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The oviposition and movement behaviour of bt resistant and bt susceptible helicoverpa armigera (hubner) (lepidoptera noctuidae) on bt cotton and non bt cotton plants

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  • The oviposition and movement behaviour of bt resistant and bt susceptible helicoverpa armigera (hubner) (lepidoptera noctuidae) on bt cotton and non bt cotton plants

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The oviposition and movement behaviour of Bt-resistant and Bt-susceptible Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) on Bt cotton and non-Bt cotton plants Luong, Thi Anh Tuyet Bsc MPh A thesis submitted for the degree of Doctor of Philosophy at The University of Queensland in 2016 The School of Biological Sciences i Abstract Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) has caused poor yields to a range of agricultural crops, particularly to cotton Pesticides have been used to control this pest with serious undesirable side effects, including the rapid development of high levels of resistance Since 1996, genetically modified cotton (Bt cotton) has been planted to control H armigera in Australia However, it is reported that surviving larvae of all sizes can be found in fields from time to time in all growing regions Research has shown that the survival of these larvae on Bt cotton is not necessarily due to physiological resistance, and behavioural resistance has been inferred Extensive work on various aspects of behaviour of H armigera in Australian has been conducted; however, to date those experiments were carried out using a Bt-susceptible H armigera strain Experiments in this thesis were undertaken with both physiologically Bt-resistant and Bt-susceptible lines of H armigera One might expect larvae that are susceptible to Bt to show differences in behaviour in comparison to Bt-resistant larvae Oviposition choice experiements consistently showed that both Bt-resistant and Bt-susceptible moths did not choose plants or plant parts that were less toxic in terms of Bt toxin on which to lay eggs There was one exception in that Bt-susceptible moths were more likely to lay eggs on squares of Bt cotton plants than those of non-Bt cotton As expected the mortality of Bt-susceptible H armigera larvae was significantly higher on structures of Bt cotton plants than on those structures of conventional cotton, and survival was greater on flowers than on other structures of Bt cotton Bt-susceptible neonates of H armigera, which were significantly heavier, could starve longer and recover better than Bt-resistant neonates Although H armigera neonates did not shift their behaviour towards Bt toxin on artificial diet before their first feeding event, Bt-susceptible neonates showed a tendency to remain on non-Bt diet and move off Bt diet These behaviours may have allowed them to survive in a Bt environment, and led to a higher percentage of survival and pupation in situations where a choice was offered The behaviour of Bt-susceptible larvae, which differed from Bt-resistant larvae, could help them avoid Bt toxin by moving off Bt substrates (drop-off behaviour) or staying on Bt substrates but eating their conspecific eggs and so survive the first instar stage (cannibalism behaviour) There was a significant difference in the numbers of Bt-susceptible larvae dropped off the two lines of cotton, more Bt-susceptible larvae dropped off Bt cotton than non-Bt cotton plants over time Significantly more Bt-susceptible larvae remained on squares and flowers, and this may offer an opportunity for them to survive on Bt-cotton plants The survival of Bt-susceptible H armigera larvae significantly improves on Bt cotton plant when they cannibalize eggs before feeding on the ii plant Cannibalism may play a significant part in the survival of Bt-susceptible H armigera larvae on Bt cotton plants Egg cannibalism could explain the relatively small, but surprising number of Bt-susceptible larvae surviving in Bt cotton fields In conclusion, subtle differences in the behaviour of Bt-susceptible female moths and first instar H armigera larvae may allow the larvae to survive on Bt cotton plants Some of Bt-susceptible larvae laid on squares or flowers that contain less Bt toxin and are generally more nutrient rich have higher chances to survive In addition, egg cannibalism contributes to the higher survival of Bt-susceptible larvae on other structures of Bt cotton plants Larvae can move off plant parts with high Bt levels and may encounter less toxic plant parts Bt-susceptible larvae can survive without food for 48 h, suggesting they have time to re-establish on less Bt toxic plant parts after moving away from higher toxic areas iii Declaration by author This thesis is composed of my original work, and contains no material previously published or written by another person except where due