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Transgenerational Effects: Nutrition, Immunity and Infection Transgenerational effects: nutrition, immunity and infection dynamics in Plodia interpunctella By Joanne Elizabeth Littlefair Submitted in partial fulfilment of the requirements of the Degree of Doctor of Philosophy at the University of London School of Biological and Chemical Sciences Queen Mary, University of London July 2015 1 Statement of originality I, Joanne Elizabeth Littlefair, confirm that the research included within this thesis is my own work or that where it has been carried out in collaboration with, or supported by others, that this is duly acknowledged below and my contribution indicated. Previously published material is also acknowledged below. I attest that I have exercised reasonable care to ensure that the work is original, and does not to the best of my knowledge break any UK law, infringe any third party’s copyright or other Intellectual Property Right, or contain any confidential material. I accept that the College has the right to use plagiarism detection software to check the electronic version of the thesis. I confirm that this thesis has not been previously submitted for the award of a degree by this or any other university. The copyright of this thesis rests with the author and no quotation from it or information derived from it may be published without the prior written consent of the author. Signature: Date: 2 Details of collaboration and publications Collaboration and support is acknowledged in more detail at the beginning of each chapter. Multiple infections result in a complex response to transgenerational immune priming. Littlefair, J; Laughton, A; Knell, R. J Functional Ecology (submitted) The development of a synthetic diet for investigating the effects of macronutrients on the development of Plodia interpunctella. Littlefair, J; Nunn, K. A; Knell, R. J Entomologia Experimentalis et Applicata (in review) 3 Abstract Phenotype can be shaped by transgenerational effects from parental and even grandparental environments. Parental resource allocation strategies can alter offspring condition to increase fitness. Alternatively, environmental influences such as stress, pollution, or pesticides, can be transmitted as non-adaptive “developmental noise”. This thesis explores the roles that nutrition and infection play as transmitted effects to offspring, using the Indian meal moth Plodia interpunctella as a model host. Parents acquire more resources to provision offspring with through their diet. However, interactions with offspring environment often go unexamined due to the complex nature of these factorial studies. Using good and poor diet qualities, I demonstrate the relative importance of parental and offspring environments and how this changes for offspring disease resistance, weight and longevity. I find that parental effects are present but are modest in comparison to the effects of the offspring’s own environment. I then develop a synthetic artificial diet to examine which macronutrients (such as protein and carbohydrate) within parental diet influence offspring phenotype. These diets allowed both the total nutritional content to be diluted, and the ratio of protein and carbohydrate to change. I find that offspring are relatively robust to extreme changes in parental dietary macronutrient content, which possibly explains the success of Plodia interpunctella as a widespread pest species. Vertical transmission of upregulated immune function has recently been demonstrated in insects, which is reliant on parental exposure but functions without an obvious acquired transmitted cue. To date, experiments have been conducted using single infections under laboratory conditions. However, this study shows that in a more ecologically realistic scenario involving multiple infections and resource limitation, 4 maternal pathogen exposure results in environmentally-contingent decreases in offspring survival in response to infection. Transgenerational immune priming may not as common as previously thought, and any benefits accrued to offspring should be viewed within an inclusive framework that includes potential costs to the parents. 5 Acknowledgements Thank you to Rob Knell for supervising me and well done for not dying in the last four years. Steve Rossiter, Steve le Comber and Christophe Eizaguirre provided fantastic encouragement, academic mentoring, and rounds of drinks. Beth Clare and Richard Buggs employed me when I had no money, and special thanks to Elisa Piccarro for help with my speech recognition stuff. Thank you to the Gen Foundation who provided me with a financial grant during the immune priming experiment. I am grateful to fantastic postdocs that helped me throughout, making this mountain much more achievable and fun. Alice Laughton helped me ace disease resistance assays and advanced gin drinking. Andres Arce injected burying beetles with me until the wee hours of the morning in Manchester, mocked my southern ways and fed me offal. Eoin O’Gorman is a brilliant friend and partner in crime and it’s safe to say that without him I would be a lot more sober but would also have given up a long time ago. My family are a huge source of support, even though they weren’t sure what I was doing (not working on bee diseases!). Jamie & Judith provided entertaining family dinners, my sister navigated my 20s with me, and Mum and Dad hosted me, the moths and their entomological friends with bemused grace. Thanks to my grandparents at Streams Farm who introduced me to a love for ecology a long time ago, provided a beautiful bolt hole out of London, and injunctions to always ‘keep my end up’. Thanks also to my brilliant group of friends. Art friends, Rebecca, and Lokadhi Lloyd are reminders that the world is much more than a PhD. The evolve gang were a great lunchtime crew; ecology office mates provided tea and distraction, Aurora helped me over the finishing line with the typing, everyone at WISE inspired me to keep plodding 6 on, Katie gave me a house to stay in, beer club fatally introduced me to poitin, and girls dinners have been a lifesaving source of gruesome medical facts and cream buns. Last but never least – my PhD would not have been possible without Alex Jones - thank you for loving me and encouraging me all this time. 7 Contents Title page .............................................................................................................................1 Statement of originality........................................................................................................2 Collaborations and publications arising from this thesis .....................................................3 Thesis abstract ......................................................................................................................4 Acknowledgements ..............................................................................................................6 Contents ...............................................................................................................................8 List of figures .....................................................................................................................12 List of tables .......................................................................................................................13 Chapter 1 General introduction 1.1 Defining transgenerational effects ...............................................................................14 1.1.1 The evolutionary significance of transgenerational effects ......................................14 1.1.2 Mechanisms of transgenerational effects ..................................................................19 1.2 Insect immunity ...........................................................................................................21 1.3 Transgenerational immune priming .............................................................................25 1.4 The biology of Plodia interpunctella ...........................................................................27 1.5 Life history traits in capital breeding Lepidoptera .......................................................29 1.6 Aims of thesis ..............................................................................................................30 Chapter 2: Methods: The development of a synthetic diet for investigating the effects of macronutrients on the development of Plodia interpunctella 2.1 Maintenance of the stock population at QMUL ..........................................................31 2.2 Artificial diets ..............................................................................................................32 2.2.1 Development of artificial diets..................................................................................34 2.2.2 Essential components of insect diets.........................................................................35 2.3 Diet development using pilot experiments ..................................................................38 2.3.1 Pilot 1: using desiccation as a method of preservation .............................................38 2.3.2 Pilot 2: using gel format diets ..................................................................................40 2.3.3 Pilot 3: low nutrients diets .......................................................................................42 8 2.4 Final method for creating artificial diet .......................................................................43 2.5 Effect of artificial diet on longevity .............................................................................45
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