Computational analysis of the biophysical controls on Southern Ocean phytoplankton ecosystem dynamics

Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Applied Ocean Science & Engineering at the Massachusetts Institute of Technology and the Woods Hole Oceanographic Institution February 2019.Southern Ocean net community productivity plays an out sized role in regulating global biogeochemical cycling and climate dynamics. The structure of spatial-temporal variability in phytoplankton ecosystem dynamics is largely governed by physical processes but a variety of competing pathways complicate our understanding of how exactly they drive net population growth. Here, I leverage two coupled, 3-dimensional, global, numerical simulations in conjunction with remote sensing data and past observations, to improve our mechanistic understanding of how physical processes drive biology in the Southern Ocean. In Chapter 2, I show how different mechanistic pathways can control population dynamics from the bottom-up (via light, nutrients), as well as the top-down (via grazing pressure). In Chapters 3 and 4, I employ a higher resolution, eddy resolving, integration to explicitly track and examine closed eddy structures and address how they modify biomass at the mesoscale. Chapter 3 considers how simulated eddies drive bottom-up controls on phytoplankton growth and finds that division rates are, on average, amplified in anticyclones and suppressed in cyclones. Anomalous division rates are predominately fueled by an anomalous vertical iron flux driven by eddy-induced Ekman Pumping. Chapter 4 goes on to describe how anomalous division rates combine with anomalous loss rates to drive anomalous net population growth. Biological rate-based mechanisms are then compared to the potential for anomalies to evolve strictly via physical transport (i.e. dilution, stirring, advection). All together, I identify and describe dramatic regional and seasonal variability in when, where, and how different mechanisms drive phytoplankton growth throughout the Southern Ocean. Better understanding this variability has broad implications to our understanding of how oceanic biogeochemisty will respond to, and likely feedback into, a changing climate. Specifically, the uncertainty associated with this variability should temper recent proposals to artificially stimulate net primary production and the biological pump via iron fertilization. In Chapter 5 I argue that Southern Ocean Iron Fertilization fails to meet the basic tenets required for adoption into any regulatory market based framework.The CESM project is supported by the National Science Foundation and the Office of Science (BER) of the U.S. Department of Energy. Computing resources were provided by the Climate Simulation Laboratory at NCAR’s Computational and Information Systems Laboratory (CISL), sponsored by the National Science Foundation and other agencies. This research was enabled by CISL compute and storage resources. TR was supported by an NDSEG graduate fellowship. TR and SCD acknowledge support from the National Aeronautics and Space Administration Ocean Biology and Biogeochemistry Program (NNX14AL86G). TR, SCD and MTK acknowledge support from the National Science Foundation Polar Programs award 1440435 (Antarctic Integrated System Science) to the Palmer LTER program. Please contact [email protected] for further questions or to access to data

Variability in the mechanisms controlling Southern Ocean phytoplankton bloom phenology in an ocean model and satellite observations

Author Posting. © American Geophysical Union, 2017. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Global Biogeochemical Cycles 31 (2017): 922–940, doi:10.1002/2016GB005615.A coupled global numerical simulation (conducted with the Community Earth System Model) is used in conjunction with satellite remote sensing observations to examine the role of top-down (grazing pressure) and bottom-up (light, nutrients) controls on marine phytoplankton bloom dynamics in the Southern Ocean. Phytoplankton seasonal phenology is evaluated in the context of the recently proposed “disturbance-recovery” hypothesis relative to more traditional, exclusively “bottom-up” frameworks. All blooms occur when phytoplankton division rates exceed loss rates to permit sustained net population growth; however, the nature of this decoupling period varies regionally in Community Earth System Model. Regional case studies illustrate how unique pathways allow blooms to emerge despite very poor division rates or very strong grazing rates. In the Subantarctic, southeast Pacific small spring blooms initiate early cooccurring with deep mixing and low division rates, consistent with the disturbance-recovery hypothesis. Similar systematics are present in the Subantarctic, southwest Atlantic during the spring but are eclipsed by a subsequent, larger summer bloom that is coincident with shallow mixing and the annual maximum in division rates, consistent with a bottom-up, light limited framework. In the model simulation, increased iron stress prevents a similar summer bloom in the southeast Pacific. In the simulated Antarctic zone (70°S–65°S) seasonal sea ice acts as a dominant phytoplankton-zooplankton decoupling agent, triggering a delayed but substantial bloom as ice recedes. Satellite ocean color remote sensing and ocean physical reanalysis products do not precisely match model-predicted phenology, but observed patterns do indicate regional variability in mechanism across the Atlantic and Pacific.NDSEG Graduate Fellowship; National Aeronautics and Space Administration Ocean Biology and Biogeochemistry Program Grant Number: NNX14L86G; NSF Poloar Programs Award Grant Number: 1440435; National Aeronautics and Space Administration Grant Number: NNX14AL86G; NDSEG; National Science Foundation Grant Number: 14404352017-11-3

