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108Soybean and Healthresistances (spinosad and pyrethroid) in H. armigera on raw soybean flour and roastedsoybean flour diets.Given that KSTI is remarkably resistant to thermal denaturation and that denaturation isreadily reversible at lower temperatures, it is likely that failure to detect resistance on a dietmade from boiled flour is due inadequate denaturation of KSTI and/or other trypsininhibitors. We observed that a soybean flour/ water mix boiled at 40 o C, which is far belowthe temperature required to permanently denature KSTI (Kunitz, 1948). It is also possiblethat boiling the soybean flour in water solubilises the tyrpsin inhibitors.KSTI and other trypsin inhibitors are known to have synergistic effects with deltaendotoxins of Bacillus thuringiensis against H. armigera and other insects (Manduca sextai,Heliothis virescens, Trichoplusia ni, Helicoverpa zea) and Leptinotarsa decemlineata (Coloradopotato beetle) ( Macintosh et al., 1990; Shao et al., 1998; Zhang et al., 2000, Zhu et al., 2007;Christtellier et al., 1992; Hubert et al., 2008). It was suggested that the mechanism ofsynergism might be prevention of activation of the toxin by gut serine proteases due to KSTIaction (Zhu et al., 2007). In Australian H. armigera, differences in detection of resistance to Bttoxin between raw and dry roasted soybean flour were attributed to synergism by dietaryKSTI in the raw flour diet. It was also suggested that greater susceptibility of cottonpopulations of H. armigera to Cry1Ac, compared to Helicoverpa punctigera, might have beendue to dietary STI inhibition of detoxificative enzymes in H. armigera (Bird & Akhurst, 2007).The present data suggest that KSTI and/or other trypsin inhibitors in soybean flour mayalso act as spinosad and pyrethroid synergists against non-specific esterase mediatedresistance in H. armigera.Non-specific esterases, derived from cell adhesion proteins, are serine hydrolases with anability to sequester, pyrethroids, spinosad and Bt toxins in Australian H. armigera (Gunninget al., 1996b; Gunning et al., 2007; Gunning & Balfe, 2002, Gunning et al., 2005;). Theseresistances can be synergised by esterase inhibitors (Gunning et al., 1999, Young et al., 2006,Gunning et al., 2007). It is possible that these non-specific esterases, found in the H. armigeramidgut, are inhibited by dietary protease inhibitors such as KSTI, found in a raw orincompletely denatured soybean flour diet.This study has indicated that rearing and bioassay of H. armigera on artificial diets that havebeen inadequately treated and thus give chronic exposure to abnormally high levels ofactive soybean protease inhibitors may be unsuitable for detection of metabolic-basedresistance to spinosad or pyrethroids (bifenthrin).5. AcknowledgementsThe authors would like to thank, Industry and Investment NSW for supporting this work.Rothamsted Research receives grant-aided support from the Biotechnology and BiologicalSciences Research Council of the United Kingdom.6. ReferencesAnon, (1970). Standard method of detection of insecticide resistance in Heliothis zea (Boddie)and Heliothis virescens (F.). Bulletin of the Entomological Society of America 16: 147 –153.Broadway, R. & Duffey, S. (1986). Plant proteinase inhibitors: mechanism of action and effecton larval growth and digestive physiology of larval Heliothis zea and Spodopteraexiqua, Journal of Insect Physiology 32: 827 - 833.
