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Investigating the Thermal Biology of Round Gobies by Justin Mark

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Investigating the Thermal Biology of Round Gobies by Justin Mark Behavioural Thermoregulation and Escape Behaviour: Investigating the Thermal Biology of Round Gobies By Justin Mark Bridgeman Bachelor of Science, 2014, Brock University Submitted in partial fulfillment of the requirements for the degree of Master of Science Brock University, Faculty of Mathematics and Science Department of Biological Sciences St. Catharines, Ontario © 2019 Abstract The invasive round goby (Neogobius melanostomus) has successfully colonized all of the Great Lakes since its discovery in the region in 1991, yet little is known about its thermal biology. The focus of this thesis was to examine the effect of acclimation to unseasonably warm temperatures on round goby behavioural thermoregulation, as well as behavioural and physiological performance during escapes with warm acute temperatures. Juvenile gobies were acclimated to either 21°C or 24°C for each set of experiments. I first examined goby thermal preference in a shuttlebox through their ability to escape from unfavourable temperatures. I found that escape temperatures were plastic following acclimation to a rise in 3°C rise in temperature (+3°C) and associated positively with acclimation temperature, even though gobies showed slightly lower-than-expected average escape temperatures in each acclimation treatment. Interestingly, acclimation to +3°C leads to lower exploratory behaviour in warm waters and lower overall activity levels during behavioural thermoregulation. In risky situations involving threat of predation, exploratory behaviour is often linked to boldness. Next I investigated exploratory swimming through two behavioural traits: ability to voluntarily enter a tunnel and subsequent swimming activity while being chased in a detour task. Detour tasks require a fish to swim down a narrow space and then detour to the left or right as they approach a barrier. A strong correlation between behavioural traits suggests the two behavioural traits are a good measure of risky swimming behaviour. With respect to the impact of warm acute temperatures, I observed an increase in strength of risky swimming behaviour following a rise in acute temperature of 6°C (+6°C). Finally, I examined burst swimming performance (C- and S-starts) during the detour task. While there was no observed effect of warm acute temperatures on most C- and S-start performance measures, there was a significant increase in S-start time to maximum velocity at +6°C. Along with the observed plasticity in strength of risky swimming behaviour, my ii findings showed that acclimation to +3°C shifted physiological performance of escapes. My findings provide a greater understanding of round goby physiological and behavioural functioning during short- and long-term rises in temperature. Keywords: Round goby; Temperature; Behavioural thermoregulation; Burst swimming; Risky swimming behaviour iii Acknowledgements I would like to give thanks to my supervisor, Dr. Glenn Tattersall, for his support and encouragement throughout my time in the Tattersall Lab. I am grateful for the many opportunities I have had to collaborate on various research projects, and it is my hope that we can continue to do so in the future. There have been many people that have helped me along the way. I would like to thank all members of my research committee for their input throughout this process. Also, thank you to all current and past members in the Tattersall Lab, including: Ian Black, Curtis Abney, Philip Bartel, Susan Wang, and Kristen Schuppe. A special thanks to Anne Yagi and the MNRF for their expertise in the collection of gobies in the field, as well as Tom Eles for always being available to answer questions regarding animal care. Finally, I would like to acknowledge my fiancée and my family for all the help they provided along the way. I would not be where I am without them. To my late grandmother, Eileen Tomlinson: you have made such a huge impact on my life and I would like to thank you for all your words of wisdom. You inspired me to believe in myself and have the courage to achieve whatever I wanted in life. Thank you for everything. iv Table of Contents Abstract………………………………………………………………………………………………… ii Acknowledgements……………………………………………………………………………..…….. iv Table of Contents………………………………………………………………………………………. v List of Tables………………………………………………………………………………...………. viii List of Figures…………………………………………………………………………………………. x Chapter 1: General Introduction………………………………………………………………………………..... 1 Background…………………………………………………………………………………………….. 1 Thermal Optimality of Performance Traits…………………………………………………………..... 2 Behavioural Thermoregulation………………………………………………………………………… 4 Shuttleboxes…………………………………………………………………………………………..... 