STUDIES on HOST-SEEKING, RESTING BEHAVIOUR and CONTROL of the DENGUE VECTOR Aedes Aegypti
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STUDIES ON HOST-SEEKING, RESTING BEHAVIOUR AND CONTROL OF THE DENGUE VECTOR Aedes aegypti Thesis submitted in accordance with the requirements of the University of Liverpool for the degree of Doctor in Philosophy Submitted by Hadura Abu Hasan September, 2014 Acknowledgements Firstly, praises to Allah for giving me health, intelligence, wealth, strength and patience to complete this study. I am personally grateful and would like to express my sincere gratitude to Dr. Philip McCall, my main supervisor, for his valuable guidance, encouragement and resources throughout the study. I would also like to thank my secondary supervisor, Professor Hilary Ranson, for her support of various kinds that facilitated my study. I gratefully acknowledge my advisory community, Professor Mike Lehane and Dr. Charles Wondji, who helped me through all my progress research. This study was financially funded by and in collaboration with Universiti Sains Malaysia (USM) and I would like to thank them for their generous support. I am also thankful for funding provided by Syngenta for the field trial on Indoor Residual Spraying (IRS) in Penang, Malaysia and also the Vector Control Research Unit (VCRU) for their cooperation and technical assistance rendered throughout my study. My highest appreciation to Professor Zairi Jaal for his support and inspiration at all times. He provided office space, vehicles, materials, lab use and advice for all areas of my research as needed during my stay in Penang, Malaysia. To the staff and fellow students at Liverpool School of Tropical Medicine (LSTM) and School of Biological Sciences, USM, I am grateful for the chance to be a part of the lab. I am grateful to everyone who has helped me in the laboratory, and especially to the volunteers who spent their valuable time under the sticky net in extremely hot and humid conditions. I am exceptionally grateful for the statistical support given by Dr. Brian Faragher from LSTM and Dr. Daniel Impoinvil from LUCINDA. I would not have made it without their technical support in data management. Many thanks go to Dr. Amy Lynd, Dr. Mark Paine and Dr. Jacob Riverón Miranda from LSTM for the guidance with experimental works and HPLC analysis, and Dr. Natalia Angarita Jaimes from Leeds University for the analysis of video recording. I am indebted to Mrs Gwen Finnegan from LSTM who helped me in most aspects of my research, such as providing support with material and administrative functions. Special thanks to my parents, Abu Hasan Yahaya and Normah Murad, parents-in-law and siblings, who always encourage me to work hard and excel in my educational endeavours. Their investment in time and love is always cherished and I give my heartfelt thanks to all of them for their prayers, help and support. Most of all, I thank my husband, Yasir Azam, for his never-ending support and love for me during this very busy period of my life. While we were both struggling with our studies, his motivation and encouragement was endless. He has been a wonderful father to our two little loves, Husna and Hamzah, and took care of our family while I was away from home with my fieldwork and studies. With immeasurable gratitude and love, this thesis is dedicated to him. i Studies on host-seeking, resting behaviour and control of the dengue vector Aedes aegypti Hadura Abu Hasan Abstract Aedes aegypti is the main vector of dengue worldwide. A highly anthropophilic and endophilic mosquito, its behaviour is a major influence on dengue epidemiology and a major challenge to vector control, which is the only dengue prevention method available. A series of studies into host-seeking and resting behaviour were carried out in the laboratory. In Penang, Malaysia, the efficacy of standard and a novel modified form of indoor residual spraying was evaluated in a field trial and the insecticide susceptibility of local vector populations was determined. Arrival patterns of female Ae. aegypti were investigated at a seated human-bait protected by an adhesive-coated net. Mosquitoes preferentially landed on the top and nearest upper vertical surfaces of the net, clustering in a region above the volunteer’s head. Although not previously reported in Ae. aegypti, this behaviour supported the proposition that a plume of potential host attractants rises from the human host. Resting preferences of unfed female Ae. aegypti were investigated using simple two- dimensional panel targets and resting boxes. Exploring the influence of colour, texture, adhesive and target height, the highest resting rates were found on black targets in a vertical configuration at 90 cm above ground. Target texture and adhesive factor did not influence target attractiveness. Data also indicated that female Ae. aegypti were randomly distributed on the panels. In laboratory tests, significantly higher numbers of mosquitoes were captured in resting boxes by raising internal humidity to over 65%. However, a field test in Malaysia did not capture any Aedes sp., although Cx. quinquefasciatus were caught. A randomised-controlled trial was conducted in Penang, Malaysia to investigate the effect of indoor residual spraying (IRS) on Ae. aegypti and Ae. albopictus populations. Two insecticides (lambda-cyhalothrin and pirimiphos-methyl) were delivered either by standard (entire interior surface sprayed) or selective IRS (upper walls and ceilings sprayed) methods. Throughout the three-month study, entomological indices fluctuated considerably and, while there was some evidence of an overall effect throughout the study area, due to a number of confounders comparison between treatments was not possible and the outcome was ultimately inconclusive. At the trial study site in Penang, the insecticide susceptibility status of local populations of Ae. aegypti, Ae. albopictus and Cx. quinquefasciatus were investigated. All were found to be resistant to lambda-cyhalothrin. For pirimiphos-methyl, Ae. aegypti and Ae. albopictus remained susceptible but Cx. quinquefasciatus was classed as ‘suspected resistance’ and potential resistance management strategies are discussed. The study has demonstrated the potential to improve traps or targets for Ae. aegypti by simple alterations to their design. The potential of IRS in the control of dengue vectors remains to be confirmed. The data on emerging insecticide resistance in the mosquito vector populations is timely and provides an evidence base for local authorities to reconsider management strategies that are currently in place for the control of dengue vectors in Malaysia. ii Table of Contents Acknowledgements ..................................................................................................... i Abstract ...................................................................................................................... ii Table of Contents ...................................................................................................... iii List of Tables........................................................................................................... viii List of Figures .......................................................................................................... xii List of Appendices ................................................................................................. xvii List of Abbreviations............................................................................................. xviii CHAPTER 1 LITERATURE REVIEW OF DENGUE, DENGUE VECTORS AND ADULT CONTROL MEASURES ......................................................................... 1 1.1 Introduction to dengue ...................................................................................... 1 1.1.1 Dengue fever (DF) and severe dengue .................................................... 2 1.1.2 Treatment of dengue infection ................................................................. 3 1.1.3 Dengue vaccine development .................................................................. 3 1.1.4 Geographical distribution and incidence of dengue ............................... 4 1.1.5 Transmission of dengue viruses .............................................................. 9 1.2 Introduction to dengue vectors ....................................................................... 11 1.2.1 Aedes aegypti (Linnaeus) ...................................................................... 11 1.2.2 Aedes albopictus (Skuse) ....................................................................... 12 1.3 Behaviour of Aedes sp. mosquitoes ................................................................ 13 1.3.1 Mating behaviour .................................................................................. 13 1.3.2 Sugar-feeding behaviour ....................................................................... 14 1.3.3 Host-seeking behaviour ......................................................................... 15 1.3.4 Blood-feeding behaviour ....................................................................... 17 1.3.5 Oviposition behaviour ........................................................................... 19 1.4 Dengue vector control..................................................................................... 21 1.4.1 Vector surveillance ................................................................................ 23 1.4.2 Control of immature stages of Aedes aegypti........................................ 24 1.4.2.1 Insecticides for larval control ................................................