Atsushi Tsuda

Synthesis of iron fertilization experiments: From the Iron Age in the Age of Enlightenment

Comparison of eight iron experiments shows that maximum Chl a, the maximum DIC removal, and the overall DIC/Fe efficiency all scale inversely with depth of the wind mixed layer (WML) defining the light environment. Moreover, lateral patch dilution, sea surface irradiance, temperature, and grazing play additional roles. The Southern Ocean experiments were most influenced by very deep WMLs. In contrast, light conditions were most favorable during SEEDS and SERIES as well as during IronEx-2. The two extreme experiments, EisenEx and SEEDS, can be linked via EisenEx bottle incubations with shallower simulated WML depth. Large diatoms always benefit the most from Fe addition, where a remarkably small group of thriving diatom species is dominated by universal response of Pseudo-nitzschia spp. Significant response of these moderate (10–30 μm), medium (30–60 μm), and large (>60 μm) diatoms is consistent with growth physiology determined for single species in natural seawater. The minimum level of “dissolved” Fe (filtrate < 0.2 μm) maintained during an experiment determines the dominant diatom size class. However, this is further complicated by continuous transfer of original truly dissolved reduced Fe(II) into the colloidal pool, which may constitute some 75% of the “dissolved” pool. Depth integration of carbon inventory changes partly compensates the adverse effects of a deep WML due to its greater integration depths, decreasing the differences in responses between the eight experiments. About half of depth-integrated overall primary productivity is reflected in a decrease of DIC. The overall C/Fe efficiency of DIC uptake is DIC/Fe ∼ 5600 for all eight experiments. The increase of particulate organic carbon is about a quarter of the primary production, suggesting food web losses for the other three quarters. Replenishment of DIC by air/sea exchange tends to be a minor few percent of primary CO2 fixation but will continue well after observations have stopped. Export of carbon into deeper waters is difficult to assess and is until now firmly proven and quite modest in only two experiments.

The decline and fate of an iron-induced subarctic phytoplankton bloom

Iron supply has a key role in stimulating phytoplankton blooms in high-nitrate low-chlorophyll oceanic waters. However, the fate of the carbon fixed by these blooms, and how efficiently it is exported into the oceans interior, remains largely unknown. Here we report on the decline and fate of an iron-stimulated diatom bloom in the Gulf of Alaska. The bloom terminated on day 18, following the depletion of iron and then silicic acid, after which mixed-layer particulate organic carbon (POC) concentrations declined over six days. Increased particulate silica export via sinking diatoms was recorded in sediment traps at depths between 50 and 125 m from day 21, yet increased POC export was not evident until day 24. Only a small proportion of the mixed-layer POC was intercepted by the traps, with more than half of the mixed-layer POC deficit attributable to bacterial remineralization and mesozooplankton grazing. The depletion of silicic acid and the inefficient transfer of iron-increased POC below the permanent thermocline have major implications both for the biogeochemical interpretation of times of greater iron supply in the geological past, and also for proposed geo-engineering schemes to increase oceanic carbon sequestration.

Functional roles of interzonal migrating mesozooplankton in the western subarctic Pacific

Grazing experiments and production estimation based on life-history analysis of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were carried out in the Oyashio region to understand the carbon flows associated with the interzonal migrating copepods. These copepods, and also Eucalanus bungii, fed on nano- and micro-sized organisms non-selectively throughout the season. However, diatoms were the dominant food resource until May and organisms, such as ciliates were the major resource after May. Daily growth rate was estimated from the Ikeda-Motoda, Huntley-Lopez and Hirst-Sheader models. Since the growth rates were considered to be overestimates for the Huntley-Lopez model and underestimates for the other two models, we applied the weight-specific growth rates previously reported for these species in the Bering Shelf. Surface biomass of Neocalanus increased rapidly in June during the appearance of C5, and a successive increase of overwintering stock was evident in the deeper layer. The deep biomass decreased gradually from September to May during the dormant and reproduction period. N. cirstatus has the largest annual mean biomass (2.3 gC m -2 ), followed by N. plumchrus (1.1) and N. flemingeri (0.4). Daily production rate of Neocalanus varied from 0.4 to 363.4 mgC m -2 day -1 , to which N. cristatus was the largest contributor. Annual production was estimated as 11.5 gC m -2 year -1 for N. cirstatus, 5.7 for N. plumchrus and 2.1 for N. flemingeri, yielding annual P/B ratio of 5 for each species. The annual production of Neocalanus accounted for 13.2% of the primary production in the Oyashio region. Their fecal pellets were estimated to account for 14.9% (0.7 gC m -2 year -1 ) of sinking flux of organic carbon at 1000-m depth. Moreover, their export flux by ontogenetic vertical migration, which is not measured by sediment trap observations, is estimated to be 91.5% (4.3 gC m -2 year -1 ) of carbon flux of sinking particles at 1000-m depth. These results suggest the important role of interzonal migrating copepods in the export flux of carbon.

