The phytoplankton bloom response to wind events and upwelled nutrients during the CoOP WEST study

In the coastal waters off northern California, seasonal wind-driven upwelling supplies abundant nutrients to be processed by phytoplankton productivity. As part of the Coastal Ocean Processes: Wind Events and Shelf Transport (CoOP WEST) study, nutrients, CO₂, size-fractionated chlorophyll, and phytoplankton community structure were measured in the upwelling region off Bodega Bay, CA, during May–June 2000, 2001 and 2002. The ability of this ecosystem to assimilate nitrate (NO₃) and silicic acid/silicate (Si(OH)₄) and accumulate particulate material (i.e. phytoplankton) was realized in all 3 years, following short events of upwelling-favorable winds, followed by periods of relaxed winds. This was observed as phytoplankton blooms, dominated by chlorophyll in cells greater than 5 μm in diameter, that reduced the ambient nutrients to zero. These communities were located over the near-shore shelf (<100 m depth) and were dominated by diatoms. An optimal window of 3–7 days of relaxed winds, following an upwelling pulse, was required for chlorophyll accumulation. The large-celled phytoplankton that result are likely important players in coastal new production and carbon cycling.     

Manganese and iron distributions off central California influenced by upwelling and shelf width

In July 2002, a combination of underway mapping and discrete profiles revealed significant along-shore variability in the concentrations of manganese and iron in the vicinity of Monterey Bay, California. Both metals had lower concentrations in surface waters south of Monterey Bay, where the shelf is about 2.5 km wide, than north of Monterey Bay, where the shelf is about 10 km wide. During non-upwelling conditions over the northern broad shelf, dissolvable iron concentrations measured underway in surface waters reached 3.5 nmol L⁻¹ and dissolved manganese reached 25 nmol L⁻¹. In contrast, during non-upwelling conditions over the southern narrow shelf, dissolvable iron concentrations in surface waters were less than 1 nmol L⁻¹ and dissolved manganese concentrations were less than 5 nmol L⁻¹. A pair of vertical profiles at 1000 m water depth collected during an upwelling event showed dissolved manganese concentrations of 10 decreasing to 2 nmol L⁻¹, and dissolvable iron concentrations of 12–20 nmol L⁻¹ in the upper 100 m in the north, compared to 3.5–2 nmol L⁻¹ Mn and 0.6 nmol L⁻¹ Fe in the upper 100 m in the south, suggesting the effect of shelf width influences the chemistry of waters beyond the shelf.These observations are consistent with current understanding of the mechanism of iron supply to coastal upwelling systems: Iron from shelf sediments, predominantly associated with particles greater than 20 μm, is brought to the surface during upwelling conditions. We hypothesize that manganese oxides are brought to the surface with upwelling and are then reduced to dissolved manganese, perhaps by photoreduction, following a lag after upwelling.Greater phytoplankton biomass, primary productivity, and nutrient drawdown were observed over the broad shelf, consistent with the greater supply of iron. Incubation experiments conducted 20 km offshore in both regions, during a period of wind relaxation, confirm the potential of these sites to become limited by iron. There was no additional growth response when copper, manganese or cobalt was added in addition to iron. The growth response of surface water incubated with bottom sediment (4 nmol L⁻¹ dissolvable Fe) was slightly greater than in control incubations, but less than in the presence of 4 nmol L⁻¹ dissolved iron. This may indicate that dissolvable iron is not as bioavailable as dissolved iron, although the influence of additional inhibitory elements in the sediment cannot be ruled out.     

Alkaline phosphatase activity of water column fractions and seagrass in a tropical carbonate estuary, Florida Bay

