Western Subarctic North Pacific Research Articles

Decadal decreasing trend in biological carbon pump estimated from 234Th in the western subarctic North Pacific

We measured thorium-234 (234Th), particulate organic carbon (POC), and chlorophyll a in the western subarctic North Pacific surface layer in 1997–2008. 234Th, POC, and chlorophyll a in the surface layer showed clear seasonal changes. As a result of approximation by using a trigonometric function, 234Th export flux estimated from the deficiency of 234Th relative to 238U and the ratio of POC to particulate 234Th (POC/234ThP) indicated constant and decreasing trends in the study area from 1997 to 2008, respectively. POC export flux estimated from 234Th decreased in the study area, showing that the strength of the biological carbon pump in this region declined in the studied decade. The ratio of POC/chlorophyll a decreased during the term. These trends were possibly caused by a decline in not the biomass but particle size of phytoplankton. Our results suggest that not only the biological carbon pump but also phytoplankton community structure change in the western subarctic Pacific Ocean in a decade.

Determination of ultra-trace Te species in open ocean waters based on Mg(OH)2 coprecipitation, anion exchange resin column separation and inductively coupled plasma sector-field mass spectrometry using a 125Te-enriched isotope spike

We present a method for the determination of ultra-trace Te species (Te(IV) and Te(VI)) in open ocean waters. The proposed method is based on Mg(OH)2 coprecipitation, anion exchange resin column separation and inductively coupled plasma sector-field mass spectrometry (ICPSFMS) using a 125Te-enriched isotope spike. The largest advantage of the method is that the use of the spike allows accurate and precise determination when it combines with either isotope dilution or recovery correction. Tellurium-IV and VI are preconcentrated in a Mg(OH)2 precipitate and separated mutually by an anion exchange resin column. Te(IV) is retained to the column, while Te(VI) passing through the column is recovered by a subsequent column procedure after reduction of Te(VI) to Te(IV). Te(IV) is successfully eluted with a small amount of 0.01 M HCl. The additional merit of using this eluent is elimination of components that result in a memory effect during the measurement of Te(IV). Possible mass spectral interference on Te(IV) can be excluded by adjusting the mass window, and the Te(IV) concentrations determined by this approach agree well with those independently obtained by an oxidation procedure which removes the interference. The accuracy of the proposed method is verified with homemade standard seawater for which the measured concentrations agree well with results calculated from the value of the standard solution. Procedural blanks for Te(IV) and Te(VI) are 1.5 ± 0.9 pg kg−1 (n = 11) and 1.3 ± 0.9 pg kg−1 (n = 11) with corresponding overall detection limits of 3.0 pg kg−1 and 2.8 pg kg−1, respectively. Using the method, we have clarified vertical profiles of Te(IV) and Te(VI) in the subarctic western North Pacific for the first time.

Distribution and stoichiometry of Al, Mn, Fe, Co, Ni, Cu, Zn, Cd, and Pb in the Seas of Japan and Okhotsk

Trace metals play an important role in marine biogeochemistry. However, the detailed distribution of trace metals in the Seas of Japan and Okhotsk remain unknown. Herein, we report the full-depth and section distributions of the dissolved (d), total dissolvable (td), and labile particulate (lp) phases of Al, Mn, Fe, Co, Ni, Cu, Zn, Cd, and Pb in seawater samples collected from the Seas of Japan and Okhotsk during the GEOTRACES-Japan program. High lpM/tdM ratios suggest active scavenging in the Seas of Japan and Okhotsk. The lpAl and lpFe species were dominant in tdAl and tdFe, showing a strong positive correlation to each other in the both the Seas of Japan and Okhotsk. The maximum lpAl, lpFe, lpMn, and lpCo concentrations were observed at σ0 = 26.8–27.0 for dense shelf water (DSW) in the Sea of Okhotsk, implying that a metal source is laterally transported to the western subarctic North Pacific. The high enrichment factors of dMn, dCo, dNi, dCu, dZn, dCd, and dPb in the surface water of the Sea of Japan were attributed to anthropogenic emissions of these metals. In addition, temporal variations of the trace metals were examined in the Sea of Japan. The distribution of dCd did not change significantly during 1984–2010; however, data suggest that lpMn and lpFe exhibited dynamic temporal and spatial variations.