The Simulated Biological Response to Southern Ocean Eddies via Biological Rate Modification and Physical Transport

We examine the structure and drivers of anomalous phytoplankton biomass in Southern Ocean eddies tracked in a global, multiyear, eddy-resolving, 3-D ocean simulation of the Community Earth System Model.We examine how simulated anticyclones and cyclones differentially modify phytoplankton biomass concentrations, growth rates, and physical transport. On average, cyclones induce negative division rate anomalies that drive negative net population growth rate anomalies, reduce dilution across shallower mixed layers, and advect biomass anomalously downward via eddy-induced Ekman pumping. The opposite is true in anticyclones. Lateral transport is dominated by eddy stirring rather than eddy trapping. The net effect on anomalous biomass can exceed 10–20% of background levels at the regional scale, consistent with observations. Moreover, we find a strong seasonality in the sign and magnitude of regional anomalies and the processes that drive them. The most dramatic seasonal cycle is found in the South Pacific Antarctic Circumpolar Current, where physical and biological processes dominate at different times, modifying biomass in different directions throughout the year. Here, in cyclones, during winter, anomalously shallow mixed layer depths first drive positive surface biomass anomalies via reduced dilution, and later drive positive depth-integrated biomass anomalies via reduced light limitation. During spring, reduced iron availability and elevated grazing rates suppress net population growth rates and drive the largest annual negative surface and depth-integrated biomass anomalies. During summer and fall, lateral stirring and eddy-induced Ekman pumping create small negative surface anomalies but positive depth-integrated anomalies. The same mechanisms drive biomass anomalies in the opposite direction in anticyclones

Eddy-Modified Iron, Light, and Phytoplankton Cell Division Rates in the Simulated Southern Ocean

We examine the effects of Southern Ocean eddies on phytoplankton cell division rates in a global, multiyear, eddy‐resolving, 3‐D ocean simulation of the Community Earth System Model. We first identify and track eddies in the simulation and validate their distribution and demographics against observed eddy trajectory characteristics. Next, we examine how simulated cyclones and anticyclones differentially modify iron, light, and ultimately population‐specific cell division rates. We use an eddy‐centric, depth‐averaged framework to explicitly examine the dynamics of the phytoplankton population across the entire water column within an eddy. We find that population‐averaged iron availability is elevated in anticyclones throughout the year. The dominant mechanism responsible for vertically transporting iron from depth in anticyclones is eddy‐induced Ekman upwelling. During winter, in regions with deep climatological mixed layer depths, anticyclones also induce anomalously deep mixed layer depths, which further supply new iron from depth via an increased upward mixing flux. However, this additional contribution comes at the price of deteriorating light availability as biomass is distributed deeper in the water column. Therefore, even though population‐averaged specific division rates are elevated in Southern Ocean anticyclones throughout most of the year, in the winter, severe light stress can dominate relieved iron stress and lead to depressed division rates in some anticyclones, particularly in the deep mixing South Pacific Antarctic Circumpolar Current. The opposite is true in cyclones, which exhibit a consistently symmetric physical and biogeochemical response relative to anticyclones

The Potential Impact of Nuclear Conflict on Ocean Acidification

We demonstrate that the global cooling resulting from a range of nuclear conflict scenarios would temporarily increase the pH in the surface ocean by up to 0.06 units over a 5-year period, briefly alleviating the decline in pH associated with ocean acidification. Conversely, the global cooling dissolves atmospheric carbon into the upper ocean, driving a 0.1 to 0.3 unit decrease in the aragonite saturation state (Ωarag) that persists for ∼10 years. The peak anomaly in pH occurs 2 years post conflict, while the Ωarag anomaly peaks 4- to 5-years post conflict. The decrease in Ωarag would exacerbate a primary threat of ocean acidification: the inability of marine calcifying organisms to maintain their shells/skeletons in a corrosive environment. Our results are based on sensitivity simulations conducted with a state-of-the-art Earth system model integrated under various black carbon (soot) external forcings. Our findings suggest that regional nuclear conflict may have ramifications for global ocean acidification

