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1.
Phytoplankton species composition was analyzed inside and outside of the iron-enriched patch during the SEEDS experiment. Before the iron-enrichment, the phytoplankton community consisted of similar proportions of pico-, nano- and micro-sized phytoplankton, and the micro-phytoplankton was dominated by the pennate diatom Pseudo-nitzschia turgidula. Although all the diatoms, except the nano-sized Fragilariopsis sp., increased during the two weeks of the observation period, the flora in the patch dramatically changed with the increase of phytoplankton biomass to a centric diatom-dominated community. Neritic diatoms, especially Chaetoceros debilis, showed higher growth rates than other diatoms, without any delay in the initiation of growth after the enrichment, and accounted for 90% of the micro-phytoplankton after day 9. In contrast, the oceanic diatoms showed distinct delays in the initiation of growth. We conclude that the responses of the diatoms to the manipulation of iron concentration were different by species, and the fast and intensive response of the phytoplankton to iron-enrichment resulted from the presence of a small amount of neritic diatoms at the study site. The important factors that determine the dominant species in the bloom are the potential growth rates under an iron-replete condition and the growth lag. Abundant species in the patch are widely distributed in the North Pacific and their relative contributions in the Oyashio area and at Stn KNOT are high from spring to summer. However, a characteristic difference of species composition between the SEEDS bloom and natural blooms was the lack of Thalassiosira and Coscinodiscus species in the patch, which usually account for a major part of the phytoplankton community under blooming conditions in the western North Pacific.  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.
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   

4.
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   

5.
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   

6.
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   

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.
Dynamics of transparent exopolymer particles (TEP) was studied during the first in situ iron-enrichment experiment conducted in the western subarctic Pacific in July–August 2001, with the goal of evaluating the contribution of TEP to vertical flux as a result of increased primary production following iron enrichment in open ocean ecosystems. Subsequent to the enhancement of phytoplankton production, we observed increase in TEP concentration in the surface layer and sedimentation of organic matter beneath it. Vertical profiles of TEP, chlorophyll a (Chl a) and particulate organic carbon (POC) were obtained from six depths between 5 and 70 m, from a station each located inside and outside the enriched patch. TEP and total mass flux were estimated from the floating sediment traps deployed at 200 m depth. Chl a and TEP concentrations outside the patch varied from 0.2 to 1.9 μg L−1 and 40–60 μg XG equiv. L−1, respectively. Inside the patch, Chl a increased drastically from day 7 reaching the peak of 19.2 μg L−1 on day 13, which coincided with the TEP peak of 189 μg XG equiv. L−1. TEP flux in the sediment trap increased from 41 to 88 mg XG equiv. m−2 d−1, with 8–14% contribution of TEP to total mass flux. This forms the basic data set on ambient concentrations of TEP in the western subarctic Pacific, and evaluation of the effect of iron enrichment on TEP.  相似文献   

