ ｨｺ06ｨｬ070502ｨｺ ｨｨ 0706ｨｰ ｨｨｨｬｨｨ ｨｬ02ｨｪｨｰｨ ｨｰｨｨ09ｨｪ06ｨｦ ｨ ｨｺｨｰｨｨ09ｨｪ0609ｨｰｨｨ (Gross, Jaenicke, 1994; Molina-Garcia, 2002), ｨ ｨｰｨ ｨｺ0302 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ ｨｺ ｨｨｨｨ 0002ｨｪ06ｨｬｨ (Fernandes et al., 2004). 01ｨ 08ｨｮ0302ｨｪｨｨ07 09 ｨｺ ｨｪ0609ｨｪ06ｨｦ ｨｨ ｨｪ020809ｨｪ06ｨｦ 09ｨｨ09ｨｰ02ｨｬｨ ｨｨ 00ｨｨ070208ｨ ｨ 08ｨｨ ﾂ0209ｨｺｨｨ01 ｨｰ08ｨ 09ｨｬｨ ｨｰ ｨｨ ﾂｨｨｨｪ06ｨｦ 00ｨｨｨ 0205ｨｨ 03ｨｨ0906ｨｰｨｪ0401 (05ｨ 05ｨｹ02ｨｬｨ ｨｪ ｨｨ 0108., 1979). 06ｨｬ0206ｨｰ ｨ ｨｪｨｪ ｨｰ06ｨｬ, ﾂｨｰ06 00ｨｨ070208ｨ ｨ 08ｨｨ ﾂ0209ｨｺｨｮ06 00ｨｨｨ 0205ｨｹ ｨ ｨｪｨｨ04ｨｬ0609 ｨｬ0603ｨｪ ｨｰ0908ｨ ｨｰｨｨｨｰｨｹ ｨ 08ｨｨｨｰ0205ｨｹｨｪ06ｨｦ 06ｨ 08ｨ ｨ 06ｨｰｨｺ06ｨｦ ｨｨ01 09ｨｰ ｨｬｨｨ ｨｬ06ｨｪｨ ｨｬｨｨ (Tanino et al., 1992) ｨｬ ｨｮ02ｨｰ 06ｨｰｨｬ02ｨｰｨｨｨｰｨｹ, ﾂｨｰ06, ｨｪ0209ｨｬ06ｨｰ0807 ｨｪｨ ｨ ｨｺｨｰｨｮｨ 05ｨｹｨｪ0609ｨｰｨｹ ｨ 0502ｨｬ04, ｨｨ ｨ 08ｨ ｨｬ02ｨｰ ｨｪ0203ｨｪ02ｨｦ ｨｨ07ｨｪｨｨ02 ｨｨｨｬ02ｨｪｨｪ06 01ｨ ｨｪｨｨ07 ｨｪｨ ｨ ｨｨ0601ｨｨｨｬｨｨ06 ｨｨ 00ｨｨ04ｨｨ ｨｨ06 03ｨｨ ｨ ｨｪｨｨ04ｨｬ0609, 09 ｨｰ06ｨｬ ﾂｨｨ ｨ 09ｨｰ02ｨｪｨｨｨｦ, ｨｨ04ｨｮ ﾂ02ｨｪ06 09 ｨｪｨ ｨｨｨｬ02ｨｪｨｹ0302ｨｦ 09ｨｰ020702ｨｪｨｨ (01ｨ ｨｬｨ 0502ｨｦ, 2004) ｨｰ06ｨｬｨｮ ｨｹ06 ｨｪｨ 0302ｨｦ 08ｨ ｨ 06ｨｰ04 ｨ ｨｪｨｨｨｰｨｹ 0102ｨｦ09ｨｰ09ｨｨ02 09ｨｪ0203ｨｪ ｨ ｨｪｨｨ07 ｨｪｨ ｨｮ05ｨｹｨｰ08ｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｮ ｨｺ0502ｨｰ06ｨｺ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ. 414

2 130005ｨｨ07ｨｪｨｨ02 00ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨｪｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｪｨｮ ｨ ｨｪｨｨ04ｨ 02ｨｨ06 ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ02ｨｦ ｨｺｨｮｨｺｨｮ08ｨｮ ｨｨ ｨｨ07ｨｪｨｨ02 09ｨｪ0203ｨｪ ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨｪｨ ｨｰ ｨｰｨｺ0609 ｨｺｨｮｨｺｨｮ08ｨｮ0404. ｨ ｪ ﾂ ﾂ ｨｪｨｨ07 01ｨ ｨｪｨｨ ｨ 08ｨ 04ｨｪｨｨ02ｨ 0105ｨｨｨｪ ｨｰｨｺ0609 ｨｬ020301ｨｮ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ04ｨｬｨｨ ｨｨ ｨｰｨｪ04ｨｬｨｨ 06ｨ 08ｨ 0402ｨ ｨｬｨｨ ｨ 0405ｨ ｨｪ ｨｰ ｨｪ06ｨｦ; ｨ ｪ ﾂ ｨｮｨｰ ｨｪｨｨ0302ｨｪｨｨ07 01ｨ ｨｪｨｨ ｨ ｨｰｨｬ ｨｪ ｨｨｨｪｨ ｨｪｨ ｨｬｨｪ06ｨｦ ｨｨ ｨｬｨｪ06ｨｦ ﾂｨ 09ｨｰ02ｨｦ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ ｨｰｨ ｨ 06ｨｺ ｨｨ 68 % 06ｨｰ ｨｺ06ｨｪｨｰ ｨｰ0902ｨｰ09ｨｰ0902ｨｪｨｪ06. 00ｨ ｨｰ0208ｨｨｨ 05 ｨｨ ｨｬ02ｨｰ0601ｨｨｨｺｨ 02ｨ ｨｲ02ｨｺｨｰ06ｨｬ ｨｨ ｨ ｨｪｨｨｨｦ 0905ｨｮ03ｨｨ05ｨｨ 4 ｪ7-01ｨｪ0209ｨｪ ｨ 09ｨｰ02ｨｪｨｨ07 ｨｺｨｮｨｺｨｮ08ｨｮ0404 Zea mays L ｨｰ ｨｪ06ｨｰｨ ｨｨ 08ｨｮｨ ｨ ｨｪｨｹ. 06ｨｰｨｨ0605ｨｨ080609ｨ ｨｪｨｪ ｨ 09ｨｰ02ｨｪｨｨ ｨ ｨｴｨｨ09ｨ 05ｨｨ09ｨｹ 00ｨｨ ｨｪｨｪ04ｨｬ ｨｬ02ｨｰ060106ｨｬ ｨｪｨ ｨｪ06ｨｦ 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 09ｨｰｨｨｨｺ ｨｬｨｨ0505ｨｨ05ｨｨｨｰ ｨｬｨｺ0609ｨｰ ｨ ｨｪｨ 05ｨｨ04ｨ 0905ｨｨ07ｨｪｨｨ07 01ｨ ｨｪｨｨ07 ｨｪｨ ｨｰ ｨｨｨｪｨｰｨ ｨｺｨｰｨｪ ｨ 09ｨｰ02ｨｪｨｨ ｨ ｨｴｨｨ09ｨ ｨｪｨｨ07 09ｨｬ0209ｨｰ02 09 ｨｺ06ｨｪｨｰ02ｨｦｨｪ020806ｨｬ 0706ｨｬ02ｨｴｨ 05ｨｨ 09 ｨｺｨ ｨｬ0208ｨｮ ｨｺ ｨ ｨｪｨｨ07 ｨｨ ｨ 05ｨｨ ｨｦ09ｨｰ09ｨｨ06 01ｨ ｨｪｨｨ07 2 ｨｨ05ｨｨ ｨ 09 ｨｰ02 ﾂ02ｨｪｨｨ02 5 ﾂ. 0806ｨｪｨｰ080605ｨｹｨｪ ｨ 08ｨ ｨｬ02ｨｴｨ 05ｨｨ 09 ｨｰｨ ｨｺｨｮ ｨｺｨ ｨｬ0208ｨｮ 0708ｨｨ ｨ ｨｰｨｬ ｨｪ06ｨｬ 01ｨ ｨｪｨｨｨｨ. 0004ｨ ｨｨ ﾂｨｨｨｪ04 01ｨ ｨｪｨｨ07, ｨ 05ｨｨ04ｨｺ06ｨｦ ｨｺ ｨｪ02ｨｦ 0008ｨ ｨｪｨｨ ｨｨ04ｨｨ ｨｨ ﾂ0209ｨｺ ｨｨｨ 07ｨ 0406ｨｪｨ, ｨ ｨｨｨｺｨｰ0609ｨ ｨｪ ｨ ｨｪｨｨ02ｨｬ ｨｮ ﾂｨｨｨｰｨｹ ｨｪｨ ｨｨｨ 0605ｨｹ03ｨｨｨｦ ｨｺ06ｨｪｨｰ08ｨ 09ｨｰ ｨｰｨｪ0401 ｨｨ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨ 08ｨ ｨ ｨｪｨｨ ｨ 09ｨ 05ｨｨ ｨｬ06ｨｴｨｹ06 07ｨｪ0209ｨｬ0609ｨｨ09ｨｰ02ｨｬ04, ｨｰ0607ｨｴ02ｨｦ ｨｨ04 ｨ ｨ ｨｪ ｨ ｨｰ ｨｮ01ｨ, ｨｮｨｺｨｰ0608ｨ 01ｨ ｨｪｨｨ07 ｨｨ ｨｺｨｰ0806ｨｺ05ｨ 07ｨ ｨｪ ｨ ﾂｨｨ ｨｨ 09ｨ ｨ 01ｨ ｨｪｨｨ07, 06ｨ ｨｲ0201ｨｨｨｪ02ｨｪｨｪ ｨｪ0209ｨｬ06ｨｬｨ 00ｨｨ09ｨｰ08ｨ 0507ｨｬｨｨ 09 ｨｺｨ ｨｬ020806ｨｦ 01ｨ ｨｪｨｨ07 (08ｨｪｨｨ09ｨｨｨｬ0609 ｨｨ 0108., 2012) ｨｰ ｨｨｨｺ09ｨ 02ｨｨ06 06ｨ 08ｨ ｨｺｨｰ0806ｨｪｨｪ06ｨｦ ｨｬｨｨｨｺ080609ｨｺ0607ｨｨｨｨ ｨｨ05ｨｨ ｨｪ ｨｰ0902ｨｪｨｪ06 09 ｨｺｨ ｨｬ ｨ ｨ ｨ 01ｨ ｨｪｨｨ ｨｰｨ 07ｨ ｨｨ09ｨ ｨｪｨｨ07 06ｨ 08ｨ ｨ ｨｪｨｨ02ｨｬ (30 ｨｬｨｨｨｪ) ｨｮ00ｨｨ01 09ｨ 08ｨｨｨ ｨｪｨｰｨ 01 00ｨｨｨｺ09ｨ 02ｨｨ ｨｨ05ｨｨ ｨ ｨ 01ｨ ｨｪｨｨ ｨ ｨｰｨｬ ｨｪ ｨｺｨｰ0806ｨｪｨｪ06-ｨｬｨｨｨｺ080609ｨｺ0607ｨｨ ﾂ0209ｨｺ ｨｨ ｨ ｨｪｨｨ07 ｨ 07ｨｨｨｺｨ 05ｨｹｨｪ ｨ 00ｨｬ02ｨｪｨｰ04 ｨｺ0608ｨｪ ｨｨｨｪ06ｨｦ 7 ｪ8 ｨｬｨｬ 00ｨｨｨｺ09ｨｨ080609ｨ 05ｨｨ 12 ﾂ %-ｨｪ06ｨｬ 08ｨ 09ｨｰ ｨｮｨｰｨ 08ｨ 05ｨｹ010200ｨｨ01ｨ ｨｪｨ ｨ ｨｰｨｪ06ｨｬ ｨ ｨｮ ( ) ｨｨ ｨｰ00ｨｨｨｺ09ｨｨ080609ｨ 05ｨｨ 2 ﾂ 09 1%-ｨｪ06ｨｬ OsO 4 ｨｪｨ ｨｰ06ｨｬ 0302 ｨ ｨｮ ｨ ｨ ｨｪｨｨ02ｨｬ 09ｨ 01ｨ (34 ｨｬ00/ｨｬ05). 02ｨ 08ｨ ｨｨ0108ｨ ｨｰｨｨ080609ｨ 05ｨｨ 09 08ｨ 09ｨｰ090608ｨ ｨｨ08ｨｰｨ ｨ 09ｨｰｨ 06ｨｴ02ｨｦ ｨｺ06ｨｪ0202ｨｪｨｰ08ｨ 02ｨｨｨｨ (30 ｪ70 %) ｨｨ ｨ 0202ｨｰ06ｨｪｨ (70 ｪ100 %) ｨｮ06ｨｴｨｨｨｬ 04ｨ ｨｬ02ｨｴ02ｨｪｨｨ02ｨｬ ｨ 0202ｨｰ06ｨｪｨ 06ｨｺｨｨ09ｨｹ ｨｨ0502ｨｪｨ (45 ｨｬｨｨｨｪ) ｨｬ02ｨｪｨｰ04 ｨｺ0608ｨｪ ｨｨｨｰ0409ｨ 05ｨｨ ｨｺ09ｨｨ01ｨｪ06ｨｦ 09ｨｬ060506ｨｦ ｨｪ-812 (010208ｨｬｨ ｨｪｨｨ07) ｨｨ ｨｨｨｬ0208ｨｨ040609ｨ 05ｨｨ 0708ｨｨ 37, 45 ｨｨ 57 縺 ﾂ ｨｮｨｰ06ｨｪｨｺｨｨ02 ｨｨ ｨｮ05ｨｹｨｰ08ｨ ｨｰ06ｨｪｨｺｨｨ ｨｮ ﾂｨ 05ｨｨ ｨｪｨ ｨｬｨｨｨｺ0806ｨｰ06ｨｬ02 LKB-III ( ｨｨ07), ｨｺ06ｨｪｨｰ08ｨ 09ｨｰｨｨ080609ｨ 05ｨｨ 08ｨ 09ｨｰ ｨｬ ｨｮ08ｨ ｨｪｨｨ05-ｨ 0202ｨｰｨ ｨｰｨ (Tander, 1990) ｨｨ 02ｨｨｨｰ08ｨ ｨｰ06ｨｬ 0909ｨｨｨｪ02ｨ (Reynolds, 1963) ｨ 08ｨ ｨｰ ｨｬｨ ｨｰ08ｨｨ09ｨ 05ｨｨ ｨｰｨｨ ﾂ0209ｨｺ06ｨｬ ｨｬｨｨｨｺ080609ｨｺ NU-2 (010208ｨｬｨ ｨｪｨｨ07) ｨｨ ｨｺｨｰ0806ｨｪｨｪ06ｨｬ ｨｬｨｨｨｺ080609ｨｺ JEM (080706ｨｪｨｨ07) ｨｰ ｨｨ05ｨｨ 09 3-ｨｺ08ｨ ｨｰｨｪ06ｨｦ ｨ ｨｨ ｨｨ ﾂ0209ｨｺ06ｨｦ ｨｰ0608ｨｪ0609ｨｰｨｨ. 01ｨ 08ｨｨ09ｨｮｨｪｨｺｨ ｨｨ ｨｪ ｨｪｨｨ02 04ｨｪｨ ﾂ02ｨｪｨｨ07 09 ｨｨ01 09ｨｰｨ ｨｰｨｨ09ｨｰｨｨ ﾂ0209ｨｺ06ｨｦ 0603ｨｨｨ ｨｺ06ｨｦ ｨｮ05ｨｹｨｰｨ ｨｰ ｨ 09ｨｪ ｨｮ ﾂ02ｨｪｨｪ04ｨｬ 01ｨ ｨｪｨｪ04ｨｬ, ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 09ｨｪ0203ｨｪ ｨ ｨｪｨｨ ｨｨ090601ｨｨ0506 ｨｺ 09ｨｪｨｨ0302ｨｪｨｨ06 09ｨｺ ｨｰｨｨ ｨｰｨ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ ｨｰ0902ｨｪｨｪ06 ﾂ ﾂ ｨｪｨ ｨｪｨｨ07 09 ｨ ｨ 0806ｨｺｨ ｨｬ ｨｪ ｨｨｨｪｨ ｨｺ0608ｨｪ07 ｨｨ 0706ｨ 0200ｨ ｨ 0405ｨ ｨｪ ｨｰ ｨｪ06 ｨｪｨｨ0302, ﾂ02ｨｬ ｨｮ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨ 09ｨｰ02ｨｪｨｨｨｦ (08ｨｨ09. 1, ｨ ), ｨｪ06 ｨｪｨ ｨｮｨｰ 08ｨ 04ｨｪｨｨ02ｨ ｨｰｨ ｨ 06ｨｺ % 06ｨｰ ｨｺ06ｨｪｨｰ ｨｨ ｨ 0405ｨ ｨｰ ｨｪ06ｨｦ (08ｨｨ09. 1, ｨ ). 08ｨｪｨ 05ｨｨ04 01ｨｨｨｪｨ ｨｬｨｨｨｺｨｨ ｨｰｨ 0909ｨｨ0102ｨｰ0205ｨｹ09ｨｰ09ｨｮ02ｨｰ 06 ｨ 0409ｨｰ0806ｨｬ ｨｨ ｨｪ06ｨｬ ｨｰｨ ｨｪ ｨｪｨｨｨｨ 09ｨｺ ｨｰｨｨ ｨｰ ｨｬ/09ｨｮｨｰ ﾂ ｨｮｨｰ ｨｺ08ｨ ｨｴ02ｨｪｨｨ ｨｦ09ｨｰ09ｨｨ07 01ｨ ｨｪｨｨ07 (08ｨｨ09. 2) ｨｮ ﾂ02ｨｪｨｪ ｨ ｨｪｨｪ ｨｨ0102ｨｰ0205ｨｹ09ｨｰ09ｨｮ06ｨｰ ｨｺ06ｨｦ ﾂｨｮ0909ｨｰ09ｨｨｨｰ0205ｨｹｨｪ0609ｨｰｨｨ ｨ ｨｰｨ 08ｨ 09ｨｰ070302ｨｪｨｨ02ｨｬ ｨｺ 00ｨｨ070208ｨ ｨ 08ｨｨｨｨ ｨｰｨｺｨｨ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨ 09ｨｰ02ｨｪｨｨｨｦ 09 ｨｺ06ｨｪ ｨｪ04 08ｨ 09ｨｰ070302ｨｪｨｨ07 (5 ｨｬｨｬ 06ｨｰ ｨｺ06ｨｪ ﾂｨｨｨｺｨ ｨｺ0608ｨｪ07) ｨｨｨｬ0205ｨｨ ｨｺ08ｨｮ07ｨｪｨｮ ｨｪｨｰ08ｨ 05ｨｹｨｪｨｮ06 09ｨ ｨｺｨｮ0605ｨｹ, ｨｨ09ｨｺ0609ｨｨ01ｨｪ06ｨｦ ｨｬ04 ｨｨ ｨｰ06ｨｪｨｺｨｨｨｦ 0708ｨｨ09ｨｰ02ｨｪｨｪ04ｨｦ ｨｦ 02ｨｨｨｰ060705ｨ 04ｨｬ04 (08ｨｨ09. 3, ｨ, ｨ ) ｨ 04ｨｬｨ 0502ｨｬｨｬｨ ｨ 0405ｨ ｨｮｨｬ020802ｨｪｨｪ ｨｪｨｨ09ｨｰ06ｨｦ ｨｨ ｨｰｨｪ ｨｨ050200ｨ 05ｨ ｨｺ ｨｺ0502ｨｰ06 ﾂ- 04ｨｨ ｨｨ07ｨｪｨｨ02 09ｨｪ0203ｨｪ ｨ ｨｪｨｨ07 ｨｪｨ 09ｨｺ ｨｰｨｹ ｨｰｨ ｨｺ0608ｨｪ02ｨｦ ｨｰｨｺ0609 ｨｺｨｮｨｺｨｮ08ｨｮ ｪ ｨｺ06ｨｪｨｰ080605ｨｹ, 2 ｪ 01ｨ ｨｪｨｨ02 4 ｨｬ03ｨ 09 ｨｰ02 ﾂ02ｨｪｨｨ02 5 ﾂ. 07ｨｺｨ 04ｨ ｨｪ ｨｪ ｨｺ ｨｰｨｹ (ｨｬ02ｨｰ0601 ｨｪｨ ｨｨｨｬ02ｨｪｨｹ03ｨｨ01 ｨｺ09ｨ 0108ｨ ｨｰ0609) ｨｨ 09ｨｺ ｨｰｨｹ ｨｰｨ ｨｮｨｰ ｨｨ ｨｪｨｨ07 01ｨ ｨｪｨｨ07. 05ｨｺ ｨｰｨｹ ｨｰｨ 09ｨｪｨｨ03ｨ 05ｨ 09ｨｹ 0708ｨｨ ｨｦ09ｨｰ09ｨｨｨｨ ｨｬ/09ｨｮｨｰ ｨｨ ｨｪ0609ｨｰｨｹ ｨｰｨ ｨｪｨ 0905ｨｨ09ｨ 05ｨ 09ｨｹ 0106 ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨｪｨ ﾂ02ｨｪｨｨｨｦ ｨｪｨ ｨｮ06ｨｴｨｨ02 09ｨｮｨｰｨｺｨｨ ｨｺ08ｨ ｨｴ02ｨｪｨｨ ｨｦ09ｨｰ09ｨｨ07.

