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ĩźţ ƂƤƳ ŹřźƟ šŚŞ ŶƗŚƈŤƯ ƵŶƃ Żř ŭƺƐſ Ʃƹř ƹ ƭƹŵ řŸƛƾƿ Ư
ÑÒ ƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ æèîçèèèéêêë ŶºǀƿƺŤƿŻřŹŚěŹƺºŞƳŻ ƾŝŚƿƱŚŝżǀƯŹŚŤƟŹźŝƾƿřŸƛƭƹŵƹƩƹřŭƺƐſŻřƵŶƃ ŶƗŚƈŤƯ ŹřźƟšŚŞǀĩźţƂƤƳ Tamarixia radiata (Hym.: Eulophidae) ƾƫLjūƲǀƯřŶưŰƯƹ*ƾƴƿŶƫřŌŚǀƋƽŶƸƯƾƿřźƣƾƬŞƤƯƲǀƯř ƱŚŬƴƀƟŹƱŚŬƴƀƟŹŪƗźƈƗƾ ƫƹƵŚĮƄƳřŵƽŻŹƹŚƄĩƽƵŶĪƄƳřŵ ƾĪƃżěƵŚǀĭƵƹźĭ [email protected]ƾĪǀƳƹźŤĪƫřŢƀ욌ŞţŚĪƯƩƺŘƀƯ* Role of volatile compounds emitted from the first and second trophic levels in host finding behavior of Tamarixia radiata (Hym.: Eulophidae) A. Moghbeli Gharaei, M. Ziaaddini* and M. A. Jalali Department of Crop Protection, Agriculture College, Vali-e-Asr University of Rafsanjan, Rafsanjan, Iran. *Corresponding author, E-mail: [email protected] ƵŶǀĪģ ŹŵŚºƷƱōŵźºĪƬưƗƂƿřżºƟřŜºūƺƯŶºƴƳřƺţİƯƹ Ŷƴƴĩƾ ƯŚƠƿřŶǀƿƺŤƿŻřŹŚěƽŚƷŹƺŞƳŻƽźĮƃƹŚĩŹŚŤƟŹŹŵřŹƾưƸƯƂƤƳŚƷƶŰ ƿřŹ ƶºŝTamarixia radiata (Waterston)ŶºǀƿƺŤƿŻřŹŚěŹƺºŞƳŻŲºſŚěƲǀºǀƘţƝŶºƷŚºŝƂƷƹĦºěƲƿřŶƳŵźĭƦƿĥƺƫƺǀŝƩźŤƴĩįŚƷƶƯŚƳźŝ ƪĪƃ YŪƴſƾƿ ŚƿƺŝŻřƵŵŚƠŤſřŚŝDiaphorina citri KuwayamašŚŞĩźƯƾƿŚǀſōƪǀƀěƾŤƀƿŻƞƬŤŴƯƪůřźƯƾƿŚǀưǀƃƽŚƷħźŰƯ ƱŚºƿźūšŶƃƹžĩƺƫæÓååŹƺƳšŶƃŶƇŹŵ ÓÍtÎÍƾŞƀƳŢŝƺƏŹŽƺǀƀƬſƽƶūŹŵÏÔtÎƽŚƯŵƾƷŚĮƄƿŚƯŻōƎƿřźƃŹŵƹ (HIPVs)ƾƿŚƤƫřŹřźƟšŚŞǀĩźţƹ (GLVs)żŞſ ƽŚƷīźŝŹřźƟšŚŞǀĩźţƂƤƳ ƲǀƴĤưƷŶƃƭŚŬƳř ƶƤǀƣŵźŝźŤǀƫƾ ƬǀƯÐÍÍƱřżǀƯƶŝřƺƷ ƁŹŚºƯšƹźºƟĚ ºƿźĭ(C. sinensis)ƪºŞĜưĩƽŚǀºƀƴƫřƹƩŚºƤţźě(Citrus aurantifolia)ƾĪƿżĪƯƺưǀƫƪƯŚƃšŚŞĩźƯŻřƮƣŹŹŚƸģ ƱŚƄƳŪƿŚŤƳŢƟźĭŹřźƣƾŝŚƿŻŹřŵŹƺƯƵŵŚƯƽƵźƄů èåƹźƳƽƵźƄů èåŵřŶƘţƽŻŚſŚƷŹŚŝ(C. aurantium)ŪƳŹŚƳƹ(C. paradise) żºǀƳƹšŚºŞĩźƯƾƿŚǀºſōƪǀºƀěƽŚƷƵŹƺěŻř ƵŶƃŶƗŚƈŤƯ ƾƿŚǀưǀƃƽŚƷħźŰƯƶŝ ƽŹřŵƾ ƴƘƯŹƺƏƶŝŶ ǀƿƺŤƿŻřŹŚě ƽƵŵŚƯįŚƷŹƺŞƳŻƶĩŵřŵ ƽŚƷħźŰƯŻř ĨƿĢ ǀƷƶŝźƳįŚƷŹƺŞƳŻƾƿƺſŻřŶƳŶƃŜƬūƚƫŚŝšřźƄůƹŚƷƵŹƺě ƽƶƿŸƜţźŧřŹŵƵŶƃŶƗŚƈŤƯ ƾƿŚƤƫřŹřźƟšŚŞǀĩźţ ƶºĩŢºſřƲƿřźĮƳŚǀŝŪƿŚŤƳŶƳŶƄƳŜƬūƾƿŚƤƫřŹřźƟšŚŞǀƧźţƹżŞſƽŚƷīźŝŹřźƟšŚŞ ǀĩźţD. citriƾĭŶƳŻƞƬŤŴƯƪůřźƯƾƿŚǀưǀƃ ƹŚºƷƵŹƺºěƽƶºƿŸºƜţźºŧřŹŵƶĩƾƿŚƤƫřŹřźƟšŚŞǀĩźţżǀƳƹšŚŞĩźƯƾƿŚǀſōƪǀƀěƽŚƷƵŹƺě ƾƿŚƿƺŝƮƿLjƗŻřŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻ ƽŚƷƵŵŚƯ ŹřźƟšŚŞǀĩźţŻřƵŵŚƠŤſřƹżŤƴſƾƿŚſŚƴƃŶƴƴĩƾ ƯƵŵŚƠŤſřŵƺųƾŝŚƿƱŚŝż ǀƯŹƺƔƴƯƶŝ ŶƳƺƃƾƯ ŶƗŚƈŤƯƱŚŝżǀƯƱŚƷŚǀĭƽƹŹƪƯŚƧšřźƄů ŶƷŵƂƿřżƟřšŚƟōİƤǀƠƬţƩźŤƴĩįŚƷƶƯŚƳźŝŹŵřŹƪƯřƺƗƲƿřŻřƵŵŚƠŤſřƾƿōŹŚĩŶƳřƺţƾƯ ƦƿĥƺƫƺǀŝƩźŤƴĩƪƯřƺƗƽƵŶƴƴĩ ŜƬū šŚŞĩźƯżŞſƽŚƷīźŝŹřźƟšŚŞ ǀĩźţƾƿŚƤƫřŹřźƟšŚŞǀĩźţDiaphorina citrišŚŞĩźƯƾƿŚǀſōƪǀƀěŪƴſƾƿ ŚƿƺŝƽŶǀƬĩƱŚĭĥřƹ Abstract Odors play an important role in the seeking behavior of wasp parasitoids and enhances their performance in biological control programs. This study was intended to determine the olfactory response of Tamarixia radiata (Waterston) to chemical cues that emanates from various stages of Asian citrus psyllid, Diaphorina citri Kuwayama, in Y-tube olfactometer in laboratory conditions (27 ± 1°C, 60 ± 10% RH, 1600 