From Elemental process studies to ecosystem models in the ocean biological pump

The biological pump controls the transport of particles from the ocean surface to the deep sea. Vertical fluxes govern chemical gradients, playing a fundamental role in the carbon dioxide (CO2) ocean-atmosphere feedback, driving physicochemical properties of seawater, and providing resources for benthic ecosystems. Particles vary in size and composition, originating in every trophic level as detritus, fecal material, biogenic carbonates or the organisms biomass. A recent concern is that the concomitant increase in anthropogenic CO2 and temperature, may alter carbon fluxes and atmospheric feedbacks, as well as ecosystem shifts driven by organisms’ physiological and biogeochemical responses. This dissertation focused on poorly characterised biological pump compartments and processes that are central to assess potential implications of future climate scenarios

Discrete Pulses of Cooler Deep Water Can Decelerate Coral Bleaching During Thermal Stress: Implications for Artificial Upwelling During Heat Stress Events

Global warming is considered to be the most severe threat to coral reefs globally, which makes it important for scientists to develop novel strategies that mitigate the impact of warming on corals and associated habitats. Artificial upwelling of cooler deep water to the surface layer may be a possible mitigation/management tool. In this study, we investigated the effect of simulated artificial upwelling with deep water off Bermuda collected at 50 m (24°C) and 100 m (20°C) on coral symbiont biology of 3 coral species (Montastrea cavernosa, Porites astreoides, and Pseudodiploria strigosa) in a temperature stress experiment. The following treatments were applied over a period of 3 weeks: (i) control at 28°C (ii) heat at 31°C, (iii) heat at 31°C+ deep water from 50 m depth, and (iv) heat at 31°C+ deep water from 100 m depth. Artificial upwelling was simulated over a period of 25 min on a daily basis resulting in a reduction of temperature for 2 h per day and the following degree-heating-weeks: 5.7°C-weeks for ii, 4.6°C-weeks for iii and 4.2°C-weeks for iv. Comparative analysis of photosynthetic rate, chlorophyll-a concentration and zooxanthellae density revealed a reduction of heat stress responses in artificial upwelling treatments in 2 of the 3 investigated species, and a stronger positive effect of 100-m water than 50-m water. These results indicate that artificial upwelling could be an effective strategy to mitigate coral bleaching during heat stress events allowing corals to adjust to increasing temperatures more gradually. It will still be necessary to further explore the ecological benefits as well as potential ecosystem impacts associated with different artificial upwelling scenarios to carefully implement an effective in situ artificial upwelling strategy in coral reefs

Modeling coral bleaching mitigation potential of water vertical translocation – an analogue to geoengineered artificial upwelling

Artificial upwelling (AU) is a novel geoengineering technology that brings seawater from the deep ocean to the surface. Within the context of global warming, AU techniques are proposed to reduce sea surface temperature at times of thermal stress around coral reefs. A computationally fast but coarse 3D Earth System model (3.6° longitude × 1.8° latitude) was used to investigate the environmental impacts of hypothetically implemented AU strategies in the Great Barrier Reef, South China Sea, and Hawaiian regions. While omitting the discussion on sub-grid hydrology, we simulated in our model a water translocation from either 130 or 550 m depth to sea surface at rates of 1 or 50 m3 s−1 as analogues to AU implementation. Under the Representative Concentration Pathway 8.5 emissions scenario from year 2020 on, the model predicted a prevention of coral bleaching until the year 2099 when AU was implemented, except under the least intense AU scenario (water from 130 m depth at 1 m3 s−1). Yet, intense AU implementation (water from 550 m depth at 50 m3 s-1) will likely have adverse effects on coral reefs by overcooling the surface water, altering salinity, decreasing calcium carbonate saturation, and considerably increasing nutrient levels. Our result suggests that if we utilize AU for mitigating coral bleaching during heat stress, AU implementation needs to be carefully designed with respect to AU’s location, depth, intensity and duration so that undesirable environmental effects are minimized. Following a proper installation and management procedure, however, AU has the potential to decelerate destructive bleaching events and buy corals more time to adjust to climate change

