survival of the fittest: florida keys vs flower garden banks...

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Survival of the Fittest: Florida Keys vs Flower Garden Banks coral, Montastraea cavernosa Joshua A. Haslun 1 , Kevin B. Strychar 1* , Paul W. Sammarco 2 , Gregory Buck 1 , Joanna Mott 1 , and James M. Cervino 3 1) Dept. of Life Sciences, Texas A&M University – Corpus Christi, Corpus Christi, TX; 2) Louisiana Universities Marine Consortium (LUMCON), Chauvin, LA; 3) Woods Hole Oceanographic Institution, Woods Hole, MA Fig 1. Hemocytometry data showing the mean temporal loss of viable (A 1 ,B 1 ), necrotic (A 2 ,B 2 ), and mitotic (A 3 ,B 3 ) zooxanthellae cells (× 10 3 mL -1 ) expelled from Montastraea cavernosa collected from Flower Garden Banks (FGB) and the Florida Keys (FKs). Error bars represent 95% confidence intervals (n i = 8); some error bars are too small to be seen. This research is supported by: For more information, contact *Kevin B. Strychar at [email protected] Introduction Mean global sea-water temperatures (SWTs) have increased by ~0.5°C over the past century and are expected to continue to rise over the next several decades (IPCC, 2007). There have been numerous studies in the Caribbean on the causes and mechanisms of coral bleaching (e.g., Baker et al., 1997; Oppen and Lough, 2009) including the probability that some corals can “cope” with heat stress better than their symbionts (Sammarco and Strychar, 2009), and that this may be related to their evolutionary age and/or their prior experiences to specific stressors (Reich and Hickey, 2005; Florida DEP, 2009). Here, we ask whether corals in the Florida Keys (FKs) are better adapted for long-term survival vs. those in the Flower Garden Banks (FGB) because of prior exposure to such stresses (e.g., bleaching) through time, in terms of frequency and level. Climate change is causing corals world-wide to bleach. We hypothesize that coral exposed to short durations of repetitive events of high heat stress are better adapted than corals lacking similar experiences. In this study, we compare populations of Montastraea cavernosa (Linnaeus, 1767) from the Florida Keys (FKs) to coral from the Flower Garden Banks (FGB). Coral from the FKs are known to experience frequent annual events of high temperatures whilst coral from the FGB have had fewer exposures to anomalous temperatures. Our data show that coral from the FKs lost significantly (ANOVA; p < 0.01) fewer viable, dead, and/or necrotic symbiont cells across all experimental temperatures than coral from the FGB. The mean percentage of viable expelled cells observed at 35ºC was significantly lower than those at 27ºC and 31ºC (p < 0.05, Games-Hollow post-hoc tests). The mean percentage of necrotic cells lost was greatest at 35ºC (p < 0.01, Games-Hollow post hoc test) culminating in the greatest loss (40%) within 24 h. There was no significant variation in the loss of apoptotic or post- apoptotic (ANOVA; p > 0.05) cells between temperatures or time. Our data indicated that M. cavernosa from FKs may be less susceptible to heat stress and bleaching than coral at the FGB. We speculate that this ‘fitness’ is a regional adaptation likely associated with numerous temperature anomalies. Abstract Results Coral collected from FKs lost significantly (ANOVA; p < 0.01) fewer viable, dead, and/or necrotic symbiont cells across all experimental temperatures than coral from the FGB. The mean percentage of viable expelled cells observed at 35ºC was significantly lower than those at 27ºC and 31ºC (p < 0.05, Games-Hollow post-hoc tests). The mean percentage of necrotic cells expelled was greatest at 35ºC (p < 0.01, Games-Hollow post hoc test) culminating in the greatest loss (40%) within 24 h. There was no significant variation in the loss of apoptotic or post-apoptotic (ANOVA; p > 0.05) cells between temperatures or time. Acknowledgments: We thank the Texas Parks and Wildlife for seawater transportation services, Texas Flower Garden Banks National Marine Sanctuary (Permit Number FGBNMS-2007-010) and the Florida Keys National Marine Sanctuary (Permit Number FKNMS 0648-0141) for permits given KB Strychar. We also thank Texas Research Development Fund for grants to KB Strychar. Special thanks to GP Schmahl and E Hickerson at NOAA (FGBNMS) for ship time and J Delaney and E. Bartels at Mote Marine Lab for collecting coral. Literature Cited: Baker, A.C., Rowan, R., Knowlton, N. 1997. Diversity of symbiotic dinoflagellates (zooxanthellae) in Scleractinian corals of the Caribbean and Eastern Pacific. Proc 8th Int Coral Reef Symp, Panama., 2, 1301–1306. Florida DEP (Department of Environmental Protection), 2009. Florida’s coral reefs, online. Last accessed May 6, 2010. Available at http://www.dep.state.fl.us/coastal/habitats/coral.htm and http://www.dep.state.fl.us/coastal/reports/AA/AA Project 2 MediaKits. IPCC (Intergovernmental Panel on Climate Change). 