10.
To verify the hypothesis that the growth of phytoplankton in the Western Subarctic Gyre (WSG), which is located in the northwest subarctic Pacific, is suppressed by low iron (Fe) availability, an in situ Fe fertilization experiment was carried out in the summer of 2001. Changes over time in the abundance and community structure of phytoplankton were examined inside and outside an Fe patch using phytoplankton pigment markers analyzed by high-performance liquid chromatography (HPLC) and flow cytometry (FCM). In addition, the abundance of heterotrophic bacteria was also investigated by FCM. The chlorophyll a concentration was initially ca. 0.9 μg l−1 in the surface mixed layer where diatoms and chlorophyll b-containing green algae (prasinophytes and chlorophytes) were predominant in the chlorophyll biomass. After the iron enrichment, the chlorophyll a concentration increased up to 9.1 μg l−1 in the upper 10 m inside the Fe patch on Day 13. At the same time, the concentration of fucoxanthin (a diatom marker) increased 45-fold in the Fe patch, and diatoms accounted for a maximum 69% of the chlorophyll biomass. This result was consistent with a microscopic observation showing that the diatom Chaetoceros debilis had bloomed inside the Fe patch. However, chlorophyllide a concentrations also increased in the Fe patch with time, and reached a maximum of 2.2 μg l−1 at 5 m depth on Day 13, suggesting that a marked abundance of senescent algal cells existed at the end of the experiment. The concentration of peridinin (a dinoflagellate marker) also reached a maximum 24-fold, and dinoflagellates had contributed significantly (>15%) to the chlorophyll biomass inside the Fe patch by the end of the experiment. Concentrations of 19′-hexanoyloxyfucoxanthin (a prymnesiophyte marker), 19′-butanoyloxyfucoxanthin (a pelagophyte marker), and alloxanthin (a cryptophyte marker) were only incremented a few-fold increment inside the Fe patch. On the contrary, chlorophyll b concentration reduced to almost half of the initial level in the upper 10 m water column inside the Fe patch at the end of the experiment. A decrease with time in the abundance of eukaryotic ultraphytoplankton (<ca. 5 μm in size), in which chlorophyll b-containing green algae were possibly included was also observed by FCM. Overall, our results indicate that Fe supply can dramatically alter the abundance and community structure of phytoplankton in the WSG. On the other hand, cell density of heterotrophic bacteria inside the Fe patch was maximum at only ca. 1.5-fold higher than that outside the Fe patch. This indicates that heterotrophic bacteria abundance was little respondent to the Fe enrichment.  相似文献   

11.
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   

12.
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   

13.
The phytoplankton community in the western subarctic Pacific (WSP) is composed mostly of pico- and nanophytoplankton. Chlorophyll a (Chl a) in the <2 μm size fraction accounted for more than half of the total Chl a in all seasons, with higher contributions of up to 75% of the total Chl a in summer and fall. The exception is the western boundary along the Kamchatka Peninsula and Kuril Islands and the Oyashio region where diatoms make up the majority of total Chl a during the spring bloom. Among the picophytoplankton, picoeukaryotes and Synechococcus are approximately equally abundant, but the former is more important in term of carbon biomass. Despite the lack of a clear seasonal variation in Chl a concentration, primary productivity showed a large seasonal variation, and was lowest in winter and highest in spring. Seasonal succession in the phytoplankton community is also evident with the abundance of diatoms peaking in May, followed by picoeukaryotes and Synechococcus in summer. The growth of phytoplankton (especially >10 μm cell size) in the western subarctic Pacific is often limited by iron bioavailability, and microzooplankton grazing keeps the standing stock of pico- and nano-phytoplankton low. Compared to the other HNLC regions (the eastern equatorial Pacific, the Southern Ocean, and the eastern subarctic Pacific), iron limitation in the Western Subarctic Gyre (WSG) may be less severe probably due to higher iron concentrations. The Oyashio region has similar physical condition, macronutrient supply and phytoplankton species compositions to the WSG, but much higher phytoplankton biomass and primary productivity. The difference between the Oyashio region and the WSG is also believed to be the results of difference in iron bioavailability in both regions. This revised version was published online in July 2006 with corrections to the Cover Date.  相似文献   