lux light intensity and constant airflow of 300 ml/min). The effect of green leaf volatiles (GLVs) and herbivore-induced plant volatiles (HIPVs) of four varieties of the rutaceous plants, Mexican lime, Citrus aurantifolia,Campbell Valencia orange, C. sinensis, Marsh grapefruit, C. Paradisi, and sour orange, C. aurantium, were also evaluated based on separately release of 30 females and 30 males of T. radiata. The results indicated that the females were significantly attracted to the volatiles emanating from D. citri nymphs and HIPVs resulted from the feeding of nymphs and adults of D. citri. The males were not responsive to the odors emanating from various stages ofD. citri, GLVs or HIPVs. It is found that the female parasitoid wasps use chemical cues emanating from the nymphs of Asian citrus psyllids and HIPVs for host seeking. Using the synthetic volatile compounds could increase the viability of the biological control agents in integrated pest management programs. Key words: olfactometer, Asian citrus psyllid, Diaphorina citri, herbivore-induced plant volatiles, green leaf volatiles, rutaceous plants ŹŵŵŹřŵŚººƷƱō ƾººƘǀŞƏƱŚƴưººƃŵƽźĮººƃƹŚĩŹŚººŤƟŹƹ źºŧřŹŵƶºĩ ƽźŧŒºƯƮºŗLjƗƾŰƐºſƶſ ƽŚƷƂƴĩ ƮƷźŝ ŶºſŹƾ ºƯƭƺºſŮƐºſŶºǀƿƺŤƿŻřŹŚěƶŝƱŚŝżǀƯŹƺƌů ƶƯŶƤƯ ƹƱŚºººƷŚǀĭŻřƵŶºººƃŶƗŚƈŤƯƾƿŚǀưǀºººƃƮºººŗLjƗ ƱřŹřƺųƵŚǀĭƾŝŚƿƱŚŝżǀƯŹŚŤƟŹŹŵ ƾưƸƯƂƤƳ ƱřŹřƺųƵŚǀĭ æèîçîçè ƁźƿŸěæèîçëéŢƟŚƿŹŵ ÑÓ źŝƾƿřŸƛƭƹŵƹƩƹřŭƺƐſŻřƵŶƃ ŶƗŚƈŤƯŹřźƟšŚŞǀĩźţƂƤƳƱřŹŚƨưƷƹƾƿřźƣƾƬŞƤƯ ƵŵƺºƫōƽŚºƷƙŚºŝ ƽŵƺŝŚºƳŜūƺƯƩŚſ ÕŚţ ÒƲǀŝ ƾƳŚƯŻ ƶºĩƾƷŚºǀĭŻřŚºƿƹƭƹŵŮƐſƶưƘƏŵƺųŻřŶƳřƺţƾƯ Żř ƾºƨƿ ƪºƣŚƳƩźºŤƴĩ ƽřźºŝ ŚºƷƂºĩƵźƄůŻřƵŵŚƠŤſř ƹŢºſřšƹŚƠŤƯŹřƺųƵŚǀĭ ƽƶƳƺĭƹƵ ŚǀĭƽƶŤƿŹřƹƶƳƺĭ řŸƫ(Halbert & Manjunath, 2004; Bové, 2006)ŵƺƃƾ Ư źºƋŚůƩŚºůŹŵŢºſřŢºǀưƷřżŗŚůŢƟōƲƿřƩźŤƴĩ ƩźŤƴĩƽřźŝƾƬưƗƹƾƯƺưƗšŹƺƇƶŝƶĩŢſř ƾƿŚƷƵřŹ (Tiwari et al., 2011)ŵƺºƃƾºƯƭŚŬƳřĬƴǀƴƿźĭƽŹŚưǀŝ ƹƾƘǀŞƏƱŚƴưƃŵŢǀƘưūƽƹŹŚƷƂĩ ƵźƄů Ōƺſšřźŧř ƹŢºƟōŢºǀƘưū ŹŵŢºƯƹŚƤƯƁźŤºƀĭŢƀƿŻ ƎǀŰƯ ŵřƺººƯŻřƽŹŚººƗšLJƺººƈŰƯŶººǀƫƺţŢººǀưƷřƲǀººƴĤưƷ ƶŤƟźºƄǀě ƽŚƷŹƺºƄĩŹŵƶºĩŢſřƵŶƃŦƗŚŝ ƾƿŚǀưǀƃ ŢºƟōƲºƿřƩźºŤƴĩŹŵĨƿĥƺƫƺǀŝƩźŤƴĩ ƪƯřƺƗŻřƵŵŚƠŤſř Ŷº ǀƿƺŤƿŻřŹŚěŹƺººŞƳŻ.ŵźº ǀĭŹřźººƣƽřƵĦ º ƿƹƶººūƺţŵŹƺººƯ ŹŵƮƸƯƪƯřƺƗƶƬưūŻřTamarixia radiata (Waterston) ƶººĩŢººſřšŚººŞĩźƯƾƿŚǀº ſōƪǀº ƀěŢº ǀƘưūƂƷŚººĩ ƦºƿĥƺƫƺǀŝƩźŤƴĩ ƽŚƷƶƯŚƳźŝŜƫŚƣ ŹŵƵŵźŤƀĭšŹƺƇƶŝ (Hoy et al., 2001; Qureshi & ŵźǀĭƾƯ ŹřźƣƵŵŚƠŤſřŵŹƺƯ ƲǀƴºſƭŚºưţŹƺºŞƳŻƲƿřƽƵŵŚƯšřźƄů Stansly, 2009) Ŷººƴƴĩƾ º ƯƶººŤƿŻřŹŚěřŹšŚººŞĩźƯƾƿŚǀº ſōƪǀº ƀěƚƫŚººŝŚƳ ƱŚºƄƳ ƽźţƂ ǀŝŮǀūźţ ƾĭŹƺěƮŬƴěƲſƶŝ ŵƺūƹƲƿřŚŝ ÔÍƽLJŚŝƮƀǀŤƿŻřŹŚěƾŤůƹ(Hoy et al., 2001)ŶƴƷŵƾ Ư (Aubert & Quilici, 1984)ŶƳŹřŵƱōƽƹŹŶƇŹŵ ƮƫŚſƱŚƷŚǀĭīźŝƾƿŚǀưǀƃƽŚƷħźŰƯƶĩ ŚŬƳōŻř ƾƿ ƽƶºƿŸƜţźºŧřźºŝƵŶºƃŶǀƫƺţ ƾƿŚºƤƫřŹřźƟŵřƺƯ (GLVs) ŵƺºųƾºƿŚǀưǀºƃƽŚºƷħźºŰƯżǀƳƹ (HIPVs)ŹřƺųƵŚ ǀĭ ƽřźºŝƱŚºŝżǀƯƾŝŚºƿŵŹƖŝŚºƴƯ ƲºƿźºţƮƸƯŻř ƾƨƿƱŚŝżǀƯ ĨƿŢǀƤƟƺƯƹŶƴƃŚŝƾƯ źĭŹŚĪƃƹŶǀƿƺŤƿŻřŹŚěšřźƄů Ʊō ƾƿŚºƳřƺţƶŝƾƐǀŰƯƎƿřźƃźŝƵƹLjƗĨ ƿĥƺƫƺǀŝƪƯŚƗ ŵŹřŵƾĮŤºººƀŝżºººǀƳŢºººƠūƹƱŚºººŝżǀƯƲŤƟŚºººƿŹŵ ƂºƤƳƂƷƹĦºěƲºƿřŹŵřŸºƫ(Dicke & Grostal, 2001) šŚºŞĩźƯƮƣŹŹŚƸģŻřƵŶƃŶƗŚƈŤƯƾƿŚǀưǀƃƽŚƷħźŰƯ ƽŚƷħźŰƯż ǀƳƹ šŚŞĩźƯ ƾƿŚǀſōƪǀƀěƶŝƵŵƺƫōƹƮƫŚſ ƱŚºŝżǀƯƪǀºƀě ƲºƿřƽŶºƃŹƞƬŤŴƯƪůřźƯ ƾƿŚǀưǀƃ ŶƃŚŝƩƹřŮƐſŵźǀĭƾƯ ŹřźƣŹřƺųƵŚǀĭƽƶƿŸƜţŵŹƺƯ ŽŚºſřźŝŚºƷƶºŰƿřŹ ƮºŗLjƗ ƽƵŶƴƷŵƪǀƨƄţ šŚŞǀƧźţ ŹŵŵřƺºƯƲºƿřƱŵźĩƪưƗ ƾ ƈƈŴţ ŜŞſšƹŚƠţ ƲǀưƷ (Dicke et al., 1999)ŵƺºƃƾ ºƯƾºƘǀŞƏƱŚƴưºƃŵŜƬū (Green Leaf Volatiles)żŞºſƽŚºƷīźºŝŹřźºƟšŚŞǀĩźţ ƹŚººƷƪººĪƫřƶººƴŝźĩÓƽŚƷŶºǀƷŶƫōźººţƂ ǀºŝƶºĩ(GLVs) ƲǀºŝƂƴºĩƮƷźºŝŹŵ ƾưƸƯƂƤƳŶƴŤƀƷŚƷƱō ƽŚƷźŤſř ƽƶºƬưůźºŧřŹŵ(Visser, 1986)ŶºƳŹřŵšřźƄůƹƵŚǀĭ ŵřƺºƯŚºƷƱō ƽżºƿŹƮºŴţŚƿƶƿŸƜţƾƏŹŵƹƱřŹřƺųƵŚ ǀĭ ŚºŝƾºƠǀƧƹ ƾưƧƑŚŰƫŻřƶĩŶƳŵźĭƾƯŶƗŚƈŤƯ ƽŹřźƟ ŶƳƺƃƾƯ ŶƗŚƈŤƯƵŚǀĭŻř ƽŵŚƗŢƫŚůŹŵƶĩ ƾƿŚƷƶŰƿ řŹ ƹ(Pare & Tumlinson, 1999)ŶƴŤºƀƷšƹŚºƠŤƯ(GLVs) (Herbivore-Induced Plant Volatiles) ƵŚǀĭ ƾƿŚƤƫř ƽŚƷƶŰƿ řŹ ƵƹLjºƗƵŚºǀĭƾƿŚƤƫř ƽŚƷƶŰƿřŹŶƳƺƃƾƯƵŶǀƯŚƳ(HIPVs) ŶºƴƳřƺţƾºƯƵŚºǀĭƽƶºƳƺĭƮºƷŵřźƟřŹŵŢƯƹŚƤƯƽŚƤƫřźŝ ƪºưƗŚƷŶǀƿƺŤƿŻřŹŚěƹŚƷźĭŹŚĪƃƽřźŝƱƺƯƺƴǀſƱřƺƴƗƶŝ ƶºĩŶƳƺºƃŹřƺųƵŚǀĭ ŢưſƶŝŚƷƱōŜƬūŜŞſƹ ŶƴƴƧ śƺºƀŰƯ ƱřŹřƺºųƵŚǀĭ ƶǀƬƗƵŚǀĭƮǀƤŤƀƯźǀƛƕŚƟŵ ƾƗƺƳ Diaphorina citri (Sabelis et al., 1999)ŵƺƃƾƯ šŚºººŞĩźƯƾƿŚǀº ºſōƪǀººƀě ŹŵƁřƾƿ ŚººƳřƺţƪºǀƫŵƶººŝKuwayama (Hem.: Liviidae) Candidatus Liberibacter americanusƽŚƷƽ źŤĩŚŝƩŚƤŤƳř ƽŹŚººưǀŝƪººƯŚƗCandidatus Liberibacter asiaticus šŚŞĩźƯżŞſƵƺ ǀƯŚƿHuanglongbing ƹ (HLB)ŚƿĬƴǀƴƿźĭ ƱřźƿřŹŵšŚŞĩźƯƽŵŚƈŤƣřƹśźŴƯŹŚǀƀŝšŚƟōŻřƾĪƿ (Halbert & Manjunath, 2004; ŵƺƃƾ ƯśƺƀŰƯƱŚƸūƹ ƶºĪƴƿřƶºŝƶºūƺţŚŝBové, 2006; Faghihi et al., 2009) šŚŞĩźƯƽƵŵřƺƳŚųƱŚƷŚ ǀĭźŨĩřƶŝšŚŞĩźƯƾƿŚǀſōƪǀƀě ŵźºţŢºƟŚŝŻřƶºƿŸƜţŚºŝƹŶºƳŻƾƯšŹŚƀų Rutaceae ƽƶƬƇŚƟŹŵ ƱŚŝżǀƯƱŚƷŚǀĭƽŚƷīźŝ ƹ ŚƷƶųŚƃ ŚƷƶƳřƺū ÑÔÎÐîçèèèƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ ƶºǀƳŚŧÒÑÐÏƹƶƤǀƣŵÎÖƶūŹŵÏÕšŚƈŤŴƯŚŝƱŚƯźĩ Źŵ.Ŷºƿŵźĭ ƾſŹźŝƮƷŚŝƢǀƠƬţŹŵƹƶƳŚĭřŶūšŹƺƇƶŝ ƪǀºƀěƚƫŚºŝšřźºƄůŶºƿŵźĭ ƽŹƹōƖưū ƽŹřŵźŝƶƳƺưƳ ŹƺºŞƳŻŜºƬūŦºƗŚŝŶƳƺºƃƾ ºƯŶƗŚƈŤƯƱŚŝżǀƯƱŚƷŚǀĭ ƶĩƾƣźƃƶǀƳŚŧ ÑÓÎÔƹƶƤǀƣŵ ÎÑƶūŹŵ ÒÕƹ ƾƫŚưƃ ƽƶººƬůźƯƲƿŶººƴģƾº ƏŶººƳŵƺŝƪǀº ƀěƲººƿřƶººŝƵŵƺººƫō ƶºƳŚŴƬĭƶºŝƩŚƤŤƳřƹƽŹƹōƖưū ŻřžěšŚŞĩźƯ ƾƿŚǀſō ƶºĩźºŤƯƾŤƳ Śſ ÒÍÒÍÒÍŵŚƘŝřƶŝ ƾƿŚƷžƠƣƪųřŵ Ǝƿřźºƃ ƶŝŚºƄƯ ƾƐƿřźƃŹŵ ŶƳŵƺŝƱŚŝżǀƯƱŚƷŚǀĭƽƹŚů šřźºƄůŻřƾƄŴŝŶƳŶƿŵźĭƽŻŚſŚƷŹ ƱŚƷŚǀĭƁŹƹźě ŵŹƺºƯ ƾºƴƬĩƒƠů ƽřźŝźĮƿŵ ƾųźŝƹŚƷƂ ƿŚƯŻō ƽřźŝ ŶƴŤƟźĭŹřźƣƵŵŚƠŤſř ŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻƾƴƬĩƪǀĪƄţ ŻřšŚºŞĩźƯƾƿŚǀºſōƪǀƀěƽƵŶƃƶŤ ƿŻřŹŚě ƽŚƷƵŹƺě ƽŹƹōƖºưūŢºƟźǀūƱŚŤſźƸƃƝřźƏřšŚŞĩźƯƽŚƷƙŚŝ źƿŻƽŚƷƩřŒſ ƶŝŲſŚěƲŤƟŚƿƂƷƹĦěƲ ƿřŻřƝŶƷƖƣřƹ 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ƽřƶºƄǀƃƁƺºěŹŵĨƿƵŚĮŤſŵƶŝƵźƄů ƹŵƶºŰƿřŹƖºŞƴƯ ƽŹřŸºĭŚūŹƺºƔƴƯƶºŝƹ ƾƬƇř ƽƹŻŚŝ ÎÔÒÐÒŵŚººƘŝřƶººŝƽřƶƳřƺŤººſřƽřƶººƄǀƃƽƶººƔƠŰƯ ƱŚƷŚǀĭƁŹƹźě ƾºĪƿżĪƯƺºưǀƫƪƯŚºƃšŚºŞĩźƯƮºƣŹŹŚºƸģŹŸŝ ƪºººŞĜưĩƽŚǀºººƀƴƫřƹƩŚºººƤţźě(Citrus