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Effect of Stress and Diapause in Two Calliphoridae Species

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Effect of stress and diapause in two Calliphoridae species. by Bobbie Johnson A thesis submitted to The University of Birmingham for the degree of DOCTOR OF PHILOSOPHY School of Biosciences The University of Birmingham January 2013 0 Abstract Cultures of two Dipteran flies (Calliphora vicina (R-D) and C. vomitoria (L.)) were established to answer questions in regards to responses to thermal and desiccation stress, effects of diapause and the mechanisms which underpin diapause. The findings are divided in to two sections. Unequivocal new findings – Calliphora vomitoria were seen to depend on water being present in culture medium for survival to be high. Furthermore, C. vomitoria were found to be less able to resist desiccation than C. vicina. Larvae of C. vicina and C. vomitoria showed different cold tolerance strategies, with C. vicina being freeze-avoiding and C. vomitoria ‘partially’ freeze- tolerant. Metabolomics, using 1H-NMR, revealed that diapause and non-diapause had distinct metabolic profiles. Diapause larvae were seen to reduce energy synthesis from the Krebs cycle and increase glycolysis. Calliphora vicina and C. vomitoria also exhibited different diapause phenotypes; C. vicina entered a maternally regulated facultative diapause as an L3 larvae, Calliphora vomitoria had a less distinct diapause, with maternal conditions having little effect. Speculative new findings - Despite the above differences in culturing, desiccation and cold tolerance, C. vicina and C. vomitoria were able to produce a viable cross, though field fresh C. vomitoria were not used, as such it cannot be confirmed if this could occur in the wild. Field studies showed that increased temperatures due to climate change may affect both phenology and survival of insects; C. vicina was seen to have a delayed induction to diapause and a reduction in the proportion entering diapause. Diapause conferred increased cold tolerance; as such those insects that overwinter not in diapause may suffer increased mortality. This study only examined one aspect of climate change – increased temperatures. To get a more holistic view an experimental design including precipitation, U.V. and thermal extremes is needed. This research shows how even closely related species can evolve different mechanisms to cope with stress, it is hoped that through RNAi and or epigenetics that this research will be furthered. 1 Acknowledgements I would like to thank NERC for sponsoring my PhD, and to my supervisors Scott Hayward and Jeff Bale for writing the project proposal and supporting me through the whole process. I was privileged to be part of a wonderful lab group at Birmingham University. Lucy and Steve offered much needed help while I was setting up the lab, and later on providing help with statistical analysis. My greatest gratitude goes to Gwen who not only provided invaluable insight in to lab life but became one of the greatest friends anyone could ask for. Thanks also go to my ‘lab partner’ Paul Coleman who kept me entertained throughout the many hours of counting and sorting smelly maggots and to Emily who was a late entrant in to ‘the dream team’. My thanks also go to Megan, Ji and Matt who not only proof-read parts of my thesis but kept me fit during the many hours of chasing a small ball around the squash courts. I really will miss our lab lunches and outings, and I thank each of you for creating a wonderful atmosphere in which to spend three and a half years. Special thanks go to John for simply being there when I needed him most and helping even with the most mundane and sole-destroying of tasks. I really can’t thank you enough! My final thanks go to my family; my brother Martyn, who endeavors to keep both my feet firmly on the floor, and especially, to my parents for convincing me to do a degree in the first place. I have loved each of my years at university; and I am truly thankful for everything my parents have done for me. 2 Table of Contents 1. Introduction ................................................................................................................................ 1 1.1. Diapause ...................................................................................................................................... 2 1.1.1. Obligatory and facultative diapause ............................................................................... 3 1.1.2. Pre-diapause stage .......................................................................................................... 4 1.1.3. Induction ......................................................................................................................... 5 1.1.4. Preparation ..................................................................................................................... 5 1.1.5. Diapause stage ................................................................................................................ 6 1.1.6. Diapause initiation .......................................................................................................... 6 1.1.7. Diapause maintenance........................................................................................................ 8 1.1.8. Diapause termination ................................................................................................... 10 1.1.9. Post diapause ................................................................................................................ 11 1.2. Alternate diapause durations ........................................................................................... 12 1.2.1. Fixed prolonged diapause ............................................................................................. 12 1.2.2. Bet hedging within the population ............................................................................... 12 1.2.3. Tropical diapause .......................................................................................................... 13 1.2.4. The role of diapauses in regulating phenology ................................................................. 14 1.3. Insect cold tolerance ......................................................................................................... 15 1.3.1. Freeze tolerance vs. Freeze avoidance ......................................................................... 16 1.3.2. Problems with Salt’s definition ..................................................................................... 17 1.3.3. Cold tolerance and diapause......................................................................................... 18 3 1.4. Measuring insect thermal tolerance ................................................................................. 20 1.4.1. SCP................................................................................................................................. 20 1.4.2. Lethal time and temperatures ...................................................................................... 21 1.4.3. Rapid cold hardening .................................................................................................... 22 1.5. Mechanisms underpinning stress adaptations ................................................................. 22 1.5.1. Cryoprotectants ............................................................................................................ 23 1.5.2. Anti-freeze proteins ...................................................................................................... 25 1.5.3. Ice nucleators ................................................................................................................ 26 1.5.4. Heat shock proteins ...................................................................................................... 29 1.5.5. Membrane adaptations ................................................................................................ 30 1.6. Desiccation stress .............................................................................................................. 33 1.6.1. Mechanisms underpinning desiccation resistance ....................................................... 33 1.6.2. Mechanisms underpinning desiccation tolerance ........................................................ 34 1.6.2.1. Aquaporins ................................................................................................................ 35 1.6.2.2. Late Embryonic Abundant Proteins .......................................................................... 35 1.6.2.3. Heat Shock Proteins .................................................................................................. 36 1.6.3. Cryoprotective dehydration .......................................................................................... 36 1.7. Metabolomics ........................................................................................................................... 37 1.8. Climate change .......................................................................................................................... 39 1.8.1. Current predictions ....................................................................................................... 40 1.8.2. The impact of climate change on insect phenology, distribution and abundance ....... 42 1.8.3. Broader impacts of disrupted insect phenology ............................................................... 44 4 1.9. Choice of study system ............................................................................................................
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