(Table S1) Age determination of sediment cores of the Gulf of Alaska

Oxygen isotope data from planktonic and benthic foraminifera, on a high-resolution age model (44 14C dates spanning 17,400 years), document deglacial environmental change on the southeast Alaska margin (59°33.32'N, 144°9.21'W, 682 m water depth). Surface freshening (i.e., d18O reduction of 0.8 per mil) began at 16,650 ± 170 cal years B.P. during an interval of ice proximal sedimentation, likely due to freshwater input from melting glaciers. A sharp transition to laminated hemipelagic sediments constrains retreat of regional outlet glaciers onto land circa 14,790 ± 380 cal years B.P. Abrupt warming and/or freshening of the surface ocean (i.e., additional d18O reduction of 0.9 per mil) coincides with the Bølling Interstade of northern Europe and Greenland. Cooling and/or higher salinities returned during the Allerød interval, coincident with the Antarctic Cold Reversal, and continue until 11,740 ± 200 cal years B.P., when onset of warming coincides with the end of the Younger Dryas. An abrupt 1 per mil reduction in benthic d18O at 14,250 ± 290 cal years B.P. likely reflects a decrease in bottom water salinity driven by deep mixing of glacial meltwater, a regional megaflood event, or brine formation associated with sea ice. Two laminated opal-rich intervals record discrete episodes of high productivity during the last deglaciation. These events, precisely dated here at 14,790 ± 380 to 12,990 ± 190 cal years B.P. and 11,160 ± 130 to 10,750 ± 220 cal years B.P., likely correlate to similar features observed elsewhere on the margins of the North Pacific and are coeval with episodes of rapid sea level rise. Remobilization of iron from newly inundated continental shelves may have helped to fuel these episodes of elevated primary productivity and sedimentary anoxia.

Stable isotope ratios of epibenthic foraminifer Cibicidoides wuellerstorfi from sediment profile GeoB9526

Studies of temporal changes of ocean circulation and deep-water ventilation often rely on d13C records of epibenthic foraminifer Cibicidoides wuellerstorfi. However, primary productivity related overprints may distort the signal and simulate a chemical age of ambient water mass that is too old and simulates poorly ventilated ambient bottom waters. To further constrain the use of C. wuellerstorfi d13C records from high-productivity areas, we analyzed a 14CAMS-dated gravity core from the upwelling regime off northwest Africa at 12°N. We compare this new record with 37 radiocarbon dated d13C records from the eastern Atlantic Ocean between 45°N and 25°S that are bathed by the same water mass. Only during Heinrich events 1 and 2, when the investigated core site off northwest Africa experienced year-round, sustained deposition of organic matter, the d13C values at this site faithfully record deep-water ventilation states. During times of predominantly seasonal deposition of fresh phytodetritus, however, d13C values were significantly lower than at the reference sites. This underscores that reconstruction of paleocirculation and deep ocean ventilation using C. wuellerstorfi d13C from regions that experienced seasonal phytodetritus deposition needs to be validated by additional proxies that are not affected by local productivity.