reference has been made in the text I have clearly stated the contribution by others to jointly-authored works that I have included in my thesis I have clearly stated the contribution of others to my thesis as a whole, including statistical assistance, survey design, data analysis, significant technical procedures, professional editorial advice, and any other original research work used or reported in my thesis The content of my thesis is the result of work I have carried out since the commencement of my research higher degree candidature and does not include a substantial part of work that has been submitted to qualify for the award of any other degree or diploma in any university or other tertiary institution I have clearly stated which parts of my thesis, if any, have been submitted to qualify for another award I acknowledge that an electronic copy of my thesis must be lodged with the University Library and, subject to the policy and procedures of The University of Queensland, the thesis be made available for research and study in accordance with the Copyright Act 1968 unless a period of embargo has been approved by the Dean of the Graduate School I acknowledge that copyright of all material contained in my thesis resides with the copyright holder(s) of that material Where appropriate I have obtained copyright permission from the copyright holder to reproduce material in this thesis iv Publications during candidature T.T.A Luong, M.P Zalucki, B Cribb, L.E Perkins and S.J Downes Oviposition site selection by adults and the survival of susceptible and resistant first instar larvae of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) on genetically modified and conventional cotton Bulletin of Entomological Research (Accepted, 2016) Publications included in this thesis Chapter T.T.A Luong, M.P Zalucki, B Cribb, L.E Perkins and S.J Downes Oviposition site selection by adults and the survival of susceptible and resistant first instar larvae of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) on genetically modified and conventional cotton Bulletin of Entomological Research (Accepted, 2016) A modified and re-formatted version of this has been incorporated into Chapter Contributor Statement of contribution Tuyet T A Luong (Candidate) Designed experiments (80%) Wrote the paper (70%) Myron P Zalucki (Principal supervisor) Designed experiments (20%) Wrote and edited paper (15%) Lynda E Perkins (Co-supervisor) Wrote and edited paper (5%) Sharon J Downes (Co-supervisor) Wrote and edited paper (5%) Bronwen Cribb (Co-supervisor) Wrote and edited paper (5%) v Contributions by others to the thesis Professor Myron P Zalucki, Lynda E Perkins, Sharon J Downes and Bronwen Cribb made significant contributions to the conception and design of the project, and provided advice and guidance regarding analyses throughout the research Statement of parts of the thesis submitted to qualify for the award of another degree None vi Acknowledgements It has been a long, tough and challenging journey and I am so grateful that I have finally finished my PhD However, this journey wouldn’t have been possible without the help, support and encouragement of so many people around me to whom I would like to take this opportunity to express my gratitude and appreciation I am grateful and would like to express my sincere thanks to my principal supervisors, Professor Myron P Zalucki for his patience and supervision of the research I greatly appreciate and value the help he provided me in developing logical thinking and teaching me on how to propose good questions and finding ways to get the answers I would like to thank my co-supervisors Dr Bronwen Cribb for her general support and helpful advice during my PhD I am eternally gratefully for Dr Lynda E Perkins for her useful advice on statistics and writing Thank you also to Dr Sharon J Downes, who, with her own practical knowledge, has given me some excellent suggestions on the design of some of my experiments, and gave me a chance to have wonderful experience on the field in Narrabri, NSW I am grateful to Vietnamese Government Scholarship (MOET) in conjunction with The University of Queensland for funding my research during my PhD studying I also acknowledge CSIRO, and The Cotton Research and Development Corporation for supplying cotton seeds and Helicoverpa armigera source for my experiments My field-based experience was wonderful due to the generous support of a handful of people from CSIRO Agriculture, Australian Cotton Research Institute, Narrabri, Australia who gave their knowledge, time and friendship to support me In particular I would like to thank Tracey Parker, who in the initial stages of my PhD gave me valuable experience in H armigera, and continuously sending cotton seeds and H armigera eggs for my experiments Thank you to my friends, both inside and outside of the university, who have made my time during this research a fun one