Control of the phytoplankton response during the SAGE experiment: A synthesis

The SOLAS Air-Sea Gas Exchange (SAGE) experiment was conducted in Sub-Antarctic waters off the east coast of the South Island of New Zealand in the late summer of 2004. This mesoscale iron enrichment experiment was unique in that chlorophyll a (chl a) and primary productivity were only 2× OUT stations values toward the end of the experiment and this enhancement was due to increased activity of non-diatomaceous species. In addition, this enhancement in activity appeared to occur without a significant build up of particulate organic carbon. Picoeukaryotes (<2 ??m) were the only members of the phytoplankton assemblage that showed a statistically significant increase, a doubling in biomass. To better understand the controls of phytoplankton growth and biomass, we present results from a series of on-deck perturbation experiments conducted during SAGE. Results suggest that the pico-dominated phytoplankton assemblage was only weakly inhibited by iron. Diatoms with high growth rates comprised a small (<1%) fraction of the phytoplankton assemblage, were likely iron limited, and potentially further limited by silicic acid and therefore did not significantly contribute to bloom dynamics. On deck experiments and comparison of SAGE with other iron addition experiments suggested that neither light availability nor deep mixed layers limited phytoplankton growth. Although no substantial increase in grazing rate or specific phytoplankton growth rate was detected, microzooplankton biomass doubled over SAGE as a result of an increase in cell size. The importance of microzooplankton grazing was highlighted by the fact that they were capable of consuming 15-49% of the total phytoplankton production per day. Removal was highest on eukaryotic picophytoplankton production with a mean value of 72% (29-143%). Patch dilution played an important role during SAGE; the mean patch net algal growth:dilution rate, 1.13 (0.4-2.2) was the lowest reported for a mesoscale iron enrichment experiment. Phytoplankton biomass, estimated by chlorophyll a, only accumulated when phytoplankton growth exceeded grazing and when net algal growth exceeded dilution rate. The SAGE results highlight the function of the smallest phytoplankton size fraction described by the ecumenical Iron Hypothesis. Thus, adding iron to HNLC-low silicic acid regions during certain times of the year may simply transfer more carbon through the microbial food web. A primary implication of this study is that any iron-mediated gain in fixed carbon with this set of environmental conditions has a high probability of being recycled in surface waters.     

The SOLAS air-sea gas exchange experiment (SAGE) 2004

The SOLAS air-sea gas exchange experiment (SAGE) was a multiple-objective study investigating gas-transfer processes and the influence of iron fertilisation on biologically driven gas exchange in high-nitrate low-silicic acid low-chlorophyll (HNLSiLC) Sub-Antarctic waters characteristic of the expansive subpolar zone of the southern oceans. This paper provides a general introduction and summary of the main experimental findings. The release site was selected from a pre-voyage desktop study of environmental parameters to be in the south-west Bounty Trough (46.5°S 172.5°E) to the south-east of New Zealand and the experiment was conducted between mid-March and mid-April 2004. In common with other mesoscale iron addition experiments (FeAX’s), SAGE was designed as a Lagrangian study, quantifying key biological and physical drivers influencing the air-sea gas exchange processes of CO₂, DMS and other biogenic gases associated with an iron-induced phytoplankton bloom. A dual tracer SF₆/³He release enabled quantification of both the lateral evolution of a labelled volume (patch) of ocean and the air-sea tracer exchange at tenths of kilometer scale, in conjunction with the iron fertilisation. Estimates from the dual-tracer experiment found a quadratic dependency of the gas exchange coefficient on windspeed that is widely applicable and describe air-sea gas exchange in strong wind regimes. Within the patch, local and micrometeorological gas exchange process studies (100 m scale) and physical variables such as near-surface turbulence, temperature microstructure at the interface, wave properties and windspeed were quantified to further assist the development of gas exchange models for high-wind environments.There was a significant increase in the photosynthetic competence (F_v/F_m) of resident phytoplankton within the first day following iron addition, but in contrast to other FeAX’s, rates of net primary production and column-integrated chlorophyll a concentrations had only doubled relative to the unfertilised surrounding waters by the end of the experiment. After 15 days and four iron additions totalling 1.1 ton Fe²⁺, this was a very modest response compared to other mesoscale iron enrichment experiments. An investigation of the factors limiting bloom development considered co-limitation by light and other nutrients, the phytoplankton seed-stock and grazing regulation. Whilst incident light levels and the initial Si:N ratio were the lowest recorded in all FeAXs to date, there was only a small seed-stock of diatoms (less than 1% of biomass) and the main response to iron addition was by the picophytoplankton. A high rate of dilution of the fertilised patch relative to phytoplankton growth rate, the greater than expected depth of the surface mixed layer and microzooplankton grazing were all considered as factors that prevented significant biomass accumulation. In line with the limited response, the enhanced biological draw-down of pCO₂ was small and masked by a general increase in pCO₂ due to mixing with higher pCO₂ waters. The DMS precursor DMSP was kept in check through grazing activity and in contrast to most FeAX’s dissolved dimethylsulfide (DMS) concentration declined through the experiment. SAGE is an important low-end member in the range of responses to iron addition in FeAX’s. In the context of iron fertilisation as a geoengineering tool for atmospheric CO₂ removal, SAGE has clearly demonstrated that a significant proportion of the low iron ocean may not produce a phytoplankton bloom in response to iron addition.     