ƹ(C. aurantifolia) paradise)ƁŹŚºƯšƹźººƟĚºƿźĭ(C. sinensis) šŚºŞĩźƯšŚºƤǀƤŰţƽƶƀſŒƯŻř (C. aurantium)ŪƳŹŚƳ ŵŚººƘŝřƶººŝƾƨǀŤºſLjěƽŚººƷƱřŶºƬĭƱƹŹŵƹƶºǀƸţŹƺºƄĩ ŢºſƺĜưĩŻř ƾƏƺƬŴƯ ƽƹŚůƶĩźŤƯƾŤƳ Śſ ÎÏÒÎÐ ƶºŝŢƿLJźěƹƱŚưƫōTerracultŢĩźƃŽŚƯŢǀěƾƫō ƎƿřźºƃŹŵƱŚºƷŚǀĭŶƳŶºƃƶŤºƃŚĩŶƳŵƺŝ ÎÏÎŢŞƀƳ ƽƶººūŹŵÏÔtÎƽŚººƯŵŶººƃŹĨººƣŚţřƽƵŶººƃƩźººŤƴĩ ÎÑƽŹƺƳƽƵŹƹŵƹ ÓÍtÎÍƾŞƀƳŢŝƺƏŹŽƺǀƀƬſ ŶºƴŤƟŚƿ ƁŹƹźěƾĪƿŹŚţŢƗŚſ ÎÍƹ ƾƿŚƴƃƹŹŢƗŚſ ŶƃƵŵŚƠŤſřƶƷŚƯŹŚƸģ ƽŚƷƩŚƸƳ ŻřŚƷƂ ƿŚƯŻōƾƯŚưţŹŵ šŚŞĩźƯƾƿŚǀſōƪǀƀěƾƴƬĩƪǀĪƄţ ƽŚºƷƙŚŝŻřšŚŞĩźƯ ƾƿŚǀſōƪǀƀě ƽƶǀƫƹřŢǀƘưū ƱŚŤſřŹŵƖƣřƹŢƟźǀūƱŚŤſźƸƃƝřźƏřšŚŞĩźƯƽŹŚŬţ ÑÕ źŝƾƿřŸƛƭƹŵƹƩƹřŭƺƐſŻřƵŶƃ ŶƗŚƈŤƯŹřźƟšŚŞǀĩźţƂƤƳƱřŹŚƨưƷƹƾƿřźƣƾƬŞƤƯ ƵŵŚºƯōžºƠƣƱƹŹŵƱŚƷŚǀĭ ƽƹŹŢƟōƪƀƳƹŵƁŹƹźě ŚºƷƱōšŚŰºƃźţ ƲǀºƴĤưƷƹ ƾĭŹƺºěƪůřźƯŶƳŶƿŵźĭ řƺºƷŶºƿŵźĭƶǀŞƘţ ƾƗźƟ ƽƹŻŚŝƹŵ ƽŚƸŤƳřŹŵźŤƯƾŤƳŚſ ƱŚƿźūšŶƃŚŝřƺƷĚưěĨƿŻř ƎŝřŹ ƽƶƫƺƫƹŵƎſƺţ ŶƃƾƯ ƵŶƷŚƄƯƱŚƷŚǀĭƽƹŹ żǀƳƶŤſƺěƹĨƬƀƗƪƯŚƃ ŪƴºſƱŚ ºƿźūĨºƿŚºŝƶºĩƶƤǀƣŵŹŵźŤǀƫƾ ƬǀƯ ÐÍÍŢŝŚŧ ƶƨƴƿřƪǀƫŵƶŝ ŶƳŶƃƾƯ ƒƠůźĮƿŵŢƟōźƷ ŚƿƹšŚŞƧźƯ ƽƶƔƠŰƯƹ ƱŚưƫōMerckŢĩźƃ(activated charcoal) ƱŚºƷŚǀĭ šŚºŞĩźƯƮƫŚſ ƽŚƷīźŝƶŝ ƾŝŚƿŢſŵŹƺƔƴƯƶŝ ƾºƿŚǀºſō ƪǀºƀěƽƶƿŸƜţŻřƁŹƹźěƩƺƏŹŵƵŶƃƶŤƃŚĩ źºţƲƀƯ ƽŚƷīźŝŚŝ ƶƀƿŚƤƯŹŵƵŻŚţ ƽŚƷƶƳřƺūƹ ŚƷīźŝ šŚºŞǀĩźţŻř ƽźţLJŚŝŢŞƀƳƵŚǀĭƽŚƷŢưƀƣ źƿŚſ żǀƳƹ šŚºŞĩźƯ ƾƿŚǀºſōƪǀƀě ƽƶƘſƺţƹŶƃŹƹŶƳŹřŵƶƿƺƳŚŧ (Catling, 1970)ŢºſřƶŤºƀŝřƹƵŻŚºţƽŚºƷƶƳřƺūƶŝż ǀƳ ƽŹŚºǀŝō (N:P:K)ƽŹŚºŬţŵƺºĩŚºŝƹ ƵŶƃŽźƷƱŚƷŚǀĭ ŹƺºƔƴƯƶºŝŵƺƃŌŚƤƫř ŶƿŶū ƽŚƷƶƳřƺū ŶǀƫƺţŚţŶƳŶƃƾ Ư Żřƭźºĭ ÐÒŹřźºƟšŚºŞǀĩźţŹřŶºƤƯƱŶƳŚſŹźŨĩřŶůƶŝ ŶºƿŵźĭƵŵŚƠŤºſřƵŵƺºƫōƹƮƫŚſšŚŞĩźƯƱřƺūƽŚƷīźŝ ƶºŝ ƵŻřŶºƳřƹƲºſźƔƳŻřŚƷŹřźĪţƹŚƷŹŚưǀţŹŵŚƷīźŝ ƱŶºƃŚºƷŹŻř ƽźǀĭƺºƬūŹƺƔƴƯƶŝ ŶƳŵƺŝƱŚƀĪƿŢŞƀƳ ƶŤºƄƛō ƲǀƟřŹŚºěŚŝƵŶƃƖƐƣ ĉīźŝ ĉīźŝƭŵŹřźƟšŚŞ ǀĩźţ ƱƹŹŵƂƿŚºƯŻō ƽřźºŝƵŵŚƠŤºſřŵŹƺºƯšřźºƄůŶƿŵźĭ ŵƺºŝƵŶƃŵƹŶƀƯƽŹƺţŚŝŚƷƱō ņŹŵƶĩƝŹƹŶƴěřƝƹźƓ ŚºƷƱōŲſŚěŚţŶƳŶƃ Ƶŵřŵ ƾĮƴſźĭŢƗŚſ ÏÓšŶƯƶŝ ƺųŹƺƔƴƯƶŝŶŝŚƿƂƿřżƟř ƾƿŚǀưǀƃƽŚƷħźŰƯ ƮƿLjƗƶŝ Ĩºƿ šřźºƄůƾŬƴºſŢºƀƿŻơŚºţřƎƿřźºƃŚŝƲŤƟźĭ ƾŬƴºſŢºƀƿŻơŚºţřƶºŝƂƿŚºƯŻōƭŚŬƳřŻřƪŞƣŢƗŚſ ƽƹŹ ƾºƄƧŢºƟōĢǀƷŻřƁŹƹźěƩƺƏŹŵŶƳŶƃƪƤŤƴƯ 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ƽřƺƷƪŝŚƤƯŹŵƮƫŚſ ƪºŝŚƤƯŹŵƪǀƀěƶŝƵŵƺƫōŪƳŹŚƳÒħŚěƽřƺƷƪŝŚƤƯŹŵ ƪºŝŚƤƯŹŵƪǀºƀěƶŝƵŵƺƫōƾĪƿżĪƯƺưǀƫÓħŚě ƽřƺƷ ƪǀºƀěƶºŝƵŵƺƫōƪŞĜưĩ ƽŚǀƀƴƫřƹƩŚƤţźěÔħŚě ƽřƺƷ ƵŵƺºƫōƁŹŚºƯšƹźƟĚ ƿźĭÕƹħŚě ƽřƺƷƪŝŚƤƯŹŵ ħŚěƽřƺƷƪŝŚƤƯŹŵƪǀƀěƶŝ ƪŞĜưĩƽŚǀƀƴƫřƹƩŚƤţźěƽŚƷīźŝƵřźưƷ ƵŵŚƯ ƽƵźƄů ƶŝ ŹřźƟšŚŞǀĩźţƶŝT. radiataŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻƽŹŚŤƟŹŲſŚě Źŵ ƵŵƺºƫōƁŹŚºƯšƹźºƟĚ ƿźĭƽŚƷīźŝƵřźưƷ ƵŵŚƯ ƶŝ ƱŚŝżǀƯ ƪǀºƀěƚƫŚŝšřźƄůæÓƹħŚě ƽřƺƷƪŝŚƤƯŹŵ Ƶŵƺƫō ƽƵźºƄů ÎÍ ƵřźºưƷƶŝźƳ ƽƵźƄůÎÍšŚŞĩźƯ ƾƿŚǀſō ħŚěƽřƺƷƪŝŚƤƯ ŚƷƵŵřŵƽŹŚƯōƪǀƬŰţ ƹƶƿżŬţ ƖŝźƯ ƽŚĩƱƺƯŻōŚŝŚƷƂ ƿŚƯŻōŻřƪƇŚů ƽŚƷƵŵřŵ ƶºƀƿŚƤƯSAS 3.1ŹřżºƟřƭźƳŻřƵŵŚƠŤſřŚŝƹ (Chi-Square) ŹřźĪţźƷƽŚƷƵŵřŵ ƽřźŝƱƺƯŻōŵŹƺƯźƠƇƉźƟŶƳŶƃ ÒÍÒÍ śŚººŴŤƳřƂƿŚººƯŻōźººƷƪººĩƽŚººƷƵŵřŵŶººƴƳŚưƷ ŵƺŝƩźŤƴĩŚƿŹŚưǀţƶŝŢŞƀƳƂƿŚƯŻōŵŹƺƯšřźƄů ŦŰŝƹŪƿŚŤƳ ŹřźƟšŚŞǀĩźţƶŝT. radiataŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻƽŹŚŤƟŹŲſŚě ƾƿŚƤƫřŹřźƟšŚŞǀĩźţƹżŞſƽŚƷīźŝ ŹƺºŞƳŻ ƽƵŵŚºƯšřźºƄůƶºĩŵźºĩƆŴƄƯŪƿŚŤƳ ŻřƵŶºƃŶƗŚºƈŤƯƾºƿŚǀưǀºƃƽŚºƷħźŰƯƶŝŶǀƿƺŤƿŻřŹŚě ƩŚºƤţźěƮƫŚºſƾºĪƿżĪƯƺºưǀƫƮƫŚºſŪƳŹŚºƳƽŚƷŹŚưǀţ ŪƳŹŚºƳƮƫŚſƁŹŚƯšƹźƟĚ ƿźĭƮƫŚſƪŞĜưĩ ƽŚǀƀƴƫřƹ ƩŚºƤţźěƪǀºƀěƶºŝƵŵƺºƫō ƾĪƿżĪƯƺưǀƫƪǀƀěƶŝƵŵƺƫō ƁŹŚºƯšƹźºƟĚ ºƿźĭƹ ƪ ǀƀěƶŝƵŵƺƫōƪŞĜưĩ ƽŚǀƀƴƫřƹ šŚºŞĩźƯƾƿŚǀºſōƪǀƀěƽŶƃŹƞƬŤŴƯƪůřźƯŻřƵŶƃŚƷŹ ƱŚºƷŚǀĭƾƿŚǀưǀºƃƽŚºƷħźŰƯŹƺƌůƭŶƗŚƿŹƺƌůŹŵ ƪǀºƀěƾƿŚǀưǀºƃƽŚºƷħźŰƯƂƤƳƲǀǀƘţŹƺƔƴƯ ƶŝ ŹřźºƟ šŚŞǀĩźţŚŝŜǀĩźţŹŵƹƾƿŚƸƴţƶŝ šŚŞĩźƯƾƿŚǀſō ŢưºſƶºŝŶºǀƿƺŤƿŻřŹŚěŹƺŞƳŻŜƬūŹŵ (HIPVs)ƾƷŚǀĭ ƶºŝƵŵŚºƯƽƵźºƄůÐÍƹźºƳƽƵźºƄů ÐÍ ŲºſŚěŢƟō ƽƵŹƺºěÏÍÎŢƟźĭŹřźƣƾſŹźŝŵŹƺƯźƿŻ ƽŚƷŹŚưǀţ ÏÍèƪǀºƀěźºƳ ƽƵźºƄůÏÍÏƪǀƀě ÑÒƲǀƴſ ĨƬºƀƗŻřƭźºĭƾ ºƬǀƯ ÎÍÑƪǀºƀě ƽƵŵŚºƯ ƽƵźƄů ƵřźưƷƶŝƵŶƃ ŮƃźţĨƬƀƗ ÒƪǀƀěƎſƺţƵŶƃŮƃźţ ĨƬºƀƗÓħŚě ƽřƺƷƪŝŚƤƯŹŵƵŵƺƫōŪƳŹŚƳƽŚƷīźŝ Źŵ Ƶŵƺºƫō ƾºĪƿżĪƯƺºưǀƫƽŚºƷīźŝƵřźưƷ ƶŝƵŶƃ Ůƃźţ ƵřźºưƷƶºŝƵŶºƃŮºƃźţĨƬºƀƗ ìħŚºě ƽřƺƷƪŝŚƤƯ ƽřƺºƷƪŝŚƤƯŹŵ ƵŵƺƫōƪŞĜưĩ ƽŚǀƀƴƫřƹƩŚƤţźěƽŚƷīźŝ ƽŚººƷīźººŝƵřźººưƷƶººŝƵŶººƃŮººƃźţĨƬººƀƗíħŚººě ÏÍîħŚºě ƽřƺƷƪŝŚƤƯŹŵ ƵŵƺƫōƁŹŚƯšƹźƟĚ ƿźĭ ŪƳŹŚºƳƽŚºƷīźŝƵřźưƷ ƶŝšŚŞĩźƯ ƾƿŚǀſōƪǀƀěƽƵŹƺě ƪǀºƀěƽƵŹƺººěÏÍæåħŚººěƽřƺººƷƪººŝŚƤƯŹŵƵŵƺºƫō Ƶŵƺºƫō ƾºĪƿżĪƯƺưǀƫƽŚƷīźŝƵřźưƷ ƶŝšŚŞĩźƯ ƾƿŚǀſō ƾƿŚǀºſōƪǀºƀěƽƵŹƺºě ÏÍææħŚºě ƽřƺƷƪŝŚƤƯŹŵ ÒÍ źŝƾƿřŸƛƭƹŵƹƩƹřŭƺƐſŻřƵŶƃ ŶƗŚƈŤƯŹřźƟšŚŞǀĩźţƂƤƳƱřŹŚƨưƷƹƾƿřźƣƾƬŞƤƯ ƵřźºưƷƶºŝƪǀºƀěƪºƯŚĩšřźƄůƹƁŹŚƯšƹźƟĚ ƿźĭ ƲƿřźǀŧŚţŢŰţŚƷƱōŹŚŤƟŹƹŶƳŶƄƳŜƬū ƪǀƀěƶŝƵŵƺƫō ƽŚºƷħźºŰƯŚºƯřŶƳŶºƃ ƁŹŚºƯšƹźºƟĚ ƿźĭ ƽƵŵƺƫō ƱŚƄƳ ƽŹřŵƾ ƴƘƯŲſŚěơƺƟƽŚƷŹŚưǀţƶŝƹÎBƪĪƃ ƾĪƿżĪƯƺưǀƫƽƵŵƺƫō ƽŚƷīźŝŪƳŹŚƳ ƽƵŵƺƫō ƽŚƷīźŝ ƽŚºƷīźºŝƹƪºŞĜưĩ ƽŚǀƀƴƫřƹƩŚƤţźě ƽƵŵƺƫō ƽŚƷīźŝ šŚŞĩźƯƾƿŚǀſōƪǀƀěƎſƺţƵŶƃŮƃźţĨƬƀƗ ƾƿŚǀưǀƃ ƩŚºƤţźěƾºĪƿżĪƯƺºưǀƫŪƳŹŚƳ ƽƵŵƺƫōƱŚƷŚǀĭƵřźưƷƶŝ šřźºƄůƽřźºŝƁŹŚºƯšƹźƟĚ ƿźĭƹƪŞĜưĩ ƽŚǀƀƴƫřƹ ŶƴŤƃřŶƳƾŤǀŝřŸūŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻƽƵŵŚƯ źǀŧŐºţŢºŰţŶºǀƿƺŤƿŻřŹŚºěŹƺŞƳŻźƳšřźƄůŹŚŤƟŹ ƽŚººƷŹŚưǀţŻřƭřŶººĩĢ ǀºƷŻřƵŶººƃƾ ºƃŚƳŹřźººƟšŚººŞǀĩźţ źºƳšřźºƄůŲſŚěÎ$ƪĪƃŢƟźĮƳŹřźƣŚƷƶŰƿ řŹ ŵƺºŝƵŵŚºƯšřźºƄůŹŚºŤƟŹƶŝŚºƄƯƮƷ ŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻ ŚºŝƂƷƹĦěŻř ŢưƀƣƲƿřŻřƵŶƯōŢſŵƶŝŪƿŚŤƳŶƳŵřŶƳ ŹƺºŞƳŻŲºſŚěƶºĩMann et al. (2010)ƂƷƹĦºěŪƿŚºŤƳ ƶºŝƵŵƺºƫōƹƮƫŚºſƱŚºƷŚǀĭƶŝřŹT. ƹMurraya radiataŶǀƿƺŤƿŻřŹŚě paniculataƲǀưºſŚūƩŚƤţźěƪƯŚƃƪǀƀě ŵŹřŵŢƤŝŚƐƯŶƳŵźĩƾſŹźŝƲǀƬƯŚƷƩŚƤţźě ƺºưǀƫŪƳŹŚºƳ ƽƵŵƺºƫōƱŚºƷŚǀĭƵřźºưƷƶŝƪ ǀƀě ƽŚƷƵŹƺě ƽŚºƷħźŰƯ ƶŝT. ƁŹŚƯšƹźƟĚ ƿźĭƹƪŞĜưĩƽŚǀƀƴƫřƹƩŚƤţźěƾĪƿżĪƯ ƞºƬŤŴƯƪºůřźƯ ƾºƿŚǀưǀºƃƽŚƷħźŰƯƶĩ ƾŤƣƹ ƁŹŚƯšƹźƟĚ ƿźĭƹƪŞĜưĩƽŚǀƀƴƫřƹƩŚƤţźěƾĪƿżĪƯ ƺưǀƫŪƳŹŚƳƽƵŵƺƫō ƱŚƷŚǀĭ ƵřźưƷƶŝƪǀƀěƪƯŚĩšřźƄů ƎºſƺţƵŶºƃŮºƃźţĨƬºƀƗƾƿŚǀưǀƃƽŚƷħźŰƯ żǀƳƹ Źřźƣ ơƺƟƽƵŵƺƫōƱŚƷŚǀĭƵřźưƷƶŝšŚŞĩźƯ ƾƿŚǀſōƪǀƀě ŶƳŶƄƳŚƷƱōŜƬūŶ ǀƿƺŤƿŻřŹŚěźƳšřźƄůƹŢƟźĮƳ ŪƿŚºŤƳŚºŝơƺºƟŢưºƀƣŻřƵŶºƯōŢºſŵƶºŝŪƿŚºŤƳ ƲǀƤƤŰƯƲƿřŵŹřŵŢƤŝŚƐƯMann et al. (2010)šŚƤǀƤŰţ ŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻ ƽƵŵŚƯšřźƄů ƶƧ ŶƳŵƺưƳƁŹřżĭ żǀƳ ƪǀºƀěƽƵŹƺºěƾƿŚǀưǀºƃƽŚºƷħźŰƯŜƬū T. radiata ƽŚºƷħźºŰƯƵřźºưƷƶºŝżǀƳƹ ƾƿŚƸƴţƶŝ šŚŞƧźƯƾƿŚǀſō ƱŚºƄƳŪƿŚºŤƳƲºƿřŶƳƺºƃƾ ºƯƱŚºŝżǀƯƱŚƷŚǀĭƾƿŚǀưǀƃ ŹƺºŞƳŻ ƽƵŵŚºƯƹźºƳšřźºƄů ƾƿŚƿƺŝŲſŚěƶĩ ŶƷŵƾƯ ƹŢſřšƹŚƠŤƯŹřźƟšŚŞǀĩźţƶŝT. radiataŶǀƿƺŤƿŻřŹŚě ƽŚºƷƵŶºƳźǀĭƱŚưŤųŚºſ ƾƀƴū ƾƬĪƃƹŵŚŝ šƹŚƠţƲƿř Ūƴºě(Onagbola et al., 2009)ŵŹřŵƍŚŞţŹřŚƷƱō ĨųŚƃ ŸºƠƴƯƱƹŶŝ ƽŶǀŗƺĪƿźţ ƽƺƯƹŵƪƯŚƃƾƀů ƽƺƯƕƺƳ ƽŸºƠƴƯŶºƴģ ƽŶǀŗƺĪƿźţ ƽƺƯĨƿAST-2ƹ AST-1 Ĩƿƹ (MPS)ƽŸƠƴƯŶƴģ ƽŶǀŗƺĩLjě ƽƺƯĨƿ(MST) ĨųŚºƃƽƹŹbasiconic capitate pegŸºƠƴƯƱƹŶŝƽƺƯ ŢſřƵŶƃƾƿŚſŚƴƃŶǀƿƺŤƿŻřŹŚěƲƿřƽƵŵŚƯƹźƳšřźƄů radiataŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻƽŹŚŤƟŹŲſŚě ŹŵšŚºŞĩźƯƾƿŚǀſōƪǀƀěƽŶƃŹƞƬŤŴƯƪůřźƯƾƿŚǀưǀƃ ƱŚŝżǀƯƱŚƷŚǀĭŹƺƌůƭŶƗŚƿŹƺƌů Ŷºƿŵźĭ ƾŝŚƿŻŹř ƾƿŚƸƴţƶŝ šŚŞĩźƯ ƾƿŚǀſōƪǀƀěƽŶƃŹ ŹƺººŞƳŻƚƫŚººŝƽƵŵŚººƯšřźººƄůƶººƧŵřŵƱŚººƄƳŪƿŚººŤƳ ƪǀƀěƽŚƷƵŹƺěŜƬūƎƤƟ ƽŹřŵƾ ƴƘƯŹƺƏƶŝŶ ǀƿƺŤƿŻřŹŚě ƽŚƷħźŰƯ źǀŧŐţŢŰţŚƷƱōŹŚŤƟŹƹ ÏAƪĪƃŶƳŶƃ ƚƫŚºŝ ƽƵŵŚºƯšřźºƄůƪǀƀěƚƫŚŝźƳšřźƄůƾƿŚǀưǀƃ ƾƿŚǀºſōƪǀºƀěƎºſƺţƵŶºƃŮºƃźţĨƬƀƗżǀƳƹƪǀƀě ŹŚººŤƟŹŶƳŶººƄƳŚººƷƱōŜººƬūƹŢººƟźĮƳŹřźººƣšŚººŞĩźƯ ƽŚºƷħźºŰƯźǀŧŐºţŢŰţŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻźƳšřźƄů ƾºƿŚǀºſōƪǀƀěƽŚƷƵŹƺě ƽŚƷŹŚưǀţŻřƭřŶĩĢ ǀƷ ƾƿŚǀưǀƃ šřźºƄůƪǀºƀěƎºſƺţƵŶºƃŮºƃźţĨƬƀƗ šŚŞĩźƯ ŢºƟźĮƳŹřźƣƪǀƀěƚƫŚŝźƳšřźƄůƹƪǀƀěƚƫŚŝ ƽƵŵŚƯ ƪǀºƀě ƾºƿŚǀưǀºƃƽŚºƷħźŰƯƶĩ ƯŚĮƴƷÏBƪĪƃ ƾ ƱŚºƷŚǀĭ ƾºƿŚǀưǀºƃƽŚƷħźŰƯƵřźưƷ šŚŞĩźƯƾƿŚǀſō ƶŝ šřźºƄůÐ ƪĪºƃ ŶºƴŤƟźĭŹřźºƣ ƾŝŚƿŻŹřŵŹƺƯƵŵƺƫō ŜººƬūƽŹřŵƾ º ƴƘƯŹƺººƏƶº ŝŶº ǀƿƺŤƿŻřŹŚěŹƺººŞƳŻƽƵŵŚººƯ ƽŚºƷīźŝŻřƭřŶĩźƷƵřźưƷƶŝƪ ǀƀě ƽŚƷƵŹƺě ƽŚƷŹŚưǀţ ƽŚƷīźŝ ƾĪƿżĪƯƺưǀƫƽƵŵƺƫō ƽŚƷīźŝŪƳŹŚƳ ƽƵŵƺƫō ƽƵŵƺºƫōƽŚºƷīźºŝƹƪºŞĜưĩƽŚǀºƀƴƫřƹƩŚƤţźě ƽƵŵƺƫō ÒÎÎÐîçèèèƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ šŚºŞǀĩźţƹżŞſƽŚƷīźŝ ƾƿŚǀưǀƃƽŚƷħźŰƯ ƶŝTamarixia radiata(B)źƳƹ(A)ƵŵŚƯŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻŲſŚě æƪĪƃ N èåšŚŞĩźƯŻřƮƣŹŹŚƸģƾƿŚƤƫřŹřźƟ Fig. 1. Response of female (A) and male (B) Tamarixia radiata to green leaf chemical cues and herbivore-induced plant volatiles (HIPVs) from four varieties of rutaceous plants (N = 30). ƲºƿřƂºƤƳ(Onagbola et al., 2009)ŶƳŹřŵŵƺųĨųŚƃ ƽŸºƠƴƯŶƴģƽŶǀŗƺĩLjěƽŚƷƺƯ (Onagbola et al., 2009) źºƋŚůƂƷƹĦºěŪƿŚºŤƳOnagbola & Fadamiro, 2008) Źŵ(Onagbola et al., 2009)ŶƳŹřŵƂƤƳƱŚŝżǀƯƾƿŚǀưǀƃ ƵŶƃŶǀƫƺţƾƀƴūƽŚƷƱƺƯƹźƟƆ ǀŴƄţƹƾƿŚſŚƴƃŚƷƺƯ (Pettersson et al., 2001; ŢºſřƵŵŚºƯšřźºƄůƎºſƺţ T. radiataŹƺŞƳŻźƳšřźƄůƶĩŢſřƲƿřƽƵŶƴƷŵ ƱŚƄƳ ƪºŰƯƲŤƟŚƿƽřźŝƱŚŝżǀƯƶŝƶŤƀŝřƹ ƾƿŚǀưǀƃšŚƗLjƏřŻř ŻřźºţƂǀŝƽŹřŵ ƾƴƘƯŹƺƏ ƶŝƵŵŚƯšřźƄůĨųŚƃƽƹŹ šŚºŞǀĩźţƆǀŴºƄţ Źŵ ŚºƷƺƯƲºƿřŢºſřźºƳšřźƄů ŵřŶƘţT. radiataźƳƽŚƷŹƺŞƳŻƵŵŚƯƽŚƷŹƺŞƳŻŚŝƶƀƿŚƤƯ Źŵ ƲºƸěƹīŹżºŝƽŸºƠƴƯŶƴģ ƽŶǀŗƺĪƿźţ ƽƺƯ ƽŵŚƿŻ ÒÏ źŝƾƿřŸƛƭƹŵƹƩƹřŭƺƐſŻřƵŶƃ ŶƗŚƈŤƯŹřźƟšŚŞǀĩźţƂƤƳƱřŹŚƨưƷƹƾƿřźƣƾƬŞƤƯ clean air D. citri nymphs Treatment * NC=3 Honeydew ÷2 = 0.04; P = 0.85 NC=3 Female D. citri NC=2 Male D. citri NC=3 60 40 ÷2 = 4.84; P = 0.03 ÷2 = 1.28; P = 0.25 ÷2 = 0.33; P = 0.56 20 0 20 40 60 80 A: Response (%) of Females ƪǀºƀěƞºƬŤŴƯƪºůřźƯƾƿŚǀưǀºƃƽŚƷħźŰƯ ƶŝTamarixia