Thesis Ebi Antony George
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Individual and social mechanisms regulating the dance activity within honey bee forager groups A Thesis Submitted to the Tata Institute of Fundamental Research, Mumbai for the degree of Doctor of Philosophy in Biology by Ebi Antony George National Centre for Biological Sciences, Bangalore Tata Institute of Fundamental Research, Mumbai October 2019 Declaration Date: 11 October 2019 iii Certificate v Publications George EA, Brockmann A. 2019 Social modulation of individual differences in dance communication in honey bees. Behav. Ecol. Sociobiol. 73, 41. (doi:10.1007/s00265-019- 2649-0) Content from the above publication has been incorporated in the thesis with permission from the publisher (License no: 4570571255954). George EA, Bröger A-K, Thamm M, Brockmann A, Scheiner R. 2019 Inter-individual variation in honey bee dance intensity correlates with expression of the foraging gene. Genes, Brain and Behavior. 1– 11. (doi:10.1111/gbb.12592) Content from the above publication has been incorporated in the thesis with permission from the publisher (License no: 4675421240663). vii Acknowledgements Charles Darwin rightly said that “it is the long history of humankind (and animal kind, too) that those who learned to collaborate and improvise most effectively have prevailed”. These words highlight an interesting parallel between the paradigm I studied (the foraging activity of the honey bee colony) and my own experience during my PhD at the National Centre for Biological Sciences, Bangalore. In both cases, any successful endeavour depended on the interaction between a diverse group of individuals, each supporting the other. Even though it is my name on the first page of this thesis, I am part of a multitude of people who have all contributed academically and otherwise to this document. I will attempt to enumerate the contributions of all these individuals below, even though space constraints will limit how much I can expound on their wonderful qualities. First and foremost, I am deeply indebted to Dr. Axel Brockmann, mentor par excellence. When I wanted to take my project in a direction that was not a focus of the lab, he was still supportive of my ideas. From finding conferences that I should attend to potential labs that I should join, Axel has always been encouraging of my career as a scientist. The frankness of his criticism and his guidance with respect to framing questions has been instrumental in shaping my thought process. I could not have asked for a better supervisor in my first major stint in academia. I would also like to thank my thesis committee members, Dr. Raghavendra Gadagkar and Dr. Uma Ramakrishnan, for all the thoughtful inputs they have provided. Dr. Raghavendra was instrumental in pointing me in the right direction with respect to hypothesis driven experiments at the very early stages of my project. Dr. Uma helped me broaden the project quite a bit, by making sure that I could always envision what the next step would be. I am also grateful to Dr. Upinder Bhalla who was part of my comprehensive examination committee. He made sure that I had the time and space to get out of my comfort zone and include a new dimension to the project, which has culminated in one whole chapter dedicated to modelling in my thesis. Dr. Ricarda Scheiner graciously hosted me for three months at the University of Würzburg, Germany. She has been extremely supportive, during the experiments that I did in Germany, as well as during the preparation of our recently published manuscript. I am also grateful to Ann-Kathrin Bröger and Markus Thamm, who helped me with the behavioural experiments and did the molecular work respectively, for the collaborative project. My stay in Germany would not have been possible without the help of Dr. Stephan Schröder-Köhne, who helped me through all the formalities part of the DAAD fellowship Of the multiple courses that I took throughout my stay at NCBS, a few of the instructors deserve special mention. My statistical and modelling knowledge is entirely built around two fantastic courses in IISc by Dr. Kavita Isvaran and Dr. ix Acknowledgements Vishwesha Guttal respectively. Dr. Rohini Balakrishnan and Dr. Maria Thaker also gave me a firm footing in animal behaviour through their course at IISc. This was especially helpful as I had been trained in biotechnology in my bachelors and had only a rudimentary grasp of behavioural studies. Dr. Geoff Hyde also deserves special mention for his wonderful research writing course. My experiments would not have been so well done without the help of a whole platoon of summer interns, masters students and project assistants. I would like to specially thank Ravi Kumar Boyapatti, Hinal Kharva and Abhishek Anand. They have helped me do most of the experiments that I describe in the first two chapters of my thesis. I am also greatly indebted to Neethu Thulasi and Patrick Kohl. Their enthusiasm for Asian honey bees led me to join them on several projects exploring the waggle dance behaviour in these native species. Supraja Rajagopal also deserves a special mention as one of the best summer interns I have had the pleasure to work with. She was instrumental in setting up the modelling aspect of my project and has continued to work on it after her internship. All the members of the Bee lab (past and present) have played important roles in shaping this thesis. The productive lab meets have helped me understand ideas in fields that I have no knowledge about. This in turn has helped me translate ideas in my fields to their projects and vice versa. I’m grateful for the opportunity to look at and analyse data from their experiments as it helped to expand my own analysis. Arumoy Chatterjee has been a great colleague and friend, always up for discussions at any time of the day. A lot of the ideas that I discuss in this thesis has stemmed from my discussions with him. Friends, both off and on campus, have been a great support during my Ph.D. I am especially thankful to Ankit and Shayan, for keeping in constant touch and for the much needed annual road trips. Harsh Ranjan has also been an extremely understanding friend, and my conversations with him have continuously reaffirmed my decision to continue in academia. I am also deeply indebted to Jaya Srivastava and Rahul Ghosh. Jaya has been an inspiration, as she transitioned away from a corporate job to academia. Rahul has always been just a call away for a meet-up in Bangalore and this has been one of the best stress-busters that I have had. I have also had the pleasure to have a great batch here at NCBS. I am eternally grateful to Pritha Ghosh, Rohit Suratekar, Jyothi V Nair, Snehal Karpe, Sreekrishna Varma and Chaitra Prabhakaran. I have had so much fun with these guys, and will forever cherish the memories of our various gatherings and board game nights. Sruthi Unnikrishnan has played a vital role in this document in more ways than one. She has been constantly giving me feedback on my manuscripts, as well as all the chapters in this thesis. Discussing science with her is extremely fruitful and has helped me focus my thoughts on various topics. She has also been a great friend, always ready to support me in any way possible. Sahana Sitaraman has been the single most important source of joy in my life for the past few years. She has always been supportive in all my endeavours. At the same time, x Acknowledgements she has kept me focussed on my project when my attention wandered too much. Her infectious enthusiasm for the project helped me tide over times when the work was too slow. She has also offered constant guidance and insights with respect to the various academic and non-academic problems that I have faced throughout the last few years. My completion of this document was dependent on her support and care to a great extent. Finally, none of this would have been possible without the help and support of my family and extended family. My extended family has always been curious about the research I do, and explaining my experiments to them has been a great learning experience. My family has been immensely supportive of my career decisions from the very beginning. They have been a source of inspiration and joy. My sister, Amala Mary George, showed me the value of persistence. She has strived hard to get to be teacher while juggling her research interests. My mother, Joysy George, helped me see that determination and hard work will always matter at the end, irrespective of the task at hand. My father, George Paul, has inspired me by his enthusiasm to continuously learn new things. He takes courses to this day and has taught me that there is no limit to what a person can learn nor the age at which a person can learn. I dedicate this document to their steadfast love and support, without which I would not have set out on this path. xi xii Table of Contents Table of Contents Declaration ......................................................................................................... iii Certificate .............................................................................................................. v Publications ........................................................................................................ vii Acknowledgements .............................................................................................. ix Table of Contents .............................................................................................. xiii Chapter 1 – Introduction ......................................................................................