Community structure and photosynthetic physiology of phytoplankton in the northwest subarctic Pacific during an in situ iron fertilization experiment (SEEDS-II)

Temporal changes in the abundance, community composition, and photosynthetic physiology of phytoplankton in surface waters were investigated during the second in situ iron (Fe) fertilization experiment in the NW subarctic Pacific (SEEDS-II). Surface chlorophyll a concentration was 0.75 mg m⁻³ on the day before the first Fe enrichment (i.e. Day 0), increased ca. 3-fold until Day 13 after two Fe additions, and thereafter declined with time. The photochemical quantum efficiency (F_v/F_m) and functional absorption cross-section (σPSII) of photosystem II for total phytoplankton in surface waters increased and decreased inside the Fe-enriched patch through Day 13, respectively. These results indicate that the photosynthetic physiological condition of the phytoplankton improved after the Fe infusions. However, the maximum F_v/F_m value of 0.43 and the maximum quantum yield of carbon fixation (φ_max) of 0.041 mol C (mol photon)⁻¹ during the development phase of the bloom were rather low, compared to their theoretical maximum of ca. 0.65 and 0.10 mol C (mol photon)⁻¹, respectively. Diatoms, which were mainly composed of oceanic species, did not bloom, and autotrophic nanoflagellates such as cryptophytes and prasinophytes became predominant in the phytoplankton community inside the Fe-enriched patch. In ferredoxin/flavodoxin assays for micro-sized (20–200 μm in cell length) diatoms, ferredoxin was not detected but flavodoxin expressions consistently occurred with similar levels both inside and outside the Fe-enriched patch, indicating that the large-sized diatoms were stressed by Fe bioavailability inside the Fe-enriched patch even after the Fe enrichments. Our data suggest that the absence of a Fe-induced large-sized diatom bloom could be partly due to their Fe stress throughout SEEDS-II.     

Distribution and photo-physiological condition of phytoplankton in the tropical and subtropical North Pacific

To better understand the vertical distribution of phytoplankton in the tropical and subtropical North Pacific, we used fast repetition rate fluorometry to investigate the photo-physiological condition of the phytoplankton assemblage in this region between February and March 2007. Along 155°E, between the equator and 24°N, the peak of fluorescence (F _m), an indication of the deep chlorophyll maximum (DCM), was deeper than the top of the nitracline and occurred at the 2.4 ± 1.3 % (mean ± SD) light depth (relative to 0 m). The photochemical efficiency (F _v/F _m) and effective absorption cross-section of photosystem II (σ_PSII) were low at the surface but increased rapidly at depths between the top of the nitracline (40–138 m) and the DCM (70–158 m), an indication that the photo-physiological condition of the phytoplankton improved below the top of the nitracline. The depth of the maximal F _v/F _m [Z(F _v/F _{m max})] was 18–32 m deeper than the DCM and corresponded to the 0.8 ± 0.2 % light depth. The values of F _v/F _m at the Z(F _v/F _{m max}) were 20 % higher than those at the DCM and averaged 0.48 ± 0.01. These results suggest that the phytoplankton assemblage beneath the DCM had a high potential photosynthetic performance capacity and was growing by using the very low ambient light in this region.     

Did dilution limit the phytoplankton response to iron addition in HNLCLSi sub-Antarctic waters during the SAGE experiment?

An in situ iron addition experiment (SAGE) was carried out in high-nitrate low-chlorophyll low-silicic acid (HNLCLSi) sub-Antarctic surface waters south-east of New Zealand. In contrast to other iron addition experiments, the phytoplankton response was minor, with a doubling of biomass relative to surrounding waters, with the temporal trends in dissolved iron and macronutrients instead dominated by physical factors such as mixing and dilution. The initial increase in patch surface area indicated a lateral dilution rate of 0.125 d⁻¹, with a second estimate from a model of the decline in peak SF₆ concentration yielding a higher lateral dilution rate of 0.16-0.25 d⁻¹. The model was tested on the SOIREE SF₆ dataset and provided a lateral dilution of 0.07 d⁻¹, consistent with previous published estimates. MODIS ocean colour images showed elevated chlorophyll coincident with the SF₆ patch on day 10 and 12, and an elevated chlorophyll filament at the SAGE experiment location 3-4 days after ship departure, which provided additional lateral dilution estimates of 0.19 and 0.128 d⁻¹. Dissolved iron at the patch centre declined by 85% within two days of the initial infusion, of which dilution accounted for 50-65%; it also decreased rapidly after the 2nd and 3rd infusions but remained elevated after the fourth infusion. Despite decreases in nitrate and silicic acid from day 7 and 10, respectively, the final nutrient concentrations in the patch exceeded the initial concentrations due to supply from lateral intrusion and mixed-layer deepening. The low Si:N loss ratio suggested that the observed limited response to iron was primarily by non-siliceous phytoplankton. Algal growth rate exceeded the minimum dilution rate during two periods (days 3-6 and 10-14), and coincided with net chlorophyll accumulation. However, as the ratio of algal growth to dilution was the lowest reported for an iron addition experiment, dilution was clearly a significant factor in the SAGE experiment recording the lowest phytoplankton response to mesoscale iron addition.     

Enhanced carbohydrate production by Southern Ocean phytoplankton in response to in situ iron fertilization

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 (F_v/F_m), 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.     

Evidence for the grazing hypothesis: Grazing reduces phytoplankton responses of the HNLC ecosystem to iron enrichment in the western subarctic pacific (SEEDS II)

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.     

Bacterioplankton responses to iron enrichment during the SAGE experiment

We studied the microbial food web in the upper 100 m of the water column in iron-limited sub-Antarctic HNLC waters south-east of New Zealand in the SAGE experiment in 2004, with focus on bacterioplankton. Samples were collected daily from inside and outside the iron enriched patch. Short term enrichment experiments were conducted on board in 4 L polycarbonate bottles with water outside the iron enriched patch to study single and combined effects of micronutrient additions on microbial food web. Low bacterial growth was recorded in the study area with community turnover times of 50 h or more during the study period. Measurements of bacterial standing stocks and production rates in the study show minor responses to the large scale iron enrichment, with increase in rates and stocks after the first enrichment and at the end of the study period after the third iron enrichment when solar radiation increased and wind mixing decreased. The average daily bacterial production rates were 31.5 and 33.7 mgCm⁻² d⁻¹ for the OUT and IN stations, respectively; thus overall there was not a significant difference between the control and the iron-enriched patch. In the bottle experiments bacterial thymidine incorporation showed responses to single iron and silicic acid enrichments and a major growth response to the combined iron and sucrose enrichments. Phytoplankton chlorophyll-a showed clear stimulation by single additions of iron and silicic acid and silicic acid enhanced the iron impact. Cobalt additions had no effect on bacteria growth and a negative effect on phytoplankton growth. Low bacterial in situ growth rates and the enrichment experiments suggest that bacteria are co-limited by iron and carbon, and that bacterial iron uptake is dependent on carbon supply by the food web. With the high iron quota (??mol Fe mol C⁻¹) bacteria may scavenge considerable amounts of the excess iron, and thus influence the relative importance of the microbial food web as a carbon sink.     