Geochemistry of Neogene sediments of ODP Hole 115-711A

Pelagic sedimentation in the northwest Indian Ocean has been studied using sediments from Hole 711A (the section from 0 to 70.5 mbsf, 0-22 Ma), a deep site (4428 m) drilled during Ocean Drilling Program Leg 115. The clay fraction of the sediments represents poorly developed pelagic deposits with considerably lower contents of Mn, Ba, Cu, Ni, Cr, and Zn than is typical for well-oxidized pelagic sediments formed far from the continents (e.g., in the central Indian or Pacific oceans). Geochemical provenance models, representing conservative mixing models with terrigenous, exhalative-volcanic, and biogenous matter as the only inputs, explain most of the compositional variations in the sediments. The models show that terrigenous matter accounts for about 96%-100% of all SiO2, Al2O3, TiO2, and Zr; about 73%-85% of all Fe2O3, V, and Ni; and about 40%-60% of the Cu and Zn abundances. Exhalative-volcanic matter delivers a large fra tion of Mn (78%-85%), some Fe (15%-219/o), and possibly some Cu (38%-51%). Biogenous deposition is generally of restricted significance; at most 6%-35% of all Cu and Zn may derive from biogenic matter. The exhalative-volcanic matter is slightly more abundant in the oldest deposits, reflecting a plate tectonic drift away from the volcanic Carlsberg Ridge. The Al/Ti ratio reveals that silicic crustal matter plays a somewhat larger role in the upper and lower part of the section studied, whereas the basaltic input is slightly higher in the intermediate levels (age 5-15 m.y.). The sediment abundances of Ba generally exceed those predicted by the models, an anomalous behavior also observed in equatorial Pacific sediments. This is possibly caused by poor knowledge of the input components. Several changes in accumulation rates seem to correlate with climatic changes (onset of monsoon-driven upwellings and sea-level regressions of about 50-100 m at 10, 15-16, and 20-21 Ma). A number of constituents show higher accumulation rates at or shortly after these regressions, suggesting an accelerated removal of fines from shallow oceanic areas. Furthermore, the SiO2/Al2O3 ratio shows a small increase in sediments younger than 10 Ma, implying an increase in biological productivity, particularly after the onset of monsoon-driven upwelling in the northwest Indian Ocean. This trend is paralleled by a general increase in the accumulation rates of Ba and CaCO3. However, these accumulation rates are generally significantly lower than under the biological high-productivity zone in the equatorial Pacific. The onset of these upwelling systems about 10 Ma is probably related to the closing of the gap between India and the main Asiatic continent, preventing free circulation around the Indian subcontinent.

Alkenones and sea surface reconstruction of sediments from, the Arabian Sea

Deep-sea sediments of two cores from the western (TY93-929/P) and the southeastern (MD900963) Arabian Sea were used to study the variations of the Indian monsoon during previous climatic cycles. Core TY93-929/P was located between the SW monsoon driven upwelling centres off Somalia and Oman, which are characterized by large seasonal sea surface temperature (SST) and particle flux changes. By contrast, core MD900963, was situated near the Maldives platform, an equatorial ocean site with a rather small SST seasonality (less than 2°C). For both cores we have reconstructed SST variations by means of the unsaturation ratio of C37 alkenones, which is compared with the delta18O records established on planktonic foraminifera. In general, the SST records follow the delta18O variations, with an SST maximum during oxygen isotope stage 5.5 (the Last Interglacial at about 120-130 kyr) and a broad SST minimum during isotope stage 4 and 3.3 (approximately 40-50 kyr). The SST difference between the Holocene and the Last Glacial Maximum (LGM) is of the order of 2°C. In both cores the SSTs during isotope stage 6 are distinctly higher by 1-2°C than the cold SST minima during the last glacial cycle (LGM and stage 3). To reconstruct qualitatively the past productivity variations for the two cores, we used the concentrations and fluxes of alkenones and organic carbon, together with a productivity index based on coccolith species (Florisphaera profunda relative abundance). Within each core, there is a general agreement between the different palaeoproductivity proxies. In the southeastern Arabian Sea (core MD900963), glacial stages correspond to relatively high productivity, whereas warm interstadials coincide with low productivity. All time series of productivity proxies are dominated by a cyclicity of about 21-23 kyr, which corresponds to the insolation precessional cycle. A hypothesis could be that the NE monsoon winds were stronger during the glacial stages, which induced deepening of the surface mixed layer and injection of nutrients to the euphotic zone. By contrast, the records are more complicated in the upwelling region of the western Arabian Sea (core TY93-929/P). This is partly due to large changes in the sedimentation rates, which were higher during specific periods (isotope stages 6, 5.4, 5.2, 3 and 2). Unlike core MD900963, no simple relationship emerges from the comparison between the delta18O stratigraphy and productivity records. The greater complexity observed for core TY93-929/P could be the result of the superimposition of different patterns of productivity fluctuations for the two monsoon seasons, the SW monsoon being enhanced during interglacial periods, whereas the NE monsoon was increased during glacial intervals. A similar line of reasoning also could help explain the SST records by the superimposition of variations of three components: global atmospheric temperature, and SW and NE monsoon dynamics.

