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Estimating the Heritability of Plasticity of Thermal Tolerance and Its Application in the Restoration of Endangered Caribbean Coral

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Estimating the Heritability of Plasticity of Thermal Tolerance and Its Application in the Restoration of Endangered Caribbean Coral University of North Florida UNF Digital Commons UNF Graduate Theses and Dissertations Student Scholarship 2020 Estimating the Heritability of Plasticity of Thermal Tolerance and its Application in the Restoration of Endangered Caribbean Coral Matz O. Indergard University of North Florida, [email protected] Follow this and additional works at: https://digitalcommons.unf.edu/etd Part of the Evolution Commons, and the Marine Biology Commons Suggested Citation Indergard, Matz O., "Estimating the Heritability of Plasticity of Thermal Tolerance and its Application in the Restoration of Endangered Caribbean Coral" (2020). UNF Graduate Theses and Dissertations. 932. https://digitalcommons.unf.edu/etd/932 This Master's Thesis is brought to you for free and open access by the Student Scholarship at UNF Digital Commons. It has been accepted for inclusion in UNF Graduate Theses and Dissertations by an authorized administrator of UNF Digital Commons. For more information, please contact Digital Projects. © 2020 All Rights Reserved Estimating the Heritability of Plasticity of Thermal Tolerance and its Application in the Restoration of Endangered Caribbean Coral by Matz Indergard A thesis submitted to the Department of Biology In partial fulfillment of the requirements for the degree of Master of Science in Biology University of North Florida College of Arts and Sciences May 2020 Unpublished work © 2018 Matz Indergard i Redacted Redacted 5/8/2020 Redacted TABLE OF CONTENTS Page List of Figures and Tables……………………………………………………...………………...iv Chapter 1 Abstract…………………………………………………………...…………………....1 Introduction………………………………………………………………………………………..2 Materials and Methods…………………………………………………………………………….5 Results……………………………………………………………………………………..……..11 Discussion………………………………………………………………………………………..13 Chapter 2 Abstract……………………………………………………………………….………20 Introduction……………………………………………………………………………..………..21 Materials and Methods…………………………….…………………………………………..…24 Results………………………………………………………………………………….………...27 Discussion………………………………………………………………………………………..28 References………………………………………………………………………………………..35 iii LIST OF FIGURES AND TABLES Page Figure 1 Average Fv/Fm values among four treatments over three timepoints ………………………………………………38 Figure 2 Average hours survived by each coral among four treatments ……………………………………………………..…………..39 Figure 3 Average difference in hours survived under treatment between ambient and heat acclimated fragments for each genotype ……………………………………….40 Figure 4 Regression of the average thermal tolerance of genotypes when acclimated to elevated and ambient temperatures……………………………….......................42 Figure 5 A photograph of a quadrat of Acropora cervicornis arranged in a randomized grid design …………….................................................................43 Figure 6 The average difference in surface area of outplanted coral fragments among three timepoints ………………………………….………….44 Figure 7 Cumulative percent mortality of A. cervicornis, A. palmata, and O. faveolata over 8 months after outplanting………………........................45 Table 1 Seven estimates of narrow sense heritability for plasticity, thermal tolerance at elevated temperature, and thermal tolerance at ambient temperature……………………………………….……….41 iv Chapter 1: Estimating the Heritability of Plasticity of Thermal Tolerance of Acropora cervicornis Abstract Rising surface temperature poses a threat to the world’s coral reefs and has been the cause of many mass bleaching events. Since coral colonies are sessile, survival of individual colonies may require them to acclimate to elevated water temperatures. Phenotypic plasticity, or the ability of an individual to express different phenotypes to survive in different environments, has been suggested as a critical first step that can buy a population time to evolve important adaptations. This chapter investigated two questions related to plasticity in high temperature tolerance in the endangered Caribbean coral, Acropora cervicornis: 1) Are coral colonies capable of acclimating to elevated water temperatures? 2) Do coral genotypes differ in their ability to acclimate to thermal stress (phenotypic plasticity of thermal tolerance)? We subjected A. cervicornis clones to different acclimation temperatures to identify the presence of plasticity in thermal tolerance, and estimated the broad (H2) and narrow-sense (h2) heritability of both thermal tolerance and plasticity of thermal tolerance. The study found that colonies exposed to a previous, short-term heat stress survived a long-term heat stress significantly longer (141 hours) than those that were acclimated at ambient temperatures. The amount of plasticity was found to depend on genotype, but not necessarily to additive effects of genotype, with H2 = 0.19 and h2 = 0. This suggests A. cervicornis lacks the capacity to evolve greater plasticity in thermal tolerance over a short period of time. However, our estimates may underrepresent the true h2 in this population, and there are other factors such as epigenetic response and holobiont composition that may play a significant role in plasticity of thermal tolerance. 