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1.
A mesoscale iron fertilization experiment was carried out in the western subarctic Pacific during summer 2004. The iron-patch was traced for 26 days after the enrichment, and the abundance and behavior of meso- and microzooplankton was compared with those outside of the patch. The surface chlorophyll-a concentration in the patch was high between days 10 and 13 (2.5 mg m−3) and decreased to the initial level after day 20. Microzooplankton grazing rates, estimated by a dilution method, was mostly balanced with phytoplankton growth rates throughout the observed period. Dominant mesozooplankton species in the upper 200 m were copepods: dominated by Eucalanus bungii, Neocalanus plumchrus and Metridia pacifica. Species composition did not change in the patch over the observation period. The copepod biomass was 3–5 times higher than in Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study (SEEDS), the previous iron-enrichment experiment in the same area, before the bloom, and exponentially increased both inside and outside the patch, which was mainly brought by the development of N. plumchrus. The development rates of N. plumchrus were not significantly different between inside and outside the patch. Estimated grazing rate suggest that the copepod grazing was main cause of the low accumulation of phytoplankton biomass, and dominance of grazing-resistant organisms such as large ciliates, large diatoms and diatoms with extremely long setae. “Arrested migration” for M. pacifica and upward shift of vertical distribution by E. bungii were observed during the bloom period, even if the accumulation of phytoplankton biomass was very low compared to other iron-enrichment experiments. These results indicate that the copepod grazing shaped the food-web structure of the lower trophic levels (biomass and species composition) in SEEDS II.  相似文献   

2.
《Journal of Oceanography》2007,63(6):983-994
A mesoscale iron-enrichment study (SEEDS II) was carried out in the western subarctic Pacific in the summer of 2004. The iron patch was traced for 26 days, which included observations of the development and the decline of the bloom by mapping with sulfur hexafluoride. The experiment was conducted at almost the same location and the same season as SEEDS (previous iron-enrichment experiment). However, the results were very different between SEEDS and SEEDS II. A high accumulation of phytoplankton biomass (∼18 mg chl m−3) was characteristic of SEEDS. In contrast, in SEEDS II, the surface chlorophyll-a accumulation was lower, 0.8 to 2.48 mg m−3, with no prominent diatom bloom. Photosynthetic competence in terms of F v/F m for the total phytoplankton community in the surface waters increased after the iron enrichments and returned to the ambient level by day 20. These results suggest that the photosynthetic physiology of the phytoplankton assemblage was improved by the iron enrichments and returned to an iron-stressed condition during the declining phase of the bloom. Pico-phytoplankton (<2 μm) became dominant in the chlorophyll-a size distribution after the bloom. We observed a nitrate drawdown of 3.8 μM in the patch (day 21), but there was no difference in silicic acid concentration between inside and outside the patch. Mesozooplankton (copepod) biomass was three to five times higher during the bloom-development phase in SEEDS II than in SEEDS. The copepod biomass increased exponentially. The grazing rate estimation indicates that the copepod grazing prevented the formation of an extensive diatom bloom, which was observed in SEEDS, and led to the change to a pico-phytoplankton dominated community towards the end of the experiment.  相似文献   

3.
To test the iron hypothesis in the subarctic Pacific Ocean, an in situ iron-enrichment experiment (SEEDS) was performed in the western subarctic gyre in July–August 2001. About 350 kg of iron (as acidic iron sulfate) and 0.48 mol of the inert chemical tracer sulfur hexafluoride were introduced into a 10-m deep surface mixed layer over an 80 km2 area. This single iron infusion raised dissolved iron levels to 2.9 nM initially. Dissolved iron concentrations rapidly decreased after the infusion, but levels remained close to 0.15 nM even at the end of the 14-day experimental period. During SEEDS there were iron-mediated increases in chlorophyll a concentrations (up to 20 μg l−1), primary production rates, biomass and photosynthetic energy conversion efficiency relative to waters outside the iron-enriched patch. The rapid and very high accumulation of phytoplankton biomass in response to the iron addition appeared to be partly attributable to shallow mixed-layer depth and moderate water temperature in the western subarctic Pacific. However, the main reason was a floristic shift to fast-growing centric diatom Chaetoceros debilis, unlike the previous iron-enrichment experiments in the equatorial Pacific and the Southern Ocean, in both of which iron stimulated the growth of pennate diatoms. The iron-mediated blooming of diatoms resulted in a marked consumption of macronutrients and drawdown of pCO2. Biological and physiological measurements indicate that phytoplankton growth in the patch became both light- and iron-limited, making phytoplankton biomass relatively constant after day 9. The increase in microzooplankton grazing rate after day 9 also influenced the net growth rate of phytoplankton. There was no significant increase in the export flux of carbon to depth during the 14-day occupation of the experimental site. The export flux between day 4 and day 13 was estimated to be only 13% of the integrated primary production in the iron-enriched patch. The major part of the carbon fixed by the diatom bloom remained in the surface mixed layer as biogenic particulate matter. Our findings support the hypothesis that iron limits phytoplankton growth and biomass in a ‘bottom up’ manner in this area, but the fate of algal carbon remains unknown.  相似文献   

