Bactrocera Cucurbitae (Diptera: Tephritidae), Sexual Behaviour
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Effects of the juvenile hormone analogue methoprene and dietary protein on the male melon fly, Bactrocera cucurbitae (Diptera: Tephritidae), sexual behaviour Dissertation submitted by Ihsan ul Haq M.Sc. (Honours) Supervisors Ao. Univ. Prof. Dipl.-Ing. Dr. Christian Stauffer Dr. Jorge Hendrichs (FAO/IAEA) Dr. Alan S. Robinson (FAO/IAEA) Institut für Forstentomologie, Forstpathologie und Forstschutz, Department für Wald- und Bodenwissenschaften Universität für Bodenkultur Wien Vienna, September 2010 Abstract The application of a juvenile hormone analogue, methoprene and access to protein in adult diet has been shown to enhance mating success of some tephritid males, supporting their incorporation as a pre-release treatment for males utilized in the sterile insect technique (SIT). The effect of protein-diet (P) (contains hydrolyzed yeast and sugar) and/or treatment with methoprene (M), (in addition to sugar and water) on genetic sexing strain (GSS) of Bactrocera cucurbitae Coquillett (Diptera: Tephritidae) was assessed in the laboratory and in field cages. The males were exposed to either 1) methoprene and protein-diet (M+P+), 2) only protein-diet (M-P+), or 3) only methoprene (M+P-), or 4) only sugar (M-P-). The effect of methoprene and protein-diet on mating behaviour of the four groups of males, on starvation survival and on body protein and lipids was studied. Total body nitrogen (TBN) and total body carbon (TBC) were measured as indicators of body protein and body lipids repectively. The M-P+ males showed increased performance at leks and higher mating success compared to M+P- and M-P- males. Application of methoprene accelerated sexual maturity and enhanced mating competitiveness only in protein-fed males, but not in sugar-fed males. More M+P+ males called and initiated and participated in lek activities than all other types of male, consequently attaining a higher mating success. No adverse effect of methoprene and/ or protein but feeding duration had significant effect on starvation survival. Protein-diet had significantly increased the male body weight, TBN and TBC as compared to only sugar-fed males. Methoprene had no significant effect on males’ TBN and TBC. The results provide a strong evidence for the benefits of exposing sterile males to methoprene and protein-diet. Treated males achieve higher sexual success, reach sexual maturity several days earlier, and are therefore much closer to sexual maturity when released as part of SIT action programmes after being held in the fly emergence and release facility. i Kurzfassung Die Applikationen vom Juvenilhormonanalog Methoprene und die Zugabe von Proteinen zur Fliegendiät können zu einer signifikanten Steigerung der Paarungshäufigkeit bei Fruchtfliegenmännchen führen. Dies ist vor allem bei der Anwendung der Sterile Insect Technique (SIT) von großem Vorteil und wird vor der Freilassung der sterilen Männchen angewandt um die Kosten in der Zucht der Tiere zu vermindern. In dieser Arbeit wurde die Auswirkungen von Protein P (hydrolyzed yeast) und die Applikation von Methoprene M auf die wirtschaftliche bedeutsame Fruchtfliegenart Bactrocera cucurbitae (Diptera: Tephritidae) analysiert. Dies wurde einerseits unter strikten Laborbedingungen, andererseits in Freilandkäfigen durchgeführt. Männchen wurden folgendermaßen behandelt: 1) Methoprene und Protein M+P+ 2) Protein M-P+ 3) Methoprene M+P- 4) Kontrolle M-P-. Der Effekt von Methoprene und Protein wurde in Bezug auf das Paarungsverhalten, auf die Überlebensdauer, dem Proteingehalt (total body protein) und dem Fettgehalt (total lipid content) analysiert. M-P+ Männchen zeigten eine bessere Leistung in den Leks und das Paarungsverhalten was signifikant höher als bei den M+P- und M-P- behandelten Männchen. Die Applikation von Methoprene hatte einen signifikanten Effekt auf die sexuelle Reifung der Männchen und verbesserte somit deren Paarungsverhalten. Die M+P+ behandeltet Männchen hatten den größten Erfolg nicht nur im Lek sondern auch in der Paarungsverhalten. Die Ergebnisse zeigen bei Bactrocera cucurbitae deutliche Vorteile von Methoprene und Protein behandelten Männchen gegenüber den Kontrollen. Durch diese Behandlung lässt sich bei dieser Fruchtfliegenart Zucht und Effizienz der Männchen im Paarungsverhalten wesentlich verbessern. Im Rahmen von SIT Programmen kann somit diese Behandlung die Zuchterfolge wesentlich verbessern, einerseits durch die gesteigerte Paarungseffizienz der behandelten Männchen nach der Freisetzung, andererseits durch die signifikante Reduktion der Zuchtdauer und somit den niedrigeren Kosten. ii Acknowledgments I thank my supervisor Christian Stauffer, for his involvement, providing a comfortable environment to conduct my