Succession of phytoplankton functional groups from spring to early summer in the central Bohai Sea using HPLC–CHEMTAX approaches

Phytoplankton is a key component in the functioning of marine ecosystems, phytoplankton community structures are very sensitive to their environment. This study was conducted in the central Bohai Sea in the spring and early summer of 2015. Spatial variations in phytoplankton functional groups were examined through high-performance liquid chromatography pigment–CHEMTAX analysis. Results suggested that the phytoplankton biomass (chlorophyll a [Chl a]) in spring was mainly derived from the diatom community and was 3.5-fold higher than that in the summer. Meanwhile, the phytoplankton in the early summer sustained more diverse marker pigments than that in the spring. Despite the overwhelming predominance of microsized phytoplankton in the spring, some smaller phytoplankton (pico- or nanosized), including flagellates, such as prasinophytes, chlorophytes, and cryptophytes, highly contributed to the total Chl a in the summer. Various physico-chemical variables were recorded, and their correlations with phytoplankton density were established by redundancy analysis. Temperature, water stratification, nutrient availability, and even nutritive proportion influenced the succession of phytoplankton functional groups from diatom dominance in the spring to flagellate (mainly haptophytes and prasinophytes) dominance in the early summer. In conclusion, our work comprehensively evaluated the phytoplankton diversity and dynamics in the central Bohai Sea and suggests the need for long-term monitoring for further investigation.     

Iron,macronutrients and diatom blooms in the Peru upwelling regime: brown and blue waters of Peru

Surface water transects and vertical profiles for dissolved iron, macronutrients, chlorophyll a (Chl a), and hydrographic data were obtained in the Peru upwelling regime during August and September 2000. The supply of the micronutrient iron, relative to that of the macronutrients nitrate, phosphate and silicic acid, is shown to play a critical role in allowing extensive diatom blooms to develop in the Peru upwelling system. The extremely high-chlorophyll “brown waters of Peru” (with Chl a concentrations between 20 and 45 μg/l) result from massive diatom blooms with maximal photochemical efficiencies (F_v/F_m >0.6) occurring in the iron-rich upwelling region observed over the broad continental shelf off northern and central Peru. The source of the upwelled water in this region is the nutrient-rich subsurface countercurrent in contact with the organic-rich shelf sediments. This subsurface shelf water is suboxic and has extremely high concentrations of dissolved Fe (>50 nM) in the near-bottom waters. In marked contrast, relatively low-chlorophyll “blue waters” (Chl a <2 μg/l) with low concentrations of dissolved Fe (<0.1 nM) and high unutilized macronutrient concentrations are observed in the coastal upwelled waters along the southern coast of Peru and in the offshore regions of the Peru Current. Southern Peru is a region without a wide shelf to serve as a source of iron and, as a result, dissolved Fe concentrations in the near-bottom suboxic waters of this region are an order-of-magnitude lower than observed off northern and central Peru. In addition, the offshore Peru Current is a broad, Fe-limited, high-nitrate, lower than expected chlorophyll region extending hundreds of kilometers offshore into the northeast region of the South Pacific subtropical gyre and northwestward into the South Equatorial Pacific.     

Primary productivity,differential size fraction and pigment composition responses in two Southern Ocean in situ iron enrichments

Two in situ iron-enrichment experiments were conducted in the Pacific sector of the Southern Ocean during summer 2002 (SOFeX). The “north patch,” established within the Subantarctic Zone (∼56°S), was characterized by high nitrate (∼21 mmol m⁻³) but low silicic acid (2 mmol m⁻³) concentrations. North patch iron enrichment increased chlorophyll (Chl) by 12-fold to 2.1 mg m⁻³ and primary productivity (PP_EU) by 8-fold to 188 mmol C m⁻² d⁻¹. Surprisingly, despite low silicic acid concentrations, diagnostic pigment and size-fraction composition changes indicated an assemblage shift from prymnesiophytes toward diatoms. The “south patch,” poleward of the Southern Boundary of the Antarctic Circumpolar Current (SBACC) (∼66°S), had high concentrations of nitrate (∼27 mmol m⁻³) and silicic acid (64 mmol m⁻³). South patch iron enrichment increased Chl by 9-fold to 3.8 mg m⁻³ and PP_EU 5-fold to 161 mmol C m⁻² d⁻¹ but, notably, did not alter the phytoplankton assemblage from the initial composition of ∼50% diatoms. South patch iron addition also reduced total particulate organic carbon:Chl from ∼300 to 100; enhanced the presence of novel non-photosynthetic, but fluorescent, compounds; and counteracted a decrease in photosynthetic performance as photoperiod decreased. These experiments show unambiguously that in the contemporary, high nitrate Southern Ocean increasing iron supply increases primary productivity, confirming the initial premise of the Martin Iron Hypothesis. However, despite a 5-fold increase in PP_EU under iron-replete conditions in late summer, the effect of iron on annual productivity in the Southern Ocean poleward of the SBACC is limited by seasonal ice coverage and the dark of polar winter.     