El Niño‐Related Vertical Mixing Enhancement Under the Winter Mixed Layer at Western Subarctic North Pacific Station K2

AbstractWe conducted a current velocity and hydrographic time series observation in the upper ocean by deploying a mooring buoy at station K2 (N, E) from July 2015 to June 2019 to understand the physical conditions that affect the acidification of the winter mixed layer (ML) water in the western subarctic North Pacific. Disturbances with substantial current velocity shear were excited by the wind changes remotely related to the 2015/2016 and 2018/2019 El Niño events. The density stratification beneath the ML was also weakened in winter, particularly the El Niño years. Based on the bulk Richardson number mixing scheme, we found that the El Niño‐related strengthening of the current velocity shear enhanced the vertical diffusivity beneath the winter ML driven by the near‐inertial current velocity variation. The weakening of the density stratification is ineffective in enhancing the vertical diffusivity. It is suggested that the El Niño‐related large vertical mixing accelerates the acidification of the winter ML water through the enhancement of the entrainment of the deep water that is rich in dissolved inorganic carbon.

Impact of mesoscale eddies on particulate organic carbon flux in the western subarctic North Pacific

Impact of mesoscale eddies on particulate organic carbon flux in the western subarctic North Pacific

Could Artificial Downwelling/Upwelling Mitigate Oceanic Deoxygenation in Western Subarctic North Pacific?

Subpolar gyre regions such as the Western Subarctic North Pacific (WSNP) contain sluggish, low-oxygen water, and are threatened by loss of oxygen (deoxygenation). Our simulations under RCP 8.5 emission scenario suggest that installing pipes to induce artificial downwelling and upwelling (AD and AU) provides short-term solutions to combat deoxygenation in the WSNP. With no engineering, the WSNP's subsurface oxygen decreases by 30–100 mmol/m3 by the year 2100. Continuous implementation of AD and AU instead counters this declining trend, and AD is more effective than AU. The oxygenation effect is primarily a consequence of how the two engineering schemes vertically redistribute oxygen via physical processes. AD directly improves oxygen at depth via advecting surface water toward the ocean interior and subsequent enhanced pycnocline mixing, and AU does so via generating compensatory downwelling outside of the pipes. Both schemes take near 40 years to complete the oxygenation. After that, oxygen reaches a new equilibrium state in the WSNP with no further improvement by the engineering. AD and AU both strongly increase primary production surrounding the deployment sites, but lead only to weak enhancement of aerobic respiration in subsurface water and thus a minor impact on the oxygenation. Other unwanted environmental side effects are negligible compared to those caused by rapid climate change within this century, including outgassing of carbon dioxide, pH decrease, and precipitation reduction.

Chemical characteristics of sulfur-containing aerosol particles across the western North Pacific and the Arctic Ocean

The characteristics of aerosol methanesulfonic acid (MSA), non-sea-salt sulfate (nss-SO42−) and MSA/nss-SO42− ratio, were measured at high-time resolution (1 h) over the East sea/sea of Japan (R1, 31° - 47°N, 121° - 145°E), western North Pacific Ocean (R2, 47° - 58°N, 145° - 176°E), subarctic western North Pacific (R3, 58° - 66°N, 176°E - 169°W) and Arctic Ocean (R4, 66° - 85°N, 150–174°W) during July–September 2018. The highest MSA levels were observed in R2, with an average of 0.22 ± 0.18 μg m−3. These levels were 3–10 times higher than in other regions, correlating with biological productivity. The concentration of nss-SO42− decreased with distance from Eurasia. The MSA/nss-SO42− ratios were 0.10 ± 0.27, 0.38 ± 0.43, 0.30 ± 0. 36 and 0.39 ± 0.48 over R1, R2, R3 and R4, respectively. Biological sources were larger contributors of nss-SO42− in R2 while nss-SO42− were affected more by continental anthropogenic sources in R1. Positive and negative relationships between the MSA/nss-SO42− ratio and temperature were observed, indicating that temperature may have a different influence on the MSA/nss-SO42− ratio. Four high level regions of MSA were observed over the R4, wind speed and temperature were critical factors that affected MSA levels over the R4. Significant depletion of chloride was observed and areas that were more affected by terrestrial sources had a greater chlorine depletion in this study, which means that the magnitude of the chlorine depletion followed the order R1 > R3 > R2 > R4.