research and his critical inputs to give a final shape to this document. I am grateful to Jorge Hendrichs for being a gentle mentor and for providing me the opportunity to join Insect Pest Control Laboratory, Joint FAO/IAEA Laboratories, Seibersdorf, Austria. I have benefitted immensely from his scholarly thoughts and also critical review of all manuscripts. Suffice it to say that interacting with him greatly influenced my development as a scientist and an individual. I would like to tribute his abilities as a scientist that I wish to have a capability to think along the same intellectual wavelength as him at the end of this journey. Special thanks to Alan Robinson for his endless patience, deep involvement in planning and execution of research and generosity for reviewing the drafts of manuscripts. I owe my thanks to Carlos Cáceres for facilitating the experimentation. His involvement guided me to define the objectives very clearly and to be on track to reach the destination with least chances of failure. I am greatly indebted to laboratory colleagues, Marc Vrerysen, Andrew Jessup, Andrew Parker, Adly Abd Ala and Cardoso Rui Pereira for their kindness and sharing their wisdom during the whole course of studies. Their rigorous involvement during the planning of experiments and deciphering on the results enabled me to conduct the studies in a systematic way and indeed made my stay more productive. Thanks to Peter Teal for his guidance for experimentation in areas of chemical ecology and analytical work. The skilled team at Seibersdorf laboratories helped me to maintain the research material used in my experiments. I owe my thanks to Viwat Wornoayporn Amirul Islam, Sohel Ahmad and Thilaqasiri Dammalage for unfailing support for fly culture maintenance and logistics, whenever needed during my stay. I would like to thank Higher Education Commission of Pakistan for financial support and Pakistan Agricultural Research Council for sparing me to do PhD studies. iii Table of Contents Abstract i Kurzfassung ii Acknowledgements iii Chapter I 1. Introduction 01 1.1 Tephritidae 01 1.1.1 Mating systems in tephritid fruit flies 02 1.1.2 Economic importance of fruit flies 06 1.2 Management of fruit flies 06 1.3 The Sterile Insect Technique (SIT) 07 1.3.1 Diet supplementation: Hydrolyzed yeast 10 1.3.2 Juvenile hormone analogue: Methoprene 12 1.4 The melon fly, Bactrocera cucurbitae: A case study for the 13 SIT 1.5 Aims of this study 16 Chapter II Conclusions 18 References 21 Chapter III Haq, I., Cáceres, C., Hendrichs, J., Teal, P.E.A., 37 Wornoayporn, V., Stauffer, C. and Robinson, A.S., 2010. Effects of the juvenile hormone analogue methoprene and dietary protein on male melon fly Bactrocera cucurbitae (Diptera: Tephritidae) mating success. Journal of Insect Physiology, 56, 1503-1509 Chapter IV Haq, I., Cáceres, C., Hendrichs, J., Teal, P.E.A., Stauffer, 45 C. and Robinson, A.S., 2010. Methoprene modulates the effect of diet on male melon fly Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae) performance at mating aggregations. Entomologia Experimentalis et Applicata, 136, 21-30. Chapter V Haq, I., Cáceres, C., Liedo, P., Soriano, D., Jessup, A., 55 Hendrichs, J., Teal, P.E.A. and Robinson, A.S., 2010. Effect of methoprene application, adult food and feeding duration on male melon fly starvation survival. Journal of Applied Entomology (in press), doi: 10.1111/j.1439- 0418.2010. 01575.x iv Chapter VI Haq, I., Mayr, L., Teal, P.E.A., Hendrichs, J., Robinson, A. 63 S., Stauffer, C. and Hood-Nowotny, R., 2010. Total body nitrogen and total body carbon as indicators of body protein and body lipids in the melon fly Bactrocera cucurbitae: effects of methoprene, a juvenile hormone analogue, and of diet supplementation with hydrolyzed yeast. Journal of Insect Physiology (in press),10.1016/j.jinsphys.2010.07.011. Curriculum Vitae 72 v Introduction r I e apt Melon fly male h C Monitoring Mass Rearing Source: Dominiakiak,, NSW, Au Aerial Release Sterilization Chapter I 1. Introduction The sterile insect technique (SIT) is the principal non-chemical, biological-based eco- friendly component of pest management of tephritid (Diptera: Tephritidae) flies (Hendrichs et al., 2002). Improvements in the quality of mass-reared sterile males being released in the field can increase the effectiveness of the SIT (Robinson et al., 2002). The objective of this study is to analyze the effect of juvenile hormone analogue, methoprene and dietary protein on male melon fly Bactrocera cucurbitae Coquillett (Diptera: Tephritidae) sexual behaviour and their application as a tool to increase the effectiveness of the SIT. Rationale More than 40% of world crop production valued at USD 750 000 million is destroyed every year by pests (Oerke et al., 1994). Considering all pests,insects caused about 14%, plant pathogens