2007: Summary for policymakers. In: Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change [Solomon, S., D. Qin, M. Manning, Z. Chen, M. Marquis, K.B. Averyt, M.Tignor and H.L. Miller (eds.)]. Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA. Oppen, M.J.H, Lough J.M. 2009. Coral bleaching: Patterns, processes, causes and consequences. Springer, 178 p. Reich C, Hickey D. 2005. Coral coring in the Flower Garden Banks National Marine Sanctuary – a collaborative effort. USGS SoundWaves. 74:1-13. Sammarco, P.W., Strychar, K.B. 2009. Effects of climate change and global warming on coral reefs: Exaptation in corals and the evolution of their zooxanthellae. Environ Bioind 4:9-45. Schmahl, G.P., Hickerson, E.L., Precht, W.F., 2008. Biology and ecology of coral reefs and coral communities in the Flower Garden Banks Region, Northwestern Gulf of Mexico. Coral reefs of the world: Coral reefs of the USA. 221-261. A B C Fig 2. Hemocytometry data showing the mean cumulative loss of viable (A 1 ,B 1 ), necrotic (A 2 ,B 2 ), and mitotic (A 3 ,B 3 ) zooxanthellae cells (× 10 3 mL -1 ) expelled from Montastraea cavernosa collected from Flower Garden Banks (FGB) and the Florida Keys (FKs). Error bars represent 95% confidence intervals (n i = 8); some error bars are too small to be seen. Note how cumulative graphs illustrate the effect of temperature on overall depletion of a coral’s Symbiodinium complement over 72 h. Fig 3. Flow cytometry data showing mean temporal loss expressed as percent (%) of viable (A 1 ,B 1 ), necrotic (A 2 ,B 2 ), apoptotic (A 3 ,B 3 ), and post-apoptotic (A 4 ,B 4 ) Symbiodinium cells expelled from Montastraea cavernosa collected from Flower Garden Banks (FGB) and the Florida Keys (FKs). Error bars represent 95% confidence intervals (n i = 8); some error bars are too small to be seen. Conclusion Montastraea cavernosa from Florida Keys (FKs) may be less susceptible to heat stress and bleaching than coral at the Flower Garden Banks (FGB). We speculate that this ‘fitness’ is a regional adaptation possibly associated with numerous temperature anomalies. Montastraea cavernosa Flow-through system Flow cytometer Considered one of the more susceptible species to thermal stress (Schmahl et al., 2008), Montastraea cavernosa (A) samples were collected from FKs and FGB and transported to Texas A&M University – Corpus Christi. A flow through sea water system (B) delivered filtered oxygenated seawater at a constant flow rate of 12.5 mL min -1 to individual incubation containers (IC), each housed within a thermally controlled water jacket. Each IC contained one coral fragment ~2.5 cm 2 . Eight coral replicates at each of three temperatures (27ºC, 31ºC, and 35ºC) were examined for each site (FCs and FGB); two experimental replicates were conducted for each site. Seawater samples (50 mL) were obtained every 6 h over 72 h from each IC and centrifuged to concentrate the samples into two 1 mL aliquots. One aliquot was used for light microscopy to determine zooxanthellar density, mitotic index, and viability. The second aliquot was examined using flow cytometry (C) to differentiate between four cell death categories: viable, necrotic, apoptotic, and post-apoptotic. Statistical analysis consisted of a Model I, repeated-measures, three-way orthogonal experimental design. All data were tested for significant variation due to sphericity using the Greenhouse-Guyser or the Huynh-Feldt correction. Since the data lacked sphericity, Tukey’s and Games-Hollow post hoc tests were applied after analysis via GLM repeated measure ANOVAs to examine differences between individual means. Data displaying significant variations over time were Bonferroni-corrected. Methods