IN SITU GRAZING RATE OF THE COPEPOD POPULATION IN THE WESTERN SUBARCTIC NORTH PACIFIC DURING SPRING

Time series sampling with a multi-layer plankton sampler was carried out in the western subarctic North Pacific during spring 1991. Neocalanus cristatus, N. flemingeri and Eucalanus bungii dominated and accounted for 88.5% of the copepod population in volume. Neocalanus spp. were distributed in the upper mixed layer, while E. bungii was mainly distributed between 120 and 300 m throughout the day and night. In contrast, Metridia pacifica, Pleuromamma scutullata and Gaetanus simplex showed clear diel vertical migration. Grazing activities were estimated simultaneously by gut fluorescence. Nocturnal grazing was observed for diel migrating species. Neocalanus spp. did not have a diel feeding rhythm and had relatively low gut fluorescence. E. bungii was considered to be dormant during the observation period. The estimated grazing rate of the copepod population on phytoplankton was 1.4 to 2.0% of the primary production while the metabolic requirement was 8.3 to 12.4% of the primary production. These facts suggest that the copepod population was unimportant as primary consumers and that microzooplankton plays a much more important role in sustaining low standing stock of phytoplankton and a high nutrient concentration in the western subarctic Pacific Ocean.

Influence of light intensity on diatom physiology and nutrient dynamics in the Oyashio region

Abstract The spring diatom bloom characterizes the plankton and nutrient dynamics in the Oyashio region, the westernmost part of the subarctic Pacific. Previous studies have shown that NO3 was not depleted during the spring bloom, and an increase in the consumption ratio of Si(OH)4 to NO3 (ΔSi(OH)4:ΔNO3) was observed as the spring bloom progressed. The increase in ΔSi(OH)4:ΔNO3 has been suggested to be caused by growth stresses of diatoms, e.g. light limitation by self-shading. In the present study, incubation experiments of sea-surface water from the Oyashio region under saturated irradiance showed that NO3 was depleted first and ΔSi(OH)4:ΔNO3 was more or less constant until the NO3 depletion occurred. The increase in ΔSi(OH)4:ΔNO3 was observed after the NO3 depletion had occurred in contrast with the field observation. This result of the increase in ΔSi(OH)4:ΔNO3 under saturated irradiance after NO3 depletion suggests that the in situ increase in ΔSi(OH)4:ΔNO3 before the NO3 depletion might be caused by light limitation for diatoms. Responses to a reduction in irradiance were examined using diatom species isolated from the Oyashio region. Variable responses to a reduced irradiance were observed for cell specific C, N, Si and chlorophyll a (Chl) contents. However, the examined diatom species showed similar tendencies for increases in Si:C and Si:N and decreases in C:Chl ratios with the reduction in irradiance. We conclude that light limitation changes the uptake ratio of nutrients and the elemental composition of diatoms and that light limitation is one of the factors influencing the physiology of diatoms and nutrient dynamics in the Oyashio region during the spring bloom.

Spawning season and migration of the myctophid fish Diaphus theta in the western North Pacific

Abstract The spawning season, spawning grounds, and migration of the myctophid fish Diaphus theta were studied in the western North Pacific, based on seasonal sampling and estimation of hatching dates. The peak abundance of larvae was observed in July in the transition waters between the Oyashio and Kuroshio fronts. The spawning season ranged from late March to early September, with a peak from May to July. Larvae and juveniles <40 mm in standard length were distributed in the transition waters, whereas larger individuals were collected in the Oyashio and the Western Subarctic waters. These results indicate that this species undergoes a horizontal spawning migration from the Oyashio and Western Subarctic waters into the transition waters crossing the Oyashio front.