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The Impact of Chlortetracycline on Drosophila Melanogaster and Aedes Aegypti Emma Victoria Ridley

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The Impact of Chlortetracycline on Drosophila Melanogaster and Aedes Aegypti Emma Victoria Ridley The impact of chlortetracycline on Drosophila melanogaster and Aedes aegypti Emma Victoria Ridley Ph.D. University of York Department of Biology May 2011 Abstract Microbial symbionts of insects have been demonstrated to play an important role in the nutrition and protection of the host; these include aphids and tsetse. Studies often use antibiotics to eliminate the symbionts but the deleterious impact of using these antibiotics is not commonly addressed. The impact of chlortetracycline treatment on Aedes aegypti and Drosophila melanogaster was investigated by assessing life-span, fecundity, development time, survival, nutrition and metabolism. The impact on microbial numbers and diversity was also determined. With Drosophila, treatment with 50 µg ml-1 and above showed a significant extension in development time and life-span, reduction in fecundity and change in nutritional content. Microbial numbers were significantly reduced at 50 µg ml-1 and above. Culturable techniques and 454 pyrosequencing, demonstrated that the microbial diversity of Drosophila was predominantly Acetobacter. Bacterial elimination through egg dechorionation yielded some similar results to chlortetracycline treatment. However, fecundity and life-span was not significantly affected. Microarray analysis established a significant reduction in the abundance of transcripts associated with immunity, particularly antimicrobial peptides. With Aedes aegypti, treatment significantly reduced the survival and also affected the life-span and nutrition of the insect. Microbial numbers of mosquito larvae were reduced at 30 and 100 µg ml-1. Colonies grew on plates supplemented with 50 µg ml-1 of chlortetracycline, indicating that the larvae bore chlortetracycline-resistant bacteria. 454 pyrosequencing demonstrated a change in diversity of bacteria found in mosquitoes +/- chlortetracycline, switching from Elizabethkingia meningoseptica to Raoultella sp with chlortetracycline. It is concluded that chlortetracycline significantly impacts the performance of the 2 insects through bacterial depletion, changes to bacterial diversity and toxicity. Nevertheless, different responses were observed with Aedes aegypti and Drosophila melanogaster. Moreover, experiments with Drosophila using egg dechorionation, emphasised the toxic impact of using antibiotics to eliminate microbes in the insect host. iii Contents Contents ................................................................................................................................. 1 Tables ..................................................................................................................................... 8 Figures .................................................................................................................................. 14 Acknowledgments ................................................................................................................ 18 Declaration ........................................................................................................................... 18 Chapter 1: Introduction........................................................................................................ 19 1.1 Commensal bacteria in vertebrate hosts ................................................................... 19 1.2 Role of bacterial symbionts in insects ........................................................................ 22 1.2.1 Role of bacteria in insect nutrition and insect performance ............................... 22 1.2.2 Bacterial symbionts and insect immunity and protection .................................. 24 1.3 Impact of tetracycline on microbes in insects ............................................................ 26 1.4 Antibiotic resistance ................................................................................................... 27 1.5 RIDL® sterile insect technique – Controlling insect pests .......................................... 29 1.6 Bacteria in Drosophila melanogaster and Aedes aegypti and the role in insect performance ..................................................................................................................... 31 1.6.1 Bacterial diversity in Drosophila melanogaster ................................................... 32 1.6.2 Role of commensal bacteria in Drosophila melanogaster................................... 32 1.6.3 Bacterial diversity in mosquitoes ......................................................................... 34 1.6.4 Role of commensal bacteria in Aedes aegypti ..................................................... 35 1.7 Thesis Objectives ........................................................................................................ 35 Chapter 2: Material and Methods ....................................................................................... 37 2.1 Materials ..................................................................................................................... 37 2.2 Insect Culture and maintenance ................................................................................ 37 2.2.1 Drosophila melanogaster rearing and maintenance ........................................... 37 2.2.2 Aedes aegypti rearing and maintenance ............................................................. 39 1 2.3 Insect Performance Experiments ............................................................................... 39 2.3.1 Performance of Drosophila melanogaster Oregon-RS ........................................ 39 2.3.2 Performance of Drosophila melanogaster Canton-S ........................................... 40 2.3.3 Performance experiments on Aedes aegypti ...................................................... 41 2.4 Microbial Culture ........................................................................................................ 43 2.4.1 Culturing microbes in Drosophila melanogaster ................................................. 43 2.4.2 Culturing microbes in Aedes aegypti ................................................................... 43 2.5 Nutrition Assays .......................................................................................................... 44 2.5.1 Wing length and area measurements ................................................................. 44 2.5.2 Tissue Preparation ............................................................................................... 45 2.5.3 Protein Assays ...................................................................................................... 46 2.5.4 Assay for Glucose, Glycogen and Trehalose ........................................................ 46 2.5.5 Assay for glycerol and triglyceride Assay ............................................................. 47 2.5.6 Total Lipid Content............................................................................................... 48 2.6 Respirometry .............................................................................................................. 48 2.7 Drosophila Gut Dissections ......................................................................................... 49 2.8 16S rRNA Gene Analysis ............................................................................................. 49 2.8.1 DNA extraction ..................................................................................................... 49 2.8.2 PCR Amplification ................................................................................................ 50 2.8.3 Wolbachia identification in Drosophila melanogaster using PCR ....................... 51 2.9 454 pyrosequencing ................................................................................................... 51 2.9.1 Sample Preparation and DNA Extraction ............................................................. 51 2.9.2 Amplification of Variable Region 2 (V2) of bacterial 16S rRNA gene ................. 52 2.9.3 Analysis of 454 sequences ................................................................................... 52 2.10 Microarray analysis .................................................................................................. 53 2.10.1 Insect rearing ..................................................................................................... 53 2 2.10.2 RNA Extraction and Sample Preparation (Method from John Ramsey) ............ 53 2.10.3 Microarray ......................................................................................................... 54 2.10.4 Microarray Analysis ........................................................................................... 54 2.10.5 Quantitative RT-PCR .......................................................................................... 57 2.11 Statistical Analysis .................................................................................................... 58 Chapter 3: Bacteria and performance of Drosophila melanogaster ................................... 60 3.1 Introduction ................................................................................................................ 60 3.2 Oregon-RS survival and development ........................................................................ 62 3.2.1 Oregon-RS survival and sex-ratio ......................................................................... 62 3.2.2 Development time to pupae and adulthood ....................................................... 64 3.2.3 Behaviour of larvae on chlortetracycline
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