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Page 1: Survival of the Fittest: Florida Keys vs Flower Garden Banks ...conference.ifas.ufl.edu/floridakeys/Presentations...Survival of the Fittest: Florida Keys vs Flower Garden Banks coral,

Survival of the Fittest: Florida Keys vs Flower Garden Banks coral, Montastraea cavernosa

Joshua A. Haslun1, Kevin B. Strychar1*, Paul W. Sammarco2, Gregory Buck1, Joanna Mott1, and James M. Cervino3

1) Dept. of Life Sciences, Texas A&M University – Corpus Christi, Corpus Christi, TX; 2) Louisiana Universities Marine Consortium (LUMCON), Chauvin, LA; 3) Woods Hole Oceanographic Institution, Woods Hole, MA

Fig 1. Hemocytometry data showing the mean temporal lossof viable (A1, B1), necrotic (A2, B2), and mitotic (A3, B3)zooxanthellae cells (× 103 mL-1) expelled from Montastraeacavernosa collected from Flower Garden Banks (FGB) andthe Florida Keys (FKs). Error bars represent 95% confidenceintervals (ni = 8); some error bars are too small to be seen.

This research is supported by: For more information, contact *Kevin B. Strychar at

[email protected]

IntroductionMean global sea-water temperatures (SWTs) have increased by~0.5°C over the past century and are expected to continue to riseover the next several decades (IPCC, 2007).

There have been numerous studies in the Caribbean on the causesand mechanisms of coral bleaching (e.g., Baker et al., 1997; Oppenand Lough, 2009) including the probability that some corals can“cope” with heat stress better than their symbionts (Sammarco andStrychar, 2009), and that this may be related to their evolutionary ageand/or their prior experiences to specific stressors (Reich andHickey, 2005; Florida DEP, 2009). Here, we ask whether corals in theFlorida Keys (FKs) are better adapted for long-term survival vs.those in the Flower Garden Banks (FGB) because of prior exposureto such stresses (e.g., bleaching) through time, in terms offrequency and level.

Climate change is causing corals world-wide to bleach. Wehypothesize that coral exposed to short durations of repetitiveevents of high heat stress are better adapted than corals lackingsimilar experiences. In this study, we compare populations ofMontastraea cavernosa (Linnaeus, 1767) from the Florida Keys (FKs)to coral from the Flower Garden Banks (FGB). Coral from the FKsare known to experience frequent annual events of hightemperatures whilst coral from the FGB have had fewer exposuresto anomalous temperatures. Our data show that coral from the FKslost significantly (ANOVA; p < 0.01) fewer viable, dead, and/ornecrotic symbiont cells across all experimental temperatures thancoral from the FGB. The mean percentage of viable expelled cellsobserved at 35ºC was significantly lower than those at 27ºC and31ºC (p < 0.05, Games-Hollow post-hoc tests). The mean percentageof necrotic cells lost was greatest at 35ºC (p < 0.01, Games-Hollowpost hoc test) culminating in the greatest loss (40%) within 24 h.There was no significant variation in the loss of apoptotic or post-apoptotic (ANOVA; p > 0.05) cells between temperatures or time.Our data indicated that M. cavernosa from FKs may be lesssusceptible to heat stress and bleaching than coral at the FGB. Wespeculate that this ‘fitness’ is a regional adaptation likely associatedwith numerous temperature anomalies.

Abstract

ResultsCoral collected from FKs lost significantly (ANOVA; p < 0.01) fewer viable, dead, and/or necrotic symbiont cellsacross all experimental temperatures than coral from the FGB. The mean percentage of viable expelled cellsobserved at 35ºC was significantly lower than those at 27ºC and 31ºC (p < 0.05, Games-Hollow post-hoc tests). Themean percentage of necrotic cells expelled was greatest at 35ºC (p < 0.01, Games-Hollow post hoc test) culminatingin the greatest loss (40%) within 24 h. There was no significant variation in the loss of apoptotic or post-apoptotic(ANOVA; p > 0.05) cells between temperatures or time.

