Pteropods are holoplanktonic gastropod molluscs found globally. Although species diversity is greater at lower latitudes, species abundance is greater at temperate and polar latitudes. Declines in pteropod populations have not only been correlated to declines of their major predators, but pteropods have also been used as bioindicators of global environmental changes such as ocean acidification. With high latitude abundances, pteropods provide significant sustenance for species such as the Atlantic salmon in the Atlantic Ocean and Pleuragramma antarcticum in the Southern Ocean. Because pteropods eat phytoplankton and other pteropods, factors that affect pteropod abundance influence many trophic levels. This dissertation explores ecological, physiological and trophodynamic relationships of pteropods when considering the influences of environmental factors observed to be altering the western Antarctic Peninsula's marine ecosystem. Over the last few decades very few studies have reported the distributions of pteropods along the western Antarctic Peninsula, in particular south of the Gerlache Strait. The ecological study provided the first detailed report of the pteropods Spongiobranchaea australis and Clione antarctica along the western Antarctic Peninsula south of the Gerlache Strait, and their local distribution was correlated to the region's major water masses and mesoscale water mass circulation. The physiological study of S. australis and C. antarctica yielded the first account of their metabolism, ratios of oxygen consumed to nitrogen excreted, proximate body composition, primary substrates oxidized, and enzymatic activities along the study's latitudinal gradient; the first report of S. australis' physiology anywhere around Antarctica. The final chapter utilized a comprehensive Ecopath with Ecosim model of the western Antarctic Peninsula's marine ecosystem. The model was used to explore the trophodynamic significance of pteropods within their polar marine ecosystem as well as changes in whole ecosystem trophodynamics by employing various climate change scenarios expected to alter the Peninsula's marine ecosystem over the next 40 years. The sum of these studies provides a foundation for exploring pteropods as bioindicators of environmental change along the western Antarctic Peninsula, a region currently experiencing considerable climate anomalies.

published_or_final_version / Biological Sciences / Master / Master of Philosophy

Harmful algal blooms (HABs) are increasing in frequency and duration worldwide. Karenia brevis, the major toxic dinoflagellate in the Gulf of Mexico, produces potent neurotoxins, known as brevetoxins. For K. brevis, only minor concentrations of brevetoxins are needed to induce toxicity and environmental conditions appear to have the most direct impact on the cellular content of these toxins. A better understanding of K. brevis biology is essential to understand the mechanisms underlying toxin production and the ecology of such HABs, as well as to better anticipate and respond to such blooms. Here we present findings on the effect of salinity and availability of carbon on cellular physiology and brevetoxin and brevenal production by K. brevis. When grown at salinities of 35 and 27, but otherwise identical conditions, total brevetoxin cellular concentration varied between 0 to 18.5 pg cell-1 and brevenal varied between 0 and 1 pg cell-1. In response to hypoosmotic stress brevetoxin production was triggered, as a result, brevetoxin production increased up to 53%, while growth rates remained unchanged. A significant hypoosmotic event of >11%, was needed to trigger the response in brevetoxin production. To determine if K. brevis was sensing changes in specific ions within seawater (K+, Cl- or Ca2+), we systematically removed one ion while keeping the remaining ions at equivalent molar concentration for salinity of 35. Dilution in seawater K+ concentrations triggered the production of brevetoxins, increasing production ≥44%. Ecosystem changes due to climate change have increased the production of toxins in other HAB species; here we examined the impact on K. brevis. We have shown that modification of pCO2 level and temperature did not influence brevetoxin production; however, predicted climate change scenarios (increased temperature and pCO2) did significantly increase the growth rate of K. brevis, by 60% at 25°C and 55% at 30°C. We suggest that K. brevis blooms could benefit from predicted increase in pCO2 over the next 100 years. Overall, our findings close a critical gap in knowledge regarding the function of brevetoxin in K. brevis by identifying a connection between brevetoxin production and osmoacclimation.

No description available.