4.
To fill temporal gaps in iron-enrichment experimental data and gain further understanding of marine ecosystem responses to iron enrichments, we apply a fifteen-compartment ecosystem model to three iron-enrichment sites, namely SEEDS (the Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study; 48.5°N, 165°E) in the western North Pacific, SOIREE (the Southern Ocean Iron RElease Experiment; 61°S, 140°E) in the Southern Ocean, and IronExII (the second mesoscale iron enrichment experiment; 3.5°S, 104°W) in the Equatorial Pacific. The ecological effects of iron in the model are represented by changing two photosynthetic parameters during the iron-enrichment period. The model results successfully reproduce the observed biogeochemical responses inside and outside the iron patch at each site, such as rapid increases in plankton biomass and biological productivity, and decreases in surface nutrients and pCO2, inside the patch. However, the modeled timing and magnitude of changes differ among the sites because of differences in both physical environments and plankton species. After the iron enrichment, the diatom productivity is strongly controlled by light at SOIREE and by silicate at IronExII and SEEDS. Light limitation due to self-shading by the phytoplankton is significant during the bloom at all sites. Sensitivity analysis of the model results to duration of the iron enrichment reveals that long-term multiple infusions over more than a week would not be effective at SEEDS because of strong silicate limitation on diatom growth. Sensitivity of the model to water temperature shows that export production is higher at lower temperatures, because of slower recycling of particulate organic carbon. Therefore, the e-ratio (the ratio of export production to primary production) is inversely correlated with temperature, and the relationship can be described with a linear function. Through this study, we conclude that ecosystem modeling is a powerful tool to help design future iron-enrichment experiments and observational plans.  相似文献   

5.
The cumulative evidence from more than a dozen mesoscale iron-enrichment studies in high nitrate low chlorophyll (HNLC) waters demonstrates that iron limitation is widespread and very likely affects atmospheric carbon dioxide and thus global climate. However, the responses of microphytoplankton (>20 μm), predominantly diatoms, vary greatly among these mesoscale experiments even though similar amounts of iron were added, making it difficult to quantitatively incorporate iron effects into global climate models. Nowhere is this difference more dramatic than between the massive bloom observed during Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study (SEEDS) I and the order of magnitude smaller ecosystem response in SEEDS II; two mesocale experiments performed in the same HNLC region of the western subarctic Pacific in different years. Deckboard incubation experiments initiated during the early, middle, and late stages of the 32-day SEEDS II experiment show that while the two iron infusions increased phytoplankton growth, diatoms remained significantly limited by iron availability, despite total dissolved Fe concentrations in the patch being well above the diffusion-limited threshold for rapid diatom growth. This iron limitation was apparent <6 days after the initial iron infusion and was not alleviated by the second, smaller iron infusion. In contrast, smaller phytoplankton (<20 μm) showed a more restricted response to further iron amendments, indicating that their iron nutrition was near optimal. Iron complexed to desferrioximine B, a commonly available siderophore produced by at least one marine bacterium, was poorly available to diatoms throughout the patch evolution, indicating that these diatoms lacked the ability to induce high-affinity iron uptake systems. These results suggest that the strong organic complexation of Fe(III) observed in the SEEDS II-fertilized patch was not compatible with rapid diatom growth. In contrast, iron associated with protoporphyrin IX, a weaker iron complexing ligand of a class hypothesized to be representative of recycled iron species, was readily available to diatoms. Our findings demonstrate that a persistence of iron limitation was the primary factor underlying the comparatively small diatom response during SEEDS II. This continued growth limitation would have increased the importance of mesozooplankton grazing as a controlling factor in the SEEDS II ecosystem response.  相似文献   