14.
The subarctic North Pacific is one of the three major high nitrate low chlorophyll (HNLC) regions of the world. The two gyres, the NE and the NW subarctic Pacific gyres dominate this region; the NE subarctic Pacific gyre is also known as the Alaska Gyre. The NE subarctic Pacific has one of the longest time series of any open ocean station, primarily as a result of the biological sampling that began in 1956 on the weathership stationed at Stn P (50°N, 145°W; also known as Ocean Station Papa (OSP)). Sampling along Line P, a transect from the coast (south end of Vancouver Island) out to Stn P has provided valuable information on how various parameters change along this coastal to open ocean gradient. The NW subarctic Pacific gyre has been less well studied than the NE gyre. This review focuses mainly on the NE gyre because of the large and long term data set available, but makes a brief comparison with the NW gyre. The NE gyre has saturating NO3 concentrations all year (winter = about 16 μM and summer = about 8 μM), constantly very low chlorophyll (chl) (usually <0.5 mg m−3) which is dominated by small cells (<5 μm). Primary productivity is low (about 300–600 mg C m−2 d−1 and varies little (2 times) seasonally. Annual primary productivity is 3 to 4 times higher than earlier estimates ranging from 140 to 215 g C m−2 y−1. Iron limits the utilization of nitrate and hence the primary productivity of large cells (especially diatoms) except in the winter when iron and light may be co-limiting. There are observations of episodic increases in chl above 1 mg m−3, suggesting episodic iron inputs, most likely from Asian dust in the spring/early summer, but possibly from horizontal advection from the Alaskan Gyre in summer/early fall. The small cells normally dominate the phytoplankton biomass and productivity, and utilize the ammonium produced by the micrograzers. They do not appear to be Fe-limited, but are controlled by microzooplankton grazers. The NW Subarctic Gyre has higher nutrient concentrations and a shallower summer mixed depth and photic zone than Stn P in the NE gyre. Chl concentrations tend to be higher (0.5 to 1.5 μg L−1) than Stn P, but primary productivity in the summer is similar to Stn P (600 mg C m−2 d−1). There are no seasonal data from this gyre. Iron enrichment experiments in October, resulted in an increase in chl (mainly the centric diatom Thalassiosira sp.) and a draw down of nitrate, suggesting that large phytoplankton are Fe-limited, similar to Stn P.  相似文献   

15.
The first iron (Fe) – fertilization experiment in the western North Pacific was carried out using SF6 to trace the Fe-fertilized water mass. A solution in 10,800 liters of seawater of 350 kg of Fe and 0.48 M of SF6 tracer was released into the mixed layer over a 8 × 10 km area. On the first underway transects through the patch after the Fe release, we observed a significant increase of dissolved Fe (ave. 2.89 nM). The fertilized patch was traced for 14 days by on-board SF6 analysis. A Lagrangian frame of reference was maintained by the use of a drogued GPS buoy released at the center of the patch. The patch moved westward at a rate of 6.8 km d−1. Mixed layer depth increased from 8.5 to 15 m during the experiment. Horizontal diffusivity was determined by the change of SF6 concentration in the patch. The horizontal diffusivity increased during the experiment. We evaluate here the fate of Fe in a Fe-fertilized patch using the dilution rate determined from sulphur hexafluoride (SF6) concentration. Dissolved Fe concentrations subsequently decreased rapidly to 0.15 nM on Day 13. However, the dissolved Fe half-life of 43 h was relatively longer than in previous Fe-enrichment studies, and we observed a larger increase of the centric diatom standing stock and corresponding drawdown of macro-nutrients and carbon dioxide than in the previous studies. The most important reason for the larger response was the phytoplankton species in the western North Pacific. In addition, the smaller diffusivity and shallower mixed layer were effective to sustain the higher dissolved Fe concentration compared to previous experiments. This might be one reason for the larger response of diatoms in SEEDS.  相似文献   

16.
We characterized the community composition of phytoplankton in the western subarctic Pacific from the pre-bloom to the decline phase of the spring bloom with special reference to decreases in the silicic acid concentration in surface waters as an index for diatom bloom development. Furthermore, responses of heterotrophic bacteria and viruses to the spring bloom were also concomitantly investigated. Under pre-bloom conditions when nutrients were abundant but the surface mixed layer depth was relatively deep, chlorophyll (Chl) a concentrations were consistently low and green algae (chlorophytes and prasinophytes), cryptophytes, and diatoms were predominant in the phytoplankton assemblages as estimated by algal pigment signatures. Together with the shallowing of the mixed layer depth and the decrease in silicic acid concentration, diatoms bloomed remarkably in the Oyashio region, though the magnitude of the bloom in the Kuroshio-Oyashio transition (hereafter Transition) region was relatively small. A total of 77 diatom species were identified, with the bloom-forming diatoms mainly consisting of Thalassiosira, Chaetoceros, and Fragilariopsis species. It has become evident that the carotenoid fucoxanthin can serve as a strong indicator of the diatom carbon biomass during the spring diatom bloom. Differences in the species richness of diatoms among stations generally enabled us to separate the Oyashio bloom stations from the Transition and the Oyashio pre-bloom stations. Relatively high values of the Shannon-Wiener index for the diatom species were also maintained during the Oyashio bloom, indicating that a wide variety of species then shared dominance. In the decline phase of the Oyashio bloom when surface nutrient concentrations decreased, senescent diatom cells increased, as inferred from the levels of chlorophyllide a. Although the cell density of heterotrophic bacteria changed little with the development of the diatom bloom, viral abundance increased toward the end of the bloom, suggesting an increased likelihood of mortality among organisms including diatoms resulting from viral infection. This is the first report on the microbial trophodynamics, including viruses, during the spring diatom bloom in the western subarctic Pacific.  相似文献   