A major goal in conservation biology is to understand the effects of short and long term environmental change on organisms. Fishes are the most valuable marine resource, however very little is known about the synergistic effect of current ocean warming and acidification, and the role of body size and local adaptation on their resilience. There is growing evidence that increased environmental temperature correlates with a reduction in ectotherm body size, suggesting a universal response to warming. To investigate the potential advantage of small body size in fish resilience, I made intra- and inter-specific comparisons of dwarf- and normal-size cleaner gobies of the genus Elacatinus. I first tested the hypothesis that smaller body size would correlate with a wider thermal tolerance by using same-age but different-size gobies reared at 'common garden' conditions. By employing critical thermal methodology, I provided empirical evidence supporting thermal biology theories that predict wider thermal tolerance windows as body size shrinks. These results provided the motivation to examine the effect of body mass on digestive performance, an indicator of fitness. Only smaller fish increased digestive metabolic scope at higher temperatures, thus suggesting that temperature increase caused by global warming will favor smaller individuals. To investigate the role of local adaptation on resilience in climate change, I compared the responses to warming and acidification between latitudinally- and morphologically-distinct populations of the little skate Leucoraja erinacea, by focusing on the most vulnerable life stages, embryos and juveniles. Embryos maintained at common garden conditions showed countergradient variation in performance curves. In juvenile skates, post-exercise metabolic curves shifted performance optima, exhibiting thermal adaptation in the two populations examined. This suggests that as skates hatch and are able to thermoregulate, they can change their temperature optima to exploit local thermal environments. Lastly, temperature and acidification levels predicted by the end of the century may reduce fitness of the northern population of skates, thus increasing vulnerability to local extinction.

Environ 30% du dioxyde de carbone produit par des activités humaines est absorbé par l’océan menant à une diminution de pH d’eau de mer et de l’état de saturation de carbonate de calcium (CaCO3). L’acidification des océans engendrera probablement de profonds changements dans les écosystèmes marins, en particulier chez les organismes marins calcifiants. Les coccolithophoridés produisent avec les foraminifères plus de 90% des carbonates pélagiques dans l’océan actuel. Les expériences de culture ont montré que la réponse des coccolithophoridés à l’acidification des océans varie au sein d’une même espèce ce qui complique l’estimation de l’impact global sur le cycle de carbone et des rétroactions sur le climat. En effet, la sensibilité des organismes et les réponses vis-à-vis de l’augmentation du CO2 dissous dans l’océan et donc de la diminution du pH de l’eau de mer sont différentes. Les conséquences de la calcification sur les coccolithophoridés sont encore très peu décrites et quantifiées. Les coccolithes sont formés à l’intérieur de la cellule dans une vésicule interne. Le pH à l’intérieur de cette vésicule est un paramètre central qui détermine la précipitation de la calcite et donc de la formation des coccolithes. Actuellement, le pH de la vésicule ne peut pas être précisément mesuré et c’est la mesure indirecte de paramètres géochimies qui nous permet d’estimer ces processus. La capacité de réguler le pH de la vésicule vis-à-vis des changements du pH d’eau de mer permet la précipitation de calcite et détermine l’adaptation potentielle de certainscoccolithophoridés à l’acidification des océans. Deux souches d’E. huxleyi ont été cultivées dans des cultures en batchs dilués dans trois conditions pH différentes afin d’évaluer les modalités de réponse aux variations du pH de l’eau de mer. Des paramètres physiologiques incluant le taux de croissance, le POC et le PIC et ont été examinés, en parallèle aux mesures de B/Ca et δ11B dans la calcite des coccolithes pour progresser sur la compréhension de ces mécanismes intracellulaire et sur l’existence d’une relation entre ces paramètres et le pH pour évaluer le potentiel de l’isotopie du bore comme proxy du paleo-pH. / About 30% of the carbon dioxide produced by human activities is absorbed by the ocean leading to a decrease of seawater pH and saturation state of calcium carbonate (CaCO3). The subsequent ocean acidification is likely to result in profound changes in marine ecosystems, in particular among the marine calcifiers. Coccolithophorides together with foraminifera produce more than 90% of the pelagic carbonate in the modern ocean. Culture experiments have shown that the response of coccolithophores to pH varies between and within species, thus complicating our understanding of the overall impact biological response on the carbon cycle and feedbacks on climate. Indeed, different sensitivities to increase dissolved CO2 and decrease seawater pH, and their consequences on calcification exist among coccolithophores, but they are still not fully described nor quantified. Calcareous coccoliths are formed inside the cell in an internal vesicle called the coccolith vesicle. The pH inside the coccolith vesicle would be a key parameter in determining calcite precipitation and therefore coccolith formation. Currently the coccolith vesicle pH cannot be accurately measured and thus estimates have to be based on indirect geochemical evidences. The capacity of the coccolith vesicle to regulate pH allowing for calcite precipitation could explain the resilience of somecoccolithophores to ocean acidification. To further explore this hypothesis, two strains of E. huxleyi were grown in batch cultures under three different pH conditions to assess their response to changing seawater pH. Physiological parameters including growth rate, POC and PIC were examine, in addition to assessing changes in the vesicle pH by measuring B/Ca and δ11B in coccolith calcite and evaluate the potential of boron for paleo-pH reconstruction.