Observations of anticyclonic eddies in the western subarctic North Pacific

We conducted a ship observation of two anticyclonic mesoscale eddies (Eddies A and B) between June and July 2016 in the offshore region of the western subarctic North Pacific to clarify the impact of anticyclonic eddies on biological production. Analysis of satellite data reveals that Eddy A formed south of the Alaskan Peninsula on May 2011 while Eddy B formed in the observation area on November 2015. Iron concentration was low both inside and outside Eddies A and B, and there were no large differences between chlorophyll a (chl-a) concentration inside and outside eddies. Satellite-derived chl-a concentration inside Eddy A had been high compared with that in the surrounding area when the eddy was located along the Aleutian Islands and just after it detached from the islands, suggesting that iron originating from the coastal region was consumed during the two-year journey from the coastal region to the observation area. Field measurements also reveal that vertical diffusivity was low both inside and outside Eddies A and B, which could be another factor underlying low iron both inside and outside Eddies A and B and small differences between chl-a concentration inside and outside the eddies.

Seasonal occurrence and vertical distribution of larval and juvenile northern smoothtongue,Leuroglossus schmidti(Pisces, Bathylagidae), in the western subarctic Pacific

AbstractWe investigated the seasonal occurrence and vertical distribution/migration of larval and juvenile northern smoothtongue,Leuroglossus schmidti(Pisces, Bathylagidae), in oceanic areas of the western subarctic Pacific. This species was the most abundant larval fish and one of the most abundant juvenile/adult fish at the study site. Larval recruitment of this species occurred in early March to mid-October. Larvae were found, however, throughout all seasons, suggesting that some had overwintered. The highest abundance (5.8 individuals m–2) of larvae was in summer. During summer and autumn, hatching mainly occurred at 100–150-m depth and larvae migrated toward the surface with growth. Once larvae reached about 20 mm in length, they moved below 100 m, and larger larvae (over 24 mm) exhibited diel vertical migration. The abundance of juveniles integrated through the water column was relatively constant (0.3 individuals m–2) throughout the study period. They were collected from below 300 m during the day, and part of the population (13–38%) swam into the epipelagic layer at night. The information on early stages ofL. schmidtipresented here provides a basis for future investigations of mesopelagic fish ecology in oceanic areas of the western subarctic North Pacific Ocean.

Response of N2O production rate to ocean acidification in the western North Pacific.

Ocean acidification induced by the increase of anthropogenic CO2 emissions has a profound impact on marine organisms and biogeochemical processes.1 The response of marine microbial activities to ocean acidification might play a crucial role in the future evolution of air-sea fluxes of biogenic gases such as nitrous oxide (N2O), a strong greenhouse gas and the dominant stratospheric ozone-depleting substance.2 Here, we examine the response of N2O production from nitrification to acidification in a series of incubation experiments conducted in subtropical and subarctic western North Pacific. The experiments show that, when pH was reduced, the N2O production rate during nitrification measured at subarctic stations increased significantly whereas nitrification rates remained stable or decreased. Contrary to what was previously thought, these results suggest that the effect of ocean acidification on N2O production during nitrification and nitrification rates are likely uncoupled. Collectively these results suggest that, if seawater pH continues to decline at the same rate, ocean acidification could increase the marine N2O production during nitrification in subarctic North Pacific by 185 to 491% by the end of the century.

Wind-driven decadal sea surface height and main pycnocline depth changes in the western subarctic North Pacific

The northward shrinkage of the North Pacific western subarctic gyre (WSAG) in the early 2000s is associated with a sea surface height (SSH) elevation and is correlated to sea surface wind stress change. By using a Rossby wave model forced by wind stress, which computes the component variations due to the barotropic and first to fourth baroclinic modes, we estimated decadal changes in SSH and main pycnocline depth in the subpolar region. Realistic decadal SSH elevation and deepening of the main pycnocline depth associated with the northward shrinkage of the western subarctic gyre from the late 1990s to the mid-2000s were reproduced by the model. The sea surface elevation was caused primarily by the barotropic Rossby wave response to the relaxation of the Ekman suction due to the attenuation of the Aleutian Low by frequent La Niña occurrences after the late 1990s in addition to the long-term weakening of the westerly wind. The northward shrinkage of the WSAG was found to be associated with the intensification of an anticyclonic circulation centered around 43–44 ∘ N, 170–175 ∘ E. The westerly wind weakening deepened the main pycnocline in the western subarctic region through the baroclinic Rossby wave mode response to the wind stress change, which mostly accounts the equivalent halocline deepening at station K2 (47 ∘ N, 160 ∘ E). While the first baroclinic mode variation of the water density significantly attenuates during propagation, the higher mode variations, particularly the second and third mode variations, are locally excited through a quasi-resonant amplification mechanism and have profound impacts on the depth of the upper main pycnocline.