Layer description, carbon and nitrogen content and stable isotope record of bulk sedimentary organic matter from Laguna Potrok Aike

An investigation of stable isotope (d13C TOC and d15N TN) and elemental parameters (TOC, TN contents and TOC/TN ratios) of bulk organic matter (<200 µm) from sediment cores recovered from the Patagonian lake Laguna Potrok Aike (Argentina) in the framework of the ICDP deep drilling project PASADO provided insights into past changes in lake primary productivity and environmental conditions in South Patagonia throughout the last Glacial-Interglacial transition. Stratigraphically constrained cluster analyses of all proxy parameters suggest four main phases. From ca 26,100 to 17,300 cal. years BP, lacustrine phytoplankton was presumably the predominant organic matter source in an aquatic environment with low primary productivity rates. At around 17,300 cal. years BP, abrupt and distinct shifts of isotopic and elemental values indicate that the lacustrine system underwent a rapid reorganization. Lake primary productivity (phytoplankton and aquatic macrophytes) shows higher levels albeit with large variations during most of the deglaciation until 13,000 cal. years BP. The main causes for this development can be seen in improved growing conditions for primary producers because of deglacial warming in combination with expedient availability of nutrients and likely calm wind conditions. After 13,000 cal. years BP, decreased d13C TOC values, TOC, TN contents and TOC/TN ratios indicate that the lake approached a new state with reduced primary productivity probably induced by unfavourable growing conditions for primary producers like strengthened winds and reduced nutrient availability. The steady increase in d15N TN values presumably suggests limitation of nitrate supply for growth of primary producers resulting from a nutrient shortage after the preceding phase with high productivity. Nitrate limitation and consequent decreased lacustrine primary productivity continued into the early Holocene (10,970-8400 cal. years BP) as reflected by isotopic and elemental values.

(Table 1) Age determination of Arctic Ocean sediment cores

Accelerator mass spectrometer 14C dated stable isotope data from Neogloboquadrina pachyerma in cores raised from the Mendeleyev Ridge and slope provide evidence for significant influx of meltwater to the western Arctic Ocean during the early part of marine oxygen isotope stage 1 (OIS 1) and during several intervals within OIS 3. The strongest OIS 3 meltwater event occurred before ca. 45 ka (conventional radiocarbon age) and was probably related to the deglaciation at the beginning of OIS 3. Major meltwater input to the western Arctic Ocean during the last deglaciation coincides closely with the maximum rate of global sea-level rise as determined from the Barbados sea-level record, demonstrating a strong link between the global record and changes in the central Arctic Ocean. OIS 2, which includes the last glacial maximum, is very condensed or absent in the cores. Abundance and d13C values for N. pachyderma in the middle part of OIS 3 are similar to modern values, indicating high productivity and seasonal ice-free areas along the Arctic margin at that time. These records indicate that the Arctic Ocean was a source of heat and moisture to the northern polar atmosphere during parts of OIS 3.