Acknowledgments: We thank the Texas Parks and Wildlife for seawater transportation services, Texas Flower Garden Banks National Marine Sanctuary (Permit Number FGBNMS-2007-010) and the Florida Keys National Marine Sanctuary (Permit Number FKNMS 0648-0141) for permits given KB Strychar. We also thank Texas Research Development Fund for grants to KB Strychar. Special thanks to GP Schmahl and E Hickerson at NOAA (FGBNMS) for ship time and J Delaney and E. Bartels at Mote Marine Lab for collecting coral. Literature Cited: Baker, A.C., Rowan, R., Knowlton, N. 1997. Diversity of symbiotic dinoflagellates (zooxanthellae) in Scleractinian corals of the Caribbean and Eastern Pacific. Proc 8th Int Coral Reef Symp, Panama., 2, 1301–1306. Florida DEP (Department of Environmental Protection), 2009. Florida’s coral reefs, online. Last accessed May 6, 2010. Available at http://www.dep.state.fl.us/coastal/habitats/coral.htm and http://www.dep.state.fl.us/coastal/reports/AA/AA Project 2 MediaKits. IPCC (Intergovernmental Panel on Climate Change). 2007: Summary for policymakers. In: Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change [Solomon, S., D. Qin, M. Manning, Z. Chen, M. Marquis, K.B. Averyt, M.Tignor and H.L. Miller (eds.)]. Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA. Oppen, M.J.H, Lough J.M. 2009. Coral bleaching: Patterns, processes, causes and consequences. Springer, 178 p. Reich C, Hickey D. 2005. Coral coring in the Flower Garden Banks National Marine Sanctuary – a collaborative effort. USGS SoundWaves. 74:1-13. Sammarco, P.W., Strychar, K.B. 2009. Effects of climate change and global warming on coral reefs: Exaptation in corals and the evolution of their zooxanthellae. Environ Bioind 4:9-45. Schmahl, G.P., Hickerson, E.L., Precht, W.F., 2008. Biology and ecology of coral reefs and coral communities in the Flower Garden Banks Region, Northwestern Gulf of Mexico. Coral reefs of the world: Coral reefs of the USA. 221-261.

A

B

C

Fig 2. Hemocytometry data showing the mean cumulative loss ofviable (A1, B1), necrotic (A2, B2), and mitotic (A3, B3) zooxanthellaecells (× 103 mL-1) expelled from Montastraea cavernosa collected fromFlower Garden Banks (FGB) and the Florida Keys (FKs). Error barsrepresent 95% confidence intervals (ni = 8); some error bars are toosmall to be seen. Note how cumulative graphs illustrate the effect oftemperature on overall depletion of a coral’s Symbiodiniumcomplement over 72 h.

Fig 3. Flow cytometry data showing mean temporal lossexpressed as percent (%) of viable (A1, B1), necrotic (A2, B2),apoptotic (A3, B3), and post-apoptotic (A4, B4) Symbiodiniumcells expelled from Montastraea cavernosa collected fromFlower Garden Banks (FGB) and the Florida Keys (FKs). Errorbars represent 95% confidence intervals (ni = 8); some errorbars are too small to be seen.

ConclusionMontastraea cavernosafrom Florida Keys (FKs)may be less susceptibleto heat stress andbleaching than coral atthe Flower Garden Banks(FGB). We speculate thatthis ‘fitness’ is a regionaladaptation possiblyassociated with numeroustemperature anomalies.

Montastraea cavernosa

Flow-through system

Flow cytometer

Considered one of the more susceptible species to thermal stress (Schmahl et al., 2008), Montastraea cavernosa (A)samples were collected from FKs and FGB and transported to Texas A&M University – Corpus Christi.

A flow through sea water system (B) delivered filtered oxygenated seawater at a constant flow rate of 12.5 mL min-1

to individual incubation containers (IC), each housed within a thermally controlled water jacket.

Each IC contained one coral fragment ~2.5 cm2. Eight coral replicates at each of three temperatures (27ºC, 31ºC, and35ºC) were examined for each site (FCs and FGB); two experimental replicates were conducted for each site.Seawater samples (50 mL) were obtained every 6 h over 72 h from each IC and centrifuged to concentrate thesamples into two 1 mL aliquots. One aliquot was used for light microscopy to determine zooxanthellar density,mitotic index, and viability. The second aliquot was examined using flow cytometry (C) to differentiate between fourcell death categories: viable, necrotic, apoptotic, and post-apoptotic.

Statistical analysis consisted of a Model I, repeated-measures, three-way orthogonal experimental design. All datawere tested for significant variation due to sphericity using the Greenhouse-Guyser or the Huynh-Feldt correction.Since the data lacked sphericity, Tukey’s and Games-Hollow post hoc tests were applied after analysis via GLMrepeated measure ANOVAs to examine differences between individual means. Data displaying significant variationsover time were Bonferroni-corrected.

Methods