Few phosphorus-depleted coastal ecosystems have been examined for their ability to hydrolyze phosphomonoesters. We examined seasonal (August 2006–April 2007) alkaline phosphatase activity in Florida Bay, a phosphorus-limited shallow estuary, using fluorescent substrate at low concentrations (≤2.0 μM). In situ dissolved inorganic and organic phosphorus levels and phosphomonoester concentrations were also determined. Water column alkaline phosphatase activity was partitioned into two particulate size fractions (>1.2 and 0.2–1.2 μm) and freely dissolved enzymes (<0.2 μm). Water column alkaline phosphatase activity was also compared to leaf and epiphyte activity of the dominant tropical seagrass Thalassia testudinum. Our results indicate: (1) potential alkaline phosphatase activity in Florida Bay is high compared to other marine ecosystems, resulting in rapid phosphomonoester turnover times (2 h). (2) Water column alkaline phosphatase activity dominates, and is split equally between particulate and dissolved fractions. (3) Alkaline phosphatase activity was highest during cyanobacterial blooms, but not when normalized to chl a. These results suggest that dissolved, heterotrophic and autotrophic alkaline phosphatase activity is stimulated by phytoplankton blooms. (4) The dissolved alkaline phosphatase activity is relatively constant, while the particulate activity is seasonally and spatially dynamic, typically associated with phytoplankton blooms. (5) Phosphomonoester concentrations throughout the bay are low, even though potential hydrolysis rates are high. We propose that bioavailable dissolved organic P is hydrolyzed by dissolved and microbial alkaline phosphatase enzymes in Florida Bay. High alkaline phosphatase activity in the bay is also promoted by long hydraulic residence times. This background activity is primarily driven by carbon and phosphorus limitation of microorganisms, and regeneration of enzymes associated with cell lysis. Pulses of inorganic phosphorus and labile organic phosphorus and nitrogen may stimulate autotrophs, particularly cyanobacteria, which in turn promote biological activity that increase alkaline phosphatase activity of both autotrophs and heterotrophs in the bay.     

WEST: A northern California study of the role of wind-driven transport in the productivity of coastal plankton communities

The “Wind Events and Shelf Transport” (WEST) program was an interdisciplinary study of coastal upwelling off northern California in 2000–03. WEST was comprised of modeling and field observations. The primary goal of WEST was to better describe and understand the competing influences of wind forcing on planktonic productivity in coastal waters. While increased upwelling-favorable winds lead to increased nutrient supply, they also result in reduced light exposure due to deeper surface mixed layers and increased advective loss of plankton from coastal waters. The key to understanding high levels of productivity, amidst these competing responses to wind forcing, is the temporal and spatial structure of upwelling. Temporal fluctuations and spatial patterns allow strong upwelling that favors nutrient delivery to be juxtaposed with less energetic conditions that favor stratification and plankton blooms. Observations of winds, ocean circulation, nutrients, phytoplankton and zooplankton off Bodega Bay and Point Reyes (38°N) were combined with model studies of winds, circulation and productivity. This overview of the WEST program provides an introduction to the WEST special issue of Deep-Sea Research, including the motivation for WEST, a summary of study components, an integrative synthesis of major research results to-date, and background on conditions during field studies in May–June 2001 (the upwelling period on which this special issue is focused).     

Winter-spring phytoplankton blooms in Dabob Bay, Washington

Scientific investigations in Dabob Bay, Washington State, USA, have been extensive since the early 1960s, but phytoplankton blooms have been studied mostly with regard to chlorophyll concentrations and little is known about the phytoplankton species themselves. Here we provide information on the species present, their abundances during blooms, their contribution to organic carbon concentrations and the ability of some phytoplankton species to produce toxic aldehydes that may impact metazoan grazers.Multiple blooms of phytoplankton, dominated by diatoms, occurred in the late winter-early spring period, with depth-integrated chlorophyll levels ranging from <20 to 230 mg m⁻² and peaks in February and April. The major bloom species included Skeletonema costatum, Thalassiosira spp. and Chaetoceros spp; Phaeocystis cf. pouchetii occurred in 2002 and 2004. Other taxa or groups of organisms that were sometimes abundant included unidentified small flagellates <10 μm in size and unidentified heterotrophic dinoflagellates. Large diatoms usually comprised most of the cell carbon, but a large, heterotrophic dinoflagellate, identified only as Gyrodinium “tear” because of its shape, was a major contributor to the microplankton carbon when present even in small numbers. Five Thalassiosira species and S. costatum were found to produce polyunsaturated aldehydes (PUA) that are known to affect copepod reproduction and hatching success. Our findings are similar to the few previous studies in the last four decades that included phytoplankton species and suggest long-term similarities and relative stability in the phytoplankton species present and their timing in Dabob Bay.     