Surface sediment data, CaCO3 and TOC

Modern sedimentary total organic carbon (TOC) content as a proxy for surface water export production was mapped on the shelf and on the upper continental slope of the Benguela upwelling system using 137 core tops. Shelf maxima in TOC can be correlated with maxima in surface water productivity. On the slope, high TOC contents are observed offshore from sites of strong modern upwelling. Estimates of modern TOC mass accumulation rates (MAR) show that approximately 85% of the total is accumulating on the shelf. TOC MAR were calculated, mapped, and budgeted for the Holocene and for the Last Glacial Maximum (LGM) using 19 sediment cores from the continental slope. During the LGM, centers of deposition and production have migrated offshore with respect to their Holocene positions. TOC accumulation on the continental slope was approximately 84% higher during the LGM than during the Holocene, possibly reflecting enhanced productivity. The TOC distribution patterns and sediment echo sounding data suggest that undercurrents strongly influence the sedimentation off Namibia. Winnowing and focusing result in great lateral heterogeneity of sedimentation rates and sediment properties. Individual cores therefore do not necessarily reflect general changes in export production. These results highlight the need for detailed regional studies based on a large number of sediment cores for highly heterogeneous high-productivity areas in order to derive general statements on total fluxes.

Communities and microhabitats of living benthic foraminifera from the tropical East Atlantic

Living (Rose Bengal stained) benthic foraminifera were collected with a multicorer from six stations between 2°N and 12°S off West Africa. The foraminiferal communities in the investigated area reflect the direct influence of different productivity regimes, and are characterized by spatially and seasonally varying upwelling activity. At five stations, foraminiferal abundance coincides well with the gradient of surface productivity. However, at one station off the Congo River, the influence of strong fresh water discharge is documented. Although this station lies directly in the center of an upwelling area, foraminiferal standing stocks are surprisingly low. It is suggested that the Congo discharge may induce a fractionation of the organic matter into small and light particles of low nutritional content, by contrast to the relatively fast-sinking aggregates found in the centers of high productivity areas. Quality and quantity of the organic matter seem to influence the distribution of microhabitats as well. The flux of organic carbon to the sea-floor controls the sequence of degradation of organic matter in sediment and the position of different redox fronts. The vertical foraminiferal stratification within sediment closely parallels the distribution of oxygen and nitrate in porewater, and reflects different nutritive strategies and adaptation to different types of organic matter. The epifauna and shallow infauna colonize oxygenated sediments where labile organic matter is available. The intermediate infauna (M. barleeanum) is linked to the zone of nitrate reduction in sediments where epifaunal and shallow infaunal species are not competitive anymore, and must feed on bacterial biomass or on metabolizable nutritious particles produced by bacterial degradation of more refractory organic matter. The deep infauna shows its maximum distribution in anoxic sediments, where no easily metabolizable organic matter is available.

Geochemistry of Fe-Mn crusts at seamounts in the Northwest Pacific

Hydrogenetic ferromanganese crusts were dredged from four seamounts in the western Pacific, OSM7, OSM2, Lomilik, and Lemkein, aligned in a NW-SE direction parallel to Pacific Plate movement. The crusts consist of four well-defined layers with distinct textural and geochemical properties. The topmost layer 1 is relatively enriched in Mn, Co, Ni, and Mo compared to the underlying layer 2, which is relatively enriched in Al, Ti, K, and Rb and Cu, Zn, and excess Ba. Textural and geochemical properties of layer 2 suggest growth conditions under high biogenic and detrital flux. Such conditions are met in the equatorial Pacific (i.e., between the Intertropical Convergence Zone (ITCZ) and equatorial high-productivity zone). Layer 2 likely formed when each seamount was beneath the equatorial Pacific along its back track path. On the other hand, layer 1 probably started to grow after seamounts moved northwest from the ITCZ. This interpretation is consistent with the thickness of layer 1 across the four crusts, which increases to the northwest. Ages of the layer 1-layer 2 boundary in each crust, a potential proxy for northern margin of the ITCZ, also increase to the northwest at 17, 11, 8, and 5 Ma for OSM7, OSM2, Lomilik, and Lemkein, respectively. Assuming Pacific Plate motion of 0.3°/Myr, the seamounts were located at 12°N, 11°N, 9°N, and 8°N at the time of boundary formation. This result suggests that the north edge of the ITCZ has shifted south since the middle Miocene in the western Pacific, which agrees with information from the eastern Pacific.

