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ƽřżººūř ĩźţǀšŚººŞ ŶººƋ Ųººƿ Śººŝ ƱŻƹ ƫƺººĪƫƺƯ ƾŚººěǀƿƲ Źŵ
æÔ ƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ æèîåèæçæìèç ƪººƯŚƧšřźººƄůŹŵƲǀƿŚººěƾƫƺººĪƫƺƯƱŻƹŚººŝŲººƿŶººƋšŚººŞǀĩźţƽřżººūř Xanthogaleruca luteola ƱƹŹŚººƳŹřƺººųīźººŝ ĨººſƺſƱřŹŸººĭƱŚŤººƀƯŻ (Col.: Chrysomelidae) æ ŹƺěŚƐƗƮƿźƯƹçźĭŻźŝƲƀŰƯ*ƹæŹƺěƾƯźŰƯŶǀƘſ ƽŵƺſƢƿŚƤƃ æ ŢºǀŝźţƵŚĮºƄƳřŵ ÏÎÑÎÎÒ ÐÐÓ ƾŤºƀěơƹŶƴºƇƱřźºƸţ ƽŻŹƹŚºƄƧƾºſŚƴƃƵźºƄůƵƹźºĭƽŻŹƹŚƄƧƽƵŶƨƄƳřŵŽŹŶƯŢǀŝźţ ƵŚĮƄƳřŵ æ ÎÑÎÎÒ ÐÐÓ ƾŤƀěơƹŶƴƇƱřźƸţ ƾƿřŸƛƖƿŚƴƇƹƭƺƬƗƵƹźĭƽŻŹƹŚƄƧƽƵŶƨƄƳřŵŽŹŶƯ [email protected]ƾĪǀƳƹźŤĪƫřŢƀ욌ŞţŚĪƯƩƺŘƀƯ* Cryoprotectant contents of overwintering adults of elm leaf beetle, Xanthogaleruca luteola (Col.: Chrysomelidae) Sh. Soudi1, S. Moharramipour1&*, M. Barzegar2 and M. Atapour1 1. Department of Entomology, Faculty of Agriculture, Tarbiat Modares University, P.O Box: 14115-336, Tehran, Iran, 2. Department of Food Science and Technology, Faculty of Agriculture, Tarbiat Modares University. P.O Box: 14115-336, Tehran, Iran. *Corresponding author E-mail: [email protected] ƵŶǀĪģ ŵƺƃƾ Ư śƺƀŰƯ ƱƹŹŚƳ ƱŚŤųŹŵ šŚƟō ƲƿźţƮƸƯ Żř Xanthogaleruca luteola (Muller) ƱƹŹŚƳ Źřƺųīźŝ Ĩſƺſ šřźǀǀƜţ ƹ Ųƿ ŶƋƽŶƴƣ šŚŞǀĩźţ ƾƿŚſŚƴƃ ŹƺƔƴƯƶŝ .Ŷƴĩƾ ƯƽźĜſ ŻƺěŚƿŵ ŢƫŚů Źŵ ƪƯŚĩ šřźƄů ƪĪƃ ƶŝ řŹ ƱŚŤƀƯŻ ƶĩ ƾƬƇřƾƬĪƫřƽŚƷŶƴƣ šŚŞǀĩźţ ƹƽŹƹōƖưū æèíí ŢƄƸŞƿŵŹř Śţ æèíì źƸƯ Żř ƶƳŚǀƷŚƯ šŹƺƇƶŝ ƪƯŚĩ šřźƄů ŚƷƱō ƾƬƈƟ ƹ żĩƺƬĭŻƺƫŚƷƽźţ ƩƺŤƿŻƺƴƿřƺƿŚƯŶƃƽźǀĭƵŻřŶƳř (HPLC)LJŚŝƾƿřŹŚĩŚŝ ƖƿŚƯƾƟřźĭƺţŚƯƹźĩƁƹŹ ƶŝ ŚƷƱō ƱŶŝ Źŵ ŵƺūƺƯ ƱřƺƴƗƶŝ ƩƺŤƿŻƺƴƿřƺƿŚƯ ŢƔƬƛ ŶƳŵƺŝ ƱřŹŸĭƱŚŤƀƯŻ ƪƯŚĩ šřźƄů ƱŶŝ Źŵ ƵŶƃƾƿ ŚſŚƴƃŲƿ ŶƋ šŚŞǀĩźţƲƿźţƮƸƯ Żř ƩƹźƀǀƬĭ ƹ ŽƺƨƘƯ ƽƶƐŝřŹ ƱƹŹŚƳ Źřƺųīźŝ Ĩſƺſ ƱřŹŸĭƱŚŤƀƯŻ šřźƄů Źŵ ƵŶƃƾƿ ŚſŚƴƃ Ųƿ ŶƋ šŚŞǀĩźţ ƲƿźţƾƬƇř Żř ƾƨƿ źŨƧřŶů ƶŝ ƵŚƯ Źŷō Źŵ Ʊō ŢƔƬƛřƺƷƱŶƃ ŵźſ Śŝ ƶƧƽŹƺƏ r = ÍÕÎÑ P = ÍÍÎÑ ŵřŵ ƱŚƄƳ ŚƯŵ šřźǀǀƜţ Śŝ ƽŹřŵƾƴƘƯ ƶŝ èå t åÑ ƪƣřŶů ƶŝ ƵŚƯ ƲƿŵŹƹźƟ Źŵ ŜǀƧźţ Ʋƿř ƶƧƾƫŚůŹŵ ŢƟŚƿ ƂƿřżƟř źţ ƱŻƹ ƭźĭ źƷ Źŵ ƩƺƯƹźƨ ǀƯ èçÓ t çì ƵŚƯ ƽŵ Źŵ ŚƷƩř ƾ Ƭě ƹƽŶƴƣ šŚŞǀĩźţ ƪƧ ƕƺưŬƯ ƲƿźţƂǀŝ ŚƯřŢƟŚƿ ƂƷŚƧ ƪƯŚĩ ƽƵźƄů źţ ƱŻƹ ƭźĭ źƷ Źŵ ƩƺƯƹźƨ ǀƯ ƩƺŤƿŻƺƴƿřƺƿŚƯ ŶſŹƾƯ źƔƳ ƶŝ ƵŶƯō Ţſŵƶŝ ŪƿŚŤƳ ŽŚſř źŝ Ŷƃ ƵŶƷŚƄƯ źţ ƱŻƹ ƭźĭ źƷ Źŵ ƩƺƯƹźƨǀƯ æååì t ìÓ ŶƴƴƧƾƯ ŚƠƿřŚƯźſƶŝƱƹŹŚƳŹřƺųīźŝ ĨſƺſƪưŰţŹŵřŹƾưƸƯƂƤƳżƧƺƬĭƹŻƺƫŚƷƽźţ ƩƺŤƿŻƺƴƿřƺƿŚƯŲƿŶƋšŚŞǀĩźţƾƳřŹŸĭƱŚŤƀƯŻ ŻƺěŚƿŵƱƹŹŚƳŹřƺųīźŝ ĨſƺſƽŶǀƬĩƱŚĭĥřƹ Abstract The elm leaf beetle, Xanthogaleruca luteola (Muller), is one of the most destructive pests of elm trees that overwinters as diapausing adult. The main cryoprotectants and their seasonal fluctuations in the elm leaf beetles were analyzed based on the monthly level of polyhydric alcohol from October 2008 through June 2009, using high performance liquid chromatography (HPLC). The compounds myoinositol, trehalose, glucose and glycerol were found to be the most important polyols in the overwintering æèíîæåìƁźƿŸěæèíîèææŢƟŚƿŹŵ ÎÕ šřźƄůŹŵƲǀƿŚěƾƫƺƨƫƺƯƱŻƹŚŝŲƿŶƋšŚŞǀƧźţƽřżūřƱřŹŚƨưƷƹƽŵƺſ adults, of which myo-inositol content exhibited an inversely significant correlation with temperature fluctuations (r = -0.814, P = 0.014). The highest level of myo-inositol was observed in December (32.6 ± 2.7 µmol g-1 fresh body weight), while its lowest level was recorded in April (3.0 ± 0.4 µmol g-1 fresh body weight) The total concentration of all cryoprotectants was 100.7 ± 7.6 µmol g-1 fresh body weights in January. The results underline the influential role of myo-inositol, trehalose and glucose compounds in enhancing the cold tolerance of elm leaf beetles and their chance of survival during freezing winter days. Key words: elm leaf beetle, diapause, overwintering, cryoprotectants, myo-inositol Ʊŵƺŝ řŹřŵ ŢƬƗƶŝ ƶĩ ŶƴŤƀƷ ƾĭźŝƲƸě ƱŚŤųŹŵ ƶƬưū Żř Ulmus ƶƯŶƤƯ spp. ƱƹŹŚƳ ƽŚƷƶƳƺĭ ƹšřźƄůŻřƶƳƺĭíÓƱƺƴĩŚţŶƳřƶŤƟźĭ ŹřźƣƶūƺţŵŹƺƯŻŚŝźƿŵŻřƾŤƘƴƇƹƾŤƀƿŻƽŚƷŢ ǀƬŝŚƣ ŚƷƱō Żř ƾưĩ ŵřŶƘţ ŚƸƴţ ƶĩ Ţſř ƵŶƃ ƁŹřżĭ Ʊřźƿř Żř ƱŚŤųŹŵ Ʋƿř ƽřŻšŹŚƀų ƽŚƷƶƴĩ śƺƀŰƯ ƱŚŤųŹŵ Ʋƿř Ĩƿ ƽƶūŹŵ ŢƟō ƱřƺƴƗƶŝ Ɩǀſƹ Ƃƴĩřźě ƽƶƴƯřŵ ƹ LJŚŝ šŹŚƀų ŢƬƗƶŝ Żř Xanthogaleruca luteola (Muller) ƱƹŹŚƳ Źřƺųīźŝ Ĩſƺſ ŚƷƱō ƱŚǀƯ Źŵ ƶĩ ŶƳƺƃƾ Ư æíåè ƩŚſ Źŵ ŹŚŝ ƲǀŤƀŴƳ ƽřźŝ ŢƟō Ʋƿř(Dahlsten, 1994) Ţſř ŹřŵŹƺųźŝ ƽźŤƄǀŝ ŢǀưƷř ŹŵżǀƳƱřźƿřŹŵƹ(Hance et al., 1994; Miller & Ware, 2001)ŚĪƿźƯōŹŵŶƴƬƿźƯŢƫŚƿřŻřƽŵLjǀƯ ƶŤƟŚƿ ŹŚƄŤƳř Ʊřźƿř ƍŚƤƳ ƾƯŚưţ Źŵ ľŚŞƿźƤţ źƋŚůƩŚů Źŵ ƹ ŶƃƁŹřżĭ ŹŚƄƟř Ǝſƺţ æèçÑ ƩŚſ ƱŚŤųŹŵ īźŝ Żř ƶƿŸƜţ Śŝ ƱƹŹŚƳ Źřƺųīźŝ Ĩſƺſ (Davatchi & Moradeshaghi, 1962) Ţſř Źŵ ŢƯƹŚƤƯ ƂƷŚĩ śƺģ ŢǀƠǀĩ ƂƷŚĩ ŜŞſ ƱŚŤųŹŵ ũŚţ ƱŵźƧ ƵźƔƴƯŶŝ źŝ ƵƹLjƗ ƱƹŹŚƳ řŻƽ ŹŚưǀŝ ƪƯřƺƗ źƿŚſ ƹ ƶƿƺƳŚŧ šŚƟō ƽƶƬưů źŝřźŝ Źŵ ŚƷƱō ƞƘƋ ƹ ƾƐǀŰƯ ƽŚƷƂƴţ źŝřźŝ ŹŵƪƀƳèŚţçƱřźƿřŹŵƾƐǀŰƯƹƾƿŚǀƟřźƜūƎƿřźƃƶŝƶŤƀŝŢƟōƲƿř(Brewer, 1973)ŵƺƃƾ Ư ƹƱŚŤųŹŵĨŤſƺěźƿŻŹŵƪƯŚĩšřźƄůšŹƺƇƶŝ řŹƱŚŤƀƯŻƹ(Arbab et al., 1998)ŵŹřŵƩŚſ (Hance et al., 1994)Ŷƴĩƾ ƯƽźĜſŚƷƵŚĮƷŚƴě źƿŚſ ƽŚƷŚƯŵ śƺƬƐƯŚƳ šřźŧř Żř ƱŚĪƯř Ŷů Śţ ŶƿŚŝ ƾƳřŹŸĭƱŚŤƀƯŻ ƽƵŹƹŵ ƩƺƏ Źŵ šřźƄů ƾƬĪƫř ƽŚƷŶƴƣ Ŷǀƫƺţ ƱřŹŸĭƱŚŤƀƯŻ šřźƄů ƽŚƷƾ ĭĦƿƹ Żř ƾĪƿ ŶƴƴĩśŚƴŤūř ƱŚƄƳŶŝ źŝ ƲǀƿŚě Źŵ šřźƄů źŝ ƵƹLjƗ ƱŚŤƀƯŻ ƽƵŹƹŵ ƩƺƏ Źŵ ŚƷŶƴƣ Ʋƿř żŤƴſ ŶƃŚŝƾ Ư Ʈĩ ƾƫƺĪƫƺƯ ƱŻƹ Śŝ (Sømme, 1979)ŚƷšƺŞĪƴƗ ƹŚƷƶƴĩ (Holmstrup, 1995)ŚƷƭźĩ źǀƔƳƵŶƳŻšřŵƺūƺƯŻřƽŵřŶƘţ ƁŹřżĭ żǀƳ (Levitt, 1980) ƱŚƷŚǀĭ Źŵ ƾŤů ƹ (Sømme, 1982; Storey & Storey, 1992) ƱŚƯŵŚě ŢſřƵŶƃ ÎÖÎÐîåçèæƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ šŚŞǀĩźţ ƩƹźƀǀƬĭ źŝ ƵƹLjƗ Ŷƿōƾ Ư ŹŚưƃƶŝ šřźƄů Źŵ ƾƬĪƫř Ŷƴƣ ƲƿźţƩƺưƘƯ ƩƹźƀǀƬĭ ƱřŹŸĭƱŚŤƀƯŻ šřźƄů Żř ƽŹŚǀƀŝ Źŵ żǀƳ ƩƺŤǀƳŚƯ ƹ żĩƺƬĭ ƩƺŤǀŝŹƺſ ŻƺƫŚƷƽźţ źǀƔƳ ƽźĮƿŵ Mamestra brassicae L. ƽƵźě Ŝƃ ƽƵź ǀƠƃ Źŵ ŻƺƫŚƷƽźţ ƹ żĩƺƬĭ(Salt, 1961) ŵƺƃƾ Ư Ŷǀƫƺţ ƪƯŚĩ šřźƄů Źŵ żĩƺƬĭ ƹ ŻƺƫŚƷƽźţ ƩƺŤƿŻƺƴƿřƺƬƿŚſ ƩƺŤƿŻƺƴƿřƺƿŚƯ (Goto et al., 2000) Źŵ ƩƺŤǀŝŹƺſ (Watanabe & Tanaka, 1998) Aulacophora nigripennis (Motschulsky)Ʀſƺſ ƹ ƩƺŤǀŝŹƺſ ƩƹźƀǀƬĭ (Sømme, 1967) Pieris brassicae L.ƮƬĩ ƽƵŶ ǀƠſƽƶƳřƹźě ƽŚƷƵź ǀƠƃ ƹ ƩƺŤƿźţ (Sømme, 1965) Panonychus ulmi (Koch) ƾƿŚěƹŹř żƯźƣ ƽƶƴĩ ƽŚƷƮŴţ Źŵ żĩƺƬĭ ƩƹźƀǀƬĭƹ(Miller & Smith, 1975)Upis ceramboides (L.)ĨſƺſƪƯŚĩšřźƄůŹŵƩƺŤǀŝŹƺſ (Morrissey & Baust, 1976)Eurosta solidaginis (Fitch)žĮƯƽŚƷƹŹLJŹŵŻƺƫŚƷƽźţ ƹƩƺŤǀŝŹƺſ ƽƹŹƶĩƽŵŶƘŤƯšŚƤǀƤŰţƮƛŹƾ ƬƗŶƳř ƵŶƃ ƾƿŚſŚƴƃŲƿŶƋƽŶƴƣšŚŞǀĩźţƲƿźţƮƸƯ ƱřƺƴƗƶŝ šŚŞǀĩźţ Żř ƾƃŹřżĭ ƱƺƴĩŚţ ƾƫƹ Ţſř ƶŤƟźĭ šŹƺƇ šřźƄů Żř ƽŹŚǀƀŝ Źŵ Ųƿ ŶƋ šŚŞǀĩźţ ŢſřƵŶǀſźƳęŚģƶŝƱƹŹŚƳŹřƺųīźŝ ĨſƺſŹŵŲƿŶƋ Śŝ ƾĮƴţŚĮƴţ ƍŚŞţŹř šřźƄů Żř ƽŹŚǀƀŝ Źŵ šŚŞǀĩźţ Ʋƿř żŤƴſ ƲǀƿŚě ƽŚƷŚƯŵ źŝ ƵƹLjƗ (Tsumuki & Kanehisa, 1978; Shimada, 1982; Furusawa et al., 1987) ŵŹřŵŻƺěŚƿŵŻŚƟƶŝŵƹŹƹ ŵřŶƿƹŹ ƾƗƺƳ ŻƺěŚƿŵ Ɩƣřƹ Źŵ ŶƳƺƃƾ Ư ŻƺěŚƿŵ ŻŚƟ ŵŹřƹ ƾƳƺƯŹƺƷ ƮŤƀǀſ Ĩưĩ Śŝ šřźƄů ƹƽŹƺƳƾŤŝƺƏŹƾƿŚƯŵŶƗŚƀƯŚƳƾƐǀŰƯƎƿřźƃƶŝŲſŚěŹŵƶĩŢſřƵŶƃƽżƿŹƶƯŚƳźŝ ƾĪǀŤƳĥ ƽŚƷŹŚĩ ƹ ŻŚſ ŻƺěŚƿŵ ƽƵŹƹŵ ƩƺƏ Źŵ (Danks, 1987) ŶŤƟřƾ Ư ơŚƠţř ƾƿřŸƛ ŵřƺƯ ƱřŶƤƟ ŵźǀĭƾ Ư šŹƺƇŚƯźſƶŝ šřźƄůƪưŰţƂƿřżƟřŚŝ ƍŚŞţř ŹŵƾƠƬŤŴƯƾĪƿĥƺƫƺƿżǀƟƹƾƿŚǀưǀƃƺǀŝ ƽƵŹƹŵ ƩƺƏ Źŵ ĨƿĥƺƫƺƿżǀƟƽŚƷŹŚĩ ƹ ŻŚſ ŻřƾĪƿ ƲǀƿŚěƾƫƺĪƫƺƯ ƱŻƹ ŚŝƽŶƴƣ šŚŞǀĩźţ żŤƴſ (Tsumuki & Kanehisa, 1978; Shimada, 1982; Furusawa et al., 1987) ŵƺƃƾ Ư śƺƀŰƯ ŻƺěŚƿŵ ƽƵŻƺěŚ ƿŵ ƽŚƷƮŴţ Źŵ ŹŚŝƲ ǀŤƀŴƳƽřźŝ šřźƄůŹŵ ƩƺŤǀŝŹƺſƹ ƩƹźƀǀƬĭƾŬƿŹŶţ ƂƿřżƟř ľLjŨƯ ƵźƄů Ʋƿř źŝ ƵƹLjƗ (Chino, 1957) Ŷƃ ƵŶƷŚƄƯ Bombyx mori L. ƮƄƿźŝř ƭźĩ ƽƶƳřƹźě (Khani et al., 2007)Cydia pomonella (L.)ŜǀſƭźĩƽƵŻƺěŚ ƿŵƽŚƷƹŹLJŹŵŻƺƫŚƷƽźţ ƂƿřżƟř Chilo suppressalis Walker ŪƳźŝ ŹřƺųƶƣŚſ ƭźĩ ƽƵŻƺěŚ ƿŵ ƽŚƷƹŹLJ Źŵ ƩƹźƀǀƬĭ ÏÍ šřźƄůŹŵƲǀƿŚěƾƫƺƨƫƺƯƱŻƹŚŝŲƿŶƋšŚŞǀƧźţƽřżūřƱřŹŚƨưƷƹƽŵƺſ ƽŚƷƹŹLJ Źŵ ŻƺƫŚƷƽźţ ƹ żĩƺƬĭ ƩƺŤǀŝŹƺſ ƩƹźƀǀƬĭ ƹ (Atapour & Moharramipour, 2009) ŶƃƁŹřżĭ(Morrisey & Baust, 1976)Scolytus intricate (Ratzeburgi)Ĩſƺſ ŚƯźſ źŝřźŝ Źŵ ƵŶƳŻ ŵƺūƺƯ ŢƓŚƠů ŜŞſ ƾƬĩ ƮƀǀƳŚĪƯ ƹŵ ƢƿźƏ Żř ơƺƟ šŚŞǀĩźţ ŵƺƃƾƯ Ų ƿ ƽŚƷƶŤƀƷ ƪǀĪƄţ ŢƗźſ ƂƷŚĩ ŜŞſšŚŞǀĩźţ Ʋƿř ƽżưſř ƅřƺųæ ŶƳƺƃƾ Ư ƽŚƷŹŚŤųŚſ ƹ ƾƫƺƬſ ŌŚƄƛ ƽŹřŶƿŚě ŦƗŚŝ Ʋƿř źŝ ƵƹLjƗ ç ƹ (Lovelock & Bishop, 1959) (Gekko & Timasheff, 1981a, 1981b; ŵŵźĭƾ Ư ƾƫƺƬſ śō Ʊŵřŵ Ţſŵ Żř ƱŚƯŻ Źŵ ƾƴǀŘţƹźě Ŷů Śţ ŵŚưŬƳř ƽƶƐƤƳ ƾŬƿŹŶţ ƂƷŚĩ Śŝ ƾĭŵŻŲ ƿ ƶŝ ŽŚƀů ƽŚƷƶƳƺĭ Crowe et al., 