Diante do quadro atual de previsões de mudanças climáticas, estudos a respeito das possíveis respostas dos organismos a estas alterações são importantes. Com a finalidade de prever e verificar se estas serão de fato deletérias ou se os organismos são capazes de lidar com elas sem alterações na sua fisiologia, e consequentemente na estrutura do ambiente, E. brasiliensis foi utilizada como modelo para estudar possíveis impactos do aumento da temperatura e acidificação dos oceanos na sua fisiologia. Para isso, espécimes foram expostos a 9 possíveis combinações de temperatura (24ºC, 28ºC e 30ºC) e pH (8.0, 7.7 e 7.3) em diferentes intervalos de tempo (1, 3, 12, 24 e 48 h). Amostras de gônadas e fluido celomático foram coletadas para avaliar a expressão das proteínas de estresse HSP70, AIF-1 e p38-MAPK, e a variação no número e viabilidade dos celomócitos. Nossos resultados mostram que o modelo é sensibilizado pelas mudanças no ambiente, através da hiper-regulação das proteínas de estresse. O cenário considerado mais extremo (30°C + pH7.3) ocasionou a morte de 100% dos organismos após 24horas. E o segundo cenário mais severo (30°C + pH7.7) desencadeou o desenvolvimento de ulceração de pele. Os efeitos são mais pronunciados nos celomócitos e a acidificação da água parece ter efeitos antagônicos com a temperatura nos celomócitos e sinérgicos nas gônadas. Embora a resposta tenha sido sistêmica, o grau e a dinâmica foram distintos em relação às diferentes amostras e estresses. Podendo causar modificações na resposta imune dos organismos e consequentemente na sobrevivência da espécie a longo prazo. / Under the current Climate Change context, studies about the potential responses of the organisms to their changing environment are of extreme importance. Recent studies point out the synergy of temperature and ocean acidification altogether. In this study, we used the sea star E. brasiliensis to assess the physiological effects of rising temperature, seawater acidification and the interaction of both factors. Independent individuals (N=225) were exposed to 9 possible combinations of temperature (24ºC, 28ºC and 30ºC) and pH (8.0, 7.7 and 7.3), for 1, 3, 12, 24 and 48 h. We compared the stress produced by these treatments measuring the expression of heat shock proteins (HSP70), the production of the allograft inflammatory factor (AIF−1) and the activation of mitogen kinases (MAPKs) at both gonad and celomic fluid. Furthermore, we assessed the quantity and quality of coelomocytes. Our results demonstrated that E. brasiliensis is vulnerable to the interaction of temperature and acidification. All the stress proteins evaluated were upregulated. The extreme scenario (30°C + pH7.3) caused the death of 100% of organisms after 24 hours, while the second most severe scenario (30°C + pH7.7) triggered skin ulceration. Nevertheless, we found that water acidification produces antagonistic effects to the temperature in coelomocytes and synergistic effects in gonad cells. Furthermore, these effects were more pronounced in the coelomocytes than in the gonads. The systemic response found in this study suggest that the interactive effects of elevated temperatures in conjunction with ocean acidification may endanger the survival of this species, and it could compromise the ecosystem functioning at long term.