夏季南极长城湾海流变化特征及其对浮游植物生物量的影响

通过对2010/2011年夏季南极长城湾叶绿素a浓度和海流等环境参数的传感器连续观测;研究其时空变化特征及海流对浮游植物生物量的影响。结果表明;12.9 m水层叶绿素a浓度从12月中旬开始增长;直至2月上旬开始维持相对稳定;期间出现两个高值点（2.74 μg/L和3.76 μg/L）。叶绿素a浓度表现出了24 h周期变化特征;每日叶绿素a浓度最高值出现在正午前后时段的概率要高于其他时段。表层海水受西北风影响;流速较大;约为60~100 cm/s;中层和底层水体流速较小;约为10 cm/s。表层的风海流使海水在长城湾发生辐散;下层海水向上涌升;驱动了水体的垂直交换。长城湾与麦克斯韦尔湾存在水体交换;水体主要从中层进入湾内;从表层输出。长城湾的叶绿素a浓度与流速呈显著负相关;表明低流速环境有利于长城湾浮游植物的生长。     

Seasonal variability of living benthic foraminifera from the outer continental shelf of the Bay of Biscay

Living benthic foraminiferal faunas of six stations from the continental shelf of the Bay of Biscay have been investigated during three successive seasons (spring, summer and autumn 2002). For the three investigated stations, bottom water oxygen concentration, oxygen penetration into the sediment and sediment organic carbon contents are all relatively similar. Therefore, we think that the density and the composition of the foraminiferal faunas is mainly controlled by the quantity and quality of organic input resulting from a succession of phytoplankton bloom events, occurring from late February to early September. The earliest blooms are positioned at the shelf break, late spring and early summer blooms occur off Brittany, whereas in late summer and early autumn, only coastal blooms appear, often in the vicinity of river outlets. In spring, the benthic foraminiferal faunas of central (B, C and D) and outer (E) continental shelf stations are characterised by strong dominance in the first area and strong presence in the second area of Nonionella iridea. In fact, station E does not serve as a major depocenter for the remains of phytoplankton blooms. If station E is not considered, the densities of this taxon show a clear gradient from the shelf-break, where the species dominates the assemblages, to the coast, where it attains very low densities. We explain this gradient as a response to the presence, in early spring, of an important phytoplankton bloom, mainly composed of coccolithophorids, over the shelf break. This observation is supported by the maximum particles flux values at stations close to the shelf break (18.5 g m^− 2 h^− 1) and lower values in a station closer to the coast (6.8 g m^− 2 h^− 1). In summer, the faunal density is maximum at station A, relatively close to more varied phytoplancton blooms that occur off Brittany until early June. We suggest that the dominant species, Nonion fabum, Cassidulina carinata and Bolivina ex. gr. dilatata respond to phytodetritus input from these blooms. In autumn, the rich faunas of inner shelf station G are dominated by N. fabum, B. ex. gr. dilatata, Hyalinea balthica and Nonionella turgida. These taxa seem to be correlated with the presence of coastal blooms phenomena, in front of river outlets. They may be favoured by an organic input with a significant contribution of terrestrial, rather low quality organic matter.     

Seasonal variations in dissolved organic carbon concentrations and characteristics in a shallow coastal bay

Dissolved organic matter (DOM) composition and dynamics in temperate shallow coastal bays are not well described although these bays may be important as local sources of organic carbon to ocean waters and are often sites of economically-important fisheries and aquaculture. In this study surface water samples were collected on a monthly to bi-monthly basis over two years from a mid-Atlantic coastal bay (Chincoteague Bay, Virginia and Maryland, USA). Dissolved organic carbon (DOC) concentrations and light absorbance characteristics were measured on sterile-filtered water, and high-molecular weight (> 1 kDa) dissolved OM (DOM) was isolated to determine stable isotope composition and molecular-level characteristics. Our time series encompassed both a drought year (2002) and a year of above-average rainfall (2003). During the dry year, one of our sites developed a very intense bloom of the brown tide organism Aureococcus anophagefferens while during the wet year there were brown tide bloom events at both of our sampling sites. During early spring of the wet year, there were higher concentrations of > 1 kDa DOC; this fraction represented a larger proportion of overall DOC and appeared considerably more allochthonous. Based upon colored dissolved organic matter (CDOM) and high-molecular weight DOM analyses, the development of extensive phytoplankton blooms during our sampling period significantly altered the quality of the DOM. Throughout both years Chincoteague Bay had high DOC concentrations relative to values reported for the coastal ocean. This observation, in conjunction with the observed effects of phytoplankton blooms on DOM composition, indicates that Chincoteague Bay may be a significant local source of “recently-fixed” organic carbon to shelf waters. Estimating inputs of DOC from Chincoteague Bay to the Mid-Atlantic Bight suggests that shallow productive bays should be considered in studies of organic carbon on continental shelves.     

Estimating direct and episodic atmospheric nitrogen deposition to a coastal waterbody

Direct deposition of atmospheric nitrogen to shallow coastal embayments is usually estimated, since insufficient field measurements are available. Using Waquoit Bay (Cape Cod, MA. USA) as a case study, and a recent review of literature, we determined reasonable bounds on wet and dry inputs of inorganic and organic N. Since precipitation and wind vary daily, we explored the potential of episodic events to stimulate phytoplankton blooms. Many coastal waterbodies like Waquoit Bay are small relative to their watersheds. Nevertheless, direct deposition of NH(3), NO(3)(-), and HNO(3) is significant in the loading budget. For Waquoit Bay, direct deposition was calculated to be 7-15.5 kg total N ha(-1)yr(-1), representing 70-150% of the atmospheric N reported to reach the bay via the watershed, and 20-45% of the total N reaching the bay from all land-based sources. Episodic events were estimated to deliver up to 65 mg N m(-2)day(-1), representing a phytoplankton stock of 12.3 mg Chl m(-2), an amount unlikely to stimulate dense blooms in shallow coastal waters in the northeastern United States.     

Thermal Stratification in Saldanha Bay (South Africa) and Subtidal, Density-driven Exchange with the Coastal Waters of the Benguela Upwelling System

A study was conducted to understand the mechanisms driving observed subtidal variability in the stratification of Saldanha Bay, located in the southern Benguela system. It was found that the 6–8 day period variability in bay stratification was caused by the inflow and outflow of cold upwelled water driven by changing baroclinic pressure gradients between the coastal and bay domains. The direction and magnitude of the pressure gradients were governed by coastal upwelling activity and a lag in the response of the bay to changes in density structure in the coastal ocean. When the pressure gradients were bayward and cold water was being driven into the bay the cycle was termed to be in an ‘ active phase ’ and the reverse was termed the ‘ relaxation phase ’. The upwelling-favourable equatorward wind stress impacted the bay stratification in two ways: on the regional scale, wind drives upwelling and governs the inflow–outflow of cold upwelled bottom water, which strengthens stratification; conversely, on the local bay scale, wind drives vertical mixing, which weakens stratification. A four-phase model is used to describe the observed variability in stratification in the bay. The associated density-driven exchange flows are capable of flushing the bay in 6–8 days, about one-third of the time for tidal exchange alone (c. 25 days). These inflows of cold bottom water are ecologically critical as they supply nutrients to the bay and thus impose a control on new production within the bay environment. Further ecological implications of this bay–ocean exchange include export of phytoplankton new production to the coast, limitation of the risk of harmful algal blooms (HABs) and the division of the system into two distinct ecosystems (bay and lagoon).     

Importance of vertical mixing for additional sources of nitrate and iron to surface waters of the Columbia River plume: Implications for biology

The influence of the Columbia River plume on the distributions of nitrate and iron and their sources to coastal and shelf waters were examined. In contrast to other large estuaries, the Columbia River is a unique study area as it supplies very little nitrate (5 μM) and iron (14–30 nM) at salinities of 1–2 to coastal waters. Elevated nitrate and dissolved iron concentrations (as high as 20 μM and 20 nM) were observed, however, in the near field Columbia River plume at salinities of 20. Surface nitrate concentrations were higher than observed in the Columbia River itself and therefore must be added by entrainment of higher nitrate concentrations from subsurface coastal waters. Tidal flow was identified as an important factor in determining the chemical constituents of the Columbia River plume. During the rising flood tide, nitrate and iron were entrained into the plume waters resulting in concentrations of 15 μM and 6 nM, respectively. Conversely, during the ebb tide the concentrations of nitrate and total dissolved iron were reduced to 0.3–3 μM and 1–2 nM, respectively, with a concomitant increase in chlorophyll a concentrations. As these plume waters moved offshore the plume drifted directly westward, over a nitrate depleted water mass (< 0.2 μM). The plume water was also identified to move southwards and offshore during upwelling conditions and nitrate concentrations in this far field plume were also depleted. Iron concentrations in the near-field Columbia River plume are sufficient to meet the biological demand. However, due to the low nitrate in the Columbia River itself, nitrate in the plume is primarily dependent on mixing with nitrate rich, cold, high salinity subsurface waters. Without such an additional source the plume rapidly becomes nitrate limited.     