6.
Several in situ iron-enrichment experiments have been conducted, where the response of the phytoplankton community differed. We use a marine ecosystem model to investigate the effect of iron on phytoplankton in response to different initial plankton conditions and mixed-layer depths (MLDs). Sensitivity analysis of the model results to the MLDs reveals that the modeled response to the same iron enhancement treatment differed dramatically according to the different MLDs. The magnitude of the iron-induced biogeochemical responses in the surface water, such as maximum chlorophyll, is inversely correlated with MLD, as observed. The significant decrease in maximum surface chlorophyll with MLD results from the difference in diatom concentration in the mixed layer, which is determined by vertical mixing. The modeled column-integrated chlorophyll, on the other hand, is the highest with intermediate MLD cases, suggesting difference in iron-induced biogeochemical responses between volume and area considerations. The iron-induced diatom bloom is severely restricted below the compensation depth due to both light limitation and grazing pressure, irrespective of the MLD. Sensitivity of the model to initial mesozooplankton (as grazers on diatoms) biomass shows that column-integrated biomass, net community production and export production are strongly controlled by the initial mesozooplankton biomass. Higher initial mesozooplankton biomass yields high grazing pressure on diatoms, which results in less accumulation of diatom biomass and may account for notably lower surface chlorophyll during SEEDS (Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study) II than during SEEDS. The initial diatom biomass is also important to the outcome of iron enrichment but is not as crucial as the MLD and the initial mesozooplankton biomass. This modeling study suggests that not only MLD but also the initial biomass of diatoms and its principle grazers are crucial factors in the response of the phytoplankton community to iron enrichments, and should be considered in designing future iron-enrichment experiments.  相似文献   

7.
During two mesoscale iron-enrichment studies in the northwestern subarctic Pacific (SEEDS in 2001 summer and SEEDS II in 2004 summer), particulate materials from the iron-induced phytoplankton bloom in the upper water column were monitored to analyze the export processes beneath the upper mixed layer, mainly with drifting sediment traps. We could not observe the total downward export process of the high accumulation of particulate organic carbon from the mixed layer induced by the large diatom bloom of SEEDS [e.g., Tsuda, A., Takeda, S., Saito, H., Nishioka, J., Nojiri, Y., Kudo, I., Kiyosawa, H., Shiomoto, A., Imai, K., Ono, T., Shimamoto, A., Tsumune, D., Yoshimura, T., Aono, T., Hinuma, A., Kinugasa, M., Suzuki, K., Sohrin, Y., Noiri, Y., Tani, H., Deguchi, Y., Tsurushima, N., Ogawa, H., Fukami, K., Kuma, K., Saino, T., 2003. A mesoscale iron enrichment in the western subarctic Pacific induces large centric diatom bloom. Science 300, 958–961] because the 2-week observation period was too short to examine the decline phase of the bloom. In contrast, in SEEDS II, the particulate organic carbon and particulate organic nitrogen were accumulated 123 and 23 mmol m−2, respectively, in the mixed layer until day-15 (days from iron-enrichment), and then ca. 90% were removed from the mixed layer by day-25. The sediment traps at 40 m depth between day-15 and day-25 accounted for at least more than 35% of these particles. There was no large variation in chemical composition in settling particles above 100 m depth throughout the experimental periods both in SEEDS and SEEDS II. The content of biogenic opal remained more than 50% of all settling particles during SEEDS, while the content of biogenic calcium carbonate was relatively high, with a low biogenic opal content of consistently less than 30% during SEEDS II. These results suggest that high standing stock of seed population of diatoms before the iron fertilization, indicated by low C/Si ratio of particulate matter, is an important factor to induce the large diatom bloom in SEEDS.  相似文献   