3 ｨ 0108ｨ 01ｨｨｨｬ0609 ｨｨ ｨｨ ｨｹｨｰ08ｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨ ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ02ｨｦ ｨｺ06ｨｪｨｰ080605ｨｹｨｪ ｨ 09ｨｰ02ｨｪｨｨｨｦ, 04ｨ 00ｨｨｨｺ09ｨｨ080609ｨ ｨｪｨｪ ｨｨ ｨ ｨｰｨｬ ｨｪ06ｨｬ 01ｨ ｨｪｨｨｨｨ (07) 01ｨｨ09ｨｺ0609ｨｨ01ｨｪ06ｨｦ ｨｬ04 (ｨ ) 08ｨ ｨｪ06 09 ｨｰ06ｨｪｨｺ06ｨｬ 0708ｨｨ09ｨｰ02ｨｪｨｪ06ｨｬ ｨｨｨｰ060705ｨ 04ｨｬ04 (ｨ ). 01ｨｨｨ ｨ 04ｨｬｨ ｨｪ02ｨｦ ｨｺｨｰ0806ｨｪｨｪ06ｨｦ ｨｰｨｪ0609ｨｰｨｨ ｨｨｨｰ ｨｬｨｪ ﾂｨｨ090502ｨｪｨｪ ｨｨ0906ｨｬ04 (09, 00) ｨｬ02ｨｪｨｰ04 05ｨｪ ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺ06ｨｦ 0902ｨｰｨｨ (0509) ｪ 02ｨｨ09ｨｰ0208ｨｪ04 ｨｨ 01ｨｨｨｺｨｰｨｨ060906ｨｬ04 ｨ 0707ｨ 08ｨ ｨｰｨ ｨｹ0103ｨｨ ｪ 08ｨ ｨｪ04 08ｨ 09ｨｪ06ｨｬ0208ｨｪ ｨ ｨｲ02ｨｬｨｮ 02ｨｨｨｰ (09, 00). 09ｨｺ ｪ 09ｨ ｨｺｨｮ0605ｨｹ, ｨｺ00 ｪ 01ｨｨｨｺｨｰｨｨ060906ｨｬ04 ｨｺ06ｨｬ070502ｨｺ09ｨ ｨｹ0103ｨｨ, ｨｺ09 ｪ ｨｺ0502ｨｰ06 ﾂｨｪｨ 07 09ｨｰ02ｨｪｨｺｨ, ｨｬ ｪ ｨｬｨｨｨｰ060106ｨｪ0108ｨｨ07. ｨｪ06ｨｦ 09ｨｰ02ｨｪｨｺ02 (08ｨｨ09. 3, ｨ, 09). 01ｨｨｨ ｨ 04ｨｬｨ ｨｨｨｬ0205ｨ ｨｪ ｨｺｨｰ0806ｨｪｨｪｨｮ ｨｰｨｪ0609ｨｰｨｹ ｨｨ ｨ 05ｨ 04ｨ 09ｨｨｨｰｨｺｨｨ ｨｨ0906ｨｬ (08ｨｨ09. 3, 09, 00). 02ｨｺ08ｨｮ ｨｰ060106ｨｺ09ｨ 05ｨｹｨｪ0402 ｨｬｨｨｨｰ060106ｨｪ0108ｨｨｨｨ, ｨｺ060806ｨｰｨｺｨｨ02 02ｨｨ09ｨｰ0208ｨｪ04 05ｨｪ ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺ06ｨｦ 0902ｨｰｨｨ (0605) ｨｨ 01ｨｨｨｺｨｰｨｨ060906ｨｬ04 ｨ 0405ｨｨ 01ｨｨ0000ｨｮ04ｨｪ06 ｨｨ 08ｨ 09ｨｪ06ｨｬ0208ｨｪ06 08ｨ ｨｪ ｨ ｨｲ02ｨｬｨｮ 02ｨｨｨｰ (08ｨｨ09. 3, ｨ ｪ00). ﾁｨｨ ﾂ0205ｨｪ06 ﾂｨｪ ｨｨｨｺｨｮ05 ｨｮ 02ｨｨ09ｨｰ0208ｨｪ 01ｨｨｨｺｨｰｨｨ060906ｨｬ ｨｪ02ｨｬｨｪ ﾂｨｨ090502ｨｪｨｪ ｨｪ060705ｨ 09ｨｰ ｨｺ0502ｨｰ06ｨｺ ｨｺ ﾂｨ 09ｨｰ ｨｬｨｨ0600ｨｨ0502ｨｪ ｨｨ ｨ 01ｨｺ08ｨｮ09ｨｰｨｨ080609ｨ ｨｪ ｨｺｨｰ0806ｨｪｨｪ ｨｰｨｪ04ｨｬ 0902ｨｴ0209ｨｰ0906ｨｬ. 08ｨｪｨ ｨｨ ﾂｨｪ ｨｴ0209ｨｰ ｨｨ ｨｹ0209 ｨｨ 0008ｨ ｨｪｨｮ ｨｹ ｨｨ ｨｰｨｨ 0202ｨｪｨｰ08ｨ 05ｨｹｨｪ06ｨｦ 09ｨ ｨｺｨｮ0605ｨｨ (08ｨｨ09. 3, ｨ, 00). 00 ｨｮ ｨｨ ｨｪ0203ｨｪ ｨ ｨｪｨｨ ｨ (30 ｨｬｨｨｨｪ) ｨｪｨ ｨ ｨ 05ｨｨ 06ｨｺ08ｨｮ000502ｨｪｨｨ (08ｨｨ09. 4, ｨ ) ｨｨ ｨｺ05ｨ 09ｨｰ0208ｨｨ04ｨ 02ｨｨ ｨｬ02ｨｪｨｰ ｨ 08ｨ ｨ ｨｪｨｨ02ｨｬ ｨｮ ﾂｨ 09ｨｰｨｺ ｨ 09ｨｰ060705ｨ 04ｨｬ04 (08ｨｨ09. 4, ｨ, 01). 01ｨ ｨ ｨｮ01ｨ ｨｪｨｨ ｨｰｨｨ 0902ｨｰｨｨ ｨ ｨｪ0204ｨｪｨ ﾂｨｨｨｰ0205ｨｹｨｪ04ｨｬ. 0201ｨｪ ｨｬ02ｨｪｨｪ ｨｨ09ｨｹ ｨｰ0502ｨｪｨｨ02 ｨｬｨ ｨｰ08ｨｨｨｺ09ｨ ｨｬｨｨｨｰ060106ｨｪ0108ｨｨｨｦ (08ｨｨ09. 4, 09) ｨｨ 08ｨ 0907ｨ ｨｨ0906ｨｬ ｨｪｨ ｨｬ06ｨｪ060906ｨｬ04 (08ｨｨ09. 4, 00, 01) ｨｨ ﾂ02ｨｪｨｨ ｨｺｨｰ0806ｨｪｨｪ06ｨｦ ｨｰｨｪ0609ｨｰｨｨ 02ｨｨｨｰ ｨ ｨｹ ｨｰ06ｨｬ 09 02ｨｨｨｰ060705ｨ 04ｨｬ02 ﾂｨｨ0905ｨ ｨｬ0205ｨｺｨｨ ｨ ｨｺｨｮ060502ｨｦ (08ｨｨ09. 4, ｨ ). 01ｨ ｨｰ08ｨ ｨｪ09-09ｨｰ060806ｨｪ02 01ｨｨｨｺｨｰｨｨ060906ｨｬ ｨｪｨ ｨｺｨ 0705ｨｨ09ｨ 05ｨｨ09ｨｹ ｨｨｨｺｨｮ ｨｺｨｰ0806ｨｪｨｪ ｨｰｨｪ04ｨｬ ｨｨｨｬ04ｨｬ (08ｨｨ09. 4, 00) ｨｮｨｺ02ｨｨ ｨｺｨｰ0806ｨｪｨｪ06ｨｦ ｨｰｨｪ0609ｨｰｨｨ ｨｬ02ｨｬｨ 08ｨ ｨｪ04 ｨｰ06ｨｪ060705ｨ 09ｨｰｨ ｨｺ ｨｨ080609ｨ 05ｨ 09 ｨｮｨｬ02ｨｪｨｹ0302ｨｪｨｨ02ｨｬ 09 09ｨ ｨｺｨｮ ｨｺ0605ｨｨ ﾂ0209ｨｰ09ｨ 0609ｨｬｨｨ0600ｨｨ05ｨｹｨｪ ｨｺ0506 ﾂ02ｨｪｨｨｨｦ (08ｨｨ09. 4, ｨ, 09, 01). 0308ｨｨ 01ｨ ｨｪｨｨｨｨ ｨ 01ｨ 08ｨ ｨｺｨｰ ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｪ0401 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨｨｦ 09 02ｨｨｨｰ , 01ｨｨｨｺｨｰｨｨ060906ｨｬｨ 01 ｨｨ ｨｬｨｨｨｰ060106ｨｪ0108ｨｨ0701 (08ｨｨ09. 5, ｨ ｪ09) ｨｬ ｨ 0405 ｨ ｨｪｨ ｨｨ ﾂ02ｨｪ ｨ 05ｨｹｨｰ0208ｨ 02ｨｨ07ｨｬ, ｨｪｨｪ04ｨｬ 0708ｨｨ 01ｨ ｨｪｨｨｨｨ ｨ (08ｨｨ09. 4, ｨ ｪ01) ｨｪｨｨ ｨｨ ﾂｨｨｨｪ04 01ｨ ｨｪｨｨ07 ｨｨｨｪ01ｨｮ02ｨｨ080609ｨ ｨｪｨｨｨｺｨｪ060902ｨｪｨｨ02 ｨ 05ｨｹｨｰ0208ｨ 02ｨｨｨｦ 09 ｨｰ06ｨｪ060705ｨ 09ｨｰ02 (08ｨｨ09. 5, 09, 09ｨｰ080205ｨｺｨｨ). 01ｨ ｨｺｨｰ0806ｨｪ060008ｨ ｨｬｨｬｨ 01 06ｨｪｨｨ ｨｨ09ｨｹ ｨｺｨ ｨｺ 06ｨ 05ｨ 09ｨｰｨｨ ｨ 02ｨｨｨｦ, ﾂｨｰ06 ｨｬ ｨｮｨｺｨ ｨ ｨｰｨｹ ｨｪｨ ｨｪｨ 08ｨｮ0302ｨｪｨｨ ｨｰｨｪ0609ｨｰｨｨ ｨｬ02ｨｬｨ 08ｨ ｨｪ ｨｪｨｰ08ｨ 05ｨｹｨｪ06ｨｦ 09ｨ ｨｺｨｮ0605ｨｨ (08ｨｨ09. 5, 09) ｨｺ ｨｪｨｨ0302ｨｪｨｨ02 00ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ ｨ ｨｰｨｬ ｨｪ ｨｨ090601ｨｨ0506 ｨｺ ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ ｨｪｨｨｨｦ ｨｰ06ｨｪ060705ｨ 09ｨｰｨ (08ｨｨ09. 6), ｨｺ0605ｨｨ ﾂ0209ｨｰ0906 ｨｺ06ｨｰ ｨ ｨｬ ｨｨ06ｨｪｨ 05ｨｹｨｪ06 08ｨ 04ｨｪｨｨ ｨ ｨｪｨｨｨｦ ｪ ｨｬｨｨｨｪｨｨｨｬｨ 05ｨｹｨｪ06ｨｦ 09 09ｨ 08ｨｨｨ ｨｪｨｰ ｨ (08ｨｨ09. 6, ｨ, 09ｨｰ080205ｨｺｨｨ) ｨｨ ｨｬｨ ｨｺ09ｨｨｨｬｨ 05ｨｹｨｪ06ｨｦ 0708ｨｨ ｨ (08ｨｨ09. 6, ｨ, 09ｨｰ080205ｨｺｨｨ). 0308ｨｨ 05ｨｰ06ｨｬ 01ｨ 08ｨ ｨｺｨｰ ｨｮ00ｨｨ01 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨｨｦ (ｨ ｨ 02ｨｨ ｨｰｨｨ ｨｨ 01ｨｨｨｺｨｰｨｨ060906ｨｬ ｨｨｨｺｨｮ05ｨ ｨｬｨｨ, ｨｰ0502ｨｪｨｨ02 ｨｬｨｨｨｰ060106ｨｪ0108ｨｨｨｦ), ｨｪｨｪ ｨｨ 00ｨｨｨｺ09ｨ 02ｨｨｨｨ 06ｨ 08ｨ ｨｮ ｨｨ ｨｪｨｪ ｨ ｨｪｨｨ07, 09ｨｮｨｴ0209ｨｰ0902ｨｪｨｪ06 ｨｪ02 ｨｨ04ｨｬ02ｨｪ (08ｨｨ09. 