Transformation dynamics and reactivity of dissolved and colloidal iron in coastal waters

We have investigated the chemical forms, reactivities and transformation kinetics of Fe(III) species present in coastal water with ion exchange and filtration methods. To simulate coastal water system, a mixture of ferric iron and fulvic acid was added to filtered seawater and incubated for a minute to a week. At each incubation time, the seawater sample was acidified with hydrochloric acid and then applied to anion exchange resin (AER) to separate negatively charged species (such as fulvic acid, its complexes with iron and iron oxyhydroxide coated with fulvic acid) from positively charged inorganic ferric iron (Fe(III)′). By monitoring the acid-induced Fe(III)′ over an hour, it was found that iron complexed by fulvic acid dissociated rapidly to a large extent (86–92% at pH 2), whereas amorphous ferric oxyhydroxide particles associated with fulvic acid (AFO-L) dissociated very slowly with the first-order dissociation rate constants ranging from 6.1 × 10^− 5 for pH 3 to 2.7 × 10^− 4 s^− 1 for pH 2. Therefore, a brief acidification followed by the AER treatment (acidification/AER method) was likely to be able to determine fulvic acid complexes and thus differentiate the complexes from the AFO-L particles (the dissolution of AFO-L was insignificant during the brief acidification). The acidification/AER method coupled with a simple filtration technique suggested that the iron–fulvic acid complexes exist in both the < 0.02 μm and 0.02–0.45 μm size fractions in our coastal water system. The truly dissolved iron (< 0.02 μm) was relatively long-lived with a life-time of 14 days, probably due to the complexation by strong ligands. Such an acid-labile iron may be an important source of bioavailable iron in coastal environments, as a significant relationship between the chemical lability and bioavailability of iron has been well recognised.     

胶州湾海域浮游植物和赤潮物种的生物多样性研究进展

胶州湾是我国的一个典型近海海湾,也是近海生态系统的缩影。历史上胶州湾海域赤潮频发,因此也是研究赤潮暴发机制(包括赤潮物种组成、时空动态变化及对环境因子响应)的理想实验场所。迄今,几代科学家对胶州湾浮游植物和赤潮物种开展了长期而系统的调查研究,获得了大量的研究成果。为了系统跟踪胶州湾海域浮游植物群落组成,尤其是赤潮物种的组成变化,本文对近84年间(1936—2019年)胶州湾海域涉及浮游植物调研的54个调查数据结果进行了系统的统计和比较分析。文献共报道了549种浮游植物,包括硅藻326种、甲藻164种、绿藻21种、金藻9种、隐藻9种、淡色藻6种、定鞭藻5种、蓝藻3种、黄藻3种、下睫虫2种、裸藻1种和针胞藻1种。其中只有不到一半(40.29%)的浮游植物的全长18S rDNA序列得到解析,表明分子标记数据库需要继续完善,促进宏条形码分析的有效应用。胶州湾海域的优势浮游植物随时间的推移发生了较大变化,但是报道的甲藻与硅藻物种数的比例没有显著的变化。统计表明,研究者在胶州湾共鉴定到153种赤潮物种,包括硅藻79种、甲藻67种、定鞭藻2种、金藻2种、淡色藻1种、黄藻1种和针胞藻1种。胶州湾海域的典型优势浮游植物均为赤潮物种,包括中肋骨条藻、短角弯角藻、旋链角毛藻、加氏星杆藻、日本星杆藻、星脐圆筛藻和尖刺拟菱形藻,其中短角弯角藻和中肋骨条藻是胶州湾海域的"常驻"赤潮物种。本文通过系统综述胶州湾浮游植物和赤潮物种的生物多样性,为利用宏条形码和宏基因组等方法跟踪研究胶州湾和其他海域的浮游植物和赤潮物种的演化奠定了基础。     

Sea level fluctuations in central California at subtidal to decadal and longer time scales with implications for San Francisco Bay, California