17.
Sulfur hexafluoride (SF6) tracer release experiments were carried out to trace the iron-fertilized water mass during the iron-fertilization experiments in the western North Pacific of Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study II (SEEDS II) in 2004. A solution of Fe and SF6 tracer was released into the surface mixed layer over an 8×8 km area, and the fertilized patch was traced by onboard SF6 analysis for 12 days during each experiment. A Lagrangian frame of reference was maintained by the use of a drogued GPS buoy released at the center of the patch to reduce the advection effect on observations. The patch moved along the contour of sea-surface height (SSH) of a clockwise mesoscale eddy for 4 days after release. Then strong easterly winds dragged the patch across the contour of SSH. The patch behavior was affected by both the mesoscale eddy and surface winds. Apparent horizontal diffusivities were determined by the change of the distribution of SF6 concentrations. The averaged apparent horizontal diffusivity was about 49 m2 s−1 during SEEDS II. It was larger than the one in SEEDS. Mixed-layer depth (MLD) was 8.5–18 m during SEEDS, and 12–33 m during SEEDS II. The larger horizontal diffusivity and deeper MLD in SEEDS II were disadvantages to maintain a high iron concentration in the surface layer compared to SEEDS. Temporal change of the MLD corresponded to the temporal change of chlorophyll-a concentration. Temporal change in the surface MLD was also important for the response of phytoplankton by iron fertilization.  相似文献   

18.
To elucidate iron regeneration and organic iron(III)-binding ligand formation during microzooplankton and copepod grazing on phytoplankton, incubation experiments were conducted in the western subarctic Pacific. During 8 days of dark incubation of ambient water and that amended with plankton concentrate, dissolved iron and organic iron(III)-binding ligands accumulated, approximately proportionally to the decrease in chlorophyll a. The observed increases in dissolved iron concentration were much greater than those expected from the consumption of phytoplankton biomass and previously reported Fe:C value of cultured algal cells, suggesting resolution from colloidal or particulate iron adsorbed onto the algal cell surface. When copepods were added to the ambient water, organic iron(III)-binding ligands accumulated more rapidly than in the control receiving no copepod addition, although consumed phytoplankton biomass was comparable between the two treatments. Bioassay experiment using filtrates collected from the incubation experiment showed that organic ligands formed during microzooplankton grazing reduced the iron bioavailability to phytoplankton and suppressed their growth. Moreover, picoplankton Synechococcus sp. and Micromonas pusilla were more suppressed by the organic ligands than the diatom Thalassiosira weissflogii. In conclusion, through microzooplankton and copepod grazing on phytoplankton, organic iron(III)-binding ligands as well as regenerated iron are released into the ambient seawater. Because the ligands lower iron bioavailability to phytoplankton through complexation and the degree of availability reduction varies among phytoplankton species, grazing by zooplankton can shift phytoplankton community structure in iron-limited waters.  相似文献   