6, 09, 00). 01ｨｨ ｨｪ06ｨｬ ｨｨ04 ｨｨ04ｨｮ ﾂｨ 02ｨｬ ｨ 08ｨｨｨ ｨｪｨｰ0609 ｨｪｨ ｨｬｨｨ ｨｪ ｨｹ ｨｪｨ 08ｨｮ0302ｨｪｨｨ ｨｰｨｪ0609ｨｰｨｨ 0705ｨ 04ｨｬｨ 0502ｨｬｨｬ04, ﾂｨｰ ｨｰ ｨ ｨｹ 06ｨｺ08ｨ 09ｨｺ06ｨｦ ｨｨｨｪ01ｨｨｨｺｨ ｨｰ060806ｨｬ ｨｰｨｪ0609ｨｰｨｨ ｨ ｨｪｨｨ ﾂｨｪ06ｨｦ ｨｬ02ｨｬｨ 08ｨ ｨｪ ｨ ｨｪ0906ｨｬ ｨｮｨ 04ｨｬ (01ｨ ｨｪｨｪ0402 ｨｪ ｨｨ ｨｪ04). 02ｨ 09ｨｮ030102ｨｪｨｨ ｨ 09ｨｪ ｨｮ ﾂ02ｨｪｨｪ04ｨｬ 01ｨ ｨｪｨｪ04ｨｬ, ｨｪｨｨ02 00ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ ｨｰ ｨｺ ｨｰ0608ｨｬ060302ｨｪｨｨ ｨｰｨ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ (08ｨｨ09. 1, 2) ｨｨｨｰ0208ｨ ｨｰｨｮ0802 ｨｨｨｪ00ｨｨｨ ｨｨ080609ｨ ｨｪｨｨ ｨｰｨ 06ｨｰｨｬ02 ﾂｨ 02ｨｰ0907 ｨｺｨ ｨｺ ｨ ｨ ｨｺ02ｨｨ07 ｨｪｨ ｨｪｨｨ ｨｬ06ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 (Bartels, Sunkar, 2005; Munns, 2011), 0708ｨｨ ﾂ02ｨｬ ﾂｨｮ0909ｨｰ09ｨｨｨｰ0205ｨｹｨｪ0609ｨｰｨｹ ｨｰｨ 0706ｨ 0200ｨ ｨｺ 0609ｨｬ06ｨｰｨｨ ﾂ0209ｨｺ06ｨｬｨｮ 09ｨｰ ｨｮ 06ｨｺｨ 04ｨ 05ｨ 09ｨｹ , ﾂ02ｨｬ ｨｮ ｨｺ0608ｨｪ07 (Bartels, Sunkar, 2005). 00 ｨｪｨ 0302ｨｬ

4 0005ｨｨ07ｨｪｨｨ02 00ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨｪｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｪｨｮ ｨ ｨｪｨｨ04ｨ 02ｨｨ06 ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ02ｨｦ ｨｺｨｮｨｺｨｮ08ｨｮ ｨｨ ｨｹｨｰ08ｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨ ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ02ｨｦ 0406ｨｪ04 08ｨ 09ｨｰ070302ｨｪｨｨ07, 04ｨ 00ｨｨｨｺ09ｨｨ080609ｨ ｨｪｨｪ ｨｨ 09ｨｪ0203ｨｪ02ｨｬ 01ｨ ｨｪｨｨｨｨ ｨ (30 ｨｬｨｨｨｪ). 02ｨｺ08ｨｮ000502ｨｪｨｨ (ｨ ) ｨ ｨｹ ｨ ｨ 02ｨｨ02ｨｦ 02ｨｨ09ｨｰ0208ｨｪ 05ｨｪ ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺ06ｨｦ 0902ｨｰｨｨ (ｨ ), ｨｰ0502ｨｪｨｨ02ｨｬ ｨｬｨ ｨｰ08ｨｨｨｺ09ｨ ｨｬｨｨｨｰ060106ｨｪ0108ｨｨｨｦ (09), 08ｨ 0907ｨ 0106ｨｬ ｨｨ0906ｨｬ ｨｨ ｨｪｨ ｨｺ ｨｪｨｨ02ｨｬ ｨｰ08ｨ ｨｪ ｨｰｨｪ ｨｨｨｺｨｮ05 (00, 01). 02ｨ 0604ｨｪｨ ﾂ02ｨｪｨｨ07 ｨｰ , ﾂｨｰ06 ｨｨ ｨｪｨ 08ｨｨ ｨｮ ﾂｨ 02 ｨｪｨ ｨ ｨ ｨｹ ｨｨ06ｨｪｨ 05ｨｹｨｪ ｨｪｨｨ0302ｨｪｨｨ02 09ｨｺ ｨｰｨｨ ｨｰ ｨｺ0608ｨｪ07 ｨｨ 0706ｨ 0200ｨ (08ｨｨ09. 1, ｨ, ｨ ), 0708ｨｨ ﾂ02ｨｬ 0708ｨｨ 01ｨ ｨｪｨｨ0701 ｨｪｨ ｨｺ ｨｪｨｨ ｨｨ ﾂｨｨｨｪ04, 0708ｨｨ ｨｺ06ｨｰ060806ｨｦ ｨｪｨ ｨ ｨ 02ｨｰ ｨｺ08ｨ ｨｴ02ｨｪｨｨ ｨｪｨｨ07 ｨｺ0502ｨｰ06ｨｺ ｨｮ 05ｨｮｨｺｨ 08ｨｨ06ｨｰ (Iwahashi et al., 2005). 06ｨｰｨｨ ｨｪｨ ｨ ｨｪｨｨ ｨｨ05ｨｨ 04ｨ ｨｺ0506 ﾂｨｨｨｰｨｹ, ﾂｨｰ ｨｨ ﾂｨｨｨｪ06ｨｦ ｨｰ0608ｨｬ060302ｨｪｨｨ ｨｰｨ 09 ｨｪｨ 03ｨｨ ｨｰｨ ｨｰ0907 ｨｨｨｪ00ｨｨｨ ｨｨ080609ｨ ｨｪｨｨ ｨｰｨ 08ｨ 09ｨｰ070302ｨｪｨｨ02ｨｬ. 08ｨｪｨ 05ｨｨ ｨｪ04 08ｨ 09ｨｰ070302ｨｪｨｨ07 ｨｺ0608ｨｪ ｨｦ09ｨｰ09ｨｨｨｰ0205ｨｹｨｪ ｨｨ05 09ｨｮｨｴ0209ｨｰ0902ｨｪｨｪ0402 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07 ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｦ ｨ ｨｪｨｨ04ｨ 02ｨｨｨｨ ｨｰｨｪ ｨ 09ｨｰ02ｨｪｨｨｨｦ 0708ｨｨ 00ｨｨ070208ｨ ｨ 08ｨｨｨｨ (08ｨｨ09. 4, 5, 6). 06ｨｰｨｨ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ ｨ 03ｨ 05ｨｨ09ｨｹ 09 ｨｮｨｬ02ｨｪｨｹ0302ｨｪｨｨｨｨ 09 ｨｺ0502ｨｰｨｺｨ ｨ ｨｪｨｨ ｨｨ0906ｨｬ 09 02ｨｨｨｰ , ｨｺ05ｨ 09ｨｰ0208ｨｨ04ｨ 02ｨｨｨｨ ｨｬ02ｨｪｨｰ ｨｪ ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺ06ｨｦ 0902ｨｰｨｨ ｨｨ 01ｨｨｨｺｨｰｨｨ060906ｨｬ ｨ 0707ｨ 08ｨ ｨｰｨ ｨｹ0103ｨｨ (08ｨｨ09. 4, ｨ, 00; 5, ｨ ｪ09). 01ｨ ｨｨ000208ｨｨｨｨ 01ｨｨｨｺｨｰｨｨ060906ｨｬ 06ｨｰｨｬ02 ﾂｨ ｨｹ ｨｪｨ ｨｺ ｨｪｨｨ02 ｨｰ08ｨ ｨｪ ｨｰｨｪ ｨｮ040408ｨｹｨｺ0609, ｨ ｨｴｨｨ ｨｺｨｰ0806ｨｪｨｪ ｨｰｨｪ ｨｴ0209ｨｰ0906 (08ｨｨ09. 5, 09) ｨｺｨｮ07ｨｪ0609ｨｰｨｹ 05ｨｰｨｨ01 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｪ0401 ｨ 05ｨｹｨｰ0208ｨ 02ｨｨｨｦ ｨｬ0209ｨｰｨｪ06 09 ｨｬ ｨｨ ﾂ0209ｨｺｨｨｨｬｨｨ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07ｨｬｨｨ 0106ｨｪ0108ｨｨ06ｨｬｨ (08ｨｨ09. 4, 09; 5, ｨ ) ｨｨ0102ｨｰ0205ｨｹ09ｨｰ09ｨｮ02ｨｰ 06 ｨｰ0608ｨｬ060302ｨｪｨｨｨｨ ｨｬ02ｨｰｨ ｨ 0605ｨｨ ﾂ0209ｨｺｨｨ , 09ｨｺ0506 ﾂｨ 07 ｨ ｨｨ0609ｨｨｨｪｨｰ0204 ｨ 0205ｨｺ0609 ｨｨ ｨｰ08ｨ 00ｨｨｨｺ 09 ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪ06ｨｦ 09ｨｨ09ｨｰ02ｨｬ02, ﾂｨｰ ｨ 09ｨｮ02ｨｰ ｨｨｨｪ00ｨｨｨ ｨｨ080609ｨ ｨｪｨｨ02ｨｬ ｨｰ (08ｨｨ09. 1, 2). 08ｨ ｨｺｨｰ, ﾂｨｰ06 06ｨ ｨｪｨ 08ｨｮ0302ｨｪｨｪ0402 ｨ 05ｨｹｨｰ0208ｨ 02ｨｨｨｨ (08ｨｨ09. 4, 5) ｨ ｨｪ0706ｨｰ ｨｨ 0102ｨｺ06ｨｬ ｨｨｨｨ (08ｨｨ09. 6), 0909ｨｨ0102ｨｰ0205ｨｹ09ｨｰ09ｨｮ02ｨｰ ｨｬ06ｨｬ 0905ｨｨ07ｨｪｨｨｨｨ 09ｨｪ0203ｨｪ ｨ ｨｪｨｨ07 ｨｪｨ ｨｮ05ｨｹｨｰ08ｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｮ 08ｨ 09ｨｰｨｨｨｰ0205ｨｹｨｪ06ｨｦ ｨｺ0502ｨｰｨｺｨｨ; 0102ｨｺ06ｨｬ ｨｨ07 ｨｮ090205ｨｨ ﾂｨｨ09ｨ 05ｨ ﾂｨ 09ｨｰ06ｨｰｨｮ ｨｪｨ 08ｨｮ0302ｨｪｨｨｨｦ ｨｬ02ｨｬｨ 08ｨ ｨｪ04 ｨｰ06ｨｪ060705ｨ 09ｨｰｨ, ｨｪ06 ｨｪ ｨｨ0705ｨ ｨｪｨ 06ｨ ｨｴｨｨｨｦ 01ｨ 08ｨ ｨｺｨｰ0208 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨｨｦ. 01ｨ ｨｨｨ ｨｨｨｪｨｰ ｨｪ04ｨｬｨｨ 09 ｨｰ06 ﾂｨｺｨｨ ｨｪｨｨ ｨｨ07ｨｪｨｨ07 09ｨｪ0203ｨｪ ｨ ｨｪｨｨ07 ｨｪｨ 03ｨｨ ｨ ｨｪｨｨ04ｨｬ ｨｰｨ ｨｰ0907 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ ｨｪｨｮｨｰ0802ｨｪｨｪ02ｨｦ 09ｨｨ09ｨｰ02ｨｬ02 02ｨｨｨｰ060705ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺｨｨ01 ｨｬ02ｨｬｨ 08ｨ ｨｪ ｨ 09ｨｪ06 ｨｬｨｪ02ｨｪｨｨ06 01ｨ ｨｬｨ (01ｨ ｨｬｨ 0502ｨｦ, 2004), 05ｨｪ0106ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪｨ ｨｰｨｹ, ｨ ｨｮ01ｨｮ ﾂｨｨ 04ｨ ｨｬｨｺｨｪｨｮｨｰ06ｨｦ 09ｨｨ09ｨｰ02ｨｬ06ｨｦ, ｨｪｨ 01ｨ 08ｨ ｨｺｨｰ0208ｨｨ040609ｨ ｨｰｨｹ0907 ｨｬｨ ｨｺ09ｨｨｨｬｨ 05ｨｹｨｪ06ｨｦ ﾂｨｮ0909ｨｰ09ｨｨｨｰ0205ｨｹｨｪ0609ｨｰｨｹ06 ｨｺ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ06 09ｨｪ0203ｨｪ ｨ ｨｪｨｨ07, ﾂｨｰ06 ｨｨ ｨｰ ｨｪｨ 03ｨｨ ｨｰｨ ﾂｨｮ0909ｨｰ09ｨｨｨｰ0205ｨｹｨｪ0609ｨｰｨｹ ｨ 03ｨ 05ｨ 09ｨｹ 09 ｨｺ05ｨ 09ｨｰ0208ｨｨ04ｨ 02ｨｨｨｨ 02ｨｨ09ｨｰ0208ｨｪ ｨｪｨ ﾂｨ 05ｨｹｨｪ06 08ｨ 09ｨｪ06ｨｬ0208ｨｪ06 08ｨ ｨｪｨｪｨ ｨ ｨｲ02ｨｬｨｮ ｨｺ0502ｨｰｨｺｨｨ 0902ｨｰｨｹ ｨｬｨｨ080609ｨ 05ｨ ｨ ｨ ｨｰ04, ﾂｨ 09ｨｰ06 09 ｨ ｨｨ05ｨｨ ｨｬ02ｨｪ ｨ 06ｨｰｨｨ ﾂｨｪ06 08ｨ ｨｪｨｪ04ｨｬｨｨ 02ｨｨ09ｨｰ0208ｨｪｨ ｨｬｨｨ (08ｨｨ09. 4, ｨ, 01; 5, ｨ ). 02ｨ 04 ﾂｨｪ06 02ｨｨ09ｨｰ0208ｨｪ ｨｰｨｨｨｺｨｮ05ｨｮｨｬｨ ｨｺ060506ｨｺｨ 05ｨｨ040609ｨ ｨｪ ｨｬ02ｨｪｨｰｨ ｨｬｨｨ ｨ ｨｺｨｰｨｨｨｪ ｨｨｨｰ0609ｨｺ020502ｨｰｨ ｨｨ ｨｬ ｨｺ09ｨ 0006ｨｪｨ 05ｨｹｨｪ06ｨｦ 0902ｨｰｨｨ 08ｨ 09ｨｪ06ｨｬ0208ｨｪ06 08ｨ ｨｪ04 09 ｨｺ0502ｨｰ06 ﾂｨｪ06ｨｬ 06ｨ ｨｲ02ｨｬ02 (Snapp et al., 2003; Sparkes, 2011). 0308ｨｨ ｨｨ ｨ ｨｪｨｨｨｨ ｨｮ05ｨｹｨｰ08ｨ ｨｰ06ｨｪｨｺｨｨ ｨｰｨ ｨｺｨ ｨｰｨｹ ｨｰ ｨｨ ｨｨ0000ｨｮ04ｨｪ06 08ｨ ｨｪｨｪ ｨ ｨｲ02ｨｬ02 ｨｺ0502ｨｰｨｺｨｨ ｨｮ070506ｨｴ02ｨｪｨｪ ｨｨ09ｨｰ0208ｨｪ (08ｨｨ09. 3, 09, 00). 0306ｨｺｨ 04ｨ ｨｪ06, ﾂｨｰ06 ｨｮ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ ｨ ｨｺｨｰ06ｨｬｨｨ0604ｨｨｨｪ0609ｨ 07 09ｨｨ09ｨｰ02ｨｬｨ ｨｨ0008ｨ 02ｨｰ 0609ｨｪ0609ｨｪｨｮ ｨｹ ｨ ｨｪｨｨ04ｨ 02ｨｨｨｨ ｨｨ 01ｨｨｨｪｨ ｨｬｨｨｨｺ ｨｬ02ｨｪｨｰ ｨｪ0106ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪ06ｨｦ 09ｨｨ09ｨｰ02ｨｬ04 ｨｨ ｨｰ08ｨ 00ｨｨｨｺ ｨｨ ｨｪ0401 ｪ ｨｰ08ｨ ｨｪ ｨｰｨｪ ｨｨｨｺｨｮ05 (Sparkes, 2011). 08ｨ ｨｺｨｰ ｨｺ05ｨ 09ｨｰ0208ｨｨ04ｨ 02ｨｨｨｨ 05ｨｪ ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺ06ｨｦ 0902ｨｰｨｨ (08ｨｨ09. 4, ｨ ; 5, ｨ ), 01ｨｨｨｺｨｰｨｨ060906ｨｬ ｨ 0707ｨ 08ｨ ｨｰｨ ｨｹ0103ｨｨ (08ｨｨ09. 4, 00) ｨｨ ｨｰ08ｨ ｨｪ ｨｰｨｪ ｨｨｨｺｨｮ05 ｨｬ060302ｨｰ ｨｮｨｺｨ ｨ ｨｰｨｹ ｨｪｨ ﾂｨｮ0909ｨｰ09ｨｨｨｰ0205ｨｹｨｪ0609ｨｰｨｹ ｨ ｨｺｨｰｨｨｨｪ06-ｨｬｨｨ0604ｨｨｨｪ ｨｺ06ｨｬ070502ｨｺ09ｨ ｨｺ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07ｨｬ 01ｨ ｨｪｨｨ07. 06ｨ ｨｺｨ 07 ﾂｨｮ0909ｨｰ09ｨｨｨｰ0205ｨｹｨｪ0609ｨｰｨｹ 0102ｨｦ09ｨｰ09ｨｨｨｰ0205ｨｹｨｪ ｨｺｨ 04ｨ ｨｪｨ ｨｪｨ ｨｺ0502ｨｰｨｺｨ 01 ｨｺｨｮ05ｨｹｨｰｨｮ0804, ｨｪ ｨｨ 01ｨ ｨｪｨｨ0701, ｨｪｨ ｨｺ ｨ 06ｨｴｨｨ01 ｨｰ02, ﾂｨｰ06 ｨｨ ｨｹ040609ｨ ｨｪ04 ｨｪｨ ｨｬｨｨ (Kawarai et al., 2006). 0805ｨ 09ｨｰ0208ｨｨ04ｨ 02ｨｨ ｨｬ02ｨｪｨｰ ｨｰｨｨ 09 06ｨ 08ｨ ｨ ｨｪｨｨ02ｨｬ ｨｮ ﾂｨ 09ｨｰｨｺ ｨ 09ｨｰ060705ｨ 04ｨｬ04 ｨｰｨｨ07ｨｨ ﾂｨｪｨ ｨｺ0802ｨｰｨｨ08ｨｮ06ｨｴｨｨ01 ｨｺ0502ｨｰ06ｨｺ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ ｨｨ 03ｨｨ0906ｨｰｨｪ0401 (Snapp et al., 2003). 08ｨ ｨｺ 0708ｨ 09ｨｨ0506, ｨｬｨｨ080609ｨ ｨｪｨｨ02 ｨｺ05ｨ 09ｨｰ ｨｰｨ ｨｺｨｨ01 ｨｺ0502ｨｰｨｺｨ 01 ｨｪｨ ｨｨｨｰ ｨｺ06ｨｪｨｰ ｨｬ ｨｺ0502ｨｰｨｺｨｨ, ｨ 09ｨ ｨｬｨｨ 06ｨｪｨｨ ｨｰｨ ｨｪ04 ｨｮ070506ｨｴ02ｨｪｨｪ04ｨｬｨｨ 02ｨｨ09ｨｰ0208ｨｪｨ ｨｬｨｨ, ｨｮ07ｨ ｨｺ0609ｨ ｨｪｨｪ04ｨｬｨｨ ｨｰｨｪ ｨ ｨｺ02ｨｰ04. 06ｨｪ06ｨｦ 01ｨ 08ｨ ｨｺｨｰ0208 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨｨｦ ｨｪ 09 ｨｪｨ 03ｨｨ ｨｰｨ ｨｬ02ｨｪｨｰ04 ｨｨ04ｨｬ02ｨｪ0705ｨｨ ｨｬ0209ｨｰ ｨｦ 0506ｨｺｨ 05ｨｨ04ｨ 02ｨｨｨｨ ｨｨ 06ｨ 08ｨ 0406-