Phytoplankton production characteristics in the southern Atlantic and the Atlantic sector of the Southern Ocean in the austral summer of 2009–2010

In December–January of 2010 the spatial distribution of the phytoplankton production characteristics was studied along transects in the vicinity of the Greenwich meridian (I) and in the Drake Passage (II). On transect I, the surface chlorophyll a concentration and primary production varied from 0.11 up to 3.57 mg/m³ and from 4.38 up to 37.47 mgC/m³ per day, respectively. The chlorophyll a in the photosynthetic layer and the integrated primary production varied from 10.7 up to 66.1 mg/m² and from 83 to 646 mgC/m² per day, respectively. On transect II in the surface layer, the chlorophyll a concentration changed within the range of 0.09–1.02 mg/m³ and the primary production ranged from 2.08 to 9.49 mgC/m³. The integrated values ranged from 6.32 to 38.29 mg/m² and from 41 to 221 mgC/m² per day, respectively. The moderate means of themaximum quantum yield (F _v/F _m) on transects I and II (0.41 and 0.35, respectively) testify to the low activity of the phytoplankton’s photosynthetic apparatus. The studied water areas in the Southern Ocean differed both in the phytoplankton biomass expressed in the chlorophyll a concentration values and in the conditions of the primary production formation.     

印尼爪哇上升流海域次表层叶绿素最大值层的浮游植物色素分布特征

Upwelling occurs on the coast of Java between June and October, forced by local alongshore winds associated with the southeasterly monsoon. This causes variations in phytoplankton community composition in the upwelling zone compared with the surrounding offshore area. Based on pigments analysis with subsequent calculations of group contributions to total chlorophyll a(Chl a) using CHEMTAX, we studied the distribution and composition of phytoplankton assemblages in the subsurface chlorophyll maximum along the south coast of Java and the influence of upwelling. Nineteen phytoplankton pigments were identified using high-performance liquid chromatography, and CHEMTAX analysis associated these to ten major phytoplankton groups. The phytoplankton community in the coastal area influenced by upwelling was characterized by high Chl a and fucoxanthin concentrations, indicating the dominance of diatoms. In contrast, in the offshore area, the Chl a and fucoxanthin concentrations declined to very low levels and the community was dominated by haptophytes represented by 19′-Hexanoyloxyfucoxanthin. Accordingly, microphytoplankton was found to be the major size class in the coastal area influenced by upwelling, while nanophytoplankton was most abundant in the offshore area. Low concentrations of other accessory pigments indicated less contribution from dinoflagellates,prasinophytes, chlorophytes and cryptophytes. Photo-pigment indices revealed that photosynthetic carotenoids(PSCs) were the largest component of the pigment pool, exceeding the proportion of Chl a, with the average PSCTP up to 0.62. These distribution trends can mainly be explained by phytoplankton adaption strategies to upwelling and subsurface conditions by changing species composition and adjusting the pigment pool.     

Grazing suppression of dimethylsulphoniopropionate (DMSP) accumulation in iron-fertilised, sub-Antarctic waters

