Dietary Tryptophan Requirements in Adult Dogs
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Dietary Tryptophan Requirements in Adult Dogs and How Optimization of the Tryptophan to Large Neutral Amino Acid Ratio Affects the Metabolism and Gut Health in Actively Training Sled Dogs. by James Russell Templeman A Thesis presented to The University of Guelph In partial fulfilment of requirements for the degree of Doctor of Philosophy in Animal Biosciences Guelph, Ontario, Canada © James R. Templeman, August 2020 ABSTRACT DIETARY TRYPTOPHAN REQUIREMENTS IN ADULT DOGS AND HOW OPTIMIZATION OF THE TRYPTOPHAN TO LARGE NEUTRAL AMINO ACID RATIO AFFECTS THE METABOLISM AND GUT HEALTH IN ACTIVELY TRAINING SLED DOGS James R. Templeman Advisor: University of Guelph, 2020 Dr. Anna-Kate Shoveller Exercise improves the health of dogs; although, extreme exertion experienced by sled dogs may lead to variable metabolic and gastrointestinal outcomes. Nutritional solutions can reduce the prevalence of these disturbances, but a deeper understanding of sled dog nutritional programs is warranted to ensure appropriate gaps in research are targeted. Supplementation of tryptophan, an indispensable amino acid for dogs, has been explored as a means to increase serotonin, a neurotransmitter involved in central and enteric nervous system function; however, there are no empirical data to support current regulatory recommendations of dietary tryptophan for adult dogs. The objectives of this thesis were to 1) critically review aspects of sporting dog nutrition, methods of estimating amino acid requirements, and tryptophan metabolism, 2) evaluate current sled dog nutritional programs, 3) estimate tryptophan requirements in adult dogs, and 4) apply those estimations via optimizing the dietary tryptophan to large neutral amino acid ratio for actively training sled dogs, with the goal of improving metabolic and gastrointestinal outcomes. Survey- based data pertaining to sled dog nutritional management revealed the need for improvements in nutrient provision, particularly protein, in order to reduce the perceived necessity of (and risks associated with) raw meat supplementation and formulation of homemade diets without scientific or clinical consideration. Using indicator amino acid oxidation, tryptophan requirements were estimated for small (miniature dachshunds), medium (beagles), and large breed (Labrador retrievers) dogs, and all estimates were higher than current regulatory recommendations. Effects of supplemental dietary tryptophan and incremental exercise on metabolic and gastrointestinal outcomes were then investigated using Siberian huskies and a diet formulated to achieve the highest tryptophan to large neutral amino acid ratio used when feeding adult dogs to their tryptophan requirements. Tryptophan supplementation improved serotonin status and fecal quality without affecting body composition in actively training dogs, demonstrating the potential benefits of tryptophan supplementation. Specifically, tryptophan supplementation may help to support gastrointestinal function in sporting dogs under stressful conditions (e.g. intensive exercise) without compromising athletic performance. Further research is warranted to investigate effects of secondary metabolic pathways and factors that may increase metabolic demand on indispensable amino acid requirements and supplementation. iv ACKNOWLEDGEMENTS When I moved nearly four thousand kilometers from my home in Heffley Creek, British Columbia, to Guelph, Ontario, in the fall of 2010, to say that I was naïve was a severe understatement. At the time, and in classic freshman fashion, I really had no idea what I wanted in life, all I knew was that in one way or another, I wanted to work with animals. I can say quite confidently though, that the idea of graduate school was nowhere near my radar. In the beginning, being not nearly the extrovert I am today, my circle was very small; however, as I sit here now nearly 10 years later, I find it hard to even fathom trying to account for all the people who have affected my life and the path I have taken to get here. If you do not find your name among those listed below, please do not interpret that as a reflection of the influence you have had on me. I cannot go any further without acknowledging Rachel Dobberstein, who has undoubtedly had a more significant and positive impact on my life than I thought would have been possible. I still do not know how I built the confidence to talk a girl who was, and still is, miles out of my league, but with the utmost certainty, I can admit that it was the best decision I have ever made. Saying that you have made me a better person is massively clichéd, but it is undeniably true, and anyone who knew me before we met can attest to that. I am not the easiest person to deal with, to say the least, I can be stubborn and cynical at times, but you have supported me through thick and thin. Most importantly though, your confidence in me is never greater than at the times when I lose belief in myself. These last six years have been challenging, but incredibly rewarding, and there is nobody I would have rather had by my side throughout it all than you. I have spent nearly seven years in the Department of Animal Biosciences, and every year the list of faculty members and graduate students that made significant impacts on my life has grown considerably, and I am grateful to have had the privilege of working with and learning from each and every one of you. To start, I must acknowledge Dave Seymour, Linaya Pot, and Natalie Asaro, the original inhabitants of office 229, aka ‘The Frat House’. We have had a lot of great times together, culminating at ASAS in Vancouver in the summer of 2018, and I consider myself incredibly fortunate to have become, and remained to this day, very close friends with you all. Wilfredo ‘Danny’ Mansilla, where do I even start. Over what was, unfortunately, a relatively short period of time, you quickly became someone I deeply admired. The combination of your staggering intelligence and down-to-earth personality is inspiring; the world needs more people v like you, and I am grateful for the time I had to learn from you. While I know I could never come close to filling your shoes as a Post Doc in the Shoveller lab, I can only hope to have the same impact on an emerging graduate student as you had on me. Vern Osborne and John Cant, you have both mentored me from day one and have continuously provided me with guidance, encouragement, and sound advice on many aspects of life. You showed me how important life outside of the office can be and encouraged me to think about things from a different perspective. I also know that without you two acting as liaisons, I would not have had the chance to meet Anna- Kate Shoveller, and my life would be very different right now. Kate, you took a chance on me after little-more than an hour-long talk over a couple of pints and a few words of support from Vern and John, and because of that I am sincerely thankful. Your contribution to my academic career is unmatched, but it is your dedication to making me an honest and well-rounded individual that is truly invaluable. Something just seems to work between us. I feel comfortable around you, but at the same time I always feel driven to exceed your expectations. I honestly did not think I would ever meet someone who is as competitive as me, or who understands the value of harnessing that competitive nature, but you proved me very wrong. I have the utmost respect for the impacts you have made and the success you have had in industry and academia, and the fact you still work tirelessly to improve yourself and those around you is inspirational. I could not imagine having a better role model in my life, and I hope that we will always get the opportunity to work together. Along with Kate, I would like to thank my advisory committee (John Cant and David Wright), as well as my qualifying examination committee (John Cant, David Wright, Jim Atkinson, and Lindsay Robinson). You all helped push me to my limits and encouraged me to pursue ideas or opportunities that were far out of my comfort zone. I would also like to thank Jim for introducing me to companion animal nutrition nearly 8 years ago, and for providing the support and gentle nudge needed for me to take a step into undergraduate teaching. Jim’s immense knowledge and charismatic approach to lecturing (or really, any situation) drew me in instantly, and I strive to model my teaching style after his. Last, but certainty not least, I would like to recognize my family. There have been times when being so far away from them has been very difficult, the most recent and burdensome example of this has been my inability to experience my nieces growing up. At times, this has vi weighed heavily on my mind and caused me to question why exactly I am doing this, but throughout all these years, knowing that I have always had their unconditional support has driven me to carry on. I would not be the person I am today, or where I am today, without their unwavering encouragement and positivity, and I am eternally grateful. vii TABLE OF CONTENTS Abstract ........................................................................................................................................... ii Acknowledgements ........................................................................................................................ iv Table of Contents .......................................................................................................................... vii List of Tables ................................................................................................................................