5 ｨ 0108ｨ 01ｨｨｨｬ0609 ｨｨ ｨｨ ｨｨ07ｨｪｨｨ02 09ｨｪ0203ｨｪ ｨ ｨｪｨｨ ｨ (30 ｨｬｨｨｨｪ) ｨｪｨ ｨｮ05ｨｹｨｰ08ｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｮ ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ02ｨｦ ｨｺｨｮｨｺｨｮ08ｨｮ ｨ 02ｨｨ07 05ｨｪ ｨ 04ｨｬｨ ｨｰｨｨ ﾂ0209ｨｺ06ｨｦ 0902ｨｰｨｨ, 01ｨｨｨｺｨｰｨｨ060906ｨｬ, ｨｰ08ｨ ｨｪ ｨｰｨｪ ｨｨｨｺｨｮ05 (ｨ ) ｨｨ ｨｰ0502ｨｪｨｨ02 ｨｬｨ ｨｰ08ｨｨｨｺ09ｨ ｨｬｨｨｨｰ060106ｨｪ0108ｨｨｨｦ (ｨ ) ｨ 06ｨｰ0907 ｨｪｨ 08ｨｮ0302ｨｪｨｨ07ｨｬｨｨ ｨｰｨｪ0609ｨｰｨｨ ｨｰ06ｨｪ060705ｨ 09ｨｰｨ (09). 02ｨ 08ｨ ｨ 00ｨｨｨｺ09ｨｨ080609ｨ ｨｪ04 09 ｨｺｨ ｨｬ ｨ ｨｪｨｨ07. 02ｨ 0604ｨｪｨ ﾂ02ｨｪｨｨ07 ｨｰ , ﾂｨｰ06 ｨｨ ｨｪｨ 08ｨｨ ｨ 05ｨｨ ｨ ｨ ｨｰ ｨｪｨ 0302ｨｬ 0905ｨｮ ﾂｨ 02 ｨ ｨ 02ｨｨ ｨｬ02ｨｪｨｰ0609 ｨｪ0609ｨｨ05ｨ, 09ｨｺ , 01ｨ 06ｨｰｨｨ ﾂｨｪ04ｨｦ ｨｨ ｨｪ02ｨｮ0708ｨ ｨｬ04ｨｦ 01ｨ 08ｨ ｨｺｨｰ ｨｪｨ 09ｨｰ0607ｨｴ ｨｬ ｨｹ 0605 ｨｺｨ ｨｺ ｨ ｨｪ , 06ｨｰ0902ｨｰ09ｨｰ0902ｨｪｨｪ06ｨｦ 04ｨ ｨ 03ｨｪ02ｨｦ03ｨｨ01 00ｨｨ04ｨｨ ｨｨ ﾂ0209ｨｺｨｨ , 09ｨｺ0506 ﾂｨ 07 09ｨｨｨｪｨｰ0204, ｨｨｨｪ00 ｨｨ ｨｰｨｰ08ｨ ｨｪ ｨｨ06ｨｪｨｪｨｮ06 ｨｬ0601ｨｨ00ｨｨｨｺｨ 02ｨｨ06 ｨ 0605ｨｹ03ｨｨｨｪ09ｨｰ09ｨ ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪ0401 ｨｨ 0902ｨｺ0802ｨｰ0608ｨｪ0401 ｨ 0205ｨｺ0609, 06ｨ ｨｴ02ｨｨ ｨｰｨｪｨ (Samali et al., 2010). 00 ｨｮ ｨｨ ｨ ｨｰ ｨｨｨｨ 09 ｨｺ0502ｨｰｨｺｨ 01 03ｨｨ0906ｨｰｨｪ0401 ｨｪｨ ｨ ｨ 02ｨｰ ｨ ｨｺ02ｨｨ07 09ｨｰ ｨ 0605, ｨｺ06ｨｰ0608ｨ ｨｦ09ｨｰ0902ｨｪｨｪｨ ｨｰｨ ｨｺ ｨｨ ｨｬｨｨ080609ｨ ｨｪｨｨｨｨ ｨｪ02ｨｦ ｨ ｨｰｨｨ09ｨｪ0401, 08ｨ ｨｺ ｨ ｨ ｨ ｨｪｨｨｨｦ ｨｨ 01ｨｨｨ ｨ 02ｨｰｨ ｨ ｨｴ02ｨｬ 0905ｨｮ ﾂｨ 02 05ｨｰｨ 0802ｨ ｨｺ02ｨｨ07 04ｨ 07ｨｮ09ｨｺｨ 02ｨｰ0907, ｨｺ060001ｨ 00ｨｨ04ｨｨ ｨｨ ﾂ0209ｨｺｨｨ02 ｨｨ05ｨｨ 09ｨｪ0203ｨｪｨｨ02 ｨｮ ｨｨ ｨｨ0706ｨｰ ｨｪｨ ｨｨｨｪ00ｨ ｨ 0205ｨｺ0609 ｨｨ ｨｪ ｨｨ05ｨｨ ｨｪ020708ｨ 09ｨｨ05ｨｹｨｪ ｨ 08ｨ ｨｪｨｪ0402 ｨ 0205ｨｺｨｨ ｨｪｨ ｨｺｨ 0705ｨｨ09ｨ 06ｨｰ ｨｰｨｨ 0902ｨｰｨｨ (Fulda et al., 2010). 02ｨｪｨ 01ｨ 08ｨ ｨｺｨｰ0208ｨｨ04ｨｮ02ｨｰ0907 ｨ ｨｺｨｰｨｨ09ｨ 02ｨｨ02ｨｦ ｨｰ ｨｰ ｨｰ : PKR ｨ ｨｪ06ｨｦ ER-ｨｺｨｨｨｪｨ (PERK), ｨ ｨｺｨｰｨｨ09ｨｨ08ｨｮ06ｨｴ 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 6 (ATF6) ｨｨ ｨｨｨｪ0604ｨｨｨｰ060504ｨ 09ｨｨ09ｨｨｨｬ ｨｬ02ｨｪｨｰｨ 1 (Ire1), ｨｺ06ｨｰ ﾂ ｨｹ ﾂ02ｨｪ ｨ ｨｺ02ｨｨ06 ｨｺ0502ｨｰｨｺｨｨ ｨｪｨ 0102ｨｪｨ ｨｰｨｮ08ｨｨ080609ｨ ｨｪｨｪ0402 ｨ 0205ｨｺｨｨ (UPR, unfolded protein reaction) ｨｬ0209ｨｰｨｪ06 ｨｰ08ｨｨ 0902ｨｰ09ｨｨ 06ｨｰ0902ｨｰｨ ｨ 0506ｨｺｨｨ08ｨｮ06ｨｰ ｨｰ08ｨ ｨｪ ｨｨ06, ｨｮ090205ｨｨ ﾂｨｨ09ｨ 06ｨｰ 05ｨｺ ｨｨ06 03ｨ ｨｪ0609 ｨｨ ｨｮ09ｨｨ05ｨｨ09ｨ 06ｨｰ ｨ 09ｨｨ09ｨｨｨｬ ｨｮｨｰｨｨ ｨ 01ｨ 02ｨｨｨｨ ｨｨ070207ｨｰｨｨ ｨｨ ｨｬ06ｨｴｨｪ0609ｨｰｨｹ 0802ｨ ｨｺ02ｨｨｨｨ UPR ｨｪ ｨｰｨ ｨｰ06 ﾂｨｪｨ ｨｰ06ｨｺ09ｨｨｨｺｨ 02ｨｨｨｨ 0902ｨｺ0802ｨｰｨｨ08ｨｮ02ｨｬ ｨｮｨｺｨｰ0609, 04ｨ 07ｨｮ09ｨｺｨ 02ｨｰ ｨｨｨｪ01ｨｮ02ｨｨ08ｨｮ02ｨｬｨ 07 00ｨｨｨ 0205ｨｹ ｨｺ0502ｨｰ06ｨｺ ｨ 09ｨｰ02ｨｪｨｨｨｦ ｨｬ060502ｨｺｨｮ050708ｨｪｨ 07 09ｨｨ00ｨｪｨ ｨｰｨｮ08ｨ ｨ 0409ｨｨｨｰｨｨ ｨ ｨｺ02ｨｨｨｨ UPR 0106 ｨｺ06ｨｪ02ｨ ｨｪ02 08ｨ 0903ｨｨ ｨ ｨｪｨ. 0306ｨｺｨ 04ｨ ｨｪ06, ﾂｨｰ06 ｨ ｨｺｨｰｨｨ09ｨ 02ｨｨ07 bzip17 ｪ ｨ ｨｪｨ ｨ ATF6 ｪ ｨｨ ｨｨｨｰ 0708ｨｨ ｨｬ 09ｨｰ (Liua, Howell, 2010) (Varadarajan et al., 2012) 0607ｨｨ09ｨ ｨｪ ｨ 0409ｨｰ0806ｨｦ ｨｨ 06ｨ 08ｨ ｨｰｨｨｨｬ06ｨｦ ｨ ｨｪｨｨ04ｨ 02ｨｨｨｨ 0605, ｨｺ06ｨｰ060804ｨｦ ｨｰ09ｨｮ02ｨｰ ｨｬｨｨ080609ｨ ｨｪｨｨ06 09ｨｰ ｨ ｨｺ02ｨｨｨｦ 09 ｨｺ0502ｨｰｨｺｨ 01 ｨｨ ｨｰｨ ｨｰ 0906ｨ 06ｨｦ ｨｬｨ 0903ｨｰｨ ｨ ｨｪ0602 ｨｨ ｨｨ00ｨｨ ﾂｨｪ ｨｬ ｨｨ080609ｨ ｨｪｨｨ02 09ｨｨ09ｨｰ02ｨｬ04 ｨｬ02ｨｬｨ 08ｨ ｨｪ 0902ｨｰｨｨ (EMR, endoplasmic reticulum membrane remodeling). 06ｨｰ06ｨｰ ｨ 08ｨ ｨｺｨｰ0208ｨｨ04ｨｮ02ｨｰ0907 ｨｺ05ｨ 09ｨｰ0208ｨｨ04ｨ 02ｨｨ02ｨｦ ｨｺ08ｨｮ07ｨｪ ｨｺ ｨｪｨｨ07 ｨ ｨｨ080609ｨ ｨｪｨｪ ｨｨ09ｨｰ0208ｨｪ ｨｦ ｨｬ ｨｨｨｨ 06ｨｪｨｨ ｨｪｨ 0706ｨｬｨｨｨｪｨ 06ｨｰ ｨ ｨ ｨｰ04 02ｨｨ09ｨｰ0208ｨｪ 0902ｨｰｨｨ, ｨｪｨｪ0402 ｨｪｨ ｨｬｨｨ 09 ｨｮ ｨｨ ｨｨ070208ｨ ｨ 08ｨｨｨｨ (08ｨｨ09. 4, ｨ ; 5, ｨ ) ｨｬｨｪ02ｨｪｨｨ06 ｨ 09ｨｰ , 05ｨｰ06ｨｰ 06ｨｰ0902ｨｰ 0902ｨｰｨｨ ｨｨ06ｨｪｨｪ06 ｨｺ06ｨｪ ｨ ｨｰｨｨ0902ｨｪ, ｨ 02ｨｰ ｨｺ08ｨ ｨｴ02ｨｪｨｨ02ｨｬ ｨｰ08ｨ ｨｪ ｨｨｨｨ ｨｨ 0902ｨｺ080202ｨｨｨｨ, ｨｪ ｨｰ05ｨｨ ﾂｨｨ02 06ｨｰ ｨｨ ｨｰｨｪ06ｨｦ 0802ｨ ｨｺ02ｨｨｨｨ 09ｨｰ ｨ 0605 (UPR) ｨｪ0609ｨｰｨｹ06 06ｨ 08ｨ ｨｰｨｨｨｬ (Varadarajan et al., 2012). 01ｨ ｨ ｨ 02ｨｬ0402 ｨｪｨ ｨｬｨｨ ｨｬ ｨｨ ﾂ0209ｨｺｨｨ02 ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07 ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ07, ｨｺ08ｨ ｨｴ02ｨｪｨｨ ｨｰｨ ｨｨ 06ｨ 08ｨ ｨｰｨｨｨｬ0609ｨｰｨｹ 00ｨｮｨｪｨｺ02ｨｨ06ｨｪｨ 05ｨｹｨｪ ｨ 08ｨ ｨｺｨｰ0208ｨｨ09ｨｰｨｨｨｺ 0708ｨｨ 0102ｨｺ06ｨｬ ｨｨｨｨ ｨ ｨｰ0902ｨｰ09ｨｰ09ｨｮ06ｨｰ 05ｨｰ06ｨｦ 06ｨ 08ｨ ｨｰｨｨｨｬ06ｨｦ 0802ｨ ｨｺ02ｨｨｨｨ 0902ｨｰｨｨ (Varadarajan et al., 2012), ｨｪ ｨｨ 00ｨｺ05ｨ 0909ｨｨ ﾂ0209ｨｺ06ｨｦ ｨ ｨｺ02ｨｨｨｨ 09ｨｰ ｨ 0605 (Fulda et al., 2010) ｨｬ060302ｨｪｨｨ02 ｨｰ08ｨ 00ｨｨｨｺｨ ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪ ｨｬｨ ｨｰ0208ｨｨｨ 05ｨ 09 08ｨ 09ｨｰｨｮｨｴｨｮ06 09ｨ ｨｺｨｮ0605ｨｹ ｨｬ060302ｨｰ ｨ 04ｨｰｨｹ 0708ｨｨ ﾂｨｨｨｪ06ｨｦ ｨｪｨｨ ｨ 02ｨｨｨｦ 09 ｨｰ06ｨｪ060705ｨ 09ｨｰ02, ﾂｨｰ ﾂ ｨｹ ｨｬ060302ｨｰ ｨｰｨｨ ｨｺ ｨｺ08ｨ ｨｴ02ｨｪｨｨ ｨｰｨ ｨｺ0502ｨｰ06ｨｺ 08ｨ 09ｨｰ070302ｨｪｨｨ02ｨｬ. 0102ｨｰ 0906ｨｬｨｪ02ｨｪｨｨｨｦ, ﾂｨｰ ｨｰｨｪ0609ｨｰｨｹ ｨｰ06ｨｪ060705ｨ 09ｨｰｨ, ｨ ｨ ｨｰ0205ｨｹｨｪ06, ｨ ｨｪ0609ｨｰｨｹ ｨｺ0502ｨｰｨｺｨｨ 0002ｨｪ0208ｨｨ080609ｨ ｨｰｨｹ ｨｰ08ｨ ｨｪ09ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪ04ｨｦ 0706ｨｰ02ｨｪ02ｨｨｨ 05 ｨｨ06ｨｪ ｨ, ｨｪ02-