radiata(B)źƳƹ(A)ƵŵŚƯŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻŲſŚě ÏƪĪƃ ƖŝźƯƽŚĩƱƺƯŻōŽŚſřźŝŹřŵƾ ƴƘƯƝLjŤųřŵƺūƹƽƵŶƴƷŵ ƱŚƄƳ *N èåDiaphorina citrišŚŞĩźƯƾƿŚǀſō (P < 0.05)ŶƃŚŝƾ Ư Fig. 2. Response of female (A) and male (B) Tamarixia radiata to chemical cues that emanating from various stages of Asian citrus psyllid, Diaphorina citri (N = 30). * Indicates significant difference by Chi-Square test (P < 0.05). řŹšŚºƗLjƏřŻřƽźţŵŚºưŤƗřƪŝŚƣƖŞƴƯŶƳƺƃƾ ƯŶƗŚƈŤƯ ƱŚºƄƳŪƿŚŤƳƲƿř ƲǀƴĤưƷŶƴƴĩƾưƳƵŵŚƠŤſř ƽźǀĭŢƠū ƾºƿŚǀºſō ƪǀƀěƽŚƷƵŹƺěƶŝŶ ǀƿƺŤƿŻřŹŚě ƽŚƷŹƺŞƳŻŜƬū ƖºŞƴƯ ƲǀºƫƹřšŚŞĩźƯ ƾƿŚǀſō ƪǀƀěƽŚƷƵŹƺě ƶƧƽŹƺƏƶŝ Ŷººƴƴĩƾ ººƯƮƷřźººƟŚƷŶººǀƿƺŤƿŻřŹŚěƾŝŚººƿƱŚººŝż ǀƯƽřźººŝ ƽřźºŝŶºƃŚŝƾºƬǀƫŵƲºƿřŶƿŚºƃƹ(Vet & Dicke, 1992) ŻřŢºſřƲºĪưƯ ƾƿŚǀưǀƃšŚƗLjƏř ƩŚůƲ ƿřŚŝ šŚŞĩźƯ ŻřŚº ƿƹƱŚººƷŚǀĭƶŤººſƺěĨƬººƀƗƱŚººŝżǀƯšřŶº ǀƫƺţ ƩƹřƽƶºƬůźƯŹŵ ŹƺºŞƳŻƲƿřƽƵŵŚƯšřźƄůƶĩŶƷŵƾƯ ŶƴƴĩƾƯƵŵŚƠŤſřƱŚŝżǀƯ ƾƿŚƿƺŝƮƿLjƗŻřŵƺųƾŝŚƿƱŚŝżǀƯ ƾƬĩŹƺºƏƶºŝŶƴŤºƀƷŶºǀƿƺŤƿŻřŹŚºěŹƺŞƳŻƽƵŶƴƴĩśŸū ƱřŹřƺºųƵŚ ºǀĭŵƺºųŻřƮǀƤŤºƀƯŹƺºƏƶºŝƶƧƾƿŚƷƥźŰƯ ÒÐÎÐîçèèèƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ ƽŶºƃŹƞºƬŤŴƯƪºůřźƯƾƿŚǀưǀºƃƽŚƷħźŰƯ ƶŝTamarixia radiata(B)źƳƹ(A)ƵŵŚƯŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻŲſŚě ÐƪĪƃ ƽƵŶƴƷŵ ƱŚƄƳ *N èåƱŚŝżǀƯƱŚƷŚǀĭƾƿŚǀưǀƃƽŚƷħźŰƯƵřźưƷ Diaphorina ƶŝ citrišŚŞĩźƯƾƿŚǀſōƪǀƀě (P < 0.05)ŶƃŚŝƾ ƯƖŝźƯƽŚĩƱƺƯŻōŽŚſřźŝŹřŵƾ ƴƘƯƝLjŤųřŵƺūƹ Fig. 3. Response of female (A) and male (B) Tamarixia radiata to chemical cues from various stages of Asian citrus psyllid, Diaphorina citri, together with host plant chemical cues (N = 30).* Indicates significant difference by Chi-Square test (P < 0.05) ÒÑ źŝƾƿřŸƛƭƹŵƹƩƹřŭƺƐſŻřƵŶƃ ŶƗŚƈŤƯŹřźƟšŚŞǀĩźţƂƤƳƱřŹŚƨưƷƹƾƿřźƣƾƬŞƤƯ ƽŚºƷƩŚƴĮ ǀºſŻřƵŵŚƠŤºſřŢºſř ƾºƿŚºƤƫřŹřźƟšŚŞǀƧźţ ŢƠūƲŤƟŚƿƽřźŝƾƫƺưƘƯƹƩƹřŶŤƯƁƹŹ ƦƿƾƿŚǀưǀƃ ŶƳźǀĭŌŚƄƴƯŚƷƱō ƲǀŝƂƴĩƮƷźŝ (Vinson, 1998, 1991; Vet & Dicke, 1992; Steiner et al., 2007) ŢººſřŶººǀƿƺŤƿŻřŹŚººěƽŚººƷŹƺŞƳŻƽƶƬǀººſƹƶº ŝƱŚººŝżǀƯƹ ŹřźºƟšŚŞǀƧźţƶĩŶƃƆŴƄƯźƋŚůƂƷƹĦěŹŵ ƾºƿŚǀưǀºƃšŚºŞǀƧźţŮºƃźţƹ ƽźºǀĭŢºƠūƾĭŶƴĩřźě ƽƵźƄůŚƿƵŹƺěƽ ƶƿŸƜţŚƯřŶƳƺƃ ƾưƳŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻ ƾƬĩŹƺººƏƶººŝ(Vinson, 1991; Ayasse et al., 2001) ŹŵƾƗƺººƴŤƯŹŚǀºƀŝƹƵŶºǀĤǀěƽŚºƷŹŚŤƟŹƱŚºǀƿŚººƄƛƩŚºŝ (Ayasse et al., 2001)ŶºƴƷŵƾºƯƱŚƄƳŵƺųŻřƾūŹŚų ƽřƵĦºƿƹ ƾƿŚƿƺŝ ƽŚƷƺƯ ƽƶƬǀſƹƶŝřŹŚƷƱƺƯƹźƟŚƷŹƺŞƳŻ ŶºƴƴĩƾºƯŢºƟŚºƿŹŵ ŵŹřŵŹřźºƣƱŚƄƿŚƷĨųŚƃ ƽƹŹƶĩ ƹŚƷƱƺƯƹźƟ (Isidoro et al., 1996; Ayasse et al., 2001) ƽŚºƷƺƯƽƶƬǀºſƹƶºŝŢºſřƲĪưƯƱŚƷŚǀĭŹřźƟšŚŞǀƧźţ ŶƳƺºººƃƾƿŚºººſŚƴƃT. radiataŹƺºººŞƳŻĨųŚºººƃƽƹŹ šŚººŞǀĩźţŢº ǀưĩƹŢº ǀƷŚƯ(Onagbola et al., 2009) ƵŚºǀĭƾºƗřŹŻƮºƣŹƹƶºƳƺĭŽŚºſřźŝ ƾƿŚƤƫř ƽŚƷƶŰƿřŹ ƶºĩƵŚºǀĭŻř ƾƿŚƷƶŤ ƿŹřƹśŚŴŤƳřŶƃŚŝšƹŚƠŤƯŶƳřƺţƾ Ư ŢǀººƇŚųƱřŹřƺººųƵŚººǀĭŻřƾººƃŚƳŜǀººſōŻřžººě ŶƴƃŚŝƶŤƃřŵƾƘǀŞƏƱŚƴưƃŵƽřźŝƽźţƂ ǀŝƾĭŶƴƴĩŜƬū ƾºƿōŹŚĩŶƳřƺţƾ ƯŢƸūƲƿřŹŵƱŚƷŚǀĭ ƾĪǀŤƳĥŭLjƇřƹ šŚºƟōƩźŤƴĩ ƽřźŝřŹ ƦƿĥƺƫƺǀŝƩźŤƴĩƪƯřƺƗŻřƵŵŚƠŤſř ŻřƵŵŚƠŤſřƹżŤƴſƾƿŚſŚƴƃŶƷŵ ƂƿřżƟřźţƂǀŝ ƶģźƷ ƽƵŶ ƿřŶƳřƺţƾ ƯŚƷŶǀƿƺŤƿŻřŹŚě ƽƵŶƴƴĩŜƬūŹřźƟšŚŞ ǀĩźţ ƲƿřŶƃŚŝƽŻŹƹŚƄĩšLJƺƈŰƯŻřŢƓŚƠů ƽřźŝ ƾŞƫŚū ƹƱŶºƃƵŶºƴĩřźěƭŶºƗŦºƗŚŝšŚºƟōƩźºŤƴĩŹŵŵźŞƷřŹ ĨºƿĥƺƫƺǀŝƩźºŤƴĩƪºƯřƺƗƽźĭƺŬŤƀūƱŚƯŻŹŵƂƷŚĩ ƂƿřżºƟřƶºĩŵƺºƃƾ ƯŵƺųŜſŚƴƯ ƽƶưƘƏƲŤƟŚƿƽřźŝ ŶºƷřƺųƵřźºưƷƶºŝřŹĨºƿĥƺƫƺǀŝƩźºŤƴĩƪºƯřƺƗ ƾƿōŹŚĩ ƁŹƹźººěľřźººŨĩřƶººƨƴƿřƶººŝƶººūƺţŚŝƾƿƺººſŻřŢººƃřŵ řŸºƫŵźºǀĭƾưƳšŹƺƇ ƾƬƇřƱŚŝżǀƯƽƹŹŚƷŶǀƿƺŤƿŻřŹŚě ŚƷŶǀƿƺŤƿŻřŹŚěƵƺŞƳřƁŹƹźěŹŵ ƾſŚſřšLjĪƄƯŻř ƾĪƿ ŚƷƱō ƾƿōŹŚĩŚƳĨƿĥƺƫƺǀŝƩźŤƴĩŹŵŚƷƱōŻřƵŵŚƠŤſř ƽřźŝ ŶºƃŚŝƾºƯ ƽŻŚºſŚƷŹŻřžěƶưƘƏƢƟƺƯ ƽƺŬŤƀūŹŵ ŻřŶºǀƿƺŤƿŻřŹŚěƽŚºƷŹƺŞƳŻŻř ƽŹŚǀºƀŝ ƶºĪƴƿřƶºŝƶūƺţŚŝ ƵŶºƿŵŜǀºſōƱŚºƷŚǀĭŚºƿƹƮƫŚſƱŚƷŚǀĭŻřƵŶƃŶƗŚƈŤƯ ŜºƬūŦºƗŚŝƾƿŚºƸƴţƶºŝšŚŞĩźƯƾƿŚǀſōƪǀƀěƎſƺţ ŜƬūƹŢŞŨƯƂƴĩřƹŜūƺƯƱŚŝżǀƯƱŚƷŚǀĭƽƹŹƪƯŚĩ ƾºƏƵŚºǀĭƾƿŚºƤƫřƽŚƷƶŰƿřŹ ŵŵźĭƾƯŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻ ƶºĩƽřƵŶºƴƴĩŚƤƫřŵřƺºƯƢƿŹżţźŧřŹŵƹƩŚƘƟŶƴƿōźƟĨƿ ŹřƺºųƵŚºǀĭšřźºƄůƽżƿŹƮŴţƖƿŚƯŚƿơřżŝŹŵľLJƺưƘƯ ƍŚºŞţŹřŚƷƱōŶǀƫƺţƹŶƳƺƃ ƾƯŶǀƫƺţƵŚǀĭŹŵŵŹřŵŵƺūƹ ŵŹřŶºƳƵŚºǀĭƾĪǀƳŚĪƯƹƾĪƿżǀƟƽŚƷŜǀſōŚŝƾưǀƤŤƀƯ źºŧřŹŵƶºĩƽŹřźƟŵřƺƯřŸƫ(Pare & Tumlinson, 1999) ƱŚºƷŚǀĭīźŝŻřƵŹƺěŚƿƪƯŚĩƽƵźƄůƱŚƯŻƮƷƽƶƿŸƜţ ŚºŝƾºƠǀĩ ƹƾºưĩƑŚºŰƫŻřŵŵźºĭƾºƯŶƗŚƈŤƯƱŚŝżǀƯ ŚƿƪƯŚĩƽƵźƄůƽ ƶƿŸƜţƱŶƃƖƐƣŻřžěƶĩƽřƶŰƿřŹ źºƔƳŻřƹŶƃŚŝšƹŚƠŤƯŶƳřƺţƾƯŵŵźĭ ƾƯŶƗŚƈŤƯƵŹƺě ŮƐºſŹŵƹŶºŝŚƿƾºƯƂƷŚºĩŚºƷƱōŢºǀưĩƮºƷƾưĩ ƭŶºƗźŝƾƬǀƫŵŶƳřƺţƾƯƲƿř ŶƳƺƃƾƯŶƗŚƈŤƯƽźţ ƲǀƿŚě ƾƳŚºƷŚǀĭƶŝŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻƲƿřƽƵŵŚƯšřźƄůŲſŚě ŚƷƱōƽƹŹƶƔŰƫƱŚưƷŹŵƹŶƳŵƺŝƵŶƃƵŵƺƫōƪŞƣŻřƶĩ ƶºŝƶºūƺţŚŝźºĮƿŵƾƿƺſŻřŶƃŚŝŵřŵƾưƳűŹƽř ƶƿŸƜţ ƾƿŚǀºſōƪǀƀěƚƫŚŝšřźƄůƾƿŚǀưǀƃƽŚƷħźŰƯƶƨƴƿř řŸºƫŶƴŤƃřŶƳřŹŶǀƿƺŤƿŻřŹŚěŹƺŞƳŻŜƬūšŹŶƣƾƿŚƸƴţƶŝ ƾƿŚºƤƫřŹřźºƟšŚºŞǀƧźţƂºƤƳƱōƾƬºƇřƪºƿLJŵŻřƾƨƿ żºǀƳŚƷŶºǀƿƺŤƿŻřŹŚěźƿŚºſŵŹƺƯŹŵƶĩŶƃŚŝƾƯ (HIPVs) ƽƶƿŸƜţ ƾƟźƏŻř(Vinson, 1991)ŢſřƵŶƿŵźĭƵŶƷŚƄƯ ƽŚºƷīźºŝƽƹŹšŚºŞĩźƯƾƿŚǀºſōƪǀƀěƽŚƷƵŹƺěƭŏƺţ ƶºŝŢŞºƀƳŶºǀƿƺŤƿŻřŹŚěƽŚºƷŹƺŞƳŻƽřźŝƱŚŝżǀƯƱŚƷŚǀĭ ƹŵƺºŝƽźţƂǀŝŢǀŝřŸūƽřŹřŵƾƿŚƸƴţ ƶŝƪǀƀěƽŚƷƵŹƺě ŹřźººƟšŚººŞǀƧźţƾŤººƀƿĥźƴǀſźººŧřƽƵŶººƴƷŵƱŚººƄƳƲººƿř ƂºƤƳƹŹřƺųƵŚǀĭƹƵŚǀĭƲǀŝƂƴĩ ƮƷźŝŻřƵŶƃŶƗŚƈŤƯ ÒÒÎÐîçèèèƱřźƿřƾſŚƴƃƵźƄůƲưŬƳřƽ ƶƯŚƳ ƾƳŚƿŚƃĨưĩ ƦƿĥƺƫƺǀŝƩźŤƴĩ ƽŚƷƶƯŚƳźŝŹŵ ŶǀƿƺŤƿŻřŹŚě ƲŤƟŚºƿżºǀƳƹƱŚºŝżǀƯƵŚĮŤƀƿŻƲŤƟŚƿƽřźŝ ƾƿŚƿƺŝƮƿLjƗ źºţƂ ǀºŝƶģźƷħŹŵƶŝĨưĩźŝƵƹLjƗŶ ǀƿƺŤƿŻřŹŚěŹƺŞƳŻ ŹŵƵŶºƳŻŹř ƽŚƷŹŚĪƷřŹŻř ƾĪƿƱŚŝżǀƯŚŝƎŞţźƯ ƾƿŚǀưǀƃ šŚƗLjƏřŶƴĩƾƯ ŚƠƿřšŚŞĩźƯ ƾƿŚǀſō ƪǀƀěƩźŤƴĩ ƽřźŝ ƽƵŶºƴƴĩŜºƬūšŚºŞǀĩźţ ƾƿŚºſŚƴƃ ƽƶƴǀƯŻŹŵ ƾƬǀưĪţ Ʋǀºŝ ƾŰƐºſƶºſƂƴĩƮƷźŝĨ ƿŹŵƾĪƿĥƺƫƺĩřƎŝřƹŹ ƂºƿřżºƟřƶºŝźŬƴƯŶƳřƺţƾ ƯŶǀƿƺŤƿŻřŹŚěŹřƺųƵŚ ǀĭƵŚǀĭ ŵŵźĭƦƿĥƺƫƺǀŝƩźŤƴĩƽŚƷƶƯŚƳźŝŹŵŶ ǀƿƺŤƿŻřŹŚěƾƿōŹŚĩ ƾƏźƃřŸƫ(Vinson, 1981)Ŷƴƴĩƾ ƯƵŵŚƠŤſřƱŚŝżǀƯŵƺų ƮƿLjƗ ƽƶƬǀſƹƶŝƁŹƹźěƪůřźƯŹŵŚƷŶ ǀƿƺŤƿŻřŹŚěƱŵźĩ ŢſřƦƿĥƺƫƺǀŝƩźŤƴĩƽŚƷƶƯŚƳźŝ ƾƿLJŚºŝƪǀºƀƳŚŤěŻřT. radiataŶºǀƿƺŤƿŻřŹŚºěŹƺŞƳŻ řŸƫ ŢſřŹřŵŹƺųźŝšŚŞĩźƯ ƾƿŚǀſōƪǀƀěƩźŤƴĩŢƸū Ʋƿř ƽƵŶƴƴĩŜƬūšŚŞ ǀĩźţ ƾŝŚƿŻŹřƹšLjƿŚưţ ƾƿŚſŚƴƃ ƖŝŚƴƯ Aubert, B. & Quilici, S. 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