Temperature Induced Physiological Reaction Norms of the Coccolithophore Gephyrocapsa oceanica and Resulting Coccolith Sr/Ca and Mg/Ca Ratios

Coccolithophores are one of the major contributors to the pelagic production of calcium carbonate and their fossilized remains are a key component of the biogeochemical cycles of calcium (Ca), magnesium (Mg), and other divalent cations present in the intracellular precipitated calcitic structures (coccoliths). The geochemical signature of coccoliths (e.g., Sr/Ca and Mg/Ca ratios) is used as paleoproxy to reconstruct past environmental conditions and to understand the underlying physiological precipitation kinetics. Here, we present the elemental fractionation of Sr and Mg in calcite of the coccolithophore Gephyrocapsa oceanica from controlled laboratory experiments applying an extended temperature gradient (12 to 27°C). The physiological reaction norm of G. oceanica, in terms of growth rate, exhibited optimum behavior while the partition coefficient of Sr (DSr) was linearly correlated with temperature and DMg indicated no specific trend. Our results indicate: (1) a presumably secondary physiological control of DSr, and (2) the importance of calibrating coccolithophore-based proxies using experiments that include the full physiological reaction norms (i.e., a possible non-linear response) to environmental drivers (e.g., temperature, salinity, and pH, etc.). The presented results contribute to an improved understanding of the underlying physiological kinetics involved in regulating coccolith elemental fractionation and give additional implications for designing future laboratory experiments to calibrate and apply coccolithophore based paleoproxies on the fossil sediment record

Physiological responses of coccolithophores to abrupt exposure of naturally low pH deep seawater

Upwelling is the process by which deep, cold, relatively high-CO2, nutrient-rich seawater rises to the sunlit surface of the ocean. This seasonal process has fueled geoengineering initiatives to fertilize the surface ocean with deep seawater to enhance productivity and thus promote the drawdown of CO2. Coccolithophores, which inhabit many upwelling regions naturally ‘fertilized’ by deep seawater, have been investigated in the laboratory in the context of ocean acidification to determine the extent to which nutrients and CO2 impact their physiology, but few data exist in the field except from mesocosms. Here, we used the Porcupine Abyssal Plain (north Atlantic Ocean) Observatory to retrieve seawater from depths with elevated CO2 and nutrients, mimicking geoengineering approaches. We tested the effects of abrupt natural deep seawater fertilization on the physiology and biogeochemistry of two strains of Emiliania huxleyi of known physiology. None of the strains tested underwent cell divisions when incubated in waters obtained from 1,000 m, artificial upwelling using shallower waters may not be a suitable approach for promoting carbon sequestration for some locations and assemblages, and should therefore be investigated on a site-by-site basis

Sinking of Gelatinous Zooplankton Biomass Increases Deep Carbon Transfer Efficiency Globally

Gelatinous zooplankton (Cnidaria, Ctenophora, and Urochordata, namely, Thaliacea) are ubiquitous members of plankton communities linking primary production to higher trophic levels and the deep ocean by serving as food and transferring “jelly‐carbon” (jelly‐C) upon bloom collapse. Global biomass within the upper 200 m reaches 0.038 Pg C, which, with a 2–12 months life span, serves as the lower limit for annual jelly‐C production. Using over 90,000 data points from 1934 to 2011 from the Jellyfish Database Initiative as an indication of global biomass (JeDI: http://jedi.nceas.ucsb.edu, http://www.bco‐dmo.org/dataset/526852), upper ocean jelly‐C biomass and production estimates, organism vertical migration, jelly‐C sinking rates, and water column temperature profiles from GLODAPv2, we quantitatively estimate jelly‐C transfer efficiency based on Longhurst Provinces. From the upper 200 m production estimate of 0.038 Pg C year−1, 59–72% reaches 500 m, 46–54% reaches 1,000 m, 43–48% reaches 2,000 m, 32–40% reaches 3,000 m, and 25–33% reaches 4,500 m. This translates into ~0.03, 0.02, 0.01, and 0.01 Pg C year−1, transferred down to 500, 1,000, 2,000, and 4,500 m, respectively. Jelly‐C fluxes and transfer efficiencies can occasionally exceed phytodetrital‐based sediment trap estimates in localized open ocean and continental shelves areas under large gelatinous blooms or jelly‐C mass deposition events, but this remains ephemeral and transient in nature. This transfer of fast and permanently exported carbon reaching the ocean interior via jelly‐C constitutes an important component of the global biological soft‐tissue pump, and should be addressed in ocean biogeochemical models, in particular, at the local and regional scale