Atmospheric Dry Deposition of Water-Soluble Nitrogen to the Subarctic Western North Pacific Ocean during Summer

To estimate dry deposition flux of atmospheric water-soluble nitrogen (N), including ammonium (NH4+), nitrate (NO3−), and water-soluble organic nitrogen (WSON), aerosol samples were collected over the subarctic western North Pacific Ocean in the summer of 2016 aboard the Korean icebreaker IBR/V Araon. During the cruise, concentrations of NH4+, NO3−, and WSON in bulk (fine + coarse) aerosols ranged from 0.768 to 25.3, 0.199 to 5.94, and 0.116 to 14.7 nmol m−3, respectively. Contributions of NH4+, NO3−, and WSON to total water-soluble N represented ~74%, ~17%, and ~9%, respectively. Water-soluble N concentrations showed a strong gradient from the East Asian continent to the subarctic western North Pacific Ocean, indicating that water-soluble N species were mainly derived from anthropogenic or terrestrial sources. During sea fog events, coarse mode NO3− was likely to be scavenged more efficiently by fog droplets than fine mode NO3−; besides, WSON was detected only in fine mode, suggesting that there may have been a significant influence of sea fog on WSON, such as the photochemical conversion of WSON into inorganic N. Mean dry deposition flux for water-soluble total N (6.3 ± 9.4 µmol m−2 d−1) over the subarctic western North Pacific Ocean was estimated to support a minimum carbon uptake of 42 ± 62 µmol C m−2d−1 by using the Redfield C/N ratio of 6.625.

Nitrate Isotope Distribution in the Subarctic and Subtropical North Pacific

AbstractNitrogen and oxygen isotopic compositions of nitrate (δ15NNitrate and δ18ONitrate) were determined along 47°N in the subarctic North Pacific and 149°E in the western North Pacific. In the western subarctic gyre, known as a high‐nutrient, low‐chlorophyll region, the δ15NNitrate and the differences between δ15NNitrate and δ18ONitrate (herein Δ(15–18)) in the intermediate water were significantly lower than in the surrounding area. The 15N‐depleted nitrate was generated by nitrified nitrate from the remineralization of organic matter produced by the partial consumption of the surface nitrate pool. Between the western subarctic gyre and the Alaskan gyre, there was a 0.2‰ increase in δ15NNitrate and a 0.3‰ increase in Δ(15–18) in the intermediate water associated with a 7.2‐μM decrease in the nitrate concentration at the surface. Assuming Rayleigh distillation kinetics, the increase in utilization of the surface nitrate pool between the western subarctic gyre and the Alaskan gyre (from 29% to 85%) corresponds to a change in δ15N of organic matter from 2.6‰ to 4.9‰. The 15N enrichment of intermediate nitrate toward the east resulted from the increase in utilization. Moreover, the 15N‐depleted nitrate in the mixed‐layer water of the western subtropical gyre was affected by N2 fixation in the Kuroshio water, whereas the 15N‐enriched nitrate in the intermediate water at the western margin of North America was affected by water column denitrification. The δ15N sediment record in the western subarctic North Pacific likely reflects past changes in the high‐nutrient, low‐chlorophyll region but may also be controlled by water column denitrification and N2 fixation.

Basin-scale distribution of NH4 + and NO2 − in the Pacific Ocean

We used more than 25,000 nutrient samples to elucidate for the first time basin-scale distributions and seasonal changes of surface ammonium (NH4 +) and nitrite (NO2 −) concentrations in the Pacific Ocean. The highest NH4 +, NO2 −, and nitrate (NO3 −) concentrations were observed north of 40°N, in the coastal upwelling region off the coast of Mexico, and in the Tasman Sea. NH4 + concentrations were elevated during May–October in the western subarctic North Pacific, May–December in the eastern subarctic North Pacific, and June–September in the subtropical South Pacific. NO2 − concentrations were highest in winter in both hemispheres. The seasonal cycle of NH4 + was synchronous with NO2 −, NO3 −, and satellite chlorophyll a concentrations in the western subtropical South Pacific, whereas it was synchronous with chlorophyll-a but out of phase with NO2 − and NO3 − in the subarctic regions.

Importance of E kman transport and gyre circulation change on seasonal variation of surface dissolved iron in the western subarctic N orth P acific

Iron (Fe) is an essential nutrient for marine phytoplankton and it constitutes an important element in the marine carbon cycle in the ocean. This study examined the mechanisms controlling seasonal variation of dissolved Fe (dFe) in the western subarctic North Pacific (WSNP), using an ocean general circulation model coupled with a simple biogeochemical model incorporating a dFe cycle fed by two major sources (atmospheric dust and continental shelf sediment). The model reproduced the seasonal cycle of observed concentrations of dFe and macro nutrients at the surface in the Oyashio region with maxima in winter (February–March) and minima in summer (July–September), although the simulated seasonal amplitudes are a half of the observed values. Analysis of the mixed-layer dFe budget indicated that both local vertical entrainment and lateral advection are primary contributors to the wintertime increase in dFe concentration. In early winter, strengthened northwesterly winds excite southward Ekman transport and Ekman upwelling over the western subarctic gyre, transporting dFe-rich water southward. In mid- to late winter, the southward western boundary current of the subarctic gyre and the outflow from the Sea of Okhotsk also bring dFe-rich water to the Oyashio region. The contribution of atmospheric dust to the dFe budget is several times smaller than these ocean transport processes in winter. These results suggest that the westerly wind-induced Ekman transport and gyre circulation systematically influence the seasonal cycle of WSNP surface dFe concentration.

Bio-availability of iron derived from subarctic first-year sea ice

Sea ice contains high concentrations of iron (Fe), and melting of sea ice is one of the possible processes that supplies Fe to surface waters (e.g., Lannuzel et al., 2007). To assess the biological availability (bio-availability) of Fe in sea ice, a shipboard bottle incubation experiment (Exp. I) was conducted with surface water from the western subarctic North Pacific and Fe derived from sea ice collected from the Sea of Okhotsk, as compared to Fe derived from inorganic Fe (FeCl3). Addition of FeCl3 solution stimulated the growth of phytoplankton relative to a control treatment. Addition of sea ice meltwater enhanced the particulate labile Fe (>0.22μm) by 112nM and 0.26nM of dissolved Fe (<0.22μm) in the incubated seawater and allowed the phytoplankton to grow, especially large (>10μm) phytoplankton, at their maximum growth rate in the incubated seawater. In contrast, addition of desferrioxamine B (DFB) plus sea ice meltwater, in which the strong ligand DFB reduced the bio-availability of Fe, significantly depressed the growth of the large and small (0.8–10μm) phytoplankton relative to the control. These results clearly demonstrated that Fe stored in the sea ice stimulates growth of the phytoplankton. We carried out another shipboard bottle incubation experiment (Exp. II) to determine the relationship between concentrations of Fe and phytoplankton growth rates by adding FeCl3 solution at a series of concentrations to the seawater. The results indicated that the amount of dissolved Fe that the incubation bottles received from addition of the sea ice meltwater in Exp. I was insufficient to saturate the growth rate of the large phytoplankton. Therefore, the maximum growth rate achieved by the large phytoplankton can be accounted for by the use of the particulate labile Fe released from the sea ice meltwater. These results indicate that Fe stored in sea ice (mainly in particulate form) is bio-available and can contribute to phytoplankton growth when it is released into surface waters in the spring.

Seasonal variations in the nitrogen isotopic composition of settling particles at station K2 in the western subarctic North Pacific

Intensive observations using hydrographical cruises and moored sediment trap deployments during 2010 and 2012 at station K2 in the North Pacific Western Subarctic Gyre (WSG) revealed seasonal changes in δ 15N of both suspended and settling particles. Suspended particles (SUS) were collected from depths between the surface and 200 m; settling particles by drifting sediment traps (DST; 100–200 m) and moored sediment traps (MST; 200 and 500 m). All particles showed higher δ 15N values in winter and lower in summer, contrary to the expected by isotopic fractionation during phytoplankton nitrate consumption. We suggest that these observed isotopic patterns are due to ammonium consumption via light-controlled nitrification, which could induce variations in δ 15N(SUS) of 0.4–3.1 ‰ in the euphotic zone (EZ). The δ 15N(SUS) signature was reflected by δ 15N(DST) despite modifications during biogenic transformation from suspended particles in the EZ. δ 15N enrichment (average: 3.6 ‰) and the increase in C:N ratio (by 1.6) in settling particles suggests year-round contributions of metabolites from herbivorous zooplankton as well as TEPs produced by diatoms. Accordingly, seasonal δ 15N(DST) variations of 2.4–7.0 ‰ showed a significant correlation with primary productivity (PP) at K2. By applying the observed δ 15N(DST) vs. PP regression to δ 15N(MST) of 1.9–8.0 ‰, we constructed the first annual time-series of PP changes in the WSG. This new approach to estimate productivity can be a powerful tool for further understanding of the biological pump in the WSG, even though its validity needs to be examined carefully.