6 130005ｨｨ07ｨｪｨｨ02 00ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨｪｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨｪｨｮ ｨ ｨｪｨｨ04ｨ 02ｨｨ06 ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ02ｨｦ ｨｺｨｮｨｺｨｮ08ｨｮ ｨｨ ｨｹｨｰ08ｨ 09ｨｰ08ｨｮｨｺｨｰｨｮ08ｨ ｨｺ0502ｨｰ06ｨｺ ｨｺ0608ｨｪ02ｨｦ 0708ｨｨ ｨ 0409ｨｰ0806ｨｬ 09ｨ ｨｪ0203ｨｪ ｨ ｨｪｨｨ ｨ ｨｰｨｬ ｨｪ06ｨｦ ｨｨ ﾂｨｨｨｪ04. ｨ, 09 ｪ 09ｨ ｨ ｨｪｨｨ ｨ 0106 ｨ ｨｰｨｬ ｨｪ060006; ｨ, 00 ｪ 09ｨ ｨ ｨｪｨｨ ｨ 0106 ｨ ｨｰｨｬ ｨｪ ﾁｨ 09ｨｰ06ｨｰｨ 08ｨ (09ｨｰ080205ｨｺｨｨ) ｨｰ06ｨｪ060705ｨ 09ｨｰｨ ｨｨ ｨｬｨ 02ｨｨｨｦ ｨｺ0502ｨｰ06 ﾂｨｪ ｨｰ02ｨｪ06ｨｺ ｨｬ ｨｨ06ｨｪｨ 05ｨｹｨｪｨ ｨ ｨｬ0705ｨｨｨｰｨｮ ｨ ｨｪｨｪ ｨ ｨｪｨｨ07. 02ｨ 08ｨ ｨ 00ｨｨｨｺ09ｨｨ080609ｨ ｨｪ ｨｨ ｨ ｨｰｨｬ ｨｪ06ｨｬ 01ｨ ｨｪｨｨｨｨ. 02ｨ 0604ｨｪｨ ﾂ02ｨｪｨｨ07 ｨｰ , ﾂｨｰ06 ｨｨ ｨｪｨ 08ｨｨ ｨ ｨｨｨｬ04ｨｦ ｨｰ0608ｨｨ ﾂｨｪ ｨｰ08ｨ ｨｪ ｨｰｨ 0609ｨｬ06ｨｰｨｨｨｺ0609, ｨｰ0907 ｨｺ08ｨｨｨｰｨｨ ﾂｨｪ04ｨｬｨｨ ｨｰ ｨｪ02 08ｨ 09ｨｰ070302ｨｪｨｨ07. 06ｨ ｨｺｨｨｨｬ 06ｨ 08ｨ 0406ｨｬ, ｨｮ090205ｨｨ ﾂ02ｨｪｨｨ02 00ｨｨ ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ ｨ 05ｨ 09ｨｰ02ｨｦ ｨｪ02ｨｦ 0008ｨ ｨｪｨｨ ｨｨ04ｨｨ ｨｨ ﾂｨｪ ｨ ｨｪｨｨｨｦ (2 ｪ4 0003ｨ ) 0708ｨｨ090601ｨｨｨｰ ｨｺ 09ｨｮｨｴ0209ｨｰ0902ｨｪｨｪ04ｨｬ ｨｨ04ｨｬ02ｨｪ02ｨｪｨｨ07ｨｬ 09 05ｨｪ0106ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪ06ｨｦ 09ｨｨ09ｨｰ02ｨｬ02, ｨｴｨｨｨｬ ｨ 05ｨｹｨｰ0208ｨ 02ｨｨｨｨ ｨ ｨｪ020505, ｨｮ ﾂｨ 09ｨｰ09ｨｮ06ｨｴｨｨ01 09 ｨｬ02ｨｬｨ 08ｨ ｨｪｨｪ06ｨｬ ｨｰ08ｨ 00ｨｨｨｺ02, ｨｨ ｨｬｨｨ080609ｨ ｨｪｨｨｨｨ 09 ｨｺ0502ｨｰｨｺｨ 01 08ｨ 09ｨｰ02ｨｪｨｨｨｦ 0802ｨ ｨｺ02ｨｨｨｨ 09ｨｰ ｨ 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 ). 0507ｨｨ0906ｨｺ 05ｨｨｨｰ0208ｨ ｨｰｨｮ ｨｪｨｨ09ｨｨｨｬ , 05ｨｮ , 05ｨｮｨｦｨｺ ｨｪｨ 09ｨｰｨｺｨ ｨｺ ｨｨ ｨ ｨｪｨｨ07ｨｬ ｨｬｨ ｨｪ0609ｨ ｨｨ07ｨｪｨｨ02ｨｬ 09ｨｰｨ ｨｰｨｨ ﾂ0209ｨｺ ｨｨ 01ｨｨｨｪｨ ｨｬｨｨ ﾂ0209ｨｺ ｨ ｨｪｨｨ07 ｨｪ ｨｰ0902ｨｪｨｪ ｨ ｨｰ ﾂｨｨｨｺ ｨ 01ｨｨ02ｨｪｨｰｨｪ ｨ ｨｰ ﾂｨｨｨｺｨｨ ｨｨ 09ｨｨ09ｨｰ02ｨｬ04. 7 : 64 ｪ67. 00ｨ ｨｦｨｪｨ ｨｨ0302ｨｪｨｨ07 ｨｮ 08ｨ 09ｨｰ02ｨｪｨｨｨｦ. 00.: 05ｨｪｨ ｨｪｨｨ ｨ ｨｬｨ 0502ｨｦ ｨ ｨｪ ｨｰｨｪｨ 07 09ｨｨ09ｨｰ02ｨｬｨ ｨｮ01ｨｨ09ｨｰ ｨ 09ｨｰ02ｨｪｨｨｨｦ. 0503ｨ.: ｨｰ0208ｨ ｨｮ0800. ｨｮｨｪ-ｨｰｨ. 424 c. 05ｨ 05ｨｹ02ｨｬｨ ｨｪ , 08ｨｮ ﾂｨｮｨｺ , 01ｨｮ080002ｨｪｨｨ ｨｪ ｨｨ070208ｨ ｨ 08ｨｨ ﾂ0209ｨｺ06ｨｦ 00ｨｨ04ｨｨ ｨｨｨｨ 09.: Abe F., Horikoshi K Hydrostatic pressure promotes the acidification of vacuoles in Saccharomyces cerevisiae. FEMS Microbiol. Lett. 130 : 307 ｪ312. Ache P., Bauer H., Kollist H., Khaled A. S., Al-Rasheid, Lautner S., Hartung W., Hedrich R Stomatal action directly feeds back on leaf turgor: new insights into the regulation of the plant water status from non-invasive pressure probe measurements. Plant J. 62 : 1072 ｪ1082. Bartels D., Sunkar R Drought and salt tolerance in plants. Critical Reviews in Plant Sciences. 24 : 23 ｪ58. Dziubinska H Ways of signal transmission and physiological role of electrical potentials in plants. Acta Soc. Bot. Pol. 72 : 309 ｪ318. Fernandes P. M., Domitrovic T., Kao C. M., Kurtenbach E Genomic expression pattern in Saccharomyces cerevisiae cells in response to high hydrostatic pressure. FEBS Lett. 556 : 153 ｪ160. Franks P. J Use of the pressure probe in studies of stomatal function. J. Exp. Bot. 54 : 1495 ｪ1504. Fulda S., Gorman A. M., Hori O., Samali A Cellular stress responses: cell survival and cell death. Int. J. Cell Biol. Article ID , p. 23. Galmｨｦs J., Ribas-Carbｨｮ M., Medrano H., Flexas J Rubisco activity in Mediterranean species is regulated by the chloroplastic CO 2 concentration under water stress. J. Exp. Bot. 62 : 653 ｪ665. Gross M., Jaenicke R Proteins under pressure. The influence of high hydrostatic pressure on structure, function and assembly of proteins and protein complexes. Eur. J. Biochem. 221 : 617 ｪ630. Iwahashi H., Odani M., Ishidou E., Kitagawa E Adaptation of Saccharomyces cerevisiae to high hydrostatic pressure causing growth inhibition. Nature. 579 : 2847 ｪ2852. Kawarai T., Arai S., Furukawa S., Ogihara H., Yamasaki M High-hydrostatic-pressure treatment impairs actin cables and budding in Saccharomyces cerevisiae. J. Biosci. Bioeng. 101 : 515 ｪ518. Knoblauch M., Peters W. S Mｨｹnch, morphology, microfluidics ｪ our structural problem with the phloem. Plant, Cell and Environment. 33 : 1439 ｪ1452. Liua J., Howell S. H Endoplasmic reticulum protein quality control and its relationship to environmental stress responses in plants. Plant Cell. 22 : 2930 ｪ2942.