1992) Ʋƿř Żř ƩƺƯ Ĩƿ ƽřŻř ƶŝ ƾĭŵŻŲ ƿ ƶŝ ŽŚƀů šřźƄů ŹŵŶƴƴĩƾ Ư śŚƴŤūř ƾĭŵŻŲ ƿ Żř ƱŚĪƯř ŶŝŚƿƾ Ư ƂƷŚĩ ŽƺǀƀƬſ ƽƶūŹŵ Ñ Śţ è ŵŚưŬƳř ƽƶƐ ƤƳ ƹ æíÒ śƹŷ ƽƶƐƤƳ šŚŞǀĩźţ ƾƫƺƬſ ŌŚƄƛ ŹŚŤųŚſ ƽŹřŶƿŚě ŜŞſ ƲǀƿŚě ƽŚƷŢƔƬƛ Źŵ šŚŞǀĩźţ Ʋƿř (Zachariassen, 1985) źţLJŚŝƽŚƷŚƯŵƾŤůƹƲǀƿŚěƽŚƷŚƯŵŹŵŚƷƲ ǀŘţƹźěŹŚŤųŚſźǀǀƜţŻřƹ(Williams, 1990)ŶƳƺƃƾƯ Źřƺųīźŝ Ĩſƺſ ƶĩŚŬƳōŻř (Storey & Storey, 1988) Ŷƴƴĩƾ Ư ƽźǀĭƺƬū ŵŚưŬƳř ƽƶƐƤƳ Żř ŶƘŝ ƩŚſ ŹŚƸŝ Źŵ ƲǀĮƴſ šřŹŚƀų ŵŚŬƿř ŜŞſ ƹ Ƶŵźĩ ƽźĜſ ŢǀƤƟƺƯ Śŝ řŹ ƱŚŤƀƯŻ ƱƹŹŚƳ ŹřŵŹƺųźŝ ƾƿLJŚŝ ŢǀưƷř Żř ŚƯźſ ƶŝ ŢƟō Ʋƿř ƪưŰţ ƹ ŻƺěŚƿŵ ŢǀƘƋƹ ƽƶƘƫŚƐƯ řŸƫ ŵƺƃ ƾ Ư Ʋƿř ƪưŰţ Źŵ ƾƿřżſƶŝ ƂƤƳ ƶĩ ƾưƸƯ Ųƿ ŶƋ šŚŞǀĩźţ ŢųŚƴƃ ŹƺƔƴƯƶŝ ƢǀƤŰţ Ʋƿř Ţſř ŢſřƵŶƃƭŚŬƳřŵŹřŵŚƯźſƶŝŢƟō ŚƷƁƹŹ ƹŵřƺƯ ƶƳƺưƳƽŹƹōƖưū ƽƵŶĪººƄƳřŵƱƹŹŚººƳƱŚººŤųŹŵƽƹŹŻřæèííŢººƄƸŞƿŵŹřŚººţæèíìźººƸƯŻřƶººƳƺưƳƽŹƹōƖººưū ƾƣźºƃ ÒΰÎÍ ' ƹƾƫŚưºƃ èÒ°ÑÑ 'ũźĩ ƱřźƸţƱŚŝƺţřŹŵƖƣřƹŽŹŶƯŢǀŝźţƵŚĮƄƳřŵƽŹŻƹŚƄĩ ƪºƿřƹřŻřƽŹřŵźºŝƶºƳƺưƳŹŵŢƫƺƸºſŹƺºƔƴƯƶŝ ŢƟźĭšŹƺƇƾƟŵŚƈţƪĪƃƶŝźŤƯ æèÒåƕŚƠţŹř ŚºƷƶºƬţƲºƿřŻřƶºƳŚƷŚƯŚºƷƶºƳƺưƳƹŶºƃƵŶºǀĤǀěƱŚºŤųŹŵƽƶƴţ ŹƹŵƶŝƾƿřƺƤƯ ƽŚƷƶƬţ ŹƺƿźƸƃ ŹŵƎſƺŤƯŹƺƏƶŝ ƶƧƱŶŝƱŻƹ ƽźǀĭƵŻřŶƳř ŻřžěƵŶƃƽŹƹō Ɩưū ƽŚƷƶƳƺưƳ .Ŷƿŵźĭ ƽŹƹōƖưū ƽŚƷśƺ ǀţƹźĪǀƯƪųřŵŵƺŝƭźĭƾƬǀƯ æåìæ ± åèìŵźƟźƷƽřŻřƶŝƽŹřŵźŝƶƳƺưƳƽƵŹƹŵ ƩƺƏ ÏÎÎÐîåçèæƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ ƵŵŚƠŤºſřŚŝżǀƳƎǀŰƯƽƶƳřŻƹŹƽŚƯŵŶƳŶƃƽŹřŵƶĮƳ ŽƺǀƀƬſƽƶūŹŵèåƽŚƯŵŹŵƹƪƿźŤſř ƪºŰƯŻřƽźŤƯƺºƬǀĩĨºƿƽƶƬºƇŚƟŹŵƶºĩźºĮŤǀģŹŵźƤŤºƀƯƾºſŚƴƃřƺƷƵŚĮŤƀƿřƽŚƷƵŵřŵ Żř ŶƃƪƇŚůŢƃřŵŹřźƣƽŹřŵźŝƶƳƺưƳ ƾƬĪƫřƽŚƷŶƴƣƽźǀĭƵŻřŶƳř ƶſŻřƪĪƄŤƯƶƳƺưƳźƷƱŶŝŹŵŵƺūƺƯƾƬĪƫřƽŚƷŶƴƣƱřżǀƯƽźǀĭƵŻřŶƳř ƹũřźŴŤſřƽřźŝ ƪųřŵ ƶŝ ƹ ƲĮưƷ ƾƴǀģ ƱƹŚƷ ƱƹŹŵ ŶƇŹŵ íå ƩƺƳŚţř źŤǀƫƾ ƬǀƯ æÒ Źŵ ƪƯŚĩ ƽƵźƄů ŢƗźſŚŝƶƤǀƣŵæÒšŶƯƽřźŝŚƷƶƳƺưƳ žĜſŶƃƪƤŤƴƯƽźŤǀƫƾ ƬǀƯçƪƿźŤſřƽŚƷśƺ ǀţƹźĪǀƯ ƪųřŵ ƹ řŶū śƺſŹ Żř ƶƳƺưƳ ƾƿLJŚŝ Ţưƀƣ Ŷƃ ĥƺǀƠƿźŤƳŚſ ƶƤǀƣŵ Źŵ Źƹŵ æçååå ŌLjų ŢŰţ Ʊƹō Źŵ ŚƷśƺ ǀţƹźĪǀƯ žĜſ ƹ Ŷƃ ƶŤŴƿŹ ƪƿźŤſř ƽŚƷśƺ ǀţƹźĪǀƯ Śţ ŶƳŶƃ Ƶŵřŵ Źřźƣ ŽƺǀƀƬſ ƽƶūŹŵ èÒ ƽŚƯŵ Źŵ (Mmert, model VS-1202V5, Germany) śƺſŹ ƶŝ źǀƐƤţ ŹŚŝ ƹŵ śō źŤǀƫƹźĪ ǀƯ çååƩLjů ƪƯŚĩ źǀŴŞţ Żř žěŵƺƃźǀŴŞţ ŚƷƱō ƩLjů ƽƶƬ ǀſƹƶŝ ŚƷƶƳƺưƳ ƵŚĮŤſŵƪųřŵ ƶŝ ƶƳƺưƳ ƢƿŹżţ Żř ƪŞƣŶƃ ƍƺƬŴƯ ľLjƯŚĩ ƹ ƶƟŚƋř ƵŶƳŚƯƾ ƣŚŝ ƵŚĮŤſŵ ƽƶƬ ǀſƹƶŝ ƾƬĪƫř ƽŚƷŶƴƣ Ŷƃ ƝŚƇ źŤƯƹźĪǀƯ åÑÒ ŸƟŚƴƯ Śŝ ƾĮƳźſźſ ƾƟŚƇ ƱƺŤſ ƶŝ żƸŬƯ (Waters, Milford, MA, USA) (HPLC) LJŚŝ ƾƿřŹŚĩ Śŝ ƖƿŚƯ ƾƟřźĭƺţŚƯƹźĩ ŶƃƽŻŚſřŶū(Ca 59305-U, 300 × 7.8 mm, Supelco-Sigma Aldrich)ŚƷšřŹŶ ǀƷƺŝźĩƽƶƿżŬţ Żř ƵŵŚƠŤſř ŵŹƺƯ ŻŚſŹŚĪƃō ƹ ƶƤǀƣŵ źŝ źŤǀƫƾ ƬǀƯ Ĩƿ ħźŰŤƯ ŻŚƟ ŢƗźſ śō ƪƯŚƃħźŰŤƯ ŻŚƟ ƽƶūŹŵ íå ƽŚƯŵ Źŵ ŚƷŶƴƣ ƽŻŚſřŶūŵƺŝ (Refractive Index Detector) ŢƀĪƃ ŜƿźƋ ƕƺƳ Ŷƃ ƾƿřŵŻŻŚĭ ƭƺǀƬƷ ŻŚĭ ƽƶƬ ǀſƹƶŝ ħźŰŤƯ ŻŚƟŶƃ ƭŚŬƳř ŻŚſŹŚĪƃō ƹ ƱƺŤſ ƽřźŝ ŽƺǀƀƬſ ƶŝ ƪƯŚĩ šřźƄů ƱŶŝ ƪĩ Żř ƵŶƃƶǀƸţ ƽŶƴƣ ƽƶƳƺưƳ Ʊō ƱŶƃŹřŶ ƿŚě ƹ ƵŚĮŤſŵ šŚưǀƔƴţ Żř žě ŶƃŢŞŧƱōƭřźĭƺţŚƯƹźĩƹƢƿŹżţHPLCƵŚĮŤſŵƶŝźŤǀƫƹźĪǀƯçåƮŬů ƾƴŰƴƯ ƮſŹ ƁƹŹ Żř ƵŶƃřŶū ƾƬĪƫř ƽŚƷŶƴƣ ƾưĩ ƽƶƿżŬţ Śƿ ƽźǀĭƵŻřŶƳř ŢƸū ƹ ƩƹźƀǀƬĭ ŻƺƫŚƷƽźţ żĩƺƬĭ ƽŚƷŵŹřŶƳŚŤſř ŹƺƔƴƯ Ʋƿř ƽřźŝ Ŷƃ ƵŵŚƠŤſř ƽŶƴŝƶūŹŵ ŵŹřŶƳŚŤſř ƾƴŰƴƯ žĜſ ƹ ƢƿŹżţ ƵŚĮŤſŵ ƶŝ ppm Ñååå Śţ æåå ƽŚƷŢƔƬƛ Śŝ ƩƺŤƿŻƺƴƿřƺƿŚƯ Ŷƴƣ ƕƺƳ ƽ ŹřŵŻŚŝ ƱŚƯŻ ƶŝ ƶūƺţ Śŝ Ŷƃ ƮſŹ ŢƔƬƛ źƷ ƽřźŝ ŢƔƬƛ źŝřźŝ Ĩǀě źƿŻ ŮƐſ ÏÏ šřźƄůŹŵƲǀƿŚěƾƫƺƨƫƺƯƱŻƹŚŝŲƿŶƋšŚŞǀƧźţƽřżūřƱřŹŚƨưƷƹƽŵƺſ ƶƳƺưƳ źƷ Źŵ Ŷƴƣ ŢƔƬƛ ŵŹřŶƳŚŤſř ƾƴŰƴƯ ƹ Ĩǀě źƿŻ ŮƐſ ƶŝ ƶūƺţ Śŝ ƹ ƶƳƺưƳ Źŵ ŵƺūƺƯ (Atapour et al., 2008)ŶƃƽźǀĭƵŻřŶƳř ƽŹŚƯōƽƶƿżŬţ ƁŹřżĭ(mean ± standard error)ŹŚǀƘƯƽŚƐųtƲǀĮƳŚǀƯšŹƺƇƶŝ ƵŶƯōŢſŵƶŝ šŚƗLjƏř ƽƶºƀƿŚƤƯƹ(one-way ANOVA)ƶºƟźƏĨ ºƿžƳŚºƿŹřƹƽƶºƿżŬţĨưĩŚŝŚƷƵŵřŵ ƽƶƿżŬţŶƃ ŹřżºƟřƭźºƳŻřƵŵŚƠŤºſřŚºŝŶºƇŹŵÒŮƐºſŹŵ ƾĩƺţƱƺƯŻōĨưĩŚŝƞƬŤŴƯ ƽŚƷŹŚưǀţƲǀĮƳŚǀƯ ŢƟźƿŸěƭŚŬƳřSPSS (15.0) ŦŰŝƹŪƿŚŤƳ ƹƪƣřŶºůƽŚºƯŵƲǀĮƳŚǀƯƱŶŝƾƿŚǀưǀƃšŚŞǀĩźţšřźǀǀƜţƽƹŹŚƯŵźǀŧŐţƾſŹźŝŹƺƔƴƯƶŝ ŢſŵƶŝƾſŚƴƃřƺƷƽŚƷ ƵŵřŵŻřƵŵŚƠŤſřŚŝ æèííŢƄƸŞƿŵřŚţæèíìźƸƯƵŚƯŻřƪƣřŶůƢƬƐƯƽŚƯŵ ŹŵŚºƯŵƪƣřŶůƲǀĮƳŚǀƯŵřŵƱŚƄƳƩŚſƵŚƯƲƿźţŵźſřŹƽŵƾſŚƴƃřƺƷƽŚƷƵŵřŵ æƪĪƃŶƯō ŵƺŝŽƺǀƀƬſƽƶūŹŵ ÒƪƣřŶůƢƬƐƯƽŚƯŵƹåçƵŚƯƽŵ æèííŢƄƸŞƿŵŹřŚţæèíìźƸƯŻřƢƬƐƯƪƣřŶůƽŚƯŵƹŚƯŵƪƣřŶůƽƶƳŚƷŚƯ ƲǀĮƳŚǀƯæƪĪƃ Fig. 1. Monthly mean of minimum temperature and absolute minimum temperature between October 2008 and June 2009. ÏÐÎÐîåçèæƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ ƩƹźƀǀƬĭƹżĩƺƬĭŻƺƫŚƷƽźţƩƺ ŤƿŻƺƴƿřƺƿŚƯŵřŵƱŚƄƳƪƯŚĩšřźƄůƱŶŝƽŚƷŶƴƣƽƶƿżŬţ ƲưƋDƹCçƪĪƃŶƳƺƃƾƯśƺƀŰƯƱƹŹŚƳŹřƺų īźŝĨſƺſŹŵŲƿŶƋƾƬƇřšŚŞǀĩźţ ŢºſŵƶºŝHPLCƵŚĮŤºſŵĨưĩƶŝƞƬŤŴƯƽŚƷƵŚƯ ƾƏżǀƳšŚŞǀĩźţƲƿřŢƔƬƛšřźǀǀƜţƶĪƴƿř ƶºĩŶƃƵŶƷŚƄƯŚƷƭřźĭƺţŚƯƹźĩŹŵż ǀƳƽźĮƿŵƽŶƴƣšŚŞǀĩźţƵŶƃźĩŷƽŚƷŶƴƣźŝƵƹLjƗŶƯō ƽřźŝƽŹřŵŻŚŝƱŚƯŻŢƃřŶƳŵƺūƹŚƷƱō ƾƿŚſŚƴƃƱŚĪƯřŚƿƹŶƳŵƺŝżǀģŚƳŹŚǀƀŝźƿŵŚƤƯƽřŹřŵŚƿ ŵƺºŝƶºƤǀƣŵæîæƹÎÒèæè ÒæåíŜºǀţźţƶŝƩƹźƀ ǀƬĭƹƩƺŤƿŻƺƴƿřƺƿŚƯżĩƺƬĭŻƺƫŚƷƽźţ Źřƺºųīźŝ ĨſƺſŹŵƮĩƾƫƺĪƫƺƯƱŻƹŚŝƽŶƴƣšŚŞǀĩźţƲƿźţƮƸƯŻř ƾĪƿBƹAçƪĪƃ ŹŵźºţƱŻƹƭźºĭźƷŹŵƩƺƯƹźƨǀƯÑåèŚţçÒšřźǀǀƜţƽƶƴƯřŵŚŝƶĩŵƺŝƩƺŤ ƿŻƺƴƿřƺƿŚƯƱƹŹŚƳ šřźºƄůF=ÎÎÕÖdf=ÔP < ÍÍÒŵřŵƱŚºƄƳřŹƽŹřŵƾ ºƴƘƯšřźºǀǀƜţƞƬŤŴƯƽŚƷƵŚƯ ƩƺŤƿŻƺƴƿřƺƿŚºƯƾĩŶºƳřźƿŵŚºƤƯƽřŹřŵƵŚºƯźºƸƯŹŵŻƺěŚºƿŵƽƶºǀƫƹřƪºůřźƯŹŵƵŶƃƽŹƹōƖưū ƹŻƺěŚºƿŵŢƟźºƄǀěŚºŝŜǀĩźţƲƿřƱřżǀƯŚƯřŶƳŵƺŝźţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯ ÓçtæÓ) èçÑìtçìƱřżǀƯƲƿźţƂ ǀŝƶŝƵŚƯƽŵŹŵƹŢƟŚƿƂƿřżƟřŪƿŹŶţƶŝż ǀƿŚěƩƺƏŹŵŚƯŵƂƷŚĩ řŹƽŹřŵƾ ºƴƘƯƝLjŤºųřƽŵƹŹŷōƽŚºƷƶƳƺưƳƶĩ ŹƺƏƶŝ ƽ ŶǀſŹźţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯ ƱōƱřżºǀƯŻƺěŚƿŵŢƀĪƃƹŚƯŵƾŬƿŹŶţƂƿřżƟřŚŝƱŚŤƀƯŻźųřƹřŹŵŶƳŵřŵƱŚƄƳŚƷƵŚƯźƿŚſŚŝ źƿŵŚºƤƯƽřŹřŵŻƺěŚƿŵžě ƽƶƬůźƯ ƲƿŵŹƹźƟŹŵƵŶƃƽŹƹōƖưūšřźƄů ŢƟŚƿƂƷŚĩƵŹŚŝƹŵ AèƪĪƃŶƳŵƺŝźţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯçîtåçƩƺŤƿŻƺƴƿřƺƿŚƯƾĩŶƳřŹŚǀƀŝ ŹŵŜºǀĩźţƲºƿřƱřżºǀƯŶƃƾƿŚſŚƴƃƪƯŚĩšřźƄůŹŵƶĩŵƺŝƽźĮƿŵŜǀĩźţŻƺƫŚƷƽź ţ šƹŚºƠţŢºƄƸŞƿŵŹřƹƲƿŵŹƹźƟƽŚƷƶƳƺưƳƎƤƟƹŢƃřŶƳ ƽŹřŵƾ ƴƘƯźǀǀƜţƱŚŤƀƯŻƹżǀƿŚěƩƺƏ ƲƿźºţƂ ǀºŝF=ÐÍÑdf=ÔP < ÍÍÒŶºƳŵřŵƱŚƄƳŚƷƵŚƯźƿŚſ ƽŚƷƶƳƺưƳ ŚŝřŹ ƽŹřŵƾ ƴƘƯ ŹŵƱōŹřŶºƤƯƲƿźºţƮºĩƹźºţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯ èçtÒÒŹŷōŹŵŻƺƫŚƷƽźţŹřŶƤƯ ŚŝżǀƳżĩƺƬĭCçƪĪƃŶƃƵŶƷŚƄƯźţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯ æåætåæƲƿŵŹƹźƟ šřźºǀǀƜţƞºƬŤŴƯƽŚºƷƵŚºƯŹŵźºţƱŻƹƭźºĭźºƷŹŵƩƺºƯƹźƨǀƯÒçŚºţîíšřźǀǀƜţƽƶƴƯřŵ ŹŵżĩƺºƬĭŹřŶƤƯƲƿźţƂ ǀŝ.BèƪĪƃF=ÑÖdf=ÔP < ÍÍÒŵřŵƱŚƄƳřŹƽŹřŵƾ ƴƘƯ ƽŚƷƶƳƺưƳŹŵƱōŹřŶƤƯƲ ƿźţƮĩƹ źţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯ ÑåçtÒ ƵŚƯƱŚŝō ƽŚƷƶƳƺưƳ żǀƳƩƹźƀǀƬĭCèƪĪƃŶƃƵŶƷŚƄƯźţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯ æçtæçŢƄƸŞƿŵŹř šřźƄůŹŵƲǀƿŚěƾƫƺƨƫƺƯƱŻƹŚŝŲƿŶƋšŚŞǀƧźţƽřżūřƱřŹŚƨưƷƹƽŵƺſ ÏÑ šřźºǀǀƜţƞºƬŤŴƯƽŚºƷƵŚºƯŹŵźºţƱŻƹƭźºĭźƷŹŵƩƺƯƹźƨǀƯÓÒŚţèèšřźǀǀƜţƽƶƴƯřŵŚŝ ŚºŝƶºƀƿŚƤƯŹŵŜǀĩźţƲƿřŹřŶƤƯŶƴģźƷF=ÒÕdf=ÔP < ÍÍÒŵřŵƱŚƄƳřŹƽŹřŵƾ ƴƘƯ DèƪĪƃŵƺŝħŶƳřŹŚǀƀŝźĮƿŵšŚŞǀĩźţ A B C D ƹ(A)ƩƹźºƀǀƬĭƹżĩƺºƬĭŻƺƫŚºƷƽźºţƽŚƷŵŹřŶƳŚŤſřƶŝƍƺŝźƯHPLCƭřźĭƺţŚƯƹźĩ çƪĪƃ ŶºƋšŚºŞǀĩźţƽŻŚſřŶūƶŝƍƺŝźƯƭřźĭƺţŚƯƹźĩƹƭřƾěƾěçåå(B)ƩƺŤƿŻƺƴƿřƺƿŚƯ (D)ƵŚƯƲƿŵŹƹźƟƹ(C)ƵŚƯƽŵƪƯŚĩšřźƄůŹŵŵƺūƺƯŲƿ Fig. 2. HPLC choromatograms of trehalose, glucose and glycerol (A) myo-inositol (B) (200 ppm) and chromatogram of cryoprotectants separation in April (C) and January (D). ÏÒÎÐîåçèæƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ P < ÍÍÒŢºƃřŵƽŹřŵƾ ºƴƘƯšřźºǀǀƜţƞºƬŤŴƯƽŚƷƵŚƯŹŵż ǀƳƽŶƴƣšŚŞǀĩźţƕƺưŬƯ ƽřŹřŵƵŚºƯƽŵƩŚºſƪºƈƟƲƿźţŵźºſŹŵƵŶƃƽŹƹōƖưūƪƯŚĩšřźƄů F=ÓÑdf=Ô ƽŚƷƶƳƺưƳƹ źţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯ æååìtìÓ)ŵƺŝƽŶƴƣšŚŞǀĩźţŹřŶƤƯƲƿźţƂ ǀŝ ƱŚºƄƳřŹźºţƱŻƹƭźºĭźƷŹŵƩƺƯ ƹźƨǀƯ èåÑtåÑšŚŞǀĩźţƲƿřŹřŶƤƯƲƿźţƮĩƲ ƿŵŹƹźƟ ÑƪĪƃŶƳŵřŵ BA DC ƩƹźºƀǀƬĭƹ(C)żĩƺºƬĭ(B) ŻƺƫŚºƷƽźţ (A)ƩƺŤƿŻƺƴƿřƺƿŚƯŢƔƬƛƲǀĮƳŚǀƯšřźǀǀƜţ èƪĪƃ æèííŢºƄƸŞƿŵŹřŚºţæèíìźºƸƯŻřƱƹŹŚƳŹřƺųīźŝĨſƺſƪƯŚĩšřźƄůŹŵ (D) ƱƺƯŻōŹŵŶƇŹŵÒŮƐſŹŵŹřŵƾ ƴƘƯƝLjŤųřŵƺūƹƭŶƗƽƵŶƴƷŵƱŚƄƳƶŝŚƄƯƝƹźů ŶƃŚŝƾ ƯTukey Fig. 3.Changes of myo-inositol (A), trehalose (B), glucose (C) and glycerol (D) content in adult of elm leaf beetle between October 2008 and June 2009. Values labeled with the same letters are not significantly different at the 5% level by Tukey’s test. ÏÓ šřźƄůŹŵƲǀƿŚěƾƫƺƨƫƺƯƱŻƹŚŝŲƿŶƋšŚŞǀƧźţƽřżūřƱřŹŚƨưƷƹƽŵƺſ ŹřƺºŴĭźŝĨſƺſƪƯŚĩšřźƄůŹŵŲƿŶƋƽŶƴƣšŚŞǀĩźţƕƺưŬƯƲǀĮƳŚǀƯšřźǀǀƜţ.