Depth-dependent and seasonal variability in archaeal community structure in the subarctic and subtropical western North Pacific

Archaea represent an important component of marine picoplankton. Marine archaeal communities are generally dominated by Marine Group I (MGI) and Marine Group II (MGII), which are affiliated to the phyla Thaumarchaeota and Euryarchaeota. We used pyrosequencing-based analysis of archaeal 16S rRNA genes to investigate archaeal community composition throughout the water column (0–5000 m) at the subarctic (station K2) and subtropical (station S1) stations in western North Pacific. Sampling was conducted on four occasions between 2010 and 2011 to examine seasonal variability in community composition. Four phylogenetic groups (MGI-α, MGI-γ, MGII-α, and MGII-β) belonging to Thaumarchaeota and Euryarchaeota were found throughout the water column, but the relative abundance of each group varied depending on depth. In the surface water, archaeal communities displayed marked seasonal variability at both stations, whereas in deeper waters (300–5000 m), temporal variability in community composition was less evident. Sequences similar to those of the uncultured Marine Benthic Group A, to date found only in marine sediments, were detected throughout the water column at station S1.

Dichothermal layer deepening in relation with halocline depth change associated with northward shrinkage of North Pacific western subarctic gyre in early 2000s

In the western subarctic North Pacific, a wind-driven cyclonic circulation, called the western subarctic gyre (WSAG), exists. We examined year-to-year changes of the gyre and hydrographic structures, applying the altimetry-based gravest empirical mode (AGEM) method to hydrographic and altimetric sea surface height (SSH) data, and relation to the in situ variation of the temperature minimum layer, i.e., the dichothermal layer, depth at station K2 (47∘ N, 160∘ E). The AGEM-based geostrophic volume transport and the streamfunction of the WSAG in the top 1000-dbar layer show that the gyre changes substantially. From the late 1990s to the mid-2000s, the gyre shrunk northward. Due to the shrinkage, the halocline bottom, which is equivalent to the top of the main pycnocline, deepens at K2 outside the central part of the gyre. The downward displacement of the dichothermal layer at K2 was found to be significantly related to that of the underlying halocline due to the northward shrinkage of the WSAG.

Glacial-to-interglacial changes in nitrate supply and consumption in the subarctic North Pacific from microfossil-bound N isotopes at two trophic levels

Reduced nitrate supply to the subarctic North Pacific (SNP) surface during the last ice age has been inferred from coupled changes in diatom-bound δ15N (DB-δ15N), bulk sedimentary δ15N, and biogenic fluxes. However, the reliability of bulk sedimentary and DB-δ15N has been questioned, and a previously reported δ15N minimum during Heinrich Stadial 1 (HS1) has proven difficult to explain. In a core from the western SNP, we report the foraminifera-bound δ15N (FB-δ15N) in Neogloboquadrina pachyderma and Globigerina bulloides, comparing them with DB-δ15N in the same core over the past 25 kyr. The δ15N of all recorders is higher during the Last Glacial Maximum (LGM) than in the Holocene, indicating more complete nitrate consumption. N. pachyderma FB-δ15N is similar to DB-δ15N in the Holocene but 2.2‰ higher during the LGM. This difference suggests a greater sensitivity of FB-δ15N to changes in summertime nitrate drawdown and δ15N rise, consistent with a lag of the foraminifera relative to diatoms in reaching their summertime production peak in this highly seasonal environment. Unlike DB-δ15N, FB-δ15N does not decrease from the LGM into HS1, which supports a previous suggestion that the HS1 DB-δ15N minimum is due to contamination by sponge spicules. FB-δ15N drops in the latter half of the Bolling/Allerod warm period and rises briefly in the Younger Dryas cold period, followed by a decline into the mid-Holocene. The FB-δ15N records suggest that the coupling among cold climate, reduced nitrate supply, and more complete nitrate consumption that characterized the LGM also applied to the deglacial cold events.