The impact of in situ iron fertilisation on the production of particulate dimethylsulphoniopropionate (DMSPp) and its breakdown product dimethyl sulphide (DMS) was monitored during the SOLAS air-sea gas exchange experiment (SAGE). The experiment was conducted in the high nitrate, low chlorophyll (HNLC) waters of the sub-Antarctic Southern Ocean (46.7°S 172.5°E) to the south-east of New Zealand, during March-April, 2004. In addition to monitoring net changes in the standing stocks of DMSPp and DMS, a series of dilution experiments were used to determine the DMSPp production and consumption rates in relation to increased iron availability. In contrast to previous experiments in the Southern Ocean, DMS concentrations decreased over the course of the 15-d iron-fertilisation experiment, from an integrated volume-specific concentration in the mixed layer on day 0 of 0.78 nM (measured values 0.65-0.91 nM) to 0.46 nM (measured values 0.42-0.47 nM) by day 15, in parallel with the surrounding waters. DMSPp, chlorophyll a and the abundance of photosynthetic picoeukaryotes exhibited indiscernible or only moderate increases in response to the raised iron availability, despite an obvious physiological response by the phytoplankton. High specific growth rates of DMSPp, equivalent to 0.8-1.2 doublings d⁻¹, occurred at the simulated 60% light level of the dilution experiments. Despite the high production rates, DMSPp accumulation was suppressed in part by microzooplankton grazers who consumed between 61% d⁻¹ and 126% d⁻¹ of the DMSPp production. Temporal trends in the rates of production and consumption illustrated a close coupling between the DMSP-producing phytoplankton and their microzooplankton grazers. Similar grazing and production rates were observed for the eukaryotic picophytoplankton that dominated the phytoplankton biomass, partial evidence that picoeukaryotes contributed a substantial proportion of the DMSP synthesis. These rates for DMSPp and picoeukaryotes were considerably higher than for chlorophyll a, indicating higher cycling rates of the DMSP-producing taxa than for the bulk phytoplankton community. When compared to the total phytoplankton community, there was no evidence of selection against the DMSP-containing phytoplankton by the microzooplankton grazers; the opposite appeared to be the case. SAGE demonstrated that increased iron availability in the HNLC waters of the Southern Ocean does not invariably lead to enhanced DMS sea-air flux. The potential suppression of DMSPp accumulation by grazers needs to be taken into account in future attempts to elevate DMS emission through in situ iron fertilisation and in understanding the hypothesised link between levels of Aeolian iron deposition in the Southern Ocean, DMS emission and global albedo.     

Strong CO2 outgassing from high nutrient low chlorophyll coastal waters off central Chile (30°S): The role of dissolved iron

Carbonate system parameters (pH and alkalinity) were used to estimate the coastal water CO₂ fluxes off central Chile (30°S) during September 2007. Coastal waters rich in nitrate and silicate were strongly CO₂ supersaturated and normally poor in chlorophyll a. MODIS satellite chlorophyll a data suggest that phytoplankton biomass remained particularly low during September 2007 although coastal waters were highly fertilized with nitrate and silicate. The phytoplankton gross primary productivity in macronutrient-rich waters was very low with the exception of shallow waters (e.g. within or near bays). Several iron-enrichment bottle experiments show that fCO₂ rapidly decreases during iron-enrichment treatments compared to controls. This suggests that iron limitation of phytoplankton growth (mainly diatoms) plays a role in maintaining high-CO₂ outgassing by preventing rapid interception of upwelled CO₂.     

Chemotaxonomic investigation of phytoplankton in the shelf ecosystem of the KwaZulu-Natal Bight,South Africa

Phytoplankton community structure was elucidated during summer and winter in the KwaZulu-Natal Bight using pigment and CHEMTAX analyses. The surface pattern in January 2010 indicated that diatoms, haptophytes and prasinophytes tended to be the most prominent groups inshore and in the southern sector of the bight, whereas Prochlorococcus and Synechococcus were more dominant in the north and towards the offshore region. At the deep chlorophyll maximum (DCM), diatoms were dominant in the inner bight and in the south, with prasinophytes being the prominent flagellates, but this changed to Prochlorococcus and pelagophytes being the important groups in the northern sector and outer part of the bight. A different pattern was observed at the surface in July 2010, where diatoms, haptophytes, prasinophytes and cryptophytes in varying proportions comprised most of the community in the inner half of the bight, whereas Synechococcus and haptophytes were the main groups in the outer sector. A similar pattern occurred at the DCM, except that Synechococcus was less prominent and pelagophytes were distributed across the bight in both the inshore and offshore zones. Observations and relationships between phytoplankton groups and environmental parameters indicated that the groups were most closely related to temperature, accounting for 24–64% of the deviance. The influence of nutrients on phytoplankton was less clear but nitrate and silicate seemed to account for some of the patchy distribution patterns.     

Responses of phytoplankton and heterotrophic bacteria in the northwest subarctic Pacific to in situ iron fertilization as estimated by HPLC pigment analysis and flow cytometry

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.     