1 Introduction Anthropogenic activities have led to increased concentrations of CO2 in the atmosphere, creating a greenhouse effect that is projected to increase global surface temperatures 1-4 oC within the century (Hoegh-Guldberg et al., 2007, IPCC 2014). Increases in surface temperature pose a threat to coral reefs because coral species tend to live at or near their thermal threshold (Jokiel and Coles 1990; Berkelmans & Willis, 1999; Hoegh-Guldberg, 1999). Elevated temperatures are the cause of many mass-bleaching events, where corals expel their symbiotic dinoflagellates as increased heat damages photosystem II and causes spillover of reactive oxygen species into coral tissues (Weis, 2008) sometimes leading to mortality (Glynn, 1983; Porter & Meier, 1992; Blaird & Marshal, 2002). To combat rising surface temperatures, corals must either relocated to cooler waters, acclimate to elevated temperatures, or evolve higher thermal thresholds. Relocation requires motility, which is only possible in a coral’s larval stage and is limited by a variety of oceanographic and physiological factors (Pechenik, 1990; 1987). Evolution requires multiple generations for selection to alter the population, and with one-third of corals facing extinction due to climate change, (Carpenter et al., 2008) evolution may not occur fast enough to stop mass extinction. Acclimation therefore may be the most probable short-term solution to rising surface temperature, as it may function as an initial step that buys time for adaptation to occur (Chevin & Lande, 2010; Snell-Rood et al., 2018). Phenotypic plasticity is a form of acclimation which improves the probability of survival in populations under stressful conditions without evolution (Pigliucci, 2005). Phenotypic plasticity, or the ability of an individual to express different phenotypes to survive in different environments (Ghalambor et al., 2010) is crucial for sessile organisms since it can increase 2 survivability in species incapable of moving to better conditions (Sultan, 1995). Phenotypic plasticity is typically observed in organisms that occupy areas with highly variable environmental conditions and where the ability to quickly respond to environmental changes is favorable (Via & Lande, 1985). Previous studies have shown prior exposure to elevated temperatures improves the survivability of some coral species to future heat stresses (Middlebrook et al., 2008; Oliver & Palumbi 2011; Maynard et al., 2008). It is unknown whether phenotypic plasticity in thermal tolerance is widespread among coral species, or if it is a trait held only by a few. Furthermore, while host genotype significantly influences thermal tolerance (Fitt et al., 2009; Weis, 2010; Csazar et al., 2010, Kenkel et al., 2015), it is unknown whether coral genets differ in their ability to acclimate to temperature changes. If variability in plasticity of thermal tolerance is found among genotypes, restoration managers may choose to propagate specific corals that are more plastic in their tolerance to thermal stress, effectively using assisted evolution to increase the frequency of plastic genotypes and bolstering the resilience of outplanted coral (Van Oppen et al., 2015). Two mechanisms have been hypothesized to explain phenotypic plasticity in coral. First, coral may alter gene expression patterns that result in physiological changes in response to environmental shifts. When experiencing thermal stress, some species increase expression of genes that transcribe antioxidant (Császár et al., 2009), anti-apoptosis, and symbiont-recognition proteins (Bellantuono et al., 2012). Increased production of antioxidants may reduce damage from free-radicals activated by photosystem II. Anti-apoptosis proteins may prolong cell life while under stress, and symbiont recognition proteins may serve to reinforce the bonding of symbionts in the endoderm, reducing the likelihood of bleaching (Bellantuono et al., 2012). 3 Other species have been observed increasing expression of heat-shock and chromoproteins (Seneca et al., 2010). Heat shock proteins help maintain conformation among structural proteins and important enzymes, while chromoproteins assist in absorbing excess solar radiation. Actin and collagen are also upregulated, suggesting that cellular integrity becomes a priority when threatened with excess heat (Kenkel et al., 2013). The same study also discovered the downregulation of genes coding
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