Sea level elevations from near the mouth of San Francisco Bay are used to describe the low-frequency variability of forcing of the coastal ocean on the Bay at a variety of temporal scales. About 90% of subtidal fluctuations in sea level in San Francisco Bay are driven by the sea level variations in the coastal ocean that propagate into the Bay at the estuary mouth. We use the 100-year sea level record available at San Francisco to document a 1.9 mm/yr mean sea level rise, and to determine fluctuations related to El Nino-Southern Oscillation (ENSO) and other climatic events. At time scales greater than 1 year, ENSO dominates the sea level signal and can result in fluctuations in sea level of 10–15 cm. Alongshore wind stress data from central California are also analyzed to determine the impact of changes in coastal elevation at the mouth of San Francisco Bay within the synoptic wind band of 2–30 days. At least 40% of the subtidal fluctuations in sea level of the Bay are tied to the large-scale regional wind field affecting sea level variations in the coastal ocean, with little local, direct wind forcing of the Bay itself. The majority of the subtidal sea level fluctuations within the Bay that are not related to the coastal ocean sea level signal are forced by an east–west sea level gradient resulting from tidally induced variations in sea level at specific beat frequencies that are enhanced in the northern reach of the Bay. River discharge into the Bay through the Sacramento and San Joaquin River Delta also contributes to the east–west gradient, but to a lesser degree.     

Matching carbon pools and fluxes for the Southern Ocean Iron Release Experiment (SOIREE)

The Lagrangian Southern Ocean Iron Release Experiment (SOIREE) allowed study of a gradually evolving iron-mediated phytoplankton bloom in water labelled with the inert tracer sulfur hexafluoride, SF₆. This article describes a pelagic carbon budget for the mixed layer in SOIREE and assesses the extent to which closure of the budget is achieved. Net community production (NCP) converted 837 mmol m⁻² of inorganic carbon to organic carbon in 12.0 d after the first iron addition. A large fraction (41%) of NCP remained as particulate organic carbon in the mixed layer of the iron-enriched patch, while 23% was lost by horizontal dispersion and 0–29% was exported. The closure of the carbon budget is hampered by the lack of measurements of dissolved organic carbon (DOC), by a major uncertainty in carbon export, and by use of empirical conversion factors in estimates of carbon biomass and metabolic rates. Lagrangian carbon-budget studies may be improved by direct measurement of all major carbon parameters and conversion factors. Carbon cycling in the SOIREE bloom resembled that in ‘natural’ algal blooms in the open Southern Ocean in some respects, but not in all. Daily NCP in the SOIREE bloom (70 mmol m⁻² d⁻¹) was higher than in natural blooms, partly because other studies did not account for horizontal dispersion, were for longer periods or included less productive areas. The build-up of POC stock and carbon export as a fraction of NCP in SOIREE were in the lower range of observations elsewhere.     

Phosphate, nitrogen and iron enrichment in the polluted Izmir Bay, Aegean Sea.

Izmir Bay is one of the most polluted estuaries in the Mediterranean Sea. The extent of pollution due to anthropogenic inputs in the Izmir Bay is investigated on the basis of nutrients together with basic oceanographic parameters. Significant phosphate, nitrogen (N) and iron (Fe) enrichment has been found in the water of inner Izmir Bay. The situation of middle Izmir Bay is also similar but not as critical as in inner Izmir Bay, whereas outer Izmir Bay shows 'typical Mediterranean coastal marine ecosystem' characteristics. The average atomic ratios of apparent oxygen utilization (AOU) and nutrient fractions (N and phosphorous [P]) have been found as AOU:P (49:1), AOU:N (48:1) and N:P (5:1). A comparative evaluation of these ratios leads to the conclusion that they are too low; e.g. such a low AOU:P ratio supports fairly high P enrichment. Principal component analysis (PCA) indicates that 31% of the variation in the data is related to enrichment parameters, i.e. dissolved inorganic phosphate, NH4+, reactive Fe, 15% is related to NO3-, pH and 13% is related to temperature and total Fe. Additionally, PCA shows that the middle Izmir Bay has an ecological transient character between the inner and outer Izmir Bay.     

An in situ iron-enrichment experiment in the western subarctic Pacific (SEEDS): Introduction and summary

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.     

Plankton distribution associated with frontal zones in the vicinity of the Pribilof Islands