In particular to Corinna L Lange and Jason Callander for their support and often giving me their critical comments on certain chapters of this thesis Finally I would like to thank my family for their love and support throughout my PhD My parents, Luong Minh Son and Nguyen Thi Trinh, gave me a childhood that inspired a love of nature and instilled a drive to further my education Thank you also to my brother, Luong Minh Tung, who encouraged me to begin my PhD, and to finish it I deeply appreciate the support of my parents-inlaw, Tran Van Muoi and Nguyen Thi Hong Phuong during my PhD Lastly, I would like to thank my husband, Tran Xuan Hiep, for his love, support and encouragement over my PhD Luckily for me Hiep has an excellent understanding of computers and has assisted me with computer skills He has also given up his down time to assist with editing most of the chapters vii Keywords Helicoverpa armigera, Bt-resistant, Bt-susceptible, Bt cotton, Bt-resistant behaviour, oviposition behaviour, time to starvation, recovery ability, Bt-detection ability, drop-off behaviour, egg cannibalism Australian and New Zealand Standard Research Classifications (ANZSRC) ANZSRC code: 060801, Animal behaviour, 70% ANZSRC code: 070308, Crop and Pasture Protection, 20% ANZSRC code: 060499, Genetics not elsewhere classified, 10% Fields of Research (FoR) Classification FoR code: 0608, Zoology, 70% FoR code: 0703, Crop and Pasture Production, 20% FoR code: 0699, Other Biological Sciences, 10% viii Table of Contents Abstract ii Acknowledgements vii Chapter Literature Review 1.1 General introduction 1.2 Host-plant selection behaviour 1.3 Oviposition preference .9 1.4 The movement behaviour of first instar H armigera 12 1.5 Bacillus thuringiensis toxins 14 1.6 Bt crops and Bt cotton 14 1.7 The expression of Bt genes on Bt cotton .16 1.8 Behavioural resistance 17 1.9 Cannibalism in natural population .18 1.10 Structure of the thesis 21 Chapter 24 Oviposition site selection and survival of Bt-resistant and Bt-susceptible larvae of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) on Bt and non-Bt cotton 2.1 Introduction 26 2.2 Materials and methods 27 2.2.1 Insects 27 2.2.2 Plants 28 2.2.3 Oviposition preference .30 2.2.4 Survival of newly hatched larvae .31 2.2.5 Data analysis 31 2.3 Results 33 2.3.1 Oviposition preference .33 2.3.2 Survival 34 2.4 Discussion 36 Chapter 39 Feeding and survival of Bt-resistant and Bt-susceptible larvae Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) when exposed to a diet with Bt-toxin 3.1 Introduction 41 3.2 Materials and Methods 43 3.2.1 Insects 43 3.2.2 How long can larvae survive starvation? 43 3.2.3 Can H armigera larvae recover after a period of starvation? 44 3.2.4 Can H armigera larvae detect Bt toxin on artificial diet? .44 3.2.5 Bt detection assay: detailed observations 45 3.2.6 Data Analysis .46 3.3 Results 47 3.3.1 How long can H armigera larvae survive starvation? 47 ix 3.3.2 Can H armigera larvae recover after a period of starvation? 48 3.3.3 Can H armigera larvae detect Bt toxin on artificial diet? .49 3.3.4 Bt detection assay: detailed observations 50 3.4 Discussion 55 Chapter 58 The drop-off behaviour of Bt-resistant and Bt-susceptible Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) larvae on Bt-cotton and non-Bt cotton plants 4.1 Introduction 60 4.2 Material and Methods 61 4.2.1 Plants 61 4.2.2 Insects 61 4.2.3 Drop-off behaviour of Bt-resistant and -susceptible H armigera neonates on artificial diet with and without Bt toxin 62 4.2.4 Drop-off behaviour of Bt-resistant and -susceptible H armigera neonates on leaves of Bt cotton and non-Bt cotton 63 4.2.5 Drop-off behaviours of Bt-resistant and -susceptible neonates of H armigera on different structures of Bt and non-Bt cotton plants 64 4.3 Results 65 4.3.1 Drop-off behaviour of Bt-resistant and Bt-susceptible H armigera larvae on artificial diet with and without Bt toxin 65 4.3.2 Drop-off behaviour of Bt-resistant and -susceptible H armigera larvae on leaves of Bt and non-Bt cotton plants .67 4.3.3 Drop-off behaviour of Bt-resistant and -susceptible H armigera neonates on different structures of Bt and non-Bt cotton plant .68 4.4 Discussion 72 Chapter 76 Egg cannibalism in Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) larvae: overcoming the plant establishment hurdle Abstract 77 5.1 Introduction 78 5.2 Materials and Methods 80 5.2.1 Egg cannibalism by Bt-susceptible H armigera neonates on different eggs in a no choice experiment 80 5.2.2 Comparison of egg cannibalism by 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Xem thêm: The oviposition and movement behaviour of bt resistant and bt susceptible helicoverpa armigera (hubner) (lepidoptera noctuidae) on bt cotton and non bt cotton plants , The oviposition and movement behaviour of bt resistant and bt susceptible helicoverpa armigera (hubner) (lepidoptera noctuidae) on bt cotton and non bt cotton plants