7 ｨ 0108ｨ 01ｨｨｨｬ0609 ｨｨ Molina-Garcia A. D The effect of hydrostatic pressure on biological systems. Biotechnol. Genet. Eng. Rev. 19 : 3 ｪ53. Munns R Plant adaptations to salt and water stress: differences and Commonalities. Adv. Bot. Res. 57 : 1 ｪ32. Reynolds E. S The use of lead citrate at high ph as an electron opaque stain in electron microscopy. Cell Biol. 17 : 208 ｪ212. Rothschild L. J., Mancinelli R. L Life in extreme environments. Nature. 409 : 1092 ｪ1101. Samali A., Gerald U. F., Deegan S., Gupta S Methods for monitoring endoplasmic reticulum stress and the unfolded protein response. Int. J. Cell Biol : 1 ｪ12. Snapp E. L., Hegde R. S., Francolini M., Lombardo F., Colombo S., Pedrazzini E., Borgese N., Lippincott-Schwartz J Formation of stacked ER cisternae by low affinity protein interactions. J. Cell Biol. 163 : 257 ｪ269. Sparkes I Recent advances in understanding plant myosin function: life in the fast lane. Mol. Plant. 4 : 805 ｪ812. Tander B Improved uranyl acetate staining for electron microscopy. J. Electron. Microsc. Techn. 16 : 81 ｪ82. Tanino K. K., Chen H. H., Fuchigami L. H., Weiser C. J Abscisic acid increases terrestrial plant cell resistance to hydrostatic pressure. Plant Physiol. 98 : 745 ｪ748. Turgeon M The puzzle of phloem pressure. Plant Physiol. 154 : 578 ｪ581. Varadarajan S., Bampton E. T., Smalley J. L., Tanaka K., Caves R. E., Butterworth M., Wei J., Pellecchia M., Mitcheson J., Gant T. W., Dinsdale D., Cohen G. M A novel cellular stress response characterised by a rapid reorganisation of membranes of the endoplasmic reticulum. Cell Death and Differentiation. 19 : 1896 ｪ ｨｰｨｮ07ｨｨ05ｨ 28 I 2013 EFFECT OF HYDROSTATIC PRESSURE ON STRUCTURAL ORGANIZATION OF THE MAIZE ROOT CELLS F. A. Abdrakhimov, 1 M. A. Suslov, A. V. Anisimov Institute of Biochemistry and Biophysics, Kazan Scientific Center RAS; 1 abdfar@mail.ru We analyzed the effect of hydrostatic pressure at 2 ｪ4 MPa level on the growth and organization of corn seedlings cells. At pressure value of 2 MPa, we observed the clarification of the mitochondrial matrix, clusterization of endoplasmic reticulum elements with formation of ergastorplasm sites and the increasing number of small vacuoles in the cytoplasm. We detected the accumulation of vesicles with electron-dense content on the trans side of dictyosomes, which indicates a change in the endomembranous system traffic. The reduction of electronic density of tonoplast was accompanied by a reduction of osmiophilic inclusions in the vacuoles. Increasing the pressure up to 4 MPa showed the emergence of destructive alterations in the tonoplast. None of the variants showed any violation of plasmalemma integrity, as confirmed by coloring with Evans blue. When we reset the hydrostatic pressure to atmospheric pressure, the frequency of structural distortions of the tonoplast membrane was proportional to the pressure difference: the minimum was at reset from 2 to 0 MPa and the maximum was at reset from 4 to 0 MPa. Thus, it has been concluded, that the endomembraneous system is among the targets of pressure effect on the plant cells. K e y w o r d s: pressure, stress, endoplasmic reticulum, corn seedlings.