Effects of Dietary Soybean Trypsin Inhibitorson Detection of Resistance toPyrethroid and Spinosad Insecticides in Helicoverpa armigera Hübner (Lepidoptera: Noctuidae) 109Christtellier, J. Laing, W., Markwick, N. & Burgess, E. (1992). Midgut protease activities in 12phytophagous Lepidopteran larvae: dietary and protease inhibitor interactions,Insect Biochemistry and Molecular Biology 22: 735 – 746.Finney, D (1971). Probit Analysis, Cambridge University Press, London.Gunning, R., Easton, C., Balfe, M. & Ferris, I. (1991). Pyrethroid resistance mechanisms inAustralian Helicoverpa armigera. Pesticide. Science 33: 473 –490.Gunning, R., Moores, G. & Devonshire, A. (1996a). Esterases and fenvalerate resistance inAustralian Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae). PesticideBiochemistry and Physiology. 54: 12 – 23.Gunning, R., Moores G. & Devonshire, A. (1996b). Insensitive acetylcholine esterase andresistance to thiodicarb in Australian Helicoverpa armigera (Hübner) (Lepidoptera:Noctuidae) Pesticide Biochemistry and Physiology 55: 21 – 28.Gunning, R., Moores, G. & Devonshire, A. (1988). Inhibition of pyrethroid resistance relatedesterases by piperonyl butoxide in Australian Helicoverpa armigera (Lepidoptera:Noctuidae) and Aphis gossypii (Hemiptera: Aphididae). In: Glynne-Jones, D. (Ed.),Piperonyl Butoxide: The Insecticide Synergist, Academic Press, 215 - 226.Gunning, R., Moores, G. & Devonshire, A. (1999). Esterase inhibitors synergise the toxicityof pyrethroids in Australian Helicoverpa armigera (Hübner) (Lepidoptera:Noctuidae). Pesticide Biochemistry and Physiology 63: 50 - 62.Gunning, R. & Ba;lfe M. (2002). Spinosad resistance in Helicoverpa armigera. Proceedings of 10 thInternational Pesticide Chemistry Conference, Basel Switzerland.Gunning, R. & Moores, G. (2002). Chlorfenapyr resistance in Australian Helicoverpa armigera.Proceedings of the British Crop Protection Council Conference – Pest and Diseases, pp. 793–798.Gunning, R. Dang, H. Kemp, F. Nicholson, I. &. Moores, G. (2005) New resistancemechanism in Helicoverpa armigera threatens transgenic crops expressing Bacillusthuringiensis Cry1Ac toxin. Applied and Environmental Microbiology. 71: 2558 – 2563.Gunning, R., Nicholson, I., Kemp, F., Borzatta, V., Cottage, E. Field L. & Moores, G. (2006).Piperonyl butoxide, restores the efficacy of Bacillus thuringiensis toxin in transgeniccotton against resistant Helicoverpa armigera. Biopesticides. International 2: 139 – 136.Gunning, R., Moores, G, Jewess, P., Boyes, A., Devonshire, A & Khambay, B. (2007). Use ofpyrethroid analogues to identify key structural features for enhanced-esteraseresistance in Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae). PestManagement Science 63: 276-281.Gunning, R. & Moores, G. (2010). The effects of diet on the detection of resistance to Cry1Actoxin in Australian Helicoverpa armigera Hubner (Lepidoptera: Noctuidae). PesticideBiochemistry and Physiology 97: 55–59.Hubert, J. Nesvorna, M. Zemek, R. Stara, J. and Stejskal V. (2008). Effects of metabolicinhibitors on activity of Cry1Ab toxin to inhibit growth of Ephestia kuehniella larvae.Pest Management Science 64: 1063 – 1068.Johnson, K., Gatehouse, J. & Anstee, J. (1993). Effects of soybean protease inhibitors on thegrowth and development of larval Helicoverpa armigera. Journal of Insect Physiology39: 657 – 664.Kunitz, M (1945). Crystallization of a trypsin inhibitor from soybean. Science 101: 668 – 669.Kunitz. M (1945). Crystalline soybean trypsin inhibitor. Journal of General Physiology 28: 149 –154.
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PrefaceWorldwide, soybean seed prot
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Soybean Products Consumption in the
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20Soybean and HealthWagner, J.D., A
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44Soybean and Healthrespectively, w
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46Soybean and HealthIn order to des
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Lunasin, a Cancer Preventive Seed P
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Insights into the Pharmacological E
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9Occurrence of Biogenic Aminesin So
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Occurrence of Biogenic Amines in So
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11From Soybean Phytosterolsto Stero
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From Soybean Phytosterols to Steroi
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17Anti-Diarrhoeal Aspectsof Ferment
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18Antioxidant and Hypocholesterolem
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19Important Minor Soybens Proteins:
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22Polyamines - The Principal Candid
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