The main aim of this thesis was to determine how marine metazoans might persist as ocean acidification (OA) conditions intensify. This was done using a combination of field surveys, field transplants and laboratory experiments with polychaetes from a site where volcanically-derived CO2 gas bubbles through the seafloor and drives the seawater pH down, resulting in a marine ecosystem representative of global OA projections for, or before, the year 2100. My first objective was to identify phenotypes, or traits, associated with OA tolerance (Chapter 2 and 5). To do this, I characterized the distribution of dominant calcifying polychaetes along natural pH gradients and used a comparative species recruitment trial to investigate life history traits underlying species’ OA tolerance, or vulnerability. I first found two dominant, closely related species of polychaete: Pileolaria militaris Claparède, 1870 and Simplaria sp. (Serpulidae, Spirorbinae). I then found that increased fecundity and rapid settlement are important traits in determining species’ abilities to persist in low pH environments (Chapter 2). Afterwhich, I investigated the life history traits of the non-calcifying polychaete, Platynereis dumerilii (Audouin & Milne Edwards, 1834), of one of the few species from the low pH site known to have broadcasting, pelagic development. I performed breeding experiments on P. dumerilii collected in both ambient and low pH sites and found that specimens from the low pH site were actually the direct developing brooder sister species, Platynereis massiliensis (Moquin-Tandon, 1869). By reanalayzing the distributions of both species at each site using genetic barcoding, I found clear evidence that direct development and brooding are dominant traits at low pH site, and for OA persistence (Chapter 5). My second objective was to use reciprocal transplant experiments to compare the relative importance of local adaptation and/or plasticity as potential mechanisms responsible for the differential tolerances of populations of the polychaete species Simplaria sp. to low pH. Laboratory transplants indicate that a local adaptation response occurred through genetic accommodation in the Simplaria sp. population from the low pH site. However, neither local adaptation nor plasticity appeared responsible for this species natural low pH persistence when assessed in situ (Chapter 3 & 4). My final objective was to create a framework using the polychaete vent model to identify other types of marine metazoans that are likely to be able to adapt to, and survive, under the predicted environmental conditions (Chapter 5). I overviewed the life history strategies of all dominant polychaetes in the low pH sites, and related trends in their life history strategies to those of other marine invertebrates. Brooding and direct development appear to be key traits for species likely to persist in future oceans pH. I conclude by summarizing how research regarding evolutionary responses may be advanced to add confidence to our projections of future marine metazoan responses.

Climate change is predicted to affect the larval stages of many marine organisms. Ocean warming can reduce larval survival and hasten larval development, whereas ocean acidification can delay larval development. Ocean acidification is especially concerning for marine organisms that develop and grow calcified shells or skeletons in an environment undersaturated with calcium carbonate minerals. This study assessed the effects of ocean warming and acidification on the fertilization and larval development of the green sea urchin, Lytechinus variegatus, a tropical species common in Florida and the Caribbean. After spawning, gametes were fertilized and embryos/larvae were reared at: 1) 28°C and pH 8.1 (control), 2) 28°C and pH 7.8 (ocean acidification scenario), 3) 31°C and pH 8.1 (ocean warming scenario), and 4) 31°C and pH 7.8 (ocean warming and acidification scenario). Exposure to acidified conditions had no effect on fertilization, but delayed larval development, stunted growth and increased asymmetry. Exposure to warm conditions decreased fertilization success at a high sperm to egg ratio (1,847:1), accelerated larval development, but had no significant effect on growth. Under exposure to both stressors (ocean warming and acidification), larval development was accelerated, but larvae were smaller and more asymmetric. These results indicate that climate change will have a serious impact on the larval development and growth of the green sea urchin, L. variegatus, and may negatively affect its persistence.