A New Ocean State After Nuclear War

Nuclear war would produce dire global consequences for humans and our environment. We simulated climate impacts of US-Russia and India-Pakistan nuclear wars in an Earth System Model, here, we report on the ocean impacts. Like volcanic eruptions and large forest fires, firestorms from nuclear war would transport light-blocking aerosols to the stratosphere, resulting in global cooling. The ocean responds over two timescales: a rapid cooling event and a long recovery, indicating a hysteresis response of the ocean to global cooling. Surface cooling drives sea ice expansion, enhanced meridional overturning, and intensified ocean vertical mixing that is expanded, deeper, and longer lasting. Phytoplankton production and community structure are highly modified by perturbations to light, temperature, and nutrients, resulting in initial decimation of production, especially at high latitudes. A new physical and biogeochemical ocean state results, characterized by shallower pycnoclines, thermoclines, and nutriclines, ventilated deep water masses, and thicker Arctic sea ice. Persistent changes in nutrient limitation drive a shift in phytoplankton community structure, resulting in increased diatom populations, which in turn increase iron scavenging and iron limitation, especially at high latitudes. In the largest US-Russia scenario (150 Tg), ocean recovery is likely on the order of decades at the surface and hundreds of years at depth, while changes to Arctic sea-ice will likely last thousands of years, effectively a “Nuclear Little Ice Age.” Marine ecosystems would be highly disrupted by both the initial perturbation and in the new ocean state, resulting in long-term, global impacts to ecosystem services such as fisheries.publishedVersio

The potential impact of nuclear conflict on ocean acidification

We demonstrate that the global cooling resulting from a range of nuclear conflict scenarios would temporarily increase the pH in the surface ocean by up to 0.06 units over a 5‐year period, briefly alleviating the decline in pH associated with ocean acidification. Conversely, the global cooling dissolves atmospheric carbon into the upper ocean, driving a 0.1 to 0.3 unit decrease in the aragonite saturation state ( &Omega;arag) that persists for &sim;10 years. The peak anomaly in pH occurs 2 years post conflict, while the &Omega;arag anomaly peaks 4‐ to 5‐years post conflict. The decrease in &Omega;arag would exacerbate a primary threat of ocean acidification: the inability of marine calcifying organisms to maintain their shells/skeletons in a corrosive environment. Our results are based on sensitivity simulations conducted with a state‐of‐the‐art Earth system model integrated under various black carbon (soot) external forcings. Our findings suggest that regional nuclear conflict may have ramifications for global ocean acidification. Key Points Nuclear conflict has the potential to increase surface ocean pH and decrease aragonite saturation state The decrease in saturation state would exacerbate shell dissolution from anthropogenic ocean acidification A regional nuclear conflict may have far‐reaching effects on global ocean carbonate chemistry</p

Proton transfer in acetaldehyde–water clusters mediated by a single water molecule

Proton transfer in aqueous media is a ubiquitous process, occurring in acid-base chemistry, biology, and in atmospheric photochemistry. Photoionization mass spectrometry coupled with theoretical calculations demonstrate that a relay-type proton transfer mechanism is operational for single-water-molecule-assisted proton transfer between two acetaldehyde molecules in the gas phase. Threshold photoionization of acetaldehyde-water clusters leads to proton transfer between the formyl groups (-CH[double bond, length as m-dash]O) of one acetaldehyde molecule to another, and the subsequent formation of cationic moieties. Density functional theory computations reveal several plausible pathways of proton transfer in mixed cluster cations. Among these pathways, water-mediated proton transfer is energetically favored. Mass spectra and photoionization efficiency curves confirm these theoretical findings and also demonstrate the increased stability of cluster cations where acetaldehyde molecules are symmetrically bonded to the hydronium ion

Temporal progression in migratory status and sexual maturation in European silver eels during downstream migration

The onset of downstream migration of European eels is accompanied by a cessation of feeding and the start of sexual maturation which stresses the link between metabolism and sexual maturation, also suggesting an important role for exercise. Exercise has been tested with eels in swim tunnels and was found to stimulate the onset of sexual maturation. In this study, we have investigated the interplay between migration and maturation in the field during the downstream migration of female silver eels. Temporal changes in migratory status and sexual maturation among silver eels of the upstream Rhine River system over 3 months of the migration season (August, September and October) were determined in biometrical parameters, plasma 17β-estradiol and calcium levels, oocyte histology and gonadal fat levels. Furthermore, the ecological relevant parameters age as determined by otolithometry and health aspects indicated by haematocrit, haemoglobin and swim-bladder parasite load were measured. Silver eels were estimated to be 14 years old. A strong temporal progression in migratory stage was shown over the months of downstream migration. Catches probably represented a mix of reproductive migrants and feeding migrants of which the ratio increased over time. Furthermore, this study confirmed our hypothesis linking the migratory stage to early maturation as indicated by enlargement of the eyes, oocyte growth and fat deposition in the oocytes, exactly the same changes as found induced by exercise but not ruling out environmental influences. Migrants show extensive fat uptake by the oocytes, probably stimulated by the swimming exercise. In addition, at least 83% of the silver eels in this spawning run may have suffered from negative effects of swim-bladder parasites on their swimming performance