8.
Little is known about the effects of iron enrichment in high-nitrate low-chlorophyll (HNLC) waters on the community composition of heterotrophic bacteria, which are crucial to nutrient recycling and microbial food webs. Using denaturing gradient gel electrophoresis (DGGE) of 16S rDNA fragments, we investigated the heterotrophic eubacterial community composition in surface waters during an in situ iron-enrichment experiment (SEEDS-II) in the western subarctic Pacific in the summer of 2004. DGGE fingerprints representing the community composition of eubacteria differed inside and outside the iron-enriched patch. Sequencing of DGGE bands revealed that at least five phylotypes of α-proteobacteria including Roseobacter, Cytophaga-Flavobacteria-Bacteroides (CFB), γ-proteobacteria, and Actinobacteria occurred in almost all samples from the iron-enriched patch. Diatoms did not bloom during SEEDS-II, but the eubacterial composition in the iron-enriched patch was similar to that in diatom blooms observed previously. Although dissolved organic carbon (DOC) accumulation was not detected in surface waters during SEEDS-II, growth of the Roseobacter clade might have been particularly stimulated after iron additions. Two identified phylotypes of CFB were closely related to the genus Saprospira, whose algicidal activity might degrade the phytoplankton assemblages increased by iron enrichment. These results suggest that the responses of heterotrophic bacteria to iron enrichment could differ among phylotypes during SEEDS-II.  相似文献   

9.
In order to detect iron (Fe) stress in micro-sized (20–200 μm) diatoms in the Oyashio region, western subarctic Pacific during spring, immunological ferredoxin/flavodoxin assays were applied to samples collected from the surface layer in May 2005. Concomitantly, the community composition of the micro-sized phytoplankton and hydrographic conditions, including dissolved Fe and macronutrient concentrations, were also examined. Chlorophyll (Chl) a concentrations were <2 mg m−3 at all sampling stations, except at a station where the Chl a level was 9.0 mg m−3 and a micro-sized diatom bloom occurred. A high abundance of ferredoxin in micro-sized diatoms was detected only at a rather near-shore station where dissolved Fe and macronutrient concentrations were higher, indicating that the micro-sized diatoms did not suffer from iron deficiency. On the other hand, flavodoxin in micro-sized diatoms was often observed at the other stations, including the bloom station, where macronutrients were replete but dissolved Fe concentration was low (0.31 nM). A significant amount of chlorophyllide a, a degradation product of Chl a, was also observed at the bloom station, suggesting a decline of the diatom bloom. The micro-sized phytoplankton species at all the stations were mainly composed of the diatoms Thalassiosira, Chaetoceros, and Fragilariopsis spp. Our study indicates that micro-sized diatoms were stressed by Fe bioavailability during the spring season in the Oyashio region  相似文献   

10.
A mesoscale iron-fertilization experiment was carried out in the western subarctic Pacific during summer 2001. The iron-patch was traced for 14 days after the fertilization, and the abundance and behavior of mesozooplankton were compared with those outside of the patch. The phytoplankton biomass in the patch rapidly increased to over 15 times the initial level by the later half of the observation period, and was composed of large-sized (>10 mm), centric diatoms. Dominant zooplankton species in the upper 200-m depth were large copepods: Neocalanus plumchrus, Neocalanus cristatus, Eucalanus bungii and Metridia pacifica. Mesozoplankton biomass as well as species composition did not change significantly in the patch over the observation period. Furthermore, no changes of vertical distribution or diel vertical migration were observed for any species or stages of mesozooplankton throughout the observation period. However, the abundance of the first copepodite stages of N. plumchrus and E. bungii increased several fold in the patch after the diatom bloom formation compared to the densities outside the patch. The increases of both species are considered to be due to lowered mortality during the egg and nauplius stages. Spawning of N. plumchrus takes place at depth using lipid storage, while spawning of E. bungii takes place in the surface layer supported by grazing. These facts suggest that the relative importance of nauplii in the diets of the large copepods was decreased in the patch by the diatom bloom. Gut-pigment contents of dominant copepods in the patch increased 4–18 times, and the maximum values were observed during the bloom peak. However, the grazing impact on phytoplankton was low throughout the experiment, especially during the bloom period (<6% of the primary production).  相似文献   

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