We studied the effect of four types of fronts, the coastal front, the middle front, the shelf partition front and the shelf break front on the quantitative distribution and the composition of plankton communities in the Pribilof area of the eastern Bering Sea shelf in late spring and summer of 1993 and 1994. The coastal fronts near St. Paul and St. George Islands and the coastal domains encircled by the fronts featured specific taxonomic composition of planktonic algae, high abundance and production of phytoplankton, as well as large numbers of heterotrophic nanoplankton. The coastal fronts also were characterized by high values of total mesozooplankton biomass, high concentrations of Calanus marshallae, as well as relatively high abundances of Parasagitta setosa and Euphausiacea compared to surrounding shelf waters. We hypothesize that wind-induced erosion of a weak thermocline in the inner part of the coastal front as well as transfrontal water exchange in subthermocline layers result in nutrient enrichment of the euphotic layer in the coastal fronts and coastal domains in summer time. This leads to prolonged high primary production and high phytoplankton biomass. In this paper a new type of front—the shelf partition front located 45–55 km to the north-east off St. Paul Island—is described, which is assumed to be formed by the flux of oceanic domain waters onto the shelf. This front features a high abundance of phytoplankton and a high level of primary production compared to the adjacent middle shelf. Near the southwestern periphery of the front a mesozooplankton peak occurred, composed of C. marshallae, with biomass in the subthermocline layer, reaching values typical for the shelf break front and the highest for the area. High abundance of phyto- and zooplankton as well as heterotrophic nanoplankton and elevated primary production were most often observed in the area adjacent to the shelf break front at its oceanic side. The phyto- and mesozooplankton peaks here were formed by oceanic community species. The summer levels of phytoplankton numbers, biomass and primary production in the shelf break frontal area were similar to those reported for the outer and middle shelf during the spring bloom and the coastal domains and coastal fronts in summer. In the environment with a narrow shelf to the south of St. George Island, the mesozooplankton peak was observed at the inner side of the shelf break front as close as 20 km from the island shore and was comprised of a “mixed” community of shelf and oceanic species. The biomass in the peak reached the highest values for the Pribilof area at 2.5 g mean wet weight m⁻³ in the 0–100 m layer. Details of the taxonomic composition and the numbers and production of phytoplankton hint at the similarity of processes that affect the phytoplankton summer community in the coastal domains of the islands, at the coastal fronts, and at the oceanic side of the shelf break front. The middle front was the only one that had no effect on plankton composition or its quantitative characteristics in June and July. Location of a variety of frontal productive areas within 100 km of the Pribilof Islands creates favorable foraging habitat for higher trophic level organisms, including sea birds and marine mammals, populating the islands.     

Submarine groundwater discharge: A large, previously unrecognized source of dissolved iron to the South Atlantic Ocean

This paper reports the initial results of a study of groundwater and coastal waters of southern Brazil adjacent to a 240 km barrier spit separating the Patos Lagoon, the largest coastal lagoon in South America, from the South Atlantic Ocean. The objective of this research is to assess the chemical alteration of freshwater and freshwater–seawater mixtures advecting through coastal permeable sands, and the influence of the submarine discharge of these fluids (SGD) on the chemistry of coastal waters. Here we focus on dissolved iron in this system and use radium isotopic tracers to quantify SGD and cross-shelf fluxes. Iron concentrations in groundwaters vary between 0.6 and 180 μM. The influence of the submarine discharge of these fluids into the surf zone produces dissolved Fe concentrations as high as several micromolar in coastal surface waters. The offshore gradient of dissolved Fe, coupled with results for Ra isotopes, is used to quantify the SGD flux of dissolved Fe from this coastline. We estimate the SGD flux to be 2 × 10⁶ mol day^− 1 and the cross-shelf flux to be 3.2 × 10⁵ mol day^− 1. This latter flux is equal to about 10% of the soluble atmospheric Fe flux to the entire South Atlantic Ocean. We speculate on the importance of this previously unrecognized iron input to regional ocean production and on the potential significance of this source to understanding variations in glacial–interglacial ocean production.     

Diel fluctuations in dissolved free amino acids and monosaccharides in Chesapeake Bay dinoflagellate blooms

The accumulation of phytoplankton biomass in recurring summer dinoflagellate blooms of Chesapeake Bay is accompanied by large pools of dissolved organic matter (DOM). Two fractions of the DOM, free amino acids (DFAA) and monosaccharides (MONO), were measured at 3 h intervals in mixed species dinoflagellate blooms (Katodinium rotundatum, Gymnodinium spp.) and related to productivity, biomass and photoperiod. Peak chlorophyll levels for the three blooms were 28, 65 and 938 μg1⁻¹. In general, DFAA and MONO concentrations increased with increasing biomass of bloom-forming species, reaching 203 and 844 μg1⁻¹. MONO appeared to accumulate during the day while there was no consistent pattern for DFAA. The accumulations of DFAA and MONO in blooms indicate that bloom production might stimulate microheterotrophy, thereby enhancing carbon and nutrient cycling in bloom-impacted regions.