19.
Bacterial biomass and production rate were measured in the surface (0–100 m) and mesopelagic layers (100–1,000 m) in the subarctic Pacific and the Bering Sea between July–September, 1997. Depth profiles were determined at stations occupied in oceanic domains including the subarctic gyres (western, Bering Sea, and Gulf of Alaska) and a boundary region south of the gyres. In the surface layer (0–100 m), both bacterial biomass and production were generally high in the western and Bering Sea gyres, with the tendency of decrease toward east. This geographic pattern was consistent with the dominant regime of phytoplankton biomass at the time of our survey. A significant portion of variation in bacterial production was explained by the concentration of chlorophyll a (r 2 = 0.340, n = 60, P < 0.001) and, to the greater extent, by the concentration of semilabile total organic carbon (SL-TOC = TOC at a given depth—TOC at 1,000 m, r 2 = 0.488, n = 59, P < 0.0001). Temperature significantly improved the regression model: temperature and chlorophyll jointly explained 60% of variation in bacterial production. These results support the hypothesis that bacteiral growth is largely regulated by the combination of temperature and the supply of dissolved organic carbon in subarctic surface waters. In the mesopelagic layer (100–1,000 m), the geographic pattern of bacterial production was strikingly different from the surface phytoplankton distribution: the production was high in the boundary region where the phytoplankton biomass was lowest. Bacterial growth appeared to be largely controlled by the supply of organic carbon, as indicated by the strong dependency of bacterial production on SL-TOC (r 2 = 0.753, n = 75, P < 0.0001). The spatial uncoupling between surface phytoplankton and mesopelagic bacterial production suggests that the supply rate of labile dissolved organic carbon in the mesopelagic zone does not simply reflect the magnitude of the particulate organic carbon flux in the subarctic Pacific.  相似文献   

20.
《Marine Chemistry》2005,93(1):33-52
Storage carbohydrates (e.g., water-extractable β-1,3-d-glucan in diatoms) are of key importance for phytoplankton growth in a variable light climate, because they facilitate continued growth of the cells in darkness by providing energy and carbon skeletons for protein synthesis. Here, we tested the hypothesis that synthesis of storage carbohydrates by phytoplankton in the Southern Ocean is reduced by low iron and light availability. During the EisenEx/CARbon dioxide Uptake by the Southern Ocean (CARUSO) in situ iron enrichment experiment in the Atlantic sector of the Southern Ocean in November 2000, we studied the dynamics of water-extractable carbohydrates in the particulate fraction over the period of 3 weeks following the iron release. The areal amount (integral between 0- and 100-m depth) of carbohydrates increased from 1400 to 2300 mg m−2 inside the iron-enriched patch, while remaining roughly constant in the surrounding waters. Most of the increase inside the patch was associated with the fraction of large (>10 μm) phytoplankton cells, consistent with the shift in the community structure towards larger diatoms. Deck incubations at 60% of the ambient irradiance revealed that the diurnal chlorophyll a (Chl a)-specific production rates of water-extractable polysaccharides were significantly higher for “in-patch” than for “out-patch” samples (0.5 vs. 0.3 μg C [μg Chl a]−1 h−1, respectively). Together with the higher photochemical efficiency of photosystem II (Fv/Fm), this indicates enhanced photosynthetic performance in response to iron fertilization. In addition, the nocturnal polysaccharide consumption rates were also enhanced by iron release, causing a striking increase in the diel dynamics of polysaccharide concentration. An iron-stimulated increase in diel dynamics was also observed in the fluorescence and size of pico- and nanophytoplankton cells (measured by flow cytometry) and is indicative of enhanced phytoplankton growth. Diurnal polysaccharide production by phytoplankton inside the patch was light-limited when they were incubated at intensities below ca. 200 μmol m−2 s−1 (daytime average). These irradiance levels correspond to those at 20- to 30-m depth in situ, whereas the upper mixed layer was frequently several-fold deeper due to storms. Therefore, these first measurements of phytoplankton carbohydrates during an in situ iron release experiment have revealed that both light and iron availability are the key factors controlling the synthesis of storage carbohydrates in phytoplankton and, hence, the development of diatom blooms in the Southern Ocean.  相似文献   

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