Biogeochemistry of the Eastern Weddell Gyre

The Southern Ocean (SO) plays a key role in modulating atmospheric CO2 via physical and biological processes. However, over much of the SO, biological activity is iron-limited. New in situ data from the Antarctic zone south of Africa in a region centered at -20°E - 25°E reveal a previously overlooked region of high primary production, comparable in size to the northwest African upwelling region. Here, sea ice together with enclosed icebergs is channeled by prevailing winds to the eastern boundary of the Weddell Gyre, where a sharp transition to warmer waters causes melting. This cumulative melting provides a steady source of iron, fuelling an intense phytoplankton bloom that is not fully captured by monthly satellite production estimates. These findings imply that future changes in sea-ice cover and dynamics could have a significant effect on carbon sequestration in the SO.

Conservation of freshwater thermal habitats for Pacific salmon in a changing climate

Thesis (Ph.D.)--University of Washington, 2016-06

Foreign manufacturing, regional agglomeration and technical efficiency in UK industries: a stochastic production frontier approach

The paper examines howfar foreign manufacturing investment in UK industries, together with the spatial agglomeration of those industries, affect technical efficiency. The paper links research on the estimation of technical efficiency,with those literatures demonstrating the economies associated with foreign direct investment and spatial agglomeration. The methodology involves estimation of a stochastic production frontier with random components associated with industry technical inefficiency, and a standard error. The paper also explores whether the degree of foreign involvement has a greater impact on technical efficiency where the domestic industry sector is characterized by comparatively high productivity and spatial agglomeration. The policy implications of the analysis are discussed.

Pichia pastoris regulates its gene-specific response to different carbon sources at the transcriptional, rather than the translational, level

Background: The methylotrophic, Crabtree-negative yeast Pichia pastoris is widely used as a heterologous protein production host. Strong inducible promoters derived from methanol utilization genes or constitutive glycolytic promoters are typically used to drive gene expression. Notably, genes involved in methanol utilization are not only repressed by the presence of glucose, but also by glycerol. This unusual regulatory behavior prompted us to study the regulation of carbon substrate utilization in different bioprocess conditions on a genome wide scale. Results: We performed microarray analysis on the total mRNA population as well as mRNA that had been fractionated according to ribosome occupancy. Translationally quiescent mRNAs were defined as being associated with single ribosomes (monosomes) and highly-translated mRNAs with multiple ribosomes (polysomes). We found that despite their lower growth rates, global translation was most active in methanol-grown P. pastoris cells, followed by excess glycerol- or glucose-grown cells. Transcript-specific translational responses were found to be minimal, while extensive transcriptional regulation was observed for cells grown on different carbon sources. Due to their respiratory metabolism, cells grown in excess glucose or glycerol had very similar expression profiles. Genes subject to glucose repression were mainly involved in the metabolism of alternative carbon sources including the control of glycerol uptake and metabolism. Peroxisomal and methanol utilization genes were confirmed to be subject to carbon substrate repression in excess glucose or glycerol, but were found to be strongly de-repressed in limiting glucose-conditions (as are often applied in fed batch cultivations) in addition to induction by methanol. Conclusions: P. pastoris cells grown in excess glycerol or glucose have similar transcript profiles in contrast to S. cerevisiae cells, in which the transcriptional response to these carbon sources is very different. The main response to different growth conditions in P. pastoris is transcriptional; translational regulation was not transcript-specific. The high proportion of mRNAs associated with polysomes in methanol-grown cells is a major finding of this study; it reveals that high productivity during methanol induction is directly linked to the growth condition and not only to promoter strength.