ÑƪĪƃ ŵƺºūƹƭŶºƗƽƵŶºƴƷŵƱŚºƄƳƶŝŚºƄƯƝƹźůæèííŢƄƸŞƿŵŹřŚţæèíìźƸƯŻřƱƹŹŚƳ ŶƃŚŝƾ ƯTukeyƱƺƯŻōŹŵŶƇŹŵÒŮƐſŹŵŹřŵƾ ƴƘƯƝLjŤųř Fig. 4. Changes of total mean content in adult of elm leaf beetle between October 2008 and June 2009. Values labeled with the same letters are not significantly different at the 5% level by Tukey’s test. ƱŚǀƯŹŵƖƿŚƃŵřŶƿƹŹĨƿƱŚŤƀƯŻŹŵŚƷƩř ƾ ƬěƹƮĩƾƫƺĪƫƺƯƱŻƹŚŝƾƬĪƫřƽŚƷŶƴƣŶǀƫƺţ šŚŞǀĩźţƲƿřƾĭŵŻŲ ƿƶŝŽŚƀůƽŚƷƶƳƺĭŹŵ (Levitt, 1980)ŵƺƃƾ ƯśƺƀŰƯƵŶƳŻšřŵƺūƺƯ ƽŚƷŚƯŵŹŵƾƴǀŘţƹźěƽŚƷŹŚŤųŚſƹƾƫƺƬſŌŚƄƛŢƓŚƠůŜŞſŵŚưŬƳř ƽƶƐƤƳƂƷŚĩźŝƵƹLjƗ źǀƔƳšřźƄůŻřƽŵřŶƘţŹŵ(Crowe et al., 1983; Quinn, 1985; Williams, 1990)ŶƳƺƃƾ ƯƲǀƿŚě żºǀƳŹLJƺºƯŪƴºěƶºŝƾƷŚºĭƱŚŤºƀƯŻŹŵŚƷƩř ƾ ƬěƹƾƬĪƫřƽŚƷŶƴƣƱřżǀƯ ŚƷŹƺŞƳŻŻřƾųźŝƹŹLJ ƮĩľŚŤŞƀƳźƿŵŚƤƯŹŵšŚŞǀĩźţƲƿřƾĭŵŻŲ ƿƶŝŽŚƀůšřźƄůźŨĩřŹŵƾƫƹ(Salt, 1961)ŶſŹƾ Ư ŽŚƀůƽřƶƳƺĭƱƹŹŚƳŹřƺų īźŝĨſƺſ (Salt, 1959; Ring, 1981)ŶƳƺƃƾ ƯŶǀƫƺţƩƺƯƹźƨǀƯ ƵŚºƯźºƸƯŹŵƵŶºƃƽŹƹōƖºưūƪºƯŚĩšřźºƄůƱŶŝŵŚưŬƳřƽƶƐƤƳƲǀĮƳŚǀƯŢſřƾĭŵŻŲ ƿƶŝ ƹæíÑtåîƶºŝŜǀţźţƶŝ ƽŵƹŹŷōŹŵƱřż ǀƯƲƿřƶĩŵƺŝŽƺǀƀƬſƽƶūŹŵæèÑtåì ŵƺºŝŹřŵƾ ºƴƘƯƽŹŚºƯōźºƔƳŻřƂƷŚºĩƲºƿřƹŢºƟŚƿƂƷŚºĩŽƺǀƀƬſƽƶūŹŵ æíçtæè ƪººƯŚĩšřźººƄůŹŵ(Soudi & Moharramipour, 2009)F=ÕÖdf=ÔP < ÍÍÍÎ ÏÔÎÐîåçèæƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ ƱřżºǀƯƲƿźºţƂ ǀŝƶŝƵŚƯƽŵŹŵƾƬĪƫřƽŚƷŶƴƣƕƺưŬƯƱƹŹŚƳŹřƺųīźŝĨſƺſƱřŹŸĭ ƱŚŤƀƯŻ ƎºſƺţŵŚưŬƳřƹśƹŷƽƶƐƤƳƂƷŚĩƶĩƾƿ ŚŬƳōŻřŶǀſŹźţƱŻƹƭźĭźƷŹŵƩƺƯƹźƨǀƯåæ ŚţèƹæíÓŜǀţźţƶŝŵŚưŬƳřƹśƹŷ ƽƶƐƤƳƂƷŚĩŢſřŚƷƱōŢƐƬƛƶŝƶŤƀŝřƹšŚŞ ǀĩźţƲƿř ƂƷŚºĩƾƬºƇřƪºƯŚƗƱřƺţƾ ưƳŢǀƘƏŚƣŚŝřŸƫ(Zachariassen, 1985)ƩƺƯźƷƽřŻřƶŝƶūŹŵÑ ƲºƿřƶºĩŶºſŹƾ ƯźƔƳƶŝŵřŵŢŞƀƳƵźƄůƲƿřŹŵŲƿŶƋšŚŞǀĩźţƱřżǀƯƶŝřŹŵŚưŬƳřƽƶƐƤƳ ƶŤºƃřŵƂºƤƳƲǀƿŚºěƽŚºƷŚƯŵŹŵƾƴǀŘţƹźěƽŚƷŹŚŤųŚſƹƾƫƺƬſŌŚƄƛƩŵŚƘţƒƠůŹŵšŚŞǀĩźţ ŢºƟōƲºƿřŹŵƱŶºŝŵŚºưŬƳřƽƶºƐƤƳƂƷŚĩƹƽŶƴƣšŚŞǀĩźţƂƿřżƟřƱŚǀƯƾƳŚƯŻƢŝŚƐţŶƴƃŚŝ ƾƳřŹŸºĭƱŚŤºƀƯŻƽĦţřźŤºſřŹŵƾƿřżºſƶºŝƂºƤƳšŚŞǀĩźţƲƿřżŤƴſƶĩŶƃŚŝƱōźĮƳŚǀŝŶƳřƺţƾ Ư Ŷƴĩƾ ƯŚƠƿřƱƹŹŚƳŹřƺųīźŝĨſƺſ ƱƹŹŚºƳŹřƺųīźŝĨſƺſŹŵƵŶƃ ŚſŚƴƃƾƬƇřšŚŞǀĩźţŻřƾĪƿƱřƺƴƗƶŝƩƺŤ ƾƿ ƿŻƺƴƿřƺƿŚƯ ƽŚºƷĨºſƺſźǀƔƳƾţřźƄůŹŵƹŵƺƃƾ ƯśƺƀŰƯƱŚƿŚěŶƴŝźƿŚſƱŚǀƯŹŵśŚƿƮĩŜ ǀĩźţĨƿ Epilachna vigintioctomaculata MotschulskyħŻƹŶƄƠĩ(Block & Sømme, 1983)Carabidae Achaearanea tepidariorum (Koch)ƾĮƳŚųšƺŞĪƴƗƹ(Kostal et al., 1996; Hoshikawa, 1987) ŲºƿŶºƋƽŶºƴƣšŚºŞǀĩźţżŤƴſĨƿźŰţŜŞſƶĩƾƿŚƯŵŢſřƵŶƃƾƿŚſŚƴƃ(Tanaka, 1995) ÒŚºţźƠºƇƽŚºƷŚƯŵŹŵƾºƫƹŵŹřŵŹřźºƣŽƺǀºƀƬſƽƶūŹŵÒŚţźƠƇƽƵŵƹŶŰƯŹŵŵƺƃƾ Ư źºŝ(Storey & Storey; 1991)ŶºŝŚƿƾ ºƯƂƿřżºƟřšŚŞǀĩźţƲƿřżŤƴſŢƗźſŽƺǀƀƬſƽƶūŹŵ ŢºƔƬƛƂƿřżºƟřŹŵƾƿřżºſƶºŝźǀŧŐºţƲǀƿŚºěƽŚƷŚƯŵŶſŹƾ ƯźƔƳƶŝƵŶƯōŢſŵƶŝŪ ƿŚŤƳŽŚſř ŹŵƩƺŤƿŻƺƴƿřƺƿŚºƯƱřżºǀƯƲƿźţƂǀŝƶƧ ƽŹƺƏ ŵŹřŵƱƹŹŚƳŹřƺųīźŝĨſƺſŹŵƩƺŤ ƶŝ ƿŻƺƴƿřƺƿŚƯ żŤƴºſĨºƿźŰţƶºƧŶºſŹƾºƯźºƔƳƶŝŶƃƵŶƷŚƄƯƵŚƯƽŵƹŹŷōƾƴƘƿƩŚſ ƽŚƷƵŚƯƲ ƿźţŵźſ ƹŽƺºƨƘƯƽƶºƐŝřŹźºƔƳƲºƿřŻřƹŶºƃŚŝƲ ǀƿŚºěƽŚƷŚƯŵŶƴƯŻŚǀƳƵźƄůƲƿřŹŵƩƺŤƿŻƺƴƿřƺƿŚƯ P=ÍÍÎÑŶºƃƵŶƷŚºƄƯƩƺŤƿŻƺƴƿřƺƿŚºƯŢºƔƬƛŚºŝƶºƳŚƷŚƯƽŚºƯŵƲǀĮƳŚºǀƯƱŚǀƯƽŹřŵƾƴƘƯ ŵƹŹƹŚŝƶƧƽŹƺƏ ŵŹřŵŻƺěŚƿŵŚŝƾĪƿŵżƳƍŚŞţŹřƩƺŤƿŻƺƴƿřƺƿŚƯšřźǀǀƜţľLJŚưŤůřr=ÍÕÎÑ ƶŝ ŹŵƹŢºƟŚƿƂƿřżƟřŪƿŹŶţƶŝƩƺŤ ƿŻƺƴƿřƺƿŚƯŢƔƬƛƵŚƯźƸƯŹŵŻƺěŚƿŵƂ ǀ쯌ƟƶŝƪƯŚĩšřźƄů