6 Thermal Acclimation………………………………………………………………………………….. .7 Escape Responses…………………………………………………………………………………….. ..8 Animal Personality - Boldness………………………………………………………………………... 12 Round Gobies…………………………………………………………………………………………. 14 Thesis Rational and Objectives………………………………………………………………………. 15 References……………………………………………………………………………………………..18 Chapter 2: Temperature avoidance behaviour in the invasive round goby: impact of warm acclimation on swimming activity during behavioural thermoregulation………………………………………... 25 2.1 Introduction……………………………………………………………………………………….. 25 2.2 Methodology………………………………………………………………………………………28 Subjects……………………………………………………………………………………………….. 28 Apparatus……………………………………………………………………………………………..29 Experimental Design…………………………………………………………………………………. 31 Data Analysis………………………………………………………………………………………….33 Statistical Methodology………………………………………………………………………………. 33 2.3 Results…………………………………………………………………………………………….. 35 Shuttles………………………………………………………………………………………………... 35 Ratio of Distance Swam Between Chambers………………………………………………………….35 Escape Temperature………………………………………………………………………………….. 36 2.4 Discussion………………………………………………………………………………………… 47 Conclusions and Perspectives…………………………………………………………………………49 References……………………………………………………………………………………………..51 v Chapter 3: Assessing the effects of short- and long-term temperature change on round goby fast-start performance…………………………………………………………………………………………. 54 3.1 Introduction……………………………………………………………………………………….. 54 3.2 Methodology……………………………………………………………………………………… 60 Subjects………………………………………………………………………………………………..60 Apparatus……………………………………………………………………………………………... 62 Experimental Design…………………………………………………………………………………. 64 Analysis of Fast-start Maximum Velocity and Time to Maximum Velocity…………………………...65 Velocity and Acceleration Measurements……………………………………………………………. 68 Statistical Analysis…………………………………………………………………………………….69 Differences in Performance between Fast-start Types at Acclimation Temperature…………….. 69 Effect of Warm Acute Temperatures on Fast-Start Performance…………………………………70 3.3 Results…………………………………………………………………………………………… .70 Direct Comparison Between C- and S-start Performance…………………………………………….70 Effect of Constant Temperatures Across Days on Fast-start Performance…………………………..71 Effect of Impact of Warm Acute Temperatures on Fast-start Performance………………………….. 71 3.4 Discussion………………………………………………………………………………………… 83 Impact of Thermal Acclimation and Differences in Performance between C- and S-starts…………. 83 Impact of Warm Acute Temperatures on Fast-start Performance…………………………………….84 Conclusion……………………………………………………………………………………………. 85 References…………………………………………………………………………..…………………87 Chapter 4: An Investigation of Exploratory Behaviour While Being Chased as a Measure of Risky Swimming Behaviour During a Detour Task………………………………………………………91 4.1 Introduction………………………………………………………………………………………. 91 4.2 Methodology………………………….…………………………………………………………...97 Subjects……………………………………………………………………………………………….97 Experimental Procedure………………………………………………………………………………97 Analysis of Fast-start Presence……………………………………………………………….……...101 Statistical Analysis…………………………………………………………………………….……..101 Relationship between Behavioural Traits and S-start Presence………………………..….……101 Impact of Warm Thermal Acclimation and Acute Temperatures on Strength of Behavioural Traits………………………………………………………………………..…...102 4.3 Results…………………………………………………………………………………………...103 Relationship between Behavioural Traits and S-start Presence…………………………………….103 Impact of Warm Thermal Acclimation and Acute Temperatures on Strength of Behavioural Traits………………………………………………………………………………...104 vi 4.4 Discussion………………………………………………………………………………………. 116 Exploratory and Risk-Taking Behaviour During the Detour Task………………………………….. 116 Occurrence of S-starts as a Measure of Freezing Behaviour……………………………………….. 118 Impact of Warm Acute Temperatures on Behavioural Traits…………..…………………………... 120 Conclusion…………………………………………………………………………………………... 121 References……………………………………………………………………………………………122 Chapter 5: Summary and Future Directions………………………………………………………………….. 126 References…………………………………………………………………………………………... 132 Appendix……………………………………………………………………………………………. 136 vii List of Tables Table 2.1: Results for the F-tests and Type II Wald Chi-square tests examining the effect of cohort on round goby escape temperature (ET), Shuttle Rate, and Ratio of Distance Swam Between Chambers during both ramping and adjustment periods…………………………... 41 Table 2.2: Results for the Type II Wald Chi-square tests examining the effect of warm thermal acclimation and Length on the number of overall shuttles (Shuttles) during the adjustment and ramping periods……………………………………………………………..42 Table 2.3: Linear model parameter estimates examining effect of warm thermal acclimation on the number of overall shuttles (Shuttles) for 21°C-acc and 24°C-acc round gobies during the adjustment
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