Changes in Caribbean paleoproductivity, diversity and benthic foraminiferal test size caused by the Neogene closing of the tropical Atlantic-Pacific ocean gateway

The purpose of this study was to test 3 hypotheses: (a) that late Miocene to early Pliocene constriction and complete closure of the Central American Seaway (CAS), connecting tropical Atlantic and East Equatorial Pacific (EEP) oceans, caused decreased productivity in the Caribbean, due to reduced coastal upwelling and an end to the connection with high-productivity Pacific waters, (b) reduced paleoproductivity resulted in decreased diversity in the Caribbean and, (c) this decreased availability of food (reduced paleoproductivity) was responsible for larger mean test size in the three most common benthic foraminiferal species Epistominella exigua, Oridorsalis umbonatus and Globocassidulina subglobosa. ^ These are tested by applying correlation analysis to 7 groups of paleoceanographic proxies, 3 indices of diversity measures and mean test size data from the Caribbean Ocean Drilling Project Site 999, to 47 core samples for the interval between 8.3-2.5 Ma. Results are compared with published Caribbean and Pacific deep-sea records. ^ The Caribbean, between 8.3-7.9 Ma, experienced reduced current velocity and lower ventilation of bottom waters. Thereafter, until 4.2 Ma, the seasonality of phytodetritus input increased and ventilation further reduced. From 4.2-2.5 Ma, paleoproductivity decreased, current velocity reduced, ventilation improved, and the seasonality of phytodetrital input decreased dramatically. The benthic foraminiferal diversity followed the same trend as paleoproductivity. Individual correlation analysis between mean test size of benthic foraminiferal species Epistominella exigua, Oridorsalis umbonatus and Globocassidulina subglobosa and paleoceanographic proxies yielded a positive and significant relationship with paleoproductivity. However, a combined datasets of all 3 species yielded a negative and significant relationship with species abundance. ^ Thus, the study concludes that (a) the gradual closure of the CAS led Caribbean diversity and paleoproductivity to decrease abruptly at 7.9 Ma, when the nutrient-rich Pacific deep waters were cut off, and then, again with the complete closure of the seaway at 4.2 Ma, (b) diversity and paleoproductivity are positively correlated in the Caribbean and (c) that the availability of food is an overriding factor that influences mean test size; lower availability of food and decreased abundance leads to larger test size. ^

Spatial variability in Florida Bay particulate organic matter composition: combining flow cytometry with stable isotope analyses

Long-term management plans for restoration of natural flow conditions through the Everglades increase the importance of understanding potential nutrient impacts of increased freshwater delivery on Florida Bay biogeochemistry. Planktonic communities respond quickly to changes in water quality, thus spatial variability in community composition and relationships to nutrient parameters must be understood in order to evaluate future downstream impacts of modifications to Everglades hydrology. Here we present initial results combining flow cytometry analyses of phytoplankton and bacterial populations (0.1–50 μm size fraction) with measurements of δ13C and δ15N composition and dissolved inorganic nutrient concentrations to explore proxies for planktonic species assemblage compositions and nutrient cycling. Particulate organic material in the 0.1–50 μm size fraction was collected from five stations in Northeastern and Western Florida Bay to characterize spatial variability in species assemblage and stable isotopic composition. A dense bloom of the picocyanobacterium, Synechococcus elongatus, was observed at Western Florida Bay sites. Smaller Synechococcus sp. were present at Northeast sites in much lower abundance. Bacteria and detrital particles were also more abundant at Western Florida Bay stations than in the northeast region. The highest abundance of detritus occurred at Trout Creek, which receives freshwater discharge from the Everglades through Taylor Slough. In terms of nutrient availability and stable isotopic values, the S. elongatus population in the Western bay corresponded to low DIN (0.5 μM NH 4 + ; 0.2 μM NO 3 − ) concentrations and depleted δ15N signatures ranging from +0.3 to +0.8‰, suggesting that the bloom supported high productivity levels through N2-fixation. δ15N values from the Northeast bay were more enriched (+2.0 to +3.0‰), characteristic of N-recycling. δ13C values were similar for all marine Florida Bay stations, ranging from −17.6 to −14.4‰, however were more depleted at the mangrove ecotone station (−25.5 to −22.3‰). The difference in the isotopic values reflects differences in carbon sources. These findings imply that variations in resource availability and nutrient sources exert significant control over planktonic community composition, which is reflected by stable isotopic signatures.