ŶǀſŹźŨĩřŶůƶŝšŚŞǀĩźţƲƿřŢƔƬƛŵźŝƾ ƯźſƶŝŻƺěŚ ƿŵƢưƗŹŵƵźƄůľLJŚưŤůřƶĩƽŵƹŹŷō ŻŚºƟƶºŝšřźºƄůŵƹŹƹƹŻƺěŚºƿŵƱŚºƿŚěŚºŝŹŚºƸŝŹŵƩƺŤƿŻƺƴƿřƺƿŚºƯƾƳŚºƸĭŚƳƂƷŚĩƲǀƴĤưƷ ÏÕ šřźƄůŹŵƲǀƿŚěƾƫƺƨƫƺƯƱŻƹŚŝŲƿŶƋšŚŞǀƧźţƽřżūřƱřŹŚƨưƷƹƽŵƺſ ƭŚưţřƽƶƳŚƄƳ ƱřƺƴƗƶŝ ŹŚƸŝŹŵŚƷƩř ƾ ƬěƹƾĪƫřƽŚƷŶƴƣƂƷŚĩŵŹřŵƾưǀƤŤƀƯƍŚŞţŹřŻƺěŚƿŵžě ƂƷŚºƧŚºŝƶºƧƽŵƺūƹŚŝ(Tsumuki, 1990)ŢſřƵŶƃƁŹřżĭšřźƄůŻřƽŹŚǀƀŝŹŵŻƺěŚƿŵ ƩƺŤƿŻƺƴƿřƺƿŚºƯŹřŶºƤƯƱƹŹŚºƳŹřƺºųīźºŝĨºſƺſƱřŹŸĭƱŚŤƀƯŻšřźƄůƱŶŝŵŚưŬƳř ƽƶƐƤƳ P=ÍÍÓÎŶºƄƳŶƷŚºƄƯŶºƇŹŵÒƽŹŚºƯōŮƐºſŹŵƽŹřŵƾƴƘƯ ƽƶƐŝřŹƾƫƹ ŢƟŚƿƂƿřżƟř r=ÍÔÖÏ ŹřƺºųīźºŝĨºſƺſƱřŹŸĭƱŚŤƀƯŻƪƯŚĩšřźƄůŹŵƶĩ ŢſřƽźĮƿŵŜǀĩźţŻƺƫŚƷƽźţ ƵŶƃƁŹřżĭżǀƳšřźƄůŻřƽŵŚƿŻŵřŶƘţŹŵŲƿŶƋŜǀĩźţĨƿƱřƺƴƗƶŝƹŶƃ ƾƿŚſŚƴƃƱƹŹŚƳ ƮƀǀƳŚĪƯ(Rains & Dimock, 1978; Shimada & Riihimaa, 1990; Kostal & Simek, 1995)Ţſř ƵŶºƃƶŤųŚƴƃƪƯŚĩŹƺƏƶŝŻƺƴƷŚƯźſƶŝƱřŹŸĭ ƱŚŤƀƯŻšřźƄůƪưŰţŹŵŻƺƫŚƷ ƽźţƢ ǀƣŵƪưƗ ƽźǀĭƺºƬūƾƿŚºƯŵƽŚƷĨƃŻřžěƲ ǀŘţƹźěƽŚƷŹŚŤųŚſŜƿźŴţŻřľLJŚưŤůřŜǀĩźţƲƿřŢƀǀƳ ŵƺºƃƾ ºƯƽƺǀţřŶǀºƀĩřƽŚºƷƂƴºĩřƹŻřƾºƃŚƳŜǀºſōźŝřźŝŹŵƩƺƬſŢƓŚƠůŜŞſƹŶƴĩƾ Ư ŶƃƵŶƷŚƄƯLJŚŝƽŚƷŢƔƬƛŹŵƱŚŤƀƯŻƹż ǀƿŚěƩƺƈƟŹŵŜǀĩźţƲƿř.(Benaroudj et al., 2001) ƽŹřŵƾ ºƴƘƯƽƶºƐŝřŹƲǀƴĤưƷ ŵƺŝŢŝŚŧľŚŞƿźƤţƹŵźĪƳƾƳřŶƴģźǀǀƜţƩƺƈƟƲƿřƾƏƱōŹřŶƤƯƹ P=ÍÎÔÕŶºƄƳƵŶƷŚºƄƯƶºƳŚƷŚƯƽŚºƯŵƲǀĮƳŚºǀƯšřźºǀǀƘţŚŝƵŵŚƯƲƿřŢƔƬƛšřźǀǀƜţƱŚǀƯ ƲǀƿŚºěƽŚºƷŚƯŵƦºƿźŰţŻřƪƤŤƀƯŜǀƧźţƲƿřżŤƴſƶƧŶſŹƾƯźƔƳƶŝ ƲƿřźŝŚƴŝr=ÍÓÐÐ ŶƃŚŝƵŵźŝLJŚŝřŹƱōŢƔƬƛƱřżǀƯŚƯźſƶŝŵƹŹƹŻřƪŞƣƵźƄůƹƵŵƺŝ ŵŹƺƯƽŚƷƵŚƯŹŵƱōƱřż ǀƯƶĩŵƺŝżĩƺƬĭŶƃƾƿŚſŚƴƃšřźƄůƲƿřŹŵƶĩƽźĮƿŵŜǀĩźţ ƹżǀƿŚºěƩƺºƈƟƶºŝƍƺºŝźƯżƧƺºƬĭŢºƔƬƛƲƿźţƂǀŝ ŵřŵƱŚƄƳřŹƽŹřŵƾ ƴƘƯšřźǀǀƜţƶƘƫŚƐƯ ŶºƄƳƵŶƷŚºƄƯŜºǀƧźţƲºƿřźƿŵŚºƤƯƱŚºǀƯƽŹřŵƾºƴƘƯƝLjŤºųřƩƺƈƟƲƿřƾƏƹŵƺŝƱŚŤƀƯŻ ŶºƄƳƵŶƷŚºƄƯƶºƳŚƷŚƯƽŚƯŵƲǀĮƳŚǀƯŚŝƽŹřŵƾ ƴƘƯƶƐŝřŹŜǀĩźţƲƿřšřźǀǀƜţŵŹƺƯŹŵƲǀƴĤưƷ ŻřƪŞƣŻƺƫŚƷƽźţŶƴƳŚưƷżǀƳŜǀƧźţƲƿřżŤƴſƶƧŶſŹ ƾƯźƔƳƶŝřŸƫ r=ÍÍÍÒP=ÍÖÖÐ ŶƃŚŝƵŶƃŻŚƛōŚƯźſƽƵŹƹŵ ĨºſƺſƱřŹŸºĭƱŚŤºƀƯŻšřźƄůŹŵřŹƱřżǀƯƲƿźţƮĩšŚŞ ǀĩźţźƿŚſƶŝŢŞƀƳƩƹźƀǀƬĭ řŹƾºƈŴƄƯŶºƳƹŹƩƹźƀǀƬĭŢƔƬƛšřźǀǀƜţƾſŹźŝŵřŵƅŚƈŤųřŵƺųƶŝƱƹŹŚƳŹřƺųīźŝ ƶºƳŚƷŚƯƽŚºƯŵƲǀĮƳŚºǀƯƹŜºǀĩźţƲºƿřŢºƔƬƛšřźºǀǀƜţƱŚǀƯƽŹřŵƾ ƴƘƯƽƶƐŝřŹƹŵřŶƳƱŚƄƳ ÏÖÎÐîåçèæƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ ƱŻƹŚºŝ ƽŶºƴƣšŚºŞǀĩźţżŤƴºſ ƾƿŚºƳřƺţ ƾƬĩŹƺºƏƶŝ r=ÍÒÑÖP=ÍçëåŶƄƳƵŶƷŚƄƯ ŚºƯŵƾŬƿŹŶºţƂƷŚºĩŚºŝƱƹŹŚºƳŹřƺųīźŝĨſƺſƱřŹŸĭ ƱŚŤƀƯŻšřźƄůŹŵƲ ǀƿŚěƾƫƺĪƫƺƯ ŶǀºſŹźŨĩřŶºůƶºŝƽŶºƴƣšŚŞǀĩźţŢƔƬƛƕƺưŬƯƩŚſƽŚƷƵŚƯƲ ƿźţŵźſŹŵƹŢƟŚƿƂƿřżƟř ƪºƧƲǀĮƳŚºǀƯƹŚºƯŵƽƶºƳŚƷŚƯƢºƬƐƯƪƣřŶºůƱŚºǀƯƽŹřŵƾƴƘƯƹŽƺƨƘƯ ƽƶƐŝřŹƶƧƽŹƺƏ ƶŝ ŹŵšŚºŞǀĩźţƲºƿřŶºſŹƾºƯźºƔƳƶŝr=ÍÕÑÓP=ÍÍÐÑŶƃƵŶƷŚƄƯŲƿŶƋšŚŞǀƧźţ ƽŚƷŹŚŤųŚºſŢºƓŚƠůŜ޺ſľLJŚºưŤůřƱƹŹŚºƳŹřƺųīźŝĨſƺſƱřŹŸĭ ƱŚŤƀƯŻƪƯŚĩšřźƄů ƹƵŶºƃƲǀƿŚºěŢŞƀƳƶŝ ƽŚƷŚƯŵŹŵƾŤůƾƿŚƯźſƽŚƷħƺƃźŝřźŝŹŵ ƾƫƺƬſŌŚƄƛƹƾƴǀŘţƹźě ŢƃřŵŹŚƔŤƳřƩŚſƽŚƷƵŚƯƲ ƿźţŵźſŹŵřŹŚƯźſƶŝŢŞƀƳšřźƄůƪưŰţźŨĩřŶůƱřƺţƾ Ư ƖŝŚƴƯ Arbab, J., Jalali, J. & Sahragard, A. 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