Phenotypic Variability in the Coccolithophore Emiliania huxleyi

Coccolithophores are a vital part of oceanic phytoplankton assemblages that produce organic matter and calcium carbonate (CaCO3) containing traces of other elements (i.e. Sr and Mg). Their associated carbon export from the euphotic zone to the oceans' interior plays a crucial role in CO2 feedback mechanisms and biogeochemical cycles. The coccolithophore Emiliania huxleyi has been widely studied as a model organism to understand physiological, biogeochemical, and ecological processes in marine sciences. Here, we show the inter-strain variability in physiological and biogeochemical traits in 13 strains of E. huxleyi from various biogeographical provinces obtained from culture collections commonly used in the literature. Our results demonstrate that inter-strain genetic variability has greater potential to induce larger phenotypic differences than the phenotypic plasticity of single strains cultured under a broad range of variable environmental conditions. The range of variation found in physiological parameters and calcite Sr:Ca highlights the need to reconsider phenotypic variability in paleoproxy calibrations and model parameterizations to adequately translate findings from single strain laboratory experiments to the real ocean

The effect of nitrate and phosphate availability on Emiliania huxleyi (NZEH) physiology under different CO2 scenarios

Growth and calcification of the marine coccolithophorid Emiliania huxleyi is affected by ocean acidification and macronutrients limitation and its response varies between strains. Here we investigated the physiological performance of a highly calcified E. huxleyi strain, NZEH, in a multiparametric experiment. Cells were exposed to different CO2 levels (ranging from 250 to 1314 μatm) under three nutrient conditions nutrient replete (R), nitrate limited (-N), and phosphate limited (-P). We focused on calcite and organic carbon quotas and on nitrate and phosphate utilization by analyzing the activity of nitrate reductase (NRase) and alkaline phosphatase (APase), respectively. Particulate inorganic (PIC) and organic (POC) carbon quotas increased with increasing CO2 under R conditions but a different pattern was observed under nutrient limitation. The PIC:POC ratio decreased with increasing CO2 in nutrient limited cultures. Coccolith length increased with CO2 under all nutrient conditions but the coccosphere volume varied depending on the nutrient treatment. Maximum APase activity was found at 561 μatm of CO2 (pH 7.92) in -P cultures and in R conditions, NRase activity increased linearly with CO2. These results suggest that E. huxleyi's competitive ability for nutrient uptake might be altered in future high-CO2 oceans. The combined dataset will be useful in model parameterizations of the carbon cycle and ocean acidification

Phenotypic variability in the coccolithophore Emiliania huxleyi

Coccolithophores are a vital part of oceanic phytoplankton assemblages that produce organic matter and calcium carbonate (CaCO3) containing traces of other elements (i.e. Sr and Mg). Their associated carbon export from the euphotic zone to the oceans' interior plays a crucial role in CO2 feedback mechanisms and biogeochemical cycles. The coccolithophore Emiliania huxleyi has been widely studied as a model organism to understand physiological, biogeochemical, and ecological processes in marine sciences. Here, we show the inter-strain variability in physiological and biogeochemical traits in 13 strains of E. huxleyi from various biogeographical provinces obtained from culture collections commonly used in the literature. Our results demonstrate that inter-strain genetic variability has greater potential to induce larger phenotypic differences than the phenotypic plasticity of single strains cultured under a broad range of variable environmental conditions. The range of variation found in physiological parameters and calcite Sr:Ca highlights the need to reconsider phenotypic variability in paleoproxy calibrations and model parameterizations to adequately translate findings from single strain laboratory experiments to the real ocean