Contrasting oceanographic conditions and phytoplankton communities on the east and west coasts of Australia

The composition and dynamics of the phytoplankton communities and hydrographic factors that control them are described for eastern and western Australia with a focus on the Eastern Australian Current (EAC) and Leeuwin Current (LC) between 27.5° and 34.5°S latitude. A total of 1685 samples collected from 1996 to 2010 and analysed for pigments by high performance liquid chromatography (HPLC) showed the average TChla (monovinyl+divinyl chlorophyll a) concentration on the west coast to be 0.28±0.16 ??g L⁻¹ while it was 0.58±1.4 ??g L⁻¹ on the east coast. Both coasts showed significant decreases in the proportions of picoplankton and relatively more nanoplankton and microplankton with increasing latitude. On both coasts the phytoplankton biomass (by SeaWiFS) increased with the onset of winter. At higher latitudes (>27.5°S) the southeast coast developed a spring bloom (September) when the mean monthly, surface chlorophyll a (chla) concentration (by SeaWiFS) was 48% greater than on the south west coast. In this southern region (27.5-34.5°S) Synechococcus was the dominant taxon with 60% of the total biomass in the southeast (SE) and 43% in the southwest (SW). Both the SE and SW regions had similar proportions of haptophytes; ∼14% of the phytoplankton community. The SW coast had relatively more pelagophytes, prasinophytes, cryptophytes, chlorophytes and less bacillariophytes and dinophytes. These differences in phytoplankton biomass and community composition reflect the differences in seasonality of the 2 major boundary currents, the influence this has on the vertical stability of the water column and the average availability of nutrients in the euphotic zone. Seasonal variation in mixed layer depth and upwelling on the west coast appears to be suppressed by the Leeuwin Current. The long-term depth averaged (0-100 m) nitrate concentration on the west coast was only 14% of the average concentration on the east coast. Redfield ratios for NO₃:SiO₂:PO₄ were 6.5:11.9:1 on the east coast and 2.2:16.2:1 on the west coast. Thus new production (nitrate based) on the west coast was likely to be substantially more limited than on the eastcoast. Short term (hourly) rates of vertical mixing were greater on the east coast. The more stable water column on the west coast produced deeper subsurface chlorophyll a maxima with a 25% greater proportion of picoeukaryotes.     

A comparative study of maximal quantum yield of photosystem II to determine nitrogen and phosphorus limitation on two marine algae

We studied the effects of nitrogen and phosphorus supply on F_v/F_m (maximal quantum yield of photosystem II) in the diatom Chaetoceros debilis and dinoflagellate Scrippsiella trochoidea in nitrogen (N) and phosphorus (P) depleted cultures to determine whether this parameter could be used to monitor N or P limitation. In the nutrient depleted experiments, no obvious decrease of cell density and chlorophyll concentration was observed except in N-depleted incubation of S. trochoidea. For C. debilis, F_v/F_m decreased quickly in periods of N- and P-depletion and re-supply of N and P induced a quick recovery of F_v/F_m. However, in S. trochoidea culture, F_v/F_m remained unchanged in N- and P-depleted conditions and addition of sufficient N and P to N- and P-depleted cultures did not affect F_v/F_m. Therefore, F_v/F_m is insensitive to N and/or P limitation in growth of S. trochoidea. The results suggested that F_v/F_m was not a robust diagnostic for nutrient limitation in dinoflagellates. The differences in the sensitivity of F_v/F_m to nutrient limitation may result from different nutrient storage abilities among algal species.     

Spring and summer phytoplankton communities in the Chukchi and Eastern Beaufort Seas

Phytoplankton pigments and size-fractionated biomass in the Chukchi and Beaufort Seas showed spatial and temporal variation during the spring and summer of 2002. Cluster analysis of pigment ratios revealed different assemblages over the shelf, slope and basin regions. In spring, phytoplankton with particle sizes greater than 5 μm, identified as diatoms and/or haptophytes, dominated over the shelf. Smaller (<5 μm) phytoplankton containing chlorophyll b, most likely prasinophytes, were more abundant over the slope and basin. Due to extensive ice cover at this time, phytoplankton experienced low irradiance, but nutrients were near maximal for the year. By summer, small prasinophytes and larger haptophytes and diatoms co-dominated in near-surface assemblages in largely ice-free waters when nitrate was mostly depleted. Deeper in the water column at 1–15% of the surface irradiance larger sized diatoms were still abundant in the upper nutricline. Phytoplankton from the shelf appeared to be advected through Barrow Canyon to the adjacent basin, explaining similar composition between the two areas in spring and summer. Off-shelf advection was much less pronounced for other slope and basin areas, which are influenced by the low-nutrient Beaufort gyre circulation, leading to a dominance of smaller prasinophytes and chlorophytes. The correlation of large-sized fucoxanthin containing phytoplankton with the higher primary production measurements shows promise for trophic status to be estimated using accessory pigment ratios.