For decades, humans have impacted marine ecosystems in a variety of ways including contamination by pollution, fishing, and physical destruction of habitats. Global change has, and will, lead to alterations in in a number of abiotic factors of our ocean in particular reduced oxygen saturation, salinity changes, elevated temperature (ocean warming or OW) and elevated carbon dioxide (ocean acidification or OA). Now and in the future, OA and OW will operate together with local anthropogenic drivers such as oil pollution. And yet, at present, very little is known about their potential combined interactive effects on physiological performance and tolerance of marine organisms. Therefore, multiple driver experiments are required if we are to understand and predict future vulnerability of species, populations and ecosystems. Early life stages of invertebrates are generally considered most vulnerable to environmental stress. However, few studies consider the combined effects OA and OW on survival and growth during early development of marine invertebrates, and to our knowledge, there is no information on the additional effects of oil pollution. Therefore, the aim of this thesis was to investigate the effects of combined exposure to OA, OW, and incorporating local drivers such as oil pollution on the development, morphology and physiology of three economically and ecologically important marine invertebrates. These are Northern shrimp Pandalus borealis, Northern krill Meganyctiphanes norvegica, and the green sea urchin Strongylocentrotus droebachiensis. All are cold-water species, assumed to have a narrower tolerance than more temperate species, and so could be particular sensitive to combined stressor affects. Both Northern krill and to a lesser extent Northern shrimp larvae survived experimental conditions, mirroring those predicted under a future global change scenario (combined OA and OW exposure). Neither was hatching success affected. Both shrimp and krill larvae exhibited accelerated developmental rates and incurred greater maintenance costs as a result of exposure to these stressors. Shrimp larvae showed accelerated developmental rates (-9 days), increased metabolic rates (+20 %), and increased feeding rates (+20 %), but reduced growth (- 9 %) when exposed to OW compared with the control. OA increased development rate but only at the control temperature. Although juvenile mortality of krill was not affected by predicted OA/OW conditions, metabolic rate increased significantly (+ 36 %), as did larval developmental rate, while number of moults, feeding rate and growth (- 67 %) decreased significantly (- 67 %, - 60 % and -8 % respectively). Accelerated development was accompanied by greater maintenance costs possibly due to experience a mismatch between energy supply and demand. Both species had an excess of food, and so growth reduction was more likely to be associated with higher metabolic demands in the future global change treatments. Food shortage in situ, due to variable food availability in the sea and/or mismatch with key prey species (algae and zooplankton) could result in more negative effects on growth and ultimately survival. Green sea urchins were also able to survive OA exposure, without detectable effects on hatching success. However, at day 44 post-fertilization, larval body length in the OA treatment was 9 % lower compared to the control. Furthermore, there was a significant tendency of urchin larvae to increase swimming activity in the OA conditions that might indicate compensatory feeding. Elevated maintenance and repair costs as a result of exposure to multi-stressors affected the energy budget of all the three species studied here resulting in reduced growth. Global drivers (OA and OW) resulted in trade-offs with more energy reallocated to swimming activity and metabolism, rather than growth. Exposure to oil reduced the acquisition of energy by reduced feeding which in turn resulted in less energy being available for growth. Both shrimp and sea urchin larvae showed reduced activity and feeding when exposed to oil. It is possible that the reduced swimming activity observed may be due to a narcotic effect of the oil. Furthermore, early stage sea urchin larvae showed increased mortality when exposed to oil while the older larvae did not, indicating a stage specific toxicity to oil for sea urchin larvae. The combination of global drivers and oil pollution acted additively on growth for both sea urchin and shrimp larvae. The impact of combined drivers on the size of shrimp larvae was equal the sum of the negative impacts observed for each driver: a 5 % reduction when exposed to OA and OW, a 9 % reduction when exposed to oil, and a cumulative 15 % reduction when exposed to all stressors. Similarly, the impact of combined drivers on the size of sea urchin larvae was equal to the sum of the negative impacts observed for each driver: a 14 % reduction when exposed to OA, a 9 % reduction when exposed to oil, and a 21 % reduction when exposed to all drivers. Therefore, the study demonstrated the additive physiological effects of OA, OW and a contaminant, and indicated that larval (sea urchin and shrimp) resilience to future changes (i.e. pollution) could be greatly reduced if larvae were already energy limited and severely stressed (reduced development) as a result of exposure to the global drivers. This study therefore shows the importance that the effective management of local drivers such as oil pollution could have against the backdrop of OA and OW, and emphasises that it is important to study impacts of toxicants, such as an oil pollution, in the context of predicted changes in the environment, as OW and OA are becoming major concerns. Finally, the fact that some local and global drivers seem to act additively should encourage local managers to act on local driver regulations, to obtain positive effects on local populations and environment and thereby rendering them more resilient to the negative impacts of future global drivers.